Sex, size and colour in a semi-terrestrial crab, Heloecius cordiformis (H. Milne Edwards, 1837)

We investigated the relationship between sex, size and colour in the little studied Australian endemic semaphore crab, Heloecius cordiformis, and related it to the crabs' social system with the aim of identifying the potential signalling function of claw colour.Equal sampling of crabs from all size classes revealed a strong relationship between sex, size and claw colour. Purple-clawed males were larger and had larger claws than pink-, orange- or green-clawed males. Male claws showed positive allometric growth: relative to body size, purple-clawed males had larger claws. The largest females had pink claws; the few with purple claws were no larger than immature green-clawed crabs. Female claws grow isometrically with the body so the relative claw size did not differ among the female colour classes. Quantitative measurements of claw colour revealed spectral differences between these subjectively described colours. The purple claws typical of large males also contrasted more strongly against the mudflat background than the other colours.Heloecius copulate outside female-owned burrows and probably within male-owned burrows. The male's waving display, in which both claws are raised and lowered, may feature in both mating strategies: as a territorial display and to attract wandering females. Large males are competitively superior so size, and potentially colour, are important in territorial disputes and may also feature in mate choice.     

Feeding Habits of Percnon gibbesi (H. Milne Edwards, 1853) in the Sicily Strait

The subtropical grapsid crab Percnon gibbesi (H. Milne Edwards, 1853) is one of the most recent alien decapods found in the Mediterranean, where it was discovered at Linosa (Pelagie Islands, Sicily Strait) in summer 1999. At present, the invasion of this species has been recorded in several other insular localities. We studied the feeding habits of the crab in an attempt to understand its success in the Mediterranean. The morphological characteristics of the chela, the feeding adaptations of the gastric mill and the analysis of stomach contents indicate that P. gibbesi is a strictly herbivorous species, a characteristic not shared with any other large-sized infralittoral Mediterranean crab.     

Reproductive biology of the Suez Canal spider crab Schizophrys aspera (H. Milne Edwards, 1834: Crustacea: Brachyura: Majidae)

A reproductive biology study of the spider crab Schizophrys aspera (H. Milne Edwards, 1834) was conducted in the Suez Canal from July 2012 to June 2013. The annual sex ratio (Male:Female) of S. aspera was female biased with values of 1:1.25. Out of the four ovarian development stages of this crab, two stages were observed in the Suez Canal throughout the whole year. The ovigerous crab’s carapace width varied from 28 to 52 mm. This crab species can spawn during most of the year in the canal water, with a peak during late spring and early winter. The fecundity of ovigerous females ranged between 2349 and 13600 eggs with a mean of 5494 ± 1486 eggs. Female crabs that reached sexual maturity exhibited a minimum carapace width varying between 22 and 46 mm, and fifty percentage of all ovigerous females showed a carapace width of 36 mm.     

A phylogenetic study of the Galappidae H. Milne Edwards 1837 (Crustacea: Brachyura) with a reappraisal of the status of the family

The systematic status of the family Calappidae and the phylogenetic relationships of its four component subfamilies are re-evaluated based on a cladistic analysis of 78 adult morphological characters. A single tree was produced (CI = 0.654). The monophyly of the Calappidae sensu lato is rejected. The data suggest that the Calappinae and Hepatinae form a single lineage which is closer to some xanthids than to the Matutinae or Orithyiinae. A close link between the Matutinae and some leucosiids and between the Orithyiinae. and some dorippids is also apparent, with a suggestion that these four taxa all belong to a single lineage. A revised classification of the Oxystomata emend , and Calappidae is proposed.     

Competing for last place: Mating behaviour in a pill-box crab, Halicarcinus cookii (Brachyura: Hymenosomatidae)

The mating strategy of Halicarcinus cookii was investigated to ascertain how males maximised their fitness through mate choice. An intertidal population at Kaikoura, New Zealand, was dominated by mature crabs of both sexes in summer and by immature crabs in the colder months. More than 95% of mature females were ovigerous with early stage and late stage broods found in almost every month, indicating that egg production and larval release is continuous. The operational sex ratio was less than 1 male/female in summer, but often more than 1.0 in the colder months. The gonosomatic index increased along with brood development so that as soon as zoeae were released, the next clutch of eggs was ready to be fertilised. Males searched for receptive females and began pre-copulatory mate guarding without any courtship display. They mated preferentially with late stage or non-ovigerous females: copulation duration was longest for stage 5 females as was post-copulatory guarding (mean 18.3 h). Late stage females were up to 14% of the female population. Mate attraction seems to be the result of an ovarian signal rather than from the developing brood. Manipulation of the sex ratio had effects upon copulation duration and post-copulatory guarding: presence of a rival male increased duration of guarding. Females showed precocious mating in the penultimate instar and were able to lay fertilised eggs after their pubertal moult in the absence of males. H. cookii females have many mates, but males attempt to ensure paternity by preferentially pursuing mature females close to egg laying and by guarding these females after copulation. These behaviours are all elements of a competitive strategy to ensure that a male loses (not wins) the race to copulate because females have a ventral seminal receptacle, giving sperm precedence to the last male to mate. Male mating behaviour is a consequence and evolutionary response to female morphology.     

Agonistic interactions between invasive green crabs, Carcinus maenas (Linnaeus), and sub-adult American lobsters, Homarus americanus (Milne Edwards)

The invasive green crab, Carcinus maenas, has recently expanded its range into the Southern Gulf of St. Lawrence, where there is potential for substantial niche overlap with juvenile American lobsters, Homarus americanus. We used two experiments to elicit, record and analyze the agonistic interactions of adult green crabs (carapace width of 63-75 mm) and sub-adult (carapace length of 55-70 mm) lobsters. The first experiment gave each animal equal access to a limited food resource. The green crabs were first to the food in significantly more trials, spent a significantly greater proportion of time with the food, and were able to successfully defend the food from attacks by the heavier lobsters. In the second experiment, we allowed the lobsters to gain possession and initiate feeding on the food before releasing the green crabs. In these trials, the lobsters spent significantly more time with the food, and were able to defend the food from the green crabs. The results of both experiments are discussed in the context of game theory. The different behaviour of the crustaceans in the two experiments is consistent with the “bourgeois” strategy in a hawk and dove game simulation. With this strategy, an animal acts like a hawk if in possession of a resource, but acts like a dove if the other animal is in possession of the resource. The fact that the green crabs were able to physically compete with, and in many cases dominate the larger, heavier lobsters supports the potential for competitive impacts of green crabs on sub-adult lobsters.     

Reproductive Aspects of a Tropical Population of the Fiddler Crab Uca annulipes (H. Milne Edwards, 1837) (Brachyura: Ocypodidae) at Costa Do Sol Mangrove, Maputo Bay, southern Mozambique

Uca annulipes(H. Milne Edwards) is the dominant fiddler crab inhabiting mangrove forests in East Africa. However, several aspects concerning its reproduction are poorly known. Such information is necessary for a better understanding of several biological aspects occurring in this species. The aim of this study was to assess the reproductive biology of Uca annulipes as well as some factors determining its breeding season. Samplings were undertaken from January to December 2002 at Costa do Sol, a tropical mangrove forest around Maputo Bay, southern Mozambique. Crabs were randomly sampled, measured for carapace width (CW) and checked for the presence of eggs on female pleopods. Like most tropical brachyurans, Uca annulipes follows a continuous reproduction throughout the year with peaks of spawning during summer season and low frequency of egg-bearing females during winter. The frequency of ovigerous females was correlated with the air temperature (r ²= 0.652, p < 0.05) and rainfall (r ²= 0.897, p < 0.001), with the last parameter being the most important factor regulating the reproductive activity of this crab.     

Laboratory experiments on the effects of variable suspended sediment concentrations on the ecophysiology of the porcelain crab Petrolisthes elongatus (Milne Edwards, 1837)

The porcelain crab Petrolisthes elongatus is a particulate suspension feeding species common to coastal areas of New Zealand (NZ). Consistent with the responses of other suspension feeding species, it is likely to be negatively influenced by elevated suspended sediment concentrations. Laboratory experiments were conducted to quantify the effect of temperature (12 °C, 15 °C and 18 °C) and suspended sediment concentration (total particulate matter (TPM): low < 100 mg L^− 1; medium 100-1000 mg L^− 1; high > 1000 mg L^− 1) on the clearance rate (CR in L h^− 1), oxygen uptake rate (VO₂ in mL h⁻¹), net absorption efficiency (AE), and net energy budget (NEB in J h^− 1) of P. elongatus across a range of sizes. Variation in CR and AE was independent of temperature and of body size, but were significantly different (P < 0.05) at low and medium suspended sediment concentrations compared with high suspended sediment concentrations. CR responded in a non-linear manner to changes in TPM, increasing with TPM up to a maximum value at medium-low concentrations (approximately 250 mg L^− 1) and then decreasing thereafter. CR had almost completely shut down at TPM concentrations of > 1000 mg L^− 1 and at particulate organic matter (POM) concentrations of > 250 mg L^− 1. AE was zero at approximate TPM and POM values of 1200 mg L^− 1 and 300 mg L^− 1, respectively. VO₂ was positively correlated with body size and with temperature, but was independent of TPM. NEB values for P. elongatus were low (approx 110 J g^− 1 h^− 1) at low sediment concentrations, were high (approx 320 J g^− 1 h^− 1) at medium sediment concentrations, and were negative (approx − 114 J g^− 1 h^− 1) at high sediment concentrations. These findings indicate that P. elongatus is likely to be food-limited at sediment concentrations of < 100 mg L^− 1, and severely negatively affected at sediment concentrations of > 1000 mg L^− 1, at least for the duration of such events which may persist for 2-3 days in coastal environments where this crab occurs.     

The potential for mitten crab Eriocheir sinensis H. Milne Edwards, 1853 (Crustacea: Brachyura) invasion of Pacific Northwest and Alaskan Estuaries

Eriocheir sinensis H. Milne Edwards, 1853 is on the list of top 100 invaders compiled by the International Union for Conservation of Nature and Natural Resources. The recent establishment of a large Chinese mitten crab population in San Francisco Bay and the potential for introductions from California, Asia and Europe pose a significant invasion potential for estuaries and rivers from California to Alaska. This alien species would place at risk the catchment areas of the Pacific Northwest including the economic and social activities that depend upon intact aquatic systems. An analysis of ecological conditions that define the mitten crab’s native and introduced range suggests that large stable estuaries with long flushing times are necessary to sustain significant populations. Most Pacific Northwest estuaries have limited salinity intrusion, estuarine habitat and short flushing times and face a reduced risk of population establishment. Large, stable estuaries, such as the Puget Sound, may support significant populations. River-dominated estuaries, such as the Columbia River, have flushing times less than the duration of larval development and wouldn’t support populations. An application of a temperature based larval development rate to near-shore and estuary sea surface temperatures suggests that estuaries in Oregon and Washington have sufficient thermal regimes to support larval development. Most estuary systems in Alaska have limited periods where water temperatures are above the mortality threshold for the larval stages and are at a low risk for the establishment of populations. A potential sea temperature rise of two degrees Celsius would permit larval development in Alaskan estuaries, where sufficient estuarine and freshwater habitats exist.     

Agonistic Behaviour and Development of Territoriality During Ontogeny of the Sea Anemone Dwelling Crab Allopetrolisthes Spinifrons (H. Milne Edwards, 1837) (Decapoda: Anomura: Porcellanidae)

Allopetrolisthes spinifrons is an ectosymbiotic crab of the sea anemone Phymactis clematis. As a consequence of low host abundance, these represent a scarce and limited resource for the crab. Additionally, the relatively small size of the sea anemone host suggests that few symbiotic crabs can cohabit on one host individual, forcing crabs to adopt a territorial behaviour. In order to examine the potential presence and ontogenetic development of territoriality, the agonistic behaviour between crabs of various ontogenetic stages (adults, juveniles, and recruits) was studied in the laboratory. Laboratory experiments demonstrated that adult or juvenile crabs aggressively defended their sea anemone hosts against adult or juvenile intruders, respectively, but both adult and juvenile crabs tolerated recruits. Adult crabs behaved indifferently towards juvenile crabs, sometimes tolerating them, sometimes expelling them. Recruits never showed agonistic behaviour among themselves. The agonistic interactions observed in the laboratory and the uniform population distribution pattern on sea anemones recently described for A. spinifrons indicate that this species exhibits territorial behaviour, which develops during ontogeny. Territoriality in this species and other symbiotic decapods may function as a density-dependent mechanism of population regulation, being mediated by the availability of hosts. Resource monopolisation behaviours may be common among other symbiotic and free-living marine invertebrates inhabiting discrete habitats that represent a limiting resource.     

Agonistic Behaviour and Development of Territoriality During Ontogeny of the Sea Anemone Dwelling Crab Allopetrolisthes Spinifrons (H. Milne Edwards, 1837) (Decapoda: Anomura: Porcellanidae)

Allopetrolisthes spinifrons is an ectosymbiotic crab of the sea anemone Phymactis clematis . As a consequence of low host abundance, these represent a scarce and limited resource for the crab. Additionally, the relatively small size of the sea anemone host suggests that few symbiotic crabs can cohabit on one host individual, forcing crabs to adopt a territorial behaviour. In order to examine the potential presence and ontogenetic development of territoriality, the agonistic behaviour between crabs of various ontogenetic stages (adults, juveniles, and recruits) was studied in the laboratory. Laboratory experiments demonstrated that adult or juvenile crabs aggressively defended their sea anemone hosts against adult or juvenile intruders, respectively, but both adult and juvenile crabs tolerated recruits. Adult crabs behaved indifferently towards juvenile crabs, sometimes tolerating them, sometimes expelling them. Recruits never showed agonistic behaviour among themselves. The agonistic interactions observed in the laboratory and the uniform population distribution pattern on sea anemones recently described for A . spinifrons indicate that this species exhibits territorial behaviour, which develops during ontogeny. Territoriality in this species and other symbiotic decapods may function as a density-dependent mechanism of population regulation, being mediated by the availability of hosts. Resource monopolisation behaviours may be common among other symbiotic and free-living marine invertebrates inhabiting discrete habitats that represent a limiting resource.     

Allopatric distribution of juvenile red-legged banana prawns (Penaeus indicus H. Milne Edwards, 1837) and juvenile white banana prawns (Penaeus merguiensis De Man, 1888), and inferred extensive migration, in the Joseph Bonaparte Gulf, northwest Australia

During October to December 1997, we trawled estuarine habitats in the Joseph Bonaparte Gulf (JBG) to determine the distribution of juvenile red-legged banana prawns, Penaeus indicus (H. Milne Edwards, 1837) and white banana prawns, Penaeus merguiensis (de Man, 1888). We made 229 beam-trawls at 185 sites, mostly over a 100-m path (3-min duration). A Global Positioning System (GPS) receiver was used to verify our location. During October to December 1998, we intensively resampled three of the rivers that were sampled in 1997 to confirm the gulf-wide distribution of P. indicus and P. merguiensis and to investigate the microhabitat use of P. indicus. We chose previously sampled and new sites in Forsyth Creek (eastern JBG), the Lyne River (Cambridge Gulf), and the Berkeley River (western JBG). We made 249 trawls at 21 sites, mostly over 100 m.Juvenile banana prawns were abundant in eastern JBG, Cambridge Gulf and western JBG. They were not abundant in southern JBG, although fewer trawls were made there, due to its inaccessibility. In eastern JBG and Cambridge Gulf, over 96% and 73% (respectively) of juvenile banana prawns were P. indicus and they were more abundant there than in the western JBG. Conversely, in the western JBG over 93% of the juvenile banana prawns were P. merguiensis and they were more abundant than in the eastern JBG and Cambridge Gulf. The Lyne River in the northwestern Cambridge Gulf seems to be the transition zone; both P. indicus and P. merguiensis are equally abundant.P. indicus are most abundant on the mangrove-lined muddy banks of waterways within mangrove forests, similar habitats to P. merguiensis. Within these habitats, they were most abundant in gutters and small creeks, rather than rivers and large creeks. Few P. indicus or P. merguiensis were caught in 100 m² trawls undertaken midriver (on the channel bottom and on emergent banks), although these habitats may be only 100 m from the mangrove-lined habitats. In all creek and river habitats, both species are most catchable at low tide (irrespective of daylight or darkness) when they move out of the mangrove forests and accumulate in the remnant water bodies.The offshore fishery for P. indicus is in northwestern JBG in waters 50-80 m deep, about 300 and 200 km, respectively, from where juveniles are abundant in their extensive inshore habitats in east JBG and in Cambridge Gulf, demonstrating a geographical separation of the juvenile and adult phases. Postlarval P. indicus, spawned offshore, must use tides and currents to travel south and east to reach nursery habitats. Emigrant subadults must migrate north and west, across relatively shallow inshore sand substrates (30-40 m deep) to reach their offshore habitats.     

Orientation and Shape Discrimination in Juveniles and Adults of the Mangrove Crab Aratus pisonii (H. Milne Edwards, 1837): Effect of Predator and Chemical Cues

Behavioral responses of juveniles and adults of the mangrove crab Aratus pisonii (H. Milne Edwards, 1837) to black geometric shapes of equal surface area was measured. Crabs were tested either in presence or absence of chemicals generated from two common predator species, the portunid crab Callinectes ornatus Ordway, and the soapfish Haemulon aurolineatum Cuvier, 1830. The present study tested the hypothesis that A. pisonii (1) has the capacity to orient to visual cues; (2) it discriminates between different visual objects based on a combination of chemical and visual information and (3) this behavior changes with age. When presented with single black targets in background water, juveniles oriented toward all shapes. This behavioral response was interpreted as visual orientation toward potential shelter. Among shapes, juveniles showed preference for the vertical rectangle, probably due to the recognition of natural visual elements like mangrove roots. In predator conditioned water, juveniles exhibited a stronger response than in background water. Thus, juveniles were able to detect by odor the potential presence of predators. Change in responsiveness between adults and juveniles was also demonstrated.     

Spermatophores, female genitalia, and courtship behaviour of two whip spider species, Charinus africanus and Damon tibialis (Chelicerata: Amblypygi)

This paper describes courtship behaviour, spermatophore morphology, and the female genitalia of the African whip spiders Charinus africanus Hansen, 1921 (Charinidae) and Damon tibialis (Simon, 1876) (Phrynichidae). In C. africanus, only the first part of courtship behaviour, up to spermatophore formation, could be observed; though different in detail, it is similar to that of many other species. The small spermatophore of C. africanus contains one large median sperm package. Charinus africanus is one of the Charinus species with thin finger-like gonopods and the first species with such gonopods of which the spermatophore is known. Spermatophores and female genitalia of D. tibialis are similar, though different in details, to those of Trichodamon and Musicodamon. They thus suggest that these two genera are correctly included in the Damoninae.     

Thorn fish Terapon jarbua (Forskål) predation on juvenile white shrimp Penaeus indicus H. Milne Edwards and brown shrimp Metapenaeus monoceros (Fabricius): the effect of turbidity, prey density, substrate type and pneumatophore density

A series of laboratory experiments was conducted at Inhaca Island Marine Biological Station, Mozambique, in order to assess the separate effects of turbidity, prey density, substrate type, pneumatophore density, and the combined effects of turbidity with the latter three, on rate of predation by the thorn fish Terapon jarbua (Forskål, 1775) on white shrimp Penaeus indicus and brown shrimp Metapenaeus monoceros.Significant interactions between turbidity and the other three factors on shrimp predation for both prey species were detected. Regardless of prey density, increasing turbidity decreased predation on P. indicus, but not on M. monoceros, for which increasing densities reduced the protective effect of turbidity. Increasing prey density increased predation on P. indicus in clear water, and increased predation on M. monoceros in low and high, but not in intermediate turbidity or clear water. The presence of a substrate suitable for burying decreased predation on M. monoceros in clear water, but not in the turbidity levels used. In clear water, solely sandy-shell substrate afforded protection to P. indicus, while in turbid water, no substrate offered significant protection and muddy substrate even increased prey vulnerability to fish probably as a result of increased preys' locomotor activity. Raising pneumatophores density seems to lower the protective value of turbidity for both species. In clear water, only low and high structure density provided a deterrent effect on predation on P. indicus; in turbid water, intermediate and higher structure density increased predation. Increasing structural complexity reduced predation on M. monoceros linearly in clear water; but in low turbid water it increased. In high turbid waters, the increase was only significant in intermediate pneumatophore density. High structural complexities impair the pursuing capacity of fish and thus decreased predation rates. The results indicate that the effective provision of shelter of different habitats depends not only on the various environmental parameters analysed, but also on the way they interact and on the behaviour of prey and predator as well.     

Flowers and floral biology of the holy grasses (Hierochloe and Anthoxanthum: Aveneae,Gramineae)

Trifloreted spikelets in species of Anthoxanthum and of Hierochloe are united by the syndrome of two lower male or neuter florets and a differentiated third floret, perfect, bistaminate, and protogynous. Differences in presentation of the floral organs from the third anthoecium separate the genera. Anthers in Anthoxanthum, longer than the f3 anthoecium, keep it open until they migrate on elongating filaments, following behind the protogynous stigma-styles. Both emerge near the apex of the upper glume which tightly encloses all three florets. In Hierochloe the uppermost anthoecium is closed over the androecium and gynoecium until lodicule-controlled anthesis when stigma-styles are exserted unhindered by glumes, and the anthers are exserted later on long filaments from the chasmogamous lower florets. In Hierochloe triandrous lower florets are constant. In Anthoxanthum the two lower florets are neuter in Eurasian species, but in species in Africa, Asia, and Malesia, sometimes f1 florets are also staminate, and very exceptionally both florets present stamens. These reflect a sexual variability in Anthoxanthum unknown in Hierochloe. Data from flowers of 30 species of Hierochloe and 17 species of Anthoxanthum are tabulated; included are ratios of f3 anthers:anthoecium and stigma-styles:the upper glume, G2, two ratios which most reliably reflect the intergeneric floral patterns. Pathways to those varied floral systems are outlined. A sectional taxonomy is proposed for Hierochloe: Sect. Hierochloe autonymum; Sect. nova Monoecia for species in South America where monoecism occurs in all taxa. Andromonoecism is the predominant sexual system in boreal and austral species of Hierochloe, but is interrupted by apomixis in northern Europe.     

The effects of sexual experience and female receptivity on courtship-rape decisions in male guppies, Poecilia reticulata (Pisces: Poeciliidae)

This paper examines the effects of prior sexual experience and of female receptivity on the probability that male guppies will court females with high or low intensity displays or attempt rape. There were no differences in behaviour between males with a history of high copulation success and those with low copulation success. Males with receptive females exhibited higher levels of high intensity displays and fewer rape attempts than males with unreceptive females, regardless of prior experience. The results are discussed in terms of sexual selection and the selective advantage of behavioural plasticity in which different sexual behaviour patterns are chosen with respect to the probability of a successful mating at a given time.     

Nectonema zealandica n. sp. (Nematomorpha: Nectonematoidea) parasitising the purple rock crab Hemigrapsus edwardsi (Brachyura: Decapoda) in New Zealand,with notes on the prevalence of infection and host defence reactions

A new species of Acusicola is described based on adults of both sexes taken from plankton samples collected in the upper reaches of the Piauí River estuary, in the northeast of Brazil. Ovigerous females were present in the plankton. The new species, Acusicola minuta n. sp., can be distinguished from its congeners by its small body size, female antennal morphology and leg setation. The male described here as A. minuta n. sp. is the first known male attributed to the genus.