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Managing a subsidized predator population: Reducing common raven predation on desert tortoises

Human communities often are an inadvertent source of food, water, and other resources to native species of wildlife. Because these resources are more stable and predictable than those in a natural environment, animals that subsist on them are able to increase in numbers and expand their range, much to the detriment of their competitors and species they prey upon. In the Mojave Desert, common ravens (Corvus corax) have benefited from human-provided resources to increase in population size precipitously in recent years. This trend has caused concern because ravens prey on juvenile desert tortoises (Gopherus agassizii), a federally threatened species. In this paper, I discuss management strategies to reduce raven predation on desert tortoises. The recommendations fall into three categories: (1) managing raven populations by reducing access to anthropogenic resources; (2) removing offending ravens or other birds in specially targeted tortoise management zones; and (3) continuing research on raven ecology, raven behavior, and methods of reducing raven predation on tortoises. I also recommend approaching the problem within an adaptive management framework: management efforts should first be employed as scientific experiments - with replicates and controls - to yield an unbiased assessment of their effectiveness. Furthermore, these strategies should be implemented in concert with actions that reduce other causes of desert tortoise mortality to aid the long-term recovery of their populations. Overall, the approaches outlined in this paper are widely applicable to the management of subsidized predators, particularly where they present a threat to a declining species of prey.

Environmental Management

Broad‐scale occurrence of a subsidized avian predator: reducing impacts of ravens on sage‐grouse and other sensitive prey

Expanding human enterprise across remote environments impacts numerous wildlife species. Anthropogenic resources provide subsidies for generalist predators that can lead to cascading effects on prey species at lower trophic levels. A fundamental challenge for applied ecologists is to disentangle natural and anthropogenic influences on species occurrence, and subsequently develop spatially explicit models to help inform management and conservation decisions. Using Bayesian hierarchical occupancy models, we mapped the broad‐scale occurrence of common ravens Corvus corax as a function of natural and anthropogenic landscape covariates using >15,000 point count surveys performed during 2007–2016 within the Great Basin region, USA. Raven abundance and distribution is substantially increasing across the American west due to unintended anthropogenic resource subsidies. Importantly, ravens prey on eggs and chicks of numerous species including greater sage‐grouse Centrocercus urophasianus , an indicator species whose decline is at the centre of national conservation strategies and land use policies.Anthropogenic factors that contributed to greater raven occurrence were: increased road density, presence of transmission lines, agricultural activity, and presence of roadside rest areas. Natural landscape characteristics included lower elevations with greener vegetation (NDVI), greater stream and habitat edge densities, and lower percentages of big sagebrush A. tridentata spp . Interactions between anthropogenic sources of nesting substrate and food subsidies suggested that raven occurrence increased multiplicatively when these resource subsidies co‐occurred. Overall, the average probability of raven occurrence estimated within sagebrush ecosystems of the study area was ~0.83. Synthesis and applications . We demonstrate how anthropogenic factors can be disentangled from natural effects when making spatially‐explicit predictions of subsidized predators occurring across expansive landscapes. This approach can guide management decisions where subsidized predators overlap sensitive prey habitats. For example, we identify areas where elevated raven occurrence coincides with breeding sage‐grouse concentration areas and appears to be largely driven by anthropogenic factors. Management applications could focus on reducing raven access to anthropogenic subsidies in these areas, while prioritizing habitat improvements for sage‐grouse elsewhere. Our approach is applicable to other species where widespread survey data are available.

Journal of Applied Ecology

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, attributed to the nonbreeding and breeding segments of the population, respectively. Furthermore, because locations of successful nests changed almost annually, consistent low-predation refugia for juvenile desert tortoises were nearly nonexistent. Consequently, anthropogenic resources for ravens could indirectly lead to the suppression, decline, or even extinction of desert tortoise populations.

California

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 development. 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 humanprovided 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.

California

Movements of juvenile common ravens in an arid landscape

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).

Journal of Wildlife Management

Selection of anthropogenic features and vegetation characteristics by nesting Common Ravens in the sagebrush ecosystem

Common Raven ( Corvus corax ) numbers and distribution are increasing throughout the sagebrush steppe, influencing avian communities in complex ways. Anthropogenic structures are thought to increase raven populations by providing food and nesting subsidies, which is cause for concern because ravens are important nest predators of sensitive species, including Greater Sage-Grouse ( Centrocercus urophasianus ). During 2007–2009, we located raven nests in southeastern Idaho and conducted a resource selection analysis. We measured variables at multiple spatial scales for 72 unique nest locations, including landscape-level vegetation characteristics and anthropogenic structures. Using generalized linear mixed models and an information-theoretic approach, we found a 31% decrease in the odds of nesting by ravens for every 1 km increase in distance away from a transmission line. Furthermore, a 100-m increase in distance away from the edge of two different land cover types decreased the odds of nesting by 20%, and an increase in the amount of edge by 1 km within an area of 102.1 ha centered on the nest increased the odds of nesting by 49%. A post hoc analysis revealed that ravens were most likely to nest near edges of adjoining big sagebrush ( Artemisia tridentata ) and land cover types that were associated with direct human disturbance or fire. These findings contribute to our understanding of raven expansion into rural environments and could be used to make better-informed conservation decisions, especially in the face of increasing renewable energy development.

Idaho

Common Raven ( Corvus corax ) kleptoparasitism at a Golden Eagle ( Aquila chyrsaetos ) nest in southern Nevada

The Common Raven ( Corvus corax ) is a ubiquitous species in the Mojave Desert of southern Nevada and California. From 5 to 24 May 2014, using remote trail cameras, we observed ravens repeatedly kleptoparasitizing food resources from the nest of a pair of Golden Eagles ( Aquila chyrsaetos ) in the Spring Mountains of southern Nevada. The ravens fed on nine (30%) of the 30 prey items delivered to the nest during the chick rearing period. Kleptoparasitic behavior by the ravens decreased as the eagle nestling matured to seven weeks of age, suggesting a narrow temporal window in which ravens can successfully engage in kleptoparasitic behavior at eagle nests. The observation of kleptoparasitism by Common Ravens at the nest suggests potential risks to young Golden Eagles from Common Ravens.

Nevada

Observations of territorial breeding common ravens caching eggs of greater sage-grouse

Previous investigations using continuous video monitoring of greater sage-grouse Centrocercus urophasianus nests have unambiguously identified common ravens Corvus corax as an important egg predator within the western United States. The quantity of greater sage-grouse eggs an individual common raven consumes during the nesting period and the extent to which common ravens actively hunt greater sage-grouse nests are largely unknown. However, some evidence suggests that territorial breeding common ravens, rather than nonbreeding transients, are most likely responsible for nest depredations. We describe greater sage-grouse egg depredation observations obtained opportunistically from three common raven nests located in Idaho and Nevada where depredated greater sage-grouse eggs were found at or in the immediate vicinity of the nest site, including the caching of eggs in nearby rock crevices. We opportunistically monitored these nests by counting and removing depredated eggs and shell fragments from the nest sites during each visit to determine the extent to which the common raven pairs preyed on greater sage-grouse eggs. To our knowledge, our observations represent the first evidence that breeding, territorial pairs of common ravens cache greater sage-grouse eggs and are capable of depredating multiple greater sage-grouse nests.

Idaho, Nevada

Landscape alterations influence differential habitat use of nesting buteos and ravens within sagebrush ecosystem: implications for transmission line development

A goal in avian ecology is to understand factors that influence differences in nesting habitat and distribution among species, especially within changing landscapes. Over the past 2 decades, humans have altered sagebrush ecosystems as a result of expansion in energy production and transmission. Our primary study objective was to identify differences in the use of landscape characteristics and natural and anthropogenic features by nesting Common Ravens ( Corvus corax ) and 3 species of buteo (Swainson's Hawk [ Buteo swainsoni ], Red-tailed Hawk [ B. jamaicensis ], and Ferruginous Hawk [ B. regalis ]) within a sagebrush ecosystem in southeastern Idaho. During 2007–2009, we measured multiple environmental factors associated with 212 nest sites using data collected remotely and in the field. We then developed multinomial models to predict nesting probabilities by each species and predictive response curves based on model-averaged estimates. We found differences among species related to nesting substrate (natural vs. anthropogenic), agriculture, native grassland, and edge (interface of 2 cover types). Most important, ravens had a higher probability of nesting on anthropogenic features (0.80) than the other 3 species (<0.10), and the probability of nesting near agriculture was greatest for ravens (0.55) followed by Swainson's Hawk (0.28). We also describe changes in nesting densities over 4 decades at this site as related to natural and anthropogenic disturbances. Since the 1970s, the composition of the raptor and raven nesting community has drastically changed with anthropogenic alterations and loss of continuous stands of sagebrush ( Artemisia spp.), favoring increased numbers of nesting ravens and fewer nesting Ferruginous Hawks. Our results indicate that habitat alterations, fragmentation, and forthcoming disturbances anticipated with continued energy development in sagebrush steppe ecosystems can lead to predictable changes in raptor and raven communities.

Idaho

Evaluation of the Raven sUAS to detect and monitor greater sage-grouse leks within the Middle Park population

Staff from the U.S. Geological Survey Fort Collins Science Center and the Colorado Parks and Wildlife Hot Sulphur Springs Office began discussions in 2011 for a proof of concept study to test the Raven RQ-11A small Unmanned Aircraft System (Raven sUAS) for its suitability to detect and monitor greater sage-grouse (Centrocercus urophasianus) breeding sites (leks). During April 2013, the Raven sUAS was flown over two known lek sites within the Middle Park population in Grand County, Colorado. Known sites were flown to determine the reaction of the greater sage-grouse to the aircraft and to determine if the technology had potential for future use of locating new leks and obtaining population counts on known, active lek sites. The Raven sUAS is a hand-launched reconnaissance and data-gathering tool developed for the U.S. Department of Defense by AeroVironment, Inc. Originally designed to provide aerial observation, day or night, at line-of-site ranges up to 6.2 miles (10 kilometers), the Raven sUAS has a wingspan of 4.5 feet (1.38 meters) and weighs 4.2 pounds (1.9 kilograms). A 60-minute lithium-ion rechargeable battery powers the system which also transmits live video (color or infrared imagery), compass headings, and location information to a ground control station. The Raven sUAS is typically operated by a three-person flight crew consisting of a pilot, mission operator, and a trained observer.

Colorado

First recorded observations of conspecific egg and nestling consumption in common ravens (Corvus corax)

We observed cannibalism, the act of consuming a conspecific, of eggs and nestlings by Common Ravens ( Corvus corax ; hereafter “raven”) by video-monitoring nests in Nevada and California. Specifically, within the sagebrush steppe of Nevada, adult ravens killed and consumed raven chicks from an active nest. Additionally, on the coast of California, we observed adult ravens consume inviable eggs from their own nest following full-term incubation. To our knowledge, these observations represent the first documented cannibalistic behavior by ravens.

California, Nevada

Evaluating common raven take for greater sage-grouse in Oregon’s Baker County Priority Conservation Area and Great Basin Region

The common raven ( Corvus corax ; raven) is a nest predator of species of conservation concern, such as the greater sage-grouse ( Centrocercus urophasianus ). Reducing raven abundance by take requires authorization under the Migratory Bird Treaty Act. To support U.S. Fish and Wildlife Service’s take decisions (e.g., those that authorize killing a specified proportion or number of individuals annually in a defined area), including the most recent one for Oregon’s Baker County Priority Area for Conservation (PAC), we modeled raven population dynamics under hypothetical scenarios with take rates ranging from below to above the maximum sustained yield (MSY; i.e., tr msy = 0.01-0.60). We fit a Bayesian state-space logistic model to estimate abundance based on the Breeding Bird Survey route-level count data for the PAC during 1997-2019 and Great Basin Region (GBR) during 1968-2019. We predicted abundance for 2019-2030 and evaluated potential take levels (PTL) for the PAC and GBR. Abundance averaged 682 (SE = 93) for the PAC during 1997-2019 and 333,027 (SE = 20,504) for the GBR during 1968-2019. With take rates between 0.41 and 0.60, predicted abundance averaged 308 (SD = 405) for the PAC and 142,258 (SD = 53,474) for the GBR during 2019-2030. With management factor F = 0.75-2 for takes ranging from below to above the MSY, the PTL 50 th percentiles were 150-401 yr -1 for the PAC and 60,457-161,219 yr -1 for the GBR. Our modeling framework is flexible and can be part of a comprehensive management strategy for ravens in the western United States.

California, Idaho, Nevada, Oregon, Utah

Sediment accumulation and water volume in Loch Raven Reservoir, Baltimore County, Maryland

Baltimore City and its metropolitan area are supplied with water from three reservoirs, Liberty Reservoir, Prettyboy Reservoir, and Loch Raven Reservoir. Prettyboy and Loch Raven Reservoirs are located on the Gunpowder Falls (figure 1). The many uses of the reservoir system necessitate coordination and communication among resource managers. The 1996 Amendment to the Safe Drinking Water Act require States to complete source-water assessments for public drinking-water supplies. As part of an ongoing effort to provide safe drinking water and as a direct result of these laws, the City of Baltimore and the Maryland Department of the Environment (MDE), in cooperation with other State and local agencies, are studying the Gunpowder Falls Basin and its role as a source of water supply to the Baltimore area. As a part of this study, the U.S. Geological Survey (USGS), in cooperation with the Maryland Geological Survey (MGS), with funding provided by the City of Baltimore and MDE, is examining sediment accumulation in Loch Raven Reservoir. The Baltimore City Department of Public Works periodically determines the amount of water that can be stored in its reservoirs. To make this determination, field crews measure the water depth along predetermined transects or ranges. These transects provide consistent locations where water depth, or bathymetric, measurements can be made. Range surveys are repeated to provide a record of the change in storage capacity due to sediment accumulation over time. Previous bathymetric surveys of Loch Raven Reservoir were performed in 1943, 1961, 1972, and 1985. Errors in data-collection and analysis methods have been assessed and documented (Baltimore City Department of Public Works, 1989). Few comparisons can be made among survey results because of changing data-collection techniques and analysis methods.

Water-Resources Investigations Report

The multiple-comparison trap and the Raven’s paradox—perils of using null hypothesis testing in environmental assessment

Detecting and quantifying environmental thresholds is frequently an important step in understanding ecological responses to environmental stressors. We discuss two statistical issues often encountered in threshold detection and quantification when statistical null hypothesis testing is used as a main analytical tool.The hidden multiple-comparison trap (leading to a much higher risk of a false detection) and Raven’s paradox(rendering a "detection" meaningless) are often obscured when statistical hypothesis testing is used as part of a more elaborate model, especially models based on computer-intensive methods. Using two examples, we show that the hidden multiple-comparison trap can be exposed using computer simulation to estimate the probability of making a false detection; Raven’s paradox can be avoided by clearly stating the null and alternative hypotheses using scientific terms to substantiate that the rejection of the null is equivalent to proving that the alternative of interest is true. The hidden multiple-comparison trap implies that a null hypothesis testing based on a computer-intensive method should be used with caution. The implication of Raven’s paradox re-quires that we focus on providing evidence supporting the proposed hypothesis or model, rather than seeking evidence against the frequently irrelevant null hypothesis. These two problems, and many others related to null hypothesis testing, suggest that statistical hypothesis testing should be used only as a component of the body of evidence, perhaps, as the devil’s advocate.

Environmental Monitoring and Assessment

A decision tool to identify population management strategies for common ravens and other avian predators

Some avian species have developed the capacity to leverage resource subsidies associated with human manipulated landscapes to increase population densities in habitats with naturally low carrying capacities. Elevated corvid densities and new territory establishment have led to an unsustainable increase in depredation pressure on sympatric native wildlife prey populations as well as in crop damage. Yet, subsidized predator removal programs aimed at reducing densities are likely most effective longer-term when conducted in tandem with subsidy control, habitat management, and robust assessment monitoring programs. We developed decision support software that leverages stage structured Lefkovitch population matrices to compare and identify treatment strategies that reduce subsidized avian predator densities most efficiently, in terms of limiting both cost and take levels. The StallPOPd (Version 4; available at https://doi.org/10.7298/sk2e-0c38.4) software enables managers to enter the area of their management stratum and the demographic properties (vital rates) of target bird population(s) of interest to evaluate strategies to decrease or curtail further population growth. Strategies explicitly include the reduction in fertility (i.e., eggs hatched) and/or the culling of hatchlings, non-breeders and/or breeders, but implicitly comprise reduction in survival or reproduction through subsidy denial. We illustrate the utilities of the software with examples using common ravens ( Corvus corax; ravens) in the Mojave Desert of California, USA. Unfortunately, the survival and reproduction effects of each unit of a particular subsidy in that system have remained elusive, though this is the priority of current research. Because the software leverages a life history representation that is known to characterize hundreds of wildlife species in addition to ravens, the work expands the suite of tools available to wildlife managers and agricultural industry specialists to abate bird damage and impacts on sensitive wildlife in habitats with persistent human subsidies.

Human-Wildlife Interactions