Olfactory communication and counter-marking in brown brocket deer <Emphasis Type="Italic">Mazama gouazoubira</Emphasis>

Scent marking by defecation and urination in numerous small latrines may be related to resource defence in brown brocket deer (Mazama gouazoubira). Both males and females seem to be territorial, and both contribute to latrines where their ranges overlap. Latrines could thus potentially function as centres of information exchange and intrasexual competition. Counter-marking occurs when animals respond to invaders' marks with a greater number of marks. The objectives of this experimental study were to determine whether brown brocket deer distinguish dung of presumed invaders from their own and whether they counter-mark such faecal deposits. Two samples of dung (from unknown males or females and the experimental animal) were introduced near the latrines of 21 captive deer (13 males and eight females), and we observed their responses, including investigative (sniffing) and marking (urination and defecation) behaviours. Males investigated the introduced dung and their own latrine significantly more when the dung was from an unknown male than when it was their own. Females investigated unknown female dung significantly more than their own. Males counter-marked the introduced dung and their latrine significantly more when the dung was from an unknown male than when it was their own. Males marked with a shorter latency and at a greater frequency than females. Our data indicated that males counter-mark most intensively dung from male ‘intruders’ which may be related to intrasexual competition and resource defence. Females showed a non-significant tendency to counter-mark same sex intruders.     

Exclusive core areas in overlapping ranges of the sleepy lizard, Tiliqua rugosa

Where animal home ranges overlap extensively, objectively identifyingexclusive areas within individual ranges has been difficult,particularly in species lacking overt territorial behaviors.By analyzing the overlap between successively smaller core areasamong individuals in a population of the long-lived Australianskink, the sleepy lizard (Tiliqua rugosa), we objectively determinedexclusive areas within animal ranges. Using 4-year radio trackingdata, we found that ranges consisted of relatively large sallyzones (mean 66–80% total range), around home ranges withmultinucleate cores strongly associated with key refuge sites.Total range and home range area varied significantly among years,being smaller in a drought year. Total ranges overlapped extensivelybetween and within sexes, but for both sexes, intrasexual overlapof inner range cores rapidly approached zero, suggesting intrasexualterritoriality. Intersexual inner core overlap reflected thisspecies socially monogamous mating system. But, male overlapof female ranges and inner cores was consistently higher thanfemale-male overlap. Refuges and/or mates may be defended resourceswithin these core areas, although aggressive behavioral interactionswere rarely observed. In the extensively overlapping sally zones,males shared space with females other than their principal partner.In productive years, with larger home ranges and more extensiveoverlap, some lizards associated with extra partners, suggestingthat males opportunistically use sally zones for polygyny. Consequently,we suggest that benefits to females from male association maychange with environmental quality, such that if food resourcesare low, monogamy may be favored if females increase foragingefficiency as a consequence of male vigilance during pairing.     

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.     

Scent marking in a territorial African antelope: I. The maintenance of borders between male oribi

Scent marking is ubiquitous among the dwarf antelope and gazelles of Africa, but its function has been the subject of debate. This study examined preorbital gland scent marking in the oribi, Ourebia ourebi, a territorial African antelope. Several hypotheses for the function of scent marking by territorial antelope were tested with observational data. Of these, the hypotheses that scent marking is driven by intrasexual competition between neighbouring males, and that marks serve as an honest advertisement of a male's ability to defend his territory from rivals, were supported best. Thirty-three territorial male oribi on 23 territories marked most at borders shared with other territorial males, and territorial males marked more often at borders shared with multimale groups than at borders shared with a single male. This suggests that males perceived neighbouring male groups as a greater threat to territory ownership than neighbouring males that defended their territories without the aid of adult subordinates. Marking rate was unrelated to territory size or the number of females on adjacent territories, but males with many male neighbours marked at higher rates than those with fewer male neighbours. These results suggest that the presence of male neighbours has a greater effect on the scent marking behaviour of territorial antelope than has been considered previously. Copyright 1999 The Association for the Study of Animal Behaviour.     

Scent-marking behaviour of the honey badger, Mellivora capensis (Mustelidae), in the southern Kalahari

We investigated sexual and seasonal patterns in scent-marking behaviour of the honey badger, by direct observations of habituated individuals (five females, four adult males, two young males). Four categories of scent-marking behaviour were identified: (1) scent marking at latrines; (2) token urination in holes along the foraging path; (3) squat marking at single-use sites; and (4) functional excretion. Females and young males used all four types of scent marking, but adult males were not observed to use token urination. A strategy of hinterland scent marking was used, as was predicted from the large home ranges of both male and female honey badgers. There were significant sexual differences in marking rate: adult males primarily used latrines and adult females favoured token urination. Latrine scent marking in adult male honey badgers provides support for the ‘scent-matching’ hypothesis. Females visited latrines when they were in oestrus. However, the low level of marking activity during a visit and the intensive smelling suggested a scent-matching function rather than reproductive advertisement. Token urination appeared to be related to the maintenance of spatiotemporal separation in females, although we also observed token urination in young males. While the placement of urine in foraging holes and its relation with successful digging attempts offer some support for the foraging efficiency hypothesis, we consider this unlikely, because we did not observe it in adult males and there was no seasonal pattern. Squat marking occurred under a wide range of conditions in both males and females and may be related to marking valuable resources. It is likely that scent marking in honey badgers has many functions.     

Seasonal Changes of Vocal Rates and Their Relation to Territorial Status in Male Galápagos Sea Lions (Zalophus wollebaeki)

Male vocal displays play an important role in sexual selection through both male–male competition and female choice. While this is supported by numerous studies in birds, much less attention has been paid to the role of such displays in mammals. We investigated the function of vocal displays in a polygynous mammal, the Galápagos sea lion (Zalophus wollebaeki). In our study population a large proportion of mature males are unable to establish a territory, providing the opportunity to compare the vocal behaviour between territorial and non‐territorial males. We examined how seasonal patterns of vocalizations differed between territorial and non‐territorial males and how the number of females present in a territory influenced behaviour of territorial males. We found that territorial males vocalized at higher rates than non‐territorial males, and territorial males vocalized more at the onset than towards the end of territory tenure. During the onset of territory‐tenure vocalizations of territorial males were directed more often towards other males than females. Furthermore, we found that vocalizations of territorial males were not only given in male–male interactions, but were also actively directed towards females. Territorial males vocalized at higher rates when more females were present in their territory. Our results suggest that vocalizations are important in male–male interactions, are relevant in territorial disputes and are used as a proxy for females to assess a male’s quality.     

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 in the pheasant (Phasianus colchicus): harem formation, mate selection and the role of mate guarding

Harem formation and mate selection were studied in the pheasant in order to determine the advantages of territorial harem defence polygyny to the two sexes. We investigated the factors affecting harem size and the advantage to a female in remaining with one territorial male during breeding.
Female group size declined during late March and early April as females moved from large overlapping ranges into smaller, more widely dispersed breeding ranges. The proportion of female groups accompanied by males increased during this period.
Some males had a disproportionate share of females. Settled females were monogamous but, because a female's nest was generally outside the male's territory, her home range was larger than his territory.
Harem members were usually from the same winter group. Harem size was not related to territory quality in terms of food supply or nesting cover. Females were loyal to one male in more than one year even if his territory position changed. Older, territory-owning males had more females, both adult and immature, than males with newly-established territories. Harem size was not correlated with territory size.
We conclude that the mating system of the pheasant is based on mate guarding which protects females not only from the risk of predation or injury, but also from excessive energy expenditure incurred through being chased by other males. When escorted by a territorial male, females spent three times as much time feeding, one-fifth as much time running, and one-tenth as much time alert, as they did when not guarded.     

Pair territoriality in the longnose filefish,oxymonacanthus longirostris

The longnose filefish,Oxymonacanthus longirostris, usually lives in heterosexual pairs, the male and female swimming together and sharing the same territory. Pair territoriality in the species was examined in detail in relation to sexual differences in territorial defense activities. Rigorous pair territoriality was maintained only during the breeding season, although pairs used their home ranges exclusively to a certain extent, during the non-breeding season. The frequency of aggression against other conspecific pairs in the breeding season was higher than in the non-breeding season. Agonistic interactions appear to be over both mates and food resources, the strict pair territoriality in the breeding season possibly being due to mutual mate guarding. In intraspecific aggressive interactions, males usually led their partner females when attacking intruders. The feeding frequency of males was much lower than that of females in the breeding season. Mate removal experiments indicated that females could not defend their original territories solitarily and their feeding frequency decreased. Conversely, males could defend territories solitarily without a decrease in feeding frequency. These results suggest that males contribute most to the defense of the pair territory, with females benefiting from territorial pair-swimming with their partner males.     

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.     

Mating system structure and population density in a polygynous lizard, Sauromalus obesus (= ater)

Increases in population density often are associated with achange in mating system structure in numerous taxa. Typically,male interactions are minimal in extremely low density populations.As density increases, males exhibit territoriality but if densitybecomes too high, the energetic cost of defending a territorywill eventually outweigh the reproductive benefits associatedwith territoriality. Consequently, males in high density populationsmay abandon territoriality and adopt dominance polygyny, lekking behavior, or scramble competition. We investigated the relationshipbetween population density and mating system structure in threepopulations of the chuckwalla, Sauromalus obesus (= ater),near Phoenix, Arizona. Densities in the Phoenix Mountains (2.7chuckwallas/ha) were lower than any population previously studied.In the Santan Mountains (10.9 chuckwallas/ha), densities weresimilar to populations studied in the Mojave Desert, and inthe South Mountains (65 chuckwallas/ha), densities were the highest yet recorded. Male mating behavior was examined by determininghome range overlap and by making direct behavioral observations.Male home range size decreased with increasing population density.There was little overlap in home ranges among males in allthree populations, whereas home ranges of males and femalesconsistently overlapped, indicating that males were strictly territorial. This conclusion was supported by behavioral observationsof interactions among individuals in a natural setting. Thenumber of females wihin male territories was correlated withfood resources (plants) in all three populations. Female homerange size appeard to be related to food resources whereasmale home ranges appeared to be related to female distribution,population density, and geology. The retention of territoriality in spite of high population densities raises new questions aboutthe relationship between density and resource defense.     

Why are Male Columbian Ground Squirrels Territorial?

Male territorial defence is a component of many vertebrate mating systems and is often regarded as a tactic for acquiring mates. Traditionally considered within the context of overt site‐specific defence, territoriality actually may have several components which encompass a variety of behavioural tactics (e.g. post‐copulatory mate‐guarding, defence of resources that females need, defence of area around females) that underlie a mating system. The purpose of our study was to evaluate such influences on the territorial behaviour of male Columbian ground squirrels in southwestern Alberta, Canada. Males were dominant and territorial if they defended a minimum convex polygon activity range by chasing other males more within the activity range than they were chased. Subordinate males had no territory and were chased throughout their ranges, but they competed for mates by increasing chases in their activity range when nearby females were oestrous. Dominant males exhibited conditional breeding tactics, tending to chase other dominant males from their territory when nearby females were oestrous, but travelling outside their activity ranges to chase subordinate males when females were not oestrous. Although females mated first with a dominant male on whose territory they resided (and in order from oldest to youngest if several territories overlapped), mating pairs were not exclusive, as females usually mated with additional males. Males also guarded females after copulation and defended females directly just before oestrus, rather than defending territory per se during those times. Thus, males possess a repertoire of behaviours that complement site‐specific territoriality, and territory ownership serves to facilitate a first mating with females that live on the territory.     

Temporal and Spatial Distribution of Male Scent Marks in the Polygynous Greater Sac-Winged Bat

Scent marks are relatively long-lived signals that can be perceived by conspecifics when the producer is absent. Therefore, it is often not obvious to whom the signal is directed. In daytime roosts of the polygynous greater sac-winged bat, males scent mark territories with facial gland secretions. Territories are a valuable resource for males, as they offer exclusive courtship opportunities, which results in increased male reproductive success and, consequently, increased male–male competition over territories. The information encoded in male scent marks could, therefore, be either directed at females as part of an olfactory courtship display or at male competitors as part of territorial behaviour. We expected territorial males to scent mark in the morning, shortly before females return to the territory and close to female roosting sites, if scent marks are directed at females as part of the courtship display. And we expected harem males to scent mark at the territory boundaries, where male–male encounters are most likely to occur, if scent marks are directed at male competitors. We found that males marked more frequently in the afternoon, at a time when all females have already left the territory, and harem males marked at the territory boundaries and not inside their territory in the area where females roost. At boundaries males fan volatiles from specialised wing sacs towards competitors outside the territory. Scent marking of male Saccopteryx bilineata might therefore be congruent with the assessment-hypothesis, which states that scent marks offer intruders the possibility to make an olfactory assessment of the territory owner without direct physical interaction. Thus, scent marks of male S. bilineata are most likely influenced by male–male competition and not by female choice.     

Scent marking in a territorial African antelope: II. The economics of marking with faeces

Faeces may be ideal substances for scent marking because they have a minimal energetic cost to the signaller. However, marking with faeces is also constrained by the animal's ability to produce faeces. This study examined whether limits on the volume of faeces produced by oribi Ourebia ourebi, in Serengeti National Park, Tanzania, caused territorial males to regulate their output and prioritize the placement of faecal marks. Territorial males marked with faeces more often, and with a smaller volume per defecation, than did juvenile males and females. Territorial males also defecated only on established dung middens along borders shared with other territorial males or on top of a female's urine and faeces. In contrast, juvenile males and females defecated randomly with regard to their location in territories. Territorial males with larger harems marked with faeces at higher rates and less volume than males with few or no females. This difference suggests that when males overmark female excretions they reduce the amount of faeces available for marking other preferred sites, such as along territory borders shared with other males. Dominant males with adult subordinates marked with faeces less often, and with a greater volume per mark, than males that defended territories without the aid of subordinates. Dominant males also reduced the volume of marks less as the number of females on their territory increased than did males without subordinates. Territories occupied by more than one adult male also were marked with faeces at higher rates, and with marks of greater volume, than territories held by single males. These results suggest that the presence of subordinate males reduced the demand on dominant males to regulate the volume and placement of faecal marks. Overall, these results suggest that territorial male oribi regulate their faecal marking behaviour in response to a limited supply of faeces. Copyright 1999 The Association for the Study of Animal Behaviour.     

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.     

Varying Ecological Quality Influences the Probability of Polyandry in White-handed Gibbons (Hylobates lar) in Thailand

Although members of the family Hylobatidae are known to be monogamous, adult white-handed gibbons ( Hylobates lar ) at Khao Yai National Park, Thailand, also show multimale groups and polyandry. A need for more than one male to successfully raise offspring cannot explain the occurrence of polyandry in these territorial primates, because direct paternal care is absent in this species. We hypothesize that polyandry is primarily related to costs/benefits for males of cooperatively defending a female and/or resources; our prediction was that polyandry would become more frequent with increasing costs of female/resource defense. We measured the ecological quality of seven gibbon home ranges over a 3-yr period (2001–2003) to investigate how resource availability affected the probability of polyandry, and found a significant negative relationship between home range quality and home range size. Larger home ranges were of lower quality. As predicted, groups living on larger, poorer home ranges also experienced longer periods of polyandry. In forest areas of comparatively low quality, acquiring and maintaining a large home range that includes enough resources for a female to reproduce steadily may surpass a single male's capacity. Our model of cooperative male polyandry was supported by preliminary data of shared territorial defense and access to the female. However, interaction proportions were strongly skewed, and female's primary male partners monopolized grooming and mating. Nevertheless, a primary male on a large territory may benefit from the presence of a secondary male with aid in territorial/female defense, whereas a secondary male may gain by avoiding high dispersal costs.     

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.     

The evolution of monogamy in primates

The evolution of primate monogamy is described as an ordered sequence of choices by generalized, hypothetical females and males. Females first choose whether or not to associate with other females. Predators encourage gregariousness in diurnal primates; however, nocturnality or scarce and evenly distributed food supplies may enforce separation. A testable group size model based on food patch size is developed and qualitatively supported.If females choose solitude, males then choose either to defend a single female and invest in her offspring, or to compete with other males for access to several females, usually by defending a territory or establishing dominance over the home ranges of several females. The decision rests on the defensibility of females and on the availability of an effective form of male parental investment. Both of these factors are dependent on local female population density. A model is developed that assumes that territorial defense is the principal form of male parental investment, and it predicts that monogamy should occur at intermediate densities: at high densities, males should switch to defense of multiple females, and at low densities there is no investment value in male territorial defense. The model is shown to be only partly adequate. Variation in local population densities prevents the establishment of obligate monogamy through territoriality in small monkeys, since male territorial behavior is inconsistent over the long run. Here, carrying of offspring by males can succeed territoriality, providing an effective and reliable form of parental investment to maintain the pair bond in the face of population fluctuations and changes in group structure. This hypothesis is supported by the scarcity of obligate monogamy among the prosimians, which frequently do not carry their young.     

The adaptive significance of non-contact mate guarding by males of the dragonfly,Nannophya pygmaea Rambur (Odonata: Libellulidae)

InNannophya pygmaea, ovipositing females were frequently disturbed by conspecific males. Disturbed females often copulated with one of these males or flew away from the pool. Females which flew away from the pool due to male disturbance often returned later the same day and mated with different males. A territorial male would guard his ovipositing mate by hovering above her, presumably trying to prevent her from moving out of his territory. A non-territorial male would also guard his mate in a similar way, both at a vacant water area which was not occupied by any territorial males, or within the territory of a resident male. In addition, both territorial and non-territorial males chased intruding males in an attempt to prevent their mates from being stolen. Territorial males defended their mates better than non-territorial males. Both males and females often mated more than once in the course of a single day. Some territorial males copulated with a new female while another mate oviposited in their territories. This observation supported the “multiple mating hypothesis” proposed by Alcock (1979) and Uéda (1979) but other evidence suggested that this is an inadequate explanation for the non-contact guarding ofN. pygmaea.     

Sex Differences in Territorial Behavior Exhibited by the Spotted Hyena (Hyaenidae, Crocuta crocuta)

Spotted hyenas ( Crocuta crocuta ) are gregarious carnivores that defend group territories against encroachment by neighboring conspecifics. Here we monitored the behavior of members of one clan of free-ranging spotted hyenas during border patrols, 'wars' with neighboring clans, and other interactions with alien intruders, to document differences between the sexes in territorial behavior in this species. We also examined the possibility that the probability or rate of attack on alien hyenas encountered within the clan's territory would vary with the sex of the intruders. Initiation and leadership of most cooperative territorial behaviors were by adult female clan members, although border patrols were occasionally conducted by groups composed exclusively of resident males. The vast majority of alien intruders into the territory of the study clan were males. Resident females were more likely to attack intruding females than intruding males, but hourly rates of aggression directed by females towards aliens did not vary with intruder sex. Resident males were more likely than resident females to attack alien males, and resident males directed significantly higher hourly rates of aggression towards intruding males than females. Although female leadership in most cooperative territorial behaviors distinguishes spotted hyenas from many mammalian carnivores, other sex differences in the territorial behavior of spotted hyenas resemble those documented in other gregarious predators. Sex differences observed in hyena territoriality are consistent with the hypothesis that male and female clan members derive different selective benefits from advertisement and defense of group territories.