Temporal variation in abundance of the egg predator Carcinonemertes epialti (Nemertea) and its effect on egg mortality of its host,the shore crab,Hemigrapsus oregonensis

An outbreak of the nemertean, Carcinonemertes epialti, was observed on Hemigrapsus oregonensis during October, 1982 at Campbell Cove, Bodega Harbor, California. Mean worm intensity (296 worms/crab) was the highest recorded for this nemertean egg predator on H. oregonensis. During the outbreak, male crabs were found to harbor more worms than both non-ovigerous and ovigerous females. Crab egg mortality was substantial; 83% of the ovigerous females experienced 75–100% brood mortality. The seasonal peak in worm abundance coincided with the seasonal low in crab reproduction at this locality. A method for estimating the impact of C. epialti on H. oregonensis natality was developed using crab size and fecundity, and worm prevalence and intensity. For a non-outbreak sampling period, a mean of 5.6% egg mortality was experienced by infested crabs for the period selected. Thus, brood mortality during the outbreak was much greater than that experienced at non-outbreak periods. Heavy fishing pressure on some commercially important crab species has been suggested as a possible factor inducing worm outbreaks and facilitating their continued persistence. These observations suggest that fisheries are not necessarily responsible for the outbreaks of nemerteans on commercially important crab species. However, fishing pressure may still be a sufficient condition to promote nemertean outbreaks.     

The invasive green crab and Japanese shore crab: behavioral interactions with a native crab species,the blue crab

Blue crabs, Callinectes sapidus (Rathbun), are an ecologically and commercially important species along the East coast of North America. Over the past century and a half, blue crabs have been exposed to an expanding set of exotic species, a few of which are potential competitors. To test for interactions with invasive crabs, juvenile C. sapidus males were placed in competition experiments for a food item with two common non-indigenous crabs, the green crab Carcinus maenas (L.) and the Japanese shore crab, Hemigrapsus sanguineus (De Haan). Agonistic interactions were evaluated when they occurred. In addition, each species’ potential to resist predators was examined by testing carapace strength. Results showed that C. maenas was a superior competitor to both C. sapidus and H. sanguineus for obtaining food, while the latter two species were evenly matched against each other. Regarding agonism, C. sapidus, was the “loser” a disproportionate number of times. C. sapidus carapaces also had a significantly lower breaking strength. These experiments suggest that both as a competitor, and as potential prey, juvenile blue crabs have some disadvantages compared with these common sympatric exotic crab species, and in areas where these exotics are common, juvenile native blue crabs may be forced to expend more energy in conflict that could be spent foraging, and may be forced away from prime food items toward less optimum prey.     

Native species vulnerability to introduced predators: testing an inducible defense and a refuge from predation

To manage the impacts of biological invasions, it is important to determine the mechanisms responsible for the effects invasive species have on native populations. When predation by an invader is the mechanism causing declines in a native population, protecting the native species will involve elucidating the factors that affect native vulnerability. To examine those factors, this study measured how a native species responded to an introduced predator, and whether the native response could result in a refuge from predation. Predation by the green crab, Carcinus maenas, has contributed to the decline in numbers of native soft-shell clams, Mya arenaria, and efforts to eradicate crabs have proven futile. We tested how crab foraging affected clam burrowing, and how depth in the sediment affected clam survival. Clams responded to crab foraging by burrowing deeper in the sediment. Clams at shallow depths were more vulnerable to predation by crabs. Results suggest soft-shell clam burrowing is an inducible defense in response to green crab predation because burrowing deeper results in a potential refuge from predation by crabs. For restoring the native clam populations, tents could exclude crabs and protect clams, but when tents must be removed, exposing the clams to cues from foraging crabs should induce the clams to burrow deeper and decrease vulnerability. In general, by exposing potential native prey to cues from introduced predators, we can test how the natives respond, identify whether the response results in a potential refuge, and evaluate the risks to native species survival in invaded communities.     

The relation between cypris ultrastructure and metamorphosis in male and female Sacculina carcini (Crustacea,Cirripedia)

Summary To elucidate current controversies on sex in rhizocephalan barnacles, broods of Sacculina carcini, infesting the shore crab Carcinus maenas, were raised to cyprids in the laboratory and followed through settlement and metamorphosis. Free-swimming cyprids were studied by transmission electron microscopy and occur in male and female morphological types, which differ in the structure of carapace cuticle, antennular cuticle, antennular glands, and the cells suspected of being the stem cells during metamorphosis. These dissimilarities are in addition to the already known differences in cypris size, in number of antennular sense organs, and in substrata settled on by morphological males and females. Metamorphosing males (trichogons) and females (kentrogons) are illustrated in interference phase-contrast micrographs. The morphological differences between male and female cyprids are directly related to their dissimilar metamorphosis. Hence, cyprids of male morphology are anatomically incapable of metamorphosing into kentrogons, while cyprids of female morphology cannot metamorphose into trichogons. The determination of sex in rhizocephalan barnacles is discussed.The results refute the hypothesis that sex in Sacculina carcini is determined environmentally, e.g., by the substratum encountered by the cyprids at settlement. It is concluded that sex is determined already in the free-swimming larvae and, most probably, already in the ovary. This agrees with the mode of sex determination in other species of the Rhizocephala Kentrogonida.     

Learning in an invasive and a native predatory crab

The invasion of the green crab Carcinus maenas in the northeastern U.S. and its competition with the native blue crab Callinectes sapidus and other native crustaceans has been well-documented and researched. Various reasons for the invader’s success against native crabs have been examined (juvenile predation, food source flexibility, etc.), but another possibility is a difference in the learning ability of invasive versus native crab species. In this study, the learning ability of C. maenas and C. sapidus was tested by their increased speed in locating hidden food over successive days. The data suggest that C. maenas possesses a learning ability significantly greater than that of C. sapidus, which may partially contribute to its success.     

Size and settling behaviour in male and female cypris larvae of the parasitic barnacle Sacculina carcini Thompson (Crustacea: Cirripedia: Rhizocephala)

Twenty-one broods from different externae of Sacculina carcini Thompson were cultured to the cypris stage. The size of the cyprids was measured and the larvae subjected to settling upon unparasitized crabs (Garcinus maenas (L.)) and small juvenile externae. The cyprids occur in two sizes that may appear singly or together in the same brood. Small cyprids are of the female sex that settle upon crabs and are infective, while large cyprids are of the male sex and only settle upon juvenile externae. These results are in agreement with other well-studied rhizocephalans.     

Exotic Spartina alterniflora provides compatible habitats for native estuarine crab Sesarma dehaani in the Yangtze River estuary

Although the impact of plant invasions on benthic communities, especially burrowing crabs, has received increasing attention, the results from past studies are mixed. The exotic plant Spartina alterniflora has become the most abundant species in the salt marshes of the Yangtze River estuary since it was first found just over a decade ago, but its effects on crabs in the salt marshes is largely unknown. To examine whether the invasions of this exotic plant affected native crabs, we compared the biomass and abundance of the dominant burrowing crab Sesarma dehaani in an exotic Spartina marsh, native Phragmites australis marsh and mudflats of the Yangtze River estuary, China. To explain the differences of S. dehaani populations between different habitats, feeding preference of S. dehaani for Spartina and Phragmites was investigated. Results showed crab abundance and biomass in the Spartina marsh were significantly greater than those in the Phragmites marsh and mudflats. Soil water content and plant community characteristics in the Spartina marsh also significantly differed in the Phragmites marsh and mudflats. Moreover, the feeding preference experiment showed that crabs consumed Spartina more than twice as much as Phragmites. In summary, this study showed that Spartina provided compatible habitats for native crab S. dehaani through offering suitable food source and moderate environmental conditions.     

Impact Scenario for an Introduced Decapod on Arctic Epibenthic Communities

The intentional introduction of a species for the enhancement of stock or establishment of new fisheries, often has unforeseen effects. The red king crabs, Paralithodes camtschaticus, which was introduced into the Barents Sea by Russian scientists, has established a self-sustaining population that has expanded into Norwegian waters. As top benthic predators, the introduced red king crabs may have possible effects upon native epifaunal scallop (Chlamys islandica) communities. These benthic communities may be a source of prey species in late spring, when the red king crabs feed most intensively. Foraging rates (consumption, killing or severely damaging) of red king crab on native prey organisms were measured by factorial manipulation of crab density (0.5, 1.5 and 3 per m ²), size classes (immature, small mature, and large mature crabs), and by evaluating prey consumption after 48 h, in order to extrapolate a scenario of the likely impacts. Foraging rates of the red king crab on scallops ranged between 150 and 335 g per m² within 48 h. These rates did not change when crab density was altered, though an increased amount of crushed scallops left uneaten at the tank floor, were correlated with high density of small mature crabs. Foraging rate changed significantly with crab size. Consequently, the susceptibility of native, shallow water epibenthic communities to red king crab predation in the early life history stages, and during the post-mating/molting spring period, must be considered significant when foraging rates are contrasted with natural scallop biomass between 400 and 1200 g scallops per m².     

Distribution and abundance of a nemertean egg predator ( Carcinonemertes sp.) on a leucosiid crab,Randallia ornata

The purple globe crab, Randallia ornata (Randall, 1839) (Decapoda: Leucosiidae) is a common crustacean found on sandy beaches from Northern California to Baja California, Mexico. An undescribed species of Carcinonemertes, which shares biological features with a partially described worm from Tasmania was recovered from R. ornata. The worm reaches 10 mm in length and secretes a sturdy tube with a distinctive spiral shape. The total prevalence of the worm on R. ornatawas 70% (85% for ovigerous female crabs, 67% for post-ovigerous female crabs, and 45% for male crabs). Intensities reached 32 worms per crab. Adult worms were found in crab egg masses, regressed adults were recovered from under the abdomen of post-ovigerous females, and encysted juveniles were found on the gill lamellae of non-ovigerous female crabs and occasionally from the gill lamellae of male crabs.     

Marine invasive species: validation of citizen science and implications for national monitoring networks

Approximately 1,000 volunteers assessed the presence of invasive (Carcinus maenas and Hemigrapsus sanguineus) and native crabs within the intertidal zone of seven coastal states of the US, from New Jersey to Maine. Identification of crab species and determination of the gender of the observed crabs was documented at all 52 sites across a 725-km coastal transect. Using quantitative measures of accuracy of data collected by citizen scientists, a significant predictor of a volunteer’s ability was determined and eligibility criteria were set. Students in grade three and seven had the ability to differentiate between species of crabs with over 80% and 95% accuracy, respectively. Determination of gender of the crabs was more challenging and accuracy exceeded 80% for seventh grade students, while 95% accuracy was found for students with at least 2 years of university education. We used the data collected by citizen scientists to create a large-scale standardized database of the distribution and abundance of the native and invasive crabs. Hemigrapsus sanguineus dominated the rocky intertidal zone from Sandy Hook, New Jersey to Boston Harbor, Massachusetts while C. maenas dominated the northern extent of the sampled coastline. A citizen scientist of this monitoring network detected a range expansion of H. sanguineus. We identified obstacles to creating a national monitoring network and proposed recommendations that addressed these issues.     

A trematode infection with no effect on reproductive success of a sand-bubbler crab

The infection effects of the parasitic digenean trematode on the body weight and reproductive success of the sand-bubbler crab were examined. Gynaecotyla squatarolae (Trematoda: Microphallidae) infects the body cavity of Scopimera globosa (Decapoda: Scopimeridae) and uses the crab as its second intermediate host. The parasites infected all reproductive crabs examined to varying degrees. Larger crabs of both sexes had more parasites than smaller ones, probably because body size reflects age, and older crabs had a longer period of exposure to infection. Males had more parasites than females, probably because of sexual difference in acting time on the surface. Ovigerous females stay in closed burrows and do not act on the surface during incubation, and so have less chance of infection than males. The quantity of infecting parasites did not explain variations in either body weight or reproductive success of individual crabs in a field experiment. The life history of this parasite, relative body size of the crabs, and cost and the possible benefit of manipulation for the parasite may explain these results.     

An invasive crab alters interaction webs in a marine community

Over the last decade, the non-native, filter-feeding crab Petrolisthes armatus invaded oyster reefs of the South Atlantic Bight at densities of thousands m⁻². Mesocosm and field experiments demonstrated that P. armatus at ∼10–75% of mean summer densities: (1) suppressed growth of small oysters, biomass of benthic microalgae, and recruitment of native mud crabs, (2) enhanced oyster, mussel, and total bivalve recruitment, macroalgal cover, and survivorship of predatory oyster drills, but (3) did not affect native taxonomic richness. Laboratory feeding assays, field tethering experiments, and population changes in field and mesocosm experiments suggest that P. armatus is a preferred prey for native mud crabs and other consumers, thus relieving predation on native species and enhancing recruitment or survival of bivalves and oyster drills. In contrast, the invasive crab can consume crustacean larvae and via this feeding may suppress recruitment of native mud crabs. Our findings should be conservative given the low densities of P. armatus seeded into experimental plots and our inability to run longer-term experiments due to controls rapidly being colonized by non-native crabs recruiting from the plankton. Invasive crabs commonly impact native communities via predation, but community impacts of this invasive crab may be as much due to its role as a preferred prey of native consumers as to its predation on native prey. Given that oysters are foundation species for shallow reefs in the South Atlantic Bight, the long-term effects of this invasion could be considerable.     

Occurrence of additional Zoea-VI larvae in the mud crab, Scylla paramamosain (Estampador), reared in the laboratory

Mud crabs, Scylla spp. , are commercially important in many Indo-Pacific countries. The larval development of mud crabs has been reported previously as five zoeal and one megalopal stages. This paper reports larval rearing experiments that revealed variability in larval developmental stages in the mud crab Scylla paramamosain, one of four mud crab species. In addition to normal five zoeal stages, an alternative pathway of developing through six zoeal stages was observed for the crab. There were evidences suggested that the appearance of the additional Zoea-VI larvae was associated with unfavourable dietary conditions, including poor quality of diet, inadequate quantity of dietary supply and a period of starvation for newly hatched larvae. Based on exuviae and larval specimens, the morphology of the additional Zoea-VI larvae was described.     

The infestation and dispersion patterns of Carcinonemertes spp. (Nemertea) on their crab hosts

Seasonal changes in the infestation and dispersion patterns of egg predatory nemerteans on their crab hosts were analyzed. Marked differences in the seasonal patterns of infestation were noted between worm species, yet, common patterns in the aggregation of the worms were found. Worm aggregation increased at the onset of the reproductive seasons of the hosts, and at the nadirs of the reproductive seasons for those hosts with year round breeding. The aggregation patterns of two worm species fluctuated with the physical environment of their estuarine hosts. Salinity changes as a result of seasonal rains may have caused changes in the underlying dispersion patterns of Carcinonemertes epialti on Hemiarapsus oregonensis, and C. mitsukurii on Portunus pelagicus. Lastly, the embryogenic cycle of the host species was significant in shaping the infestation and aggregation patterns of C. epialti on Cancer anthonyi and C. regicides on Paralithodes camtschaticus. Worm immigration and emigration were linked to crab embryogenesis and directly influenced the dispersion patterns of the worms.     

Impact scenario for the invasive red king crab <Emphasis Type="Italic">Paralithodes camtschaticus</Emphasis> (Tilesius, 1815) (Reptantia,Lithodidae) on Norwegian,native, epibenthic prey

Large invasive predators like the king crab, Paralithodes camtschaticus, deserve particular attention due to their potential for catastrophic ecological impact on recipient communities. Conspicuous, epibenthic prey species, such as the slow growing commercial scallop Chlamys islandica, are particularly exposed to the risk of local extinction. A research program integrating experiments and field monitoring is attempting to predict and track the impact of invasive king crab on scallop beds and associated fauna along the north Norwegian coast. The claw gape of the crab shows no limitations in handling the flat-bodied scallop. However, the potential impact of the crab on scallop may depend on the availability of other calcified prey associated with scallop beds, such as the sea star, sea urchin, and blue mussel, all species recorded in the diet of P. camtschaticus. To address this issue, a laboratory experiment on foraging behaviour of P. camtschaticus was conducted. The experimental results show that all size classes of red king crab prefer scallops, but small juveniles and medium sized crabs demonstrate active selection for starfish (Asterias rubens) that equals or surpasses the electivity of the large crab. The selection of sea urchin (Strongylocentrotus droebachiensis) and blue mussel (Mytilus edulis) is slightly positive or neutral for the three crab size classes. These results suggest that scallop beds with a rich associated fauna are less vulnerable to red king crabs predation and possibly more resilient than beds with few associated species. Also, crab size distribution is likely relevant for invasion impact, with increasing abundance of small and medium sized crabs being detrimental for alternative calcified prey associated with scallop beds. Successive stages of crab invasion will see an acceleration of scallop mortality rates associated with (i) decreasing availability of alternative prey, due to protracted predation pressure intensified by recruitment of juvenile crabs, and (ii) increased number of large crabs. Estimates of crab density and intake rates suggest that the accelerated loss rates will eventually endanger scallop beds persistence.     

Larval Development in the Rhizocephalan Barnacle Sacculina pilosella

We studied, under laboratory conditions, the larval development of a rhizocephalan barnacle Sacculina pilosellaVan Kampen et Boschma, 1925, which parasitizes the kelp crab Pugettia quadridens(de Haan) in Vostok Bay, Sea of Japan. It is shown that at 22–23°C, the whole cycle of larval development takes about 3 days. The larvae of S. pilosellaare lecithotrophic; their development, like in other rhizocephalans, comprises five naupliar instars. Like the larvae of all sacculinids, the nauplii of S. pilosellahave no flotation collar. In their structure, the larvae of S. pilosellaare similar to the nauplii of the typical sacculina, S. carcini(elongated body outline, long furcal branches, and weakly pronounced segmentation of the abdomen). On the other hand, the characteristic outgrowth inbetween the furcal branches that is characteristic of stages IV and V in S. carciniand S. polygeneais absent in the larvae of S. pilosella.The first seta on the antennula of S. pilosellacompletely disappears only at stage IV; however, at stage III, it is already significantly reduced. No morphological differences have been revealed between male and female larvae of S. pilosellaexcept certain size differences.     

Host range and specificity of an Argentinean isolate of the aquatic fungus <Emphasis Type="Italic">Leptolegnia chapmanii</Emphasis> (Oomycetes: Saprolegniales), a pathogen of mosquito larvae (Diptera: Culicidae)

An isolate from Argentina of the fungal mosquito pathogen Leptolegnia chapmanii (ARSEF 5499), was tested against 12 species of mosquito larvae and on species of non-target aquatic invertebrates and vertebrates. The mosquito species tested were Aedes aegypti, Anopheles sp., Culex apicinus, Cx. castroi, Cx. dolosus, Cx. pipiens, Cx. renatoi, Isostomyia paranensis, Ochlerotatus albifasciatus, Oc. crinifer, Psorophora cyanescens, and P. ferox. Mosquito larvae of 10 species were susceptible, with mortality rates from 10–100%. Two mosquito species Cx. renatoi and I. paranensis were not infected by Leptolegnia. None of the non-target fauna treated was infected by L. chapmanii with exception of members of the Family Chironomidae which were susceptible at low infection rates. Researcher CIC (Committee for Scientific Research of Buenos Aires State).     

Selective crab predation on native and introduced bivalves in British Columbia

Experiments were conducted to determine whether locally abundant crab species prefer co-occurring littleneck clams, Protothaca staminea (Conrad, 1837) and Tapes philippinarum (A. Adams and Reeve, 1850), relative to a recently introduced species, the varnish clam, Nuttallia obscurata, (Reeve, 1857). Prey preference, handling time, pick-up success, profitability and consumption rates were investigated for two crab species, Dungeness crab, Cancer magister (Dana, 1852) and red rock crab, Cancer productus (Randall, 1839) crabs. Both crab species preferred varnish clams over the native species. This may be attributable to the lower handling time, higher pick-up success and increased profitability of consuming varnish clams. Handling time appeared to be a factor not only in species preference, but also in the degree of preference, with shorter handling times corresponding to stronger preference values. Both native and introduced bivalves burrow into the substratum, with the varnish clam burrowing deepest. When feeding on clams in limited substratum both crab species preferred the varnish clam. In the unlimited substratum trials Dungeness crabs preferred varnish clams (although to a lesser degree) while red rock crabs preferred littleneck clams. This was likely due to the significantly deeper burial of the varnish clam, making it less accessible. Although the morphology (i.e. thin shell, compressed shape) of the invader increases its vulnerability to predation, burial depth provides a predation refuge. These results demonstrate how interactions between native predators and the physical characteristics and behaviour of the invader can be instrumental in influencing the success of an invasive species.     

Pathogenicity of fungi isolated from the larvae of the mangrove crab,Scylla serrata, in Indonesia

Heavy mortality reaching almost 100% occurred in the larvae of the mangrove crab,Scylla serrata, from July to December 1997 at the hatchery of Gondol Research Station for Coastal Fisheries in Bali, Indonesia. Mortality was observed in larvae after hatching from the eggs. The affected larvae were whitish and filled with numerous aseptate hyphae. Three fungi belonging to the order Lagenidiales,Lagenidium callinectes, Haliphthoros milfordensis, andHalocrusticida sp., were isolated from the infected larvae. Pathogenicity tests of the infected fungi against the larvae of mangrove crab demonstrated that all isolates were pathogenic.     

The trimorphic flea-beetle, <Emphasis Type="Italic">Alagoasa extrema</Emphasis>, not suitable for biocontrol of <Emphasis Type="Italic">Lantana camara</Emphasis> in Africa

The biology and host range of the flea-beetle, Alagoasa extrema (Harold) (Coleoptera: Chrysomelidae: Galerucinae: Alticini), was studied under quarantine laboratory conditions to evaluate the insect’s suitability for release as a biological control agent for the noxious weed, Lantana camara L. (Verbenaceae), in South Africa. Identification of this species proved noteworthy. The adults of A. extrema display three freely interbreeding and very distinct colour morphs, which can be confused with no less than 11 species of Alagoasa found in Mexico and the southwestern U.S. Initially, specimens were identified as two species of the genus Alagoasa Bechyné, i.e. A. quadrilineata (Harold) and A. extrema. Thirteen additional Alagoasa species and Kushelina petaurista (F). that can be confused with A. extremabased on external morphology of adults or larvae are discussed. Favourable biological characteristics included long-lived adults, several overlapping generations per year, and high adult and larval feeding rates. Observations from the insect’s native Mexican range and studies in South Africa suggest that A. extrema would probably be more suited to subtropical, rather than temperate areas in Africa. Host-specificity studies showed A. extrema to be an oligophagous species, capable of feeding and developing on several non-target species, especially two indigenous, African Lippia species (Verbenaceae). The host suitability of these indigenous species was only marginally lower than that of L. camara, and the potential risk to them was deemed to be too high to warrant release. It was therefore recommended that A. extrema be rejected as biocontrol agent for lantana in Africa.