Evidence for division of labor in the social caterpillar Eucheira socialis (Lepidoptera: Pieridae)

The caterpillars of Eucheira socialis westwoodi cooperatively spin and maintain a hollow silken nest and an elaborate network of silken foraging trails on their host plant, madrone (Arbutus spp.: Ericaceae). Nests typically contain several hundred larvae. Two populations are known to harbor a sex ratio distorter. The primary sex ratio in these two populations for four generations has been exceedingly male biased (64–79% male). Lepidoptera larvae are easily sexed using external morphology, allowing us to uniquely mark male and female larvae and to assemble larval groups of particular sex ratios. We report here the results of experiments on sex-specific larval behavior and physiology and the effect of colony sex ratio on individual behavior. We found that male larvae spent more time spinning silk on the nest and less time feeding than female larvae. Males were the first to emerge from the nest and the first to venture out along trails to feed. Male-biased nests had a significantly greater amount of silk deposited on their surfaces than female-biased nests. In the field, male-biased nests produced heavier male and female pupae than female-biased nests. Male and female larvae in 75% male nests became active earlier than males and females in other sex ratio treatments. Received: 11 September 1998 / Received in revised form: 24 February 1999 / Accepted: 27 March 1999     

Local mate competition in the solitary parasitoid wasp <Emphasis Type="Italic">Ooencyrtus kuvanae</Emphasis>

Local mate competition (LMC) occurs when brothers compete with each other for mating opportunities, resulting in selection for a female-biased sex ratio within local groups. If multiple females oviposit in the same patch, their sons compete for mating opportunities with non-brothers. Females, in the presence of other females, should thus produce relatively more sons. Sex ratio theory also predicts a more female-biased sex ratio when ovipositing females are genetically related, and sex-ratio responses to foundress size if it differentially affects fitness gains from sons versus daughters. The mating system of the parasitoid wasp Ooencyrtus kuvanae meets assumptions of LMC. Females insert a single egg into each accessible egg of gypsy moth, Lymantria dispar, host egg masses. Wasps complete development inside host eggs and emerge en masse, as sexually mature adults, resulting in intense competition among brothers. We tested the hypothesis that O. kuvanae exhibits LMC by manipulating the number of wasp foundresses on egg masses with identical numbers of eggs. As predicted by LMC theory, with increasing numbers of wasp foundresses on an egg mass, the proportions of emerging sons increased. In contrast, the presence of a sibling compared to a non-sibling female during oviposition, or the size of a female, did not affect the number or sex ratio of offspring produced. The O. kuvanae system differs from others in that larvae do not compete for local resources and thus do not distort the sex ratio in favor of sons. With no resource competition among O. kuvanae larvae, the sex ratio of emergent son and daughter wasps is due entirely to the sex allocation by ovipositing wasp foundresses on host egg masses.     

Origin of male-biased sex allocation in orphaned colonies of the termite, <Emphasis Type="Italic">Coptotermes lacteus</Emphasis>

In eusocial insects, sex allocation often constitutes a ground for intracolonial conflicts. This occurrence provides ideal opportunities to test kin-selection theory. A vast literature on this topic is available for social Hymenoptera, but the same field remains almost untouched in termites. A preeminent case is that of some species of Coptotermes, where the sex-allocation ratio in nymphs shifts from near equity to all-male when the primary reproductives are replaced by neotenics. To shed light on the developmental origin of this shift, we compared the sex ratio of the various castes and instars in primary- and neotenic-headed mature colonies of Coptotermes lacteus. The male-biased sex allocation in the latter type of colony results from two concurrent events: first, the sex ratio of the youngest instars (larvae) is male-biased by a 3:1 ratio; and second, all female larvae become workers, while a large fraction of the male larvae proceed to the nymphal and alate stages. Colony-founding experiments showed that inbreeding by itself cannot account for the male bias at hatching. We suggest that both genetic factors, due to the reproductive behaviour of neotenics, and environmental factors (colony condition and resource availability) may influence this process. Their exact nature and respective impact have not yet been clarified.     

Adult sex ratio affects divorce rate in the monogamous endoparasite Schistosoma mansoni

“Divorce” (mate switching) rate is known to vary largely both between and within socially monogamous species. Although the adult sex ratio can have an important influence on mating patterns, very few studies have investigated the influence of sex ratio on divorce rate in monogamous species, and even less so from an experimental point of view. In addition, most studies on the causes and consequences of divorce have been performed on vertebrate species, whereas data for invertebrate monogamous species remain scarce. Schistosoma mansoni is a monogamous endoparasite with a complex life cycle characterized by asexual reproduction in the intermediate host and sexual reproduction in the definitive host. In the wild, populations of S. mansoni inside their definitive hosts are characterized by a male-biased sex ratio. We studied the influence of experimentally varying the adult sex ratio on divorce rate in S mansoni, using controlled infections of hosts with clonal populations. The more male-biased the sex ratio was, the more the divorce rate increased, whereas no such effect was observed under a female-biased sex ratio. In this study and for the first time, we showed, by handling the sex ratio, that the divorce rate increases in adult male-biased sex ratio conditions in a monogamous species.     

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.     

Why are larvae of the social parasite wasp <Emphasis Type="Italic">Polistes sulcifer</Emphasis> not removed from the host nest?

A challenge for parasites is how to evade the sophisticated detection and rejection abilities of potential hosts. Many studies have shown how insect social parasites overcome host recognition systems and successfully enter host colonies. However, once a social parasite has successfully usurped an alien nest, its brood still face the challenge of avoiding host recognition. How immature stages of parasites fool the hosts has been little studied in social insects, though this has been deeply investigated in birds. We look at how larvae of the paper wasp obligate social parasite Polistes sulcifer fool their hosts. We focus on cuticular hydrocarbons (CHCs), which are keys for adult recognition, and use behavioral recognition assays. Parasite larvae might camouflage themselves either by underproducing CHCs (odorless hypothesis) or by acquiring a chemical profile that matches that of their hosts. GC/MS analyses show that parasite larvae do not have lower levels of CHCs and that their CHCs profile is similar to the host larval profile but shows a reduced colony specificity. Behavioral tests show that the hosts discriminate against alien conspecific larvae from different colonies but are more tolerant towards parasite larvae. Our results demonstrate that parasite larvae have evolved a host larval profile, which overcomes the host colony recognition system probably because of the lower proportion of branched compounds compared to host larvae. In some ways, this is a similar hypothesis to the odorless hypothesis, but it assumes that the parasite larvae are covered by a chemical blend that is not meaningful to the host.     

Intraspecific competition: the role of lags between attack and death in host-parasitoid interactions

Many natural enemies do not immediately kill their host, and the lag this creates between attack and host death results in mixed populations of uninfected and infected hosts. Both competition and parasitism are known to be major structuring forces in ecological communities; however, surprisingly little is known about how the competitive nature of infected hosts could affect the survival and dynamics of remaining uninfected host populations. Using a laboratory system comprising the Indian meal moth, Plodia interpunctella, and a solitary koinobiont parasitoid, Venturia canescens, we address this question by conducting replicated competition experiments between the unparasitized and parasitized classes of host larvae. For varying proportions of parasitized host larvae and competitor densities, we consider the effects of competition within (intraclass) and between (interclass) unparasitized and parasitized larvae on the survival, development time, and size of adult moths and parasitoid wasps. The greatest effects were on survival: increased competitor densities reduced survival of both parasitized and unparasitized larvae. However, unparasitized larvae survival, but not parasitized larvae survival, was reduced by increasing interclass competition. To our knowledge, this is the first experimental demonstration of the competitive superiority of parasitized over unparasitized hosts for limiting resources. We discuss possible mechanisms for this phenomenon, why it may have evolved, and its possible influence on the stability of host-parasite dynamics.     

Oviposition height increases parasitism success by the robber fly Mallophora ruficauda (Diptera: Asilidae)

For parasitoids, host finding is a central problem that has been solved through a variety of behavioural mechanisms. Among species in which females do not make direct contact with hosts, as is the case for many dipteran parasitoids, eggs must be laid in an appropriate part of the host habitat. The asilid fly Mallophora ruficauda lays eggs in clusters on tall vegetation. Upon eclosion, pollen-sized larvae fall and parasitize soil-dwelling scarab beetle larvae. We hypothesized that wind dissemination of M. ruficauda larvae is important in the host-finding process and that females lay eggs at heights that maximize parasitism of its concealed host. Through numerical and analytical models resembling those used to describe seed and pollen wind dispersal, we estimated an optimal oviposition height in the 1.25- to 1.50-m range above the ground. Our models take into account host distribution, plant availability and the range over which parasitic larvae search for hosts. Supporting our findings, we found that the results of the models match heights at which egg clusters of M. ruficauda are found in the field. Generally, work on facilitation of host finding using plants focuses on plants as indicators of host presence. We present a case where plants are used in a different way, as a means of offspring dispersal. For parasitoids that carry out host searching at immature stages rather than as adults, plants are part of a dissemination mechanism of larvae that, as with minute seeds, uses wind and a set of simple rules of physics to increase offspring success.     

Test of the adaptiveness of sex ratio manipulation in a parasitoid wasp

In behavioral ecology it is generally assumed that behavior is adaptive. This assumption is tested here for sex ratio manipulation in response to host size in the parasitoid wasp Spalangia cameroni. Females produce a greater proportion of daughters on larger hosts. If this behavior is adaptive, it is not through a positive effect of host size on the fitness of daughters, as theory suggests and as found for other species. Females that developed on larger hosts were not more successful at drilling into hosts, were not more successful at interspecific competition for hosts, and did not have greater dispersal ability as measured by wing loading (weight/area of wing and thorax). The possibility that S. cameroni's sex ratio manipulation may be adaptive through a negative effect of host size on the fitness of sons cannot be ruled out. Relative to males from larger hosts, males from smaller hosts had lower wing loading and thus potentially greater dispersal ability. The actual effect of wing loading on fitness remains to be tested.     

Offspring sex ratio response to host size in the parasitoid wasp Spalangia endius

The host size model, an adaptive model for maternal manipulation of offspring sex ratio, was examined for the parasitoid wasp Spalangia endius. In a Florida strain, as the model predicts, daughters emerged from larger hosts than sons, but only when mothers received both small and large hosts simultaneously. The pattern appeared to result from the mother's ovipositional choice and not from differential mortality of the sexes during development. If sex ratio manipulation is adaptive in the Florida strain, it appears to be through a benefit to daughters of developing on large hosts rather than through a benefit to sons of developing on small hosts. Both female and male parasitoids were larger when they developed on larger hosts. For females, developing on a larger host (1) increased offspring production, except for the largest hosts, (2) increased longevity, (3) lengthened development, and (4) had no effect on wing loading. For males, development on a larger host had no effect on any measure of male fitness – mating success, longevity, development duration, or wing loading. In contrast, a strain from India showed no difference in the size of hosts from which daughters versus sons emerged, although both female and male parasitoids were larger when they developed on larger hosts. These results together with previous studies of Spalangia reveal no consistent connection between host-size-dependent sex ratio and host-size-dependent parasitoid size among strains of S. endius or among species of Spalangia. Received: 28 October 1998 / Received in revised form: 20 May 1999 / Accepted: 30 May 1999     

Individual sex ratios and offspring emergence patterns in a parasitoid wasp, Melittobia australica (Eulophidae), with superparasitism and lethal combat among sons

Since the mating of the parasitoid wasp Melittobia australica occurs on their eclosed hosts, the sex ratio is predicted to follow the local mate competition (LMC) theory. However, while LMC models predict that the sex ratio will increase from female-biased toward a 1:1 ratio with an increase in the number of foundresses, the observed female-biased sex ratios (1–5% males) show little increase in response to an increased foundress number. Lethal combat among adult males may serve as an explanation for this observed phenomenon. Using a microsatellite DNA marker, we first examined the individual sex ratio of two foundresses who had sequentially parasitized the same host. Both foundresses produced an extremely female-biased clutch and the sex ratios of the second foundress were only slightly less biased than that of the first. A small number of sons from both foundresses emerged at a constantly low rate over a prolonged period, resulting in a temporal mixture of emerging males derived from both the foundresses. Second, we conducted a one-on-one arena experiment to examine the combat level in relation to the relatedness of the opponents. Almost all the later-emerging males were killed by previously eclosed males irrespective of whether they were sibs or non-sibs. These results suggest that each foundress should not produce males in a single burst, but continue to produce male eggs at a constantly low rate in order to avoid the high mortality of her own sons by lethal male-male combat. This combat may be one of factors in explaining the extremely female-biased sex ratio even with an increasing foundress number.Communicated by R.F.A. Moritz     

Induction of host specificity in larvae of Manduca sexta: chemical dependence controlling host recognition and developmental rate

Summary. The tobacco hornworm, Manduca sexta L. (Lepidoptera: Sphingidae) is a specialist on Solanaceae. This host specificity is induced as the larva feeds on solanaceous foliage, so that solanaceous-reared larvae will refuse to feed on other plants. Experiments were designed to determine the role of dietary constituents on the induction of host specificity and the effects of these on development of M. sexta. Choice assays using leaf discs of cowpea, Vigna sinensis (Fabaceae), were used to monitor the isolation of relevant chemical cues from foliage of potato. An aqueous extract of potato foliage strongly stimulated feeding. This extract was partitioned with n-butanol under alkaline conditions to obtain a highly active butanol extract. Reversed phase flash chromatography with a water-methanol gradient gave an active fraction that was used as a supplement for wheat germ-based artificial diet. Larvae reared on this s-diet became dependent on potato allelochemicals for initiation and continuation of feeding activity. These larvae also developed faster than larvae reared on the control p-diet, but no effect on adult mass was detected. Further flash chromatography of the active fraction under alkaline conditions provided a highly active sub-fraction, and semi-preparative HPLC using gradients of water and acetonitrile resulted in the isolation of a single compound. Bioassays indicated that this compound alone can account for host recognition by solanaceous-reared larvae. The results suggest that the mechanism of induced host specificity in M. sexta involves development of dependence on this compound. Received 21 December 1999; accepted 14 March 2000     

Host choice and offspring sex allocation in a solitary parasitic wasp

Summary Females of the parasitic wasp Antrocephalus pandens can detect differences in the quality of their hosts (pupae of Corcyra cephalonica, a stored-product moth) and allocate offspring of either sex accordingly. Larger and younger hosts are accepted more often in both dead and live hosts; more female offspring emerge from the perceived better hosts, while more males emerge from the smaller, older ones. These patterns are consistent with a sex allocation strategy by the mother, since females from a given size host tend to be larger than males and larger females produce more eggs. However, when wasps lay their eggs in groups of hosts of different size and age rather than encountering them one at a time, no difference in number or sex ratio of offspring is detected between groups. This result and evidence from the change in offspring sex ratio with female age and with numbers of females foraging on a group of hosts are interpreted and discussed in the context of sex allocation (Charnov 1979) and local mate competition (LMC, Hamilton 1967) theories.     

Physiological benefits of feeding in the spring by Lymantria dispar caterpillars on red oak and sugar maple leaves: nutrition versus oxidative stress

The rapid growth of insects that feed on tree leaves in the spring is believed to be due to high nutritional quality. This study tested the hypothesis that both high nutritional quality and low levels of oxidative stress (i.e., toxicological effects) benefit caterpillars that feed in the spring. Fourth-instar larvae of Lymantria dispar (Lepidoptera: Lymantriidae) were used to bioassay the leaves of two contrasting host plants in the spring and summer: red oak (Quercus rubra), a high-quality host, and sugar maple (Acer saccharum), a low-quality host. On spring foliage, the combined effects of rapid consumption rate, efficient nutrient assimilation, and high nutritional quality allowed larvae to grow rapidly and attain larger body mass. Ellagitannins, a major source of oxidative stress in the midgut, were at higher concentrations in the spring than in the summer in maple leaves, but were at negligible levels throughout the growing season in oak. Thus, the impact of phenolic defenses (measured as semiquinone free radicals and oxidized glutathione in the midgut) was not decreased in spring-feeding larvae. Instead, oxidative stress in larvae on maple remained at elevated levels in the spring and summer. By contrast, larvae that fed on oak had consistently low levels of oxidative stress. We conclude that oak and maple were better host plants in the spring because of their higher nutritional quality, and not because of a lower effectiveness of their chemical defenses. This work emphasizes the need to measure not only foliar nutritional and phenolic chemistry but also specific physiological responses in the herbivore, such as oxidative stress. These physiological mechanisms add to our understanding of why spring-feeding life-history strategies have evolved in some insect herbivores.     

Secondary sex ratios do not support maternal manipulation: extensive data from laboratory colonies of spiny mice (Muridae: Acomys)

Spiny mice of the genus Acomys (Muridae) represent a very suitable mammalian model for studying factors influencing the secondary sex ratio (SSR). The maternal effort in these rodents is extremely biased in favour of the prenatal period and, therefore, maternal manipulation of the SSR is potentially more advantageous. We studied the SSR in four populations/species of spiny mice kept in family groups consisting of two closely related females, one non-relative male and their descendants. The groups were established from founding animals aged about 3 months (maturing age) and were allowed to breed freely for several months. Each litter was sexed after birth, and relevant data were thoroughly recorded. Altogether, data were collected on 1684 litters: 189 of Acomys sp. from Iran, 203 of A. cilicicus, 875 of A. cahirinus, and 417 of A. dimidiatus. We recorded the sex of 4048 newborns of which 1995 were males and 2053 were females. The overall sex ratio was close to 1:1 (49.2%). Generalized linear mixed models and/or generalized linear models were constructed to evaluate the effect of four life history and eight social variables on the sex ratio. No consistent effects of these variables on the sex ratio were found and, interestingly, none of the variables associated with maternal life history had any effect on the sex ratio. Three factors associated with group composition (i.e. the number of immature males, the number of immature females and the number of breeding females) did have significant effects on the sex ratio, but these effects were not consistent across the studied species. In conclusion, our evaluation of this large dataset revealed that the sex ratio in spiny mice is surprisingly stable.     

Permanence of Euscorpius carpathicus (L.) larvae on the mother's back (Scorpiones,Chatidae)

Summary The tendency of Euscorpius carpathicus larvae to remain on immobilized live or dried scorpions of the same or different species (Euscorpius flavicaudis) has been tested. Dried scorpions are less attractive than live ones, which are equally attractive regardless of species, sex or reproductive phase. The larvae find old (one year) dried specimens less attractive than freshly or recently (15 days) killed females, which are equally as attractive as live mothers (Table 1). A substratum treated with a chloroform extract of the mother's cuticle is consistently preferred by the larvae. The maternal behaviour (tolerating the larvae on the back) usually disappears about a week after the moult of the larvae, but it can be protracted if they are continually replaced by younger larvae (Table 2). The survival rate of larvae on live mothers is higher than on dried specimens.     

Absence of sex-differential brood raising by workers in Diacamma sp. from Japan

To optimally allocate resources between workers, reproductive females, and males, ant workers have to be able to identify the sex of larvae and raise them differently. The ability of workers to discriminate between the sexes in the brood was tested in colonies of queenless ponerine ants, Diacamma sp., from Japan. The ratio of male eggs in the egg pile was increased experimentally. This manipulation resulted in a corresponding increase in the ratio of adult males, suggesting that Diacamma workers do not raise the sexes differently. Received: 4 November 1997 / Accepted after revision: 14 March 1998     

Ecological costs on local adaptation of an insect herbivore imposed by host plants and enemies

Herbivore populations may become adapted to the defenses of their local hosts, but the traits that maximize host exploitation may also carry ecological costs. We investigated the patterns and costs of local adaptation in the pine processionary moth, Thaumetopoea pityocampa, to its host plants, Pinus nigra and P. sylvestris. The two hosts differ in needle toughness, a major feeding impediment for leaf-eating insects. We observed a west-to-east gradient of increasing progeny size in the Italian Alps, matching the pattern in toughness of their respective local host plant. Eastern populations that feed on the native P. nigra with tough needles had larger eggs, and neonate larvae with larger head capsules, than western populations that feed on the native P. sylvestris and the introduced P. nigra with softer foliage. In a reciprocal transfer experiment that involved the eastern-most and the western-most populations of T. pityocampa from this region, and excluded natural enemies, we found evidence for local adaptation to the host plant. Specifically, larvae from the western population only performed well when raised on their local hosts with soft needles, and they suffered near-complete mortality on the tough foliage at the eastern site. In contrast, larvae from the eastern population survived equally well at both sites. Local adaptation involved a trade-off between progeny size and the number of offspring. We hypothesized that an additional cost, imposed by natural enemies, may be associated with increased egg size: we also observed a west-to-east gradient of increased egg parasitism. We tested this hypothesis in a common garden by exposing eggs of both populations to parasitism by two native egg parasitoids, Ooencyrtus pityocampae and Baryscapus servadeii. The eastern population suffered a higher level of parasitoid attack by O. pityocampae than the western population, and performance of hatched adults of both parasitoids was enhanced in large eggs. Thus, increased neonate quality (larger eggs yielding larger larvae) confers an advantage on tough foliage but incurs the ecological cost of increased parasitism, which may constrain further adaptation by this herbivore.     

Mother really knows best: host choice of adult phytophagous insect females reflects a within-host variation in suitability as larval food

Non-random distribution patterns of specialized phytophagous insects on their hosts may depend on intraspecific differences in plant tissue quality, including nutrients and secondary compounds. Secondary compounds are involved in plant resistance, but are also important for the recognition and acceptability of plants as resources by specialized insects. If individuals within a plant species vary in their content of such secondary substances, there may also be qualitative differences between them. In such cases, natural selection will favor insects with the ability to distinguish and prefer the more suitable plants. In Sweden, the leaf beetle Gonioctena linnaeana Schrank (Coleoptera, Chrysomelidae) is highly specialized on one host, the native willow Salix triandra L (Salicaceae). Field observations reveal that some host plants in a population harbor many feeding larvae, causing severe defoliation, whereas neighboring plants may have few or no feeding larvae. Our hypothesis is that the distribution pattern of G. linnaeana larvae in this population results from qualitative differences between individual host plants in combination with the ability of G. linnaeana females to distinguish between plants that are suitable and not suitable for offspring performance. We examine whether larval survival differs depending on diet and whether the content of secondary chemical compounds explains female preference. Based on the higher survival rate of larvae reared on leaves from preferred hosts, we conclude that G. linnaeana females have evolved a behavior that maximizes offspring performance and thus positively affects female fitness. A chemical survey of the plants indicates that luteolin-7-glucoside and an unidentified flavonoid are important for separating the preferred from the non-preferred plants.     

Dispersal: an alternative mating tactic conditional on sex ratio and body size

Summary Small male milkweed beetles are less successful at obtaining mates than are larger males. Larger males usually win fights and prevent smaller males from obtaining mates and from choosing larger more fecund females as mates. When sex ratios are male-biased, smaller males are particularly likely to experience these mating disadvantages. It follows that smaller males should be especially responsive to their local competitive environment and behave so as to minimize the mating disadvantages of their smaller size. This paper tests the hypothesis that smaller males disperse from host plant patches with male-biased sex ratios and remain in patches with female-biased sex ratios more readily than larger males.Results show both larger and smaller males disperse from patches with male-biased sex ratios more frequently than from patches with femalebiased sex ratios. As predicted, however, small males are more likely to disperse from patches with male-biased sex ratios and remain in patches with female-biased sex ratios than are larger males.The data also show that smaller males dispersing from patches with male-biased sex ratios obtain more matings than non-dispersing males.For milkweed beetles, moving between patches can be viewed as an alternative mating tactic conditional on male body size and local sex ratio.