Non-breeding behaviour of Magenta Petrels Pterodroma magentae at Chatham Island, New Zealand

Magenta Petrels Pterodroma magentae were caught at light-attraction stations on southwest Chatham Island, New Zealand, and most were fitted with transmitters. Of 52 captured since 1993, 71% were males, and all 36 tracked adequately proved to be non-breeders in the breeding season of capture. Our data indicated no sex bias in their probability of being captured at lights. Males provided 86% of trackings, and 87% of trackings of birds flying over the breeding area were males. Males landed 118 times; females 13 times. Only males were found on the ground, by night and day, apparently unassociated with burrows (three with and ten without transmitters), but subsequently digging burrows ( n  = 8). Of 19 birds banded as fledglings up to 2000, males were first recaptured nearing 4 years old (at lights and on the ground) and a female nearing 6 years old (in burrow). Among 37 fledglings, the sex ratio was even. Nine tracked males occupied burrows, as did two females, but the latter were older recaptures (10+ and 25+ years old). It appears that only males claimed existing, or dug new, burrows. They then attracted a mate to the burrow by means unknown, but from among females frequenting an inshore courtship area near the colony, or occasionally flying over the colony, at night. Females established in burrows, but then losing their mate, were able to re-mate there, by calling from near the burrow or by attracting a mate in flight or from the postulated inshore courtship area. Both sexes sometimes took years to pair or re-mate, possibly reflecting the dearth of available mates.     

Burrow ornamentation in the fiddler crab (Uca leptodactyla): female mate choice and male–male competition

Non-biological ornamentation is found in the nests and burrows of different kinds of animals. We evaluated here whether sand hoods constructed by male fiddler crabs (Uca leptodactyla) are one of the signals used by males to attract females during courtship. We observed females when they were walking among the males, and we quantified the proportion of females that visited male burrows with and without ornamentation and the choice to stay in a male’s burrow. Females visited more burrows with hoods than burrows without hoods, and they chose significantly more builder males. Male investment in ornamentation nevertheless decreased when the proportion of females increased in the area. Male investment was not correlated with the proportion of non-builder males nearby, but was positively correlated with overall density. The density sex ratio, however, was more male-biased in high-density than in low-density areas suggesting that even if building attracts females, the function could be related to male competition for mates.     

Lekking in Gryllotalpa major, the Prairie Mole Cricket (Insecta: Gryllotalpidae)

Gryllotalpa major is a rare, burrowing insect native to the tallgrass prairie of the south-central United States and is known to exhibit 'lek-like' behavior during mating. Here I report on a study carried out in the field that demonstrates that the prairie mole cricket meets all criteria defining a classical lekking species. Males construct specialized acoustic burrows from which they call to attract females for mating. I show that these burrows, which seem to serve no purpose other than for sexual advertisement and mating, are aggregated spatially on at least three scalar levels. Females fly through the aggregation of burrows and drop to the ground in the vicinity of calling males, and are, thus, not constrained in choosing a mate. Females enter the males' acoustic burrows, but I argue that the burrows are not used as oviposition sites, and that the males do not otherwise sequester resources important to females. Although the term 'lek' is useful for the discussion of mating systems, its definition remains ambiguous. I discuss the current usage of the term and suggest extensions.     

Chorusing Behaviour,a Density-dependent Alternative Mating Strategy in Male Common Toads (Bufo bufo)

Male chorusing behaviour was studied in a population of common toads (Bufo bufo) on the island of Öland south Sweden, and the functional role of male advertisement calling in this species was experimentally examined. Calling males were larger and heavier than non-calling males (t = 2.41, p < 0.025 and t = 2.36, p < 0.025, respectively). However, small males were also found to call. This indicates that large males persisted in calling for longer and/or called more often. The proportion of calling males decreased as population size increased during the breeding season, indicating that calling is a low density strategy. Females responded more readily to calls than males. There were insufficient data to determine if the dominant frequencies of advertisement calls were inversely correlated with male body size, however, this relationship was found for the similar release calls. Females were found not to discriminate between high and low frequency calls, but when given a choice between two calls of different sound pressure levels (SPL), females were attracted to the louder calls. Thus, the function of chorusing is to advertise the position of males to mate-seeking females when the population density is low.     

Reproductive behaviour of the South American fruit fly Anastrepha fraterculus (Diptera: Tephritidae): laboratory and field studies

Abstract. The mate choice, courtship and oviposition behaviour of laboratory-reared and field-collected Anastrepha fraterculus (Wied.) were compared. In laboratory cultures in Southampton the duration of male calling activity in small leks increased gradually from 1-2h at 5 days old to up to 7 h at 10 days. This finding correlates with previous reports on the time at which male salivary glands, which are believed to produce sex pheromone, are fully developed. Wild flies which emerged from infested fruits in Brazil began to oviposit on the day they mated, whereas in laboratory flies oviposition began 1 day following the first mating. Both types of fly usually defended their position on a particular fruit throughout the day, and re-mated with either virgin or mated males. There was no significant difference in mating duration. Females did not copulate before the mean age (±SE) of 16.8±0.9 days. For both types of flies mating initiation occurred in the first 2h of photophase, with virgin females choosing mainly mated males. The average number of matings in the laboratory was three for females and four for males, and the interval between matings in females was significantly increased after the second mating. It is suggested that the tendency of virgin females to mate with mated males will lead to increased fitness, as males are on average 48 days old at their second mating. The potential life span of around 200 days for both sexes would allow adults to bridge the gap between seasonally available fruits in warm-temperate and sub-tropical South America.     

Nocturnal Calling Behavior in Mantids

Females of Mantis religiosa and Empusa pennata were video taped for several 24 h periods to determine if they showed behaviors associated with pheromone release. In the photophase the abdomen of both species was motionless and rested in continuous contact with the wings. However, at the beginning of the scotophase the females bent the abdomen ventrally so that the space between the abdomen and the wings increased significantly with respect to the daytime posture. Calling behavior (abdominal bending) was maintained throughout the 8 h scotophase and ended abruptly at lights on. Females of M. religiosa did not start calling until they were 30 days of age. Calling disappeared in mated females, but it reappeared two weeks later. Males stayed motionless in response to the odors emitted by other males or by noncalling females, but walked when a calling female was placed in the air flow. These observations suggest that female mantids bend their abdomens at night to release a sex pheromone. The adaptive function of nocturnal sex pheromone release in sexually cannibalistic species that rely strongly on visual cues for mating is discussed.     

Differential dispersal and female-biased sex allocation in a parasitic wasp

1. Differential dispersal of males and females from a population is predicted to result in biased sex-allocation decisions, even in the absence of sibmating.
2. Mated Bracon hebetor Say (Hymenoptera: Braconidae) females produce distinctly female-biased sex ratios (≈ 30% male), yet sibmating is not a feature of the mating biology of this species. Therefore the dispersal behaviour of male and female B. hebetor from caged subpopulations was examined.
3. A higher proportion of females than males dispersed from the caged subpopulations. Furthermore, females dispersed earlier than males. This suggests that the level of competition for mates experienced by males is higher than the level of competition for hosts experienced by sisters.
4. Roughly half of the dispersing females left after they had mated. Females generally mate once in their lifetimes, suggesting that competition between brothers for mates may be high.     

Familiarity and flexible mating strategies of a solitary rodent, Dipodomys ingens

Because mating is a product of individual reproductive strategies that may vary with changing conditions, we predicted variable mating behaviour in an arid-adapted, territorial rodent, the giant kangaroo rat, Dipodomys ingens. We also predicted that familiarity would facilitate nonaggressive courtship and mating in this solitary rodent. Through direct observations in the field, we found that mating varied from exclusive to multiple partners. Where densities were low, and on nights when multiple females were in oestrus, each animal mated with one member of the opposite sex. In conditions where the operational sex ratio was skewed towards multiple males, males footdrummed and competed for females. Males were able to mate with one or two females in adjacent territories, and they successfully competed for these same females throughout the breeding season. Females that mated exclusively with one male had more pups emerge from the burrow compared with females that experienced male competition. Females allowed nearest neighbour males to enter their burrows, and they engaged in more nonaggressive contact with close neighbours than with other males. Paired encounters in the field showed less aggression towards neighbours than strangers. In laboratory tests, females were less aggressive towards and allowed more contact with familiar than unfamiliar males. These results show that D. ingens can alter mating strategies as conditions change. Familiarity is an important factor in nonaggressive interactions between males and females and may be important to mate preferences in females during reproduction. The less aggressive behaviour to neighbours than to strangers (‘dear enemy’ phenomenon) is consistent with other solitary animals that defend multipurpose territories. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Limited flexibility in female Pacific field cricket (Teleogryllus oceanicus) exploratory behaviors in response to perceived social environment

How populations adapt, or not, to rapid evolution of sexual signals has important implications for population viability, but is difficult to assess due to the paucity of examples of sexual signals evolving in real time. In Hawaiian populations of the Pacific field cricket (Teleogryllus oceanicus), selection from a deadly parasitoid fly has driven the rapid loss of a male acoustic signal, calling song, that females use to locate and evaluate potential mates. In this newly quiet environment where many males are obligately silent, how do phonotactic females find mates? Previous work has shown that the acoustic rearing environment (presence or absence of male calling song) during late juvenile stages and early adulthood exposes adaptive flexibility in locomotor behaviors of males, as well as mating behaviors in both sexes that helps facilitate the spread of silent (flatwing) males. Here, we tested whether females also show acoustically induced plasticity in walking behaviors using laboratory‐reared populations of T. oceanicus from Kauai (HI; >90% flatwings), Oahu (HI; ~50% flatwings), and Mangaia (Cook Islands; no flatwings or parasitoid fly). Though we predicted that females reared without song exposure would increase walking behaviors to facilitate mate localization when song is rare, we discovered that, unlike males, female T. oceanicus showed relatively little plasticity in exploratory behaviors in response to an acoustic rearing environment. Across all three populations, exposure to male calling song during development did not affect latency to begin walking, distance walked, or general activity of female crickets. However, females reared in the absence of song walked slower and showed a marginally non‐significant tendency to walk for longer durations of time in a novel environment than those reared in the presence of song. Overall, plasticity in female walking behaviors appears unlikely to have facilitated sexual signal loss in this species.     

The energetic cost of calling in the variable field cricket, Gryllus lineaticeps

Abstract. Female preferences for conspicuous male calls have been documented in many groups. However, relatively few studies have examined the metabolic costs associated with the production of call types preferred by females. We measured the oxygen consumption of calling male Gryllus lineaticeps Stål crickets using closed chamber respirometry. Calling song was recorded concurrently. The average increase in mass-specific oxygen consumption during calling was 2.7 times basal rates of oxygen consumption, and calling males consumed approximately 1.2ml O₂g^-lh^-1. Oxygen consumption increased with increasing chirp rate and pulse duration, but not with increasing chirp duration. Females of this species prefer higher chirp rates, thus some call types that increase the male's attractiveness to females require more metabolic energy to produce.     

The natural history of a monogamous coral-reef fish, Valenciennea strigata (Gobiidae): 2. behavior, mate fidelity and reproductive success

This study examined the behavior and reproduction of a monogamous coral-reef fish, Valenciennea strigata, to determine mate fidelity and the proximate causes of monogamy. Most fish were found in monogamous pairs that remained together over several rounds of reproduction. Pairs stayed within close proximity to each other and their burrows. Females fed at a higher rate than their mates, while males spent more time maintaining burrows. Females spawned every 13 days; males guarded eggs in the burrow for 2–3 days. Although females limited the RS of males, males did not mate polygynously under natural conditions. Reproductive success (RS) was affected primarily by survival, and secondarily by size. Both sexes enforced monogamy by guarding their mates. Three factors facilitated mate guarding: (1) all males were able to hold a nest site, (2) both sexes showed strong site fidelity, and (3) residents had an advantage in contests over mates. Thus, mates were economically defensible. Additionally, females formed a crescent of dark pigments on their abdomen that resembled a gravid condition; these marks may enhance continuation of the pair bond. Both sexes preferred large mates, and pairs were positively assorted by size. Males benefited from guarding large females because fecundity increased with size. Females may benefit from the burrowing of males, and larger males should be better burrowers.     

Social organization in the dusky-footed woodrat (Neotoma fuscipes): A field and laboratory study

The dusky-footed woodrat has been characterized as solitary and asocial. In this research I sought to determine some of the parameters of woodrat social organization. In the field, I live trapped, marked and released members of a distinct group of woodrats at bi-weekly intervals from February to June. Twenty-two residents were identified and 21 out of 34 houses were found to be occupied. Visiting occurred regularly, most often at non-occupied houses. Males were unlikely to be visited at their house and were more likely to visit occupied houses than females. Females were visited at home and visited each other. In June the population was removed to the laboratory where paired encounters with familiar and unfamiliar woodrats were used to examine the effect of sex and familiarity on social interactions. Both females and males interacted little with familiar same sex conspecifics, with the behaviour of one member of such pairs being very inhibited. Females with strange females were more interactive and spent more time in contact. Neither sex clearly differentiated between familiar and strange conspecifics of the opposite sex. Males were agonistic or affiliative in encounters with females. The type of response was consistent for a given male, and females responded differently to the two types. It is concluded that woodrats differentiate between same sex conspecifics and show sufficient individual variability to make individual recognition possible and adaptive.     

Pillar Function in the Fiddler Crab Uca beebei (II): Competitive Courtship Signaling

Male Uca beebei court and attract females into burrows they defend on muddy sand flats in the intertidal zone on the Pacific coast of the tropical Americas. Mating, oviposition and incubation (breeding) occur underground in males' burrows. Some courting males build mud pillars (2 cm high) at the entrance of their burrow. The purpose of this field study was to assess the role of pillars in competitive courtship signaling among males. I studied the effect of pillars on female behavior by recording the responses of wandering females to courtship from males resident at burrows with and without pillars. I also caught females, released them individually in a circular arena with an equal number of empty burrows with and without pillars around its circumference, and chased the females with a simulated avian predator. Females moved to burrows of both types more often when they were courted (82%) than when they were chased (67%). Receptive females were attracted to the burrows of the males that courted them significantly more often (97%) when these burrows had pillars than when they lacked pillars (66%). However, once females entered males' burrows they were equally likely to remain, mate and breed in both types of burrows. Females also more often moved to burrows with pillars (66%) than to burrows without pillars when they ran from the simulated predator. Both male courtship displays and pillars probably provide cues females use to locate males' burrows. The visual similarity between pillars and a display courting males give immediately before they enter their burrows suggests that pillars are icons of the display. The effect of pillars on female behavior, the timing of pillar building relative to when females choose mates, and contrasts in the behavior of males that do and those that do not build pillars suggest that pillar building has evolved due to competition among males to attract females into their burrows.     

Laboratory breeding studies of freshwater crayfish, Austropotamobius pallipes (Lereboullet)

SUMMARY. 1. Mature crayfish, collected from an Irish lake before breeding had started, were held in breeding combinations and their mating and brooding activities observed.
2. All mating attempts were initiated by the male. A single mating led to spawning within 6 days but a subsequent mating cancelled the effects of the first. Males mated more often when there were more females present. Males lacking a major cheliped mated less often than did normal males.
3. Larger males mated more often than did smaller males, and although males showed no female size preference, matings were less frequent and generally unsuccessful when males were much larger than females; the female was usually killed. Large females mated successfully with smaller males.
4. Females held at high densities with a larger male mated earlier than at low densities. However, aggression also increased with density; at high densities males fought and killed females.
5. Males held in pairs without females fought; in occasional mating attempts spermatophores were not positioned correctly. Paired females rarely fought; all spawned normally although unmated. Although their eggs soon died and were removed during grooming, brooding behaviour continued for at least 2 months.
6. Brooding females held in pairs shed pleopodal eggs during aggressive encounters. Females held singly showed a lower initial rate of egg loss.     

Factors affecting female sexual receptivity in the planthopper, Prokelisia dolus

Abstract. The mating system of Prokelisia dolus Wilson (Homoptera: Delphacidae) was characterized by determining: if males and females multiply mate; when transitions occur in female sexual receptivity, what triggers sexual refractoriness; and what behaviours characterize unreceptive virgins, receptive virgins, and unreceptive mated females. Males copulated with up to six females in less than 1 h, but completely inseminate, on average, only the first four females. Females rarely mated more than once, unless males were depleted of sperm due to previous copulations or if copulation was interrupted (if duration was<2 min). Male and female calling was associated (100% and 91%, respectively) with sexual receptivity and resultant matings. The transition from unreceptive virgin to receptive (calling) mature virgin occurred 48 h posteclosion, and all were mated by day 4. Females that were sexually immature and those completely inseminated did not call. Rejection of males by females included walking away from approaching males (65%), female kicking (7%), and abdominal lifting (5%). Rejection of males was observed by immature, mature and calling, and mated females. Sexual refractoriness was not triggered by acoustic and visual stimuli or mechanical stimulation of genitalia. Refractoriness was also not triggered by reception of small quantities of sperm because some females laid a few viable eggs yet calling was not terminated. Sexual refractoriness was activated by a substance in the ejaculate as demonstrated by injection into the haemocoel of male accessory glands or testes and homogenates of seminal vesicles. This is the first study that documents the role of male ejaculate in inhibiting female sexual receptivity in Hemiptera (Homoptera).     

Calling behavior of the female german cockroach,Blattella germanica (Dictyoptera: Blattellidae)

Virgin German cockroach females, Blattella germanica(L.), were observed, for the first time, to exhibit a characteristic calling behavior during which females emit a volatile sex pheromone. Under a photoperiod of 12L12D, the percentage of 7-day-old virgin females that exhibited this behavior peaked before the end of the scotophase in a similar pattern to the diel periodicity of mating. A clear relationship was evident between calling and stages of sexual receptivity during successive gonotrophic cycles. Females initiated calling 5–6 days after the imaginal molt, when their basal oocytes were 1.6 mm long. If not mated, females continued to exhibit bouts of calling during the next 3–4 days until 24 h before ovulation. Calling was completely suppressed by mating as well as the presence of an egg case in the genital atrium in both virgin and mated gravid females. We suggest that calling and the emission of a volatile sex pheromone serve to attract males from a distance as well as to potentiate responses to contact sex pheromone in aggregations.     

Sex differences in developmental plasticity and canalization shape population divergence in mate preferences

Sexual selection of high-quality mates can conflict with species recognition if traits that govern intraspecific mate preferences also influence interspecific recognition. This conflict might be resolved by developmental plasticity and learned mate preferences, which could drive preference divergence in populations that differ in local species composition. We integrate field and laboratory experiments on two calopterygid damselfly species with population genetic data to investigate how sex differences in developmental plasticity affect population divergence in the face of gene flow. Whereas male species recognition is fixed at emergence, females instead learn to recognize heterospecifics. Females are therefore more plastic in their mate preferences than males. We suggest that this results from sex differences in the balance between sexual selection for high-quality mates and selection for species recognition. As a result of these sex differences, females develop more pronounced population divergence in their mate preferences compared with males. Local ecological community context and presence of heterospecifics in combination with sex differences in plasticity and canalization therefore shape population divergence in mate preferences. As ongoing environmental change and habitat fragmentation bring formerly allopatric species into secondary contact, developmental plasticity of mate preferences in either or both sexes might facilitate coexistence and prevent local species extinction.     

Male behaviour over the season in a wild population of the field cricket Gryllus campestris L.

Abstract.  1. Cricket model systems have greatly enhanced our knowledge about mate choice and sexual selection. However, no studies exist that take seasonal dynamics into account and that follow individual males over the complete adult stage in the wild. In the present study, all adult field crickets ( Gryllus campestris L.) were individually marked and measured in a population in central Germany and their behaviour was followed over their life from May to July 2000.
2. The population reached a maximum of 108 males within 10 days and continuously declined thereafter. Daily mortality was positively correlated with burrow-changing activity, which was highest at high mean temperatures. Median burrow occupation time was short at 1 day but males stayed up to 34 days in one burrow. Heavy males stayed longer in burrows and achieved a higher age.
3.  Male displacement was on average 2.8 m per day and up to 284 m during their life. Displacement and lifespan were negatively correlated.
4. Burrows were highly aggregated at the meadow in areas of vegetation cover <85%. Aggregation of males increased with advancing season and declining population size. Consequently, nearest-neighbour distances remained stable at 5 m until the 50th day of the study.
5. About half of the 159 detected matings occurred 4–6 h after sunrise. An increased mating frequency was observed between the 30th and 50th day after adult moult.     

Intraspecific acoustic communication and mechanical sensitivity of the tympanal ear of the gypsy moth Lymantria dispar

Tympanal ears of female gypsy moths Lymantria dispar dispar (L.) (Lepidoptera: Erebidae: Lymantriinae) are reportedly more sensitive than ears of conspecific males to sounds below 20 kHz. The hypothesis is tested that this differential sensitivity is a result of sex‐specific functional roles of sound during sexual communication, with males sending and females receiving acoustic signals. Analyses of sounds produced by flying males reveal a 33‐Hz wing beat frequency and 14‐kHz associated clicks, which remain unchanged in the presence of female sex pheromone. Females exposed to playback sounds of flying conspecific males respond with wing raising, fluttering and walking, generating distinctive visual signals that may be utilized by mate‐seeking males at close range. By contrast, females exposed to playback sounds of flying heterospecific males (Lymantria fumida Butler) do not exhibit the above behavioural responses. Laser Doppler vibrometry reveals that female tympana are particularly sensitive to frequencies in the range produced by flying conspecific males, including the 33‐Hz wing beat frequency, as well as the 7‐kHz fundamental frequency and 14‐kHz dominant frequency of associated clicks. These results support the hypothesis that the female L. dispar ear is tuned to sounds of flying conspecific males. Based on previous findings and the data of the present study, sexual communication in L. dispar appears to proceed as: (i) females emitting sex pheromone that attracts males; (ii) males flying toward calling females; and (iii) sound signals from flying males at close range inducing movement in females, which, in turn, provides visual signals that could orient males toward females.     

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.