Kinship, familiarity and social status modulate social learning about “micropredators” (biting flies) in deer mice

Animals can learn to recognize and respond to dangerous, threatening factors through either individual or social learning, whereby an individual learns and acquires the defensive behaviors and avoidance responses of another. Here we show that kinship, familiarity, and relative dominance of the interacting individuals affect social learning of defensive responses to micropredators in deer mice, Peromyscus maniculatus. Brief exposure of individual male deer mice to biting flies (stable fly, Stomoxys calcitrans) induced a decrease in pain sensitivity, or analgesia, and active self-burying avoidance responses. The defensive analgesic responses and their fear/anxiety/stress associated correlates facilitate the display of behavioral avoidance responses. Preparatory analgesia and avoidance responses were evident 1–3 days later when the mice were exposed to biting flies that were altered to be incapable of biting but were not displayed to similar-sized non-biting house flies. These anticipatory avoidance and analgesic responses to biting flies were also acquired through social learning—without direct individual aversive experience with biting flies. Fly-naive mice (observers) that witnessed other mice (demonstrators) being attacked by biting flies but themselves were not bitten did not display any avoidance responses. However, when exposed 1–3 days later to altered flies with biting mouth parts removed, the observers displayed socially acquired analgesic and self-burying avoidance responses. Observers whose demonstrators were either kin (siblings) and, or members of a familiar pair (kin or non-kin) displayed enhanced social learning of defensive responses. Also, within the familiar pairs, social status affected learning with subordinate observers displaying better social learning than dominant observers. These findings indicate that kinship, familiarity and social status modulate social learning of defensive responses to, and the recognition of, dangerous and threatening stimuli, likely including that of predators.Communicated by G. Wilkinson     

A fungus infecting domestic flies manipulates sexual behaviour of its host

The Enomopathogenic fungus Entomophthora muscae infects and kills domestic flies Musca domestica, and rates of transmission are higher at high population densities. I tested whether features of parasitized hosts increased the likelihood of transmission of the fungus in a series of experiments. Individual flies contacting an infected conspecific had a much higher probability of becoming infected than had flies contacting a dead fly without signs of infection. Males especially were attracted to and behaved sexually towards dead, infected flies even when a choice was available between a dead, infected fly and a dead, uninfected individual. The abdomen of infected flies swelled considerably as a consequence of infection, and uninfected female flies with larger abdomens are more fecund than the average female and maybe more attractive. Experiments in which abdomens of infected and uninfected flies were cut off and glued back on other flies demonstrated that the abdomens of infected flies were highly attractive to males. When the size of the abdomen of infected and uninfected individual flies was held constant, infected dead flies were still more attractive than corpses of uninfected flies. This result suggests that features other than the size of the swollen abdomen enhanced the attractivity of infected flies. Features of infected hosts therefore increased the likelihood of transmission of the parasitic fungus.     

Socially facilitated egglaying behavior in Mediterranean fruit flies

We examined the behavior of individual mature female Mediterranean fruit flies (medflies), Ceratitis capitata (Wiedemann), transferred from a holding cage without fruit to a clean host kumquat fruit already occupied by another medfly female engaged in ovipositional behavior. A significantly greater proportion of transferred (=test) ovipositionally naive females initiated ovipositor boring into a fruit in the presence than in the absence of an occupying medfly. Moreover, test females commenced boring significantly sooner in the presence than in the absence of another medfly. Interestingly, similar results occurred when a test medfly was transferred to a kumquat occupied by an ovipositing female oriental fruit fly, Bactrocera dorsalis Hendel. In contrast, there was no enhancement of propensity to bore when a test medfly was transferred to a kumquat occupied by a female Biosteres arisanus (Sonan), a parasitoid attacking medfly eggs. The principal proximate stimulus giving rise to the higher tendency to oviposit of a test medfly in the presence of an occupying medfly or oriental fruit fly appeared to be wing-waving by the occupant during an encounter. We consider our findings to be good evidence of socially facilitated ovipositional behavior in ovipositionally naive medflies, and suggest that such females may benefit from acquisition of a cue demonstrating the acceptability of a host for oviposition. Received: 7 July 1997 / Accepted after revision: 30 August 1997     

Influence of male harassment and female competition on female feeding behaviour in a sexual–asexual mating complex of mollies (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, <Emphasis Type="Italic">P. formosa</Emphasis>)

Differential interests between the sexes regarding the number of copulations can result in sexual harassment. Hence, females may have less time available for foraging. Male sexual harassment often leads to fitness reduction in females. We used the mating complex of the bisexual fish Poecilia mexicana and the co-occurring all-female Poecilia formosa to study sexual harassment and its incurred cost on female feeding efficiency. P. formosa is a sperm-dependent parthenogen that requires mating with host males to induce embryogenesis, but the male genes are not used. We therefore predicted P. mexicana males to prefer conspecific females. Hence, costs of male sexual harassment should not occur in unisexuals. While P. formosa are at a disadvantage compared to P. mexicana females due to male mate choice (leading to sperm limitation), this could be traded-off by suffering less from sexual harassment. In our experiment, we found males to direct significantly more pre-copulatory mating behaviour towards conspecific females, whereas actual mating attempts did not differ between species. Contrary to our prediction, both types of females started feeding later and spent less time feeding in the presence of a male partner compared to the time spent feeding with another female, suggesting that females of both species suffer from male harassment. The focal females' feeding time declined with increasing body size of the female competitor, and the same pattern was found when a male was present. We discuss that—besides sexual harassment—other factors such as food competition and female mate choice may affect female feeding efficiency.     

Effects of parasites on foraging and defense behavior of a termitophagous ant,Pheidole titanis Wheeler (Hymenoptera: Formicidae)

Summary Pheidole titanis Wheeler, an ant that occurs in desert and deciduous thorn forest in the southwestern United States and western Mexico, is a predator on termites. In the dry season well-coordinated raids against termite foraging parties occur early in the morning or late in the afternoon, whereas in the wet season most raids occur at night. This seasonal shift in the timing of raids is due to the increased activity of a fly (Diptera: Phoridae) that is a specialist parasitoid on P. titanis workers and soldiers. When parasitic flies discover P. titanis nest entrances or raiding columns, workers stop foraging and defend themselves against oviposition attacks. Flies are only active during the day and never interfere with foraging at night. However, P. titanis does not increase the frequency of raids at night and, as a result, colonies collect less food in the wet season compared to the dry season. Presence of parasitic flies also interferes with normal defense behavior of P. titanis against conspecific and heterospecific enemy ants. Dissections of P. titanis workers and soldiers suggest that the parasitism rate by flies is less than 2% and observations indicate that parasitic flies are much rarer than their host workers and soldiers. Nonetheless, these parasites exert a strong ecological impact on their host.     

Innate responses to male sexual harassment in female mosquitofish

Male mosquitofish are very persistent in their sexual activity and harass any female they encounter. Gravid females pay a large tribute to this intense male sexual activity in terms of reduced foraging efficiency. Previous observations have demonstrated that gravid females, when chased by a male, dilute male harassment by moving closer to other females to form shoals. They also approach other males to promote male competition, and when males differ in size, they preferentially target large males, whose harassment is less intense. In this study, we tested whether the modulation of females’ social preferences in response to male harassment is innate or learned. We tested social preference in three groups of females that differed in experience of sexual harassment and in the factors affecting it. Females of the first group were reared without any sexual experience, and pregnancy was induced through artificial insemination. The second group was composed of naive females kept singly with a male; these females experienced sexual harassment but were prevented from experiencing the effects of male–male competition and shoaling on the amount of male sexual harassment. In the third group (controls), females were reared in multi-male, multi-female groups and could experience the modulating effects of social interactions on sexual harassment. When exposed to a harassing male, females of the three groups immediately reduced their distance from another female, approached a group of males or moved toward the larger of two available males. Moreover, the results for these three groups of females were similar to those obtained in wild-caught females that were tested in the same three tests in a previous study (Dadda et al. An. Behav., 70:463–471, 2005). This suggests that the strategies adopted by females in response to male sexual harassment do not need to be learned through specific experience of the social contexts.     

Manganese toxicity in Drosophila melanogaster: extension of the life span by resveratrol

The effect of resveratrol was studied on the life span and motor activity of Drosophila melanogaster treated with manganese (Mn). Two days after emerging from the pupa, male wild type D. melanogaster were fed for 13 days with corn media containing 15?mM Mn. Thereafter, Drosophila were divided into six groups of 300 flies each: (1) the flies remained treated with Mn; (2) began treatment with 0.43?mM resveratrol (Mn-resveratrol group); (3) received no additional treatment (Mn-no treatment group); (4) simultaneously fed with Mn and resveratrol (Mn?+?resveratrol group). In addition, a control (5) with no treatment and another group (6) fed only with resveratrol after emerging from the pupa were included. All Mn-treated flies (group 1) were dead on the 25th day. The life span in the resveratrol group was 91?±?0.33 days (mean?±?S.E.M.) and in Mn-resveratrol flies was 83?±?2 days. These two values were significantly higher than those detected in the control (5) and Mn-no treatment (group 3) flies whose life span were 68?±?0.33 and 67?±?2.31 days, respectively. The Mn?+?resveratrol-fed flies had a markedly higher life span (31?±?1.53 days) than Mn-fed flies (23?±?0.88 days). In the flies that received Mn (Mn and Mn?+?resveratrol groups), the motor activity decreased significantly with respect to control (groups 5) and the Mn–resveratrol and resveratrol groups. In conclusion, resveratrol increased significantly the life span of Mn-treated D. melanogaster.     

Calling characteristics of parasitized and unparasitized populations of the field cricket Teleogryllus oceanicus

Summary Examination of three populations of the field cricket Teleogryllus oceanicus revealed the presence of an acoustically-orienting parasitoid fly, Ormia ochracea (Tachinidae), in the population of crickets that has been introduced to the Hawaiian Islands. The cricket is native to Australia and the Pacific, and the fly is native to North America but has also been introduced to Hawaii. Up to 27% of males and 7% of females in Hawaii were infested with fly larvae. Song structure in the parasitized Hawaiian population was distinct from that of the other two groups, with the Hawaiian crickets showing several reduced song parameters. In addition, onset and cessation of calling at dusk and dawn were more abrupt in the Hawaiian population. These results are consistent with selective pressure from the phonotactic flies to decrease risky calling. Silent males were present in all three populations, suggesting that these noncallers may not represent a unique adaptation to the parasitoid. Correspondence to: M. Zuk     

The response of an insect parasitoid,Ormia ochracea (Tachinidae), to the uncertainty of larval success during infestation

Ormia ochracea is a parasitoid fly which lays its larvae on its hosts, the field crickets Gryllus integer and Gryllus rubens, in two distinct modes: (1) directly on the host and (2) around the host. In the field, 12.7% of male crickets were parasitized and 3.2% were super-parasitized. Despite the disadvantages of parasitizing infested hosts, there was no evidence that O. ochracea avoided superparasitism. This and other experiments suggest that the host assessment ability of O. ochracea is less than that reported for many hymenopteran parasitoids. by manipulating the number of larvae in each cricket, we determined that four to five larvae per host resulted in the largest number of adult flies. However, as larval number per host increased from one to six, pupal size, and hence adult size, declined. In the field, hosts were found with a mean of 1.7 ± 1.0 (SD) larvae per cricket, suggesting that there may be some selection pressure against larger clutch sizes. Nevertheless clutch sizes larger than the host can support were sometimes found in the field. During the first mode of larviposition, gravid flies deposited no more than three larvae directly onto the host. Larvae deposited directly on the host had a high probability of infesting it. During the second mode of larviposition, gravid flies laid a larger number of larvae around the host (6.1 ± 5.2). Larvae that were laid around the host were less likely to infest a cricket than were larvae that were deposited directly onto it. O. ochracea is unique in that its two different modes of larviposition have different probabilities of larval success. Even though the success rate for larvae laid during the second mode of larviposition was low, the possibility of parasitizing more hosts appears to have selected for flies laying more larvae (e.g. increasing clutch size) than is optimal if all the larvae successfully entered a single host.     

Polychlorinated biphenyl (PCB) residues in European roe deer (Capreolus capreolus) and red deer (Cervus elaphus) from north-western Poland

The purpose of this study was to evaluate polychlorinated biphenyls (PCBs) contamination levels in roe and red deer from north-western Poland and to assess environmental pollution in this area. A quantitative analysis was conducted using a capillary gas chromatography/mass spectrometry method. The mean concentrations of ΣPCBs (sum of PCBs: 28, 52, 101, 138, 153, 180) in liver samples were 30.24±12.35 ng·g^?1 of lipid weight (l.w.) in roe deer and 60.13±14.23 ng·g^?1 l.w. in red deer, compared with 24.21±10.02 and 45.22±9.77 ng·g^?1 in the lungs of roe and red deer, respectively. PCBs 138, 153 and 180 were the dominant congeners in the liver samples of the analysed animals, whereas PCB 138 and 153 in the lungs. TEQs levels calculated for only dioxin-like PCBs were low: 0.32 and 0.29 pg WHO-PCB-TEQ·g^?1 fat in liver of red deer and roe deer, respectively. The mean PCB concentrations obtained in our study for organs of roe deer and red deer were several times lower than those reported elsewhere. These findings show that the investigated roe and red deer originated from an area with low levels of PCB contamination.     

Should consumptive sustainable use of Sika Deer (Ceruus nippon) in Dilijan Reserve,NE Armenia be instituted?

SUMMARY

Numbers of sika deer (Ceruus nippon), resident in the Dilijan Reserve in 1969 (52 animals), 1970 (50), 1972 (1), and 1976 (28) dramatically increased in number until 1981, when the population reached a ceiling, being the number which can subsist in the area—300 individuals (density—11–12.5 deer per 1000 ha). Thus, Dilijan Reserve's C. nippon population develops in a way which resembles an S-like log curve equation.

Actually, Dilijan Reserve no longer has the status of a protected area—nowadays lumbering activities predominate and, inevitably, the unique broadleaf deciduous forest ecosystem will run down to extinction. Such economically valuable species as oak, beech, and sika deer are likely to be the first victims of this large-scale devastation.

The present article does not reject exploitation of any resource (including sika deer), if necessary. The key question is: how C. nippon can be sustainably utilized, so that the number of culled animals is replenished within the same year and and so that eventually the population size will remain constant?     

Differences in the electroantennal responses of apple- and hawthorn-infesting races of Rhagoletis pomonella to host fruit volatile compounds

Summary. Domestic apple (Malus pumila)- and hawthorn (Crataegus sp.)-infesting races of Rhagoletis pomonella, Walsh (Diptera: Tephritidae) provide an excellent model to examine the role that host plant specificity plays during sympatric speciation (i.e., divergence in the absence of geographic isolation). Previous work has shown that these races differ in their propensities to accept apple and hawthorn fruits in behavioral choice assays, and that this discrimination translates into "host fidelity" in the field (i.e., apple flies tend to mate on and oviposit into apples and hawthorn flies on hawthorns). ?We present the results of a study examining possible physiological factors contributing to host choice differences in R. pomonella. We tested whether apple and hawthorn flies differ in their electroantennogram (EAG) responses to biologically relevant volatile compounds emitted from apples and hawthorns. Significant differences were found in the relative EAG responses of apple and hawthorn flies to host fruit compounds at five of six paired study sites across the eastern United States. The geographic pattern of EAG variation was complex, however, with local populations of apple and hawthorn flies tending to be more similar to one another than to flies of the same race at distant sites. This pattern was largely due to EAG responses for several compounds showing longitudinal or latitudinal clines, the latitudinal clines being similar to those observed for allozyme loci in the host races. We also found evidence for sex-related differences, as males tended to have higher mean EAG responses to compounds than females. Host-associated differences were therefore nested within geographic and sex-related differentiation in R. pomonella.?Further behavioral studies are needed to distinguish whether the EAG differences are responsible for, as opposed to being a consequence of, host-plant fidelity and adaptation. Crosses are also required to establish a genetic basis for the EAG responses, although we did find significant correlations between EAG scores for several compounds and the allozymes NADH-Diaphorase-2 and Hydroxyacid dehydrogenase at one of the study sites. Questions therefore remain concerning the evolutionary significance of the EAG response differences between apple and hawthorn fly races. Nevertheless, these differences raise the possibility that antennal responses to fruit-related volatile compounds contribute to host plant discrimination in R. pomonella. Regardless, the EAG responses represent another set of traits, in addition to diapause/eclosion time phenotypes and allozyme frequencies, differing between apple and hawthorn host races of R. pomonella. Received 17 March 1998; accepted 21 September 1989.     

Effects of male sexual harassment on female time budgets,feeding behavior,and metabolic rates in a tropical livebearing fish (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>)

Sexual harassment by males has the potential to affect almost any aspect of female behavior and life history. Using Atlantic mollies (Poecilia mexicana) as a model—a species in which males do not court but almost constantly try to forcefully mate with females—we asked whether and how male harassment influences (a) females’ time budgets and (b) feeding rates (e.g., through frequent flight from male approaches), and (c) whether metabolic rates are increased as a response to stress. Field observations in a natural P. mexicana population revealed that males (average feeding rate 15%) spent far less time feeding than females (60%), and clearly traded off frequent pursuit of females with foraging. Most importantly, females’ feeding times were dramatically reduced when being pursued by a harassing male. Also in standardized lab experiments, females spent significantly less time feeding when accompanied by a male as compared to being in the presence of another female. This effect was also observed when partner fish (male or female) were presented only visually, but could not interact physically with the focal female. It seems, therefore, that females increase vigilance when a harassing male is around, which keeps them from feeding even before males actually approach them. Based on the latter result, we asked whether a stress-induced increase in metabolic rates would be discernible. We measured oxygen consumption and gill ventilation frequencies (opercular rates) of females in different social contexts (alone, with another female, or a male). The predicted, strong body mass dependency of both physiological parameters was uncovered, but no evidence for an effect of social context was detected. We argue that male harassment represents such a constant (but non-lethal) stressor for poeciliid females that their metabolic stress responses have adapted to this through habituation.     

Influence of dung volatiles on the process of resource selection by coprophagous beetles

Summary. Most dung beetles colonize the faeces of several vertebrate species without much discrimination, and are thus often considered as polyphagous. Recent studies have provided evidence for clear feeding preferences in scarab beetles colonizing dung of herbivore species, but little is known about these insects’ abilities to discriminate among odours from faeces of various herbivores. In this study, trophic preferences were examined using blocks of pitfall traps baited with dung from four different herbivore species, i.e., sheep, cattle, horse, and red deer, in a mountainous area of south-central France. 4941 coprophagous scarabs, belonging to 27 species, were captured. Beetles were more attracted to dung of sheep (2257 individuals) than that of cattle (1294 individuals), followed by deer dung (768 individuals) and horse dung (622 individuals). Eleven of the 27 beetle species collected had significant feeding preferences for one of the four dung types. For each insect species, trophic habits did not vary between the two different sites of trapping, an open pasture and a wooded habitat. In laboratory olfactometer bioassays, scarab beetles orientated preferentially towards the dung volatiles from the dung type they preferred in the field. Trypocopris pyrenaeus, Anoplotrupes stercorosus, and Aphodius rufipes were more attracted to volatile compounds from sheep dung, Onthophagus fracticornis significantly preferred horse dung volatiles, and Aphodius haemorrhoidalis responded positively to deer dung odours. The role of dung olfactory cues in the process of resource selection by dung beetles is discussed.     

Effects of the Presence of Official‐Looking Volunteers on Harassment of New Zealand Fur Seals

Abstract: An increased number of tourists viewing animals in the wild have increased stress on these animals (hereafter wildlife). Many wildlife‐viewing locations rely on voluntary compliance with posted regulations to protect animals from tourists because of the expense of employing on‐site enforcement personnel. Voluntary compliance, however, is ineffective. The presence of official‐looking volunteers may decrease the incidence of wildlife harassment by tourists. To test this possibility, we observed tourists interacting with 5‐ to 12‐month‐old New Zealand fur seals (Arctocephalus forsteri) at the popular Ohau Stream waterfall while in the absence or presence of a young woman in plain sight wearing a neon vest (i.e., observer) and when an observer was not present. We observed 254 tourist groups at the waterfall when young seals were present. The percentage of groups in which at least one person harassed (approached, touched, or threw objects) a young seal was two‐thirds lower when the official‐looking observer was present. Frequency of harassment was inversely related to observer presence. Programs in which volunteers work at tourist sites are popular in countries with high tourism rates, such as New Zealand. Our results show that a relatively inexpensive and effective tourism‐management strategy may be to post such volunteers as observers at sites where tourists view wildlife.     

Temporal changes of Collembola population and alterations of life history parameters and acetylcholinesterase and superoxide dismutase activities in Cyphoderus javanus (Collembola) as biomarkers of fly ash pollution in lateritic soil

Fly ash is applied in agricultural fields to improve soil quality and crop yield; however, there are concerns regarding environmental hazards and toxicity to ecologically important soil organisms. The soil microarthropod fauna is a vital component of detritus food web, and major groups like Collembola are sensitive indicators of soil quality; however, information is scanty on their biomarker potentials against xenobiotics in tropical soils. The present study was aimed to evaluate temporal changes of Collembola population in fly ash amended field plots, and assess the biomarker potentials of life history parameters and biochemical responses such as acetylcholinesterase (AChE) and superoxide dismutase (SOD) activities in Cyphoderus javanus Borner (Collembola, Insecta), exposed to fly ash treated soil in microcosms. The field study using 5% (50 t ha^?1) and 20% (200 t ha^?1) doses of fly ash revealed dose-dependent and persistent decline in the density and relative abundance of Collembola population in sandy loam lateritic soil. The microcosm experiments showed negligible lethal effect of fly ash on C. javanus, but major life history parameters namely survival success, fecundity, and molting were significantly inhibited by fly ash treatments. The activity of AChE was downregulated, whereas activity of SOD was upregulated within 7 days of exposure of C. javanus to fly ash treated soil. These biological and biochemical parameters in Collembola are potential biomarkers, and therefore, the effects of fly ash are significant in C. javanus, an ecologically relevant species in the tropical soils of India.     

A female evolutionary response when survival is at risk: male harassment mediates early reallocation of resources to increase egg number and size

One unexplored area in sexual conflict studies is the female physiological costs and possible resource reallocation that accompany evolutionary costs due to male harassment. Using females of the damselfly Hetaerina americana, I first investigated whether male harassment affected female mating rate and survival and explored whether such effects induced a resource allocation from immunity (in the form of phenoloxidase activity) and muscular fat reserves to egg number and size. Using two seasons that differed in male harassment, it was found that the higher the male harassment, the fewer are the female matings and the lower is the female survival. These results were corroborated using an experimental approach in which a situation of high male harassment was induced. It was also found that when the first mating takes place and at high male harassment, females had more reduced phenoloxidase activity and fat reserves and tended to lay most of the eggs they produce in their lifetime and these were considerably large. However, at low male harassment, egg number and size were more equally produced across matings. Females under high male harassment seemed to suffer the survival costs but may show a plastic evolutionary response of reallocating resources to egg traits to maximize fitness.     

Interference by wood ant influences size selection and retrieval rate of prey by Formica fusca

Summary Dominance relations between the territorial wood ant Formica polyctena and the submissive F. fusca generate testable predictions on the mechanisms of their coexistence. Here I tested the influence of interference competition by the dominant F. polyctena on the foraging of F. fusca. In the presence of F. polyctena, the activity and the retrieval rate of items of F. fusca decreased significantly. When F. fusca were given a choice between small chironomids and flies a hundred times heavier they selected nearly always flies in the absence of wood ants; when disturbed F. fusca took proportionately more chironomids. In nature, irrespective of distance from the wood-ant mound the size distribution of potentially available food items was the same. F. fusca collected smaller items close to the wood-ant mound, where the density of interfering wood ants was high, than far from it. Also, F. polyctena carried larger protein items from the outskirts of its territory than from the center. The items of F. fusca were on average smaller than those of F. polyctena although the item-size overlap was substantial. When encountering a F. polyctena, those F. fusca workers carrying a fly always lost their booty to the dominant but always managed to bring the chironomids to the nest. F. fusca and F. polyctena were equally efficient in detecting single chironomids placed on the surface of ground, but the presence of either species decreased the discovery rate of the other. This implies mutual exploitation competition between the species for the locally most abundant protein resource. F. polyctena found single flies faster than F. fusca. In an earlier study I showed that close to the wood-ant mound the colony size and production of sexual offspring of F. fusca were reduced. I suggest that the suppressed colony success of F. fusca is attributable to the diminished size and decreased retrieval rate of prey items close to the woodant mound abounding with interfering wood-ant foragers.     

Tests of the harassment-reduction function and frequency-dependent maintenance of a female-specific color polymorphism in a damselfly

Color polymorphisms have provided classical examples of how frequency-dependent selection maintains genetic variation in natural populations. Here we tested for the first time, the hypothesized adaptive function of a female-specific color polymorphism in odonates to lower male harassment towards females generally. Under conditions controlling for sex ratio, population density and morph frequency, we also tested two major frequency-dependent selection hypotheses for the maintenance of the polymorphism. Using groups of captive Enallagma hageni, whose females are either green or a male-like blue, we varied morph frequency at two sex ratios. We quantified sexual harassment towards females by visual observations, and by the presence of dust on females that was transferred from dusted males. Per capita harassment rate for the female-monomorphic treatments did not differ from that of the female-polymorphic treatments. At a male-biased sex ratio, per capita harassment rate towards blue, but not green females increased with morph frequency, providing partial support for frequency-dependent selection resulting from male learning of female morphs. Even at high frequency, green females were not harassed more than blue, contrary to the prediction that males should always recognize green females as mates. Moreover, frequency-dependent harassment towards blue females was not detectable using harassment measured with dust evidence, which greatly underestimated the incidence of sexual harassment. Our findings identified problems with the use of insectaries and the dusting technique to quantify male sexual harassment towards females, as well as with a past insectary experiment on Ischnura elegans that failed to demonstrate frequency-dependent harassment.     

The cost of sexual coercion and heterospecific sexual harassment on the fecundity of a host-specific, seed-eating insect (Neacoryphus bicrucis)

Two species of seed-eating true bugs, Neacoryphus bicrucis (Heteroptera: Lygaeidae) and Margus obscurator (Heteroptera: Coreidae) co-occur on ragwort, Senecio tomentosus, in southern Georgia, USA. Males of both species sometimes engage in chases and protracted grappling with females that flee initial mountings. Sometimes genital coupling occurs while the wriggling female is restrained in the male's grasp. Chases, grappling, and mounting attempts are misdirected toward heterospecific females, heterospecific males, or conspecific males. In a laboratory study, confinement of mated N. bicrucis females with either conspecific or heterospecific males reduced fecundity by approximately one-half relative to mated females confined only with other females. Perhaps as a consequence of this, N. bicrucis females frequently leave areas of high host plant density, where they prefer to oviposit, when males are abundant. The abundance of each species is positively correlated with host plant density but the two species rarely occur together on the same plants. This may be an effect of heterospecific courtship which induces the flight of N. bicrucis more than the flight of M. obscurator. The laboratory results suggest that copulations following chases and grappling represent sexual harassment, not a mechanism of active female choice for a vigorous mate. As sexual harassment imposes high fitness costs that favor abandonment of host plants, it may, when misdirected, incidentally limit habitat use by ecologically similar species. Received: 3 October 1998 / Received in revised form: 21 January 1999 / Accepted: 14 February 1999