Alternative reproductive tactics in the territorial damselflyCalopteryx maculata: sneaking by older males

Summary We conducted daily censuses on a marked population of the damselflyCalopteryx maculata for two complete breeding seasons to document the reproductive tactics of individual males. Overall, 78% of the 600 males studied defended territories and 14% of those territorial males were also observed engaged in sneaking behaviour on some days. When sneaking, males did not defend territories but attempted to steal females from other males' territories. Sneakers were usually previously successful territorial males who emerged early in the season and began sneaking relatively late in their lives (Fig. 2). Thus the adoption of sneaking was conditional and related to both male age and population density. We suggest that sneaking, in this species, is a “make the best of a bad situation” tactic adopted when intense male-male competition forced older males to abandon territoriality. Thus, sneaking allowed males with declining resource holding potential to prolong their reproductive careers after they were no longer able to hold a territory.     

Sex ratios in high-density populations of the montane vole,Microtus montanus,and the behavior of territorial males

Summary Montane voles (Microtus montanus nanus, Rodentia: Muridae) were studied in unconfined populations and in an exclosure in open fields in Wyoming, USA. Field work involved capture-mark-recapture grids, tagging select individuals with irradiated wires, subcutaneous implantation of dye pellets, and behavioral observations.Territorial males were found in both high and low density populations. They were characterized as having scrotal testes, actively defended territories, residency over time, association with females, few scars, and scent-marking behavioral patterns.Males maintaining territories in high-density populations were nearly the only males in breeding condition. Five high-density populations were trapped in late fall, and age distributions were determined by an eye lens technique. There were two age cohorts of territorial males: those born earlier in the season and those which had overwintered. For each grid, the distribution of males by age was discontinuous at about 95 days. The ratio of older males to older females of comparable ages in these five populations varied between 1:2.1 and 1:3.8. The operational sex ratio was even more unequal.The observations of territoriality at such high densities did not support the hypothesis that territoriality and social rank are two poles of a behavioral continuum related to density.The system of polygyny found, wherein the male has a territory overlapping those of several females, does not neatly fit previous schemata considered favorable for the evolution of polygyny, but is considered adaptive in view of female movements and the labile response of female reproduction to the male.     

Lek structure and territoriality in the chryxus arctic butterfly,Oeneis chryxus (Satyridae)

Summary The territorial and mating systems of the satyrid butterfly Oeneis chryxus were studied for 3 years on individually marked populations in algonquin Provincial Park, Ontario. Males defended territories in a dry open forest clearing by means of chases and spiral flights with conspecifies. Defended areas did not contain food or oviposition sites required by females. Territories were not uniformly distributed over the study area; the highest density of territories was clustered over bare ground covered with sand and small stones rather than areas of vegetation. This pattern of distribution of territories occurred during each year of the study. The patterns of occupancy of territories showed a high degree of consistency from one day to the next, some males defending a given territory for as many as 11 consecutive days. Removal experiments showed that a surplus of non-territorial males did not exist. Vacant territories in the high-density area were absorbed by neighboring males; those in the low-density area remained empty. Male size was not correlated with territory position in the study are. Wing patterning (i.e. size of spots) differed between males occupying territories on sandy/stony versus vegetated areas. The territorial and mating systems of O. chryxus resemble lek polygyny, and are more similar to some vertebrate lek systems than to hilltopping or landmark territoriality.     

Social spacing in the temperate marine goby Coryphopterus nicholsi

Coryphopterus nicholsi is a temperate marine goby that occupies rock rubble in protected subtidal areas along the Pacific coast of NOrth America. In field populations, fish of all sizes and both sexes are found in the same habitat. All observed fish over 25 mm standard length defended space during both the reproductive and the non-reproductive seasons, with territory size being directly correlated with fish size. There was peripheral overlap of territories, particularly between adjacent adult conspecifics of different size classes, but also with smaller fish that had territories centered in the interstices of larger territories. However, zones of overlap were used at different times by space-sharing fish so that defended areas remained temporally discrete. Access to areas of overlap appeared to be determined by dominance rank. More dominant (usually larger) fish used areas of overlap at will; the defense and use of shared areas by less dominant fish was contingent upon the absence of larger fish. Social organization in C. nicholsi appears to be defined by a combination of territorial behavior and dominance relationships among adjacent individuals, permitting the coexistence of all members of the population under circumstances of limited habitat availability.     

Patterns of use of space in a guild of territorial reef fishes

There exists on Heron Reef, at the southern end of the Great Barrier Reef, an 8-species guild of ecologically very similar, territorial, herbivorous reef fishes. All individuals of these species maintain territories on rubble substrata throughout juvenile and adult life. Territories are defended from all other guild members. Three rubble patches, each containing residents of 3 guild species: Pomacentrus apicalis, P. wardi, and Abudafduf lachrymatus, have been monitored for 12 to 18 months. This paper examines the patterns of use of space as shown by the antecedent and subsequent histories of sites chosen by 43 new colonists, and sites vacated through the loss of 34 residents on the rubble patches. In addition, variation through time is examined in the total amount of space held in territories on each rubble patch. The 3 species show similar preferences for space as colonists, although adult and juvenile colonists behave differently. Numbers of colonists detected on rubble patches are not proportional to the resident populations of the 3 species. Residents of the 3 species are equal in their abilities to enter spaces vacated through mortality, although they differ slightly in methods used to enter sites. They do not respond preferentially to sites previously occupied by any particular species. P. wardi shows a higher rate of mortality than the other species, and residents of this species are more often dislodged by new colonists. The total amount of space held on any rubble patch did not vary during the year from September, 1972 to October, 1973. The data are interpreted as supporting the hypothesis that the 3 species do not differ in their requirements for space on rubble patches, and that they are in competition for a short supply of such space. Differences exist in their strategies for obtaining and holding such space. These differences are important for explaining the continued presence of P. wardi on rubble patches. P. apicalis and a. lachrymatus are both specialists in holding territories on the upper reef slope. No differences have been detected in their requirements or competitive abilities. P. wardi is a fugitive species on the upper reef slope, coexisting because it maintains a refuge from competition by occupying some un-preferred sites off rubble patches.     

Seasonal variation in territorial response,and other aspects of the ecology of the Australian temperate pomacentrid fish Parma microlepis

The Australian temperate pomacentrid fish Parma microlepis Gunther maintains permanent, interspecifically defended territories. Breeding occurs in late spring (October to December), and juveniles after a pre-settling period of 4 to 6 weeks take up territories in the adult habitat. P. microlepis feed in their territories on benthic algae and invertebrates. Density of fish is significantly correlated with the availability of suitable shelter. There is a variation in the territorial response directed at different species, and a significant correlation exists between intensity of response to a species and its similarity to P. microlepis in use of living space. No significant correlation exists between intensity of response and dietary similarity. P. microlepis defends larger areas in the breeding than in the non-breeding season against those species which represent a possible threat to its breeding success. It shows a reduced tendency to attack juvenile conspecifics at the time of year when they would be settling from the plankton. Responses to species not likely to interfere with breeding success do not vary through the year. P. microlepis is thus seen to have complex territorial responses, specific to the species of intruder and time of year. In this it is quite similar to other territorial pomacentrids (chiefly tropical) for which information exists.     

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.     

Pair territoriality of wintering stonechats : behaviour,function and hormones

Migratory stonechats (Saxicola torquata) spend the winter in the northern Negev desert in Israel. After arrival in October most birds defend territories throughout the winter either as heterosexual pairs or as single birds. In this population we examined: (1) aggressive behaviour of pairs; (2) the functional significance of pair territoriality; and (3) hormonal correlates of territorial behaviour. Pairs were aggressive towards both conspecific intruders and stonechat dummies, particularly in autumn when territories were established. Males were more aggressive than females and aggression was primarily intrasexual. More frequent than intraspecific interactions were interspecific encounters, with 94% involving the local mourning wheatear (Oenanthe lugens). With one exception these encounters consisted of attacks of mourning wheatears against stonechats, mainly males. Two functions of winter territoriality of migrant pairs are proposed: (a) a reduction of individual defense costs associated with the maintenance of a territory; (b) an increase of the net vigilance to reduce the risk of attacks by competitors or predators. The levels of testosterone (males and females) and estradiol (females only) were close to or below the lower detection limit of the assay throughout the winter. Therefore, high levels of gonadal steroids are not required for pair formation and aggression in the non-reproductive context. The results indicate that both the function and the proximate control mechanisms of territoriality during the non-breeding season in winter are different from those during the breeding season in spring.     

Territorial behavior of the black and yellow rockfish and gopher rockfish (Scorpaenidae,Sebastes)

Tagged black-and-yellow rockfish Sebastes chrysomelas and gopher rockfish S. carnatus were observed in situ for 1 yr at 3 study sites off California, USA in order to determine the nature of their home ranges and territories. Three basic patterns of space use were identified: (1) a shelter hole located within a larger area of activity (in which the fish presumably fed, since its size increased with size of fish and decreased with prey density); (2) discontinuous home ranges, in which the shelter holes and feeding areas were separte; and (3) dispersed home ranges with little or no exclusive area. Fish in the first category were regarded as territorial, since their home ranges were defended and relatively exclusive. Because individuals did not patrol borders, however, territories often overlapped at their peripheries. In addition, some territory holders appeared to tolerate the sheltering of other fish within their territories, as long as these intruders left the territory during periods of activity. Some fish with discontinuous home ranges (commuters) may have been territorial, but others may have been forced to occupy suboptimal feeding areas. Fish in the third category (floaters) were regarded as nonterritorial; their activities were apparently limited to areas where they could escape the aggression of territory holders. The varying patterns appeared to develop through defense and accommodation during the turnover of individuals, with each fish securing shelter and feeding areas as these became available.     

Correlation between nectar supply and aggression in territorial honeyeaters: causation or coincidence?

Summary A fundamental prediction of food-based economic models of territoriality is that animals will not defend territories if food is so abundant that defense will not improve access to food. Several studies of nectar-feeding birds support this prediction, with territoriality being rare or absent in years when nectar was particularly abundant. However, these results could potentially be an artefact of changes in bird density with nectar availability, and in at least some cases the correlations between territory defense and nectar availability could be purely coincidental. This paper reports the first experimental test of whether cessation of territory defense in nectar-feeding birds results from a direct response to abundance of nectar. New holland honeyeaters Phylidonyris novaehollandiae and white-cheeked honeyeaters P. nigra show pronounced changes in their levels of territorial aggression over the 7–8 months that they breed. These changes are predictable from economic considerations in that the birds are least aggressive in the months when nectar is extremely abundant. I tested whether the birds were responding to changes in nectar availability by providing sugar-water feeders at neutral locations that were easily accessible to territory holders, but far enough away from territories that intrusion rates were unaffected. I tested for responses at two time scales feeders were put out for 48-h periods in 1987, and were left out continuously from January to October 1988. The only effect was that territory holders visited feeders instead of flowers when floral nectar was scarce. They continued to defend their territories aggressively at those times, showed seasonal changes in aggressiveness similar to birds on a site without feeders, and did not shift their territories toward feeders. I conclude that the observed changes in aggressiveness are not responses to changes in nectar availability, and suggest alternative explanations.     

Long-term mate and territory fidelity in neotropical buff-breasted wrens (Thryothorus leucotis)

Among sedentary species, mate and site fidelity may be imposed by year-round territoriality and high annual survivorship, both of which theoretically lead to few opportunities to move between mates and breeding territories. We examined mate and territory fidelity in sedentary neotropical buff-breasted wrens (Thryothorus leucotis) in relation to the opportunities to divorce (vacancies on neighbouring territories), breeding experience and site of origin (within vs outside local breeding population). Overall, buff-breasted wrens showed high rates of mate and territory fidelity between years, with individuals significantly more faithful to their territories than to their partners. Mate and territory fidelity was not forced upon buff-breasted wrens, as most individuals had opportunities to switch mates and territories. However, this was not uniform among all individuals. Individuals that had bred together displayed 100% mate and territory fidelity despite the fact that males and females in almost all experienced pairs had opportunities to switch mates and territories. Local recruits (philopatric offspring that acquired territories in the local breeding population) also displayed mate and territory fidelity, but fidelity occurred because recruits rarely had the opportunity to divorce prior to initiating their first breeding attempt. By contrast, almost half of all territorial individuals that originated outside the local breeding population (immigrants) had opportunity to divorce, with approximately two thirds of them switching mates and territories. Divorces occurred within the first 5 months of pairing, suggesting that immigrants may have sampled mates and territories before forming permanent partnerships.     

Territory and group sizes in Eurasian beavers (Castor fiber): echoes of settlement and reproduction?

According to current theories of territoriality, an animal is expected to defend the smallest area that can provide resources for maximisation of reproduction, known as the economically defendable area. In group territorial species however, the strategies behind resource defence are likely to be more complex with corporate territoriality, cooperative breeding, delayed dispersal and intra-group competition all potentially playing a role. Here we examined group territoriality in a social herbivorous rodent, the Eurasian beaver Castor fiber. Beavers in our study do not inhabit economically defendable territories. Instead the sequence of arrival of pairs into unoccupied areas seems to play a more important role in determining the size of the territory, whereas group size is determined by past reproductive success. We argue that the settlement pattern and reproductive history have a lasting impact in the territorial system of beavers due to a combination of the low adult mortality, high dispersal costs, and avoidance of resource depletion.R.D. Campbell and F. Rosell contributed equally to this work     

Satellites and sneakers: opportunistic male breeding tactics in pupfish (Cyprinodon pecosensis)

Summary Manipulations of population density, availability of oviposition substrate, and size of breeding habitat affected the proportion of pupfish (Cyprinodon pecosensis) males adopting territorial and satellite breeding tactics. Satellites occupied stations above breeding substrates defended by territorial males. Both territorial and satellite males developed a bright blue breeding coloration. A third breeding tactic, sneak-spawning, was occasionally observed. Sneak-spawning males retained the cryptic female coloration and occasionally spawned on territories. The lower reproductive success of satellites and sneak-spawning males suggested that both are conditional breeding tactics adopted by competitively inferior males. Satellites were present in all treatments that favored a territorial breeding system, but were absent when the breeding system was a dominance hierarchy. Satellites were associated with territorial males that had higher reproductive success and larger territories than males without satellites. Satellites functioned as parasites rather than as mutualists, since they disrupted spawnings and stole copulations from territorial males. The occurrence and frequency of the conditional breeding behaviours in pupfish represent facultative responses of males to changes in the intensity of competition for breeding sites and females.     

Space use and social structure in meadow voles,Microtus pennsylvanicus

Summary Free-ranging, sexually mature meadow voles (Microtus pennsylvanicus) were tracked by using radiotelemetry from June through August in Front Royal, Virginia, U.S.A. Estimates of intraspecific spacing were derived from the concurrent movements of up to 16 voles. Positions were recorded hourly for 24 h, twice per week. A total of 16 male and 15 female voles were studied during sixteen 24-h sessions.The daily ranges of males (192.3±109.7 m²) were larger and more variable than those of females (68.6±39.4 m²). Males also changed locations more frequently (Fig. 2).Adult females usually maintained territories free of other females; males overlapped considerably among themselves (Fig. 2). Males temporarily moved into the areas occupied by estrous females, indicating intrasexual competition among males for access to receptive females (Fig. 3). M. pennsylvanicus appears to be promiscuous, is socially organized into territorial, maternal-young units during the breeding season, and fits the female territorial model of population regulation.     

Responses to conspecific scent marks and the ontogeny of territorial marking in immature terrestrial salamanders

We investigated the ontogeny of responses to scent marks in immature terrestrial salamanders (Salamandra lanzai) that inhabit the southwest Alps. In this species, sexual maturity is usually reached at about 8 years, and adults exhibit territoriality. One should expect territoriality to take place largely before the acquisition of sexual maturity if sexual competition is not the main force driving territoriality. However, both the difficulties for inexperienced animals to find a suitable territory and size-related competition may delay the acquisition of territoriality in juveniles. We performed choice tests with juveniles belonging to two age groups (1±1 years old versus 4±1 years old). Each focal animal was offered, in random order, the choice between two shelters, one without scent and the other containing one of the following scents: own, same-aged animal, a juvenile belonging to the other age group, or adult female. We also performed choice tests with adult females for which the scents of two juveniles belonging to a different age group were successively offered in a random order. Older juveniles were strongly attracted toward their own shelter and mostly avoided the shelters that contained the scent of juveniles of about the same age and of adult females. Adult females avoided the scents of older juveniles but not younger juveniles. These results therefore suggest that older juveniles use territorial marking. Conversely, young juveniles behaved randomly with respect to their own scents and to those of a same-aged juvenile, and they were significantly attracted towards the odor of an older animal, especially adult females. Both older juveniles and adult females displayed a random behavior toward the scents of young juveniles. Our results suggest that young juveniles do not defend territories but use spaces occupied by older individuals.Communicated by A. Mathis     

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.     

Population and individual consequences of breeding resource availability in the European bitterling (Rhodeus amarus)

Resource availability may affect both individual fitness and population demography and the effects can interact. We used two experiments to test how breeding resource abundance and its spatial distribution, combined with female abundance, affected male reproductive behavior, population spawning rate, and embryo development and recruitment in the European bitterling (Rhodeus amarus), a small cyprinid fish that lays its eggs in living unionid mussels. In the first experiment, we found that at the population level the abundance of breeding resources (freshwater mussels) was more important for bitterling recruitment than resource spatial distribution (clumped or regular). In contrast at the individual level, (variability in reproductive success) the spatial distribution of resources was more important, but only when resource abundance was not limiting. Territorial males obtained almost exclusive access to fertilizations when resources were abundant and distributed regularly, but were unable to defend large clusters of resources (when rival abundance was always high) and abandoned territoriality. Surprisingly, territorial males remained aggressive and successfully defended their territories when resources were grouped into a single cluster, but at a low abundance. In the second experiment, more rapid embryo development and larger juvenile body size at the end of the growing season was detected at high resource abundance and low female abundance, indicating that early hatched juveniles survived better and hence investment in offspring production early in the season yields a higher fitness pay-offs. The abundance of females in spawning condition was the best overall predictor of the intensity of male reproductive behavior in both experiments.     

Plasticity in spawning visits of female damselfish, Stegastes nigricans : effect of distance to mates

Both sexes of the damselfish Stegastes nigricans hold individual territories in which they feed on filamentous algae. At dawn, females visit males' territories to spawn, and the males guard the eggs until hatching. We examined how females' spawning behavior varied according to the distances to their mates. Females usually mated with a single male per morning. The distance to the territory of a mate (0.7–12.8 m) did not affect a female's total spawning time per morning, but affected the number of her spawning visits with that male. Females made many repeated spawning visits when spawning with males at short distances, while they spawned the entire clutch in one visit when spawning with males at long distances. This plasticity in female behavior appears to be related to two costs during the spawning visits: (1) intrusions by other fish to feed on algae in the female's territory during her absence, which may cause the female to return repeatedly to her territory for defense, and (2) attacks on the female by other territorial fish, which increased with the distance to their mates' territories. To minimize the sum of both costs, females should change the number of spawning visits depending on the distance to the males' territories. Received: 30 September 1996 / Accepted after revision: 17 March 1997     

Host plant density and territorial behavior of the seed bug,Neacoryphus bicrucis (Hemiptera: Lygaeidae)

Summary Adults of the seed bug, Neacoryphus bicrucis, are most numerous in large host plant patches and where host plants are most dense within patches. Males patrol small territories composed of host plants and expel other males and unreceptive females. Larger males aggressively exclude smaller males from high quality territories where both host plants and females are most dense. Consequently, large males mate more frequently than small males. Males remain for longer time in areas where host plants are more dense. However, mating experience influences tenure within a host plant patch such that males tend to remain where they have recently mated. Courtship is aggressive and male territorial behavior is similar to courtship behavior. Preference for territories in areas of high host plant density appears to have been selected to increase the rate of encounter with females which preferentially oviposit in areas of high host plant density.     

Territory quality and territorial behavior in two sympatric species of hummingbirds

Summary Changes in territorial behavior of blackchinned hummingbirds (Archilochus alexandri) in response to experimental changes in territory quality were investigated using artificial feeders and simultaneous, pair-wise observations of owners. Some of the responses of A. alexandri were similar to those documented by a previous study of the Anna's hummingbird (Calypte anna): A. alexandri defending rich territories spent more time on their territories, encountered a greater percentage of intruders, and chased more intruders than did A. alexandri defending poor territories. In contrast to C. anna, A. alexandri supplemented chases with energetically inexpensive threat vocalizations more extensively when territory quality was increased. This difference may be related to A. alexandri's more tenuous control of rich territories. When both species were observed on very poor territories, A. alexandri chased a greater percentage of intruders, consumed a greater proportion of available food, and obtained a greater net energy gain from their territories. When observed simultaneously on territories with ad lib food, both species defended highly exclusive territories but A. alexandri suffered higher intrusion pressure and spent more time and energy in defense. These interspecific differences in territorial behavior may facilitate sympatric coexistence of the two species through aggressive partitioning of flower patches according to productivity: the greater payoff received by C. anna on rich territories and A. alexandri on poor territories should favor a corresponding monopolization of rich areas by C. anna and poor areas by A. alexandri.