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Estimating transition probabilities for stage-based population projection matrices using capture-recapture data

In stage—based demography, animals are often categorized into size (or mass) classes, and size—based probabilities of surviving and changing mass classes must be estimated before demographic analyses can be conducted. In this paper, we develop two procedures for the estimation of mass transition probabilities from capture—recapture data. The first approach uses a multistate capture—recapture model that is parameterized directly with the transition probabilities of interest. Maximum likelihood estimates are then obtained numerically using program SURVIV. The second approach involvesa modification of Pollock's robust design. Estimation proceeds by conditioning on animals caught in a particualr class at time i, and then using closed models to estimate the number of these that are alive in other classes at i + 1. Both methods are illustrated by application to meadow vole, Microtus pennsylvanicus, capture—recapture data. The two methods produced reasonable estimates that were similar. Advantages of these two approaches include the directness of estimation, the absence of need for restrictive assumptions about the independence of survival and growth, the testability of assumptions, and the testability of related hypotheses of ecological interest (e.g., the hypothesis of temporal variation in transition probabilities).

Ecology↗

The Husting dilemma: A methodological note

Recently, Gill (1985) discussed the interpretation of capture history data resulting from his own studies on the red-spotted newt, Notophthalmus viridescens , and work by Husting (1965) on spotted salamanders, Ambystoma maculatum. Gill (1985) noted that gaps in capture histories (years in which individuals were not captured, preceded and followed by years in which they were) could result from either of two very different possibilities: (1) failure of the animal to return to the fenced pond to breed (the alternative Husting (1965) favored), or (2) return of the animal to the breeding pond, but failure of the investigator to capture it and detect its presence. The authors agree entirely with Gill (1985) that capture history data such as his or those of Husting (1965) should be analyzed using models that recognize the possibility of 'census error,' and that it is important to try to distinguish between such 'error' and skipped breeding efforts. The purpose of this note is to point out the relationship between Gill's (1985:347) null model and certain capture-recapture models, and to use capture-recapture models and tests to analyze the original data of Husting (1965).

Ecology↗

Geomys bursars burrowing patterns: influence of season and food patch structure

The relationship between belowground food resources and the pattern and pace of soil excavation by Geomys bursarius, the plains pocket gopher, was assessed during spring and autumn using replicated 18 x 18 m enclosures containing uniform arrays of food plants (Daucus carota) at spacings assumed to result in burrowing being energetically profitable (25 and 50 cm) or unprofitable (100 cm and empty). Soil excavation rate, measured in terms of either daily additions to tunnel length or volume, or the maximum volume excavated during a 3—d interval, varied greatly among individuals. No significant effect due to season or density of belowground plant biomass was detected when burrowing was profitable. Individuals readily initiated aboveground travel in unprofitable treatments. Soil bulk density was negatively related and animal body mass positively related to excavation rate. Individuals deposited from 41 to 87% of excavated soil on the ground surface; the remainder was used to backfill previously created tunnels or chambers. Plant size, but not plant spacing, affected the ratio of the volumes deposited aboveground and belowground. There was no evidence that animals in enclosures disturbed soil in a manner quantitatively or qualitatively different from free—ranging adults. The large proportions and absolute quantities (up to 30 L/d) of soil involved suggest that backfilling may affect ecosystem structure and functioning as much as surface mound—building.

Ecology↗

Assessing habitat selection when availability changes

We present a method of comparing data on habitat use and availability that allows availability to differ among observations. This method is applicable when habitats change over time and when animals are unable to move throughout a predetermined study area between observations. We used maximum—likelihood techniques to derive an index that estimates the probability that each habitat type would be used if all were equally available. We also demonstrate how these indices can be used to compare relative use of available habitats, assign them ranks, and assess statistical differences between pairs of indices. The set of these indices for all habitats can be compared between groups of animals that represent different seasons, sex or age classes, or experimental treatments. This method allows quantitative comparisons among types and is not affected by arbitrary decisions about which habitats to include in the study. We provide an example by comparing the availability of four categories of sea ice concentration to their use by adult female polar bears (Ursus maritimus), whose movements were monitored by satellite radio tracking in the Bering and Chukchi Seas during 1990. Use of ice categories by bears was nonrandom, and the pattern of use differed between spring and late summer seasons.

Ecology↗

The abundance and diversity of the herpetofaunas of tropical forest litters

Quantitative and qualitative samples of amphibian and lizard faunas were taken from forest litter in Costa Rica and compared with similar samples collected in Borneo, the Philippines, and Panama. Animal abundance is about ten times greater in Costa Rican lowland wet forest than in Borneo. Radically different routes and rates of energy flow are postulated to account for the difference. In a series of upland (1010-1425 m) samples in the Philippines, the number of litter lizards and frogs increases with elevation. A single upland (1200 m) Costa Rican sample contains about three times as many animals per 100 m2 as two lowland sites. This increase with altitude correlates well with hypotheses that overall tropical forest productivity is greatest at intermediate elevations. Herpetofaunal densities are greater in wet areas compared to dry sites and flat terrain compared to slopes. These observations are linked to the greater variation and lesser total litter fall in dry sites and/or slopes. The number of species of amphibians and lizards regularly inhabiting the litter is similar in all wet lowland forests studied. Increasing elevation and decreasing rainfall correlate with faunas having fewer species. The former is seen as the result of the differential ability of lowland species to invade uplands, and the latter as a decrease in the kind and duration of frog-spawning sites. Reduced equitability with increasing elevation is principally due to the much greater proportional increase of the commonest species. In the evolution of American and Bornean-Philippine wet lowland faunas, frogs with direct development (Eleutherodactylus) have radiated in the former in the same fashion that skinks have in the latter.

Biotropica↗

Relating residue in raccoon feces to food consumed

Feeding tests were conducted with captive raccoons ( Procyon lotor ) to permit more meaningful interpretation of food habit data obtained from fecal analysis. Ten diverse types of natural foods were offered in 20 tests. Digestibility coefficients were calculated that ranged from 3.6 for dry sunflowers, where considerable residue was recovered, to infinity for earthworms and boned meat where no residue was recovered. The influence of differences in both food and animal behavior on digestibility coefficients was significant (ANOVA, F<0.001). The use of digestibility coefficients to adjust quantitative estimates of fecal residue or to predict biomass consumed is of questionable value with raccoons due to variability in foods consumed and behavior of individual animals.

American Midland Naturalist↗

The use of auxiliary variables in capture-recapture and removal experiments

The dependence of animal capture probabilities on auxiliary variables is an important practical problem which has not been considered in the development of estimation procedures for capture-recapture and removal experiments. In this paper the linear logistic binary regression model is used to relate the probability of capture to continuous auxiliary variables. The auxiliary variables could be environmental quantities such as air or water temperature, or characteristics of individual animals, such as body length or weight. Maximum likelihood estimators of the population parameters are considered for a variety of models which all assume a closed population. Testing between models is also considered. The models can also be used when one auxiliary variable is a measure of the effort expended in obtaining the sample.

Biometrics↗

Estimation of population trajectories from count data

Monitoring of changes in animal population size is rarely possible through complete censuses; frequently, the only feasible means of monitoring changes in population size is to use counts of animals obtained by skilled observers as indices to abundance. Analysis of changes in population size can be severely biased if factors related to the acquisition of data are not adequately controlled for. In particular we identify two types of observer effects: these correspond to baseline differences in observer competence, and to changes through time in the ability of individual observers. We present a family of models for count data in which the first of these observer effects is treated as a nuisance parameter. Conditioning on totals of negative binomial counts yields a Dirichlet compound multinomial vector for each observer. Quasi-likelihood is used to estimate parameters related to population trajectory and other parameters of interest; model selection is carried out on the basis of Akaike's information criterion. An example is presented using data on Wood thrush from the North American Breeding Bird Survey.

Biometrics↗

Endocrine function and reproductive impairment in an irradiated population of the lizard Uta stansburiana

This study describes gonadal changes in lizards ( Uta stansburiana ) exposed to chronic low levels of γ radiation (1.5-10 R/day) under field conditions and attempts to evaluate endocrine involvement in these changes. Reproductive impairment in irradiated males is demonstrated during the breeding season by reduced testes weights and by seminiferous tubules devoid of germ cells. Ultrastructural morphology of Leydig cells, plasma testosterone levels, and developed accessory sex structures indicate normal steroidogenesis in such males; cytology of the pituitary gonadotropes and pituitary gonadotropin content indicate normal gonadotropin levels. These findings suggest that low levels of radiation affect the spermatogenic process directly, rather than through damage to the pituitary. Comparable irradiation causes complete resorption of ovarian tissue in some female Uta . The oviducts are atrophic and the pituitary gonadotropes hyperactive. Resemblance of gonadotropes to those of castrated animals and reduced pituitary hormone content suggest elevated levels of gonadotropin production, probably due to the absence of ovarian steroid feedback in affected animals. The continued mitotic activity of oogonia in adult reptiles and their limited number contribute to the vulnerability of germinal tissue in irradiated female lizards. Destruction of the germ cells is accompanied by resorption of all ovarian tissue including steroidogenic elements. Thus, in females as in males, radiation appears to damage gametogenesis rather than pituitary function.

Nevada↗

Back-pack unit for capturing waterfowl and upland game by night-lighting

A night-lighting unit, designed as a light weight back-pack, proved successful for capturing waterfowl pairs, pheasants ( Phasianus colchicus ), and cottontail rabbits ( Sylvilagus floridanus ) during the spring and summer when most breeding populations are widely dispersed. Eighty ducks of seven species were captured in 48 hours (1.7 ducks per hour) of night-lighting in marsh habitat. Similarly, 30 pheasants were trapped in 25 hours (1.2 birds per hour) and 63 cottontail rabbits were either observed at close range (6-12 ft) or captured during night-lighting operations in upland habitat. Catch per hour of effort increased for all species as their night habitat requirements and reaction to night-lights became known. The mobile unit proved well suited for intensive use on small areas where other methods of capture were unfeasible and where representative coverage of various habitat types was desired. Besides its utility for capturing animals, the unit provided a method for studying nocturnal movements, behavior, and habitat use of marked animals.

South Dakota↗

Analysis of radiotracking data using digitized habitat maps

A method is described that provides a rapid and accurate analysis of habitat used by radio-equipped animals. The digitizer (basically an X-Y plotter in reverse) converts maps into digital form by describing each habitat unit as a polygon that closely approximates the actual shape of the unit. The coordinates of each polygon are then stored on magnetic tape. Habitat classification data and other information are coded and combined with the proper polygon coordinates. This results in one file containing all habitat data. A computer program with inputs of tracking data and habitat data provides a listing of the habitat used by the animals studied. Analysis of habitat used by radio-equipped ducks is demonstrated using this method.

Minnesota↗

Metabolic indicators of habitat differences in four Minnesota deer populations

Blood samples were collected from 40 white-tailed deer ( Odocoileus virginianus ) from 4 winter yards in northeastern Minnesota from 17 March 1974 through 23 April 1975. The results of 26 blood assays were examined for the effects of age, sex, capture date, capture method, disease and location. Age-related effects were found for serum chloride, calcium, gamma globulin, creatine phosphokinase (CPK), lactic dehydrogenase (LDH), and alkaline phosphatase. The only sex difference was lower CPK in males. Date of collection effects were found for erythrocyte count, mean corpuscular volume (MCV), serum glucose, and nonesterified fatty acids (NEF A). Capture method affected serum glucose, acid base balance, and serum enzymes. Effects related primarily to capture location or habitat differences were found for erythrocyte count, MCV, mean corpuscular hemoglobin concentration (MCHC), serum urea, cholesterol, LDH, thyroxine, and NEF A. Animals whose assays indicated the poorest nutritional status inhabited wintering areas with the oldest vegetation. Habitat differences can be detected by measuring the physiological status of the local animal populations.

Minnesota↗

Feeding Ecology of Breeding Blue-Winged Teals

A 5-year investigation of factors influencing the selection of foods consumed by blue-winged teals ( Anas discors ) during the breeding season in the glaciated prairie region of south-central North Dakota showed that birds first arriving on the breeding grounds consumed a diet consisting of 45 percent invertebrates. The proportion of animal foods in the diet increased to 95 percent at the onset of the nesting season. The quality and quantity of foods selected at any given time were influenced by the biological demands and morphological adaptations of the bird, the behavior and ecology of the invertebrates selected as foods, and the general nature of the aquatic ecosystems as determined by the hydrology and geology of the area and modified by land use and weather. Feeding activities changed significantly when food availability within the aquatic ecosystem changed. During the spring and early summer, temporary and seasonal wetlands, if not severely disturbed, were of paramount importance to breeding blue-winged teals since they provided abundant and readily available, high protein, animal foods. Later in the summer when seasonal wetlands began to dry up, insects began to emerge in the semipermanent ponds and lakes, and feeding intensity shifted to these more permanent waters. This trend, however, was often reversed temporarily during the early summer following heavy precipitation that refilled shallow water areas and again stimulated invertebrate development.

North Dakota↗

A double survey estimate for sighting probability of sea otters in California

We developed an estimator for the probability of sighting sea otters ( Enhydra lutris ) by shore- based counters, based on simultaneous double-surveys. We then estimated probability of sighting sea otters in California and evaluated the estimator's principal assumptions. The overall probability of sighting sea otters on 5 replicated double-surveys at each of 6 study areas was 0.945. Estimated probability of sighting did not vary (P > 0.05) among study areas or over time and was not correlated (P > 0.05) with distance of otters from observers. Probability of sighting was affected (P < 0.05) by sea otter activity and group size. Activity-specific probabilities of sighting were: resting = 0.990, foraging = 0.769, and other = 0.885. All animals missed by 1 observer team were in groups of ≤ 2 otters, whereas 52% of the animals sighted during the study were in groups ≥ 3. Observer teams may have varied slightly in sighting ability, but this variation did not substantially affect population estimates.

California↗

Effects of sample size on kernel home range estimates

Kernel methods for estimating home range are being used increasingly in wildlife research, but the effect of sample size on their accuracy is not known. We used computer simulations of 10-200 points/home range and compared accuracy of home range estimates produced by fixed and adaptive kernels with the reference (REF) and least-squares cross-validation (LSCV) methods for determining the amount of smoothing. Simulated home ranges varied from simple to complex shapes created by mixing bivariate normal distributions. We used the size of the 95% home range area and the relative mean squared error of the surface fit to assess the accuracy of the kernel home range estimates. For both measures, the bias and variance approached an asymptote at about 50 observations/home range. The fixed kernel with smoothing selected by LSCV provided the least-biased estimates of the 95% home range area. All kernel methods produced similar surface fit for most simulations, but the fixed kernel with LSCV had the lowest frequency and magnitude of very poor estimates. We reviewed 101 papers published in The Journal of Wildlife Management (JWM) between 1980 and 1997 that estimated animal home ranges. A minority of these papers used nonparametric utilization distribution (UD) estimators, and most did not adequately report sample sizes. We recommend that home range studies using kernel estimates use LSCV to determine the amount of smoothing, obtain a minimum of 30 observations per animal (but preferably a?Y50), and report sample sizes in published results.

Journal of Wildlife Management↗

True metabolizable energy for wood ducks from acorns compared to other waterfowl foods

Acorns of bottomland red oaks (Quercus spp.) are an important food of North American wood ducks (Aix sponsa). Barras et al. (1996) demonstrated that female wood ducks selected willow oak (Q. phellos) acorns over other species. We measured true metabolizable energy (TME) derived by captive, wild-strain, adult female wood ducks from acorns of willow oak, water oak (Q. nigra), cherrybark oak (Q. pagoda), and pin oak (Q. palustris) to determine whether female wood ducks ' preference for willow oak acorns was related to TME. Estimates of TME within acorn species were relatively precise, yet we did not detect variation in TME among acorn species (P=0.31); hence, we estimated TME across species (2.76±0.033 [SE] kcal/g dry mass; n=34). We concluded that TME apparently did not explain female wood ducks ' preference for willow oak acorns and hypothesized that morphological characteristics of willow oak acorns may be proximate cues related to selection by wood ducks . We also summarized known TME estimates for acorns fed to wood ducks and mallards (Anas platyrhynchos), and natural and agricultural foods fed to mallards, northern pintails (A. acuta), blue-winged teal (A. discors), and Canada geese (Branta canadensis). We found that acorns and moist-soil plant seeds and tubers provided, on average, about 76% of the TME in agricultural seeds. Thus, bottomland-hardwood and moist-soil habitats have potential to provide significant amounts of dietary energy , as well as greater diversity of foods and nutrients than croplands. Researchers should continue to determine TME of common foods (plant and animal) of waterfowl , and use TME in estimating waterfowl habitat carrying capacity (e.g., Reinecke et al. 1989). Additionally, large-scale, reliable estimates of plant and animal food availability in bottomland-hardwood and moist-soil habitats are needed to evaluate carrying capacity of landscapes important to waterfowl , such as the Mississippi Alluvial Valley (MAV).

Journal of Wildlife Management↗

Autocorrelation of location estimates and the analysis of radiotracking data

The wildlife literature has been contradictory about the importance of autocorrelation in radiotracking data used for home range estimation and hypothesis tests of habitat selection. By definition, the concept of a home range involves autocorrelated movements, but estimates or hypothesis tests based on sampling designs that predefine a time frame of interest, and that generate representative samples of an animal's movement during this time frame, should not be affected by length of the sampling interval and autocorrelation. Intensive sampling of the individual's home range and habitat use during the time frame of the study leads to improved estimates for the individual, but use of location estimates as the sample unit to compare across animals is pseudoreplication. We therefore recommend against use of habitat selection analysis techniques that use locations instead of individuals as the sample unit. We offer a general outline for sampling designs for radiotracking studies.

Journal of Wildlife Management↗

Estimation of brood and nest survival: Comparative methods in the presence of heterogeneity

The Mayfield method has been widely used for estimating survival of nests and young animals, especially when data are collected at irregular observation intervals. However, this method assumes survival is constant throughout the study period, which often ignores biologically relevant variation and may lead to biased survival estimates. We examined the bias and accuracy of 1 modification to the Mayfield method that allows for temporal variation in survival, and we developed and similarly tested 2 additional methods. One of these 2 new methods is simply an iterative extension of Klett and Johnson's method, which we refer to as the Iterative Mayfield method and bears similarity to Kaplan-Meier methods. The other method uses maximum likelihood techniques for estimation and is best applied to survival of animals in groups or families, rather than as independent individuals. We also examined how robust these estimators are to heterogeneity in the data, which can arise from such sources as dependent survival probabilities among siblings, inherent differences among families, and adoption. Testing of estimator performance with respect to bias, accuracy, and heterogeneity was done using simulations that mimicked a study of survival of emperor goose ( Chen canagica ) goslings. Assuming constant survival for inappropriately long periods of time or use of Klett and Johnson's methods resulted in large bias or poor accuracy (often >5% bias or root mean square error) compared to our Iterative Mayfield or maximum likelihood methods. Overall, estimator performance was slightly better with our Iterative Mayfield than our maximum likelihood method, but the maximum likelihood method provides a more rigorous framework for testing covariates and explicity models a heterogeneity factor. We demonstrated use of all estimators with data from emperor goose goslings. We advocate that future studies use the new methods outlined here rather than the traditional Mayfield method or its previous modifications.

Journal of Wildlife Management↗