In organisms with complex life cycles, physiological stressors during early life stages may have fitness-level impacts that are delayed into later stages or habitats. We tested the hypothesis that body size and date of metamorphosis, which are highly responsive to aquatic stressors, influence post-metamorphic survival and movement patterns in the terrestrial phase of an ephemeral pond-breeding frog by examining these traits in two populations of northern red-legged frogs (Rana aurora aurora). To increase variation of body size at metamorphosis, we manipulated food availability for 314 of 1045 uniquely marked tadpoles and estimated the probability that frogs survived and emigrated using concentric rings of drift fencing surrounding ponds and Bayesian capture-recapture modeling. The odds of surviving and emigrating from the ponds to the innermost drift fences, approximately 12 m, increased by factors of 2.20 (95% credibility intervals 1.39-4.23) and 2.54 (0.94-4.91) with each millimeter increase in snout-vent length and decreased by factors of 0.91 (0.85-0.96) and 0.89 (0.80-1.00) with each day's delay in metamorphosis for the two ponds. The odds of surviving and moving to the next ring of fencing, 12 m to approximately 40 m from the ponds, increased by a factor of 1.20 (0.45-4.06) with each millimeter increase in size. Our results demonstrated that body size and timing of metamorphosis relate strongly to the performance of newly metamorphosed frogs during their initial transition into terrestrial habitat. Carryover effects of aquatic stressors that reduce size and delay metamorphosis may have population-level impacts that are not expressed until terrestrial stages. Since changes in both aquatic and terrestrial systems are implicated in many amphibian declines, quantifying both immediate and delayed effects of stressors on demographic rates is critical to sound management.

Batrachochytrium dendrobatidis is a fungal pathogen that is receiving attention around the world for its role in amphibian declines. Study of its occurrence patterns is hampered by false negatives: the failure to detect the pathogen when it is present. Occupancy models are a useful but currently underutilized tool for analyzing detection data when the probability of detecting a species is <1. We use occupancy models to evaluate hypotheses concerning the occurrence and prevalence of B. dendrobatidis and discuss how this application differs from a conventional occupancy approach. We found that the probability of detecting the pathogen, conditional on presence of the pathogen in the anuran population, was related to amphibian development stage, day of the year, elevation, and human activities. Batrachochytrium dendrobatidis was found throughout our study area but was only estimated to occur in 53.4% of 78 populations of native amphibians and 66.4% of 40 populations of nonnative Rana catesbeiana tested. We found little evidence to support any spatial hypotheses concerning the probability that the pathogen occurs in a population, but did find evidence of some taxonomic variation. We discuss the interpretation of occupancy model parameters, when, unlike a conventional occupancy application, the number of potential samples or observations is finite.

Summary 1.We estimated age-specific probability of breeding for black brant ( Branta bernicla nigricans Lawrence) (hereafter brant) at the Tutakoke River Colony, Alaska, during a series of years in which local breeding density and juvenile mortality both increased. We use these analyses to examine demographic responses to increased population density. 2. Estimates of breeding probability were based on Kendall & Nichols (1995) fulllikelihood modification of Pollock's (1982) robust design applied to observations of uniquely colour-banded brant. 3. Breeding probability varied with female age but not across years, indicating lack of response to increased population density. Annual probability of breeding increased from 0·67 ± 0·042 for 2-year-old females to 0·90 ± 0·024 for females 5 years old or older. 4. Annual breeding probability of males ≥ 5 years old was 0·78 ± 0·03. Lower breeding probability of males compared to females probably reflects dispersal of males whose mates died. 5.Lower juvenile survival appears to be a general response to increased density in large herbivorous birds and mammals, while changes in adult survival and reproductive investment may be more variable.

Summary 1.Patterns of temporary emigration (associated with non-breeding) are important components of variation in individual quality. Permanent emigration from the natal area has important implications for both individual fitness and local population dynamics. 2. We estimated both permanent and temporary emigration of black brent geese ( Branta bernicla nigricans Lawrence) from the Tutakoke River colony, using observations of marked brent geese on breeding and wintering areas, and recoveries of ringed individuals by hunters. We used the likelihood developed by Lindberg, Kendall, Hines & Anderson 2001 (Combining band recovery data and Pollock's robust design to model temporary and permanent emigration. Biometrics , 57 , 273-281) to assess hypotheses and estimate parameters. 3. Temporary emigration (the converse of breeding) varied among age classes up to age 5, and differed between individuals that bred in the previous years vs. those that did not. Consistent with the hypothesis of variation in individual quality, individuals with a higher probability of breeding in one year also had a higher probability of breeding the next year. 4. Natal fidelity of females ranged from 0·70 ± 0·07-0·96 ± 0·18 and averaged 0·83. In contrast to Lindberg et al . (1998), we did not detect a relationship between fidelity and local population density. Natal fidelity was negatively correlated with first-year survival, suggesting that competition among individuals of the same age for breeding territories influenced dispersal. Once females nested at the Tutakoke River, colony breeding fidelity was 1·0. 5. Our analyses show substantial variation in individual quality associated with fitness, which other analyses suggest is strongly influenced by early environment. Our analyses also suggest substantial interchange among breeding colonies of brent geese, as first shown by Lindberg et al . (1998).