Ecological niche and phylogeny: the highly complex echolocation behavior of the trawling long-legged bat, Macrophyllum macrophyllum

Bats produce echolocation signals that reflect the sensory tasks they perform. In open air or over water, bats encounter few or no background echoes (clutter). Echolocation of such bats is the primary cue for prey perception and varies with the stage of approach to prey, typically comprising search, approach, and terminal group calls. In contrast, bats that glean stationary food from rough surfaces emit more uniform calls without a distinct terminal group. They use echolocation primarily for orientation in space and mostly need additional sensory cues for finding food because clutter echoes overlap strongly with food echoes. Macrophyllum macrophyllum is the only Neotropical leaf-nosed bat (Phyllostomidae) that hunts in clutter-poor habitat over water. As such, we hypothesized that, unlike all other members of its family, but similar to other trawling and aerial insectivorous bats, M. macrophyllum can hunt successfully by using only echolocation for prey perception. In controlled behavioral experiments on Barro Colorado Island, Panamá, we confirmed that echolocation alone is sufficient for finding prey in M. macrophyllum. Furthermore, we showed that pattern and structure of echolocation signals in M. macrophyllum are more similar to aerial and other trawling insectivorous bats than to close phylogenetic relatives. Particularly unique among phyllostomid bats, we found distinct search, approach, and terminal group calls in foraging M. macrophyllum. Call structure, however, consisting of short, multiharmonic, and steep frequency-modulated signals, closely resembled those of other phyllostomid bats. Thus, echolocation behavior in M. macrophyllum is shaped by ecological niche as well as by phylogeny.     

Mating system of a Neotropical roost-making bat: the white-throated, round-eared bat, Lophostoma silvicolum (Chiroptera: Phyllostomidae)

The vast majority of bats strongly depend on, but do not make, shelters or roosts. We investigated Lophostoma silvicolum, which roosts in active termite nests excavated by the bats themselves, to study the relationship between roost choice and mating systems. Due to the hardness of the termite nests, roost-making is probably costly in terms of time and energy for these bats. Video-observations and capture data showed that single males excavate nests. Only males in good physical condition attracted females to the resulting roosts. Almost all groups captured from excavated nests were single male-multifemale associations, suggesting a harem structure. Paternity assignments based on ten polymorphic microsatellites, revealed a high reproductive success of 46% by nest-holding males. We suggest that the mating system of L. silvicolum is based on a resource-defense polygyny. The temperatures in the excavated nests are warm and stable, and might provide a suitable shelter for reproductive females. Reproductive success achieved by harem males appears to justify the time and effort required to excavate the nests. Reproductive success may thus have selected on an external male phenotype, the excavated nests, and have contributed to the evolution of an otherwise rare behavior in bats.Communicated by G. Wilkinson     

The sensory basis of prey location by the California leaf-nosed bat Macrotus californicus (Chiroptera: Phyllostomidae)

Summary Macrotus californicus, an insectivorous bat, captures prey on the ground, and shows great sensory flexibility in hunting for prey: it uses high frequency, low intensity, frequency modulated echolocation to locate prey in total darkness, however data from this study suggest that it uses vision preferentially, and switches off its echolocation when adequate illumination is available. When souncs of prey are available it exploits these also. It uses echolocation only 50% of the time at 4.2x10^-2 mL, comparable to ground luminance on a brightly moonlit night, and employs vision even at 10^-3 mL.     

Flexible bat echolocation: the influence of individual,habitat and conspecifics on sonar signal design

Acoustic signals which are used in animal communication must carry a variety of information and are therefore highly flexible. Echolocation has probably such functions and could prove as flexible. Measurable variabitlity can indicate flexibility in a behaviour. To quantify variability in bat sonar and relate to behavioural and environmental factors, I recorded echolocation calls of Euderma maculatum, Eptesicus fuscus, Lasiurus borealis and L. cinereus while the bats hunted in their natural habitat. I analysed 3390 search phase calls emitted by 16 known and 16 unknown individuals foraging in different environmental and behvioural situations. All four species used mainly multiharmonic signals that showed considerable intra- and inter-individual variability in the five signal variables I analysed (call duration, call interval, highest and lowest frequency and frequency with maximum energy) and also in the shape of the sonagram. A nested multivariate analysis of variance identified the influences of individual, hunting site, close conspecifics and of each observation on the frequency with maximum energy in the calls, and on other variables measured. Individual bats differed in multiple comparisons, most often in the main call frequency and least often in call interval. In a discriminant function analysis with resubstitution, 56–76% of a species' calls were assigned to the correct individual. Distinct individual call patterns were recorded in special situations in all species and the size of foraging areas in forested areas influenced temporal and spectral call structure. Echolocation behaviour was influenced by the presence of conspecifics. When bats were hunting together, call duration decreased and call interval increased in all species, but spectral effects were less pronounced. The role of morphometric differences as the source of individually distinct vocalizations is discussed. I also examined signal adaptations to long range echolocation and the influence of obstacle distance on echolocation call design. My results allow to discuss the problems of echo recognition and jamming avoidance in vespertilionid bats.     

Relatedness structure and kin-biased foraging in the greater horseshoe bat (Rhinolophus ferrumequinum)

Female greater horseshoe bats (Rhinolophus ferrumequinum) exhibit strong natal philopatry to their maternity roost over many years, leading to the aggregation of matrilineal kin. Maternity colonies may, therefore, be expected to comprise highly related individuals, and, as such, provide conditions suitable for the evolution of kin-selected behaviours. To test these predictions, we examined relatedness and behaviour among matrilineal kin within a colony in south-west Britain. Genetic analysis of 15 matrilines, identified from microsatellite genotyping and long-term ringing surveys, revealed average relatedness levels of 0.17 to 0.64. In contrast, background relatedness among colony females approximated to zero (0.03). These results suggest that inclusive fitness benefits may only be accrued through discriminate cooperation within matrilines, and not at the wider colony level. To examine whether the potential for such benefits is realised through kin- biased cooperation during foraging, females from two matrilines were radio-tracked simultaneously over 3 years. Pairwise home-range overlap correlated significantly with Hamilton's relatedness coefficient. The greatest spatial associations were observed between females and their adult daughters, which shared both foraging grounds and night roosts, sometimes over several years. Tagged females, however, generally foraged and roosted alone, suggesting that kin-biased spatial association probably does not result from either information-transfer or cooperative territorial defence. Such patterns may instead result from a mechanism of maternal inheritance of preferred foraging and roosting sites.     

The economics of harem maintenance in the sac-winged bat, Saccopteryx bilineata (Emballonuridae)

Saccopteryx bilineata has a polygynous mating system in which males defend females in a harem territory. Harem defense and courtship include energetically costly flight maneuvers and hovering displays. We tested if (1) harem males have a greater field metabolic rate than non-harem males or females and if (2) the field metabolic rate of harem males is correlated with the number of females in a harem territory. We measured the energy budget in 32 S. bilineata with the doubly labeled water method and compared these estimates with behavioral observations in the daytime roost. Among adult bats, field metabolic rate varied with body mass by an exponent of approximately two. We found no significant difference in field metabolic rate or mass-specific field metabolic rate between harem and non-harem males. The mass-specific field metabolic rate of harem-males increased with harem size. The latter finding supports the hypothesis that the energy costs of courtship display and territorial defense influence the energy budget of harem males. Overall, field metabolic rates of S. bilineata were lower than those of similarly sized bats of the temperate zone and only 2.3 times above the basal metabolic rate recorded for this species. We suggest that male S. bilineata did not take advantage of their metabolic capacity because a prudent allocation of energy to activities of harem maintenance is an adaptive strategy for males in this mating system.     

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     

Response of mud crab (Panopeus herbstii) megalopae to cues from adult habitat

During the summer of 1995, we investigated the response of mud crab megalopae (Panopeus herbstii Milne-Edwards) to cues associated with adult habitat. Natural rock/shell substratum and the rock-associated seaweed, Fucus vesiculosus L., both induced metamorphosis of competent megalopae; natural sand substratum did not induce metamorphosis. Structural mimics of preferred substrata induced metamorphosis only when covered with natural biofilm. Clean mimics did not induce metamorphosis. Water-soluble exudates from preferred substrata showed weak induction of metamorphosis. Exudates from adult P. herbstii elicited a similar weak response. Exudates from another species of crab and from two fish predators did not induce metamorphosis. We conclude that water-soluble cues associated with major components of preferred adult habitat induce the metamorphic molt of P. herbstii megalopae. The three-dimensional structure of the substratum is not important in induction of metamorphosis, but the biofilm associated with preferred substratum plays a critical role. It is not clear whether the biofilm produces the water-soluble cue or simply provides a tactile stimulus that works in conjunction with the soluble cue.     

Climate, habitat, and species interactions at different scales determine the structure of a Neotropical bat community

Climate, habitat, and species interactions are factors that control community properties (e.g., species richness, abundance) across various spatial scales. Usually, researchers study how a few properties are affected by one factor in isolation and at one scale. Hence, there are few multi-scale studies testing how multiple controlling factors simultaneously affect community properties at different scales. We ask whether climate, habitat structure, or insect resources at each of three spatial scales explains most of the variation in six community properties and which theory best explains the distribution of selected community properties across a rainfall gradient. We studied a Neotropical insectivorous bat ensemble in the Isthmus of Panama with acoustic monitoring techniques. Using climatological data, habitat surveys, and insect captures in a hierarchical sampling design we determined how much variation of the community properties was explained by the three factors employing two approaches for variance partitioning. Our results revealed that most of the variation in species richness, total abundance, and feeding activity occurred at the smallest spatial scale and was explained by habitat structure. In contrast, climate at large scales explained most of the variation in individual species' abundances. Although each species had an idiosyncratic response to the gradient, species richness peaked at intermediate levels of precipitation, whereas total abundance was very similar across sites, suggesting density compensation. All community properties responded in a different manner to the factor and scale under consideration.     

Foraging areas and foraging behavior in the notch-eared bat,Myotis emarginatus (Vespertilionidae)

Summary Field observations in a maternity colony of Myotis emarginatus (Vespertilionidae) were made during the summers of 1986 and 1987 in southern Germany. The nursery colony consisted of about 90 adult and 30 juvenile bats which roosted in a dimly lit and relatively cool church attic. Telemetry data from six adult M. emarginatus disclosed that some individuals also use secondary day roosts in trees or small buildings located close to their foraging areas. During the night, radiotagged individuals spent most of the time on the wing in forested areas (Fig. 2). Stationary bouts lasted no longer than 63 min. Individual bats returned to the same foraging areas on consecutive nights. All major foraging areas were situated in or at the fringes of forests, at distances as far as 10 km from the nursery roost. During commuting flights to the forests, M. emarginatus avoided open fields and preferred flight paths which offered cover such as orchards, hedges, overhanging foliage along creeks, etc. On the way to the forests, the bats started to forage within buildings, in open spaces where aggregations of insects were present, and around or within the foliage of various types of trees at the level of tree tops or the upper third of the foliage. At these transient foraging areas close to the maternity roost, M. emarginatus displayed flexible foraging strategies: (1) They gleaned prey (mainly flies and spiders) from the substrate, (2) seized insects in aerial pursuit, and (3) occasionally hovered in front of foliage and walls.Our observations confirm the conclusion from morphometric data on the wings that M. emarginatus is a predominantly gleaning bat and contradict the suggestion that it makes only brief flights of short distances. On the contrary, our field data suggest that M. emarginatus spends most of the night on the wing and commutes over distances of at least 10 km. Offprint requests to: D. Krull     

Foraging behaviour and echolocation in the rufous horseshoe bat (Rhinolophus rouxi) of Sri Lanka

Summary In October 1984 foraging areas and foraging behaviour of the rufous horseshoe bat, Rhinolophus rouxi, were studied around a nursery colony on the hill slopes of Sri Lanka. The bats only foraged in dense forest and were not found in open woodlands (Fig. 1). This strongly supports the hypothesis that detection of fluttering prey is by pure tone echolocation within or close to echo-cluttering foliage. During a first activity period after sunset for about 30–60 min, the bats mainly caught insects on the wing. This was followed by a period of inactivity for another 60–120 min. Thereafter the bats resumed foraging throughout the night. They mainly alighted on specific twigs and foraged in flycatcher style. Individual bats maintained individual foraging areas of about 20x20 m. They stayed in this area throughout the night and returned to the same area on subsequent nights. Within this area the bats generally alighted on twigs at the same spots. Foraging areas were not defended against intruders. The bats echolocated throughout the night at an average repetition rate of 9.6±1.4 sounds/s. While hanging on twigs they scanned the surrounding area for flying prey by turning their bodies continuously around their legs. On average they performed one brief catching flight every 2 min and immediately returned to one of their favourite vantage points. Echolocation sounds may consist of up to three parts, a brief initial frequency-modulated (FM) component, a long constant frequency (CF) part lasting for about 40–50 ms, and a final FM part again (Fig. 4b, c). Adult males and females emitted pure tone frequencies in separate bands, the males from 73.5–77 kHz and the females from 76.5–79 kHz (Fig. 5). During scanning for prey from vantage points, the bats mostly emitted pure tones without any FM component (Fig. 4a). The last few pure tones emitted before take-off were prolonged to about 60 ms duration. The final FM part was therefore not an obligatory component of the echolocation signals in horseshoe bats. During flight and especially during emergence from the cave, most sounds consisted of a pure tone and loud initial and final FM sweeps. We therefore suggest that the initial FM part might also be relevant for echolocation. From our observations we conclude that the FM components are especially important during obstacle avoidance. In most sounds emitted in the field a fainter first harmonic was present. It was usually up to 30 dB fainter than the second harmonic, but in some instances it was as loud or even distinctly louder than the second one (Fig. 6a). Even within one sound the intensity relationship between the two harmonics may be reversed. We therefore suggest that the first harmonic is an integral part of the signal and relevant for information analysis in echolocation.     

Spatio-temporal use of the urban habitat by feral pigeons (Columba livia )

Feral pigeons are descendants of wild rock pigeons that have adapted to the urban habitat. They have partially conserved the foraging behaviour of their wild ancestors (flights to agricultural areas) but have also developed new habits. Previous studies on the foraging strategies of feral pigeons have given various results, e.g. maximum distances reached by the pigeons (measured in a straight line from the resting places) differed between 0.3–0.5 km and 18–25 km. This study focuses on the spatio-temporal activity of feral pigeons in the urban habitat. We equipped 80 free-living feral pigeons from Basel, Switzerland with GPS receivers. We found three different foraging strategies for pigeons in Basel: (1) in the streets, squares and parks near the home loft, (2) in agricultural areas surrounding the city, (3) on docks and railway lines in harbours. The maximum distance reached by a pigeon was 5.29 km. More than 32% of the pigeons remained within 0.3 km of the home lofts and only 7.5% flew distances of more than 2 km. Females covered significantly longer distances than males, preferring to fly to more abundant and predictable food sources. Temporal activity patterns showed to be influenced by sex, breeding state and season. In contrast to wild rock pigeons and to feral pigeons in other cities, pigeons in Basel showed a clear bimodal activity pattern for breeding birds only. The differences between our results and those of other studies seem to be partly method-dependent, as the GPS-technique allows to record the pigeons’ localisations continuously in contrast to other methods. Other differences might be due to different kinds of food supply in the various cities. Our study shows that feral pigeons have individual foraging strategies and are flexible enough to adapt to different urban environments.Electronic supplementary material Supplementary material is available for this article at     

Ontogenetic habitat shift and risk of cannibalism in the common chameleon (Chamaeleo chamaeleon)

Ontogenetic habitat shifts have been documented in numerous fish and amphibians and in some reptiles. Intraspecific competition together with differential predation, prey size, social interactions, size-related thermal requirements, and morphological constraints on movement are often implicated in this ontogenetic habitat separation. In the current study, we combined field observation with experiments in seminatural arenas to test various hypotheses regarding the ontogenetic habitat shift that we have documented in the common chameleon. Juveniles (mean, 1 g) occupied low grasses and the adults (mean, 35 g) were found on bushes and trees. Overlap in habitat use between these two age classes was minimal. Our field experiments showed that juveniles actively avoid the presence of adults by concealment or flight. Adults readily attacked and consumed juveniles, regardless of their own mass. These results suggest that the risk of cannibalism towards juveniles is an important selective force behind the ontogenetic habitat shift observed in the common chameleon and may be important in other species too.     

Physiological responses to fluctuation in temperature of salinity in invertebrates. Adaptations ofAlpheus viridari (Decapoda,Crustacea),Terebellides parva (Polychaeta) andGolfinigia cylindrata (Sipunculida) to the mangrove habitat

The snapping shrimpAlpheus viridari (Armstrong, 1949), the polychaeteTerebellides parva Solis-Weiss, Fauchald and Blankensteyn 1990, and the sipunculanGolfingia cylindrata (Keferstein, 1865) are commonly found in the same mangrove habitat, where they experience frequent, acute fluctuations in temperature and salinity. Ecological studies indicate a temporal variation, including occasional absence, in the distribution of bothG. cylindrata andT. parva; this fed us to examine the physiological adaptations of the three species (collected at Western Bay, Twin Cays, Belize in 1985, 1986 and 1988). Each was subjected to acute, repeated exposure to either control (35 S) and decreased (25 S) salinity or to control and increased (45 S) salinity. Ability to regulate water and ion content (g H₂O or mol g^-1 solute free dry wt) was examinedA. viridari behaved as a hyperosmotic conformer at decreased salinity but as an osmoconformer at increased salinity. Regardless of direction of salinity change,A. viridari regulated water content through change in Na⁺, K⁺, and Cl^– contents. In contrast,G. cylindrata behaved as an osmoconformer and did not demonstrate ability to regulate water content.T. parva behaved as an osmoconformer, showed incomplete regulation of water content via change in Na⁺, K⁺, and Cl^– contents but had limited survival following exposure to 45 S. Each species was also exposed to change in temperature. Species were subjected to acute, repeated exposure either to control (28°C) and decreased (21°C) temperature or to control and inereased (35°C) temperatureA. viridari regulated water and ion content under both experimental conditions. In contrast,T. parva did not regulate water and ion content under either experimental temperature.G. cylindrata did not regulate water and ion content during exposure to decreased temperature and did not survive exposure to increased temperature. ForA. viridari, weight specific oxygen uptake rates (mg O₂ g^-1 ash-free dry wt) were determined. Exposure to decreased salinity or to increased temperature resulted in a small sustained elevation in O₂ uptake. It is concluded that, unlikeA. viridari, T. parva andG. cylindrata are only marginally adapted to withstand the salinity and temperature stresses, respectively, of the mangrove habitat. The inability ofT. parva andG. cylindrata to fully adapt to extremes in the mangrove habitat could well explain the temporal variation seen in the distribution of these two species.Contribution number 380. Caribbean Coral Reef Ecosystems, National Museum of Natural History, Smithsonian Institution     

Foraging habitat and echolocation behaviour of Schneider's leafnosed bat, Hipposideros speoris, in a vegetation mosaic in Sri Lanka

The Hipposideridae and Rhinolophidae are closely related families of bats that have similar echolocation (long-duration pure-tone signal, high duty cycle) and auditory systems (Doppler-shift compensation, auditory fovea). Rhinolophid bats are known to forage in highly cluttered areas where they capture fluttering insects, whereas the foraging habitat of hipposiderid bats is not well understood. Compared to rhinolophids, hipposiderid calls are shorter in duration, have lower duty cycles, and they exhibit only partial Doppler-shift compensation. These differences suggest that the foraging habitat of the two families may also differ. We tested this hypothesis by studying foraging and echolocation of Hipposideros speoris at a site with a range of vegetation types. Bats foraged only while in flight and used all available closed and edge habitats, including areas adjacent to open space. Levels of clutter were high in forest and moderate in other foraging areas. Prey capture (n=42) occurred in edge vegetation where it bordered open space. Echolocation signals of H. speoris lacked an initial upward frequency-modulated sweep and were of moderate duration (5.1-8.7 ms). Sequences had high duty cycles (23-41%) and very high pulse repetition rates (22.8-60.6 Hz). Variation in signal parameters during search phase flight across foraging habitats was low. H. speoris showed a greater flexibility in its use of foraging habitat than is known for any rhinolophid species. Our study confirmed that there are differences in habitat use between hipposiderid and rhinolophid bats and we suggest that this divergence is a consequence of differences in their echolocation and auditory systems.     

Growth response of Lebanon cedar (Cedrus libani) plantations to thinning intensity in Western Turkey

This paper presents the growth response of 25 yr old Lebanon cedar (Cedrus libani A. Rich.) plantation to thinnings of different intensities in Isparta in western Turkey. The thinning intensity was measured by using the residual basal area (%) as parameter. In spring of 2005, three treatments were tested; light, moderate and heavy thinning with respectively 10, 25 and 35% of basal area removed. The statistical design of the experiment was a randomized incomplete block with two blocks and three treatments. Variables such as diameter at breast height (diameter) and height were measured. Growth rate ratios of diameter in moderately thinned and heavily thinned stands were 1.02 and 1.03, respectively. Basal area growth rates in moderately thinned and heavily thinned plots were 0.93 and 1.05, respectively. The largest values for the mean tree were observed with the heaviest thinning treatment. Absolute diameter increment was positively correlated with initial diameter in all plots. Relative diameter growth was negatively correlated with initial diameter. Growth rate interpretations were supported by analysis of variance using Duncan's test of range multiple. The results obtained show significant differences between treatments for tree height growth, for the two inventories carried out (2005, 2008). However diameter basal area and volume were no found between treatments for tree.     

Brood defence in the great tit (Parus major): the influence of life-history and habitat

Summary We examined the extent to which parental investment, as measured by brood defence, is determined by life-historical selection in a shortlived bird, the great tit (Parus major). Pairs tending first (n=20) and second (n=21) broods in the same Scots pine woods in 1983 were used to test predictions of a cost/benefit model of brood defence based on the species' average demography in coniferous forest. Furthermore, the differences in demography between pine and deciduous forest permitted us to test whether habitat-specific life-history would affect the seasonal pattern of defence. In the model, benefit was defined as the brood's potential contribution to a parent's fitness, and the cost as the potential loss if the defender dies in the act of defence. Univariate and multivariate procedures were applied to six measures of defence response to a live owl (Glaucidium perlatum) plus a taped mobbing chorus. Results proved three of the model's predictions to be false. In coniferous forest, neither the overall strength nor the individual variance of defence behaviour differed among first and second broods, nor was there any consistent difference in defence strength between pairs living in coniferous (n=54) or deciduous (n=84) forest as revealed by comparisons within each brood. These failures could be reconciled with the model by assuming that selection in the past had acted via the average demography of both types of habitat. The model received direct support from defence strength increasing with age of young and, more forcefully, becoming more influenced by brood size in second broods, regardless of habitat.A difference in the strength of defence by the male and female suggests two more functions of behaviour: In first broods, the male risks more than the female as measured by five of six variables. This suggests that defence is facultatively linked to the need for territorial protection from predators all the year round. In the female's presence, the male, taking an additional risk, approaches the owl to half the distance of that of a single, though paired, male, suggesting an additional, social role of defence behaviour.Taken together, anti-predator defence in the great tit serves to protect the brood, the home range and, in the male, the female mate. The magnitude of the benefits envisaged varies among the sexes.