Same-sex sexual behaviour among mammals is widely observed, yet seldomly reported: Evidence from an online expert survey.

Same-sex sexual behaviour (SSSB) occurs in most animal clades, but published reports are largely concentrated in a few taxa. Thus, there remains a paucity of published reports for most mammalian species. We conducted a cross-sectional expert survey to better understand the underlying reasons for the lack of publications on this topic. Most respondents researched Primates (83.6%, N = 61), while the rest studied Carnivora (6.9%, N = 5), Rodentia (4.1%, N = 3), Artiodactyla (2.7%, N = 2), and Proboscidea (2.7%, N = 2). Most respondents (76.7%, N = 56) had observed SSSB in their study species, but only 48.2% (N = 27) collected data on SSSB, and few (18.5%, N = 5) had published papers on SSSB. Of the unique species identified as engaging in SSSB in the survey, 38.6% (N = 17) have no existing reports of SSSB to the knowledge of the authors. In both the survey questions and freeform responses, most respondents indicated that their lack of data collection or publication on SSSB was because the behaviours were rare, or because it was not a research priority of their lab. No respondents reported discomfort or sociopolitical concerns at their university or field site as a reason for why they did not collect data or publish on SSSB. Multiple logistic regressions were performed to assess whether taxa studied, education level, or identification within the LGBTQ+ community predicted observing, collecting data on, or publishing on SSSB, but none of these variables were significant predictors. These results provide preliminary evidence that SSSB occurs more frequently than what is available in the published record and suggest that this may be due to a publishing bias against anecdotal evidence.

When population growth intensifies intergroup competition, female colobus monkeys free-ride less.

Intergroup aggression often results in the production of public goods, such as a safe and stable social environment and a home range containing the resources required to survive and reproduce. We investigate temporal variation in intergroup aggression in a growing population of colobus monkeys (Colobus vellerosus) to ask a novel question: "Who stepped-up to produce these public goods when doing so became more difficult?". Both whole-group encounters and male incursions occurred more frequently as the population grew. Males and females were both more likely to participate in whole-group encounters when monopolizable food resources were available, indicating both sexes engaged in food defence. However, only females increasingly did so as the population grew, suggesting that it was females who increasingly produced the public good of home range defence as intergroup competition intensified. Females were also more active in male incursions at high population densities, suggesting they increasingly produced the public good of a safe and stable social environment. This is not to say that males were chronic free-riders when it came to maintaining public goods. Males consistently participated in the majority of intergroup interactions throughout the study period, indicating they may have lacked the capacity to invest more time and effort.

Coping with extreme free cortisol levels: Seasonal stress axis changes in sympatric North American flying squirrels.

Most environments exhibit predictable yearly changes, permitting animals to anticipate them. The hypothalamic-pituitary-adrenal (HPA) axis is a key physiological pathway that enables animals to cope with such changes. Monitoring glucocorticoid (the end products of the HPA axis) levels in wild animals throughout the year can improve our understanding of how this pathway responds to different conditions. For this study, we collected 18 months of data on two species of North American flying squirrels (Glaucomys sabrinus and G. volans) living in a southern Ontario forest where temperature and food availability fluctuate dramatically throughout the year. These squirrels are active year-round, nest communally, and rely on scatter hoarded foods in the winter months. Flying squirrels have extremely high levels of free plasma cortisol relative to other mammals, but it is unknown how these levels are affected by environmental and reproductive factors. For both species, our goals were to (1) validate an enzyme immunoassay (EIA) to measure their fecal glucocorticoid metabolite (FGM) concentrations and (2) assess yearly differences, seasonal changes, and the influence of sex, reproduction, and ambient temperature on FGM concentrations in each species. In the lab, we successfully validated the use of antibody 5α-pregnane-3ß, 11ß, 21-triol-20-one EIA for FGM analysis in both species. In the field, neither sex nor reproductive status (breeding condition or not) were linked to FGM concentrations in either species. FGM concentrations were higher in autumn compared to the spring and summer. There were no other seasonal differences. We discuss possible explanations for the autumn peak in FGM concentrations (increased energy expenditure and social nesting changes), as well as outline possible avenues for future research. Understanding how individuals and populations respond to environmental change is a critical goal in evolutionary ecology, particularly in the context of a rapidly changing Anthropocene.

How can we apply decision-making theories to wild animal behavior? Predictions arising from dual process theory and Bayesian decision theory.

Our understanding of decision-making processes and cognitive biases is ever increasing, thanks to an accumulation of testable models and a large body of research over the last several decades. The vast majority of this work has been done in humans and laboratory animals because these study subjects and situations allow for tightly controlled experiments. However, it raises questions about how this knowledge can be applied to wild animals in their complex environments. Here, we review two prominent decision-making theories, dual process theory and Bayesian decision theory, to assess the similarities in these approaches and consider how they may apply to wild animals living in heterogenous environments within complicated social groupings. In particular, we wanted to assess when wild animals are likely to respond to a situation with a quick heuristic decision and when they are likely to spend more time and energy on the decision-making process. Based on the literature and evidence from our multi-destination routing experiments on primates, we find that individuals are likely to make quick, heuristic decisions when they encounter routine situations, or signals/cues that accurately predict a certain outcome, or easy problems that experience or evolutionary history has prepared them for. Conversely, effortful decision-making is likely in novel or surprising situations, when signals and cues have unpredictable or uncertain relationships to an outcome, and when problems are computationally complex. Though if problems are overly complex, satisficing via heuristics is likely, to avoid costly mental effort. We present hypotheses for how animals with different socio-ecologies may have to distribute their cognitive effort. Finally, we examine the conservation implications and potential cognitive overload for animals experiencing increasingly novel situations caused by current human-induced rapid environmental change.

Examining postmounting grooming in male golden snub-nosed monkeys to investigate the functions of same-sex mounts.

Same-sex mounting behavior has been documented across primate species and several biological functions, that are often difficult to differentiate, have been proposed. Postmounting affiliative behaviors (e.g., embracing and grooming) have been partly overlooked and their performance may more clearly reveal the function of same-sex mounts for different age-sex classes. Here, considering postmounting grooming behaviors (PMG), we tested the affiliation hypothesis for same-sex mounts in an all-male unit (AMU) of captive golden snub-nosed monkeys (Rhinopithecus roxellana) (n = 7-8) and the needing to learn hypothesis in the juveniles of the AMU (n = 4) and the juvenile male from the neighboring one-male unit in Shanghai Wild Animal Park, China from November 2014 to June 2015. A total of 1986 same-sex mounts were recorded from the AMU individuals and the juvenile of the neighboring one-male unit. We found that neither dyadic proximity-based association nor grooming-based affiliation predicted the occurrence of mounts in the AMU, and PMG was more likely to occur in nontense than tense social contexts, providing no support for the affiliation hypothesis. Although dyadic grooming-based affiliation predicted the occurrence of PMG, it exerted no influence on the occurrence of mounts, which was necessary to support the hypothesis. However, consistent with the needing to learn hypothesis, from ages two to four, juveniles' mounting duration increased and they performed pelvic thrusting during mounting more and more often as they grew older. Reciprocated series mounts were much more common among juveniles than other mounting dyads, providing learning opportunities for both participants. The mounter was more likely to groom the mountee at the end of a mount among juveniles than other mounting dyads, suggesting that juvenile mounters may profit from mounting and groom mountees in return for providing them with opportunities to learn copulatory skills. A hypothesis that merits further investigation.

Monkeys who experience more feeding competition utilize social information to learn foraging skills faster.

Animals must learn foraging skills to successfully survive and reproduce but the sources of interindividual variation in learning are poorly understood. For example, there is little consensus on the role motivation plays, even though it is a key factor impacting learning outcomes in humans. Here, we conduct a field experiment on a wild primate to investigate whether an individual's vulnerability to feeding competition impacts their motivation to learn a beneficial foraging technique. We provided a group of monkeys with a food reward (i.e., a half banana) that needed to be retrieved from a box. The monkeys discovered an efficient technique that consistently allowed them to retrieve the banana quickly, decreasing the risk of food loss to competitors. We found that individuals who frequently experienced feeding competition learned this efficient technique significantly faster than individuals who rarely foraged in the presence of a dominant competitor. They appeared to use social learning to learn faster as they were more attentive to the handling techniques others used and improved their foraging skills after opportunities to observe a skilled demonstrator. These findings support that an individual's vulnerability to feeding competition impacts their motivation to learn foraging skills that reduce food loss to competitors.

No food left behind: foraging route choices among free-ranging Japanese macaques (Macaca fuscata) in a multi-destination array at the Awajishima Monkey Center, Japan.

Animals must make route choices every day when moving through their habitat while foraging. Choosing an optimal route can be cognitively costly, and primates and other animals have been shown to use simple heuristics, "rules of thumb", to make foraging route choices. We investigated the potential use of heuristics among foraging free-ranging Japanese monkeys (Macaca fuscata) during solitary foraging trials. We also investigated the potential influence of individual variables (age and sex) and social variables (presence in the central group, presence of potential inter- and intraspecific competitors), on the use of heuristics, route length and trial time. We used a multi-destination foraging experiment with 6 platforms in a (4 m × 8 m) Z-array, completed by 29 Japanese macaques in 155 runs at the Awajishima Monkey Center in Japan. Our results showed that the macaques chose routes consistent with heuristics (e.g. nearest neighbour heuristic 19.4%, convex hull heuristic 4.5%) and selected optimal routes (shortest path in 23.9% of the trials). We also identified a potential new heuristic that was used most frequently, that we termed the "sweep heuristic" (27.1% of trials), which we interpreted as a strategy to deal with competitive foraging trade-offs - choosing routes to prioritize not leaving isolated food pieces behind. Age was significantly related to trial time; juvenile macaques were faster than adults and young adults, using speed to gain access to resources. Solitary trials with conspecifics present took significantly longer routes. Our results suggest that contextual factors led to variation in Japanese macaque decision-making, and we suggest that the preferential use of a sweep heuristic may have been a response to high intragroup competition.

Non-Reproductive Sexual Behavior in Wild White-Thighed Colobus Monkeys (Colobus vellerosus).

Rare behaviors are often missing from published papers, hampering phylogenetic analyses. Here, we report, for the first time, masturbation and same-sex sexual behavior (SSB) in both male and female black-and-white colobus monkeys. We recorded these behaviors during 32 months of observation (1573 h of focal animal sampling) on Colobus vellerosus collected at the Boabeng-Fiema Monkey Sanctuary in Ghana. Males were observed masturbating and involved in SSB more than females. Subadult males were the age-sex class that engaged in both of these behaviors most often and a third of all SSB observed in young males occurred when they were forming an all-male band (AMB), which are temporally transient social groups in this species. Our data support that masturbation in males may be a sexual outlet for individuals that do not have a current sexual partner, while in females it may function in mate attraction by advertising receptivity. SSB may occur as an evolutionary byproduct but given the temporal clustering of observed events in males prior to AMB formation, our data best support the hypothesis that these behaviors facilitate male-male bonding (i.e., act as social glue). Within AMB's, males engage in coalitionary behavior to take over social groups containing females and strong bonds are important for success and later access to females, which could have selected for SSB in C. vellerosus.

Mild movement sequence repetition in five primate species and evidence for a taxonomic divide in cognitive mechanisms.

When animals forage, they face complex multi-destination routing problems. Traplining behaviour-the repeated use of the same route-can be used to study how spatial memory might evolve to cope with complex routing problems in ecologically distinct taxa. We analyzed experimental data from multi-destination foraging arrays for five species, two cercopithecine monkeys (vervets, Chlorocebus pygerythrus, and Japanese macaques, Macaca fuscata) and three strepsirrhines (fat-tailed dwarf lemurs, Cheirogaleus medius, grey mouse lemurs, Microcebus murinus, and aye-ayes, Daubentonia madagascariensis). These species all developed relatively efficient route formations within the arrays but appeared to rely on variable cognitive mechanisms. We found a strong reliance on heuristics in cercopithecoid species, with initial routes that began near optimal and did not improve with experience. In strepsirrhines, we found greater support for reinforcement learning of location-based decisions, such that routes improved with experience. Further, we found evidence of repeated sequences of site visitation in all species, supporting previous suggestions that primates form traplines. However, the recursive use of routes was weak, differing from the strategies seen in well-known traplining animals. Differences between strepsirrhine and cercopithecine strategies may be the result of either ecological or phylogenetic trends, and we discuss future possibilities for disentangling the two.

Temporal patterns in the social network of core units in Rwenzori Angolan colobus monkeys: Effects of food availability and interunit dispersal.

Multi-level societies are complex, nested social systems where basic social groups (i.e., core units) associate in a hierarchical manner, allowing animals to adjust their group sizes in response to variables such as food availability, predation, or conspecific threat. These pressures fluctuate over time and examining the extent to which this variation affects the clustering of core units into different tiers may be instrumental in understanding the evolution of multi-level societies.The goal of our study was to determine the degree of temporal variability in interunit associations in a multi-level society of Rwenzori Angolan colobus monkey (Colobus angolensis ruwenzorii), and to determine the social and ecological factors that underlie association patterns. The C. a. ruwenzorii multi-level society consists of at least three tiers, with core units clustering into clans that share a home range in a band tier.We performed social network analyses on 21 months of association data from 13 core units (totaling 139 identifiable individuals) at Lake Nabugabo, Uganda. We described the patterns of variation in core-unit associations over time and investigated how changes in rainfall, food availability, and interunit dispersals were correlated with these associations over the short-term (month to month) and long-term (year to year).Although clans were relatively stable, larger-scale changes in association patterns included the formation of an all-male unit and the transfer of one core unit between clans (within the band tier). Seasonally, core units associated significantly more when fruit, their preferred food source, was abundant (i.e., social networks were denser and more clustered) and there was no direct effect of rainfall seasonality or young leaf availability. Male dispersals also occurred more during periods of high fruit availability, suggesting that greater band cohesion allowed males to prospect and transfer between core units. Once males transferred, their previous and new units associated significantly more with one another than with other core units for 1-2 months postdispersal. The dispersal of five males from one core unit to another in a different clan co-occurred with this core unit switching its clan affiliation.By examining temporal shifts in social network structure among core units, this study shows the interconnected roles that food availability and dispersal have in shaping the C. a. ruwenzorii multi-level social system. Our findings highlight how ecological conditions can drive association patterns, impact interunit relationships, and influence social organization.

High levels of infant handling by adult males in Rwenzori Angolan colobus (Colobus angolensis ruwenzorii) compared to two closely related species, C. guereza and C. vellerosus.

Infant handling (holding or carrying) by adult males is rare in mammals; however, high levels have been reported in some primates. Though infant handling is a costly behaviour, there are many benefits that male handlers can accrue. Infant handling by males is most conspicuous in platyrrhines and tends to be uncommon in catarrhines. In the latter species, research on male-infant interactions has focused on low-cost behaviours, such as proximity and grooming. However, to better understand the evolution of infant handling by males, more data on its occurrence across the Primate order are essential, even in species where it is relatively uncommon. We compare the occurrence of infant handling by males in three closely related species of catarrhine: Colobus vellerosus, C. guereza, and C. angolensis ruwenzorii. We collected focal animal samples on infants to quantify infant handling rates and durations, and found that adult male C. a. ruwenzorii handled infants much more frequently and for much longer than males in the other two species. We discuss how C. a. ruwenzorii's unique social organization may explain high levels of infant handling by adult males in this species. More long-term and detailed comparisons of infant handling across species and populations will shed light on how sociality has shaped the evolution of this behaviour in the Primate order.

Diet and Activity Budget in Colobus angolensis ruwenzorii at Nabugabo, Uganda: Are They Energy Maximizers?

INTRODUCTION: Colobine monkeys are specialized folivores that use foregut fermentation to digest leaves. The slow process of fermentation forces them to spend a lot of time resting and to minimize their energy expenditure to subsist on a lower-quality diet. METHODS: We recorded the diet and activity budget of Colobus angolensis ruwenzorii, which form a three-tiered multi-level society, at Lake Nabugabo, Uganda, over 12 months using scan sampling on adults and subadults, to determine whether they utilize the energy minimization strategy typical of colobines. RESULTS: We found that the annual diet was primarily comprised of high-quality food resources (young leaves 65% and fruit 31%), and fruits were the only plant part the monkeysselected when available. Both the fruits and young leaves of some species were preferred food items in some months, and mature leaf consumption correlated negatively with preferred food availability. Mature leaves appear to be a fallback food for this population but are rarely relied upon (3%). The C. a. ruwenzoriiat Nabugabo spent less time resting (40%) and more time moving (25%) than is typical for other species of black-and-white colobus. DISCUSSION/CONCLUSION: The high-quality diet of this population appears to allow them to utilize an energy maximization strategy. Their reliance on food items that tend to be clumped in space and time likely explains the frequent fission-fusion behaviour that we observe between core units. Our findings demonstrate that the foraging strategies of colobines may be more flexible than was previously thought and illustrate how food availability and distribution can impact primate social organization.

Anogenital distance as a measure of male competitive ability in Rwenzori Angolan colobus.

Anogenital distance (AGD) is positively correlated to fetal androgen exposure and developmental masculinization in mammals. Independent of overall body size, AGD shows a strong positive correlation with male fertility and in rodents, AGD is a good indicator of male competitive ability and is associated with female choice. We hypothesized that AGD will also predict male competitive ability in non-human primates. To test this, we measured AGD noninvasively with a parallel laser in a wild population of Angolan colobus monkeys (Colobus angolensis ruwenzorii) in Uganda and correlated to it to their social structure. C. angolensis ruwenzorii form a multilevel society with both one-male/multifemale units (OMUs) and multimale/multifemale units (MMUs). We compared AGD in males from five OMUs and six MMUs and related it to their fecal androgen metabolite concentrations, dominance rank and body size, and to the number of females in their unit. Males in OMUs had greater access to females, so were predicted to have longer AGDs, but this was not found. AGD also did not correlate overall with mean fecal androgen metabolites in MMUs. However, AGD was correlated with dominance rank in MMUs, demonstrating that higher-ranking males in these multimale units had longer AGDs. Body size did not show the same relationship with dominance rank, suggesting that male rank was not just a reflection of absolute male size. Our findings indicate that AGD predicts male competitive ability in this species and that it may be a useful correlate throughout the non-human primates. These results also support the idea that prenatal androgen exposure increases the likelihood of the expression of behaviors that maintain high dominance rank.

Animal cognition in the field: performance of wild vervet monkeys (Chlorocebus pygerythrus) on a reversal learning task.

Increasingly, researchers are moving animal cognitive research into wild field settings. A field-based approach offers a valuable complement to laboratory-based studies, as it enables researchers to work with animals in their natural environments and indicates whether cognitive abilities found in captive subjects are generalizable to wild animals. It is thus important to field-based research to clarify which cognitive tasks can be replicated in wild settings, which species are suitable for testing in the wild, and whether replication produces similar results in wild animals. To address these issues, we modified a well-known lab test for field applications. The transfer index (TI) is a reversal learning task that tests whether animals rely on more associative or rule-based learning strategies (Rumbaugh in Primate behavior: developments in field and laboratory research. Academic Press, Inc., New York, pp. 2-66, 1970). In this paper, we detail changes needed to use a TI-like task in the field, here referred to as the Field Reversal Index (FRI). We tested a sample of nine wild vervet monkeys (Chlorocebus pygerythrus) on the FRI task at Lake Nabugabo, Uganda. We show that wild primates can successfully be tested on reversal learning paradigms, and present findings that reinforce previous conclusions from captive experiments. Our results indicate that vervets, like other cercopithecoids, rely on associative learning rather than rule-based learning. Further, our results are consistent with previous research that reports improved performance post-reversal in younger individuals relative to older individuals. The FRI enables researchers to test animals both in the wild and in captivity to facilitate direct comparisons between the learning abilities of captive and wild animals.

A multi-level society comprised of one-male and multi-male core units in an African colobine (Colobus angolensis ruwenzorii).

Several mammalian species exhibit complex, nested social organizations, termed multi-level or modular societies. Multi-level societies comprise stable core units that fission and fuse with one another in a hierarchical manner, forming groups that vary in size over time. Among nonhuman primates, these social systems have been confirmed in several African papionin and Asian colobine species. We use data from August 2017 to July 2018 on individually-recognized Rwenzori Angolan colobus living near Lake Nabugabo, Uganda to document the first multi-level society in an African colobine. The study band comprised up to 135 individuals organized into 12 socially and spatially distinct core units that ranged in size from 4 to 23 individuals. These core units showed a strong affinity to one another, spending roughly 75% of their time together. Core units fissioned and fused non-randomly with one another throughout the day, leading to different combinations of core units being observed. Using association indices between core units, we employed hierarchical cluster analyses and permutation tests to show that some core units associated preferentially into clans. Thus, we confirm three tiers of social organization for Rwenzori Angolan colobus: core unit, clan, and band. The social organization of this subspecies is unlike any reported previously in a nonhuman primate, with about half the core units containing a single adult male and the others containing multiple reproductive adult males. The discovery of a unique primate multi-level society in a novel lineage could allow for a better understanding of the evolution of these complex social systems across the Animal Kingdom. Preliminary data show males transfer within the band and females transfer outside of the band, which is proposed for hominin multi-level societies. This subspecies could thus also provide insight into the selective pressures underlying multi-level societies in our own lineage.

Playing it safe? Solitary vervet monkeys (Chlorocebus pygerythrus) choose high-quality foods more than those in competition.

An important goal in foraging ecology is to determine how biotic and abiotic variables impact the foraging decisions of wild animals and how they move throughout their multidimensional landscape. However, the interaction of food quality and feeding competition on foraging decisions is largely unknown. Here we examine the importance of food quality in a patch on the foraging decisions of wild vervet monkeys (Chlorocebus pygerythrus) at Lake Nabugabo, Uganda using a multidestination platform array. The overall nutritional composition of the vervet diet was assessed and found to be low in sodium and lipids, thus we conducted a series of experimental manipulations in which the array was varied in salt and oil content. Although vervets prioritized platforms containing key nutrients (i.e., sodium and lipids) overall, we found that solitary vervets prioritized nutrient-dense platforms more strongly than competing vervets. This finding was opposite to those in a similar experiment that manipulated food site quantity, suggesting that large, salient rewards may be worth competing over but slight differences in nutritional density may be only chosen when there are no potentially negative social consequences (i.e., aggression received). We also found that vervets chose platforms baited with oil-only, and oil combined with salt, but not salt-only, suggesting that energy was an important factor in food choice. Our findings demonstrate that when wild vervets detect differences in feeding patches that reflect nutritional composition, they factor these differences into their navigational and foraging decisions. In addition, our findings suggest that these nutritional differences may be considered alongside social variables, ultimately leading to the complex strategies we observed in this study.

Navigation strategies in three nocturnal lemur species: diet predicts heuristic use and degree of exploratory behavior.

Humans generally solve multi-destination routes with simple rules-of-thumb. Animals may do the same, but strong evidence is limited to a few species. We examined whether strepsirrhines, who diverged from haplorhines more than 58 mya, would demonstrate the use of three heuristics used by humans and supported in vervets, the nearest neighbor rule, the convex hull, and a cluster strategy, when solving a multi-destination route. We hypothesized that the evolution of these strategies may depend on a species' dietary specialization. Three nocturnal lemur species were tested on an experimental array at the Duke Lemur Center. Frugivorous fat-tailed dwarf lemurs (Cheirogaleus medius) were expected to follow paths most consistent with distance-saving navigational heuristics because fruit trees are stationary targets. Gray mouse lemurs (Microcebus murinus) and aye-ayes (Daubentonia madagascariensis), which rely on more mobile and ephemeral foods, were expected to use fewer paths consistent with these heuristics and be more exploratory. Our data supported all of these hypotheses. Dwarf lemurs used paths consistent with all three heuristics, took the shortest paths, and were the least exploratory. Mouse lemurs were quite exploratory but sometimes used paths consistent with heuristics. Aye-ayes showed no evidence of heuristic use and were the most exploratory. Distinguishable patterns of inter- and intra-individual variation in ability to solve the route, speed, and behavior occurred in each species. This research suggests that these simple navigational heuristics are not part of a readily available set of cognitive tools inherited by all primates but instead evolve due to need in each lineage.

Proximity and grooming patterns reveal opposite-sex bonding in Rwenzori Angolan colobus monkeys (Colobus angolensis ruwenzorii).

Close proximity and social grooming are important bonding mechanisms in primates. These behaviors show the social structure of a species and many studies have found positive correlations between the degree of kinship and grooming and proximity. We used 1 year of data collected via instantaneous scan sampling on a large "supertroop" of Colobus angolensis ruwenzorii at Lake Nabugabo, Uganda, to examine partner preferences for grooming and nearest neighbors in each age-sex class. Little is known about this species, so we based our hypotheses on congeners. Of the five species of black-and-white colobus, data on sex-biased dispersal patterns are available for three (C. guereza, C. vellerosus, and C. polykomos), all of which show male-biased dispersal with occasional female dispersal. We thus predicted that female C. a. ruwenzorii would be more strongly bonded than males, showing greater proximity and grooming. We did not expect bonding between the sexes since congeners do not show this pattern. We found that among adult dyads, males and females were more likely to be found in loose proximity, and to groom, than would be expected given group composition. Conversely, both males and females had relatively weak same-sex relationships. Between the sexes, adult males had higher proximity and grooming indices with adult females without infants than with females with infants. These observations indicate that this subspecies is cross-bonded and that both sexes may disperse. Furthermore, our findings suggest that the social organization and social structure of C. a. ruwenzorii differ greatly from other black-and-white colobus species.

Alpha male replacements in nonhuman primates: Variability in processes, outcomes, and terminology.

Alpha male replacements occur in all primates displaying a dominance hierarchy but the process can be extremely variable. Here, we review the primate literature to document differences in patterns of alpha male replacements, showing that group composition and dispersal patterns account for a large proportion of this variability. We also examine the consequences of alpha male replacements in terms of sexual selection theory, infanticide, and group compositions. Though alpha male replacements are often called takeovers in the literature, this term masks much of the variation that is present in these processes. We argue for more concise terminology and provide a list of terms that we suggest more accurately define these events. Finally, we introduce the papers in this special issue on alpha male replacements in the American Journal of Primatology and discuss areas where data are still lacking.

The influence of male takeovers on female dispersal in Colobus vellerosus.

Male takeovers affect male tenure, female mate choice and ultimately, individual reproductive success in group-living primates. In social systems with female philopatry and high male reproductive skew, male takeovers largely determine female mate choice, whereas in species with female dispersal, females have the option of deserting a new male. We focused on a species with facultative female dispersal to investigate which factors promote female desertion of males after takeover, using 15 cases (12 for which we have complete data on the takeover process and the female dispersal outcome). These cases took place in nine groups of Colobus vellerosus between 2001 and 2013 at the Boabeng-Fiema Monkey Sanctuary, Ghana. Quick takeovers were usually achieved by single adult males and were never followed by female dispersal. Slow takeovers involved several males, and these takeovers were regularly accompanied by female emigration. Infant attacks and infanticide by males occurred during both kinds of takeovers, but females with dependent offspring never dispersed, regardless of whether their infant was attacked or killed by the new male(s). Subadult females, who were not constrained by the presence of infants, dispersed more often after slow takeovers than after quick takeovers. Whether female dispersal post-takeover is an expression of female mate choice, or occurs to avoid the social upheaval surrounding slow takeovers, remains to be investigated. Am. J. Primatol. 79:e22436, 2017. © 2015 Wiley Periodicals, Inc.