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     

Ecological and social determinants of spacing behaviour in desert baboon groups

This paper investigates the determinants of individual spacing behaviour in a desert baboon population (Papio cynocephalus ursinus). Patterns of neighbour proximity and neighbour density were examined among adults in four groups under different ecological and social conditions (through instantaneous sampling during focal follows). Initial analysis of these data shows that (1) the use of vertical substrates (refuges such as tall trees and cliff faces) can confound patterns of spacing, and (2) individual differences in spacing can depend on the spatial scale over which it is measured. To minimise these substrate and scale effects, this analysis focuses on animals which are off refuges and examines spacing behaviour through its underlying statistical `dimensions' (identified through factor analysis). Analysis of these dimensions indicates that sex, group size, activity-habitat and female reproductive state can all have independent effects on spacing: (1) males are more dispersed than females in smaller groups, (2) male and female dispersion increases with time spent in foraging habitats, and (3) female dispersion is reduced during lactation. According to the hypotheses tested, these results indicate that feeding competition only affects spacing behaviour during foraging while predation risk plays little or no role in spacing. Most aspects of spacing behaviour are best explained by male reproductive strategies and their social repercussions. Received: 25 May 1998 / Accepted after revision: 18 July 1998     

Disappearance and dispersal of Spermophilus elegans juveniles in relation to behavior

Summary Patterns of disappearance and dispersal of Spermophilus elegans juveniles during the first 6-weeks postemergence were compared for 1977 and 1978 and related to quantitative and qualitative changes in social interaction involving juveniles.Juvenile disappearance (emigration or mortality) and dispersal within the study site varied between the sexes within and between years. Female disappearance and dispersal were significantly greater in 1977, and male losses in 1978 significantly exceeded male losses in 1977. Greater female loss in 1977 resulted in total male — female losses being equivalent, whereas in 1978 juvenile loss was strongly biased toward males by the end of the 6-week period.Greater female loss in 1977 was attributed primarily to increased aggression between female juveniles in that year because of larger average litter size with more females per litter. Increased disappearance of males in 1978 showed no correlation with litter size or relative increase in number of males per litter. Male young interacted with individuals of several age/sex classes, and a possible behavioral influence on male disappearance was increased aggressiveness by yearling males toward juvenile males in 1978.Behavior appeared to act as a proximate factor in juvenile disappearance and dispersal, and the observed differences between how male and female juveniles interacted in 1977 and 1978 were hypothesized to reflect the operation of different selective pressures to increase individual male or female fitness.     

Nomadic behavior of army ants in a desert-grassland habitat

Summary Nomadic behavior of the army ant Neivamyrmex nigrescens was studied in a desert-grassland habitat. Six colonies were followed through eight nomadic phases (94 nomadic days) while direction and distance of emigrations, growth of larvae, number of adults and larvae per colony were determined.In all colonies, the nomadic phase began when newly eclosed adults and small larvae were present, and ended when the larvae were fully grown. Average emigration distance was positively related to number of larvae in the colony. These findings support Schneirla's theory that brood stimulation is a proximate cause of the nomadic phase.Failures to emigrate were equally likely at all points in the nomadic phase, and there was no systematic increase in emigration distance as the phase progressed. These findings do not support Schneirla's version of brood-stimulative theory.Number of adults per colony was positively related to the directionality of the nomadic phase; however, both the direction and distance of emigrations varied unpredictably from one nomadic day to the next, in marked contrast to predictions from optimal foraging theory.Schneirla's theory is useful in predicting phase differences in colony behavior, but it does not account for characteristics such as frequency, direction, or distance of emigrations within the nomadic phase. These aspects of nomadic behavior are more closely related to characteristics of the habitat such as prey density and availability of nest sites. Army ant nomadism in this habitat may depart from the optimum because of high prey density, small colony size, or lack of nesting sites.     

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.     

Signaling in multiple modalities in male rhesus macaques: sex skin coloration and barks in relation to androgen levels, social status, and mating behavior

The past decade has seen an increasing shift in animal communication towards more studies that incorporate aspects of signaling in multiple modalities. Although nonhuman primates are an excellent group for studying the extent to which different aspects of condition may be signaled in different modalities, and how such information may be integrated during mate choice, very few studies of primate species have incorporated such analyses. Here, we present data from free-ranging male rhesus macaques on sex skin coloration (modeled to receiver perception), bark vocal signals, androgen levels, morphometric variables, dominance status, and female mate choice. We show that, consistent with data on females, most intra- and interindividual variation in sex skin appearance occurs in luminance rather than color. Sex skin luminance was significantly correlated across different skin regions. Sex skin luminance did not correlate with the majority of bark parameters, suggesting the potential for the two signals to convey different information. Sex skin appearance was not related to androgen levels although we found some evidence for links between androgen levels and bark parameters, several of which were also related to morphometric variables. We found no evidence that either signal was related to male dominance rank or used in female mate choice, though more direct measures of female proceptive behavior are needed. Overall, the function of male sex skin coloration in this species remains unclear. Our study is among the first nonhuman primate studies to incorporate measurements of multiple signals in multiple modalities, and we encourage other authors to incorporate such analyses into their work.     

Costs and consequences of variation in the size of ruff leks

Summary We studied 13 ruff leks in a small region on the island of Gotland (Sweden) to investigate the effect of lek size on the costs and benefits of lekking for individual males. Male ruffs occur in two behaviourally and morphologically distinct forms, independents (residents plus marginals) and satellites, whose costs and benefits we have assessed separately. These ruff leks had from 1–10 resident (territory-holding) males and were visited daily by satellites, marginals and females from 5–25 May, when most copulations occurred. We used the average number of independent males, counted during censuses taken every 5 min during 2-h observation periods at each lek, as an index of mean lek size. Per independent male, the numbers of both satellites and females increased significantly with mean lek size. Female arrival rate and attendance (total female-minutes) also increased significantly with mean lek size as did the average per capita rate of mating success for resident males (that of satellites was not quite significant). Thus, the dispersion of both of these male categories did not appear to fit an ideal free distribution with respect to mating success. In addition, the number of independent-independent fights per independent and the rate of satellite-resident dyad formation per resident increased significantly with mean lek size. These results suggest that ruffs on larger leks enjoy higher mating success than those on smaller leks but also that costs increase with lek size. We suggest that independent males distribute themselves so as to maximize their own net benefits and that this factor can account for both the occurrence of ruff leks and the variation in their size. Correspondence to: J. Höglund     

Male aggression varies with consortship rate and habitat in a dolphin social network

Behavioral Ecology and Sociobiology - In many species, outcomes of male duels determine access to females and, ultimately, male reproductive success. Ritualization of behavior in male contests can...     

Egg composition in relation to social environment and maternal physiological condition in the collared flycatcher

Offspring survival can be influenced by resources allocated to eggs, which in turn may be affected by the environmental factors the mother experiences during egg formation. In this study, we investigated whether experimentally elevated social interactions and number of neighbouring pairs influence yolk composition of collared flycatchers (Ficedula albicollis). Social challenge was simulated by presentation of a conspecific female. Experimental females spent more time near the cage and produced eggs with higher androgen concentration, but local breeding density did not affect yolk androgen level. Moreover, we found that females exposed to more intra-specific interactions and those that bred at higher density produced eggs with smaller yolk. These females may be more constrained in foraging time due to more frequent social encounters, and there might be increased competition for food at areas of higher density. In contrast, the present study did not reveal any evidence for the effect of social environment on yolk antioxidant and immunoglobulin levels. However, we found that yolk lutein and immunoglobulin concentrations were related to the female’s H/L ratio. Also, yolk lutein and α-tocopherol levels showed a seasonal increase and were positively related to the female’s plasma carotenoid level. Mothers may incur significant costs by transferring these compounds into the eggs, thus only females in good physiological condition and those that lay eggs later, when food is probably more abundant, could allocate higher amounts to the eggs without compromising their defence mechanisms. Our results suggest that environmental circumstances during egg formation can influence conditions for embryonic development.     

Predatory behavior and prey selection by army ants in a desert-grassland habitat

Summary Colonies of Neivamyrmex nigrescens conduct extensive nocturnal raids on other ants and termites in the desert-grassland of Arizona-New Mexico. We collected quantitative data on several aspects of raiding to pinpoint differences due to colony size and behavioral phase. In the nomadic phase, colonies began raiding at sunset and continued until dawn. Larger colonies covered more area, discovered more prey sites, and collected more booty than smaller colonies, but there were no systematic changes in raid intensity over the course of the nomadic phase. In the statary phase, raiding occurred less frequently and was less intense when it occurred; however, at the end of this phase, raiding was similar to nomadic phase raids in extent, duration, and booty captured. N. nigrescens preyed exclusively on termites and ants, and appeared to select certain species of Pheidole in preference to other ants. Pheidole was the most abundant genus, but was preyed upon twice as often as expected based on relative colony density. Pheidole attempted to avoid predation by fleeing or defending their nest, but rarely succeeded. Because they are about the same size as army ants and lack defensive chemicals, Pheidole made comparatively easy prey. N. nigrescens ignored or was repelled by other ants (Pogonomyrmex, Novomessor, Iridomyrmex, Myrmecocystus) during the early summer, when Pheidole was abundant; however, in late summer when Pheidole was less available, the army ants preyed upon Novomessor cockerelli. N. harrisi raided in close proximity to N. nigrescens, but preyed exclusively on Solenopsis xyloni. Selection of prey and partitioning of resources are now indicated in several army ant species; these processes have probably been important factors in the evolution of the ants' predatory behavior.     

Nectar-receiver behavior in relation to the reward rate experienced by foraging honeybees

Since forager honeybees change their food-unloading behavior according to nectar-source profitability, an experiment was performed in order to analyze whether food-receivers modify their within-hive tasks related to different reward conditions. We offered individual foragers two reward conditions at a rate feeder while an additional feeder offered a constant reward and was of free access to the rest of the hive. Both feeders were the only food sources exploited by the colony during the assays since a flight chamber was used. After receiving nectar, hive bees performed processing cycles that involved several behaviors and concluded when they returned to the delivery area to receive a new food sample. During these cycles, receivers mainly performed oral contacts offering food, or inspected cells, and often both. In the latter case, both behaviors occurred simultaneously and at the same distance from the hive entrance. When they performed a single task, either the occurrence of cell inspections increased or contact offerings decreased for the highest reward rate offered to the donor-forager. Receivers also begged for food more often after interacting with low-profit foragers. Thus, the profitability of the food source exploited by nectar-forager honeybees could affect receiver behaviors within the hives based on individual-to-individual interactions.Communicated by R.F.A. Moritz     

Distribution of deep-water gorgonian corals in relation to benthic habitat features in the Northeast Channel (Atlantic Canada)

The distribution and abundance of deep-water gorgonian corals were investigated along 52 transects at 183–498 m depth in the Northeast Channel, between Georges Bank and Browns Bank in the northwest Atlantic, using a remotely operated vehicle and a towed video-camera system. Three species (Paragorgia arborea, Primnoa resedaeformis, and Acanthogorgia armata) were observed. Primnoa occurred on 35 transects below 196 m depth, with highest local abundance in stands of 104 colonies per 100 m². Paragorgia was present on 21 transects deeper than 235 m, with highest local abundance of 49 colonies per 100 m². Acanthogorgia was observed at only four transects between 231 m and 364 m, with a local maximum abundance of 199 colonies per 100 m². The maximum abundance averaged for whole transects was 19.2 and 6.2 colonies per 100 m² for Primnoa and Paragorgia, respectively. The corals were more common in the outer part of the channel along the shelf break and slope than on the shelf in the inner part. All three species showed a patchy distribution with no signs of competitive exclusion at any spatial scale. Transects with high abundance of corals were characterised by depths greater than 400 m, maximum temperatures less than 9.2°C, and a relatively high percentage coverage of cobble and boulder (more than 19% and 6%, respectively). High temperatures probably control the upper depth limit of the corals, and Primnoa seems to tolerate slightly higher temperatures than Paragorgia. Abundance of both species was negatively correlated with average temperature and positively with cobbles. Together, temperature, percentage cobble and salinity accounted for 38% of the variance of Primnoa. The comparable figure for Paragorgia was 15%. The observed distribution indicated that the abundance of coral is controlled by additional factors such as larger-scaled topographic features governing the current regimes and thus also the supply of food and larvae.Communicated by R. J. Thompson, St. Johns     

Determinants of male spacing behavior in microtines: an experimental manipulation of female spatial distribution and density

Theory on microtine mating systems predicts that male spacing behavior will be related to female spatial and temporal distribution. However, data from a natural population of field voles, Microtus agrestis, indicated a potential influence of female density on the spacing behavior of males. Therefore, I experimentally investigated the relative importance of female density and female spatial distribution for the spacing behavior of males in M. agrestis. Males were radio-tracked in enclosed natural habitats in which females at different densities were placed in two different spatial arrangements: clumped versus even distribution. Female density was the main factor determining male spacing behavior. At the high female density males had smaller home ranges and moved shorter distances between radio-tracking recordings. Also, home ranges were more exclusive at high female density. However, since there was a significant positive correlation between home range size and range overlap, range exclusiveness seemed to be influenced by female density indirectly through the effect of range size. Female spatial distribution, on the other hand, had no influence on male home range exclusiveness.     

Sex differences in the foraging behavior of squirrel monkeys in a seasonal habitat

Summary The effects of sex and seasonal changes in food abundance on foraging behavior was studied in squirrel monkeys (Saimiri oerstedi) in Costa Rica over an eleven-month period. Females searched for and ate food at significantly greater frequencies than did males throughout the study. The frequency of the specific foraging techniques used occasionally differed significantly within seasons, but not across the study period. Few differences were found in the foraging behaviors of nonreproductive sexually mature females compared to females that were pregnant or lactating. The major exception was that during the month following parturition reproductive females foraged for flowers and fruits more frequently than did non-reproductive females. The reduction of time spent by males in foraging activities gives them more time for other activities, especially anti-predator vigilance. Foraging techniques and the proportions of different food types in the diet changed seasonally. Foraging for arthropods was most frequent in the season when arthropod abundance was lowest, resulting in the amount of time spent eating arthropods to vary less across the seasons. Fruits and flowers were not eaten in a direct relationship to availability, but were used more than expected relative to availability when arthropod abundance was reduced. Individuals were more dispersed when foraging compared to other activities. Overall, there was little evidence of any direct foraging benefits for a squirrel monkey from being social.     

Relationships between spacing behavior and growth rates: A field study of lizard feeding territories

Summary Relationships between spacing behavior and growth rates were investigated in a field experiment with juvenile lizards, Anolis aeneus. The behavioral variable most closely related to juvenile growth was distance moved per unit time. This variable had a curvilinear relationship with growth, such that juveniles moving approximately 400 cm/h grew more rapidly than those traveling either larger or shorter distances per unit time. Daily fluctuations in arthropod abundance were also related to growth rates, with restricted growth during periods of low food availability. Temporal fluctuations in prey and distance traveled per unit time had independent effects on growth; together these two variables accounted for 43% of the variance in growth rate for the juveniles in this study.Territory size, overlap and social status appeared to affect growth indirectly, by influencing distance traveled per unit time. Optimal travel distances of around 400 cm/h were most likely when a juvenile had a relatively exclusive territory of about 0.5 m². High ranking juveniles were more apt to achieve this spacing pattern than were low ranking juveniles, but some high ranking juveniles had very large territories, extensive overlap with subordinates, supraoptimal travel distances and relatively low growth rates. Low ranking juveniles tended to fall into two groups: subordinates, with a small home range overlapping that of a more dominant individual and low travel distances, and floaters, with a large home range overlapping several more dominant individuals and high travel distances. Although a few low ranking juveniles achieved travel distances permitting high growth rates, most had either supra or suboptimal travel distances and relatively low growth rates.     

Multiple mating and the evolution of social behavior in the hymenoptera

Summary The data on the frequency of mating by queens of eusocial Hymenoptera are reviewed.It is pointed out that the issue of sperm clumping is probably irrelevant to the evolution of eusociality.The hypothesis is presented that multiple mating is an adaptation for maintaining large colonies. In ants there is a significant relation between the size of the colony and the frequency of mating.The effect of multiple mating on the spread of a gene for worker behavior is explored. If a female mates twice, the effective number of matings is less than two except in the case of identical sperm contribution by the males.Sperm bias is defined as the contribution of unequal amounts of sperm by the males that mate with a queen. Sperm bias can be produced as a sampling phenomenon, by inter-male competition for females and by sperm competition.The relation between the ergonomic efficiency of the workers at the production of reproductives and the number of matings that is consistent with the evolution of eusociality is derived. If workers are only about 10% more efficient at producing reproductives within a eusocial colony than they are solitarily, then two matings by the queen will still produce a selective advantage to eusocial behavior.