Survey and census of howler monkeys (Alouatta palliata) in the rain forest of “Los Tuxtlas,” Veracruz,Mexico

Howler monkey troops were censused at the biological reserve “Los Tuxtlas” in Veracruz, Mexico. The reserve includes 700 ha of rain forest. Twenty howler monkeys were also trapped, measured, marked, and released. Censuses were conducted for a period of 26 months, and they indicated the existence of 17 troops. The mean troop size was 9.12 (SD ± 2.93), and mean troop composition was 3.0 adult males, 4.12 females, 1.56 juveniles, and 1.54 infants. Ecological density was 0.23 howlers/ha or 23.29 howlers/km². The male to female ratio was 1:1.37. No discrete seasonality in births was noted. Howler monkeys in this locality inhabit the northernmost limit of the neotropical rain forest. The population parameters fall within those reported for Alouatta palliata at other sites.     

Population of the black howler monkey (Alouatta pigra) in a fragmented landscape in Palenque,Chiapas, Mexico

Little is known about the population characteristics of Alouatta pigra under conditions of forest fragmentation-information that is important to understanding its tolerance to habitat loss. In this work we present data on forest loss and on troop size, age, and sex composition for a population of black howler monkeys existing in the fragmented landscape surrounding the Mayan site of Palenque, Chiapas, Mexico. Two aerial photos (1:70,000) of the study area (261 km(2)) taken in 1984 and 2001 were examined to assess forest loss. Between June and December 2001 and January and March 2002 we surveyed 44 forest fragments for the presence of howler monkeys. Examination of aerial photos showed that 33% of the forest present in 1984 had disappeared by 2001, and detected an increment in the number of forest fragments present in the landscape. We discovered a total of 115 howler monkeys living in 22 of the 44 forest fragments studied, of which 107 were members of 18 troops. The rest were solitary males or small groups of males living in isolated forest fragments. Troop size ranged from two to 15 individuals (mean 5.9+3.0 ind). 31% and 15% of individuals in the troops were juveniles and infants, respectively, suggesting continued reproductive activity. Howler monkey troops in the forest fragments were on average smaller (5.9+/-3.0 ind) than troops in the nearby protected forest of the Mayan site (7.0+/-2.8 ind). The mean density of howlers in the forest fragments was 119+/-82.9 ind/km(2). The establishment of corridors is suggested as a possible conservation scenario for the fragmented howler population investigated, and as a conservation measure to connect this population with the howler population found in the protected forest of the Mayan site.     

Survey of black howler (<Emphasis Type="Italic">Alouatta pigra</Emphasis>) and spider (<Emphasis Type="Italic">Ateles geoffroyi</Emphasis>) monkeys in the Mayan sites of Calakmul and Yaxchilán,Mexico and Tikal,Guatemala

Surveys of populations of spider and howler monkeys were conducted at the Mayan sites of Calakmul and Yaxchilán, Mexico and Tikal, Guatemala. The forests in which these sites are found are part of the largest landmass of tropical rain forests present in Mesoamerica, encompassing about 4 million ha. Triangulation of monkey vocalization combined with ground surveys was used to determine the presence of howler and spider monkey groups. Howler monkey mean troop size at these sites varied from 6.6±2.1 individuals in Yaxchilán to 7.5±1.9 in Calakmul to 8.7±2.2 in Tikal. Density estimates varied from 12.8 individuals/km² in Yaxchilán to 15.2 individuals/km² in Calakmul to 17.8 individuals/km² in Tikal. Mean spider monkey subgroup size varied from 4.7±2.6 individuals in Tikal to 5.6±3.0 individuals in Yaxchilán to 7.7±3.8 individuals in Calakmul. Spider monkey density varied from 17.0 individuals/km² in Yaxchilán to 17.2 individuals/km² in Calakmul to 56.4 individuals/km² in Tikal. All sightings of both howler and spider monkeys at the three sites were in undisturbed rain forest vegetation and spider monkeys in general were more frequently sighted at higher tree heights than howlers. We discuss the value of further acquiring data on howler and spider monkey populations existing in extensive forest tracts and on the conservation value for both primate species of the forests surrounding the Mayan ruins found in this area of Mesoamerica.     

Survey of Helminth Parasites in Populations of <Emphasis Type="Italic">Alouatta palliata mexicana</Emphasis> and <Emphasis Type="Italic">A. pigra</Emphasis> in Continuous and in Fragmented Habitat in Southern Mexico

The 2 howler species that occur in southern Mexico, Alouatta palliata mexicana and Alouatta pigra are endangered, mainly as a result of habitat loss and fragmentation from human activity. Little is known about the gastrointestinal parasite communities affecting their populations, and lack of baseline information for populations of howler species in continuous forest habitats, makes evaluations of gastrointestinal parasite prevalence in populations in fragmented landscapes difficult. We report the results of a one-time broad survey of gastrointestinal parasites in fecal samples of individuals from several demographically stable populations of Alouatta palliata mexicana and A. pigra existing in continuous and/or protected forests. We further report similar data for populations of both species in human-fragmented landscapes. We detected 6 parasites for each howler monkey species, but only 3 of them (Trematode I, Controrchis biliophilus, Trypanoxyuris sp.) were common to both species. While parasitic prevalence in populations of both howler species was, in general, higher in the fragmented habitat than in continuous and/or protected forests. The difference is only marginally significant in Alouatta pigra. Some parasites (Coccidia and Strongylid) only appeared in populations in fragmented landscapes. Preliminary data suggest that adult males tended to have higher parasite prevalence values than those of adult females in both howler species. Parasite prevalence is associated to average group size, but not to population density in Alouatta pigra.     

Influence of the landscape matrix on the abundance of arboreal primates in fragmented landscapes

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.     

Ecology and behavior of the wild proboscis monkey,Nasalis larvatus (Wurmb), in Sabah,Malaysia

Proboscis monkeys,Nasalis larvatus (Wurmb), were investigated in the Padas Bay and Kinabatangan River areas, Sabah, from July 1968 to March 1969. The proboscis monkey is a common, widely distributed species of coloboids in Sabah, living in the various vegetations that extend from the coastal swamp to the inland plain. It appears to have a terrestrial tendency, compared with other coloboids, moving around on the ground in the forest and extending its activity, sometimes, to the open land along the river beach. The number of monkey in a troop, we observed, ranged from 11 to 32, and almost all troops were composed of several adult males, many more adult females, and immature animals. The adult males appear to play the role of leaders in the troop through their characteristic behavior, attacking, threatening, warning, watching, etc., other troop members or invaders. Each troop always emitted various boisterous vocalizations which were divided into 14 types.     

Current status of the habitat and population of the black howler monkey (Alouatta pigra) in Balancán, Tabasco, Mexico

We evaluated the habitat and populations of the black howler monkey (Alouatta pigra) in the municipality of Balancán, Tabasco, southeastern Mexico, using a combination of field surveys and remotely sensed data. We identified 21,937 ha of remnant vegetation composed of 1,348 fragments. Fragments separated by up to 200 m were grouped into "clusters" of fragments in accordance with the maximum observed open distance crossed by A. pigra. A total of 11% or 84 of the 772 clusters identified through remote sensing were selected at random, and for these we determined the vegetation type, canopy height, area, and distance to the closest human settlement. In these same 84 clusters, which included a total area of 9,817 ha, from October to June of 2006 we located a total of 1,064 black howler monkeys, including 228 troops and 49 solitary monkeys. A. pigra was found in 62 (74.7%) of all clusters visited, with a cumulative area of 6,032 ha. Troops varied in size from 2 to 15 individuals (average 6.0+/-2.9 ind/troop). Adults were 67% (n=716) of detected individuals, whereas juveniles were 20.5% (n=218) and infants were 12.5% (n=133). We found black howlers to occur at an ecological density of 10.8 ind/km(2), which is low in comparison with A. pigra in other fragmented and conserved sites. We found a statistically significant relationship between the area of clusters and the abundance of howler monkeys (r(2)=0.2, F=10.47, gl=3, P=0.002). In addition, the probability of finding A. pigra was greater in secondary vegetation, riparian vegetation, tropical dry forest, undisturbed tropical oak forest, and palm forest (F=12, gl=3, P<0.0001), as compared with disturbed tropical oak forest. Our results provide data on the distribution, abundance, and population structure of black howler monkeys in a fragmented landscape in the southeast of Mexico. These data are a necessary prerequisite for conservation planning for this species.     

Habitat selection and seasonal patterns of activity and foraging of mantled howling monkeys(Alouatta Palliata) in Northeastern Costa Rica

I conducted a 15- month ecological study of habitat preferences and activity and foraging patterns of two troops of mantled howling monkeys, Alouatta palliata,in a lowland rain forest at La Selva Biological Reserve in northeastern Costa Rica. The two troops specialized on different habitats in spite of the fact that both of them had all habitats available and were not constrained by neighboring troops since the population density of howlers is low (7- 15 howlers/km ² ).Troop 1 spent the majority of time in primary forest (80%) followed by secondary forest (10%), while troop 2 spent the majority of time in undisturbed riparian habitat (60%) followed by primary forest (30%). Habitat sampling indicates that neither the total number of stems, species, or families nor the diversity (Shannon index) or evenness is a good indicator for howler habitat selection. Instead the density of trees from the 12 species most commonly consumed by each troop is the most important factor. Activity and foraging patterns were not dependent upon the season as has been described for howling monkeys in forests with a more pronounced dry season at Barro Colorado Island, Panama, and La Pacifica in northwestern Costa Rica. This is likely a result of the more constant food supply at La Selva, combined with less intraspecific competition due to the low howler density. The intraspecific variability of foraging patterns and troop- specific habitat specialization observed in Alouatta palliatashould be considered in the conservation biology of primates. Primate relocation programs should include not only an ecological assessment of the release site but also a comparison of the release site with the habitat that the groups currently occupy.     

Breeding behaviors in the black howler monkey (Alouatta pigra) of belize

Adulterous breeding between a female howling monkey (Alouatta pigra) of one troop with a male of an adjacent troop occurred despite territorial defense between the two troops. The specific behaviors are described as well as a synopsis of daily events which occurred between the female and two males from adjacent troops. A discussion of how this interaction and the behavior patterns relate to what is known about breeding in howler monkeys and related species follows.     

Ecology and behavior ofPresbytis thomasi in Northern Sumatra

A field study of 23 bisexual troops, ranging in size from 3 to 21 members ( =8), and two all-male groups of the Thomas's leaf monkey (Presbytis thomasi) was conducted in North Sumatra from November 1981 to April 1984. Most troops (N=19 or 82.6%) contained only one adult male. Two troops and one group were most intensively studied. The home range was 12.3–15.7 ha for the two bisexual troops, and only 1.7 ha for the one, all-male, ten-member group. Fruits composed more than 50% of their diets. Vocalizations were classified into 13 types. Births occurred at any time of the year. Among three males of a bisexual troop, serious fights were observed: two males died of wounds and the former beta male became the new alpha male. After this social change, the home range area of this troop gradually shifted eastward. But, 3.5 months after the social change, a 9-month-old infant male stayed alone in the western part of its former range. Thereafter, he became a solitary male and sometimes went into the riverine area of the Bohorok river. The occurrence of male replacement suggests instability of multi-male organization in bisexual troops. Moreover, the different mortality rate between males and females and the unequal sex ratios forced by the formation of one-male troops, maintained high tension levels among males competing on females.     

Survey of Alouatta caraya, the black-and-gold howler monkey, and Alouatta guariba clamitans, the brown howler monkey, in a contact zone, State of Rio Grande do Sul, Brazil: evidence for hybridization

Sympatry and natural hybridization between howler monkey taxa (Alouatta spp.) has only recently being confirmed in the wild. Surveys in areas of potential contact between the distribution of two taxa have shown that sympatry is rare, although more common than previously known. Here we report the results of a survey conducted in a contact zone between the only two sexually dichromatic howler monkey taxa, Alouatta caraya and A. guariba clamitans, in São Francisco de Assis, State of Rio Grande do Sul, Brazil. Our survey, covering an area of about 400 ha at the Cerro dos Negros (29°33′50″–29°35′10″S, 54°58′40″–54°59′50″W; ~100–279 m a.s.l.), was successful in locating seven black-and-gold and one brown howler monkey social groups living syntopically. Black-and-gold group size ranged from 5 to 15 individuals, whereas the brown group was composed of 7 individuals. The pelage color of three adult males belonging to different black-and-gold groups and another adult male belonging to the brown howler group presented a mosaic of red or rufous and black. These adult males and an adult female living in another black-and-gold group are putative hybrids. Therefore, it appears that pre-zygotic reproductive isolation has not evolved, at least not completely, between these howler monkey species, corroborating previous reports for these and other Alouatta taxa. Future genetic studies need to confirm the occurrence of hybridization in this contact zone, and to determine the viability and fertility of hybrids and their possible offspring. In addition, there is no evidence supporting the existence of significant segregation in habitat and resource utilization by black-and-gold and brown howler monkeys.     

Male life history in natural populations of Japanese macaques: Migration, dominance rank, and troop participation of males in two habitats

This paper compares male life history parameters of two populations of Japanese macaques (Macaca fuscata Blyth, 1875), studied without provisioning: Yakushima (M. f. yakui), a subtropical forest habitat in southwestern Japan, and Kinkazan (M. f. fuscata), a temperate, deciduous forest habitat in northeastern Japan. The males of the two sites experienced similar life histories with respect to several traits. Age at natal dispersal was at about 5 years. Average troop residence was about three years. Most males joined troops at the bottom of the rank order, although a few males joined troops at the top rank. Dominance ranks of males tended to rise with the death or departure of higher ranking males. Visiting males accounted for about 41% of observed mating at both sites. However, the two sites differed in the sex ratio of troops, partly because a larger proportion of males apparently lived outside of troops in the Kinkazan site compared to Yakushima. In particular, non-natal young males were absent from the main study troop at Kinkazan. Large within-species variation may exist in the degree to which males associate with troops.     

Status of Trachypithecus geei in a Rubber Plantation in Western Assam, India

Golden langurs (Trachypithecus geei) are an endangered species in a limited area of Northeast India and Bhutan. They are confined to a forest belt in western Assam between the Manas River in the east, Sankosh in the west and Brahmaputra in the south along the Indo-Bhutan border. Due to habitat destruction, their populations are restricted to fragmented forest pockets, especially in India. We compared data on an isolated population of golden langurs in the rubber plantation of Nayakgaon in the Kokrajhar district of Assam, May 2002 with an earlier census in 1997. We counted a total of 52 individuals in 3 troops in 2002. The average troop size is 17.3 (r = 7–26). The ratio of adult male to adult females is 1:3.16. Of the total population 9.6% were adult males, 21.1% were adult females, 17.3% subadult males, 7.7% subadult females, 19.2% juveniles and 25% infants. In 1997 Srivastava et. al., recorded 38 individuals in 5 troops with an average troop size of only 7.6 . Apparently troop fusion has occurred. The increased population is encouraging.     

Intermale relations and troop male membership changes in langurs (Presbytis entellus) in Nepal

Data on intermale social relations and troop membership changes in one Nepalese high-altitude population of free-ranging langurs (Presbytis entellus)are reported here. Data were collected from six troops by three observers and cover 32 months of observations. The predominantly multi-male troops indicate an alternating pattern of exclusions and introductions with gradual adult male replacement. Takeovers and infant killing were not observed. Analysis of adult social behavior records show qualitative and quantitative differences in intrasexual relations, with primarily agonistic social contacts occurring between males. Agonistic encounters between females and between males differ in frequency of occurrence, types of be-haviors used, cause, and consistency in direction of threats between individuals. Individual adult male frequency of interaction with females and immatures varied significantly, with the majority of these interactions occurring between the dominant troop male and other troop members. Data indicate that intermale dominance is a major factor in determining male access to fertile females: This appears to be achieved by either directly excluding males from the troop or effectively “controlling” their inter-actions with troop females. Data from these studies are compared with data from other Presbytis entellusinvestigations. Review of these data suggests that intraspecific variability in intermale social dynamics and type of troop male membership change are correlated with the percentage of nontroop males. It is suggested that environmental pressures resulting in social crowding can be critical in determing the occurrence of takeovers in some populations of Presbytis entellus.     

Demography of Lemur catta at Berenty Reserve,Madagascar: Effects of Troop Size,Habitat and Rainfall

We censused Lemur catta within a 1 km² study area at Berenty Reserve, Madagascar, during the September–October birth season for 19 years between 1963 and 2000, a total of 290 troop counts (266 with age and sex). The non-infant population was 155 in 1972–5, fell to 105 in 1985, and rose to a maximum of 282 in 1997, while troops increased from 12 in 1972–1985 up to 25 in 1998–2000. Local density varies between habitat types from 1 per ha to ca. 6 per ha. Troops fission at ca. 15–25 individuals, or 6–10 females. Adult sex ratio has no apparent correlation with fissions, birth rate or survival. Birth rate falls steeply with number of adult females, from 80–100% in 2-female troops to about 50% in 8–10 female troops. The penalty for large troop size is greater in the dense, rich areas, but nonetheless troops there are also larger. One-year-survival does not vary with troop size, and is lower in the sparse, dry zone. Troop size is too large for optimal birth rate, but fissioning to much lower size might make troops too small for optimal adult survival, given the intense intertroop competition. This reflects Sibley's (1983) conjecture that troop sizes may not reach stable optima. Rainfall per lemur-year (beginning Oct 1) varied from 265 to 894 mm. Drought followed by rain can eliminate >90% of a cohort, especially in the dryest zone. Possibly this results from fruit failure in years following drought. It is unknown whether food supplementation of some Berenty troops is dangerous for the forest, or helpful for an isolated and vulnerable ring-tailed lemur population.     

The variable social organization of hanuman langurs(Presbytis entellus), infanticide,and the monopolization of females

Data from 24 wild populations of hanuman langurs (Presbytis entellus)in south Asia are used to test hypotheses seeking to explain variation in troop structure and the incidence of infanticide. The occurrence of infanticide is associated with a one-male troop structure and not with a high density. The density, predation, and economic-advantage hypotheses, as explanations for the occurrence of one-male and multimale troops, are not supported by the review. However, the monopolization hypothesis is not contradicted; the number of adult males per troop is significantly correlated with troop size and with the number of adult females per troop. Therefore it is suggested that a one-male troop structure will arise if a male is able to monopolize a group of females, a multimale troop if he cannot. One-male troops may predispose to infanticide because of high variance in male mating success and high intermale competition between groups rather than within troops. If female dispersion determines troop structure, it is speculated that females could manipulate males to form a multimale society if the advantages in terms of infant survival and intertroop conflict exceeded the costs in terms of not producing infanticidal “sexy sons.”     

Male Relationships and Infant Deaths in Red Howler Monkeys (Alouatta seniculus)

For decades howler monkeys were considered to be among the most peaceful animals. Recent investigators, however, have noted that, although male aggression is rare, it may be extremely severe; the criteria used to quantify howler relationships should therefore be different from those used for other primates. This paper describes the pattern of male relationships in the red howler monkey Alouatta seniculus, and examines its effect on reproductive success within the troop and the ability of troop males to resist intrusion by competing males. 4 of the 8 infants born during the study period were killed or disappeared when change occurred in relative status between two adult males.     

Chasing behavior between males within a howler monkey troop

On Barro Colorado Island, Panama, a mature male howler monkey (Alouatta palliata) was observed chasing a young adult male howler, both members of the same troop. Detailed observations on births, deaths and solitary males before and after the chase suggest a strong correlation between an unstable social structure within the troop and overt aggression between different-aged males. Support for this field work was received from the Environmental Sciences Program, Smithsonian Institution; and the National Zoological Park, Washington, D.C.     

Social interactions among adult male langurs (Presbytis entellus) at Rajaji Wildlife Sanctuary

A population of langurs (Presbytis entellus)at the Rajaji Wildlife Sanctuary in northern India was investigated for 1820 hr throughout a 10-month period in 1978. Data were collected from four bisexual troops and the adult males that ranged outside of bisexual troops. Most (60%) of the observation hours occurred with a main study troop from which social and ecological data were collected. The langur population at Rajaji shows pronounced birth and mating seasons. The population density is high (ca. 80/km ²), with about 75% of the adult males living outside of bisexual troops, which typically are large and multimale. Males outside of bisexual troops occur in small all-male bands or as isolates. Relations between bisexual troops and all-male bands are characterized by relatively low levels of aggression, and members of all-male bands are able to associate with bisexual troops for prolonged periods during the mating season. As a result of these associations, nontroop males are about as successful as troop males in achieving reproductive access to troop females. These associations between bisexual troops and all-male bands occurred with a minimal amount of agonistic behavior and without mortality or injury to troop females or immatures.     

Adult male replacement and social change in two troops of hanuman langurs (Presbytis entellus) at Jodhpur,India

I studied the process of adult male replacement and social change in two one- male troops (B20 and B21) of hanuman langurs (Presbytis entellus)at Jodhpur, India. Male-male competition lasted for about 6 months before the successful takeover of one troop (B20). During that period, five adult males from three neighboring bands (AMB7, AMB9, and AMB10) and a resident male of a neighboring troop (B21) were involved in taking over the troop. The latter male also copulated with six females during his interim residency, which suggests that he may have opportunistically maximized his mating chances with females of a neighboring group. During an intertroop interaction, a 14-month-old female infant of the other troop (B21) was fatally attacked by an adult female of the first troop and the infant eventually died. The attacker may have taken advantage of the disorganization created by male-male competition, perhaps to eliminate a future food competitor. In addition, the first troop gained an additional feeding area from the other troop’s range; it included a sleeping site and a waterhole, indicating that territorial fights during social instability may have led to the expansion of the winner’s resource area.