Are fission–fusion dynamics consistent among populations? A large‐scale study with Cape buffalo

Fission–fusion dynamics allow animals to manage costs and benefits of group living by adjusting group size. The degree of intraspecific variation in fission–fusion dynamics across the geographical range is poorly known. During 2008–2016, 38 adult female Cape buffalo were equipped with GPS collars in three populations located in different protected areas (Gonarezhou National Park and Hwange National Park, Zimbabwe; Kruger National Park, South Africa) to investigate the patterns and environmental drivers of fission–fusion dynamics among populations. We estimated home range overlap and fission and fusion events between Cape buffalo dyads. We investigated the temporal dynamics of both events at daily and seasonal scales and examined the influence of habitat and distance to water on event location. Fission–fusion dynamics were generally consistent across populations: Fission and fusion periods lasted on average between less than one day and three days. However, we found seasonal differences in the underlying patterns of fission and fusion, which point out the likely influence of resource availability and distribution in time on group dynamics: During the wet season, Cape buffalo split and associated more frequently and were in the same or in a different subgroup for shorter periods. Cape buffalo subgroups were more likely to merge than to split in open areas located near water, but overall vegetation and distance to water were very poor predictors of where fission and fusion events occurred. This study is one of the first to quantify fission–fusion dynamics in a single species across several populations with a common methodology, thus robustly questioning the behavioral flexibility of fission–fusion dynamics among environments.     

Spatial and temporal habitat use of kob antelopes (Kobus kob kob, Erxleben 1777) in the Comoé National Park, Ivory Coast as revealed by radio tracking

Spatial and temporal habitat use of kob antelopes (Kobus kob kob) have been investigated in the Comoé National Park (Ivory Coast, West Africa) by use of radio telemetry. A total of 23 kob were equipped with radio collars and radio tracked for up to 15 months. Home ranges of males were smaller and those of females larger than expected from theoretical models. Adult males used smaller areas than adult females and did not show seasonal home range shifts. Daily distances travelled did not differ between sexes. Kob walked less during the night than by day and covered shorter distances in the wet season. Whereas an increase in home range overlap between females resulted in higher rates of association among individuals, association of adults of mixed sexes was not correlated with the degree of home range overlap. Territorial behaviour of males and predator avoidance by females are suggested to explain the sex‐specific differences in home range size of adults and the deviation from the predicted sizes. Predator avoidance is presumed as the main reason for the reduced walking distances at night as well as in the wet season. Reproductive behaviour and feeding ecology are assumed to determine the degree of association of conspecifics.     

Herd composition,kinship and fission–fusion social dynamics among wild giraffe

A variety of social systems have evolved as a consequence of competition and cooperation among individuals. Giraffe (Giraffa camelopardalis sp.) societies are an anomaly because the dearth of long‐term data has produced two polar perspectives: a loose amalgamation of non‐bonded individuals that sometimes coalesce into a herd and a structured social system with a fission–fusion process modifying herd composition within a community. We analysed 34 years of data collected from a population of Thornicroft's giraffe (G. c. thornicrofti, Lydekker 1911) residing in South Luangwa, Zambia, to establish the nature of giraffe society. Our sample consisted of 52 individually recognized animals. We found that giraffe herd composition is based upon long‐term social associations that often reflect kinship, with close relatives significantly more likely than non‐relatives to establish herds. Mother/offspring dyads had the strongest associations, which persisted for years. Giraffe live in a complex society characterized by marked flexibility in herd size, with about 25% of the variance in herd composition owing to kinship and sex. We suggest that giraffe herds share many characteristics of fission–fusion social systems and propose that sophisticated communication systems are a crucial component regulating subgroup dynamics.     

Male and female range use in a group of white‐bellied spider monkeys (Ateles belzebuth) in Yasuní National Park,Ecuador

Spider monkeys (Ateles sp.) live in a flexible fission–fusion social system in which members of a social group are not in constant association, but instead form smaller subgroups of varying size and composition. Patterns of range use in spider monkeys have been described as sex‐segregated, with males and females often ranging separately, females utilizing core areas that encompass only a fraction of the entire community range, and males using much larger portions of the community range that overlap considerably with the core areas of females and other males. Males are also reported to use the boundary areas of community home ranges more often than females. Spider monkeys thus seem to parallel the “male‐bonded” patterns of ranging and association found among some groups of chimpanzees. Over several years of research on one group of spider monkeys (Ateles belzebuth) in Yasuní National Park, Ecuador, we characterized the ranging patterns of adult males and females and evaluated the extent to which they conform to previously reported patterns. In contrast to ranging patterns seen at several other spider monkey sites, the ranges of our study females overlapped considerably, with little evidence of exclusive use of particular areas by individual monkeys. Average male and female home range size was comparable, and males and females were similar in their use of boundary areas. These ranging patterns are similar to those of “bisexually bonded” groups of chimpanzees in West Africa. We suggest that the less sex‐segregated ranging patterns seen in this particular group of spider monkeys may be owing to a history of human disturbance in the area and to lower genetic relatedness between males, highlighting the potential for flexibility some aspects of the spider monkeys' fission–fusion social system. Am. J. Primatol. 72:129–141, 2010. © 2009 Wiley‐Liss, Inc.     

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.     

Leadership of herd progressions in the Thornicroft's giraffe of Zambia

In cohesive social groups, travel progressions are often led by dominant or older individuals, but the leadership traits of individuals residing in flexible social systems are poorly known. Giraffe reside in herds characterized by fission–fusion dynamics frequently mediated by kinship. We analyzed 41 years (1971–2012) of longitudinal data collected from a community of Thornicroft's giraffe (Giraffa camelopardalis thornicrofti) living around South Luangwa National Park, Zambia, to assess the characteristics of herd leaders. Movement of giraffe in a single file progression was not associated with either season or time of day, but progressions were significantly more likely to occur when giraffe traveled in open areas. The oldest female in a herd was significantly more likely to be at the front position than expected, occupying the leadership niche on 79% of observations. We reason that matriarchal leadership in giraffe, as in African elephants, Loxodonta africana, is associated with resource learning. Giraffe societies are constructed on a heretofore unrecognized foundation that integrates relatedness and familiarity with matriarchal leadership in herd movement.     

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.     

The influence of male mating tactics on habitat use in mountain gorillas (Gorilla gorilla beringei)

Previous research has shown that both ecological and social factors influence mountain gorilla habitat use. New data on habitat use by a male gorilla and by a group confirm that male mating competition influences short- and long-term habitat use patterns, and show that its influence can supersede that of ecological factors on a long-term basis. When solitary, the male regularly approached and sometimes followed groups. His monthly home range size and equitability of home range use were directly proportional to the number of such interactions per month. His relationships with other groups became more conservative after he gained females, and, contrary to expectations based on metabolic needs, he used a much smaller area. The group considered here gradually expanded its home range and shifted its areas of intensive use throughout a three-year period. It then made a complete home range shift after three dramatic interactions, during which it was temporarily fragmented and two females emigrated. The group shared its home range with many other social units; overlap with most of these decreased after the shift. The degree of overlap and the lack of site fidelity by males and their groups support the argument that transfer is not ecologically costly to mountain gorilla females.     

Beyond habitat requirements: individual fine-scale site fidelity in a colony of the Galapagos sea lion (<Emphasis Type="Italic">Zalophus wollebaeki</Emphasis>) creates conditions for social structuring

Site fidelity has been widely discussed, but rarely been related explicitly to a species’ social context. This is surprising, as fine-scale site fidelity constitutes an important structural component in animal societies by setting limits to an individual’s social interaction space. The study of fine-scale site fidelity is complicated by the fact that it is inextricably linked to patterns of habitat use. We here document fine-scale site fidelity in the Galapagos sea lion (Zalophus wollebaeki) striving to disentangle these two aspects of spatial behaviour. Regardless of sex and age, all individuals used small, cohesive home ranges, which were stable in size across the reproductive and non-reproductive season. Home ranges showed a large individual component and did not primarily reflect age- or sex-specific habitat requirements. Site specificity could be illustrated up to a resolution of several metres. Long-term site fidelity was indicated by home range persistence over 3 years and the degree of site fidelity was unaffected by habitat, but showed seasonal differences: it was lower between reproductive and non-reproductive periods than between reproductive seasons. We further examined static and social interaction within mother–offspring pairs, which constitute a central social unit in most mammalian societies. Regardless of the occupied habitat type, adult females with offspring had smaller home range sizes than non-breeding females, demonstrating the importance of spatial predictability for mother–offspring pairs that recurrently have to reunite after females’ foraging sojourns. While social interaction with the mother dropped to naught in both sexes after weaning, analysis of static interaction suggested female-biased home range inheritance. Dispersal decisions were apparently not based on habitat quality, but determined by the offspring’s sex. We discuss the implication of observed fine-scale site fidelity patterns on habitat use, dispersal decisions and social structure in colonial breeding pinnipeds.     

Whole-day follows of striped mice (Rhabdomys pumilio), a diurnal murid rodent

Understanding mammal social systems and behaviour can best be achieved through observations of individuals in their natural habitat. This can often be achieved for large mammals, but indirect methods have usually been employed for small mammals. I performed observations of the striped mouse (Rhabdomys pumilio) during the breeding season in the succulent karoo, a desert of South Africa. The open habitat and the diurnal habit of striped mice, together with the use of radio-telemetry, made it possible to collect data on activity patterns and social interactions over an entire activity period (whole-day follow). The striped mouse in the succulent karoo has been reported to form groups of one breeding male, two to four breeding females, juvenile and adult offspring of both sexes, and several litters. Accordingly, daily range size did not differ between males and females, but females spent more time foraging whereas males spent more time patrolling territory boundaries. Captive R. pumilio display biparental care, and in this study both sexes visited the nesting site during the day, possibly engaging in parental care. Mice travelled more than 900 m/day, mainly during the morning and afternoon, and rested in bushes during the hottest times of the day.     

Seasonal and sex differences in the fission–fusion dynamics of spider monkeys (<Emphasis Type="Italic">Ateles geoffroyi yucatanensis</Emphasis>) in Belize

Spider monkeys (Ateles sp.) are characterized by high fission–fusion dynamics, meaning their social grouping pattern is fluid and consists of subgroups that vary in size, composition, and spatial cohesion over time. In this study, we quantify the fission–fusion dynamics of a group of spider monkeys at Runaway Creek Nature Reserve in Belize by measuring subgroup size, spatial cohesion, and stability using data spanning 5 years. We then test whether variation in these three subgroup measures differ according to season, subgroup sex composition, and the reproductive status of female subgroup members. Our results show that subgroups were larger in size and less stable in membership during the wet season compared to the dry season. All-female subgroups were less spatially cohesive but more stable in membership than all-male subgroups. Finally, we report that subgroups with one or more non-lactating females (i.e., without nursing young) were smaller on average than subgroups containing lactating females with nursing young. These data contribute to a growing body of research documenting the ecological and social dimensions along which grouping patterns might vary.     

Spatial structuring and dispersal in a high density population of the California vole Microtus californicus

Space use patterns of California voles are described from intensive live-trapping data obtained during the late breeding season in a peak density year. Operational sex ratios were strongly female biased. Breeding males had mutually exclusive home ranges that overlapped the home ranges of one to several females. The average male home range was larger than the average female home range and tended to encompass female home ranges in their entirety. Breeding females had home ranges that often overlapped extensively with the home ranges of other breeding females. The degree of overlap, however, tended to be either very high or very low, indicating that groups of females may act territorially among themselves. Family groups that may include members of sequential litters formed at high density. Males on experimental removal grids had larger home ranges than males on the high density grids and some became wanderers while most females had home ranges comparable to those on the high density grids.
Dispersers onto low density grids were not a random sample of the high density population; large breeding adults and small juveniles were under-represented. No sex differences in dispersal were noted. Immigration into the high density populations was probably negligible.     

Dispersal,philopatry, and the role of fission‐fusion dynamics in bottlenose dolphins

In this quantitative study of locational and social dispersal at the individual level, we show that bottlenose dolphins (Tursiops sp.) continued to use their natal home ranges well into adulthood. Despite substantial home range overlap, mother–offspring associations decreased after weaning, particularly for sons. These data provide strong evidence for bisexual locational philopatry and mother–son avoidance in bottlenose dolphins. While bisexual locational philopatry offers the benefits of familiar social networks and foraging habitats, the costs of philopatry may be mitigated by reduced mother–offspring association, in which the risk of mother–daughter resource competition and mother–son mating is reduced. Our study highlights the advantages of high fission–fusion dynamics and longitudinal studies, and emphasizes the need for clarity when describing dispersal in this and other species.     

Jaguar spacing, activity and habitat use in a seasonally flooded environment in Brazil

Between July 1980 and February 1984, six jaguars (two males, four females) were fitted with radio-collars and monitored for a cumulative total of 105 months, in the Pantanal region of southwestern Brazil. Mean home range size (minimum convex polygon) for five of them (one male, four females) was 142- 1 km². Mean home range size during the dry season was 54·3 km², whereas in the wet season it was significantly smaller, 12·8 km². Mean home range overlap of the four females was 42%. Use of gallery forest and forest patches exceeded the availability of these habitat types in the animals' home ranges, whereas open forest and grassland were used less than expected on the basis of their availability. Mean distance moved between locations on consecutive days was 2·4 km (0.2-10·4 km). The mean one-day movement of the male jaguar was significantly (P<0·001) larger than that of the females. Mean distance travelled by all animals during one-day intervals in the dry season was significantly greater (P<0·001) than that travelled in other months. Jaguars were more active during daytime than night-time (P<0·001). Overall level of activity for the wet season did not differ from that of the dry season. With their larger body size, jaguars in the Pantanal may require more food than jaguars in other areas of the species' range. This, in addition to differences in prey availability and the annual availability of dry land, appear to influence the difference in home range size between jaguar populations.     

Collective decision‐making and fission–fusion dynamics: a conceptual framework

Sociality exists in an extraordinary range of ecological settings. For individuals to accrue the benefits associated with social interactions, they are required to maintain a degree of spatial and temporal coordination in their activities, and make collective decisions. Such coordination and decision‐making has been the focus of much recent research. However, efforts largely have been directed toward understanding patterns of collective behaviour in relatively stable and cohesive groups. Less well understood is how fission–fusion dynamics mediate the process and outcome of collective decisions making. Here, we aim to apply established concepts and knowledge to highlight the implications of fission–fusion dynamics for collective decisions, presenting a conceptual framework based on the outcome of a small‐group discussion INCORE meeting (funded by the European Community's Sixth Framework Programme). First, we discuss how the degree of uncertainty in the environment shapes social flexibility and therefore the types of decisions individuals make in different social settings. Second, we propose that the quality of social relationships and the energetic needs of each individual influence fission decisions. Third, we explore how these factors affect the probability of individuals to fuse. Fourth, we discuss how group size and fission–fusion dynamics may affect communication processes between individuals at a local or global scale to reach a consensus or to fission. Finally, we offer a number of suggestions for future research, capturing emerging ideas and concepts on the interaction between collective decisions and fission–fusion dynamics.     

Mother–offspring distances reflect sex differences in fine‐scale genetic structure of eastern grey kangaroos

Natal dispersal affects life history and population biology and causes gene flow. In mammals, dispersal is usually male‐biased so that females tend to be philopatric and surrounded by matrilineal kin, which may lead to preferential associations among female kin. Here we combine genetic analyses and behavioral observations to investigate spatial genetic structure and sex‐biased dispersal patterns in a high‐density population of mammals showing fission–fusion group dynamics. We studied eastern grey kangaroos (Macropus giganteus) over 2 years at Wilsons Promontory National Park, Australia, and found weak fine‐scale genetic structure among adult females in both years but no structure among adult males. Immature male kangaroos moved away from their mothers at 18–25 months of age, while immature females remained near their mothers until older. A higher proportion of male (34%) than female (6%) subadults and young adults were observed to disperse, although median distances of detected dispersals were similar for both sexes. Adult females had overlapping ranges that were far wider than the maximum extent of spatial genetic structure found. Female kangaroos, although weakly philopatric, mostly encounter nonrelatives in fission–fusion groups at high density, and therefore kinship is unlikely to strongly affect sociality.     

Seasonal movements of Gyrfalcons Falco rusticolus include extensive periods at sea

Little information exists on the movements of Gyrfalcons Falco rusticolus outside the breeding season, particularly amongst High Arctic populations, with almost all current knowledge based on Low Arctic populations. This study is the first to provide data on summer and winter ranges and migration distances. We highlight a behaviour previously unknown in Gyrfalcons, in which birds winter on sea ice far from land. During 2000–2004, data were collected from 48 Gyrfalcons tagged with satellite transmitters in three parts of Greenland: Thule (northwest), Kangerlussuaq (central‐west) and Scoresbysund (central‐east). Breeding home‐range size for seven adult females varied from 140 to 1197 km² and was 489 and 503 km² for two adult males. Complete outward migrations from breeding to wintering areas were recorded for three individuals: an adult male which travelled 3137 km over a 38‐day period (83 km/day) from northern Ellesmere Island to southern Greenland, an adult female which travelled 4234 km from Thule to southern Greenland (via eastern Canada) over an 83‐day period (51 km/day), and an adult female which travelled 391 km from Kangerlussuaq to southern Greenland over a 13‐day period (30 km/day). Significant differences were found in winter home‐range size between Falcons tagged on the west coast (383–6657 km²) and east coast (26 810–63 647 km²). Several Falcons had no obvious winter home‐ranges and travelled continually during the non‐breeding period, at times spending up to 40 consecutive days at sea, presumably resting on icebergs and feeding on seabirds. During the winter, one juvenile female travelled over 4548 km over an approximately 200‐day period, spending over half that time over the ocean between Greenland and Iceland. These are some of the largest winter home‐ranges ever documented in raptors and provide the first documentation of the long‐term use of pelagic habitats by any falcon. In general, return migrations were faster than outward ones. This study highlights the importance of sea ice and fjord regions in southwest Greenland as winter habitat for Gyrfalcons, and provides the first detailed insights into the complex and highly variable movement patterns of the species.     

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.     

Social and demographic influences on the feeding ecology of giraffe in the Luangwa Valley,Zambia: 1973–2014

Body size influences metabolic rate, which impacts feeding ecology. Body mass differs by sex in size‐dimorphic species, such as giraffes, and also by age. Giraffes reside in a fission–fusion social system, which influences feeding ecology due to frequent changes in group size and composition. We analysed 40 years of feeding records collected from a population of Thornicroft's giraffes (Giraffa camelopardalis thornicrofti) living in the Luangwa Valley, Zambia. We examined the influence of herd composition, age and sex on diet. Solitary males and herd males did not differ in diet. Dietary diversity was comparable for females and males, with sex differences in plant species eaten present during the dry season. Age differences in feeding ecology were pronounced, with juveniles often feeding on bushes and smaller trees, while adults tended to feed upon taller trees. Both sexes have evolved foraging strategies that maximize nutrient and energy intake commensurate with their reproductive strategies, with male metabolic requirements sometimes greater, and sometimes less, than that of females. We propose that females are not exchanging foraging efficiency with reproductive tactics by feeding on smaller trees in the open, but are increasing prospects for their calves to survive when confronted with interspecific competition by browsers.     

Home range dynamics,habitat selection,and survival of Greater Roadrunners

ABSTRACT Greater Roadrunners (Geococcyx californianus) are common, poorly studied birds of arid and semi‐arid ecosystems in the southwestern United States. Conservation of this avian predator requires a detailed understanding of their movements and spatial requirements that is currently lacking. From 2006 to 2009, we quantified home‐range and core area sizes and overlap, habitat selection, and survival of roadrunners (N= 14 males and 20 females) in north‐central Texas using radio‐telemetry and fixed kernel estimators. Median home‐range and core‐area sizes were 90.4 ha and 19.2 ha for males and 80.1 ha and 16.7 ha for females, respectively. The size of home range and core areas did not differ significantly by either sex or season. Our home range estimates were twice as large (x?= 108.9 ha) as earlier published estimates based on visual observations (x?= 28–50 ha). Mean percent overlap was 38.4% for home ranges and 13.7% for core areas. Male roadrunners preferred mesquite woodland and mesquite savanna cover types, and avoided the grass‐forb cover type. Female roadrunners preferred mesquite savanna and riparian woodland cover types, and avoided grass‐forb habitat. Kaplan‐Meier annual survival probabilities for females (0.452 ± 0.118[SE]) were twice that estimated for males (0.210 ± 0.108), but this difference was not significant. Mortality rates of male roadrunners were higher than those of females during the spring when males call from elevated perches, court females, and chase competing males. Current land use practices that target woody‐shrub removal to enhance livestock forage production could be detrimental to roadrunner populations by reducing availability of mesquite woodland and mesquite savanna habitat required for nesting and roosting and increasing the amount of grass‐forb habitat that roadrunners avoid.