Offensive and defensive sperm competition roles in the dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae)

Sperm competition is predicted to generate opposing selection pressures on males. On one hand, selection should favour ‘defensive’ adaptations that protect a male’s ejaculate from displacement, while, on the other hand, selection should favour ‘offensive’ adaptations that overcome paternity assurance mechanisms of rivals. Here, we use the sterile male technique to assess sperm precedence when a male dung beetle Onthophagus taurus mates in both a defensive (first male) and an offensive (second male) role. Significant variation in a male’s sperm precedence (both P ₁ and P ₂) was detected, and an individual’s defensive (P ₁) and offensive (P ₂) abilities were positively correlated. Thus, it appears that sexual selection simultaneously selects for ‘defensive’ and ‘offensive’ adaptations in O. taurus. We discuss a variety of male traits in O. taurus that potentially contribute to a male’s ability to be successful when mating in an ‘offensive’ and a ‘defensive’ role.     

Correlations of food distribution and patch size with agonistic interactions in female vervets (Chlorocebus aethiops) and patas monkeys (Erythrocebus patas) living in simple habitats

Food distribution is hypothesized to be important in determining the nature of female relationships within social groups of primates. When food limits female reproductive success, spatially clumped foods are expected to produce strong, linear dominance hierarchies within groups, whereas more spatially dispersed foods are expected to produce weaker or non-existent dominance hierarchies. The association between food distribution and competitive relationships presumably occurs because clumped foods are usurpable but dispersed foods are not. We examined the spatial distribution of food patches (trees) and patch size relative to feeding behavior and agonistic interactions in vervets and patas monkeys, two closely related and sympatric species that nonetheless differ in the strength of the female dominance hierarchy. Food patches of both patas monkeys and vervets were small in size and randomly distributed in Acacia drepanolobium habitat. In contrast, in A. xanthophloea woodland, the habitat type that was exclusively used by vervets, food patches were larger and more spatially clumped. These similarities and differences between and within species were correlated with similarities and differences in the strength and linearity of their dominance hierarchies. Patas monkeys and vervets in A. drepanolobium habitat had dominance hierarchies that were weakly defined because there were relatively few agonistic interactions between females. By contrast, in A. xanthophloea habitat, vervets had a stronger, linear dominance hierarchy characterized by a higher rate of agonistic interactions over food. The covariation of agonistic interactions with patch size is discussed in relation to depletion time, another characteristic that may covary with food distribution, and resource renewal rate, an important determinant of agonistic interactions in insectivorous birds, fishes, insects, and mammals. Received: 18 February 2000 / Revised: 5 September 2000 / Accepted: 26 September 2000     

Sperm competition and evidence of sperm fertilization patterns in the carrion ball-roller beetle Canthon cyanellus cyanellus LeConte (Scarabaeidae: Scarabaeinae)

Field observation and laboratory experiments were conducted to investigate sperm competition and patterns of sperm fertilization under different experimental conditions in the carrion ball-roller beetle Canthon cyanellus cyanellus. Males in nature can mate with females whose spermathecae contains fertile sperm from other males. Sperm precedence was investigated using a visible genetic marker. The progeny of red (homozygous recessive) virgin females mated once with a red male and afterward, once with a green beetle (homozygous dominant) and vice versa, revealed that regardless of its color, the last male to mate fertilized c.a. 50% of the eggs, suggesting strong sperm competition. Males were able to achieve higher levels of paternity (more than 80%) when mated ad libitum with previously mated females, although large amount of variance in paternity does not exclude the possibility of first male sperm precedence or female cryptic choice. These results suggest that repeated mating and sperm replacement are the mechanisms by which last males achieve sperm precedence.     

Sexual selection in the northern pintail (Anas acuta): the importance of female choice versus male-male competition in the evolution of sexually-selected traits

We experimentally studied the relative importance of plumage, dominance status, and courtship behavior in determining male pairing success in the northern pintail Anas acuta and assessed whether these traits function in female choice, male-male competition or both. In an experiment (experiment IA) that eliminated the confounding effects of male-male competition and social courtship, females chose males with pure white breasts and colorful scapular feathers. When the same group of birds were free to interact (experiment 1B), male behavior was more important: females chose males that courted them intensely and were attentive to them, although preferred males again had whiter breasts and more colorful scapulars. In a second experiment (experiment 2), testing the effect of age on pairing success, females showed a significant preference for 2-year-old males over yearlings: 2-year-old males courted more and were more attentive to the female than yearlings; they were also more colorful than yearlings in a number of plumage measurements. Although males (in both experiments 1B and 2) were aggressive to one another while courting the female and dominant males were sometimes able to exclude subordinates from social courtship, contrary to expectation, we found no relationship between initial dominance rank and pairing success or dominance rank and age. In addition, dominance was not correlated with any of the morphological traits measured. Once chosen, however, subordinate males typically initiated fights with the higher-ranked male(s) and quickly achieved dominance. These results suggest that (1) females choose males based on a suite of morphological and behavioral characteristics, (2) male dominance relationships do not constrain active female choice, (3) a male's position in a dominance hierarchy is largely a result rather than a cause of female choice, and (4) female choice plays a more significant role than male-male competition in the evolution of several secondary sexual traits in male northern pintails.     

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.     

Maintenance of variance in sperm transfer rates in a scorpionfly: food availability,genetic basis,and heritability

Variation in traits closely related to fitness is expected to be low. This is because these traits are under directional selection and the best genotype should prevail. However, there have been a number of studies demonstrating the existence of considerable variance in sexually selected traits, which is generally known as the lek paradox. Accordingly, earlier studies found substantial variation in sperm transfer rates in Panorpa vulgaris. Aiming at finding the mechanism that maintains this variation, we analyzed the condition dependence and the narrow sense heritability of sperm transfer rates. Food deprivation in the larval/adult phase caused a decrease in the males’ capability of saliva secretion resulting in shorter copulations and a reduced number of transferred sperm. There was a positive correlation between mean sperm transfer rates and mean body mass. Additionally, intermale variation in sperm transfer rates decreased with increasing food availability. Hence, we suggest that sperm transfer rates in P. vulgaris are influenced by adult feeding history. Heritability analyses of sperm transfer rates did not provide significant results, which is consistent with the general hypothesis that additive genetic variance in traits closely related to fitness is small. Since a trait’s potential to respond to selection is proportional to the amount of contained additive genetic variance, the ascertained small heritability provides a satisfying explanation for the maintenance of substantial variation in sperm transfer rates.     

Male care of infants in a siamang (Symphalangus syndactylus) population including socially monogamous and polyandrous groups

While male parental care is uncommon in mammals, siamang (Symphalangus syndactylus) males provide care for infants in the form of infant carrying. I collected behavioral data from a cohort of five wild siamang infants from early infancy until age 15–24 months to identify factors affecting male care and to assess the consequences of male care for males, females, and infants in a population including socially monogamous groups and polyandrous groups. There was substantial variation in male caring behavior. All males in polyandrous groups provided care for infants, but males in socially monogamous groups provided substantially more care than males in polyandrous groups, even when the combined effort of all males in a group was considered. These results suggest that polyandry in siamangs is unlikely to be promoted by the need for “helpers.” Infants receiving more care from males did not receive more care overall because females compensated for increases in male care by reducing their own caring effort. There was no significant relationship between indicators of male–female social bond strength and male time spent carrying infants, and the onset of male care was not associated with a change in copulation rates. Females providing more care for infants had significantly longer interbirth intervals. Male care may reduce the energetic costs of reproduction for females, permitting higher female reproductive rates.     

Influences of moonlight,ambient temperature,and food availability on the diurnal and nocturnal activity of owl monkeys (<Emphasis Type="Italic">Aotus azarai</Emphasis>)

The study of activity rhythms, their potential zeitgebers and masking factors among free-ranging primates has received relatively little attention in the past. Most primates are diurnal, a few of them nocturnal, and even fewer are cathemeral. Owl monkeys (Aotus azarai azarai) regularly show diurnal, as well as nocturnal, activity in the Argentinean and Paraguayan Chaco. The goal of this study was to examine how changes in activity patterns in owl monkeys of Formosa, Argentina are related to daily, monthly, and seasonal changes in temperature, light and food availability . During 1 year, I collected activity data from five groups followed continuously from dawn to dusk, dusk to dawn or uninterruptedly during 24 or 36 h for approximately 1,500 h. I kept hourly and daily records of temperature and light conditions, and I gathered monthly information on the density, distribution and abundance of food resources available to the monkeys. I found that the area of study is highly seasonal, and characterized by significant fluctuations in rainfall, temperature, photoperiod, and food availability. Owl monkeys had on average 5 h of activity during the day and 4 h during the night. The amount of diurnal activity remained fairly constant through the year despite seasonal changes in exogenous factors. Owl monkeys did not show changes in their activity patterns that could be attributed to changes in food availability. Nocturnal activity increased as the amount of moonlight increased, whereas diurnal activity decreased following a full-moon night. Ambient temperature was a good predictor of activity only when the moon was full. These results argue convincingly for an interaction between ambient temperature and moonlight in determining the observed activity pattern. It is then highly advisable that any evaluation of diurnal activity in cathemeral animals be analyzed controlling for the possible effects of moonlight during the previous night.Communicated by P. Kappeler     

Fertilisation success in the fly Dryomyza anilis (Dryomyzidae): effects of male size and the mating situation

Male fertilisation success in relation to male size and the mating situation (ordinary pair formation with a single, nonvirgin female vs. take overs) was examined in the fly Dryomyza anilis. In ordinary matings, large males achieved higher fertilisation success than small ones when they were the second to mate with the female. Take overs differ from ordinary pair formation in that the second male experiences intensified sperm competition. This is because in take overs the female is not able to discharge any of the sperm inseminated by the first male as she usually does before a new mating. Compared with ordinary matings, take overs reduced the fertilisation success of the second male by 8–10%, whereas that of the first male was 7–14% higher in take overs. Even though the intruder was always larger than the paired male his superior fertilisation success did not compensate for the effect of the sperm already present in the female. In D. anilis, males can increase their fertilisation success by tapping the female's external genitalia with their claspers or having several copulation bouts per mating. Thus, in a take over, the intruder could respond to the intensified sperm competition by performing more tapping sequences per copulation bout or more copulation bouts per mating. In matings observed in the wild, males performed more tapping sequences after a take over than after pair formation with a single female, although the difference was not significant. The results show that there are differences in fertilisation success between males of different size. In addition, different mating situations can result in considerable variation in the fertilisation success of an individual male. Higher fertilisation success for the first male after a take over may be significant, in particular, for the reproductive success of small males, which frequently lose their females to large males.     

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.     

Species,population and individual specific odors in urine of mole rats (Spalax ehrenbergi) detected by laboratory rats

Summary Urine odors from 4 species of blind subterranean mole rats belonging to theSpalax ehrenbergi superspecies in Israel were evaluated to determine whether there were discriminable differences in the odors that were specific to different individuals, populations, and species of mole rats. Trained laboratory rats assessed these differences in an automated olfactometer using an operant conditioning paradigm. They demonstrated the discriminability of the urine odors in the 3 categories by their correct responses in generalization trials. These characteristic species, population and individual odors may provide these solitary, territorial and blind rodents a means for diverse chemical communication in spacing behavior, reproduction, species isolation and speciation.     

Copper tolerance in a population of Silene vulgaris ssp. maritima (A. & D. Love) at Dolfrwynog Bog near Dolgellau,North Wales

Clones of Silene vulgaris ssp. maritima individuals from Dolfrwynog Bog, North Wales and from a coastal control site were propagated under standardised environmental conditions. After five months, cuttings were taken and tested for copper tolerance by means of root extension analysis in solutions of various different copper concentrations. Indices of tolerance suggested that individuals from the Dolfrwynog population were more resistant to elevated copper concentrations than were the control population individuals. Due to the initial five months under standardised conditions, it was assumed that the cloned plant material would have lost all plastic adaptive responses to its native environment. If this assumption was valid then the differences in copper tolerance observed between the two populations had a genetic basis and the Dolfrwynog Bog population is a copper tolerant ecotype.     

Manipulation of nest-box density affects extra-pair paternity in a population of blue tits (<Emphasis Type="Italic">Parus caeruleus</Emphasis>)

We tested the effect of manipulation of breeding density on the occurrence of extra-pair paternity in a blue tit (Parus caeruleus) population during two consecutive years. In a homogeneous oak forest, nest-box manipulation provided a high density plot (plot A, 1.10 and 1.32 pairs/ha) and a low density plot (plot B, 0.43 and 0.46 pairs/ha). Microsatellite analysis on 91 broods revealed a higher proportion of extra-pair paternity in broods in plot A (mean of 17.2%) than in plot B (mean of 11.4%). A correlative approach showed that the proportion of extra-pair young in broods was affected by the number of breeding neighbours within 100 m around the nest-box, by the distance to the nearest breeding neighbour and by an additional plot effect. However, the nearest neighbours accounted for only 39.3% of extra-pair paternities and distance to extra-pair fathers was significantly higher than the nearest neighbour distance in both plots. This implies that the effect of density on the occurrence of extra-pair paternities is associated with active female choice to enhance the brood fitness. Although there were more extra-pair young in broods when density was high, the number of extra-pair fathers did not increase and stayed close to one. We suggest that density increases the cost of mate guarding by males, thereby increasing the possibility for females to solicit extra-pair paternities to the cuckolding male they have chosen. Finally, we discuss why correlatives approaches do not always show evidence for an effect of breeding density on extra-pair paternity occurrence.Communicated by M. Soler     

Social transmission of food handling in the context of triadic interactions between adults and young canaries (<Emphasis Type="Italic">Serinus canaria</Emphasis>)

We studied the factors that enhance food recognition and consumption in young canaries when confronted with adults. In contrast to previous studies on canaries, in which social transmission of food habits was studied in the context of dyadic interactions (one juvenile–one adult), we proposed a more realistic framework in which young canaries were studied in the context of triadic interactions, free or not, with adults of both sexes. We found that during free interactions, the young bird only eats with a familiar male and that this association enhances the social transmission of seed handling. When the juvenile was separated from the adults by a transparent partition, it only learned to husk seed if it was present at the feeder at the same time as a familiar adult acting as a demonstrator. The presence of adults that are familiar but do not act as demonstrators does not facilitate social transmission of handling. However, the presence of a familiar, demonstrating female had also no effect on this transmission. Coordination of the actions of the experienced bird and of the naive subject is required for social transmission to occur. Action coordination does not depend solely on the level of familiarity between partners but also on the role played by the demonstrator (here, the adult male) that looks after the juvenile during its transition towards independence.     

Oily substances from gastral intersegmental glands of the antPachycondyla tridenta (Ponerinae): Lack of pheromone function in tandem running and antibiotic effects but further evidence for lubricative function

Summary Gas chromatographic analyses of volatizable material from gaster intersegmental complex glands ofP. tridentata revealed the presence of linoleic acid, palmitic acid, methyl oleate, and several long-chain hydrocarbons as major constituents, which form an oily fluid mixture. The nest relocation communication ofP. tridentata is tandem running. Tandem following is mediated by pheromones as demonstrated by dummy experiments with isolated gasters and thoraces and with glass balls. The secretions of all gaster complex glands, as well as hind gut contents and metapleural gland secretions, were unable to evoke tandem following. Morphology and position of the glands, lack of pheromonal function, oily properties, low volatility, and lack of antibiotic effects of these secretions strongly suggest a function as lubricants for the ants' gastral segments.     

Identifying key life-traits for the dynamics and gene flow in a weedy crop relative: Sensitivity analysis of the GeneSys simulation model for weed beet (Beta vulgaris ssp. vulgaris)

The benefits of genetically modified herbicide-tolerant (GMHT) sugar beet (Beta vulgaris) varieties stem from their presumed ability to improve weed control and reduce its cost, particularly targeting weed beet, a harmful annual weedy form of the genus Beta (i.e. B. vulgaris ssp. vulgaris) frequent in sugar beet fields. As weed beet is totally interfertile with sugar beet, it is thus likely to inherit the herbicide-tolerance transgene through pollen-mediated gene flow. Hence, the foreseeable advent of HT weed beet populations is a serious threat to the sustainability of GM sugar beet cropping systems. For studying and quantifying the long-term effects of cropping system components (crop succession and cultivation techniques) on weed beet population dynamics and gene flow, we developed a biophysical process-based model called GeneSys-Beet in a previous study. In the present paper, the model was employed to identify and rank the weed life-traits as function of their effect on weed beet densities and genotypes, using a global sensitivity analysis to model parameters. Monte Carlo simulations with simultaneous randomization of all life-trait parameters were carried out in three cropping systems contrasting for their risk for infestation by HT weed beets. Simulated weed plants and bolters (i.e. beet plants with flowering and seed-producing stems) were then analysed with regression models as a function of model parameters to rank processes and life-traits and quantify their effects. Key parameters were those determining the timing and success of growth, development, seed maturation and the physiological end of seed production. Timing parameters were usually more important than success parameters, showing for instance that optimal timing of weed management operations is more important than its exact efficacy. The ranking of life-traits though depended on the cropping system and, to a lesser extent, on the target variable (i.e. GM weeds vs. total weed population). For instance, post-emergence parameters were crucial in rotations with frequent sugar beet crops whereas pre-emergence parameters were most important when sugar beet was rare. In the rotations with frequent sugar beet and insufficient weed control, interactions between traits were small, indicating diverse populations with contrasted traits could prosper. Conversely, when sugar beet was rare and weed control optimal, traits had little impact individually, indicating that a small number of optimal combinations of traits would be successful. Based on the analysis of sugar beet parameters and genetic traits, advice for the future selection of sugar beet varieties was also given. In climatic conditions similar to those used here, the priority should be given to limiting the presence of hybrid seeds in seed lots rather than decreasing varietal sensitivity to vernalization.