Ranging and territoriality in red-shanked doucs (Pygathrix nemaeus) at Son Tra Nature Reserve, Vietnam.

The home ranges of odd-nosed colobines as reported by a number of studies can vary greatly both within and between taxa. Evaluating Pygathrix, particularly within the context of the odd-nosed colobines, has proven challenging due to the comparable dearth of published data on all three species-Pygathrix nigripes, Pygathrix cinerea, and Pygathrix nemaeus. This dearth has also proven challenging when assessing territoriality within Pygathrix and contextualizing this behavior to odd-nosed colobines. For this study, we set out to assess home range, daily path length, and territoriality in a population of P. nemaeus at Son Tra Nature Reserve in Vietnam. We collected a total of 259 h of behavioral data using 10-min focal follows with 1-min instantaneous scans, including GPS recordings for each follow. We collected phenology and weather data, which were then used with Pearson's correlations to test whether ecological factors influenced variation in ranging or daily path length at both the monthly and seasonal levels. Finally, we used the Mitani-Rodman defendability index to assess territoriality in P. nemaeus. We found that our population of P. nemaeus had a home range of 36.00 ha with a core area of 8.02 ha and an average daily path length of 509.11 ± 244.87 m. Daily path length was correlated to precipitation such that daily path lengths were shorter during increases in rainfall (r(17) = 0.52, p = .03). These measures resulted in a defendability index of 0.75, suggesting that the population of P. nemaus on Son Tra does not travel far enough each day to actively defend territorial borders.

Food availability, plant diversity, and vegetation structure drive behavioral and ecological variation in Endangered Coimbra-Filho's titi monkeys.

There is wide variability in primate behavior and ecology. Understanding how frugivorous primates behave under different habitat fragmentation levels is key for effective conservation and management of species and their habitats. We evaluated the seasonality in activity budget, diet, and ranging behavior of two groups of Endangered Coimbra-Filho's titi monkeys (Callicebus coimbrai). One group inhabited a 14-ha forest fragment, whereas the other lived in a 522-ha fragment. We measured the monthly density of trees and lianas available as food sources over 8 months. We also collected behavioral and group location data every 5 min, from dawn to dusk, using the scan sampling method. The two forest fragments differed seasonally in the number of fruiting food-resource available. In the 14-ha fragment, we found that the time spent by titi monkeys feeding, foraging, resting, and traveling differed seasonally. In the 522-ha fragment, titi monkeys exhibited seasonal differences in time spent sleeping, socializing, foraging, and revisiting food sources. In both titi monkey groups, diets varied seasonally. Our findings indicate that Coimbra-Filho's titi monkeys can exhibit behavioral flexibility in their activity budgets, diets, and movement patterns. Such flexibility is important for this species to survive in fragmented habitats and may be linked to three key factors: species-specific resource availability, plant species diversity, and the vegetation structure of each forest fragment.

Ranging behavior and the potential for territoriality in pair-living titi monkeys (Plecturocebus discolor).

Patterns of ranging behavior and space use are key for evaluating current ideas about the evolution and maintenance of pair-living and sexual monogamy as they provide insights into the dispersion of females, the potential for territoriality, and whether males are limited to defending an area that can support only one female and her offspring. We examined ranging behavior and space use to evaluate the potential for territoriality in five groups of red titi monkeys (Plecturocebus discolor) during a 10-year study in Ecuadorian Amazonia. Mean home range size, calculated using a time-sensitive local convex hull estimation procedure, was 4.0 ± 1.4 ha. Annual home ranges of neighboring groups overlapped, on average, 0%-7%. Mean daily path length was 670 ± 194 m, resulting in defendability indices of 2.2-3.6 across groups. Groups visited, on average, 4 of 12 sections of their home range border area per day, but that was not more often than would be expected by chance, and intergroup encounters were infrequent. We did not find evidence of active monitoring for intruders in border areas, in that groups did not travel either faster or slower when at the border than when in central areas of their range. The absence of overt monitoring might be compensated for by engaging in loud calls, which the study groups did throughout their home ranges; these calls may serve as an advertisement of occupancy and a deterrent to intruding conspecifics. Our finding that red titis have a high potential for territoriality is consistent with several of the main hypotheses proposed to explain pair-living in mammals.

Sex-Segregated Range Use by Black-and-White Ruffed Lemurs (Varecia variegata) in Ranomafana National Park, Madagascar.

Ranging behavior is one important strategy by which nonhuman primates obtain access to resources critical to their biological maintenance and reproductive success. As most primates live in permanent social groups, their members must balance the benefits of group living with the costs of intragroup competition for resources. However, some taxa live in more spatiotemporally flexible social groups, whose members modify patterns of association and range use as a method to mitigate these costs. Here, we describe the range use of one such taxon, the black-and-white ruffed lemur (Varecia variegata), at an undisturbed primary rain forest site in Ranomafana National Park, Madagascar, and characterize sex differences in annual home range area, overlap, and daily distances traveled. Moreover, we characterize seasonal variability in range use and ask whether ranging behaviors can be explained by either climatic or reproductive seasonality. We found that females used significantly larger home ranges than males, though sexes shared equal and moderate levels of home range overlap. Overall, range use did not vary across seasons, although within sexes, male range use varied significantly with climate. Moreover, daily path length was best predicted by day length, female reproductive state, and sex, but was unrelated to climate variables. While the patterns of range use and spatial association presented here share some similarities with "bisexually bonded" community models described for chimpanzees, we argue that ruffed lemurs best conform to a "nuclear neighborhood" community model wherein nuclear (core) groups share the highest levels of home range overlap, and where these groups cluster spatially into adjacent "neighborhoods" within the larger, communally defended territory.

Ranging behavior and potential for territoriality in equatorial sakis (Pithecia aequatorialis) in Amazonian Ecuador.

OBJECTIVES: Territoriality refers to the consistent defense of an area within the home range (HR) against intrusions of conspecifics. It implies exclusive space use with low degree of overlap among neighboring groups, high site fidelity, specific ranging behavior such as high mobility relative to HR size and frequent visits of territory borders, and monitoring behavior. We examined ranging behavior and use of space to evaluate territoriality in Pithecia aequatorialis in Ecuador. MATERIALS AND METHODS: Between 2005 and 2015, we monitored one main study group continuously and five additional groups for shorter periods (5 months to 2.5 years) at the Tiputini Biodiversity Station, in eastern Ecuador. We scored the location of the study groups at 20 min intervals during, on average, 5 days per month. We estimated saki HRs and core areas (CAs) using the fixed kernel density method (95 and 50%, respectively). RESULTS: The average HR size was 57 ha and the average CA 14 ha. The degree of overlap between HRs of neighboring groups was low (2-9%). For the main study group, the average overlap between annual HRs was 82%. Mean daily path length across groups was 1,151 m; the defensibility index varied between 1.1 and 2.3 (values >1 are suggestive of territoriality), and the fractional monitoring rate varied between 0.06 and 0.15 (values >0.08 are suggestive of territoriality). Groups did not visit their HR borders (100 m inner buffer) more often than would be expected by chance. Travel speed and directness were comparable between the borders and the centers of groups' HRs. DISCUSSION: Our multiyear study suggests that equatorial sakis show low degree of range overlap and high site fidelity and have the potential to be territorial, given their high mobility relative to HR size that allows for frequent border monitoring. Nevertheless, their movement patterns in border areas did not reveal evidence for monitoring behavior.

Surviving in the urban-rural interface: Feeding and ranging behavior of brown howlers (Alouatta guariba clamitans) in an urban fragment in southern Brazil.

Wild primates that live in urban areas face extreme threats that are less frequent in nonurban fragments, such as the presence of dangerous matrix elements (e.g., roads, power lines, buildings, and a high density of domestic dogs near food patches), that could influence their movements, feeding behavior, and survival. However, the scarcity of studies addressing this issue hinders our understanding of the behavioral adjustments that favor the survival of primates in urban areas. For 12 months, we studied a six-individual group of brown howlers (Alouatta guariba clamitans) in an urban fragment to determine (i) their diet richness and its relationships with food availability, (ii) their daily path length (DPL) and the matrix elements used during movement, and (iii) the main ecological drivers of the DPL. Sampling effort totaled 72 days, 787 hr, and 3,224 instantaneous scans. We found that the diet of brown howlers contained 35 plant species (including seven cultivated crops) belonging to 33 genera and 21 families. The consumption of fruits and young leaves was directly related to their temporal availability. The average DPL (446 m) was smaller than that reported for groups of howlers inhabiting large nonurban fragments. To move between food patches, animals used three main matrix elements: trees, power lines, and roofs. The number of plant species used during the day was the main driver of DPL. Our findings highlighted that the generalist-opportunistic diet of brown howlers and their ability to move across the anthropogenic matrix using artificial elements such as power lines, roofs, and wildlife crossings represent a remarkable part of their behavioral repertory in Lami. However, there are potential costs associated with these strategies, namely, electrocution and predation by domestic dogs. Thus, it is urgent to further investigate how these behaviors could affect the long-term survival of these animals.

Long-term home range use in white-handed gibbons (Hylobates lar) in Khao Yai National Park, Thailand.

Ranging behavior is an important element of how nonhuman primates obtain sufficient resources to ensure biological maintenance and reproductive success. As most primates live in permanent social groups, group members must balance the benefits of group living with the costs of intragroup competition for resources. One way to mitigate the cost of intragroup feeding competition is to increase foraging-related travel, thereby increasing the number of patches visited. As a result we might expect home range size to increase as a function of group size. On the other hand, for perennially territorial species, ranging behavior may be constrained by the ranging requirements of territorial defense or by the location of neighboring territories, which would result in long-term stability in the size and location of a group's home range. In this study, we examined changes in range-use characteristics in one well-habituated group of white-handed gibbons (Hylobates lar) during three study periods over a 10-year span. Group size changed from five members, two adults, two juveniles, and one infant, in 1994, to two adults in 2002, and to three adults and one sub-adult in 2004. Despite inter-annual changes in core area use we found that home range location was highly stable across years. Nevertheless, home range size was larger and daily path length significantly longer in 2002 relative to 1994 when a dependent infant was present in the group. The percentage of time adults spent resting was also significantly greater in 1994 when the infant was present. These findings highlight the importance of considering group composition, in addition to group size, when evaluating the determinants of ranging behavior. We also consider the influence of individual and shared knowledge on home range stability.

Ranging behavior drives parasite richness: A more parsimonious hypothesis.

Parasitism is a ubiquitous interspecific interaction that may play an important role in the evolution of hosts and parasites, molding many aspects of their behavior and ecology. Detecting behavioral changes of hosts infected with parasites is not a straightforward task. Extrapolating from individual-level responses to group-level decision-making is still a much more complex challenge. The ranging behavior of hosts that live in social groups is a good example. Many hypotheses of the cause-effect relationship between this behavior and parasite diversity and load have been proposed. For instance, Brockmeyer et al. [2015, Am. J. Primatol. 77:1036-1048] recently suggested that the richness of protozoan parasites influences the daily path length of free-ranging mandrills. We believe that this explanation for the relationship contains several implicit assumptions. Therefore, we offer an alternative, more parsimonious hypothesis in which daily path length is the driver of parasite richness rather than its consequence. Our hypothesis only assumes that ranging farther exposes animals to a richer parasite diversity. We discuss the data required to test these alternative hypotheses and recall empirical evidence and theoretical modeling results supporting or rejecting their assumptions. We also propose a model of the expected outcomes in terms of species richness, load, intensity of infection, and within-group community similarity of non-lethal environmentally transmitted parasites in social animal groups showing distinct patterns of range use. Am. J. Primatol. 78:923-927, 2016. © 2016 Wiley Periodicals, Inc.

Impact of cost distance and habitat fragmentation on the daily path length of Rhinopithecus bieti.

An understanding of primate movement patterns in response to natural and anthropogenically induced changes in habitat heterogeneity, food availability, and plant species distribution is essential for developing effective management and conservation programs. Therefore, from July 2013 to June 2014, we examined the effects of landscape configuration on the ranging behavior (daily path length, DPL) of the Endangered Yunnan snub-nosed monkey (Rhinopithecus bieti) in the Baimaxueshan National Nature Reserve (27°34'N, 99°17'E) in Gehuaqing, China. Given the extreme difficulties in following the study group across high altitude mountainous terrain across an elevation of 2,500-4,000 m, we were only able to collect DPL using 3-4 GPS points per day on 21 individual days. We found that R. bieti traveled the shortest DPL in winter (1,141.31 m), followed by spring (2,034.06 m) and autumn (2,131.19 m). The cost distance, a statistical tool designed to estimate the difficulty of a species moving across its distributional range, was lowest in autumn (205.47), followed by spring (225.93) and winter (432.59) (one-way ANOVA: F = 3.852, P = 0.026, df = 2). The habitat fragmentation index (HFI), which measures the density of forest patches, indicated areas visited in the winter were more fragmented (HFI = 2.16) compared to spring (HFI = 1.83) or autumn (HFI = 1.3). Although our results should be considered preliminary, they suggest that both the availability of suitable travel routes and habitat fragmentation, driven by high-intensity human disturbance, constrain the movement of R. bieti. We found that undisturbed areas of the bands' range contained a high density of lichens, which represent a nutritious and abundant and year-round food source for Yunnan snub-nosed monkeys. In order to protect this Endangered species, we recommend that researchers construct detailed maps of landscape heterogeneity, particularly habitat connectivity, forest fragmentation, and seasonal variation in the location of major food patches in order to better understand and mitigate the effects of seasonal habitat change on patterns of R. bieti habitat utilization and population viability.

Ranging behavior of eastern hoolock gibbon (Hoolock leuconedys) in a northern montane forest in Gaoligongshan, Yunnan, China.

Generally, food abundance and distribution exert important influence on primate ranging behavior. Hoolock gibbons (genus Hoolock) live in lowland and montane forests in India, Bangladesh, Myanmar and China. All information about hoolock gibbons comes from studies on western hoolock gibbons (Hoolock hoolock) living in lowland forest. Between August 2010 and September 2011, we studied the ranging behavior of one habituated group of eastern hoolock gibbon (H. leuconedys) living in a seasonal montane forest in Gaoligongshan, Yunnan, China. Results show that the study group did not increase foraging effort, calculated in this study as the daily path length, when fruit was less available. Instead, the gibbons fed more on leaves and decreased traveling to conserve energy. They relied heavily on a single food species in most study months which was patchily distributed within their total (14-month) home range, and during most months they used only a small portion of their total home range. In order to find enough food, the group shifted its monthly home range according to the seasonal availability of food species. To satisfy their annual food requirements, they occupied a total home range of 93 ha. The absence of neighboring groups of gibbons and the presence of tsaoko cardamom (Amomum tsaoko) plantations may also have influenced the ranging behavior of the group. Further long-term studies of neighboring groups living in intact forests are required to assess these effects.