Minimum sampling effort for reliable non-invasive estimations of excretion abundance of Elaphostrongylus cervi L1 in red deer (Cervus elaphus) populations

Faecal surveys are commonly used as non-invasive means to evaluate population abundance of animals as well as comparable indexes of prevalence and intensity of diseases between populations, especially macroparasites. While faecal surveys are among one of the simplest means to perform these evaluations, they are time consuming and labour intensive. The present study evaluated 80 red deer (Cervus elaphus) faecal samples collected in two study sites for the presence and abundance of first-stage larvae of the nematode Elaphostrongylus cervi and established pools of samples for epidemiological analysis. The analysis of 20-30 individual samples would produce a reliable estimate of the 'true' prevalence, and the error of the smaller sample size only doubled that of the 80 reference samples. The analysis of 5 pools of 5 pellet groups each, or of 4 pools of 10 pellet groups each, provided a reliable estimation of the 'true' excretion abundance estimated by the analysis of all 80 samples. These pools comprise a cost-effective and dependable alternative for measuring and obtaining comparable abundances of parasitic faecal excretion stages in red deer populations, which is very valuable for epidemiological and ecological research as well as for management purposes.     

Estimation of roe deerCapreolus capreolus and mouflonOvis aries densities, abundance and habitat use in a mountainous Mediterranean area

The abundance, density, and habitat use of roe deerCapreolus capreolus (Linnaeus, 1758) and mouflonOvis aries Linnaeus, 1758 were studied in a confined Mediterranean area in Greece with a dung survey based on the faecal accumulation rate (FAR) technique. Estimated density was modelled with generalized additive models using altitude, habitat type, and slope as potential covariates. Model selection among the set of candidate models was conducted based on their generalized cross-validation score. Roe deer had an estimated mean density of 13.9 ind./km² and the best model included slope and habitat type as covariates. The mean density of mouflon in the study area was 22.1 ind./km² and the best model used altitude and habitat type as covariates. For both species, the highest densities were encountered in abandoned cultivations and glades, followed by conifer forests, while the lowest densities were observed in maquis. However, use of open habitats by mouflon was much greater than it was for roe deer. The strong preference of mouflon (a grazer species) for open habitats that were abundant with grasses probably reflected food availability and contrasted with the more diverse habitat use by roe deer (a selective browser).     

The disappearance of muntjac (Muntiacus reevesi) and roe deer (Capreolus capreolus) pellet groups in a pine forest of lowland England

Reliable knowledge of the disappearance rate of faecal pellets is essential for converting pellet density to deer density when using standing-crop pellet-group counts. Disappearance of muntjac and roe deer pellet groups was monitored in four growth stages of a pine forest of lowland England over a 15-month period. Time to disappearance of the pellet groups (days) of both species significantly differed between habitats and months; it was shorter in late summer to early autumn and in habitats with more ground vegetation. Muntjac pellet groups disappeared more quickly than roe deer pellet groups. Time to disappearance of roe deer pellet groups was negatively correlated with air and grass temperature in pre-thicket and pre-fell habitats, while time to disappearance of muntjac pellet groups was negatively correlated with frequency of rainfall and positively correlated with the run of wind (air passage over a site within a 24-h period measured in km) in pre-thicket habitats. It is the time of the standing-crop pellet-group counts and the disappearance rate of pellet groups deposited in different months and habitats that determine the appropriate method for conversion of pellet-group density to deer density.     

Modeling the Relationship Between Fecal Pellet Indices and Deer Density

Abstract: Although fecal pellet counts have been widely used to index changes in deer abundance in forests, few studies have modeled the relationship between the indices and deer density. We examined the relationships between 3 fecal pellet indices (total pellets, pellet groups, and pellet frequency) and the density of deer (primarily red deer [Cervus elaphus scoticus]) in 20 enclosures in the North and South islands of New Zealand. In each enclosure we estimated the 3 indices on 30 randomly located 150-m transects, with each transect having 30 circular plots of 3.14 m². We developed 4 candidate models (1 linear and 3 nonlinear) to describe the relationship between the indices and deer density. We used a Bayesian analysis to account for uncertainty in the estimates of deer abundance and to facilitate fitting models that included random transect effects. The 4 models explained the relationship between the 3 indices and deer density similarly well. The slopes of the linear relationships between the 3 indices and deer density were positive. Our results suggest that fecal pellet counts may be useful indices of deer abundance.     

小兴安岭马鹿粪便的分解速率

粪便分解速率是粪便堆在自然条件下消失所用的时间,利用粪便距离取样法估算鹿科动物种群数量,是种群数量管理研究的重要基础。为揭示高纬度温带大陆性气候条件下的鹿科动物粪便分解速率,解决在我国普遍存在的目标种相对隐蔽、种群密度低、粪便分散导致的原地回访观测耗时耗力难以完成的问题,本研究在温带大陆性气候的小兴安岭范围内,选择铁力林业局建设林场为研究区,于2015年1月收集38堆马鹿(Cervus elaphus)的新鲜粪便,将这些粪便堆集中移至方便回访的一处马鹿代表性生境——阳坡针阔混交林,记录每堆粪便的原始粪粒数,之后间隔一定时间进行回访并记录每堆粪便剩余的粪粒数和各粪便堆是否分解完全,共回访8次。利用R 4.1. 2中的逻辑斯蒂回归得出回归方程的截距b0和斜率b1,再利用MATLAB R2020b计算粪便分解速率。结果表明,马鹿粪便的平均分解速率为(646±53) d。本研究将粪便人工移动集中放置于一处进行观测实验,为研究低种群密度的动物粪便分解速率提供了可行途径,丰富了我国鹿科动物粪便分解速率研究,从而减小了利用粪便距离取样法估算种群数量的阻碍。     

Estimating roe deer abundance from pellet group counts in Spain: An assessment of methods suitable for Mediterranean woodlands

Despite of the generalized expansion of wild ungulates in Europe, roe deer (Capreolus capreolus) is experiencing contrasted population trends; it is expanding in some regions while declining in others likely due to the expansion of other deer species. In both extremes, reliable methods to estimate population abundance are required. We evaluated different methods of estimating deer abundance in Mediterranean woodlands based on pellet group counts. Distance Sampling applied to pellet counts and a new easier and cost-effective method based on strip-variable transect counts (FST) were assessed comparing their estimates (pellet group density) with the abundance indices obtained from traditionally used reference methods (faecal standing crop) in 61 localities (n = 183 surveys). The average roe deer density estimated from faecal standing crop was 5.56 ± 0.75 (range 0.01–20.74) deer per 100 ha. Distance Sampling and FST estimates correlated with reference methods. As a first conclusion it may be noted that all indirect methods used here can be used to estimate roe deer abundance. The selection of a given method based on pellet counts to estimate roe deer population abundance should take into account the specific objectives of the research, resources available, and the timescale in which the information is required. Among them, Distance Sampling may be used when human resources and skills are enough but FST is a rapid and efficient alternative to estimate pellet group density when they are not.     

Pellet group count methods to estimate red deer densities: Precision,potential accuracy and efficiency

Pellet group count methods are key tools for wildlife conservation and management. Therefore, it is essential to clarify the performance and suitability of the different pellet group count methods at distinct realities. This issue has been discussed by other researchers but with inconsistent conclusions, leaving open the necessity of additional studies. Aiming to fill this need, we used a combination of field data and simulation models to evaluate the density estimates, precision and potential accuracy of the results obtained through each pellet group count method, and also the sampling effort and efficiency associated with them. The methods evaluated were standing crop plot counts (FSCP), clearance plot counts (FAR), standing crop strip transect counts (ST) and standing crop line transect counts (LT). Deer density estimates by the four methods were statistically similar at all effort levels simulated. The analyses of CV trends reveal a better precision supplied by FSCP than by FAR for the same effort, while ST and LT yielded comparable values in analogous situations. The time required to make a survey is a key factor in the choice of a field technique. LT with distance sampling was the most efficient method to count pellet groups, while FAR seems to be the less proficient method. Attending to the limitations usually inherent to field surveys, like time, technicians and budget, LT appeared more efficient than the other methods, providing great precision and accuracy in less time. Nevertheless, at high densities and pronounced habitat heterogeneity, FAR became more efficient than FSC, do not requiring the decay rates, allowing accurate estimates in a few months when applied with the proper effort and environmental conditions. This study highlights the importance of carrying out pilot studies and simulation models to evaluate the sampling design prior to its implementation in the field.     

新疆哈密山区天山马鹿种群数量和冬季生境选择

2010年12月至2011年2月,在新疆哈密山区,采用截线抽样法和遥感技术,对野生天山马鹿（Cervus elaphus songaricus）种群现状和冬季生境选择进行了研究。在不同区域和不同类型栖息地共布设了28条样线,样线总长度达60.1 km。其中,16条样线上发现马鹿共233头,调查区域平均种群密度（2.83±1.01）头/km²,种群数量（1 684.56±379.71）头,与1993年的调查结果相比有所上升。雌雄性比为2.24：1,幼体和亚成体总数多于成体和老体总数,种群数量呈增长趋势。根据野外考察GPS数据并解译天山马鹿分布生境2006秋季的LANDSAT TM/ETM+遥感影像,将生境要素分为山地针叶林、草甸、灌木丛、农田和戈壁5种类型,其中,草甸与山地针叶林为天山马鹿冬季适宜生境。     

Morphological criteria to identify faecal pellets of sympatric ungulates in West African savanna and estimates of associated error

Indirect surveys may prove to be useful tools in complementing classical direct counts when monitoring ungulate populations and may also promote better understanding of the precise structure and functioning of the rich ungulate communities of African savannas. However, the identification of faecal pellets can be difficult where several sympatric species occur. This study develops simple field criteria for distinguishing between pellets among ten sympatric West African ungulates. A discriminant analysis was performed, using the mean of measurements of pellet groups from different species to pinpoint and characterize the most useful morphological criteria for separation between them. The mean diameter of pellets within each pellet group proved to be the most valuable variable for species segregation, whilst the second axis separated species by mean indent depth. The pellet groups of six of the ten designated species could be identified with a minimum misclassification error. However, no simple morphological variables emerged to permit discrimination between hartebeest and topi, or between bushbuck and Bohor reedbuck pellets. Once pellet groups have been identified, their density and spatial distribution may provide useful information on the use of space and habitat of sympatric species, over given periods.     

Estimating abundance of introduced Chinese muntjac Muntiacus reevesi and native roe deer Capreolus capreolus using portable thermal imaging equipment

• 1 Managing the conservation impacts of deer requires knowledge of their numbers. However, estimating densities in forested areas is problematic, with pellet‐group‐based methods subject to error and uncertainty in estimating defecation and decay rates in addition to sampling variance. Use of thermal imaging equipment allows direct census by distance sampling.
• 2 Densities of introduced Chinese muntjac Muntiacus reevesi and native roe deer Capreolus capreolus were estimated in 12.8 km² of conifer forest in eastern England by thermal imaging distance transects. Estimated density of introduced muntjac (±95% confidence intervals [CI]: 20 ± 8 km⁻²) exceeded that of native roe deer (16 ± 6 km⁻²); 95% CI of c. 40% were achieved in 10 days of fieldwork.
• 3 Density estimates were not sensitive to the number of width bands applied during analysis. Detectability functions differed, with narrower effective strip widths (ESW) for muntjac (74 m) than roe deer (123 m). Thus, it is important to discriminate between species when censusing mixed assemblages. Detectability and ESW also differed among plantation growth stages. Stratification by habitat may therefore improve accuracy of density estimates but would require additional survey effort.

     

Landscape structure affects food quality of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) evidenced by fecal nitrogen levels

Evaluating the quality of wildlife habitat is essential for understanding and predicting population dynamics in heterogeneous environments. We used fecal nitrogen levels as an indicator of habitat quality of sika deer (Cervus nippon) and explored important landscape elements influencing nitrogen levels, taking deer density into account. We established 92 plots differing in deer density and landscape structure on the Boso Peninsula, central Japan, and collected fecal samples along a 1-km transect at each plot. The regression models involving two independent variables, i.e., deer density and the length of forest edge within an area of 100 or 200 m from the transect, were selected based on the Akaike’s Information Criterion (AIC). Levels of fecal nitrogen were positively correlated with the length of the forest edge and negatively correlated with population density of deer. The area of 100 or 200 m from the transect most likely reflected the behavioral scale of the deer. Coverage of palatable understory vegetation increased with proximity to forest edge and decreased with deer density. Variability in the level of fecal nitrogen could thus be explained by food availability in the landscape. These results suggest that landscape alterations increase the carrying capacity of sika deer and thereby increase impacts upon the ecosystem.     

新疆哈密山区天山马鹿(Cervus elaphus songaricus)的种群数量调查*

查清新疆东部哈密山区野生天山马鹿(Cervus elaphus songaricus)种群的现状,可为有效保护与合理利用野生马鹿资源提供科学依据。我们通过截线抽样法和遥感技术,弄清哈密山区天山马鹿的栖息地范围,估算马鹿的种群数量及种群密度。在哈密山区野生天山马鹿分布区域共设计了28条样线,总长度60.1 km,遇见马鹿233头,通过计算得到该地区马鹿天山亚种种群平均密度为(2.83?1.01)头/km2。栖息地面积374.35 km2,估计马鹿总数为(1 057.56?379.71)头左右。雌雄比例2.24:1,幼体和亚成体的总数量多于成体和老体总数。研究区域内马鹿的种群密度和资源总数量比往年的调查结果有所上升;马鹿的分布海拔高度比较一致,但种群的大小具有不均匀性;从性比和年龄结构来看该研究区域的马鹿种群面向着数量的增长趋势。     

Seasonal changes and altitudinal variation in deer fecal pellet decay

Surveying dung pellet groups to estimate population size requires knowledge of the dung decay rates in different environmental conditions. We evaluated seasonal and elevational differences in the decay rates of fresh deer pellets in the Ashio–Nikko Mountains, Japan. At each of four sites (1,500, 1,200, 900, and 600 m above sea level), we set out four replicates of 50 fresh deer pellets each month from September 2008 to August 2009. Dung pellet groups were evaluated after 24 h and again monthly. We also monitored the dung beetle fauna monthly using pitfall traps. Dung beetle activity was positively correlated with temperature, and the rate of dung decay was strongly correlated with dung beetle dry weight. Decay was most rapid during summer and at lower altitudes; pellets set out during winter did not decay until the following spring. Resource managers who use dung surveys to estimate population sizes in mountainous regions should consider the decay rates at different altitudes.     

Spatial and temporal habitat use and selection by red deer: The use of direct and indirect methods

Given the importance of red deer Cervus elaphus for hunting and conservation purposes, understanding the interactions between this species and its habitats in the Mediterranean region is a crucial step for the sustainable management of this species. Aiming to compare pellet group counts and direct observations methods to study the habitat use and selection by red deer, the results obtained by both methods were compared, and their advantages and disadvantages were discussed. To understand the temporal patterns of habitat use and selection, the survey was conducted at three different seasons, birth period, rut season and winter. The habitat use and selection were studied in relation to land cover, watercourse, roads, ecotone zones and other topographic features (altitude, slope and aspect), using generalized linear models and selection ratios. The similarity of the results provided by pellet group counts and direct observations indicate that both methods may constitute useful tools to study the habitat use and selection by red deer. Globally, red deer seemed to select habitats that provide simultaneously food and some cover, as shown by its preference for shrublands, independently of the sampling season. The positive selection of ecotone zones embodies the need for open spaces. Males and females showed a similar use of shrubland, but selected patches with different characteristics therein. The spatial and temporal patterns exhibited by our results suggest that red deer balance their habitat requirements in respect to each phase of their reproductive cycle. Pellet group counts and direct observations seem to be useful methods to analyze habitat use and selection, and may provide helpful knowledge to the management and conservation of red deer.     

Applicability of a Bayesian state-space model for evaluating the effects of localized culling on subsequent density changes: sika deer as a case study

At the landscape scale, localised culling is often conducted to achieve various deer management aims. However, few studies have assessed the effects of localised culling on deer population dynamics, owing to the spatially and temporally insufficient datasets of deer abundance that are derived from limited survey efforts. In this study, we estimated the population dynamics of a sika deer (Cervus nippon) population in the Tanzawa Mountains, central Japan, by Bayesian state-space modelling using spatiotemporally insufficient abundance indices and evaluated the effects of unit-specific culls on subsequent density changes in 56 units. The responses of deer density to unit-specific culls differed greatly among units, and a very weak correlation was observed between the intensities of unit-specific culls and the reduction in density. Deer populations in some units tended to resist density decreases despite high culling pressure, whereas those in other units were susceptible to density decreases with little to no culling pressure. Because the spatial scales of each unit were relatively small, annual density changes in each unit were largely influenced by deer movement in this estimation. The obscured effects of unit-specific culls, which were probably derived from deer movement among units in this case study, re-emphasized that deer migration should be incorporated into the planning of localised culling and that deer management should be coordinated over a wide area beyond landscape components and landownerships. Thus, we conclude that Bayesian state-space modelling is valuable for practical deer management programs at a large spatial scale even if different abundance indices are used.     

Monitoring ungulates in steep non-forest habitat: a comparison of faecal pellet and helicopter counts

Faecal pellet counts have been widely used to monitor the abundances of introduced ungulates in New Zealand, but ground-based sampling cannot be conducted safely in the steep non-forest habitats that are common in New Zealand's Southern Alps. Helicopter counts may be an effective technique for monitoring ungulates in steep non-forest habitat. We evaluated the relationship between faecal pellet and helicopter counts of ungulates (primarily feral goat Capra hircus) at 12 non-forest sites in the Southern Alps. Within each site we counted the numbers of ungulates from a helicopter on three occasions and the number of intact faecal pellets along 30 transects. Mean observed densities of feral goats derived from helicopter counts ranged from 0.0 to 20.2 km^?2. There was a positive curvilinear (concave down) relationship between faecal pellet and helicopter counts. Compared with faecal pellet counts, helicopter counts were cheaper, could identify ungulate species and provided estimates of absolute density. Helicopter counts are a cost-effective method for monitoring ungulates in the steep non-forest habitats of New Zealand's Southern Alps.     

Faecal pellet group counting as a promising method of wild boar population density estimation

The wild boar is a native animal species of the Czech Republic that has significantly increased its population density in recent years. Objective methods of population density estimation are required to inform the intensity of regulation needed for the sustainable management of the wild boar. Population estimation and regulation of the abundance of the wild boar are problematic in comparison to those of other ungulates, and the only widely used method of control is intensive hunting. The purpose of this study was to test if faecal pellet group (FPG) counting was a reliable method of wild boar population censusing in a forest environment. The density of wild boars was evaluated in an area of 22.56 km², circumscribed by both natural and man-made barriers that restrict wild boar migration. As it is essential for proper application of FPG count methods, a separate trial was carried out to determine the daily defecation rate of wild boars. The distribution of faeces in different types of habitat in a forest environment was tested. As the period of faeces accumulation fell within the intensive hunting season, the result is biased by the fact that the calculation includes FPGs produced by animals removed from the population through hunting during winter. The average daily defecation rate of adults and piglets was 4.29 and 5.82, respectively. The corrected (without shot animals) average density of wild boars in the study area estimated by the FPG count at the time of the census was 6.08 individuals (ind.)/km². We have confirmed that an FPG count conducted in winter is applicable for estimation of wild boar abundance on a forested area.     

Using spotlight observations to predict resource selection and abundance for white-tailed deer

Spotlight surveys for white-tailed deer (Odocoileus virginianus) can yield large presence-only datasets applicable to a variety of resource selection modeling procedures. By understanding how populations distribute according to a given resource for a reference area, density and abundance can be predicted across new areas assuming the relationship between habitat quality (measured by an index of selection) and species distribution are equivalent. Habitat-based density estimators have been applied to wildlife species and are useful for addressing conservation and management concerns. Although achieving reliable population estimates is a primary goal for spotlighting studies, presence-only models have yet to be applied to spotlight data for estimating habitat selection and abundance for deer. From 2012 to 2017, we conducted spring spotlight surveys in each of 99 counties in Iowa, USA, and collected spatial locations for 20,149 groups of deer (n = 71,323 individuals). We used a resource selection function (RSF) based on deer locations to predict the relative probability of use for deer at the population level and to estimate statewide abundance. The number of deer observed statewide increased significantly with increasing RSF value for all years and the mean RSF value along survey transects explained 59% of the variability in county-level deer counts, indicating that a functional response between habitat quality and deer distribution existed at landscape scales. We applied our RSF to a habitat-based density estimator (extrapolation) and zero-inflated Poisson (ZIP) and negative binomial (ZINB) count models to predict statewide abundance from spotlight counts. Population estimates for 2012 were variable, indicating that atypical weather conditions may affect spotlight counts and population estimates in some years. For 2013–2017, we predicted a mean population of 439,129 (95% CI ∼ ± 55,926), 440,360 (∼ ± 43,676), and 465,959 (∼ ± 51,242) deer across years for extrapolation, ZIP, and ZINB models, respectively. Estimates from all models were not significantly different than estimates from an existing deer population accounting model in Iowa for 2013 and 2016, and differed by <76,000 deer for all models from 2013–2017. Extrapolation and ZIP models performed similarly and differed by <2,897 deer across all years, whereas ZINB models showed inconsistencies in model convergence and precision of estimates. Our results indicate that presence-only models are capable of producing reliable and precise estimates of resource selection and abundance for deer at broad landscape scales in Iowa and provide a tool for estimating deer abundance in a spatially explicit manner. © 2019 The Wildlife Society.     

The consequences of using indirect signs that decay to determine species' occupancy

Statistical models of species' distributions rely on data on species' occupancy, or use, of sites across space and/or time. For rare or cryptic species, indirect signs, such as dung, may be the only realistic means of determining their occupancy status across broad spatial extents. However, the consequences of sign decay for errors in estimates of occupancy have not previously been considered. If signs decay very rapidly, then false‐negative errors may occur because signs at an occupied site have decayed by the time it is surveyed. On the other hand, if signs decay very slowly, false‐positive errors may occur because signs remain present at sites that are no longer occupied. We addressed this issue by quantifying, as functions of sign decay and accumulation rates: 1) the false‐negative error rate due to sign decay and, 2) the expected time interval prior to a survey within which signs indicate the species was present; as this time interval increases, false‐positives become more likely. We then applied this to the specific example of koala Phascolarctos cinereus occupancy derived from faecal pellet surveys using data on faecal pellet decay rates. We show that there is a clear trade‐off between false‐negative error rates and the potential for false‐positive errors. For the koala case study, false‐negative errors were low on average and the expected time interval prior to surveys that detected pellets indicate the species was present within less than 2–3 yr. However, these quantities showed quite substantial spatial variation that could lead to biased parameter estimates for distribution models based on faecal pellet surveys. This highlights the importance of observation errors arising from sign decay and we suggest some modifications to existing methods to deal with this issue.     

Factors affecting the selection of and displacement within core areas by female mule deer (Odocoileus hemionus) in the Chihuahuan Desert,Mexico

Core activity areas are the most important sites within an animal’s home range as they contain the greatest density of critical resources. Because these areas are small and intensively used, they can affect the total distances covered by animals searching for resources. Our objective was to document which habitat variables affect core area selection by female mule deer (Odocoileus hemionus) in an arid environment, and to learn which variables affect their displacement within these areas. Using radiotelemetry, from 2012 to 2014 we monitored seven female mule deer. We estimated their core areas, and with this information we identified and classified the location records as inside or outside of the core area. For each record, we also recorded variables related to time (month, time of day), environmental conditions (temperature, relative humidity), and habitat characteristics (plant association, distance to the nearest water body, slope), and used them to predict the presence of the deer in the core area using Classification and Regression Trees (CART). The same variables were analyzed using a GLM to determine which factors explain variations in displacement distance. The most important variable in core area selection was the distance to the nearest body of water (<933.71 m at the population level and <1375.11 m on average for individual deer), followed by two plant associations that included the dominant species Prosopis glandulosa, Pleuraphis mutica, Larrea tridentata, Opuntia rastrera and Fouquieria splendens in areas with a gentle slope or none. Displacement distances within the core area were shorter than those outside and were affected by time of day and month. Our results are the first to elucidate the use of the core area by female mule deer in the Chihuahuan Desert, and allow us to start understanding the connection between the habitat resources essential to the survival of these deer and their movement under desert conditions.