Early comers occupy popular territories in a lycaenid butterfly,Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae)

Territorial activities of a lycaenid butterfly, Chrysozephyrus smaragdinus, were observed throughout its adult season to investigate the territorial dynamics. The territories occupied earlier in the season were abandoned later than later‐occupied territories, indicating that the site preference of male C. smaragdinus was largely maintained during the adult season. Three indexes of territory quality based on animal preference, occupation duration (OD), initial occupation date (IOD) and the number of contests (NOC), were highly correlated. Therefore, these preference‐based indexes are consistent and reliable, although they are indirect estimates of benefits obtained from territories. Males arriving at territory sites early in the season were more likely to hold territories of better quality when NOC was used as the index of territory quality. The positive relationship between territory quality and male arrival date is congruent with the result of a previous study showing that early comers tend to hold a territory whereas late comers do not. However, male arrival date was not related to territory quality when OD or IOD was used as an index of territory quality. One possible reason for this apparent discrepancy is that the coefficient of variation of NOC is larger than those of OD and IOD, and therefore NOC could evaluate territory quality more quantitatively than the other two indexes. To date, studies of territoriality in butterflies have not considered the quality of territories. More studies considering the quality of territories will provide new insights into butterfly territoriality.     

Effect of competitor abundance on feeding territoriality in a grazing fish,the ayuPlecoglossus altivelis

The grazing fish, ayu,Plecoglossus altivelis Temminck & Schlegel, establishes feeding territorialiry during the young stage. The population density fluctuates from year to year by more than a hundredfold, but the determinant of territory size is less well known. The feeding territoriality of ayu was examined under simulated habitat conditions where fish density was manipulated and food resources were renewable. Fish competed for algae attached to the substrata and were divided into residents with territories, and floaters without territories. By experimental alteration of fish density the number of residents increased with density and rerritory size decreased with density. Floaters intruded into territories in a school to feed on algae, which induced overt aggression of the resident and reduced the productivity of algae growing there. Both the intruding frequency of floaters over territorial areas and their feeding pressure on algae increased at higher floater density. Floaters functioned to shift cost-benefit relationships for various territory sizes. They acted as food competitors to restrict territory size below a maximum through competitive interference. Although the growth rate of residents was inversely related to fish density, residents grew faster than floaters in each group. Under a given set of competitor abundances, economic defensibility determined territory size.     

Harem structure and female territoriality in the dwarf hawkfish <Emphasis Type="Italic">Cirrhitichthys falco</Emphasis> (Cirrhitidae)

We investigated the territoriality and the spatial and mating relationships of the haremic hawkfish, Cirrhitichthys falco, on a reef off Kuchierabu-jima Island in southern Japan. Each individual maintained a territorial home range which was defended against same-sex conspecifics at the boundary of the home range. The territory of each male encompassed the territories of 2–3 females, allowing the male to completely monopolize mating opportunities with those females. Based on our observations, we classified the harem type of C. falco as a territorial female type. Large juveniles maintained independent home ranges outside the female territories. In contrast, small juveniles were allowed to cohabit within the territory of an adult female. Stomach contents analysis revealed that the smallest size class of C. falco fed primarily on copepods. In contrast, all other size classes fed primarily on decapods. Together, these results suggest that female territoriality plays an important role in defending food resources.     

Territory quality determines social group composition in Ethiopian wolves Canis simensis

1. We contrast the value of four different models to predict variation in territory size as follows: resource density (the ideal free distribution), population density, group size and intruder pressure (relative resource-holding potential). In the framework of the resource dispersion hypothesis, we test the effect of resource abundance and spatial variation in resource distribution on the age/sex composition of social groups. 2. We explore these drivers of territory size and group size/composition in Ethiopian wolves Canis simensis in the Bale Mountains, Ethiopia, using fine-scale distribution maps of their major prey species based on satellite-derived vegetation maps. 3. The number of adult males is correlated with territory size, while prey density, wolf population density and intruder pressure are not associated with territory size. On average, each additional adult male increases territory size by 1.18 km(2). 4. Prey abundance increases with territory size (average biomass accumulation of 6.5 kg km(-2)), and larger territories provide greater per capita access to prime foraging habitat and prey. 5. The age/sex composition of wolf packs is more closely related to territory quality than territory size. Subordinate adult females are more likely to be present in territories with greater proportions of prime giant molerat Tachyoryctes macrocephalus habitat (i.e. >80% of Web Valley territories and >20% in Sanetti/Morebawa), and more yearlings (aged 12-23 months) occur in territories with greater overall prey biomass. 6. Wolf packs with restricted access to good foraging habitat tend to defend more exclusive territories, having a lower degree of overlap with neighbouring packs. 7. The greater per capita access to prey in large groups suggests a strong evolutionary advantage of collaborative territorial defence in this species, although the relative costs of territorial expansion vs. exclusion depend upon the spatial distribution of resources. We propose a model whereby territory size is determined by the number of adult males, with the presence of subordinate females and yearlings dependent on the quality of habitat, and the abundance and distribution of prey, incorporated within territory boundaries.     

INTRASEXUAL COMPETITION ALONE FAVORS A SEXUALLY DIMORPHIC ORNAMENT IN THE RUBYSPOT DAMSELFLY HETAERINA AMERICANA

I studied the sex-limited red spots on the wings of male rubyspot damselflies (Hetaerina americana) in relation to territoriality and fitness in the wild. Both observational and experimental (wing spot manipulation) studies indicated that wing spots were selected through competition among males for mating territories, not through female choice or direct competition for females. Males with naturally or artificially large wing spots were more successful at holding territories and consequently mated at higher rates than males with relatively small wing spots. In contrast, sexual selection on male body size appeared to operate among nonterritorial males at the clasping stage of the mating sequence, perhaps because larger males were better at clasping females forcibly. Of four models proposed to explain the evolution of ornaments through territory competition, only the agonistic handicap model makes predictions consistent with the results of this study.     

The Behavioral Ecology of Libellula luctuosa (Burmeister) (Anisoptera: Libellulidae): I. Temporal Changes in the Population Density and the Effects on Male Territorial Behavior

A single population of a common pond dragonfly, Libellula luctuosa, was studied at a site where the density of males increased dramatically during the breeding season. Early in the summer one active male was found on each territory on the pond. Satellite males were only occasionally found on the territories. Later in the season the number of males per territory increased so that two or more males simultaneously defended on many of the territories, and several satellite males occupied each of the territories. The number and rate of female visitations per day did not change over the summer. These factors resulted in a change in the operational sex ratio with variations in male density. Male behavior was also altered with increasing population density. As male density increased, males were less likely to be seen perching on their territories and more likely to be seen performing aggressive acts such as chasing nearby territorial males and chasing intruders. At high male density, the duration of territorial behaviors was shorter than at low male density. Thus, the percent of a time budget spent in any one activity did not change despite the change in number of males present. Male activity in L. luctuosa is not strictly determined by the opportunity for aggression. Costs of aggression associated with territoriality are minimized by maintaining flexible territorial behaviors.     

An Extreme Case of Interspecific Territoriality: Male Anthidium manicatum (Hymenoptera,Megachilidae) Wound and Kill Intruders

Anthidium manicatum males defend patches of flowering plants as mating territories and copulate with females that enter the territory to obtain nectar or pollen. Territorial males defend their territories not only against conspecific males but also against many other insect species that attempt to utilize the flowers inside the territory. In our study area, a territorial male defended its territory against conspecific males on average only twice per h but it attacked other species of insects on average 70 times per h. Territorial males distinguished between different species of intruders and attacked them unequally. During the hour following experimental removal of the territory holder the number of intruders of some insect species tripled but the number of intruders of those species that were rarely attacked remained approximately the same. A. manicatum males, but not females, bear long spines on the last two segments of the abdomen. Territorial males rammed intruding insects at high speed and shortly before the moment of impact they curved the abdomen forward to hit with the spines. Intruders could be seriously damaged or killed by these attacks. Abdominal spines of A. manicatum males may have evolved as weapons to increase the effectiveness of interspecific territoriality.     

Pair territoriality in the longnose filefish,oxymonacanthus longirostris

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

Why are Male Columbian Ground Squirrels Territorial?

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

Predation Risk,Resource Quality,and Reef Structural Complexity Shape Territoriality in a Coral Reef Herbivore

For many species securing territories is important for feeding and reproduction. Factors such as competition, habitat availability, and male characteristics can influence an individual’s ability to establish and maintain a territory. The risk of predation can have an important influence on feeding and reproduction; however, few have studied its effect on territoriality. We investigated territoriality in a haremic, polygynous species of coral reef herbivore, Sparisoma aurofrenatum (redband parrotfish), across eight reefs in the Florida Keys National Marine Sanctuary that were either protected or unprotected from fishing of piscivorous fishes. We examined how territory size and quality varied with reef protection status, competition, predation risk, and male size. We then determined how territory size and quality influenced harem size and female size to understand the effect of territoriality on reproductive potential. We found that protected reefs trended towards having more large predatory fishes and that territories there were smaller but had greater algal nutritional quality relative to unprotected reefs. Our data suggest that even though males in protected sites have smaller territories, which support fewer females, they may improve their reproductive potential by choosing nutritionally rich areas, which support larger females. Thus, reef protection appears to shape the trade-off that herbivorous fishes make between territory size and quality. Furthermore, we provide evidence that males in unprotected sites, which are generally less complex than protected sites, choose territories with higher structural complexity, suggesting the importance of this type of habitat for feeding and reproduction in S. aurofrenatum. Our work argues that the loss of corals and the resulting decline in structural complexity, as well as management efforts to protect reefs, could alter the territory dynamics and reproductive potential of important herbivorous fish species.     

Conflicting Territory Use in Males and Females of a Monogamous Ungulate,the Kirk's Dikdik

In monogamous mammals it is often unclear why males do not defend larger territories to attract more than one female. I investigated the territoriality of the monogamous Kirk's dikdik, Madoqua kirki, a dwarf antelope, in which food resources increase with territory size and some males defend enough resources for more than one female. Yet, all males are paired monogamously. When males were removed from small territories, their female partners spent more time outside of their territories than females in large ones. When females were removed, their male partners almost never left. Pairs in small territories spent more time together than pairs in large ones. Paired males left mostly together with their females, apparently not on their own initiative. Presumably because females in small territories left more often, their males spent more time outside in the female's company than males in large territories. I argue that males in smaller territories can keep better track of their females and that they can effectively reduce their females' time outside. Male intrusion pressure was unrelated to territory size, but it increased in the presence of unguarded females. If large territories decrease the ability to mate guard, and if unguarded females attract competing males, then defending large territories may be uneconomical, even it they could attract more than one female. On the other hand, territories must be large enough to satisfy the requirements of a single female.     

Survival and population persistence in the critically endangered Tuamotu kingfisher

A wide range of threats affect populations of Pacific island birds and conservationists have been challenged to identify factors upon which to focus management. The Tuamotu kingfisher (Todiramphus gambieri) is one of the most endangered vertebrate species in the world, yet little has been published about basic biology or causes of the population decline. We used 4 years of mark-resight and territory resource information to model survival in juvenile and adult Tuamotu kingfishers. Annual survival of adult males (ϕ = 0.77) was similar to congeneric species, whereas survival of adult females (ϕ = 0.40) and juveniles (ϕ = 0.12) was much lower. The best-supported survival model indicated adult female survival was positively related to territory size, whereas adult male survival was negatively related to atoll forest within territories. We used parameter estimates from survival models in a life-stage simulation analysis to evaluate the relative influence of vital rates and territory habitats on population processes. Results indicated that adult female and juvenile survival had the greatest impact on Tuamotu kingfisher populations, accounting for 58% and 32% of variation in the finite rate of population increase, respectively. Nocturnal predation by rats (Rattus spp.) on incubating kingfishers may explain the lower survival of females than males, whereas juvenile birds may be especially vulnerable to predation by cats (Felis catus). Thus, conservation management for Tuamotu kingfishers should include use of metal guards on cavity trees to protect nests and incubating females, and control predator access during fledging periods. © 2012 The Wildlife Society.     

Determinants of territory size in the pomacentrid reef fish,Parma victoriae

Summary Factors governing the size of territories defended by the pomacentrid reef fish, Parma victoriae, were investigated, prompted by contradictory predictions in the literature concerning the effects of food supply and competitors. Observations were carried out over the non-breeding period (March–October) on a medium density population in which territories were partially contiguous. The territory size of adult fish varied between 3 and 26 m², and was inversely correlated with local densities of conspecifics. The same range in territory size was found for both males and females, which did not differ in the time they spent on territory defence and foraging activities. No correlation existed between territory size and the abundance of algal food, body size, age or time spent on territory defence. Also, there was little variation in territory size over time, despite seasonal changes in the abundance of food algae.Experimental reduction of food supplies on isolated territories of males and females had no effect on territory size. In a higher density habitat an experiment was carried out in which population density and food abundance were simultaneously manipulated. This showed that territory size was primarily determined by intraspecific interactions, as territories exhibited considerable increases in size upon removal of neighbours. No changes in the size of defended areas resulted from either artificial increases or decreases of food levels. There were also no changes in the time spent on defence of territories, foraging time or feeding rates associated with food manipulations or territory expansion, which suggested that food was not a limited resource. This conflicted with current theories proposed to explain territory defence and expansion. It is hypothesized that intraspecific interactions constrain territory size well below the optimum in terms of the abundance of preferred food algal species.     

Delayed maturation in female bank voles: optimal decision or social constraint?

1. Density, maturation and survival of female bank vole ( Clethrionomys glareolus ) in the northern taiga of Finnish Lapland were studied using long-term capture–mark–recapture data from two large grids, one food-addition grid and one control grid, in 1982–94.
2. The density on the food grid was consistently higher than the density on the control grid.
3. Females born early in the breeding season usually matured, except at very high densities. Those born later in the summer season commonly delayed maturation to the following spring.
4. Winter survival of sub-adult (having delayed maturation) females was significantly higher than survival of adult (breeding) females. However, empirical values of sub-adult and adult survival, as well as difference between them, were not consistent with survival values assumed in theoretical models on optimal deferred breeding.
5. There was a density-dependent relationship between the maturation rate of young voles and the density of already established breeding females (both bank voles and all Clethrionomys together; C. rutilus and C. rufocanus occasionally occurred on the study grids). This density dependence was different for the two grids (weaker on the food-addition grid).
6. These findings are discussed within an evolutionary context: we have, on the basis of these findings, no evidence suggesting that the observed delayed maturation represents an evolutionary optimal strategy. Rather, there is evidence suggesting that the delay is due to social constraints.     

Prey availability affects territory size,but not territorial display behavior,in green anole lizards

The availability of food resources can affect the size and shape of territories, as well as the behaviors used to defend territories, in a variety of animal taxa. However, individuals within a population may respond differently to variation in food availability if the benefits of territoriality vary among those individuals. For example, benefits to territoriality may differ for animals of differing sizes, because larger individuals may require greater territory size to acquire required resources, or territorial behavior may differ between the sexes if males and females defend different resources in their territories. In this study, we tested whether arthropod abundance and biomass were associated with natural variation in territory size and defense in insectivorous green anole lizards, Anolis carolinensis. Our results showed that both male and female lizards had smaller territories in a habitat with greater prey biomass than lizards in habitats with less available prey, but the rates of aggressive behaviors used to defend territories did not differ among these habitats. Further, we did not find a relationship between body size and territory size, and the sexes did not differ in their relationships between food availability and territory size or behavioral defense. Together, these results suggest that differences in food availability influenced male and female territorial strategies similarly, and that territory size may be more strongly associated with variation in food resources than social display behavior. Thus, anole investment in the behavioral defense of a territory may not vary with territory quality.     

Influences on variation in territorial tenures of male white-faced dragonflies (Leucorrhinia intacta) (Odonata: Libellulidae)

Some individuals in species with extended periods of territorial occupancy may change territory locations within a single bout of territorial activity. Length of occupancy of mating territories among males in a local population of white-faced dragonflies (Leucorrhinia intacta) varied from more than 6 h to 15 min or less. Males with short tenures often established territories in several locations on the pond during a day. Several hypotheses have been proposed to explain shifting territorial sites rather than remaining in a single site during one bout of territoriality. We attempted to test the hypothesis that males shift to leave low-quality sites. Site quality may be affected by costs of defense in relation to intruder rate and the mating benefits of holding the territory. To test whether variation in these possible effects of benefits and costs of territoriality influenced tenure, we manipulated local quality of oviposition substrate and perch density. The quality of oviposition substrate, but not perch density, influenced both potential benefits and costs of territoriality. Female density was higher in areas with good substrate, but so were rates of males intruding into the territories, rates of chasing by territorial males, and local density of territorial males. More matings occurred in areas with good substrate, but among males with tenures of 15 min or more, mating success per male and tenure lengths did not differ statistically among treatments. Defense costs were low for all treatments and perhaps were not an important influence on tenure duration. Territorial males in this population probably adjusted local density to expected mating success by initial choice of site rather than by varying tenure length. Variation in tenure length at a site resulted, in part, from stochastic external factors, such as predation attempts.     

Winter territoriality and its implications for the breeding ecology of White-throated Dippers Cinclus cinclus

Capsule: Pairs of White-throated Dippers Cinclus cinclus which defended winter territories bred earlier than non-territorial individuals, but there was no difference in reproductive success.

Aims: The effect of winter territoriality on breeding ecology has rarely been studied in resident birds. We carried out a preliminary investigation of whether winter territorial behaviour and territory size affect the timing of reproduction, breeding territory size and reproductive success in a riverine bird, the White-throated Dipper.

Methods: We monitored an individually marked population of White-throated Dippers in the UK. Wintering individuals were classified as either territorial or ‘floaters’ according to their patterns of occurrence and behaviour, and their nesting attempts were closely monitored in the subsequent months. Winter and breeding territory sizes were measured by gently ‘pushing’ birds along the river and recording the point at which they turned back.

Results: All birds defending winter territories did so in pairs, but some individuals changed partners before breeding. Territorial pairs that were together throughout the study laid eggs significantly earlier than pairs containing floaters and those comprising territorial birds that changed partners. However, there were no significant differences in clutch size, nestling mass or the number of chicks fledged. There was no relationship between winter territory length and lay date or any measure of reproductive success, although sample sizes were small. Winter territories were found to be significantly shorter than breeding territories.

Conclusion: Winter territoriality may be advantageous because breeding earlier increases the likelihood that pairs will raise a second brood, but further study is needed. Territories are shorter in winter as altitudinal migrants from upland streams increase population density on rivers, but this may also reflect seasonal changes in nutritional and energetic demands.     

Effects of Exogenous Androgen and Antiandrogen on Territorial and Nonterritorial Red-winged Blackbirds (Aves: Icterinae)

We investigated the role of testosterone (T) in territory establishment and maintenance in male red-winged blackbirds (Agelaius phoeniceus) by implanting exogenous T or pharmacological agents that block the action of T in nonterritorial floaters and territory owners. Floaters with artificially elevated plasma T levels were unable to obtain territories. Territory owners implanted with T did not expand their territories, although they engaged in more aggressive behavior than did control males. Flutamide, which binds to T receptors in target areas, had no effect on territoriality. However, most territory owners given a combination of flutamide and ATD, an aromatization blocker, lost portions of their territories even though they actively defended them. Our results 1) suggest that, although T influences aggressive behavior, elevated plasma levels alone are insufficient to overcome previously-established social relationships between territory owners or between owners and floaters; 2) indicate that impairing the action of T leads to reduced abilities of territory owners to maintain territories against vigorous challenges: and 3) support recent findings that T acts on reproductive behavior in birds through both androgenic and estrogenic metabolites.     

Spawning dynamics of orange roughy,Hoplostethus atlanticus,in mid-slope waters of New Zealand

Synopsis The social and reproductive biology of the sand tilefish,Malacanthus plumieri (Malacanthidae), was studied at Glover's Reef, Belize, where this species occurs in colonies over sand-rubble flats. Individuals each occupy a home burrow refuge and a surrounding home range. Home range overlap among adjacent fish of the same sex is low, and individuals defend exclusive use of much of their home range against all conspecifics except mates (i.e., territoriality). Areas defended by males overlap the territories of up to 6 females; and male territory area is positively related to the number of female residents. Males maintain dominance over females within their territories by aggression, including intervention into some female disputes. Females spawn pelagically-dispersed eggs as frequently as every day. Each female spawns near her burrow, almost exclusively with the male whose defended area encompasses her territory (harem polygyny). Tilefish colonies therefore consist of a mosaic of female territories over which adjacent male territories are superimposed. Histological evidence and observation of behavioral sex change in one female revealed thatM. plumieri is capable of protogynous sex reversal. Females did not change sex in response to removal of one male. Occurrence of small transitional fish indicates that the onset of sex change is controlled by factors other than size-related social hierarchies within harems or colonies.     

Competition for landmark territories among male Polistes canadensis (L.) (Hymenoptera: Vespidae): large-size advantage and alternative mate-acquisition tactics

Alternative male mating tactics of insects at landmarks (leks)have only rarely been investigated. Some males of the paperwasp, Polistes canadensis (L.), were territorial at small treesalong the crests of dry ridges in Santa Rosa National Park,Costa Rica. Territories did not contain nests or resources forwhich females foraged. Contrary to other "hilltopping" species,male P. canadensis competed most intensely for territories insaddles along these ridges rather than at the highest points.Nonterritorial males patrolled small areas of the ridge line,following a path that took them to a number of territories.Many males switched between territoriality and patrolling, suggestingthat both size-related tactics belong to one conditional strategy.Males that were territorial on 2 or more days were larger thanthose that were territorial on only 1 day, and these in turnwere larger than permanent patrollers. Moreover, the mean sizeof territorial males was positively correlated with two measuresof territory attractiveness, suggesting that larger males monopolizepreferred sites. Mean age of territorial males was also relatedto territory attractiveness, but males of intermediate age claimedthe most attractive territories.