Territory establishment in lekking marine iguanas, Amblyrhynchus cristatus: support for the hotshot mechanism

The territory establishment of male marine iguanas and their subsequent mating success were analysed to identify spatial spillover (hotshot) and temporal spillover effects on lek formation. Males started to establish small display territories 2 months ahead of the mating season. Males did not establish territories in temporal synchrony and did not settle at sites where the probability of encountering females was highest. However, males arriving later preferentially established their territories in the neighbourhood of already established territories independently of the density of female-sized iguanas in these territories. Although settling in close proximity, there were no fights between those males. The number of fights between territorial males increased towards, and peaked during, the mating season. Fights did not result in the transfer of space, indicating that space per se was no resource. Instead, fights were directed towards central (hotshot) males. These central males had higher mating success than marginal males. Female density during the time of territory establishment did not predict the mating success of males, because females changed their spatial preferences between early establishment and mating periods. Similarly, the areas where males achieved the highest numbers of copulations changed during 4 years of our study. Thus, there was no evidence for temporal spillover between subsequent seasons. However, most male-male interactions served to distract successful males and may lead to spatial spillover of females into territories of unsuccessful males. In marine iguanas, territorial establishment appears largely governed by hotshot processes.     

Neighbours, strangers and male-male aggression as a determinant of lek size

Interactions between males on leks may play an influential role in lek formation and the regulation of lek size. In this paper I present the results of a playback experiment that simulated de novo settlement at sites adjacent to currently existing display territories of the ochre-bellied flycatcher, Mionectes oleagineus. In the study population, males displayed both solitarily and at small leks. A large proportion of males held no display territory at all. A stranger's song was played to both solitary and lekking males from 10 m outside their territorial boundaries. In separate playbacks, lekking males were also played neighbour's song. Both lekking and solitary territorial males reacted to the playback by decreasing their song rate, approaching the playback speaker and, on occasion, attacking the model. Solitarily displaying males responded more aggressively to playback of stranger's song than did lek males. Lek males were able to distinguish between their neighbour's and a stranger's song and did so irrespective of whether it was played from the neighbour's territory or from outside the lek. In addition to distinguishing between neighbours and strangers, lek males modified their responses to these different playbacks depending on where the playback originated. These results suggest that male-male interactions can be influential in structuring leks. In M. oleagineus, interactions between males are aggressive and act to limit rather than augment lek size. Received: 6 March 1996 / Accepted after revision: 9 December 1996     

Spatial and temporal dynamics at manakin leks: reconciling lek traditionality with male turnover

Leks, display grounds where males congregate and females visit to copulate, are typically traditional in location, despite often high turnover of individual males. How leks can persist in face of male turnover is not well understood, in part due to a lack of detailed field data allowing for a clear understanding of lek dynamics. We followed the fate of individual males at 11 to 15 leks of the blue-crowned manakin Lepidothrix coronata across four breeding seasons to gain insights on how leks are formed and changed in space and time. Between years, leks were traditional in location despite changes in territory ownership due to male disappearance and recruitment. New males were equally likely to recruit by taking over existing territories or by establishing new territories. Recruitment was influenced by age, as recruits were more likely to be adults than subadults. Lek size did not affect the probabilities of a male recruiting or persisting at a territory, and vocalization rate, a correlate of mating success in this population, did not affect male persistence. We used our field data to model changes in lek size and composition over longer periods of time (100 years) to understand how lek traditionality can be reconciled with high male turnover. Our simulations showed that leks in our population rapidly stabilize in size despite changes in territory ownership and that rates of male recruitment and disappearance compensate each other, such that leks have the potential to persist for several decades after the original males have disappeared from them.     

Density dependence and habitat quality modulate the intensity of display territory defence in an exploded lekking species

We evaluated the effect of conspecific abundance and habitat quality of leks on the territorial behaviour of males in an exploded lekking species, the Little Bustard (Tetrax tetrax). The hypothesis that males more intensely defend territories with higher conspecific abundance and better habitat quality was evaluated experimentally analysing the agonistic response of experimental males to male decoys placed on their displaying areas. Decoy experiments showed that the intensity of display territory defence by little bustard males is density dependent. The time experimental males took to return to their display sites after decoy placement decreased with abundance of both males and females. The strength of their final response was positively associated to local male and female abundance in the vicinity of their display sites. Habitat quality also influenced males’ display territory defence since the intensity of male response increased with the degree of natural vegetation cover. Habitat quality was particularly relevant in explaining variation of experimental males’ snort call rate, which decreased with the degree in plough cover and increased with the number of fields in the lekking area. Snort call rate decreased with the level of male aggregation and was lowest in males exhibiting the strongest aggressive response to decoys. These results add new evidence for the density dependence of species’ breeding territorial behaviour, supporting density-dependent models of lek formation and reinforcing the role of resources defence in exploded lek mating systems.     

Passing the buck: resource defence,lek breeding and mate choice in fallow deer

Summary Lek breeding systems, where males defend small, clustered mating territories, are thought to occur where the distribution of females is heavily clumped but males are unable to defend resources used by females. In this paper, we describe a breeding system in fallow deer where males are able to defend resources used by females but the most successful bucks instead defend small territories on a traditional mating ground; where the lek is sited in an area not heavily used by females at other times of year and is visited primarily by females in or close to oestrus; and where mating success on the lek is related to territory position and to male phenotype but not to the resources available on different lek territories. Comparisons with other ungulates suggest that lek breeding species fall into two groups: those where leks are regularly visited by herds of females many of which are not in oestrus and those, like fallow deer, where leks are visited primarily by oestrous females. In the latter species, it is unlikely that females visit the lek for ecological reasons.     

Mating system shift in a pronghorn population

Summary At the National Bison Range (western Montana, USA), pronghorn (Antilocapra american) maintained a territorial mating system from as early as 1965 through 1978. Preliminary observations in 1981 suggested that the mating system had changed. Data from the ruts, 1982–1984 revealed a progressive decay in territoriality. In this paper, data from the territorial years 1969–1978 are contrasted with data from the decay years 1982–1984, with results as follow: 1. In 1982–84, fewer territories were defended than in 1969–78. This was attributable to a smaller proportion of males defending territories in 1982–84, not to a smaller number of males. 2. In 1982–84, most territory owners either abandoned their territories early in ruts, or lost control of females on them following frequent, persistent intrusions by non-territorial males. Abandonment and loss of control did not occur in 1969–78. 3. In 1982–84, territorial males that maintained control of females on their territories did so by shrinking their zones of defense to small areas around female groups. 4. In 1982–84, following the disruption or severe disturbance of all territories, many females left territories, and mated elsewhere, with non-territorial males. In 1969–78, most females remained on territories throughout rut, and mated with territory owners. —The mating system change followed catastrophic winter mortality, 1978–79, that removed 75% of the males, including all males older than 5 years, and all male fawns, from the population. In 1982–84, the number of males present was not different from the number of males in 1969–1978, but the frequency distribution of male ages was strongly shifted toward younger ages. The small number of older males, 1982–84, likely resulted in smaller proportions of males initially defending territories, and in less effective territory defense. When females then clustered on the few territories where defense was at first successful, they attracted large numbers of non-territorial males. The resulting high rates of raids on these territories, coupled with reduced defense radii by territorial males, allowed females only 12% reclining time (summer percentage was 39%). This increased energy cost, plus an apparently greater risk of injury on weakly defended territories, appeared to prompt many females to seek calmer matings elsewhere. Also, if female pronghorn practiced mate selection based upon horn or body size, they may have reduced their efforts to remain on territories in 1982–84. Males from 1969–78 were larger than males from 1982–84, and showed greater variance in horn size. At least two conditions appear to influence the tendency of males to be territorial. First, males must be at least three years old before they attempt to defend a territory. Second, the declining proportion of males defending territories, 1982–84, that coincided with an increasing number of males three years and older, suggests that males also decide whether or not to attempt territory defense based upon the frequency of territorial defense in the population.     

Role of female territoriality in social and mating systems of Canthigaster valentini (Pisces: Tetraodontidae): evidence from field experiments

Sharpnose puffers, Canthigaster valentini (Pisces: Tetraodontidae) at Lizard Island, Australia, live in made-dominated haremic social and mating systems. The hypothesis was that mature females are restricted in their movements and can be monopolized by some males. Field experiments at Lizard Island, Great Barrier Reef, between January and March 1983 showed that mature females were still territorial in the absence of males and movements of females were not controlled by males. Males abandoned their territories when their females were removed. The territorial behavior of those males with access to females (territorial males) restricted the access of other males (bachelor males) to them. Bachelor males took over harems and became territorial males when established territorial males were removed. The results of the experiments thus supported the hypothesis.     

Uganda kob mating success does not increase on larger leks

One explanation for the movement of sexually receptive females to clusters of male territories in lek-breeding species is that larger clusters provide females with higher-quality mating partners, as would be the case if males were distributed between leks in an ideal free distribution for unequal competitors. This ideal free model of lek evolution predicts that male competitive ability and mating success will be greater on larger leks than smaller ones. I tested these predictions by comparing the mean number of males on 19 different Uganda kob leks with the sex ratio, the availability of oestrous females, mating rates, male fighting rates and male turnover rates. Contrary to the predictions of the model, numbers of females, receptive females and fights increased proportionally with lek size, but were no greater per male on larger leks. Multivariate analyses of male and female numbers on leks showed that male numbers were associated with female numbers, female numbers in the past, and a variety of habitat variables which may have related to the costs of holding a lek territory, but female numbers varied only with male numbers and female density in the area. These data do not provide evidence that females gain access to superior males by mating on larger leks, though they do support the possibility that lekking may be promoted by a tendency for larger leks to retain females longer.     

Size and spacing of capercaillie leks in relation to social behavior and habitat

Summary Spring territories of 17 adult capercaillie (Tetrao urogallus L.) cocks were 10–79 ha in extent; they varied inversely in size with the relative proportion of mature forest within them. The number of resident cocks at leks increased with the amount of mature forest within a 1 km radius of each lek center. Leks in two areas (n=46) were regularly spaced with mean distances to nearest neighbors of 1.98 and 2.07 km, corresponding to the territorial space occupied by adult cocks of adjoining leks. In one area with intensive logging, interlek distance increased with decreasing amounts of mature forest between them. Spring home ranges of 18 adult females averaged 51.3 ha±8.2 SE. The spatial relationships did not fit recent models of interlek spacing based on female spacing behavior. Instead, the results suggested that spacing of leks may be related to the territorial requirements of males.     

Female grouping best predicts lekking in blackbuck (Antilope cervicapra)

The study of intraspecific variation can provide insights into the evolution and maintenance of behavior. To evaluate the relative importance of ecological, demographic and social conditions thought to favor lekking, I studied variation in mating behavior among and within populations of the blackbuck, Antilope cervicapra, an Indian antelope. Rather than viewing lekking as a discrete mating strategy, I took a continuous approach and treated lekking as a question of the clustering of mating territories, with leks representing one extreme in a range of territory distributions. I surveyed nine blackbuck populations, which differed in population density and in habitat conditions. For each population, I described the mating system in terms of the clustering of mating territories, and measured various factors suggested to favor lekking. I found that large-scale, among-population variation in territory clustering was most strongly related to female group size. Territory clustering was not related to population density. Female group size, in turn, was best explained by habitat structure. Interestingly, these among-population patterns were repeated at a finer spatial scale within one intensively studied population. These findings suggest that territorial males respond to local patterns in female distribution (represented by group size) when making decisions regarding territory location. Finally, although female distribution may explain territory clustering at the population level and more locally within a population, other selective factors (e.g., female preference, male competition, male harassment) are likely to shape the clustering and size of territories at even finer scales.Electronic Supplementary Material Supplementary material is available in the online version of this article at .Communicated by T. Czeschlik     

Song repertoires and density assessment in red-winged blackbirds: further tests of the Beau Geste hypothesis

Summary The Beau Geste hypothesis proposes that song repertoires are advantageous in territory defense because nonterritorial males, who are prospecting for territories, use the number of song types they hear to assess the density of territorial males, and then avoid densely-settled areas. A territorial male can then inflate the apparent density of singers on his territory by singing several distinct song type. This hypothesis assumes (1) a positive correlation between the density of song types and the density of territorial males, and (2) a negative correlation between the rate of trespassing and the densities of both song types and territorial males. We studied the behavior of male red-winged blackbirds (Agelaius phoeniceus) and found support for the first, but not the second, assumption. Results showing a positive correlation between density of song types and density of territorial males show that the proposed method of density assessment is feasible in this species. However, we could find no support for the assumption that nonterritorial male red-winged blackbirds avoid densely settled areas. In contrast to the assumed negative correlation, rate of trespass per territory was not consistently correlated with male density, and trespass rate per unit area was positively correlated with male density. Further, these relationships were retained when we controlled statistically for the effects of territory quality. We conclude that prospecting male red-winged blackbirds to not attempt to avoid densely settled areas, and that, although they do avoid territories defended by many song types, they do not use song type density to assess the density of territorial males. It is thus unlikely that the Beau Geste hypothesis adequately accounts for the evolution of song repertoires in the red-winged blackbird.     

Correlates of male mating success in two species of Anthidium bees (Hymenoptera : Megachilidae)

Summary Field observations were made on the mating behavior of two congeneric species of solitary bees, Anthidium porterae in an arid grassland and A. palliventre in a coastal sand dune habitat. Males of both species exhibited resource defence polygyny and defended hostplants to gain access to females foraging for nectar and pollen. The mating frequencies of marked and measured resident (territorial) males were monitored during periods of continuous observation, following which measurements of territory size and floral resources were obtained. Mating success of A. palliventre males was strongly influenced by territory characteristics: Males that defended small areas with a few rich hostplant patches mated more often than males that held larger territories containing many hostplant patches of low floral density. Large males generally held high-quality sites and thus had a mating advantage over smaller individuals. In A. porterae, on the other hand, male mating success was unrelated to any measure of territory quality. Copulation frequency and male size were positively correlated, however, apparently due to the increased ability of large males to seize and hold females for mating. The two species also differed in the incidence of non-territorial, sneaky males. While absent in A. palliventre, sneaky males accounted for 12% of all mating observed in A. porterae. Males of A. porterae that displayed sneaky tactics mated, on average, as often as resident males. Offprint requests to: E.M. Villalobos     

Effectiveness of territorial polygyny and alternative mating strategies in northern fur seals, <Emphasis Type="Italic">Callorhinus ursinus</Emphasis>

We conducted a 6-year longitudinal behavioral and genetic investigation of a highly polygynous pinniped, the northern fur seal (Callorhinus ursinus), to determine the contribution of terrestrial polygyny to male fertilization success and to assess the occurrence of alternative mating strategies. Genetic samples from 37 adult males, 50 adult females, and 85 pups were collected and genotyped using five polymorphic microsatellite loci. Pup paternity was assigned using Cervus 2.0 at 99% confidence level. Paternity of 83 pups (98%) was assigned to terrestrial males who held territories or stayed temporarily in the study area during the breeding season when fertilization occurred. For 56 pups of which attendance records of their mothers were available, paternity of 45 pups (80%) was assigned to the associate males in whose territory their mothers stayed during the perioestrus period. In addition to defending breeding territories, territorial males have often been observed attempting to forcibly abduct adult females from adjacent territories (female stealing): We observed a total of 95 such cases, in which the stealers had significantly fewer females than the territorial males from whose territories they stole females. Our results indicate that terrestrial resource-defense polygyny is the major mating system in this species and that nonassociated paternity occurs mostly as a result of alternative mating strategies of less successful males. Male northern fur seals thus appear to adopt conditional alternative strategies that depend on their current social status to maximize their life-time reproductive success.     

Correlates of copulatory success in a fallow deer lek

Summary We studied the behavior and copulatory success of fallow deer (Cervus dama) males at a lek. We recorded 471 copulations in 3 years. Most males did not copulate. The top three males accounted for between 60 and 90% of copulations each year. Lek attendance time was the major determinant of male copulatory success, but territory location also affected the number of copulations achieved. Copulatory success was correlated with the number of females in a male's territory and possibly with dominance status, but not with fighting success or fighting rate. All males that defended lek territories were 5 years of age or older. Copulatory success may improve with age. Body condition appears to be an important determinant of male copulatory success, because only males in superior condition could defend a lek territory for up to 2 weeks. Males do not feed while defending lek territories. Foraging ability during the year probably determines condition at the onset of the rut. Females appear to choose mates at least partially on the basis of location, preferring males located near traditional routes. Females may ultimately select mates in the best body condition.     

The significance of hotspots to lekking topi antelopes (<Emphasis Type="Italic">Damaliscus lunatus</Emphasis>)

Controversy has surrounded the question of why lek-breeding has evolved in certain ungulate species. Can the behavior be explained simply by males mapping onto a female distribution that is determined by factors unrelated to mating? Or are leks created because estrous females distinguish between males and favor males who cluster? Here I address these questions by looking at spatial distribution in lekking topi antelopes (Damaliscus lunatus). Contrary to the predictions of a model assuming male clustering in the zone of maximum female range overlap, territories were highly clustered also within this zone, and lek size correlated positively with population density. In support of models derived from the ideal free distribution of males onto female dispersion, leks were in areas with high female density during the rut. However, models not taking into account both individual variation in male quality and female mate preferences failed to explain the extreme male clumping also within high density areas, which was revealed by a strongly male-biased sex-ratio on leks. Additional support for the female preference-based model came from the finding that estrous females concentrated onto leks. Female preference for clustered males may develop if males initially follow an ideal free distribution of unequal competitors with high quality males slightly clustered at density hotspots; positive feedback between female benefits of preference for clustered males and male benefits of clustering could lead to contraction of the territorial network and lek behavior. Thus only the female preference-based model correctly predicted a negative correlation between male mating rate and resource density.     

Body morphologies shape territorial dominance in the satyrine butterfly <Emphasis Type="Italic">Lethe diana</Emphasis>

Male butterflies compete over mating territories via aerial interactions. It has often been suggested that flight performance, and thus flight-related traits, would play an important role in butterflies’ contest resolution; however, most studies have not demonstrated a clear relationship between these traits and territory ownership in butterflies. Males of the satyrine butterfly Lethe diana compete over territories via linear chases, a behavior that is uncommon among butterflies: rather, contests of most butterfly species consist of circling flights. In the present study, some morphological and physiological traits of L. diana were compared between territory owners and intruders to search for traits correlated with resource holding power (RHP). Territory owners had higher flight-muscle ratio (FMR) and were heavier than intruders, indicating that flight performance plays a key role in their contest resolution. Contest duration was not affected by the owners’ FMR or body mass, and thus I obtained no evidence supporting the hypothesis that contestants assess their relative RHP before making a decision to retreat. I speculate that the fact that these traits had an effect on territorial status in L. diana was a result of the characteristics of the contest behavior of this butterfly. During their contests, males L. diana fly faster in their linear flights than males of most other butterflies, which engage in slower circling flights. Therefore, males of L. diana probably need higher flight performance capabilities in order to win their contests.     

How reliable are behavioral cues for assessment of male mating status in polyterritorial wood warblers,Phylloscopus sibilatrix?

It has been suggested that polyterritoriality in birds, i.e. mated males defending two spatially separated territories, deceives females into accepting polygyny against their best interest. Mated males singing on secondary territories should make it difficult for new arriving females to assess the mating status of unmated and mated males advertising to females. However, there have been objections to the deception hypothesis, suggesting that differences in territorial behavior between males of different mating status may make correct assessment possible. Polyterritorial males frequently leave their secondary territory to visit the female in the primary territory, thereby revealing their mating status to females in search of mates. The aim of this study is to investigate how reliable behavioral cues are in assessing the mating status of males in polyterritorial wood warblers. Our study shows that singing activity of mated males on secondary territories and unmated males was similar in the morning, while there was a significant difference in the afternoon. Unmated males spent, on average, more time on their territories than mated males spent on their secondary territories. However, there was considerable variation in both unmated and mated males in the frequency in which they left their territories. For instance, unmated males left their territories quite frequently early in the season compared to later. Hence, females must be able to interpret singing behavior and a male leaving his territory differently depending on time of the season and time of the day in order to correctly assess the mating status of advertising males. Estimating the increase in probability of finding an unmated male by using behavioral cues showed that females could increase the probability of finding an unmated male compared to random choice with respect to mating status. The uncertainty was, however, still considerable. The most important factor affecting the probability of finding an unmated male was the proportion of mated males on secondary territories compared to unmated males. Our study suggests that presence on a territory is not a reliable cue for the assessment of male mating status in polyterritorial wood warblers.     

A dynamic model of mating behavior in digger wasps: the energetics of male-male competition mimic size-dependent thermal constraints

I develop a state-based dynamic model of behavior to demonstrate that size-dependent differences in temperature tolerances are not necessary to account for the activity of small male digger wasps late in the day. In the model, males defend or patrol the nesting area, wait near nests, or feed away from the nesting area depending on time of day, energy reserves and size rank. I assume a large male competitive advantage, so mating opportunities decrease with size rank for territorial or patrolling males and are rare for all waiting males; the costs of patrolling or defense are higher than the costs of waiting. If energy reserves of all males are initially small, all males alternate feeding and territorial or patrolling behavior. If energy reserves are initially large, large males patrol or maintain territories until they risk starvation and leave the area to feed. At this time, smaller males that have conserved their resources by waiting and feeding may defend territories or patrol. I simulate the behavior of three populations representing two species of Microbembex by assuming large initial energy reserves for populations in which males were territorial and small initial reserves for populations in which males patrolled, and then convert the predicted time of activity to temperature using local regressions from field studies. Temporal patterns in the activity of large and small males were similar to those actually observed, and relationships between size and temperature predicted by the model corresponded to most observations and were sometimes positive. Thus, the delayed activity of smaller males does not correspond to activity at higher temperatures and is probably not attributable to size-dependent thermal tolerances, but may represent a temporal displacement of mating activity due to intra-sexual competition and mediated by energetics. The model makes testable predictions on the timing of feeding and depletion of energy reserves in relation to size and initial energy state, and suggests how differences among species may influence the temporal and spatial organization of male mating behavior. Received: 27 February 1997 / Accepted after revision: 26 July 1997     

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

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

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

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