Factors associated with black bear density and implications for management

Wildlife density estimates are important to accurately formulate population management objectives and understand the relationship between habitat characteristics and a species’ abundance. Despite advances in density and abundance estimation methods, management of common game species continues to be challenged by a lack of reliable population estimates. In Washington, USA, statewide American black bear (Ursus americanus) abundance estimates are predicated on density estimates derived from research in the 1970s and are hypothesized to be a function of precipitation and vegetation, with higher densities in western Washington. To evaluate current black bear density and landscape relationships in Washington, we conducted a 4-year capture-recapture study in 2 areas of the North Cascade Mountains using 2 detection methods, non-invasive DNA collection and physical capture and deployment of global positioning system (GPS) collars. We integrated GPS telemetry from collared bears with spatial capture-recapture (SCR) data and created a SCR-resource selection model to estimate density as a function of spatial covariates and test the hypothesis that density is higher in areas with greater vegetative food resources. We captured and collared 118 bears 132 times and collected 7,863 hair samples at hair traps where we identified 537 bears from 1,237 detections via DNA. The most-supported model in the western North Cascades depicted a negative relationship between black bear density and an index of human development. We estimated bear density at 20.1 bears/100 km², but density varied from 13.5/100 km² to 27.8 bears/100 km² depending on degree of human development. The model best supported by the data in the eastern North Cascades estimated an average density of 19.2 bears/100 km², which was positively correlated with primary productivity, with resulting density estimates ranging from 7.1/100 km² to 33.6 bears/100 km². The hypothesis that greater precipitation and associated vegetative production in western Washington supports greater bear density compared to eastern Washington was not supported by our data. In western Washington, empirically derived average density estimates (including cubs) were nearly 50% lower than managers expected prior to our research. In eastern Washington average black bear density was predominantly as expected, but localized areas of high primary productivity supported greater than anticipated bear densities. Our findings underscore the importance that black bear density is not likely uniform and management risk may be increased if an average density is applied at too large a scale. Disparities between expected and empirically derived bear density illustrate the need for more rigorous monitoring to understand processes that affect population numbers throughout the jurisdiction, and suggest that management plans may need to be reevaluated to determine if current harvest strategies are achieving population objectives. © 2019 The Wildlife Society.     

空间直观景观模型的验证方法

空间直观景观模型已是当前景观生态学研究的一大热点。空间景观模型模拟空间格局变化。其模拟结果包含非空间数据和空间数据。空间直观景观模型的验证除进行非空间数据的验证外,还需要进行空间数据的验证。本文回顾了空间直观模型发展历程,总结现有的空间直观模型验证方法。包括主观评价、图形比较、偏差分析、回归分析、假设检验、多尺度拟合度分析和景观指数分析,同时提出今后空间直观景观模型验证方法研究的重点方向。     

像元尺度上不确定性对空间景观直观模型模拟的影响

LANDIS模型是模拟自然和人为干扰下森林景观变化的空间直观景观模型。模型把景观概念化为由相同大小的像元或样地组成的格网。在每一个像元上,模型要求输入物种和年龄组信息。但是,由于研究区一般由成千上百万个像元构成,不可能通过实际调查获取每一个像元上的物种和年龄组信息。因此,采用了一种基于小班的随机赋值法从森林调查数据中获取每一个像元的物种和年龄组信息。该方法是一种基于概率的方法,会在LANDIS模型模拟的物种和年龄组信息的输入中引入不确定性。为了评价由基于小班的随机赋值法所引入像元尺度上的不确定性对模型模拟结果的影响,用蒙特卡罗模拟法进行不确定性分析。对LANDIS模型模拟的每一个物种,用众数年龄组发生频率来定量化单个像元上年龄组信息的不确定性,用所有像元上的众数年龄组平均发生频率来定量化年龄组信息在像元尺度上总的不确定性。平均发生频率越高,不确定性越低。为了评价基于小班的随机赋值法对景观尺度上模型模拟结果的影响,计算了每一个物种在整个研究区内的面积百分比和聚集度指数。变异系数越大,不确定性越高。对所有物种,年龄组信息不确定性在模型模拟的初期是比较低的(平均发生频率大于10)。种子传播、建群、死亡和火干扰使模型结果的不确定性随模拟时间增加而增加。最后,不确定性达到稳定状态,达到平衡状态的时间与物种寿命接近。此时,初始的物种和年龄组信息不再对模型结果有影响。在景观尺度上,物种分布面积百分比和由聚集度指数所定量化的空间格局并未受像元尺度上不确定性增加的影响。因为LANDIS模型模拟研究的目的在于预测总的景观格局变化,而不是单一的事件,所以,基于小班的随机赋值法可用于LANDIS模型的参数化。     

Spatial variation in density of American black bears in northern Yellowstone National Park

The quality and availability of resources are known to influence spatial patterns of animal density. In Yellowstone National Park, relationships between the availability of resources and the distribution of grizzly bears (Ursus arctos) have been explored but have yet to be examined in American black bears (Ursus americanus). We conducted non-invasive genetic sampling during 2017–2018 (mid-May to mid-July) and applied spatially explicit capture-recapture models to estimate density of black bears and examine associations with landscape features. In both years, density estimates were higher in forested vegetation communities, which provide food resources and thermal and security cover preferred by black bears, compared with non-forested areas. In 2017, density also varied by sex, with female densities being higher than males. Based on our estimates, the northern range of Yellowstone National Park supports one of the highest densities of black bears (20 black bears/100 km²) in the northern Rocky Mountains (6–12 black bears/100 km² in other regions). Given these high densities, black bears could influence other wildlife populations more than previously thought, such as through displacement of sympatric predators from kills. Our study provides the first spatially explicit estimates of density for black bears within an ecosystem that contains the majority of North America's large mammal species. Our density estimates provide a baseline that can be used for future research and management decisions of black bears, including efforts to reduce human–bear conflicts.     

A spatially explicit model of patchy stomatal responses to humidity

Stomata of leaves can exhibit either temporally stable, spatially homogeneous behaviour or complex spatial and temporal dynamics, depending on environmental and physiological conditions. To test the ability of accepted physiological mechanisms to describe these patterns, we developed a simple, spatially explicit model of stomatal responses to humidity that incorporated hydraulic interactions among stomata. Model results showed qualitative agreement with experimental evidence for a number of phenomena: (1) at high humidities, whole-leaf steady-state conductance is a monotonic function of humidity; (2) the initial stomatal response following a perturbation in humidity is in the direction opposite to the final response, and (3) spatial dynamics include patch formation and self-organization similar to that observed in actual leaves. These comparisons do not eliminate other explanations, but do suggest that novel mechanisms need not be invoked to explain the diversity of spatial and temporal patterns of stomatal behaviour in leaves.     

Home ranges in adult Scandinavian brown bears (Ursus arctos): effect of mass, sex, reproductive category, population density and habitat type

Annual home-range size indices for 36 male and 52 female adult brown bears Ursus arctos in two study areas in central and northern Scandinavia were estimated to evaluate factors believed to influence home-range size. Male home ranges were larger than home ranges of lone females after controlling for the sexual size dimorphism acting on metabolic needs. Further, home ranges of females with cubs were smaller than home ranges of lone females and females with yearlings. Thus, differences in metabolic need were not able to explain the variation in range size among females of different reproductive categories or between males and females, suggesting roaming behaviour of males in this promiscuous species. Home-range size in both males and females was inversely related to population density along a density gradient that was not linked to food availability. This contradicts the hypothesis that females use the minimum areas that sustain their energy requirements. However, on a large geographical scale a negative relationship between range size and food availability was evident. The annual home ranges in inland boreal environments in Scandinavia are the largest reported for brown bears in Eurasia, and similar to those in inland boreal and montane environments in North America.     

Estimates of Abundance and Harvest Rates of Female Black Bears Across a Large Spatial Extent

American black bears (Ursus americanus) are an iconic wildlife species in the southern Appalachian highlands of the eastern United States and have increased in number and range since the early 1980s. Given an increasing number of human-bear conflicts in the region, many management agencies have liberalized harvest regulations to reduce bear populations to socially acceptable levels. Wildlife managers need reliable population data for assessing the effects of management actions for this high-profile species. Our goal was to use DNA extracted from hair collected at barbed-wire enclosures (i.e., hair traps) to identify individual bears and then use spatially explicit capture-recapture methods to estimate female black bear density, abundance, and harvest rate. We established 888 hair traps across 66,678 km² of the southern Appalachian highlands in Georgia, North Carolina, South Carolina, and Tennessee, USA, in 2017 and 2018, arranged in 174 clusters of 2–9 traps/cluster. We collected 9,113 hair samples from those sites over 6 weeks of sampling, of which 1,954 were successfully genotyped to 462 individual female bears. Our spatially explicit estimator included a percent forest covariate to explain inhomogeneous bear density across the region. Densities ranged up to 0.410 female bears/km² and regional abundance was 5,950 (95% CI = 4,988–7,098) female bears. Based on hunter kill data from 2016 to 2018, mean annual harvest rates for females were 12.7% in Georgia, 17.6% in North Carolina, 17.6% in South Carolina, and 22.8% in Tennessee. Our estimated harvest rates for most states approached or exceeded theoretical maximum sustainable levels, and population trend data (i.e., bait-station indices) indicated decreasing growth rates since about 2009. These data suggest that the increased harvest goals and poor hard mast production over a series of prior years reduced bear population abundance in many states. We were able to obtain reasonable population abundance and density estimates because of spatially explicit capture-recapture methods, cluster sampling, and a large spatial extent. Continued monitoring of bear populations (e.g., annual bait-station surveys and periodic population estimation using spatially explicit methods) by state jurisdictions would help to ensure that population trajectories are consistent with management goals. © 2021 The Wildlife Society.     

A spatially explicit capture–recapture estimator for single‐catch traps

Single‐catch traps are frequently used in live‐trapping studies of small mammals. Thus far, a likelihood for single‐catch traps has proven elusive and usually the likelihood for multicatch traps is used for spatially explicit capture–recapture (SECR) analyses of such data. Previous work found the multicatch likelihood to provide a robust estimator of average density. We build on a recently developed continuous‐time model for SECR to derive a likelihood for single‐catch traps. We use this to develop an estimator based on observed capture times and compare its performance by simulation to that of the multicatch estimator for various scenarios with nonconstant density surfaces. While the multicatch estimator is found to be a surprisingly robust estimator of average density, its performance deteriorates with high trap saturation and increasing density gradients. Moreover, it is found to be a poor estimator of the height of the detection function. By contrast, the single‐catch estimators of density, distribution, and detection function parameters are found to be unbiased or nearly unbiased in all scenarios considered. This gain comes at the cost of higher variance. If there is no interest in interpreting the detection function parameters themselves, and if density is expected to be fairly constant over the survey region, then the multicatch estimator performs well with single‐catch traps. However if accurate estimation of the detection function is of interest, or if density is expected to vary substantially in space, then there is merit in using the single‐catch estimator when trap saturation is above about 60%. The estimator's performance is improved if care is taken to place traps so as to span the range of variables that affect animal distribution. As a single‐catch likelihood with unknown capture times remains intractable for now, researchers using single‐catch traps should aim to incorporate timing devices with their traps.     

Combining genetic non-invasive sampling with spatially explicit capture-recapture models for density estimation of a patchily distributed small mammal

Estimating the size of animal populations is essential for understanding the demography and conservation status of species. Genetic Non-Invasive Sampling (gNIS) combined with Spatially Explicit Capture-Recapture (SECR) modelling may provide a practical tool to obtain such estimates. Here, we evaluate for the first time the potential and limitations of this approach to estimate population densities for small mammals inhabiting patchily distributed habitats, focusing on the endemic Iberian Cabrera vole (Microtus cabrerae). Using 11 highly polymorphic microsatellites and two sex-linked introns, we compared population estimates in November/December 2011 based on live-trapping and gNIS and assessed the impact of distinct consensus criteria to differentiate unique genotypes. Live-trapping over 21 days captured 31 individuals, while gNIS over 5 days recorded 65–69 individuals. SECR models indicated that individual detectability was positively affected by live-trapping capture success on the previous occasion, while for gNIS, it was mainly affected by genotyping success rates and patch size. Live-trapping produced the lowest density estimates (mean ± SE) of 16.6?±?3.2 individuals per hectare of suitable habitat (ind/ha). Estimates based on gNIS were higher and varied slightly between 25.2?±?4.0 and 28.8?±?4.5 ind/ha depending on assuming one or two genotyping errors, respectively, when differentiating individual genetic profiles. Results suggest that live-trapping underestimated the vole population, while the larger number of individuals detected through gNIS allowed better estimates with lower field effort. Overall, we suggest that gNIS combined with SECR models provides an effective tool to estimate small mammal population densities in fragmented habitats.     

Modelling food web complexity: The consequences of individual-based, spatially explicit behavioural ecology on trophic interactions

We present a prototype simulator that enables one to explore the influence of individual behaviour on the dynamics and structural complexity of food webs. In the simulations, individuals act according to simple, biologically plausible rules in a spatially explicit setting. We present the results of a series of simulation experiments on artificial, tri-trophic level food chains used to calibrate the simulator against real-world systems and to demonstrate the simulators promise for ecological modelling. Our primary objective was to discover the biological features leading to stability of artificial food chains over ecological time and under different conditions of trophic efficiency. This involved a qualitative analysis of food chains comprised of a plant, a herbivore and a carnivore species. We explored the consequences of allowing individual heterotrophs to make active choices about resource selection (perception and intentional behaviour) under high and low degrees of trophic efficiency. We found that individuals had to adopt realistic behavioural ecological strategies, such as active resource selection, for systems to persist, especially under conditions in which trophic efficiencies were of the magnitude observed in real systems (e.g. 10%). Our results reaffirm previous convictions that a better understanding of food web interactions in real-world systems will require approaches that blend animal behavioural ecology with population and community ecology. However, the evidence comes from a new mathematical perspective.     

青海沙蜥种群密度调查的一种新方法

在2002年9月,分别利用洞穴深度法、标志重捕法和洞口计数法对分布在若尔盖草原荒漠中的青海沙蜥(Phrynocephalus vlangalii)的种群密度进行调查,所得的结果分别是190.4、76.8和250.7只.1000 m-2。通过对3种结果的比较与分析,证明洞穴深度法有较高的可靠性,这种方法的理论依据是:沙蜥(Phrynocephalus)是一种变温动物,不能长时间处在温度低于致死低温(-2.5℃)以下的环境中,为了能够成功地度过漫长而寒冷的冬季,它必须居住在深度达到最大冻土层之下的洞穴中。这是沙蜥躲避低温伤害的一种行为机制。该方法可以适用于分布在中国的沙蜥属的其它物种密度调查。     

空间直观景观模型LANDIS在大兴安岭呼中林区的应用

应用空间直观景观模型(LANDIS),研究有采伐和无采伐预案下大兴安岭呼中林区的森林景观的长期变化。用APACK计算每一个物种及各年龄级的分布面积。为了研究物种分布格局的变化,计算了物种分布的聚集度指数。研究结果如下：(1)在无采伐预案下。火干扰模式为低频率大面积高强度火烧;在有采伐预下,火干扰模式为高频率小面积低强度火烧;(2)在无采伐预案下,火会造成各种群分布面积的强烈波动,但是对种群的年龄结构没有很大影响;在有采伐预案下,火对种群分布面积和年龄结构都没有很大的影响;(3)采伐能完全改变各种群的年龄结构。降低种群分布的聚集度,但是对各种群的分布面积并没有很大影响;(4)在有采伐预案下,各种群为增长型种群,增长量通过采伐取走,群落处于演替的干扰顶极状态;在无采伐预案下,各种群为稳定型种群(樟子松和偃松除外),大面积高强度火烧使群落产生较大的波动。结果表明,在呼中林业局,在没有人为干扰情况下,火干扰是森林景观变化的主导因素。自从有了人为干扰,采伐开始逐渐取代火干扰成为影响森林景观变化的主导因素。空间直观景观模型的一个挑战是模型的验证。由于缺乏详细的空间数据及模型模拟中的随机性,很难通过模型模拟结果与实地调查或遥感数据的比较进行验证。通过对火模拟、物种分布和物种组成的生态或生物学实现对模型进行验证。     

Estimating animal population density using passive acoustics

Reliable estimation of the size or density of wild animal populations is very important for effective wildlife management, conservation and ecology. Currently, the most widely used methods for obtaining such estimates involve either sighting animals from transect lines or some form of capture‐recapture on marked or uniquely identifiable individuals. However, many species are difficult to sight, and cannot be easily marked or recaptured. Some of these species produce readily identifiable sounds, providing an opportunity to use passive acoustic data to estimate animal density. In addition, even for species for which other visually based methods are feasible, passive acoustic methods offer the potential for greater detection ranges in some environments (e.g. underwater or in dense forest), and hence potentially better precision. Automated data collection means that surveys can take place at times and in places where it would be too expensive or dangerous to send human observers. Here, we present an overview of animal density estimation using passive acoustic data, a relatively new and fast‐developing field. We review the types of data and methodological approaches currently available to researchers and we provide a framework for acoustics‐based density estimation, illustrated with examples from real‐world case studies. We mention moving sensor platforms (e.g. towed acoustics), but then focus on methods involving sensors at fixed locations, particularly hydrophones to survey marine mammals, as acoustic‐based density estimation research to date has been concentrated in this area. Primary among these are methods based on distance sampling and spatially explicit capture‐recapture. The methods are also applicable to other aquatic and terrestrial sound‐producing taxa. We conclude that, despite being in its infancy, density estimation based on passive acoustic data likely will become an important method for surveying a number of diverse taxa, such as sea mammals, fish, birds, amphibians, and insects, especially in situations where inferences are required over long periods of time. There is considerable work ahead, with several potentially fruitful research areas, including the development of (i) hardware and software for data acquisition, (ii) efficient, calibrated, automated detection and classification systems, and (iii) statistical approaches optimized for this application. Further, survey design will need to be developed, and research is needed on the acoustic behaviour of target species. Fundamental research on vocalization rates and group sizes, and the relation between these and other factors such as season or behaviour state, is critical. Evaluation of the methods under known density scenarios will be important for empirically validating the approaches presented here.     

Genetic estimates of annual reproductive success in male brown bears: the effects of body size, age, internal relatedness and population density

1. We studied male yearly reproductive success (YRS) and its determinants (phenotypic characteristics, age, population density) in two Scandinavian brown bear populations, using molecular techniques to determine paternity. 2. We found a significant difference in male YRS between the study areas, with lower YRS in the south than in the north. 3. In general, older and larger males had higher YRS. Older males may be more experienced in competition for reproduction (male dominance). Large body size is of direct benefit in male-male competition and of advantage in endurance competition for the access to females. 4. Age was relatively more important for YRS in the north and body size was more important in the south, due perhaps to differences in male age structure due to illegal killing. A single old male dominated the reproduction in the north during the study, which resulted most probably in the relatively higher importance of age in the north. In the south, with a more even male age structure, no single male was able to dominate, probably resulting in a more intense competition among males, with body size as the deciding factor. 5. Male YRS was correlated positively with population density. This may be related to the structure of the expanding bear population, with female densities declining towards the population edge. 6. Internal relatedness, a measure of genetic heterozygosity, was correlated negatively with YRS, suggesting that outbred individuals have a higher YRS. Individual heterozygosity at key or many loci may reflect male physical qualities and condition-sensitive traits, which may benefit males directly in contest or in sperm competition.