Size-dependent Mating Behaviours of Male Sand-bubbler Crab,Scopimera globosa: Alternative Tactics in the Life History

Factors leading to the separation of mating behaviours were investigated in the sand-bubbler crab, Scopimera globosa. The crabs mated on the surface (surface copulation, SC) and underground (UC). UC males were large (old) whilst SC males were small (young). Burrowless females bred in the UC males' burrows. These females accepted UC in exchange for access to a burrow. UC occurred much more frequently than SC in the burrow area in which females oviposited. Most SC occurred in the water-saturated area affording a rich diet. SC was accepted by most large and small females in both areas and most UC by small females in the burrow area. SC was an alternative to UC for males in that there was a size dependence between types of copulation. These two mating behaviours involved different degrees of interaction with neighbouring males. Males attempting to carry a female to their burrows for UC were more often disturbed by other males than were males attempting SC. In the interaction for both UC and SC, larger males were likely to resist the disturbances. UC males needed their own burrows, but these burrows were not enlarged before mating. UC males have a higher paternity of eggs than SC males, because SC males' sperm is often displaced by other males. Thus, UC was a behaviour with relatively higher costs and benefits for male crabs than SC behaviour. Alternative mating behaviours in male S. globosa are conditional, and explained by intrasexual interactions and a male life history strategy with a trade-off between growth and reproduction. It is not likely that the size dependence of male mating behaviour is caused by mate preference of females for UC males in the burrow area.     

Inter-male competition and mating success in the field cricket,Gryllus bimaculatus (de Geer)

Males of the field cricket Gryllus bimaculatus compete aggressively for the acquisition of burrows and females. The effect of population density on the nature of encounters between males, and factors affecting individual competitive ability were examined. Aggression was reduced in high population densities, both in terms of the proportion of encounters leading to aggression and in the intensity of aggressive disputes. The addition of burrows and females increased aggressiveness and removed population density effects. Individual competitive ability was determined primarily by body size and secondly, by an individual's past experience of winning (‘confidence’). Competitive ability, prior residence in burrows and the presence of females all appear to influence the outcome of aggressive disputes. Variation in calling frequency of males was influenced both by population density and competitive ability; males who called more often had a greater encounter rate with females. Females were more likely to mate with burrow residents. However, amongst residents, larger males gained more matings. The potential for female choice in this species is discussed.     

Sexual selection for structure building by courting male fiddler crabs: an experimental study of behavioral mechanisms

Males of the fiddler crab Uca musica sometimes build sand hoodsat the entrances of their burrows, to which they attract femalesfor mating with claw waving and other displays. Females significantlymore often approached males with hoods than males without hoods,but once at a burrow, they were just as likely to stay andmate whether the male had a hood or not. To determine how hoodsaffect male attractiveness, we conducted experiments that controlledfor other differences in courtship behavior between buildersand nonbuilders; we removed hood builders' hoods and we addedhood models to nonbuilders' burrows. We then measured the attractivenessof hood builders and nonbuilders with and without hoods. Neithermanipulation measurably affected male courtship behavior. Thepresence of a hood did not increase male—female encounterrates, suggesting that hoods do not attract distant femalesinto a male's courtship range. However, once a male courteda female, she was significantly more likely to approach ifhe had a real or model hood. We obtained direct evidence thatfemales orient to hoods by replacing them with hood modelspositioned about 3 cm away from the openings to males' burrows.Females approached the models, not the courting males, about27% of the time. We conclude that hood building is sexuallyselected because courted females differentially approach hoods,not because hoods attract distant females and not because femalesprefer to mate with hood builders.     

Burrow architecture and digging activity in the Cape dune mole rat

While females are traditionally thought to invest more time and energy into parental care than males, males often invest more resources into searching and displaying for mates, obtaining mates and in male–male conflict. Solitary subterranean mammals perform these activities in a particularly challenging niche, necessitating energetically expensive burrowing to both search for mates and forage for food. This restriction presumably affects males more than females as the former are thought to dig longer tunnels that cover greater distances to search for females. We excavated burrow systems of male and female Cape dune mole rats Bathyergus suillus the, largest truly subterranean mammal, to investigate whether male burrows differ from those of females in ways that reflect mate searching by males. We consider burrow architecture (length, internal dimensions, fractal dimension of tunnel systems, number of nesting chambers and mole mounds on the surface) in relation to mating strategy. Males excavated significantly longer burrow systems with higher fractal dimensions and larger burrow areas than females. Male burrow systems were also significantly farther from one another than females were from other females' burrow systems. However, no sex differences were evident in tunnel cross-sectional area, mass of soil excavated per mound, number of mounds produced per unit burrow length or mass of soil excavated per burrow system. Hence, while males may use their habitat differently from females, they do not appear to differ in the dimensions of the tunnels they create. Thus, exploration and use of the habitat differs between the sexes, which may be a consequence of sex differences in mating behaviour and greater demands for food.     

Location of long‐term communal burrows of a threatened arid‐zone lizard in relation to soil and vegetation

The great desert skink (Liopholis kintorei) of the Egerniinae subfamily (Reptilia: Scincidae) is a communal burrowing lizard that inhabits arid spinifex grasslands in central Australia. Great desert skink activity is centred in and around the burrows which are inhabited for many years. However, it is not known whether skinks select burrow sites with specific attributes or how continuing occupancy of burrows is influenced by the surrounding habitat; especially post‐fire, when plant cover is reduced. Here, we test whether great desert skink burrows in areas burnt 2 years previously and in longer unburnt areas are associated with particular habitat attributes, and whether there are differences between occupied and recently abandoned burrow sites. Vegetation composition, cover and soil surface characteristics at 56 established great desert skink burrows, including occupied and recently unoccupied burrows, were compared with 56 random nearby non‐burrow control sites. Burrow sites had higher plant cover compared with the surrounding landscape in both recently burnt and longer unburnt areas and were more likely to be associated with the presence of shrubs. Soil stability and infiltration were also higher at burrow sites. However, we found no evidence that burrows with lower cover were more likely to be abandoned. Our results suggest that great desert skinks may actively select high cover areas for burrow construction, although differences between burrow and control sites may also partly reflect local changes to plant cover and composition and soil properties resulting from burrow construction and long‐term habitation of a site. Further research should determine if burrows with shrubs or higher plant cover provide greater protection from predators, more structural stability for burrow construction, increased prey abundance or other benefits. We recommend that maintenance of areas with relatively higher plant cover be prioritized when managing great desert skink habitat.     

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.     

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.     

Elevated predation risk changes mating behaviour and courtship in a fiddler crab

The fiddler crab, Uca beebei, lives in individually defended burrows, in mixed-sex colonies on intertidal mud flats. Avian predation is common, especially of crabs unable to escape into burrows. Mating pairs form in two ways. Females either mate on the surface at their burrow entrance (''surface mating'') or leave their own burrow and sequentially enter and leave (''sample'') courting males'' burrows, before staying in one to mate underground (''burrow mating''). We tested whether perceived predation risk affects the relative frequency of these mating modes. We first observed mating under natural levels of predation during one biweekly, semi-lunar cycle. We then experimentally increased the perceived predation risk by attracting grackles (Quiscalus mexicanus) to each half of the study site in two successive biweekly cycles. In each experimental cycle, crabs were significantly less likely to mate on the side with more birds. Moreover, on the side with elevated predation risk, the number of females leaving burrows to sample was greatly reduced relative to the number of females that surface-mated. Males waved less and built fewer mud pillars, which attract females, when birds were present. We discuss several plausible proximate explanations for these results and the effect of changes in predation regime on sexual selection.     

Strategies of mate finding in the European field cricket (Gryllus campestris) at different population densities: a field study

Abstract. 1. Members of a field population of Gryllus campestris L. varied in their walking and calling activity. In both sexes, some individuals occupied burrows whereas others walked around in the observation area. Males at burrows could be either silent or calling.
2. In the course of one summer, population density decreased and the initial balanced sex ratio changed to a large surplus of males.
3. At high population density, there were equal numbers of non-calling males at burrows, calling males at burrows and walking males, while walking females predominated over females at burrows. Non-calling males at burrows achieved more encounters with females than did calling and walking males. Females met males by walking through the population and by waiting at burrows. Thus, calling and phonotaxis were not essential for mate finding and calling was less effective than previously thought.
4. At low population density calling males predominated. Calling males at burrows achieved the most encounters with females. Females met males only by walking around in the population area. Calling was thus more important in mate finding than at high population density.
5. Changes in sex ratio and population density may cause the flexibility in mate finding behaviour of individual crickets.     

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.     

Leishmaniasis in the Jordan Valley. V. Dispersal characteristics of the sandfly Phlebotomus papatasi

Several characteristics of dispersing and non-dispersing Phlebotomus papatasi (Scopoli) were quantified and compared. The majority of dispersing sandflies, trapped crossing fallow fields, were females (68.5% v. 51.1%); of the dispersing females, 55.4% were parous, 48.1% were inseminated and 11.6% were gravid. In the population of sandflies sampled exiting from burrows of the sand rat Psammomys obesus Cretschmar, these categories, respectively, represented 39%, 90% and 26% of the females examined. Leishmania promastigotes were found in 9% of females exiting from P. obesus burrows, and in 2.7% of the dispersing females. The anthrone test established that the reason for activity of gravid females is sugar feeding. These females do not disperse and are suitable targets for future control measures.     

Selection of food patches by sympatric herbivores in response to concealment and distance from a refuge

Small herbivores face risks of predation while foraging and are often forced to trade off food quality for safety. Life history, behaviour, and habitat of predator and prey can influence these trade‐offs. We compared how two sympatric rabbits (pygmy rabbit, Brachylagus idahoensis; mountain cottontail, Sylvilagus nuttallii) that differ in size, use of burrows, and habitat specialization in the sagebrush‐steppe of western North America respond to amount and orientation of concealment cover and proximity to burrow refuges when selecting food patches. We predicted that both rabbit species would prefer food patches that offered greater concealment and food patches that were closer to burrow refuges. However, because pygmy rabbits are small, obligate burrowers that are restricted to sagebrush habitats, we predicted that they would show stronger preferences for greater cover, orientation of concealment, and patches closer to burrow refuges. We offered two food patches to individuals of each species during three experiments that either varied in the amount of concealment cover, orientation of concealment cover, or distance from a burrow refuge. Both species preferred food patches that offered greater concealment, but pygmy rabbits generally preferred terrestrial and mountain cottontails preferred aerial concealment. Only pygmy rabbits preferred food patches closer to their burrow refuge. Different responses to concealment and proximity to burrow refuges by the two species likely reflect differences in perceived predation risks. Because terrestrial predators are able to dig for prey in burrows, animals like pygmy rabbits that rely on burrow refuges might select food patches based more on terrestrial concealment. In contrast, larger habitat generalists that do not rely on burrow refuges, like mountain cottontails, might trade off terrestrial concealment for visibility to detect approaching terrestrial predators. This study suggests that body size and evolutionary adaptations for using habitat, even in closely related species, might influence anti‐predator behaviors in prey species.     

Lekking and the small-scale distribution of the sexes in the Caribbean fruit fly,Anastrepha suspensa (Loew)

Male and female Anastrepha suspensa(Loew) had a clumped distribution in the foliage of their guava host plants. Males were no closer to other males than they were to females or than females were to other females. Flies were often found in roughly the same locations over time. However, contemporaries (flies present at the same time) were closer to each other than subsequent flies were to their predecessors. Males were more likely to be found near spots previously occupied by males than they were to spots used previously by females. Some trees had more flies than others, but there was no regional (northwest, etc.) preference within trees. Females were no more likely to be found in the vicinity of clumped (lekking) males than they were by isolated males. About a third of the females taken from inside leks had sperm in their spermathecae, and it is not clear if their motive for being in these areas was sexual. In pairs of males (within 15 cm of each other), the larger fly tends to be in a position farther up the branch, suggesting that larger males may control preferred territories. It seems possible that males attempting to intercept females accumulate in favorable microhabitats where females are likely to be concentrated and that leks have evolved from such clumping.     

Not all who wander are lost: nest fidelity in <Emphasis Type="Italic">Xylocopa virginica</Emphasis> examined by mark recapture

Relocation to novel nests (sometimes called drifting) in flying Hymenoptera is often interpreted as the result of navigation error and guard bees erroneously admitting foreign individuals into the nest. We studied nest fidelity and nest relocation of both females and males in a nesting aggregation of Xylocopa virginica in southern Ontario, Canada, where females can nest either solitarily or socially. Adult female and male bees were trapped at nest entrances, individually paint marked, and then released. Subsequent recapture patterns were used to assess nest fidelity: that is, how faithful individuals were to their home nest and how often they moved to another nest. Bees were considered to have relocated if they were recaptured in a nest different from the one in which they were initially trapped, indicating that they had spent at least one night in a new nest. Some females were only captured in one nest, some occasionally moved to new nests, temporarily or permanently, and a few were never caught in the same nest twice. In addition, females relocated to nests that were further away in 2007 when population density was low, suggesting that they seek out and claim nesting spaces when they are available. Males relocated more frequently than females, with most drifting from nest to nest in no obvious pattern. This indicates that males spend the night wherever space is available or in nests nearest to their territories. This study reveals that for both female and male X. virginica, nest membership is not as stable as once thought.     

Evolutionarily stable nesting strategy in a digger wasp.

Two alternative “strategies” will not coexist in a population unless on average they are equally successful. The most likely way for such an equilibrium to be maintained is through something equivalent to frequency-dependent selection. Females of the digger wasp Sphex ichneumoneus (Sphecidae) nest in underground burrows. They usually dig and provision these by themselves but occasionally a nest is jointly occupied. The two wasps fight whenever they meet and in the end only one of the two females lays an egg in the shared nest. Two models based on the theory of mixed evolutionarily stable strategies were developed and tested on comprehensive field data from two North American populations of these wasps. The first model proposes two strategies called founding and joining. Founders start burrows alone, but they are more successful when they are joined by a joiner. At equilibrium founders and joiners are equally successful, which amounts to an amicable, sharing relationship. The predictions of this amicable model are decisively rejected by the data. The second model proposes two strategies called digging and entering. Diggers dig their own burrows but they often have to abandon these burrows because of temporary unsuitability. Enterers move in later, thereby exploiting abandoned burrows as a valuable resource. They do not distinguish an adandoned burrow from one that is still occupied. Therefore sharing of burrows arises as an unfortunate by product of selection for entering abandoned burrows, and Model 2 is not an amicable model. Its quantitative predictions are impressively fulfilled in one population, though not in another population. This is one of the only examples yet known of a mixed evolutionarily stable strategy in nature. Yet the word strategy itself can confuse, and this paper tries the experiment of substituting “decision”, defined as a moment at which the animal commits future time to a course of action.     

Behaviour, population changes and dispersal of mountain hares (Lepus timidus) in Scotland

Mountain hares in north-east Scotland spent the day in forms in long heather and moved downhill in the evening to feed in hill pastures. Here they grazed intensively, often in groups of 4–6 individuals. In June and July adult females, then pregnant or lactating, grazed in daylight on the pastures. Leverets spent less time grazing, and more in play and exploration. They spent the day in cover near the feeding areas, and if disturbed during grazing crouched or went into cover, while adults fled. During 1982 and 1983 males predominated in groups of grazing hares, but in 1984 both sexes were equally represented. These changes in sex ratio were reflected in the increased proportion of females trapped for marking between February and late July during 1982–86. There was a dominance order related to weight among male hares and dominant hares approached more females. There was no firm evidence of mate-guarding. Males approached females regardless of their oestrous state and were usually rebuffed with varying degrees of intensity including striking and chasing. There were no interactions between females. Adults of both sexes chased leverets for short distances but leverets joined groups of feeding adults. Neither leverets nor first-winter hares showed evidence of dispersal. Mountain hares avoided sheep and cattle and there were fewer hares after the arrival of sheep in May.     

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.     

Pelage color variation in pocket gophers (Rodentia: Geomyidae) in relation to sex,age and differences in habitat

Pelage coloration is a phenotypic characteristic in mammals that could be associated with an individual's survival and fitness; thus coloration gains adaptive importance. In geomyids, pelage coloration shows a relationship with the color of soils on which they live, mainly freshly dug soil of their burrows. This characteristic could be due to a camouflage adaptation to avoid predators. Pocket gophers disperse aboveground to establish a territory before they reach reproductive condition, and males disperse longer distances than females. The aim of this study was to evaluate pelage color variation in pocket gophers (Thomomys anitae) in relation to habitat differences, sex, and age, and determine its association to the color of the soil on which they live. Brightness of T. anitae's dorsal pelage coloration and that of soil samples from five different habitats in the Baja California peninsula, Mexico were measured to test four hypotheses: (1) Subadults show a wide coloration range, but extreme colors are lost in adulthood due to natural selection. (2) Males are more vulnerable to depredation than females; therefore, males’ coloration is more homogenous as a protective camouflage. (3) In open habitats pocket gophers are more exposed to being detected by predators, therefore their pelage coloration pattern is less variable than that of individuals from habitats with more vegetation cover. (4) Pelage coloration better matches soil coloration in moist conditions similar to that of freshly dug soil of their burrows. The results confirmed our predictions; however, selection does not impose an equal pressure on pelage coloration on the five habitats evaluated; other factors such as population density and predator presence need to be assessed. The strongest effects are found in the most open habitat, and there is less strong support for predictions in habitats where predator assemblage is diminished.     

Responses of the endangered pygmy bluetongue lizard to conspecific scats

Many animals use chemical signals for communication between conspecifics and for territory marking. The pygmy bluetongue lizard is normally solitary, focussing activity around the entrance of its burrow, from where it ambushes prey, and rarely contacts other individuals. In this paper we examined whether lizards in laboratory experiments alter their behaviour in the presence of scats from conspecifics. In the first experiment, when lizards were offered a choice of two vacant burrows with or without a scat close to the entrance, they tongue flicked more often at the burrow entrance when the scat was present, and more often chose to occupy the burrow with the scat. An interpretation is that lizards use scat signals to recognise burrows that may be suitable because they have previously been occupied by a conspecific, but that they approach those burrows cautiously in case a resident is still present and likely to resist a takeover. Scats from male lizards were inspected (by both sexes) for longer than scats of female lizards. In the second experiment, when resident lizards were presented with scats outside of their burrows, they inspected and tongue flicked at those scats more often if the scat came from a male than a female lizard, but there was no definitive evidence from our experiments that lizards differentiated in their response to scats from lizards that were found close to or far from the test lizard. The results were consistent with a communication system in which lizards use scats to advertise their presence, independent of any direct contact.     

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.