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Podocotyle is a genus of marine opecoelid digeneans, widespread in fishes and invertebrates (gastropods and crustaceans) in the neritic and littoral zones of mainly the Northern Hemisphere. The phylogeny, taxonomy and evolution of digeneans of this genus are quite mysterious due to the wide distribution of these parasites and their species richness. For a number of representatives of Podocotyle spp. a wide range of phenotypic variability has been described, which prevents species identification based solely on morphological characteristics. In this study, we apply molecular and morphological techniques to identify Podocotyle spp. from vertebrate and invertebrate hosts inhabiting the White Sea. Specimens of Podocotyle spp. were collected from pleuronectid and zoarcid fishes, as well as from periwinkles of the intertidal zone, and were studied using light microscopy, scanning electron microscopy, and molecular phylogenetic methods. A combination of molecular and morphological approaches revealed Podocotyle angulata and three species-level taxa awaiting nomenclatural proposals, Podocotyle sp. of Denisova et al. 2023, P. atomon sensu lato and Podocotyle cf. angulata of Sokolov et al. 2023. Conspecificity of one type of cercariae released from infected Littorina obtusata and L. saxatilis with adult digeneans identified as P. atomon sensu lato was established. We provide detailed descriptions of the cotylomicrocercous larvae of P. atomon sensu lato and Podocotyle sp. of Denisova et al. 2023, focusing on the features of the surface morphology of their tegument and associated structures in order to determine the general and diagnostic characteristics of these species. [ABSTRACT FROM AUTHOR]

Chromosomal rearrangements can lead to the coupling of reproductive barriers, but whether and how they contribute to the completion of speciation remains unclear. Marine snails of the genus Littorina repeatedly form hybrid zones between populations segregating for multiple inversion arrangements, providing opportunities to study their barrier effects. Here, we analyzed 2 adjacent transects across hybrid zones between 2 ecotypes of Littorina fabalis ("large" and "dwarf") adapted to different wave exposure conditions on a Swedish island. Applying whole-genome sequencing, we found 12 putative inversions on 9 of 17 chromosomes. Nine of the putative inversions reached near differential fixation between the 2 ecotypes, and all were in strong linkage disequilibrium. These inversions cover 20% of the genome and carry 93% of divergent single nucleotide polymorphisms (SNPs). Bimodal hybrid zones in both transects indicated that the 2 ecotypes of Littorina fabalis maintain their genetic and phenotypic integrity following contact. The bimodality reflects the strong coupling between inversion clines and the extension of the barrier effect across the whole genome. Demographic inference suggests that coupling arose during a period of allopatry and has been maintained for > 1,000 generations after secondary contact. Overall, this study shows that the coupling of multiple chromosomal inversions contributes to strong reproductive isolation. Notably, 2 of the putative inversions overlap with inverted genomic regions associated with ecotype differences in a closely related species (Littorina saxatilis), suggesting the same regions, with similar structural variants, repeatedly contribute to ecotype evolution in distinct species. [ABSTRACT FROM AUTHOR]

Marine snails (genus Littorina) have recently emerged as a model for speciation research, with repeated formation of different types of barriers to gene flow in contact zones between ecotypes and taxa. Inversion polymorphisms are found in an increasing number of species, including Littorina snails, and have been suggested to promote both local adaptation, ecotype formation and speciation, but their roles can vary. In Littorina , their role in later stages of speciation remains particularly unclear. A one-dimensional speciation continuum is frequently used to frame our understanding of the speciation process. However, a number of recent studies have argued that no single measure adequately summarises reproductive isolation and, therefore, the speciation process is best viewed in multiple dimensions. The many Littorina contact zones provide empirical data to illustrate how this principle can be applied. Speciation is a key evolutionary process that is not yet fully understood. Combining population genomic and ecological data from multiple diverging pairs of marine snails (Littorina) supports the search for speciation mechanisms. Placing pairs on a one-dimensional speciation continuum, from undifferentiated populations to species, obscured the complexity of speciation. Adding multiple axes helped to describe either speciation routes or reproductive isolation in the snails. Divergent ecological selection repeatedly generated barriers between ecotypes, but appeared less important in completing speciation while genetic incompatibilities played a key role. Chromosomal inversions contributed to genomic barriers, but with variable impact. A multidimensional (hypercube) approach supported framing of questions and identification of knowledge gaps and can be useful to understand speciation in many other systems. [ABSTRACT FROM AUTHOR]

The intertidal gastropod Littorina saxatilis is a model system to study speciation and local adaptation. The repeated occurrence of distinct ecotypes showing different levels of genetic divergence makes L. saxatilis particularly suited to study different stages of the speciation continuum in the same lineage. A major finding is the presence of several large chromosomal inversions associated with the divergence of ecotypes and, specifically, the species offers a system to study the role of inversions in this divergence. The genome of L. saxatilis is 1.35 Gb and composed of 17 chromosomes. The first reference genome of the species was assembled using Illumina data, was highly fragmented (N50 of 44 kb), and was quite incomplete, with a BUSCO completeness of 80.1% on the Metazoan dataset. A linkage map of one full-sibling family enabled the placement of 587 Mbp of the genome into 17 linkage groups corresponding to the haploid number of chromosomes, but the fragmented nature of this reference genome limited the understanding of the interplay between divergent selection and gene flow during ecotype formation. Here, we present a newly generated reference genome that is highly contiguous, with a N50 of 67 Mb and 90.4% of the total assembly length placed in 17 super-scaffolds. It is also highly complete with a BUSCO completeness of 94.1% of the Metazoa dataset. This new reference will allow for investigations into the genomic regions implicated in ecotype formation as well as better characterization of the inversions and their role in speciation. [ABSTRACT FROM AUTHOR]

Simple Summary: Nowadays, it is well known that the gut bacterial microbiome is crucially important for the adaptation of multicellular organisms to their environment. In this study, we aimed to identify the differences between the bacteriomes of two closely related marine snail species living in sympatry but feeding in different micro-niches. Although there were significant interspecies differences detected during the summer season, we did not observe this dissimilarity during the cold season. Moreover, the diversity of bacterial communities associated with snails decreased in autumn. We suggest that bacteria predominantly associated with one species degrade its toxic feeding substrate. These results help to understand the role of bacteriomes in the adaptation and divergence of closely related species. Symbiotic microorganisms may provide their hosts with abilities critical to their occupation of microhabitats. Gut (intestinal) bacterial communities aid animals to digest substrates that are either innutritious or toxic, as well as support their development and physiology. The role of microbial communities associated with sibling species in the hosts' adaptation remains largely unexplored. In this study, we examined the composition and plasticity of the bacteriomes in two sibling intertidal gastropod species, Littorina fabalis and L. obtusata, which are sympatric but differ in microhabitats. We applied 16S rRNA gene metabarcoding and shotgun sequencing to describe associated microbial communities and their spatial and temporal variation. A significant drop in the intestinal bacteriome diversity was revealed during the cold season, which may reflect temperature-related metabolic shifts and changes in snail behavior. Importantly, there were significant interspecies differences in the gut bacteriome composition in summer but not in autumn. The genera Vibrio, Aliivibrio, Moritella and Planktotalea were found to be predominantly associated with L. fabalis, while Granulosicoccus, Octadecabacter, Colwellia, Pseudomonas, Pseudoalteromonas and Maribacter were found to be mostly associated with L. obtusata. Based on these preferential associations, we analyzed the metabolic pathways' enrichment. We hypothesized that the L. obtusata gut bacteriome contributes to decomposing algae and detoxifying polyphenols produced by fucoids. Thus, differences in the sets of associated bacteria may equip their closely phylogenetically related hosts with a unique ability to occupy specific micro-niches. [ABSTRACT FROM AUTHOR]

We studied the reproductive behaviour associated with mate choice by size in three species of White Sea molluscs of the genus Littorina—L. saxatilis, L. obtusata and L. fabalis. Three behavioural patterns were revealed after a comparative analysis of the behaviour of the male snails. The males of L. obtusata, regardless of their size, could mate with all females but preferred to mate with larger ones. The males of L. fabalis chose females of their own size or larger. The males of L. saxatilis mated with females of approximately their own size. In the first case (L. obtusata), there is a free exchange of genes between all the size groups in the population. In the second case (L. fabalis), the gene transfer from small males to females of any size is free but the gene transfer from large males to small females is limited. In the third case (L. saxatilis), the gene flow is limited between groups of individuals of dramatically different size. Thus, in the case of L. saxatilis and L. fabalis, to varying degrees restriction of random mating associated with size assortativity in choosing a mating partner was expressed, which under certain circumstances (divergence of individuals in different econiches, the presence of ecotypes differing in size, lack of stronger natural selection in the other direction) can favour sympatric speciation. The males of all the three periwinkles' species preferred larger females but sexual selection in favour of larger females was found only in L. obtusata. In the other two species, there were no statistically significant size differences between copulating and non-copulating females. This appears to be due to the occurrence of size-assortative mating in these species, which limits the ability of males to choose larger female partners. [ABSTRACT FROM AUTHOR]

Microbes can play a prominent role in the evolution of their hosts, facilitating adaptation to various environments and promoting ecological divergence. The Wave and Crab ecotypes of the intertidal snail Littorina saxatilis is an evolutionary model of rapid and repeated adaptation to environmental gradients. While patterns of genomic divergence of the Littorina ecotypes along the shore gradients have been extensively studied, their microbiomes have been so far overlooked. The aim of the present study is to start filling this gap by comparing gut microbiome composition of the Wave and Crab ecotypes using metabarcoding approach. Since Littorina snails are micro‐grazers feeding on the intertidal biofilm, we also compare biofilm composition (i.e. typical snail diet) in the crab and wave habitats. In the results, we found that bacterial and eukaryotic biofilm composition varies between the typical habitats of the ecotypes. Further, the snail gut bacteriome was different from outer environments, being dominated by Gammaproteobacteria, Fusobacteria, Bacteroidia and Alphaproteobacteria. There were clear differences in the gut bacterial communities between the Crab and the Wave ecotypes as well as between the Wave ecotype snails from the low and high shores. These differences were both observed in the abundances and in the presence of different bacteria, as well as at different taxonomic level, from bacterial OTU's to families. Altogether, our first insights show that Littorina snails and their associated bacteria are a promising marine system to study co‐evolution of the microbes and their hosts, which can help us to predict the future for wild species in the face of rapidly changing marine environments. [ABSTRACT FROM AUTHOR]

Renicolid digeneans parasitize aquatic birds. Their intramolluscan stages develop in marine and brackish-water gastropods, while metacercariae develop in molluscs and fishes. The systematics of renicolids is poorly developed, their life cycles are mostly unknown, and the statuses of many species require revision. Here, we establish based on integrated morphological and molecular data that adult renicolids from gulls Larus argentatus and Larus schistisagus and sporocysts and cercariae of Cercaria parvicaudata from marine snails Littorina spp. are life-cycle stages of the same species. We name it Renicola parvicaudatus and synonymized with it Renicola roscovitus. An analysis of the cox1 gene of R. parvicaudatus from Europe, North America and North Asia demonstrates a low genetic divergence, suggesting that this species has formed quite recently (perhaps during last glacial maximum) and that interregional gene flow is high. In Littorina saxatilis and L. obtusata from the Barents Sea, molecular analysis has revealed intramolluscan stages of Cercaria littorinae saxatilis VIII, a cryptic species relative to R. parvicaudatus. In the molecular trees, Renicola keimahuri from L. schistisagus belongs to another clade than R. parvicaudatus. We show that the species of this clade have cercariae of Rhodometopa group and outline morphological and behavioural transformations leading from xiphidiocercariae to these larvae. Molecular analysis has revealed 3 main phylogenetic branches of renicolids, differing in structure of adults, type of cercariae and host range. Our results elucidate the patterns of host colonization and geographical expansion of renicolids and pave the way to the solution of some long-standing problems of their classification. [ABSTRACT FROM AUTHOR]

Divergence of ecological niches in phylogenetically closely related species indicates the importance of ecology in speciation, especially for sympatric species are considered. Such ecological diversification provides an advantage of alleviating interspecies competition and promotes more efficient exploitation of environmental resources, thus being a basis for ecological speciation. We analyzed a group of closely related species from the subgenus Neritrema (genus Littorina, Caenogastropoda) from the gravel‐bouldery shores. In two distant sites at the Barents and Norwegian Sea, we examined the patterns of snail distribution during low tide (quantitative sampling stratified by intertidal level, presence of macrophytes, macrophyte species, and position on them), shell shape and its variability (geometric morphometrics), and metabolic characteristics (metabolomic profiling). The studied species diversified microbiotopes, which imply an important role of ecological specification in the recent evolution of this group. The only exception to this trend was the species pair L. arcana / L. saxatilis, which is specifically discussed. The ecological divergence was accompanied by differences in shell shape and metabolomic characteristics. Significant differences were found between L. obtusata versus L. fabalis and L. saxatilis / L. arcana versus L. compressa both in shell morphology and in metabolomes. L. saxatilis demonstrated a clear variability depending on intertidal level which corresponds to a shift in conditions within the occupied microhabitat. Interestingly, the differences between L. arcana (inhabiting the upper intertidal level) and L. compressa (inhabiting the lower one) were analogous to those between the upper and lower fractions of L. saxatilis. No significant level‐dependent changes were found between the upper and lower fractions of L. obtusata, most probably due to habitat amelioration by fucoid macroalgae. All these results are discussed in the contexts of the role of ecology in speciation, ecological niche dynamics and conservatism, and evolutionary history of the Neritrema species. [ABSTRACT FROM AUTHOR]

The presence of shell bands is common in gastropods. Both the marine snails Littorina fabalis and Lttorina saxatilis are polymorphic for this trait. Such polymorphism would be expected to be lost by the action of genetic drift or directional selection, but it appears to be widespread at relatively constant frequencies. This suggests it is maintained by balancing selection on the trait or on a genetically linked trait. Using long time series of empirical data, we compared potential effects of genetic drift and negative frequency‐dependent selection (NFDS) in the two species. The contribution of genetic drift to changes in the frequency of bands in L. fabalis was estimated using the effective population size estimated from microsatellite data, while the effect of genetic drift in L. saxatilis was derived from previously published study. Frequency‐dependent selection was assessed by comparing the cross‐product estimator of fitness with the frequency of the polymorphism across years using a regression analysis. Both studied species showed patterns of NFDS. In addition, in L. fabalis, contributions from genetic drift could explain some of the changes in banding frequency. Overdominance and heterogeneous selection did not fit well to our data. The possible biological explanations resulting in the maintenance of the banding polymorphism are discussed. [ABSTRACT FROM AUTHOR]

Trematodes of the genus Himasthla are usual parasites of coastal birds in nearshore ecosystems of northern European seas and the Atlantic coast of North America. Their first intermediate hosts are marine and brackish-water gastropods, while second intermediate hosts are various invertebrates. We analysed sequences of partial 28S rRNA and nad1 genes and the morphology of intramolluscan stages, particularly cercariae of Himasthla spp. parasitizing intertidal molluscs Littorina spp. in the White Sea, the Barents Sea and coasts of North Norway and Iceland. We showed that only three Himasthla spp. are associated with periwinkles in these regions. Intramolluscan stages of H. elongata were found in Littorina littorea, of H. littorinae, in both L. saxatilis and L. obtusata, and of Cercaria littorinae obtusatae, predominantly, in L. obtusata. Other Himasthla spp. previously reported from Littorina spp. in North Atlantic are either synonymous with one of these species or described erroneously. Based on a comparison of newly generated 28S rDNA sequences with GenBank data, rediae and cercariae of C. littorinae obtusatae were identified as belonging to H. leptosoma. Some previously unknown morphological features of young and mature rediae and cercariae of the three Himasthla spp. are described. We provide a key to the rediae and highlight characters important for identification of cercariae. Genetic diversity within the studied species was only partially determined by their specificity to the molluscan host. The nad1 network constructed for H. leptosoma lacked geographical structure, which is explained by a high gene flow owing to highly vagile definitive hosts, shorebirds. [ABSTRACT FROM AUTHOR]

Sympatric coexistence of recently diverged species raises the question of barriers restricting the gene flow between them. Reproductive isolation may be implemented at several levels, and the weakening of some, e.g. premating, barriers may require the strengthening of the others, e.g. postcopulatory ones. We analysed mating patterns and shell size of mates in recently diverged closely related species of the subgenus Littorina Neritrema (Littorinidae, Caenogastropoda) in order to assess the role of premating reproductive barriers between them. We compared mating frequencies observed in the wild with those expected based on relative densities using partial canonical correspondence analysis. We introduced the fidelity index (FI) to estimate the relative accuracy of mating with conspecific females and precopulatory isolation index (IPC) to characterize the strength of premating barriers. The species under study, with the exception of L. arcana, clearly demonstrated preferential mating with conspecifics. According to FI and IPC, L. fabalis and L. compressa appeared reliably isolated from their closest relatives within Neritrema. Individuals of these two species tend to be smaller than those of the others, highlighting the importance of shell size changes in gastropod species divergence. L. arcana males were often found in pairs with L. saxatilis females, and no interspecific size differences were revealed in this sibling species pair. We discuss the lack of discriminative mate choice in the sympatric populations of L. arcana and L. saxatilis, and possible additional mechanisms restricting gene flow between them. [ABSTRACT FROM AUTHOR]

Abstract The rough periwinkle Littorina saxatilis is characterised for showing shell phenotypic variation associated to environmental clines and this is found in parallel across shores. In order to disentangle the additive genetic effects and the environmental factors involved in this variation, we analysed natural populations subjected to a gradient in wave exposure and crab predation in 3 different Rias (primary estuaries) from Galicia (NW Spain). In addition to previously described "Crab" and "Wave" ecotypes, we studied a new phenotype ("Crab+") associated to the most sheltered habitats of the gradient. We obtained estimates of heritability and Q st (between population genetic differentiation) in embryos and P st (between population phenotypic differentiation) in adults for shell morphology traits. Heritability estimates were close to one for shell size and relative shell aperture, suggesting a low impact of environmental factors in embryos. Estimates of Q st between ecotypes were significant and typically much higher than published F st estimates from neutral markers, indicating a role for selection on shaping additive genetic variation. Estimates of P st between ecotypes for adults were substantially larger than embryos´ Q st , suggesting also a role of phenotypic plasticity in ecotype differentiation. Our results suggest that P st vs F st comparisons should be taken with caution because they might not reflect real additive genetic effects, but P st vs Q st comparisons could shed light into the role of phenotypic plasticity. Highlights • Morphometric analysis of Littorina saxatilis is carried out in adults and embryos. • We compared phenotypic (P ST) and genetic divergence (Q ST , F ST) between ecotypes. • High heritabilities are shown for shell traits (size and shell aperture) in embryos. • Q ST vs F ST comparisons show a role for natural selection in ecotype differentiation. [ABSTRACT FROM AUTHOR]

Abstract Dislodgment, in particular by breaking waves, is a widely acknowledged source of stress, if not mortality, in rocky shores intertidal gastropods. This raises the question of understanding if snails dislodged from a specific habitat have the ability to navigate back to their original substrate, and how this is achieved. In this context, we investigated the consequences of dislodgement on a population of Littorina littorea inhabiting two proximate habitats (a seawall ranging from the low to the high intertidal zone and the related low intertidal rocky platform) located on an isolated offshore reef over a three-month period. Using tagged undisturbed, disturbed, translocated and transplanted L. littorea , we assessed a potential tidal height preference and found that L. littorea showed a clear tendency to remain on their original tidal height, with recapture rates ranging between 100% and 88%, respectively 1 and 78 days after dislodgment. We subsequently measured the recapture rates of individuals dislodged from three distinct tidal levels on the seawall and from a horizontal platform. Dislodged L. littorea showed a clear tendency to return to their original tidal level, with return rates ranging between 32 and 48% 78 days after dislodgement; snails from the horizontal platform were never found on the proximate seawall. In addition, high frequency observations conducted on individually tagged dislodged L. littorea during the first 8 successive low tides of our survey indicate that individuals originating from the seawall consistently showed higher movement rates and directionality towards the seawall, while individuals originating from the platform showed no directionality and reduced migration rate. These results show that dislodged L. littorea navigate back to their original substrate and tidal height. Putative processes include a combination of geotaxis, chemotaxis and rheotaxis. At relatively short time scales (i.e. days to months), L. littorea show both tidal and habitat partitioning that is distinct from the movement patterns previously reported in littorinids over a range of temporal scales. [ABSTRACT FROM AUTHOR]

Littorina Férussac, 1822 is an abundant genus of small gastropods found in the upper littoral zone of rocky seashores worldwide. Although ecologically important, shell-based species identification in this genus is challenging due to phenotypic variation in shell morphology and lack of diagnostic characters among morphologically similar species. In this study, we revised the taxonomy of Korean Littorina species using morphological characters (shell and radula) and cox1 mitochondrial DNA sequences for three Korean species: L. brevicula, L. sitkana, and L. horikawai. Results suggest that L. sitkana was erroneously reported as L. kasatka in a previous study. A new record for Littorina horikawai (Matsubayashi & Habe in Habe, 1979), previously unknown from Korea, is described, which can be distinguished from L. sitkana by the presence of alternating white and brown spiral ribs on each whorl. Comparison of the mtDNA cox1 gene sequences shows very low intraspecific variation even between geographically distant populations. A phylogenetic tree supports a close relationship between L. horikawai and L. sitkana, consistent with earlier phylogenetic studies. [ABSTRACT FROM AUTHOR]

The evolution of complement system in invertebrates is poorly investigated. While the repertoire of complement genes in several Ecdysozoa lineages is found substantially different from that of Deuterostomia, the composition and function of the complement in the second protostome lineage, Lophotrochozoa, remains unclear. Here we report the general description of new transcriptomic data on the common periwinkle, Littorina littorea , and trace the evolutionary trajectories of the ancestral proto-complement repertoire. The repertoire is defined as immune cascade providing the minimum set of C3-associated molecules required for C3b amplification, opsonization of the targets and their phagocytosis: thioester protein (TEP) C3, serine protease C2/factor B (Bf) and complement receptors (CR). The reference transcriptome of L. littorea was built from the dual-species RNA-seq experiment with the periwinkle and its tissue digenean parasite Himasthla elongata . Five TEPs, including the ortholog of the C3, are found expressed in the in the mollusk's inflamed tissues. The homolog of the complement receptors CR1/CR2 is also expressed, however the ortholog of Bf is not. The extensive phylogenetic analysis showed that the C3 ortholog and the complement receptors are retained in all key lophotrochozoan taxa: Mollusca, Annelida and Brachiopoda. However, the Bf ortholog was lost at least three times independently in different lineages: i) Cephalopoda, ii) a common ancestor of all Gastropoda and iii) one of the Annelida lineage, Clitellata. Both C3 and Bf molecules were retained in bivalve species, brachiopods and annelid worms from the Polychaeta lineage. Hypothetically, the function of the lost Bf in these animals can be compensated by Factor L (Lf) – the serine protease first found in L. littorea and homologous to both, the Bf and the arthropod factor C (Cf). The contrast differences in proto-complement repertoire between the sister mollusk’ taxa, Bivalvia and Gastropoda (the conserved and modified sets, respectively), can underlie differences in their susceptibility to digenean infection. [ABSTRACT FROM AUTHOR]

With both global surface temperatures and the incidence and intensity of extreme temperature events projected to increase, the assessment of species' sensitivity to chronic and acute changes in temperature has become crucial. Sensitivity predictions are based predominantly on adult responses, despite the fact that early life stages may be more vulnerable to thermal challenge. Here, we compared the sensitivity of different life history stages of the intertidal gastropod Littorina obtusata using thermal death time curves, which incorporate the intensity and duration of heat stress, and used these to calculate upper critical thermal limits (CTmax) and sensitivity to temperature change (z). Early (larval) life stages had both a lower CTmax and a lower z than adults, suggesting they are less good at withstanding short-term extreme thermal challenges but better able to survive moderate temperatures in the long term. This result supports the predicted trade-off between acute and chronic tolerance to thermal stress, and is consistent with the different thermal challenges that these stages encounter in the intertidal zone. We conclude that different life history stages employ different thermal strategies that may be adaptive. Our findings caution against the use of predictions of the impact of global warming that are based on only adult responses and, hence, which may underestimate vulnerability. [ABSTRACT FROM AUTHOR]

With both global surface temperatures and the incidence and intensity of extreme temperature events projected to increase, the assessment of species' sensitivities to chronic and acute changes in temperature has become crucial. Sensitivity predictions are based predominantly on adult responses, despite the fact that early life stages may be more vulnerable to thermal challenge. Here, we compared the sensitivity of different life history stages of the intertidal gastropod Littorina obtusata using thermal death time curves, which incorporate the intensity and duration of heat stress, and used these to calculate upper critical thermal limits (CTmax) and sensitivity to temperature change (z). Early (larval) life stages had both a lower CTmax and z than adults, suggesting they are less good at withstanding short term extreme thermal challenges, but better able to survive moderate temperatures in the long term. This result supports the predicted trade-off between acute and chronic tolerance to thermal stress, and is consistent with the different thermal challenges that these stages encounter in the intertidal zone. We conclude that different life history stages employ different thermal strategies that may be adaptive. Our findings caution against the use of predictions of the impact of global warming that are based on only adult responses and, hence, which may underestimate vulnerability. [ABSTRACT FROM AUTHOR]

Aim: The present study aims to focus on the role of common marine snails (Littorina littorea) as a vector for some trematode parasites. Materials and Methods: A total of 327 marine water L. littorea snails were collected during the summer of 2016 from a Qarun lake in the EL-Fayoum Governorate, Egypt. The snails were investigated for infection by trematode parthenitae through induction of cercarial shedding by exposure to light and crushing the snails. The species were stored in Search Laboratory of Zoology Department, Faculty of Science, Fayoum University. Results: Three species of Strigeid littorina cercaria were identified from the infected snails. They are described here and they identified in relation to close-up morphological features and linked to its snail hosts. They give the following names: Cercaria strigeid littorina type 1, C. strigeid littorina type 2, and C. strigeid littorina type 3. The incidence of infection by these cercariae was 33%, 25.7%, and 2.4%, respectively. Conclusion: This study is clarifying the importance of this marine snail as intermediate hosts for new trematode species. [ABSTRACT FROM AUTHOR]

There has been a significant increase in the literature regarding the effects of warming and acidification on the marine ecosystem. To our knowledge, there is very little information on the potential effects of both combined stressors on marine grazer-seaweed interactions. Here, we evaluated, for the first time several phenotypic responses (e.g periwinkle survival, condition index, consumption rates, seaweed photosynthetic activity and oxidative stress) of the temperate periwinkle Littorina obtusata (grazer) and the brown seaweed Ascophyllum nodosum (prey) to such climate change-related variables, for 15 days. Increased temperature (22 °C, pH 8.0) elicited a significant lethal effect on the periwinkle within a short-term period (mortality rate > 90%). Acidification condition (18 °C, pH 7.6) was the one that showed lower mortality rates (≈20%), reflected by lower impact on periwinkle fitness and consumption rates. Under a scenario of increased temperature and lowered pH the antioxidant defences of L. obtusata seemed to be supressed increasing the risk of peroxidative damage. The seaweed evidenced signs of cellular damage under such conditions. These results suggest that: i) lower pH per se seems to benefit the interaction between grazer and seaweed while, ii) a combined scenario of increased temperature and lowered pH may be negative for the interaction, due to the unbalance between periwinkle mortality rates and consumption rates. But most importantly, since grazing often plays an important role on structuring natural communities, such predator-prey disturbances can elicit cascading effects on the remaining community structure and functioning of the temperate rocky-shore ecosystems. [ABSTRACT FROM AUTHOR]

Cadmium (Cd) is one of the most harmful metals, being toxic to most animal species, including marine invertebrates. Among marine gastropods, the periwinkle (Littorina littorea) in particular can accumulate high amounts of Cd in its midgut gland. In this organ, the metal can elicit extensive cytological and tissue-specific alterations that may reach, depending on the intensity of Cd exposure, from reversible lesions to pathological cellular disruptions. At the same time, Littorina littorea expresses a Cd-specific metallothionein (MT) that, due to its molecular features, expectedly exerts a protective function against the adverse intracellular effects of this metal. The aim of the present study was, therefore, to assess the time course of MT induction in the periwinkle's midgut gland on the one hand, and cellular and tissue-specific alterations in the digestive organ complex (midgut gland and digestive tract) on the other, upon exposure to sub-lethal Cd concentrations (0.25 and 1 mg Cd/L) over 21 days. Depending on the Cd concentrations applied, the beginning of alterations of the assessed parameters followed distinct concentration-dependent and time-dependent patterns, where the timeframe for the onset of the different response reactions became narrower at higher Cd concentrations compared to lower exposure concentrations. [ABSTRACT FROM AUTHOR]

The paper describes the conditions of formation of large accumulative forms on the southeastern coast of the Baltic Sea and presents the results of a comparative analysis of these structures. It has been established that Baltic barrier-lagoon systems have a complex geological and geomorphologic structure. Sandy barriers, which are, in general, marine accumulative formations, often comprise various fragments of different genesis (moraine remnants, glaciolacustrine and deltaic plains, etc.). The formation and development of large accumulative forms in the southeastern Baltic occurred against the background of sea level fluctuations in the Littorina time following a unified scenario for the entire coast. [ABSTRACT FROM AUTHOR]

After the resolution of the 3D structure of the Cd9-aggregate of the Littorina littorea metallothionein (MT), we report here a detailed analysis of the metal binding capabilities of the wild type MT, LlwtMT, and of two truncated mutants lacking either the N-terminal domain, Lltr2MT, or both the N-terminal domain, plus four extra flanking residues (SSVF), Lltr1MT. The recombinant synthesis and in vitro studies of these three proteins revealed that LlwtMT forms unique M9-LlwtMT complexes with Zn(II) and Cd(II), while yielding a complex mixture of heteronuclear Zn,Cu-LlwtMT species with Cu(I). As expected, the truncated mutants gave rise to unique M6-LltrMT complexes and Zn,Cu-LltrMT mixtures of lower stoichiometry with respect to LlwtMT, with the SSVF fragment having an influence on their metal binding performance. Our results also revealed a major specificity, and therefore a better metal-coordinating performance of the three proteins for Cd(II) than for Zn(II), although the analysis of the Zn(II)/Cd(II) displacement reaction clearly demonstrates a lack of any type of cooperativity in Cd(II) binding. Contrarily, the analysis of their Cu(I) binding abilities revealed that every LlMT domain is prone to build Cu4-aggregates, the whole MT working by modules analogously to, as previously described, certain fungal MTs, like those of C. neoformans and T. mesenterica. It is concluded that the Littorina littorea MT is a Cd-specific protein that (beyond its extended binding capacity through an additional Cd-binding domain) confers to Littorina littorea a particular adaptive advantage in its changeable marine habitat. [ABSTRACT FROM AUTHOR]

The Littorina Sea stage (past c. 7000 years) development of the northern, and its interactions with the central Baltic Sea have been influenced by spatially different but in the north very strong glacio-isostatic uplift. Here we investigate the impact of the isostatic readjustment on the northern Baltic Sea environment and on the water exchange with the central Baltic Sea using geochemical and microfossil records from well-dated sediment cores of the Bothnian Sea/Bay that are compared to sedimentary records of the Gotland Basin and Landsort Deep. Benthic foraminifera and manganese carbonate layers in the sediments from northern Baltic Sea sub-basins imply oxic marine intrusions from c. 7.0 cal. ka BP. Related water-column stratification together with enhanced primary productivity caused oxygen-depleted bottom waters and deposition of organic-rich laminated sediments during the Holocene Thermal Maximum. Comparable sedimentary/geochemical records in early Littorina Sea sediments of e.g. the eastern Gotland Basin therefore imply a low impact of isostatic uplift on the northern Baltic Sea environment. A climate-controlled decline in hypoxia induced a gradual deposition of rather homogenous sediments in the central and northern Baltic Sea during the mid-Littorina Sea stage. Progressive isostatic uplift caused a diverging environmental development in both basins during the mid- to late Littorina Sea stage. The Åland sill shoaled markedly allowing only strong inflows to enter the northern Baltic Sea as late as 2.3 cal. ka BP. The restricted deep-water exchange with the central Baltic Sea probably resulted in a pronounced freshening and oxic bottom waters. This, together with an uplift-related sediment re-deposition induced accumulation of rather homogenous sediments throughout the remaining Littorina Sea stage. By contrast, a climate-controlled depositional environment in the central Baltic Sea during this time caused the formation of homogeneous sediments during cold phases and laminated sediments during warm phases. [ABSTRACT FROM AUTHOR]

The idea of parasitic systems, formulated by V.N. Beklemishev 70 years ago, represents a conceptual tool for the analysis of populational and biocoenotic roles of parasites. The questions concerning longterm stable persistence of host-parasite systems in communities can be discussed meaningfully only within this concept. Importantly, the set of terms elaborated within the parasitic system concept is applicable not only to parasitology, but also contributes to the general knowledge of life cycles of organisms and differences in the environment. This concept provides an opportunity for comprehensive analysis of systems, based on any type of stable biocoenotic interactions in the community (predation, commensalism, competition, etc.). Trematode-based parasitic systems, involving populations of intertidal mollusks of the genus Littorina, allow demonstrating how the strong 'negative' effect of parasites on hosts at the individual level (complete parasitic castration) can be compensated at the population level. Such compensation functions as a prerequisite for maintaining long-term stable interactions between populations of parasites and their hosts within parasitic system (the ecosystem, biocoenotic level). [ABSTRACT FROM AUTHOR]

The hypothesis on monogenic inheritance of white longitudinal shell bands in the snail Littorina obtusata was tested. The data showed that a single gene with two alleles was responsible for the presence of the bands on the snail shell. The presence of bands was the dominant character. [ABSTRACT FROM AUTHOR]

L. obtusata and Trivia sp. shells were systematically used for personal ornamentation by groups who occupied northern Iberia during the Mesolithic. The shells from El Mazo and El Toral III (Asturias, Spain) offer a unique opportunity for investigating raw material procurement, selection strategies, and manufacture processes developed by Asturian Mesolithic societies for beads production. By combining taphonomic, morphometric, and microscopic analyses, our results show that the shells were introduced and transformed in the caves. Mollusk consumption at the sites and bead manufacture indicate that the sites were occupied for both economic and symbolic purposes. The use of similar shell beads by contemporaneous societies located in different environments (coastal and interior) and relying on drastically different subsistence strategies mirrors the complex circulation network developed by Mesolithic foraging societies. [ABSTRACT FROM AUTHOR]

Littorina brevicula is one of the most common gastropods in the supralittoral zone around Japan. The northernmost population of this species is around Hokkaido and the determinant of this northern limit is likely seawater and air temperature. To reconstruct an evolutionary history of this species, we investigated genetic differentiation among 12 populations (three from Hokkaido, six from Honshu and three from Kyushu) using a mitochondrial DNA marker (partial sequence of the NADH dehydrogenase 6 gene). The haplotype network showed shallow genetic divergence within the species, suggesting a bottleneck followed by population expansion. One major haplotype that occurred in 70.5% of all individuals examined was the most frequent in every population sampled. A second major haplotype was abundant around Kyushu but not found in Hokkaido. This skewed haplotype distribution resulted in significant genetic differentiation along the north-south axis of Japan. The importance of the southern clade, which included the second major haplotype, was supported by population genetic analyses of datasets that excluded either the southern clade or the northern clades. The north-south differentiation remained when datasets that excluded the northern clades were used, but disappeared when datasets that excluded the southern clade were used. The combined evidence of shallow divergence and the north-south population structure suggests that the L. brevicula population around Japan once declined and then expanded and colonized northward. Although the time of population reduction and recolonization could not be precisely estimated, the observation that this species is absent further north in Japan suggests that it would have been unable to survive in northern Japan during the last glacial maximum (LGM) and therefore recolonization likely occurred after the LGM, probably from south to north. [ABSTRACT FROM AUTHOR]

Calidris maritima (Purple Sandpiper) is a shorebird found along northern coasts of Europe and eastern North America during the winter. While its winter diet is well known in Europe, the only North American study examining diet included 5 individuals and suggested a narrow diet focused on Littorina spp. (periwinkles). We examined the diet of Purple Sandpipers wintering at several sites in Nova Scotia, Canada, during 2013, and found that the birds consumed a broad variety of items, though principally marine invertebrates, dominated by periwinkles and Mytilus edulis (Blue Mussel), but also many arthropods. Our study confirms the importance of mollusks in the diet of this species and also indicates that Purple Sandpipers in North America have a similar winter diet to those studied in Europe. [ABSTRACT FROM AUTHOR]

The ingestion of microplastics has been shown for a great variety of marine organisms. However, benthic marine mesoherbivores such as the common periwinkle Littorina littorea have been largely disregarded in studies about the effects of microplastics on the marine biota, probably because the pathway for microplastics to this functional group of organisms was not obvious. In laboratory experiments we showed that the seaweed Fucus vesiculosus retains suspended microplastics on its surface. The numbers of microplastics that adhered to the algae correlated with the concentrations of suspended particles in the water. In choice feeding assays L. littorea did not distinguish between algae with adherent microplastics and clean algae without microplastics, indicating that the snails do not recognize solid nonfood particles in the submillimeter size range as deleterious. In periwinkles that were feeding on contaminated algae, microplastics were found in die stomach and in the gut. However, no microplastics were found in the midgut gland, which is the principle digestive organ of gastropods. Microplastics in the fecal pellets of the periwinkles indicate diat the particles do not accumulate rapidly inside the animals but are mostly released with the feces. Our results provide the first evidence that seaweeds may represent an efficient pathway for microplastics from the water to marine benthic herbivores. [ABSTRACT FROM AUTHOR]

Local adaptation may optimize an organism's investment in defenses in response to the risk of infection by spatially heterogeneous parasites and other natural enemies. However, local adaptation may be constrained if recruitment is decoupled from selective pressure experienced by the parent generation. We predicted that the ability of three intertidal littorinid snail species to defend against trematode parasites would depend on prior levels of population exposure to parasites and on larval dispersal mode, a proxy for population openness. In a common garden experiment, for two snail species with direct development and localized recruitment ( Littorina obtusata and Littorina saxatilis), hosts from sites with high trematode infection risk were less susceptible to infection than hosts from low-risk sites. However, this relationship was not apparent for a third host species with broadcast larvae ( Littorina littorea), suggesting that broad larval dispersal can impede local adaptation; alternatively, the lack of response in this species could owe to other factors that limited experimental infection in this host. Our findings support that locally recruiting hosts can adapt their defenses to scale with localized infection risk. [ABSTRACT FROM AUTHOR]

Our knowledge of the effects of consumer species loss on ecosystem functioning is limited by a paucity of manipulative field studies, particularly those that incorporate inter-trophic effects. Further, given the ongoing transformation of natural habitats by anthropogenic activities, studies should assess the relative importance of biodiversity for ecosystem processes across different environmental contexts by including multiple habitat types. We tested the context-dependency of the effects of consumer species loss by conducting a 15-month field experiment in two habitats (mussel beds and rock pools) on a temperate rocky shore, focussing on the responses of algal assemblages following the single and combined removals of key gastropod grazers (Patella vulgata, P. ulyssiponensis, Littorina littorea and Gibbula umbilicalis). In both habitats, the removal of limpets resulted in a larger increase in macroalgal richness than that of either L. littorea or G. umbilicalis. Further, by the end of the study, macroalgal cover and richness were greater following the removal of multiple grazer species compared to single species removals. Despite substantial differences in physical properties and the structure of benthic assemblages between mussel beds and rock pools, the effects of grazer loss on macroalgal cover, richness, evenness and assemblage structure were remarkably consistent across both habitats. There was, however, a transient habitat-dependent effect of grazer removal on macroalgal assemblage structure that emerged after three months, which was replaced by non-interactive effects of grazer removal and habitat after 15 months. This study shows that the effects of the loss of key consumers may transcend large abiotic and biotic differences between habitats in rocky intertidal systems. While it is clear that consumer diversity is a primary driver of ecosystem functioning, determining its relative importance across multiple contexts is necessary to understand the consequences of consumer species loss against a background of environmental change. [ABSTRACT FROM AUTHOR]

Species face multiple selective pressures that may require opposing responses to mitigate. On rocky shorelines, fitness of the intertidal snail Littorina littorea is determined by both parasitism and predation. We experimentally demonstrated that L. littorea was at greatest risk of infection from trematode parasites high in the intertidal zone where it was in closest proximity to abundant gull feces (the vector for the snail's parasites). However, because of extreme, size-selective predation pressure at low tidal elevations, small snails often live high in the intertidal until they have grown sufficiently large. By prolonging their exposure to infection higher on the shore, ontogenetic responses to predation risk accentuate parasite risk. Counterintuitively, snails exhibited the highest trematode prevalence at the lowest tidal elevations where they had almost no risk of contracting infection. By carrying contracted infections into the lowest tidal zones, the larger, predation-resistant snails invert hotspots of infection risk and prevalence, underscoring that size-dependent selection pressures can decouple infection process and pattern even over small scales. [ABSTRACT FROM AUTHOR]

The ancient Ventspils Lagoon is an area where human settlements have been located on former coastal landforms and sediments. However, the chronology of its development has so far not been studied in detail. Since the ancient lagoon is dominated by sand deposition, optically stimulated luminescence dating (OSL) was used to determine when the ancient coast was formed. Twenty-one OSL samples gave a wide age range, between 8.57 ± 0.65 ka and 73.8 ± 8.3 ka. Twelve of the ages fall in the Late Glacial and Holocene and document the development of the lagoon between the Baltic Ice Lake stage (∼16 ka) and the Littorina Sea stage (∼8 ka). The new dates correspond to some extent with previously modelled shorelines for the Ventspils Lagoon and to radiocarbon ages from archaeological sites in the area. One date, 16.0 ± 1.3 ka, falls in the period before the Baltic Ice Lake and represents either a very early Baltic Ice Lake stage or, less likely, the Venta glacial lake prior to the Baltic Ice Lake. One date, 5.09 ± 0.42, is likely underestimated. Seven samples yielded dates between 21.5 ± 1.5 ka and 73.8 ± 8.3 ka, which corresponds to the Early and Middle Weichselian in the Baltic region. The samples in this latter group are likely overestimating the true age due to insufficient exposure to sunlight during deposition. • Depositional time frame of the ancient Ventspils Lagoon. • Part of OSL datings fall in interval between the Baltic Ice Lake stage and post-maximum Litorina Sea stage. • Partial correspondence of OSL and radiocarbon datings with the previously modelled shorelines for the Ventspils Lagoon. • Coastal sediment deposition took place between 12.6 ± 2.8 ka and 5.09 ± 0.42 ka. • Older dates suffer from incomplete bleaching because of the skewness of dose distributions and large overdispersion. [ABSTRACT FROM AUTHOR]

The common periwinkle Littorina littorea was introduced in the list of models for comparative immunobiology as a representative of phylogenetically important taxon Caenogastropoda. Using Illumina sequencing technology, we de novo assembled the transcriptome of Littorina littorea hemocytes from 182 million mRNA-Seq pair-end 100 bp reads into a total of 15,526 contigs clustered in 4472 unigenes. The transcriptome profile was analyzed for presence of carbohydrate-binding molecules in a variety of architectural contexts. Hemocytes' repertoire of lectin-like proteins bearing conserved carbohydrate-recognition domains (CRDs) is highly diversified, including 11 of 15 lectin families earlier described in animals, as well as the novel members of lectin family found for the first time in mollusc species. The new molluscan lineage-specific domain combinations were confirmed by cloning and sequencing, including the fuco-lectin related molecules (FLReMs) composed of N-terminal region with no sequence homology to any known protein, a middle Fucolectin Tachylectin-4 Pentaxrin (FTP) domain, and a C-terminal epidermal growth factor (EGF) repeat region. The repertoire of lectin-like molecules is discussed in terms of their potential participation in the receptor phase of immune response. In total, immune-associated functions may be attributed to 70 transcripts belonging to 6 lectin families. These lectin-like genes show low overlap between species of invertebrates, suggesting relatively rapid evolution of immune-associated genes in the group. The repertoire provides valuable candidates for further characterization of the gene functions in mollusc immunity. [ABSTRACT FROM AUTHOR]

The responses of the osphradium in the freshwater mollusc Viviparus sp. and single osphradial neurons in the pond snail Lymnaea stagnalis to L-glutamine and L-asparagine as well as the changes in these responses under the effect of heavy metals (Cu, Cd) were recorded electrophysiologically. The functional connections of the osphradium with the identified neurons of the central pattern generator of respiratory movements and cardiac activity as well as the modification of these connections under the effect of short-term application of HgCl solution onto the snail osphradium were investigated. The heart rate (HR) in the mollusc Littorina littorea under the effect of Cu ions was registered non-invasively in long-term experiments. The dose-dependent short-term effects of heavy metals injected into the mantle cavity on cardiac activity in intact molluscs and the absence of CR changes after osphradium damage were revealed. The implication of osphradium in adaptive reactions of the cardiac system in aquatic molluscs to the environmental heavy metal pollution is suggested. The dependence of HR on the degree of accumulation of copper ions in the mollusc tissues was detected. The results obtained are essential for unraveling neural mechanisms and pathways allowing heavy metals to affect the functional state of hydrobionts, particularly, the cardiac activity frequency characteristics of which are widely used as informative biomarkers to assess physiological condition of aquatic invertebrates. [ABSTRACT FROM AUTHOR]

We investigated in a gastropod mollusk Littorina obtusata ( L. obtusata) the inheritance of back-ground shell coloration of the shell, which arises on the basis of three pigments: purple, orange, and yellow. We found that the hypothesis on polyallelic inheritance, as in the genus Cepaea, cannot explain the inheritance of shell color in periwinkles. We demonstrated that a separate genetic system is responsible for incorporation of each pigment into the shell. The composition of these genetic systems includes at least two genes each in the case of the yellow and purple pigments. Our analysis shows that caution should be applied when extending the results obtained in the studies of the Cepaea genus to the other species of gastropods. [ABSTRACT FROM AUTHOR]

All semelparous short-lived gastropods studied so far for the experimental transplantation of trematode parthenitae, from one specific host to another, showed more or less successful acceptance of implanted parasites. We implanted echinostomatid rediae, Himasthla elongata, to the specific iteroparous long-living host, coenogastropod Littorina littorea. Using simple and low-invasive implantation techniques we have tested 680 snails injected with 75 redia microhemipopulations (MHP) harvested from naturally infected snails. Neither young nor mature rediae were able to survive in the recipient periwinkles in the course of 30 days post-implantation. A strong immune response from the host was already evident within the first week after implantation: initial inactivation of motile rediae with toxic humoral immune factors, following encapsulation of the implants and increased hemocyte counts. In contrast, rediae from the same MHPs showed perfect survival rates in primary in vitro axenic cultures. The failure of the transplantation experiments is explained in terms of the compatibility matching phenotype model. In the studied host-parasite combination all periwinkles are potentially susceptible and all rediae MHPs are potentially infective, however the probability of the compatible phenotypes matching is virtually low. Low investment in L. littorea annual reproduction would result in increased investment in self maintenance and immune mechanisms, causing the general resistance to the trematode infestation. Presumably, this resistance is relatively higher in long-lived iteroparous gastropods compare to semelparous short-lived mollusks such as pulmonates. [ABSTRACT FROM AUTHOR]

Background: Array comparative genomic hybridization (aCGH) is commonly used to screen different types of genetic variation in humans and model species. Here, we performed aCGH using an oligonucleotide gene-expression array for a non-model species, the intertidal snail Littorina saxatilis. First, we tested what types of genetic variation can be detected by this method using direct re-sequencing and comparison to the Littorina genome draft. Secondly, we performed a genome-wide comparison of four closely related Littorina species: L. fabalis, L. compressa, L. arcana and L. saxatilis and of populations of L. saxatilis found in Spain, Britain and Sweden. Finally, we tested whether we could identify genetic variation underlying “Crab” and “Wave” ecotypes of L. saxatilis. Results: We could reliably detect copy number variations, deletions and high sequence divergence (i.e. above 3%), but not single nucleotide polymorphisms. The overall hybridization pattern and number of significantly diverged genes were in close agreement with earlier phylogenetic reconstructions based on single genes. The trichotomy of L. arcana, L. compressa and L. saxatilis could not be resolved and we argue that these divergence events have occurred recently and very close in time. We found evidence for high levels of segmental duplication in the Littorina genome (10% of the transcripts represented on the array and up to 23% of the analyzed genomic fragments); duplicated genes and regions were mostly the same in all analyzed species. Finally, this method discriminated geographically distant populations of L. saxatilis, but we did not detect any significant genome divergence associated with ecotypes of L. saxatilis. Conclusions: The present study provides new information on the sensitivity and the potential use of oligonucleotide arrays for genotyping of non-model organisms. Applying this method to Littorina species yields insights into genome evolution following the recent species radiation and supports earlier single-gene based phylogenies. Genetic differentiation of L. saxatilis ecotypes was not detected in this study, despite pronounced innate phenotypic differences. The reason may be that these differences are due to single-nucleotide polymorphisms. [ABSTRACT FROM AUTHOR]

Interactions between grazers and autotrophs are complex, including both top-down consumptive and bottom-up facilitative effects of grazers. Thus, in addition to consuming autotrophs, herbivores can also enhance autotroph biomass by recycling limiting nutrients, thereby increasing nutrient availability. Here, we evaluated these consumptive and facilitative interactions between snails {Littorina littorea) and seaweeds (Fucus vesiculosus and Ulva lactuca) on a rocky shore. We partitioned herbivores' total effects on seaweeds into their consumptive and facilitative effects and evaluated how community context (the presence of another seaweed species) modified the effects of Littorina on a focal seaweed species. Ulva, the more palatable species, enhanced the facilitative effects of Littorina on Fucus. Ulva did not modify the consumptive effect of Littorina on Fucus. Taken together, the consumptive and facilitative effects of snails on Fucus in the presence of Ulva balanced each other, resulting in no net effect of Littorina on Fucus. In contrast, the only effect of Fucus on Ulva was to enhance consumptive effects of Littorina on Ulva. Our results highlight the necessity of considering both consumptive and facilitative effects of herbivores on multiple autotroph species in order to gain a mechanistic understanding of grazers' top-down and bottom-up roles in structuring communities. [ABSTRACT FROM AUTHOR]

Unravelling the form of selection acting on hybrids of ecotypes undergoing ecological speciation is essential to understand the mechanisms behind the evolution of reproductive isolation in the face of gene flow. Shell phenotype is known to be affected by natural selection and is involved in the fitness of the marine snail Littorina saxatilis. Here, we studied the association between shell traits and fitness in hybrids in order to determine the relative role of exogenous and endogenous selection in this hybrid zone of L. saxatilis. We show that directional selection is the predominant mode of selection among hybrids. We also show its heterogeneity, affecting different shell traits, within populations at the level of the microhabitat. Therefore, endogenous selection mechanisms are most probably lacking in this hybrid zone and exogenous barriers (pre- and post-zygotic) are possibly one of the main forces behind the evolution of barriers to gene flow between these ecologically divergent ecotypes. This study shows how this barrier might represent an important type of reproductive isolation within ecological speciation, and this should be taken into account in future studies of speciation in hybrid zones. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111, 391-400. [ABSTRACT FROM AUTHOR]

Climate change may cause ecosystems to become trophically restructured as a result of primary producers and consumers responding differently to increasing CO2 and temperature. This study used an integrative approach using a controlled microcosm experiment to investigate the combined effects of CO2 and temperature on key components of the intertidal system in the UK, biofilms and their consumers (Littorina littorea). In addition, to identify whether pre-exposure to experimental conditions can alter experimental outcomes we explicitly tested for differential effects on L. littorea pre-exposed to experimental conditions for two weeks and five months. In contrast to predictions based on metabolic theory, the combination of elevated temperature and CO2 over a five-week period caused a decrease in the amount of primary productivity consumed by grazers, while the abundance of biofilms increased. However, long-term pre-exposure to experimental conditions (five months) altered this effect, with grazing rates in these animals being greater than in animals exposed only for two weeks. We suggest that the structure of future ecosystems may not be predictable using short-term laboratory experiments alone owing to potentially confounding effects of exposure time and effects of being held in an artificial environment over prolonged time periods. A combination of laboratory (physiology responses) and large, long-term experiments (ecosystem responses) may therefore be necessary to adequately predict the complex and interactive effects of climate change as organisms may acclimate to conditions over the longer term. [ABSTRACT FROM AUTHOR]

Abstract: Thallium is a highly toxic metal whose biogeochemical behaviour in the marine environment is poorly understood. The dominant species predicted in sea water (Tl+) is also chemically and biologically analogous to the potassium ion. In this study, the accumulation of Tl(I) by the littorinid snail, Littorina littorea, was examined under carefully controlled laboratory conditions for a period of five days. The snail was exposed to sea water amended with a sub-phytotoxic concentration of Tl (5 μg L−1) and to a food source (the green macroalga, Ulva lactuca) that had been pre-contaminated by Tl in sea water at the same concentration. L. littorina accumulated Tl from sea water relative to food in a ratio of about 20:1 and there were no significant differences in Tl concentrations among the different tissues analysed (head, foot, visceral mass, gonads) for each treatment. When availabilities of Tl in each vector were compared, however, it was found that only 2% of sea water Tl was accumulated compared with about 15% of dietary Tl. Uptake of Tl appears to proceed via both the gills and gut, while at the cellular level accumulation likely involves binding with biomolecules and exchange with K+ in potassium-bearing mineralised granules. [Copyright &y& Elsevier]

Abstract: The gastropod mollusk, Littorina littorea L., is a common inhabitant of the intertidal zone along rocky coastlines of the north Atlantic. This species has well-developed anoxia tolerance and freeze tolerance and is extensively used as a model for exploring the biochemical adaptations that support these tolerances as well as for toxicological studies aimed at identifying effective biomarkers of aquatic pollution. This article highlights our current understanding of the molecular mechanisms involved in anaerobiosis and freezing survival of periwinkles, particularly with respect to anoxia-induced metabolic rate depression. Analysis of foot muscle and hepatopancreas metabolism includes anoxia-responsive changes in enzyme regulation, signal transduction, gene expression, post-transcriptional regulation of mRNA, control of translation, and cytoprotective strategies including chaperones and antioxidant defenses. New studies describe the regulation of glucose-6-phosphate dehydrogenase by reversible protein phosphorylation, the role of microRNAs in suppressing mRNA translation in the hypometabolic state, modulation of glutathione S-transferase isozyme patterns, and the regulation of the unfolded protein response. [Copyright &y& Elsevier]

Arsenic is naturally found in the tissues of marine animals, usually as the non-toxic arsenical arsenobetaine, but exposure to elevated arsenic concentrations in the environment may alter the arsenic species distribution within tissues of the organism. This study examined the arsenic species in the tissues of the marine periwinkle (Littorina littorea) along an arsenic concentration gradient in the sediment. The arsenicals in L. littorea were examined using the complementary analytical methods high performance liquid chromatography coupled with inductively coupled plasma mass spectrometry (HPLC–ICPMS) and X-ray absorption spectroscopy (XAS). Total arsenic concentrations in the periwinkle tissues ranged from 56 to 840mg·kg−1 dry weight (equivalent to 13 to 190mg·kg−1 wet weight). Inorganic arsenicals were found to be positively correlated with total arsenic concentrations (R2 =0.993) and reached 600mg·kg−1 dry weight, the highest reported to date in marine organisms. These high inorganic arsenic concentrations within this low trophic organism pose a potential toxicological risk to higher trophic consumers. [ABSTRACT FROM AUTHOR]

Hybrid zones of ecologically divergent populations are ideal systems to study the interaction between natural selection and gene flow during the initial stages of speciation. Here, we perform an amplified fragment length polymorphism ( AFLP) genome scan in parallel hybrid zones between divergent ecotypes of the marine snail Littorina saxatilis, which is considered a model case for the study of ecological speciation. Ridged-Banded ( RB) and Smooth-Unbanded ( SU) ecotypes are adapted to different shore levels and microhabitats, although they present a sympatric distribution at the mid-shore where they meet and mate (partially assortatively). We used shell morphology, outlier and nonoutlier AFLP loci from RB, SU and hybrid specimens captured in sympatry to determine the level of phenotypic and genetic introgression. We found different levels of introgression at parallel hybrid zones and nonoutlier loci showed more gene flow with greater phenotypic introgression. These results were independent from the phylogeography of the studied populations, but not from the local ecological conditions. Genetic variation at outlier loci was highly correlated with phenotypic variation. In addition, we used the relationship between genetic and phenotypic variation to estimate the heritability of morphological traits and to identify potential Quantitative Trait Loci to be confirmed in future crosses. These results suggest that ecology (exogenous selection) plays an important role in this hybrid zone. Thus, ecologically based divergent natural selection is responsible, simultaneously, for both ecotype divergence and hybridization. On the other hand, genetic introgression occurs only at neutral loci (nonoutliers). In the future, genome-wide studies and controlled crosses would give more valuable information about this process of speciation in the face of gene flow. [ABSTRACT FROM AUTHOR]

Anti-herbivory defenses support persistence of seaweeds. Little is known, however, about temporal dynamics in the induction of grazer-deterrent seaweed traits. In two induction experiments, consumption rates of the periwinkle Littorina obtusata (L.) on the brown seaweed Ascophyllum nodosum (L.) Le Jolis were measured in 3-d intervals. Changes in palatability of directly grazed A. nodosum were tested every 3 d with feeding assays using fresh and reconstituted seaweed pieces. Likewise, assays with fresh A. nodosum assessed changes in seaweed palatability in response to water-borne cues from nearby grazed conspecifics. Consumption rates of L. obtusata varied significantly during the 27-d induction phase of each experiment. Direct grazing by L. obtusata lowered palatability of fresh and reconstituted A. nodosum pieces to conspecific grazers after 15 d as well as after 6 and 12 d, respectively. After 12, 18, and 24 d, fresh A. nodosum located downstream of L. obtusata-grazed conspecifics was significantly less palatable than A. nodosum located downstream of ungrazed conspecifics. Changes in L. obtusata consumption rates and A. nodosum palatability during both induction experiments suggest temporal variation of grazer-deterrent responses, which may complicate experimental detection of inducible anti-herbivory defenses. [ABSTRACT FROM AUTHOR]