Allopatric differentiation in the acoustic communication of a weakly electric fish from southern Africa, <Emphasis Type="Italic">Marcusenius macrolepidotus</Emphasis> (Mormyridae,Teleostei)

A few species of the weakly electric snoutfish, the African freshwater family Mormyridae, have been reported to vocalise. However, allopatric populations of a single species were never compared. Members of three allopatric Marcusenius macrolepidotus populations, originating from the Upper Zambezi River in Namibia, the Buzi River (Mozambique), and the Incomati River system in South Africa, vocalised with pulsatile growl- and tonal hoot sounds in dyadic confrontation experiments. A high rate of growling accompanied territorial and agonistic interactions and also non-threatening interactions between males and females, which in one pair appeared to be courtship. Growl sound characteristics of M. macrolepidotus from the Incomati system differed from those of the Upper Zambezi in a significantly higher frequency of the first harmonic (mean, 355 Hz vs 266 Hz). The two vocalising males from the Buzi River generated growls about twice as long as the other fish. Furthermore, the growl pulse period was about 4 ms in M. macrolepidotus from the Upper Zambezi River and from the Incomati system, but 6 ms in M. macrolepidotus from the Buzi River. Hoots were only observed in agonistic encounters. Hoot oscillograms showed a sinusoidal waveform, and the mean duration of this sound was similar in Incomati system fish (mean, 161 ms), Upper Zambezi fish (172 ms) and Buzi fish (103 and 145 ms for the two vocalising individuals). The mean frequency of the first hoot harmonic was higher in Incomati system fish (326 Hz) than in Upper Zambezi fish (245 Hz). Both growl and hoot occurred only in the presence of conspecifics, probably signalling the presence and condition of an opponent, territory owner or potential mate. This is the first evidence for (1) sound production and acoustical communication in another species and genus, M. macrolepidotus, from southern Africa to be (2) geographically differentiated.     

Size assessment in simulated territorial encounters between male green frogs (Rana clamitans)

We examined the ability of male green frogs to assess the size of an opponent based on the dominant frequency of their advertisement call, which is negatively correlated with size, using synthetic stimuli to simulate intruders of different sizes. In one field playback experiment, we broadcast a pair of stimuli representing a small and a large male; in a second experiment, we broadcast calls of a medium and a large male. In both experiments, males produced calls with significantly lower dominant frequencies in response to each stimulus. Contrast analyses revealed that males lowered the dominant frequency of their calls more in response to the large-male stimulus than in responses to the small- and medium-male stimuli. In the second experiment, males also responded to the large-male stimulus by calling at higher rates. There were no differences in mean note duration or the number of moves made toward or around the playback speaker in response to any stimulus. Thus, the frequency of an opponent's calls elicits a differential modification of calling behavior, primarily in the form of differential dominant frequency alteration, suggesting that males use dominant frequency to assess the size of opponents during aggressive encounters. Received: 17 April 1998 / Accepted after revision: 7 October 1998     

Scent-marking behavior in wild groups of common marmosets (Callithrix jacchus)

Scent-marking in mammals has been frequently related to within-group social and reproductive dominance and to defense of territory and resources. We studied the scent-marking behavior of five wild groups of common marmosets, Callithrix jacchus, during 5 months of the fruiting season in northeastern Brazil. Circumgenital marking was the most common type of marking. Marks were distributed throughout the home range and were deposited mainly during travel and intergroup encounters. Although marks were commonly deposited at gum trees, there was no evidence that the animals used scent marks to label fruiting trees or sleeping sites. Contrary to expectations, reproductively dominant females did not mark more than reproductively subordinate females. Moreover, during intergroup encounters, reproductively subordinate females displayed higher frequencies of scent-marking than the reproductively dominant females of their group. Our results suggest that scent-marking is not strictly tied to reproductive dominance or territorial (or resource) defense in common marmosets. Because marks provide information about individual identity and reproductive condition, scent marks could serve different functions when used by different individuals. The high frequency of marking by reproductively subordinate females during intergroup encounters suggests that scent-marking might be used to signal to individuals of neighboring groups. Our data highlight the importance of social and ecological variables in scent-marking behavior. Received: 26 October 1998 / Received in revised form: 20 May 1999 / Accepted: 30 May 1999     

Female territoriality and the function of scent-marking in a monogamous antelope (Oreotragus oreotragus)

In monogamous species, females often choose between males according to the quality of the territories they defend, but the extent to which females themselves contribute to territory defence is frequently underestimated. Here we test for differences in male and female roles during paired scent-marking bouts, a key component of territorial defence, in a monogamous antelope. In two populations (Kenya, Zimbabwe) of klipspringer, Oreotragus oreotragus, both males and females usually scent-marked at the same site, but there were significant differences between sexes in terms of investment within bouts. Females initiated most bouts, thus dictating the marking strategy of the pair. Males initiated relatively few bouts, but deposited more scent marks per bout than females and were usually the last to scent-mark before leaving the site; they marked on the same branches as the female and thus overmarked her scent. Both sexes deposited more marks during paired than solo visits. Immediately preceding and following scent-marking bouts, males approached females and females left males more often than expected. Female scent-marking rates were higher when they were receptive than at other times, and this increase was matched by elevated marking rates of males. Females may increase marking rates when they are receptive in order to test the quality of their mate or to incite male competition. However, these ideas are unlikely to explain female scent-marking behaviour outside the mating season, which appears to be related primarily to territorial defence. We suggest that these differences in investment in scent-marking bouts are consistent with predictions that females may be autonomously territorial and that overmarking of female scent by males is a form of mate-guarding. Received: 17 November 1999 / Received in revised form: 24 February 2000 / Accepted: 13 March 2000     

Site fidelity and patterns of short- and long-term movement in the brilliant-thighed poison frog <Emphasis Type="Italic">Allobates femoralis</Emphasis> (Aromobatidae)

We studied movement and site fidelity of males and females of the territorial frog Allobates femoralis (Aromobatidae) in a population in the Nature Reserve “Les Nouragues” in French Guiana, South America. Observations during 3 months in 2006 ascertained intra-seasonal site fidelity for males and females. Males actively defend large multi-purpose territories whereas females retreat to small resting sites from where they commute to neighbouring males for courtship and mating. Female short-term movement corroborates the previous assumption of a polygynous or promiscuous resource-defence mating system. Year-to-year recaptures from 2005 until 2008 revealed distinct patterns of inter-annual movement for males and regional site fidelity for females. Males abandon their territories and have to re-negotiate them when reproduction starts again at the end of the dry season. Females are not subject to intra- or inter-sexual territorial competition and as a result move significantly less between reproductive seasons than males. Male long-term movement reflects spatial structure and prevailing social interactions and is a reliable indicator for tadpole deposition sites. The combined effects of intra- and inter-seasonal movement promote the diversity of mates for both sexes.     

Familiarity with adults,but not relatedness,affects the growth of juvenile red-backed salamanders (<Emphasis Type="Italic">Plethodon cinereus</Emphasis>)

The relative roles of kinship and familiarity in affecting an individual’s growth and fitness are not easy to disentangle. Not only is an individual more likely to have prior behavioral interactions with conspecifics in close proximity, it may also be related (in terms of kinship) to those nearby conspecifics. While some studies have inferred that kin discrimination affects fitness correlates, other studies found that familiarity alone can reduce aggressive interactions, thus increasing fitness. These studies have all focused on intra-age class pairs or groups. However, many animals interact with conspecifics from different cohorts. In many populations, adults of Plethodon cinereus territorially defend rocks and logs that retain moisture and food resources. We investigated whether juveniles of P. cinereus grew more in the presence of adults that were relatives or familiar. We collected pairs of juveniles and adults found under the same cover objects in the forest (familiar) and pairs of juveniles and adults found under different cover objects, approximately 10 m apart (unfamiliar). We determined parentage and relatedness of the adult–juvenile pairs and then placed these pairs in semi-natural mesocosms for 17 days. We found that juveniles housed with familiar adults had significantly greater increases in mass and snout–vent length than juveniles housed with unfamiliar adults in 2006 but not in 2007. Relatedness had no effect on growth. In addition, juveniles cohabitating with adults were not more likely to be their offspring. At least under certain environmental conditions, familiarity with adults, independent of relatedness or parentage, increased the growth of juvenile salamanders.     

Quantitative variation in ecological and hormonal variables correlates with spatial organization of pronghorn (Antilocapra americana) males

Whereas variation in pronghorn (Antilocapra americana) spatial organization is well documented, underlying ecological or physiological explanations are not well understood. This study quantitatively describes spacing systems of pronghorn males and correlates of their spatial organization. I collected behavioral data from two populations in South Dakota (Wind Cave) and Montana (Bar Diamond) to determine if males differed in space use, response to intruders, and behavior patterns indicative of area defense. I measured sex ratio and population density, and I examined characteristics of food resources, including forb species diversity, richness, coverage, biomass, and nitrogen content, and how they changed during the growing season. I also collected and analyzed fecal samples to determine if males differed in testosterone concentrations. Pronghorn males at Wind Cave were more territorial than males at Bar Diamond, although males at Bar Diamond became more territorial during the second year. The forb community at Wind Cave was more diverse, contained greater amounts of forbs later in summer, and had a higher nitrogen content later in summer. Population density was lower at Wind Cave, although density dropped at Bar Diamond during the second year, and sex ratios were skewed toward males at Bar Diamond. Finally, males at Wind Cave had higher testosterone concentrations than did Bar Diamond males, although differences were not statistically significant. With lower population density and higher forb abundance and quality, food resources were more economically defensible at Wind Cave, and males were more territorial there. Analyses using these and other pronghorn populations revealed that population density and sex ratio correlated weakly with spatial organization, whereas precipitation correlated most strongly, which suggests plant productivity has a powerful role in determining pronghorn territoriality. Received: 16 June 1999 / Received in revised form: 21 September 1999 / Accepted: 31 December 1999     

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.     

Inbreeding and competitive ability in the common shrew (<Emphasis Type="Italic">Sorex araneus</Emphasis>)

Common shrews (Sorex araneus) maintain a foraging territory for most of their immature life. Possessing a high-quality territory is vital for overwinter survival in the harsh boreal climate, and hence, competitive ability in territorial disputes is expected to be an important component of individual fitness. To test possible association between individual inbreeding and fitness, we used neutral arena trials to assess the competitive performance of young common shrews. The experiment involved pairs of individuals originating from small island populations, where breeding must often occur between related individuals, and from large outbred mainland populations. The percentage of neutral arena tests that an individual won was highly significantly explained by internal relatedness, a surrogate measure of individual inbreeding, measured using ten microsatellite markers. Body size, sex, learning, and population type (mainland vs island) made no significant contributions. Even a low level of individual inbreeding may lead to significant adverse consequences in multiple territorial contests, which may represent a significant cause of inbreeding depression in many wild vertebrate populations.     

Patterns of participation and free riding in territorial conflicts among ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Cooperation in animal social groups may be limited by the threat of free riding, the potential for individuals to reap the benefits of other individuals actions without paying their share of the costs. Here we investigate the factors that influence individual contributions to group-level benefits by studying individual participation in territorial defense among female ringtailed lemurs (Lemur catta). To control for potentially confounding factors, particularly group size, we studied two semi-free-ranging groups at the Duke University Primate Center. First, we used a combination of experimental and observational methods to investigate the costs and benefits of territorial defense for individual lemurs. We found three indications of costs: physical contact occurred during inter-group encounters, participation in territorial defense was negatively correlated with ambient temperature, and rates of self-directed behaviors increased during encounters. Benefits were more difficult to quantify, but observational and experimental tests suggested that individuals shared the gains of territorial defense by foraging in defended territories. Thus, during experiments in which one of the groups was prevented from defending its territory, the free-ranging group made more frequent incursions into the other groups territory. Second, we examined variation in participation in territorial defense. Individuals varied significantly in their rates of aggression and genital marking during inter-group encounters. The extensive variation documented among individuals was partially accounted for by dominance rank, kinship and patterns of parental care. However, we found no evidence to suggest that participation was enforced through punishment (policing) or exchange of benefits involving grooming. In conclusion, this study provides further insights into cooperative behavior in mammalian social groups by revealing how the costs and benefits of territoriality influence patterns of individual participation in the context of shared (collective) goods.Communicated by P. Kappeler     

Territorial vocal behavior in hybrid smooth froglets, Geocrinia laevis complex (Anura: Myobatrachidae)

Males of the parapatically distributed myobatrachid frogs Geocrinia laevis and G. victoriana have highly divergent advertisement calls. Furthermore, the two species differ strongly in the complexity of their vocal repertoires, with males of G. victoriana possessing, and those of G. laevis lacking, a distinct territorial vocalization (encounter call). We investigated the territorial vocal behavior of males in a persistent natural hybrid population. Most hybrid males possessed encounter calls functionally equivalent to those of G. victoriana, that were produced following exposure to playback of recorded advertisement calls presented at >110 dB peak sound pressure level. The territorial acoustic responses were not associated with an index of hybridity derived from the structure of the advertisement call, suggesting genetic and functional decoupling of the two components of the vocal repertoire; i.e., advertisement calls and encounter calls. This decoupling may be the result of sexual selection favouring those hybrids with pronounced territorial behavior and the associated vocalization, regardless of the structure of their advertisement calls.Communicated by A. Mathis     

Multimodal signals,imperfect information,and identification of sex in red-backed salamanders (<Emphasis Type="Italic">Plethodon cinereus</Emphasis>)

Theoretical treatments and empirical studies both suggest that signals that occur in multiple sensory modes have superior detectabilities, discriminabilities, and memorabilities. There is also an intuitive link between signal detectability, discriminability, and memorability and the quality of information that is transferred via the signaler. We investigated the role of information quality and sensory modality in the sexual identification of intruding conspecifics by territorial male red-backed salamanders (Plethodon cinereus). Our results imply that these salamanders are able to discriminate between intruding male and female conspecifics (based on the allocation of aggression) in bimodal scenarios (vision and olfaction) even under situations in which the information available about the intruders sexual identities is reduced in quality. In unimodal scenarios (olfaction only), male residents exhibited heightened levels of aggression toward male secretions and reduced levels of aggression toward female secretions. In unimodal scenarios where information pertaining to the sex of the stimulus was reduced in quality, male residents displayed intermediate levels of aggression relative to the responses that male and female stimuli received respectively. Although resident males touched bimodal stimuli significantly more than they touched unimodal stimuli, we were unable to find support for the notion that sensory modality greatly influences how male resident red-backed salamanders allocate aggression toward intruding male versus female stimuli.Communicated by H. Kokko     

The significance of male-male aggression for yearling dispersal in roe deer (Capreolus capreolus)

The significance of adult buck aggression as a proximate factor driving yearling male dispersal in roe deer, Capreolus Capreolus, was studied in Sweden during the dispersal season (April–June) in 1988–1991. In roe there is a well established positive correlation between yearling antler size and dispersal tendency. I tested the prediction that yearlings with large antlers will experience more agonism from adult bucks than those with smaller antlers. Additionally, I studied the relationship between antler size and level of sexual maturity, by recording date of velvet shedding, and sampling testes weights from culled yearlings during the breeding season. The amount of agonism experienced by yearlings was positively correlated with the size of their antlers during April and May, whereas in June no such relationship was found. Aggression towards all yearlings tended to increase as the season progressed. A positive correlation between antler size and testes weight was evident, and the larger the antlers of yearlings the earlier the velvet shedding occured. These results indicate: firstly, that adult buck agonism is an important proximate factor underlying yearling male dispersal; and secondly, that agonism is mainly directed towards the sexually most mature yearlings. This, in turn, suggests mate competition to be the ultimate cause of male-male agonism, and thus male dispersal in this species.     

A comparative study of aggressiveness in eastern North American field cricket species (genus <Emphasis Type="Italic">Gryllus</Emphasis>)

To understand the variation in aggressiveness and factors important for contest outcome, we quantified and compared agonistic interactions of four field cricket species in eastern North America: Gryllus fultoni (Orthoptera; Gryllidae), G. vernalis, G. pennsylvanicus, and G. rubens. The most aggressive behavior that we observed, the grapple, was frequently displayed in agonistic trials of G. pennsylvanicus and G. rubens. By contrast, we never observed this behavior in trials involving G. fultoni and G. vernalis. Consequently, species was the only significant variable affecting the variation in aggressiveness, whereas size difference, age difference, and contest duration had no significant effect. In species with high levels of aggressiveness, G. pennsylvanicus and G. rubens, the factors that might be indicative of resource holding potential such as size or age difference seemed to be significant for contest outcome. In species with low levels of aggressiveness, G. fultoni and G. vernalis, however, there were indications that contest outcomes were determined by aggressiveness itself rather than the size and age differences between opponents. Markov chain analyses revealed that the difference in aggressiveness between species with high and low levels of aggressiveness lay in a sequence of escalating behaviors: antennal fencing, mandible flare, and grapple. The escalated state of aggressiveness characterized by this behavioral sequence in G. pennsylvanicus and G. rubens seems to be the ancestral state in the North American Gryllus phylogeny. We argue that the loss of a tendency to use burrows in G. fultoni and G. vernalis might be related to low levels of aggressiveness in these species.     

The functional significance of infanticide in a population of California ground squirrels (Spermophilus beecheyi)

Naturally-occurring infanticide was observed in a population of California ground squirrels (Spermophilus beecheyi). In four seasons, 40 infanticides were observed. All victims were post-emergent pups. Of 37 killings in which the killer was sexed 36 were by females. All infanticidal females were mothers at the time they killed, but in no case was a mother seen to kill or harm her own young. The victim was cannibalized in 22 cases and taken immediately into the killer’s burrow in 16 others. In no case did killers gain access to the victimized mother’s burrow or territory and female pups were not killed preferentially over males. In light of evolutionary explanations, infanticide in this population may best fit the resource exploitation hypothesis, in which killers commit infanticide to gain a nutritional benefit. Resource competition is a possible auxiliary explanation, since any time a female kills unrelated young she is eliminating possible competitors to her own offspring. This behavior could confer a fitness advantage on killers or it could be an adaptively neutral, alternative feeding strategy. More data are necessary to distinguish between these hypotheses. Received: 26 January 1995/Accepted after revision: 9 September 1995     

Neighbor–stranger discrimination in Audubon's shearwater (Puffinus l. lherminieri) explained by a “real enemy” effect

Neighbor–stranger discrimination (NSD) occurs when animals respond with more aggression to strangers than to territorial neighbors. NSD has been reported in many species that defend multi-purpose territories for breeding and foraging, but it is rare among species that defend other types of territories. For birds that defend only their nest sites, there is no experimental evidence for NSD, and observational studies have provided mixed results. In a colony of Audubon's shearwaters (Puffinus l. lherminieri), I played back the calls of a neighbor and a stranger to males defending nest sites. Subjects responded with longer calls to playbacks of strangers than to those of neighbors. In shearwater colonies, strangers are often birds looking for future breeding sites. In contrast, there is no evidence that established breeders compete with their neighbors for any resources. Shearwaters should benefit from NSD because strangers represent a “real enemy” and established neighbors do not.     

Behavioral diversification in a young species flock of pupfish (<Emphasis Type="Italic">Cyprionodon</Emphasis> spp.): shoaling and aggressive behavior

Increasing empirical and theoretical evidence supports the idea that sympatric speciation is operating, for example, in species flocks comprising several closely related fish species within one lake. Divergent natural selection (promoting spatial and food niche partitioning) and sexual selection (assortative mating) have been identified as key selection factors in intralacustric adaptive radiations. However, the evolution of social behaviors accompanying such adaptive radiations is less well understood. Using a phylogenetically young species flock of pupfish (Cyprinodon spp.) as a model, we examined differences among six sympatric species and compared their shoaling, aggressive, and territorial behaviors with that of a sister species (C. artifrons). Despite an estimated age of the species flock of less than 8,000 years, pronounced behavioral differentiation was found. C. simus, the smallest species in the flock, shoaled more than the other species and was less aggressive and less territorial than C. beltrani. F₁-hybrids between C. simus males and C. beltrani females showed an intermediate expression of shoaling and aggressive behavior. Niche partitioning among the members of this species flock appears to be accompanied by rapid divergent evolution of social behaviors. We discuss the potential role of phenotypic plasticity and within-species variation of social behaviors for such rapid behavioral diversifications in sympatric speciation processes.     

The timing of chemical signaling with urine in dominance fights of male lobsters (Homarus americanus)

Chemicals carried in the urine are a rich source of information about the identity, sex, aggressive motivation, and other attributes of an animal. In agonistic interactions, animals should be careful about disclosing such information, since it can be used by receivers to the disadvantage of the sender. By adjusting the timing of urine release, the signaler may still influence the behavior of the receiver to its own benefit. Here we investigate the urine signaling of American lobsters (Homarus americanus), necessary for the maintenance of dominance, using a catheter technique with high temporal resolution. We hypothesize that urine release in lobster fights is not continuous but restricted to moments that may elicit a reaction in the receiver beneficial to the sender. We found that eventual winners signaled more readily in the initial phase of the fight. Eventual losers did not show such initial urination. This suggests that a confident lobster may use urine signals to influence the behavior of its opponent into giving up the fight at an early stage. For both winners and losers, urine release is linked to offensive behavior and increases as the intensity of agonistic behavior increases. The coupling of urine release to offensive behaviors appears to improve both the reliability and efficacy of the chemical signal. Releasing urine under the increasing risk of inflicting injury during the fight provides a handicap that may ensure reliability of the threat signal. Furthermore, the immediate coupling of urine components to offensive behavior may help to consolidate the receiver"s memory of the signaler"s individual scent and thus facilitate future recognition of the dominant animal. Received: 6 October 1999 / Accepted 28 August 2000