Males are attracted by their own courtship signals

Courting male fiddler crabs Uca terpsichores (1 cm carapace width) sometimes build mounds of sand called hoods at the entrances to their burrows. Males wave their single enlarged claws to attract females to their burrows for mating. It was shown previously that burrows with hoods are more attractive to females and that females visually orient to these structures. In this study, we test whether males also use their hoods to find their burrows. We first determined the maximum distance that males can see and find a burrow opening without a hood. Males were removed from their burrows and placed on the sand at a range of distances from a burrow opening. If they were more than about 8 cm (seven units of eye-height) away, they were unable find the burrow. In contrast, males that were burrow residents used a non-visual path map to return to their burrows from much greater distances. To determine if hoods help males find their burrows when there are errors in their path maps, we moved residents 1–49 cm on sliding platforms producing errors equal to the distances they were moved. Males with self-made hoods or hood models at their burrows relocated their burrows at significantly greater distances than did males with unadorned burrows. Hood builders also relocated their burrows faster. Hence, hoods have two functions: they attract females and they provide a visual cue that males use to find their burrows quickly and reliably when their path maps fail. An erratum to this article can be found at     

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.     

Factors limiting distribution and activity patterns of the soldier crab Dotilla myctiroides in Phuket, South Thailand

This study was carried out between January and March 1995 on the intertidal sand flats of Tang Khen Bay, Phuket, South Thailand, where the soldier crab Dotilla myctiroides (H. Milne-Edwards) occurs in densities of up to 120 m⁻². In this bay, long, ribbon-like sand waves (wavelength 40 m, height 0.4 m) are interspersed with shallow pools, running approximately parallel to the shore. During daylight low-tides, exposure of the sand waves is followed 15 to 20 min later by the emergence of the crabs which have been buried under the sediment surface during high tide. Their subsequent burrowing and feeding activity results in the production of large numbers of sand pellets on the sediment surface. Most crabs retreat down their burrows, and some also plug the burrow entrance, prior to being covered by the incoming tide. The crab burrows have a distinct distribution on the sand waves. Burrows are most dense at the top of each sand wave, and a band of unburrowed sediment adjoins the adjacent tidal pools. Crabs are most abundant between mean high-water neap-tide level and mean low-water neap-tide level, where the median particle size of the surface sediment is ∼2 . Measurements of water-table depth below the sand waves and the exposure time of the sediment indicated that, where sediment size is suitable, the main factor controlling crab distribution is the duration of daytime exposure. This observation is in contrast to those of many previous studies, which have suggested that water-table height and sediment drainage are the main factors controlling the distribution of D. myctiroides. Received: 14 January 1998 / Accepted: 6 May 1999     

Significance of mangrove–mudflat boundaries as nursery grounds for the mud crab, Scylla paramamosain

Fisheries managers frequently try to protect juveniles in order to preserve stocks. Juveniles can be protected by either implementing changes designed to avoid catching immature animals (e.g. increasing mesh size or altering fishing techniques) or protecting nursery grounds. To prevent the capture of immature animals, an estimate of size at maturity is required as well as a knowledge of both fishing methods and the exact location of the nursery grounds. Strong demand for juvenile mud crabs to stock aquaculture ponds has resulted in development of fisheries targeting crabs of all sizes from instar 1 to mature individuals. Using five different fishing methods, different stages in the life cycle of Scylla paramamosain were followed for a period of 16 months in an estuarine population in the Mekong Delta, Vietnam. Mangrove habitat utilisation begins when crabs settle out from the plankton at instar 1 [modal internal carapace width (ICW), 0.5 cm] amongst the pneumatophores at the mangrove fringe. Increasingly larger crabs were found deeper into the mangrove but they were still living on the surface (modal ICW size class, 1.5 cm). As their size increases, the crabs either dig burrows (modal ICW size class, 4.5 cm) or they live in the sub-tidal zone, migrating into the mangrove with each tide to feed (modal ICW size class, 4.5 cm). Larger crabs were caught offshore (modal ICW size class, 12.5 cm) where females accounted for 60% of the catch although of these, only 63% were mature. Recruitment of early instars was continuous but peaked in December to February. Subsequent peaks in the catch rates of larger size classes indicated the development of a single cohort with an estimated growth rate of 2.0 cm ICW per month. On the basis of abdominal width, females were estimated to mature at 10.2 cm ICW although at 9.7 cm ICW, 50% of females had disengaged abdomens. Abdominal disengagement occurred in males at the slightly smaller size of 9.1 cm ICW. Allometric relationships between chela height and carapace width suggested 50% of males acquire mature chelae at 10.2 cm ICW. These results demonstrate the close linkage between early life stages of S. paramamosain and certain specific niches within mangrove habitats, with the main adult population found to be living sub-tidally at some distance from the mouth of the estuary. The study also highlights the special importance of the mangrove fringe, the border between the mangrove forest and the sea, an area which is particularly vulnerable to physical and anthropogenic impacts.     

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.     

Activity pattern and spatial strategy in Pachygrapsus marmoratus (Decapoda: Grapsidae) from Mediterranean and Atlantic shores

The spatial strategies and activity patterns of two populations of Pachygrapsus marmoratus (Fabricius), from Italy and Portugal, were studied in the field. The Mediterranean site was a non-tidal splash zone about 30 cm wide, while the Atlantic site was a 30 m wide intertidal belt. Almost all P. marmoratus of both populations were active on the emerged rocks, rarely entering the sea. The Mediterranean crabs actively fed on the algal turf throughout the 24 h, while in Portugal active crabs were more numerous at night. In the Italian population, all crabs were packed together, although large males had larger activity areas than females and smaller males. In Portugal, the large males were more concentrated in the sublittoral fringe, while both small males and females were confined to the eulittoral and littoral fringe. The Italian P. marmoratus performed feeding loops within areas smaller than 9 m². Although most Portuguese crabs showed the same spatial strategy, some of them performed feeding migrations to the sublittoral fringe, covering distances of up to 20 m. Environmental factors (e.g. temperature), physical factors (e.g. refuge availability) and biotic factors (e.g. competitors and predators) of the different shores probably determine the behavioural plasticity of␣P.␣marmoratus. Received: 1 February 1998 / Accepted: 13 November 1998     

Adaptive droving behavior observed in the fiddler crab Uca vocans vocans

Droving is conspicuous in Uca vocans vocans in summer. The crabs burrowing on the upper habitat have a higher tendency to wander compared to the ones burrowing on the lower habitat. Most of the wandering crabs captured on the low tide levels are relatively large and male. Larger crabs and males prefer to burrow on the upper zones of U. vocans habitat, but the smaller ones and females prefer the lower habitat. The upper level of the U. vocans habitat has relatively low N-content compared to the water's edge. Therefore droving is advantageous in crabs that have burrows on the upper level. Female and smaller resident crabs have faster feeding motions than male and larger ones, and can satisfy their feeding demands more rapidly. Therefore, for males and larger crabs it is advantageous to move away from the burrow area and forage in areas of higher food content near the low tide level where the number of feeding motions increases.     

Spontaneous colour preferences and colour learning in the fruit-feeding butterfly,Mycalesis mineus

Behavioral Ecology and Sociobiology - In sand fiddler crabs, Uca pugilator, contests between residents and intruders for breeding burrows often escalate to forceful pinching. In these contests, we...     

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.     

岷江上游大沟流域驱动植被退化的人为干扰体研究

研究了岷江上游大沟流域内的人为干扰体,分析了干扰的类型、强度、频率及其时空变化特征,揭示了不同干扰类型及其特性对植被退化的作用、干扰体与人口增长、植被退化等的关系．结果表明：大沟流域人为干扰类型多达８种,但优势的干扰类型是砍伐薪材和牲畜放牧．从近３０ａ的干扰强度变化看,以８０年代初的干扰强度最大,并在近２０ａ内保持很高的干扰强度水平;干扰强度的年内变化表现为春末、夏、秋大而冬、早春小．近４０ａ来的干扰频率愈趋增高,年内表现为春秋季干扰频率最高,夏季中等,而冬季最小．人为干扰的空间差异十分明显,反映出人为干扰的现状及其对植被的破坏强度差异,即由村寨向沟尾和高海拔区、由近及远、由低到高,形成“钟形”干扰格局．认为大沟流域植被的严重退化是所有人为干扰长期共同驱动的结果,与该区人口及其需求迅速增长密切相关．结果还表明,不同性质的干扰对植被的影响作用具协同性,同一干扰体长期连续对植被的影响具累加和放大的作用     

Population dynamics of the commensal spider crab Inachus phalangium (Decapoda: Maiidae)

The ecology of the spider crab Inachus phalangium (Fabricius, 1775) (Decapoda: Maiidae) was studied in the field. I. phalangium inhabits the sublittoral on the sea anemone Anemonia sulcata Pennant. From July 1981 to April 1984 in the Mediterranean (Banyuls sur Mer, southern France) more than 3000 anemones were examined and ca 1800 I. phalangium were found on them. The population dynamics' generation cycles, reproductive activities and the dynamics of the sex ratio were investigated. The density of juveniles (crabs before the pubertal moult) on anemones changed in a yearly cycle from low in the first six months to very high in the second six months. The first occasional young crabs of a generation appeared in March/April (3rd and 4th decapodite stages) on the anemones. Their density increased enormously in the following months. The generation grew gradually on the anemones and moulted into puberty in September-January. Density of adults (crabs after the pubertal moult) on anemones changed in a yearly cycle from low to high from the summer to winter months. A new adult generation was recruited every autumn through the pubertal moult and disappeared in the following summer. Female reproductive activity continued throughout the year. Females carried several broods in succession, but the frequency of breeding females fluctuated on a yearly cycle. The highest percentage of egg-carrying females, i.e. the peak of the reproductive season, lay in the first half of the year. The maximal life span of a generation, from the hatching of the first larvae to the disappearance of the last adults, lasted 1.5 to 2 years. Males moulted into puberty ca one month later than females. The moulting distribution of adult males had roughly the same course as in females. An adult male generation diet out about one to two months before the female generation. Life expectancy was therefore 14 to 17 months for females and 12 to 15 months for males. The sex ratio of juveniles shortly before the pubertal moult was balanced. The sex ratio of adults shifted from 1:1 at the beginning of the reproductive period to ca 1:9 in favour of females at its end.     

Observations on the burrows and burrowing behaviour of two mud-dwelling decapod crustaceans,Nephrops norvegicus and Goneplax rhomboides

The burrows of the Norway lobster Nephrops norvegicus (L.) and of the crab Goneplax rhomboides (L.) were studied in Loch Torridon, Scotland. Polyester resin casts of burrows in the sea were made by divers to reveal their subsurface form. Tunnels made by N. norvegicus were usually simple, with two or more openings on the mud surface, and penetrated to a depth of about 30 cm. G. rhomboides burrows did not descend more than about 15 cm beneath the surface, but were usually more complex than the lobster burrows and had several openings. The methods of burrow construction used by the two crustaceans are described from aquarium observations. Neither N. norvegicus nor G. rhomboides show obvious morphological adaptations for burrowing, and it is suggested that the fossorial habit was adopted very early by decapods. The burrows of N. norvegicus do not seem to have assumed any functions in addition to the original one of providing refuge from predators. There is not sufficient known of the biology of the crab to indicate whether the same is true in its case.     

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     

Behaviour and distribution of the sand-beach amphipod Orchestoidea corniculata

The field distribution of Orchestoidea corniculata Stont a talitrid amphipod inhabiting the upper intertidal zone of sandy beaches, was measured by pitfall trap and subsurface core transects. Variations in the distribution of juveniles and adults were recorded. At Coal Oil Point (Goleta, California, USA), the population was concentrated on the leeward side of the Point and, with increasing distance from the Point, both the number and size of beachhoppers decreased. The amphipods burrowed in a 5 to 7 m wide band during the day, and were active over a 15 to 25 m area at night. When they emerged from their burrows, a net seaward movement of 3 m was observed for the population. Behavioural tests suggested that sand penetrability is an important factor determining the location of burrow sites. Diurnal orientation was investigated, and it was found that the amphipods oriented in a landward direction despite overcast sky conditions, beach slope, moisture gradients, or displacement to a new beach.     

Behavioural ecology of the mud-burrowing crab Goneplax rhomboides

Diving studies at Fishguard harbour in South Wales (UK) combined with laboratory studies, provided information on the burrowing behaviour of Goneplax rhomboides (L.). Polyester resin casts indicated that burrows had from 1 to 6 openings and tunnels were horizontal at about 10 cm depth, with branches usually at right angles to each other. Burrowing was studied in the laboratory and related to variations in burrow casts taken in the field. Preliminary laboratory experiments suggest that social interactions occur between crabs, but analysis of the spatial distribution of burrows in the field suggested randomness. Burrows in the field showed some similarity of orientation, although the reasons for this were uncertain.     

Significance of burrow-opening diameter as a flood-prevention mechanism for air-filled burrows of small intertidal arthropods

Small air-breathing terrestrial arthropods (e.g. beetles, ants, crabs) living within the intertidal zone of mud and sand flats construct burrows with a narrow opening to the substrate surface. Despite frequently remaining open, these burrows are not flooded by the incoming tide. We examine, from a theoretical standpoint, the rôle of burrow-opening diameter in preventing burrow flooding. We determine a critical burrow diameter of 3 mm above which burrows are likely to flood unless the entrance is plugged by the burrow's inhabitant(s) prior to tidal inundation. We examine water penetration via the burrow entrance and seepage through the burrow walls. A practical mathematical solution is provided which can be used to examine the extent to which water may penetrate a given animal burrow in the field.     

Feeding ecology of the loggerhead sea turtle Caretta caretta in the Northwestern Gulf of Mexico

Digestive tract contents collected from carcasses of 82 loggerhead sea turles (Caretta caretta) found on the south Texas coast from 1986 through 1988 were examined. Benthic invertebrates were the predominant prey. Sea pens (Virgularia presbytes), crabs, and mollusks accounted for 94% of the dry weight of the digestive tract samples. Temporal changes in the percent occurrence and percent dry weight of sea pens, crabs, and mollusks in the digestive tract samples were significant. Loggerheads fed primarily on sea pens during spring, then primarily on crabs during summer and fall. The increase of crabs in the loggerhead diet paralleled the annual increase in the abundance of crabs in the Gulf of Mexico. Sea pens were located nearshore at depths of 6 to 12 m and had a disjunct distribution. Present address: Texas A&M University, Department of Biology, College Station, Texas 77843-3258, USA     

Temperature tolerance of the estuarine prawn Upogebia africana (Anomura,Crustacea)

Continuous temperature measurements were made in a typical South East African estuary. Mean summer (November to March) temperatures were in the range 19° to 24°C, and in winter (June to August) from 13° to 16°C. Large daily temperature fluctuations of 6° to 10°C occurred in summer; these appear to result from tidal movement of cool sea water into the estuary. In winter, temperature fluctuations were much smaller (3° to 5°C). The burrowing prawn Upogebia africana (Obtmann) was found to have an upper lethal temperature of 29°C in both winter and summer. The resistance time of prawns to temperatures above 30°C was much greater in summer than in winter. It was possible to acclimate winter prawns and increase their resistance time to a level comparable to that of summer individuals. A latent period of 40 h occurred before acclimation effects were detectable. Long-term exposure of prawns to high temperatures did not increase their resistance above that of summer prawns. Water at a temperature above this upper lethal temperature is not pumped through the burrows. This avoidance behaviour considerably increases the ability of U. africana to withstand short-lived temperature extremes.     

Mate competition in the velvet swimming crab Necora puber: effects of perceived resource value on male agonistic behaviour

Male velvet swimming crabs [Necora puber (L.)] were observed to engage in long and vigorous agonistic interactions to gain possession of a sexually receptive female. The role of agonistic behaviour in male mate competition in this species was examined by investigating the effect of the perceived presence of sexually receptive females on interactions between males. Exposing male crabs to water conditioned by sexually receptive females resulted in prolonged interactions, with a greater incidence of potentially injurious behaviour than interactions between control crabs, exposed only to sea water. Maleconditioned water also resulted in more offensive behaviour, but these interactions were of shorter duration and not significantly different from sea water controls. Agonistic superiority was strongly correlated with contestant relative size when males were exposed to male-conditioned water or sea water, but not when exposed to female-conditioned water. Overall, the results conformed with the general predictions of game theoretic models in relation to the influence of resource value on agonistic behaviour. Crabs were more persistent and probably incurred greater fitness costs in the perceived presence of a sexually receptive female, when interactions may have been resolved on the basis of factors more closely related to actual fighting ability than the relative body sizes of contestants.