Zebra mussels decrease burrowing ability and growth of a native snail, <Emphasis Type="Italic">Campeloma decisum</Emphasis>

Invasive species can drive native organisms to extinction by limiting movement and accessibility to essential resources. The purpose of this study was to determine if zebra mussels (Dreissena polymorpha) affect the burrowing ability and growth rate of a native snail, Campeloma decisum. Snails with and without zebra mussels were collected from Douglas Lake, MI, and burrowing depths were studied in both the laboratory and Douglas Lake. Growth rates were calculated as the amount of shell growth from 2004 to 2005. Both the tendency of snails to burrow and the depth to which they burrowed into the substrate were significantly decreased by the presence of zebra mussels on snail shells in both laboratory and lake experiments. There was no difference in the percentage of snails that exhibited growth as a function of zebra mussel presence. However, for those snails that grew, there was a 50% higher growth rate for snails without zebra mussels compared to snails with zebra mussels. These negative effects of zebra mussels on growth and burrowing ability will likely lead to decreases in snail population densities in the future. Handling editor: S. Wellekens     

Push and bite: trade‐offs between burrowing and biting in a burrowing skink (Acontias percivali)

Trade‐offs are thought to be important in constraining evolutionary divergence, as they may limit phenotypic diversification. Limbless animals that burrow head‐first have been suggested to be evolutionarily constrained in the development of a large head size and sexual head shape dimorphism because of potential trade‐offs associated with burrowing. Here we use an acontiine skink (Acontias percivali) to test for the existence of trade‐offs between traits thought to be important in burrowing (speed and force). As head size dimorphism has been shown to be limited in acontiine lizards, thus suggesting constraints on head size and shape, we additionally explore the potential for trade‐offs between burrowing and biting. Our data show that A. percivali uses a burrowing style different from those previously described for caecilians and amphisbaenians, which relies on the use of extensive lateral and dorsoventral head movements. Our data also show that animals use their entire bodies to generate force, as peak force was determined by total length only. Additionally, both bite force and the time needed to burrow into the substrate were principally determined by relative head width, suggesting a trade‐off between biting and burrow speed. Performance data were indeed suggestive of a correlation between bite force and the time needed to burrow, but additional data are needed to confirm this pattern. In summary, our data suggests that trade‐offs may exist, and may have been of crucial importance in shaping the evolution of head shape in A. percivali, and burrowing lizards more generally. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 91–99.     

Burrowing mechanisms and sculptures in Recent gastropods

Savazzi, Enrico 1989 01 15: Burrowing mechanisms and sculptures in Recent gastropods. Lethaia , Vol. 22, pp. 31–48. Oslo. ISSN 0024–1164.
Burrowing was observed in 32 gastropod species, belonging to 8 families, from Italy and the Philippines. Most species burrow by repeating a three-phase sequence: (1) digging with the foot, (2) dragging the shell forward and downward, and (3) rocking the shell around its longitudinal axis. Minor specific differences in the burrowing dynamics are common, and totally different mechanisms also occur. Burrowing sculptures consisting of terraces or asymmetrical tubercles are observed in the majority of the studied species. Characteristics of the burrowing process explain some cases of apparent divergence of burrowing sculptures from the paradigm. Burrowing sculptures in the Gastropoda should be expected to occur mostly among medium-slender, rather than markedly high-spired, shell morphologies. □ Mollusca, Gastropoda, Cerithiidae, Nassariidae, Mitridae, Costellariidae, Conidae, Terebridae, Turridae, Hydro-biidae, shell, sculpture, burrowing, functional morphology, ecology, behaviour, Holocene, Indo-Pacific. Philippines, Mediterranean, Italy .     

Interactions between locomotion,feeding, and bodily elongation during the evolution of snakes

Information from lizard lineages that have evolved a highly elongate (snake‐like) body form may clarify the selective forces important in the early evolution of snakes. Lizards have evolved bodily elongation via two distinct routes: as an adaptation to burrowing underground or to rapid locomotion above ground. These two routes involve diametrically opposite modifications to the body plan. Burrowing lizards have elongate trunks, small heads, short tails, and relatively constant body widths, whereas surface‐active taxa typically have shorter trunks, wider heads, longer tails, and more variable body widths. Snakes resemble burrowing rather than surface‐active (or aquatic) lizards in these respects, suggesting that snakes evolved from burrowing lizards. The trunk elongation of burrowing lizards increases the volume of the alimentary tract, so that an ability to ingest large meals (albeit consisting of small individual prey items) was present in the earliest snakes. Subsequent shifts to ingestion of wide‐bodied prey came later, after selection dismantled other gape‐constraining morphological attributes, some of which may also have arisen as adaptations to burrowing through hard soil (e.g. relatively small heads, rigid skulls). Adaptations of snake skulls to facilitate ingestion of large prey have evolved to compensate for the reduction of relative head size accompanying bodily elongation; relative to predator body mass, maximum sizes of prey taken by snakes may not be much larger than those of many lizards. This adaptive scenario suggests novel functional links between traits, and a series of testable predictions about the relationships between squamate morphology, habitat, and trophic ecology. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 293–304.     

Dislodgement force and shell morphology vary according to wave exposure in a tropical gastropod (Cittarium pica)

Wave exposure has strong influences on population density, morphology and behaviour of intertidal species in temperate zones, but little is known about how intertidal organisms in tropical regions respond to gradients in wave exposure. We tested whether dislodgement force and shell shape of a tropical gastropod, Cittarium pica, differs among shores that vary in wave exposure. After adjusting for body size, we found that C. pica from exposed shores required greater dislodgement force to remove them from the shore, had slightly larger opercula (the closure to the shell aperture), and were slightly squatter in shape (reduced in shell height relative to shell width) than C. pica from sheltered shores. These morphological adjustments are consistent with those observed in temperate gastropods, which are argued to represent adaptive responses to the risk of mortality associated with dislodgement.     

PATTERNS OF ORIENTATION IN UNIONIDS AS A FUNCTION OF RIVERS WITH DIFFERING HYDROLOGICAL VARIABILITY

As unionids can become dislodged with high flows, it may provebeneficial for an individual to minimize its exposure to theflow. This can be accomplished by either burrowing as deep aspossible or orienting in a way that effectively reduces thedrag exerted on the mussel by the flow. The patterns of orientationwere examined in unionids with respect to hydrological variability.The orientation of mussels to flow was measured at four sitesalong an event river (hydro-logically variable) and a stableriver (hydrologically stable). Burrowing depth was measuredat a reference site in each river. Most individuals in bothriver types were oriented with their siphons pointing upstream.Mussel orientation differed significantly between the two rivertypes with mussels in the event river orienting more parallelto the flow than those in the stable river. Mussels in the eventriver were significantly larger than those in the stable riverbut the size of a mussel did not determine its orientation withina river. Burrowing depth did not differ for mussels betweenevent and stable rivers. The observed differences in orientationsamong river types are likely a function of differences in thepattern of orientation of the mussel community as a whole, withineach drainage. This burrowing behaviour may be an attributethat enhances the adaptations mussels have for remaining burrowedin the sediment. (Received 2 December 1996; accepted 24 February 1997)     

Identification of ecophenotypic trends within three European freshwater mussel species (Bivalvia: Unionoida) using traditional and modern morphometric techniques

Most species of freshwater mussels (Unionoida) show a wide variability in shell form and size but an understanding of which factors determine unionoid morphology is poor. We identified ecophenotypic trends in shell and internal characters within three unionoid species from two habitat types (marinas and river) of the River Thames, UK, using traditional and modern morphometric techniques. In marinas, all species grew to larger maximum sizes than in the river, which might be a result of higher temperatures and phytoplankton densities in marinas. Unio pictorum in marinas was more elongated than in the river and Fourier shape analysis revealed a trend from dorsally arched river specimens to straight dorsal and pointed posterior margins in marina individuals. The degree of shell elongation and shape of dorso‐posterior margin were not associated with sediment composition, but were associated with the different hydrological characters of the two habitat types. Relative shell width was a poor indicator of collection site and influenced by allometric growth. Unlike U. pictorum, a difference in shell elongation of marina and river mussels could not be detected in Unio tumidus and Anodonta anatina. However, all three species showed the same trends regarding the shape of the dorso‐posterior shell margin. This shell character may thus have broad ecological significance and could have considerable utility to palaeontologists, taxonomists, and conservation biologists. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 814–825.     

The effects of latitude,body size,and sexual selection on wing shape in a damselfly

Under natural selection, wing shape is expected to evolve to optimize flight performance. However, other selective factors besides flight performance may influence wing shape. One such factor could be sexual selection in wing sexual ornaments, which may lead to alternative variations in wing shape that are not necessarily related to flight performance. In the present study, we investigated wing shape variations in a calopterygid damselfly along a latitudinal gradient using geometric morphometrics. Both sexes show wing pigmentation, which is a known signal trait at intra‐ and interspecific levels. Wing shape differed between sexes and, within the same sex, the shape of the hind wing differed from the front wing. Latitude and body size explained a high percentage of the variation in wing shape for female front and hind wings, and male front wings. In male hind wings, wing pigmentation explained a high amount of the variation in wing shape. On the other hand, the variation in shape explained by pigmentation was very low in females. We suggest that the conservative morphology of front wings is maintained by natural selection operating on flight performance, whereas the sex‐specific differences in hind wings most likely could be explained by sexual selection. The observed sexual dimorphism in wing shape is likely a result of different sex‐specific behaviours. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 263–274.     

Morphological trends and environment: a case study on Early Cretaceous Myophorelloidea (Bivalvia,Trigoniida) from Neuquén Basin,Argentina

Trigoniide clams were conspicuous components of bivalve faunas in shallow seas during most of the Mesozoic. Morphological trends in different features of shell shape and ornamentation of three related species of myophorelloids from the Agrio Formation (Early Cretaceous of Neuquén Basin, Argentina) that inhabited environments of different hydrodynamic energy conditions are explored in this study by means of a detailed morphometric analysis. Four hydrodynamic energy categories were established for depositional environments, from most (1), to least (4), energetic. General shell shape was characterized by measurements of height, length and width. New quantitative methods were developed to evaluate the distribution of costae along the marginal carina and the divergence of costae along the flank. Morphometric variables were detrended with respect to size, to avoid ontogenetic variability in further analyses. Relative width of the shell increases as environments become higher in hydrodynamic energy; this unusual trend may be interpreted as an anchoring/stabilizing strategy developed by the lineage in shallow marine environments. Trends in ornamentation correspond to more densely distributed costae in environments of higher hydrodynamic energy, a pattern that improves substrate penetration.     

Movements and burrowing activity in the Antarctic bivalve molluscs<Emphasis Type="Italic"> Laternula elliptica</Emphasis> and<Emphasis Type="Italic"> Yoldia eightsi</Emphasis>

Burrowing was investigated in two Antarctic infaunal bivalve molluscs, Laternula elliptica and Yoldia eightsi, representing amongst the least and most active members of the class Bivalvia in the Southern Ocean. Burrowing rate was expressed via the Burrowing Rate Index (BRI=[³wet weight/time to bury]×10⁴), and produced values of 0.1–10.6 for L. elliptica and 8.8–49.8 for Y. eightsi. These compare with values ranging from 3 to 2,000 for N. American bivalves (mean=222, SE=42.6, n=81), and 200 to 2,200 for Hong Kong bivalves (mean=1,140, SE=346, n=6). Values for the Antarctic species are, therefore, low compared to warmer-water bivalves, and the values below 1 for large L. elliptica are the lowest on record by around ×5. There is no compensation of burrowing activity for low temperature in these species. The relative BRI values for L. elliptica and Y. eightsi reflect the differences in their mode of life, with the former being large, sedentary and suspension-feeding, and the latter being smaller, mobile, ploughing through the sediment and feeding on sediment-surface organic matter. Burrowing in L. elliptica is unexpected, because other members of the Laternulidae do not burrow. This ability is most probably a response to the regular disturbance of sediments in Antarctica by ice, and the strong selective advantage to being able to resume a protected position after disturbance. The burrowing cycle in L. elliptica is composed of three main phases: (1) foot extension and sediment penetration; (2) foot dilation to form an anchor; (3) the drawing down of the shell by contraction of the pedal retractor muscles. Burrowing in Y. eightsi also has three phases: (1) foot extension and penetration of the sediment (digging); (2) rocking movements in the upright position; (3) shell anchorage. In excess of burrowing activity, L. elliptica exhibits a unique suite of movements when exposed at the surface. These comprise levering, where the tips of the siphons are pressed against the sediment to lift the shell from the substratum, looping, where the siphons are extended and rotated and, in the process, translocate the whole animal across the sediment, and jetting, where water is ejected forcibly through the siphons while their tips are directed towards the sediment, lifting part or all of the animal clear of the substratum. In the field, following exhumation by icebergs, these activities serve to place the animal in a favourable position for reburial, which is a clear advantage in disturbed polar environments where predatory nemerteans and asteroids are abundant.     

Influence of genotype and geography on shell shape and morphometric trait variation among North Atlantic blue mussel (Mytilus spp.) populations

The influence of geography and genotype on shell shape (outline) and trait (morphometric) variation among North Atlantic blue mussels and their hybrids has been examined. Shape differences among reference taxa (Mytilus trossulus, Mytilus edulis and Mytilus galloprovincialis) were consistent with an association between taxon‐specific genes and shape genes. Newfoundland M. edulis × M. trossulus populations and northern Quebec M. trossulus populations exhibited an uncoupling of taxon‐specific genes from shape genes, whereas Nova Scotia M. trossulus populations and SW England M. edulis × M. galloprovincialis populations exhibited an association between taxon‐specific genes and shape genes. We found no evidence of a geographic effect (NE versus NW Atlantic) for shape variation, indicating that the genotype effect is stronger than any geographic effect at macrogeographic scales. Pronounced differences were observed in trait variability consistent with an association between taxon‐specific genes and trait genes in European populations, and trait divergence of New York M. edulis from all European mussels. Trait variability in mussels from Newfoundland, Nova Scotia and northern Quebec indicated an uncoupling of taxon genes from trait genes, whereas trait variability in SW England M. edulis × M. galloprovincialis populations was consistent with background genotype, indicating a strong association between taxon genes and trait genes. A pronounced macrogeographic split (NE versus NW Atlantic) regardless of taxonomy was observed, indicating that geography exerts a greater influence than genotype on trait variation at the macrogeographic scale. This is consistent with pronounced within‐taxon genetic divergence, indicative of different selection regimes or more likely of different evolutionary histories of mussels on either side of the North Atlantic. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96 , 875–897.     

Heart rate and burrowing behavior in the mussel Anodontites trapesialis (Bivalvia: Mycetopodidae) from lotic and lentic sites

Burrowing rate index, diurnal variations in heart rate and heart rate alterations associated with behavioral patterns were studied in Anodontites trapesialis from the Pardo River (PR) and the Galo Bravo Dam (GBD). In both populations the pattern immobile/buried was associated with the lowest (basal) values of heart rate, which increase significantly during burrowing remaining steady in its phases I and II and at the first 10 min of phase III. A decrease in the frequency of digging cycles was significantly correlated with the gradual decrease in heart rate during the phase III of burrowing. Nevertheless, mussels from PR burrow faster than those of GBD and possess higher values of basal heart rate which present a diurnal variation with the higher values occurring during the light phase. Yet, the basal heart rate in mussels from GBD remained stable over the course of 24 h. Also, the increase in heart rate associated with burrowing was higher in mussels from PR and compatible with the highest burrowing index previously detected. Since mussels possess a plastic phenotype it is probable that the cited functional differences are phenotypic adaptations, but we cannot exclude the possibility of these adaptations being genotypic in nature.     

Commensals of Mytilus galloprovincialis in the Black Sea: Urastoma cyprinae (Turbellaria) and Polydora ciliata (Polychaeta)

The intensity of infection of the mussel Mytilus galloprovincialis Lamarck in the Black Sea by the turbellarian Urastoma cyprinae (Graff), which lives on its gills, was found to be higher in larger hosts, reaching a maximum in mussels of 50–70 mm length. Greater numbers occured in mussels inhabiting a silty bottom than in cultivated mussels suspended above the bottom. Over the period 1982–1987, U. cyprinae was most numerous in winter and especially so in years that were colder. The spionid polychaete Polydora ciliata (Johnston) also infects M. galloprovincialis, burrowing into the shell. Young spionids of up to 1 mm length occured in mussels with a shell length of 35 mm. Numbers of this commensal reached a maximum in mussels of intermediate size.     

First trace and body fossil evidence of a burrowing, denning dinosaur

A fossil discovery in the mid-Cretaceous Blackleaf Formation of southwest Montana, USA, has yielded the first trace and body fossil evidence of burrowing behaviour in a dinosaur. Skeletal remains of an adult and two juveniles of Oryctodromeus cubicularis gen. et sp. nov. a new species of hypsilophodont-grade dinosaur, were found in the expanded distal chamber of a sediment-filled burrow. Correspondence between burrow and adult dimensions supports Oryctodromeus as the burrow maker. Additionally, Oryctodromeus exhibits features of the snout, shoulder girdle and pelvis consistent with digging habits while retaining cursorial hindlimb proportions. Association of adult and young within a terminal chamber provides definitive evidence of extensive parental care in the Dinosauria. As with modern vertebrate cursors that dig, burrowing in Oryctodromeus may have been an important adaptation for the rearing of young. Burrowing also represents a mechanism by which small dinosaurs may have exploited the extreme environments of polar latitudes, deserts and high mountain areas. The ability among dinosaurs to find or make shelter may contradict some scenarios of the Cretaceous-Paleogene impact event. Burrowing habits expand the known range of nonavian dinosaur behaviours and suggest that the cursorial ancestry of dinosaurs did not fully preclude the evolution of different functional regimes, such as fossoriality.     

Identification and evaluation of the Atlantic razor clam (Ensis directus) for biologically inspired subsea burrowing systems

In this article, we identify and analyze a subsea organism to serve as a model for biologically inspired burrowing technology to be used in applications such as anchoring, installation of cables, and recovery of oil. After inspecting myriad forms of life that live on or within ocean substrates, the Atlantic razor clam, Ensis directis, stood out as an attractive basis for new burrowing technology because of its low-energy requirements associated with digging (0.21?J/cm), its speed and depth of burrrowing (～1?cm/s and 70?cm, respectively), and its size and simplicity relative to man-made machines. As anchoring is a prime application for the technology resulting from this work, the performance of an Ensis directus-based anchoring system was compared to existing technologies. In anchoring force per embedment energy, the E. directus-based anchor beats existing technology by at least an order of magnitude. In anchoring force per weight of device, the biologically inspired system weighs less than half that of current anchors. The article concludes with a review of E. directus's digging strategy, which involves motions of its valves to locally fluidize the substrate to reduce burrowing drag and energy, and the successful adaptation of E. directus's burrowing mechanisms into an engineering system: the RoboClam burrowing robot, which, like the animal, uses localized fluidization to achieve digging energy that scales linearly with depth, rather than depth squared, for moving through static soil.     

Do sturgeon limit burrowing shrimp populations in Pacific Northwest Estuaries?

Green sturgeon, Acipenser medirostris, and white sturgeon, Acipenser transmontanus, are frequent inhabitants of coastal estuaries from northern California, USA to British Columbia, Canada. An analysis of stomach contents from 95 green sturgeon and six white sturgeon commercially landed in Willapa Bay, Grays Harbor, and the Columbia River estuary during 2000–2005 revealed that 17–97% had empty stomachs, but those fish with items in their guts fed predominantly on benthic prey items and fish. Burrowing thalassinid shrimp (mostly Neotrypaea californiensis) were important food items for both white and especially for green sturgeon taken in Willapa Bay, Washington during summer 2003, where they represented 51% of the biomass ingested (84.9% IRI). Small pits observed in intertidal areas dominated by these shrimp, are likely made by these sturgeon and we present evidence from exclusion studies and field observation that the predator making the pits can have a significant cumulative negative effect on burrowing shrimp density. These burrowing shrimp present a threat to the aquaculture industry in Washington State due to their ability to de-stabilize the substrate on which shellfish are grown. Despite an active burrowing shrimp control program in these estuaries, it seems unlikely that current burrowing shrimp abundance and availability as food is a limiting factor for threatened green sturgeon stocks. However, these large predators may have performed an important top down control function on shrimp populations in the past when they were more abundant.     

Survival,behaviour and spatial distribution of shell morphs in a population of the snail Brephulopsis bidens (Pulmonata)

Summary Microspatial variation of banded and unbanded shell morphs frequencies as well as number of individuals per m², mortality, migration and burrowing into the ground were examined in a population of snail Brephulopsis bidens found in South Crimea mountains (USSR). Differential values of relative survival of morphs were determined by their thermotolerance. The relative survival of the banded morph was lower at the west sites of population area (W=0.273), and increased gradually up to 1 at the east sites. Survival of the banded morph was dependent on its burrowing activity. Differences in relative survival of morphs decreased parallel with increasing general mortality of snails.Burrowing activity and intensity of migration of the banded morph were significantly higher than that in unbanded. In experiments with artificial shaded sections, the banded morph preferred shaded sections, whereas unbanded chose illuminated sites. All these differences in behaviour probably form the main factors for microspatial variation of morph frequencies.     

Adulthood and phylogenetic analysis in gastropods: character recognition and coding in shells of Lavigeria (Cerithioidea, Thiaridae) from Lake Tanganyika

In the study of gastropod shell morphology, determination of comparable ontogenetic stages is crucial, because all the states that various shell features go through during ontogeny are preserved on the shell. The protoconch/teleoconch transition and marks of episodic growth are among the few ways of defining discrete, comparable, growth stages. In gastropods with determinate growth the attainment of adulthood may provide additional shell markers permitting comparison among individuals and taxa. Adulthood is reflected in shell morphology in ways as diverse as shell deposition covering all the previous whorls and radically changing the shape of the shell through to slight changes in the trajectory of the suture. While the very prominent adulthood‐related changes of shell morphology have been used as systematic characters, the more moderate changes have not been studied in detail and their potential systematic value has been ignored. In this paper we give a detailed account of adult modifications of the shell appearing with cessation of growth. Our study group comprises eight closely related species of Lavigeria from Lake Tanganyika. We show that the ways adulthood is manifested are quite diverse. We describe eight characters of the aperture, the suture and the sculptural ornamentation. Character occurrence varies greatly among species. We show that characters appear in suites and that in many cases their appearance is connected to size. We use size as a proxy for adulthood and test whether character occurrence alone or its connection to size can help resolve species relationships. In both cases our characters confirm the monophyly of our ingroup and yield cladograms with various degrees of resolution of ingroup relationships. The coding method that yields the greater character congruence is the one that takes into consideration the connection between appearance of a character and size. This study demonstrates that ontogenetically correlated character transformations may nevertheless be phylogenetically independent. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140 , 223?240.     

Burrowing behavior of Dermatonotus muelleri (Anura,Microhylidae) with reference to the origin of the burrowing behavior of Anura

Dermatonotus muelleri is a forelimbs-head-first burrowing frog that uses its forelimbs for soil removal, and it is the second anuran species known to arch its head downwards at an angle of almost 90° to the longitudinal axis of its body when burrowing. The burrowing behavior of D. muelleri is divided in three stages: head burrowing, body burrowing, and chamber construction. Burrowing in D. muelleri includes construction of a subterranean chamber used for estivation during the dry season. Phylogenetic analysis based on literature survey of burrowing behavior suggested that head-first burrowing behavior has evolved several times in anuran history, forming a convergence complex, and that hindlimbs-first burrowing is a basal behavior.     

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.