Territorial bequeathal by red squirrel mothers

Red squirrels (Tamiasciurus hudsonicus) defend individual, food-basedterritories year round. These territories are crucial for overwintersurvival, yet some female red squirrels in northern populationsbequeath their territories to offspring and search for a newone. Bequeathal involves active, strategic dispersal by breedingfemales. Our study documents this unusual behavior and investigatesits correlates in two red squirrel populations in northern Canada.Thirty percent of breeding females dispersed. Bequeathal wasrelated to breeding date, with late-breeding females more oftendispersing, but was not related detectably to female conditionor territory quality. As an underlying trade-off, early-bornjuveniles more likely acquired a territory independently, butearly-breeding females lost mass and may have increased theirrisk of territory loss. We suggest that bequeathal has evolvedas a consequence of condition dependence in breeding date coupledwith limited resources.     

Nest and mate choice in the red bishop (Euplectes orix) : female settlement rules

We investigated female settlement in a colony of red bishops(Euplectes orix), a territorial and highly polygynous weaverbirdwidely distributed over sub-Saharan Africa. An earlier studyshowed that male reproductive success is mainly determinedby the number of nests a male builds in his territory, whichappeared to be a good indicator of male quality. Because malesprovide no parental care or food resources within the territory,females sharing a territory do not compete for material resourcesand might therefore be expected to settle preferentially interritories of males that build many nests to gain the possiblegenetic benefit of high-quality offspring. An analysis of femalesettlement, however, revealed that females did not show a preferencefor territories of males with many nests and that the distribution of female breeding attempts with regard to the number of vacantnests within a territory could be explained best by randomfemale settlement in 3 out of 4 years. Females settled moreoften than expected by chance (in 3 out of 4 years) in territoriesalready containing occupied nests, indicating that residentfemales did not discourage settlement of additional females.However, sharing a territory with other females might imposecosts in terms of an increased predation risk because nestsin territories that contained other nests with young sufferedfrom higher predation than nests in territories that did notcontain other nests with young. Females therefore might tradethe possible benefits of settling in territories of males withmany nests against the costs of sharing a territory with otherfemales. This might result in the mating pattern found withrandom female settlement and male reproductive success beingdirectly proportional to the number of nests built. We discuss possible implications of this mating pattern for sexual selectionon males.     

Uncertainty in territory quality affects the benefits of usurpation in a Mediterranean wrasse

Individuals should defend sites when the expected benefits ofthe territory exceed the cost of defense. However, if territoryqulaity is unpredictable or difficult to assess, the expectedpattern of territorial behavior is less clear. In a Mediterraneanwrasse, Symphodus ocellatus, mating success is skewed with 2%of nesting males getting more than 20% of the spawning success.Yet, variation in mating success is not explained by any knownphysical characteristic of males or their territories. Instead,females prefer nests with a recent history of mating successbecause males are less likely to desert the offspring she leavesbehind. Thus, territory quality is transient and determinedby interactions between the sexes. I measured the frequencyof territorial takeovers and the uncertainty in mating successamong days at a nest. Observations indicated that S. ocellatusmales usurped their neighbor's successful nests when males wereunsuccessful and larger than their successful neighbor. Sitesthat achieved mating success had a significantly higher probability(0.84) of remaining sucessful between consecutive days thanunsuccessful territories had of becoming successful (0.30).Unsuccessful males obtained higher and more certain fitnessreturns if they usurped a successful neighbor's territory. Interactionswithin and between the sexes drive uncertainty in success, whichinfluences territorial behavior in this species.     

Black holes, mate retention, and the evolution of ungulate leks

In some ungulates living in unstable herds, females in estrusleave their usual groups and join males defending mating territories.During the 12-24 h before mating, females commonly move severaltimes between males. These movements are commonly caused byharems being disrupted by young males or by overenthusiasticcourtship by the territorial male. When females leave a territorythey typically move to its nearest neighbor. Under these circumstances,clusters of territories can retain estrous females until theymate and leave the lek. This paper develops a model to investigatethe benefits of defending clustered versus dispersed territoriesto males and the consequences of variation in the rate of femalemovement between territories (P_m) and the tendency for femalesto move from one territory to a neighboring territory (a). Wherefemales move between territories at least once every 24 h (P_m< 0.04) and usually move to neighboring territories (a <0.5), the mating success of males is inversely related to thedistance from their territory to its nearest neighbor, and malesdefending clustered territories have higher mating rates thanthose defending dispersed territories. This process may be importantin the initial evolution of ungulate leks, which may resembleblack holes, attracting and retaining estrous females untilthey mate and their estrus ceases. It provides one possibleexplanation of the evolution of ungulate leks that does notrely on female preferences for mating with particular phenotypiccategories of males.     

The use of territorial gardening versus kleptoparasitism by a subtropical reef fish (Kyphosus cornelii) is influenced by territory defendability

Many territorial herbivorous animals increase the productivityor quality of the prey community on their territory. However,this gardening tactic may be vulnerable to kleptoparasitismunless gardeners can exclude others from the patch. We investigatedthe influence of territory defendability on the decision ofwestern buffalo bream (Kyphosus cornelii) to defend an algalgarden or to leave their territories and invade those of conspecifics.Fish monitored while away from their territories did kleptoparasitizethe gardens of conspecifics. We manipulated the structural complexityof territories through the addition of short or tall simulatedvegetation. Increasing the height of simulated vegetation onthe territory decreased the defendability of territories butincreased their safety for fish; territories with tall vegetationwere more easily invaded by conspecifics, but fish returnedmore rapidly to these territories after being disturbed. Ina paired experiment, fish spent more time away from their territories(roving) when tall vegetation was added than when short vegetationwas added. However, this effect was influenced by depth, witha greater difference between treatments on deeper territories,and by body size, with a greater difference for larger fish.These results support the prediction that territory-holdingindividuals faced with a trade-off between defending their owngarden or kleptoparasitizing from others adjust their use ofthese tactics based on the defendability of resources.     

Accepting unrelated broods helps replacement male yellow-headed blackbirds attract mates

Replacement male yellow-headed blackbirds (Xanthocephalus xanthocephalus)did not destroy broods sired by the previous territorial maleand they showed no aggression toward females with unrelatedbroods. To test whether their tolerance of unrelated young wasmisdirected normal parental care, we removed males from experimentalterritories after primary nests were completed but before secondarynests were initiated. Replacement males fed young that theypresumably had sired in secondary nests and ignored foster youngin primary nests, whereas control males fed young in their primarynests. To identify potential benefits of accepting unrelatedyoung, we analyzed patterns of within-season breeding dispersaland of female settlement on territories following nest lossesto predators. Although some female yellowheads do renest onthe same territory following nest failures, the number of nestsinitiated on territories after a predation event was significantlylower than the number initiated on territories without predationover the same period of time. This implies that late-settlingfemales use the number of active or failed nests and/or thenumber of females on a territory when choosing where to breed.If replacement males that accepted unrelated offspring attractmore new females in the remainder of the current breeding seasonthan infanticidal males, then tolerance of unrelated young byreplacement males may be adaptive in some polygynous birds.     

Territorial bequeathal by red squirrel mothers: A dynamic model

Female red squirrels (Tamiasciurus hudsonicus) sometimes bequeath their territory to their offspring. Late-breeding females are more likely to leave than are early-breeding females. Early-breeding females tend to lose weight during reproduction while late-breeding females do not. Early-born juveniles are more successful at winning territories than are those born late. I use dynamic programming to investigate the trade-offs involved in territorial bequeathal from the female's perspective as a function of breeding date, litter size and female body condition. The model predicts that two classes of females are more likely to bequeath: those in good condition early in the season, and those in poor condition at the season's end. Only versions of the model incorporating a function improving the female's ability to win a territory over the season mimicked the pattern of bequeathal observed in the field. A seasonal effect is likely an important factor in driving the pattern of strategies observed.     

Male Seychelles warblers use territory budding to maximize lifetime fitness in a saturated environment

In cooperatively breeding species, helping at the nest and buddingoff part of the natal territory have been advanced as strategiesto increase fitness in an environment that is saturated withterritories. The importance of helping or territory buddingas a determinant of lifetime reproductive success (LRS) hasbeen debated because the potential benefits of both strategiescould not be separated. Here we test the causes and the immediateand future fitness consequences of single dispersal decisionstaken by male Seychelles warblers (Acrocephalus sechellensis).Males breeding in high-quality territories (high food abundance)have significantly higher LRS than similar-aged males buddingoff part of the parental territory. Initially, budders havea low reproductive success (because of limited food resourcesor absence of a breeding partner). However, they have a longlife span and inherit high-quality territories through sitedominance, by which they gain higher LRS than breeders on low-qualityterritories, helpers, or floaters. Experimental creation ofmale breeding territory vacancies showed that most young malesbecame budders because of intense competition for high-quality territories. The translocation of warblers to the previouslyunoccupied Aride Island shows that males behave according tothe expected fitness benefits of each dispersal strategy. Inthe absence of competition for territories on Aride, all youngmales bred in high-quality territories. However, after saturationof high-quality habitat with territories, most males becamebudders rather than breeders on low-quality habitat, helpers,or floaters.     

Territory quality drives intraspecific patterns of extrapair paternity

Despite the potential reproductive benefits of extrapair matings,extrapair paternity rates in many avian species often vary greatlyamong populations. Although ecological factors have been shownto influence intraspecific patterns of extrapair paternity insome species, for cooperatively breeding species living in familygroups, social/demographic factors may also play a role. Thisstudy examined how ecological factors related to territory quality(vegetation cover, insect abundance) and social/demographicfactors (group size, number of breeding pairs, genetic relatedness)influenced intraspecific patterns of extrapair paternity incooperatively breeding superb starlings, Lamprotornis superbus.Superb starlings inhabit spatiotemporally variable African savannaswhere high temporal variability drives reproductive decisions(adoption of breeding roles, offspring sex allocation) and whereterritories suitable for breeding are limited. Although extrapairpaternity rates were only 14% of offspring and 25% of nests,they varied greatly among groups, ranging from 4% to 32% ofoffspring and from 7% to 60% of nests. These among-group differencesin extrapair paternity were not related to social/demographicfactors but instead to territory quality; extrapair paternitywas higher on lower quality territories (lower vegetation coverand grasshopper abundance) than on higher quality territories(higher vegetation cover and grasshopper abundance). These resultssuggest that even in a heterogeneous landscape where suitablebreeding territories are limited, subtle differences in habitatquality can have profound effects on reproductive decisionsand patterns of extrapair paternity. Understanding the interactionbetween spatial (habitat heterogeneity) and temporal (temporalvariability) environmental variation will be important for determininghow environmental and social factors drive avian reproductiveand mating decisions.     

Establishment Success of Experimentally Delayed Juvenile Marsh Tits Parus palustris

By experimentally manipulating the time when young marsh tits, Parus palustris, could become established, I tested the influence of size, hatching date, social dominance in caged situations, and time of season on establishment success. Individuals that managed to become established were divided into two groups: those that became established in new territories and those that were found in the same territory as the one where they were caught. In no circumstances did size, hatching date or social dominance influence establishment success. The only factor tested that had a significant effect on establishment success, was when the juveniles were released; those that established themselves in new territories had been released earlier than unsuccessful ones. Thus, prior occupancy rather than hatching date per se is the factor determining establishment in new territories. Juveniles that became re-established in their former territory had been released significantly later than those not becoming re-established. As the season progresses, increasingly fewer unestablished juveniles are present to fill vacancies. Thus, late-released juveniles stood a better chance of finding their flock position still vacant compared with juveniles released early, whose previous positions would already have been occupied by other juveniles.     

Territorial songs indicate male quality in the sac-winged bat Saccopteryx bilineata (Chiroptera, Emballonuridae)

Defense of territories in many animal species involves the advertisementof territory holder quality by acoustic signaling. In the sac-wingedbat Saccopteryx bilineata, males engage in territorial countersingingwhen reoccupying their day-roost territories in the morningand in the evening before abandoning the roost for the night.Females roost mainly in male territories, and territory holdersare reproductively more successful than nonterritorial males.In territorial songs of male S. bilineata, we distinguished6 syllable types and parameterized their acoustic properties.The analysis of 11 microsatellite loci allowed assignments ofjuveniles to their parents. Males had a higher reproductivesuccess both when they uttered more territorial songs per dayand when their long buzz syllables had a lower end frequencyof the fundamental harmonic. Long buzzes had a harsh qualitydue to a pulsation of the fundamental frequency at the syllableonset and also had the highest sound pressure level of all syllabletypes in most territorial songs. Territorial songs and especiallylong buzz syllables are thus likely to advertise territory holderquality and competitive ability.     

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.     

Survival of the Orange-tufted Sunbird,Nectarinia osea

Cape Sugarbirds are southern African endemics, found in fynbos vegetation of the Western and Eastern Cape Provinces of South Africa, and they feed almost exclusively on the nectar of Protea inflorescences. Breeding male Sugarbirds are highly territorial and defend resources for themselves, their mates and their offspring. Sugarbirds, however, often leave their breeding territories in search of food during the dry season and return to the same breeding sites each year, thus experiencing an annual cycle of movement from one food source to another. Male territory size as well as breeding success, were determined over two breeding seasons for a population in the Helderberg Nature Reserve in the Western Cape Province. Sugarbird fledgling success was significantly greater for males defending large territories. Males moved or increased the size of their territories between seasons, possibly to improve breeding success, but no changes in territory size were observed during the breeding season.     

Caribbean damselfish with varying territory quality: correlated behaviors but not a syndrome

The behavioral syndrome hypothesis suggests that individualanimals within a population behave differently due to specificbehavioral types, and these should be consistent across behaviorsor in different contexts. In contrast, for animals that livewithin an environment in which territory quality can changeover time, natural selection should have favored behavioralflexibility and modulation of the cost of defense in relationto territory quality. This would require assessment of the territoryfollowed by displays of appropriate types and intensities ofbehavior. We examined the territorial behavior of male beaugregorydamselfish (Stegastes leucostictus) by enhancing territory qualityusing artificial breeding sites and comparing their behaviorto males on lower quality natural sites. When male fish weredefending high-quality artificial territories, they had higherlevels of aggression toward male conspecifics and courtshiptoward females than when on low-quality natural territories.We also found that aggression and courtship behaviors were correlatedon natural sites but not on artificial sites. Behaviors werenot correlated within individuals when males switched from naturalto artificial territories or from artificial to natural territories.These results indicate that males assess their current territoriesand adjust behaviors accordingly and that courtship and aggressivebehaviors are not linked within a permanent behavioral syndrome.     

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.     

Red coloration of male northern cardinals correlates with mate quality and territory quality

I investigated how mate quality and territory quality influencean extravagant ornament in a socially monogamous species thatdefends multipurpose territories. Northern cardinals (Cardinaliscardinalis) are a highly dichromatic, socially monogamous species,and males are a brilliant red. I conducted a 3-year field studyof northern cardinals and found that redder males produced moreoffspring in a breeding season. Two selective factors mediatedthis fitness gain. Redder males were paired with earlier breedingfemales, an established measure of mate quality in birds. Second,redder males obtained territories of higher quality, as measuredby vegetation density. Interactions among these factors werealso important in explaining variance in male reproductive success.Multivariate analysis indicated that earlier breeding increasedreproductive success independent of territory quality. In turn,territory quality contributed to male reproductive success throughits effect on nest survival and possibly through its role in attractingan earlier breeding female.     

Competitor intrusions and mate-search tactics in a territorial marine fish

Recent theoretical studies have drawn attention to the importanceof understanding the costs females experience during mate choiceand mating and how females resolve the trade-offs presentedby such costs. In garibaldi damselfish (Hypsypops rubicundus)females defend permanent feeding and shelter territories andmust leave their territories to search for and spawn with nestingmales. Both mate searching and spawning occur in bouts, separatedby returns to the territory. I used focal female observationsto test predictions of the hypothesis that such behavior isa means of minimizing the amount of food lost to competitorsthat enter the female's territory while she is searching foror spawning with males. Consistent with this hypothesis, therewas a strong likelihood that the number of intruders enteringthe territory would increase linearly, and hence their totalimpact exponentially, with time away. In addition, the averageduration of searching bouts and the duration of a spawning boutwere significantly inversely related to a measure of territorialintruder pressure. Most intruders were heterospecific food competitors;incidence of intrusion by conspecifics was rare, and intrudingconspeciflcs did not contest the owner upon her return. In addition,the time females spent in their territories before beginninganother bout of searching was not positively related to theduration of the previous bout. Thus, there, is no support forthe alternative hypotheses that returning to the territory isa means of preventing takeover of the territory by conspedflcsor is the result of fatigue.     

Territory switching behavior in a sedentary tropical passerine, the dusky antbird (Cercomacra tyrannina)

Demographic data from an 8-year study of a marked populationshowed that switching territories and mates is common in bothgenders of dusky antbirds (Cercomacra tyrannina), a sedentaryneotropical passerine with year-round territories and pairbonds.We conducted 22 experimental removals and followed six naturaldisappearances to examine territory switching. Antbirds quicklyabandoned territories and mates to move to openings created by experimental removals. Pairing with the resident on a newterritory was rapid. Unmated birds attracted new mates by singinga gender-specific song that differed from songs given by matedbirds. There were no gender differences in replacement timeor rate. Some vacancies, experimental and natural, were notfilled, suggesting that floaters were rare. Territory and mateswitching were not related to immediate enhancement of reproductive success because the probability of reproducing successfullywas equally poor on all territories. Territory switching maybe an overlooked but common tropical form of territorialitythat increases individual survivorship during periods of lowfood abundance (dry season). We suggest that switching is favoredwhen low annual reproductive success enhances selection fora long lifespan as the primary means to increase reproductivesuccess.     

Form,function and consequences of density dependence in a long‐distance migratory bird

The density dependence of demographic parameters and its implications for population regulation have long been recognized. Recent work has revealed potential effects of density on mating systems and sexual selection, but few studies concurrently assess the consequences of density on both demography and sexual selection. Such an approach is important because population processes and individual behaviors can interact to influence population growth and evolutionary trajectories. In this study, we tested the density dependence of breeding success, extra‐pair paternity, and the opportunity for sexual selection in a population of American redstarts Setophaga ruticilla using two different measures of density. To evaluate temporal patterns, we analyzed annual territory density, based on the total number of territories at our study site each year. To evaluate spatial patterns, we analyzed local territory density within years, based on the number of territories surrounding a focal territory. Greater annual density was associated with fewer offspring fledged per female, a reduced mean population rate of fledging success, and a lower relative contribution of extra‐pair paternity to male fitness. Greater local density was associated with fewer offspring fledged, reduced fledgling success, higher rates of nest loss, and higher rates of paternity loss on focal territories. Interestingly, greater local density was also associated with greater nestling mass on focal territories, which could imply that more densely‐packed territories contain superior resources. Overall, our results suggest that the effects of crowding via greater territory density reduce fecundity through increased nest predation, rather than reduced food availability, and increase rates of extra‐pair paternity. Thus, the selective pressures faced by individuals and their reproductive behaviors are likely to differ based on the annual and local density they experience, which may then feed back into population demography.     

Parent-offspring conflict over duration of parental care and its consequences in tawny owls Strix aluco

In species defending territories, fitness of newly independent juveniles could depend on phenotypic quality or early access to vacant resources as a result of rapid dispersal. In the first case, parent-offspring conflict will arise when parents cease feeding maturing offspring while these still demand provisioning. In the latter case, the young should decide when to stop begging to search for vacant territories. I radio-tracked 72 juvenile tawny owls from independence to sexual maturity to investigate whether parents or offspring decided the timing of the onset of independence and compare effects of age versus date of independence on survival and reproductive status. Juvenile owls stopped begging when 90–123 d old. This was synchronous within broods, independent of gender or age rank. Independence age of cross-fostered young varied across foster nests but was independent of hatching nest. After independence, young roosted in their parents' territory for 18 d on average before dispersal. This suggests that juvenile owls would rather extend dependency than disperse early. As predicted from this behaviour, age of independence had a positive influence on survival and reproductive status at maturity, whereas there were no effects of date of independence.