The cause of size-assortative mating in the leaf beetle Trirhabda canadensis (Coleoptera : Chrysomelidae)

Summary Examples of positive assortative mating by body size are abundant but its causes remain controversial. I show that size-assortative mating occurs in the chrysomelid beetle Trirhabda canadensis and I test a series of alternative hypotheses to explain how this mating pattern comes about. Results suggest that assortative mating in this beetle is due to the greater ease with which size-matched pairs can achieve intromission, and not due to size-biased skews in the availability of mates or mate choice favoring large individuals. There was no correlation between male and female elytron length (a measure of body size) at the initiation of courtship, but pairs assorted positively by size at the onset of intromission. Moreover, in the laboratory, there was a negative correlation between male and female size for pairs engaged in courtship that terminated without mating. Assortative mating was not associated with a large-male mating advantage and there was no evidence of female choice of large males. Nor was there unequivocal evidence for male choice of large females; although mating females were slightly larger and considerably heavier than solitary females, males did not differ in the frequency with which they rejected large and small females. Assortative mating in T. canadensis appeared to be caused by the lower ability of mismatched pairs to achieve intromission after an encounter, both when males were larger and when they were smaller than the female.     

The effect of multiple mating on genetic relatedness in larval aggregations of the imported willow leaf beetle (Plagiodera versicolora,Coleoptera: Chrysomelidae)

Summary The effect of multiple mating on the average genetic relatedness within socially interacting sib groups was studied in the imported willow leaf beetle. Using an electrophoretically determined genetic marker, it was demonstrated that between 49 and 96% of wild caught females produce egg clutches expressing multiple sires. Following theoretical predictions of Wade (1982), this information was used to estimate the harmonic mean number of sires per family and then the average relatedness within families. Average relatedness was estimated to be 0.38. An alternate form of data analysis suggested by Pamilo and Crozier (1982) did not require any knowledge of paternity and resulted in a slightly higher estimate of relatedness, 0.44.     

Prolonged mating in the milkweed leaf beetle Labidomera clivicollis clivicollis (Coleoptera: Chrysomelidae): a test of the “sperm-loading” hypothesis

Summary Female milkweed leaf beetles (Labidomera clivicollis clivicollis) frequently mate with more than one male, and pairs form mating associations which last for up to 42 h in the field. I tested the hypothesis that males remaining with females for long periods of time benefit by numerically overwhelming the sperm of their competitors. Male L.c.clivicollis copulated intermittently with females throughout an 11 hour period in the laboratory. When virgin females were allowed a single copulation, 94.3% of the sperm they received were located in the spermatheca immediately afterward. Males were not sperm-depleted, for they had large numbers of sperm available after one copulation (mean=230,000±43,200); the maximal number of sperm a male transferred to a female in 24 h was 30,500. There was a positive linear relationship between the number of sperm transferred and time up to 24 h after mounting (r ²=0.178, P<0.003). These data suggest that males transfer increasing numbers of sperm throughout a 24-h-period. Mating duration was the most important determinant of paternity when females were placed with one male for 24 h and another male for 6 hours. Females whose first matings were longer showed first male sperm predominance (as determined by starch-gel electrophoresis), while females whose second matings were longer showed last male sperm predominance. In view of these data, it is puzzling that males do not inseminate with large numbers of sperm immediately after mounting the female. It is possible that female refractory behaviors make insemination difficult and favor prolonged mating by male milkweed leaf beetles.     

Sperm competition in the damselfly Enallagma hageni Walsh (Odonata: Coenagrionidae): benefits of multiple mating to males and females

Summary Sperm competition was investigated in the non-territorial damselfly, Enallagma hageni. Using irradiated (sterile) male techniques, the last male to mate was found to fertilize up to 95% of the eggs of the first clutch laid after mating. Dissection of females collected before, during, and after copula showed that a male removes a maximum of 87% of the sperm by volume of a previous mate. These data verify an earlier estimate of lifetime reproductive success in this species which was based on mating success, and suggest that indirect dissection methods offer minimum estimates of sperm precedence. Male E. hageni have ample opportunity to benefit from sperm precedence, since at least 10% of the receptive females encountered had already mated once that day, but still contained complete or partial clutches of eggs. Female E. hageni benefit directly from high sperm precedence because it allows them to exchange matings for guarding service by males during oviposition bouts under water.     

The benefits of multiple mating to female seaweed flies, Coelopa frigida (Diptera: Coelpidae)

Female seaweed flies, Coelopa frigida, have the potential to benefit from mating more than once. Single matings result in low fertility so females may benefit directly from multiple copulations by sperm replenishment. A chromosomal inversion associated with larval fitness, with heterokaryotypic larvae having higher viability than homokaryotypes, means that polyandrous homokaryotypic females have a higher probability of producing genetically fit offspring than monandrous homokaryotypic females. We allowed females to mate only once, repeatedly four times to the same male, or polyandrously four times to four different males. Multiply mated and polyandrous females laid more eggs and produced more offspring than singly mated and monandrous females, respectively. Polyandrous females laid more eggs, had higher egg-to-adult survival rates and produced more offspring than repeatedly mated females. Fertility rates did not differ between treatments. The observed fitness patterns therefore resulted from increased oviposition through multiple mating per se, and a further increase in oviposition coupled with higher egg-to-adult offspring survival benefits to polyandry. Daily monitoring of individual females over their entire life spans showed that multiple copulations induced early oviposition, with polyandrous females ovipositing earlier than repeatedly mated females. Singly mated and polyandrous females incurred a longevity cost independent of egg production, whereas repeatedly mated females did not. This suggests that repeatedly mating with the same male may counteract a general cost of mating. Longevity, however, was not correlated with overall female fitness. Our data are discussed in the overall context of the seaweed fly mating system.Communicated by G. Wilkinson     

Volatile emissions from the flea beetle Altica litigata (Coleoptera: Chrysomelidae) associated with invasive Ludwigia hexapetala

The flea beetle Altica litigata (Chrysomelidae) is an insect herbivore to plants within the families Lythraceae and Onagraceae, including ornamentals such as crape myrtle, Lagerstroemia spp. This insect is important both as a pest species and as a naturally occurring biological control agent due to its aggregate feeding behavior, which typically results in severe defoliation of the host plant. Despite the negative economic impact to ornamentals and contrary benefits as a biological control agent, there are few reports on the semiochemical communication of this family of insects. Uruguayan primrose-willow (Ludwigia hexapetala) is an invasive aquatic weed in California and serves as a host to A. litigata. To better characterize this association, the volatile emissions of A. litigata were collected while the flea beetles were: in containers by themselves, in containers with L. hexapetala leaves, in situ on L. hexapetala leaves in a growth chamber, and in situ on L. hexapetala leaves in the field. For comparison, the volatile emissions of A. litigata associated with two subspecies of creeping water primrose (L. peploides) were also evaluated. Two himachalene-type sesquiterpenes, showing the same carbon skeleton as compounds previously reported from Aphthona flava and Epitrix fuscula, were detected as volatiles from A. litigata.     

Alternative mating tactics in males of Polistes dominulus (Hymenoptera: Vespidae)

Summary In summer, males of Polistes dominulus form large aggregations at sunny landmarks. We identified two size-correlated behavioural categories: residents (R) and transient (T). R males, which constitute 20%–25% of the total population, are larger than T males, territorial, aggressive, and more site-faithful, while T males range more widely, are non-aggressive, and show little site tenacity. Field and laboratory data suggest that R males have an advantage in mating, particularly if they engage in frequent flights while on their territories. These alternative mating tactics within the same population are combined with behavioural flexibility in some individuals, which switch from one option to the other.     

Fitness benefits of multiple mating versus female mate choice in the cellar spider (Pholcus phalangioides)

Recent studies have demonstrated that mating with multiple males can be beneficial for females and her offspring even if males contribute nothing but sperm. This was mainly established for species in which sperm from several males mix in the reproductive tract of the female, thus allowing sperm competition and/or female sperm choice. However, in species with last male sperm precedence, female re-mating decides against the previous male by strongly limiting his reproductive success. We tested the effect of female re-mating behaviour using the cellar spider Pholcus phalangioides, which shows strong last males sperm precedence and moderate levels of polyandry under natural situations. We predicted that females prevented from remating even though they are receptive would show reduced reproductive success compared to females that accept two copulations and females that reject a second male, since the latter two treatments were allowed to behave according to their decisions. However, if the number of matings per se had an effect on oviposition or on offspring performance, double-mated females should perform better compared to both treatments of once-mated females. We measured female fecundity and fertility over a period of 140 days, comparable to the species' natural reproductive peak season. Two thousand one hundred and fifty-two offspring from 67 first egg sacs were reared under two feeding levels. We registered development time and survival, and measured offspring adult size and mass. We found a positive effect of double mating, as in this treatment, oviposition probability was higher compared to the other treatments. Interestingly, adult female offspring of the DM treatment that were raised under low food level had a higher condition index compared to those from FS and RM, but development time, size and mass at adulthood were not affected by mating treatment. Female choice only seemed to affect hatching latency of the offspring. Overall, the main predictor of female reproductive output and success was female body size.     

Trade-off between fitness components in males of the polygynous butterfly Callophrys xami (Lycaenidae): the effect of multiple mating on longevity

The differential costs of mating paid by males and females influence the nature and strength of sexual selection. In butterflies, males invest a relatively large amount of time and resources in each mating, but male survival costs of mating have not been demonstrated. I present the results of experiments designed to measure the effect of different aspects of mating on male longevity in the polygynous butterfly Callophrys xami. In experiment 1, I compared the longevity of pairs of males that produced similar amounts of spermatophore, but that mated at different rates, a different numbers of times, and that produced spermatophores at different rates, and found that the longevity of ”low-mating-rate” males was not different from that of ”high-mating-rate” males. In experiment 2, the longevity of virgin males was not significantly different from that of multiply mated males. In experiment 3, I used resource-limited males resulting from experimental food limitation of last-instar larvae; resource-limited virgin males lived significantly more days than resource-limited multiply mated males. Since ecological costs of mating (e.g., disease transmission, predation risk) were excluded in the experiment, diminished male longevity was a product of physiological costs of sexual interactions. These results suggest that the cost of ejaculate production is an important cause of longevity reduction when there are resource limitations; however, the role of other possible physiological costs of mating in longevity reduction is still unknown. Received: 21 March 2000 / Accepted: 26 August 2000     

Behavior and ecology of Old World Luperini beetles of the genusAulacophora (Coleoptera: Chrysomelidae)

Summary The attraction of Old World leaf beetles in the genusAulacophora to kairomones and parakairomones which are effective lures for New WorldDiabrotica andAcalymma were investigated. Beetles captured on sticky traps baited with single and multicomponent lures were no different from the control traps for two species ofAulacophora. Yellow colored traps and squash blossoms are attractive toAulacophora beetles which detect sub-microgram quantities of cucurbitacins on silica gel. Leaf feeding behavior and flight activity data are correlated with varietal preference of threeAulacophora species. The common response byDiabrotica andAulacophora to cucurbitacins reinforces the two groups' coevolutionary association with the Cucurbitaceae. The apparent lack of a common response toCucurbita blossom volatiles suggests recent evolutionary pathways are substantially different for these two groups of beetles.     

Male mating success and paternal care in Nicrophorus vespilloides (Coleoptera: Silphidae)

Summary If no female is present, male burying beetles Nicrophorus vespilloides (Coleoptera: Silphidae) co-operate in the burial of a corpse. Once a female has arrived, the males fight with one another. The defeated male stays near the corpse and to copulate with the female. Laboratory experiments using sterilised males showed that the defeated male was able sometimes to father some of the offspring raised on the corpse. Male N. vespilloides almost always participate in defence and feeding of the brood. This is not affected by the size of the male. Males quickly leave or are driven from 5 g corpses. Males feed the larvae as often as females do, and larvae raised by males alone are not significantly different in weight from larvae raised by females alone or by both parents. Males which cared for a succession of broods in the laboratory did not differ significantly in median lifespan from males which were removed from their corpses after eggs had been laid. Non-caring males weighed significantly more than caring males over a sequence of corpses, but the caring males did not differ significantly in weight from non-breeding controls.     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

Quantitative analysis of pyrrolizidine alkaloids sequestered from diverse host plants in Longitarsus flea beetles (Coleoptera, Chrysomelidae)

Summary. Several species of the flea beetles genus Longitarsus are able to sequester pyrrolizidine alkaloids (PAs) from their host plants. In five Longitarsus species we compare the concentration of PAs present in their host plants belonging to the Asteraceae or Boraginaceae with those found in the beetles. To get an estimate of the intrapopulation variability, three samples of five beetles each and five individual plants were analyzed for each comparison. A strong intrapopulation variability could be detected both among plant and beetle samples. The total concentration found in the beetles varied strongly between species. The local host plant and its phenology influence the concentrations present in the beetles as evidenced in comparisons of a single beetle species from two different hosts and of one beetle species collected at the same site at different times of the year. In addition, different beetle species apparently vary in their capacity to sequester the alkaloids, at the lowest extreme the mean PA concentration in the beetles (0.034 μg PA/mg dry weight) was 1/30 of the mean concentration found in the plant leaves (L. aeruginosus from Eupatorium cannabinum), at the highest extreme (2.098 μg PA/mg dw) the concentration in the beetles was a 1000 fold higher than in the plant leaves (L. nasturtii from Symphytum officinale). The highest mean concentration found in the beetles was 3.446 μg/mg dw (L. exoletus from Cynoglossum officinale). The absolute concentrations found in the beetles are comparable to other insects which have been shown to be effectively defended against their potential predators. Received 22 June 1999; accepted 25 August 1999     

Differential attraction of females to dominant males: Olfactory discrimination and mating preference in the brown lemming (Lemmus trimucronatus)

Summary This study investigated differential attraction of estrous brown lemmings (Lemmus trimucronatus) to conspecific males recently exposed to each other for a 10-min agonistic encounter. In tests conducted 5 min, 1 h and 24 h after agonistic encounters, females preferred the oder of dominant males to that of defeated males when both odors were presented simultaneously in a Y-maze olfactometer. Defeat in an agonistic encounter did not reduce the propensity of male lemmings to initiate sexual behavior. In one-male, one-female tests conducted 5 min after agonistic encounters, dominant males achieved higher mount and thrust scores while defeated males obtained higher scores for attempted mounts. The sexual behavior of dominant and defeated males did not differ significantly in similar tests conducted 1 h and 24 h later. In contrast, females readily mated with dominant males and tended to avoid defeated males in two-male tethering tests conducted 5 min after agonistic encounters. In these tests, females still showed a preference for dominant males 1 h and 24 h after male agonistic encounters.     

Extrapair copulations in western bluebirds (Sialia mexicana): female receptivity favors older males

. Extrapair paternity has been the focus of numerous studies of socially monogamous birds, but the dynamics of extrapair male-female interactions have been explored in relatively few. In western bluebirds (Sialia mexicana), a species with 20% extrapair paternity, two to four observers monitored extrapair copulations (EPCs) occurring on the female's home territory while her social mate was detained in a cage near the nest. Detention elevated the frequency and visibility of EPCs, so we could score receptivity of females to identifiable extrapair males without the interference of intermale aggression. Most intruding extrapair males were neighboring breeders visiting outside their own mate's fertile period. Local male relatives of the detained male attempted nonincestuous EPCs, but neighboring sons did not attempt to copulate with their mothers and, in some cases, chased away intruders. Female receptivity was not influenced by whether the detained mate was visually present or hidden behind cloth and did not increase with the persistence or relative size of the extrapair male. Female receptivity increased with the age difference between the extrapair male and the female's social mate, suggesting that females prefer older males as extrapair partners. This study suggests that costs of resistance are relatively unimportant in determining female receptivity in western bluebirds, but identifies preferential receptivity to older males as a potential factor in the outcome of inter- and intrasexual conflict over paternity.     

Attraction of the potential biocontrol agent <Emphasis Type="Italic">Galerucella placida</Emphasis> (Coleoptera: Chrysomelidae) to the volatiles of <Emphasis Type="Italic">Polygonum orientale</Emphasis> (Polygonaceae) weed leaves

Galerucella placida Baly (Coleoptera: Chrysomelidae) is a potential biocontrol agent of the rice-field weed Polygonum orientale L. (Polygonaceae). The volatile organic compound (VOC) profiles from undamaged and mechanically damaged plants, and from plants 12- and 36-h following continuous feeding of female G. placida adults and 2nd instar larvae were identified and quantified by GC–MS and GC-FID analyses. Twenty-four and 21 compounds were identified in volatiles of undamaged and insect feeding plants, respectively; whereas 22 compounds were detected in volatiles of mechanically damaged plants. Decanal and 1-dodecanol were unique to undamaged plants, and linalool was detected in volatiles of undamaged and mechanically damaged plants, but not in volatiles of insect damaged plants. However, the beetles are not attracted by none of these volatile components, when tested individually in Y-shaped glass tube olfactometer bioassays. In all plants, methyl jasmonate was predominant. 1-Undecanol was the least amount in undamaged plants, and plants 12-h after feeding by G. placida adults and larvae; whereas 1-tridecanol was the least abundant in plants 36-h after feeding by G. placida adults and larvae, and mechanically damaged plants. The beetles showed significant preference to the whole volatile blends from plants 12-h after feeding by larvae and plants 36-h after feeding by either larvae or adults compared to those of undamaged plants. Further, G. placida responded to individual synthetic compounds, 3-hexanol, 1-octen-3-ol, nonanal, and geraniol at 7, 1.38, 3.75 and 4.5 µg/25 µL CH₂Cl₂, respectively, and provide a basis for attraction of the potential biocontrol agent in the field.     

Post-copulatory aggression toward their mates by males of the rove beetle Leistotrophus versicolor (Coleoptera: Staphylinidae)

Summary Rove beetles (Leistotrophus versicolor) forage and mate at dung and carrion in the riparian forest of northwestern Costa Rica. After copulating, males often launch a post-copulatory attack against their recent partner. We test four hypotheses on the adaptive value of male behavior: (1) The sperm competition hypothesis proposes that the behavior may be the functional equivalent of mate-guarding, (2) the sperm-transfer signal hypothesis states that males bite their mates after copulating to signal that they have successfully passed sperm, (3) the feeding competition hypothesis argues that male aggression toward mates occurs to drive away competitors for fly prey, and (4) the redirected aggression hypothesis is that male attacks after mating occur when threatened males redirect their aggression onto their partners. Only the sperm competition hypothesis withstands testing. As required by this hypothesis, females are usually receptive while at dung, and will mate with more than one male in a morning. In addition, males are more likely to attack a mate when they have fought earlier in the day with other males, an indicator of the presence of rival males and the risk of sperm competition. Contrary to the sperm-transfer signal hypothesis (2), biting of mates does not occur after nearly 40% of all copulations; it seems unlikely that mating males so often fail to transfer sperm. Whether males have fed or not prior to mating has no effect on the probability of post-copulatory attack, a result that contradicts the food competition hypothesis (3). Finally, the occurrence of attacks by males on females in the absence of an immediate threat from a rival argues against the redirected aggression hypothesis (4).     

Role of female territoriality in social and mating systems of Canthigaster valentini (Pisces: Tetraodontidae): evidence from field experiments

Sharpnose puffers, Canthigaster valentini (Pisces: Tetraodontidae) at Lizard Island, Australia, live in made-dominated haremic social and mating systems. The hypothesis was that mature females are restricted in their movements and can be monopolized by some males. Field experiments at Lizard Island, Great Barrier Reef, between January and March 1983 showed that mature females were still territorial in the absence of males and movements of females were not controlled by males. Males abandoned their territories when their females were removed. The territorial behavior of those males with access to females (territorial males) restricted the access of other males (bachelor males) to them. Bachelor males took over harems and became territorial males when established territorial males were removed. The results of the experiments thus supported the hypothesis.