Antipredatory traits of the ammonoid shell — Indications from Jurassic ammonoids with sublethal injuries

More than 1500 Jurassic ammonoids with sublethal injuries from the H.Keupp collection (Berlin) were examined. They came from the Toarcian of Southern France, Southern Germany and England, from the Callovian of Villers-sur-Mer (Normandy, France) and from the Oxfordian of Sacaraha (Madagascar). Most sublethal injuries observed can not be referred to specific predators but display the intrinsic factors like tolerance against irritations and disturbances caused by predatory attacks. The types of shell breakages are strongly dependent on shell shape and specific sculpture. Some of the observed types occur exclusively in specific morphotypes. Average range of observed injuries is highest in weakly ornamented shells and lowest in highly ornamented shells. Maximum observed aboral range of shell breakages occurred in longidomes, the lowest in brevidomes and mesodomes. Shell form, sculpture and the length of the living chamber influence susceptibility to lethal injuries, confirmed by the frequency of multiple injuries, which is apparently a good proxy of the vulnerability towards shell crushing predators. The antipredatory traits were in an adaptational conflict with traits demanded for high manoeuvrability and streamlining.      

Shape differentiation and characterization in the two morphotypes of the Antarctic limpet Nacella concinna using Elliptic Fourier analysis of shells

The Antarctic limpet Nacella concinna (Strebel 1908) presents two ecotypes (littoral and sublittoral) differing in morphological and behavioral characteristics and more recently discovered, in physiological traits and genetic population structure. Previous works, based on traditional morphometry, used only three measurements (length, width and height of the shell) and their relationships. However, this methodology could not describe in detail the shape of the morphotypes. In the present work, Elliptic Fourier analysis (EFA) of shells was used to study the shape of the two morphotypes in six localities along the Antarctic Peninsula. The use of EFA confirmed the morphometric differences. The littoral morphotype has higher and globose shells with the apex displaced to the anterior part; the sublittoral individuals are more flat and pointed, and have the apex very well defined. Low allometric effect was detected in SL individuals, whereas L specimens did not show an allometric relationship within the examined size range. Differences in shell shape among localities were recorded. EFA analysis reflected the overall shell shape and allowed to characterize the main differences in shell shape between ecotypes that were difficult to quantify using the standard morphometric approach.     

Influence of shell morphometry,microstructure, and thermal conductivity on thermoregulation in two tropical intertidal snails

Tropical intertidal organisms tolerate large fluctuations in temperature and high desiccation rates when exposed during low tide. In order to withstand the short‐term heat stress, intertidal organisms adopt behavioral responses to maximize their survival. Our previous research showed that tropical littorinids found at the upper and lower intertidal shores in Singapore exhibited different behavioral adaptations during low tide. Most of the upper‐shore Echinolittorina malaccana kept a flat orientation, with the aperture against the substrate and the long axis of the shell towards the sun, whereas a majority of the lower‐shore individuals of Echinolittorina vidua stood with the edge of the aperture perpendicular to the substrate on the rocky shore during low tide. This prompted analyses of the shells of these two species to determine whether the differences in the shell morphometry, microstructure, and thermal conductivity of shells of E. malaccana and E. vidua were associated with their respective behavioral responses to thermal stress. Analyses of shell morphometry and thermal conductivity showed that shells of E. malaccana were more likely to minimize heat gain, despite having a higher thermal conductivity on the outer surface, due to their light‐gray, elongated shell. By contrast, the dark‐colored, globose shells of E. vidua probably gain heat more readily through solar radiation. Scanning electron microscopy images of the shells of both littorinid species further revealed that they have cross‐lamellar structure; however, only individuals of E. vidua showed the presence of disjointed rod layers and a pigmented inner shell surface. Individuals of E. malaccana had a rough outer shell surface with holes that inter‐connect to form water‐trapping channels that probably aid cooling. Individuals of E. vidua, however, had a smooth outer surface with rows of kidney‐shaped depressions as microsculptures which probably help to stabilize shell shape. In both Echinolittorina species, behavioral responses were used to overcome thermal stress during low tide that was associated with shell morphometry and shell thermal conductivity. Such combined adaptations increase survivability of the littorinids at their respective tidal levels.     

Leaf morphological responses to variation in water availability for plants in the <Emphasis Type="Italic">Piriqueta caroliniana</Emphasis> complex

Distribution of plants and the expression of traits associated with environmental variation can be affected by both average conditions and the variance in conditions including extreme climatic events. We expect that these same factors should affect the distribution of plants in hybrid zones between ecologically distinct species where the hybrids should occupy ecotones or intermediate habitats. We evaluated water availability and leaf morphological differences among parental and hybrid populations of herbaceous perennial plants in the Piriqueta caroliniana complex along environmental gradients in Southeastern North America. We focus on two taxa in this group; the viridis morphotype, which occurs in southern Florida, and the caroliniana morphotype, which is distributed from northern Florida to southern Georgia. Advanced-generation hybrid derivatives of these morphotypes occupy a broad geographic region that extends across much of central Florida. Overall, we found that hybrid populations occurred in significantly drier locations, indicating that their habitat requirements are transgressive (i.e., exceeding parental values) rather than intermediate to the parental morphotypes. Water availability differed between the two sampling years, and plants displayed morphological changes in response to these changes in moisture. During the drier year, leaves were narrower and more hirsute, corroborating experimental results that these leaf traits are plastic, and confirming that plasticity occurs in natural habitats. Hybrids exhibited intermediate leaf traits (shape and size) across both years, and displayed transgressive (hair density) leaf traits during the drier year. The apparent canalization of the hybrids’ leaf morphological traits may contribute to their tolerance of variable environmental conditions and may partially explain why they have displaced the caroliniana morphotype in central Florida.     

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.     

Adaptive responses of independent traits to the same environmental gradient in the intertidal snail Bembicium vittatum

The snail Bembicium vittatum occupies a wide range of intertidal habitats in the Houtman Abrolhos Islands, Western Australia. Allozyme variation reflects patterns of connectivity, which are independent of local habitat. In contrast, heritable differences in shell shape among 83 shore sites vary with habitat, indicating local adaptation. Here we examine dimorphisms of colour and spotting of the shell in the same populations, as a test of consistency and complexity of patterns of local adaptation. Within populations, the frequency of spotted shells is higher in dark shells. Despite this association, spatial variations of colour and spotting are only weakly correlated. As predicted for traits associated with local adaptation, subdivision is greater for colour, spotting and shape than for allozymes. Colour and shape are associated with local habitat, such that populations on vertical shores have higher frequencies of dark and relatively flatter shells than those on gently sloping shores. These associations are repeatable between three separate groups of islands. Spotting shows a weaker, but significant association with the same gradient. Although shape does not differ between colour morphs within populations, the proportion of dark shells is strongly associated with shape. Thus, the independent shell traits are apparently adapted to a common, biologically significant gradient, even though the adaptive mechanisms probably differ for colour and shape. The parallel variations of independent traits highlight both the complexity of local adaptation and the potential to reveal evolutionarily significant environmental contrasts by examining adaptively relevant traits.     

Crushing behavior of tropical and temperate crabs

The Guamunian xanthids Carpilius maculatus (L.), C. convexus (Forskal), and Eriphia sebana (Shaw & Nodder), and the parthenopid Daldorfia horrida (L.), possess large master claws with molariform teeth than are used to crush the shells of hermit crabs and snails. These crabs typically sever the spire of their prey, or make a gash in the body whorl. They tend to employ sustained pressure on the prey shell, and, except for Eriphia, rarely attack the outer lip, so that the outer lip of the shell typically remains undamaged, except in shells near the critical size, i.e., the maximum size of vulnerability to predation. Temperate species of Cancer (C. productus Randall and C. oregonensi Rathbun) may also crush shelled prey in the larger of their two claws, but more commonly they use both claws together in breaking open their victims. Sustained pressure is applied for only short periods by these crabs.Gastropod adaptations conferring resistance to crushing by crabs include a thick shell, narrow or otherwise small aperture, thickened outer lip, strong sculpture, and a low spire. Emphasis on these traits lowers the critical size of the prey, i.e., permits escape from cushing at a smaller size. An equatorward increase in the expression of the characteristics of crushing-resistance parallels an increase in crushing power of the crabs.     

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

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

Palaearctic gastropod gains a foothold in the dominion of endemics: range expansion and morphological change of Lymnaea (Radix) auricularia in Lake Baikal

In and around the endemic-dominated Lake Baikal, palaearctic species are generally restricted to shallow, sheltered bays and in- and out-flowing river floodplains. However, we observed populations of the palaearctic snail Lymnaea (Radix) auricularia on the steep, rocky littoral of Lake Baikal proper. We compared the morphology of 542 shells sampled from this new habitat with potential source populations from conventional habitats. A size-free Discriminant Analysis indicated a strong morphological differentiation of the newly established populations from their likely sources. The new populations had a more compact shell shape with a wide aperture, which may be advantageous in wave-exposed habitats where a firm attachment to the substrate is needed. Shells from the conventional habitats were more elongated, with a narrow aperture, which may be advantageous in habitats that have a dry period where retreating into the mud is required and water loss should be limited. These results may suggest that selection is acting on shell shape in Lake Baikal. The apparent recent arrival of this pandemic gastropod in a habitat previously dominated by endemics constitutes a potential ecological threat and an alert to possible ecological change.     

Morphological Variation in Wild Marmosets (Callithrix penicillata and C. geoffroyi) and Their Hybrids

Evolutionary theory and observation predict wider phenotypic variation in hybrids than parental species. Emergent phenotypic novelty in hybrids may in turn drive new adaptations or speciation by breaking parental phenotypic constraints. Primate hybridization is often documented through genetic evidence, but knowledge about the primate hybrid phenotype remains limited due to a small number of available studies on hybrid primate morphology. Here, we examine pelage and morphometric variation in two Brazilian marmoset species (Callithrix penicillata and C. geoffroyi) and their hybrids. Hybrids were sampled in an anthropogenic hybrid zone in the municipality of Viçosa, Minas Gerais state, Brazil. We analyzed hybrid facial and body pelage color variation, and compared 13 morphometric measures between hybrids and parental species. Five different hybrid facial morphotypes were observed, varying from intermediate to parental-like. Hybrid facial morphotypes were biased towards C. penicillata, suggesting that the pelage of this species may be dominant to that of C. geoffroyi in this context, and indicating that mate preference, and therefore gene flow/introgression, may be biased towards C. penicillata within the hybrid zone. Hybrid morphometric features were on average intermediate to parental species traits, but transgressive hybrids were also observed, suggesting that morphometric variation for the studied traits is consistent with Rieseberg’s complementary allele model. Finally, we observed a decoupling of facial patterning and size/shape in hybrids, relative to parent phenotypes, suggesting that an important factor driving phenotypic novelty within the Viçosa marmoset hybrid zone might be the loosening of evolutionary constraints on phenotypic trait integration.     

Quantifying phenotypic gradients in freshwater snails: a case study in Lithasia (Gastropoda: Pleuroceridae)

Many authors have described a pattern of morphological variation in freshwater bivalves where shells taken from lentic and lotic environments, or headwaters and main stem reaches, appear to exhibit phenotypic gradients in size and shape. For example, mussels taken from headwater reaches tend to possess smooth, less inflated shells compared to the more obese, sculptured individuals downstream. Others observed similar relationships in certain freshwater gastropods, but this variation has not been quantified nor its existence explained in an ecological or evolutionary context. Geometric morphometrics indicated freshwater snails shells from the pleurocerid genus Lithasia from the Duck River, Tennessee, USA, show phenotypic gradients similar to those in freshwater mussels. Shells from upstream areas were narrow and less sculptured on the posterior portions of their body whorls, while downstream shells were more inflated and possessed significantly more sculpture. This phenotypic variation may reduce predation or damage due to dislodging. The nature of the observed plasticity suggests an unidirectional environment similar to that proposed by the river continuum concept. Handling editor: K. Martens     

Shell morphometry in barnacles: quantification of shape and shape change in Balanus

Geometric design in the barnacle genus Balanus has been studied in relation to variation in adult shell form, that includes differences among species, and size-related changes in shape. The genus comprises 40 Recent species, and as a group these display a more or less constant morphology over an extraordinary size range (10 to 200 mm in basal length).
Linear and volumetric measurements were collected from 232 adult individuals of 14 species representing the variation in size, shell form and shell design thought to occur in the genus. Specimens were chosen to represent the size ranges of the species. Only isolated individuals growing on planar surfaces were used; shells were complete, undamaged and undistorted.
Shell form differs among taxa, and no two species scale alike; intraspecific variation for five ratio variables shows strong allometry over the adult size range of each species. As size increases, there is a trend for the basis and orifice to maintain their shapes or to become slightly more elliptical, and for shells to become more conical and proportionately taller.
Throughout their size ranges, species can be described by these geometries: paraboloid (6 species), frustum of an ellipsoidal cone (5 species), frustum of a cone (2 species) and a cone (1 species). Shell geometry is not a function of species size, but there does appear to be a correlation between shell geometry and shell volume. Species with relatively small shell volumes are described by a frustum of an ellipsoidal cone, or by a cone, while those with a relatively large shell volume are described by a paraboloid, or by the frustum of a cone.     

Shell morphology and color of the subtidal whelk Buccinum undatum exhibit fine‐scaled spatial patterns

Geographical patterns in morphology can be the result of divergence among populations due to neutral or selective changes and/or phenotypic plasticity in response to different environments. Marine gastropods are ideal subjects on which to explore these patterns, by virtue of the remarkable intraspecific variation in life‐history traits and morphology often observed across relatively small spatial scales. The ubiquitous N‐Atlantic common whelk (Buccinum undatum) is well known for spatial variation in life‐history traits and morphology. Previous studies on genetic population structure have revealed that it exhibits significant differentiation across geographic distances. Within Breiðafjörður Bay, a large and shallow bay in W‐Iceland, genetic differentiation was demonstrated between whelks from sites separated by just 20 km. Here, we extended our previous studies on the common whelk in Breiðafjörður Bay by quantifying phenotypic variation in shell morphology and color throughout the Bay. We sought to test whether trait differentiation is dependent on geographic distance and/or environmental variability. Whelk in Breiðafjörður Bay displayed fine‐scale patterns of spatial variation in shape, thickness, and color diversity. Differentiation increased with increasing distance between populations, indicating that population connectivity is limited. Both shape and color varied along a gradient from the inner part of the bay in the east to the outer part in the west. Whelk shells in the innermost part of Breiðafjörður Bay were thick with an elongate shell, round aperture, and low color diversity, whereas in the outer part of the bay the shells were thinner, rounder, with a more elongate aperture and richer color diversity. Significant site‐specific difference in shell traits of the common whelk in correlation with environmental variables indicates the presence of local ecotypes and limited demographic connectivity.     

Coarse- versus fine-grained water stress in Arabidopsis thaliana (Brassicaceae)

The effects of coarse vs. fine-grained variation in water regime on 13 accessions of Arabidopsis thaliana were compared and it was found that the scale of environmental variation can lead to differences in which traits are affected and in the magnitude and direction of effect. Coarse-grained (between individual) variation was studied by comparing plants that received high or low amounts of water on a daily basis. Fine-grained (within individual) variation was distributed across the coarse-grained treatments; plants in the fine-grained treatment received the same amount of water per week as the corresponding high and low coarse-grained treatments; however, in the fine-grained treatments, plants received their entire weekly water allotment over 2 days. The number of leaves produced, days to bolting, length of bolt, number of basal and lateral branches, and survival to 30 days after bolting were measured and analyzed by ANOVA. Plants grown in high water generally produced more leaves, taller main axes, more branches and had increased survival, relative to those in low water. Bolting date showed a significant genotype specific effect of water level. Plants grown under fine-grained variation in watering had taller main axes relative to constant water. There were significant interactions between both types of variation for bolting date, main axis height, and survival. Principal components for all traits were loaded such that the components themselves were also independently affected by genotype, water level, variability of water level (grain), and level-by-grain interactions in ways that were largely predicted by their component traits. Overall, the effects of fine-grained variation were subtler than those of coarse-grained variation: fewer traits responded, and the magnitude of the responses was smaller. Nevertheless, the responses to fine-grained variation are distinct from those of coarse-gained variation, differing with respect to which traits respond and in the direction of response, and must therefore be treated as independent sources of environmental variation.     

Morphotypes of the lacustrine ostracod <Emphasis Type="Italic">Limnocythere rionegroensis</Emphasis> Cusminsky &amp; Whatley from Patagonia,Argentina, shaped by aquatic environments

This study describes and quantifies morphological valve traits of the ostracod Limnocythere rionegroensis from Patagonian steppe lakes and explores their association with lake water characteristics. Surface ornamentation was examined by scanning electron and transmitted light microscopy, and valve size and shape were analyzed using morphometric techniques. Limnocythere rionegroensis shows remarkable variations in surface ornamentation, based on which three morphotypes (MI, MII, and MIII) were identified. Valves of morphotypes MI and MIII are larger, show slight to moderate external reticulation, and a higher shape variability, whereas MII is characterized by a very conspicuous reticulation, lower shape variability, and smaller valves. Outline analysis yielded a great shape disparity related to the dorsal margin slope. MI was found in sexual populations from euhaline to mesohaline ephemeral lake; MII occurs in parthenogenetic populations from mesohaline to oligohaline permanent or ephemeral lakes; and MIII, from both sexual and asexual populations, inhabits a broad range of environmental conditions in terms of salinity and stability. Limnocythere rionegroensis intraspecific variations may be caused by environmental parameters and genetic factors associated to reproductive strategies. These results contribute to the knowledge of extant L. rionegroensis morphological variability and provide additional clues to improve the environmental interpretation of fossil assemblages.     

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)     

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.     

Ecomorphological variation in shell shape of the freshwater turtle Pseudemys concinna inhabiting different aquatic flow regimes

Populations of species that inhabit a range of environments frequently display divergent morphologies that correlate with differences in ecological parameters. The velocity of water flow (i.e., flow velocity) is a critical feature of aquatic environments that has been shown to influence morphology in a broad range of taxa. The focus of this study was to evaluate the relationship between flow velocity and shell morphology for males and females of the semi-aquatic freshwater turtle Pseudemys concinna. For both sexes, the carapace and plastron show significant morphological differences between habitats characterized by slow-flowing (i.e., lentic) and fast-flowing (i.e., lotic) water. In general, the most prominent pattern for both sexes is that the shells of individuals from lotic habitats are more streamlined (small height-to-length ratio) than the shells of individuals from lentic habitats. Of the two shell components (carapace and plastron), the carapace shows greater divergence between habitats, particularly for males. These results are consistent with adaptations to flow velocity, and suggest that variation in shape may be more constrained in females. I also provide empirical evidence for an adaptive benefit of the observed shape change (i.e., drag reduction) and a brief comment on the relative roles of genetic divergence and phenotypic plasticity in generating shape differences observed in this species.