Hans Van Gossum

Frequency-dependent male mate harassment and intra-specific variation in its avoidance by females of the damselfly Ischnura elegans

Abstract. We focused on male harassment on different female color morphs of the damselfly Ischnura elegans and on variation in morph-specific mating avoidance tactics by females. In I. elegans, one of the female morphs is colored like the conspecific male (andromorphs) while the other morphs are not (gynomorphs). Our first goal was to quantify morph-specific male mating attempts, hence male harassment, in populations with manipulated population parameters (densities, sex ratios, and proportion of andromorphs). Second, we examined the females perspective by looking for potential differences in morph-specific mating avoidance tactics and success of those tactics in a natural population. Differences in population conditions did influence the number of male mating attempts per morph. The less frequent female morph was always subject to fewer mating attempts, which contradicts earlier hypotheses on mimicry, but supports those that assume that males learn to recognize female morphs. Gynomorphs occupy less open habitat and often fly away when a male approaches, while andromorphs use more open habitat, do not fly large distances and directly face approaching males. Female morphs did not differ in the proportion of successful mating-avoidance attempts. Our results suggest that the maintenance of the color polymorphism is most probably the result of interactive selective forces depending on variation in all population conditions, instead of solely density- or frequency-dependent selection within populations.

Male choice for female colour morphs in Ischnura elegans (Odonata, Coenagrionidae): testing the hypotheses

The occurrence of different conspecific female colour morphs, with one of the morphs resembling the male, is supposed to have consequences for mate choice. There are two hypotheses linking mate choice and female colour polymorphism. First, males may mate predominantly with female morphs that differ from the male because they do not recognize androchrome females as females (male mimic hypothesis). Second, males may be more attracted to the most common morph in the population (habituation hypothesis). We tested these hypotheses in five populations of the same species, Ischnura elegans, with a range of androchrome frequencies. In each population we performed binary choice experiments in small cages. Males did not consistently prefer gynochrome females but mated predominantly with the most common morph in the population. Moreover, a reanalysis of the available damselfly data in the literature also supported the habituation hypothesis. Copyright 1999 The Association for the Study of Animal Behaviour.

Variation in female morph frequencies and mating frequencies: random, frequency-dependent harassment or male mimicry?

Female-limited colour polymorphisms occur in a variety of species, where often one female morph (androchrome) resembles the body coloration of the conspecific male, whereas the other (gynochrome) does not. We tested predictions of two frequency-dependent hypotheses that are commonly invoked to explain the maintenance of these polymorphisms for multiple populations of the damselfly Ischnura elegans: (1) that males prefer mating with the most abundant female morph (LMR) or (2) that androchromes are functional male mimics (MM). We also asked whether variation in social and abiotic environments account for interpopulation variation in female morph frequencies. Contrary to predictions of the LMR hypothesis, morph mating frequency was not lower than expected based on morph frequency when a morph was uncommon or higher when a morph was most common. In support of the MM hypothesis, androchrome mating frequency was lower than predicted based on population morph frequency. In addition, as predicted, androchrome mating frequency increased with increasing ratio of androchromes to males, but we could not disentangle whether this was a consequence of the changing mimic/model ratio or because of rising androchrome frequency. Although variation in female morph frequencies between populations was not random across our study area, this could not be explained by geographical variation in frequencies and densities of males and females. Androchrome frequencies were higher in populations to the north and west where ambient temperatures were lower. Abiotic conditions such as temperature may need to be considered for understanding the maintenance of female-limited polymorphisms.

Observations on rearing damselflies under laboratory conditions

Rearing damselflies under laboratory conditions is a promising means of solving a variety of biological questions. Therefore, in order to improve the success of future researchers we felt the need to indicate potential difficulties in carrying out rearing experiments. Laboratory crosses were obtained using virgin animals originating from natural populations in Belgium and Spain. Resulting offspring was maintained, under laboratory conditions, in small aquaria until emergence and in insectaries as adults. Our results show that keeping damselflies during their entire life cycle under artificial conditions can be very difficult. We suggest that future researchers should change water regularly, supply sufficient food, and rear animals at low density or even individually. Furthermore, suggestions are given on type of food, advisable laboratory conditions and female oviposition methodology.

Lethal and sublethal costs of autotomy and predator presence in damselfly larvae

Abstract We studied the costs of lamellae autotomy with respect to growth and survival of Lestes sponsa damselfly larvae in field experiments. We manipulated predation risk by Aeshna cyanea dragonfly larvae and lamellae status of L. sponsa larvae in field enclosures and compared differences in numbers, size and mass of survivors among treatments. In the absence of a free-ranging A. cyanea larva, about 29% of the L. sponsa larvae died. This was probably due to cannibalism. The presence of a free-ranging A. cyanea reduced larval survival by 68% compared to treatments in which it was absent or not permitted to forage on L. sponsa damselflies. Across all predator treatments, lamellae autotomy reduced survival by about 20%. The mean head width and mass of survivors was lower in the enclosures with a free-ranging A. cyanea compared to the other two predator treatments. This suggested that larvae grew less in the presence of a free-ranging predator, indicating that increased antipredator behaviours were more important in shaping growth responses than reduced population density. Mass, but not head width, of survivors was also reduced after autotomy. The fitness consequences of these effects for the adults may be pronounced. In general, these field data strongly suggest that lamellae autotomy affects population regulation of damselflies.

Reflectance spectra and mating patterns support intraspecific mimicry in the colour polymorphic damselfly Ischnura elegans

Coexistence of female colour morphs in animal populations is often considered the result of sexual conflict, where polymorphic females benefit from reduced male sexual harassment. Mate-searching males easily detect suitable partners when only one type of female is present, but become challenged when multiple female morphs coexist, which may result in frequency-dependent mate preferences. Intriguingly, in damselflies, one female morph often closely resembles the conspecific male in body coloration, which has lead to hypotheses regarding intra-specific male-mimicry. However, few studies have quantitatively evaluated the correspondence between colour reflectance spectra from males and male-like females, relying instead on qualitative visual assessments of coloration. Using colour analyses of reflectance spectra, we compared characteristics of the body coloration of ontogenetic male and female colour morphs of the damselfly Ischnura elegans. In addition, we evaluated whether males appear to (1) discriminate between immature and mature female colour morphs, and (2) whether male-like females experience reduced male mating attention and low mating frequencies as predicted from male-mimicry. Spectral reflectance data show that immature female morphs differ substantially in coloration from mature individuals. Mating frequencies were much lower for immature than mature female morphs. For the male-like female morph, measures of colour were statistically indistinguishable from that of both immature and mature conspecific males. Mating frequencies of male-like females were lower than those of other mature female morphs under field and experimental conditions. Together, our results indicate that males may use the observed spectral differences in mate choice decisions. Furthermore, male-like females may be regarded as functional mimics that have reduced attractiveness and lowered rates of sexual harassment by mate-searching males.

Negative frequency-dependent selection or alternative reproductive tactics: maintenance of female polymorphism in natural populations

BackgroundSex-limited polymorphisms have long intrigued evolutionary biologists and have been the subject of long-standing debates. The coexistence of multiple male and/or female morphs is widely believed to be maintained through negative frequency-dependent selection imposed by social interactions. However, remarkably few empirical studies have evaluated how social interactions, morph frequencies and fitness parameters relate to one another under natural conditions. Here, we test two hypotheses proposed to explain the maintenance of a female polymorphism in a species with extreme geographical variation in morph frequencies. We first elucidate how fecundity traits of the morphs vary in relation to the frequencies and densities of males and female morphs in multiple sites over multiple years. Second, we evaluate whether the two female morphs differ in resource allocation among fecundity traits, indicating alternative tactics to maximize reproductive output.ResultsWe present some of the first empirical evidence collected under natural conditions that egg number and clutch mass was higher in the rarer female morph. This morph-specific fecundity advantage gradually switched with the population morph frequency. Our results further indicate that all investigated fecundity traits are negatively affected by relative male density (i.e. operational sex ratio), which confirms male harassment as selective agent. Finally, we show a clear trade-off between qualitative (egg mass) and quantitative (egg number) fecundity traits. This trade-off, however, is not morph-specific.ConclusionOur reported frequency- and density-dependent fecundity patterns are consistent with the hypothesis that the polymorphism is driven by a conflict between sexes over optimal mating rate, with costly male sexual harassment driving negative frequency-dependent selection on morph fecundity.

Behaviour of damselfly larvae (Enallagma cyathigerum) (Insecta, Odonata) after long-term exposure to PFOS.

Perfluorooctane sulfonic acid (PFOS) is a persistent and ubiquitous environmental contaminant that has been detected in organisms worldwide. Here, we evaluate whether long-term (1 and 4 months) exposure to PFOS contamination affects the behavioural performance of freshwater larvae of the damselfly Enallagma cyathigerum (Insecta: Odonata). Our results show reduced behavioural performance with increasing PFOS concentration. In 1 month exposed larvae, no observed effect concentrations (NOECs) were 100 microg/L for general activity. In 4 months exposed larvae, NOECs were 10 microg/L, for each behavioural trait, except swimming acceleration of male larvae where the NOEC was 100 microg/L. When faced with PFOS concentrations above the NOEC, E. cyathigerum larvae were less active, less capable to escape a simulated predator attack and less efficient in foraging. Together, our results show that damselfly larvae suffer reduced survival-related behavioural performance.

Ontogenetic shifts in male mating preference and morph-specific polyandry in a female colour polymorphic insect

BackgroundSexual conflict over mating rates may favour the origin and maintenance of phenotypes with contrasting reproductive strategies. The damselfly Ischnura elegans is characterised by a female colour polymorphism that consists of one androchrome and two gynochrome female morphs. Previous studies have shown that the polymorphism is genetic and to a high extent maintained by negative frequency-dependent mating success that varies temporally and spatially. However, the role of learning in male mating preferences has received little attention. We used molecular markers to investigate differences in polyandry between female morphs. In addition, we experimentally investigated innate male mating preferences and experience-dependent shifts in male mating preferences for female morphs.ResultsField and molecular data show that androchrome females were less polyandrous than gynochrome females. Interestingly, we found that naïve males showed significantly higher sexual preferences to androchrome than to gynochrome females in experimental trials. In contrast, experienced males showed no preference for androchrome females.ConclusionsThe ontogenetic change in male mate preferences occurs most likely because of learned mate recognition after experience with females, which in this case does not result in a preference for one of the morphs, but rather in the loss of an innate preference for androchrome females.

Exposure to perfluorooctane sulfonic acid (PFOS) adversely affects the life-cycle of the damselfly Enallagma cyathigerum.

We evaluated whether life-time exposure to PFOS affects egg development, hatching, larval development, survival, metamorphosis and body mass of Enallagma cyathigerum (Insecta: Odonata). Eggs and larvae were exposed to five concentrations ranging from 0 to 10000 microg/L. Our results show reduced egg hatching success, slower larval development, greater larval mortality, and decreased metamorphosis success with increasing PFOS concentration. PFOS had no effect on egg developmental time and hatching or on mass of adults. Eggs were the least sensitive stage (NOEC=10000 microg/L). Larval NOEC values were 1000 times smaller (10 microg/L). Successful metamorphosis was the most sensitive response trait studied (NOEC<10 microg/L). The NOEC value suggests that E. cyathigerum is amongst the most sensitive freshwater organisms tested. NOEC for metamorphosis is less than 10-times greater than the ordinary reported environmental concentrations in freshwater, but is more than 200-times smaller than the greatest concentrations measured after accidental releases.