Amy A. Yackel Adams

NEST SURVIVAL ESTIMATION: A REVIEW OF ALTERNATIVES TO THE MAYFIELD ESTIMATOR

Abstract Reliable estimates of nest survival are essential for assessing strategies for avian conservation. We review the history of modifications and alternatives for estimating nest survival, with a focus on four techniques: apparent nest success, the Mayfield estimator, the Stanley method, and program MARK. The widely used Mayfield method avoids the known positive bias inherent in apparent nest success by estimating daily survival rates using the number of exposure days, eliminating the need to monitor nests from initiation. Concerns that some of Mayfields assumptions were restrictive stimulated the development of new techniques. Stanleys method allows for calculation of stage-specific daily survival rates when transition and failure dates are unknown, and eliminates Mayfields assumption that failure occurred midway through the nest-check interval. Program MARK obviates Mayfields assumption of constant daily survival within nesting stages and evaluates variation in nest survival as a function of biologically relevant factors. These innovative methods facilitate the evaluation of nest survival using an information-theoretic approach. We illustrate use of these methods with Lark Bunting (Calamospiza melanocorys) nest data from the Pawnee National Grassland, Colorado. Nest survival estimates calculated using Mayfield, Stanley, and MARK methods were similar, but apparent nest success estimates ranged 1– 24% greater than the other estimates. MARK analysis revealed that survival of Lark Bunting nests differed between site–year groups, declined with both nest age and time in season, but did not vary with weather parameters. We encourage researchers to use these approaches to gain reliable and meaningful nest survival estimates. Estimación de la Supervivencia de Nidos: Una Revisión de las Alternativas del Estimador Mayfield Resumen. Es esencial contar con estimaciones confiables de la supervivencia de nidos para evaluar las estrategias de conservación de las aves. Revisamos la historia de modificaciones y las alternativas para estimar la supervivencia de nidos, enfocándonos en cuatro técnicas: éxito aparente del nido, el estimador de Mayfield, el método de Stanley y el programa MARK. El método de Mayfield, ampliamente usado, evita el conocido sesgo positivo inherente al éxito aparente del nido mediante la estimación de tasas de supervivencia diaria usando el número de días de exposición, eliminando así la necesidad de monitorear los nidos desde el inicio. Las preocupaciones de que algunos de los supuestos del método de Mayfield son restrictivos estimularon el desarrollo de nuevas técnicas. El método de Stanley permite el cálculo de tasas de supervivencia diarias específicas para cada etapa cuando las fechas de transición y fracaso son desconocidas, y elimina el supuesto del método de Mayfield que sostiene que el fracaso ocurre en el medio del intervalo de monitoreo del nido. El programa MARK elimina el supuesto del método de Mayfield sobre supervivencia diaria constante dentro de las etapas de nidificación y evalúa la variación en la supervivencia de nidos como función de factores biológicamente relevantes. Estos métodos innovadores facilitan la evaluación de la supervivencia de nidos usando un enfoque teórico-informativo. Ilustramos el uso de estos métodos con datos de nidos de Calamospiza melanocorys provenientes de Pawnee National Grassland, Colorado. Las estimaciones de supervivencia de los nidos calculadas usando los métodos de Mayfield, Stanley y MARK fueron similares, pero las estimaciones del éxito aparente de los nidos fueron entre 1–24% mayores que las otras estimaciones. Los análisis con MARK revelaron que la supervivencia de los nidos de C. melanocorys difirió entre grupos de sitio-año, disminuyó con la edad del nido y el tiempo de la estación, pero no varió con parámetros climáticos. Estimulamos a los investigadores a usar estos enfoques para obtener estimaciones de supervivencia de nidos confiables y válidas.

MODELING POST-FLEDGING SURVIVAL OF LARK BUNTINGS IN RESPONSE TO ECOLOGICAL AND BIOLOGICAL FACTORS

We evaluated the influences of several ecological, biological, and methodological factors on post-fledging survival of a shortgrass prairie bird, the Lark Bunting (Calamospiza melanocorys). We estimated daily post-fledging survival (n = 206, 82 broods) using radiotelemetry and color bands to track fledglings. Daily survival probabilities were best explained by drought intensity, time in season (quadratic trend), ages < or = 3 d post-fledging, and rank given drought intensity. Drought intensity had a strong negative effect on survival. Rank was an important predictor of fledgling survival only during the severe drought of 2002 when the smallest fledglings had lower survival. Recently fledged young (ages < or = 3 d post-fledging) undergoing the transition from nest to surrounding habitat experienced markedly lower survival, demonstrating the vulnerable nature of this time period. Survival was greater in mid and late season than early season, corresponding to our assumptions of food availability. Neither mark type nor sex of attending parent influenced survival. The model-averaged product of the 22-d survival calculated using mean rank and median value of time in season was 0.360 +/- 0.08 in 2001 and 0.276 +/- 0.08 in 2002. Survival estimates that account for age, condition of young, ecological conditions, and other factors are important for parameterization of realistic population models. Biologists using population growth models to elucidate mechanisms of population declines should attempt to estimate species-specific of post-fledging survival rather than use generalized estimates.

MOVEMENTS AND SURVIVAL OF LARK BUNTING FLEDGLINGS

Abstract We quantified post-fledging pre-independence behavior and survival in Lark Buntings (Calamospiza melanocorys) using radio-telemetry. Brood division was recorded in six broods and was maintained throughout the observed fledgling care period. Chicks were capable of short flights (up to 25 m) by fledgling day 6 and longer flights (to 100 m) by fledgling day 13. During the first three weeks after fledging, juveniles moved as far as 800 m from nests. Nine of 23 (39%) monitored fledglings died within 15 days of fledging, primarily due to predation by raptors. Daily survival rates were 0.953 ± 0.019 for fledgling days 0–9, 0.955 ± 0.038 for fledgling days 10–20, and 0.953 ± 0.015 for fledgling days 0–20. The probability of surviving fledgling days 0–20 was 0.367. More quantification of juvenile survival is clearly needed to understand the role of post-fledging mortality in source-sink dynamics. Los Movimientos y Supervivencia de los Volantones de Calamospiza melanocorys Resumen. Cuantificamos la conducta y la supervivencia de volantones de Calamospiza melanocorys antes de independizarse de sus padres usando telemetría de radio. La división de la nidada se registró en seis nidadas y se mantuvo a través del período del cuidado de los volantones. Tras seis días de haber abandonado el nido, los polluelos eran capaces de realizar vuelos cortos (de hasta 25 m) y para el día trece ya realizaban vuelos más largos (a 100 m). Durante las primeras tres semanas después de salir del nido, los juveniles se movieron hasta 800 m de los nidos. Nueve de 23 (39%) volantones se murieron en los primeros 15 días fuera del nido, principalmente debido a depredación por aves rapaces. Las tasas diarias de supervivencia fueron de 0.953 ± 0.019 para los días 0 a 9, 0.955 ± 0.038 para los días 10 a 20, y 0.954 ± 0.015 para días 0 a 20. La probabilidad de sobrevivir entre los días 0 y 20 fue de 0.367. Se necesita más cuantificacion de la sobrevivencia en la etapa juvenil para entender el papel de la mortalidad tras la salida del nido en la dinámica de fuente-sumideros.

Weather effects on avian breeding performance and implications of climate change.

The influence of recent climate change on the worlds biota has manifested broadly, resulting in latitudinal range shifts, advancing dates of arrival of migrants and onset of breeding, and altered community relationships. Climate change elevates conservation concerns worldwide because it will likely exacerbate a broad range of identified threats to animal populations. In the past few decades, grassland birds have declined faster than other North American avifauna, largely due to habitat threats such as the intensification of agriculture. We examine the effects of local climatic variations on the breeding performance of a bird endemic to the shortgrass prairie, the Lark Bunting (Calamospiza melanocorys) and discuss the implications of our findings relative to future climate predictions. Clutch size, nest survival, and productivity all positively covaried with seasonal precipitation; yet relatively intense daily precipitation events temporarily depressed daily survival of nests. Nest survival was positively related to average temperatures during the breeding season. Declining summer precipitation may reduce the likelihood that Lark Buntings can maintain stable breeding populations in eastern Colorado although average temperature increases of up to 3 degrees C (within the range of this study) may ameliorate declines in survival expected with drier conditions. Historic climate variability in the Great Plains selects for a degree of vagility and opportunism rather than strong site fidelity and specific adaptation to local environments. These traits may lead to northerly shifts in distribution if climatic and habitat conditions become less favorable in the drying southern regions of the Great Plains. Distributional shifts in Lark Buntings could be constrained by future changes in land use, agricultural practices, or vegetative communities that result in further loss of shortgrass prairie habitats.

Distribution, density, and biomass of introduced small mammals in the Southern Mariana Islands.

Abstract: Although it is generally accepted that introduced small mammals have detrimental effects on island ecology, our understanding of these effects is frequently limited by incomplete knowledge of small mammal distribution, density, and biomass. Such information is especially critical in the Mariana Islands, where small mammal density is inversely related to effectiveness of Brown Tree Snake (Boiga irregularis) control tools, such as mouse-attractant traps. We used mark-recapture sampling to determine introduced small mammal distribution, density, and biomass in the major habitats of Guam, Rota, Saipan, and Tinian, including grassland, Leucaena forest, and native limestone forest. Of the five species captured, Rattus diardii (sensu Robins et al. 2007) was most common across habitats and islands. In contrast, Mus musculus was rarely captured at forested sites, Suncus murinus was not captured on Rota, and R. exulans and R. norvegicus captures were uncommon. Modeling indicated that neophobia, island, sex, reproductive status, and rain amount influenced R. diardii capture probability, whereas time, island, and capture heterogeneity influenced S. murinus and M. musculus capture probability. Density and biomass were much greater on Rota, Saipan, and Tinian than on Guam, most likely a result of Brown Tree Snake predation pressure on the latter island. Rattus diardii and M. musculus density and biomass were greatest in grassland, whereas S. murinus density and biomass were greatest in Leucaena forest. The high densities documented during this research suggest that introduced small mammals (especially R. diardii) are impacting abundance and diversity of the native fauna and flora of the Mariana Islands. Further, Brown Tree Snake control and management tools that rely on mouse attractants will be less effective on Rota, Saipan, and Tinian than on Guam. If the Brown Tree Snake becomes established on these islands, high-density introduced small mammal populations will likely facilitate and support a high-density Brown Tree Snake population, even as native species are reduced or extirpated.

Potential misuse of avian density as a conservation metric.

Effective conservation metrics are needed to evaluate the success of management in a rapidly changing world. Reproductive rates and densities of breeding birds (as a surrogate for reproductive rate) have been used to indicate the quality of avian breeding habitat, but the underlying assumptions of these metrics rarely have been examined. When birds are attracted to breeding areas in part by the presence of conspecifics and when breeding in groups influences predation rates, the effectiveness of density and reproductive rate as indicators of habitat quality is reduced. It is beneficial to clearly distinguish between individual- and population-level processes when evaluating habitat quality. We use the term reproductive rate to refer to both levels and further distinguish among levels by using the terms per capita fecundity (number of female offspring per female per year, individual level) and population growth rate (the product of density and per capita fecundity, population level). We predicted how density and reproductive rate interact over time under density-independent and density-dependent scenarios, assuming the ideal free distribution model of how birds settle in breeding habitats. We predicted population density of small populations would be correlated positively with both per capita fecundity and population growth rate due to the Allee effect. For populations in the density-dependent growth phase, we predicted no relation between density and per capita fecundity (because individuals in all patches will equilibrate to the same success rate) and a positive relation between density and population growth rate. Several ecological theories collectively suggest that positive correlations between density and per capita fecundity would be difficult to detect. We constructed a decision tree to guide interpretation of positive, neutral, nonlinear, and negative relations between density and reproductive rates at individual and population levels.

NEST SURVIVAL RELATIVE TO PATCH SIZE IN A HIGHLY FRAGMENTED SHORTGRASS PRAIRIE LANDSCAPE

Abstract Understanding the influences of habitat fragmentation on vertebrate populations is essential for the protection and ecological restoration of strategic sites for native species. We examined the effects of prairie fragmentation on avian reproductive success using artificial and natural nests on 26 randomly selected, privately owned patches of shortgrass prairie ranging in size from 7 to 454 ha within a cropland matrix in Washington County, Colorado, summer 2000. Survival trends of artificial and natural nests differed. Daily survival of artificial nests increased with patch size up to about 65 ha and differed little at larger patch sizes, whereas daily survival of Lark Bunting (Calamospiza melanocorys) and Horned Lark (Eremophila alpestris) nests decreased with increasing size of the grassland patch. We hypothesize that our unexpected findings of lower survival of natural nests with increasing patch sizes and different trends between artificial and natural nests are due to the particular structure of predator communities in our study area and the ways in which individual predators respond to artificial and natural nests. We recommend that the value of small habitat patches in highly fragmented landscapes not be overlooked.

POPULATION-SPECIFIC DEMOGRAPHIC ESTIMATES PROVIDE INSIGHTS INTO DECLINES OF LARK BUNTINGS (CALAMOSPIZA MELANOCORYS)

Abstract Many North American prairie bird populations have recently declined, and the causes of these declines remain largely unknown. To determine whether population limitation occurs during breeding, we evaluated the stability of a population of prairie birds using population-specific values for fecundity and postfledging survival. During 2001-2003, we radiomarked 67 female Lark Buntings (Calamospiza melanocorys) to determine annual fecundity and evaluate contributing factors such as nest survival and breeding response (number of breeding attempts and dispersal). Collectively, 67 females built 112 nests (1.67 ± 0.07 nests female−1 season−1; range: 1–3); 34 were second nests and 11 were third nests. Daily nest survival estimates were similar for initial and later nests with overall nest survival (DSR19) of 30.7% and 31.7%, respectively. Nest predation was the most common cause of failure (92%). Capture and radiomarking of females did not affect nest survival. Lark Bunting dispersal probabilities increased among females that fledged young from initial nests and females that lost their original nests late in the season. Conservative and liberal estimates of mean annual fecundity were 0.96 ±0.11 and 1.24 ± 0.09 female offspring per female, respectively. Given the fecundity and juvenile-survival estimates for this population, annual adult survival values of 71–77% are necessary to achieve a stable population. Because adult survival of prairie passerines ranges between 55% and 65%, this study area may not be capable of sustaining a stable population in the absence of immigration. We contrast our population assessment with one that assumes indirect values of fecundity and juvenile survival. To elucidate limiting factors, estimation of population-specific demographic parameters is desirable. We present an approach for selecting species and areas for evaluation of population stability. Las Estimaciones Demográficas Poblacionales Específicas Brindan Pistas sobre la Disminución de Calamospiza melanocorys

Do predators control prey species abundance? An experimental test with brown treesnakes on Guam

The effect of predators on the abundance of prey species is a topic of ongoing debate in ecology; the effect of snake predators on their prey has been less debated, as there exists a general consensus that snakes do not negatively influence the abundance of their prey. However, this viewpoint has not been adequately tested. We quantified the effect of brown treesnake (Boiga irregularis) predation on the abundance and size of lizards on Guam by contrasting lizards in two 1-ha treatment plots of secondary forest from which snakes had been removed and excluded vs. two 1-ha control plots in which snakes were monitored but not removed or excluded. We removed resident snakes from the treatment plots with snake traps and hand capture, and snake immigration into these plots was precluded by electrified snake barriers. Lizards were sampled in all plots quarterly for a year following snake elimination in the treatment plots. Following the completion of this experiment, we used total removal sampling to census lizards on a 100-m2 subsample of each plot. Results of systematic lizard population monitoring before and after snake removal suggest that the abundance of the skink, Carlia ailanpalai, increased substantially and the abundance of two species of gekkonids, Lepidodactylus lugubris and Hemidactylus frenatus, also increased on snake-free plots. No treatment effect was observed for the skink Emoia caeruleocauda. Mean snout-vent length of all lizard species only increased following snake removal in the treatment plots. The general increase in prey density and mean size was unexpected in light of the literature consensus that snakes do not control the abundance of their prey species. Our findings show that, at least where alternate predators are lacking, snakes may indeed affect prey populations.

Evaluating Abundance Estimate Precision and the Assumptions of a Count‐Based Index for Small Mammals

Abstract Conservation and management of small mammals requires reliable knowledge of population size. We investigated precision of mark–recapture and removal abundance estimates generated from live-trapping and snap-trapping data collected at sites on Guam (n = 7), Rota (n = 4), Saipan (n = 5), and Tinian (n = 3), in the Mariana Islands. We also evaluated a common index, captures per unit effort (CPUE), as a predictor of abundance. In addition, we evaluated cost and time associated with implementing live-trapping and snap-trapping and compared species-specific capture rates of selected live- and snap-traps. For all species, mark–recapture estimates were consistently more precise than removal estimates based on coefficients of variation and 95% confidence intervals. The predictive utility of CPUE was poor but improved with increasing sampling duration. Nonetheless, modeling of sampling data revealed that underlying assumptions critical to application of an index of abundance, such as constant capture probability across space, time, and individuals, were not met. Although snap-trapping was cheaper and faster than live-trapping, the time difference was negligible when site preparation time was considered. Rattus diardii spp. captures were greatest in Haguruma live-traps (Standard Trading Co., Honolulu, HI) and Victor snap-traps (Woodstream Corporation, Lititz, PA), whereas Suncus murinus and Mus musculus captures were greatest in Sherman live-traps (H. B. Sherman Traps, Inc., Tallahassee, FL) and Museum Special snap-traps (Woodstream Corporation). Although snap-trapping and CPUE may have utility after validation against more rigorous methods, validation should occur across the full range of study conditions. Resources required for this level of validation would likely be better allocated towards implementing rigorous and robust methods.