Effect of acanthocephalan parasites on the behaviour and coloration of the mud crab Macrophthalmus hirtipes (Brachyura: Ocypodidae)

In the field, the numbers of cystacanths of the parasitic acanthocephalan Profilicollis spp. harboured by crabs are relatively high and correlate with carapace width. In a field experiment, the responses of crabs to the simulated approach of a bird predator (the parasite's definitive host) was not influenced by the number of acanthocephalans they harboured. Crabs that were exposed at the surface of the sediments during receding high tide, however, tended to harbour more parasites than nearby crabs hidden in burrows. An analysis of colour patterns on the carapace of crabs showed that infection levels did not influence carapace pigmentation, and thus did not affect the conspicuousness of a crab relative to the background environment. However, the likelihood of a male crab winning a ritualized fight against a conspecific in the field was associated with its infection level, but in a way that suggests that this finding is a consequence of pathology rather than an adaptation of the parasite to increase its transmission rate. Although only weak evidence was found indicating that Profilicollis manipulates the behaviour or colour of its host to its own benefit, the high infection levels observed suggest that the crab population acts as a major reservoir for larval stages of this parasite that are infective to birds.     

Mound building in Ilyoplax pingi (Crustacea: Brachyura: Ocypodidae)

The ocypodid crab Ilyoplax pingi, observed in Kanghwa I., Korea in 1992, builds a mound at the burrow entrance, by piling mud dug out from the burrow. The mounds were made by both waving and non-waving males, and by both ovigerous and non-ovigerous females. The burrow diameter at the widest part tended to be larger in crabs with mounds than in crabs without mounds, whereas the burrow depth was not different between them. The mound density increased in the late exposure period, when the crabs' surface activities declined. Mound removal and rebuilding experiments revealed that the presence of the mounds has the effect of keeping neighboring crabs away.     

Neighbor burrow-plugging in Ilyoplax pusillus (Crustacea: Brachyura: Ocypodidae)

The burrow living ocypodid crab Ilyoplax pusillus (De Haan, 1835) sometimes plugs the burrow of neighbors situated from 1.0 to 8.3 cm away with surface mud, while the neighbor, termed the pluggee, is in the burrow. Most pluggers were large males, whereas pluggees were smaller than pluggers and had a sex ratio close to 1:1. After being plugged, most pluggees usually reemerged on the surface within 5 min, but occasionally took up to 77 min to do so. The plugger always foraged or performed waving displays around the burrow of the pluggee while the pluggee stayed inside the burrow. Although the activity site of the pluggee was originally directed toward the burrow of the plugger or the adjacent area, after reemergence its activity site was oriented toward another direction. From these observations, it is suggested that neighbor burrow-plugging is a behavior adopted mainly by large males as a means of maintaining the area of their surface activities against smaller neighbors. My observations were made from June 1984 to June 1985 at Fukuro River Estuary, central Japan.     

Life-history patterns in two forms of Macrophthalmus japonicus (Crustacea: Brachyura)

Two behaviorally distinct forms — courtship signal — (Form L and V) of Macrophthalmus japonicus (De Haan) are known. Monthly samples of each form were collected from two areas in Japan: Form L from Amakusa, western Kyushu, between April 1985 and February 1986; Form V from Fukuoka, northern Kyushu, between November 1982 and December 1983. Life-history traits were compared between two populations and discussed. Individuals from the Amakusa population grew more slowly and matured at a smaller size than those from the Fukuoka population. Maturation period and longevity did not differ significantly. In the first year of reproduction, females in the Fukuoka population produced more, and larger broods than those in the Amakusa population and crab mortality was higher in every age group. Egg size did not differ significantly. The major environmental factors — air-temperature, nitrogen content of the surface mud and period of aerial exposure were similar in both areas, and it is suggested that the differences in life-history exhibited by these crabs are genotypic, and perhaps related to their different form.     

Factors affecting the burrowing behaviour of Helice tridens (Grapsidae) and Macrophthalmus japonicus (Ocypodidae) in an estuary of northeast Japan

Helice tridens (De Haan) (Grapsidae) and Macrophthalmus japonicus (De Haan) (Ocypodidae) build separate burrows in reed marshes and muddy tidal flats, respectively, in the brackish-water estuaries of northeast Japan (38°11N, 141°48E). Habitat segregation between burrows of these two species was analysed by comparing the density of burrows of both species in their natural habitats, and on tidal flats subjected to various types of artificial treatment, in the summer of 1981 and 1982. Baskets, containing many stones, were placed on the ground in the reed marsh, tidal flat and creek and attracted many individuals of H. tridens, but not M. japonicus. When stones were placed on an area of tidal flat, H. tridens frequently formed burrows at the border between the stones and mud, suggesting that burrowing of H. tridens was related to the presence of solid substances lying on the ground, such as stones and shoots of the reed Phragmites australis (Cav.) Trin. In closed systems on the tidal flat and reed marsh, H. tridens and M. japonicus were able to construct burrows in both substrata regardless of high or low frequency. Moreover, it was recognized that H. tridens prevented the burrowing of M. japonicus in these closed systems. These results suggested that habitat segregation between H. tridens and M. japonicus burrows was caused primarily by an exclusive interaction between individuals of these two species.     

Feeding and mouthpart morphology in the semaphore crabHeloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

The mouthparts of the Australian semaphore crabHeloecius cordiformis (H. Milne-Edwards, 1837) are described. Male and female semaphore crabs were observed and collected from mangrove swamps around the south side of Botany Bay, Australia between 1983 and 1987. Semaphore crabs inhabit mangrove mud/sand flats between the midlittoral intertidal zone and the supralittoral fringe of bordering saltflats. Scanning electron micrographs provide detailed pictures of mouthpart morphology and the setae present on the third, second, and first maxillipeds; the second and first maxillae; and the mandibles. Amongst other setal types, spoon-tipped, corn, and bottle-brush setae are described. While semaphore crabs are primarily deposit feeders (microphagous), they have well developed mandibles and are, therefore, also able to eat larger plant and animal material (macrophagous). A brief comparison is made between the mouthparts of semaphore crabs and fiddler crabs (Uca spp.), since the two genera are closely related, occupy similar niches, and resemble each other in appearance. This comparison reveals that while the second maxilliped setal morphology differs between the two genera (Heloecius possesses corn setae,Uca possesses spoontipped setae) these setae may have similar functions. The feeding behaviour ofH. cordiformis is related to substrate composition, burrow construction and various mouthpart specialisations. It is conlcuded that, within the context of the present study, certain structural e.g. mouthpart modifications allowing both scavenging and deposit feeding, and behavioural modifications e.g. mound building in wet muddy habitats, enable semaphore crabs to occupy a wide range of habitat types within the mangrove.Please address all correspondence and requests for reprints to Dr Maitland at his present address at the University of the Witwatersrand     

Carapace and branchial water circulation,and water-related behaviours in the semaphore crabHeloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

Water uptake, branchial water circulation and various water-related behaviours and anatomical specialisations, are described for the Australian semaphore crabHeloecius cordiformis (H. Milne-Edwards 1837). Male and female semaphore crabs were observed and collected from mangrove swamps around the south side of Botany Bay, Australia between 1983 and 1987. Semaphore crabs are active at low tide in air. Their branchial chambers are modified for both air breathing and water circulation. Water is held within the branchial chambers and is used for feeding and, to a certain extent, respiration. Crabs carry between 0.35 and 0.5 ml of water (7 g live body wt). Branchial water is continuously pumped out by the scaphognathites over specialised setae-bearing regions of the carapace and back through the branchial chambers over the gills, in a manner similar to that found in many grapsid crabs. Setae associated with water circulation are described. Water falling ventrally is collected by the abdominosternal cavity (formed between the abdomen and sternum) from where it returns to the branchial chambers via branchiosternal canals. Several novel water-related behaviours are described: Water shoogling — branchial water is drawn out between the walking legs where it is worked by scissor-like up and down movements of the legs. Abdomen flap — branchial water volume is adjusted by the extension and flexion of the abdomen against the cephalothorax. Water dumping — by tilting forwards and kissing the substrate, branchial water is voided to allow an entirely fresh load to be taken up. Statuesque posing — crabs remain erect and motionless for long periods. Branchial water is absent and external body surfaces dry out. Posing behaviour may function in thermoregulation, osmoregulation or as an anti-fouling mechanism against settling organisms. The significance of these various behaviours is discussed.     

Mating behaviour and reproductive cycle of Archaster typicus (Echinodermata: Asteroidea)

This study examines the reproductive cycle, the mechanism of male-on-female pairing behavior and the spawning behavior of Archaster typicus Müller et Troschel. Field studies were conducted in the intertidal zone of the sand beaches at Penghu, Taiwan (23 32N; 119 33E) at ebb-tide in 1984 and 1985. The pairing behavior of A. typicus is a reproductive behavior which leads to simultaneous spawning, increasing the probability of fertilization. As the breeding season approaches, sea stars, especially males, display increased mobility. Because only males tend to mount another individual and because males can detect the sex of another individual by contact with the side of their arms, a male-on-female pair is formed when a male encounters a female. Eighty-five percent of the sea stars observed were paired during the pairing season. Spawning by a paired female is closely followed by spawning of its paired male; male spawning, however, does not induce spawning in its paired female. During spawning, the male turns slightly so that its arms overlap the arched arms of the female. The gonad volume of males is much less than that of females. This may result from the high efficiency of fertilization in this species, which does not require a large amount of gametes to be released, or from the higher energetic demands made on the males.     

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.     

Hatching rhythms in Uca pugilator (Decapoda:Brachyura)

Gravid female Uca pugilator (Bosc) maintained in conditions of 14h L: 10h D light regime and constant temperature without simulated tides showed a significant positive correlation between time of hatch and time of the nighttime high tide. Light: Dark experiments showed no entrainment by LD cues. Larval hatching was delayed until dark when high tide occurred between 1900 and 2130 hrs. Hatch time was a positive linear function of the time of the nighttime high tide when high tide occurred after 2130 hrs. Time of hatch was delayed by the light period on days lacking a nighttime high tide. Experiments conducted under constant light (LL) and temperature, without simulated tides, indicated a persistent free-running rhythm of approximately 25 h 03 min, suggesting an endogenous timer. Rhythmic larval release on the nighttime high tide may be adaptive as a predator-avoidance mechanism for adult gravid females.Contribution No. 366 from the Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina     

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.     

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.     

The effect of food availability on the semilunar courtship rhythm in the fiddler crab<Emphasis Type="Italic"> Uca lactea</Emphasis> (de Haan) (Brachyura: Ocypodidae)

Semilunar or lunar reproductive rhythms that follow tidal cycles are common in marine animals. For brachyuran crabs, an adaptive explanation for the synchronous release of larvae in phase with the tidal cycle is that females release larvae when their offspring are best able to escape predators. As a corollary to this hypothesis, the synchronous male reproductive cycle is selected to be in phase with female receptivity. As an alternative hypothesis, we propose that variation in food abundance influences the onset and intensity of the semilunar courtship cycle. We tested this hypothesis in male fiddler crabs ( Uca lactea) by experimentally manipulating food availability for 4 weeks. Food-supplemented males built more semidomes and waved for more days than did food-deprived males or untreated control males. Moreover, food-supplemented males began courtship earlier and the median courtship day advanced with an increase in food. Courtship intensity was not related to crab size. These results provide the first evidence in marine animals that changes in food abundance due to the tides, and hence in phase with the semilunar cycle, influence male reproductive rhythm.     

Mating behaviour of the tiger blue butterfly (Tarucus theophrastus): competitive mate-searching when not all females are captured

Summary Females of the tiger blue butterfly (Tarucus theophrastus) fly upwind in search of the larval hostplant. Males perch or patrol the downwind edges, searching for incoming females or those already on the bush. A model of competitive mate-searching is developed for the case where not all receptive females are contacted by searching males: the model differs from the earlier ones of Parker (1970, 1974), particularly when few males are involved. Observed distributions of males upon bushes of different sizes agree better with the predictions of the game theory model than with a random distribution of males. The described model may be widely applicable to populations where females remate, but frequently evade detection by searching males.     

Mating behavior of Centropages typicus (Copepoda: Calanoida)

Observations were made on approximately 300 mating couples of Centropages typicus Krøyer. Mating behavior consists of 4 distinct stages: pre-copulation, spermatophore extrusion, copulation, and post-copulation. Successful copulation requires proper positioning of the spermatophore complex on the female urosome and is dependent upon precise movements and orientation of the male during the mating encounter. Sub-optimal mating encounters, resulting in the improper placement of the spermatophore complex, were observed and adaptations tending to reduce spermatophore wastage are discussed.Contribution No. 62, Harbor Branch Foundation, Inc.     

Mating behavior of Labidocera aestiva (Copepoda: Calanoida)

The calanoid copepod Labidocera aestiva Wheeler exhibits a ritualized mating behavior. Sensory hairs and spines and integumental pores on the male and female are believed to play a regulatory role in the mating encounter. Scanning electron microscopy (SEM) revealed approximately 20 unusual pore structures (pit-pores) on the right ventrolateral surface of the female's abdominal segment, the area where the spermatophore coupling device is attached. Prior to spermatophore transfer, the male vigorously strokes this area with the modified endopod of his left fifth leg to prepare the pit-pore region for attachment of a new spermatophore. Transmission electron microscopy (TEM) indicated that the cells underlying each pit-pore may have a secretory function. We hypothesize that the pit-pores secrete a substance which dissolves the cement binding the coupling device to the abdomen and aids the female to remove a discharged spermatophore from her urosome.Contribution No. 116, Harbor Branch Foundation, Inc.     

Larval development of Pilumnoides perlatus (Brachyura: Xanthidae) under laboratory conditions

Larvae of the xanthid crab Pilumnoides perlatus (Poeppig, 1836) have been reared in the laboratory at 3 different temperatures (10.2°, 15° and 20°C) from hatching to megalopa stage. The 5 zoea stages and the megalopa, as well as the setation of the functional appendages are described and illustrated. The main characteristics useful to differentiate the larval stages of P. perlatus from those of Homalaspis plana, the other Chilean species of the same family so far reared, are discussed. Data on duration of zoea development, length of moulting intervals, and mortality at the 3 test temperatures are also given.This study was partially supported by the Chilean National Commission for Scientific and Technological Research (CONICYT).     

Larval development of Ocypode quadrata (Brachyura: Crustacea) under laboratory conditions

The larvae of Ocypode quadrata (Fabricius) have been reared in the laboratory, from hatching to megalopa stage, at 35 S, 25°C. The five zoeal stages and the megalopa are described, including functional appendages of each stage. On the basis of morphological characteristics, the first zoeal and megalopa stages of O. quadrata can be distinguished from similar stages of closely related Ocypodinae. At 25°C, the megalopa appeared in a minimum of 34 days following hatching.