Muscle esterase variation and size variation in the sand launce Ammodytes dubius

Esterase variation detected in homogenates of muscle tissue taken from the sand launce Ammodytes dubius in April, 1970 and 1971 is described. Eight alleles were found segregating at a single locus, and the frequencies of these alleles were homogenous in population samples taken on the Emerald Bank, off Nova Scotia, Canada, in successive years. When ordered by their electrophoretic mobility, the profile of the frequencies of these alleles was unimodal and symmetric. In both of the population samples, there were significant excesses of homozygotes and deficencies of heterozygotes. There was an association between genotype and size of individuals among homozygous genotypes; individual homozygous for the most common, intermediately-migrating allele were larger than homozygotes bearing alleles with relatively fast or slow mobilities.     

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.     

Testis size variation in frogs: testing the alternatives

While sperm competition may be a major factor affecting relative testis size in vertebrates as a whole, additional hypotheses have not been given much attention in frogs. This is important because sperm competition is relatively uncommon in frogs and relative testis size varies in frogs that do not have multiple-male breeding systems. This paper tests two additional hypotheses for differences in relative testis size among frogs: relative clutch size (number of eggs/snout vent length) and androgen level. Testis size was measured in 90 species of frogs belonging to five families. Relative testis size was found to be positively correlated with relative clutch size in species that lack sperm competition. Mean androgen levels of species also positively covaried with relative testis size. However, there was no correlation between relative testis size and level of male agonistic behavior among species, despite other work indicating that testosterone levels are positively correlated with agonistic behavior in at least some species. These findings suggest that a number of factors in addition to sperm competition are important in the evolution of testis size in male frogs. Received: 17 January 1997 / Accepted after revision: 23 June 1997     

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.     

Individual variation in the relationship between vigilance and group size in eastern grey kangaroos

The mean vigilance of animals in a group often decreases as their group size increases, yet nothing is known about whether there is individual variability in this relationship in species that change group sizes frequently, such as those that exhibit fission–fusion social systems. We investigated variability in the relationship between group size and vigilance in the eastern grey kangaroo (Macropus giganteus) by testing whether all individuals showed decreased vigilance with increased group size, as has been commonly assumed. We carried out both behavioural observations of entire groups of kangaroos and focal observations of individually recognised wild female kangaroos. As in other studies, we found a collective group-size effect on vigilance; however, individuals varied in their vigilance patterns. The majority (57%) of the identified individual kangaroos did not show significant group-size effects for any of the recorded measures of vigilance. The females that did not show a negative group-size effect were, on average, more vigilant than those females that did show a group-size effect, but this difference was not significant. We propose that some females exhibit higher levels of social vigilance than others, and that this social vigilance increases with group size, cancelling out any group-size effect on anti-predator vigilance for those females. Our results therefore suggest that only some prey individuals may gain anti-predator benefits by reducing their time spent scanning when in larger groups. The large amount of variation that we found in the vigilance behaviour of individual kangaroos highlights the importance of collecting and analysing vigilance data at the individual level, which requires individual recognition.     

Reproductive consequences of variation in the composition of howler monkey (Alouatta spp.) groups

At least three general categories of environmental pressure - predation, resource distribution, and demographics - shape the costs and benefits of group-living for animals. Among the demographic factors that influence individual survival and reproduction, the composition of social groups can play an important role. Census data drawn from 26 populations of howler monkeys (Alouatta spp.) were used to determine if the composition of groups explained variation in their reproductive performance. Each group's reproductive performance was estimated by calculating the difference between the observed number of immatures and the number expected from its population average. Of four group structure variables tested, only one - the residual of the adult and subadult sex ratio - was a consistent correlate of reproductive performance across the howler monkey populations. Groups with a greater proportion of adult and subadult males contained more juveniles than expected from the population average. I propose that the survival or retention of immatures within howler monkey groups depends in part on the behavior of resident males. Of particular importance, the relative proportions of resident males and females were more informative than the absolute number of males or females. On this basis, I evaluate the possible role of males in protection from predation, conspecific aggression, and resource competition. The techniques used here can also be used to forecast major changes in demographic structure within populations.     

Correlates of male mating success in the ruff Philomachus pugnax,a lekking shorebird

Summary Male ruffs (Philomachus pugnax), a lekking shorebird, can generally be divided into two morphs according to behavior and plumage coloration. Predominantly dark-colored, independent male ruffs defend small territories on a lek, whereas lighter colored satellite males are non-territorial and less site-faithful to a particular lek. The traits associated with the mating success of independent male ruffs were studied at two leks during two seasons on the island of Öland, Sweden in May and June of 1985 and 1987. Using multivariate analyses, three characteristics were found to be significantly related to mating success: high frequency of visits by satellites to an independent male's residence, consistency of lek attendance, and low rates of aggressive behavior. In contrast, mating success was unrelated to the degree of darkness of the independent male nuptial plumage, territory location on the lek, or rate of courtship displays. The use of multivariate analyses helped to resolve conflicting conclusions from previous studies which employed simple statistical analyses, or none at all. Experimental manipulations are suggested which will help to further determine the influence of female mate choice in this lekking system.     

Life history consequences of variation in age at primiparity in northern elephant seals

Summary The age when female northern elephant seals, Mirounga angustirostris, bear their first young varies from 2 to 6 years. At Año Nuevo, California, a group of 77 females, primiparous at age 3, had a lower survivorship rate to each successive year up to age 8 than a group of 98 females that deferred initial pupping until age 4. The difference in survivorship appears to be due to the greater relative energetic costs of gestation and lactation incurred by the earlier breeding females during a period in their development when growth is rapid. An alternate hypothesis for the difference in survivorship — that young primiparous females are in poor condition from birth-is untenable; females that pupped early in life were larger at weaning age (a correlate of condition) than females that were primiparous 1 year later.Models based on the data show that differential survival of seals that vary in age at primiparity has important consequences for population growth and life history strategies. The effect of age at primiparity on the rate of increase of populations varies with colony density and juvenile survivorship. The optimal life history strategy for female elephant seals under most conditions existing today, including those at Ano Nuevo during the study period, is to bear the first offspring at age 4. Primiparity at age 3 is projected to be favored when harem density is very low and weaning success and juvenile survivorship are high; postponement of first breeding to age 5 is expected at high harem densities with intense competition for breeding space. Offprint requests to: B.J. Le Boeuf     

Active mate choice at cock-of-the-rock leks: tactics of sampling and comparison

Summary The importance of active mate choice as a component of sexual selection remains controversial, despite recent studies identifying traits that correlate with male mating success. We analyzed the sampling sequences of individual female Guianan cock-of-the-rock, providing quantitative documentation of active female choice in this lekbreeding species. We compared observed sampling patterns with null models of random female movement and of movement weighted by the observed distribution of male success. The courts actually visited by individual females were significantly more clumped and centrally located than predicted by computer simulations of either random or weighted movement. Contrary to simplified hypotheses of female choice, hen cock-of-the-rock were not limited either to sequential comparisons or to threshold decision rules when selecting mates. Instead, they repeatedly sampled a subset of potential mates, using the pool-comparison tactic. Within this pool, females preferentially mated with the males ranked highest in terms of total courtship visits received, and rejected low-ranking males. Our results suggest complex interactions between patterns of female sampling and male dispersion in lek mating systems, and also provide insights into the constraints on female sampling in lek species.     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

Spatial and temporal variation in the size at maturity of the western rock lobster Panulirus cygnus George

The sizes at which female and male western rock lobster Panulirus cygnus become mature were examined over 32 years from records at six localities along the coast of Western Australia. The size of males at maturity was estimated from a subset of these data by a morphometric and a physiological method, with both producing statistically similar results. Males were larger at first maturity than females at the same sites. For both sexes, the sizes at first maturity at each location correlated (P<0.05) with the mean annual water temperature at that location, decreasing from south to north along the mainland coast and being smallest at the offshore Abrolhos Islands. Smaller sizes at maturity were recorded for both sexes than have been published previously. One certain explanation for these differences is that management measures protecting females with ovigerous setae have distorted size compositions and the ratio of immature to mature females, thereby increasing the likelihood of capturing small mature females. However, these fishing effects cannot fully account for the progressive decline in CL₅₀ observed over the past 20 years. Other possible hypotheses considered include increases in water temperature over this period, as well as whether this change could be consistent with a genotypic response caused by the selective removal of large lobsters combined with high exploitation rates.     

Sexual selection and cues for female choice in leks of Jackson's widowbird Euplectes jacksoni

Summary Sexual selection through female mate choice was investigated in the lekking Jackson's widowbird by applying multivariate selection analysis to observational data from four leks. Males perform a stereotyped jump display on small display courts (dance rings) constructed by the males in open grassland. Females visit the lek solely for mating and nest on their own, away from the lek area. Few cases of interference during courtship and absence of position effects on mating success indicated that female choice within the leks was not pre-empted by male-male competition. In a set of 11 male traits with mating success as the dependent fitness measure, significant selection differentials (covariances) were found for the length of the conspicuous tail and the rate of the jump display, suggesting sexual selection of these traits. They also showed the largest selection gradients (partial effects) and thereby seem to be the cues on which females base their choice. The success of males in obtaining copulations appears to depend on two components: display rate and lek attendance affect the number of female visits, whereas tail length seems to primarily influence the chance of copulating with a visiting female. Tail length was positively related to a measure of body condition, which is of interest with regard to the suggestions that sexual ornaments may serve as indicators of male viability.     

Imperfect female choice and male mating skew on leks of different sizes

We present a model of error-prone female choice on leks, and investigate the effects of different degrees of error on the distribution of mating success among males present at leks of different sizes. At higher levels of error, the best male is predicted to gain a smaller share of matings, while low-ranking males gain a larger share. Males who are of high rank but not the most desirable on the lek do best at intermediate levels of error, since the top-ranked male does not then claim all the matings, but assessment is still sufficiently accurate for females to discriminate between high-ranking and low-ranking competitors. The effects of error are shown to be more pronounced on larger leks, due to smaller expected differences in mating value between males of adjacent ranks. This interaction between lek size and error suggests that observed negative relationships between lek size and mating skew need not be attributed solely to intrasexual competition, as previously suggested, but could also be a result of imperfect choice. Received: 20 February 1998 / Accepted after revision: 25 October 1998     

Spatial proximity loggers for recording animal social networks: consequences of inter-logger variation in performance

Social network analysis has become an increasingly popular method to link individual behaviour to population level patterns (and vice versa). Technological advances of recent years, such as the development of spatial proximity loggers, have enhanced our abilities to record contact patterns between animals. However, loggers are often deployed without calibration which may lead to sampling biases and spurious results. In particular, loggers may differ in their performance (i.e., some loggers may over-sample and other loggers may under-sample social associations). However, the consequences of inter-logger variation in logging performance has not been thoroughly considered or quantified. In this study, proximity loggers made by Sirtrack Ltd. were fitted to 20 dairy cows over a 3-week period. Contact records resulting from field deployment demonstrated variability in logger performance when recording contact duration, which was highly consistent for each logger over time. Testing loggers under standardised conditions suggested that inter-logger variation observed in the field was due to a combination of intrinsic variation in devices, and environmental/behavioural effects. We demonstrate the potential consequences that inter-logger variation in logging performance can have for social network analysis; particularly how measures of connectivity can be biased by logging performance. Finally, we suggest some approaches to correct data generated by proximity loggers with imperfect performance, that should be used to improve the robustness of future analyses.     

Geographic variation in maternal investment: acidity affects egg size and fecundity in Rana arvalis

Environmental-stress-mediated geographic variation in reproductive parameters has been little studied in natural vertebrate populations outside the context of climatic variation. Based on life-history theory, an increase in the degree of environmental stress experienced by a population should lead to (1) a shift in reproductive allocation from fecundity to offspring quality, (2) stronger trade-offs between reproductive parameters, and (3) changes in the relationship between female phenotype and maternal investment. To test these predictions, we investigated geographic variation in maternal investment of moor frogs (Rana arvalis) in relation to breeding site acidity (pH 4-8). We found that mean egg size increased and clutch size and total reproductive output (TRO) decreased with increasing acidity among 19 Swedish moor frog populations. Tests for variation and co-variation in maternal investment and female size and age in 233 females from a subset of four acid origin (AO) and four neutral origin (NO) populations revealed that clutch size and TRO increased with female size in both acid and neutral environments. However, in AO populations, egg size also increased with female size, and clutch size and TRO with female age, whereas in NO populations, egg size increased with female age. The strength of the egg-size-clutch-size tradeoff tended to be stronger in AO than in NO females as expected if the former experience stronger environmental constraints. All in all, these results suggest that environmental acidification selects for investment in larger eggs at a cost to fecundity, imposes negative effects on reproductive output, and alters the relationship between female phenotype and maternal investment.     

Offspring size variation within broods as a bet-hedging strategy in unpredictable environments

Offspring size is strikingly variable within species. Although theory can account for variation in offspring size among mothers, an adaptive explanation for variation within individual broods has proved elusive. Theoretical considerations of this problem assume that producing offspring that are too small results in reduced offspring viability, but producing offspring that are too large (for that environment) results only in a lost opportunity for increased fecundity. However, logic and recent evidence suggest that offspring above a certain size will also have lower fitness, such that mothers face fitness penalties on either side of an optimum. Although theory assuming intermediate optima has been developed for other diversification traits, the implications of this idea for selection on intra-brood variance in offspring size have not been explored theoretically. Here we model the fitness of mothers producing offspring of uniform vs. variable size in unpredictably variable environments and compare these two strategies under a variety of conditions. Our model predicts that producing variably sized offspring results in higher mean maternal fitness and less variation in fitness among generations when there is a maximum and minimum viable offspring size, and when many mothers under- or overestimate this optimum. This effect is especially strong when the viable offspring size range is narrow relative to the range of environmental variation. To determine whether this prediction is consistent with empirical evidence, we compared within- and among-mother variation in offspring size for five phyla of marine invertebrates with different developmental modes corresponding to contrasting levels of environmental predictability. Our comparative analysis reveals that, in the developmental mode in which mothers are unlikely to anticipate the relationship between offspring size and performance, size variation within mothers exceeds variation among mothers, but the converse is true when optimal offspring size is likely to be more predictable. Together, our results support the hypothesis that variation in offspring size within broods can reflect an adaptive strategy for dealing with unpredictably variable environments. We suggest that, when there is a minimum and a maximum viable offspring size and the environment is unpredictable, selection will act on both the mean and variance of offspring size.     

Experimental evidence for the effects of chronic anthropogenic noise on abundance of Greater Sage-Grouse at leks

Increasing evidence suggests that chronic noise from human activities negatively affects wild animals, but most studies have failed to separate the effects of chronic noise from confounding factors, such as habitat fragmentation. We played back recorded continuous and intermittent anthropogenic sounds associated with natural gas drilling and roads at leks of Greater Sage-Grouse (Centrocercus urophasianus). For 3 breeding seasons, we monitored sage grouse abundance at leks with and without noise. Peak male attendance (i.e., abundance) at leks experimentally treated with noise from natural gas drilling and roads decreased 29% and 73%, respectively, relative to paired controls. Decreases in abundance at leks treated with noise occurred in the first year of the study and continued throughout the experiment. Noise playback did not have a cumulative effect over time on peak male attendance. There was limited evidence for an effect of noise playback on peak female attendance at leks or male attendance the year after the experiment ended. Our results suggest that sage-grouse avoid leks with anthropogenic noise and that intermittent noise has a greater effect on attendance than continuous noise. Our results highlight the threat of anthropogenic noise to population viability for this and other sensitive species.     

Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort

We examined how mating success varied in relation to age, weight, body size, and display behavior among great bustard Otis tarda males. The estimated mating success was strongly skewed, with 45% of adult males being involved in copulation attempts and only 9.7% actually seen copulating successfully. Unlike most birds, body size continued increasing in great bustards several years after reaching sexual maturity. Age, weight, and display effort were all significant and independent predictors of male mating success. The higher display effort involved performing longer full-display bouts. Older males could detach from the male flock earlier in the season as well as on each day and spend longer seasonal and daily periods displaying as solitary birds, which contributed to increase their mating success. In contrast, males weighing more did not invest more in display, which suggests that they could be recognized as dominants by other males and selected by females through assessment of their plumage sexual traits. In contrast to most other bird species, the system described for great bustards resembles that found in some lek-mating ungulates, where social rank is a complex trait determined by both age and mass, and as in these mammals, it suggests that sexual selection continues to favor a high male weight in this extremely sexually dimorphic species.     

The evolution of leks through female choice: differential clustering and space utilization in six sympatric manakins

Summary The degree to which lekking and non-lekking male manakins select display sites in order to maximise proximity to females was examined by contrasting movements of females with male dispersion. Data on female visiting patterns, male courtship disruption, and mating skew were also collected over three successive breeding seasons. For the five lek-breeding species, female home-ranges were 3–7 times larger than those of adult males. Female movements were concentrated around leks, fruiting places and stream bathing sites. None of the females monitored by radio-tracking expanded her normal range in order to visit males on leks. On the contrary, feeding bouts of females frequently preceded a visit to potential mates at neighboring leks. Despite small sample sizes, significant correlations were found between female home-range size and male clustering (distances between neighboring leks and distances between neighboring males), as predicted by the female choice model and the hotspot model. Adult and immature male home-range sizes were not significantly correlated with male dispersion or female ranges. On the other hand, males and females of the only non-lekking species exhibited similar use of space and home-range size. Male settlement at sites with high levels of female traffic showed that the hotspot model is adequate to explain differences in male dispersion among sympatric lekking species. Comparisons with other studies suggest that apparent female choice could be overidden by past and present male-male interactions or female mate-comparison tactics. In fact, both the hotspot model and the attractiveness hypothesis appear to shape male dispersion on leks: males appear to settle under hotspot conditions with despotic rules generated through bias in female choice or male-male interference. It is proposed that the evolution of leks is ecologically motivated by the spatio-temporal distribution of trophic resources, initially leading to a dispersed male-advertisement polygyny. Following this, a foraging ecology that promotes high mobility by females and the magnetic effect of mating skew in particular males may have favored clustering on exploded leks. Later, the development of male-male interference and the increasing female home-range size could have led to the evolution of classical leks.