Temporal variation in parasite loading in relation to life history patterns of Littorina obtusata and L. fabalis

Parasitic infection of Littorina obtusata (L.) and Littorina fabalis Turton by larval digeneans was studied in summer (August) and winter (February) on a sheltered shore at St Michael's Island, Isle of Man. Sampling was timed to investigate parasite loading at different stages of the littorinids' life history. Nine species of digeneans were identified and infection rates were similar between the species, with a slightly higher prevalence in females for L. obtusata and males in L. fabalis. Temporal differences in infection were small, although metacercariae and degenerate infections were greater in female L. obtusata in February. Analysis of shell parameters and size of the penis in males and the pallial oviduct in females revealed morphological differences between parasitised and uninfected individuals. Parasitic stunting of the penis was evident in both species and the pallial oviduct was also smaller in parasitised females. There was also a tendency for parasitised individuals to have larger shells; although size also varied with season for L. fabalis.     

Ontogeny of shell morphology and shell strength of the marine snails Littorina obtusata and Littorina mariae: different defence strategies in a pair of sympatric, sibling species

The sibling marine snails Littorina obtusata (L.) and Littorina mariae Sacchi & Rastelli are sympatrically distributed and the shells of both species are subject to similar breaking forces by predatory crabs. Nevertheless, the two species exhibit rather different growth and defence strategies. To determine growth patterns, we measured changes in five morphological variables with increasing shell length: body whorl thickness at the point of crushing force application, shell height (related to globosity), shell mass, body mass, and apertural lip thickness. We also measured ontogenetic changes in the ability to withstand shell crushing. For most morphological variables, L. mariae showed uniformly allometric growth of juveniles into adults. In contrast, L. obtusata usually exhibited a distinct change in growth pattern upon reaching maturity. As adults, L. mariae showed a more sustained increase in overall shell mass and in body whorl thickness (defence against crushing attacks) and also had proportionally thicker apertural lips (defence against peeling attacks). Littorina obtusata , however, grew to a larger size and their shells could accommodate larger bodies at all sizes. Furthermore, the strength of L. obtusata shells increased faster than could be accounted for by either overall shell mass or thickness at the point of force application, suggesting strengthening by other means such as changes in shell microstructure or shape (other than globosity). These results illustrate the viability of two contrasting antipredator strategies, despite a highly similar phylogenetic history and selective regime.     

Maintenance of zonation patterns in two species of flat periwinkle,Littorina obtusata and L. mariae

The zonation patterns of Littorina obtusata (L.) and Littorina mariae Sacchi et Rastelli were shown to be quite distinct on a sheltered rocky shore. L. obtusata was found at all the heights sampled; it reached peak numbers at mid shore on the alga Ascophyllum nodosum L. (Le Jol). There was no difference in the tidal height occupied by adults or juveniles; or in the mean size of L. obtusata along the vertical gradient of the shore. In contrast L. mariae occurred exclusively low on the shore, on Fucus serratus L. Translocation of the two species within their respective levels resulted in random movement after 4 days, although initial movements after 1 and 2 days were sometimes directional. Animals transplanted to the normal level of the other species showed directional movement towards their home zone; this was most pronounced after 4 days. There was no difference in the distance moved by the two species, although the distance moved did vary with tidal height, both species moving further at mid shore than low shore. Distances moved by littorinids at replicate areas in the low shore were similar but those at mid shore did vary. There was an interaction between the species and the different tidal heights which revealed that transplanted species moved further than translocated species at the same tidal level. However, this was only significant in the case of L. mariae. It is suggested that the close relationship between the winkles and their host algae may direct the homing behaviour of displaced individuals.     

Differential snail predation by an exotic crab and the geography of shell-claw covariance in the Northwest Atlantic

Here we investigate if predation by the European green crab (Carcinus maenas) differs between two congeneric snails in the northwest Atlantic (Littorina littorea and L. obtusata), and ask if differential predation can help explain the geography of claw and shell forms among geographically separated populations. First, correlations between crusher-claw size and shell mass -- tested across a wide size range of animals -- were highly significant among populations of C. maenas and L. obtusata, whereas only a small number of significant correlations were found between C. maenas and L. littorea, and these were limited to the smaller size classes of snails and crabs. Moreover, among populations, L. obtusata shells were more frequently scarred than those of L. littorea, and L. obtusata were attacked and killed more frequently than L. littorea during field- and laboratory-predation experiments. Combined, results suggest L. obtusata is currently under greater selection by C. maenas than L. littorea for more crab-resistant shell forms. One possible explanation for these patterns is that L. littorea may have interacted with green crabs for centuries (in Europe) prior to their reintroduction to green crabs in America, thus predator-resistance may had already evolved.     

Changes in the wildfowl populations wintering on the Severn Estuary

Monthly winter counts of seven major wildfowl species show that, since 1960, Bewick's swan, shelduck, wigeon, teal and pintail have significantly increased in both the Severn and Britain as a whole; European White-fronted geese have shown no trend on the Severn but have significantly declined in Britain; mallard show no trend on the Severn but have increased in Britain. All except shelduck show a correlation between the trends in Britain and the Severn Estuary, while all but mallard and pintail rank as internationally important on the Severn. In recent years the Severn has assumed a considerable importance for pochard and tufted ducks. The status of many species in the Severn Estuary is dependent on the management of key areas.     

The Severn Estuary's changing wader populations during the last two decades

Many of the changes in population size which have occurred in the wading birds of the Severn Estuary reflect changes that have taken place in Britain as a whole. An important exception is the dunlin, which has maintained its numbers in the Severn despite undergoing a significant decline nationally. The fluctuations of several species show evidence of the effects of the hard winters of 1979, 1982 and 1986, the dry conditions of 1976 and perhaps of good breeding seasons in the high Arctic. Bridgwater Bay has declined in importance for nearly all species, with many individuals appearing to have switched to the Rumney on the northern shore.     

ALLOZYME SIMILARITY OF ATLANTIC AND PACIFIC SPECIES OF LITTORINA (GASTROPODA: LITTORINIDAE)

Four Littorina species from the Pacific (L. sitkana, L. brevicula,L. mandshurica, L. squalida) and four from the Atlantic (L.mariae, L. obtusata, L. saxatilis and L. littorea) were comparedelectrophoretically using 17 loci. Analysis of genetic identities showed that there were threegroups. The first group comprises L. sitkana, L. mariae, L.obtusata and L. saxatilis The second group consists of L. breviculaand L. mandshurica and the third one includes L. squalida andL. littorea. Our results support Reid's hypothesis of the originof Atlantic littorines from two Pacific ancestors. (Received 18 December 1991; accepted 30 March 1992)     

Environmentally induced variation in uric acid concentration and xanthine dehydrogenase activity in Littorina saxatilis (Olivi) and L. arcana Hannaford Ellis

Littorina saxatilis and Littorina arcana collected from a boulder field low in the intertidal zone had a uric acid concentration significantly higher than snails collected from nearby cliff crevices that were at the upper limit of their vertical range on the shore. The absolute concentrations varied with different collections, suggesting a possible seasonal fluctuation in uric acid. L. arcana had a greater concentration of uric acid than did L. saxatilis when both were from the boulder field, but the two did not differ significantly when taken from crevices. Samples from the two sites were marked and transplanted; by four weeks the transplanted animals showed a tendency to develop a uric acid concentration similar to that of natives of the site, suggesting a physiological rather than a genetic adaptation. Xanthine dehydrogenase activity differed between animals from the two sites in the same manner as the uric acid concentration.     

Spatial and temporal variation of trematode infection in coexisting populations of intertidal gastropods Littorina saxatilis and L. obtusata in the White Sea

Trematode infection was studied in sympatric populations of the periwinkles Littorina saxatilis and L. obtusata in 2 regions of Kandalaksha Bay of the White Sea to assess host-parasite interactions at the population level. Twenty-seven spatially separated populations were each surveyed in 1984-1994; 2 heavily infected populations were investigated annually over a 16 yr period. Ten trematode species were found in the periwinkle populations. The closest association in spatial distribution and temporal dynamics was observed between 3 ecologically and morphologically similar trematodes of the 'pygmaeus' group: Microphallus piriformes, M. pygmaeus and M. pseudopygmaeus. For these 3 species, the prevalences were closely associated in the 2 host species when spatially separated sites from the 2 studied regions were considered, while in the 2 populations studied over the 16 yr period, a correlation was only observed between the infection levels of L. saxatilis and L. obtusata by either M. piriformes and immature microphallids. Likewise, within each host species, significant correlations were revealed between the prevalence of the different microphallids of the 'pygmaeus' groups. However, they were fewer and weaker when the long-term dynamics of infection in the 2 heavily infected populations were considered. Most other trematodes did not show significant association in prevalence either within or between the 2 host species on spatial and temporal scales.     

Phenotypic clines, plasticity, and morphological trade-offs in an intertidal snail

Understanding the genetic and environmental bases of phenotypic variation and how they covary on local and broad geographic scales is an important goal of evolutionary ecology. Such information can shed light on how organisms adapt to different and changing environments and how life-history trade-offs arise. Surveys of phenotypic variation in 25 Littorina obtusata populations across an approximately 400-km latitudinal gradient in the Gulf of Maine revealed pronounced clines. The shells of snails from northern habitats weighed less and were thinner and weaker in compression than those of conspecifics from southern habitats. In contrast, body size (as measured by soft tissue mass) followed an opposite pattern; northern snails weighed more than southern snails. A reciprocal transplant between a northern and southern habitat revealed substantial plasticity in shell form and body mass and their respective measures of growth. Southern snails transplanted to the northern habitat produced lighter, thinner shells and more body mass than controls raised in their native habitat. In contrast, northern snails transplanted to the southern site produced heavier, thicker shells and less body mass than controls raised in their native habitat. Patterns of final phenotypic variation for all traits were consistent with cogradient variation (i.e., a positive covariance between genetic and environmental influences). However, growth in shell traits followed a countergradient pattern (i.e., a negative covariance between genetic and environmental influences). Interestingly, body growth followed a cogradient pattern, which may reflect constraints imposed by cogradient variation in final shell size and thickness. This result suggests the existence of potential life-history trade-offs associated with increased shell production. Differences in L. obtusata shell form, body mass, and their respective measures of growth are likely induced by geographic differences in both water temperature and the abundance of an invading crab predator (Carcinus maenas). Water temperatures averaged 6.8 degrees C warmer during the transplant experiment and C. maenas abundance is greater in the southern Gulf of Maine. Because both increased water temperature and crab effluent affect shell form in the same way, future experiments are needed to determine the relative importance of each. Nevertheless, it is clear that phenotypic plasticity has an important role in producing geographic variation in L. obtusata shell form. Moreover, the evolution of phenotypic plasticity in L. obtusata and other marine gastropods may be driven by architectural constraints imposed by shell form on body mass and growth.     

The diet of juvenile and adult twaite shad Alosa fallax fallax (lacépède) from the rivers Severn and Wye (Britain)

The diet of juvenile and adult twaite shad was studied at a number of freshwater and estuarine sites in the rivers Severn and Wye and in the coastal waters of Cardigan Bay (West Wales). In the Severn estuary and in the freshwater reaches of the Severn and Wye, adult twaite shad consumed little during their pre-spawning migration. In the Severn Estuary post-spawning adults were present during the summer months where they fed actively on mysids, gammarids and shrimps. Mysids dominated the diet of the adults caught in coastal waters during the autumn. The diet of one year old twaite shad was studied in the Severn Estuary. In May the one year olds from the lower estuary consumed mainly gammarids. During the summer months, this age-class moved into the inner estuary where they fed predominantly on mysids.The larvae and juveniles (age 0⁺) fed mainly on chironomid larvae and pupae and Simuliidae larvae, while in fresh water. In the estuary copepods, cladocerans and mysids dominated the diet of the juveniles. There was some suggestion of an increase in prey size during their period of residency in the estuary. The contrasting distribution of the juveniles in two river systems is discussed in relation to the effects of channelization.     

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.     

Not an ancient relic: the endemic Livistona palms of arid central Australia could have been introduced by humans

Livistona mariae is an endemic palm localized in arid central Australia. This species is separated by about 1000 km from its congener L. rigida, which grows distantly in the Roper River and Nicholson-Gregory River catchments in northern Australia. Such an isolated distribution of L. mariae has been assumed to have resulted from contraction of ancestral populations as Australia aridified from the Mid-Miocene (ca 15 Ma). To test this hypothesis at the population level, we examined the genetic relationships among 14 populations of L. mariae and L. rigida using eight nuclear microsatellite loci. Our population tree and Bayesian clustering revealed that these populations comprised two genetically distinct groups that did not correspond to the current classification at species rank, and L. mariae showed closest affinity with L. rigida from Roper River. Furthermore, coalescent divergence-time estimations suggested that the disjunction between the northern populations (within L. rigida) could have originated by intermittent colonization along an ancient river that has been drowned repeatedly by marine transgression. During that time, L. mariae populations could have been established by opportunistic immigrants from Roper River about 15 000 years ago, concurrently with the settlement of indigenous Australians in central Australia, who are thus plausible vectors. Thus, our results rule out the ancient relic hypothesis for the origin of L. mariae.     

The relationship between flat periwinkle life histories and digenean infections

Larval digenean parasites were studied in Littorina obtusata (L.) and L. mariae Sacchi & Rastelli at Sawdern Point in West Wales. Shell parameters, ovipositor and shell colour, penis morphology and sex ratios were scored, and the influence of parasitism studied. A total of 7 species of parasites were found, although the prevalence was very low in L. mariae, especially in the females. The parasitic gigantism that has been described in other species of gastropod was not found in this study. Parasitic castration does occur in some infected male L. obtusata, resulting in severe stunting of the penis. However, this phenomena was never observed in L. mariae. The winkle species are congeneric and inhabit broadly similar niches on the shore; their life histories are however quite different — one being annual and the other perennial. This probably affects exposure to infection, and might explain the differing prevalence of the parasites.     

Fish fauna of the Severn Estuary. Are there long-term changes in abundance and species composition and are the recruitment patterns of the main marine species correlated?

Fish were collected from the intake screens of the Oldbury Power Station in the Severn Estuary in each week between early July 1972 and late June 1977 and at least twice monthly between early January 1996 and late June 1999. The annual catches, after adjustment to a common sampling effort, demonstrate that the abundance of fish at Oldbury was far greater in the 1990s than 1970s, mainly due to marked increases in the numbers of certain marine species, such as sand goby, whiting, bass, thin-lipped grey mullet, herring, sprat and Norway pout. These increases may reflect the great improvement that occurred in the water quality of the Severn Estuary between these decades. The only species that declined markedly in abundance was poor cod. Modest declines in flounder and River lamprey paralleled those occurring elsewhere in the UK. The species composition in the two decades also differed, reflecting changes not only in the relative abundances of the various marine estuarine-opportunistic species, which dominated the ichthyofauna, but also in those of the suite of less abundant species in the estuary. The cyclical changes undergone each year by the species composition of the fish fauna of the Severn Estuary reflect sequential intra-annual changes in the relative abundances of species representing each of the marine, diadromous and freshwater categories. New approaches have been developed to test whether or not large sets of correlations between patterns of recruitment amongst abundant marine species (internal correlations), and between those patterns and salinity and water temperature within the estuary (cross-correlations), were significant. The correlation profile analyses found no evidence that the annual recruitment strengths of these species were either intercorrelated, or correlated with either one or a combination of both of the above environmental variables. Yet, the timings of the recruitment of these species into the estuary were intercorrelated, i.e. a slightly earlier or later than normal immigration by one species in a given year was paralleled by the same trend in other species. However, this association in recruitment times could be linked neither to salinity nor water temperature within the estuary, nor to a combination of these two variables. These results indicate that, while the factors that influence the annual recruitment strengths of the juveniles of different marine species vary, inter-annual differences in the phasing of events that regulate spawning times and/or larval dispersal influence, in the same direction, the times when marine species are recruited into the estuary.     

Benthic macrofauna and sediments in the Severn Estuary

In the Severn Estuary and Bristol Channel, a multivariate analysis indicated eight groupings of macrobenthic fauna, each associated with a different physical habitat. Samples were faunistically impoverished, especially in sublittoral areas, but greater species richness was associated with reefs of Sabellaria alveolata , which extended well into the estuary.     

Reproductive strategies of two invasive tilapia species Oreochromis mossambicus and Tilapia mariae in northern Australia

The reproductive biology of two invasive tilapia species, Oreochromis mossambicus and Tilapia mariae, resident in freshwater habitats in north-eastern Australia was investigated. Oreochromis mossambicus exhibited plasticity in some of its life-history characteristics that enhanced its ability to occupy a range of habitats. These included a shallow, weed-choked, freshwater coastal drain that was subject to temperature and dissolved oxygen extremes and water-level fluctuations to cooler, relatively high-altitude impoundments. Adaptations to harsher conditions included a decreased total length (L(T) ) and age (A) at 50% maturity (m50), short somatic growth intervals, early maturation and higher relative fecundities. Potential fecundity in both species was relatively low, but parental care ensured high survival rates of both eggs and larvae. No significant difference in the relative fecundity of T. mariae populations in a large impoundment and a coastal river was found, but there were significant differences in relative fecundities between several of the O. mossambicus populations sampled. Total length (L(T) ) and age at 50% maturity of O. mossambicus populations varied considerably depending on habitat. The L(Tm50) and A(m50) values for male and female O. mossambicus in a large impoundment were considerably greater than for those resident in a small coastal drain. Monthly gonad developmental stages and gonado-somatic indices suggested that in coastal areas, spawning of O. mossambicus and T. mariae occurred throughout most of the year while in cooler, high-altitude impoundments, spawning peaked in the warmer, summer months. The contribution these reproductive characteristics make to the success of both species as colonizers is discussed in the context of future control and management options for tilapia incursions in Australia.     

Inheritance of longitudinal shell bands in the snails Littorina obtusata and Littorina saxatilis (Gastropoda, Prosobranchia)

The hypothesis of a monogenic inheritance of dark longitudinal bands on the shell in the gastropods Littorina obtusata and L. saxatilis was checked. One gene having two alleles proved to be responsible for the shell banding pattern in both of the species. The presence of bands was a dominant character in either case.     

Distribution of the species of rough periwinkle (Littorina) in Great Britain

The status of the four currently recognised species in the Littorina saxatilis species-complex, i.e. Littorina nigrolineata Gray, L. arcana Hannaford Ellis, L. saxatilis (Olivi) and L. neglecta Bean is reviewed briefly, with notes on their characteristic features and location on the shore. Since the taxonomy of these rough periwinkles has only become stable relatively recently much of the previously published information on distribution is of little or no use. In this paper their distribution around the coastline of England and Wales (with some notes on Scotland) is described and discussed. L. saxatilis is found in a wide range of habitats from exposed peninsulas to estuaries on all shores where there is a suitable rocky or stony substrate; also in salt marsh pools. L. arcana has a more restricted distribution and is notably absent along much of the south English coast, central Cardigan Bay and possibly northern Scotland; it is not found in estuaries. L. nigrolineata has an even more restricted distribution, although it occurs both on exposed coasts and in estuaries; it has only been found by us in one locality on the east coast. L. neglecta is probably fairly widely distributed but we have few details so far. The implications of the different patterns of distribution are discussed.     

Growth and abundance of O-group herrings, Clupea harengus L., in the Severn Estuary

Collections made from the cooling water intake screens of Power Stations in the Severn Estuary and the Bristol Channel have been used to investigate the biology of O-group herrings in this region. The abundance of herrings at Oldbury, the principal sampling site, was low in most years and in five successive seasons between July 1972 and December 1976, large numbers were caught only between the summer of 1975 and the spring of 1976. The herrings were first observed in July, when many had still not completed metamorphosis, and reached peak numbers in September and October during which period they were clearly growing. Numbers declined in December before rising dramatically in January at which time the length-frequency curves displayed a pronounced bimodality. On the basis of data from other parts of the Bristol Channel and Severn Estuary, it is suggested that this bimodality is due to the influx of some animals from other regions where the growth rate was greater. Although movement is apparently taking place at this time and the numbers decline markedly in subsequent months, the last remnants of the population do not leave the estuary until early May. Post-pelvic scute (K₂) and vertebral counts (VS) of 13.82 and 55.24 respectively, indicate that the Severn Estuary herrings are the product of a spring spawning stock, a view entirely consistent with length and field data.