Natal emigration by both sexes in the La Pacifica population of mantled howlers: when do some stay?

We have reported previously that all male and female mantled howlers emigrate from natal groups at Hacienda La Pacifica, Costa Rica. In the years since that report, a small number of juveniles have stayed in the natal group without experiencing a solitary phase. Here, we present a post hoc analysis on juvenile emigration in six groups of howlers under observation for varying amounts of time between 1972 and 2005. Our records revealed 139 juveniles for whom emigration status was certain, and 125 of these did emigrate. There was a significant association between presence of mother and emigration: juveniles without mothers were more likely to remain in their natal group (chi(1)(2) = 53.1, P<.0001). The mean age of emigration for all juveniles (n = 125) was 2.47 years (SD = 0.9, range = 1.5-6.5). There was no difference in age of emigration by adult male composition (one-male, multi-male, both), but juveniles of unknown sex emigrated younger than either known males or females (F(2,116) = 4.4, P<.02). For emigrating juveniles of known sex (n = 99), both males and females without mothers left at a later age than those with mothers (F(1,95) = 6.5, P<.02). Although philopatry or delayed emigration occurs in a few motherless animals, most males and females do emigrate from their natal groups at ages consistent with those reported for other species of howlers.     

The effect of prey abundance and foraging tactics on the population dynamics of a social, territorial carnivore, the spotted hyena

We used naturally occurring spatial and temporal changes in prey abundance to investigate whether the foraging behavior of a social, territorial carnivore, the spotted hyena ( Crocuta crocuta ), conformed to predictions derived from the ideal free distribution. We demonstrate that hyenas in the Ngorongoro Crater, Tanzania, redistributed themselves from less profitable to more profitable areas, even when this required them to undertake foraging trips to areas beyond their clan territory boundary, or required normally philopatric females to emigrate. As expected for a system with rank related access to food resources in the territory, females of low social status foraged more often outside their territory and were more likely to emigrate than dominant females. Probably because Crater hyenas regularly foraged outside their territories, there was no correlation between clan size and prey density within territories, suggesting that clan sizes may have exceeded the carrying capacity of territories. A substantial decline of the hyena population in the Crater from 385 adults in the mid 1960s to 117 in 1996 was most likely due to a substantial decline of their main prey. The decline in the hyena population was associated with a decline in the size of clans but not in the number of clans. The number of clans probably remained constant due to emigration by females from large clans into vacant areas or clans with no adult females, and because hyenas regularly fed in areas containing concentrations of prey beyond their territory boundary. Between 1996 and 2003 annual recruitment rates of Crater hyenas consistently exceeded annual mortality rates, resulting in an almost doubling of the adult population. This increase was most likely due to an increase in prey abundance, a relatively low level of predation on hyenas by lions ( Panthera leo ), and an absence of high levels of disease related mortality.     

Social organization of the Alaotran gentle lemur (Hapalemur griseus alaotrensis)

Knowledge of the social organization of lemurs is still limited for most species. Where there is sufficient information, it has been shown that lemur social organization differs in essential points from that of other primates. In the field study reported here, demographic structure and life-history processes were investigated in order to characterize the social organization of the Alaotran gentle lemur (Hapalemur griseus alaotrensis). Data were obtained through captures and observations. Alaotran gentle lemurs were found in small groups of up to nine individuals. Although most groups contained just one breeding female, a substantial proportion of groups (35%) had two breeding females. Therefore, Alaotran gentle lemurs cannot be classed as being organized in monogamous family groups. An extended birth season was found, and groups with two breeding females had significantly higher breeding output per adult than groups with a single adult female. Limited data suggest that females emigrate from their natal group while still subadult, whereas males can stay in the natal group until they are fully grown and disperse as adults. Variability in group composition, significantly higher reproductive output per adult in groups with two breeding females, and delayed dispersal of males suggest that Alaotran gentle lemurs pursue a resource-defense mating strategy, rather than a female-defense mating strategy. The suggestion that extant social lemurs may have evolved from a monogamous system, could explain the differences between lemur social systems and those of other primates.     

The social organization of feral peafowl

Peafowl are usually reported to have a mating system based on harem defence by adult males. In a small feral population near Oxford, males defended small (<1 ha) territories while females remained in one flock that ignored male territory boundaries. After mating, females become solitary. At no time did a female associate selectively with one male or remain within his territory, nor did males attempt to follow or guard female groups. Two out of four males were seen to mate. These differed from the other two in being neither very old nor very young; they held territories smaller than that of the young male and were no larger or longer-tailed. However, they spent more time displaying. We suggest that peafowl have a mating system similar to a lek: males defending small, clumped territories visited by females for mating.     

Variation in dispersal mortality and dispersal propensity among individuals: the effects of age, sex and resource availability

1.  Dispersal of individuals between habitat patches depends on both the propensity to emigrate from a patch and the ability to survive inter-patch movement. Environmental factors and individual characteristics have been shown to influence dispersal rates but separating the effects of emigration and dispersal mortality on dispersal can often be difficult. In this study, we use a soil mite laboratory system to investigate factors affecting emigration and dispersal mortality.
2.  We tested the movement of different age groups in two-patch systems with different inter-patch distances. Differences in immigration among age groups were primarily driven by differences in emigration but dispersal mortality was greater for some groups. Immigration declined with increasing inter-patch distance, which was due to increasing dispersal mortality and decreasing emigration.
3.  In a second experiment, we compared the dispersal of recently matured males and females and tested the impact of food availability during the developmental period on their dispersal. Dispersal was found to be male biased but there was no significant sex bias in dispersal mortality. There was some evidence that food availability could affect emigration and dispersal mortality.
4.  These results demonstrate that both emigration and dispersal mortality can be affected by factors such as individual age and resource availability. Understanding these effects is likely to be important for predicting the fitness costs and population consequences of dispersal.     

Turnover and dispersal in a Peregrine Falco peregrinus population

In south Scotland, most Peregrines returned to the same territories to breed in successive years, though a few females changed territory from one year to the next.
Annual mortality among breeding birds was at most 9% among females (or 11% in both sexes combined). There may have been considerable annual variation, however, and excluding one exceptional year out of five reduced the estimate for females to 7%. These estimates are maxima, but are still considerably lower than those obtained from ring recoveries of dead birds reported by members of the public.
Among trapped birds, four males first bred at age two years, one at three and another at four or five; two females first bred at one year, 13 at two years old and one at three. Five other females which were seen to be in first-year plumage but were not trapped, also laid eggs, and 12 other such paired females held territory but did not lay. Only one paired male held territory in first-year plumage.
In their movements between natal and breeding territories, some females moved further than males, with median distances of 83 and 58 km respectively. In addition, of birds trapped breeding in the study area, a greater proportion of the males than of the females had been born locally, despite an equal sex ratio among fledglings; this was also consistent with a greater dispersal of females. In general, Peregrines made much longer movements in their first year of life than subsequently. Movements were in any direction.     

Sexual conflict over breeding substrate causes female expulsion and offspring loss in a cichlid fish

Females of the Lake Tanganyika cichlid Lamprologus callipterusexclusively breed in empty snail shells that males collect intheir territories. Male–male competition for shells issevere, leading to frequent shell stealing and territory takeover.As a consequence, males have breeding females in their shellsthat spawned with competitors. In this field study, we investigatedboth naturally occurring and experimentally induced encountersof territorial males with females that had spawned with othermales. We found that the breeding success of females that weretaken over by a different male was significantly reduced. Behavioralobservations after experimental shell relocation further showedthat males recognized females that they had not spawned with:males directed more exploration and manipulation behavior towardsuch shells compared with controls. Reoccupation rate of emptiedexperimental shells was significantly higher than that of unmanipulatedempty shells. This indicates that shell stealing and nest takeover,followed by female expulsion, contribute to the reproductivesuccess of L. callipterus males. We also found that female matechoice reduces expulsion risk: females preferred to mate withlarge males, and male size correlated with dominance. We concludethat the limited availability of breeding substrate is a keydeterminant of both intrasexual competition and intersexualconflict in this species.     

Reproductive behavior and social organization of some Pacific hawkfishes (Cirrhitidae)

Ten species of Pacific hawkfishes occur in small social groups consisting of a single male and one or more females. Facultative monogamy occurred regularly in two obligate coral dwelling species while the remaining species were largely haremic. Males had larger body sizes than females within groups but not necessarily between groups. Males defended territories in which females had home areas. Females defended courtship sites within a male’s territory against neighboring females of the same social group. Courtship was paired and sequential within groups, commencing just prior to or after sunset and concluding at dusk. Spawning, when observed, was pelagic.     

Behaviour and breeding biology of the cooperatively breeding Grey-backed Fiscal Shrike Lanius excubitorius in Kenya

Grey-backed Fiscal Shrikes Lanius excubitorius were studied over a 2j year period near Lake Naivasha, Kenya. Grey-backs are cooperative breeders, with group sizes ranging from two to II. Only one pair breeds per group, with all other group members aiding in the rearing of young. The study population ranged from 64 to 79 individuals that occurred in from 13 to 16 groups. Non-breeding helpers made up to 66% of the population, with male helpers being more numerous overall than females. The annual survival rate was 65%, with no differences detected between the survival of males and females, or of breeders and helpers. Only male helpers were observed to acquire breeding status within the natal territory. Some female helpers acquired breeding positions in territories adjacent to their natal territories. Group territorial displays occurred throughout the year but were most pronounced prior to breeding during rainy periods. Reproductive success was very low, with only 14.5% of the recorded breeding attempts leading to fledged young. Large groups (four or more birds) had greater reproductive success than small groups, but many factors other than, or in addition to, group size may have influenced this pattern. The breeding male contributed the most food to the incubating female and to the nestlings. Male helpers and the breeding female contributed more to nestlings than did female helpers. Observations on the post-fledging period indicate that socialization and establishment of dominance may be of importance in group-living species living in a restricted ecological and social setting.     

Some evidence for different ecological pressures that constrain male and female body size

The dwarf morph of the Lake Tanganyika cichlid Telmatochromis temporalis uses empty snail shells as shelters and breeding sites in shell beds, in which many empty shells exist. Here, we assessed selection forces regulating body size in this fish. Field observations showed that large males tended to have a greater number of females in their territories, suggesting that sexual selection favours large males. Nonetheless, a transplant experiment suggested that male body size was limited by the ability to hide in empty shells from large piscivorous fish. In females, the number of ovarian eggs increased with body size, suggesting that fecundity selection favours large females. However, females are smaller than males. Females spawned eggs close to the apices inside the shells. The small space there would decrease the risk of egg predation by egg predators, and small body size of females may be a result of adaptation to spawn eggs in the small, safe spaces. This study provides support for the idea that male and female body sizes have been limited by different ecological pressures (predation on adult fish in males, predation on eggs in females), which has not been reported previously in any animal.     

Sex-biased movement in the guppy (<Emphasis Type="Italic">Poecilia reticulata</Emphasis>)

The movement strategies of birds and mammals are often closely linked to their mating system, but few studies have examined the relationship between mating systems and movement in fishes. We examined the movement patterns of the guppy (Poecilia reticulata) in the Arima river of Trinidad and predicted that sexual asymmetry in reproductive investment would result in male-biased movement. Since male guppies maximize their reproductive success by mating with as many different females as possible, there should be strong selection for males to move in search of mates. In agreement with our prediction, the percentage of fish that emigrated from release pools was higher for males than females (27.3% vs. 6.9%, respectively). Sex ratio was highly variable among pools and may influence a male's decision to emigrate or continue moving. We also detected a positive relationship between body length and the probability of emigration for males and a significant bias for upstream movement by males. Among the few females that did emigrate, a positive correlation was observed between body length and distance moved. Sex-biased movement appears to be related to mating systems in fishes, but the evidence is very limited. Given the implications for ecology, evolution, and conservation, future studies should explicitly address the influence of sex and mating systems on movement patterns.     

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.     

Associations between gastropod shell characteristics and egg production in the hermit crab Pagurus longicarpus

Summary Two populations (salt marsh and seagrass bed) of the hermit crab Pagurus longicarpus were sampled to examine associations between shell characteristics and egg production. Multivariate statistical analyses controlled for crab size and time of year, variables that otherwise could be confounded with shell effects. Although correlations between shell characteristics and reproduction existed in both populations, generalizations could not be made because associations varied within and between populations. Shell species was not associated with a female's reproductive state (i.e., whether or not she was barren when sampled) in either population. In the seagrass population, medium-large and large females occupying severely damaged or fouled shells were half as likely to be reproductive as females occupying other shell conditions. However, there was no association between shell condition and reproductive state for small and small-medium females in the seagrass population or among all females in the salt marsh population. In the seagrass population, small through medium-large reproductive females occupied shells more similar to their predicted shell size, regardless of whether the occupied shell was relatively small or large, than nonreproductive females. In contrast, relative shell sizes of reproductive and nonreproductive females were similar for large females in the seagrass population and all females in the salt marsh population. Clutch sizes were enhanced for females occupying Polinices duplicatus shells or shells larger than their predicted shell size in the seagrass population. Relative shell size also was associated with clutch size in the salt marsh population, but crabs occupying shells similar to their predicted shell size had the largest clutches.     

The effect of food abundance on territory size and population density of juvenile steelhead trout (<Emphasis Type="Italic">Oncorhynchus mykiss</Emphasis>)

Optimal territory size models predict a decrease in territory size with increasing food abundance. However, most of these models may not be applicable to juvenile salmonids in streams, because they defend contiguous territories at high densities. The optimal size of a contiguous territory is predicted to (1) be independent of food abundance when food is rare and (2) decrease only when food abundance is high enough to induce a reduction in territory size below the contiguous optimum. To test these predictions, we raised equal densities of juvenile steelhead trout in outdoor stream channels over a 32-fold range of food abundance in the absence of emigration for 25 days. Increasing competition for scarce food resulted in increasing mortality, higher willingness to emigrate, higher variance in body mass, lower growth, lower population density and lower biomass. The size of territories decreased with increasing local population density, and increased with increasing body size. However, territory size did not change with food abundance, a result consistent with the prediction of a contiguous territory size model. On average, total salmonid biomass increased 5.7 times in response to the 32-fold increase in food abundance. Our data provide strong support for an earlier quantitative relationship between the abundance of stream salmonids and their food.     

Community responses to experimental food depletion: resource tracking by stream invertebrates

The regulation of population processes for most organisms depends upon the strength and rate of feedback between resources and consumers. We conducted an experimental manipulation of leaf packs in stream channels, a patchy and ephemeral resource, which is consumed by a number of detritivorous invertebrates. We reduced the number of available food patches (red alder leaf packs) by half and then measured a variety of community responses, including emigration rate, aggregation on remaining food patches, decomposition rate of food patches, and species-specific differences in these responses. Replacement of removed leaf packs with polyester mimics resulted in no statistical difference in emigration rates or aggregation on remaining resources when compared to those removal channels without replacement. These results indicate that leaf packs are not used primarily for refuge. In the removal channels (including those with leaf pack mimics) emigration rate nearly doubled relative to control channels. Those invertebrates that did not emigrate from removal channels aggregated on remaining leaf packs, which led to more rapid decomposition of leaf packs relative to control channels. The increase in emigration rate only became apparent 2-3 days after the manipulation, presumably because animals colonized the remaining leaf packs and did not emigrate until food patch value per individual had been reduced by higher densities or due to increased discharge. Discharge through the channels increased slightly starting 3 days after the manipulation, resulting in increased emigration rates in all channels. Despite the increase in discharge, the effect of the manipulation remained strong. These results show that stream invertebrates colonizing leaf packs responded in predictable ways to a short-term reduction in food resources which would be adaptive in a system which is heterogeneous in space and time.     

Field evidence for bi-directional sex change in the polygynous gobiid fish Trimma okinawae

The social condition of bi-directional sex change in the gobiid fish Trimma okinawae was investigated at Akamizu Beach, Kagoshima, Japan. Social groups of T. okinawae usually consisted of a large male and one or more smaller females. The number of females in the group was positively correlated with male body size and groups were usually separated from each other by 1–3 m. In total, 22 instances of female-to-male sex change and three instances of male-to-female sex change were observed during the 16 months that social groups were monitored. Two individuals changed sex twice: female to male and back to female. Female-to-male sex change occurred when the male disappeared from a group. Either the largest remaining female changed sex to male or a large female from another group immigrated and changed sex to male. Larger individuals appear to benefit from becoming male because they can monopolize the breeding opportunities with several females, as reported in other protogynous fishes. Sex change from male-to-female only occurred when a solitary male joined another group as a subordinate. Mortality rates are high in these small fish, therefore joining another group and reproducing as a female is likely to increase the reproductive value of a solitary male.     

War and the evolution of belligerence and bravery

Tribal war occurs when a coalition of individuals use force to seize reproduction-enhancing resources, and it may have affected human evolution. Here, we develop a population-genetic model for the coevolution of costly male belligerence and bravery when war occurs between groups of individuals in a spatially subdivided population. Belligerence is assumed to increase an actor's group probability of trying to conquer another group. An actor's bravery is assumed to increase his group's ability to conquer an attacked group. We show that the selective pressure on these two traits can be substantial even in groups of large size, and that they may be driven by two independent reproduction-enhancing resources: additional mates for males and additional territory (or material resources) for females. This has consequences for our understanding of the evolution of intertribal interactions, as hunter-gatherer societies are well known to have frequently raided neighbouring groups from whom they appropriated territory, goods and women.     

Longevity factor in hominoid social organization

In most primate groups emigration of the maturing young of one or the other sex tends to serve as an incest avoidance mechanism. Among most primate species it is the males who change groups. This supports the theory that, in terms of reproductive success, males should compete for mates and females should compete for resources. In hominoids the combination of increased longevity and greater female discrimination in mate selection seems responsible for female emigration. This may relate to the high frequency of patrilocality and male control of resources among human groups.     

Spawning behavior and haremic mating system in the corallivorous butterflyfish,Chaetodon trifascialis, at Kuroshima Island, Okinawa

Territorial and spawning behavior ofChaetodon trifascialis were investigated on a small patch of reef at Kuroshima Island, Okinawa, Japan. Three males and 8 females inhabited the reef, each individual defending a territory against conspecifics of the same sex. Each male territory included 2 or 3 female territories. In the daytime, each male frequently visited the females living in its territory. At dusk in the full or new moon periods, courtship began within the female territories, pair spawning subsequently occurring within or near those territories. When a male actively courted a female in the territory of a second male, the latter male immediately chased off the intruder. Thus, mating occurred only between a male and females living in former's territory. This is the first report of a haremic mating system among butterflyfishes (Chaetodontidae).     

Intraspecific competition and the maintenance of monogamy in tree swallows

Intraspecific competition for access to breeding resources maylimit male mating success typically monogamous birds. We examinedthe potential for intraspecific competition to limit polygynyin tree swallows at Beaverhill Lake, Alberta, Canada. In thispopulation, polygynous males raisedmore fledglings than monogamousmales, and there was little or no cost to females from nestingpolygynously. Under these conditions one might expect polygynyto be more common than that observed(8% of males). We foundthat females were most aggressive toward conspecific intrudersearly in the breeding season. This aggression was associatedwith (1) females settling farther apart than expected underrandom settlement, (2) later settlement by secondary than bymonogamous females, and (3) no relationship between female settlementdate and male territory size instead of the negative correlationexpected if females settled randomly without competition. Earlyin the season, males also settled farther apart than expectedif they had settled randomly, and among males with two or morenest boxes on their territory, males with widely separated nestboxes were more likely to be polygynous. Monogamy is probablythe most common pairing association in this population becauseintraspecific competition for nest sites prevents most malesfrom gaining a territory with nest sites far enough apart topermit two females to breed without one female excluding theother. Females appeared to be defending an area surroundingtheir nest box to limit nest usurpation or intraspecific broodparasitism, rather than to limit any loss of male parental carefrom polygyny.