Rapid Sustainability Modeling for Raptors by Radiotagging and DNA-Fingerprinting

Abstract: Sustainable use of wildlife is crucial to ensuring persistence of natural resources. We used age-specific survival and breeding data to parameterize a demographic model for a harvested Kazakh saker falcon (Falco cherrug) population by radiotagging juveniles and estimating adult turnover with DNA-fingerprinting during 1993–1997. We gathered similar data during 1990–1998 to model populations of British buzzards (Buteo buteo), and during 1980–1998 to model populations of Swedish goshawks (Accipiter gentilis). Leg-bands and implanted microtransponders provided ways to test for bias and to estimate the harvest of sakers for falconry. Despite an estimated minimum first-year survival of only 23%, the observed productivity of 3.14 young per clutch would sustain a saker population (i.e., λ = 1) with a breeding rate (at laying) of only 0.63 for adults or with a residual juvenile yield of 37% if all adults breed. Higher first-year survival rates for goshawks and buzzards correlated with juvenile yields of up to 71%, but no more than half as many individuals if adults also were harvested. An annual population decline of 40% for sakers in southern Kazakhstan could be explained by observed productivity of only 0.71 young per clutch if there was also an estimated harvest of 55% of adults. This study shows that demographic models such as these can now be built rapidly if nestlings are fitted with reliable and safe radiotags and adult turnover is estimated from genetic analyses or other techniques.     

Optimizing Radio Retention and Minimizing Radio Impacts in a Field Study of Upland Sandpipers

Abstract: Two challenges in wildlife telemetry are optimizing the duration of transmitter attachment and minimizing the impacts of radios on the behavior and demography of the study animal. We tested 4 methods of radio attachment for a breeding population of upland sandpipers (Bartramia longicauda) under natural conditions at a tallgrass prairie site in Kansas, USA. To estimate radio retention and weekly survival rates, we used the nest survival model of Program MARK. Radio retention was lowest at the start and the end of the breeding period. The expected duration of radio retention was 1.8 years for a leg-loop harness, 40 days for radios glued to clipped feathers, 26 days for radios glued directly to feathers, and 7 days for radios glued to bare skin. Few radiomarked birds died during our study, but 4 of 8 mortality events were discovered within one week of radiomarking. Both glue and harnesses increased predation risk immediately after radio attachment. The weekly probability of survival was high after a 1-week acclimation period, and the expected survival for a 10-week breeding period was similar in males and females. Attachment of radios with glue had no effect on annual return rates. However, attachment of radios with leg harnesses resulted in lower return rates among radiomarked birds than birds without radios. Radios attached with glue were shed in <1 year but radios attached with harnesses were retained for up to 1-2 years. Our results indicate a tradeoff between optimizing radio retention and minimizing impacts on demography. Glue techniques had retention rates that were suitable for only short-term studies, but attachment with glue had no long-term effect on annual return rates. Leg harnesses provided effective radio retention that had little effect on survival rates during the stationary breeding period, but resulted in lower annual return rates. Robust estimates of radio retention and survival will assist researchers in selecting attachment techniques that best meet the study goals of future telemetry projects.     

A Bayesian estimate of harbour seal survival using sparse photo-identification data

Survival rates have rarely been estimated for pinniped populations due to the constraints of obtaining unbiased sample data. In this paper, we present an approach for estimating survival probabilities from individual recognition data in the form of photographic documentation of pelage patterns. This method was applied to estimate adult (age 2+) survival for harbour seals in the Moray Firth, NE Scotland. An astronomical telescope was used to obtain digital images of individual seals, and high-quality images were used to document the annual presence or absence of individuals at a single haul-out site over a 4-year period. A total of 95 females, 10 males and 57 individuals of unknown sex were photographically documented during the study period. Survival and recapture probabilities were estimated using Jolly–Seber mark–recapture models in a Bayesian statistical framework. Computer-intensive Markov Chain Monte Carlo methods were used to estimate the probability distributions for the survival and recapture probabilities, conveying the full extent of the uncertainty resulting from unavoidably sparse observational data. The deviance information criterion was used to identify a best-fitting model that accounted for variation in the probability of capture between sexes, with constant survival. The model estimated adult survival as 0.98 (95% probability interval of 0.94–1.00) using our photo-identification data alone, and 0.97 (0.92–0.99) with the use of an informative prior distribution based on previously published estimates of harbour seal survival. This paper represents the first survival estimate for harbour seals in the UK, and the first survival estimate using photo-identification data in any species of pinniped.     

On the sustainability of a reintroduced Crested Ibis population in Qinling Mountains,Shaanxi, Central China

Reintroduction projects aim to reestablish a self‐sustaining population of an endangered species within its historical range. Adequate post‐release monitoring by gathering demographic data is important to evaluate the success of a reintroduction. Survival and reproduction rates of a reintroduced population can be compared with a self‐sustaining wild population to evaluate the success of a reintroduction. In early 2007, Nipponia nippon (Crested Ibis) was reintroduced into the Qinling Mountains (Shaanxi, Central China). In this study, we attempt to evaluate the demographic status of the reintroduced population. Age‐specific survival rates of 56 released adults and 77 wild‐born fledglings were estimated using mark‐recapture data obtained from 2007 to 2014. Survival rates for the yearlings (0.599, with 95% confidence interval [CI]: 0.467–0.719) were lower than the estimates from a wild population in Yangxian County, but the survival rates of the adults (0.678, with 95% CI: 0.603–0.745) were similar. The number of breeding pairs gradually increased since 2008, although breeding success (52.5%) was somewhat less than that of the wild population (67.6%). The stochastic estimation of population growth rate (1.084 with 95% CI: 1.069–1.098) and population size (5‐fold increase) estimated from an age‐classified Leslie matrix indicate that the reintroduced population of the Crested Ibis is more likely in regulation phase over the next 25 years. We conclude that the reintroduction of the Crested Ibis in Qinling Mountains has great promise, and progress toward a self‐sustaining population has been made under some interventions. Governments, local communities, and scientists need to facilitate habitat restoration for the long‐term survival of this endangered species.     

Factors affecting duckling survival of Eiders Somateria mollissima in northeast Scotland

Mortality factors affecting Eider ducklings Somateria mollissima and their impact with respect to age were studied on the Ythan estuary, Aberdeenshire, Scotland. Survival from hatching to fledging averaged 10.4% over 13 years; numbers declined steeply during the first three to four weeks and slowly thereafter. Growth of tagged young varied considerably between broods. Mortality was not correlated with numbers of ducklings or adults.
Predation in 1974 averaged 13.6% per day during the first two weeks, and accounted for most mortality. Herring Gulls Larus argentatus were the principal predators. Predation was 2.5 times as high in rainy, windy weather as in fine, calm weather. Weather, primarily mean rainfall and wind speed, during early life accounted for over 90% of all variation in survival during eight of nine years. Predation was considered the proximate factor in most weather-related mortality. High survival in 1971 was not explained by weather. An estimated 25–40% of mortality during the second week of life was from renal coccidiosis Eimeria somateriae. Reproductive output of the Ythan population probably limits the rate of population growth, but numbers of breeding birds seem to be regulated by other factors acting after fledging.     

SURVIVAL ESTIMATES FOR FLORIDA MANATEES FROM THE PHOTO-IDENTIFICATION OF INDIVIDUALS

We estimated adult survival probabilities for the endangered Florida manatee ( Trichechus manatus latirostris ) in four regional populations using photoidentification data and open-population capture-recapture statistical models. The mean annual adult survival probability over the most recent 10-yr period of available estimates was as follows: Northwest - 0.956 (SE 0.007), Upper St. Johns River - 0.960 (0.011), Atlantic Coast - 0.937 (0.008), and Southwest - 0.908 (0.019). Estimates of temporal variance independent of sampling error, calculated from the survival estimates, indicated constant survival in the Upper St. Johns River, true temporal variability in the Northwest and Atlantic Coast, and large sampling variability obscuring estimates for the Southwest. Calf and subadult survival probabilities were estimated for the Upper St. Johns River from the only available data for known-aged individuals: 0.810 (95% CI 0.727–0.873) for 1st year calves, 0.915 (0.827–0.960) for 2nd year calves, and 0.969 (0.946–0.982) for manatee 3 yr or older. These estimates of survival probabilities and temporal variance, in conjunction with estimates of reproduction probabilities from photoidentification data can be used to model manatee population dynamics, estimate population growth rates, and provide an integrated measure of regional status.     

Behaviour in the post-nestling dependence period of radio-tagged Common Buzzards Buteo buteo

Behaviour of 26 young Common Buzzards Buteo buteo was studied by systematic radiotracking during July and August 1991. After hatching between 11 May and 18 June, the young buzzards fledged when they were 43–54 days old. Distances travelled from the nest increased abruptly after birds were 65 days old, when their flight feathers had completed growth: buzzards were located more than 500 m from the nest in only 2% of records within 65 days of hatching but in 26% of records when they were older. Before their 65th day, there was an increase with age in distance from the nest, time spent flying and time spent calling, especially for buzzards with continuous woodland around the nest. The increases in distance and areas covered were greatest for broods where parents were most often present, which was at nests with the most grass and arable farmland nearby. Between their 65th and 100th days, buzzards showed no increase in activity with age and called less, especially where there was extensive woodland, and travelled farthest from nests with least neighbouring grassland; broods with few young most often had parents nearby. Young buzzards associated strongly with each other between leaving the nest and completing feather growth, but some broods later became much less cohesive. Variation in activity was not linked to sex or to the presence of 30 g back-pack radio-tags compared with 12-g leg-mounted radio-tags.     

Density, survival and dispersal of Anopheles gambiae complex mosquitoes in a West African Sudan savanna village

Abstract. To obtain information on adult populations of Afrotropical malaria vector mosquitoes, mark-release-recapture experiments were performed with Anopheles females collected from indoor resting-sites in a savanna area near Ouagadougou, Burkina Faso, during September 1991 and 1992. Results were used to estimate the absolute population densities, daily survival rates, and dispersal parameters of malaria vectors in that area.
In 1991 a total of 7260 female Anopheles were marked and released, of which 106 were recaptured in the release village and 6 in the neighbouring villages, a total recapture rate of 1.5%. The following year 13, 854 female Anopheles were released and 116 recaptured in Goundri and 8 in the neighbouring villages, a total recapture rate of 0.9%. Recaptures were found in three of eight villages near Goundri. Nearly all of the recaptured mosquitoes were An.gambiae s.l. Of these, molecular determination revealed that An.gambiae s.s. and An.arabiensis were present in a ratio of -2:3.
Two simple random models of dispersal were simulated and the parameters of the models determined by searching for the least-squared fit between simulated and observed distributions. The mean distance moved by individual mosquitoes, estimated in this way, ranged 350–650 m day^-1, depending on die model and the year considered. Population densities were estimated using the Lincoln Index, Fisher-Ford and Jolly's methods. The estimates of population size had high standard errors and were not particularly consistent. A 'consensus' value of 150,000–350,000 mosquitoes is believed to apply for ht An.gambiae s.l. female population. Survival was estimated to be 80–88% per day.     

A STAGE-BASED MODEL OF MANATEE POPULATION DYNAMICS

A stage-structured population model for the Florida manatee ( Trichechus manatus latirostris ) was developed that explicitly incorporates uncertainty in parameter estimates. The growth rates calculated with this model reflect the status of the regional populations over the most recent 10-yr period. The Northwest and Upper St. Johns River regions have growth rates (λ) of 1.037 (95% interval, 1.016–1.056) and 1.062 (1.037–1.081), respectively. The Southwest region has a growth rate of 0.989 (0.946–1.024), suggesting this population has been declining at about 1.1% per year. The estimated growth rate in the Atlantic region is 1.010 (0.988–1.029), but there is some uncertainty about whether adult survival rates have been constant over the last 10 yr; using the mean survival rates from the most recent 5-yr period, the estimated growth rate in this region is 0.970 (0.938–0.998). Elasticity analysis indicates that the most effective management actions should seek to increase adult survival rates. Decomposition of the uncertainty in the growth rates indicates that uncertainty about population status can best be reduced through increased monitoring of adult survival rate.     

Estimating survival of the shortfin mako Isurus oxyrinchus (Rafinesque) in the north-west Atlantic from tag-recapture data

Survival was estimated for shortfin mako Isurus oxyrinchus in the north-west Atlantic from tag-recapture data. The data used in this study were collected by the National Marine Fisheries Service Cooperative Shark Tagging Programme from 1962 to 2003. In total, 6309 shortfin mako sharks were tagged, of which 730 were recaptured. The high recapture rate of 11·6% for this species provided adequate recapture data to carry out survival analyses. Estimates of survival were generated with the computer software MARK, which provided a means for estimating parameters from tagged animals when they were recaptured. The results of several models are presented with various combinations of constant and time-specific survival and recovery rates. A parametric bootstrap and the median variance inflation factor ( ) approach were used to test the fit of the general model to the data. The estimated indicated a very good model fit. The models with time invariant survival rate had the most support from the data and no group or time period effects were found. Recovery rate ( f ) appeared to increase from 0·043 in the early years to 0·056 in the later years. The nominal survival rate of 0·59 year⁻¹ was adjusted with an estimated tag-shedding rate of 0·26 year⁻¹ to generate a final corrected annual survival estimate of 0·79 with a 95% CI of 0·71–0·87.     

Winter site-fidelity and survival of Redshank Tringa totanus at Cardiff,south Wales

The site-fidelity of adult Redshank Tringa totanus was investigated using colour-ringing and radiotracking at Cardiff Bay, a wintering site in south Wales, in 1996/97 and 1997/98. Between 7 and 20% of colour-ringed adults were seen away from the bay in winter, although only at a distance of 4 km. Radiotracking also suggested that adults were largely faithful either to the bay or to this neighbouring site. Return rates of 89% and 83% were estimated over two successive summers, indicating site-fidelity between winters. Survival rates of 91% and 100% were estimated over the two preceding (four-month) winter periods. Annual survival rates, calculated as the product of the winter survival and over-summer return rates, were therefore 81% and 83%. These estimates are comparable to those previously obtained for Redshank, in studies both on breeding and wintering grounds. Analyses of ring recovery data from throughout the UK complemented the colour-ringing and radiotracking studies in showing that the majority of Redshank are site-faithful both within and between winters. Ring recovery analyses also indicated that first-year Redshank had similar recovery patterns to adults.     

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.     

Sex-biased survival and breeding dispersal probability in a patchy population of the Rock Sparrow Petronia petronia

Demographic parameters of the polygynous Rock Sparrow Petronia petronia were investigated in a small patchy population in the Italian Alps. The population included two distinct breeding patches that differed in altitude and breeding success. Survival parameters were estimated by capture–recapture analysis of 170 individually marked animals. At the whole population level (Cormack–Jolly Seber model), no sex difference in local survival probability was detected. We then used a multisite capture–recapture approach (Arnason–Schwarz model) to investigate patch-specific survival probability and between-patch dispersal rate conditional on survival. Female local survival in the higher-altitude patch (mean ± se: 0.54 ± 0.04) was significantly greater than in the other patch (0.37 ± 0.04), probably because permanent emigration from the study area was greater. In the higher-altitude patch, breeding dispersal was constrained by the altitude limit and breeding movements were directed toward the patch at lower altitude. The probability of changing patch in the next breeding season was significantly higher for females (range 0.16–0.21) than for males (0.01–0.03). Breeding success varied between years and patches, being lower in the patch where frequency of polygamy and female local mortality were higher.     

Black Brant Harvest,Density Dependence,and Survival: A Record of Population Dynamics

ABSTRACT We analyzed 53 years of banding and band recovery data along with estimates of harvest and population size to assess the role of harvest and density dependence in survival patterns and population dynamics of black brant (Branta bernicla nigricans) over the period 1950–2003. The black brant population has declined steadily since complete annual surveys began in 1960, so the role of harvest in the dynamics of this population is of considerable interest. We used Brownie models implemented in Program MARK to analyze banding data. In some models, we incorporated estimated sport harvest to test hypotheses about the role of harvest in survival. We also examined the hypothesis of density-dependent regulation of mortality by incorporating estimates of population size as a covariate into models of survival. For a shorter period (1985–2003), we also assessed hypotheses about the role of subsistence harvest and predation as sources of mortality. The best supported model of variation in survival and band recovery allowed survival rates to vary among 2 age classes (juv, second-yr plus ad brant) and the 2 sexes. We constrained survival probabilities to be constant within decades but allowed them to vary among decades. We also constrained band recovery rates to be constant within decades and to vary in parallel among age and sex classes. We were limited to decade-specific estimates of survival and band recovery rates because some years before 1984 lacked any banding, and banding in some other years was sparse. A competitive model constrained survival estimates to be the same for males and females. No model containing harvest or population size was competitive with models lacking these covariates (relative quasi-Akaike's Information Criterion adjusted for small sample size [βQAIC_c] > 13). In the best supported model, band recovery rates declined from 0.038 ± 0.0028 (F) and 0.040 ± 0.0031 (M) to 0.007 ± 0.0007 (F) and 0.007 ± 0.0007 (M) between the 1950s and 2000s, a clear indication that harvest rates declined over this period. Survival rates increased from 0.70 ± 0.02 and 0.71 ± 0.02 for adult males and females, respectively, in the 1950s to 0.88 ± 0.009 and 0.88 ± 0.01 for males and females, respectively, in the 1990s. Survival rates in the 1990s were among the highest estimated for brant and did not increase in the 2000s with additional reductions in sport harvest. For the shorter data set from 1985 to 2003, models containing covariates for either sport or subsistence harvest were less competitive than models lacking these terms (βQAIC_c > 3). For the best model containing subsistence harvest, the estimate of β linking subsistence harvest to survival, although imprecisely estimated, was near zero (β = −0.04 ± 0.30), consistent with the hypothesis that subsistence harvest had little impact on survival during this period. We conclude that while harvest likely influenced survival and population dynamics in earlier decades, it is most likely that continued population decline at least since 1990 is a result of low recruitment.     

Adult survival and numbers in a coastal breeding population of Redshank Tringa totanus in northwest England

The results of a long-term capture-mark-recapture ringing programme carried out on a coastal population of breeding Redshanks Tringa totanus between 1974 and 1988 are presented.
Both sexes were equally likely to be recaptured in subsequent years, as were birds captured for the first time compared with those that had been captured previously. Older birds were more frequently recaptured than were young birds. There was no significant difference in male and female adult survival rates, with a mean of 72% of females and 75% of males surviving each year.
The breeding population fluctuated annually with estimated breeding densities of 122–285 pairs per km². Variation in breeding numbers (males), but not survival, was partially attributable to winter air temperature.     

Abundance of Snowy and Wilson's Plovers in the lower Laguna Madre region of Texas

ABSTRACT.   Snowy Plovers ( Charadrius alexandrinus ) and Wilson's Plovers ( C. wilsonia ) are shorebird species of increasing conservation concern, with populations apparently declining in North America. However, estimates of current populations are needed before initiating long-term monitoring or planning. In 2004, we estimated abundance of breeding Snowy and Wilson's plovers in the lower Laguna Madre region of Texas using occupancy abundance estimation. We made repeated visits to survey plots from April to June, recording the number of adults of both species observed and the amount of suitable breeding habitat within each plot. We considered Bayesian occupancy abundance models with and without habitat covariates to explain the abundance of both species. For both Snowy and Wilson's plovers, the number of birds counted in each plot was influenced by the amount of suitable breeding habitat within the plot (Snowy Plover α_habitat= 0.52, SD = 0.10, 95% CI = 0.33–0.71; Wilson's Plover α_habitat= 0.48, SD = 0.12, 95% CI = 0.24–0.71). Using the habitat covariate models for each species, we estimated that 416 adult Snowy Plovers (95% CI = 394–438) and 279 adult Wilson's Plovers (95% CI = 262–296) were present in our study area. Our results illustrate the use of a relatively new method for abundance estimation, and indicate that the lower Laguna Madre region of Texas is an important breeding area for both Snowy and Wilson's plovers. Given the documented and suspected population declines for Snowy and Wilson's plovers, we recommend protection of their breeding habitats along the coast of Texas from development and degradation resulting from unregulated use.     

Climate and density influence annual survival and movement in a migratory songbird

Assessing the drivers of survival across the annual cycle is important for understanding when and how population limitation occurs in migratory animals. Density‐dependent population regulation can occur during breeding and nonbreeding periods, and large‐scale climate cycles can also affect survival throughout the annual cycle via their effects on local weather and vegetation productivity. Most studies of survival use mark–recapture techniques to estimate apparent survival, but true survival rates remain obscured due to unknown rates of permanent emigration. This is especially problematic when assessing annual survival of migratory birds, whose movement between breeding attempts, or breeding dispersal, can be substantial. We used a multistate approach to examine drivers of annual survival and one component of breeding dispersal (habitat‐specific movements) in a population of American redstarts (Setophaga ruticilla) over 11 years in two adjacent habitat types. Annual survival displayed a curvilinear relation to the Southern Oscillation Index, with lower survival during La Niña and El Niño conditions. Although redstart density had no impact on survival, habitat‐specific density influenced local movements between habitat types, with redstarts being less likely to disperse from their previous year's breeding habitat as density within that habitat increased. This finding was strongest in males and may be explained by conspecific attraction influencing settlement decisions. Survival was lowest in young males, but movement was highest in this group, indicating that apparent survival rates were likely biased low due to permanent emigration. Our findings demonstrate the utility of examining breeding dispersal in mark–recapture studies and complement recent work using spatially explicit models of dispersal probability to obtain greater accuracy in survival estimates.     

Estimating demographic parameters for loggerhead sea turtles using mark–recapture data and a multistate model

The survival for adult loggerhead sea turtles from a saturation tagging study on Bald Head Island, NC, USA, was estimated using a multistate model with unobservable states to relax assumptions that are violated when survival is estimated from multistate models and produce more accurate estimates of survival, recapture, and breeding transition probabilities. The influence of time, trap dependence, and low site fidelity to the study nesting beach on survival and recapture were examined. The best model given the data included an imprecise site-fidelity effect on survival, constrained the reproductive cycle to 4 years, and contained a time effect on recapture rates. The estimate of annual survival for adult females was of 0.85, producing the highest estimate in the literature for loggerhead sea turtles. Multistate models should be applied to other nesting beach data for sea turtles to improve survival estimates and in turn the ability to model and manage populations.     

Negative population trend for Chapada Flycatchers (Suiriri islerorum) despite high apparent annual survival

ABSTRACT Population size and trends are demographic parameters that can be used to help determine the threat of extirpation or extinction of bird populations and in determining management strategies. Chapada Flycatchers (Suiriri islerorum) are endemic to the Cerrado region of Brazil, dependent on open cerrado habitat, defend large territories, and their populations seem to be declining. From 2003 to 2007, we analyzed demographic parameters of a declining population of Chapada Flycatchers, using the rate at which territories became vacant and the size of the breeding population to ascertain the status of the population, and evaluated the relative contributions of apparent annual survival and recruitment rates to these trends. Territories became vacant at a mean rate of 13% per year, and 32% of all territories were vacated during our study. The current breeding population (20 pairs) was at least seven times smaller than the estimated carrying capacity of the reserve (141 territorial pairs). Estimated apparent annual survival probabilities (0.77 for breeders; 0.67 for nonbreeders) based on model averaging were comparable to those reported for other Neotropical passerines. Survival rates did not differ between the sexes and the estimated recruitment rate was 0.21. In many species, adult survival is the factor that most strongly influences population growth rates. However, the population of Chapada Flycatchers we studied is declining despite high annual survival rates, with low and variable breeding success probably causing the decline. Our results improve our understanding of the possible role of adult survival and breeding success in the decline of populations of small passerines in isolated reserves in the tropics.     

Biases in the estimation of the demographic parameters of a snowshoe hare population

1. Survival rates and natalities for a population of snowshoe hares in the Yukon were estimated independently of and simultaneously with estimates of population change during the increase phase of a hare cycle.
2. Simple demographic models are used to show that even though the estimated survival rates and natalities were high relative to previously published estimates, the observed demographic parameters are unable to explain the extent of population increase, and we conclude that some of these parameters must be underestimates.
3. A sensitivity analysis is used to examine the potential influence of changes in these demographic parameters on the population growth rate. During most years of the hare cycle the population growth rate is potentially most sensitive to changes in juvenile postweaning survival. Only during crash years is adult survivorship likely to be a more important determinant of the rate of population change.
4. Examination of previously published data sets on two full population cycles suggests that while survival rates are positively correlated with population growth rates, their incorporation into demographic models results in frequent underestimation of the rate of population increase.