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Ecological neighborhoods as a framework for umbrella species selection

Umbrella species are typically chosen because they are expected to confer protection for other species assumed to have similar ecological requirements. Despite its popularity and substantial history, the value of the umbrella species concept has come into question because umbrella species chosen using heuristic methods, such as body or home range size, are not acting as adequate proxies for the metrics of interest: species richness or population abundance in a multi-species community for which protection is sought. How species associate with habitat across ecological scales has important implications for understanding population size and species richness, and therefore may be a better proxy for choosing an umbrella species. We determined the spatial scales of ecological neighborhoods important for predicting abundance of 8 potential umbrella species breeding in Nebraska using Bayesian latent indicator scale selection in N-mixture models accounting for imperfect detection. We compare the conservation value measured as collective avian abundance under different umbrella species selected following commonly used criteria and selected based on identifying spatial land cover characteristics within ecological neighborhoods that maximize collective abundance. Using traditional criteria to select an umbrella species resulted in sub-maximal expected collective abundance in 86% of cases compared to selecting an umbrella species based on land cover characteristics that maximized collective abundance directly. We conclude that directly assessing the expected quantitative outcomes, rather than ecological proxies, is likely the most efficient method to maximize the potential for conservation success under the umbrella species concept.

Biological Conservation↗

Feather isotope analysis reveals differential patterns of habitat and resource use in populations of white-winged doves

The white-winged dove ( Zenaida asiatica ) serves an important ecological role as a diurnal pollinator of the saguaro cactus in the Sonoran desert and an economic role as a highly sought after game bird in North America. White-winged doves are intimately linked to anthropogenic changes on the landscape and because of this, have experienced dramatic population fluctuations over the last 75 years in response, both positively and negatively, to anthropogenic changes on the landscape. To understand the factors driving population growth and decline of migratory species like the white-winged dove, it is imperative we study resource use on both their breeding and wintering grounds. To understand how populations are distributed on the wintering grounds, we tested an alternative to band recovery approaches by using stable isotope analysis. Before we could use isotope analysis to link breeding and wintering locations for this species, we first needed to determine if hydrogen (δ 2 H) and carbon (δ 13 C) stable isotopes in feather tissue (δ 2 H f and δ 13 C f , respectively) could differentiate among populations of white-winged doves across their breeding range in Texas, New Mexico, and Arizona. δ 2 H f and δ 13 C f not only differentiated between populations of white-winged doves that breed in the United States, but δ 2 H f also provided further differentiation in white-winged doves that breed in native Sonoran Desert and agricultural habitats in the western portion of their range. Ecological processes associated with desert resources and anthropogenic influences, specifically saguaro cacti and irrigated crops, largely determined δ 2 H f in some white-winged doves in Arizona whereas δ 2 H of precipitation (δ 2 H p ) largely determined δ 2 H f of doves in New Mexico and Texas. This study highlights the usefulness of stable isotope analysis to differentiate populations of animals across the landscape and the insight isotopes can provide into habitat and resource use. Published 2015. This article is a U.S. Government work and is in the public domain in the USA.

Arizona, California, New Mexico, Texas↗

A serosurvey of diseases of free-ranging gray wolves (Canis lupus) in Minnesota

We tested serum samples from 387 free-ranging wolves ( Canis lupus ) from 2007 to 2013 for exposure to eight canid pathogens to establish baseline data on disease prevalence and spatial distribution in Minnesota's wolf population. We found high exposure to canine adenoviruses 1 and 2 (88% adults, 45% pups), canine parvovirus (82% adults, 24% pups), and Lyme disease (76% adults, 39% pups). Sixty-six percent of adults and 36% of pups exhibited exposure to the protozoan parasite Neospora caninum . Exposure to arboviruses was confirmed, including West Nile virus (37% adults, 18% pups) and eastern equine encephalitis (3% adults). Exposure rates were lower for canine distemper (19% adults, 5% pups) and heartworm (7% adults, 3% pups). Significant spatial trends were observed in wolves exposed to canine parvovirus and Lyme disease. Serologic data do not confirm clinical disease, but better understanding of disease ecology of wolves can provide valuable insight into wildlife population dynamics and improve management of these species.

Minnesota↗

An overview of the Southern Nevada Agency Partnership science and research synthesis

Maintaining and restoring the diverse ecosystems and resources that occur in southern Nevada in the face of rapid socio-economic and ecological change presents numerous challenged to Federal land managers. Rapid population growth since the 1980s, the land uses associated with that growth, and the interactions of those uses with the generally dry and highly variable climate result in numerous stresses to ecosystems, species, and cultural resource. In addition, climate models predict that the rate of temperature increase and, thus, changes in ecological processes, will be highest for ecosystems like the Mojave Desert. The Southern Nevada Agency Partnership (SNAP; http:www.SNAP.gov) was established in 1999 to address common issues pertaining to public lands in southern Nevada. Partners include the Bureau of Land Management, National Park Service, U.S. Fish and Wildlife Service, and USDA Forest Service and they work with each other, the local community, and other partners. SNAP agencies manage more than seven million acres of public lands in southern Nevada (95% of the land area). Federal land includes two national recreation areas, two national conservation area, four national wildlife refuges, 18 congressionally designated wilderness areas, five wilderness study areas, and 22 areas of critical environmental concern. The partnership's activities are mainly centered in Southern Nevada's Clark County (fig. 1.1), but lands managed by SNAP partner agencies also include portions of Lake Mead National Recreation Area in Mohave County, Arizona, U.S. Fish and Wildlife Service, and USDA Forest Service-managed lands in Lincoln and Nye Counties, Nevada, and all lands and activities managed by the Southern Nevada District Office of the Bureau of Land Management. These lands encompass nine distinct ecosystem types (fig. 1.2), support multiple species of management concern an 17 listed species, and are rich in cultural and historic resource. This introductory executive summary discusses the Science and Research Strategy developed by the SNAP agencies, the Science and Research Report, and need for science-based management in southern Nevada.

Nevada↗

Consideraciones para la estimacion de abundancia de poblaciones de mamiferos. [Considerations for the estimation of abundance of mammal populations.]

Estimation of abundance of mammal populations is essential for monitoring programs and for many ecological investigations. The first step for any study of variation in mammal abundance over space or time is to define the objectives of the study and how and why abundance data are to be used. The data used to estimate abundance are count statistics in the form of counts of animals or their signs. There are two major sources of uncertainty that must be considered in the design of the study: spatial variation and the relationship between abundance and the count statistic. Spatial variation in the distribution of animals or signs may be taken into account with appropriate spatial sampling. Count statistics may be viewed as random variables, with the expected value of the count statistic equal to the true abundance of the population multiplied by a coefficient p. With direct counts, p represents the probability of detection or capture of individuals, and with indirect counts it represents the rate of production of the signs as well as their probability of detection. Comparisons of abundance using count statistics from different times or places assume that the p are the same for all times or places being compared (p= pi). In spite of considerable evidence that this assumption rarely holds true, it is commonly made in studies of mammal abundance, as when the minimum number alive or indices based on sign counts are used to compare abundance in different habitats or times. Alternatives to relying on this assumption are to calibrate the index used by testing the assumption of p= pi, or to incorporate the estimation of p into the study design.

Mastozoologia Neotropical / Journal of Neotropical↗

Ecology of Florida black bears in the Okefenokee-Osceola ecosystem

The population status of the Florida black bear ( Ursus americanus floridanus ) is problematic within many portions of its range and its potential listing as a federally threatened species has been the subject of legal debate. We studied Florida black bears in 2 areas in the Okefenokee-Osceola ecosystem in southeast Georgia (i.e.,Okefenokee) and north Florida (i.e., Osceola) from 1995 to 1999 to evaluate relationships between population characteristics, habitat conditions, and human activities. Bears in Okefenokee were hunted and those in Osceola were not. We captured 205 different black bears (124M:81F) 345 times from June 1995 to September 1998. We obtained 13,573 radiolocations from 87 (16M:71F) individual bears during the study. In Okefenokee, black gum ( Nyssa sylvatica ) and saw palmetto ( Serenoa repens ) fruits were the most important foods for bears based on scat analysis. In Osceola, corn from white-tailed deer ( Odocoileus virginianus ) feeders was the most stable food source but saw palmetto was heavily used when available. Corn from deer feeders was not available in Okefenokee. Adult bears in Osceola were 29% heavier than those in Okefenokee ( t 82 = 3.55, P <0.001). The mean annual home-range size for Osceola females ( x̄ =30.3 km 2 &plusmn; 4.0 [SE], n =53) varied little seasonally or annually and was almost half that of Okefenokee females (55.9 km 2 &plusmn; 6.9, n = 69; Z = &ndash;2.47, P = 0.014). In contrast, radiocollared females in Okefenokee expanded their home ranges during years of poor black gum production. That expansion was most apparent between autumn 1998 and 1999, when mean home-range size for Okefenokee females increased from 14.5 km 2 to 78.4 km 2 , respectively, and included a larger proportion of upland areas open to sport hunting. As a result, 5 females were harvested in the Okefenokee study area during the 1999 bear hunting season compared with only 7 harvested from 1996 to 1998. Home ranges of adult female bears were located in areas with disproportionately high loblolly bay ( Gordonia lasianthus ) and gum-bay-cypress ( Taxodium spp.) vegetation associations in Okefenokee and gum-bay-cypress associations in Osceola. The pine vegetation association ranked lower than most other associations within the home ranges of bears in both study areas even though much of the summer and autumn diets of bears included food items found almost exclusively in pine. Sixteen mortalities of radiocollared bears were documented in Okefenokee; hunting accounted for 11 (68.8%) of these deaths. The annual survival rate of radiocollared males in Okefenokee was 0.71 (95% CI = 0.53&ndash;0.88) whereas survival of females in Okefenokee was higher ( Z =18.87, P <0.001) at 0.89 (95% CI = 0.83&ndash;0.95). The survival rate for females in Osceola was 0.97 (95% CI = 0.92&ndash;1.00). Overall, 67 bears (51M:16F) were killed by hunters in the Okefenokee study area from 1995 to 1999. Based on all radiocollared bears in Okefenokee, many of which resided within areas closed to hunting, we estimated an annual harvest rate of 0.22 (95% CI = 0.03&ndash;0.37) for males and 0.07 (95% CI = 0.01&ndash;0.12) for females. When we excluded those bears that were not in areas open to hunting, however, the annual harvest rate rose to 0.37 (95% CI = 0.07&ndash;0.58) for males and 0.39 (95% CI = 0.09&ndash;0.58) for females. Following a black gum shortage in autumn 1995, only 1 of 15 radiocollared females in Okefenokee produced cubs in winter 1996. That low reproductive rate was in contrast to winter 1997, which followed heavy black gum production, when 21 of 22 radiocollared females produced cubs. Reproductive output was more consistent in the Osceola study area, with 46 cubs being produced from 8, 5, and 9 litters in 1997, 1998, and 1999, respectively. To estimate population size, we maintained 88 and 94 barbed-wire hair traps during 1999 in the Okefenokee and Osceola study areas, respectively. Using DNA collected at the hair traps, mark&ndash;recapture models produced estimates of 71 bears (95% CI = 59&ndash;91) in Okefenokee and 44 bears (95% CI = 40&ndash;57) in the Osceola study area during 1999. The estimated densities in the Okefenokee and Osceola study areas were 0.12 and 0.14 bears/km 2 , respectively. Alternative density estimates based on the amount of time bears spent within study area boundaries were similar (0.11 and 0.14 bears/km 2 on Okefenokee and Osceola, respectively). We used a population model to estimate the effect of harvest in the Okefenokee bear population. Excluding harvest, bears at Osceola experienced higher average annual population growth (&lambda; = 1.184 &plusmn; 0.002) than those at Okefenokee (1.064 &plusmn; 0.002; t 18 = 3.93, P = 0.001), most likely due to protection from hunting and higher recruitment. Including the effects of emigration and immigration, we estimated an average annual sustainable harvest at Okefenokee of approximately 9 bears (12.6%), which was slightly less than the average 1995&ndash;1999 annual harvest of 9.4. That level of hunting in Okefenokee is sustainable but likely represents the highest exploitation rate in the region. Our mark&ndash;recapture data from Osceola suggest a high dispersal rate by subadult bears, and our population modeling data support this hypothesis; we documented bears in Okefenokee that originated from Osceola but not the reverse. We speculate that bears from the interior of the Okefenokee National Wildlife Refuge (ONWR), and to some extent northern Florida, served as a source to the population sink caused by hunting mortality in Okefenokee and in the surrounding Georgia counties. Corn from deer feeders was the most probable reason for smaller home-range sizes and greater body masses and reproductive output at Osceola. Changes in management to eliminate or reduce baiting for deer with corn would negatively affect the Osceola bear population. On Okefenokee, sporadic black gum and palmetto production influenced harvest rates and cub production and, thus, governed bear population dynamics. The U.S. Fish and Wildlife Service (USFWS) concluded in 1998 that listing the Florida black bear as threatened under the Endangered Species Act of 1973 was not warranted. That decision was largely based on the stability and protection afforded to a few subpopulations within the range of the subspecies, which includes the Okefenokee-Osceola subpopulation; our results support that conclusion. However, we suggest that metapopulation processes among the various subpopulations be given greater consideration, with the ultimate goal of managing the sub-species as a unit rather than as an assemblage of independent components. Our study illustrates the importance of travel corridors for maintaining metapopulation processes.

Florida, Georgia↗

The importance of functional form in optimal control solutions of problems in population dynamics

Optimal control theory is finding increased application in both theoretical and applied ecology, and it is a central element of adaptive resource management. One of the steps in an adaptive management process is to develop alternative models of system dynamics, models that are all reasonable in light of available data, but that differ substantially in their implications for optimal control of the resource. We explored how the form of the recruitment and survival functions in a general population model for ducks affected the patterns in the optimal harvest strategy, using a combination of analytical, numerical, and simulation techniques. We compared three relationships between recruitment and population density (linear, exponential, and hyperbolic) and three relationships between survival during the nonharvest season and population density (constant, logistic, and one related to the compensatory harvest mortality hypothesis). We found that the form of the component functions had a dramatic influence on the optimal harvest strategy and the ultimate equilibrium state of the system. For instance, while it is commonly assumed that a compensatory hypothesis leads to higher optimal harvest rates than an additive hypothesis, we found this to depend on the form of the recruitment function, in part because of differences in the optimal steady-state population density. This work has strong direct consequences for those developing alternative models to describe harvested systems, but it is relevant to a larger class of problems applying optimal control at the population level. Often, different functional forms will not be statistically distinguishable in the range of the data. Nevertheless, differences between the functions outside the range of the data can have an important impact on the optimal harvest strategy. Thus, development of alternative models by identifying a single functional form, then choosing different parameter combinations from extremes on the likelihood profile may end up producing alternatives that do not differ as importantly as if different functional forms had been used. We recommend that biological knowledge be used to bracket a range of possible functional forms, and robustness of conclusions be checked over this range.

Ecology↗

Marine ecoregion and Deepwater Horizon oil spill affect recruitment and population structure of a salt marsh snail

Marine species with planktonic larvae often have high spatial and temporal variation in recruitment that leads to subsequent variation in the ecology of benthic adults. Using a combination of published and unpublished data, we compared the population structure of the salt marsh snail, Littoraria irrorata , between the South Atlantic Bight and the Gulf Coast of the United States to infer geographic differences in recruitment and to test the hypothesis that the Deepwater Horizon oil spill led to widespread recruitment failure of L. irrorata in Louisiana in 2010. Size-frequency distributions in both ecoregions were bimodal, with troughs in the distributions consistent with a transition from sub-adults to adults at ~13 mm in shell length as reported in the literature; however, adult snails reached larger sizes in the Gulf Coast. The ratio of sub-adults to adults was 1.5–2 times greater in the South Atlantic Bight than the Gulf Coast, consistent with higher recruitment rates in the South Atlantic Bight. Higher recruitment rates in the South Atlantic Bight could contribute to higher snail densities and reduced adult growth in this region. The ratio of sub-adults to adults in Louisiana was lower in 2011 than in previous years, and began to recover in 2012–2014, consistent with widespread recruitment failure in 2010, when large expanses of spilled oil were present in coastal waters. Our results reveal an important difference in the ecology of a key salt marsh invertebrate between the two ecoregions, and also suggest that the Deepwater Horizon oil spill may have caused widespread recruitment failure in this species and perhaps others with similar planktonic larval stages.

Ecosphere↗

Invasive Brook Stickleback Culaea inconstans minimally alters the trophic ecology of four native fishes in Wyoming, USA

Invasive species introductions are a primary threat facing populations of native freshwater fishes. There are multiple mechanisms by which an invader can affect native species, with competition for food resources being one mechanism that can lead to declines in the distribution and abundance of native species. Invaders that are trophic generalists may cause shifts in the trophic ecology of native species and may be better suited for long-term persistence amid environmental stochasticity. Therefore, trophic studies can provide valuable information on the risk an invader poses to native species. Brook Stickleback Culaea inconstans is an invasive fish species in Wyoming whose effect on native fish assemblages is poorly understood. Our goal was to understand the potential for competitive interactions between Brook Stickleback and native fishes. We used stable isotopes of carbon (ẟ 13 C) and nitrogen (ẟ 15 N) to evaluate the feeding ecology of Brook Stickleback relative to four native fishes, and to explore whether native fish isotopic niches changed in sympatry with Brook Stickleback. We hypothesized that the isotopic niche of Brook Stickleback would be larger than that of native fishes, suggesting broader resource use. Additionally, we hypothesized that the isotopic niche of native fish populations sympatric with Brook Stickleback would contract. We did not find support for our hypotheses as the isotopic niche of Brook Stickleback was not substantially different from that of native fishes. Further, the isotopic niche of native fishes was not substantially affected by Brook Stickleback presence. As a result, we do not currently see evidence of Brook Stickleback altering the trophic ecology of native fish species. Our results provide insight to the effects of a small-bodied invasive fish species on native fishes in a previously unstudied region, and can help managers prioritize management actions to conserve native fishes.

Wyoming↗

Moving forward in circles: Challenges and opportunities in modeling population cycles

Population cycling is a widespread phenomenon, observed across a multitude of taxa in both laboratory and natural conditions. Historically, the theory associated with population cycles was tightly linked to pairwise consumer–resource interactions and studied via deterministic models, but current empirical and theoretical research reveals a much richer basis for ecological cycles. Stochasticity and seasonality can modulate or create cyclic behaviour in non-intuitive ways, the high-dimensionality in ecological systems can profoundly influence cycling, and so can demographic structure and eco-evolutionary dynamics. An inclusive theory for population cycles, ranging from ecosystem-level to demographic modelling, grounded in observational or experimental data, is therefore necessary to better understand observed cyclical patterns. In turn, by gaining better insight into the drivers of population cycles, we can begin to understand the causes of cycle gain and loss, how biodiversity interacts with population cycling, and how to effectively manage wildly fluctuating populations, all of which are growing domains of ecological research.

Ecology Letters↗

Diverse patterns of migratory timing, site use, and site fidelity by Alaska-breeding Whimbrels

Birds that conduct long-distance migrations exhibit varied patterns of consistency in migratory timing and site use. Understanding variation in these traits among populations can help uncover mechanisms driving migratory behaviors and identify potential population threats. Whimbrels ( Numenius phaeopus ) are long-distance migratory shorebirds with a Holarctic breeding distribution, and recent studies have documented population-specific migrations that vary in duration (short to long) and frequency of stops (none/few to multiple). Factors driving these population-specific differences are unclear. We studied the migration ecology of Whimbrels breeding in Alaska, USA, using satellite transmitters deployed from 2006 to 2010 and tracked through 2015. Whimbrels moved entirely within the Pacific Americas Flyway, and some conducted nonstop flights that exceeded seven days across ~ 8700 km. Birds dispersed across numerous sites throughout the flyway, often using agriculture or aquaculture habitats. Whimbrels generally exhibited fidelity to breeding and non-breeding sites, but typically only exhibited fidelity to staging sites used prior to long, nonstop migratory flights. The duration of migration for Whimbrels at more southern non-breeding locations was longer than for those at more northern non-breeding sites, and birds at more southern sites also terminated southbound migration later and initiated northbound migration earlier than birds at more northern sites. Alaska-breeding Whimbrels exhibited greater variation in migratory behaviors than those in other populations in the species’ range. We attribute this within-population diversity to the extensive breadth of non-breeding distributions (~ 70° latitude across ~ 8600 km), a range that naturally shaped individual responses to unequal migratory demands.

Alaska↗

Pleistocene glacial cycles drove lineage diversification and fusion in the Yosemite toad (Anaxyrus canorus)

Species endemic to alpine environments can evolve via steep ecological selection gradients between lowland and upland environments. Additionally, many alpine environments have faced repeated glacial episodes over the past two million years, fracturing these endemics into isolated populations. In this “glacial pulse” model of alpine diversification, cycles of allopatry and ecologically divergent glacial refugia play a role in generating biodiversity, including novel admixed (“fused”) lineages. We tested for patterns of glacial pulse lineage diversification in the Yosemite toad ( Anaxyrus [ Bufo ] canorus ), an alpine endemic tied to glacially influenced meadow environments. Using double‐digest RADseq on populations densely sampled from a portion of the species range, we identified nine distinct lineages with divergence times ranging from 18 to 724 thousand years ago (ka), coinciding with multiple Sierra Nevada glacial events. Three lineages have admixed origins, and demographic models suggest these fused lineages have persisted throughout past glacial cycles. Directionality indices supported the hypothesis that some lineages recolonized Yosemite from east of the ice sheet, whereas other lineages remained in western refugia. Finally, refugial niche reconstructions suggest that low‐ and high‐elevation lineages have convergently adapted to similar climatic niches. Our results suggest glacial cycles and refugia may be important crucibles of adaptive diversity across deep evolutionary time.

California↗

Estimating functional connectivity of wildlife habitat and its relevance to ecological risk assessment

Habitat fragmentation is a major threat to the viability of wildlife populations and the maintenance of biodiversity. Fragmentation relates to the sub-division of habitat into disjunct patches. Usually coincident with fragmentation per se is loss of habitat, a reduction in the size of the remnant patches, and increasing distance between patches. Natural and anthropogenic processes leading to habitat fragmentation occur at many spatial scales, and their impacts on wildlife depend on the scales at which species interact with the landscape. The concept of functional connectivity captures this organism-based view of the relative ease of movement or degree of exchange between physically disjunct habitat patches. Functional connectivity of a given habitat arrangement for a given wildlife species depends on details of the organism's life history and behavioral ecology, but, for broad categories of species, quantities such as home range size and dispersal distance scale allometrically with body mass. These relationships can be incorporated into spatial analyses of functional connectivity, which can be quantified by indices or displayed graphically in maps. We review indices and GIS-based approaches to estimating functional connectivity, presenting examples from the literature and our own work on mammalian distributions. Such analyses can be readily incorporated within an ecological risk framework. Estimates of functional connectivity may be useful in a screening-level assessment of the impact of habitat fragmentation relative to other stressors, and may be crucial in detailed population modeling and viability analysis.

ASTM Selected Technical Papers↗

Velocity-based movement modeling for individual and population level inference

Understanding animal movement and resource selection provides important information about the ecology of the animal, but an animal's movement and behavior are not typically constant in time. We present a velocity-based approach for modeling animal movement in space and time that allows for temporal heterogeneity in an animal's response to the environment, allows for temporal irregularity in telemetry data, and accounts for the uncertainty in the location information. Population-level inference on movement patterns and resource selection can then be made through cluster analysis of the parameters related to movement and behavior. We illustrate this approach through a study of northern fur seal ( Callorhinus ursinus ) movement in the Bering Sea, Alaska, USA. Results show sex differentiation, with female northern fur seals exhibiting stronger response to environmental variables.

Alaska↗

Incorporating citizen science data in spatially explicit integrated population models

Information about population abundance, distribution, and demographic rates is critical for understanding a species’ ecology and for effective conservation and management. To collect data over large spatial and temporal extents for such inferences, especially for species with low densities or wide distributions, citizen science can be an efficient approach. Integrated models have also emerged as an important methodology to estimate population parameters by combining multiple types of data, including citizen science data. We developed a spatially explicit integrated model that combines opportunistically collected presence–absence (PA) data, commonly collected in citizen science efforts, with systematically collected spatial capture–recapture (SCR) data, which are often limited to small spatial and temporal extents. We conducted single and multi‐season simulations with parameters informed by North American black bear ( Ursus americanus ) populations, to evaluate the influence of varying amounts of opportunistic PA data collected at larger spatial and temporal extents on the estimation of population‐level parameters. Integrating opportunistic PA data increased the precision and accuracy of posterior estimates of abundance, and survival and recruitment rates. In some cases, adding PA locations improved abundance estimates more than increasing PA detection probability. Posterior estimates were as precise and unbiased as when higher quality, but sparse, SCR data were available. We also applied the integrated model to SCR and citizen science PA data collected on black bears in New York, with results consistent with our simulations. Our findings indicate that citizen science in integrated models can be a cost‐efficient way to improve estimates of population parameters and increase the spatiotemporal extent of inference. Continued developments with integrated models and citizen science data will offer additional ways to improve our understanding of population structure and demographics.

Ecology↗

Complex trophic interactions in kelp forest ecosystems

The distributions and abundances of species and populations change almost continuously. Understanding the processes responsible is perhaps ecology’s most fundamental challenge. Kelp-forest ecosystems in southwest Alaska have undergone several phase shifts between alga- and herbivore-dominated states in recent decades. Overhunting and recovery of sea otters caused the earlier shifts. Studies focusing on these changes demonstrate the importance of top-down forcing processes, a variety of indirect food-web interactions associated with the otter-urchin-kelp trophic cascade, and the role of food-chain length in the coevolution of defense and resistance in plants and their herbivores. This system unexpectedly shifted back to an herbivore-dominated state during the 1990s, because of a sea-otter population collapse that apparently was driven by increased predation by killer whales. Reasons for this change remain uncertain but seem to be linked to the whole-sale collapse of marine mammals in the North Pacific Ocean and southern Bering Sea. We hypothesize that killer whales sequentially "fished down" pinniped and sea-otter populations after their earlier prey, the great whales, were decimated by commercial whaling. The dynamics of kelp forests in southwest Alaska thus appears to have been influenced by an ecological chain reaction that encompassed numerous species and large scales of space and time.

Bulletin of Marine Science↗

Connectivity of Mojave Desert tortoise populations—Management implications for maintaining a viable recovery network

Executive Summary The historic distribution of Mojave desert tortoises ( Gopherus agassizii ) was relatively continuous across the range, and the importance of tortoise habitat outside of designated tortoise conservation areas (TCAs) to recovery has long been recognized for its contributions to supporting gene flow between TCAs and to minimizing impacts and edge effects within TCAs. However, connectivity of Mojave desert tortoise populations has become a concern because of recent and proposed development of large tracts of desert tortoise habitat that cross, fragment, and surround designated conservation areas. This paper summarizes the underlying concepts and importance of connectivity for Mojave desert tortoise populations by reviewing current information on connectivity and providing information to managers for maintaining or enhancing desert tortoise population connectivity as they consider future proposals for development and management actions. Maintaining an ecological network for the Mojave desert tortoise, with a system of core habitats (TCAs) connected by linkages, is necessary to support demographically viable populations and long-term gene flow within and between TCAs. There are four points for wildlife and land-management agencies to consider when making decisions that could affect connectivity of Mojave desert tortoise populations (for example, in updating actions in resource management plans or amendments that could help maintain or restore functional connectivity in light of the latest information): Management of all desert tortoise habitat for persistence and connectivity . Desert tortoise populations continue to decline within most TCAs, and it is unlikely that trends are better in populations outside protected areas. Fragmentation exacerbates negative population trends by breaking large continuous populations into smaller isolated populations. Connectivity within large populations can enhance resilience to localized disturbances due to rescue by neighboring individuals. In contrast, smaller fragmented populations are resistant to rescue by their isolation and thus could suffer irreversible declines to extirpation from a variety of threats and stochastic events. Enhanced threat reduction to reverse declines within TCAs and to maintain occupied habitat in the surrounding matrix would help reduce the variability in population growth rates and improve the resilience of protected populations even while implementing efforts to improve connectivity. Each TCA has unique strengths and weaknesses regarding its ability to support minimum sustainable populations based on areal extent and its ability to support population increases based on landscape connection with adjacent populations. Considering how proposed projects (inside or outside of TCAs) affect connectivity and the ability of TCAs to support at least 5,000 adult tortoises (the numerical goal for each TCA) could help managers to maintain the resilience of TCAs to population declines. The same project, in an alternative location, could have very different impacts on local and regional populations. For example, within the habitat matrix surrounding TCAs, narrowly delineated corridors may not allow for natural population dynamics if they do not accommodate overlapping home ranges along most of their widths so that tortoises reside, grow, find mates, and produce offspring that can replace older tortoises. In addition, most habitat outside TCAs may receive more surface disturbance than habitat within TCAs. Therefore, managing the entire remaining matrix of desert tortoise habitat for permeability may be better than delineating fixed corridors. These concepts apply, especially given uncertainty about long-term condition of habitat, within and outside of TCAs under a changing climate. Ultimately, questions such as “ What are the critical linkages that need to be protected ?” could be better framed as “ How can we manage the remaining habitat matrix in ways that sustain ecological processes and habitat suitability for special status species ?” Land-management decisions made in the context of the latter question may be more conducive to maintenance of a functional ecological network. Limitations on landscape-level disturbance across habitat managed for the desert tortoise Clearly delineating habitat linkages and differentiating them from non-delineated areas by the uses that are permitted or prohibited within them by specific management guidelines can help achieve functional connectivity. Such guidelines would be most effective if they considered and accounted for all surface disturbances (for example, temporary disturbances such as fiberoptic lines or off-highway vehicle routes, right-of-ways, utility-scale solar development, urbanization) to the extent possible. A weighted framework that varies with the permanence or severity of the disturbance, and can be additive to quantify cumulative effects, could be useful (Xiong, 2020). For example, minor roads can alter tortoise movements independently of other features (Peaden and others, 2017; Hromada and others, 2020), but if the isolated dirt road is accompanied by a powerline that encourages raven predation (Xiong, 2020), then the two features together may be additive. Ignoring minor or temporary disturbance on the landscape could result in a cumulatively large impact that is not explicitly acknowledged (Goble, 2009); therefore, understanding and quantifying all surface disturbance on a given landscape is prudent. In California, the Bureau of Land Management established 0.1–1.0 percent caps on new surface-disturbance for TCAs and mapped linkages that address the issues described in number 1 of this list. Nevada, Utah, and Arizona currently do not have surface-disturbance limits. Limits comparable to those in the Desert Renewable Energy Conservation Plan (DRECP) would be 0.5 percent within TCAs and 1 percent within the linkages modeled by Averill-Murray and others (2013). Limits in some areas of California within the Desert Renewable Energy Conservation Plan, such as Ivanpah Valley, are more restrictive, at 0.1 percent. Continuity across the state line in Nevada could be achieved with comparable limits in the adjacent portion of Ivanpah Valley, as well as the Greater Trout Canyon Translocation Area and the Stump Springs Regional Augmentation Site. These more restrictive limits would help protect remaining habitat in the major interstate connectivity pathway through Ivanpah Valley and focal areas of population augmentation that provide additional population connectivity along the western flank of the Spring Mountains. In a recent study that analyzed 13 years of desert tortoise monitoring data, nearly all desert tortoise observations were at sites in which 5 percent or less of the surrounding landscape within 1 kilometer was disturbed (Carter and others, 2020a). To help maintain tortoise habitability and permeability across all other non-conservation-designated tortoise habitat, all surface disturbance could be limited to less than 5-percent development per square kilometer because the 5-percent threshold for development is the point at which tortoise occupation drops precipitously (Carter and others, 2020a). However, although individual desert tortoises were observed at development levels up to 5 percent, we do not know the fitness or reproductive characteristics of these individuals. This level of development also may not allow for long-term persistence of healthy populations that are of adequate size needed for demographic or functional connectivity; therefore, a conservative interpretation suggests that, ideally, development could be lower. Lower development levels would be particularly useful in areas within the upper 5th percentile of connectivity values modeled by Gray and others (2019). Reducing ancillary threats in places where connectivity is restricted to narrow strips of habitat, for example, narrow mountain passes or vegetated strips between solar development, could enhance the functionality of these vulnerable linkages. In such areas, maintaining multiple, redundant linkages could further enhance overall connectivity. Minimization of mortality from roads and maximization of passage under roads . Roads pose a significant threat to the long-term persistence of local tortoise populations, and roads of high traffic volume lead to severe population declines, which ultimately fragments populations farther away from the roads. Three points (a.–c.) pertain to reducing direct mortality of tortoises on the many paved roads that cross desert tortoise habitat and to maintaining a minimal level of permeability across these roads: Tortoise-exclusion fencing tied into culverts, underpasses, overpasses, or other passages below roads in desert tortoise habitat, would limit vehicular mortality of tortoises and provide opportunities for movement across the roads. Installation of shade structures on the habitat side of fences installed in areas with narrow population-depletion zones would limit overheating of tortoises that may pace the fence. Passages below highways could be maintained or retrofitted to ensure safe tortoise access, for example, by filling eroded drop-offs or modifying erosion-control features such as rip-rap to make them safer and more passable for tortoises. Wildlife management agencies could work with transportation departments to develop construction standards that are consistent with hydrologic/erosion management goals, while also incorporating a design and materials consistent with tortoise survival and passage and make the standards widely available. The process would be most effective if the status of passages was regularly monitored and built into management plans. Healthy tortoise populations along fenced highways could be supported by ensuring that land inside tortoise-exclusion fences is not so degraded that it leads to degradation of tortoise habitat outside the exclusion areas. For example, severe invasive plant infestations inside a highway exclusion could cause an increase of invasive plants outside the exclusion area and degrade habitat; therefore, invasive plants inside road rights of way could be mown or treated with herbicide to limit their spread into adjacent tortoise habitat and minimize the risk of these plants carrying wildfires into adjacent habitat. Adaptation of management based on new information . Future research will continue to build upon and refine models related to desert tortoise population connectivity and develop new ones. New models could consider landscape levels of development and be constructed such that they share common foundations to support future synthesis efforts. If model development was undertaken in partnership with entities that are responsible for management of desert tortoise habitat, it would facilitate incorporation of current and future modeling results into their land management decisions. There are specific topics that may be clarified with further evaluation: The effects of climate change on desert tortoise habitat, distribution, and population connectivity; The effects of large-scale fires, especially within repeatedly burned habitat, on desert tortoise distribution and population connectivity; The ability of solar energy facilities or similar developments to support tortoise movement and presence by leaving washes intact; leaving native vegetation intact whenever possible, or if not possible, mowing the site, allowing vegetation to re-sprout, and managing weeds; and allowing tortoises to occupy the sites; and The design and frequency of underpasses necessary to maintain functional demographic and genetic connectivity across linear features, like highways.

Arizona, California, Nevada↗

At-sea feeding ecology of parasitic lampreys

This paper synthesizes information on the at-sea ecology of ten anadromous lampreys, with emphasis on trophic ecology. The at-sea ecology of these lampreys concerns the juvenile stage, in which growth is most rapid. Anadromous lampreys can be categorized into four groups, based on feeding modalities: 1) scavenger (Caspian lamprey, Caspiomyzon wagneri ); 2) parasite-predator (Pacific lamprey, Entosphenus tridentatus ); 3) predators (western river lamprey, Lampetra ayresii ; European river lamprey, L. fluviatilis ; Arctic lamprey, Lethenteron camtschaticum ; pouched lamprey, Geotria australis ; and Argentinian pouched lamprey, G. macrostoma ); and 4) parasites (sea lamprey, Petromyzon marinus ; Chilean lamprey, Mordacia lapicida ; and short-headed lamprey, M. mordax ). This paper discusses direct evidence for lamprey feeding ecology, as observed through lamprey-induced wounds on hosts and prey, and lamprey attachments on hosts and prey; and indirect evidence for feeding ecology, via analyses of fatty acids, stable isotopes, contaminants, and bioenergetics modelling. A part of the information presented on feeding ecology is from landlocked sea lamprey, and in some instances this information can be generalizable to anadromous populations. For most anadromous lampreys, but particularly for Southern Hemisphere taxa, little is known about their feeding ecology at sea. Duration of the trophic marine phase and habitat use are still subjects of debate. Species identified as lamprey hosts can be demersal or pelagic, possibly reflecting marine habitat preferences. To unlock understanding of the marine phase of anadromous lampreys, direct evidence of feeding ecology should be coupled with natural (i.e., biomarkers) and artificial (e.g., biologgers) markers to identify habitat use, movement patterns and dispersal.

Journal of Great Lakes Research↗