Interspecific attractiveness of structures built by courting male fiddler crabs: experimental evidence of a sensory trap

Male fiddler crabs Uca musica sometimes build sand hoods and male Uca beebei sometimes build mud pillars next to their burrows to which they attract females for mating. Mate-searching females preferentially approach these structures and subsequently mate with structure builders. Here we show that the preference for structures is not species-specific and argue that it may not have evolved for mate choice. When not near burrows, many species of fiddler crabs approach and temporarily hide near objects, suggesting that hoods and pillars may attract females because they elicit this general predator-avoidance behavior. To test this sensory trap hypothesis we individually released female U. musica, U. beebei and Uca stenodactylus, a non-builder, in the center of a circular array of empty burrows to which we added hoods and pillars and then moved a model predator toward the females. All species ran to structures to escape the predator and the two builders preferred hoods. Next, we put hood replicas on male U. beebei burrows and pillar replicas on male U. musica burrows. When courted, females of both species preferentially approached hoods as they did when chased with a predator. However, males of both species with hoods did not have higher mating rates than males with pillars perhaps because hoods block more of a male's visual field so he sees and courts fewer females. Sexual selection may often favor male signals that attract females because they facilitate general orientation or navigation mechanisms that reduce predation risk in many contexts, including during mate search.     

Hood construction as a spacing mechanism inCleistostoma kuwaitense (Crustacea: Ocypodidae)

The ocypodid crabCleistostoma kuwaitense inhabits the upper shores of the mudflats of Kuwait, where it constructs a semi-permanent mud hood over the entrance to its burrow. Between September and November 1986, data were collected on the distribution and orientation of the hoods and their openings to investigate the crab's social spacing system. Irrespective of density, the distribution of burrow hoods is non-random, tending towards a regular dispersion. A contributory factor for this dispersion pattern is that the surface hoods tend to increase the distance between adjacent burrows. The hood openings are randomly distributed with respect to compass direction but avoid directly facing those of nearest neighbours. In the absence of burrow plugging and eviction as used by other ocypodid crabs, construction of elongated hoods appears to be a good alternative for ensuring social spacing.     

Daring females,devoted males,and reversed sexual size dimorphism in the sand-dwelling spider <Emphasis Type="Italic">Allocosa brasiliensis</Emphasis> (Araneae,Lycosidae)

Sexual selection theory predicts that a higher investment in offspring will turn females into the selective sex, while males will compete for accessing and courting them. However, there are exceptions to the rule. When males present a high reproductive investment, sex roles can reverse from typical patterns, turning males into the choosy sex, while females locate males and initiate courtship. In many spiders, males are smaller than females, wandering in search of sedentary females and maximizing the number of copulations. In the present study, we present findings on the sand-dwelling wolf spider, Allocosa brasiliensis, evidencing a reversal in typical courtship roles reported for the first time in spiders. Males were bigger than females. Females located males and initiated courtship. Copulation always occurred in male burrows and took place mainly in long burrows. Males donated their burrows to the females after copulation, closing the entrance before leaving with female cooperation from inside. Males would provide females with a secure place for ovipositing, being exposed to predation and diminishing their future mating possibilities until constructing a new burrow. The cost of vacating the burrow and losing the refuge in an unpredictable habitat, such as sand dunes, would explain the courtship roles reversal in this spider species. Results turn A. brasiliensis as a promising model for discussing the determinants of sex roles and the pressures that drive their evolution and maintenance. Electronic supplementary material The online version of this article (doi: ) contains supplementary material, which is available to authorized users.     

How do weaponless male fiddler crabs avoid aggression?

Mimicry of females enables weaker males in many species to avoid intrasexual aggression. In fiddler crabs (Uca annulipes), males use their major claw in aggressive interactions to acquire and defend a territory. Males that have autotomised their major claw will be disadvantaged in fighting, but might use their temporary resemblance to females to avoid costly aggressive encounters with other males. We investigated whether: (1) courting males mistake clawless male fiddler crabs for females; (2) clawless males are able to acquire, defend and retain territories as successfully as intact males; and (3) clawless males are more cautious than intact males. Clawless and intact males differed in burrow acquisition methods and fighting behaviour, but were equally successful at acquiring and retaining burrows. While courting males treated clawless males as female, we found no evidence that clawless males mimic the behaviour of females, or that they exploit the advantage of their mistaken identity. Clawless males further appear to avoid male aggression by altering their territorial strategies to minimise the potential for conflict.     

Sand disposal of the painted ghost crab Ocypode gaudichaudii (Decapoda: Ocypodidae): A possible role in courtship

In 1991/1992, we studied the sand disposal behavior of the painted ghost crabs Ocypode gaudichaudii on the Pacific coast of Panamá. O. gaudichaudii either kick, dump or tamp sand they excavate from their burrows. Here we relate these three kinds of sand disposal to burrow structure and distribution, as well as to crab size and sex. Our objective was to determine whether tamping may be a male courtship signal. Burrows whose owners tamped sand were on average longer, deeper, and higher on the beach than were burrows whose owners kicked or dumped sand. Five burrow shapes were distinguished, with half-spiral and spiral shapes being most common among tamped burrows. All crabs excavated from tamped burrows were males. Tamped burrows peaked in abundance around full and new moons. These observations, together with what is known about mating and breeding behavior of other ghost crabs, suggest that tamping may be involved in O. gaudichaudii courtship.     

Biparental investment and reproductive success in a subsocial desert beetle: the role of maternal effort

Parastizopus armaticeps is a nocturnal subsocial detritivorous desert tenebrionid that produces very few offspring per brood. The two environmental factors that constrain reproduction, rapid sand desiccation rate and food scarcity, are countered by biparental effort. Males dig and extend breeding burrows, maintaining their moisture level; females forage on the surface at night for high-quality detritus, the larval food. This was shown to be a scarce and unpredictable resource for which there is high competition. When food was supplemented in a field experiment, offspring number and survivorship doubled and burrow failure due to desiccation dropped from approximately half, the typical failure rate for unsupplemented burrows, to zero. Food supplementation did not, however, increase larval foodstore size and there was no difference in the size of the offspring produced. Supplemented females reallocated their time, foraging less and digging more with the male. This change in maternal behaviour patterns resulted in deeper burrows which remained moist longer, thus extending the larval production period. Female foraging efficiency, particularly food retrieval speed, determined how much time females could allocate to digging, consequently increasing the reproductive success of the pair. Burrow depth and sand moisture level at the burrow base were the major correlates of reproductive success, but the scarcity and unpredictability of high-quality food on the surface and the competition for this resource influenced the number of offspring indirectly through their effect on female behaviour. Received: 29 November 1996 / Accepted after revision: 7 December 1997     

Cannibalize or care? The role of perceived paternity in the sand goby, <Emphasis Type="Italic">Pomatoschistus minutus</Emphasis>

Parental care is a costly part of reproduction. Hence, natural selection should favor males which avoid caring for unrelated young. However, the decision to abandon or reduce care requires cues which are evaluated to give information on potential reproductive value of the offspring. The prediction that male sand gobies, Pomatoschistus minutus, care for foreign eggs as long as they were spawned in their own nest and at least some of such cues are fulfilled was tested. Egg-guarding males that had recently taken part in a spawning event were given a clutch of eggs that was sired either by themselves or another male, in either their own or another male’s aquarium. Males that had not taken part in a spawning event were used as controls and were given eggs sired by another male. We measured the amount of filial cannibalism and nest building. Control group males did not care for eggs and ate them all before rebuilding the nest. In the other treatments, there were no significant effects of paternity, though males moved to another male’s aquarium increased their clutch area threshold and completely consumed larger clutches than males that were not moved. There was no intermediate response in any treatment in the form of increased partial filial cannibalism or less well-constructed nests. Our results suggest that egg-guarding males cannot distinguish between eggs sired by themselves and those sired by other males but are able to react to cues indicating paternity state. Males do not adopt eggs to attract females in P. minutus.     

A conspicuous courtship signal in the fiddler crab Uca perplexa: female choice based on display structure

The fiddler crab Uca perplexa has a conspicuous male courtship signal that is directed at females to attract them to the male’s burrow for mating. The signal involves waving the unflexed large claw up and down. To determine whether the spatiotemporal structure of the wave is under selection by female choice, we examined whether females had a preference for any particular features of the wave. Females respond to a waving display by either visiting the male’s burrow entrance or by electing to pass without visiting the burrow. We filmed mate-searching females and the waving males that they visited or passed. We documented the wave structure of these males using frame-by-frame analysis. Males produce a two-part wave with component A preceding component B. Both components have an upstroke, a pause at the apex and a downstroke. The tip of the claw was raised much higher in B than in A. Visited males had a shorter delay between the two wave components than did males that the females passed without visiting. Visited males also produced component B waves that had a slower upstroke than those of passed males. There was a significant correlation between the relative height of the raised claw and the duration of the upstroke of component B. Females were selecting males that raised their major claw to the highest position (two to three times as high as the carapace width). Passed males brought down their major claw earlier and from a lower position than did visited males. The data suggests that wave structure has evolved through female choice. Male display rate and body size were not female choice cues. An earlier study showed that display duration was also not used by females in selecting mates.     

Temporal constraints and female preference for burrow width in the fiddler crab, <Emphasis Type="Italic">Uca mjoebergi</Emphasis>

We studied sampling behaviour and mate choice in the fiddler crab Uca mjoebergi. Once a female selects a mate, she copulates in his burrow and remains there until releasing her aquatic larvae. U. mjoebergi occurs in habitats that are inundated only by the highest amplitude spring tides. Females can only release their larvae during these tides, and release before or after will result in complete failure of reproductive effort. Matings occur over a 5-day period near the end of neap tides. Our results suggest that within the mating period, females adjust their larval developmental rates by selecting specific burrows in which to incubate their clutches. We found that at the start of the mating period, females chose larger males as mates. Since male size was positively correlated to burrow width, females were selecting wide burrows and effectively incubating at lower temperatures. This would slow down the developmental rates of larvae. In contrast, females that mated late in the mating period selectively chose small males. By incubating in narrower, warmer burrows, these females may increase the developmental rates of larvae. We propose that females are selecting burrows to influence incubation rate and ensure timely release of their larvae. Female U. mjoebergi appear to adjust their preference for the direct benefits of mate choice to increase their reproductive success.     

Perceived predation risk and mate defense jointly alter the outcome of territorial fights

Virtually all animal conflicts occur over access to mates or resources that affect survival, the two key components of fitness. In this paper, we report that predation risk and mate defense jointly affect the outcomes of contests between male sand crabs (Scopimera globosa) for burrows in which crabs mate and take shelter from predators. We observed the contests under three different conditions: (1) the natural condition of low predation risk and without the presence of a female; (2) the first experiment in which we imposed upon only intruding males the perception of predation risk—by digging them from their burrows, capturing and handling them, and placing them into other males’ burrows—to increase the value of the burrows for the intruders as shelter, and (3) the second experiment in which we repeated this treatment but increased the resource value of the burrow to the resident by placing a female in his burrow. The difference in body size between contestants was the main determinant of victory in all analyses. However, perceived predation risk also partly affected the outcomes of the fights: The motivated intruders were likely to win even when they were a little smaller than the residents. In addition, defense of a female had a significant effect on the outcomes of fights: The motivated residents won more fights than the motivated intruders, indicating that these two treatments caused asymmetric increases of the resource value. This is the first report of two external factors simultaneously raising resource value, affecting motivation of contestants, and altering the outcome of fights.     

Natal dispersal in Belding's ground squirrels (Spermophilus beldingi)

Summary Dispersal from the natal site was documented in two populations of Belding's ground squirrels (Spermophilus beldingi) living at different altitudes in the Sierra Nevada of North America. Distance dispersed and age at dispersal were monitored by a combination of observation, trapping, radio telemetry, and examination of road kills. Dispersal was sexually dimorphic in both populations (Tables 1 and 2). All surviving males emigrated before they were 55 weeks of age, with most dispersing midway through the juvenile summer (Fig. 1). By contrast, most females remained within the boundaries of their mothers' home range (Fig. 2 and 3). Those very few females that did emigrate moved distances from their natal burrows similar to those travelled by dispersing males (200–450 m; Table 2), but females tended to disperse at a slightly older age. Significant differences between the study populations were found in distances moved by juveniles of both sexes (Fig. 2). Body weights of juvenile male dispersers were significantly greater than were those of juvenile males of equivilent ages that had not yet dispersed (Fig. 4). The results were considered in light of Shields' (1982) discussion of dispersal and inbreeding. I concluded that if dispersal is related to reproductive success, then dispersal distances selected for use in tests of evolutionary hypotheses should be measured just prior to the subject animals' first reproduction. Finally, although not tested in the present study, the evolutionary hypothesis most consistent with my data suggests that dispersal in S. beldingi may function to minimize nuclear family incest.     

Sex-specific differences in site fidelity and the cost of dispersal in yellow-headed blackbirds

Male migratory birds tend to be more faithful than females to previous breeding sites, suggesting sex differences in costs or benefits of dispersal. In Illinois, greater site fidelity by male yellow-headed blackbirds (Xanthocephalus xanthocephalus) was associated with reduced reproductive success the following year for dispersers relative to non-dispersers. Dispersing females suffered no reduction in reproductive success the following year. Males that attracted few social mates, and thus had low reproductive success, were more likely to disperse, whereas females dispersed in response to low-patch reproductive success, regardless of their individual performance. Males that dispersed appeared to be successful acquiring territories because none was observed as a floater. The rate of dispersal by males in this low-density population was greater than in more dense populations where dispersing males may be less successful at acquiring territories. Despite success at obtaining territories, males that dispersed acquired territories on the periphery of wetlands where fewer females nested, resulting in lower reproductive success. In the second year after dispersing, however, males moved onto more central territories where they acquired larger harems. Thus, dispersal by males may be a long-term strategy requiring at least 2 years for benefits to be realized. Long-term success was enhanced because dispersing males moved to wetlands on which reproductive success was higher than on the wetlands they left. In addition to demonstrating that both individual and patch reproductive success affect dispersal decisions, these data indicate that when evaluating costs and benefits of dispersal, researchers should use a time frame beyond 1 year.     

Reproductive biology of a wood-boring isopod, Sphaeroma terebrans, with extended parental care

The isopod Sphaeroma terebrans Bate, 1866 burrows in aerial roots of the red mangrove Rhizophora mangle L. The burrows serve as shelter and as a reproductive habitat, and females are known to host their offspring in their burrows. I examined the reproductive biology of S. terebrans in the Indian River Lagoon, a shallow lagoon stretching for ∼200 km along the Atlantic coast of Florida, USA. Reproductive isopods were found throughout the year, but reproductive activity was highest in the fall and during late spring/early summer. During the latter periods, large numbers of subadults established their own burrows in aerial roots. The average numbers of S. terebrans per root were high during the fall, but decreased during the winter and reached lowest levels at the end of the summer. Females reached maturity at a larger size than males, but also grew to larger sizes than the males. The average size of females varied between 8 and 10 mm, the average size of males between 6.5 and 8.5 mm. The number of embryos female⁻¹ was strongly correlated with female body length. No indication for embryo mortality during development was found. Parental females (i.e. with juveniles in their burrows) hosted on average between 5 and 20 juveniles in their burrows (range 1 to 59). Most juveniles found in female burrows were in the manca stage and 2 to 3 mm in body length. Juveniles did not increase in size while in the maternal burrow, and juveniles of similar sizes could also be found in their own burrows. Males did not participate in extended parental care, since most of them left the females after copulation. Many females that were born in the summer produced one brood in the fall and a second during winter/early spring. Females that were born in the fall produced one brood during spring/early summer, but then probably died. Extended parental care in S. terebrans is short compared to other peracarid crustaceans. It is concluded that this reproductive strategy in S. terebrans serves primarily to shelter small juveniles immediately after they emerge from the female body, when their exoskeleton is still hardening and their physiological capabilities are still developing. Thus, in S. terebrans, extended parental care probably aids to protect small juveniles from adverse physical conditions in their subtropical intertidal habitat. Received: 9 December 1998 / Accepted: 24 June 1999     

Mating system of the burrowing crab Neohelice granulata (Brachyura: Varunidae) in two contrasting environments: effect of burrow architecture

Few studies conducted in crustaceans have demonstrated how habitat features could shape the mating systems. Here, the burrow of Neohelice granulata was considered as a resource used for mating, and its architecture was characterized in two contrasting study sites: Mar Chiquita Lagoon (MCL), an estuary composed of muddy sediments, and San Antonio Oeste (SAO), a marine bay composed of gravel sediment. Burrow features differed between study sites and occupant gender. Large males constructed burrows with a chamber in MCL and with a widened entrance in SAO, while small males constructed the same narrow burrows as females at both study sites. Field experiments demonstrated that burrows with chambers or widened entries are places used for copulation, although successful post-copulatory guarding was displayed only in those with chambers. The intensity of the agonistic encounters and the success of males in winning resources (burrows/females) also depend on the habitat characteristics. N. granulata shows a resource defense mating system where males employ different mating strategies according to the burrow architecture to ensure mating success.     

Natal philopatry in bannertailed kangaroo rats

Summary I describe prolonged retention of offspring in natal home ranges, or natal philopatry, in the bannertailed kangaroo rat Dipodomys spectabilis. Though weaning occurs at about one month of age, offspring shared natal burrows with their mothers for three to seven months, and 39% of surviving offspring remained within natal home ranges through reproductive maturity. Males as well as females were philopatric. Data on the availability of burrows and on patterns of resettlement suggest that natal philopatry in this species may be a means of providing juveniles with access to essential resources, in this case food caches and large complex burrow systems, that are not readily available outside natal home ranges.     

Differences in context and function of two distinct waving displays in the fiddler crab, <Emphasis Type="Italic">Uca perplexa</Emphasis> (Decapoda: Ocypodidae)

To respond appropriately to communication signals, animals must have the ability to decipher signal meaning. At a basic level, interpreting the difference between territorial and courtship signals can be vital for the survival and reproduction of social animals. Male and female fiddler crabs communicate with claw-waving displays, but the function of these waves remains uncertain. Species differ in the context in which they wave: Some wave during courtship, some during territorial defence and some during both. In this paper, we provide evidence that males of an Australian species of fiddler crab, Uca perplexa, use two different types of claw waving display, lateral and vertical. Lateral waves are employed solely in a courtship context, whilst vertical waves are employed during courtship as well as territorial interactions. Using video recordings, we show that lateral waves were produced spontaneously (broadcast), and their frequency increased only in the presence of female wanderers. Vertical waves were not broadcast but were elicited by male wanderers during agonistic interactions and female wanderers during close range courtship. Male resident U. perplexa were able to discriminate the sex of wandering crabs on the mudflat at distances of 32 cm. During all resident–wanderer interactions, residents attempted to maintain a position directly between the wanderer and the home burrow and orient themselves to face females and to present the major claw towards males. To our knowledge, this is the first demonstration of the multiple use of waving displays in a fiddler crab species.     

Dispersal,population structure and burrow shape of Ocypode cursor

The distribution of the sand crab Ocypode cursor (L.), as indicated by the number of burrows, was studied for 2 years in a 50×50 m sand beach area in northern Israel. A definite relationship was established between the distribution pattern from the seashore inwards towards the sand dunes, and the degree of sand moisture as it changed seasonally. During autumn, more crabs were found at a distance of 15 to 25 m from the sea where sand moisture was about 14%. At the beginning of winter crabs dispersed evely, disappearing with advancing winter. Crabs reappeared in spring, although in smaller numbers, dispersing in a pattern similar to that in autumn. At the beginning of summer and later on, more crabs appeared and concentrated closer to the sea (5 to 10 m). The population structure was analysed directly by measuring the crab's dimensions, and indirectly by counting burrows and measuring the diameter of their openings. Direct analysis revealed two distinct sizeage groups: smaller crabs 0.5 to 3 cm long, and larger ones over 4 cm long. The smaller burrows were inhabited by the first group and were mostly found closer to the sea; the second group was found more landwards. Three main types of burrow shapes are described.     

Indirect evidence indicates female semiochemicals release male precopulatory behaviour in the snow crab,Chionoecetes opilio (Brachyura: Majidae)

Summary Experiments were conducted in aquaria to test the hypothesis that females of the snow crab,Chionoecetes opilio, release a sex pheromone to attract mates. Males exhibited significantly increased activity to water from a source aquarium containing recently-moulted pubescent females, egg-stripped multiparous females and recently-moulted immature females than to water from an aquarium containing berried multiparous females, eggs alone, adolescent males or an empty aquarium. Males without their cephalic chemoreceptors maintained low activity levels in the presence of recently-moulted pubescent females, whereas maxilla-ablated males reacted as strongly as intact males. These findings are discussed within the context of the ecdysteroid-sex pheromone hypothesis, proposed by Kittredgeet al. (1971) and Kittredge & Takahashi (1972), but subsequently rejected. We propose that ecdysteroids from both pubescent and multiparous females may elicit male search-and-clasp behaviour inC. opilio and that the reproductive biology of species used to refute the hypothesis was inappropriate to test the role of ecdysteroids as a cue in the mating process.     

Diurnal variations in the behaviour of the hagfish Eptatretus burgeri

The Japanese hagfish Eptatretus burgeri may be found in relatively shallow water, and is thus accessible to field observations by SCUBA diving. A population of hagfish living in a bay at a depth of about 10 m in Koajiro Bay, Kanagawa Prefecture, Japan has been observed periodically throughout a year by day and night diving. During the day, with rare exceptions, the hagfish remains within its burrow in the soft, muddy substrate; its anterior tip is sometimes just visible within the single opening of the burrow. There are no heaps of mud associated with the burrows, which seem to be relatively impermanent structures. During the night, many hagfish emerge from their burrows, either partially or completely. Partially emerged individuals protrude as much as a third of their length beyond the burrow opening, while completely emerged individuals either rest uncovered on the substrate surface or swim actively near the bottom. An emerged hagfish prefers to reenter a preexisting burrow; when one is not available, however, it quickly excavates a new burrow by swimming vigorously down into the mud. No hagfish were found in the investigated locality from the middle of June to the middle of October.