Long-Term Habitat Use by Mountain Gorillas (Gorilla gorilla beringei). 1. Consistency, Variation, and Home Range Size and Stability

Mountain gorillas are highly folivorous. Food is abundant and perennially available in much of their habitat. Still, limited research has shown that single gorilla groups heavily used areas where food biomass and quality were relatively high and where they met daily nutritional needs with relatively low foraging effort. Also, ecological factors influenced solitary males less than groups with females. Long-term data on habitat use by multiple mountain gorilla social units and more extensive data on variation in food distribution, presented here, confirm that food distribution influences areal occupation densities across groups and over time. These data also confirm the group/solitary male distinction and show that food distribution became more important for one male once he acquired females. Groups used 25 km ² , and inter-annual home range and core area overlap was often low. Annual home range and core area size varied considerably within groups and across years. It bore no simple relationship to group size and estimated group biomass. Core areas were biased samples of total home ranges and were relatively good foraging areas. One group abruptly shifted its home range in response to male mating competition. Home ranges of two others expanded from 1981 to 1987, though at a decreasing rate. Data on one such group, which varied considerably in size, are consistent with arguments that costs of scramble competition are low except in unusually large groups. Low site fidelity, low scramble costs, and high home range overlap should decrease the ecological costs of female transfer.     

Strategies of habitat use by mountain gorillas

Patterns of home range use by a mountain gorilla group are examined here in relation to variation in food abundance and quality, diet quality and rainfall, to interactions between different social units and to the distribution of hazards (poaching risk). Variation in habitat quality influenced both long-term area occupation densities and shorter-term movement parameters (day journey length and the distance moved between consecutive feeding sites). Observed home range use patterns increased the gorillas' foraging efficiency. Rainfall influenced the evenness of home range use slightly, but this was the only seasonal variability. Interactions with other gorillas and poaching risk had proximate effects superimposed on the fundamental influence of food distribution patterns. Data from a larger sample of groups show a positive influence of group size on day journey length, but this was apparent only at very small and relatively large group sizes. The absolute distance travelled per day tends to be short even for large groups. The data support arguments that the costs of social foraging are low for mountain gorillas.     

Activity rhythm and the ranging of a solitary male mountain gorilla (Gorilla gorilla beringei)

To clarify the advantages of solitary life in gorilla males, a lone silverback mountain gorilla (Gorilla gorilla beringei) was studied for nine months in the natural habitat of the Virunga volcanoes. While the time budget for each activity and daily activity cycle were similar to those of groups, his daily journey distance and ranging patterns differed from those of groups. His movements were little influenced by the distribution and abundance of foods, which strongly influence the movements of groups. He notably increased his day journey distances when he encountered neighboring groups. He persistently followed the groups for days and went out of his usual range area. These encounters shifted his monthly range from his natal group's range to that of other groups. When the silverbacks of the encountered groups noticed his presence, they usually gave hoots and chest-beats and sometimes fought violently with him, while females and immatures did not show positive responses towards him. Lone males could have more chance to contact females and to lure them away from their groups than silverbacks within groups. The lone male stage, accompanied by frequent contacts with different groups, probably provides maturing males with useful knowledge of neighboring groups and areas.     

Space use patterns of mountain hare (<Emphasis Type="Italic">Lepus timidus</Emphasis>) on the Alps

Populations on the limits of species’ distribution can show different behavioral adaptations to strong ecological pressure than in the central part of the range. We investigated space use patterns of alpine mountain hare (Lepus timidus) at two areas on the southern edge of the species’ range. We monitored 34 hares between 2005 and 2008, estimating home range size, overlap, and site fidelity, and compared our results with space use in Scottish and North-European populations. Home ranges of mountain hares did not differ between two study areas with different habitat types. Subadult animals used larger ranges than adults and both age groups reduced home range size in autumn, a period that might be critical for hares due to changes in diet and/or high energy expenditure during the previous breeding season. Home ranges in these alpine populations were smaller than in Scandinavian populations but within the range of populations in different habitat types in Scotland. Seasonal home ranges overlapped considerably, but differed among the sexes: male–female overlap was higher than same sex (male–male and female–female) spatial overlap. Seasonal shifts of home ranges were small, and site fidelity remained high over the seasons, suggesting that resource distribution remained constant throughout the year and that the knowledge of an intensively frequented area is an important element of habitat quality. We concluded that habitat structure and availability of mates interact in affecting mountain hare space use in alpine habitats.     

Long-term Site Fidelity and Individual Home Range Shifts in <Emphasis Type="Italic">Lophocebus albigena</Emphasis>

We investigated long-term site fidelity of gray-cheeked mangabey (Lophocebus albigena) groups in Kibale National Park, Uganda. Concurrently, we monitored shifts in home range by individual females and subadult and adult males. We documented home range stability by calculating the area of overlap in successive years, and by recording the drift of each group’s monthly centroid from its initial location. Home ranges remained stable for 3 of our 4 groups (overlap over 10 yr >60%). Core areas were more labile, but group centroids drifted an average of only 530 m over the entire decade. Deviations from site fidelity were associated with dispersal or group fission. During natal dispersal, subadult males expanded their home ranges over many months, settling ≤4 home ranges away. Adult males, in contrast, typically dispersed within a few days to an adjacent group in an area of home range overlap. Adult males made solitary forays, but nearly always into areas used by their current group or by a group to which they had previously belonged. After secondary dispersal, they expanded their ranging in the company of their new group, apparently without prior solitary exploration of the new area. Some females also participated in home range shifts. Females shifted home ranges only within social groups, in association with temporary or permanent group splits. Our observations raise the possibility that male mangabeys use a finder-joiner mechanism when moving into new home ranges during secondary dispersal. Similarly, females might learn new resource locations from male immigrants before or during group fission.     

Spatial and temporal interactions of free-ranging pacas (<Emphasis Type="Italic">Cuniculus paca</Emphasis>)

Although the paca is the most frequently hunted game species throughout the Neotropics, its behavioral and ecological requirements remain poorly understood. Here, we describe ranging behavior, spatio-temporal intraspecific interactions, and cavity use within a mosaic landscape in Central Belize, based on radio-tracking of four males and two females. This study is the first to investigate social interactions and spatial structure within a paca population in unprotected marginal habitat. We detected extensive home range overlap between males and between sexes. Male-male overlap was less extensive within core areas, while female core ranges were almost entirely occupied by one or more males. On average, pacas used at least six cavities within their home range. The majority of cavities were in the core areas and we did not detect simultaneous co-habitation of the same location. On average, females occupied cavities that were closer together, and closer to water bodies, than those of males. Overall, our study suggests a general tolerance during nocturnal foraging activities, but exclusive use of core areas and associated cavities. The larger ranges of males than females and the extensive overlap between conspecifics suggest a polygamous or promiscuous mating system in this landscape.     

Intra-specific variation in social organization of gorillas: implications for their social evolution

We analysed intra-specific variation in the social organization of gorillas and ecological and social factors influencing them, based on recent data on diet, day journey length, home range size, group size and proportion of multi-male groups in three subspecies [western lowland gorillas (WLG); eastern lowland gorillas (ELG); mountain gorillas (MG)]. Median group size was similar across subspecies and across habitats, but the extraordinarily large group including >30 gorillas was only found in habitat with dense terrestrial herbaceous vegetation. Within-group competition may determine the upper limit of group size in frugivorous WLGs and ELGs in lowland habitats with scarce undergrowth. A frugivorous diet may be a causal factor of subgrouping in multi-male groups of WLGs and ELGs, while a folivorous diet may prevent subgrouping in multi-male groups of MGs. Social factors, rather than ecological factors, may play an important role in the formation of multi-male groups and their cohesiveness in MGs. High gregariousness of female gorillas and their prolonged association with a protector male are explained by their vulnerability to both infanticide (MGs) and predators (ELGs). Comparison of long-term changes in group composition and individual movements between ELGs in Kahuzi and MGs in the Virungas suggest that the occurrence of infanticide may promote kin-male association within a group. Threat of infanticide may stimulate MG females to transfer into multi-male groups to seek reliable protection and maturing MG males to stay in their natal groups after maturity. By contrast, the absence of infanticide may facilitate ELG females to associate with infants and other females at transfer and ELG males to establish large groups in a short period by taking females from their natal groups, by luring females from neighbouring groups, or by takeover of a widow group after the death of its leading male. These conditions may prevent ELG and WLG maturing males from remaining to reproduce in their natal groups and possibly result in a rare occurrence of multi-male groups in their habitats. Similar reproductive features of MG and ELG females suggest both female strategies have been adaptive in their evolutionary history.     

Spatial organization of leopards Panthera pardus in Taï National Park, Ivory Coast: is rainforest habitat a 'tropical haven'?

Between June 1992 and July 1994, two female leopards and one male were radio-tracked. Regular locations of the leopards, the use of a phototrap, and spoor data, provided the first detailed ecological data about this elusive felid in tropical rainforest habitat. The home range of the male was 86 km², those of the two females were 29 km² and 22 km², respectively. One female's home range was fully included within that of the male. Home ranges of neighbouring residents were not exclusive. Population density is estimated at one leopard per 9-14 km². Intraspecific interactions were rare and predominantly involved mating. The large size of the home ranges and a relatively high population density imply large overlap between adjacent resident leopards' ranges. Differences in the leopard's land tenure system between the rainforest and the savanna are discussed. Doubt is cast on the validity of the often-quoted estimate of one leopard per 1 km² in tropical rainforest habitat.     

Relations between group size and composition and feeding competition in mountain gorilla groups

The intensity and patterning of interference competition for food within mountain gorilla (Pan gorilla beringei) social groups is influenced in several ways by group size and composition and varies within and across age/sex classes. Data on 251 spatial displacemtnts associated with feeding, collected during a 17-month study of mountain gorilla feeding ecology, show that overall displacement rates and displacement rates for individuals were positively correlated with social group size. Silverback males were responsible for a disproportionately high number of displacements. Adult females also were involved in competitive interactions over food more often than expected from their representation in groups, and had feeding bouts interrupted disproportionately often, principally by other females and by silverbacks. Competitive relationships between females varied in association with female dominance rank and age, but were not clearly associated with relatedness between females. The results support the argument that social foraging entails a cost which is proportional to group size and which falls particularly on adult females. The comparatively low rates of competitive interactions, however, suggest that this cost is relatively low, and that female mountain gorillas sacrifice little in terms of feeding efficiency by living in social groups.     

Movement patterns and herd dynamics among South African giraffes (Giraffa camelopardalis giraffa)

Giraffes reside in a fission–fusion social system, with sex, age proximity, kinship and home range overlap accounting for some of the variance in herd composition, while season, sex, age and time of day influence diet, home range size and distance travelled. To increase our knowledge of habitat use and fission–fusion herd dynamics, we placed GPS devices on eight adult female South African giraffes (Giraffa camelopardalis giraffa) living in the Khamab Kalahari Nature Reserve (South Africa). We tested four predictions about how season, kinship, home range and travel patterns influence habitat use and herd dynamics. Our two key findings were that females with a greater degree of home range overlap were more likely to form herds, but the degree of overlap was independent of the amount of time that they spent together in a herd, and that on the day prior to herd formation, females travelled about twice as far as their daily average and tended to move directly towards their future herd mate. We conclude that habitat use and movement patterns regulating fission–fusion dynamics reflect an interaction of ecological, social and reproductive factors operating in tandem, not independently.     

Social group dynamics and composition of managed wild vicuñas (<Emphasis Type="Italic">Vicugna vicugna vicugna</Emphasis>) in Jujuy,Argentina

The vicuña is a high Andean wild camelid that lives year-long in groups. We analyzed the social organization of 98 marked vicuñas after capture and live shearing, focusing on group composition, spatial patterns, territoriality and habitat use. The social units analyzed were families, bachelor groups and solo animals. Location fixes of 54 males and 36 females were registered over a 2-year period, and home range was estimated. All females belonged to family groups, and 8–12 month yearlings changed their home ranges prior to first reproduction at 2 years. Female yearlings dispersed to a new family group, while male yearlings dispersed to a bachelor group. Solitary males appeared during the interval between leaving bachelor or family groups, and vice versa. Our study showed that vicuña polygyny was based on female grazing area requirements. Females that stayed with the same males were defined as loyal, and had a mean 18.9 ha home range. Females that changed both to a different family and male; and grazed over the same home range size, were considered disloyal. Family groups selected the habitat that had a low stratum and the highest coverage of palatable plants, while bachelors used their habitat randomly.     

Habituation of western gorillas: the process and factors that influence it

Habituation of western gorillas to human presence is generally an expensive, lengthy and difficult process. Here we describe the habituation process for two groups of western gorillas at the Mondika Research Center, with the hope that the lessons we learned will facilitate future gorilla studies. We expand upon earlier studies by describing the process through complete habituation for both males and females, and for more than one group. The major obstacle to habituation was developing sufficient tracking skills to follow gorilla trail on a daily basis. Once this was achieved, the silverback became semi-habituated (i.e. ignoring human presence during half of contacts) within a year, although the majority of group females continued to avoid humans. As female presence at contacts increased, a period of male recidivism followed, requiring an additional year before his complete habituation was reached. Habituating the females took longer than the male, but we found, contrary to earlier studies, that it consisted of the same stages, including avoidance, aggression, and curiosity before habituation. We compare results between groups and across sites and discuss how factors such as tracking abilities, group size and cohesion, population density and home range overlap, and the manner of approaching gorillas during contacts influence the habituation process.     

Seasonality in the Ecology and Life Histories of Mountain Gorillas (Gorilla gorilla beringei)

The abundance of food, especially that of fruit and often that of young leaves, varies considerably over time for most primates. This variation can depend on or be independent of seasonality in rainfall. Mountain gorillas (Gorilla gorilla beringei) in the Virungas are exceptional: their habitat contains almost no edible fruit, and they mostly eat perennially available herbs and vines that are densely and evenly distributed in much of their habitat. Earlier studies documented little consistent temporal variation in mountain gorilla diets and habitat use, except for seasonal use of bamboo by some groups, and documented no birth seasonality. Long-term data ( 7 years) on 6 mountain gorilla groups confirm these results for habitat use, except that they show some seasonality in use of the upper altitudinal extremes of the gorillas' home ranges for unclear reasons. Relatively low and inconsistent variation in habitat quality over time should lower the costs of grouping for gorillas compared to other apes. Long-term data also confirm the absence of seasonality in births and conceptions. However, they show that mortality rates and risk of respiratory infections vary directly with rainfall. These relationships are probably causal and may be mediated through thermoregulatory stress.     

Infanticide and social flexibility in the genus <Emphasis Type="Italic">Gorilla</Emphasis>

Based on the cases of infanticide by male mountain gorillas reported from the Virunga volcanic region, the socioecological and life history features of gorillas satisfy the conditions for which infanticide may be expected. However, there are considerable variations in the occurrence of infanticide between habitats. We analyze the recent reports of infanticides that were directly observed or are suspected based on field evidence in two populations of eastern and western lowland gorillas (Kahuzi and Mbeli Bai, respectively) along with previous reports on mountain gorillas, and consider which social features are linked with and which factors influence the occurrence of infanticide in the gorilla populations. All victims were suckling infants and most of them were killed by males who seemed unrelated to them. Dependent infants are most vulnerable to infanticide when the protector male (its putative father in most cases) is absent, and so male protection ability seems to be important in determining female transfer decisions. Two cases observed in Kahuzi suggest that the infanticidal male may discriminate between infants to accept and those to kill according to his previous interactions with their mothers. Mating for a prolonged period prior to parturition is necessary for immigrant females to avoid infanticide by the new male of the group that they join. Infanticide was usually associated with female transfer, and the patterns of female association at transfer may shape variations in social structure between populations. Female mountain gorillas prefer large groups with multiple males and tend to transfer alone in order to seek more protection against infanticide in Virunga. By contrast, female eastern and western lowland gorillas tend to transfer with other females to small groups or solitary males, and maturing silverbacks take females to establish new groups through group fission in Kahuzi and Mbeli Bai. These differences may result in more multi-male and larger groups in the Virungas than in Kahuzi and Mbeli Bai. Rapid changes in density of gorilla social units and their relations following drastic environmental changes caused by recent human disturbances may also increase the probability of infanticide.     

Dispersed male networks in western gorillas

Although kin-selection theory has been widely used to explain the tendency of individuals to bias beneficial behaviors towards relatives living within the same social group, less attention has focused on kin-biased interactions between groups. For animal societies in which females emigrate, as is the case for mountain gorillas (Gorilla beringei beringei), encounters between males in different groups often involve aggressive displays that can escalate to physical violence and fatal injuries. However, recent findings on the little-studied western gorilla (Gorilla gorilla) indicate that interactions between social groups occur more frequently than they do in mountain gorillas and are often, although not always, surprisingly nonaggressive. We investigated the pattern of genetic relationships between individuals of different groups and found evidence suggesting a previously unrecognized "dispersed male network" social structure in western gorillas in which the single males leading social groups were usually related to one or more nearby males. We propose that this provides a basis for extra-group, kin-biased behaviors and may explain the reported peaceful intergroup interactions. Furthermore, these results suggest that a patrilocal social structure, in which males remain in their natal region and potentially benefit from kin associations, is a feature unifying African apes and humans.     

Sex difference in the communicatory significance of localized defecation sites in Arabian gazelles (Gazella arabica)

Mammalian scent marking in localized defecation sites (latrines) has often been interpreted in the context of (male) territory defense. However, latrines could have different functions in males and females, especially where territorial males monopolize groups of females with stable social alliances and pronounced home range overlap. We investigated the communicatory significance of latrines in wild Arabian gazelles (Gazella arabica) and assessed the spatial distribution of latrines within home ranges. Latrine density and utilization was highest in the center of female group home ranges, and less frequent in peripheral home range sections, pointing towards communication within groups rather than towards territoriality. When considering male home ranges, latrine densities and utilization were higher in non-overlap zones, contradicting a territorial function. This pattern appears to be caused by more females than territorial males per given area establishing latrines. A subsequent survey of latrine utilization, based on camera trapping, suggests that males use latrines for territory defense: males visited latrines in overlap zones disproportionally more often than females, and successions of two males prevailed. Our study thus highlights that male territorial marking can be masked when males and females use the same marking system for different purposes.     

Woodland fragmentation affects space use of Eurasian red squirrels

When habitats become fragmented, variation in patch size and quality are expected to impose changes on the spacing pattern and social organization of animal populations. General theory predicts different possible responses including shrinking home ranges (fission response), increasing range overlap (fusion) and incorporation of multiple patches in the home range (expansion response) as fragmentation increases. We studied space use and social organization in a metapopulation of red squirrels (Sciurus vulgaris) in 15 woodland fragments differing in size and tree species composition. Home ranges and core areas of males were larger than females, and fragmentation had different and complex effects on the spacing pattern of both sexes. In food-supplemented patches, high densities led to increased intra-sexual overlap. In linear-shaped patches, squirrels used smaller home ranges and core areas and had lower male–male and male–female overlap levels, independent of patch quality or size. Home range and core area size of males increased with patch size, and male core areas overlapped extensively those of other males and females. Hence males seemed to show a fission response only in some patches. In contrast, home range and core area size of females was not related with patch size, but decreased with habitat quality, supporting predictions of a fusion response and intra-sexual defense of food-based core areas. Hence, where patch size and shape strongly affected space use of male red squirrels, social organization of females was only affected in small, food-supplemented patches, suggesting that the basic spatio-social organization of adult females is very resistant to fragmentation.     

Observations on the home range of one group of mountain gorillas (Gorilla gorilla beringel)

Factors influencing the home range of a typical group (ten to seventeen individuals) of mountain gorilla (Gorilla beringei) over a 60-month period are discussed. Three other gorilla groups lived nearby and shared the group's home range. Vegetation zones were defined in terms relevant to the gorillas' needs, and the home range was divided into equal grid squares, each of one sixteenth of a square kilometre. The observer's contacts with the group within each square provide data on the relative utilization of different areas. The physical aspects of the six vegetation zones are discussed, and five types of travel within the various zones are defined. The current expansion of the home range is described. The presence or absence of other gorilla groups and/or individuals appeared to be the strongest determinant of the range patterns of the study group, though food distribution may have played some role in movements between vegetation zones or parts of the range.     

Staying close to home: Ecological constraints on space use and range fidelity in a mountain ungulate

Understanding patterns of animal space use and range fidelity has important implications for species and habitat conservation. For species that live in highly seasonal environments, such as mountain goats (Oreamnos americanus), spatial use patterns are expected to vary in relation to seasonal changes in environmental conditions and sex‐ or age‐specific selection pressures. To address hypotheses about sex, age, and seasonality influence on space‐use ecology, we collected GPS location data from 263 radio‐collared mountain goats (males, n = 140; females, n = 123) in coastal Alaska during 2005–2016. Location data were analyzed to derive seasonal and sex‐specific fixed‐kernel home range estimates and to quantify the degree of seasonal range and utilization distribution overlap. Overall, we determined that home range size was smallest during winter, expanded coincident with the onset of green‐up and parturition, and was largest during summer. Home range size of males and females did not differ significantly during winter, but females had larger home ranges than males during summer, a relationship that was switched during the mating season. Pairwise comparisons involving individual females across subsequent years indicated home ranges were significantly smaller during years when they gave birth to offspring. Mountain goats exhibited a strong degree of range fidelity, and 99% (n = 138) of individual animals returned to their previous year''s seasonal range with an average annual Bhattacharyya''s affinity utilization distribution overlap index of 68%. Similarity of seasonal home range utilization distributions varied in relation to sex and season in some respects. Home range overlap was highest during the summer vegetation growing season, particularly among females. These findings advance our understanding about how environmental variation and sex‐ and age‐related reproductive constraints influence space use and range fidelity among alpine ungulates. Documentation of the high degree of range fidelity among mountain goats has important conservation implications in landscapes increasingly altered by anthropogenic activities.     

Social relationships of immigrant and resident female mountain gorillas. I. Male-female relationships

Adult males are important social partners for all females in mountain gorilla social groups, but male-female relationships can vary in association with variation in female residence status, male age and mating status, and relatedness. Such variation occurred in a large group observed over a 3-year period. All females associated and interacted affiliatively relatively often with a young silverback with whom all mated. Long-term resident females also did so with an old, non-breeding male to whom most were related, but recent immigrants spent little time near him and had few non-aggressive interactions with him. The old male made agonistic interventions to support relatives; interventions in female conflicts by the younger male tended to ameliorate competitive differentials that immigrants faced and may have helped him to retain them as mates. Males aggression toward females was common, most probably served as a mating tactic, and did not involve resource competition. Extensive grooming by an adolescent male suggests that males may also use affiliative behavior to develop mating relationships. Females may have competed for proximity with and social access to the younger silverback; competitive success could influence how well females and their offspring are protected by silver-backs, but the extent of such competition was not clear. Large group size may have heightened differentiation of male-female relationships and competition among females, but comparative data from smaller groups generally corroborate the findings from the large group. © 1992 Wiley-Liss, Inc.