Spring wildflowers spectacular in 2003 at the Desert Tortoise Natural Area
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Tortoise Tracks↗
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The tortoise and the antilocaprid: Adapting GPS tracking and terrain data to model wildlife walking functions
Context The relationship between slope and terrestrial animal locomotion is key to landscape ecology but underexplored across species. This is partly due to a lack of scalable methodology that applies to a diversity of wildlife. Objectives This study investigates the slope-speed relationship for two species, Texas tortoise ( Gopherus berlandieri ) and pronghorn ( Antilocapra americana ), through the combined application of remote sensing, GPS tracking, behavior models, and parametric distribution. While using readily available Digital Elevation Models (DEM) for pronghorn, we explore the use of very high-resolution lidar Digital Terrain Models (DTM) from Unoccupied Aerial Systems (UAS) to characterize tortoise movements at micro-scales. Methods After classifying animal behavior with GPS tracking data and Hidden Markov Models (HMMs), we analyzed the relationship between the speed of the animals and the slope of the terrain using a 30-m DEM for pronghorn, and a fine-scale UAS DTM for Texas tortoise, and three nonlinear models: Laplace, Gauss, and Lorentz. Results High-resolution DTM, coupled with GPS tracking, accurately models the relationship of speed and slope at a micro-scale, while a DEM is suitable for a larger scale. Laplace models best predicted the speed of both the Texas tortoise and pronghorn. Models showed tortoises, which are not known for rapid and agile movement like the pronghorn, have a broader tolerance for varying slopes at a fine scale. Conclusions These findings enhance understanding of species-specific movement offering valuable insights for habitat management and conservation tailored to species’ behaviors and capabilities.
Colorado, Texas, Wyoming↗
Desert tortoises ( Gopherus agassizii ) are selective herbivores that track the flowering phenology of their preferred food plants
Previous studies of desert tortoise foraging ecology in the western Mojave Desert suggest that these animals are selective herbivores, which alter their diet according to the temporal availability of preferred food plants. These studies, however, did not estimate availability of potential food plants by taking into account the spatial and temporal variability in ephemeral plant abundance that occurs within the spring season. In this study, we observed 18 free-ranging adult tortoises take 35,388 bites during the spring foraging season. We also estimated the relative abundance of potential food plants by stratifying our sampling across different phenological periods of the 3-month long spring season and by different habitats and microhabitats. This methodology allowed us to conduct statistical tests comparing tortoise diet against plant abundance. Our results show that tortoises choose food plants non-randomly throughout the foraging season, a finding that corroborates the hypothesis that desert tortoises rely on key plants during different phenological periods of spring. Moreover, tortoises only consumed plants in a succulent state until the last few weeks of spring, at which time most annuals and herbaceous perennials had dried and most tortoises had ceased foraging. Many species of food plants—including several frequently eaten species—were not detected in our plant surveys, yet tortoises located these rare plants in their home ranges. Over 50% of bites consumed were in the group of undetected species. Interestingly, tortoises focused heavily on several leguminous species, which could be nutritious foods owing to their presumably high nitrogen contents. We suggest that herbaceous perennials, which were rare on our study area but represented ~30% of tortoise diet, may be important in sustaining tortoise populations during droughts when native annuals are absent. These findings highlight the vulnerability of desert tortoises to climate change if such changes alter the availability of their preferred food plants.
California↗
Survival and movements of head‐started Mojave desert tortoises
Head‐starting is a conservation strategy in which young animals are protected in captivity temporarily before their release into the wild at a larger size, when their survival is presumably increased. The Mojave desert tortoise ( Gopherus agassizii ) is in decline, and head‐starting has been identified as one of several conservation measures to assist in recovery. To evaluate the efficacy of indoor head‐starting, we released and radio‐tracked 68 juvenile tortoises from a 2015 cohort in the Mojave National Preserve, California, USA. We released 20 tortoises at hatching (control) in September 2015, and reared 28 indoors and 20 outdoors in predator‐proof enclosures for 7 months before releasing them in April 2016. We monitored tortoises at least weekly after release until 27 October 2016, and documented survivorship, movement, and surface activity. We estimated survivorship by treatment and evaluated effects of treatment, proximity to a raven ( Corvus corax ) nest (predator) coincidentally established after release, distance moved between monitoring events, surface activity, and release size on individual fate in a generalized linear model. Although indoor head‐start tortoises reached the size of 5–6‐year‐old wild tortoises by release at 7 months of age, survival did not differ significantly among the 3 treatment groups. Combined annual survival was 0.44 (95% CI = 0.34–0.58). Tortoises that were closer to an active raven nest were significantly more likely to die, as were those seen more often outside their burrows and active aboveground. Predicted estimates for short‐term probability of survival approached 1.0 as distance from a raven nest exceeded approximately 1.6 km. Rearing treatment, movement distance, and body size were not significant predictors of fate over the 1‐year monitoring period. Head‐started tortoises released ≥1.6 km from areas of raven activity will likely have higher short‐term survival. Population recovery through head‐starting alone is unlikely to be successful if systemic ecosystem‐level issues, such as habitat degradation and conditions that promote human‐subsidized predators, are not ameliorated. © 2019 The Wildlife Society.
California↗
Habitat selection by juvenile Mojave Desert tortoises
Growing pressure to develop public lands for renewable energy production places several protected species at increased risk of habitat loss. One example is the Mojave desert tortoise ( Gopherus agassizii ), a species often at the center of conflicts over public land development. For this species and others on public lands, a better understanding of their habitat needs can help minimize negative impacts and facilitate protection or restoration of habitat. We used radio-telemetry to track 46 neonate and juvenile tortoises in the Eastern Mojave Desert, California, USA, to quantify habitat at tortoise locations and paired random points to assess habitat selection. Tortoise locations near burrows were more likely to be under canopy cover and had greater coverage of perennial plants (especially creosote [ Larrea tridentata ]), more coverage by washes, a greater number of small-mammal burrows, and fewer white bursage ( Ambrosia dumosa ) than random points. Active tortoise locations away from burrows were closer to washes and perennial plants than were random points. Our results can help planners locate juvenile tortoises and avoid impacts to habitat critical for this life stage. Additionally, our results provide targets for habitat protection and restoration and suggest that diverse and abundant small-mammal populations and the availability of creosote bush are vital for juvenile desert tortoises in the Eastern Mojave Desert.
California↗
Aggregated space use by soft-released translocated Gopher Tortoises (Gopherus polyphemus)
Translocated herpetofauna can exhibit irregular space use and movement patterns when compared with resident conspecifics. In Florida, USA, Gopher Tortoises ( Gopherus polyphemus ) are translocated throughout the state to mitigate habitat loss due to development. The postrelease space use of translocated Gopher Tortoises within soft-release pens can affect population dynamics and population monitoring efficacy, and understanding spatial patterns can aid wildlife managers with population management. We used a combination of time-lapse cameras, animal tracking devices, and burrow distribution surveys to investigate translocated tortoise space use at Eglin Air Force Base, Florida, where tortoises have been translocated since 2015. We investigated 10 soft-release pens that varied in size (4–41 ha) and shape (due to landscape configuration and existing infrastructure). Time-lapse cameras and burrow distribution surveys showed that tortoises used habitat within 20 m of soft-release pens (silt fences) significantly more than the interior of pens. In most pens, the selection of pen-edge habitat resulted in a clustering effect that lessened upon subsequent surveys, after fences were removed. Additionally, our tracking data showed mixed evidence for clustering, where three of the seven tortoises used edge area significantly more than the interior of pens. Such clustering can affect the efficacy of population survey methods while potentially having negative impacts on the health of translocatees by increasing local density.
Florida↗
Protection benefits desert tortoise ( Gopherus agassizii ) abundance: the influence of three management strategies on a threatened species
We surveyed an area of ∼260 km 2 in the western Mojave Desert to evaluate relationships between condition of Agassiz's Desert Tortoise populations ( Gopherus agassizii ) and habitat on lands that have experienced three different levels of management and protection. We established 240 1-ha plots using random sampling, with 80 plots on each of the three types of managed lands. We conducted surveys in spring 2011 and collected data on live tortoises, shell-skeletal remains, other signs of tortoises, perennial vegetation, predators, and evidence of human use. Throughout the study area and regardless of management area, tortoise abundance was positively associated with one of the more diverse associations of perennial vegetation. The management area with the longest history of protection, a fence, and legal exclusion of livestock and vehicles had significantly more live tortoises and lower death rates than the other two areas. Tortoise presence and abundance in this protected area had no significant positive or negative associations with predators or human-related impacts. In contrast, the management area with a more recent exclusion of livestock, limited vehicular traffic, and with a recent, partial fence had lower tortoise densities and high death rates. Tortoise abundance here was negatively associated with vehicle tracks and positively associated with mammalian predators and debris from firearms. The management area with the least protection—unfenced, with uncontrolled vehicle use, sheep grazing, and high trash counts—also had low tortoise densities and high death rates. Tortoise abundance was negatively associated with sheep grazing and positively associated with trash and mammalian predator scat. cat.
California↗
Feral burros and other influences on desert tortoise presence in the western Sonoran Desert
Across the globe, conflicting priorities exist in how land and resources are managed. In the American West, conflicts are common on public lands with historical mandates for multiple uses. We explored the impacts of multiple uses of land in a case study of Agassiz's Desert Tortoises ( Gopherus agassizii ), a federally threatened species, in the western Sonoran Desert. The tortoise has declined for many reasons, most of which relate to management of land and habitat. Frequently cited causes are livestock grazing, roads, vehicle-oriented recreation, predators, and disease. In spring of 2009, we conducted a survey to evaluate relationships between desert tortoises, vegetation associations, topography, predators, and anthropogenic uses. We sampled a 93-km 2 area with 200 independent 1-ha plots. Density (± SE) of adult tortoises was low, 2.0 ± 1.0/km 2 , and the annualized death rate for adults during the 4 yr preceding the survey was high, 13.1%/yr. We observed tortoise sign, most of which was recent, on 22% of the 200 plots, primarily in the southwestern part of the study area. More tortoise sign occurred on plots with Brittlebush ( Encelia spp.) vegetation at higher elevations. Most plots (91.0%) had ≥1 human-related impacts: feral burro scat ( Equus asinus ; 84.0%), recent vehicle tracks and trails (34.0%), trash (28.0%), burro trails and wallows (26.5%), and old vehicle tracks (24.0%). We used a multimodel approach to model presence of tortoise sign on the basis of 12 predictor variables, and calculated model-averaged predictions for the probability of tortoise presence. Importance values revealed two apparent top drivers: feral burros and vegetation association. This is the first study to identify a negative association between presence of desert tortoises and feral burros.
Arizona, California↗
Animal movement in the absence of predation: environmental drivers of movement strategies in a partial migration system
Animal movement strategies including migration, dispersal, nomadism, and residency are shaped by broad-scale spatial-temporal structuring of the environment, including factors such as the degrees of spatial variation, seasonality and inter-annual predictability. Animal movement strategies, in turn, interact with the characteristics of individuals and the local distribution of resources to determine local patterns of resource selection with complex and poorly understood implications for animal fitness. Here we present a multi-scale investigation of animal movement strategies and resource selection. We consider the degree to which spatial variation, seasonality, and inter-annual predictability in resources drive migration patterns among different taxa and how movement strategies in turn shape local resource selection patterns. We focus on adult Galapagos giant tortoises Chelonoidis spp. as a model system since they display many movement strategies and evolved in the absence of predators of adults. Specifically, our analysis is based on 63 individuals among four taxa tracked on three islands over six years and almost 10 6 tortoise re-locations. Tortoises displayed a continuum of movement strategies from migration to sedentarism that were linked to the spatio-temporal scale and predictability of resource distributions. Movement strategies shaped patterns of resource selection. Specifically, migratory individuals displayed stronger selection toward areas where resources were more predictable among years than did non-migratory individuals, which indicates a selective advantage for migrants in seasonally structured, more predictable environments. Our analytical framework combines large-scale predictions for movement strategies, based on environmental structuring, with finer-scale analysis of space-use. Integrating different organizational levels of analysis provides a deeper understanding of the eco-evolutionary dynamics at play in the emergence and maintenance of migration and the critical role of resource predictability. Our results highlight that assessing the potential benefits of differential behavioral responses first requires an understanding of the interactions among movement strategies, resource selection and individual characteristics.
Oikos↗
Drawing a line in the sand: Effectiveness of off-highway vehicle management in California's Sonoran desert
Public land policies manage multiple uses while striving to protect vulnerable plant and wildlife habitats from degradation; yet the effectiveness of such policies are infrequently evaluated, particularly for remote landscapes that are difficult to monitor. We assessed the use and impacts of recreational vehicles on Mojave Desert washes (intermittent streams) in the Chemehuevi Desert Wildlife Management Area (DWMA) of southern California. Wash zones designated as open and closed to off-highway vehicle (OHV) activity were designed in part to protect Mojave desert tortoise ( Gopherus agassizii ) habitat while allowing recreation in designated areas. OHV tracks were monitored in washes located near access roads during winter and early spring holidays – when recreation is typically high – and at randomly dispersed locations away from roads. Washes near access roads had fewer vehicle tracks within closed than open zones; further away from roads, OHV tracks were infrequent and their occurrence was not different between wash designations. Washes were in better condition in closed zones following major holidays as indicated by less vegetation damage, presence of trash, and wash bank damage. Furthermore, the frequency of washes with live tortoises and their sign was marginally greater in closed than open wash zones. Collectively, these results suggest that low impacts to habitats in designated closed wash zones reflect public compliance with federal OHV policy and regulations in the Chemehuevi DWMA during our study. Future monitoring to contrast wash use and impacts during other seasons as well as in other DWMAs will elucidate spatial and temporal patterns of recreation in these important conservation areas.
California↗
Movement ecology
(Yackulic) At first glance, the decision to study movement in Galapagos tortoises seems curious. Given the slow speed of tortoises and tendency to forage and rest as they move, it seems implausible that tortoises would string their slow bursts of activity together to accomplish large-scale movements. Nonetheless, as early as 1815 (Porter 1815), visitors to Galapagos have noted the propensity for tortoises to walk long distances along well used trails, leading to seasonal changes in the distribution of tortoises on the various islands of the Archipelago. In recent years, advances in the technology used to track animals have led to a better understanding of movement and the diversity of movement strategies among many animal taxa. This chapter focuses on the application of these technologies to study of movement in Galapagos tortoises. Recent work has shown not only the diversity of movement strategies employed by Galapagos tortoises, but also illustrated how movement can both cause and be a consequence of interaction between reproductive and foraging ecologies of tortoises and the presence of strong environmental gradients. Understanding how critical unimpeded movement is to the persistence of Galapagos tortoises also informs efforts to maintain connectivity to, and suitable habitat within, tortoise range outside of protected areas.
Book chapter↗
Eastern Indigo snake (Drymarchon couperi) shelter site use In peninsular Florida, USA, and implicatIons for habItat conservatIon
Shelters are critical for many species as protection from predators and extreme temperatures. Successful conservation of reptiles requires understanding both shelter site requirements and availability. The Eastern Indigo Snake (EIS; Drymarchon couperi ) is endemic to the southeastern U.S. and is federally listed. Recovery has focused on maximizing unfragmented landscapes, with less attention on fine-scale features such as shelter sites. In the northern EIS range, Gopher Tortoise ( Gopherus polyphemus ) burrows are used extensively for shelter. Although EIS in peninsular Florida often shelter in tortoise burrows, they also use other shelters where tortoise burrows are scarce or absent. Solely focusing EIS survey and management efforts where Gopher Tortoises are present may overlook occupied habitats and misallocate resources. We investigated the importance of different shelter sites in central Florida using data from radio-tracked EIS. We modeled the use of shelter categories as a function of sex, season, and habitat using Bayesian multinomial Generalized Linear Models. Results showed that EIS in peninsular Florida used Gopher Tortoise burrows across all seasons and habitats. Tortoise burrow use was highest in xeric habitats and lowest in mesic habitats where burrows are most and least abundant, respectively. There was less variability in shelter site use in disturbed habitats and flatwoods. Tortoise burrow use by EIS in the cool season across sexes and habitats in our study was much lower than in southern Georgia. Our results indicate that EIS are less dependent on Gopher Tortoise burrows in peninsular Florida and that suitable habitats with few or no tortoise burrows could still provide conservation value for EIS.
Florida, Georgia↗
Variations in climate drive behavior and survival of small desert tortoises
In the Mojave Desert, timing and amounts of precipitation profoundly affect availability of water and annual plant foods necessary for the threatened Agassiz’s desert tortoise ( Gopherus agassizii ) to survive, especially during prolonged droughts. As part of recovery actions to increase declining populations, we translocated 83 juvenile and young desert tortoises raised in head-start pens for 4–10 years to a new location 15 km away during fall of 2013 and 2014. We tracked them for 9 years during a megadrought, during multiple years of low rainfall and a few years when precipitation neared or exceeded long-term norms. We evaluated behaviors and how precipitation and forage availability affected survival. At the end of the study, 21.6% of tortoises were alive and 6 had grown to adulthood. Annual models of survival indicated that tortoise size was the driving variable in most years, followed by number of repeatedly used burrows during periods of temperature extremes. Other variables affecting survival in ≥1 year were vegetation, movements during the first 2 years post-translocation, and condition index, a measure of health. Tortoises moved more, expanded home ranges, and grew rapidly in years when winter rainfall approached or exceeded long-term norms and annual plants were available to eat. During dry years, movements and growth were limited. Exceptions to this pattern occurred in the last year of study, a dry year: tortoises grew, moved more, and home ranges increased. The increase in size and approaching adulthood may have stimulated greater travelling. Some left the study area, indicating a need for large release areas. We may have aided survival by offering water twice yearly when handling, because some tortoises drank and increased in mass up to 40%. Prolonged droughts and hotter temperatures can limit recovery of populations, reduce survival of young tortoises, and increase the time to maturity.
California↗
Nesting ecology of a population of Gopherus agassizii at a utility-scale wind energy facility in southern California
We investigated the annual nesting ecology of a population of Desert Tortoises (Gopherus agassizii) inhabiting a utility-scale renewable energy (USRE) facility in southern California and compared our results with populations inhabiting relatively undisturbed sites. In 2000, 15 radio-tracked females produced 29 clutches, and 24 nests were monitored to examine nest-site selection, nest predation, hatching success, date of emergence of hatchlings, and hatchling mass and carapace length. Overall, the nesting ecology of the population inhabiting the USRE facility was very similar to other populations of Desert Tortoises inhabiting relatively undisturbed habitats. Oviposition occurred from 12 May to 8 July, which was similar to other sites. Nest depths (11.1 cm), nest predation (12%), hatchling emergence date (7 August and 29 September), and hatchling morphometrics (i.e., MCL: 44.5 mm; mass: 23 g) were all within ranges reported in other populations. Unlike within other populations, we observed no relationship between hatchling size and either maternal body size or egg width. We found no evidence of females selecting for a particular burrow for oviposition of eggs based on environmental or anthropogenic variables. Most nests were located in or near burrows, and nest depth was greater for nests near the entrance than those deeper in the burrow. Although this study suggests that the nesting ecology of the Desert Tortoise population we studied was not adversely affected by the USRE facility, this relationship is only correlative because our study was not a before-after-control-impact (BACI) study, which would establish a cause and effect relationship. As pointed out in a recent review, BACI studies are critically needed to address the wildlife impacts of utility-scale renewable energy development.
California↗
Environmental variation structures reproduction and recruitment in long-lived mega-herbivores: Galapagos giant tortoises
Migratory, long-lived animals are an important focus for life-history theory because they manifest extreme trade-offs in life-history traits: delayed maturity, low fecundity, variable recruitment rates, long generation times, and vital rates that respond to variation across environments. Galapagos tortoises are an iconic example: they are long-lived, migrate seasonally, face multiple anthropogenic threats, and have cryptic early life-history stages for which vital rates are unknown. From 2012 to 2021, we studied the reproductive ecology of two species of Galapagos tortoises ( Chelonoidis porteri and C. donfaustoi ) along elevation gradients that coincided with substantial changes in climate and vegetation productivity. Specifically, we (1) measured the body and reproductive condition of 166 adult females, (2) tracked the movements of 33 adult females using global positioning system telemetry, and monitored their body condition seasonally, (3) recorded nest temperatures, clutch characteristics, and egg survival from 107 nests, and (4) used radiotelemetry to monitor growth, survival, and movements of 104 hatchlings. We also monitored temperature and rainfall from field sites, and remotely sensed primary productivity along the elevation gradient. Our study showed that environmental variability, mediated by elevation, influenced vital rates of giant tortoises, specifically egg production by adult females and juvenile recruitment. Adult females were either elevational migrants or year-round lowland residents. Migrants had higher body condition than residents, and body condition was positively correlated with the probability of being gravid. Nests occurred in the hottest, driest parts of the tortoise's range, between 6 and 165 m elevation. Clutch size increased with elevation, whereas egg survival decreased. Hatchling survival and growth were highest at intermediate elevations. Hatchlings dispersed rapidly to 100–750 m from their nests before becoming sedentary (ranging over <0.2 ha). Predicted future climates may impact the relationships between elevation and vital rates of Galapagos tortoises and other species living across elevation gradients. Resilience will be maximized by ensuring the connectivity of foraging and reproductive areas within the current and possible future elevational ranges of these species.
Galapagos Islands, Santa Cruz Island↗
An uncertain future for a population of desert tortoises experiencing human impacts
We evaluated the status of a population of Mojave Desert Tortoises ( Gopherus agassizii ), a threatened species, in the El Paso Mountains of the northwestern Mojave Desert in California, USA. The study area lies north of and adjacent to a designated critical habitat unit for the species, is adjacent to a state park, and is a short distance from the Desert Tortoise Research Natural Area. We randomly sampled 373 1-ha plots from a 239.1-km 2 area in the mountain range to determine demographic attributes of the population, vegetation associations, predator presence, and human uses. Live and dead G. agassizii and sign (burrows, scats, tracks) occurred on 35.7% of plots. Densities of adults were higher than in adjacent critical habitat, and threats (traumatic injuries, infectious and other diseases) were similar to those reported elsewhere in the geographic range. Signs of human use were evident on 98.4% of plots. We used a multimodel approach to determine distribution of G. agassizii in relation to vegetation, anthropogenic, and predator variables. Vegetation, predators, trash, mining activity, and vehicles were important factors affecting the distribution and intensity of tortoise sign. We concluded that this population is in a downward trend, like other populations in the western Mojave Desert. The high death rate of adults, low population density, high human visitor use, and ongoing decline in the adjacent critical habitat unit indicate that a viable population is unlikely to persist in the study area. The future for the population found in the El Paso Mountains might depend on survival in the adjacent roadless El Paso Mountains Wilderness Area.
California↗
Status of the desert tortoise in Red Rock Canyon State Park
We surveyed for desert tortoises, Gopherus agassizii, in the western part of Red Rock Canyon State Park and watershed in eastern Kern County, California, between 2002 and 2004. We used two techniques: a single demographic plot (~4 km2 ) and 37 landscape plots (1-ha each). We estimated population densities of tortoises to be between 2.7 and 3.57/km2 and the population in the Park to be 108 tortoises. We estimated the death rate at 67% for subadults and adults during the last 4 yrs. Mortality was high for several reasons: gunshot deaths, avian predation, mammalian predation, and probably disease. Historic and recent anthropogenic impacts from State Highway 14, secondary roads, trash, cross-country vehicle tracks, and livestock have contributed to elevated death rates and degradation of habitat. We propose conservation actions to reduce mortality.
California Fish and Game↗
Influence of anthropogenic subsidies on movements of common ravens
Anthropogenic subsidies can benefit populations of generalist predators such as common ravens (ravens; Corvus corax ), which in turn may depress populations of many types of species at lower-trophic levels, including desert tortoises (Gopherus agassizii ) or greater sage-grouse (Centrocercus urophasianus). Management of subsidized ravens often has targeted local breeding populations that are presumed to affect species of concern and ignored “urban” populations of ravens. However, little is known about how ravens move, especially in response to the presence of anthropogenic subsidies. Therefore, subsidized ravens from distant populations that are not managed may influence local prey. To better understand this issue, we deployed global positioning system – global system for mobile communications transmitters to track movements of 19 ravens from September to December 2020 relative to 2 land cover types that provide subsidies: developed areas and cultivated crops. On average, ravens moved 41.5 km (±30.5) per day, although daily movement distances ranged from 0.13– 206.1 km. Raven movement among cover types during the non-breeding season varied widely, with 100% of individuals each using land cover types that provide subsidy and other types at least once in the season. On 100% of days ravens used areas that did not provide subsidy, on 86.7% of days they used developed areas, and on 20.5% of days they used cultivated crops. Although on some days a raven would stay exclusively in areas that did not provide subsidy, there were no days in which a single raven ever stayed exclusively in developed or cultivated crops. Ravens moved shorter distances on days when they used subsidies more frequently. Further, time spent in developed areas and cultivated crops increased when ravens roosted closer to them, although this effect was greater for developed areas than for cultivated crops. Individual ravens were not associated exclusively with either of the subsidy-providing landscapes we considered, but instead all birds used both subsidized and other landscapes. Our research suggests that management of ravens during the non-breeding season and possibly during the breeding season, intended to reduce risk of predation on desert tortoises, will be most effective if conducted on a broad scale because of distances the birds travel and the lack of separation between putative “urban” and “natural” populations of ravens.
California↗