Demographic population structure of black howler monkeys in fragmented and continuous forest in Chiapas, Mexico: Implications for conservation.

For wild primates, demography studies are increasingly recognized as necessary for assessing the viability of vulnerable populations experiencing rapid environmental change. In particular, anthropogenic changes such as habitat loss and fragmentation can cause ecological and behavioral changes in small, isolated populations, which may, over time, alter population density and demographic structure (age/sex classes and group composition) in fragment populations relative to continuous forest populations. We compared our study population of Endangered black howler monkeys (Alouatta pigra) in 34 forest fragments around Palenque National Park (PNP), Mexico (62 groups, 407 individuals), to the adjacent population in PNP, protected primary forest (21 groups, 134 individuals), and to previous research on black howlers in fragments in our study area (18 groups, 115 individuals). We used χ2 and Mann-Whitney U tests to address the questions: (a) what is the current black howler demographic population structure in unprotected forest fragments around PNP? (b) How does it compare to PNP's stable, continuous population? (c) How has it changed over time? Compared to the PNP population, the fragment populations showed higher density, a significantly lower proportion of multimale groups, and significantly fewer adult males per group. The population's age/sex structure in the fragmented landscape has been stable over the last 17 years, but differed in a higher proportion of multifemale groups, higher density, and higher patch occupancy in the present. In the context of conservation, some of our results may be positive as they indicate possible population growth over time. However, long-term scarcity of adult males in fragments and associated effects on population demographic structure might be cause for concern, in that it may affect gene flow and genetic diversity. The scarcity of adult males might stem from males experiencing increased mortality while dispersing in the fragmented landscape, whereas females might be becoming more philopatric in fragments.

Agonism and dominance in female blue monkeys.

Agonistic behavior features prominently in hypotheses that explain how social variation relates to ecological factors and phylogenetic constraints. Dominance systems vary along axes of despotism, tolerance, and nepotism, and comparative studies examine cross-species patterns in these classifications. To contribute to such studies, we present a comprehensive picture of agonistic behavior and dominance relationships in wild female blue monkeys (Cercopithecus mitis), an arboreal guenon, with data from 9 groups spanning 18 years. We assessed where blue monkeys fall along despotic, tolerant, and nepotistic spectra, how their dominance system compares to other primates, primarily cercopithecines, and whether their agonistic behavior matches socioecological model predictions. Blue monkeys showed low rates of mainly low-intensity agonism and little counter-aggression. Rates increased with rank and group size. Dominance asymmetry varied at different organizational levels, being more pronounced at the level of interactions than dyad or group. Hierarchies were quite stable, had moderate-to-high linearity and directional consistency and moderate steepness. There was clear maternal rank inheritance, but inconsistent adherence to Kawamura's rules. There was little between-group variation, although hierarchy metrics showed considerable variation across group-years. Overall, blue monkeys have moderately despotic, moderately tolerant, and nepotistic dominance hierarchies. They resemble other cercopithecines in having significantly linear and steep hierarchies with a generally stable, matriline-based structure, suggesting a phylogenetic basis to this aspect of their social system. Blue monkeys most closely match Sterck et al.'s [1997] Resident-Nepotistic-Tolerant dominance category, although they do not fully conform to predictions of any one socioecological model. Our results suggest that socioecological models might better predict variation within than across clades, thereby incorporating both ecological variables and phylogenetic constraints. Our findings also highlight the need for clearer definitions of socioecologically relevant dominance categories, which would ideally derive from quantitative measures of dominance behavior. Intraspecific and methodological variation may, however, be a challenge.

Effects of variation in forest fragment habitat on black howler monkey demography in the unprotected landscape around Palenque National Park, Mexico.

Habitat loss and fragmentation are leading threats to biodiversity today, and primates are particularly vulnerable to anthropogenic habitat disturbance. However, few studies have examined how differential effects of variation in forest fragment characteristics on males and females in a primate population may affect demography and population persistence. We quantified the effects of variation in forest fragment characteristics on the within-fragment demography of black howler monkeys (Alouatta pigra) in forest fragments around Palenque National Park, Mexico, and how these effects differed between adult males and females. We quantified forest loss in the landscape between 2000 and 2017, and used a redundancy analysis to examine the effects of 15 variables quantifying fragment dimensions, forest composition and physical structure, and isolation on fragment population size and density, the proportion of adult males and females in the fragment population, and the mean number of adult males and females per group in 34 fragments (N = 393 monkeys). We hypothesized that (i) population size is positively correlated with fragment area, while population density is negatively correlated, and (ii) the composition of fragment populations results from differential effects of fragment variables on adult males and females. Forest cover decreased by 23.3% from 2000 to 2017. Our results showed a significant effect of fragment variables on population demography in fragments, accounting for 0.69 of the variance in the demographic response variables. Population size increased with fragment area and connectivity, while density decreased. Larger, less isolated fragments with better connectivity, characteristics indicative of abundant secondary growth, and those with more diverse vegetation but lower Simpson's evenness indices tended to have more adult females per group and a higher proportion of adult females in the population. In contrast, fragments that were largely similar in characteristics of forest composition and structure, but that were more isolated from nearby fragments, had more adult males per group and a higher proportion of adult males. These results may stem from black howler females preferentially remaining in natal groups and fragments when possible, and dispersing shorter distances when they disperse, while males may be more likely to disperse between fragments, traveling longer distances through the matrix to more isolated fragments. These differential effects on males and females have important conservation implications: if females are more abundant in larger, less isolated fragments, while males are more abundant in more isolated fragments, then to effectively conserve this population, both landscape connectivity and fragment areas should be maintained and increased.

Priority Questions and Horizon Scanning for Conservation: A Comparative Study.

Several projects aimed at identifying priority issues for conservation with high relevance to policy have recently been completed in several countries. Two major types of projects have been undertaken, aimed at identifying (i) policy-relevant questions most imperative to conservation and (ii) horizon scanning topics, defined as emerging issues that are expected to have substantial implications for biodiversity conservation and policy in the future. Here, we provide the first overview of the outcomes of biodiversity and conservation-oriented projects recently completed around the world using this framework. We also include the results of the first questions and horizon scanning project completed for a Mediterranean country. Overall, the outcomes of the different projects undertaken (at the global scale, in the UK, US, Canada, Switzerland and in Israel) were strongly correlated in terms of the proportion of questions and/or horizon scanning topics selected when comparing different topic areas. However, some major differences were found across regions. There was large variation among regions in the percentage of proactive (i.e. action and response oriented) versus descriptive (non-response oriented) priority questions and in the emphasis given to socio-political issues. Substantial differences were also found when comparing outcomes of priority questions versus horizon scanning projects undertaken for the same region. For example, issues related to climate change, human demography and marine ecosystems received higher priority as horizon scanning topics, while ecosystem services were more emphasized as current priority questions. We suggest that future initiatives aimed at identifying priority conservation questions and horizon scanning topics should allow simultaneous identification of both current and future priority issues, as presented here for the first time. We propose that further emphasis on social-political issues should be explicitly integrated into future related projects.

Effect of unknown relationships on linearity, steepness and rank ordering of dominance hierarchies: simulation studies based on data from wild monkeys.

The presence of unknown dyadic relationships is a common problem in constructing dominance hierarchies for groups of social animals. Although previously acknowledged, the influence of unknown relationships on hierarchy measures like linearity and steepness has not been studied in detail. Using real data-sets from four groups of wild monkeys, we illustrate how unknown relationships affect linearity and steepness of hierarchies and the consistency of rank ordering based on de Vries' I&SI method. Monte Carlo simulations revealed significant negative linear relationships between the proportion of unknown relationships and both linearity and steepness. These simulations over-estimated steepness and linearity indices relative to additional real-data input matrices. Rank orders became inconsistent at 26-38% unknown relationships, depending on the group. Group size and the specific input matrix substantially affected how much unknown relationships influenced steepness and linearity, the values of these indices and the point at which rank order became inconsistent. We recommend caution in characterizing the dominance structure of a group with many unknown relationships, and in drawing conclusions about hierarchy linearity and steepness based on few input matrices, especially if they contain many unknown relationships. Quantitative characterizations of hierarchies are perhaps best viewed as a somewhat fluid range rather than fixed values.