Promiscuous tendencies in the round-eared sengi: a male’s perspective

Although monogamy is interpreted as risk-adverse strategy by reducing intra-sexual conflicts, most pair-living males increase their reproductive success by engaging in extra-pair copulations. However, little is known about costs involved in such extra-pair attempts from the male’s perspective. We investigated promiscuous tendencies of paired Macroscelides proboscideus (round-eared elephant-shrew or round-eared sengi), a pair-living small mammal occurring in southern Africa. In particular, we measured potential costs of extra-pair attempts for the males. For this, we conducted laboratory experiments, involving interactions between neighbouring male–female pairs. Data collection included direct behavioural observations and establishing the morphological and physiological characteristics of males. Both sexes intruded into the neighbouring area, and initiated sexual behaviour with neighbours of the opposite sex. Males which displayed a higher marking frequency in the neighbouring area received more sexually motivated behaviour from neighbouring females. Resident males attacked intruding males. Aggression experienced by and marking behaviour of intruding males was positively correlated with body mass losses. Furthermore, glucocorticoid levels, determined from analyses of faeces and urine samples, correlated positively with body mass losses and marking behaviour of intruding males, indicating costs of intruding. We conclude that male as well as female round-eared sengis have promiscuous tendencies. Although interactions with neighbouring females may offer potential benefits for males in terms of enhanced reproductive success, males also seem to pay substantial costs, as indicated by body mass losses and elevated stress hormone levels.     

Spawning dynamics of orange roughy,Hoplostethus atlanticus,in mid-slope waters of New Zealand

Synopsis The social and reproductive biology of the sand tilefish,Malacanthus plumieri (Malacanthidae), was studied at Glover's Reef, Belize, where this species occurs in colonies over sand-rubble flats. Individuals each occupy a home burrow refuge and a surrounding home range. Home range overlap among adjacent fish of the same sex is low, and individuals defend exclusive use of much of their home range against all conspecifics except mates (i.e., territoriality). Areas defended by males overlap the territories of up to 6 females; and male territory area is positively related to the number of female residents. Males maintain dominance over females within their territories by aggression, including intervention into some female disputes. Females spawn pelagically-dispersed eggs as frequently as every day. Each female spawns near her burrow, almost exclusively with the male whose defended area encompasses her territory (harem polygyny). Tilefish colonies therefore consist of a mosaic of female territories over which adjacent male territories are superimposed. Histological evidence and observation of behavioral sex change in one female revealed thatM. plumieri is capable of protogynous sex reversal. Females did not change sex in response to removal of one male. Occurrence of small transitional fish indicates that the onset of sex change is controlled by factors other than size-related social hierarchies within harems or colonies.     

Home range and territory in the striped plateau lizard (Sceloporus virgatus)

Observations of home range sizes, home range locations, and behavioural interactions were used to determine the spatial distribution of a population of striped plateau lizards (Sceloporus virgatus). A comparison of home ranges and nearest-neighbour distances indicates that females have clearly defined territories. In contrast, the home ranges of males overlap extensively. Although males have similar nearest-neighbour distances to those of females, their home range sizes are larger. Much of the overlap among males appears to be associated with the courtship of females, suggesting that males have exchanged defence of exclusive territories for an increased home range area and a larger number of potential mates. Aggressive interactions among males were considerably more conspicuous than those among females, and the clear territorial pattern of females as opposed to males was unexpected on the basis of the frequency of behavioural interactions alone.     

Social organization and space use in California voles: seasonal,sexual, and age-specific strategies

We intensively monitored space use and movement in Microtus californicus over a 2-year period that included 1 year of high density (maximum 618/ha) and one of low (minimum 5/ha); historically this population has exhibited cycles of 2 or 4 years. Adults of both sexes dispersed at the start of the breeding season, culminating in the establishment of intrasexually exclusive territories. In females, these territories persisted throughout life, except that many young females recruiting during the breeding season established contiguous, overlapping, or adjacent home ranges with their mothers. This female philopatry explains the conclusion of previous workers that females of this species are non-territorial. In the dry (non-breeding) season, females had smaller ranges that often overlapped and were clustered. Adult males moved breeding territories at a modal interval of 6 weeks; this is consistent with their avoidance of inbreeding with philopatric daughters. Ranges overlapped 1–4 adult females at any one time, and a cohort of 7 long-lived males overlapped an average of 16.4 females during their tenure on the grid. The period of maximum overlap with adult females varied among individual males, and did not correlate with the time of maximum body weight. Ranges of males in the dry season overlapped extensively, with persistent associations among some individuals. In the lowdensity year, ranges of some adults failed to overlap intersexually. Juvenile males dispersed gradually between 3 and 13 weeks of age (half before 9 weeks), with some leaving after reaching sexual maturity; a few remained philopatric. Of juvenile females, 47% remained philopatric with the rest disappearing before 9 weeks of age. New understanding of vole social behavior, dispersal, and space use is achieved by focusing on the seasonal dynamics of spatial relationships among individuals with respect to age, sex, and relatedness.     

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.     

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.     

The social ecology of the Pyrenean desman (Galemys pyrenaicus) (Insectivora: Talpidae), as revealed by radiotelemetry

The social organization, spatial utilization and movement patterns of free-ranging desmans ( Galemys pyrenaicus G.) were studied by radiotracking individuals in the French Pyrenees. During the period of study, May to July, single adult male and female desmans formed a pair and defended their shared territory from neighbouring conspecifics by vigilance and scent-marking behaviour. Other adults did not form pair bonds and remained solitary. In such pairs, the territory of the adult male was always larger than, and completely enclosed, that of the paired female. Neighbouring pairs occupied contiguous territories of similar length. The territories of neighbouring males overlapped to a small extent, but the territory of one male and that of a neighbouring female were never seen to overlap. Juveniles were observed to utilize the territory of resident pairs, prior to their dispersal. Paired adults and juveniles exploited their ranges on a regular daily basis, whilst those of solitary adults, being larger than those of the former, were utilized on a 48–hour basis. Paired males appeared to invest more time in defending the borders of their shared territory than did females which, in contrast, spent most of their time within the centre of the territory. Despite some degree of territorial overlap between neighbouring conspecifics, their mutual avoidance, achieved through a system of temporal range utilization, resulted in few agonistic encounters.     

Exclusive core areas and intrasexual territoriality in Eurasian red squirrels (<Emphasis Type="Italic">Sciurus vulgaris</Emphasis>) revealed by incremental cluster polygon analysis

When animal home ranges overlap extensively in species lacking overt territorial behaviours, identifying exclusive core areas within individual ranges can be difficult. By analysing the size and overlap of successively smaller core areas among individual Eurasian red squirrels (Sciurus vulgaris), we determined exclusive areas within the home ranges of resident males and females. Possible effects of habitat composition and food supplies were explored by monitoring squirrels in different conifer forests and during years with low and high tree seed production. Using outlier-exclusive cores (OEC) revealed that the total ranges consisted of large sally zones (on average, 35% of the total minimum convex polygon [MCP] range) around home ranges with multi-nucleate cores. The mean OEC home range size did not differ between the sexes but was larger with poor food availability. Home ranges (99% incremental cluster polygons [ICP]) overlapped extensively between sexes (average overlap high food–low food: males by females 21–40%, females by males 43–45%) and among males (males by males 26–44%), while intrasexual overlap among females was low (9–10%). The overlap of inner cores among females rapidly approached zero, suggesting the intrasexual territoriality of 75% core areas. This was not the case among male squirrels, for which intrasexual overlap averaged only 4% at 50% but 18% at 75% core areas. Even the smallest inner cores had some degree of intersexual overlap, indicating that complete territoriality did not occur in this species. Female home ranges were more strongly affected by annual fluctuations in food supplies than male ranges. Females reduced the size of their food-based intrasexual territories when food availability increases. Males probably benefit from using larger home ranges and core areas, which overlap with the ranges of several females, by increasing their probability of successful mating.     

Habitat selection and home range size of field voles<Emphasis Type="Italic">Microtus agrestis</Emphasis> in Słowiński National Park,Poland

Habitat preference, home range size and intra-specific overlap were investigated in summer 1998 among field volesMicrotus agrestis (Linnaeus, 1761) in Słowiński National Park (N Poland). Eight individuals (2 females, 6 males) were radio-tracked for one week in August. Field voles were shown to exhibit a marked preference for meadow and the ecotone between grassland and habitats with common reed, while avoiding alder forest and proper reedbeds. No significant differences between night and day in habitat-use of voles were noted. The results suggest that, at the end of the breeding season, it was food resources, rather than the risk of predation, played an important role in the voles’ utilisation of space. The home ranges of males were larger and more diverse than those of females; their sizes being correlated with body mass, such that heavier males had larger home ranges. This further suggests that intra-sexual competition exists between males for females. The low number of females influenced their spatial behaviour, as females had completely exclusive home ranges. Four males (out of six) had overlapping home ranges with other males; three of the overlaps were of less than 20%. Attributes of promiscuity (such as a 3.5:1 operational sex ratio of males to females, intra-sexual competition between males and the territorial exclusivity of females) influenced the social system. However, the period of radio-tracking during this study was too short to define accurately the social system in the field vole population.     

The ecology of Hypogeomys antimena, an endemic Madagascan rodent

The subfamily Nesomyinae comprises 11 species of rodents endemic to the island of Madagascar. They show considerable ecological and anatomical diversity but little is known of their biology and they have not been studied to any extent in the wild. We present here the results of a nine-week field study of Hypogeomys antimena , the largest of the Nesomyinae, found only in a restricted area of dry deciduous forest in western Madagascar. Radiotracking data gave a mean home range estimate of 3.5+0.71 ha for 10 animals (five males, five females). Individuals from the same burrow had highly overlapping ranges but ranges of animals from neighbouring burrows showed minimal overlap. We suggest that H. antimena lives in social units, probably consisting of a pair plus their offspring, which occupy exclusive territories. A mark/recapture study gave a minimum population estimate of 48 animals/km² in the study site. The conservation of the species is discussed with reference to its limited distribution and current habitat degradation in the area.     

Social organization of Galagos in Kenyan coastal forests: I. Galago zanzibaricus

The social organization of Galago zanzibaricus was studied for the first time and the study included data from two different sites in the coastal forests of Kenya. A combination of mark-recapture and radio-tracking techniques was used to investigate patterns of inter- and intrasexual home range overlap. Associations in sleeping groups indicated social ties between individuals. Patterns of range use were established by radio-tracking focal individuals. Adult males generally had nonoverlapping ranges, shared with one or two females and their offspring with which the male regularly slept. Young females remained longer in their natal ranges than males and reproduced within their natal ranges. Range size and distance traveled per night were similar between the sexes; both sexes probably defended territories. None of the other galago (bushbaby) species studied to date show the degree of close male-female association found in G. zanzibaricus. This study thus extends the variety of social organization documented in nocturnal prosimians.     

Reversed Sex Change in The Haremic Protogynous Hawkfish Cirrhitichthys falco in Natural Conditions

Bi‐directional sex change has recently been reported in a range of reef fishes, including haremic species that were earlier thought to be protogynous (female to male). However, the occurrence of this phenomenon and the social conditions driving the reversion of males to females (reversed sex change) have been poorly documented under natural conditions. Reversed sex change is predicted to occur in low‐density populations where facultative monogamy is common. However, few studies have evaluated this over a long period in such populations. We documented the occurrence of bi‐directional sex change during a 3‐yr demographic survey of a population characterised by small harem sizes in haremic hawkfish Cirrhitichthys falco. New males were derived following a change in sex of functional females (secondary males; n = 3) and juveniles always matured first as females (n = 3). Thus, C. falco exhibited a typical protogynous sexual pattern, consistent with a range of haremic fish species. We observed reversed sex change in two males. In both cases, all the females disappeared from their harems and the neighbouring males expanded their territories to encompass the territories of the sex changers. However, bachelor males did not always revert to females. A dominant male experienced bachelor status twice but regained mating opportunities following the immigration of a female into his territory or by taking a female from a neighbouring harem. Thus, we conclude that bachelor males use reversed sex change as a facultative tactic to regain reproductive status in a haremic mating system. In addition, we discuss the influence of harem size upon occurrence of reversed sex change.     

Monogamy and sex role reversal in the pipefish Corythoichthys haematopterus

We observed the mating pattern and social behaviour of the pipefish Corythoichthys haematopterus in temperate waters of Japan during three successive breeding seasons. Males cared for a clutch in their brood pouch for 9-19 days until hatching and had several broods in the season with nonbrooding intervals of only 1 or 2 days. The population sex ratio was female biased and some females were always excluded from reproduction. Although males were sometimes courted by unmated females together with their regular partners, they always mated with the latter. The pair bond was maintained until the next season if both members survived. When males lost their partners, they remated with neighbouring unmated females within a few days. In contrast, widowed females remained unmated for a long time. Females had larger home ranges and were more active in courtship displays than males. This pipefish provides the first example of sex role reversal among monogamous syngnathid fish. We suggest that mate guarding by females is a primary proximate factor for maintenance of monogamy in this fish. Copyright 2001 The Association for the Study of Animal Behaviour.     

Space Use and Social Mating System of the Hantavirus Host, <Emphasis Type="Italic">Oligoryzomys longicaudatus</Emphasis>

The long-tailed mouse, Oligoryzomys longicaudatus (Cricetidae: Sigmodontinae), is the major host of Andes hantavirus, the etiological agent of hantavirus pulmonary syndrome in the south of Argentina and Chile. Studying the ecology of this species is necessary to understand how Andes hantavirus is maintained in nature. In this study, we examine the home range size and intra- and intersexual overlap degree of male and female O. longicaudatus in order to elucidate the mating system of this species. To our knowledge, this research provides the first documentation, obtained from a specific design, of spacing and mating systems in this species in Argentina. The study was conducted seasonally from April (autumn) 2012 to October (spring) 2013 in a shrubland habitat of Cholila, Andean region, Argentina. We studied spacing patterns using 59 and 51 home ranges established by adult males and females, respectively, in two 3.24 ha capture-marked and recapture grids. Significant differences between sexes in home range size and overlap degree were found. Male home ranges were always larger than those of females. We observed exclusive space use both among males and females (13.15 ± 18.67, and 3.60 ± 3.43%, respectively). Considering only those males that get access to receptive females (40%), average intersexual overlap value was about 30.82 ± 19.73%. Sexual differences in home range sizes and the spatial avoidance between breeding males, that would reflect intrasexual competition for receptive females, allows us to propose a polygynous mating system for O. longicaudatus.     

The Evolution of <Emphasis Type="Italic">Eulemur</Emphasis> Social Organization

Recent comparative studies on the evolution of mammalian sociality came to opposite conclusions regarding the direction and drivers of evolutionary transitions in social organization, particularly concerning the evolution of pair-living among primates. Because the genus Eulemur is one of the few exceptional primate genera that contain both group- and pair-living species, eulemurs offer a rare opportunity to illuminate the processes driving social diversity among species with very similar life histories and ecologies. The aims of our study were therefore 1) to reconstruct the evolution of social organization in eulemurs with new demographic and phylogenetic data and 2) to evaluate several hypotheses about the evolution of pair-living with data from our long-term study of Eulemur rufifrons and published data on other eulemurs. Our review of published data confirmed that E. mongoz and E. rubriventer are pair-living, whereas the 10 other species of Eulemur live in groups with multiple adult males and females. Furthermore, pair-living evolved at least once among eulemurs from group-living ancestors. The available comparative evidence indicated that this evolutionary transition was probably not driven by male infanticide, social instability, or singular ecological factors. Instead, we propose that female competition has favored small female group size and that this effect may have been exacerbated by interspecific competition with sympatric congeners in the ancestors of the pair-living species. In conclusion, we confirmed that the evolution of pair-living from group-living ancestors among true lemurs is exceptional, but not unique, among mammals, and that commonly invoked hypotheses for the evolution of pair-living do not appear to be supported for this genus.     

Endocrine correlates of contrasting male mating strategies in the European badger (Meles meles)

In mammals, testosterone secretion is known to respond rapidly to changes in males' immediate social environment. However, such changes do not take testosterone levels below the baseline required to maintain spermatogenesis, and so do not usually affect males' fertility. In this paper, we show that individual males' patterns of testicular activity reflect their social roles in a group-living carnivore, the European badger ( Melcs metes ), leading to individual and population differences in the seasonal patterns of both testosterone secretion and, apparently, spermatogenesis. In one badger population, some males left their natal groups to become breeding males in neighbouring territories. These males had higher plasma testosterone levels, and appeared to sustain spermatogenesis for a longer period, than males which remained in their natal territories. In contrast, in a second (higher density) population, males rarely transferred away from their natal territories, and appeared not to defend access to groups of females. Instead, males made forays to mate with females in neighbouring territories. In this population males had a shorter period of testicular activity, and there were no differences in testicular activity between immigrant and natal males.     

Spacing behaviour and its implications for the mating system of a precocial small mammal: an almost asocial cavyCavia magna ?

Socioecological models for small mammals attempt to explain the causal relations between the spatiotemporal distributions of food resources, females and males. We tested their predictions for a wild population of Cavia magna, a grazing, precocial rodent, by analysing spacing behaviour in relation to various demographic features. Between May 1999 and January 2001 we collected capture-recapture data on 309 individuals and monitored 55 females and 49 males by radiotelemetry in periodically inundated wetland in Uruguay. Cavies showed a nonstationary use of space: monthly home ranges drifted over the whole study site. Female home ranges overlapped with those of several others. Females were randomly distributed and we found no evidence for socially mediated reproductive synchrony. Males ranged over larger areas than females, showing even less site fidelity, and also overlapped with several rivals. This basic spacing system remained stable over a wide range of densities and sex ratios. Independent of sex, animals used overlap zones randomly with respect to each other. Significant dynamic (spatiotemporal) interaction was most frequent between males and females. However, interaction analyses revealed no evidence for stable social bonds between animals, regardless of sex. We suggest that unpredictable female locations prevent males monopolizing females spatially. Because females are solitary, males could monopolize only one female by maintaining close proximity, rendering a roaming mating tactic more successful. Our findings point to a solitary ‘social’ system and overlap promiscuity as the likely mating system for the C. magna population studied. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Social organization in the Chinese water deer, <Emphasis Type="Italic">Hydropotes inermis</Emphasis>

The social organization of Chinese water deer was studied in a zoological park. Most adults lived together in the mixed zone, although other habitats were available. The overlap between individual areas was largest in females. In the mating season alone, less than half of the males established territories which overlapped in small spots where most encounters occurred. Females travelled freely throughout the entire available area, but non-territorial males stayed between the territories. Except in males during mating season, very few physical contacts other than sniffing took place between individuals regardless of age, sex or status. Each animal lived alone and did not show attraction or aggressiveness towards congeners. Grouping was temporary with no durable link between individuals, not even between mothers and daughters older than 5 months. The solitary life of individuals on a common ground and the establishment of seasonal territories make the water deer unique among ruminants. The species appears to be no more social than the water chevrotain, an “ancient” species. This fits well with other characteristics of the species.     

Polygamous mating system and protogynous sex change in the gobiid fish <Emphasis Type="Italic">Fusigobius neophytus</Emphasis>

Whereas mating behaviors and social structure have been studied extensively in monogamous hermaphroditic gobiid species, such studies are relatively limited for polygamous gobiid species. To investigate the reproductive strategy of polygamous gobies, mating groups of the common fusegoby Fusigobius neophytus were observed on reefs of Kuchierabu-jima Island, southern Japan. Males established mating nests on flat-rock surfaces within their territorial home ranges on sandy rubble flats. Females maintained independent home ranges outside the male home ranges during nonreproductive periods, but they shifted their home ranges to overlap with male ranges and actively visited male mating nests during their reproductive periods (1–3 days at ca. 7-day intervals). Females often changed mates during their serial mating. The mating system used by the common fusegoby fits with the definition of male-territory-visiting polygamy. The sex ratio within the study population was female-biased. Nest-holding males were significantly larger than females and were polygynous (mating with up to eight females). These characteristics fit well with the prediction of protogyny by the size-advantage model. Some of the females were observed to undergo functional sex changes to nest-holding males. In addition, small floating males demonstrated sneaking behavior. None of the floating males were derived from females that had changed sex, suggesting a diandric life-history pathway for F. neophytus.     

Socio-spatial organization of urban stone martens

Stone martens Martes foina are common occupants of urban environments across continental Europe. We predicted that like other 'urban adapters', urban stone martens should exhibit smaller home ranges, more extensive home-range overlap and higher population densities than their rural counterparts. We radio-tracked 13 stone martens (10 females, three males) in two towns in southern Luxembourg during a 2-year period, in order to investigate nocturnal ranging behaviour. All martens maintained stable, year-round territories almost wholly within the urban perimeter of the respective study towns. Socio-spatial organization was based on intra-sexual territoriality. Territories (100% minimum convex polygon) were small in size (males: 112.6±24.8 ha; females: 37.0±22.2 ha) and population densities moderately high (4.7–5.8 adults km⁻²) by comparison with previous studies of rural populations. We conclude that although stone martens are clearly well adapted to urban conditions, urbanization has surprisingly little impact on their socio-spatial organization by comparison with its effects on other mesocarnivores. This apparent inflexibility in the spacing patterns of stone martens lends support to the view that spacing in martens ( Martes spp.) is a predominantly phylogenetic trait that is under limited environmental influence.