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     

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.     

Fruit detection and discrimination by small fruit-eating bats (Phyllostomidae): echolocation call design and olfaction

We studied the role of echolocation and other sensory cues in two small frugivorous New World leaf-nosed bats (Phyllostomidae: Artibeus watsoni and Vampyressa pusilla) feeding on different types of fig fruit. To test which cues the bats need to find these fruit, we conducted behavioral experiments in a flight cage with ripe and similar-sized figs where we selectively excluded vision, olfaction, and echolocation cues from the bats. In another series of experiments, we tested the discrimination abilities of the bats and presented sets of fruits that differed in ripeness (ripe, unripe), size (small, large), and quality (intact(infested with caterpillars). We monitored the bats' foraging and echolocation behavior simultaneously. In flight, both bat species continuously emitted short (<2 ms), multi-harmonic, and steep frequency-modulated (FM) calls of high frequencies, large bandwidth, and very low amplitude. Foraging behavior of bats was composed of two distinct stages: search or orienting flight followed by approach behavior consisting of exploration flights, multiple approaches of a selected fruit, and final acquisition of ripe figs in flight or in a brief landing. Both bat species continuously emitted echolocation calls. Structure and pattern of signals changed predictably when the bats switched from search or orienting calls to approach calls. We did not record a terminal phase before final acquisition of a fruit, as it is typical for aerial insectivorous bats prior to capture. Both bat species selected ripe over unripe fruit and non-infested over infested fruit. Artibeus watsoni preferred larger over smaller fruit. We conclude from our experiments, that the bats used a combination of odor-guided detection together with echolocation for localization in order to find ripe fruit and to discriminate among them.     

Neocembrene A, a major component of the trail-following pheromone in the genus Prorhinotermes (Insecta, Isoptera, Rhinotermitidae)

Summary. The diterpene neocembrene A or (1E,5E,9E,12R)-1,5,9-trimethyl-12-(1-methylethenyl)-1,5,9-cyclotetradecatriene, known as the trail-following pheromone of the advanced Termitidae Nasutitermitinae Nasutitermes exitiosus and Trinervitermes bettonianus, has been identified after SPME-GC/MS as the major component of the trail-following pheromone of the Rhinotermitidae Prorhinotermitinae, Prorhinotermes canalifrons and P. simplex. In all the other Rhinotermitidae studied until now, the major component of their trail pheromones is dodecatrienol ((3Z,6Z,8E)-dodeca-3,6,8-trien-1-ol). This biochemical data further add to the anatomical and molecular characteristics that give a special status to the taxon Prorhinotermes among Rhinotermitidae. In Prorhinotermes canalifrons and P. simplex, neocembrene A was the only secretory compound specific to the sternal gland surface that could be detected after SPME. It elicited orientation as well as recruitment behavioral effects. However, the comparison of the respective biological activities triggered by neocembrene A and by sternal gland secretion suggests that minor components of the latter are acting in synergy with neocembrene A.     

Testosterone-induced depression of male parental behavior in the barn swallow: female compensation and effects on seasonal fitness

In birds, many aspects of male socio-sexual and parental behavior are influenced by androgens, most notably testosterone (T). We report the effects of subcutaneous T-implants in male barn swallows (Hirundo rustica) on male and female parental behavior and on seasonal reproductive success. Males were assigned to one of three experimental groups: (i) implanted with a T-filled Silastic tube; (ii) implanted with an empty Silastic tube; and (iii) not implanted. T-implanted males provided a smaller proportion of feedings (number of feedings by the male/total number of feedings by both parents) and fed nestlings less frequently (number of feedings/h) than males of the other two groups. Females paired to T-implanted males fed nestlings significantly more often than females paired with unimplanted males. Females almost fully compensated for their mates' shortfall, and this resulted in similar combined feeding efforts among treatments. Reproductive success in their first broods or during the entire breeding season was unaffected by T- treatment. These results confirm earlier reports of the suppressive effects of T on male parental behavior. However, they are inconsistent with current ESS models that predict partial compensation as the optimal response by one individual to reduction of parental effort by its mate in monogamous, biparental systems.     

Mating patterns and a test for female mate choice in Etheostoma spectabile (Pisces,Percidae)

Neither size nor breeding color correlated with spawning success of male orangethroat darters, Etheostoma spectabile (Pisces, Percidae), under natural field conditions. When females were presented experimentally with a simultaneous choice they spent no more time in proximity to large than small males, and were subsequently no more likely to spawn with large than with small males. Females also displayed no preference for bright versus dull males. Males and females did not differ significantly in size. Etheostoma spectabile may lack sexual size dimorphism as a result of the lack of female choice for size and the ineffectiveness of male attempts to monopolize females, or selection may be for increased size of females. Males are not dwarfs because of sperm competition. Contest competition among males appeared to be important in initiating spawnings but many males obtained spawnings by participating in ongoing spawning events. Etheostoma spectabile is an example of a sexually dimorphic species with no evident female preference for male size or color.     

Antennal responses of four species of tree-killing bark beetles (Coleoptera: Scolytidae) to volatiles collected from beetles,and their host and nonhost conifers

Summary. Host selection in tree-killing bark beetles (Coleoptera: Scolytidae) is mediated by a complex of semiochemical cues. Using gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectrometric analyses, we conducted a comparative study of the electrophysiological responses of four species of tree-killing bark beetles, the Douglas-fir beetle, Dendroctonus pseudotsugae, Hopkins, the mountain pine beetle, D. ponderosae Hopkins, the spruce beetle, D. rufipennis Kirby, and the western balsam bark beetle, Dryocoetes confusus Swaine, to volatiles captured by aeration of 1) bole and foliage of four sympatric species of conifers, Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, lodgepole pine, Pinus contorta var. latifolia Engelm., interior spruce, Picea engelmannii x glauca, and interior fir, Abies lasiocarpa x bifolia, and 2) con- and heterospecific beetles at three stages of attack. We identified 13 monoterpenes in the conifers and nine compounds in the volatiles of beetles that elicited antennal responses. There was no qualitative difference in the terpene constitution of the four species of conifers and very little difference across beetle species in their antennal response to compounds from conifers or beetles. The lack of species-specific major or minor components in conifers suggests that beetles would need to detect differences in the ratios of different compounds in conifers to discriminate among them. Attraction to hosts and avoidance of nonhost conifers may be accentuated by perception of compounds emitted by con- and heterospecific beetles, respectively. The 22 compounds identified are candidate semiochemicals with potential behavioural roles in host location and discrimination.     

The cost of success: reproductive effort in male southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>)

Reproductive effort is a key parameter of life history because it measures the resources allocated to reproduction at the expense of growth and maintenance. Male reproductive effort always had a minor role with respect to female effort both in the development of theories and in field research. Elephant seals are an ideal subject for reproductive effort studies because they fast during the breeding season, splitting the phase of energy acquisition from the phase of energy use for breeding. In this paper, we present results on male reproductive effort (weight loss estimated by photogrammetry) in southern elephant seals (Mirounga leonina), the most dimorphic and polygynous of all mammal species. We show that total reproductive effort increases with age, with no sign of late decrease or senescence. Male reproductive effort in this species depends mostly on behavioral factors, i.e., the success in competition with other males, and the intensity of interaction with females. A large effort results in large gains in both mating success and fertilizations. The large reproductive success that a few males are able to achieve come at a big cost in terms of energy expenditure, but this cost does not seem to affect the likelihood of survival to the following breeding season.     

Divergence in female calling song discrimination between sympatric and allopatric populations of the southern wood cricket Gryllus fultoni (Orthoptera: Gryllidae)

Two species of closely related wood cricket, Gryllus fultoni (Orthoptera: Gryllidae) and Gryllus vernalis, occur together in some parts of the eastern United States and have a similar calling song structure, consisting of three-pulse chirps. A previous study revealed that chirp rate in G. fultoni was highest (greatest difference vis-à-vis chirp rate in G. vernalis) in sympatric populations, intermediate in near allopatric populations that were located close to the sympatric zone, and lowest in allopatric populations. A similar trend was observed in pulse rate, but the mean values of this trait showed much more convergence than chirp rate at the low end of the range of calling temperatures. In this study, we investigated the song discrimination of females from sympatric and allopatric populations of G. fultoni at about 23°C, which is near the middle of the normal range of calling temperatures. We used both single-stimulus and two-stimulus playback experiments to learn if geographical differences in song preferences paralleled those in calling songs. Stimuli presented were representative of calling songs in three classes of G. fultoni populations (sympatric, near allopatric, and far allopatric), a calling song of G. vernalis, and three calling songs with parameter values that were intermediate with respect to those of the songs of far allopatric G. fultoni and G. vernalis. In the single-stimulus playbacks, females of all G. fultoni populations responded poorly if at all to the heterospecific stimulus. Females of sympatric and near allopatric populations responded poorly to all intermediate stimuli, but females of far allopatric populations frequently responded to these sounds. In the two-stimulus playbacks, females of sympatric and near allopatric populations generally discriminated against intermediate and heterospecific stimuli. However, females of far allopatric populations often did not discriminate against intermediate stimuli, whose characteristics resembled the calling songs of G. vernalis. The divergent pattern of female phonotactic discrimination between sympatric and far allopatric populations was thus generally congruent with the pattern of divergence in chirp and pulse rates and would be expected to significantly reduce heterospecific mating in sympatry. These geographical patterns of female song discrimination and male calling songs conform to a commonly used definition of reproductive character displacement.     

Alternative reproductive tactics in male common shrews: relationships between mate-searching behaviour,sperm production,and reproductive success as revealed by DNA fingerprinting

The common shrew (Sorex araneus) is a solitary small mammal with a promiscuous mating system. Previous studies of this species suggest that females typically mate multiply, and that males may adopt alternative mate-searching tactics. We studied two generations of common shrews in a population near Oxford, England. Males were found to adopt two different mate-searching tactics. Those classed as type A occupied relatively small exclusive ranges during March, and made repeated long-distance movements to visit female ranges around the time of first oestrus in April. Males classed as type B established large overlapping ranges in areas of relatively high female density during March, and maintained these ranges throughout April. Type B males were larger than type A males at an early stage of sexual maturation, but there was no difference in the adult body size of the two types of male. Type A males had significantly higher epididymal sperm counts than type B males. Paternity analyses of litters born during the first year of the study reveal that the mean number of offspring fathered by type B males was greater than the mean number fathered by type A males. It is concluded that different mate-searching tactics may be conditional upon the timing of sexual maturation. Differences in sperm production are discussed in relation to sperm competition theory.     

Diet-induced chemical phytomimesis by twig-like caterpillars of Biston robustum Butler (Lepidoptera: Geometridae)

Summary. Polyphagous caterpillars of the giant geometer Biston robustum resemble the twigs of their respective food sources in color and shape. Common predatory ants, including Lasius and Formica, were often observed to freely prowl directly on caterpillars bodies, even after antennal contact. This suggests that the cuticular chemicals of the caterpillars resemble those of the twigs of the foodplants, so we analyzed both by GC and GC-MS. The chemical compositions differed among caterpillars fed on a cherry, Prunus yedoensis, a chinquapin Castanopsis cuspidata, and a camellia Camellia japonica. The cuticular chemicals of the caterpillars resembled those of their corresponding food sources. When the caterpillar diets were switched from the cherry to camellia or chinquapin at the 4^th instars, the caterpillars cuticular chemicals changed after molting to resemble those of their respective foods. Caterpillars also changed their cuticular chemicals when they perched on cherry twigs and fed on camellia or chinquapin leaves, but not when they perched on camellia or chinquapin twigs and fed on cherry leaves. The chemical similarities between the caterpillars and the twigs were due to the digestion of host leaves, which indicates that this is a diet-induced adaptation.     

The effect of helpers on artificially increased brood size in sociable weavers (Philetairus socius)

In cooperatively breeding birds, the presence of helpers is expected to increase the reproductive success of the breeding pair. However, some studies fail to find this effect. A positive effect of helpers may be restricted to cases in which a breeding pair has a poor likelihood of raising the entire brood on its own, as would be the case under stressful environmental conditions or with enlarged brood sizes. We conducted brood size manipulations in a cooperative breeder, the sociable weaver, Philetairus socius, to investigate the relationship between the difficulty of raising nestlings and the effort and impact of helpers. Overall, sociable weavers did not work harder to raise the enlarged broods. However, the presence of helpers significantly increased the feeding rates at enlarged nests, but not controls. This was insufficient to prevent generalised brood reduction in enlarged broods, whether attended by pairs alone or with helpers. Nonetheless, the presence of helpers was associated with decreased nestling mortality and an increase in the numbers of young fledged. Our results suggest that groups are better able to respond to the needs of enlarged broods than pairs alone and that the presence of helpers has a beneficial effect on overall reproductive success.Communicated by J. Dickinson     

Oviposition-stimulating activity of (E)-capsaicin identified in Capsicum annuum fruit and related compounds towards Helicoverpa assulta (Lepidoptera: Noctuidae)

Summary. The oviposition-stimulating activity of (E)-capsaicin identified in the fruits of red pepper, Capsicum annuum L. (Solanaceae), towards the oriental tobacco budworm, Helicoverpa assulta (Guenée), was examined in the laboratory by using choice and no-choice bioassays. Results were compared with those of the commercially available capsanthin and dihydrocapsaicin, another known constituents of C. annuum fruit, and (Z)-capsaicin. In no-choice oviposition bioassays at 10.5 μg/cm², (E)-capsaicin was the most active oviposition stimulant. Weak stimulatory activity was observed with capsanthin, (Z)-capsaicin and dihydrocapsaicin. In choice oviposition bioassays at 2.6 μg/cm², there were significant differences in oviposition-stimulating activity between (E)-capsaicin and each of the other three alkaloids. Structure-activity relationships indicate that the presence of double bond and its trans configuration appeared to play a crucial role in the oviposition-stimulating activity. On the basis of our results, (E)-capsaicin acts as a contact oviposition stimulant of female H. assulta.     

Heritable pheromone blend variation in a local population of the butterbur borer moth Ostrinia zaguliaevi (Lepidoptera: Crambidae)

Summary. In a local population of Ostrinia zaguliaevi Mutuura & Munroe (Lepidoptera: Crambidae), extensive variation was found in the blend ratio of three sex pheromone components, (Z)-9-tetradecenyl acetate (Z9-14:OAc, 10.2-63.8%), (Z)-11-tetradecenyl acetate (Z11-14:OAc, 32.2-86.8%), and (E)-11-tetradecenyl acetate (E11-14:OAc, 2.1-11.9%). The variation was observed over a three-year period (2002-2004). Mother-daughter regression analyses have shown that although the heritability of the minor component E11-14:OAc was not significant, the heritability of the proportions of Z9-14:OAc and Z11-14:OAc were substantial (0.5–0.6). In artificial selection experiments, the mean % Z9-14:OAc in the sex pheromone changed significantly within three generations (37% in the control line, 48 and 52% for two lines selected for increase, and 23 and 30% for two lines selected for decrease). Despite these changes, the amounts of fatty acyl pheromone precursors, (Z)-9-, (Z)-11- and (E)-11-tetradecenoate, in the pheromone gland were not significantly affected by the selection. Taken together, variation in the pheromone blend of O. zaguliaevi is likely to be attributable to a few genes involved in the reduction or acetylation of fatty acyl pheromone precursors, the last two steps in pheromone biosynthesis.     

Mating system of the cooperatively breeding noisy miner Manorina melanocephala,as revealed by DNA profiling

In this study we examined parentage within broods of the cooperatively breeding noisy miner using multi-locus DNA profiling. Previous studies of noisy miners described them as highly promiscuous, leading to the suggestion that promiscuous mating behaviour was a tactic used by females to recruit males as provisioners to their nests (Dow 1978). At our study site in south-east Queensland, Australia, we found that both multiple and extra-group paternity (i.e. a female mating with a male outside the group of male provisioners at her nest) were rare. In nests where multiple paternity was possible (i.e. clutch size > 1) 97% of 31 broods were sired by only a single male. Overall, 96.5% of all nestlings (n = 85) were the result of monogamous matings. Also, at the vast majority of nests, the male that sired the nestlings was also the main provisioner among all male nest attendants. Our results show that the mating system of the noisy miner can no longer be considered cooperative polyandry (Brown 1987) but is clearly genetic monogamy. We discuss the implications of this finding for understanding the complex social system of this species.     

Regulation of worker reproduction in bumblebees (Bombus terrestris): workers eavesdrop on a queen signal

In the annual bumblebee Bombus terrestris, the onset of queen-worker conflict over male production is seasonally and socially constrained. Workers will do better if they start to reproduce (the so-called competition phase) only after ascertaining that larvae are committed to gyne development but before the season ends because they gain more by rearing sister-gynes than their own sons. Here, we tested two nonmutually exclusive hypotheses as to what triggers the onset of worker reproduction: Workers can directly monitor larval development and/or workers eavesdrop on the queen signal that directs gyne development. Exposing workers to gyne larvae through a double mesh did not advance the competition phase compared to control colonies. However, when workers, but not the queen, were allowed contact with gyne larvae, both the competition phase and gyne production were advanced. Thus, while larvae do not emit a volatile pheromone that discloses their developmental route, the physical contact of workers with such larvae triggers early competition phase. However, workers exclusively exposed to worker larvae (colonies prevented from producing gyne larvae) started to reproduce at the same time as control colonies. Replacing the resident queen with an older queen (from gyne-rearing colonies) advanced the competition phase, irrespective of worker age. The results are consistent with the hypothesis that workers eavesdrop on the queen pheromones. This is adaptive because it allows workers a broader time-window for reproduction and thus to gain fitness from rearing both sister-gynes and sons before the season ends without affecting colony development.     

Sex pheromone of the large aspen tortrix, Choristoneura conflictana(Lepidoptera: Tortricidae)

Summary. Three components that elicited antennal response from male Choristoneura conflictana were found from female gland extracts analyzed using a coupled gas chromatographic-electroantennographic detector system. The main component in gland extracts was (Z)-11-tetradecenal (Z11-14:Ald). Two minor components also elicited antennal response: (E)11-tetradecenal (E11-14:Ald) and (Z)-11- tetradecen-1-ol (Z11-14:OH). Analysis of effluvia indicated that calling virgin females release mostly Z11-14:Ald and trace amounts of Z11-14:OH. Field and wind tunnel behavioral studies showed that Z11-14:Ald alone attracted male moths in a dose response pattern. Tests comparing male response to blends of components detected in gland extracts showed that addition of 1.8% of E11-14:Ald to Z11-14:Ald did not influence male moths in the wind tunnel, but resulted in significantly lower trap captures in the field. The threecomponent blend [Z11-14:Ald (100), E11-14:Ald (1.8), Z11-14:OH (11)], was less attractive than Z11-14:Ald alone in both field and wind tunnel studies. Traps baited with two virgin female moths were equally attractive to males as traps baited with the three-component synthetic blend but less attractive than traps baited with Z11-14:Ald alone. Field tests of various blends of the two components (Z11-14:Ald, Z11-14:OH) detected in the females’ effluvia showed that the addition of 1–10% Z11-14:OH to Z11-14:Ald did not affect the males’ response to Z11-14:Ald. Our data demonstrate that female C. conflictana release sex pheromone components in a different ratio than they are stored in the pheromone gland. The sex pheromone is comprised of a single component, Z11-14:Ald, that can be used to monitor mated and virgin male C. conflictana throughout their flight period.     

Asymmetric costs favor female desertion in the facultatively polyandrous northern flicker (Colaptes auratus): a removal experiment

Previous studies of biparental care in birds have focused on the males contribution and experimental removal of males. Woodpeckers, with a high level of paternal care including nocturnal incubation and brooding by males, offer a meaningful system in which to examine the importance of care by females. I studied the reproductive performance of 17 widowed male and 8 widowed female northern flickers (Colaptes auratus) relative to biparental (control) pairs. Of these single parents all widowed shortly after hatching, only one female abandoned its nest. Single parents boosted their provisioning rates to achieve 83% the provisioning rate of control broods, but reared significantly fewer young and young of poorer quality. However, single males, with 85% of the reproductive success of controls, were more successful than females with 43% the success of controls. Among widowed birds, a not significant lower survival was observed, but the chance of re-pairing with the same partner in a subsequent year was only 16% in the natural population, so long-term costs of desertion may be small. Although females seem to have the incentive and ability to desert, a lack of available males may constrain opportunities. This study demonstrates that when parental care roles are reversed in altricial birds, asymmetric benefits can favor female and not male desertion.     

Asexuals looking for sex: conflict between species and mate-quality recognition in sailfin mollies (Poecilia latipinna)

When two closely related species are sympatric the process of species recognition (identifying conspecifics) and mate-quality recognition (increased fitness benefits) can yield a conflict when heterospecifics resemble high-quality conspecifics. Conflict in species versus mate-quality recognition may serve as a possible mechanism for the persistence of unisexual, gynogenetic Amazon mollies (Poecilia formosa). Amazon mollies require sperm from closely related species (e.g., sailfin mollies, P. latipinna) to start embryogenesis but inheritance is strictly maternal. When choosing mates, male sailfin mollies from populations sympatric with Amazon mollies may rely on traits indicating species identity rather than those indicating mate quality. Conversely, males from allopatric populations may rely more on traits indicating mate quality. Previous work has found that male sailfin mollies in sympatry exhibit a significantly greater mating preference for female sailfin mollies over Amazon mollies compared to males in allopatry. In addition, male sailfin mollies prefer to associate with and produce more sperm in the presence of larger conspecific females, which are more fecund. We hypothesized that male sailfin mollies experience a conflict in species recognition and mate-quality recognition in the presence of Amazon mollies that are relatively larger than female sailfin mollies. To test this hypothesis, we paired males from sympatric and allopatric populations with a larger Amazon molly and a smaller female sailfin molly. We scored the number of mating attempts that males directed to conspecific and heterospecific females. Males in most sympatric and allopatric populations demonstrate no clear preference for conspecifics. In addition, we found some evidence for a difference in mating preference between allopatric and sympatric populations with males from allopatry showing a greater heterospecific mate preference. These results indicate a conflict between species and mate-quality recognition. In sympatry this conflict may contribute to the persistence of gynogenetic Amazon mollies.