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Long-term field studies are critical for our understanding of animal life history and the processes driving changes in demography. Here, we present long-term demographic data for the northernmost population of mantled howler monkeys (Alouatta palliata) residing in a highly anthropogenically fragmented landscape in Los Tuxtlas, Mexico. We carried out 454 monthly group visits to 10 groups of mantled howler monkeys between 2000 and 2011. The population remained relatively stable over the 11-year study period, with an overall increase in the total number of individuals. Birth rates and inter-birth intervals were comparable to those of howler monkeys at non-fragmented sites, suggesting that living in a fragmented landscape did not affect the reproductive output of our study population. Moreover, despite the landscape, dispersal events were commonplace, including many secondary dispersals (individuals emigrating from groups that they had previously immigrated into). Finally, we found a marked effect of seasonality on the dynamics of our study population. In particular, the period of lowest temperatures and resource scarcity between November and March was associated with higher mortality and reproductive inhibition, while the period of resource abundance between April and May was associated with the majority of conceptions and weaning of offspring. This, in turn, could be influencing dispersal patterns in our study area, as male howler monkeys seem to time some of their immigrations into new groups to coincide with the start of the period of higher fertility, while females preferentially joined new groups several months before the onset of this period. These data have important implications for the conservation and management of howler monkeys in fragmented landscapes, as well as for our understanding of the effect of seasonality over howler monkey dispersal, reproduction and survival.

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The rapid loss, fragmentation and degradation of tropical forests threaten the survival of many animal species. However, the way in which these phenomena affect animal health has been poorly explored, thus limiting the design of appropriate conservation strategies. To address this, here we identified using linear mixed models the effect of proximal (diet, activity pattern, hunting and logging) and distal (sum of the basal areas of fruiting-tree species [SBAFS], landscape forest cover and degree of forest fragmentation) variables over fecal glucocorticoid metabolite (fGCM) levels-hormones associated with animal health and fitness-of six groups of spider monkeys (Ateles geoffroyi) inhabiting six landscapes with different spatial structures in Mexico. Proximal variables showed a stronger predictive power over fGCMs than distal. In this sense, increases in travel time, the occurrence of hunting, and reductions in rest time and fruit consumption resulted in higher fGCM levels. Regarding distal variables, increases in SBAFS were negatively related to fGCM levels, thus suggesting that food scarcity increases stress hormone levels. Nevertheless, contrary to theoretical expectations, spider monkeys living in smaller tracts of forest spent less time travelling, but the same time feeding on fruit as those in more forested areas. The lower net energy return associated with this combination of factors would explain why, contrary to theoretical expectations, increased forest cover was associated with increased levels of fGCMs in these groups. Our study shows that, at least in the short term, spider monkeys in fragmented landscapes do not always present higher levels of stress hormones compared to those inhabiting continuous forest, and the importance of preserving fruit sources and controlling hunting for reducing the levels of stress hormones in free ranging spider monkeys.

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Understanding the response of species to changes in landscape configuration is required to design adequate management and conservation strategies. Yet, the most appropriate spatial scale (i.e., landscape size) to assess the response of species to changes in landscape configuration (so-called "scale of effect") is largely unknown. In this paper, we assess the impact of landscape forest cover, forest fragmentation, edge density, and inter-patch isolation distance on the diet and behavior of six communities of spider monkeys (Ateles geoffroyi) in the fragmented Lacandona rainforest, Mexico. We evaluated the strength of the relationship between each landscape predictor and each response variable within ten different-sized landscapes (range = 50-665 ha) to identify the landscape size that best predicted changes in diet and behavior. The strength of most associations varied across spatial scales, with the 126-ha landscape showing the strongest relationships between landscape predictors and response variables in many cases. Yet forest cover represented the main driver of the diet and behavior of spider monkeys, being positively associated with time traveling and time feeding on wood, but negatively related to time resting and time feeding on leaves. Although weaker, the impact of edge density was opposite to forest cover for most response variables. Forest fragmentation and isolation distance showed the weakest associations with the diet and behavior of this species. Our findings thus indicate that different landscape attributes operate on different response variables at different spatial scales. Therefore, the scale of effects cannot be generalized to all response variables and to all predictors, and a multi-scale analysis will be required to accurately assess the impact of landscape configuration on species' responses.

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There are a growing number of reports of antibiotic resistance (ATBR) in bacteria living in wildlife. This is a cause for concern as ATBR in wildlife represents a potential public health threat. However, little is known about the factors that might determine the presence, abundance and dispersion of ATBR bacteria in wildlife. Here, we used culture and molecular methods to assess ATBR in bacteria in fecal samples from howler monkeys (Alouatta palliata), spider monkeys (Ateles geoffroyi), tapirs (Tapirus bairdii) and felids (jaguars, Panthera onca; pumas, Puma concolor; jaguarundis, Puma yagouaroundi; and ocelots, Leopardus pardalis) living freely in two regions of the Mexican state of Veracruz under different degrees of human influence. Overall, our study shows that ATBR is commonplace in bacteria isolated from wildlife in southeast Mexico. Most of the resistances were towards old and naturally occurring antibiotics, but we also observed resistances of potential clinical significance. We found that proximity to humans positively affected the presence of ATBR and that ATBR was higher in terrestrial than arboreal species. We also found evidence suggesting different terrestrial and aerial routes for the transmission of ATBR between humans and wildlife. The prevalence and potential ATBR transfer mechanisms between humans and wildlife observed in this study highlight the need for further studies to identify the factors that might determine ATBR presence, abundance and distribution.

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The Mexican howler monkey (Alouatta palliata mexicana) is a critically endangered primate, which is paleoendemic to Mexico. However, despite the potential significance of genetic data for its management and conservation, there have been no population genetic studies of this subspecies. To examine genetic diversity in the key remaining forest refuge for A. p. mexicana, the Selva Zoque, we amplified full-length mitochondrial control region sequences (1,100 bp) from 45 individuals and found 7 very similar haplotypes. Haplotype diversity (h = 0.486) and nucleotide diversity (π = 0.0007) were extremely low compared to other Neotropical primates. Neutrality tests, used to evaluate demographic effects (Tajima's D = -1.48, p = 0.05; Fu's F s = -3.33, p = 0.02), and mismatch distribution (sum of squares deviation = 0.006, p = 0.38; raggedness index = 0.12, p = 0.33) were consistent with a recent and mild population expansion and genetic diversity appears to be historically low in this taxon. Future studies should use a combination of mitochondrial and nuclear markers to fully evaluate genetic diversity and to better understand demographic history in A. p. mexicana. These studies should be undertaken throughout its geographic range in order to evaluate population structure and identify management units for conservation. Due to the limited distribution and population size of A. p. mexicana, future conservation strategies may need to consider genetic management. However, a more detailed knowledge of the population genetics of the subspecies is urgently recommended to maximise the conservation impact of these strategies.

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Animals' responses to potentially threatening factors can provide important information for their conservation. Group size and human presence are potentially threatening factors to primates inhabiting small reserves used for recreation. We tested these hypotheses by evaluating behavioral and physiological responses in two groups of mantled howler monkeys (Alouatta palliata mexicana) at the "Centro Ecológico y Recreativo El Zapotal", a recreational forest reserve and zoo located in the Mexican state of Chiapas. Both groups presented fission-fusion dynamics, splitting into foraging subgroups which varied in size among, but not within days. Neither subgroup size nor number of people had an effect on fecal cortisol. Out of 16 behavioral response variables tested, the studied factors had effects on six: four were affected by subgroup size and two were affected by number of people. With increasing subgroup size, monkeys increased daily path lengths, rested less, increased foraging effort, and used more plant individuals for feeding. As the number of people increased, monkeys spent more time in lower-quality habitat, and less time engaged in social interactions. Although fecal cortisol levels were not affected by the factors studied, one of the monkey groups had almost twice the level of cortisol compared to the other group. The group with higher cortisol levels also spent significantly more time in the lower-quality habitat, compared to the other group. Our results suggest that particular behavioral adjustments might allow howler monkeys at El Zapotal to avoid physiological stress due to subgroup size and number of people. However, the fact that one of the monkey groups is showing increased cortisol levels may be interpreted as a warning sign, indicating that an adjustment threshold is being reached, at least for part of the howler monkey population in this forest fragment.

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In order to better understand how patterns of parasitism in howler monkeys are affected by forest fragmentation, we carried out a 1 year survey of gastrointestinal parasites in fecal samples from three groups of Mexican howler monkeys inhabiting different forest fragments in the Los Tuxtlas Biosphere Reserve, Mexico. The study groups were chosen because the conditions in which they lived suggested a potentially negative gradient for parasite richness and a positive gradient for levels of parasitism. We report for the first time the presence of Entamoeba coli in Alouatta palliata mexicana and of hookworms (Family Ancylostomidae) in A. palliata. A reduction in home range size and an increase in disturbance was associated with a loss of parasite richness, which in general was high. Parasite prevalence and the proportion of contaminated samples in which each parasite taxon was present was also high in general and there were no differences between groups. A factor related to the generally high levels of parasitism in our study groups could be the high humidity in the study area, because this favors the survival of parasitic free forms and increases the chances of infection. This would also account for the tendency towards higher levels of parasitism observed in the rainy season. Finally we did not find a pattern relating sex and parasitism.

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The threat that forest fragmentation and habitat loss presents for several Alouatta taxa requires us to determine the key elements that may promote the persistence of howler monkeys in forest fragments and to evaluate how changes in the availability of these elements may affect their future conservation prospects. In this study we analyzed the relationship between the availability of both big trees of top food taxa (BTTFT) (diameter at breast height>60) and fruit of top food taxa (FrTFT) in the home ranges of two groups of Alouatta palliata mexicana occupying different forest fragments in Los Tuxtlas, Mexico, and their diet and activity pattern. Both study groups preferred big trees for feeding and the group with lower availability of BTTFT in their home range fed from more, smaller food sources. Furthermore, both study groups also increased the number of food sources when their consumption of fruit decreased, and the group with lower availability of FrTFT in their home range fed from more food sources. The increase in the number of food sources used under such conditions, in turn, set up a process of higher foraging effort and lower rest. In summary, our results support other studies that suggest that the availability of big trees and fruit may be two important elements influencing the persistence of howler monkeys in forest fragments.

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This paper represents the results of a long-term study (1996-2003) on the demographic changes over time of a Mexican mantled howler (Alouatta palliata mexicana) group in a rainforest fragment (40 ha) in Los Tuxtlas, Mexico, with a follow-up census 3 years later (2006). In addition to demographic and life history parameters, we describe six dispersal events. Our results suggest that this group has been expanding during the study period, growing from six to 12 individuals, with an annual average intrinsic growth rate of 0.07, an infant survivorship of 67%, and an average immature to female ratio of 0.90. This increase in size is probably related to the high food availability in their home range. However, fragment isolation may be negatively affecting the dispersal patterns typical of the species, which could result in a loss of genetic variability over time.

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The activity patterns and diet of howler monkeys (Alouatta palliata mexicana) were studied in 3 forest fragments in south-eastern Mexico: Playa Escondida (PLA), Agaltepec Island (AGA) and Arroyo Liza (LIZ). Intersite differences offered the opportunity to investigate the foraging adaptations of howler monkeys in response to population and habitat size. In the largest fragment (PLA), the howlers' diet was based on high-quality items (fruit and young leaves). In AGA, where the density of howlers was the highest, their diet was mostly folivorous with a marked exploitation of uncommon food items such as vines, lianas, shrubs and herbs. The dietary differences in AGA were accompanied by more time spent travelling and less time spent resting. Although LIZ was the smallest fragment and had a high howler density, the small group size and the use of energy-minimizing strategies (less time spent travelling and more time spent resting) probably allowed howlers to maintain a frugivorous diet.

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Accelerated deforestation is causing the rapid loss and fragmentation of primary habitat for primates. Although the genus Alouatta is one of the most studied primate taxa under these circumstances, some results are contradictory and responses of howlers to habitat fragmentation are not yet clear. In this paper, we conduct a cross-study of the available researches on mantled howlers (Alouatta palliata) in forest fragments in Los Tuxtlas, Mexico, to (1) describe the diet and activity pattern of howlers; (2) analyze the similarity in the diet across studies; and (3) relate both fragment size and howler population density with different characteristics of their diet, home range size, and activity pattern. Howlers consumed 181 plant species belonging to 54 families. Ficus was the most important taxa in the howlers' diet, followed by primary species such as Pterocarpus rohrii, Nectandra ambigens, Poulsenia armata, and Brosimum alicastrum. Secondary and non-secondary light-demanding plant species, which are representatives of disturbed habitat, contributed with a high percentage of their feeding time. Only 23% of the species consumed were the same across all the studies, suggesting that howlers adapt their diet to the food availability of their respective habitats. Population density is the best predictor of howlers' ecological and behavioral changes in response to forest fragmentation, probably owing to its relationship with food availability. Howlers respond to the increase in population densities by increasing the (1) diversity of food species in the diet; (2) consumption of non-tree growth forms; and (3) consumption of new plant items. Home range size is also predicted by population density, but fragment size is a better predictor, probably owing to the fact that howler groups can overlap their home ranges. Our results emphasize the importance of conserving the larger fragments and increasing the size of small and medium-sized ones.

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This research focuses on identifying the principal habitat characteristics that influence the presence and abundance of mantled howlers in forest fragments. We provide information on the demography of several fragmented Alouatta palliata mexicana subpopulations at Los Tuxtlas, Mexico, and relate this to the biogeographical and floristic characteristics of the forest fragments inhabited. The most important habitat characteristics related to the presence and abundance of howlers in the fragments were fragment size and floristic diversity. On the other hand, some evidence suggests that given the conditions under which howlers in our study area live (i.e., small and degraded fragments with high densities), secondary vegetation may be beneficial for the survival of the howlers. Finally, we discuss the possibility that the very low immature-to-female ratio (IFR) in the groups, and the lack of juveniles found in many of the study groups may be due to high mortality rates in immatures. A reduction in food availability because of the high population densities of these groups may be responsible for this process.

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