Ontogeny and evolution within the Myophorellidae (Bivalvia): Paedomorphic trends

The role of heterochronic phenomena in molluscan evolution is insufficiently understood but potentially significant. The aim of this paper is to explore some paedomorphic trends in the evolution of the Myophorellidae (Bivalvia: Trigoniida). Early ontogeny of general shell shape and ornamentation of one species of Steinmanella was analyzed and compared to data obtained for three species of Myophorella: two belonging to the subgenus M. (Promyophorella) (one from the Jurassic and one from the Cretaceous) and one belonging to the Jurassic M. (Myophorella). For general shell shape, a geometric morphometric analysis was performed on lateral views of the shells. Regarding ornamentation, flank costal disposition on the marginal carina, tubercle separation and relative development of the sub-commarginal subset of flank costae were quantified. A qualitative analysis was also performed. A two-trend shell shape development is considered as primitive. The first trend is marked by a relative reduction of the posterior margin together with a relative elongation of the shell. A tangential opisthogyrate growth component characterizes the second trend. There is a transitional stage where both trends interact. Early flank ornamentation is characterized by two or three sub-commarginal costae, continuous through the area, after which oblique costae with fine tubercles start to form. The subgenus M. (Myophorella) evolved by paedomorphic retention of juvenile shell shape and ornamentation, resulting in a large shell with coarse tubercles. Shell morphology in Steinmanella evolved by paedomorphic suppression of the primitive second trend in the development of the shell, resulting in an orthogyrate shell shape, and the retention of juvenile ornamentation (coarse tubercles, more sub-commarginal costae, juvenile rates of costal disposition). The paedomorphic (most likely by deceleration) retention of juvenile shell morphology within the Myophorellidae seems to have been recurrent within the group, resulting in many cases of convergence, and obscuring the phylogenetic relationships among its species.     

Effects of shell shape,size, and sculpture in burrowing and anchoring abilities in the freshwater mussel Potamilus alatus (Unionidae)

Because of the sedentary lifestyle of freshwater mussels, studies examining their movement capabilities are scarce. However, the ability to burrow into the substrate and the ability to remain stationary are likely crucial components of their behavioural repertoire. The performance of these different tasks is likely to be affected by the presence of the shell ornamentation characteristic of many mussel species. Previous studies have suggested that shell ornamentation results in a trade‐off between burrowing ability and remaining stationary when an extrinsic force attempts to dislodge it from the substrate once buried. We examined the effect of morphology and shell ornamentation on burrowing performance and anchoring ability by artificially creating shell ornamentation on a relatively smooth‐shelled species (Potamilus alatus). Burrowing behaviours and performance and the force required to dislodge mussels (anchoring ability) were quantified with and without ornamentation. Interestingly, we found that the artificial shell ornamentation had no significant effect on burrowing behaviours and performance or dislodgement force. Burrowing and dislodgement, however, were both highly influenced by shell size and shape. All of the available information suggests that shell size, shape, and sculpture influence burrowing and anchoring in complex ways that needs further examination. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 136–144.     

Finite element modeling of shell shape in the freshwater turtle Pseudemys concinna reveals a trade‐off between mechanical strength and hydrodynamic efficiency

Aquatic species can experience different selective pressures on morphology in different flow regimes. Species inhabiting lotic regimes often adapt to these conditions by evolving low‐drag (i.e., streamlined) morphologies that reduce the likelihood of dislodgment or displacement. However, hydrodynamic factors are not the only selective pressures influencing organismal morphology and shapes well suited to flow conditions may compromise performance in other roles. We investigated the possibility of morphological trade‐offs in the turtle Pseudemys concinna. Individuals living in lotic environments have flatter, more streamlined shells than those living in lentic environments; however, this flatter shape may also make the shells less capable of resisting predator‐induced loads. We tested the idea that “lotic” shell shapes are weaker than “lentic” shell shapes, concomitantly examining effects of sex. Geometric morphometric data were used to transform an existing finite element shell model into a series of models corresponding to the shapes of individual turtles. Models were assigned identical material properties and loaded under identical conditions, and the stresses produced by a series of eight loads were extracted to describe the strength of the shells. “Lotic” shell shapes produced significantly higher stresses than “lentic” shell shapes, indicating that the former is weaker than the latter. Females had significantly stronger shell shapes than males, although these differences were less consistent than differences between flow regimes. We conclude that, despite the potential for many‐to‐one mapping of shell shape onto strength, P. concinna experiences a trade‐off in shell shape between hydrodynamic and mechanical performance. This trade‐off may be evident in many other turtle species or any other aquatic species that also depend on a shell for defense. However, evolution of body size may provide an avenue of escape from this trade‐off in some cases, as changes in size can drastically affect mechanical performance while having little effect on hydrodynamic performance. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Buckman's Paradox: variability and constraints on ammonoid ornament and shell shape

Buckman's Law of Covariation states that ammonoid shell shape and ornamentation are typically correlated, such that compressed, involute forms have light ornament while more inflated, evolute forms have heavier ornament. Such covariation has been observed in many ammonoid groups, and implies a link between the morphogenesis of shell shape and ornamentation. However, other evidence suggests that while ornament growth is controlled by the genetic‐developmental program of the ammonoid, shell shape is strongly influenced by environmental factors. These differing viewpoints lead to Buckman's Paradox – are ornamentation and shell shape tightly linked, as implied by Buckman's covariation, or is the morphogenesis of ornament controlled genetically, while shell shape is controlled environmentally? To address this issue, the variability of shell shape and rib morphology has been compared for a group of endemic acanthoceratid ammonites from the Cretaceous Western Interior Seaway of North America. If Buckman's Law holds due to a morphogenetic connection between shell shape and ornamentation, we would expect taxa with more variable shell shapes to also show more variable rib features and growth. Morphometric analysis of seven shell shape and two rib characters for the Western Interior acanthoceratids finds no such correlation, suggesting that shell shape and rib growth are controlled by different processes. Indeed, rib growth appears to be more constrained than shell shape, consistent with the view that ornamentation is more tightly controlled by the developmental‐genetic growth program of the ammonoid. These results emphasize the complexity of ammonoid morphogenesis and highlight our limited understanding of the causes underlying Buckman's Law.     

PUNCTICULATA VERSIFORMIS SP. NOV. AND CYCLOTELLA KATHMANDUENSIS SP. NOV. (BACILLARIOPHYTA), NEW FOSSIL SPECIES FROM MIDDLE PLEISTOCENE LACUSTRINE SEDIMENTS,KATHMANDU, NEPAL HIMALAYA1

New fossil species, Puncticulata versiformis sp. nov. and Cyclotella kathmanduensis sp. nov., are described from lacustrine sediments in the Kathmandu Basin on the southern slope of the Nepal Himalaya. They were dominant in the Middle Pleistocene. Both LM and SEM observations reveal their unique morphological features. Puncticulata versiformis is characterized by (1) various valve outlines (circular, elliptical, or oval), (2) a tangentially undulate central area, (3) a complex alveolar structure composed of three kinds of costae (thick costae, thin costae, and thin and short costae), (4) well‐developed spines (Y‐shaped and tapering) located on one side of the valve face/mantle area junction, and (5) a valvocopula with an extremely undulate margin. The species‐specific feature of C. kathmanduensis is the presence of two kinds of alveolate zones in a single valve: type‐1 zone composed only of normal costae, and type‐2 zone composed of both normal costae and recessed costae bearing fultoportulae. Changes in valve ornamentation occur in these two species from initial valves to vegetative valves. In P. versiformis, the arrangement of areolae with internal domed cribra and fultoportulae in the central area changes from radial rows in the initial valve to groups in the vegetative valve. In the initial valve of C. kathmanduensis, the type‐1 alveolate zone is generally absent.     

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)     

Syn vivo hydrostatic and hydrodynamic properties of scaphitid ammonoids from the U.S. Western Interior

Scaphitid ammonoids were ubiquitous and significant components of the Western Interior Seaway during the Late Cretaceous. This group is characterized by a recurved hook at maturity that deviates from the juvenile whorls. Such a modification seems counterproductive to active locomotion and to manage a biologically effective orientation that facilitates efficient feeding and swimming. Virtually reconstructed 3D hydrostatic models reveal that the examined mature scaphitids had the capacity for neutral buoyancy while assuming a stable, upward-facing orientation in the water column during life. Models of juvenile Hoploscaphites nicolletii suggest that scaphitid apertures were oriented only slightly more horizontal than adults. The hydrostatic influence of sexual dimorphism was explored with the species Hoploscaphites crassus. The inflated macroconch has a lower stability and higher hydrodynamic drag compared to its microconch counterpart. The effect of shell compression was investigated by comparing H. crassus and the more compressed H. nicolletii. The latter species has a relatively high stability and much less hydrodynamic drag during movement. The mature U-shaped body chamber distributes organismal mass in a way that increases stability, and simultaneously orients the soft body so that propulsive energy is efficiently transmitted into horizontal backwards movement with minimal rocking. Swimming velocities computed from hydrodynamic drag experiments suggest that scaphitids were relatively slow swimmers with compressed forms attaining slightly higher velocities (when scaled by mass). Hydrodynamic lift was investigated with computational fluid dynamics simulations. These experiments revealed that the overall shape of the shell is responsible for significant lift in the upwards direction, which is not heavily influenced by ornamentation. This explains how a reduced soft body can overcome and manage a slightly negatively buoyant condition during life. Therefore, the seemingly cumbersome shape and orientation of the scaphitid morphotype may not have been a hindrance during locomotion.     

Sexual dimorphism in shells of Cochlostoma septemspirale (Caenogastropoda, Cyclophoroidea, Diplommatinidae, Cochlostomatinae)

Sexual dimorphisms in shell-bearing snails expressed by characteristic traits of their respective shells would offer the possibility for a lot of studies about gender distribution in populations, species, etc. In this study, the seven main shell characters of the snail Cochlostoma septemspirale were measured in both sexes: (1) height and (2) width of the shell, (3) height and (4) width of the aperture, (5) width of the last whorl, (6) rib density on the last whorl, and (7) intensity of the reddish or brown pigments forming three bands over the shell. The variation of size and shape was explored with statistical methods adapted to principal components analysis (PCA) and linear discriminant analysis (LDA). In particular, we applied some multivariate morphometric tools for the analysis of ratios that have been developed only recently, that is, the PCA ratio spectrum, allometry ratio spectrum, and LDA ratio extractor. The overall separation of the two sexes was tested with LDA cross validation.The results show that there is a sexual dimorphism in the size and shape of shells. Females are more slender than males and are characterised by larger size, a slightly reduced aperture height but larger shell height and whorl width. Therefore they have a considerable larger shell volume (about one fifth) in the part above the aperture. Furthermore, the last whorl of females is slightly less strongly pigmented and mean rib density slightly higher. All characters overlap quite considerably between sexes. However, by using cross validation based on the 5 continuous shell characters more than 90% of the shells can be correctly assigned to each sex.     

Segmental growth in planulate ammonites: inferences on costal function

Checa, Antonio & Westermann, Gerd E. G. 1989 01 15: Segmental growth in planulate ammonites: inferences on costal function. Lethaia , Vol. 22, pp. 95–100. Oslo. ISSN 0024–1164.
In planulate Ammonitina, the directions of costae and their parts coincide with the growth lines, both being accreted in parallel and subtriangular increments, i.e. segments. This implies that the growing margin was consistently corrugated (plicated) parallel to the edge of the peristome. The adventral separation of the ribs caused by the coiling increased with whorl involution and was compensated for by costal division into primaries and secondaries. We propose that costation of the shallow-water, nekto-benthic plandates reduced the stiffness and increased effective thickness of their growing peristome. Ribs functioned (primarily or secondarily) as a protection for the immature, fragile aperture against predators and impacts.□ Ammonite shell, Jurassic, morphology, growth lines, ornamentation, costae .     

Is there a link between shell morphology and parasites of zebra mussels?

The shell morphology of zebra mussels, Dreissena polymorpha, was analyzed to determine if alterations in shell shape and asymmetry between valves were related to its infection status, i.e. infected or not by microparasites like ciliates Ophryoglena spp. or intracellular bacteria Rickettsiales-like organisms (RLOs), and by macroparasites like trematodes Phyllodistomum folium and Bucephalus polymorphus. For microparasites, two groups of mussels were observed depending on shell measurements. Mussels with the more concave shells were the most parasitized by ciliates. This could be more a consequence than a cause and we hypothesized that a modification of the water flow through the mantle cavity could promote the infection with a ciliate. There were more RLOs present in the most symmetrical individuals. A potential explanation involved a canalization of the left-right asymmetry as a by-product of the parasite infection. Trematode infections were associated with different responses in valve width. Females infected by P. folium displayed significantly higher symmetry in valve width compared with non-infected congeners, whereas the infection involved an opposite pattern in males. B. polymorphus was also linked to a decrease in valve width asymmetry. This study suggested that a relationship exists between parasitism and shell morphology through the physiological condition of host zebra mussels.     

Leaf anatomy of a rheophyte,Dendranthema yoshinaganthum (Asteraceae), and of hybrids between D. yoshinaganthumand a closely related non-rheophyte,D. indicum

The rheophyte Dendranthema yoshinaganthum(Makino ex Kitam.) Kitam. is endemic to a region along the Naka River in Shikoku, Japan. Resembling other rheophytes that have adapted to environments along rivers, D. yoshinaganthumhas narrow leaves. In the native habitat of D. yoshinaganthumin Tokushima, Japan, natural hybridization with Dendranthema indicum (L.) Des Moul. has resulted in a species complex with conspicuous variation in leaf shape and, in particular, in leaf width. In this study, D. yoshinaganthum and hybrids with natural variation in leaf shape were cultivated under identical conditions and the anatomy of their leaves was examined. Variation in leaf width in the species complex was attributable, for the most part, to variation in the number of cells per leaf lamina and also, to some extent, to variation in cell size. This is the first report of natural variation in leaf width that is attributable to variation in both the size and the number of leaf cells.     

Buckman's first law of covariation–a case of proportionality

Buckman's first law of covariation states that in ammonoids, coarseness of ribs and other ornamentation correlates negatively with degree of compression (whorl height/width ratio) and involution. This ‘law’ has escaped several attempts at functional or physiological explanation. However, Buckman's first law becomes self‐evident, and almost collapses to a tautology, when we reword it as follows: Size of lateral and ventral ornamentation correlates with size of the aperture (its width and height, respectively). In a shell that is strongly compressed laterally, the diameter of the aperture, and hence the soft parts within, is small in the lateral direction. For such a shell, coarse lateral ribbing would imply impossibly strong relative compression and expansion of the soft body through the formation of a rib. Conversely, if the shell is depressed (compressed dorsoventrally), the diameter of the soft parts is small in the dorsoventral direction, and the ventral ribbing correspondingly small. Buckman's first law is thus simply a statement of proportionality that needs no special explanation. On the contrary, any departure from this ‘law’ would imply strong deformations of the soft body as ornamentation was formed. Measurements on the Triassic ammonite Pseudodanubites halli show that ventral ribs get relatively stronger on compressed specimens, in accordance with this view of Buckman's law. ? Ammonoids, intraspecific variation, morphometry, ornamentation.     

VARIATION IN ANATOMICAL AND MATERIAL PROPERTIES EXPLAINS DIFFERENCES IN HYDRODYNAMIC PERFORMANCES OF FOLIOSE RED MACROALGAE (RHODOPHYTA)1

Over the last two decades, many studies on functional morphology have suggested that material properties of seaweed tissues may influence their fitness. Because hydrodynamic forces are likely the largest source of mortality for seaweeds in high wave energy environments, tissues with material properties that behave favorably in these environments are likely to be selected for. However, it is very difficult to disentangle the effects of materials properties on seaweed performance because size, shape, and habitat also influence mechanical and hydrodynamic performance. In this study, anatomical and material properties of 16 species of foliose red macroalgae were determined, and their effects on hydrodynamic performance were measured in laboratory experiments holding size and shape constant. We determined that increased blade thickness (primarily caused by the addition of medullary tissue) results in higher flexural stiffness (EI), which inhibits the seaweed’s ability to reconfigure in flowing water and thereby increases drag. However, this increase is concurrent with an increase in the force required to break tissue, possibly offsetting any risk of failure. Additionally, while increased nonpigmented medullary cells may pose a higher metabolic cost to the seaweed, decreased reconfiguration causes thicker tissues to expose more photosynthetic surface area incident to ambient light in flowing water, potentially ameliorating the metabolic cost of producing these cells. Material properties can result in differential performance of morphologically similar species. Future studies on ecomechanics of seaweeds in wave‐swept coastal habitats should consider the interaction of multiple trade‐offs.     

Putative chromosomal rearrangements are associated primarily with ecotype divergence rather than geographic separation in an intertidal,poorly dispersing snail

Littorina saxatilis is becoming a model system for understanding the genomic basis of ecological speciation. The parallel formation of crab‐adapted ecotypes that exhibit partial reproductive isolation from wave‐adapted ecotypes has enabled genomic investigation of conspicuous shell traits. Recent genomic studies suggest that chromosomal rearrangements may enable ecotype divergence by reducing gene flow. However, the genomic architecture of traits that are divergent between ecotypes remains poorly understood. Here, we use 11,504 single nucleotide polymorphism (SNP) markers called using the recently released L. saxatilis genome to genotype 462 crab ecotype, wave ecotype and phenotypically intermediate Spanish L. saxatilis individuals with scored phenotypes. We used redundancy analysis to study the genetic architecture of loci associated with shell shape, shape corrected for size, shell size and shell ornamentation, and to compare levels of co‐association among different traits. We discovered 341 SNPs associated with shell traits. Loci associated with trait divergence between ecotypes were often located inside putative chromosomal rearrangements recently characterized in Swedish L. saxatilis. In contrast, we found that shell shape corrected for size varied primarily by geographic site rather than by ecotype and showed little association with these putative rearrangements. We conclude that genomic regions of elevated divergence inside putative rearrangements were associated with divergence of L. saxatilis ecotypes along steep environmental axes—consistent with models of adaptation with gene flow—but were not associated with divergence among the three geographical sites. Our findings support predictions from models indicating the importance of genomic regions of reduced recombination allowing co‐association of loci during ecological speciation with ongoing gene flow.     

新疆18种木灵藓属(Orthotrichum Hedw.)植物解剖结构研究

采用徒手切片法对新疆木灵藓属(Orthotrichum Hedw.)18种植物孢蒴上的气孔、茎和叶进行了比较解剖学观察。研究结果表明,木灵藓属植物茎横切结构形状稳定,由表皮和皮层组成,无中轴细胞分化;除半裸木灵藓(Orthotrichum hallii Sull.et Lesq.)外,其它种叶上部均为单层细胞;叶中部边缘外卷,叶细胞具有高度不同的分叉疣或单疣。在确定气孔显、隐型的基础上,茎和叶横切面的颜色、茎表皮和皮层细胞组成及细胞壁加厚程度、叶细胞疣的高度、叶中肋横切面的形状、中肋的宽度、细胞组成以及芽孢形态等特征可作为该属种间分类的依据。还根据18种木灵藓属植物孢蒴上的气孔显、隐型,及茎、叶的比较解剖学特征,编制了新疆木灵藓属植物的分种检索表。     

新疆18种木灵藓属(Orthotrichum Hedw.) 植物解剖结构研究

采用徒手切片法对新疆木灵藓属（Orthotrichum Hedw.）18种植物孢蒴上的气孔、茎和叶进行了比较解剖学观察。研究结果表明,木灵藓属植物茎横切结构形状稳定,由表皮和皮层组成,无中轴细胞分化;除半裸木灵藓（Orthotrichum hallii Sull.et Lesq.）外,其它种叶上部均为单层细胞;叶中部边缘外卷,叶细胞具有高度不同的分叉疣或单疣。在确定气孔显、隐型的基础上,茎和叶横切面的颜色、茎表皮和皮层细胞组成及细胞壁加厚程度、叶细胞疣的高度、叶中肋横切面的形状、中肋的宽度、细胞组成以及芽孢形态等特征可作为该属种间分类的依据。还根据18种木灵藓属植物孢蒴上的气孔显、隐型,及茎、叶的比较解剖学特征,编制了新疆木灵藓属植物的分种检索表。     

青藏高原东缘西藏沙棘(Hippophae tibetana)果实性状变异

西藏沙棘是沙棘属(Hippophae L.)中分布于青藏高原和喜马拉雅高海拔地区的特有类群之一,其果实大小、形状、颜色以及条纹长等性状表现出了高度的多样性。对分布于青藏高原东缘西藏沙棘7个天然居群的果长、果宽、果形系数(果长/果宽)、果柄长、纹长等果实性状变异进行了统计分析,探讨了其果实变异水平及地理变异式样。结果表明,西藏沙棘5个果实性状在居群间和居群内均存在着极显著差异(p0.01)(除纹长在居群间存在着显著差异(p0.05)),居群间变异系数(CV)在11.19%~45.03%之间,果柄长的变异幅度最大(CV为45.03%),其次为纹长(CV为31.97%),变异幅度最小的是果宽(CV为11.19%);表型分化系数(Vst)在0.213 3~0.629 9之间,平均值为0.489 5,居群间和居群内变异约各占果实变异的一半。相关性分析显示,除果长与果柄、果宽与纹长外,其它各性状间均表现出极显著相关(p0.01),除果长与海拔、果宽与经纬度外,其它各性状与地理因子之间均达到极显著相关(p0.01),呈现出地理梯度变异,聚类分析表明7个居群按照地理距离远近聚类。     

Length and shape variants of the bacteriophage T4 head: mutations in the scaffolding core genes 68 and 22.

The shape and size of the bacteriophage T4 head are dependent on genes that determine the scaffolding core and the shell of the prohead. Mutants of the shell proteins affect mainly the head length. Two recently identified genes (genes 67 and 68) and one already known gene (gene 22), whose products are scaffold constituents, have been investigated. Different types of mutants were shown to strongly influence the proportion of aberrantly shaped particles. By model building, these shape variants could be represented as polyhedral bodies derived from icosahedra, through outgrowths along different polyhedral axes. The normal, prolate particle is obtained by elongation along a fivefold axis. The mutations of the three core genes (genes 67, 68, and 22) affect the width mainly by lateral outgrowths of the prolate particle, although small and large isometric particles are also found. Many of the aberrant particles are multitailed, suggesting a correlation between tail attachment sites and shape.     

Large‐scale evolutionary trends of Acrochordiceratidae Arthaber, 1911 (Ammonoidea,Middle Triassic) and Cope’s rule

Abstract: Directed evolution of life through millions of years, such as increasing adult body size, is one of the most intriguing patterns displayed by fossil lineages. Processes and causes of such evolutionary trends are still poorly understood. Ammonoids (externally shelled marine cephalopods) are well known to have experienced repetitive morphological evolutionary trends of their adult size, shell geometry and ornamentation. This study analyses the evolutionary trends of the family Acrochordiceratidae Arthaber, 1911 from the Early to Middle Triassic (251–228 Ma). Exceptionally large and bed‐rock‐controlled collections of this ammonoid family were obtained from strata of Anisian age (Middle Triassic) in north‐west Nevada and north‐east British Columbia. They enable quantitative and statistical analyses of its morphological evolutionary trends. This study demonstrates that the monophyletic clade Acrochordiceratidae underwent the classical evolute to involute evolutionary trend (i.e. increasing coiling of the shell), an increase in its shell adult size (conch diameter) and an increase in the indentation of its shell suture shape. These evolutionary trends are statistically robust and seem more or less gradual. Furthermore, they are nonrandom with the sustained shift in the mean, the minimum and the maximum of studied shell characters. These results can be classically interpreted as being constrained by the persistence and common selection pressure on this mostly anagenetic lineage characterized by relatively moderate evolutionary rates. Increasing involution of ammonites is traditionally interpreted by increasing adaptation mostly in terms of improved hydrodynamics. However, this trend in ammonoid geometry can also be explained as a case of Cope’s rule (increasing adult body size) instead of functional explanation of coiling, because both shell diameter and shell involution are two possible paths for ammonoids to accommodate size increase.     

Physiological variation in the dog-whelk Nucella lapillus, L. either side of a cline in allozyme and karyotype frequencies

Genetic constitution in the intertidal gastropod Nucella lapillus influences variation in shell shape and growth rate which in turn are correlated with such habitat variables as wave action and temperature. We have investigated the response to hyperosmotic stress of samples from a cline in karyotype and allozyme frequencies and shell shape. Animals with a shell shape associated with environments where temperature and desiccation stress are important respond less to hyperosmotic stress than animals living in a high wave energy environment. With regard to the interaction between shell shape, physiology and habitat, animals with elongate shells associated with protected shores are shown to exhibit a reduced response to hyperosmotic stress compared to animals with a more spherical shell shape; this is discussed in relation to the production of an adaptive phenotype.