William I. Boarman

Cryptic genetic variation and paraphyly in ravens

Widespread species that are morphologically uniform may be likely to harbour cryptic genetic variation. Common ravens (Corvus corax) have an extensive range covering nearly the entire Northern Hemisphere, but show little discrete phenotypic variation. We obtained tissue samples from throughout much of this range and collected mitochondrial sequence and nuclear microsatellite data. Our study revealed a deep genetic break between ravens from the western United States and ravens from throughout the rest of the world. These two groups, the ‘California clade’ and the ‘Holarctic clade’ are well supported and over 4% divergent in mitochondrial coding sequence. Microsatellites also reveal significant differentiation between these two groups. Ravens from Minnesota, Maine and Alaska are more similar to ravens from Asia and Europe than they are to ravens from California. The two clades come in contact over a huge area of the western United States, with mixtures of the two mitochondrial groups present in Washington, Idaho and California. In addition, the restricted range Chihuahuan raven (Corvus cryptoleucus) of the south–west United States and Mexico is genetically nested within the paraphyletic common raven. Our findings suggest that the common raven may have formerly consisted of two allopatric groups that may be in the process of remerging.

Spatial pattern of risk of common raven predation on desert tortoises

Common Ravens (Corvus corax) in the Mojave Desert of California, USA are subsidized by anthropogenic resources. Large numbers of nonbreeding ravens are attracted to human developments and thus are spatially restricted, whereas breeding ravens are distributed more evenly throughout the area. We investigated whether the spatial distribution of risk of predation by ravens to juveniles of the threatened desert tortoise (Gopherus agassizii) was determined by the spatial distribution of (1) nonbreeding ravens at human developments (leading to “spillover” predation) or (2) breeding individuals throughout developed and undeveloped areas (leading to “hyperpredation”). Predation risk, measured using styrofoam models of juvenile desert tortoises, was high near places attracting large numbers of nonbreeding ravens, near successful nests, and far from successful nests when large numbers of nonbreeding ravens were present. Patterns consistent with both “spillover” predation and “hyperpredation” were thus observed, attr...

Diet composition of common ravens across the urban-wildland interface of the West Mojave Desert

Abstract Common ravens (Corvus corax) are human-subsidized scavengers and predators in the Mojave Desert. They have increased dramatically in number and have been implicated as contributors to the decline in desert tortoise (Gopherus agassizii) populations. Known patterns of increased fledging success near human developments suggested that food was the most likely resource subsidy received by ravens. Because ravens are opportunistic foragers with a generalist diet, we predicted that the types of resource subsidy provided by different kinds of human developments should be reflected in measures of diet composition of breeding ravens. We estimated diet composition from contents of raven pellets collected at nests and related diet composition to distance of the nests from roads and point sources of resource subsidies, such as towns or landfills. Ravens that nested close to point subsidies far from major roads had the greatest incidence of trash in their diets. Ravens that nested close to roads but far from point subsidies had a low incidence of trash and a higher incidence of presumably road-killed mammals and reptiles. Ravens far from both roads and point subsidies had more plant material and arthropods, and ravens close to both roads and point subsidies had more birds and amphibians. Diet diversity was not related to distance from roads or developments. Fledging success was correlated with diet composition, such that birds with diets consistent with trash or road-kill subsidies fledged the greatest number of chicks. Our results suggest that ravens forage opportunistically on foods available near their nests, and different kinds of human developments contribute different foods. Improved management of landfills and highway fencing to reduce road-kills may help slow the growth of raven populations in the Mojave.

COMMON RAVEN JUVENILE SURVIVAL IN A HUMAN-AUGMENTED LANDSCAPE

Abstract Anthropogenic resource subsidies have contributed to the dramatic increase in the abundance of Common Ravens (Corvus corax) in the western Mojave Desert, California, during the past 30 years. To better understand the effects of these subsidies on raven demography, we examined whether survival to juvenile departure from the natal territory could be predicted by a set of environmental and morphological variables, such as nest proximity to anthropogenic resources and juvenile condition. We captured 240 juvenile ravens over 2 years and marked them prior to fledging. Nest proximity to anthropogenic resources and earlier fledging dates significantly predicted raven juvenile survival to departure from the natal territory. The best-fitting mark-recapture models predicted postdeparture survival as a function of time since fledging, nest proximity to anthropogenic resources, and year hatched. The positive effect of nest proximity to anthropogenic resources influenced postdeparture survival for at least 9 months after fledging, as revealed by the mark-recapture analysis. Annual survival was 47% for first-year, 81% for second-year, and 83% for third-year birds. Our results support the hypothesis that anthropogenic resources contribute to increasing raven numbers via increased juvenile survival to departure as well as increased postdeparture survival. We expect raven numbers to grow in concert with the growing human presence in the Mojave Desert unless raven access to anthropogenic resources is diminished. La Sobrevivencia Juvenil de Corvus corax en un Paisaje Mejorado por Humanos Resumen. Los subsidios de recursos antropogénicos han contribuido al aumento dramático de la abundancia del cuervo Corvus corax en el occidente del desierto de Mojave durante los últimos 30 años. Para entender los efectos de estos subsidios en la demografía del cuervo, examinamos si la sobrevivencia hasta la salida de los juveniles del territorio natal podría ser predicha por un conjunto de variables morfológicas y ambientales, tales como su cercanía a los recursos antropogénicos y la condición de los juveniles. En dos años capturamos 240 cuervos juveniles y los marcamos antes que desarollaran plumas de vuelo. La cercanía del nido al punto de subsidio más cercano y las fechas tempranas de emplumamiento predijeron significativamente la sobrevivencia de los juveniles hasta su salida del teritorio natal. Los modelos de marcaje y recaptura más adecuados predijeron la sobrevivencia posterior a la salida del territorio como una función del tiempo desde el momento de emplumamiento, la cercanía al punto más cercano de subsidio y el año de nacimiento. Según el análisis de marcaje y recaptura, el efecto positivo de la cercanía a subsidios antropogénicos influye en la sobrevivencia después de la salida del territorio hasta por lo menos 9 meses después del emplumamiento. La sobrevivencia anual fue de 47% en el primer año, de 81% en el segundo año y de 83% para aves en su tercer año. Nuestros resultados apoyan la hipótesis de que los recursos antropogénicos contribuyen al crecimiento de los números de cuervos aumentando la sobrevivencia juvenil antes y después de su salida de los territorios natales. Esperamos que los números de cuervos aumenten en relación con la presencia humana en el desierto de Mojave a menos que su acceso a los recursos antropogénicos sea disminuido.

EFFECTS OF ANTHROPOGENIC DEVELOPMENTS ON COMMON RAVEN NESTING BIOLOGY IN THE WEST MOJAVE DESERT

Subsidized predators may affect prey abundance, distribution, and demography. Common Ravens (Corvus corax) are anthropogenically subsidized throughout their range and, in the Mojave Desert, have increased in number dramatically over the last 3-4 decades. Human-provided food resources are thought to be important drivers of raven population growth, but human developments add other features as well, such as nesting platforms. From 1996 to 2000, we examined the nesting ecology of ravens in the Mojave Desert, relative to anthropogenic developrhent. Ravens nested disproportionately near point sources of food and water subsidies (such as towns, landfills, and ponds) but not near roads (sources of road-killed carrion), even though both sources of subsidy enhanced fledging success. Initiation of breeding activity was more likely when a nest from the previous year was present at the start of a breeding season but was not affected by access to food. The relative effect of environmental modifications on fledging success varied from year to year, but the effect of access to human-provided resources was comparatively consistent, suggesting that humans provide consistently high-quality breeding habitat for ravens. Anthropogenic land cover types in the desert are expected to promote raven population growth and to allow ravens to occupy parts of the desert that otherwise would not support them. Predatory impacts of ravens in the Mojave Desert can therefore be considered indirect effects of anthropogenic development.

Fine-Scale Analysis Reveals Cryptic Landscape Genetic Structure in Desert Tortoises

Characterizing the effects of landscape features on genetic variation is essential for understanding how landscapes shape patterns of gene flow and spatial genetic structure of populations. Most landscape genetics studies have focused on patterns of gene flow at a regional scale. However, the genetic structure of populations at a local scale may be influenced by a unique suite of landscape variables that have little bearing on connectivity patterns observed at broader spatial scales. We investigated fine-scale spatial patterns of genetic variation and gene flow in relation to features of the landscape in desert tortoise (Gopherus agassizii), using 859 tortoises genotyped at 16 microsatellite loci with associated data on geographic location, sex, elevation, slope, and soil type, and spatial relationship to putative barriers (power lines, roads). We used spatially explicit and non-explicit Bayesian clustering algorithms to partition the sample into discrete clusters, and characterize the relationships between genetic distance and ecological variables to identify factors with the greatest influence on gene flow at a local scale. Desert tortoises exhibit weak genetic structure at a local scale, and we identified two subpopulations across the study area. Although genetic differentiation between the subpopulations was low, our landscape genetic analysis identified both natural (slope) and anthropogenic (roads) landscape variables that have significantly influenced gene flow within this local population. We show that desert tortoise movements at a local scale are influenced by features of the landscape, and that these features are different than those that influence gene flow at larger scales. Our findings are important for desert tortoise conservation and management, particularly in light of recent translocation efforts in the region. More generally, our results indicate that recent landscape changes can affect gene flow at a local scale and that their effects can be detected almost immediately.

As the raven flies: using genetic data to infer the history of invasive common raven (Corvus corax) populations in the Mojave Desert

Common raven (Corvus corax) populations in Mojave Desert regions of southern California and Nevada have increased dramatically over the past five decades. This growth has been attributed to increased human development in the region, as ravens have a commensal relationship with humans and feed extensively at landfills and on road‐killed wildlife. Ravens, as a partially subsidized predator, also represent a problem for native desert wildlife, in particular threatened desert tortoises (Gopherus agassizii). However, it is unclear whether the more than 15‐fold population increase is due to in situ population growth or to immigration from adjacent regions where ravens have been historically common. Ravens were sampled for genetic analysis at several local sites within five major areas: the West Mojave Desert (California), East Mojave Desert (southern Nevada), southern coastal California, northern coastal California (Bay Area), and northern Nevada (Great Basin). Analyses of mtDNA control region sequences reveal an increased frequency of raven ‘Holarctic clade’ haplotypes from south to north inland, with ‘California clade’ haplotypes nearly fixed in the California populations. There was significant structuring among regions for mtDNA, with high FST values among sampling regions, especially between the Nevada and California samples. Analyses of eight microsatellite loci reveal a mostly similar pattern of regional population structure, with considerably smaller, but mostly significant, values. The greater mtDNA divergences may be due to lower female dispersal relative to males, lower Ne, or effects of high mutation rates on maximal values of FST. Analyses indicate recent population growth in the West Mojave Desert and a bottleneck in the northern California populations. While we cannot rule out in situ population growth as a factor, patterns of movement inferred from our data suggest that the increase in raven populations in the West Mojave Desert resulted from movements from southern California and the Central Valley. Ravens in the East Mojave Desert are more similar to ones from northern Nevada, indicating movement between those regions. If this interpretation of high gene flow into the Mojave Desert is correct, then efforts to manage raven numbers by local control may not be optimally effective.

Movements of Juvenile Common Ravens in an Arid Landscape

Abstract Movement patterns of juvenile birds are poorly understood, yet critically important ecological phenomena, especially for species with a prolonged juvenile period. We evaluated postfledging movements of juvenile common ravens (Corvus corax) in a western Mojave Desert landscape composed of a mosaic of natural and anthropogenic elements. Generally, ravens do not begin breeding until after their fourth year. We marked 2 annual cohorts of juvenile ravens and followed them from dispersal from their natal territory for up to 33 months. Movements of juvenile common ravens were similar for males and females. Conspecifics and confined livestock feeding operations represented important resources for juvenile ravens, and juveniles were rarely located in open desert. However, initial movements from the natal territory to the nearest communal point subsidy rather than the closest anthropogenic resource suggested juvenile dispersal was influenced by the combination of conspecifics and anthropogenic resources, rather than the distribution of those resources. Land managers concerned with growing raven populations should reduce access to concentrated anthropogenic resources such as landfills and dairies, which serve as important resources for juveniles. Because juvenile ravens rarely venture into open desert, reducing their numbers by lethal removal or other means is unlikely to lessen raven predation of desert tortoises (Gopherus agassizii).

Catching Large Groups of Ravens: A Note on Procedures using Rocket Nets

ABSTRACT Capturing Common Ravens (Corvus corax) is very difficult. Several methods are currently used, but none effectively catch large numbers (>25 birds) of ravens at one time. Efficient capture of large numbers of ravens is needed for some autecology studies. We describe and evaluate the effectiveness of using a prebaited rocket net trapping method for simultaneously capturing large numbers of ravens. The study sites were within 2 landfills in Californias Mojave Desert, one at Edwards Air Force Base and the other at Fort Irwin National Training Center. We captured 283 ravens on 5 trapping occasions between 1995 and 1997, with an average of 57 birds per trapping occasion. We observed greater numbers of ravens at the bait sites with increasing bait duration, and these numbers appeared to level off after 25 to 30 days of baiting. Longer bait durations may habituate ravens to the resource and compensate for their wariness, which could increase capture success. More than half of the ravens captured (55%) were adults, and subadults composed the remaining age class (42%). Only 3% of the ravens captured were hatch-year birds, a result of trapping early in the breeding season. Using rocket nets is a safe and effective method to capture large numbers of ravens.