Fluctuating asymmetry, call structure and the risk of attack from phonotactic parasitoids in the bushcricket Sciarasaga quadrata (Orthoptera: Tettigoniidae)

The acoustically orienting parasitoid Homotrixa alleni (Diptera: Tachinidae) is a significant mortality factor of calling male Sciarasaga quadrata (Orthoptera: Tettigoniidae), with less than 10% of males escaping parasitism within the duration of the calling season. This study examined fluctuating asymmetry (FA) (small, random deviations from perfect symmetry) in five morphological traits in S. quadrata and its relation to the survival of calling males in the field. A significant relationship was demonstrated in only a single trait; across three consecutive field seasons and among each of the collections per season, parasitised males were shown to have significantly more asymmetric hind tibia than unparasitised males. Since the morphological development of males is complete prior to parasitoid attack, this relationship is not caused by the activities of the parasitoid. Further experimentation revealed that there was no relationship between the level of FA in the hind tibia and the success of parasitism, with males artificially parasitised in the laboratory showing no difference in FA from those where parasitism was unsuccessful. Instead, FA was shown to be negatively related to chirp length, an aspect of call structure that significantly affects the survival of calling males under parasitism risk in the field. We explore the possible reasons why more asymmetric males produce calls of shorter chirp length and hypothesise that the difference in hind tibia asymmetry seen between parasitised and unparasitised males in the field is related to host location in H. alleni rather than the inability of S. quadrata to defend itself against parasitoid attack. Received: 14 November 1997 / Accepted: 20 April 1998     

Stage-specific sex differences in Drosophila immunity to parasites and pathogens

Arguments from life-history theory predict that other things being equal females are likely to invest more in defence against parasites and pathogens than males. This is either because males and females differ in behaviour or, more importantly, because the variance in mating success is typically higher in males than in females. Such effects are likely to be most pronounced in those developmental stages where sex differences are greatest. In most organisms, but especially in holometabolous insects, this will be the adult stage. We explored sex-specific resistance to four natural enemies of Drosophila melanogaster that attack the insect at different developmental stages: the larval parasitoid Asobara tabida, the pupal parasitoid Pachycrepoideus vindemiae; and the adult pathogens Beauvaria bassiana (a fungus) and Tubulinosema kingi (a microsporidian). Measures of resistance were designed to reflect the four species’ natural history. Female larvae were able to defend themselves more strongly against A. tabida than males and there was weak evidence that adult females suffered less from microsporidian attack than males. No differences were found for the other two species. Our results provide some support for lower investment in defences in males, and we discuss why the strongest effect was found at the larval rather than the adult stage contrary to our prediction.     

Factors influencing egg parasitism in sub‐social insects: insights from the treehopper Alchisme grossa (Hemiptera,Auchenorrhyncha, Membracidae)

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Defence strategies against a parasitoid wasp in Drosophila: fight or flight?

Hosts may defend themselves against parasitism through a wide variety of defence mechanisms, but due to finite resources, investment in one defence mechanism may trade-off with investment in another mechanism. We studied resistance strategies against the parasitoid wasp Leptopilina boulardi in two Drosophila species. We found that D. melanogaster had significantly lower physiological resistance against L. boulardi than D. simulans, and hypothesized that D. melanogaster might instead invest more heavily in other forms of defence, such as behavioural defence. We found that when given a choice between clean oviposition sites and sites infested with wasps, both D. melanogaster and D. simulans detected and avoided infested sites, which presumably limits later exposure of their offspring to infection. Unlike D. simulans, however, D. melanogaster laid significantly fewer eggs than controls in the forced presence of wasps. Our findings suggest that D. melanogaster relies more heavily on behavioural avoidance as defence against wasp parasitism than D. simulans, and that this may compensate for a lack of physiological defence.     

Oviposition behavior of an ant-parasitizing fly,Neodohrniphora curvinervis (Diptera: Phoridae), and defense behavior by its leaf-cutting ant hostAtta cephalotes (Hymenoptera: Formicidae)

This study examines the oviposition behavior of the phorid parasitoid Neodohrniphora curvinervisand the antiparasitoid defense behavior of its leafcutting ant host Atta cephalotes. N. curvinervisfemales are diurnal sit- and- wait parasitoids that attack only outbound foragers of head width 1.6 mm or greater. Females deposit a single egg through the foramen magnum of each host successfully parasitized. Pursuit of hosts is usually initiated when an outbound forager of acceptable size passes by a parasitoid perch site. Individual foragers defend themselves against pursuing parasitoids by outrunning them along the foraging trail or by standing their ground and fending them off with their legs,antennae, and mandibles. At the colony level, susceptible foragers are protected against parasitism by a shift in the forager size distribution toward smaller unsusceptible sizes during the day when parasitoids are active and toward larger sizes at night when parasitoids are inactive. The frequency of parasitism of susceptible foragers was 15%, which is more than five times the frequency found in another system involving the phorid parasitoid Apocephalus attophilusand the leafcutting ant host Atta colombica.We offer several possible explanations for such differences in the frequency of parasitism and also examine reasons for the high incidence of superparasitism (19%) observed in the system studied.     

A defensive endosymbiont fails to protect aphids against the parasitoid community present in the field

1. The value of protective mutualisms provided by some facultative endosymbionts has been well demonstrated in the laboratory, yet only recently has their effectiveness in the field been studied. ‘Candidatus Hamiltonella defensa’ is known to defend aphids from parasitoid wasps in laboratory trials. However, the efficacy of this defence varies among parasitoids, suggesting that protection will vary spatially and temporally depending on parasitoid community composition. 2. This demonstrated specificity and a dearth of studies on Hamiltonella in the field prompted the authors to quantify parasitism rates of Hamiltonella‐infected and ‐uninfected Aphis craccivora Koch aphid colonies in a manipulative field study. 3. It was found that A. craccivora in central Kentucky alfalfa were parasitised by Lysiphlebus testaceipes (Cresson) and Aphelinus sp. Surprisingly, Hamiltonella infection did not lower successful parasitism by the naturally occurring parasitoid wasps. Whether Hamiltonella was effective against L. testaceipes was subsequently tested in a controlled laboratory assay, and no effect on parasitism rate was found. 4. This study emphasises the fact that defensive symbionts sometimes provide no tangible defensive benefits under field conditions, depending on parasitoid community composition. It is hypothesised that the protective mutualism may be beneficial in geographically localised areas. When the symbiosis is effective against a local parasitoid community, aphid clones may experience eruptive population growth and rapidly disperse across a large area, allowing spread to habitats with different parasitoid communities where the mutualism is an ineffective defence.     

Apple sawfly Hoplocampa testudinea (Hym., Tenthredinidae) and its parasitoid Lathrolestes ensator in Dutch apple orchards (Hym., Ichneumonidae, Ctenopelmatinae)

Abstract: Changes in population densities of the apple sawfly Hoplocampa testudinea (Klug) and its parasitoid Lathrolestes ensator (Brauns) were monitored in 15 apple orchards for a period of up to 4 years. The parasitoid species was found in all orchards except one, and was more numerous in plantings on sandier soils. Post‐bloom insecticide applications against other pests, and carbaryl for fruit thinning, often decimated both host and parasitoid in integrated pest management orchards. In organic orchards, where synthetic pesticides are banned, and Quassia is the only remedy against sawfly, the pest is more problematic. The low levels of parasitism in all organic orchards, except one, were possibly due to the application of wettable sulphur during the parasitoid flight period. The sawfly usually recovers more quickly than its parasitoid when chemical control is discontinued, because propagation of L. ensator is limited in various ways. The parasitoid is time limited, because suitable second instar host larvae are rarely available for more than a week on a single apple variety. Moreover, parasitism levels stay moderate because the parasitoid females do not avoid superparasitism. Finally, relatively more sawflies than parasitoids emerge after 10 months underground, because the incidence of prolonged diapause is more elevated in L. ensator than in the sawfly. It is suggested that both the high incidence of prolonged diapause and the inability to avoid superparasitism are useful in reducing the risk of local extinction. Elevated sawfly attack in a single early apple variety would reduce exploitation of suitable host larvae in other nearby varieties, in as far as the parasitoid is not able to distinguish fruitlets with accessible second instar host larvae from those with inaccessible older larvae. Although the former are available for a limited time, the latter may keep the parasitoids from moving to the less abundant second instar larvae in late(r) varieties. Reduction of elevated host density in such an early variety by a properly timed application of a short‐lived pesticide, such as Quassia, increases parasitism levels proportionally, and is expected to promote parasitoid movement to host larvae in other varieties nearby.     

Effect of mating combinations on the host parasitisation and sex allocation in solitary endoparasitoid,Aenasius arizonensis (Hymenoptera: Encyrtidae)

Aenasius arizonensis (Girault) is an important solitary endoparasitoid of Phenacoccus solenopsis Tinsley. To optimise the mass production of high-quality females, it is important to assess the influence of mating regimes on the progeny fitness and sex allocation. We, therefore, hypothesise that mating combinations in A. arizonensis adults emerged from different host instars may influence parasitism and sex allocation in the subsequent generation. Therefore, we compared three nymphal instars (1st, 2nd and 3rd) and adults host stages of P. solenopsis for parasitism and sex allocation by A. arizonensis. Further, F₁ female progeny of the parasitoid emerged from different host instars was henceforth evaluated for its fitness in six mating combinations. A. arizonensis females parasitised all the host stages except the 1st instar nymphs. The parasitised 2nd instar nymphs yielded only males, while the sex ratio in the later host instars was strongly female-biased. The parasitoid females preferred 3rd instar nymphs with respect to higher parasitism (74.0–84.0%) and produced more females in the F₁ progeny as compared to other host stages. F₁ females that emerged from 3rd instar nymphs produced significantly higher parasitism (74.0–79.0%). These mating combinations also yielded more female progeny in the F₂ generation. However, parasitism by F₁ females was significantly lower (9.0–12.0%) when mated with males that emerged from 2nd instar P. solenopsis nymphs. Moreover, latter combinations yielded only male progeny in F₂ generation. These findings can be used in laboratory mass rearing of this parasitoid vis-à-vis biological control of P. solenopsis.     

Interactions between the Parasitoid Rhopus nigroclavatus (Ashmead) (Hymenoptera: Encyrtidae) and its Mealybug Hosts Tridiscus sporoboli (Cockerell) and Trionymus sp. (Homoptera: Pseudococcidae)

This research investigated age-class-specific parasitism rates of the buffalograss mealybugs Tridiscus sporoboli (Cockerell) and Trionymus sp. by Rhopus nigroclavatus (Ashmead) (Hymenoptera; Encyrtidae), size class preference of this parasitoid, and mealybug–parasitoid interactions through choice and no-choice studies. In the no-choice studies, the mean rates of parasitism by R. nigroclavatus were 45, 20, 0, and 0%, respectively, for mealybugs adult female, third and fourth instars, first and second instars, and eggs. Choice studies indicated that rate of parasitism increased with host size. The mean rates of parasitism on mealybugs in the choice studies were 100% for adult females, 24% for third and fourth instars, 0% for first and second instars, and 0% for eggs. A second set of choice studies investigating mealybug/parasitoid behavior revealed that R. nigroclavatus oviposits in all post-egg mealybug age classes, but first and second instars were less often parasitized than older mealybugs.     

Signal transduction downstream of salicylic and jasmonic acid in herbivory-induced parasitoid attraction by Arabidopsis is independent of JAR1 and NPR1

Plants can defend themselves indirectly against herbivores by emitting a volatile blend upon herbivory that attracts the natural enemies of these herbivores, either predators or parasitoids. Although signal transduction in plants from herbivory to induced volatile production depends on jasmonic acid (JA) and salicylic acid (SA), the pathways downstream of JA and SA are unknown. Use of Arabidopsis provides a unique possibility to study signal transduction by use of signalling mutants, which so far has not been exploited in studies on indirect plant defence. In the present study it was demonstrated that jar1‐1 and npr1‐1 mutants are not affected in caterpillar (Pieris rapae)‐induced attraction of the parasitoid Cotesia rubecula. Both JAR1 and NPR1 (also known as NIM1) are involved in signalling downstream of JA in induced defence against pathogens such as induced systemic resistance (ISR). NPR1 is also involved in signalling downstream of SA in defence against pathogens such as systemic acquired resistance (SAR). These results demonstrate that signalling downstream of JA and SA differs between induced indirect defence against herbivores and defence against pathogens such as SAR and ISR. Furthermore, it was demonstrated that herbivore‐derived elicitors are involved in induced attraction of the parasitoid Cotesia rubecula     

The reproductive strategy of the gregarious parasitoid,Pteromalus puparum (Hymenoptera: Pteromalidae)

Summary Pteromalus puparum is a gregarious parasitoid of lepidopterous pupae. To determine in which phase of a host outbreak superparasitism occurs, field investigations were carried out on seasonal prevalence of the host, Papilio xuhtus, and parasitism by P. puparum in a citrus grove in Fukuoka, Japan in 1974. Host pupation occurred from May to November and the parasitoid attacked the host throughout this period. Pupal density increased rapidly after mid-August and the percentage parasitism decreased during this period. A high level of parasitism was attained after one or more parasitoid generations in mid-September. The superparasitism was observed after mid-September when the parasitoid attained extremely high density, and resulted in an increase in the proportion of males, high mortality, and a decrease in the size of the progeny.     

Parasitism of the grasshopper Melanoplus sanguinipes by a sarcophagid fly, Blaesoxipha atlanis: influence of solitary and gregarious development on host and parasitoid

Blaesoxipha atlanis (Aldrich) (Diptera: Sarcophagidae) is a common parasitoid of the grasshopper Melanoplus sanguinipes (F.) (Orthoptera: Acrididae) in western Canada. We tested the hypothesis that B. atlanis can develop as either a solitary or a gregarious parasitoid, and assessed the influence of parasitism on the growth and survival of infected grasshoppers. Males and females of M. sanguinipes were parasitized manually with one, two, or three first-instar larvae of B. atlanis in the laboratory. Parasitism was more deleterious to males than females of M. sanguinipes; females are larger than males. Host survival and longevity declined with the number of larvae per host in a sex-specific manner. In females, 39%, 24%, and 8% of hosts containing, respectively, one, two, and three sarcophagid larvae survived parasitism. Although 41% of single-parasitized males survived, all males containing more than one larva died. Variations in host quality as measured by dry mass explained much of the response to parasitism in male, but not female, hosts. Parasitoid larvae, apparently, did not cause significant physical damage to host organs and tissues but instead functioned as a metabolic sink. The greater metabolic activity associated with egg production could account for the relatively higher tolerance to parasitism of female, as opposed to male, grasshoppers. Developmental time, adult size, and percentage survival of B. atlanis declined with the intensity of parasitism, especially in parasitoids developing in male hosts. Females developing gregariously contained fewer ovarioles at eclosion than counterparts developing as solitary larvae. The mean body size of field-collected B. atlanis did not differ from that of laboratory-reared parasitoids developing singly in a host. Gregarious development is an alternative strategy to solitary development that may enable B. atlanis to maintain population numbers during periods of grasshopper scarcity.     

Asymmetric size effect of sexes on reproductive fitness in an aphid parasitoid Aphidius ervi (Hymenoptera: Aphidiidae)

Most studies on size–fitness relationships focus on females and neglect males. Here, we investigated how body size of both sexes of an aphid parasitoid, Aphidius ervi Haliday, affected the reproductive fitness. Reproductive fitness was generally positively correlated with body size for both sexes in this species. Large individuals of both sexes had greater longevity, large males fathered more progeny, and large females had higher fecundity, parasitism, and greater ability in host searching and handling. We demonstrated in this study that size effects of males and females were asymmetric on different reproductive fitness parameters. With increasing body size females gained more than males in longevity and fecundity while males gained more than females in the number of female progeny. Regardless of female size, large males sustained a female-biased population longer than small males. These results suggest that male body size should also be considered in the quality control of mass-rearing programs and the evaluation of parasitoid population growth.     

Response of a parasitoid fly, Gymnosoma rotundatum (Linnaeus) (Diptera: Tachinidae) to the aggregation pheromone of Plautia stali Scott (Hemiptera: Pentatomidae) and its parasitism of hosts under field conditions

We studied the attraction of a tachinid fly, Gymnosoma rotundatum (Linnaeus) to the male-produced aggregation pheromone of the brown-winged green bug, Plautia stali Scott, its parasitism on the bug, and its seasonal occurrence in the field. The tachinid fly was continuously attracted to the aggregation pheromone from spring to autumn and utilized the bugs as hosts. Our field experiment to clarify the effect of the pheromone on parasitism demonstrated that parasitism occurred only in female bugs baited with synthetic aggregation pheromone and did not occur in females without the pheromone. The parasitoid flies therefore appeared to use the bug’s pheromone as a host-finding kairomone. The pheromone attracted not only female flies but also males. Male flies may increase their chance of encountering pheromone-attracted females by waiting near pheromone sources. The tachinid develops multiple generations in active hosts from spring to autumn and overwinters in dormant hosts. Thus, G. rotundatum seems to be highly adapted to using P. stali as its host, and it is a potentially important biological control agent of P. stali populations in the field.     

Reproductive biology of the microtype tachinid fly Zenillia dolosa (Meigen) (Diptera: Tachinidae)

Reproductive biology including mating, adult longevity, fecundity and development of the tachinid fly Zenillia dolosa was investigated for optimizing rearing procedures using Mythimna separata as a host in the laboratory. Females lay microtype eggs containing a first instar larva on food plants of the host and then the eggs must be ingested by the host for parasitization. Mating success was 58.5% with mating duration of 80.7 min. Mating was most successful when day 0–1 females were kept with day 2–4 male flies. Female body size was positively correlated with its fecundity but not with longevity. However, females that survived longer produced more eggs during their lifetime. Parasitoids successfully developed in 4th to 6th instar host larvae. Host instars at the time of parasitoid egg ingestion significantly influenced development time of the immature parasitoid, but did not affect body size of the emerging parasitoid. We suggest that pairing newly emerged females with day 2–4 males should result in higher mating success and using the last instar hosts for parasitization should minimize development time of the parasitoid for rearing.     

Evaluation of Diachasmimorpha longicaudata (Hymenoptera: Braconidae) as a mortality factor of Ceratitis capitata (Diptera: Tephritidae) infesting Citrus species under laboratory and field-cage conditions

Ceratitis capitata (or medfly) is one of the major pests currently affecting fruit crops in northwestern Argentinian Citrus-producing areas. Medfly populations are sustained in large exotic fruits, such as Citrus paradisi, Citrus aurantium and Citrus sinensis, which are known to hinder the activity of almost all native parasitoid species. Therefore, a feasible approach to controlling medfly involves the use of exotic parasitoids such as Diachasmimorpha longicaudata. In this study, the prediction that parasitoid females would be proficient at finding medfly larvae infesting the Citrus species mentioned earlier was tested. Particularly, the variation in fruit species preference by parasitoid females, the efficacy of the parasitoid to kill medfly and the effect of host density on parasitoid performance were determined. Parasitoids were allowed to forage for 8 h on grapefruits and oranges artificially infested with medfly larvae under controlled (laboratory) and uncontrolled (field cage) environmental conditions. Fruit choice and no-choice tests were performed. Results were assessed by comparing the number of female visits to and ovipositor insertions into the fruit, and parasitoid emergence, parasitism and additional host mortality percentages. Parasitoid preference for visiting larger fruits (sour orange and grapefruit) may be related to the increased fruit surface area. Ovipositional activity on fruit was influenced by the variation of the larval host density per unit of fruit surface. The higher parasitism rates recorded from sweet orange would be mainly a result of both increased host density and fruit physical features. Nevertheless, D. longicaudata showed the capacity to parasitise hosts in all Citrus species tested.     

Male Rock Sparrow (Petronia petronia) Nest Defence Correlates with Female Ornament Size

We investigated the relationship between male nest defence and female breast patch size in an alpine population of rock sparrow (Petronia petronia) in northern Italy. We presented a mounted weasel (Mustela nivalis), a common nest predator, to 28 pairs breeding in nest boxes, with 12–13‐d‐old nestlings, and measured the intensity of male and female defence reaction. We measured the frequency of attack flights, intensity of alarm calling and total time spent in view, and then combined these for each individual, in a single defence factor by principal component analysis. All the females arrived to defend the nest while only 21 males arrived, and females defended the nest more intensely than males. We analysed, by stepwise regression, the relationship of male defence factor to female behaviour and phenotype (breast patch size, a measure of quality) and brood properties (size, mass, phenology). Male defence factor was significantly related only to female breast patch size. We argue that male rock sparrows apparently make parental investment decisions according to their mate's quality, and examine possible alternative hypotheses.     

Active defence of herbivorous hosts against parasitism: Adult parasitoid mortality risk involved in attacking a concealed stemboring host

Phytophagous insects have several defence strategies to defend themselves against attack by parasitic wasps. Larval lepidopteran hosts can defend themselves actively to prevent oviposition by the parasitoid. Among the aggressive kinds of behaviour exhibited by hosts against parasitoids are violent wriggling, biting and spitting. The behaviour of the braconid parasitoid Cotesia sesamiae attacking stemboring larvae inside their feeding tunnel in the plant stem was investigated in maize and sugarcane stem pieces and transparent artificial tunnels. Attacking a defending stemborer host inside the confined space of a tunnel was shown to be risky for the female parasitoid. A considerable proportion (25%) of female wasps were killed in their attempt to attack the spitting and biting host. No difference was found in the behaviour of C. sesamiae attacking the suitable host Sesamia calamistis or the unsuitable host Eldana saccharina. The consequences of this high mortality risk involved in each host attack is discussed in relation to the ecology of the parasitoid.     

Vine mealybug sex pheromone increases citrus mealybug parasitism by Anagyrus sp. near pseudococci (Girault)

The present study was aimed at elucidating the role of lavandulyl senecioate (LS), the sex pheromone of Planococcus ficus, in host selection of the parasitoid Anagyrus sp. near pseudococci. Field trials were carried out in Portugal, Italy and Israel. The effect of LS on the parasitism rate of the wasp was determined by exposing sentinel mealybugs combined with pheromone dispensers impregnated with LS, in comparison with other baits: lavandulyl isovalerate (LI); planococcyl acetate (PAc); and unbaited control traps. In addition, in order to study the host location behavior of A. sp. near pseudococci, pheromone dispensers were placed at three different distances: inside the trap, 30 or 60 cm away from the trap. The number of parasitoid females inside the traps, the number of parasitized mealybugs, and the number of days required for the first parasitoid emergence were recorded. The response of A. sp. near pseudococci females to different doses of LS (25–1350 μg) was also evaluated using sticky plate traps. The rate of mealybug parasitism by A. sp. near pseudococci was significantly increased by LS in the three parasitoid populations. PAc and LI had no significant effect on the wasp parasitism rate in most of the trials. However, the Italian population of the parasitoid responded to PAc, showing apparently a different behavioral pattern. The number of parasitoid females trapped did not significantly differ between tested doses of LS. The use of LS as an arrestant in host location by the A. sp. near pseudococci female is discussed.     

ECOLOGY AND EVOLUTION OF MATING SYSTEMS OF FIDDLER CRABS (GENUS UCA)

1. General accounts of the natural history and behaviour of fiddler crabs suggest there exist two broad mating patterns in the genus. Most western and Indo-Pacific species mate on the surface of intertidal substrates near burrows females defend. The sexes associate only briefly during courtship and mating. In contrast, males of many American species court from and defend burrows to which females come for mating. Copulation occurs underground in burrows plugged at the surface; the sexes usually remain together for at least several hours. Here we summarize and contrast recent detailed field studies of the mating systems of U. pugilator, an American species, and U. vocans, a species widely distributed in the western and Indo-Pacific. We indicate how differences in the breeding ecology of these two species may account for basic differences in modes of sexual selection leading to the two broad mating patterns in the genus. 2. U. pugilator burrows in protected sandy substrates in the upper intertidal and supratidal zone. During ebb tide, nonbreeding crabs leave burrows they occupy during high tide to forage on food-rich substrates in the lower intertidal zone. Reproductively active males remain in the burrow zone where they fight for and defend burrows from which they court. Large males win most fights for burrows and tend to defend burrows high on the elevation gradient, especially during periods with relatively high tides. Females usually approach and descend the burrows of several males before choosing their mates by remaining in males' burrows. Males remain underground with their mates for 1–3 days until after they oviposit their eggs. Some males then emerge and leave their burrows while others sequester their mates in the chambers where mating and oviposition has occurred, dig new chambers and resume courtship, perhaps attracting additional females. In either case, females remain underground for approximately 2 weeks, finally emerging to release their planktonic larvae. Burrows that do not collapse due to tidal inundation or flooding by groundwater are best for breeding and usually are located relatively high on the elevation gradient. Females choose mates indirectly by preferring to breed in burrows that will remain intact while they oviposit and incubate their eggs. Large males mate more often than small males because they are better able to defend burrows at locations females prefer to breed. The mating system of U. pugilator may be classified as resource-defence polygyny. 3. U. vocans burrows in open muddy substrates in the mid- to lower intertidal zone. At a site near Chunda Bay, Australia, where the reproductive behaviour of this species has been studied in depth, both sexes feed near burrows they defend. Females tend to occupy their burrows for longer periods and move shorter distances than do males. Mating occurs on the surface near the burrows that females defend. Females accept both resident and wandering males as mates. They show no preference for mating with larger males. Female choice may be based on other male morphological or behavioural characteristics. Females oviposit their eggs either while on the surface or in their burrows. They produce relatively small clutches and are active on the surface throughout their breeding periods. Males fight both their neighbours and wandering males. Large males tend to win fights and defend burrows in areas where large females, which produce relatively many eggs, are most dense. Such areas may offer greater protection from predators than areas occupied by smaller females. Small males mate about as often as large males but may father fewer larvae. The mating system of U. vocans is resource-free and promiscuous. 4. The mating systems of U. pugilator and U. vocans differ fundamentally in that female U. pugilator require access to a specific microenvironment to breed successfully, while female U. vocans do not. We suggest this difference occurs because of contrasts in clutch sizes and the mobility and movement patterns of feeding females. Female U. pugilator produce relatively large clutches and probably experience more intense selection from factors that can cause egg loss and mortality than do U. oocans, which produce clutches of sufficiently small volume to be protected by their abdominal flaps. Hence, the range of suitable breeding environments for U. pugilator is small compared to that for U. vocans. In addition, U. pugilator burrows in areas that are relatively food-poor, leading to daily migrations to and from food-rich substrates in the lower intertidal zone, preventing female defence of an area suitable for both breeding and feeding. U. vocans, however, burrows in areas sufficiently rich to support feeding, leading to relatively low female mobility and defence of burrows that are also suitable breeding sites. 5. Adaptive radiation of the genus Uca in the Americas is manifest by trends toward smaller adult size, higher population densities, more frequent microgeographic sympatry and increased terrestriality, compared to species in the western and Indo-Pacific regions. We outline the general features of the selection mechanisms tying each of these trends to the evolution of resource—defence mating systems. Intraspecific variation in the courtship behaviour and site of mating in U. lactea and U. vocans supports our contention that resourse—defence behaviour tends to occur at high population densities. Additional data are needed to evaluate the other hypotheses critically.