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1.
Rivers represent natural edges in forests, serving as transition zones between landscapes. Natural edge effects are important to study to understand how intrinsic habitat variations affect wildlife as well as the impact of human-induced forest fragmentation. We examined the influence of riparian and anthropogenic edge on mantled howler, white-faced capuchin, Central American spider monkeys, and vegetation structure at La Suerte Biological Research Station (abbreviated as LSBRS), Costa Rica. We predicted lower monkey encounter rate, tree species richness, and median dbh at both edge types compared to interior and that monkeys would show species-specific responses to edge based on size and diet. We expected large, folivorous–frugivorous howler monkeys and small, generalist capuchins would be found at increased density in forest edge, while large, frugivorous spider monkeys would be found at decreased density in forest edge. We conducted population and vegetation surveys along interior, riparian, and anthropogenic edge transects at LSBRS and used GLMM to compare vegetation and monkey encounter rate. Tree species richness and median dbh were higher in forest interior than anthropogenic edge zones. Although spider monkey encounter rate did not vary between forest edges and interior, howler monkeys were encountered at highest density in riparian edge, while capuchins were encountered at highest density in anthropogenic edge. Our results indicate that diverse forest edges have varying effects on biota. Vegetation was negatively affected by forest edges, while monkey species showed species-specific edge preferences. Our findings suggest that riparian zones should be prioritized for conservation in Neotropical forests.  相似文献   

2.
The function of long calling is a subject of interest across animal behaviour study, particularly within primatology. Many primate species have male‐specific long‐distance calls, including platyrrhines like the folivorous howler monkey (Alouatta spp.). Howler monkeys may howl to defend resources such as feeding trees or areas of rich vegetation from other monkey groups. This study tests the ecological resource defence hypothesis for howling behaviour in the mantled howler monkey (Alouatta palliata) and investigates how anthropogenic forest fragmentation may influence howling behaviour. More specifically, this study examines how howling bout rate, duration, precursors and tree species richness, DBH, and canopy cover vary in 100 m anthropogenic edge and interior forest zones at La Suerte Biological Research Station (LSBRS), a fragmented tropical rainforest in Costa Rica. Results show that tree species richness and canopy cover are higher in forest interior at this site, suggesting that monkeys should howl at greater rates in the interior to defend access to these higher‐quality vegetation resources. Overall, our results supported the ecological resource defence hypothesis. The main howl precursor was howling from neighbouring groups. Although howling rate did not differ between forest zones, howling bouts from forest interior were longer, had a greater number of howls per bout and were preceded by different precursors than howls from anthropogenic edge zones, including more howls from neighbouring groups. Our findings provide some of the first evidence for behavioural edge effects in primate vocal communication behaviour.  相似文献   

3.

Anthropogenic forest fragmentation impacts many aspects of animal behaviour, including feeding ecology. With forests increasingly fragmented in tropical regions due to human development, the proportion of forest edge (≤?100 m from clear-cut regions) is higher relative to forest interior. Forest edges differ in vegetation from interior, making it important to better understand how anthropogenic edges impact the feeding behaviour of primates such as mantled howler monkeys (Alouatta palliata). We predicted that howler monkeys would feed on higher-quality plant resources, from a larger number of tree families, and from larger trees in forest interior compared to anthropogenic forest edge. We surveyed howler monkey feeding behaviour across forest zones in a fragmented rainforest in Costa Rica, La Suerte Biological Research Station. We observed individual monkeys for 30-minute periods, collecting data on their feeding behaviour and tree use at 2-minute intervals. We measured feeding trees and recorded the plant parts and taxonomy of resources consumed. Monkeys consumed more leaves and fewer stems and fed from a smaller number of tree families in the forest interior, while they consumed fewer leaves and more stems and fed from a larger number of tree families in the forest edge. Monkeys also fed from larger, taller trees in the forest interior than the edge. The differences in howler monkey feeding behaviour between forest zones attest to the impact of human disturbance on howler monkey feeding ecology.

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4.
Across the globe, primates are threatened by human activities. This is especially true for species found in tropical dry forests, which remain largely unprotected. Our ability to predict primate abundance in the face of human activity depends on different species' sensitivities as well as on the characteristics of the forest itself. We studied plant and primate distribution and abundance in the Taboga Forest, a 516-ha tropical dry forest surrounded by agricultural fields in northwestern Costa Rica. We found that the density of white-faced capuchins (Cebus capucinus) at Taboga is 2–6 times higher than reported for other long-term white-faced capuchin sites. Using plant transects, we also found relatively high species richness, diversity, and equitability compared with other tropical dry forests. Edge transects (i.e., within 100 m from the forest boundary) differed from interior transects in two ways: (a) tree species associated with dry forest succession were well-established in the edge and (b) canopy cover in the edge was maintained year-round, while the interior forest was deciduous. Sighting rates for capuchins were higher near water sources but did not vary between the edge and interior forest. For comparison, we also found the same to be true for the only other primate in the Taboga Forest, mantled howler monkeys (Alouatta palliata). Year-round access to water might explain why some primate species can flourish even alongside anthropogenic disturbance. Forest fragments like Taboga may support high densities of some species because they provide a mosaic of habitats and key resources that buffer adverse ecological conditions.  相似文献   

5.
We linked primary dispersal by spider monkeys (Ateles geoffroyi) and howler monkeys (Alouatta pigra) to post‐dispersal seed fate by studying the effects of dung type and defecation pattern on secondary seed dispersal by dung beetles. First, we described the defecation patterns for both primate species. Howler monkeys generally defecated in groups (88% of observed defecations), with each individual producing on average 31 g of dung, resulting in a large area of the forest floor (31 m2) covered by large amounts of dung (clumped spatial pattern). Spider monkeys generally (96% of observed defecations) defecated individually, each individual producing an average of 11 g of dung, resulting in a small area of the forest floor (2 m2) covered by small amounts of dung (scattered spatial pattern). Secondly, we captured dung beetles using as bait the dung of both primate species, to detect differences in the assemblages of these secondary seed dispersers attracted to the dung of both primates. More individual dung beetles, but not more species, were attracted to howler monkey dung than to spider monkey dung. Finally, we assessed experimentally (using plastic beads as seed mimics) how dung type (Ateles vs. Alouatta) and defecation pattern (scattered vs. clumped) affect secondary seed dispersal by dung beetles. We found that post‐dispersal seed fate was affected by dung type, with more seeds being buried when present in howler monkey dung, than in spider monkey dung, but was not affected by defecation pattern. It is important to consider post‐dispersal processes, such as secondary seed dispersal by dung beetles, when comparing species of primary dispersers.  相似文献   

6.
Understanding how threatened species adapt their behavior to landscapes shaped by humans is increasingly important to ensuring they persist in a changing world. Matrix habitats can be shared spaces where human and nonhuman primates coexist. We set out to determine how an endemic, threatened forest specialist, the frugivorous, arboreal samango monkey (Cercopithecus albogularis labiatus), has responded to a matrix habitat made up of residential gardens and commercial plantations in Eastern Cape province, South Africa. We followed two groups from dawn to dusk for a mean of 3 days/mo for 12 mo (February 1, 2011 to January 31, 2012) using scan sampling to collect data on their diet, activity, and ranging patterns. We used resource abundance transects to describe the groups’ home ranges and monitored tree phenology to calculate fruit and seed availability indices. Monkeys from both groups consumed large quantities of exotic plant species, accounting for >50% of their overall annual diet, with seeds of the invasive black wattle (Acacia mearnsii) the most commonly consumed exotic species followed by acorns of two oak species (Quercus robur and Q. palustris.). However, monkeys responded to the availability of indigenous rather than exotic fruits and seeds and increased their consumption of exotics when indigenous fruits were less available. Although monkeys spent less time moving when feeding on exotic species compared to indigenous species, eating exotics did not free up monkeys’ time to rest or socialize, as additional time was required to process exotic foods. To offset the possible negative consequences of the monkeys’ reliance on exotic seeds, including escalating conflict between monkeys and people in gardens, we suggest gradual removal of exotic plant species in the habitat and replacement with indigenous species as one mitigation strategy.  相似文献   

7.
Variation at both the patch and landscape scale is known to influence the distribution and abundance of arboreal monkeys in rainforest fragments. However, few studies have examined the factors associated with these different scales of focus simultaneously. Using stepwise logistic-regression and generalized linear models (GLMs), howler monkey Alouatta palliata distribution and abundance were examined as a function of patch quality (fragment area, shape, tree DBH and canopy height) and landscape connectivity (isolation, total forest area, fragment and road abundance, corridor abundance and length) in 119 rainforest fragments in northern Chiapas, Mexico. The positive correlation observed between monkey distribution (presence/absence) and both fragment area and abundance may be explained by increased resources within larger fragments and those whose proximity allows greater exploration by monkeys. In contrast, GLM analysis indicated that monkey abundance in inhabited fragments was positively correlated with corridor abundance, canopy height and fragment area. These relationships could be explained by greater reproductive investments by monkeys in forest fragments whose size, degree of perturbation and degree of connectivity with other fragments suggest greater overall resource availability. Future studies should explicitly include a multi-scale approach to understanding the factors affecting patterns of monkey distribution and abundance, particularly as this relates to measures of and interactions between patch connectivity and resource availability.  相似文献   

8.
Historical records of Ateles chamek (black-faced black spider monkey) suggest that the species range extends further south of the known species distribution, within an ecotonal region between the Amazonia, Cerrado and Pantanal biomes in Brazil. Ecotones are zones of habitat transition with high species richness that remain undersampled as conservationists often prioritize biodiversity hotspots. Thus, distribution ranges may be inaccurately measured when species occur in ecotonal zones. We report the first precise records of A. chamek in 24 new localities surveyed in the ecotonal zone of the Upper Paraguay River Basin, and we present subgroup encounter rates in the 11 largest patches (>70 ha) along 207 km of the line transects surveyed. The new records represent an expansion of the distribution of A. chamek approximately 200 km to the south, increasing the known extent of its occurrence by 10.8%. Local tributaries may not be barriers for spider monkeys, which are able to swim and cross slow-moving rivers. However, the dry forests of the Cerrado and the flooded areas of the Pantanal, formed by grassland and scarce trees, may be habitat barriers for A. chamek. The populations living in this ecotonal zone are relatively abundant (1.1–6.67 subgroup sightings/10 km) compared to the heavily hunted continuous forests of northern Amazonia. Furthermore, these values are similar to those for other Ateles spp. inhabiting forests with low or no hunting pressure. We highlight the need for specific conservation action to protect the spider monkeys living in these landscapes, which are threatened by agriculture expansion.  相似文献   

9.
Mineral licks--also known as "salados," "saladeros," or "collpas"--are specific sites in tropical and temperate ecosystems where a large diversity of mammals and birds come regularly to feed on soil. Although the reasons for vertebrate geophagy are not completely understood, animals are argued to obtain a variety of nutritional and health benefits from the ingestion of soil at mineral licks. We studied the temporal patterns of mineral lick use by white-bellied spider monkey (Ateles belzebuth) and red howler monkey (Alouatta seniculus) in a lowland rain forest in Amazonian Ecuador. Using camera and video traps at four different mineral licks, combined with behavioral follows of one group of spider monkeys, we documented rates of mineral lick visitation by both primate species and the relative frequency and intensity of mineral lick use by spider monkeys. On the basis of 1,612 days and 888 nights of mineral lick monitoring, we found that A. belzebuth and A. seniculus both visit mineral licks frequently throughout the year (on average ~14% of days for both species), and mineral lick visitation was influenced by short-term environmental conditions (e.g. sunny and dry weather). For spider monkeys, the area surrounding the lick was also the most frequently and most intensively used region within the group's home range. The fact that spider monkeys spent long periods at the lick area before coming to the ground to obtain soil, and the fact that both species visited the lick preferentially during dry sunny conditions (when predator detectability is presumed to be relatively high) and visited simultaneously more often than expected by chance, together suggest that licks are indeed perceived as risky areas by these primates. We suggest that howler and spider monkeys employ behavioral strategies aimed at minimizing the probability of predation while visiting the forest floor at risky mineral lick sites.  相似文献   

10.
A survey of the population of the black howler monkey (Alouatta pigra) present at the Mayan site of Palenque was conducted during 2000. A total of 911 man/hours, spread over 112 days were spent surveying the 600 ha area of pristine forest at the site for howler troops. We detected the presence of 136 individuals of which 131 were members of 20 troops, the rest were 3 solitary adult males and 2 adult males travelling as a pair. Ecological density was estimated at 23 individuals/km2. Mean troop size was 7.0 individuals and it ranged from 2–12 individuals; 60% of the troops were multimale. All sighting of howler monkeys were in evergreen rain forest and 75% were in trees ≥20 m in height. The reported densities and mean troop size are higher than those reported for the species in Guatemala and in central Quintana Roo, Mexico. The vegetation of the forest contains tree species reported to be used by species ofAlouatta in the Moraceae, Sapotaceae, Leguminosae, and Lauraceae plant families. Protection of a large perimeter area (ca 1700 ha) around the archeological site by the Mexican government ensures the conservation of the forest and of the black howler monkey population present at the site.  相似文献   

11.
Composition of the landscape matrix of surrounding forest fragments is thought to be critically important to the survival of arboreal primates because it offers structures that help the animals move between fragments and other foraging sites. However, little is known about the composition of the matrix used by these animals. The aim of this study was to quantitatively assess the importance of the landscape matrix and its effects on primate abundance, using black howler monkeys (Alouatta pigra) living in a landscape fragmented by the expansion of agriculture and pastures for livestock in southeastern Mexico. In 2008, a complete census of the monkeys was carried out across the 2000-ha landscape matrix, and for every site where we observed monkeys, we recorded canopy height, tree basal area, food-source abundance, and distance to the nearest fragment. A total of 244 howler monkeys, distributed among 48 groups (including six solitary males) were counted in the matrix. Mean troop size was 5.6 ± 2.8 individuals, and the mode was three individuals. The highest number of troops and greatest howler monkey abundance were recorded in the isolated trees, the eucalyptus plantation, and orchards. A generalized linear model revealed that monkey abundance tended to be higher in matrix elements with higher canopy height, greater food availability, and closest to rainforest fragments. These results suggest that it is necessary to take into account the many elements of the landscape when drawing up conservation and habitat management plans, particularly in order to establish connectivity among the fragments and elements of the matrix with native trees.  相似文献   

12.
伊朗稀疏橡木林片段对草本植物物种多样性和土壤特性的边缘影响 温带和热带森林中的森林边缘现象已经得到了很好的研究,但在稀疏的橡木林片段中的相关研究却较为缺乏。本文研究了稀疏橡木林片段对植物物种多样性和土壤特性的边缘影响。本研究沿着伊朗克尔曼沙赫省3个小型(<10 ha)和3个大型(>10 ha)橡木林片段的3个横断面收集了从边缘到内部的相 关数据,测量了0(森林边缘)、25、50、100和150 m处的草本植物(高度<0.5 m)和土壤特性。使用香农指数量化了物种多样性,使用稀疏标准化方法比较了两个大小不同片段中的物种丰富度,并应用了非度量多维测度排序研究了物种组成的变化。通过随机化测试估算了边缘影响的距离,并利用Tukey HSD事后检验法的广义线性混合模型评估了距边缘距离和片段大小对多样性和土壤特性的影响。研究结果表明,大小片段边缘具有较高的物种丰富度、多样性和均匀度,而大片段边缘的土壤氮和有机碳含量则较内部更低(边缘50 m范围内的变化最大)。大小片段的物种组成、土壤有机碳和氮总量都存在显 著差异。本研究关于这些稀疏森林对草本植物和土壤特性产生显著边缘影响的发现,对于边缘研究,尤其是边缘和草本植物的相关研究具有重大贡献。  相似文献   

13.
Howler monkey capture is an arduous and expensive task requiring trained and specialized professionals. We compared strategies and methods to most efficiently capture Alouatta guariba clamitans in remnants of the Atlantic Forest in Rio de Janeiro and its bordering states of Minas Gerais and São Paulo. We tested whether or not the success of expeditions in the forest with anesthetic darts, nets, and baited traps differed with and without the support of an information network, a contact chain built with key institutions and inhabitants to continuously monitor howler monkey presence. The influence of forest conditions (vegetation type and fragment size) upon darting success was also evaluated. We captured 24 free‐living A. guariba clamitans. No howler monkey was caught with traps, probably due to the predominantly folivore feeding to high local plant diversity providing a great variety of food options. Captures based on an information network were significantly more efficient in terms of numbers of caught monkeys than without it. Captures with darts were considerably more efficient when performed in semideciduous forests and small forest fragments as opposed to ombrophilous forests or large woods. Although we walked great distances within the forest searching for howler monkeys, all but one animal were captured at the forest fringes. Hindrances to search and the darting method in the Atlantic Forest, for example, the steep terrain, high tree canopies, hunt pressure, and low A. guariba clamitans population density, were mitigated with the use of the information network in this monkey capture. Moreover, the information network enhanced the surveillance of zoonotic diseases, which howler monkeys and other nonhuman primates are reservoirs in Brazil, such as malaria and yellow fever.  相似文献   

14.
We report the feeding behavior and food preferences of a troop of red howler monkeys (Alouatta seniculus) over two annual cycles in primary tropical rain forest in French Guiana. The monkeys used 195 plant species from 47 families as food. Major food categories were young leaves (54%), mature fruits (21.5%), and flowers (12.6%). Other food categories included old leaves, immature fruits, termitarium soil, bark, and moss. The monkeys were less selective than other howler groups, since 19 plant species contributed 1% to their diet and accounted for only 35.7% of their total diet. The Sapotaceae was the most frequently eaten plant family and represented >10% of the total diet.  相似文献   

15.
Aims Forest fragmentation and the associated augmentation of forest edge zones are increasing worldwide. Forest edges are characterized by altered plant species richness and community composition. As the tree layer and its species composition has been shown to influence herb layer composition, changes in tree species composition or richness may weaken or strengthen edge effects in forest ecosystems. We studied effects of the edge–center transition, tree species composition and their potential interaction on the understory vegetation in the Hainich National Park, Germany's largest connected deciduous forest, allowing to cover large edge–center transects.Methods We established 12 transects in an area of 75 km 2 of continuous forest, 6 beech-dominated and 6 in multispecies forest stands. Each transect reached from the forest edge up to 500 m into the forest interior. Vegetation relevés were conducted in regular, logarithmic distances along each transect.Important findings Herb species richness was influenced by an interaction of edge effects and tree diversity level. With increasing distance from the forest edge, herb species richness remained constant in multispecies forest stands but rapidly decreased in beech-dominated forest stands. Further, herb richness was higher in the interior of multispecies forest stands. Percent forest specialists increased and percent generalists decreased with distance from the edge and this contrasting pattern was much more pronounced in beech-dominated transects. By using structural equation modeling, we identified litter depth mediated by tree species composition as the most important driver of herb layer plant species richness.  相似文献   

16.
为研究人工林群落的边缘效应特征, 本文以川西周公山森林公园的柳杉(Cryptomeria fortunei)人工林破碎化大斑块为对象, 以植株平均胸径、平均高度、平均密度、丰富度指数(D)和Shannon-Wiener指数(H)来综合衡量边缘效应深度。在实地踏查的基础上, 从林缘向林内(梯度1至梯度5)设置5条样带(宽度为10 m), 在每条样带中设置4个10 m × 10 m的小样方进行调查。结果表明: (1)从物种组成上看, 在总面积为2,000 m2的20个小样方中共记录到111个物种, 隶属于54科96属, 物种数从林缘至林内递减。(2)从群落结构上看, 乔木层的平均胸径从林缘至林内呈减小趋势, 平均密度则相反, 平均高度无显著变化; 灌木层的平均密度从林缘向林内减小, 平均高度无显著变化; 草本层的平均密度和平均高度均呈减小趋势。(3)从物种多样性上看, 总体上各层次的丰富度指数和Shannon-Wiener指数均从林缘向林内呈减小趋势, 其中灌木层和草本层的变化趋势最明显; 同时, 林内各梯度与梯度1 (林缘)的共有种和相似性系数从林缘向林内递减。(4)分析各项群落特征发现, 平均高度、平均密度和相似性系数的数值在梯度2向梯度3过渡时的起伏变化最明显, 推断本研究中柳杉人工林斑块的边缘深度可达林内20 m。  相似文献   

17.
Arboreal fauna living in tropical ecosystems may be particularly affected by roads given their dependency on forest cover and the high vulnerability of such ecosystems to changes. Over a period of 4 yr, we followed subgroups of spider monkeys living in a regenerating dry tropical forest with 8.2 km of roads within their home range. We aimed to understand whether roads shaped the home range of spider monkeys and which road features affected their movement. Only 18 percent (3 km) of the spider monkeys’ home range perimeter bordered with roads; these roads had greater habitat disparity between road sides than roads inside the home range. Although monkeys were reluctant to be close to roads, and roadside habitat contained low proportions of mature forest, spider monkeys crossed roads at 69 locations (7.5 crossings per kilometer). The main road characteristic affecting crossings was canopy opening size, with greater probability of crossing where canopy openings were smaller. Our findings support the importance of canopy opening size for road crossing of arboreal taxa, but they also indicate the relevant role roadside forest structure may have. Minimizing canopy opening size and forest disturbance along roads can facilitate the movement of arboreal fauna and preserve the important role of spider monkeys and other arboreal taxa in seed dispersal and thus the maintenance and regeneration of forest diversity.  相似文献   

18.
A geographical survey of two Mnais damselfly species in the Kinki area of Japan showed evidence for character displacements when the two species were found in sympatry. Mnais costalis, a species that has polymorphic male mating types of orange-winged territorial and clear-winged non-territorial morphs (hereafter abbreviated to orange and clear morphs respectively) in allopatry often shifted to having monomorphic orange morphs in sympatry. The mean body size of orange morphs was consistently larger than that of clear morph in allopatry. The mean body size of the sympatric orange morphs was even larger than that of allopatric orange morphs. By contrast, Mnais pruinosa, a species that also has two morphs of large orange and small clear morphs in allopatry, shifted to having monomorphic clear morphs in sympatry. The mean body size of the sympatric clear morphs was smaller than that of allopatric clear morphs. Divergence was also detected in the preference for habitat insolation conditions between sympatric Mnais damselflies. Both species in allopatric regions prefer half-light forest habitats, while in sympatric regions they showed diversified habitat preference: M. costalis preferred sunny habitats while M. pruinosa preferred shady habitats. Multiple character displacements in signal traits and habitat preference emerged in heterogeneous forest light environments are likely to have synergistic effects on the reproductive isolation of the two species.  相似文献   

19.
Forest edges are known to consist of microenvironments that may provide habitat for a different suite of species than forest interiors. Several abiotic attributes of the microenvironment may contribute to this change across the edge to center gradient (e.g., light, air temperature, soil moisture, humidity). Biotic components, such as seed dispersal, may also give rise to changes in species composition from forest edge to interior. We predicted that abiotic and biotic measures would correlate with distance from forest edge and would differ among aspects. To test these predictions, we measured abiotic and biotic variables on twelve 175 m transects in each of two 24 ha forest fragments in east-central Illinois that have remained in continuous isolation for upwards of 100 years. Both univariate and multivariate techniques were used to best describe the complex relationships among abiotic factors and between abiotic and biotic factors. Results indicate that microclimatic variables differ in the degree to and distance over which they show an edge effect. Relative humidity shows the widest edge, while light and soil moisture have the steepest gradients. Aspect influences are evidenced by the existence of more pronounced edge effects on south and west edges, except when these edges are protected by adjacent habitat. Edges bordered by agricultural fields have more extreme changes in microclimate than those bordered by trees. According to PCA results, species richness correlates well with microclimatic variation, especially light and soil moisture; however, in many cases species richness had a different depth of edge influence than either of these variables. The herbaceous plant community is heavily dominated by three species. Distributions of individual species as well as changes in plant community composition, estimated with a similarity index, indicate that competition may be influencing the response of the vegetation to the edge to interior gradient. This study indicates that edge effects must be considered when the size and potential buffering habitat of forest preserves are planned.  相似文献   

20.
Range defensibility is defined as the ability of animals to efficiently move over an area to monitor and defend it. Therefore, range defensibility can help us understand the spatial structure of animal territoriality. We used howler monkeys (Alouatta spp.), a genus for which no agreement on the extent of their territoriality exists, to investigate the factors mediating range defensibility. We compared the defensibility index (D) across 63 groups of howler monkeys, representing 8 different species, based on a literature review. All species, except Alouatta palliata, were classified as potentially territorial according to D, although there was high variability within and among species. Group size had a positive effect on D, probably owing to the greater ability of groups to defend a territory as they become larger. Study area had a negative effect on D, perhaps suggesting that unlike small areas, large areas allow groups to have territories that do not require significant defense from neighbors. However, population density was the factor with the strongest effect on D, with greater monitoring of home ranges under high levels of competition. Our results suggest that howler monkeys are theoretically capable of maintaining a territory and suggest that animals can show a gradient in territoriality, which can be mediated by the competitive context in which it occurs.  相似文献   

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