Foraging Habitat Distributions Affect Territory Size and Shape in the Tuamotu Kingfisher

I studied factors influencing territory configuration in the Tuamotu kingfisher (Todiramphus gambieri). Radiotelemetry data were used to define territory boundaries, and I tested for effects on territory size and shape of landscape habitat composition and foraging patch configuration. Tuamotu kingfisher territories were larger in areas with reduced densities of coconut plantation foraging habitat, and territories were less circular in the study site that had a single slender patch of foraging habitat. Maximum territory length did not differ between study sites, however, which suggested that the size of Tuamotu kingfisher territories might be bounded by the combined influence of maximum travel distances and habitat configurations. Results also suggested that birds enlarge territories as they age. Together, results supported previous work indicating that territory configurations represent a balance between the costs of defending a territory and gains from territory ownership

Territoriality, Prospecting, and Dispersal in Cooperatively Breeding Micronesian Kingfishers (\u3ci\u3eTodiramphus cinnamominus reichenbachii\u3c/i\u3e)

We investigated territoriality, prospecting, and dispersal behavior in cooperatively breeding Pohnpei Micronesian Kingfishers (Todiramphus cinnamominus reichenbachii) throughout the annual cycle using radiotelemetry and color-band resights. Mean home-range size was 6.3 ha and territories were 8.1 ha. Within territories, Micronesian Kingfishers shared 63% of their home-range space with coterritorial occupants, and 3% was shared with extraterritorial conspecifics. Birds on cooperative territories had larger home ranges that overlapped more with coterritory occupants’ home ranges than birds in pair-held territories. Despite evidence suggesting that resources necessary for survival and reproduction occurred on each territory, Micronesian Kingfishers of all age and sex classes made extraterritorial prospecting movements. Prospecting was rare; it comprised only 4.3% of our observations. When birds departed on forays, they were gone for ∼1.9 h and returned to home territories before sunset. Prospecting by dominant birds was temporally correlated with courtship and nest initiation, and birds were observed at neighboring nest sites with opposite-sex conspecifics during the period when females were available for fertilization. Juveniles and helpers prospected throughout the year and made repeated homesteading movements to dispersal destinations before dispersing. Mean dispersal distance for radiomarked individuals was 849 m. Results suggest that prospecting in Micronesian Kingfishers is a complex behavior that provides information for dispersal decisions and familiarity with dispersal destinations. Additionally, extraterritorial movements may provide covert opportunities for reproduction, which have potential to profoundly influence the distribution of fitness among helper and dominant Micronesian Kingfishers

Territoriality, Prospecting, and Dispersal in Cooperatively Breeding Micronesian Kingfishers (\u3ci\u3eTodiramphus cinnamominus reichenbachii\u3c/i\u3e)

We investigated territoriality, prospecting, and dispersal behavior in cooperatively breeding Pohnpei Micronesian Kingfishers (Todiramphus cinnamominus reichenbachii) throughout the annual cycle using radiotelemetry and color-band resights. Mean home-range size was 6.3 ha and territories were 8.1 ha. Within territories, Micronesian Kingfishers shared 63% of their home-range space with coterritorial occupants, and 3% was shared with extraterritorial conspecifics. Birds on cooperative territories had larger home ranges that overlapped more with coterritory occupants’ home ranges than birds in pair-held territories. Despite evidence suggesting that resources necessary for survival and reproduction occurred on each territory, Micronesian Kingfishers of all age and sex classes made extraterritorial prospecting movements. Prospecting was rare; it comprised only 4.3% of our observations. When birds departed on forays, they were gone for ∼1.9 h and returned to home territories before sunset. Prospecting by dominant birds was temporally correlated with courtship and nest initiation, and birds were observed at neighboring nest sites with opposite-sex conspecifics during the period when females were available for fertilization. Juveniles and helpers prospected throughout the year and made repeated homesteading movements to dispersal destinations before dispersing. Mean dispersal distance for radiomarked individuals was 849 m. Results suggest that prospecting in Micronesian Kingfishers is a complex behavior that provides information for dispersal decisions and familiarity with dispersal destinations. Additionally, extraterritorial movements may provide covert opportunities for reproduction, which have potential to profoundly influence the distribution of fitness among helper and dominant Micronesian Kingfishers

Population demography, resource use, and movement in cooperatively breeding Micronesian Kingfishers

Island systems and species are susceptible to extinction because of their small population size and an ecological naiveté from an evolutionary past lacking strong competition and predation. For example, only one-fifth of the world’s bird species occur on islands, yet more than 90% of the avian extinctions witnessed during historic times were island forms. Introduced predators and competitor species are among the major conservation issues facing insular systems. On the island of Guam, brown tree snakes (Boiga irregularis) are responsible for the local extinction of twelve native forest birds. The endangered Guam Micronesian Kingfisher (Todiramphus cinnamominus cinnamominus) is one of the species affected by the introduced snake, as the bird remains only in captive breeding institutions on the U.S. mainland. In addition to Guam, the islands of Pohnpei and Palau host endemic subspecies of Micronesian Kingfisher (T. c. reichenbachii and T. c. pelewensis respectively) that are similarly threatened with extinction. Previous investigations into the behavioral ecology of the Pohnpei subspecies of Micronesian Kingfisher yielded observations of cooperative social behaviors. Over the past three decades, much research has focused on cooperatively breeding species, which are commonly characterized by non-breeding individuals that delay dispersal and assist others with reproduction. Research addressing cooperative breeding suggests that the behavior is a complex response to interacting factors including life history characteristics, demography, resources, movement, and behavior. The dearth of information available about critically endangered Micronesian Kingfishers, combined with their potential to provide new insights into cooperative social behaviors, inspired the research presented in this dissertation. The aims were two-fold; results were intended to bolster our understanding of cooperative social behaviors while simultaneously providing vital information to conservation practitioners. Methodology for determining the sex of study individuals is presented in chapter two, which facilitated investigations that followed. Chapter three addressed the interaction between kingfishers and resources at both the landscape and home range scale. Higher population densities are associated with lowland mangrove, marsh forested habitats, and open vegetation types at the landscape scale. Results further indicated that at the home range scale, birds selectively used late succession forested habitats in higher proportions than their availability, and forest areas were entirely utilized in study areas where territories were packed boundary-to-boundary. Together these suggested that forested areas and the resources they contain might be limited for Micronesian Kingfishers. Movement and space use in Micronesian Kingfishers were the focus of chapter four. Within territories, the home ranges of birds overlapped, although not entirely. Birds of all ages and social classes made extraterritorial prospecting movements, but they appear to serve different functions. Juveniles and helpers were observed dispersing from natal areas, but only after repeated extraterritorial homesteading movements. The timing and destinations of adult prospecting suggested that the behavior might provide opportunities for covert reproduction. Population demography was addressed in chapters five and six, which concluded with the development of a population projection model that will be useful in kingfisher conservation efforts throughout the Pacific. Nestlings on cooperative territories had higher estimated survival rates than those on pair territories. Further, the timing of nestling disappearances and a modified nestling mandible suggested that mortalities were caused by siblicidal nest-mates. In chapter six, post-fledging vital rates were estimated for Micronesian Kingfishers and a population projection matrix model was developed. Vital rate parameters were then varied, and the model was used in a simulation analysis to evaluate the apparent influence of each parameter on population dynamics across a range of potential values. The exercise was intended to lend insight into the dynamics of Micronesian Kingfisher populations and to form a base model for management of the other eleven endangered Pacific Todiramphus species. In summary, information presented in this dissertation lends insight into factors important to understanding population demography, resource use, and movement in cooperatively breeding Micronesian Kingfishers. Results illustrate that, like other cooperatively breeding species, the birds on Pohnpei are highly territorial and dispersal options may be limited by territory vacancies and forest resources. Extraterritorial prospecting movements have been observed in many cooperative species, and these results illustrate that they may serve multiple purposes. Siblicide is also a phenomenon present in resource-limited species and its occurrence in Pohnpei Micronesian Kingfishers underscores the importance of resources to the evolutionary history of the birds. Results from demographic analyses and modeling suggest that conservation efforts for Micronesian Kingfishers, and the other eleven Todiramphus species, should be broadly focused on all life history stages.Keywords: Todirhamphus cinnamominus, endangered species, Halcyon cinnamomina, Todiramphus cinnamominus, cooperative breeding, conservatio

Sex Determination of Pohnpei Micronesian Kingfishers Using Morphological and Molecular Genetic Techniques

Conservation-oriented studies of Micronesian Kingfishers (Todiramphus cinnamominus) have been hindered by a lack of basic natural history information, despite the status of the Guam subspecies (T. c. cinnamominus) as one of the most endangered species in the world. We used tissue samples and morphometric measures from museum specimens and wild-captured Pohnpei Micronesian Kingfishers (T. c. reichenbachii) to develop methods for sex determination. We present a modified molecular protocol and a discriminant function that yields the probability that a particular individual is male or female. Our results revealed that females were significantly larger than males, and the discriminant function correctly predicted sex in 73% (30/41) of the individuals. The sex of 86% (18/21) of individuals was correctly assigned when a moderate reliability threshold was set. Sex determination using molecular genetic techniques was more reliable than methods based on morphology. Our results will facilitate recovery efforts for the critically endangered Guam Micronesian Kingfisher and provide a basis for sex determination in the 11 other endangered congeners in the Pacific Basin

Functional visual sensitivity to ultraviolet wavelengths in the Pileated Woodpecker (\u3ci\u3eDryocopus pileatus\u3c/i\u3e), and its influence on foraging substrate selection

Most diurnal birds are presumed visually sensitive to near ultraviolet (UV)wavelengths, however, controlled behavioral studies investigating UV sensitivity remain few. Although woodpeckers are important as primary cavity excavators and nuisance animals, published work on their visual systems is limited. We developed a novel foraging-based behavioral assay designed to test UV sensitivity in the Pileated Woodpecker (Dryocopus pileatus). We acclimated 21 wild-caught woodpeckers to foraging for frozen mealworms within 1.2 m sections of peeled cedar (Thuja spp.) poles.We then tested the functional significance of UV cues by placing frozen mealworms behind UV-reflective covers, UV-absorptive covers, or decayed red pine substrates within the same 1.2 m poles in independent experiments. Behavioral responses were greater toward both UV-reflective and UV-absorptive substrates in three experiments. Study subjects therefore reliably differentiated and attended to two distinct UV conditions of a foraging substrate. Cue-naïve subjects showed a preference for UV-absorptive substrates, suggesting that woodpeckers may be pre-disposed to foraging from such substrates. Behavioral responses were greater toward decayed pine substrates (UV-reflective) than sound pine substrates suggesting that decayed pine can be a useful foraging cue. The finding that cue-naïve subjects selected UV-absorbing foraging substrates has implications for ecological interactions of woodpeckers with fungi.Woodpeckers transport fungal spores, and communication methods analogous to those of plant-pollinator mutualisms (i.e. UV-absorbing patterns) may have evolved to support woodpecker-fungus mutualisms

Visual cues for woodpeckers: light reflectance of decayed wood varies by decay fungus

The appearance of wood substrates is likely relevant to bird species with life histories that require regular interactions with wood for food and shelter. Woodpeckers detect decayed wood for cavity placement or foraging, and some species may be capable of detecting trees decayed by specific fungi; however, a mechanism allowing for such specificity remains unidentified.We hypothesized that decay fungi associated with woodpecker cavity sites alter the substrate reflectance in a species-specific manner that is visually discriminable by woodpeckers. We grew 10 species of wood decay fungi from pure cultures on sterile wood substrates of 3 tree species. We then measured the relative reflectance spectra of decayed and control wood wafers and compared them using the receptor noise-limited (RNL) color discrimination model. The RNL model has been used in studies of feather coloration, egg shells, flowers, and fruit to model how the colors of objects appear to birds. Our analyses indicated 6 of 10 decayed substrate/control comparisons were above the threshold of discrimination (i.e., indicating differences discriminable by avian viewers), and 12 of 13 decayed substrate comparisons were also above threshold for a hypothetical woodpecker. We conclude that woodpeckers should be capable of visually detecting decayed wood on trees where bark is absent, and they should also be able to detect visually species-specific differences in wood substrates decayed by fungi used in this study. Our results provide evidence for a visual mechanism by which woodpeckers could identify and select substrates decayed by specific fungi, which has implications for understanding ecologically important woodpecker–fungus interactions. El aspecto de los sustratos de madera posiblemente sea relevante para especies de aves que tienen historias de vida que dependen de interacciones regulares con la madera para alimentaci´on y resguardo. Los pa´jaros carpinteros detectan la madera degradada para establecer sus cavidades o para forrajear, y algunas especies podr´ıan ser capaces de detectar a´rboles que son degradadas por alg´un hongo en particular. Sin embargo, a´un no se identifica un mecanismo que permita identificar tal especificidad. Nuestra hip´otesis es que los hongos xil ´ofagos asociados a sitios con cavidades para carpinteros alteran la reflectancia del sustrato en una manera espec´ıfica a especie que es visualmente discernible para los carpinteros. Cultivamos 10 especies de hongos xil ´ofagos a partir de cultivos puros en sustratos est´eriles de madera de tres especies de a´rboles. A continuaci´on, medimos el espectro de reflectancia de la madera de la madera degradada y trozos de madera control, y las comparamos entre s´ı usando el modelo de discriminaci´on de color del receptor de ruido limitado (RNL, por sus siglas en ingl´es). El modelo RNL ha sido utilizado en estudios de coloraci´on de plumas, cascar´on de huevo, flores y frutos para modelar c´omo perciben las aves el color de los objetos. Nuestros ana´lisis indican que 6 de 10 comparaciones sustrato/control estuvieron por encima del umbral de discriminaci´on (e.g., indicando diferencias discernibles por observadores aviares) y que las comparaciones de 12 de los 13 sustratos degradados estuvieron por encima del umbral para un carpintero hipot´etico. Concluimos que los carpinteros deben ser capaces de detectar visualmente la madera degradada en a´rboles donde la corteza esta´ ausente y tambi´en deben detectar visualmente diferencias espec´ıficas a especie en los sustratos de madera degradada por los hongos utilizados en este estudio. Nuestros resultados proveen evidencia de un mecanismo visual por medio del cual los pa´jaros carpinteros pueden identificar y seleccionar los sustratos degradados por hongos espec´ıficos, lo cual tiene implicaciones en nuestro entendimiento de las importantes interacciones entre carpinteros y hongos

Phenological mismatch in Arctic-breeding shorebirds: Impact of snowmelt and unpredictable weather conditions on food availability and chick growth

The ecological consequences of climate change have been recognized in numerous species, with perhaps phenology being the most well-documented change. Phenological changes may have negative consequences when organisms within different trophic levels respond to environmental changes at different rates, potentially leading to phenological mismatches between predators and their prey. This may be especially apparent in the Arctic, which has been affected more by climate change than other regions, resulting in earlier, warmer, and longer summers. During a 7-year study near Utqiaġvik (formerly Barrow), Alaska, we estimated phenological mismatch in relation to food availability and chick growth in a community of Arctic-breeding shorebirds experiencing advancement of environmental conditions (i.e., snowmelt). Our results indicate that Arctic-breeding shorebirds have experienced increased phenological mismatch with earlier snowmelt conditions. However, the degree of phenological mismatch was not a good predictor of food availability, as weather conditions after snowmelt made invertebrate availability highly unpredictable. As a result, the food available to shorebird chicks that were 2–10 days old was highly variable among years (ranging from 6.2 to 28.8 mg trap−1 day−1 among years in eight species), and was often inadequate for average growth (only 20%–54% of Dunlin and Pectoral Sandpiper broods on average had adequate food across a 4-year period). Although weather conditions vary among years, shorebirds that nested earlier in relation to snowmelt generally had more food available during brood rearing, and thus, greater chick growth rates. Despite the strong selective pressure to nest early, advancement of nesting is likely limited by the amount of plasticity in the start and progression of migration. Therefore, long-term climatic changes resulting in earlier snowmelt have the potential to greatly affect shorebird populations, especially if shorebirds are unable to advance nest initiation sufficiently to keep pace with seasonal advancement of their invertebrate prey

Factors affecting female space use in ten populations of prairie chickens

Conservation of wildlife depends on an understanding of the interactions between animal movements and key landscape factors. Habitat requirements of wide-ranging species often vary spatially, but quantitative assessment of variation among replicated studies at multiple sites is rare. We investigated patterns of space use for 10 populations of two closely related species of prairie grouse: Greater Prairie-Chickens (Tympanuchus cupido) and Lesser Prairie-Chickens (T. pallidicinctus). Prairie chickens require large, intact tracts of native grasslands, and are umbrella species for conservation of prairie ecosystems in North America. We used resource utilization functions to investigate space use by female prairie chickens during the 6-month breeding season from March through August in relation to lek sites, habitat conditions, and anthropogenic development. Our analysis included data from 382 radio-marked individuals across a major portion of the extant range. Our project is a unique opportunity to study comparative space use of prairie chickens, and we employed standardized methods that facilitated direct comparisons across an ecological gradient of study sites. Median home range size of females varied ~10-fold across 10 sites (3.6–36.7 km²), and home ranges tended to be larger at sites with higher annual precipitation. Proximity to lek sites was a strong and consistent predictor of space use for female prairie chickens at all 10 sites. The relative importance of other predictors of space use varied among sites, indicating that generalized habitat management guidelines may not be appropriate for these two species. Prairie chickens actively selected for prairie habitats, even at sites where ~90% of the land cover within the study area was prairie. A majority of the females monitored in our study (>95%) had activity centers within 5 km of leks, suggesting that conservation efforts can be effectively concentrated near active lek sites. Our data on female space use suggest that lek surveys of male prairie chickens can indirectly assess habitat suitability for females during the breeding season. Lek monitoring and surveys for new leks provide information on population trends, but can also guide management actions aimed at improving nesting and brood-rearing habitats

Microclimate and nest-site selection in Micronesian Kingfishers

We studied the relationship between microclimate and nest-site selection in the Pohnpei Micronesian Kingfisher (Todiramphus cinnamominus reichenbachii) which excavates nest cavities from the mudlike nest structures of arboreal termites (Nasutitermes sp.) or termitaria. Mean daily high temperatures at termitaria were cooler and daily low temperatures were warmer than at random sites in the forest. Results also indicate that termitaria provided insulation from temperature extremes, and that temperatures inside termitaria were within the thermoneutral zone of Micronesian Kingfishers more often than those outside. No differences were identified in temperatures at sites where nest termitaria and nonnest termitaria occurred or among the insulation properties of used and unused termitaria. These results suggest that although termitaria provide insulation from thermal extremes and a metabolically less stressful microclimate, king-fishers did not select from among available termitaria based on their thermal properties. Our findings are relevant to conservation efforts for the critically endangered Guam Micronesian Kingfisher (T. c. cinnamominus) which is extinct in the wild and exists only as a captive population. Captive breeding facilities should provide aviaries with daily ambient temperatures ranging from 22.06 °C to 28.05 °C to reduce microclimate-associated metabolic stress and to replicate microclimates used by wild Micronesian Kingfishers