A trade-off between prey- and predator-induced polyphenisms in larvae of the salamander Hynobius retardatus

Organisms in natural habitats participate in complex ecological interactions that include competition, predation, and foraging. Under natural aquatic environmental conditions, amphibian larvae can simultaneously receive multiple signals from conspecifics, predators, and prey, implying that predator-induced morphological defenses can occur in prey and that prey-induced offensive morphological traits may develop in predators. Although multiple adaptive plasticity, such as inducible defenses and inducible offensive traits, can be expected to have not only ecological but also evolutionary implications, few empirical studies report on species having such plasticity. The broad-headed larval morph of Hynobius retardatus, which is induced by crowding with heterospecific anuran (Rana pirica) larvae, is a representative example of prey-induced polyphenism. The morph is one of two distinct morphs that have been identified in this species; the other is the typical morph. In this paper, we report that typical larval morphs of Hynobius can respond rapidly to a predatory environment and show conspicuous predator-induced plasticity of larval tail depth, but that broad-headed morphs cannot respond similarly to a predation threat. Our findings support the hypothesis that induction or maintenance of adaptive plasticity (e.g., predator-induced polyphenism) trades off against other adaptive plastic responses (e.g., prey-induced polyphenism). For a species to retain both an ability to forage for larger prey and an ability to more effectively resist predation makes sense in light of the range of environments that many salamander larvae experience in nature. Our results suggest that the salamander larvae clearly discriminate between cues from prey and those from predators and accurately respond to each cue; that is, they adjust their phenotype to the current environment.     

Factors which affect the occurrence of cannibalism and the broad-headed "cannibal" morph in larvae of the salamander Hynobius retardatus

Cannibalism in amphibian larvae may be suppressed among siblings in comparison to distant relatives or nonkin, even when a "cannibal" morph that can consume conspecifics shows adaptive advantages. Two experimental studies were undertaken to analyze factors which affect the frequency of cannibalism and the occurrence of the broad-headed "cannibal" morph in larvae of the salamander Hynobius retardatus. The first experiment investigated whether or not the broad-headed "cannibal" morph is only induced after performing cannibalism. Because the broad-headed "cannibal" morph was observed in larval populations that showed no cannibalism, it was concluded that the "cannibal" morph could be induced without actual cannibalism. Second, possible factors affecting the occurrence of the broad-headed "cannibal" morph were analyzed with respect to the density of larvae, level of food supply, and kinship among larvae, alone or in combination. Appearance of the broad-headed "cannibal" morph was affected by interactive effects of density2kinship: although the morph was induced significantly more often at a higher density if the conspecifics were distantly or not related, it was strongly suppressed even at high density if larval kinship was very close or among siblings. In contrast, the frequency of cannibalism was independently affected by larval density, level of food supply, and kinship among larvae: it was significantly larger at high than at low larval densities, at low than at high food supply, and in mixed-sibling groups than in pure sibships. These results suggest that the "cannibal" morphs in H. retardatus are induced even without cannibalism at a high density of conspecifics in mixed-sibling groups, but the occurrence of this morph, which continuously consumes conspecifics, is suppressed among siblings.     

Impact of cannibalism on predator-prey dynamics: size-structured interactions and apparent mutualism

Direct and indirect interactions between two prey species can strongly alter the dynamics of predator-prey systems. Most predators are cannibalistic, and as a consequence, even systems with only one predator and one prey include two prey types: conspecifics and heterospecifics. The effects of the complex direct and indirect interactions that emerge in such cannibalistic systems are still poorly understood. This study examined how the indirect interaction between conspecific and heterospecific prey affects cannibalism and predation rates and how the direct interactions between both species indirectly alter the effect of the cannibalistic predator. I tested for these effects using larvae of the stream salamanders Eurycea cirrigera (prey) and Pseudotriton ruber (cannibalistic predator) by manipulating the relative densities of the conspecific and heterospecific prey in the presence and absence of the predator in experimental streams. The rates of cannibalism and heterospecific predation were proportional to the respective densities and negatively correlated, indicating a positive indirect interaction between conspecific and heterospecific prey, similar to "apparent mutualism." Direct interactions between prey species did not alter the effect of the predator. Although both types of prey showed a similar 30% reduction in night activity and switch in microhabitat use in response to the presence of the predator, cannibalism rates were three times higher than heterospecific predation rates irrespective of the relative densities of the two types of prey. Cumulative predation risks differed even more due to the 48% lower growth rate of conspecific prey. Detailed laboratory experiments suggest that the 3:1 difference in cannibalism and predation rate was due to the higher efficiency of heterospecific prey in escaping immediate attacks. However, no difference was observed when the predator was a closely related salamander species, Gyrinophilus porphyriticus, indicating that this difference is species specific. This demonstrates that cannibalism can result in the coupling of predator and prey mortality rates that strongly determines the dynamics of predator-prey systems.     

Interspecific aggression in Leucorrhinia dragonflies: a frequency-dependent discrimination threshold hypothesis

Summary Male Leucorrhinia dragonflies defend territories from conspecific and heterospecific intruders. Defense against heterospecifics is surprising, as mating and oviposition are the only activities that occur on the territories, and heterospecific males are not expected to pose a reproductive threat. L. frigida and L. intacta males respond aggressively with equal frequency and intensity against intrusions by conspecifics and heterospecifics. In contrast, L. proxima males respond more aggressively against conspecifics. The apparent lack of species discrimination shown by L. frigida and L. intacta males may result because territorial males that assess intruders (as do L. proxima) suffer a tactical disadvantage from hesitating when an intruder flies in. This assessment process may lead to reduced fighting success by the territorial male. Thus there is a tradeoff between assessment cost incurred when the intruder is a conspecific and benefit gained from avoiding conflict when the intruder is a heterospecific. Given this assessment cost, males of species that only rarely encounter conspecific intruders are more likely to evolve species discrimination than males of species that commonly encounter conspecific intruders.     

Can male-male competition stabilize speciation? A test in Lake Victoria haplochromine cichlid fish

It has been suggested that sympatric speciation can be driven by sexual selection on male mating traits alone. However, a fundamental problem for this process is the lack of ecological differentiation that would stabilize the coexistence of incipient species through frequency-dependent selection. Such selection can also occur if male aggression is primarily directed towards similar rather than towards dissimilar phenotypes, so that rare male phenotypes would enjoy a negatively frequency-dependent fitness advantage. We experimentally tested such an aggression bias in two recently diverged, ecologically and anatomically similar sympatric cichlid species pairs of the genus Pundamilia from Lake Victoria. Territorial males of a pair of partially reproductively isolated species with red and blue nuptial coloration, respectively, studied in the laboratory were confronted simultaneously with both colour types enclosed in transparent tubes. Red males were more aggressive to red stimuli under white light but not when colour differences were masked under green light. Blue males were equally aggressive to both stimuli in both light conditions. Males of two apparently fully reproductively isolated species, again one with red and one with blue nuptial coloration, studied in the field, both directed more aggressive behaviour towards conspecific than towards heterospecific stimulus males. The differential allocation of aggression would create an advantage for males of the less abundant phenotype or species, thereby potentially supporting stable coexistence of the phenotypes. The finding that this effect was less clear in the partially reproductively isolated species pair than in the fully isolated species pair is discussed.     

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.     

The effects of three sympatric ladybird species on oviposition by Menochilus sexmaculatus (Coleoptera: Coccinellidae)

Oviposition site selection has a strong impact on the spatial distribution and sustenance of a species. This site selection by ladybirds (Coleoptera: Coccinellidae) is known to be well mediated amongst other factors by the oviposition deterring pheromones present in conspecific larval tracks. It would be also useful to the organism to be sensitive to heterospecific larval tracks, to protect from intraguild predation; however, this has not yet garnered much attention and there are only a few scattered studies with ambiguous results. The present study thus investigates the oviposition deterring effects of heterospecific presence and semiochemicals of heterospecific life stages (eggs, fourth instar larvae, pupae, adult males and females) of ladybird species, viz. Coccinella septempunctata, C. transversalis, Propylea dissecta on females of Menochilus sexmaculatus. Results reveal that the presence of all heterospecific life stages inhibits oviposition of M. sexmaculatus and the inhibition is density dependent. Semiochemicals of eggs, larval and adult tracks also show density-dependent inhibitory effects. The semiochemical-induced oviposition inhibition was stage specific with respect to the degree of inhibition. Of all heterospecific species tested, maximum and minimum oviposition deterrence of M. sexmaculatus was against C. septempunctata and P. dissecta, respectively. Comparison of our previously published results on oviposition inhibition in M. sexmaculatus due to conspecific presence with the current ones revealed increased inhibition in conspecific presence than heterospecific indicating ability to differentiate.     

Use of chemical cues in detection of conspecific predators and prey by newts, <Emphasis Type="Italic">Notophthalmus viridescens</Emphasis>

Summary. Detection of heterospecific predators and prey via chemical cues is well known, but only a few studies have examined the potential for such discrimination in cannibalistic systems. In newts, Notophthalmus viridescens, adults are opportunistic predators of conspecific larvae. I used a laboratory bioassay to determine whether larvae and adults distinguish between chemical stimuli from members of the different age classes. Larvae distinguished between chemical stimuli from larvae and adults by decreasing their activity only following exposure to stimuli from adults. Decreased activity is consistent with an antipredator response in many prey species, including larval newts. In contrast, adults increased their activity and increased time spent in open areas in response to stimuli from larvae, but not to stimuli from adults. Increased activity is consistent with a feeding response; adults also showed increased activity and increased time in open areas in response to chemical stimuli from familiar heterospecific prey (brine shrimp). The proximate cue that allowed the newts to distinguish between the different age classes is not associated with short-term dietary differences because all stimulus animals were fed the same prey. Stimulus strength was controlled by diluting the stimulus solutions according to the volume of the stimulus animal. Therefore, there appear to be intrinsic differences in the chemical signatures of larval and adult newts.     

Effect of crowding on larval viability in Tisbe holothuriae (Copepoda: Harpacticoida)

Larval Tisbe holothuriae Humes exposed to anionic detergent (0.5 mg l^-1 sodium dodecyl benzene sulphonate) were found to survive better than control larvae. The detergent appears to enhance viability by relaxing inhibitory interactions between nauplii. This conclusion is inferred from the following experimental observations: (1) crowding depresses larval viability; (2) the effect of crowding on viability is indirect, and results from behavioural changes which reduce contact with food; (3) detergent exposure delays or suppresses the behavioural response to crowding. The nature of the interactions which modify behaviour is unknown, but evidence suggests that close encounters between nauplii are important, and that they may lead to the impairment of distance-chemoreception. Naupliar antagonism could have evolved as a mechanism for staggering larval development or as a means of reducing the relative fitness of neighbouring individuals.     

Assessment of patch quality by ladybirds: relative response to conspecific and heterospecific larval tracks a consequence of habitat similarity?

Summary. Aphid colonies can reach high levels of abundance but last for short periods of time. The larvae of aphidophagous ladybirds (Coleoptera: Coccinellidae) that feed on these colonies might therefore suffer from starvation, which favours the occurrence of cannibalism and intraguild predation. Thus, the assessment of patch quality becomes crucial and it has been shown that female ladybirds refrain from laying eggs in the presence of an oviposition deterring semiochemical deposited by their larvae. Adalia bipunctata (L.), Adalia decempunctata (L.) and Coccinella septempunctata L. are 3 sympatric species of ladybirds, which can co-occur in aphid colonies. As a consequence, their eggs and larvae are under threat, not only from cannibalism but also intraguild predation. Females should, therefore, also use the tracks deposited by heterospecific larvae to assess the quality of aphid colonies as oviposition sites. The expectation is that: 1- the strength of the reaction to each other’s larval tracks should be correlated with percentage habitat overlap and that 2- the reaction to conspecific larval tracks should be stronger than to heterospecific tracks. In order to test these hypotheses, females’ oviposition behaviour was analysed and a chemical analysis of the tracks of their larvae undertaken. The results show that oviposition behaviour is not related to habitat overlap. Both species of Adalia react to tracks of their own larvae and those of C. septempunctata, but A. decempunctata reacted more strongly than A. bipunctata. C. septempunctata reacted very slightly to its own tracks but not to those of either species of Adalia. The larval tracks are mainly composed of alkanes. Those of the two species of Adalia are qualitatively 100% similar and 60% so when the quantitative results are compared. They are, however, only 24–29% similar to those of Coccinella septempunctata.     

The morphology and behavior of dimorphic males in Perdita portalis (Hymenoptera : Andrenidae)

Summary In Perdita portalis, a ground nesting, communal bee, males are clearly dimorphic. The two male morphs are easily distinguished based on head size and shape into (1) a flight-capable, small-headed (SH) morph that resembles the males of other closely related species and (2) a flightless, large-headed (LH) morph that possesses numerous derived traits, such as reduced compound eyes, enlarged facial foveae and fully atrophied indirect flight muscles. The SH morph occurs exclusively on flowers while the LH morph is found only in nests with females. While on flowers, SH males are aggressive, fighting with conspecific males and heterospecific male and female bees, and they mate frequently with foraging females. Using artificial observation nests placed in the field, I observed the behavior of females and LH males within their subterranean nests. LH males are aggressive fighters; males attacked each other with mandibles agape, and male-male fights always ended in the death of one male. LH males are highly attentive to the reproductive behavior of females; they spend increasing amounts of time near open cells during cell provisioning, and mating only takes place immediately prior to oviposition when females are forming the accumulated pollen and nectar into a ball. Based on larvae reared to adulthood in the laboratory, the two male morphs occur in equal proportions. The behavior of males in closely related species, especially P. texana, and the origin and maintenance of male dimorphism are discussed.     

Adult female hamsters avoid interspecific mating after exposure to heterospecific males

When females mate with a heterospecific male, they do not usually produce viable offspring. Thus, there is a selective pressure for females to avoid interspecific mating. In many species, females innately avoid heterospecific males; females can also imprint on their parents to avoid later sexual interactions with heterospecific males. However, it was previously unknown whether adult females can learn to discriminate against heterospecific males. We tested the hypothesis that adult females previously unable to avoid interspecific mating learn to avoid such mating after being exposed to heterospecific males. Syrian hamster (Mesocricetus auratus) females not previously exposed to Turkish hamster (Mesocricetus brandti) males can discriminate between odors of conspecific and heterospecific males, but they mate with either type of male. However, when we exposed adult females to both a conspecific male and a heterospecific male through wire-mesh barriers for 8 days, and then paired them sequentially with the two males, females were more receptive to conspecific males and more aggressive to heterospecific males. When females were paired with the heterospecific male first and the conspecific male second, no female was receptive and all were aggressive to heterospecific males. When females were paired with the conspecific male first, only 43% of females were then aggressive toward the heterospecific male. That is, interactions with conspecific males may decrease a female’s ability to properly avoid heterospecific males. Our study clearly shows for the first time that females can learn during adulthood to avoid interspecific mating just by being exposed to stimuli from heterospecific males.     

Fluctuating asymmetry,interspecific aggression and male mating tactics in two species of Japanese scorpionflies

Summary Fluctuating asymmetry (FA) is defined as small, random deviations from bilateral symmetry in a generally perfect bilaterally symmetrical morphological trait. FA in forewing length affects the outcomes of interspecific contests for food items (dead arthropods) between two species of Japanese scorpionflies, Panorpa nipponensis and P. ochraceopennis, in nature. FA differences between same-sex contestants are more important than either body size or ownership of food in determining the outcomes of interspecific contests; for both sexes, winners statistically significantly more often have relatively low FA. Two condition-dependent mating tactics are used by the males of each species: (a) a male may defend a dead-arthropod nuptial gift, or (b) a male without such an arthropod may wait near a male with one. In both tactics, males release long-distance sex pheromones. Groups of pheromone-releasing males are made up of one male with a nuptial gift and his satellites; the males in a group may be conspecifics or heterospecifics. Males that lose contests for nuptial gifts often become satellites of the contest winners whether or not winners are conspecific. Satellite males have statistically significantly greater FA than males with nuptial gifts in heterospecific male display groups. Satellite males mate infrequently and briefly compared to resource-holding males. Satellites of heterospecific males copulate with conspecific females displaced from nuptial gifts by the resource-holding males of the other species. In both species, the largest and smallest individuals have the greatest FA, and intermediate-sized individuals have the least; this same pattern often occurs in other animals.     

Species recognition and species isolation in wandering spiders (Cupiennius spp.; Ctenidae)

Summary One of the roles ascribed to spider courtship is that of mate recognition and reproductive isolation. We describe behavioral mechanisms underlying these functions in three species of wandering spiders of the genus Cupiennius (Ctenidae). C. salei and the two sympatric species C. getazi and C. coccineus discriminate conspecifics and heterospecifics in a multistage process which covers three principal phases of courtship: (1) Chemical phase: male spiders initiate vibratory courtship communication less often upon contact with heterospecific than with conspecific female silk. (2) Vibratory phase: females respond with their own vibration less often and with increased delay to the vibratory courtship signals of heterospecific males than to those of conspecific males. (3) Tactile phase: in heterospecific pairings males and females interrupt the contact phase if their courtship has progressed thus far. The product of the probabilities of passing through each of the different stages of courtship is between 0.44 and 0.88 for conspecific pairings, but zero for most heterospecific pairings. This is to a greater extent due to the behavioral barriers of the vibratory and the tactile phase than to that of the chemical phase. Male taxis on tracks of female silk does not depend on the species-specificity of the silk. Female vibrations are not essential for species recognition: males increase their signal rates, orient toward and approach responding females in both con- and heterospecific pairings. Female attacks on males are similar to attacks on prey; males defend themselves with outstretched front legs and are caught and killed by the female in only 5% of the cases. In all phases of courtship, female C. salei are least and female C. getazi most selective. When female C. coccineus and C. getazi are involved in heterospecific pairings no copulation is attempted. In contrast, 13 copulations of C. salei with males of the other two species were observed; egg sacs never resulted from these copulations. Among males, those of C. getazi were least selective. The likelihood of heterospecific pairs passing through all phases of courtship is higher in case of the allopatric than in that of the sympatric species. This finding applies to the courtship as a whole, but not necessarily to each of its phases.Offprint requests to: F.G. Barth     

Territory characteristics and coexistence with heterospecifics in the Dartford warbler <Emphasis Type="Italic">Sylvia undata</Emphasis> across a habitat gradient

The study of successional gradients may help to understand the relative influence of habitat structure and competition on territory characteristics. Here, we evaluate the effects of vegetation cover, conspecific and heterospecific densities, and distance to the nearest neighbor on territory size, shape, and overlap in insectivorous birds. We studied these effects along a gradient of postfire habitat regeneration in which foliage cover and densities of focal species varied several-fold. We delineate 197 territories (minimum convex polygons) of the shrub-dwelling Dartford warbler (Sylvia undata) and 255 of the syntopic Sardinian (Sylvia melanocephala), subalpine (Sylvia cantillans) and melodious (Hippolais polyglotta) warblers at three plots in NE Catalonia (Spain and France) in 1987–2005. After accounting for the effect of the number of locations used to delineate polygons, generalized linear mixed models (GLMM) showed a reduction in territory area of the Dartford warbler as conspecific density increased and distance to nearest neighbor decreased, in accordance with the contender pressure hypothesis for territory size regulation. Heterospecific density was not included in the final model of territory size and the effect of habitat structure was marginal. Territory roundness was positively correlated with its size and with conspecific density, probably in relation to energetic constraints, and negatively with heterospecific density. Territorial exclusion was almost complete among Dartford warblers, whereas interspecific territory overlap was extensive and tended to increase with heterospecific density and with structural diversity along the gradient. Our results support the hypothesis that Mediterranean warbler coexistence derives from ecological segregation and not from interspecific territoriality.     

No cultural transmission of species recognition between parents and offspring in free-living great tits and blue tits

Imprinting plays a key role in the development of species recognition, with young imprinting upon the morphological characters of their parents. However, the potential role that cultural transmission might play in species recognition remains largely uninvestigated. Great tits (Parus major) and blue tits (Cyanistes caeruleus) do not normally perceive each other as sexual competitors or potential partners. However, after reciprocal interspecific cross-fostering, both species may perceive individuals of the foster species as potential rivals or mates. Although the experience of being raised by heterospecifics clearly has affected the species recognition of cross-fostered birds, some of them breed naturally with conspecifics. The offspring of such cross-fostered birds (OCF) are hence raised by parents that look like ordinary conspecifics but display deviant species recognition as compared to controls in terms of aggressive response towards rivals. Comparing the aggressive behavior of OCF, cross-fostered birds and controls towards territorial intruders may thus help tease apart the influence of morphological vs behavioral cues of parents in the development of offspring species recognition. To this end, we compared birds from all three treatments with respect to their aggressive response to territorial intruders of both species during the breeding season. OCF and controls did not differ in their pattern of response towards heterospecific and conspecific stimuli. Compared to cross-fostered birds, OCF and controls showed less aggression towards heterospecific intruders, while the response towards conspecific intruders did not differ between treatments. These results demonstrate that both tit species imprint on the morphological characters of their parents, but that parental behavior is not important for the development of species recognition in terms of aggressive response towards territorial intruders.     

Strange parental decisions: fathers of the dyeing poison frog deposit their tadpoles in pools occupied by large cannibals

Parents may increase the probability of offspring survival by choosing suitable rearing sites where risks are as low as possible. Predation and competition are major selective pressures influencing the evolution of rearing site selection. Poison frogs look after their clutches and deposit the newly hatched tadpoles in bodies of water where they remain until metamorphosis. In some species, cannibalism occurs, so parents deposit their tadpoles singly in very small pools. However, cannibalism also occurs in species that deposit tadpoles in larger pools already occupied by heterospecific or conspecific larvae that could be either potential predators or competitors. Here, I test the hypothesis that, given the choice, males of Dendrobates tinctorius would deposit their newly hatched tadpoles in low-risk sites for their offspring. I characterised the pools used by D. tinctorius for tadpole deposition, conducted experiments to determine the larval traits that predict the occurrence of and latency to cannibalism, and tested whether parents deposit their tadpoles in low-risk pools. I found that (1) neither pool capacity nor the presence of other larvae predict the presence/absence or number of tadpoles; (2) cannibalism occurs often, and how quickly it occurs depends on the difference in size between the tadpoles involved; and (3) the likelihood of males depositing their tadpoles in occupied pools increases with the size of the resident tadpole. I suggest that predation/cannibalism is not the only factor that parents assess when choosing deposition sites, and that the presence of larger conspecifics may instead provide information about pool quality and stability.     

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     

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

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 female's 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.     

Female mate choice determines reproductive isolation between sympatric butterflies

Animal courtship rituals are important for species recognition, and a variety of cues might be utilized to recognize conspecific mates. In this paper, we investigate different species-recognition mechanisms between two sympatric butterfly sister species: the wood white (Leptidea sinapis) and Real’s wood white (Leptidea reali). We show that males of both species frequently court heterospecific females both under laboratory and field conditions. The long-lasting elaborate courtships impose energetic costs, since the second courtship of males that were introduced to two subsequent conspecific females lasted on average only one fourth as long as the first courtship. In this paper, we demonstrate that premating reproductive isolation is dependent on female unwillingness to accept heterospecific mates. We studied female and male courtship behavior, chemical signaling, and the morphology of the sexually dimorphic antennae, one of the few male traits visible for females during courtship. We found no differences in ultraviolet (UV) reflectance and only small differences in longer wavelengths and brightness, significant between-species differences, but strongly overlapping distributions of male L. sinapis and L. reali antennal morphology and chemical signals and minor differences in courtship behavior. The lack of clear-cut between-species differences further explains the lack of male species recognition, and the overall similarity might have caused the long-lasting elaborate courtships, if females need prolonged male courtships to distinguish between con- and heterospecific suitors.