Spacing of leks in relation to female home ranges,habitat requirements and male attractiveness in the great snipe (Gallinago media)

Summary In an attempt to examine three main hypotheses on the evolution of leks, data on female home ranges, distance between leks and male site fidelity in the great snipe (Gallinago media) were obtained. In a 30 × 40 km area in central Sweden, the locations of 12 leks were identified. Six were within the study area (7 × 11 km) in which probably all leks were known. In the study area, nearest neighbor distances between leks were longer than the predicted drawing area (diameter of female home ranges plus the detection range of leks). Furthermore, in only 1 of 13 radio-tracked females did the estimated home range enclose many leks. These data do not fit the prediction from the hotspot hypothesis on the dispersion of leks, namely, that the distance between leks should be less than the drawing area and hence an average female home range should enclose more than one lek. The observed distance between leks was about the same as predicted by the female preference hypothesis. This hypothesis also predicts females visit mainly one lek; however, we found they sometimes visit two and nest close to a third. We propose that males may first settle according to hotspot rules, but females will resettle according to preferences for certain males and/or larger leks. In this way some hotspot leks are abandoned, and the distance between leks is increased as males become more tightly clumped. However, when all males and females have settled, the leks still existing would be on hotspots. In the female preference hypothesis, males are assumed to aggregate on leks because females prefer clustered males as they can be more easily compared than if they are dispersed. Alternatively, in the attractiveness hypothesis, it is suggested that females prefer certain males, and hence unattractive males surround the attractive are forced to join leks in order to come close to females. Our data on male site fidelity support the attractiveness hypothesis because successful and dominant males return to the same lek and territory, both within and between years, whereas unsuccessful males move to other leks. Attractiveness of certain males may explain why according to hotspot rules in this species males are more clumped than expected. Offprint requests to: J. Höglund     

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     

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.     

Space use and social structure in meadow voles,Microtus pennsylvanicus

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

Testing hotspot models of lek evolution: data from three species of ungulates

Summary Hotspot models propose that leks have evolved because certain sites are associated with extremely high female encounter rates. In this paper we test five predictions of hotspot models, using data from studies of three species of lekking ungulates. As predicted, we found that lekking species generally had large female home ranges. However, in contrast to other predictions of hotspot models, ungulate leks did not occur at sites of maximum adult female density, lekking was not associated with low overall female density, and the adult sex ratio and the proportion of females that were in oestrus both differed substantially between leks and off-lek areas. Our empirical results therefore confirm recent theoretical assessments of the hotspot process, suggesting that while hotspots may help explain broad patterns of male dispersion, further mechanisms are needed to generate the extent of territory clustering seen at leks. Correspondence to: A. Balmford     

Bat predation and the evolution of leks in acoustic moths

Theories of lek evolution generally invoke enhanced mating success experienced by males signalling in aggregations. Reduced predation has also been acknowledged as a potential factor driving lek formation, but its role is more ambiguous. Although lekking is a complex behaviour, few empirical studies have investigated the role of both claims. We studied the potential pressures imposed by mating success and predation in an acoustic moth, Achroia grisella, in which males gather in leks and broadcast a calling song attractive to females. We exploited the ability to manipulate the distribution of singing males in laboratory arenas to create different-sized leks and tested female preferences for these aggregations. Because A. grisella are vulnerable to predation by bats while in flight and on the substrate, we also tested the responses of a potential predator, Rhinolophus ferrumequinum, a bat species that feeds on moths, to the experimental leks. We found that the per capita attractiveness of A. grisella males to females rose with increasing lek size. R. ferrumequinum also oriented toward experimental A. grisella leks, but this attraction did not increase at larger leks. Thus, a male’s per capita exposure to predation risk declined as more moths joined the lek. A. grisella males appear to benefit from advertising in larger leks in terms of both increased mate attraction and reduced predation risk. Our results support the idea that multiple factors operating simultaneously may maintain lekking behaviour.     

An estimate of female mate searching costs in the lekking butterfly Coenonympha pamphilus

We estimated the cost to females of the lekking butterfly Coenonympha pamphilus of visiting males on leks instead of taking off and soliciting courtship from males passing them outside leks, as occurs in the non-lekking congener C. tullia. We followed released virgin females of C. pamphilus in the field until they mated. We estimated the time they lost by remaining perched and not approaching males that passed them, and observed how often and at what distance virgin females were passed by males in the field. We then calculated how much faster these females would have been detected if they had taken off and approached these males, using probabilities of detection as a function of passing distance derived from field observations on C. tullia females. C. pamphilus females on average lost 201 min by not approaching males on their way to leks. To estimate what this time loss meant in terms of fitness, we measured how the age at mating affected fecundity. Using field measures of fecundity and mortality, the time loss translated into an average 2.8% reduction in fecundity as a best estimate, and an average 1.3% reduction in fecundity as a lower estimate. This fitness cost is larger than has been reported earlier for a lekking bird, but is probably too small to eliminate the possibility of indirect benefits of mating with males on leks. Received: 15 August 1996 / Accepted after revision: 24 January 1997     

Influence of territoriality on adult density in two rockfishes of the genus Sebastes

Nonterritorial Sebastes carnatus and S. chrysomelas existed, along with territorial individuals, at 3 tagging sites off southern California, USA, which were monitored for nearly 1 yr. To test the hypothesis that territoriality affected adult density in these species, territorial fish were removed and the subsequent utilization of vacated territories by other fish was monitored. Intrusion of other fish into vacated territories increased significantly in 90% of the removals. Other fish colonized both the feeding and sheltering parts of the vacated territories, indicating that the previous owners had successfully defended both parts of their territories. Many of the colonizers had already possessed territories; they expanded their territories or moved into presumably better havitat. Several previously-nonterritorial fish also moved into vacated areas, and at least some of them appeared to establish territories. These fish, then, had previously been capable of establishing territories, but were prevented from doing so by resident territory holders. Thus territoriality, rather than such other factors as predation or low recruitment, limited the number of territorial fish at each site. However, territorial fish did not inhibit the settlement of larval recruits, and the relative mortality rates of older territorial vs nonterritorial fish were not determined. Thus the question of whether territoriality was a major factor controlling total density remains unresolved.     

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.     

Dominions and site-fixed aggressive behavior in breeding male Leucorrhinia intacta (Odonata: Libellulidae)

Summary We report aggressive spacing behavior in male dragonflies, Leucorrhinia intacta, that is characterized by variations in the probability of chasing conspecific male intruders within a defined area around a male's perch. The chase probability depends on the total intruder pressure and the behavior and distance of the intruder from the perched male (Fig. 2). This nonexclusive, site-fixed area has been called a dominion.We also examined the distribution of intruders among the various behavior-distance categories. Chase rates (per 15 min) were correlated mostly with number of intruders that hovered (rather than flew) close to the territorial male. Finally, we examined the impact of the changing chase probabilities on the variation in aggressive interactions as a function of intruder pressure. We conclude that, to some extent, territorial males stabilized defense costs across a variety of intruder pressures by maintaining dominions.     

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.     

Abnormal reproduction of the sand crab Emerita analoga in the vicinity of a nuclear generating station in Southern California

Female sand crabs (Emerita analoga) were sampled from 12 beaches along a 350 km stretch of Southern California coastline during the reproductive season in July, 1982. The percentage of mature females with egg masses and the condition of those egg masses were examined at each site. The proportion of mature females with egg masses varied only slightly between beaches. However, at sites within about 20 km of the San Onofre Nuclear Generating Station, a high percentage of females carried egg masses consisting of empty egg shells. A transplant experiment was conducted. Females with empty egg masses were taken from the 6.5 km N study site, where only 5% of the mature females carried normal egg masses, and transferred to running unfiltered seawater in Santa Barbara. After 2 wk, about a third of the transplanted females carried normal egg masses, while at the same time the proportion of mature females with normal egg masses remained at 5% on the beach of origin. Heated water discharged from the nuclear generating station is probably not the cause of the abnormal reproduction of sand crabs along the nearby coast. More likely causes are: (1) runoff of agricultural pesticides from a creek 3 km north of the nuclear generating plant; (2) release of metals from corroding cooling pipes, and/or (3) increased turbidity of the nearshore waters.     

Dispersion of displaying male sage grouse

Summary The degree to which male sage grouse select lek sites and females select nesting sites to maximize proximity to the other sex was examined by contrasting male dispersions with the dispersions and movements of females in the months preceeding incubation. Wintering females exhibit highly overlapping ranges due to shared use of central refuging areas. In late winter and early spring, females move an average 9 km from wintering areas to select nest sites and males begin occupying leks. Pooled evidence suggests that females select nest sites independently of male dispersion whereas males adjust lek occupation so as to maximize proximity to females. Relevant observations include females visiting nest sites before leks, moving further to select a nest site than to select a lek, and increasing their distance to leks as a result of selecting nest sites. In addition, males avoid leks until females have moved to within 5 km of the arenas, abandon early season leks as local female densities drop, and exhibit dispersions in which mean ratios of females/male are similar across leks. Contrasts between predicted and observed dispersions of males showed that hotspot settlement models are adequate to explain male dispersions on very coarse scales (2 km or greater); on finer scales, habitat preferences of males and tendencies for males to cluster tightly must be invoked in addition to hotspots to explain specific lek sitings.     

Scales of similarity in soft sediment epibenthic assemblages: implications for marine protected area design

Remote videography was used to investigate relationships between biological similarity and distance in Moreton Bay, Australia at site (1 km) and local (10 km) scales, and for separate biotic groups. Numerical analysis using Mantels tests to compare distance and similarity matrices showed that at both scales there was a negative correlation between similarity and distance, in that sites further apart were less similar than sites close together. The relationship, although significant (p <0.001), was quite weak (R²=5%) at the site-scale, with no significant (ANOVA with Tukeys pairwise comparisons, p >0.05) decline in similarity up to distances of 2.1–2.6 km. At the local-scale, between-site similarity was high (mean Bray-Curtis similarity >30% for 4th root transformed data) at scales of 10 km or less, and declined markedly with increasing distance. Scales of similarity for different broad taxonomic and functional groups within Moreton Bay were broadly consistent between groups and with the complete dataset. There was evidence of patchiness in the distributions of seagrass and anthozoans at scales less than 16 km. In other biotic groups there was an essentially monotonic decline in similarity with distance. The study showed that the spatial classification approach to habitat mapping is valid in this case, and that site spacing of less than 10 km is necessary to capture important components of biological similarity. Site spacing of less than 2.5 km does not appear to be warranted to capture additional components of biological similarity at the scales studied.Communicated by G.F. Humphrey, Sydney     

Mating success on white-bearded manakin (Manacus manacus) leks: male characteristics and relatedness

Morphological, territorial, and behavioural characteristics were measured on white-bearded manakin (Manacus manacus) males from two leks in two consecutive years. These data were combined with data on marker-inferred relatedness to study possible co-variation with mating success. In one year, male size and male condition were correlated with mating success. In both years, males holding courts nearer the lek centre gained more matings. No observed male display behaviour appeared to be an independently important factor in explaining variance in male mating success. Successful males made more aggressive displays than non-successful males and more displays were between close relatives. Number of aggressive displays increased as the distance between male courts decreased. Mating success in the white-bearded manakin is most likely mediated by a combination of morphological and behavioural characteristics, influencing both male-male competition and female choice. Females could potentially use centrality on the lek as an indicator of male characteristics. However, levels of relatedness may influence spatial arrangement of males on a lek thereby affecting male-male interactions and ultimately influencing patterns of mating success.     

Territorial User Rights for Fisheries as Ancillary Instruments for Marine Coastal Conservation in Chile

Territorial user rights for fisheries have been advocated as a way to achieve sustainable resource management. However, few researchers have empirically assessed their potential as ancillary marine conservation instruments by comparing them to no‐take marine protected areas. In kelp (Lessonia trabeculata) forests of central Chile, we compared species richness, density, and biomass of macroinvertebrates and reef fishes among territorial‐user‐right areas with low‐level and high‐level enforcement, no‐take marine protected areas, and open‐access areas in 42 100‐m subtidal transects. We also assessed structural complexity of the kelp forest and substratum composition. Multivariate randomized permutation tests indicated macroinvertebrate and reef fish communities associated with the different access regimes differed significantly. Substratum composition and structural complexity of kelp forest did not differ among access regimes. Univariate analyses showed species richness, biomass, and density of macroinvertebrates and reef fishes were greater in highly enforced territorial‐user‐right areas and no‐take marine protected areas than in open‐access areas. Densities of macroinvertebrates and reef fishes of economic importance were not significantly different between highly enforced territorial‐user‐right and no‐take marine protected areas. Densities of economically important macroinvertebrates in areas with low‐level enforcement were significantly lower than those in areas with high‐level enforcement and no‐take marine protected areas but were significantly higher than in areas with open access. Territorial‐user‐right areas could be important ancillary conservation instruments if they are well enforced. Derechos de Usuario Territoriales para Pesquerías como Instrumentos Accesorios para la Conservación Marina Costera en Chile     

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.     

Examination of model predictions at different horizontal grid resolutions

While fluctuations in meteorological and air quality variables occur on a continuum of spatial scales, the horizontal grid spacing of coupled meteorological and photochemical models sets a lower limit on the spatial scales that they can resolve. However, both computational costs and data requirements increase significantly with increasing grid resolution. Therefore, it is important to examine, for any given application, whether the expected benefit of increased grid resolution justifies the extra costs. In this study, we examine temperature and ozone observations and model predictions for three high ozone episodes that occurred over the northeastern United States during the summer of 1995. In the first set of simulations, the meteorological model RAMS4a was run with three two-way nested grids of 108/36/12 km grid spacing covering the United States and the photochemical model UAM-V was run with two grids of 36/12 km grid spacing covering the eastern United States. In the second set of simulations, RAMS4a was run with four two-way nested grids of 108/36/12/4 km grid spacing and UAM-V was run with three grids of 36/12/4 km grid spacing with the finest resolution covering the northeastern United States. Our analysis focuses on the comparison of model predictions for the finest grid domain of the simulations, namely, the region overlapping the 12 km and 4 km domains. A comparison of 12 km versus 4 km fields shows that the increased grid resolution leads to finer texture in the model predictions; however, comparisons of model predictions with observations do not reveal the expected improvement in the predictions. While high-resolution modeling has scientific merit and potential uses, the currently available monitoring networks, in conjunction with the scarceness of highly resolved spatial input data and the limitations of model formulation, do not allow confirmation of the expected superiority of the high-resolution model predictions.The U.S. Governments right to retain a non-exclusive royalty-free licence in and to any copyright is acknowledged.     

Sexual selection favours small and symmetric males in the polygynous greater sac-winged bat Saccopteryx bilineata (Emballonuridae, Chiroptera)

We investigated how morphological traits of territorial males in the polygynous bat Saccopteryx bilineata were related to their reproductive success. Because of the frequency of aerial courtship displays and defence manoeuvres, and the high energetic costs of flight, we expected small and symmetric males to be better able to court females on the wing and to monopolize copulations with females in their harems. We predicted that small and symmetric males would sire more offspring within the colony and a larger portion of the young born within their harem than large or asymmetric males. We measured size and fluctuating asymmetry of 21 territorial males and analysed their reproductive success in 6 offspring cohorts (n=209 juveniles) using 11 microsatellite loci. As predicted, small and symmetric males had, on average, a higher reproductive success in the colony than large and asymmetric males. The percentage of young sired by males within their harem increased as males decreased in size, but was not influenced by fluctuating asymmetry. As fluctuating asymmetry of males correlated with their reproductive success within the colony but not within their harems, we infer that fluctuating asymmetry is probably related to female choice, whereas male size is probably important for harem defence on the wing.Communicated by G. Wilkinson     

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