Kinship and social tolerance among female and juvenile wild house mice: kin bias but not kin discrimination

Kin-biased social tolerance among house mice has been interpreted in terms of kin discrimination. However, several lines of evidence suggest it may instead be an incidental artifact of group member discrimination. This leads to very different predictions about the social consequences of relatedness within and between social groups. Social interactions between wild-stock adult female and juvenile house mice (Mus domesticus) established in neighbouring territorial groups within enclosures reveal relatedness to dominant males within groups as the major factor determining social tolerance of juveniles by females. Relatedness to the female herself had no significant independent effect on responses indicating tolerance. Females were generally more aggressive toward neighbouring-group juveniles (all unrelated to females) compared with those from their own group (all related to females), but were most aggressive toward neighbouring juveniles sired by the neighbouring dominant male. They were also more aggressive toward their own-group juveniles that had been sired by the neighbouring dominant but only when encountered in the neighbouring territory and with a greater bias against female juveniles. Females were least aggressive toward own-group juveniles sired by their own-group dominant male. The sire-bias in tolerance among females is similar to that reported among the dominant males themselves in an earlier study. As a result of the combined sire-bias in tolerance by adult males and females, juveniles sired by their own-group dominant males become less likely to intrude into a neighbouring territory with time. Overall, the results suggest that differences in social tolerance reflect discrimination on the basis of social group membership rather than relatedness between interactants and thus provide strong experimental evidence in support of incidental kin bias rather than kin discrimination.     

Territorial counter-singing in male sac-winged bats (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>): low-frequency songs trigger a stronger response

In many animal species, individuals compete for resources but avoid escalated conflicts by threat displays, i.e. a mutual signalling behaviour that enables the opponents to predict the outcome of the conflict without the necessity of actual fighting. For example, territory holders may use acoustic signals to communicate not only their own identity and the borders of their territory but also their competitive quality, fighting ability and motivation. Here, we show that male sac-winged bats, Saccopteryx bilineata, adjust their vocal territorial displays according to the fundamental frequency of territorial songs of their opponents. In playback experiments with territorial males, low-frequency stimuli elicited a higher territorial song rate and length than high-frequency stimuli. Male S. bilineata that sing more often and with lower fundamental frequencies have been shown to sire more offspring than their competitors. Fundamental frequency of territorial songs, hence, may reveal male quality and, consequently, the resulting threat posed to competing males. We argue that this is reflected in the increased response of competitors to low-frequency territorial songs shown here. Such competitive signalling behaviour has been shown in a few mammal species like red deer and baboons but, thus far, not in bats. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Kin discrimination in the spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>): nepotism among siblings

We examined patterns of affiliation, association, and aggression to inquire whether spotted hyenas (Crocuta crocuta) can distinguish among various groups of maternal and paternal siblings. If so, and if these animals conform to predictions of kin selection theory, then behavioral interactions among hyenas should vary with relatedness. We also considered familiarity-based recognition and phenotype matching as mechanisms hyenas might use to recognize kin. Patterns of affiliative behavior indicated that hyenas favored full-sibling littermates over half-sibling littermates or any other group of half-siblings. Rates of dyadic aggression generally did not vary with kinship. Hyenas associated more closely with half-sibling littermates than with non-littermate half-siblings, and hyenas affiliated more with maternal half-siblings than with paternal half-siblings, suggesting that familiarity-based cues might mediate discrimination among these sibling classes. In addition, operation of a phenotype-matching mechanism was suggested by the preference hyenas demonstrated during affiliative interactions for full- over half-sibling littermates, and by their lack of preference in these interactions for half-sibling littermates over non-littermate half-siblings. Phenotype matching was also suggested by our observation that paternal half-siblings cooperated more, and fought less, than did non-kin. Our data indicate that hyenas can discriminate among various types of siblings, that their social behavior conforms to predictions of kin selection theory, and that they recognize kin using mechanisms of both familiarity and phenotype matching.Communicated by S. Alberts     

A test of the "dear enemy effect" in the strawberry dart-poison frog (<Emphasis Type="Italic">Dendrobates pumilio</Emphasis>)

Some territorial animals exhibit a form of social recognition, commonly termed the "dear enemy effect", in which territory residents display lower levels of aggression toward familiar neighbors compared to unfamiliar individuals who are non-territorial "floaters". Despite the widespread occurrence of territorial social systems and use of acoustic signals for communication in anuran amphibians, only two previous studies have demonstrated vocally mediated dear enemy behavior in a territorial frog. In this study, I conducted neighbor-stranger discrimination playback experiments in a third species of territorial frog, the strawberry dart-poison frog, Dendrobates pumilio (Anura, Dendrobatidae). In the first experiment (n=24), I broadcast the calls of a subject's nearest neighbor and the calls of an unfamiliar individual from the approximate midpoint between the subject's and the neighbor's territories. Although males responded to the stimuli, they did not exhibit differential responses to the calls of neighbors and strangers. In a second experiment (n=22), I broadcast the calls of a neighbor and a stranger to subjects through a speaker located in the approximate center of the neighbor's territory. Males also responded to the playback, although less intensely than in the first experiment, but no discrimination between the calls of neighbors and strangers was found. Thus, territorial males of the strawberry dart-poison frog appear not to discriminate behaviorally between the advertisement calls of neighbors and strangers. Several proximate and ultimate-level hypotheses for this lack of vocally mediated neighbor-stranger discrimination are discussed.Communicated by T. Czeschlik     

Differential aggressiveness between fire salamander (<Emphasis Type="Italic">Salamandra infraimmaculata</Emphasis>) larvae covaries with their genetic similarity

Responding differentially to kin and non-kin is known to be adaptive in many species. One example is the inclusive fitness benefits of reducing aggression toward closer relatives. Little is known, however, about the ability of animals to assess differential degrees of genetic relatedness and to respond accordingly with differential levels of aggression. In the present study, we tested whether aggressiveness between body mass-matched pairs of fire salamander (Salamandra infraimmaculata) larvae covaried with the genetic similarity between them. We quantified aggressiveness at three levels of genetic similarity by selecting pairs within and across pools from recently genotyped populations. We also assessed aggression between pairs of siblings. Aggression and associated injuries decreased as genetic similarity increased across the groups. These findings suggest that cannibalistic salamanders can assess their degree of genetic relatedness to conspecifics and vary their behavioral responses depending on the degree of similarity between them along a genetic relatedness continuum.     

Genetic relatedness and eusociality: parasite-mediated selection on the genetic composition of groups

Summary Contrary to the expectations of kin selection theory, intracolony relatedness in eusocial insects is often low. We examined the idea that associations of low relatedness (high genetic variability) may be advantageous because of negative frequency-dependent selection on common host phenotypes by rapidly evolving parasites and pathogens. Using the natural host-parasite system of the bumble bee Bombus terrestris and its intestinal trypanosome Crithidia bombi, we investigated the transmission properties of parasites in host groups. Within naturally infested nests and in artificially constructed groups of workers, prevalence of infestation increased with time of exposure (Table 1). The susceptibility of isolated groups of workers to the parasites to which they were exposed differed with identity and natural infestation of their nest of origin (Table 2). In addition, those workers that were related to the individual introducing an infection to their group were more likely to become infested than were unrelated workers (Table 3). Although the bumble bee workers in experimental boxes appeared to differ in behavior toward kin and non-kin, making more physical contacts with kin, we found no discernible relationship between number of physical contacts and prevalence of infestation in a group. Therefore, we conclude that differences in parasite transmission reflected interactions among different host and parasite phenotypes. This system thus demonstrates the factors necessary for negative frequency-dependent selection by parasites on common host phenotypes - variability for susceptibility and infectiousness in host and parasite populations and similarity for these traits among related individuals. If, as we show here, high genetic relatedness within groups enhances parasite transmission, kin directed altruism may increase the risk of contracting parasites and infectious diseases. Therefore, parasites and pathogens may be an important force moderating the genetic structure of social groups. Offprint requests to: J.A. Shykoff at the present address     

Playback and speaker-replacement experiments on song-based neighbor,stranger, and self discrimination in male Red-winged Blackbirds

Summary Two experiments were conducted to deterine whether territorial male Red-winged Blackbirds discriminate among neighbor, stranger, and self songs. In the first experiment, recordings of the three classes of songs were played to territorial males. Responses to playback of stranger song, as measured by song rate, intensity of Song Spread display, and closest approach, were significantly stronger than to playback of neighbor song. In addition, stranger song elicited significantly more intense Song Spread displays than did self song.In the second experiment, territorial males were removed from their territories and each was replaced with speakers broadcasting recorded songs of a stranger, and of the removed male. Broadcasts of the removed male's own songs were more effective in discouraging trespassing by neighboring male redwings than were broadcasts of songs of a stranger. However, the two broadcasts did not differ in their ability to deter nonneighboring male redwings from flying through or trespassing within the speaker territory.We suggest that neighbor-stranger discrimination in the Red-winged Blackbird develops as a consequence of associative learing.     

Do kin always make better neighbours?: The effects of territory quality

Summary The effects of territory quality on kin-biased territorial defence behaviour and fitness of juvenile rainbow trout (Oncorhynchus mykiss) were examined by manipulating food and predation risk levels in an artificial stream channel. Groups of related (full sibling) or unrelated fry were observed in the channel under one of four treatments: (1) high food-low predator (high territory quality); (2) high food-high predator; (3) low food-low predator; and (4) low food-high predator (low territory quality). Fish within kin groups always initiated fewer aggressive interactions, defended smaller territories, exhibited higher proportions of threat type territorial defence behaviours (as opposed to overtly aggressive behaviours) and had a higher mean weight increase than non-kin groups. Within both kin and non-kin groups, decreasing territory quality significantly increased the frequency of aggressive interactions, the size of territories and decreased growth. It is argued that while kin-biased territorial defence behaviours are always present, decreased availability of resources and increased predation risk serve to reduce the magnitude of this kin bias. However, even in low-quality territories, the benefit of associating with kin may increase the probability of overwintering survival (increased fitness) of the young fish by increasing body size.     

Experimental evidence that bright coloration is not important for territory defense in purple martins

Summary I tested the hypothesis that bright breeding plumage in territorial males acts as a badge of fighting ability or aggressive motivation to intimidate intruders. Territorial male purple martins (Progne subis) whose iridescent blue plumage was lightened to mimic the appearance of subadult males did not suffer an increase in intruder pressure or loss of territory compared with control males. Bright plumage color itself did not deter intruders and was not important for successful territory defense. Furthermore, a bright coloration of owners was not associated with an increased level of aggression toward intruders. Results from parallel studies on this species suggest that bright coloration is important in territory acquisition. The effectiveness of badges of fighting ability and aggressive motivation in territory defense is limited by whether intruders benefit from assessing these traits in owners. Differences in signaling systems between species are due in part to differences in floater tactics and the mode of territory acquisition.     

Land or lover? Territorial defence and mutual mate guarding in the crimson-breasted shrike

Crimson-breasted shrikes produce duets which are used in interactions with neighbours and intruders. We examined two major hypotheses explaining duetting, the territorial defence and mate guarding hypotheses, using playback experiments. The responses of 12 pairs towards solo male, solo female and stereo duet playback were analysed using principal component analysis and ANOVA. The measures of response intensity revealed in most cases highly coordinated responses of the pair partners. Although these joint responses could suggest cooperative territorial defence or mutual mate defence, responses to the treatments differed between males and females in terms of three variables. Females increased their answer rates significantly in response to unpaired female intruders, compared to unpaired male intruders. During duet playback, females directed their responses equally at both intruding pair members. In contrast, males were equally responsive towards unpaired male intruders and intruding pairs, but in the latter case directed their responses at the male pair member. In the presence of unpaired female intruders, males increased their solo song rates, but in the presence of intruding pairs, they increased overlapping rates and duetted with their mates. These divergent responses suggest that females use duets for mate guarding against unpaired females, whereas males use solo songs for mate guarding and both solos and duets in territorial defence.     

Neighbours, strangers and male-male aggression as a determinant of lek size

Interactions between males on leks may play an influential role in lek formation and the regulation of lek size. In this paper I present the results of a playback experiment that simulated de novo settlement at sites adjacent to currently existing display territories of the ochre-bellied flycatcher, Mionectes oleagineus. In the study population, males displayed both solitarily and at small leks. A large proportion of males held no display territory at all. A stranger's song was played to both solitary and lekking males from 10 m outside their territorial boundaries. In separate playbacks, lekking males were also played neighbour's song. Both lekking and solitary territorial males reacted to the playback by decreasing their song rate, approaching the playback speaker and, on occasion, attacking the model. Solitarily displaying males responded more aggressively to playback of stranger's song than did lek males. Lek males were able to distinguish between their neighbour's and a stranger's song and did so irrespective of whether it was played from the neighbour's territory or from outside the lek. In addition to distinguishing between neighbours and strangers, lek males modified their responses to these different playbacks depending on where the playback originated. These results suggest that male-male interactions can be influential in structuring leks. In M. oleagineus, interactions between males are aggressive and act to limit rather than augment lek size. Received: 6 March 1996 / Accepted after revision: 9 December 1996     

A model for the evolution of communal foraging in hierarchically structured populations

A kin selection model is described for populations in which groups of interacting individuals (trait groups, sensu Wilson 1975) are spatially situated within larger aggregations. The model predicts the optimal foraging strategy when resources are shared with other trait group members and there is an individual risk in foraging. The ecological mechanism of variation in group fitness, differential resource accumulation, is explicitly incorporated into the model. The optimal foraging rate obtained from this model depends on the product of a benefit-to-cost ratio and a relatedness parameter. The appropriate definition of relatedness for the evolution of communal foraging is determined by the details of the ecological interaction between consumers and resources. When competition is purely intra-specific, the genetic correlation among interactants relative to other members of the local aggregation defines the relatedness parameter applicable to selection on foraging propensity. When competition is primarily inter-specific, the genetic correlation among trait group members relative to the entire population defines relatedness.     

Body morphologies shape territorial dominance in the satyrine butterfly <Emphasis Type="Italic">Lethe diana</Emphasis>

Male butterflies compete over mating territories via aerial interactions. It has often been suggested that flight performance, and thus flight-related traits, would play an important role in butterflies’ contest resolution; however, most studies have not demonstrated a clear relationship between these traits and territory ownership in butterflies. Males of the satyrine butterfly Lethe diana compete over territories via linear chases, a behavior that is uncommon among butterflies: rather, contests of most butterfly species consist of circling flights. In the present study, some morphological and physiological traits of L. diana were compared between territory owners and intruders to search for traits correlated with resource holding power (RHP). Territory owners had higher flight-muscle ratio (FMR) and were heavier than intruders, indicating that flight performance plays a key role in their contest resolution. Contest duration was not affected by the owners’ FMR or body mass, and thus I obtained no evidence supporting the hypothesis that contestants assess their relative RHP before making a decision to retreat. I speculate that the fact that these traits had an effect on territorial status in L. diana was a result of the characteristics of the contest behavior of this butterfly. During their contests, males L. diana fly faster in their linear flights than males of most other butterflies, which engage in slower circling flights. Therefore, males of L. diana probably need higher flight performance capabilities in order to win their contests.     

Investment in territorial defence depends on rearing environment in brown trout (Salmo trutta)

In many animals, territoriality will arise or cease depending on environmental factors such as intruder rate and resource availability. We investigated the effect of rearing environment on territorial behaviour in ~1.5-month-old brown trout. In the laboratory, wild-caught (reared at a low density) and hatchery-reared (high density) trout were allowed to defend a territory against a size-matched intruder reared in the same or the other environment. Because territorial behaviour should be relaxed at high-rearing densities, we hypothesized that hatchery-reared trout should value their territories less and therefore invest less in defence compared with wild-caught trout. However, in all cases, territory owners were more likely to win the contest and hatchery-reared trout were just as likely as wild-reared to win mixed contests. Furthermore, pairs of hatchery-reared trout initiated contests sooner, fought longer and were more aggressive during the contest compared with pairs of wild trout. When hatchery-reared owners met wild intruders, the contest ended sooner compared with when the roles were reversed. We conclude that territorial behaviour in brown trout is largely innate, but that the hatchery environment has promoted more aggressive individuals. These results suggest that hatchery-reared trout invest more time and energy to obtain the same contest success as wild trout. In conclusion, the lack of experience of territorial defence in a high-density rearing environment seems to reduce the efficiency of territorial behaviour. In turn, this may have negative consequences for the performance of released hatchery fish in the wild.     

Relatedness structure and kin-biased foraging in the greater horseshoe bat (Rhinolophus ferrumequinum)

Female greater horseshoe bats (Rhinolophus ferrumequinum) exhibit strong natal philopatry to their maternity roost over many years, leading to the aggregation of matrilineal kin. Maternity colonies may, therefore, be expected to comprise highly related individuals, and, as such, provide conditions suitable for the evolution of kin-selected behaviours. To test these predictions, we examined relatedness and behaviour among matrilineal kin within a colony in south-west Britain. Genetic analysis of 15 matrilines, identified from microsatellite genotyping and long-term ringing surveys, revealed average relatedness levels of 0.17 to 0.64. In contrast, background relatedness among colony females approximated to zero (0.03). These results suggest that inclusive fitness benefits may only be accrued through discriminate cooperation within matrilines, and not at the wider colony level. To examine whether the potential for such benefits is realised through kin- biased cooperation during foraging, females from two matrilines were radio-tracked simultaneously over 3 years. Pairwise home-range overlap correlated significantly with Hamilton's relatedness coefficient. The greatest spatial associations were observed between females and their adult daughters, which shared both foraging grounds and night roosts, sometimes over several years. Tagged females, however, generally foraged and roosted alone, suggesting that kin-biased spatial association probably does not result from either information-transfer or cooperative territorial defence. Such patterns may instead result from a mechanism of maternal inheritance of preferred foraging and roosting sites.     

Territory defence in tropical birds: are females as aggressive as males?

Much of our knowledge concerning the functions of territorial behaviour and how territories are defended by individuals comes from research on birds. The vast majority of this work has focused on temperate zone breeding territoriality in which territories are defended most obviously by males. Our understanding of the female role in territory defence is limited because they are less conspicuous and much harder to observe. We studied sex roles in territory maintenance and defence in a duetting, resident neotropical passerine, the white-bellied antbird (Myrmeciza longipes). This species maintains territories and pair bonds year round and both sexes sing and actively participate in territory defence. We performed a series of playback experiments throughout the dry (non-breeding) and wet (breeding) seasons. We exposed territorial pairs to three types of stimuli including: (1) single sex, male only songs, (2) single sex, female only songs, and (3) both sex songs/duets. Contrary to findings for most other tropical species, individuals defended their territories with equal levels of aggression regardless of stimuli. Furthermore, sex roles were very different, with males responding more aggressively than females to all stimuli throughout both seasons. Both males and females consistently responded more aggressively to territorial intrusions during the dry season than during the wet season, likely because food abundance is low in the dry season and territory value is high. Our analysis of duetting behaviour suggests that duets do not serve a significant role in mate guarding, or territory defence.     

Effects of Roads on Movements by Understory Birds in Mixed-Species Flocks in Central Amazonian Brazil

Abstract: Roads through tropical forest create linear disturbances that have unknown consequences for forest birds. We studied how a narrow, rarely used road through otherwise undisturbed Amazonian forest affected the movements and area requirements of understory birds that form mixed-species flocks. Differences in road maintenance led to two distinct treatments along the same road. Trees along the "closed" road formed a partial canopy connecting the two sides of the road, although the roadway itself was kept open. The "open" road was regularly maintained, making a complete opening 10–30 m wide.We followed 15 flocks, 5 each in interior forest, along the open road, and along the closed road. These flocks were led by Thamnomanes antshrikes, and each flock had a discreet, permanent territory. Flock territory size (mean = 8.5 ha) did not differ among the three locations. The open road formed the territorial boundary for all five flocks, although birds moved within a few meters of the edge of the road. The closed road was less of a barrier: 2 of 5 flocks used both sides of the closed road. Playback experiments showed that flocks readily crossed the closed road to approach agonistic vocalizations. Along the open road, even though birds responded to playback by becoming agitated and moving to the extreme edge of the roadside vegetation, they were less likely to cross the road and did so only after a longer duration of playback. Our results suggest that flocks respond to a road as they would to a long linear gap. They use the vegetation along the edges of the road, but because they are unwilling to cross the open area, it becomes a flock territory boundary. Similarly, as in forest gaps, successional change along the closed road produced suitable habitat for flocks. Although this suggests that roads are a trivial problem, we caution that this result applies only to narrow roads that are not accompanied by deforestation or other disturbance.     

No cultural transmission of species recognition between parents and offspring in free-living great tits and blue tits

Imprinting plays a key role in the development of species recognition, with young imprinting upon the morphological characters of their parents. However, the potential role that cultural transmission might play in species recognition remains largely uninvestigated. Great tits (Parus major) and blue tits (Cyanistes caeruleus) do not normally perceive each other as sexual competitors or potential partners. However, after reciprocal interspecific cross-fostering, both species may perceive individuals of the foster species as potential rivals or mates. Although the experience of being raised by heterospecifics clearly has affected the species recognition of cross-fostered birds, some of them breed naturally with conspecifics. The offspring of such cross-fostered birds (OCF) are hence raised by parents that look like ordinary conspecifics but display deviant species recognition as compared to controls in terms of aggressive response towards rivals. Comparing the aggressive behavior of OCF, cross-fostered birds and controls towards territorial intruders may thus help tease apart the influence of morphological vs behavioral cues of parents in the development of offspring species recognition. To this end, we compared birds from all three treatments with respect to their aggressive response to territorial intruders of both species during the breeding season. OCF and controls did not differ in their pattern of response towards heterospecific and conspecific stimuli. Compared to cross-fostered birds, OCF and controls showed less aggression towards heterospecific intruders, while the response towards conspecific intruders did not differ between treatments. These results demonstrate that both tit species imprint on the morphological characters of their parents, but that parental behavior is not important for the development of species recognition in terms of aggressive response towards territorial intruders.     

Habitat preference increases territorial defence in brown trout (Salmo trutta)

The striking ability of territory owners to repel intruders has generated a number of theoretical explanations as well as experimental studies in many animal species. However, effects of individual habitat preferences on territorial defence have rarely been studied. From the territory value hypothesis, we predicted that owners of preferred habitats should invest more resources in defence than owners of non-preferred habitats. We tested this prediction with young territorial brown trout in a two-stage experiment. First, trout were allowed to choose individually between gravel and a uniform bright substrate. As expected, they showed a significant (79%) preference for gravel. However, there was considerable variation between individuals in substrate preference, with a few fish preferring the bright substrate. Half of the tested fish were then transferred to a gravel substrate and the rest to a bright substrate, manipulating habitat type in relation to preference. Territory owners were then staged against size-matched intruders whereupon contest aggression was observed and the winner of each contest determined. Overall, owners won most of the contests. Satisfied owners won 86% and owners of less preferred territories, 74% of the contests. Furthermore, more satisfied owners attacked sooner and were more aggressive relative to the intruders. We conclude that brown trout show individual variation in habitat preference, which appears to be linked with their investment in territorial defence. These results suggest that understanding and modelling of animal contests could benefit from considering how territorial defence is influenced by individual habitat preference and specialisation. Received: 25 February 2000 / Revised: 22 May 2000 / Accepted: 25 June 2000     

Territory ownership and familiarity status affect how much male root voles (Microtus oeconomus) invest in territory defence

Neighbour–stranger discrimination occurs when individuals respond with more aggression to strangers than to territorial neighbours—a phenomenon termed the “dear enemy phenomenon” (DEP). We investigated the DEP with male and female root voles (Microtus oeconomus Pallas 1776) using field dyadic arena tests conducted in enclosures where we could test for the effects of familiarity (familiar versus stranger), ownership (resident versus intruder status) and resource-holding potential (body mass) on territorial behaviours. The results showed that males put more effort into territorial defence than females, and males could discriminate between neighbours and strangers. In males, aggressiveness was influenced by a significant two-way interaction between treatment and ownership. Male residents were more aggressive towards stranger intruders than towards neighbour intruders, while male intruders were less aggressive towards stranger residents than towards neighbour residents. In females, neither treatment nor ownership status had a significant effect on aggressiveness. Familiar males performed more social behaviours but less non-social behaviours than stranger males. Furthermore, there was a clear dominance hierarchy between residents and intruders in stranger dyads, with the male territory holders dominating the intruder in pairwise interactions. To our knowledge, these results demonstrate for the first time DEP in a small mammal with a known pedigree and present the first evidence for “prior resident advantage” in voles. We argue that both ownership status and familiarity status affect how much an individual invests in territory defence. The benefits of neighbour–stranger discrimination for male root voles and the absence of neighbour–stranger discrimination in female root voles are discussed.