Using neutral landscapes to identify patterns of aggregation across resource points

Many organisms are aggregated within resource patches and aggregated spatially across landscapes with multiple resources. Such patchy distributions underpin models of population regulation and species coexistence, so ecologists require methods to analyse spatially‐explicit data of resource distribution and use. I describe a method for analysing maps of resources and testing hypotheses about how resource distribution influences the distribution of organisms, where resource patches can be described as points in a landscape and the number of organisms on each resource point is known. Using a mark correlation function and the linearised form of Ripley's K‐function, this version of marked point pattern analysis can characterise and test hypotheses about the spatial distribution of organisms (marks) on resource patches (points). The method extends a version of point pattern analysis that has wide ecological applicability, it can describe patterns over a range of scales, and can detect mixed patterns. Statistically, Monte Carlo permutations are used to estimate the difference between the observed and expected values of the mark correlation function. Hypothesis testing employs a flexible neutral landscape approach in which spatial characteristics of point patterns are preserved to some extent, and marks are randomised across points. I describe the steps required to identify the appropriate neutral landscape and apply the analysis. Simulated data sets illustrate how the choice of neutral landscape can influence ecological interpretations, and how this spatially‐explicit method and traditional dispersion indices can yield different interpretations. Interpretations may be general or context‐sensitive, depending on information available about the underlying point pattern and the neutral landscape. An empirical example of caterpillars exploiting food plants illustrates how this technique might be used to test hypotheses about adult oviposition and larval dispersal. This approach can increase the value of survey data, by making it possible to quantify the distribution of resource points in the landscape and the pattern of resource use by species.     

Patterns of resource tracking by avian frugivores at multiple spatial scales: two case studies on discordance among scales

Scaling is relevant for the analysis of plant‐frugivore interaction, since the ecological and evolutionary outcomes of seed dispersal depend on the spatial and temporal scale at which frugivory patterns emerge. We analyse the relationship between fruit abundance and frugivore activity at local and landscape spatial scales in two different systems composed, respectively, by the bird‐dispersed woody plants Juniperus communis and Bursera fagaroides, and their frugivore assemblages. We use a hierarchical approach of nested patchiness of fruit‐resource, where patches are defined by individual plants within site, at the local scale, and by sites within region, at the landscape scale. The structure of patches is also described in terms of contrast (differences in fruit availability among patches) and aggregation (spatial distribution of patches). For J. communis, frugivore activity was positively related to fruit availability at the landscape scale, this pattern seldom emerging at the local scale; conversely, B. fagaroides showed a general trend of positive local pattern that disappeared at the landscape scale. These particular trends might be partially explained by differences in contrast and aggregation. The strong contrast among plants within site together with a high aggregation among sites would promote the B. fagaroides pattern to be only local, whereas in J. communis, low aggregation among sites within region would favour a sharp landscape‐scale pattern. Both systems showed discordant patterns of fruit‐resource tracking among consecutive spatial scales, but the sense of discordance differed among systems. These results, and the available multi‐scale frugivory data, suggest that discordance among successive scales allows to link directly frugivory patterns to resource‐tracking mechanisms acting at particular scales, resulting, thus, more informative than concordance observational data, in which landscape patterns might result from accumulated effect of local mechanisms. In this context, we propose new methodological approaches for a better understanding of the hierarchical behavioural mechanisms underpinning the multi‐scale resource tracking by frugivores.     

Do spatial effects play a role in the spatial distribution of desert‐dwelling Acacia raddiana?

Abstract. We investigated the spatial pattern of A. raddiana in the Negev desert of Israel in order to gain insights into the factors and processes driving the dynamics of this species. Using a scale‐dependent measure, the ring statistic, we analysed both patterns observed in the field and time series of spatial tree distributions produced by a simulation model. In the field, random spacing was the predominant pattern observed. However seedlings were clumped on small scales. We ran the model under two contrasting scenarios representing hypotheses that explain the clumping of seedlings and the random distribution of trees. One hypothesis is that there is spatial heterogeneity in seed distribution, germination and seedling mortality, but that these heterogeneities are not correlated with each other in space. The second hypothesis assumes a correlation between these heterogeneities leading to areas suitable for establishment. However, the suitability of the sites is temporally variable. Furthermore, the second hypothesis assumes density‐dependent tree mortality due to competition. Both hypotheses lead to spatial distributions that are in qualitative agreement with the patterns observed in the field. Therefore, the classical view that a clumped seedling distribution and a random pattern of older trees is due to clumped regeneration and density‐dependent mortality may not hold for Acacia trees in the Negev.     

Spatial Ecological Hierarchies: Coexistence on Heterogeneous Landscapes via Scale Niche Diversification

Spatially heterogeneous environments are generally characterized by nested landscape patterns with resource aggregations on several scales. Empirical studies indicate that such nested landscape patterns impose selection constraints on the perceptive scales of animals, but the underlying selection mechanisms are unclear. We investigated the selection dynamics of perceptive scale within a spatial resource utilization model, where the environment is characterized by its resource distribution and species differ in their perceptive scales and resource preemption capabilities. Using three model landscapes with various resource distributions, we found that the optimal perceptive scale is determined by scale-specific attributes of the landscape pattern and that the number of coexisting species increases with the number of characteristic scales. Based on the results of this model, we argue that resource aggregations on different scales act as distinct resources and that animal species of particular perceptive scales are superior in utilizing resource aggregations of comparable spatial extent. Due to the allometric relationship between body size and perceptive scale, such fitness difference might result in discontinuous body mass distributions.     

Spatial distributions of multiple plant species affect herbivore foraging selectivity

Spatial distribution of food resources is an important factor determining herbivore foraging. Previous studies have demonstrated that clumped distribution of preferred species increases its consumption by herbivores in single‐ or two‐species systems. However, the potential impact of distribution pattern of less preferred species on foraging was ignored. In natural grasslands with high species diversity and complexity, the spatial distribution of preferred species impacts on herbivore foraging may be strongly correlated with the distribution of less preferred species. Our aims were to determine the effect of distribution of both preferred and other plant species on herbivore foraging under conditions close to a native, multi‐species foraging environment, and conceptualize the relationships between spatial distribution of food resources and herbivore consumption. We hypothesized that random distribution of non‐preferred species reduces herbivore consumption of preferred species because the dispersion of less preferred species likely disturbs herbivore foraging. We conducted an experiment using three species with five combinations of clumped and random distribution patterns. Three species Lathyrus quinquenervius, Phragmites australis and Leymus chinensis, were of high, intermediate and low preferences by sheep, respectively. Results showed that distribution of low preferred species, but not that of high preferred one, affected the consumption of preferred species. Sheep obtained higher consumption of high preferred species when low preferred species followed a clumped distribution than a random distribution. Distance between aggregations of high and low preferred species did not affect sheep foraging. It was concluded that the effects of spatial distribution of preferred species on its consumption are dependent on herbivore foraging strategy, and sheep can consume more preferred species when there is a consistent spatial pattern between preferred species and the entire food resource, and that the random dispersion of low preferred species in grassland may reduce herbivore consumption of high preferred species, thus minimizing selective grazing.     

Adopting a spatially explicit perspective to study the mysterious fairy circles of Namibia

The mysterious ‘fairy circles’ are vegetation‐free discs that cover vast areas along the pro‐Namib Desert. Despite 30 yr of research their origin remains unknown. Here we adopt a novel approach that focuses on analysis of the spatial patterns of fairy circles obtained from representative 25‐ha aerial images of north‐west Namibia. We use spatial point pattern analysis to quantify different features of their spatial structures and then critically inspect existing hypotheses with respect to their ability to generate the observed circle patterns. Our working hypothesis is that fairy circles are a self‐organized vegetation pattern. Finally, we test if an existing partial‐differential‐equation model, that was designed to describe vegetation pattern formation, is able to reproduce the characteristic features of the observed fairy circle patterns. The model is based on key‐processes in arid areas such as plant competition for water and local resource‐biomass feedbacks. The fairy circles showed at all three study areas the same regular spatial distribution patterns, characterized by Voronoi cells with mostly six corners, negative correlations in their size up to a distance of 13 m, and remarkable homogeneity over large spatial scales. These results cast doubts on abiotic gas‐leakage along geological lines or social insects as causal agents of their origin. However, our mathematical model was able to generate spatial patterns that agreed quantitatively in all of these features with the observed patterns. This supports the hypothesis that fairy circles are self‐organized vegetation patterns that emerge from positive biomass‐water feedbacks involving water transport by extended root systems and soil‐water diffusion. Future research should search for mechanisms that explain how the different hypotheses can generate the patterns observed here and test the ability of self‐organization to match the birth‐ and death dynamics of fairy circles and their regional patterns in the density and size with respect to environmental gradients.     

Optimal movement between patches under incomplete information about the spatial distribution of food items

If the food distribution contains spatial pattern, the food density in a particular patch provides a forager with information about nearby patches. Foragers might use this information to exploit patchily distributed resources profitably. We model the decision on how far to move to the next patch in linear environments with different spatial patterns in the food distribution (clumped, random, and regular) for foragers that differ in their degree of information. An ignorant forager is uninformed and therefore always moves to the nearest patch (be it empty or filled). In contrast, a prescient forager is fully informed and only exploits filled patches, skipping all empty patches. A Bayesian assessor has prior knowledge about the content of patches (i.e. it knows the characteristics of the spatial pattern) and may skip neighbouring patches accordingly by moving to the patch where the highest gain rate is expected. In most clumped and regular distributions there is a benefit of assessment, i.e. Bayesian assessors achieve substantially higher long-term gain rates than ignorant foragers. However, this is not the case in distributions with less strong spatial pattern, despite the fact that there is a large potential benefit from a sophisticated movement rule (i.e. a large penalty of ignorance). Bayesian assessors do also not achieve substantially higher gain rates in environments that are relatively rich or poor in food. These results underline that an incompletely informed forager that is sensitive to spatial pattern should not always respond to existing pattern. Furthermore, we show that an assessing forager can enhance its long-term gain rate in highly clumped and some specific near-regular food distributions, by sampling the environment in slightly larger spatial units.     

基于Ripley L和O-ring函数的森林景观空间分布格局及其关联性

采用空间点格局分析中的Ripley L和O-ring函数,以大兴安岭盘古林场森林资源二类调查数据为例,对比分析2种方法在森林景观类型的空间分布格局及其关联性研究中的差异.结果表明: 2种方法获得的景观空间分布格局在整体趋势上反映一致,都是在中小尺度上呈聚集分布,之后随着尺度的增大主要表现为随机分布特征;景观类型间的关联性存在显著差异,其中,Ripley L函数结果表明,天然落叶松林、天然白桦林分别与针阔混交林在中小尺度上呈负相关,而在更大尺度上整体表现出无关联性或正关联性的趋势,其余景观类型间在所有研究尺度上呈显著负相关; O-ring函数结果则表明,4种森林景观类型两两之间均呈现出相似的关联性变化趋势,即在小尺度上均表现为负相关,在中等尺度上均呈现出不相关性,但在更大尺度上呈正相关趋势;对同一景观类型(或景观类型组),2种方法在不同尺度等级上的空间分布格局及其关联性的判别一致率存在较大差异,其一致性判别率在所有研究尺度上整体呈现出基本不变、先下降后增加和一直下降3种趋势.     

Optimal movement between patches under incomplete information about the spatial distribution of food items

If the food distribution contains spatial pattern, the food density in a particular patch provides a forager with information about nearby patches. Foragers might use this information to exploit patchily distributed resources profitably. We model the decision on how far to move to the next patch in linear environments with different spatial patterns in the food distribution (clumped, random, and regular) for foragers that differ in their degree of information. An ignorant forager is uninformed and therefore always moves to the nearest patch (be it empty or filled). In contrast, a prescient forager is fully informed and only exploits filled patches, skipping all empty patches. A Bayesian assessor has prior knowledge about the content of patches (i.e. it knows the characteristics of the spatial pattern) and may skip neighbouring patches accordingly by moving to the patch where the highest gain rate is expected. In most clumped and regular distributions there is a benefit of assessment, i.e. Bayesian assessors achieve substantially higher long-term gain rates than ignorant foragers. However, this is not the case in distributions with less strong spatial pattern, despite the fact that there is a large potential benefit from a sophisticated movement rule (i.e. a large penalty of ignorance). Bayesian assessors do also not achieve substantially higher gain rates in environments that are relatively rich or poor in food. These results underline that an incompletely informed forager that is sensitive to spatial pattern should not always respond to existing pattern. Furthermore, we show that an assessing forager can enhance its long-term gain rate in highly clumped and some specific near-regular food distributions, by sampling the environment in slightly larger spatial units.     

毛乌素沙地油蒿种群点格局分析

油蒿(Artemisia ordosica)是我国北方农牧交错带的重要固沙植物,研究其种群格局对理解种群生态过程和改善流沙治理技术具有重要意义。点格局分析法是20世纪末发展起来的多尺度空间格局分析方法。通过研究油蒿种群的点格局,发现油蒿种群的空间分布格局和空间关联性同空间尺度、植株形体大小以及生境3种因素有密切联系。在较小的空间尺度上,油蒿种群倾向于非随机分布(集群分布比均匀分布常见),个体间有较强的空间关联(正关联比负关联常见);当空间尺度大于临界值后,油蒿种群倾向于服从随机分布,同时种群的空间关联性减弱。幼小油蒿植株具有明显的集群分布趋势,高大植株则表现出聚集强度的降低趋势;形体大小的差异越大,植株间的正关联关系越弱,或者负关联关系越强。与固定沙地相比,半固定沙地油蒿种群的集群分布现象更加明显,同时种群的空间正关联关系更强。研究结果表明,当通过移栽油蒿成体治理流动沙地时,应尽量将其栽种成集群分布而非均匀分布的形式,以提高植株成活率。     

A test for spatial relationships between neighbouring plants in plots of heterogeneous plant density

Abstract. Maps of plant individuals in (x, y) coordinates (i.e. point patterns) are currently analysed through statistical methods assuming a homogeneous distribution of points, and thus a constant density within the study area. Such an assumption is seldom met at the scale of a field plot whilst delineating less heterogeneous subplots is not always easy or pertinent. In this paper we advocate local tests carried out in quadrats partitioning the plot and having a size objectively determined via a trade‐off between squared bias and variance. In each quadrat, the observed pattern of points is tested against complete spatial randomness (CSR) through a classical Monte‐Carlo approach and one of the usual statistics. Local tests yield maps of p‐values that are amenable to diversified subsequent analyses, such as computation of a variogram or comparison with co‐variates. Another possibility uses the frequency distribution of p‐values to test the whole point pattern against the null hypothesis of an inhomogeneous Poisson process. The method was demonstrated by considering computer‐generated inhomoge‐neous point patterns as well as maps of woody individuals in banded vegetation (tiger bush) in semi‐arid West Africa. Local tests proved able to properly depict spatial relationships between neighbours in spite of heterogeneity/clustering at larger scales. The method is also relevant to investigate interaction between density and spatial pattern in the presence of resource gradients.     

Determinants of Spatial Distribution in a Bee Community: Nesting Resources,Flower Resources,and Body Size

Understanding biodiversity distribution is a primary goal of community ecology. At a landscape scale, bee communities are affected by habitat composition, anthropogenic land use, and fragmentation. However, little information is available on local-scale spatial distribution of bee communities within habitats that are uniform at the landscape scale. We studied a bee community along with floral and nesting resources over a 32 km² area of uninterrupted Mediterranean scrubland. Our objectives were (i) to analyze floral and nesting resource composition at the habitat scale. We ask whether these resources follow a geographical pattern across the scrubland at bee-foraging relevant distances; (ii) to analyze the distribution of bee composition across the scrubland. Bees being highly mobile organisms, we ask whether bee composition shows a homogeneous distribution or else varies spatially. If so, we ask whether this variation is irregular or follows a geographical pattern and whether bees respond primarily to flower or to nesting resources; and (iii) to establish whether body size influences the response to local resource availability and ultimately spatial distribution. We obtained 6580 specimens belonging to 98 species. Despite bee mobility and the absence of environmental barriers, our bee community shows a clear geographical pattern. This pattern is mostly attributable to heterogeneous distribution of small (<55 mg) species (with presumed smaller foraging ranges), and is mostly explained by flower resources rather than nesting substrates. Even then, a large proportion (54.8%) of spatial variability remains unexplained by flower or nesting resources. We conclude that bee communities are strongly conditioned by local effects and may exhibit spatial heterogeneity patterns at a scale as low as 500–1000 m in patches of homogeneous habitat. These results have important implications for local pollination dynamics and spatial variation of plant-pollinator networks.     

Effects of human‐induced prey depletion on large carnivores in protected areas: Lessons from modeling tiger populations in stylized spatial scenarios

Prey depletion is a major threat to the conservation of large carnivore species globally. However, at the policy‐relevant scale of protected areas, we know little about how the spatial distribution of prey depletion affects carnivore space use and population persistence. We developed a spatially explicit, agent‐based model to investigate the effects of different human‐induced prey depletion experiments on the globally endangered tiger (Panthera tigris) in isolated protected areas—a situation that prevails throughout the tiger's range. Specifically, we generated 120 experiments that varied the spatial extent and intensity of prey depletion across a stylized (circle) landscape (1,000 km²) and Nepal's Chitwan National Park (~1,239 km²). Experiments that created more spatially homogenous prey distributions (i.e., less prey removed per cell but over larger areas) resulted in larger tiger territories and smaller population sizes over time. Counterintuitively, we found that depleting prey along the edge of Chitwan National Park, while decreasing tiger numbers overall, also decreased female competition for those areas, leading to lower rates of female starvation. Overall our results suggest that subtle differences in the spatial distributions of prey densities created by various human activities, such as natural resource‐use patterns, urban growth and infrastructure development, or conservation spatial zoning might have unintended, detrimental effects on carnivore populations. Our model is a useful planning tool as it incorporates information on animal behavioral ecology, resource spatial distribution, and the drivers of change to those resources, such as human activities.     

Consequences of multiple simultaneous opportunities to exploit others’ efforts on free riding

Individuals within a group do not all act in the same way: Typically, the investors (or producers) put efforts into producing resources while the free riders (or scroungers) benefit from these resources without contributing. In behavioral ecology, the prevalence of free riders can be predicted by a well‐known game‐theoretical model—the producer–scrounger (PS) model—where group members have the options to either search for resources (producers) or exploit the efforts of others (scroungers). The PS model has received some empirical support, but its predictions, surprisingly, are based on the strict assumption that only one resource can be exploited at a time. Yet, multiple simultaneous opportunities to exploit others’ efforts should frequently occur in nature. Here, we combine analytic and simulation approaches to explore the effect of multiple simultaneous scrounging opportunities on tactic use. Our analyses demonstrate that scrounging rates should increase with the number of simultaneous opportunities. As such, the amount and spatial distribution (i.e., clumped vs. dispersed) of resources as well as the risk of predation are key predictors of scrounging behavior. Because scroungers contribute to reducing the speed of resource exploitation, the model proposed here has direct relevance to the exploitation and sustainability of renewable resources.     

Resources driving landscape‐scale distribution patterns of grazers in an African savanna

In order to effectively manage and conserve indigenous herbivores, a good understanding is needed of how resources drive their distribution patterns. This study employed a unique dataset to test a range of ecological theories and hypotheses on free‐ranging grazers. Using aerial census data collected over 14 yr across the 2 million ha Kruger National Park (South Africa), this study employs spatial autologistic regression models to explore the spatial relationships that exist between the distribution of eight indigenous grazer species and a set of resource variables. It was found that ecological theories relating to feeding guild, water‐dependence, allometric scaling, gut‐morphology and vulnerability to predation could explain most of the grazer distribution patterns observed in relation to surface‐water, forage quality, forage quantity and habitat openness. All the grazers studied were water‐dependent and occurred close to a permanent source of water in the dry season. This was ascribed to the lack of moisture in the diet of grazers during the dry season. Most ruminants’ distribution patterns were significantly associated with areas of high forage quality whereas hind‐gut fermentors were neutral towards forage quality. Average forage quantity was not a significant predictor of long‐term, landscape‐scale distribution patterns for any of the grazer species studied. Most small‐ and medium‐bodied grazers preferred open habitats above closed habitats, probably due to higher visibility and a lower predation risk. Large‐bodied grazers did not bias their distribution patterns towards open habitats. The way in which grazers distribute themselves with respect to different resources can potentially inform management actions on appropriate scales.     

Energy benefits and emergent space use patterns of an empirically parameterized model of memory‐based patch selection

Many species frequently return to previously visited foraging sites. This bias towards familiar areas suggests that remembering information from past experience is beneficial. Such a memory‐based foraging strategy has also been hypothesized to give rise to restricted space use (i.e. a home range). Nonetheless, the benefits of empirically derived memory‐based foraging tactics and the extent to which they give rise to restricted space use patterns are still relatively unknown. Using a combination of stochastic agent‐based simulations and deterministic integro‐difference equations, we developed an adaptive link (based on energy gains as a foraging currency) between memory‐based patch selection and its resulting spatial distribution. We used a memory‐based foraging model developed and parameterized with patch selection data of free‐ranging bison Bison bison in Prince Albert National Park, Canada. Relative to random use of food patches, simulated foragers using both spatial and attribute memory are more efficient, particularly in landscapes with clumped resources. However, a certain amount of random patch use is necessary to avoid frequent returns to relatively poor‐quality patches, or avoid being caught in a relatively poor quality area of the landscape. Notably, in landscapes with clumped resources, simulated foragers that kept a reference point of the quality of recently visited patches, and returned to previously visited patches when local patch quality was poorer than the reference point, experienced higher energy gains compared to random patch use. Furthermore, the model of memory‐based foraging resulted in restricted space use in simulated landscapes and replicated the restricted space use observed in free‐ranging bison reasonably well. Our work demonstrates the adaptive value of spatial and attribute memory in heterogeneous landscapes, and how home ranges can be a byproduct of non‐omniscient foragers using past experience to minimize temporal variation in energy gains.     

帽儿山地区1983—2016年森林景观空间点格局及其关联动态性

研究森林景观空间格局及其动态,对于维持森林生态系统的稳定性以及森林经营方案的制定具有重要意义.基于帽儿山实验林场1983、1993、2004和2016年共4期二类调查数据,利用Programita软件(2010版)中的O-ring统计方法对该林场的森林景观空间格局及其关联性动态进行定量分析.结果表明: 1983—2016年间,区域内软阔混交林所占面积比例均呈下降趋势,天然蒙古栎林景观所占面积比例先增加后降低,而硬阔混交林、落叶松人工林、樟子松人工林所占面积比例呈增加趋势.研究期间,软阔混交林、硬阔混交林、天然蒙古栎林的空间聚集尺度均呈减小趋势,聚集分布的范围由0～7 km缩小到0～3 km,之后随着尺度范围的增大逐渐转变为随机分布或均匀分布;落叶松人工林在较小尺度范围均表现为聚集分布,而随着尺度范围增大表现为随机分布或均匀分布,且均匀分布的尺度范围逐渐增大、随机分布的尺度范围逐渐变小;樟子松人工林聚集分布的尺度虽然也集中在小尺度范围内(0～4 km),但分布范围呈显著增大趋势.总体上,大部分景观类型之间的空间关联性在小尺度范围内呈现负关联性,随着尺度增大逐渐变为无关联性或正关联性;仅少数景观类型间(如1983年落叶松人工林与樟子松人工林)的关联性在小尺度范围内为正关联性,随着尺度范围增大逐渐变为无关联性.1983—2016年间,各景观类型之间的空间关联性动态呈现不同的变化规律,是帽儿山地区各项经营管理措施的综合体现.     

红松存在/不存在数据的多尺度空间分布格局

红松属小兴安岭地区地带性植被优势种,该地区也是其分布的北缘。在景观尺度上开展红松的分布格局研究有利于进一步了解红松分布机理、未来迁移过程等问题,对其经营和保护有重要意义。将景观指数法与点格局分析法结合,设定8个空间尺度,利用红松存在/不存在数据,通过计算各空间尺度上红松聚集程度和景观指数,分析小兴安岭地区红松种群在多尺度上的分布格局。研究结果表明,小尺度上红松聚集分布明显,随机分布区多处于其聚集分布区的边缘,均匀分布区则散布在其聚集分布区内。景观指数研究表明,通过景观指数可判断红松聚集分布格局趋势,而不能判断均匀分布、随机分布格局趋势,因为它们在多尺度下景观指数波动大,不能用景观指数来描述分布格局。研究得出如下结论:1)红松主要分布在其分布区的核心区域内,在分布区边缘和过渡带上呈随机分布,2)存在/不存在数据能够用来分析种群的多尺度空间分布格局,3)空间尺度的变化会引起树种分布格局的变化,随机分布随尺度增加,边缘化程度加强,4)单一尺度上,景观格局指数不能完全描述种群分布格局;而在多尺度上,变化趋势稳定的景观指数表明聚集分布存在,而波动剧烈的景观指数常与随机分布和均匀分布联系在一起,5)地形因子中,红松对坡度和海拔两个因子变化敏感。     

Spatial Distribution and Exploitation of Trees Gouged by Common Marmosets (Callithrix jacchus)

Resource distribution shapes many aspects of primate behavioral ecology. Though the spatial patterning of fruits, leaves, and insects has been explored among primate foods, comparatively less is known about exudate distributions. Tree exudates are a renewable resource, provide long-term evidence of exploitation, and may be selectively exploited to manipulate spatial distribution. We assessed the spatial patterning of trees gouged by common marmosets (Callithrix jacchus) to determine if they exhibit a uniform, random, or clumped distribution. We also asked whether marmosets selectively gouge trees in home range centers, which may afford them exclusive access to exudates. We explored whether spatial or physical characteristics of trees predict how intensely gouged trees were exploited. The mean nearest neighbor distance of gouged trees was significantly closer than expected for a random distribution and Ripley’s K-function showed that gouged trees were clumped across all spatial scales in our study area. Clumping may enable marmosets to reduce day and home ranges and facilitate repeated gouging of trees. Gouged trees were not closer to marmosets’ home range centers than peripheries, nor were centrally located trees more intensely gouged. Increased gouging intensity was associated with larger tree circumferences, although this effect was primarily driven by interspecific differences in circumference. Although marmosets may benefit from exploiting clumped exudates, they do not concentrate gouging in areas where they are more likely to gain exclusive access. Species-specific tree characteristics such as exudate quality and/or bark properties may play a larger role in determining gouging patterns than intergroup feeding competition.