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.     

Seasonal Changes of Vocal Rates and Their Relation to Territorial Status in Male Galápagos Sea Lions (Zalophus wollebaeki)

Male vocal displays play an important role in sexual selection through both male–male competition and female choice. While this is supported by numerous studies in birds, much less attention has been paid to the role of such displays in mammals. We investigated the function of vocal displays in a polygynous mammal, the Galápagos sea lion (Zalophus wollebaeki). In our study population a large proportion of mature males are unable to establish a territory, providing the opportunity to compare the vocal behaviour between territorial and non‐territorial males. We examined how seasonal patterns of vocalizations differed between territorial and non‐territorial males and how the number of females present in a territory influenced behaviour of territorial males. We found that territorial males vocalized at higher rates than non‐territorial males, and territorial males vocalized more at the onset than towards the end of territory tenure. During the onset of territory‐tenure vocalizations of territorial males were directed more often towards other males than females. Furthermore, we found that vocalizations of territorial males were not only given in male–male interactions, but were also actively directed towards females. Territorial males vocalized at higher rates when more females were present in their territory. Our results suggest that vocalizations are important in male–male interactions, are relevant in territorial disputes and are used as a proxy for females to assess a male’s quality.     

Male dimorphism, territoriality and mating success in the tropical damselfly, Paraphlebia zoe Selys (Odonata: Megapodagrionidae)

The tropical damselfly Paraphlebia zoe has two male morphs: a black-winged (BW) male which is associated with territorial defense of oviposition sites; and a hyaline-winged (HW) male similar in appearance to females, and, compared to the black morph, less frequently found defending territories. In a wild population of this species, we first assessed the relationship between phenotypic traits [male morph, size and territorial status (being territorial or non-territorial)], their role on mating success, and the degree to which a particular territory may contribute to male mating success. Second, to relate a physiological basis of being territorial we compared both morphs in terms of muscular fat reserves and thoracic muscle, two key traits related to territory defense ability. Males of both morphs defended territories although the BW males were more commonly found doing this. BW males were larger than HW males and size predicted being territorial but only within HW males (territorial males were larger) but not in BW males. Male mating success was related to territorial status (territorial males achieved a higher mating success), but not to morph or size. Furthermore, territory identity also explained mating success with some territories producing more matings than others. The BW morph stored more fat reserves which may explain why this morph was more likely to secure and defend a place than the HW morph. However, the HW morph showed higher relative muscle mass which we have interpreted as a flexible strategy to enable males to defend a territory. These results are distant to what has been found in another male dimorphic damselfly, Mnais pruinosa, where the advantage of the non-territorial morph relies on its longevity to compensate in mating benefits compared to the territorial morph.     

The role of male territories in the mating system of the Yellow-shouldered Widowbird Euplectes macrourus

The territorial system and breeding biology of the Yellow-shouldered Widowbird Euplectes macrourus (Ploceidae) was investigated in western Kenya. Yellow-shouldered Widowbirds had a resource-defence polygynous mating system: males defended large (mean = 0.95 ha) territories and built the coarse framing for the nests in tall grass. Males had up to five females nesting per territory. Females provided nearly all parental care except for a territorial male seen feeding a fledgling: the first observation of paternal care in the wild for this genus. There was considerable variation in territory size, but the cause of this variation remains unknown: territory size was not related to potential indicators of territory quality, such as grass height and abundance, did not relate to male morphology (mass, size and ornament size) or territorial behaviour (boundary displays and singing) and did not affect female preferences. Although resources (territories and nests) were defended by the males, observations that males frequently fed outside their territories and formed communal roosts during the breeding season suggest that this species represents a transitional stage between typical resource-defence polygyny and lek breeding.     

Incorporating territory compression into population models

The ideal despotic distribution, whereby the lifetime reproductive success a territory's owner achieves is unaffected by population density, is a mainstay of behaviour-based population models. We show that the population dynamics of an island population of Seychelles warblers ( Acrocephalus sechellensis ) cannot be modelled with an ideal despotic distribution, and suggest the effects of both territory shrinkage and territorial disputes on reproductive success must be included to adequately model the population dynamics of this species. To do this we introduce two different approaches. The first is reductionist, using data on how population density affects individuals' reproductive success to predict population growth rates. Because such a model is mechanistic, it can be used to predict population dynamics in novel environments, making it a desirable long-term solution. However, because territorial populations are typically tightly regulated, birth and death rate data at low population densities are often unavailable. Hence, our second approach statistically infers the relationship between population density and per territory reproductive success, and thus provides a stop-gap solution for the shorter term. Our analysis indicates that although the Cousin population of Seychelles warblers is highly resilient to environmental stochasticity, the degree of resilience is considerably underestimated by approaches that ignore intrinsic regulation through territory shrinkage and territorial disputes.     

The territorial system of the kob antelope (Kobus kob kob) in the Comoé National Park, Côte d'Ivoire

The territorial system of kob antelopes in the Comoé National Park, Côte d'Ivoire, was studied from March 1993 until April 1997. Whereas size and shape of territories remained constant between 1993 and 1994, the number of territories decreased from 1994 onwards. A decline of the population density probably led to the decline in territory numbers and also to variations in size and shape of the remaining territories. The decline of kob numbers is the result of heavy poaching in the study area. Whereas a lekking system was present in the study area prior to 1975 when the population density was approximately 14.4 kobs/km², males switched to the resource defence territorial system present today after the population density decreased to 12.5/km² in 1993 and finally to 2.3/km² in 1997. Compared to other research areas with higher population densities, territories in the Comoé National Park are larger, almost all adult males are capable to defend a territory, fights over territories are rare and loss of territory tenureship is almost exclusively due to the death of the territorial male. We suggest that those differences can all be related to the low population density in the study area.     

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.     

Scent marking in a territorial African antelope: I. The maintenance of borders between male oribi

Scent marking is ubiquitous among the dwarf antelope and gazelles of Africa, but its function has been the subject of debate. This study examined preorbital gland scent marking in the oribi, Ourebia ourebi, a territorial African antelope. Several hypotheses for the function of scent marking by territorial antelope were tested with observational data. Of these, the hypotheses that scent marking is driven by intrasexual competition between neighbouring males, and that marks serve as an honest advertisement of a male's ability to defend his territory from rivals, were supported best. Thirty-three territorial male oribi on 23 territories marked most at borders shared with other territorial males, and territorial males marked more often at borders shared with multimale groups than at borders shared with a single male. This suggests that males perceived neighbouring male groups as a greater threat to territory ownership than neighbouring males that defended their territories without the aid of adult subordinates. Marking rate was unrelated to territory size or the number of females on adjacent territories, but males with many male neighbours marked at higher rates than those with fewer male neighbours. These results suggest that the presence of male neighbours has a greater effect on the scent marking behaviour of territorial antelope than has been considered previously. Copyright 1999 The Association for the Study of Animal Behaviour.     

Sex difference in the communicatory significance of localized defecation sites in Arabian gazelles (Gazella arabica)

Mammalian scent marking in localized defecation sites (latrines) has often been interpreted in the context of (male) territory defense. However, latrines could have different functions in males and females, especially where territorial males monopolize groups of females with stable social alliances and pronounced home range overlap. We investigated the communicatory significance of latrines in wild Arabian gazelles (Gazella arabica) and assessed the spatial distribution of latrines within home ranges. Latrine density and utilization was highest in the center of female group home ranges, and less frequent in peripheral home range sections, pointing towards communication within groups rather than towards territoriality. When considering male home ranges, latrine densities and utilization were higher in non-overlap zones, contradicting a territorial function. This pattern appears to be caused by more females than territorial males per given area establishing latrines. A subsequent survey of latrine utilization, based on camera trapping, suggests that males use latrines for territory defense: males visited latrines in overlap zones disproportionally more often than females, and successions of two males prevailed. Our study thus highlights that male territorial marking can be masked when males and females use the same marking system for different purposes.     

Seasonal changes in territorial behaviour of hazel grouse ( Bonasa bonasia) in a temperate forest of South Korea

We studied the seasonal changes in territorial behaviour of hazel grouse (Bonasa bonasia) from December 1999 to July 2001 in a temperate forest of South Korea. The study was based on the use of radio telemetry. Territory size and degree of overlap increased after the breeding period, and index of advertisement and border disputes decreased. In spring, the territory was defended by both males and females. During autumn and winter, this behaviour was relaxed to such a degree that hazel grouse cannot be considered territorial, but as forming a home range.     

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.     

Territorial behavior of a green hairstreak Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae): site tenacity and wars of attrition

Males of Chrysozephyrus smaragdinus were active from late morning to late afternoon, during which they showed territorial behavior, perhaps for mating. The territorial male stayed in a particular area and occasionally flew around it, referred to hereafter as the inspection area. When other male intruded into this area, the territorial male rushed to him. Then, they engaged in a circling flight regarded as a "war of attrition". During this flight, the two males sometimes strayed far away from the territory. After the circling flight, the resident returned to his territory in almost all cases (98%). Despite such intrusions, many residents defended their territory for several successive days. This suggests strongly the "effect of prior residence". We recorded the circling flights with a high-speed video camera, and confirmed that the male that ceased the circling flight first was the loser. This finding gave some validity to consider circling flight as wars of attrition. In a few cases, the territorial male mated with a female that came to the territory. These once mated males held the territory no longer, suggesting that mating experience should restrict the next mating opportunity in this species.     

Territory Availability at Different Spatial Scales does not Affect Fighting Investment in Males of the Lycaenid Butterfly Strymon mulucha

In territorial species, rivals investment in fights over territories may increase when the availability of suitable areas for defense is low. This should occur because low territory availability may increase the costs to maintain and acquire territories. Although such process occurs in small spatial scales (local scale), territory availability in larger scales (regional scale) may also affect fighting investment, as losers should incur additional dispersing costs to find new territories. In this study, we used males of the hilltopping butterfly Strymon mulucha to evaluate the hypothesis that males should invest more in territorial fights when the costs to find new territories are higher (both at local and at regional scale). We timed male–male contests for territories located in 12 hilltops and measured male density per territory in each hilltop (local scale). We also quantified the distance between hilltops containing suitable areas for territories (regional scale). Male–male contests lasted 21 s on average, and copulations did not occur during the observations. The duration of contests was unrelated to the male density per territory or to the distance among hilltops, indicating that the investment in fights was unaffected by the availability of territorial sites, independent of the spatial scale. As male–male contests in S. mulucha are longer than the mean contest duration in other butterfly species and mating is extremely rare, we suggest that the value of each territory may be high enough to favor males that always invest as much as possible in contests.     

Annual variation in the mating system of the dragonfly Paltothemis Kneatipes (Anisoptera: Libellulidae)

The libellulid dragonfly Paltothemis lineatipes exhibits annual variation in male mate-locating behaviour, variation that appears to be related to changes in the number of males competing in an area. In a year when males were numerous, as reflected in a high rate of male-male interactions, individuals defended small streamside territories, regularly raided neighbouring territories in an attempt to steal females there, and often were forced off their territory after a few hours of daily ownership. In this year, temporal partitioning of a given site was common, with the same males taking it in turns to defend the location for part of each day. In a year when males were much less numerous, as seen in a very reduced rate of aggressive encounters, males defended territories more than twice as large on average than in the high-density year; they rarely took females from neighbours, and they much less frequently 'shared' ownership of a site with another male on a given day. The changes between low- and high-density years were not associated with a change in average daily copulatory success of territorial males, although the variance in mating success was significantly greater in the high-density year.     

Territory Area as a Determinant of Mating Systems

Territoriality is an integral part of breeding behavior in manyanimals. Because the reproductive success of territorial malesis often limited by access to females, breeding males shouldbehave as "area maximizers" when the basis of female choiceis either the abundance of resources within the territory orterritory area per se. Being reproductively more limited byenergy, territorial females should be "energy maximizers." Aseries of simple analytical models of territory area for suchforagers is developed to explore how changes in local food productionand/or local competitor density affect both the probabilityof a territorial male securing more than one mate (polygyny)and the probability of his and his mates' reproductive successincreasing. Two cases are modeled (only males territorial vs.both sexes territorial), each for various sets of assumptionsregarding interactions between food production, feeding efficiency,and competitor density. Concurrent responses in territory area,territory food reserves, net energy gain, and time budgetingprovide testable sets of predictions for each scenario. Whereonly males are territorial (Case I), changes in food productioncan have different (indeed, opposite) effects upon an individualmale's probability of becoming polygynous, depending upon whetherthe basis of female choice is the abundance of food within theterritory or another factor positively correlated with territoryarea. Increases in competitor density usually decrease the probabilityof polygyny regardless of the basis of female choice. Whereboth sexes are territorial and territories overlap intersexually(Case II), the mating system becomes a function of the numberof female territories within each male's territory, which varieswith the ratio of male to female territory areas. In this case,the probability of polygyny occurring will increase if foodproduction for both sexes increases without concurrent increasesin competitor density, and will decrease if competitor densityfor both sexes increases without concurrent increases in feedingefficiency. Few data are presently available to test eitherthese general predictions or numerous sets of secondary predictionstabulated in the text. Available evidence is largely consistentwith the models, but mostly circumstantial. This is becausethe predictions of these and other models of territory areaare strongly assumption dependent, and few published studieshave investigated these assumptions. These analyses demonstratethat to accurately assess the mechanisms by which environmentalfactors affect territory area, and thus mating systems, testsof the underlying assumptions of models are essential.     

The evolution of monogamy in primates

The evolution of primate monogamy is described as an ordered sequence of choices by generalized, hypothetical females and males. Females first choose whether or not to associate with other females. Predators encourage gregariousness in diurnal primates; however, nocturnality or scarce and evenly distributed food supplies may enforce separation. A testable group size model based on food patch size is developed and qualitatively supported.If females choose solitude, males then choose either to defend a single female and invest in her offspring, or to compete with other males for access to several females, usually by defending a territory or establishing dominance over the home ranges of several females. The decision rests on the defensibility of females and on the availability of an effective form of male parental investment. Both of these factors are dependent on local female population density. A model is developed that assumes that territorial defense is the principal form of male parental investment, and it predicts that monogamy should occur at intermediate densities: at high densities, males should switch to defense of multiple females, and at low densities there is no investment value in male territorial defense. The model is shown to be only partly adequate. Variation in local population densities prevents the establishment of obligate monogamy through territoriality in small monkeys, since male territorial behavior is inconsistent over the long run. Here, carrying of offspring by males can succeed territoriality, providing an effective and reliable form of parental investment to maintain the pair bond in the face of population fluctuations and changes in group structure. This hypothesis is supported by the scarcity of obligate monogamy among the prosimians, which frequently do not carry their young.     

Host plant-based territoriality in the white peacock butterfly,Anartia jatrophae (Lepidoptera: Nymphalidae)

Mate locating behavior of Anartia jatrophaewas studied near Ochopee, Collier County, Florida. Males were individually marked, and focal animal samples were used to determine activity, residency, and interactions with other butterflies. We recaptured more than half of 53 marked males within the study area on subsequent days. Marked males restricted non-feeding activity to roughly circular areas typically less than 15 m in diameter. Males showed considerable site fidelity and chased both conspecifics and a variety of other insects. Typically, encountered individuals flew away, and the resident peacock male returned to perch near his previous perch. Encounters with conspecific males averaged 38 sec, three times longer than with other insects. Such site fidelity combined with aggressive interactions indicates territorial behavior. When male density more than doubled, site fidelity decreased, resident turnover increased, and areas that had not been defended at lower density were defended. Without exception, defended sites contained the larval host, Bacopa monnieri.In butterfly mating systems, defense of host plants appears to be much less common than defense of natural features such as landmarks.     

Should I Stay or Should I Go Now: Late Establishment and Low Site Fidelity as Alternative Territorial Behaviors

In species in which males defend territories for breeding, males may differ in territorial behavior; alternative behaviors among territorial males are not well understood. In our long‐term study of partially‐migratory song sparrows, we have observed that most territorial males establish territories before females begin nesting and remain site‐faithful both within and between breeding seasons; however, some males establish territories later in the season (late establishers) and/or change territory locations either within or between seasons (movers). Whether late establishment or moving are equally successful strategies for territory defense, or best‐of‐bad‐job options, is not known. Here, we compare the frequencies of these behaviors to demographic variables over a 9‐yr period and compare lifetime tenure and early season nesting success for males who differ in site fidelity and timing of territory establishment. Across years, late establishing was negatively correlated with the return rate of previously territorial males; moving was positively correlated with the number of occupied territories at the start of the breeding season (territory density). While moving was independent of number of years on territory, late territory establishment only occurred in a male’s first year as a territory holder. Of 88 males, 25% established their first territory late, primarily in undefended space; 31% moved. Late and early establishers did not differ in lifetime tenure; movers, however, had longer lifetime tenure than site‐faithful males. Among early establishers, movers and non‐movers did not differ in the number of successful early nests/year or number of young fledged/year; among late establishers, however, movers had significantly higher early nesting success by both measures. Late establishers who moved had higher early season nesting success and higher early season nesting success/year than site‐faithful early establishers. Thus, individual variation in the timing of territory establishment and site fidelity may be facultative alternative territorial strategies.     

Social stimulation of dawn singing in Dusky Flycatchers: a serendipitous experiment

Hypotheses regarding dawn singing in birds remain largely unsupported by quantitative data, especially for suboscine passerines (suborder Tyranni). During a study of the singing behavior of a suboscine, the Dusky Flycatcher (Empidonax oberholseri), in Alberta, Canada, spring storms in 2002 caused the disappearance and presumed mortality of several territorial males and some of their replacements, creating a serendipitous experiment. Overall, 15 males disappeared and, as a result, the number of territorial males in our study area declined from 13 prior to the storms to six afterward. Only one male maintained the same territory throughout the breeding season. During the period of inclement weather and social instability (20 May to 9 June), males sang at high rates during the predawn period. From 10 to 19 June, following this period of unstable weather and turnover in territorial males, males began dawn singing later in the morning, and sang shorter bouts at lower rates. The proportion of males engaging in dawn singing also decreased between the two periods. In contrast, daytime singing activity of paired males was low during both time periods. Playback of dawn songs to territorial males between 20 June and 21 July caused a resumption of some dawn singing. Singing rates were higher on the day of playback than on the day before playback, and the times when dawn singing was initiated were earlier on each of 2 d after playback than on the day before playback. In addition, the proportion of males engaging in dawn singing increased between the day before and the day after playback. Both the decrease in dawn singing when population density was reduced and its partial restoration by playback suggest that dawn singing in this species functions in male–male interactions and support the social‐dynamics hypothesis as an explanation for the dawn chorus in this species.     

Consistent individual variations in aggressiveness and a behavioral syndrome across breeding contexts in different environments in the Black-tailed Gull

Individual behaviors of animals do not evolve separately; they do so in association with other behaviors caused by single shared genetic or physiological constraints and/or favored by selection. Thus, measuring behavioral syndromes—suites of correlated behaviors across different contexts—leads to a better understanding of the adaptive significance of variations in behaviors. However, relatively few studies have examined behavioral syndromes in wild animal populations in changing environments. We investigated a potential behavioral syndrome across antipredator nest defense, territorial defense, chick provisioning, and mating behaviors of male Black-tailed Gulls Larus crassirostris in two successive years under different conspecific territorial intrusion risks and food conditions. Males that presented high levels of antipredator nest defense (aggressive antipredator defenders) against a crow decoy (crows are egg predators) defended their territories against conspecific intruders more frequently than did other males (nonaggressive antipredator defenders), independent of the risk of intrusion. Aggressive antipredator defenders also fed their chicks more frequently than nonaggressive males, but only in a year of low food availability. Taken together, this indicates that males show consistent aggressiveness regardless of breeding context (antipredator and territorial defense), but can regulate food provisioning according to food availability.