Ecomorphology ofCrassostrea cucullata (Born, 1778) (Ostreidae) in a mangrove creek (Gazi,Kenya)

Variations of shell form and shell length were studied for oysters growing in the mangroves of Gazi Creek, Kenya, and related to different environmental factors.For the study of the form, Fourier analysis was performed on the circumference of 85 oysters. The resulting coefficients were compared among specimens using cluster analysis. The correspondence between this classification and substrate diameter is virtually perfect (only one misclassification out of 85 oysters). For the clusters based on height above chart datum, 9 specimens, all on intermediate height levels, were misclassified. Orientation with respect to tidal current had 13 misclassifications. Mangrove species seemed to influence form only marginally, if at all.In the study of the size ofCrassostrea cucullata, the length of 956 oysters, growing along two transects were measured, and correlated with several environmental factors. Oyster length was not related to substrate diameter or its orientation with respect to the main current. Length was not influenced by density up to a cover of 70%. For densities higher than 70%, there was a fairly strong negative correlation (r ² = 0.634,n = 217). Length was not correlated with height above bottom (base of the tree) for heights lower than 20 cm, while oysters growing closer to the bottom were smaller. The correlation with height above chart datum was negative but very low (r ² = 0.060,n = 957). However, if all measurements of oysters closer than 20 cm to the bottom, and all from a density of more than 70% cover are deleted from the data set, the correlation with height increased dramatically, the slope still being negative (r ² = 0.859,n = 543).     

Microgeographical variation in shell strength in the flat periwinkles Littorina obtusata and Littorina mariae

The strength of molluscan shells has been shown to vary in adaptive ways in a number of species and one of the main factors thought to be involved is shell-crushing by predators. A recent study found that the sibling species of flat periwinkle Littorina obtusata and Littorina mariae showed significant differences in the rates at which shell strength increased with shell length in specimens which had been collected from the same location, where the species were sympatric. This paper describes differences between the shells of the two species from a number of localities around Milford Haven in Dyfed, Wales, and local geographical variation in the shells. Littorina mariae, which is normally found at lower tidal levels than L. obtusata, matures at a smaller shell length. Both species reinforce the shell as they grow since shell strength, determined as the maximum force applied by a hydraulic tensile testing machine before the shell cracked, is strongly positively allometric; it increases at a rate close to the cube of shell length whilst isometric growth would result in strength increasing in proportion to the square of shell length. Because L. mariae matures earlier and reinforces the shell at a smaller size, the mature shell of L. mariae is substantially stronger on average than that of a similar sized but immature L. obtusata. At maturity the shell strengths of the two species are not very different despite the substantial difference in mean shell length. Strength varies significantly from shore to shore, and with the level of the shore from which the animals were collected. Strength increases down the shore in both species. Shell strength decreases with exposure to wave action in L. mariae but increases with exposure in L. obtusata; there is also substantial shore-to-shore variation which is not explained by exposure. Path analysis was used to explore the relationship between shell strength and other measured shell parameters (mass, length, height, thickness). The best predictor of shell strength in both species is a parameter which is heavily positively loaded on LN (shell mass) and strongly offset by negative loadings on LN (shell length) and LN (shell height). This is logical because for a given shell length a heavier shell will be thicker and stronger, whilst for a given shell mass a bigger shell will be thinner and therefore weaker. Such differential variation of shell mass and shell length explains most of the geographical variation observed in shell strength; shells are stronger in snails collected from one place than from another because, for the same shell length they are heavier or, to put it the other way, because at the same shell mass, they are smaller.     

Within- and among-lake variation in shell morphology of the freshwater clam Elliptio complanata (Bivalvia: Unionidae) from south-central Ontario lakes

To compare the relative magnitude of variation in shell morphology within and among lakes, Elliptio complanata were collected from low and high exposure areas in each of four small lakes in south-central Ontario. Nested ANOVA's on shell length, height, width and weight revealed that shell morphology varied much more between sites of differing exposure within a lake than among lakes of differing alkalinity. Canonical variates analysis showed that clams from high exposure areas had larger and proportionately taller and heavier shells than those from low exposure areas. There was no relationship between alkalinity of lakes and shell morphology. These results suggest that the use of unionid shell morphology to predict long-term whole lake water chemistry (e.g. alkalinity) requires sampling designs which take into account within-lake variation in shell morphology.     

Disentangling the effects of intraspecies variability,phylogeny, space,and climate on the evolution of shell morphology in endemic Greek land snails of the genus Codringtonia

Extensive variation in land snail shell morphology has been widely documented, although few studies have attempted to investigate the ecological and evolutionary drivers of this variation. Within a comparative phylogenetic framework, we investigated the temporal and spatial evolution of the shell morphology of the Greek endemic land snail genus Codringtonia. The contribution of both inter‐ and intraspecies shell differentiation in the overall shell variability is assessed. The effect of climate, space, and evolutionary history on the shell variability was inferred using a variance partitioning framework. For Codringtonia species, intraspecies divergence of shell traits contributes substantially to the overall shell variability. By decomposing this variability, it was shown that the overall shell size of Codringtonia clades is phylogenetically constrained, related to early speciation events, and strongly affected by large‐scale spatial variability (latitudinal gradient). The effect of climate on shell size cannot be disentangled from phylogeny and space. Shell and, to a larger extent, aperture shape are not phylogenetically constrained, and appear to be mostly related to conspecific populations divergence events. Shell shape is substantially explained by both climate and space that greatly overlap. Aperture shape is mainly interpreted by medium to small‐scale spatial variables. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 796–813.     

Tuning in to multiple predators: conflicting demands for shell morphology in a freshwater snail

1. We examined the response to chemical cues from fish and crayfish, two predators with contrasting feeding modes, and their single and combined effect on shell morphology in the freshwater snail Radix balthica. 2. Snails were subjected to four treatments: tench (Tinca tinca), signal crayfish (Pacifastacus leniusculus), a combination of tench and signal crayfish and no predators (control). Shell shape, crushing resistance and shell thickness were quantified. We also analysed whether shape or shell thickness contributes most to crushing resistance. 3. Chemical cues from the fish induced a rounder shell shape in R. balthica, a thicker shell and a higher crushing resistance, whereas crayfish chemical cues had no effect on shell morphology, shell thickness or crushing resistance. Shell shape contributed more to crushing resistance than shell thickness. 4. The combined predator treatment showed an intermediate response between the fish and crayfish treatments. Shell roundness was reduced compared with the fish treatment, but the reduced crushing resistance that comes with a less rounded shell was compensated by an increased investment in extra shell material, exceeding that of the fish treatment. 5. Our study extends previous studies of multipredator effects on phenotypically plastic freshwater snails by showing that the snails are able to fine‐tune different elements of morphology to counter predator‐specific foraging modes.     

SPATIAL PATTERNS OF SHELL SHAPE OF THREE SPECIES OF CO-EXISTING LITTORINID SNAILS IN NEW SOUTH WALES, AUSTRALIA

The shape and relative weight of the shell have been shown tovary intraspecifically and interspecifically in a number ofspecies of gastropods, including many different littorinids.These differences give rise to different shell forms in differenthabitats. In those species which have non-planktotrophic development,differences in shell form among shores have been usually explainedin terms of natural selection because exposure to waves supposedlyfavours light shells with large apertures, while predation bycrabs on sheltered shores favours elongated, thick shells withsmaller apertures. Differences in shell shape among speciesfound at different heights on the shore have been explainedin terms of resistance to desiccation and temperature. Suchvariables would tend to act on a relatively broad-scale, i.e.causing differences among heights on a shore or among shores.Rates of growth, which might vary at much smaller scales withina shore, have also been shown to affect the shapes of many shells. In this study, the shape and relative weight of shells of threespecies of co-existing littorinids (Littorina unifasciata, Bembiciumnanum and Nodilittorina pyramidalis) were measured. These speciesall haveplanktotrophic development and they are found on manyshores where there is no evidence that they are preyed uponby crabs. Before explanations of shell shape are proposed, itis necessary that patterns of variation, within different partsof ashore and among different shores are clearly documented.These patterns were measured at a number of different spatialscales within and among replicate shores with different amountsof wave exposure. Large and small specimens were included toallow intraspecific comparisons among snails of different sizesfound at different heights on the shore. The results showedsignificant differences among shores in shape and relative weightof shells, but these differences could not be explained by exposureto waves. In addition, snails of different sizes and differentspecies did not show the same patterns although they were collectedfrom the same sites. Importantly, the shell shape of Liuorinaunifasciata varied significantly among sites at approximatelythe same height within a shore. These differences could notbe clearly correlated with density, mean size nor exposure towaves. The only consistent pattern was a decrease in relativeaperture size in specimens living higher on the shore. Modelsthat have commonly been proposed to explain shape and relativeweight of shells in other species of gastropods are not adequateto explain the small- and large-scale variation of the measurementsdescribed here. It is proposed that any selective advantageof shell morphology and the effects of any variables on thedevelopment of shell morphology in these species can only beidentified after appropriately designed and replicated fieldexperiments. (Received 4 March 1994; accepted 13 September 1994)     

l-3,4-Dihydroxyphenylalanine (l-DOPA) secreted by oyster (Crassostrea gigas) mantle cells: functional aspects

l-DOPA is present in many molluscan periostraca and is thought to play an important role in the shell protein sclerotization mechanism. We analyzed l-DOPA content in organic membranes produced by the oyster (Crassostrea gigas) mantle as a reaction against shell damage, such as induced by Polydora sp. infestation, an artificial perforation of the shell, and in normal oysters mantle. When oysters are secreting the protective organic membrane against a shell perforation, the area of mantle close to the repairing region had a greater l-DOPA content than the remaining mantle. Mantle border as a whole had the same l-DOPA content as the central mantle. Nevertheless there is a variation pattern in mantle border l-DOPA content, the area near the umbo having a higher content and decreasing towards the frontal region. l-DOPA content in younger oysters was greater than that in older oysters. The results of perforation experiment suggest a good correlation between organic membrane synthesis and l-DOPA mantle border content. Concerning normal oysters, a higher l-DOPA content in mantle border than in central mantle would be expected but we couldn’t detect any differences. From all the results there is some evidence that mantle l-DOPA is related to other functions in addition to participating in the sclerotization mechanism.     

Evaluation of the sample size used for the rapid bioassessment of rivers using macroinvertebrates

Experiments were designed to investigate selective predation by medium (40–55 mm carapace width: CW) and large (55–70 mm CW) Carcinus maenas when feeding on four bivalves of contrasting shell morphology. Size-selection was examined by presenting individual crabs with a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas and Cerastoderma edule. Medium-sized crabs preferred mussels 5–15 mm shell length (maximum shell dimension: SL) and cockles 5–10 mm SL, whereas large crabs preferred mussels 15–25 mm and cockles 10–20 mm SL. Crabs generally showed no preference for any particular size of either oyster species. Species-selection was examined by presenting individual crabs with paired combinations of the four bivalves in various proportions. When offered mussels and oysters simultaneously, both size categories of crabs consistently selected mussels, and food choice was independent of prey relative abundance. By contrast, C. maenas selected mussels and cockles as expected by the frequency in which each size category of crab encountered the preferred size ranges of prey. Crab preference clearly paralleled the rank order of prey profitability, which in turn was mainly determined by prey biomass, suggesting that active selection takes place at some point of the predation cycle. Experiments with epoxy resin models showed that initial reluctance of crabs to attack oysters was not associated with the ultimate energy reward. Moreover, they suggest that foraging decisions are partly based on evaluations of overall prey shape and volume, and that the minimum dimension of the shell constitutes an important feature which crabs recognise and associate with prey value.     

INTRA-POPULATION VARIATION OF THE SHELL OF LITTORINA RUDIS (MATON) (MOLLUSCA: PROSOBRANCHIA)

Shell variation within a single population of Litlorina rudis(Maton), collected near the Biological Station at Trondheim,Norway, was studied. The variation of the shell features studied, which includedthe shape of the basal part of the outer lip, and the relativeheight of the spire, width of the shell and of the aperture,were found to be at least partially related to shell height.Furthermore, the shell shape in the population investigated,besides varying with age, also varies due to shell damage. Nosignificant differences were found between the shapes of theshells of males and females. The shell characteristics of different populations of L. rudisalso vary greatly, one of the causes being differences in theenvironmental conditions. When studying this variation betweendifferent populations, however, it must be emphasized that onlyundamaged shells of individuals of about the same age shouldbe compared, in order to obviate the effects of the variabilitydue to age differences and shell damage within each individualpopulation. ^*Contribution from Trondhjem Biological Station no. 199 (Received 18 February 1980;     

Environmental Heterogeneity and Morphological Variability in Pisidium subtruncatum (Sphaeriidae,Bivalvia)

On the example of P. subtruncatum, a widely distributed bivalve, we studied the relationship between environmental and morphological variability. The main questions addressed were: (1) are there differences in shell morphology between different populations, (2) are they dependent on sediment composition, hydrology or water chemistry, (3) are there differences in morphological variability between stable and unstable environments. Measurements of specimen from nine sampling sites showed high variability in shell shape and size. Length‐height relation, symmetry, width and dry weight of the shell all varied significantly. Shell shape was significantly correlated with sediment composition and dependent on hydrological connectivity, with higher, asymmetric shells occurring in sandy sediments and lower, rounded shells in muddy sediments. Analysis of intrapopulation morphometric variation showed that variability is higher in habitats with high temporal and spatial environmental variability and lower in habitats with low or reduced environmental heterogeneity. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Spatial variation of shell morphometrics in the subantarctic land snail <Emphasis Type="Italic">Notodiscus hookeri</Emphasis> from Crozet and Kerguelen Islands

The land snail Notodiscus hookeri, widely distributed in subantarctic islands, shows a large intraspecific variation in shell morphology. In the present work, shell size and form of individuals from populations located in Crozet and Kerguelen archipelagos were investigated by means of multivariate statistics. Variation in shell morphometrics was analysed after the partitioning of the overall variation into size and shape components by means of a principal component-based approach. Shell size shows a significant spatial heterogeneity, which seems essentially related to environmental pressures. Previous works pointed to a greater conchological variation between populations from Kerguelen but present observations show that intra-island variances are not significantly different in the two islands studied. Variation in shell shape splits the populations into two main entities because of different allometric relationships between two shell height components and all other measurements. However, using geographical affinities of populations as instrumental variables shows that more complex environmental features interfered in population clustering.     

Character displacement in Hydrobia

Summary Fenchel's study of size variation in Hydrobiid snails in the Limfjord, Denmark, provides one of the most convincing cases of ecological character displacement available. In order to assess the generality of the phenomena within the Hydrobia genus, allopatric and sympatric Hydrobia ventrosa, H. neglecta and H. ulvae were collected from 24 coastal sites around Eastern England in July and October, 1982. Shell heights of 5,850 snails from 55 samples were recorded. These data were analysed for intraspecific allopatric-sympatric, and interspecific height differences. Such differences were not significant for H. ventrosa/H. neglecta, but the species exhibited parallel variation at sympatric sites. H. ventrosa/H. ulvae showed only significant interspecific differences, but in both allopatry and sympatry. The pattern of mean size variation for these species resembles that in the Limfjord. However the statistical analyses fail to provide support for character displacement. Differences in character states attributed to the process of character displacement may result from a number of other causes. Environmental conditions at sympatric and allopatric sites may act differentially on the heights of H. ulvae and H. ventrosa. The conditions that prevail at sympatric H. ulvae sites appear to lead to increased size in this species irrespective of the presence of H. ventrosa.     

Comparative ecophysiology ofLarix kaempferi (Lamb.) Carr. andAbies veitchii Lindl. II. Mechanisms of higher drought resistance of seedlings ofL. kaempferi as compared withA. veitchii

The underlying mechanisms of drought resistance in seedlings ofLarix kaempferi as compared with those inAbies veitchii were investigated. First, seedlings of the two species were desiccated in air of differing water potential, and their survival rates were compared to detect any difference in desiccation tolerance between them. Contrary to differences in their drought resistance,A. veitchii showed higher survival. Next, the shoot heights and root depths of the two species were compared to detect any difference in desiccation avoidance. Although the shoot height differed little between the two species, the roots ofL. kaempferi seedlings were much deeper than those ofA. veitchii seedlings, and the death rate under progressing drought was well correlated with the root extensibility of each species. It was thus concluded thatL. kaempferi seedlings are able to avoid desiccation by extending their roots in deep soil faster thanA. veitchii seedlings, thereby resisting to drought. This paper is dedicated to Prof. Toshiro Saeki to mark his fruitful career in plant ecology.     

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.     

Morphology of the Bivalve Mollusks Crenomytilus grayanus and Mytilus coruscus in Relation to Their Spatial Distribution in the Upper Subtidal Zone

The morphology of the shell and byssus threads was studied in two closely related mussel species Crenomytilus grayanus and Mytilus coruscus. The two species differ significantly from each other in the shell shape and in the degrees of development and deformation of byssus threads. These differences, in turn, determine (either directly or indirectly) the differences in strength of the byssal attachment and are discussed in terms of their functional morphology with respect to the spatial distribution of the mussels in marine coastal zones.     

Scale-dependence in geographic variation in a freshwater gastropod across the Palearctic

ABSTRACT

The reality of spatial clinal variation in morphological traits of freshwater pulmonate snails (Gastropoda: Pulmonata) has repeatedly been questioned or totally discounted. There is a lack of sound statistical evidence in the articles hitherto published on this subject supporting these claims. Here, by means of different analytical methods (analysis of spatial autocorrelation, linear regression analysis, canonical correlation analysis and others), we demonstrate that shell variation in the dwarf pond snail, Galba truncatula, is patterned in space throughout the northern and central Palearctic, with latitudinally-oriented clines in body size and in some shell proportions. Shell size in G. truncatula decreases with latitude and temperature, representing a special case of converse Bergmann cline. However, the temperature itself is hardly the main driver of shell size variation. It is argued that the shorter growing seasons at high latitudes may represent a better explanation for the observed trend. Shell proportions in the dwarf pond snails vary weakly at the macrogeographic scale, being spatially patterned at lower (mesogeographic) scales around 1200–1500?km. In general, spatial variation in G. truncatula shell size is decoupled from variation in shell shape, demonstrating clear scale-dependence similar to that found in different species of terrestrial (non-aquatic) pulmonate snails.     

黄河口西南侧小岛河河口天然牡蛎礁的牡蛎种群结构

牡蛎礁是生态系统服务价值高、但退化最严重及受关注度最高的海洋生境之一,牡蛎礁修复亦成为国际海洋生态修复的热点。掌握牡蛎自然种群状况及动态变化是评估牡蛎礁修复效果的基础。目前,我国天然牡蛎礁的牡蛎种群状况相关的背景资料较为缺乏。在黄河口西南侧的小岛河河口新发现天然活体牡蛎礁,但该牡蛎礁曾被大规模的商业采捕,亟需推进针对性的保护和修复研究工作。基于2021年11月对该牡蛎礁开展的牡蛎种群生态调查,分析其种类组成、年龄结构及生长特征。结果显示:该牡蛎礁分布有近江牡蛎(Crassostrea ariakensis)和长牡蛎(Crassostrea gigas)。牡蛎礁上以活体牡蛎为主,死亡牡蛎壳体数仅占6.1%-6.7%。活体牡蛎的密度和生物量分别为(2811±778)个/m²和(21.97±30.43)kg/m²,近江牡蛎较多,其密度和生物量分别占比55.7%和76.4%。近江牡蛎和长牡蛎的年龄分别介于0+-4+ 龄和0+-2+ 龄,它们都以壳高介于30-40 mm及壳质量<5 g的0+ 龄个体数量居多(>80%)。近江牡蛎的壳体形态参数均值都高于同龄组长牡蛎的相应值。两种牡蛎壳体均呈负异速增长,不同龄级的壳体延展方向不同。拟合von Bertalanffy生长方程得到,近江牡蛎和长牡蛎的渐近壳高分别为286 mm和173 mm,估算的拐点年龄分别为5.47 龄和2.56 龄,两种牡蛎的生长曲线分布存在极显著差异(P<0.001)。以上结果表明,小岛河河口的天然牡蛎礁的牡蛎自然种群资源较丰富,具有高密度、低龄和低死亡率等特点,有较好的活力和扩张潜力,有利于被采捕后的礁体的恢复。两种牡蛎中,近江牡蛎因其具有较高的生物量和较长的生长年龄,对礁体形成和扩繁可能更为重要。建议对该天然牡蛎礁及牡蛎种群开展周期>3年的原位保护、修复和连续监测计划。     

Adaptive responses and invasion: the role of plasticity and evolution in snail shell morphology

Invasive species often exhibit either evolved or plastic adaptations in response to spatially varying environmental conditions. We investigated whether evolved or plastic adaptation was driving variation in shell morphology among invasive populations of the New Zealand mud snail (Potamopyrgus antipodarum) in the western United States. We found that invasive populations exhibit considerable shell shape variation and inhabit a variety of flow velocity habitats. We investigated the importance of evolution and plasticity by examining variation in shell morphological traits 1) between the parental and F₁ generations for each population and 2) among populations of the first lab generation (F₁) in a common garden, full‐sib design using Canonical Variate Analyses (CVA). We compared the F₁ generation to the parental lineages and found significant differences in overall shell shape indicating a plastic response. However, when examining differences among the F₁ populations, we found that they maintained among‐population shell shape differences, indicating a genetic response. The F₁ generation exhibited a smaller shell morph more suited to the low‐flow common garden environment within a single generation. Our results suggest that phenotypic plasticity in conjunction with evolution may be driving variation in shell morphology of this widespread invasive snail.     

Spatial distribution patterns of Littoraria species in Hong Kong mangroves

Variation in the abundance and distribution of two species of mangrove littorinid was investigated using a nested sampling design at different spatial scales, in the two dominant seasons, in Hong Kong. The abundance of Littoraria melanostoma, which was less abundant than L. ardouiniana, showed large-scale spatial variation whilst abundance of L. ardouiniana varied on both large and smaller spatial scales and interacted with seasons, indicating that the abundance of this species varied between mangroves in summer and winter. Small-scale variation suggested a patchy distribution of littorinids within a mangrove, whilst the large-scale variation might reflect changes in physical factors possibly associated with habitat fragmentation. Investigation of the vertical distribution of the littorinids on the mangrove trees revealed that L. melanostoma were located at a lower overall level than L. ardouiniana. This zonation pattern may be a result of the morphological differences of the two species, as L. melanostoma has a thick shell, whilst L. ardouiniana has a thinner, but colour polymorphic shell. Both species of mangrove littorinid showed patchy distributions at a variety of scales and these patterns highlight the importance of using a hierarchical sampling approach when investigating spatially fragmented habitats.