Multiple scale patterns of shell and anatomy variability in land snails: the case of the Sicilian Marmorana (Gastropoda: Pulmonata, Helicidae)

Certain major aspects of phenotypic diversity are still largely unexplained. When phenotypic patterns do not relate to habitat variables, fine analysis of morphological patterns and their distribution sheds light on the origin of diversity. Among invertebrates, snails are an ideal model for studying the roles of the neutral processes and selection involved in creating diversity. To understand patterns and processes of variability on different scales (regional: areas; local: sites), morphological variability of two sets of characters (shell and genitalia) was quantified in a group of rock-dwelling land snails of the genus Marmorana (Pulmonata, Helicidae). To analyse shell variability, partitioning of the overall variation into size and shape components was analysed by a principal component-based approach. Shell shape and size variability is not significantly influenced by any environmental pressure. Variability at site scale is mainly attributed to shell size, which is a trait demonstrated to have a high degree of phenotypic plasticity. No sharp changes were observed for genitalia. Moreover, allometries between shell size and genitalia measurements involve a few populations. The observed multiple scale patterns are in line with the hypothesis that genital variance may be selectively controlled to maintain function.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 359–370.     

Geometric morphometrics, neural networks and diagnosis of sibling Taterillus species (Rodentia, Gerbillinae)

The lack of diagnostic traits in sibling species means that great quantities of biogeographical and ecological data held in museum collections cannot be utilized effectively, leading to underestimates of biodiversity. In this study we applied neural networks (NN) and canonical variates analyses (CVA) to landmark measurements on the skulls of the West African species Taterillus arenarius , T. petteri , T. gracilis and T. pygargus in an attempt to discriminate species previously identified unambiguously from their karyotypes. Among suggested differences, the relationship between inflation of the tympanic bullae and lower population density is discussed. Cross-validated classification rates did not exceed 73%. Two hypotheses are proposed to explain such high phenotypic similarity: morphological plasticity limited by environmental constraints and the possibility that speciation has been too recent to allow significant morphological divergence.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 319−327.     

Correlating head shape with ecological variables in rock-dwelling haplochromines (Teleostei: Cichlidae) from Lake Victoria

An exploratory analysis was made of the head shape of cichlid fishes. A 3-D head truss was used, which allowed approximations of the volumes of three compartments of the head. We applied the method to six species of rock-dwelling haplochromine cichlids, of which we studied 12 populations from four rocky islands in the southern part of Lake Victoria. Head shape was correlated with eight environmental variables. Truss distances and compartment volumes correlated with these variables, e.g. volumes of the compartments containing the gills correlated (negatively) with oxygen levels; truss distances, including the oral jaws and their musculature, correlated with food composition; and eye size correlated with width of the transmission spectrum. Another finding was the likely architectonic interactions between anatomical elements. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 39–48.     

Phenotype and gene flow in a marsupial (Antechinus flavipes) in contrasting habitats

Ecological factors are important drivers of phenotypic divergence, which may lead to incipient speciation. A variety of habitats should be preserved to maintain evolutionary potential. We used the marsupial, the yellow-footed antechinus ( Antechinus flavipes ) as a model species for investigating phenotypic differentiation between animals inhabiting two habitat types in south-eastern Australia: flood-plain river red gum and box–ironbark forests. All tested phenotypic characteristics varied between years at the same sites and therefore were not useful for investigating morphological specialization that may lead to speciation. Males generally were significantly heavier when antechinus densities were lower, but exceptions were found, possibly related to food availability. Teat-number variation recently has been shown to be associated with habitat specialization and incipient speciation within Antechinus agilis . We investigated genetic differentiation associated with this trait in A. flavipes . Population genetic analyses of microsatellite genotypes and mitochondrial DNA haplotypes revealed that sympatric 12-, 13- and 14-teat females in Chiltern forest were part of one freely interbreeding population. Our parentage analyses found two cases where 13-teat mothers produced 12-teat daughters. This suggests either plasticity or paternal genetic influence on the offspring's teat-number phenotype. Laboratory matings may be required to resolve the extent to which teat number is heritable in A. flavipes .  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 303–314.     

Incipient speciation driven by phenotypic plasticity? Evidence from sympatric populations of Arctic charr

Recent models suggest that the existence of environmentally induced polymorphisms within a single population (especially those related to foraging) facilitates the process of evolutionary divergence within a single gene pool by generating distinct phenotypic modes that are exposed to differential selection. In order to test a prediction of the phenotypic plasticity model of divergence, we used a well-documented polymorphism to disentangle the relative effects of morph and rearing environment in generating phenotypic variance. We reared first-generation offspring of two sympatric morphs of Arctic charr Salvelinus alpinus in the laboratory and compared their head morphology with that of their wild parents. Morphological characters with a known functional role in foraging were highly plastic. Rearing environment accounted for the largest component of the variation in expressed phenotype, but this environmental effect overlaid a clear (but small) genetic effect. We conclude that phenotypic plasticity has played a significant role in the evolution of this trophic polymorphism, but that the evolutionary process has progressed to the point that the gene pool is now segregated.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 611–618.     

Habitat-specific differences in thermal plasticity in natural populations of a soil arthropod

When populations experience substantial variation in environmental conditions, they may evolve phenotypic plasticity in response to these varying selection pressures. Evolutionary theory predicts differentiation in the level of phenotypic plasticity among different habitats. We evaluated temperature-induced phenotypic responses in juvenile growth rate in natural populations of the springtail Orchesella cincta , inhabiting forest and heathland. These habitats typically co-occur but differ strongly with respect to, for example, thermal regime, relative humidity, and structure. Offspring of females from the two habitats were reared at different temperatures in climate rooms and the temperature response of juvenile growth rate and egg size was measured. We found a habitat-specific difference in plasticity of juvenile growth rate. The reaction norms of the forest populations were steeper than the reaction norms for heath populations at two replicated sampling sites. Egg weight itself was demonstrated to be a plastic trait with a higher egg weight at low temperatures, but the thermal response did not differ between habitats. We conclude that these populations have diverged due to strong local natural selection. Our results support the argument that the level of phenotypic plasticity itself can be under selection and that differentiation in reaction norms can occur even in neighbouring habitats with no barrier to gene flow.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 265–271.     

Gut length plasticity in perch: into the bowels of resource polymorphisms

Resource polymorphisms, intraspecific variation in morphology due to differential resource use, are common across a wide range of animal taxa. The focus in studies of such polymorphisms has been on external morphology, but the differential use of food resources could also influence other phenotypic traits such as the digestive performance. In the present study, we experimentally demonstrate that Eurasian perch ( Perca fluviatilis L.) display adaptive plasticity in gut length when exposed to different food types. Perch fed a less digestible food type developed relatively longer guts compared to fish fed a more easily digested food type. This divergence in gut length was also apparent under natural conditions because perch inhabiting the littoral and pelagic habitats of a lake differed in resource use and relative gut length. Despite that the digestive system in perch is plastic, we found that individuals switching to a novel food type might experience an initial fitness cost of the diet switch in the form of a temporary reduction in body condition. These results show the importance of gut length plasticity for an ontogenetic omnivore but also a cost that might prevent diet switching in polymorphic populations.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 517–523.     

Local adaptation in four Iris species tested in a common-garden experiment

Local adaptation is a commonly observed result of natural selection acting in heterogeneous environment. Common-garden experiments are a method of detecting local adaptation, as well as studying phenotypic plasticity and gradients of traits. The present study aimed to analyse reaction norms of four closely-related Iris species of section Oncocyclus and to identify a role of environmentally-specific natural selection in their plastic responses. The plant vegetative and phenological, as well as performance traits were measured in a full factorial common-garden experiment with three levels of water amount and three soil types. We found a significant effect of species identity on all traits measured. Water amount and soil type affected many of the traits, but soil type did not affect the performance. There was no significant difference in the effect of water amount and soil type on performance as reflected by rhizome growth; in other words, there was no significant genotype × environment interaction for performance. Plasticity levels and directions of response were also similar among the species. We conclude that phenotypic differences among species are of genetic origin, although no adaptive value was demonstrated for them at the time and life-stages 'frame' of this experiment.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 98 , 267–277.     

Resistance to temperature extremes in sub-Antarctic weevils: interspecific variation, population differentiation and acclimation

Much of the work on the responses of terrestrial arthropods to high and low temperatures has been done on model organisms such as Drosophila . However, considerable variation in thermotolerance is partitioned at the family level and above, raising questions about the broader applicability of this work to other taxa. Here we investigate resistance to high and low temperatures, following different temperature treatments, in ten species and 31 populations of weevils found on sub-Antarctic Heard Island and Marion Island, which have substantially different climates. In these weevils there is considerable interspecific and among-population variation in critical thermal minimum (CTmin) and critical thermal maximum (CTmax), but most of this variation in critical limits can be ascribed to phenotypic plasticity. We find no relationship between CTmin and CTmax at the species level, and this is true also of populations and of responses to the temperature treatments. In general, plastic (acclimation) changes in CTmin are larger than those in CTmax. Our data therefore provide support for the idea that resistance to heat and to cold are decoupled in terrestrial arthropods. Furthermore, our results suggest that investigations of physiological limits to species borders should incorporate the effects of phenotypic plasticity on physiological capabilities.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 401–414.     

Parasite-mediated sexual selection and species divergence in Lake Victoria cichlid fish

We investigate the role of parasite-mediated sexual selection in the divergence of two species of Lake Victoria cichlids. Pundamilia pundamilia and Pundamilia nyererei represent a common pattern of male nuptial colour divergence between haplochromine sister species: metallic grey–blue in P. pundamilia and bright yellow and red in P. nyererei . Female mating preferences for different male colours maintain the genetic and phenotypic differentiation of the two species in clear water. Previous work indicated that the red coloration of P. nyererei males, which is subject to directional sexual selection, may be a carotenoid-dependent signal of parasite infestation rate. In the present study, we find a parallel result for P. pundamilia : bright blue males are infected with fewer species of parasites. We also find that parasite infestation rates differ quantitatively between the two species in a way that is consistent with species differences in diet and microhabitat. We conclude that parasite-mediated sexual selection may have contributed to the divergence of female mating preferences between P. pundamilia and P. nyererei , and may currently strengthen reproductive isolation between these species.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94 , 53–60.     

Plasticity, its cost, and phenotypic selection under water and nutrient stress in two annual grasses

A comparative approach can prove to be a useful tool for studying phenotypic plasticity, if applied to specific traits involved in adaptation to particular environment in more than one species across co-located populations. The present study tested whether two annual grasses, Hordeum spontaneum and Avena sterilis , belonging to the same guild, having similar stature, seed dispersal mechanism, breeding system, and genetic variation, and sampled in exactly the same environmentally specific locations, differed with respect to: (1) plasticity in traits involved in adaptation, namely the onset of reproduction and maternal investment involving the number of inflorescences, spikelets per inflorescence, the weight of individual spikelets, and abortion rate; (2) the cost of this plasticity, and (3) the pattern of phenotypic selection on the above traits. The two species exhibited highly differing amounts of phenotypic plasticity in the onset of flowering and several reproductive traits (number of inflorescences, spikelets per inflorescence, abortion rate), but no plasticity costs in any experimental environment. The two species demonstrated a decreasing similarity in the regulation of reproduction in four experimental environments: benign, water, nutrients and water × nutrient deficient. Correlational selection appears to contribute, although not solely, to the observed species differences with respect to the regulation of reproduction.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 581–593.     

Plastic resource polymorphism: effects of resource availability on Arctic char (Salvelinus alpinus) morphology

Resource polymorphism has been suggested to be a platform for speciation. In some cases resource polymorphism depends on phenotypic plasticity but in other cases on genetic differences between morphotypes, which in turn has been suggested to be the ongoing development of a species pair. Here we study environmentally induced morphological differences in two age classes of Arctic char ( Salvelinus alpinus ) influencing char performance and diet in relation to resource availability. We found that structurally complex habitats with relatively lower zooplankton densities gave rise to individuals with a deeper body, and a downward positioned tip of the snout compared with individuals from structurally simple habitats with relatively higher zooplankton densities for both age classes. Environment also had an effect on foraging efficiency on zooplankton, with fish from structurally simple habitats had a higher foraging rate than fish from structurally complex habitats. Diet analyses showed that resource use in char mainly depends on the relative abundance of different resources. Therefore, to gain further understanding of resource polymorphism we suggest that future studies must include population dynamic feedbacks by the resources on the consumers.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 341–351.     

Performance benefits of growth-form plasticity in a clonal red seaweed

Phenotypic plasticity may be adaptive if the phenotype expressed in a focal environment performs better there relative to alternative phenotypes. Plasticity in morphology may particularly benefit modular organisms that must tolerate environmental change with limited mobility, yet this hypothesis has rarely been evaluated for the modular inhabitants of subtidal marine environments. We test the hypothesis for Asparagopsis armata , a clonal red seaweed whose growth-form plasticity across light environments is consistent with the concept of foraging behaviour in clonal plants. We manipulated the light intensity to obtain clonal replicates of compact, densely branched ('phalanx') phenotypes and elongate, sparsely branched ('guerrilla') phenotypes, which we reciprocally transplanted between inductive light environments to explore the performance consequences of a poor phenotype–environment match. Consistent with the hypothesis of adaptive plasticity, we found that performance (as relative growth rate) depended significantly on the interaction between growth form and environment. Each growth form performed better in its inductive environment than the alternative form, implying that this type of plasticity, thought to be adaptive for clonal plants, may also benefit photoautotrophs in marine environments. Given the prevalence and diversity of modular phyla in such systems, they offer a relatively unexplored opportunity to broaden our understanding of the evolutionary ecology of phenotypic plasticity.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 80–89.     

Effects of genetics and light environment on colour expression in threespine sticklebacks

The genetic basis of traits that are under sexual selection and that are involved in recognizing conspecific mates is poorly known, even in systems in which the phenotypic basis of these traits has been well studied. In the present study, we investigate genetic and environmental influences on nuptial colour, which plays important roles in sexual selection and sexual isolation in species pairs of limnetic and benthic threespine sticklebacks ( Gasterosteus aculeatus species complex). Previous work demonstrated that colour differences among species correlate to differences in the ambient light prevalent in their mating habitat. Red fish are found in clear water and black fish in red-shifted habitats. We used a paternal half-sib split-clutch design to investigate the genetic and environmental basis of nuptial colour. We found genetic differences between a red and a black stickleback population in the expression of both red and black nuptial colour. In addition, the light environment influenced colour expression, and genotype by environment interactions were also present. We found evidence for both phenotypic and genetic correlations between our colour traits; some of these correlations are in opposite directions for our red and black populations. These results suggest that both genetic change and phenotypic plasticity underlie the correlation of male colour with light environment.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 663–673.     

Genetic and environmental components of phenotypic variation in body shape among populations of Atlantic cod (Gadus morhua L.)

A common-garden experiment was conducted on larvae to test for genetic differences in body shape among populations of Atlantic cod ( Gadus morhua ). Offspring from four north-west Atlantic regions were reared from hatching to postmetamorphosis at two temperatures (7 ± 1 °C and 11 ± 1 °C) and two food levels (1500 and 4500 prey L⁻¹). Body shape differed between populations and treatments. Population differences were greatest between south-west Scotian Shelf cod and those further north; the former were characterized by a deeper body, larger head, and longer caudal peduncle than cod from the other populations. Significant differences were also observed between two putative populations on the south-west Scotian Shelf, suggesting genetic divergence between spawning aggregations at small spatial scales (< 100 km). Temperature and food supply also influenced body shape, with the effect of the former being more pronounced. Individuals reared at the higher temperature or food level had a deeper body and a larger head than those reared at the lower temperature or food supply. Phenotypic responses to changes in the rearing environment also differed among populations, indicating genetic differences in phenotypic plasticity. Differences between populations in morphology and in phenotypic plasticity suggest genetic divergence at both large (> 1000 km) and small (< 100 km) spatial scales. The genetic differences at large spatial scales counteracted the expected effects of temperature differences in the wild, suggesting countergradient variation in morphology among these populations.  © 2006 Her Majesty the Queen in Right of Canada. Journal compilation © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 351–365.     

Phylogeography and morphological variability in land snails: the Sicilian Marmorana (Pulmonata, Helicidae)

Land snails have long been recognized as suitable organisms for studying phenotypic differentiation and phylogeny in relation to geographical distribution. Morphological data (shell and anatomy biometry on different geographical scales) and partial sequences from mitochondrial genes (cytochrome oxidase subunit I , 16S rDNA) were used to test whether morphological patterns match phylogeny in a diversified group of Sicilian rock-dwelling land snails belonging to the genus Marmorana . The taxonomic implications of the three character sets (shell and anatomical biometry and molecular data) were also considered. The inferred phylogenetic relationships do not match morphological (shell and genitalia) patterns. This result may significantly modify the current taxonomy. Mitochondrial based reconstructions define several supported clades well correlated with geographic distribution and populations were found to be distributed parapatrically. The progressive decline in mitochondrial DNA sequence similarity over a distance of 250 km is consistent with a model of isolation by distance, a pattern previously recognized for other groups of land snails. For one clade of Marmorana , colonization along Mediterranean trade routes appears to be a possibility.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 809–823.     

Two new late Carboniferous Neuropteris species (Medullosales) from Saarland, Germany and their palaeobiogeographical significance

Two new species ( Neuropteris terminiscus and N. kneuperi ) are described from the Bolsovian ('Westphalian C') of Saarland. N. terminiscus had previously been misidentified as Paripteris pseudograndinioides , but the pinnae are clearly imparipinnate and thus cannot belong to that morphogenus. N. kneuperi had previously been misidentified as Neuropteris obliqua and N. heterophylla , both which appear to be absent from Saar-Lorraine. This is considered further evidence of the distinctive nature of the Westphalian vegetation of the Saar-Lorraine Basin, especially among the seed-plants.  © 2002 The Linnean Society of London , Botanical Journal of the Linnean Society , 2002, 139 , 193–205.     

DOES PHENOTYPIC PLASTICITY FOR ADULT SIZE VERSUS FOOD LEVEL IN DROSOPHILA MELANOGASTER EVOLVE IN RESPONSE TO ADAPTATION TO DIFFERENT REARING DENSITIES?

Recent studies with Drosophila have suggested that there is extensive genetic variability for phenotypic plasticity of body size versus food level. If true, we expect that the outcome of evolution at very different food levels should yield genotypes whose adult size show different patterns of phenotypic plasticity. We have tested this prediction with six independent populations of Drosophila melanogaster kept at extreme densities for 125 generations. We found that the phenotypic plasticity of body size versus food level is not affected by selection or the presence of competitors of a different genotype. However, we document increasing among population variation in phenotypic plasticity due to random genetic drift. Several reasons are explored to explain these results including the possibility that the use of highly inbred lines to make inferences about the evolution of genetically variable populations may be misleading.