Field evidence of trait-mediated indirect interactions in a rocky intertidal food web

Studies on the implications of food web interactions to community structure have often focused on density-mediated interactions between predators and their prey. This approach emphasizes the importance of predator regulation of prey density via consumption (i.e. lethal effects), which, in turn, leads to cascading effects on the prey's resources. A more recent and contrasting view emphasizes the importance of non-lethal predator effects on prey traits (e.g. behaviour, morphology), or trait-mediated interactions. On rocky intertidal shores in New England, green crab ( Carcinus maenas ) predation is thought to be important to patterns of algal abundance and diversity by regulating the density of herbivorous snails ( Littorina littorea ). We found, however, that risk cues from green crabs can dramatically suppress snail grazing, with large effects on fucoid algal communities. Our results suggest that predator-induced changes in prey behaviour may be an important and under-appreciated component of food web interactions and community dynamics on rocky intertidal shores.     

Characterization of microsatellite loci in the European green crab (Carcinus maenas)

Carcinus maenas (Decapoda: Portunidae) has proven a highly successful invasive marine species whose potential economic and ecological impacts are of great concern worldwide. Here, we characterize 14 polymorphic microsatellite loci in C. maenas and its sister species Carcinus aestuarii. These markers will prove useful for fine‐scale genetic analyses of native and introduced populations, for assessment of the sources and routes of invasion and for evaluation of post‐invasion population dynamics.     

Intertidal distribution and species composition of brachyuran crabs at two rocky shores in Central Portugal

The objectives of the present study are to describe and compare the brachyuran community of rocky shores within the Central Portuguese coast and to examine the zonation patterns of the most representative species. For this, randomly placed transects were surveyed to obtain crab counts according to microhabitat and intertidal level. Repeated sampling in two different shores during two different seasons provided spatial and temporal replication for zonation analyses. Seven species were registered: Pachygrapsus marmoratus, Eriphia verrucosa, Xantho incisus, Carcinus maenas, Necora puber, Pirimela denticulata and Pilumnus hirtellus. Species density rankings are the same at both localities, but the less exposed shore presents higher diversity. While most species are mainly confined to specific microhabitats in the lower level, P. marmoratus and E. verrucosa can exploit the whole intertidal range. Regardless of shore and season, E. verrucosa is more abundant in the lower intertidal levels, while no such zonation patterns were recorded for P. marmoratus. Initial predictions concerning the effect of wave exposure and temperature on the zonation of those species are not validated after analysing the factorial model proposed. Between-shore contrasts were found instead, with higher densities recorded in the more exposed locality for both species. Possible causes of the observed patterns are discussed.     

Amplified fragment length polymorphism data provide a poor solution to the Littorina littorea puzzle

The status of Littorina littorea as an exotic species in North America has been debated in the biological literature for over a century. In recent decades, molecular data have been used to supplement historical and archaeological evidence that has suggested that the snail may have persisted in North America well before European settlers arrived. However, these earlier molecular studies have since been questioned due to incomplete sampling of both the geographical range of L. littorea and the genome of the species itself. Here we consider an amplified fragment length polymorphism screen of the nuclear genome, which provides a large number of independent dominant markers that can be used in resolving this historical puzzle. Although these data appear to refute earlier allozyme studies of L. littorea as indicating long-term divergence between European and American populations, the data are also problematic in that they include clear artefacts. We discuss these artefacts and suggest future approaches to definitively resolving the debate over L. littorea's introduction to North America.     

泉州湾岩相潮间带底栖生物生态研究Ⅰ.物种多样性与分布特征

2001年11月至2002年8月在泉州湾浮山(R1)、下洋(R2)、垵头(R3)和大坠岛(R4) 选择4条岩相潮间带断面对底栖生物分布进行了调查,共采集了6687号标本.经分析鉴定共有324种,其中有藻类17种,多毛类102种,软体动物96种,节肢动物62种,棘皮动物11种,其他动物共有36种.岩相断面种类的水平分布受制于潮流和海浪,不同断面的种数排序是浮山(206种)＞下洋(197种)＞大坠岛(193种)＞垵头(166种).垂直分布主要受制于潮汐和海浪.各断面不同潮区的共同特征是中潮区的物种最多,在129-156种之间;低潮区居第二,在87-116种之间;高潮区的种类最少,在9-20种之间.种数的季节变化反映生物对水温的适应性,不同季节种数排序为夏季(190种)＞春季(164种)＞秋季(158种)＞冬季(150种).     

Genetic patterns across multiple introductions of the globally invasive crab genus Carcinus

The European green crab Carcinus maenas is one of the world's most successful aquatic invaders, having established populations on every continent with temperate shores. Here we describe patterns of genetic diversity across both the native and introduced ranges of C. maenas and its sister species, C. aestuarii, including all known non‐native populations. The global data set includes sequences from the mitochondrial cytochrome c oxidase subunit I gene, as well as multilocus genotype data from nine polymorphic nuclear microsatellite loci. Combined phylogeographic and population genetic analyses clarify the global colonization history of C. maenas, providing evidence of multiple invasions to Atlantic North America and South Africa, secondary invasions to the northeastern Pacific, Tasmania, and Argentina, and a strong likelihood of C. maenas × C. aestuarii hybrids in South Africa and Japan. Successful C. maenas invasions vary broadly in the degree to which they retain genetic diversity, although populations with the least variation typically derive from secondary invasions or from introductions that occurred more than 100 years ago.     

Occurrence of epifaunshers on the periwinkle,Littorina littorea (L.), and interactions with the the polychaete Polydora ciliata (Johnston)

A population of Littorina littorea on the northshore of Southampton Water, UK, has been examined forthe occurrence of epifauna. Epifaunal cover was notaffected by shore level below MTL but was greatest onlarger shells, which were commoner on the lower shoreclose to MLWS. Epifaunal components includedbarnacles, serpulids and bryozoans. The polychaetePolydora ciliata occurred most frequently on thelower shore where it excavates galleries in periwinkleshells. Heaviest infestations were found in largershells. P. ciliata settles preferentially in orbeside irregularities on the shell surface. Settlementof barnacles close to the shell aperture, where newshell growth takes place, produces shell distortionwhich creates sites for settlement of P. ciliata. Theangle between serpulid tubes and theshell surface also attracts P. ciliata. It issuggested that colonisation by barnacles and serpulidsfacilitates settlement by P. ciliata, and thatthe latter decreases survival of the host byprogressively destroying the shell.     

Physiological responses to emersion in the intertidal green crab,Carcinus maenas (L.)

The green shore crab, Carcinus maenas, undergoes on average 6?h periods of emersion during each low-tide cycle during the summer months. Under those conditions, the crab is cut off from its normal water environment and is exposed to potential stress from a suite of environmental and physiological changes: dehydration, compromised gas exchange and resultant internal hypoxia and hypercapnia, thermal stress, and ammonia toxicity. This study examined the comprehensive responses of the green crab in water and to a 6?h emersion period laboratory simulation of a tidal cycle followed by a 1?h re-immersion period, measuring indicators of dehydration, hemolymph osmolality, oxygen uptake, hemolymph acid–base status, heart and ventilatory rate, and hemolymph ammonia and ammonia excretion. Green crabs showed physiological responses of varying magnitude to 6?h of emersion. Individuals were found in the field exclusively under rocks and large clumps of seaweed where temperatures were approximately half those of exposed surfaces and relative humidity was about twice as high as ambient air. During emersion, crabs lost less than 5% of their wet weight, and hemolymph osmolality did not increase significantly. Oxygen uptake continued in air at about 50% of the control, aquatic values; and the gills continued to be ventilated by the scaphognathite, albeit at lower rates. Hemolymph lactate concentrations increased, indicating a shift to a greater reliance on anaerobic metabolism to support energetic needs. A slight acidosis developed in the hemolymph after 1?h of emersion, but it did not increase thereafter. Ammonia concentrations in the hemolymph were unchanged, as ammonia was volatilized by the gills and excreted into the air as NH₃ gas. These results show that the green crab copes with emersion by seeking refuge in microhabitats that mitigate the changes in the physical parameters of intertidal emersion. Physiologically, desiccation is avoided, cardio-respiratory processes are maintained at reduced levels, and hemolymph acid–base balance is minimally affected. Ammonia toxicity appears to be avoided by a shift to excreting NH₃ gas directly or indirectly to air.     

The close relationship between estimated divergent selection and observed differentiation supports the selective origin of a marine snail hybrid zone

To study the role of divergent selection in the differentiation of the two morphs in a hybrid zone of the intertidal snail Littorina saxatilis, we compared the strength of the divergent selection acting on a series of shell characters (as estimated by the viability of snails in a reciprocal transplant experiment) with the contribution of these characters to the phenotypic differences between the morphs. We found a close correlation between selection and differentiation, which suggests a cause-effect relationship, i.e. that all present differentiation is the result of past divergent selection. In addition, divergent selection was a very important component of the total natural selection acting on shell measures. These novel results support previous evidence, based on allozyme analysis, of a parapatric origin for this hybrid zone. We discuss possible limitations of this interpretation and the circumstances under which allopatric differentiation would produce the same results. Phenotypic analysis of divergent selection may be a useful method of investigating the evolutionary mechanisms involved in differentiation processes.     

Space Partitioning within Populations of Sea Anemones (Genus: Anthopleura) in the California Rocky Intertidal Zone

The spatial patterns of sea anemones (genus: Anthopleura) were analyzed off the rocky intertidal of California. A. elegantissima partitions space in a regular or even pattern and interferes with conspecifics. Its associate A. xanthogrammica shows no signs of interference with conspecifics or congenerics and displays a clumped spatial distribution.     

Phenotypic plasticity in two marine snails: constraints superseding life history

In organisms encountering predictable environments, fixed development is expected, whereas in organisms that cannot predict their future environment, phenotypic plasticity would be optimal to increase local adaptation. To test this prediction we experimentally compared phenotypic plasticity in two rocky-shore snail species; Littorina saxatilis releasing miniature snails on the shore, and Littorina littorea releasing drifting larvae settling on various shores, expecting L. littorea to show more phenotypic plasticity than L. saxatilis. We compared magnitude and direction of vectors of phenotypic difference in juvenile shell traits after 3 months exposure to different stimuli simulating sheltered and crab-rich shores, or wave-exposed and crab-free shores. Both species showed similar direction and magnitude of vectors of phenotypic difference with minor differences only between ecotypes of the nondispersing species, indicating that plasticity is an evolving trait in L. saxatilis. The lack of a strong plastic response in L. littorea might be explained by limits rather than costs to plasticity.