Sinistral snail shells in the sea: developmental causes and consequences

The paucity of sinistral (left-coiling) relative to dextral (right-coiling) species of gastropods in the marine realm is an enigma. In Conus , one of the most diverse marine animal genera, sinistral shell coiling has evolved as a species-wide character only once. Fossils of this species, Conus adversarius , are found in Upper Pliocene and lowermost Pleistocene deposits in the southeastern USA. Conus adversarius had nonplanktonic larval development; this may have been a critical factor for the early establishment of the species, as well as sinistral marine species in other clades. Notably, most specimens of aberrantly sinistral modern Conus are derived from typically dextral species that have nonplanktonic development. If C. adversarius was reproductively isolated from dextral conspecifics, then this species may provide an example of nearly instantaneous sympatric speciation in the fossil record. Furthermore, the common and widespread – while geologically short-lived – fossil shells of C. adversarius show large amounts of variability in form and this variation may be related, at least in part, to a pleiotropic effect associated with the reversed coiling direction of this species.     

Chiral speciation in terrestrial pulmonate snails

On the basis of data in the literature, the percentages of dextral versus sinistral species of snails have been calculated for western Europe, Turkey, North America (north of Mexico), and Japan. When the family of Clausiliidae is represented, about a quarter of all snail species may be sinistral, whereas less than one per cent of the species may be sinistral where that family does not occur. The number of single-gene speciation events on the basis of chirality, resulting in the origin of mirror image species, is not closely linked to the percentage of sinistral versus dextral species in a particular region. Turkey is nevertheless exceptional by both a high percentage of sinistral species and a high number of speciation events resulting in mirror image species. Shell morphology and genetic background may influence the ease of chirality-linked speciation, whereas sinistrality may additionally be selected against by internal selection. For the Clausiliidae, the fossil record and the recent fauna suggest that successful reversals in coiling direction occurred with a frequency of once every three to four million years.     

Reproductive character displacement by inversion of coiling in clausiliid snails (Gastropoda, Pulmonata)

In land snails, a change in the direction of coiling, being associated with a shift in the position of the genital apparatus, may act as a barrier against hybridization between sympatric species. Putative reproductive character displacement by an inversion in chirality has been reported in only a few land snails, based on observations in the field and interbreeding experiments. In this study, we present a new case of possible reproductive character displacement in the direction of coiling, in the clausiliid snail Isabellaria dextrorsa . This species is dextral, in contrast with its nearest relatives, including I. torifera and I. lophauchena , which share plesiomorphic sinistral coiling. Whereas I. dextrorsa occurs in sympatry and even syntopically with I. lophauchena throughout most of its range, the sinistral species have a mosaic distribution. Phylogenetic analyses of mitochondrial cytochrome c oxidase subunit I (COI) sequences demonstrated that I. dextrorsa constitutes a clade with I. torifera . In this clade, a shift in coiling direction occurred at least twice, maybe triggered by the presence of a sympatric congeneric sinistral species. The analyses separated the sequences of all I. dextrorsa samples from those of sympatric and syntopic I. lophauchena samples. The failure to demonstrate gene flow between these species is consistent with the hypothesis of genetic isolation by reproductive character displacement.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 155–164.     

Whole-body enantiomorphy and maternal inheritance of chiral reversal in the pond snail Lymnaea stagnalis

Sinistral and dextral snails have repeatedly evolved by left-right reversal of bilateral asymmetry as well as coiling direction. However, in most snail species, populations are fixed for either enantiomorph and laboratory breeding is difficult even if chiral variants are found. Thus, only few experimental models of chiral variation within species have been available to study the evolution of the primary asymmetry. We have established laboratory lines of enantiomorphs of the pond snail Lymnaea stagnalis starting from a wild population. Crossing experiments demonstrated that the primary asymmetry of L. stagnalis is determined by the maternal genotype at a single nuclear locus where the dextral allele is dominant to the sinistral allele. Field surveys revealed that the sinistral allele has persisted for at least 10 years, that is, about 10 generations. The frequency of the sinistral allele showed large fluctuations, reaching as frequent as 0.156 in estimate under the assumption of Hardy-Weinberg equilibrium. The frequency shifts suggest that selection against chiral reversal was not strong enough to counterbalance genetic drift in an ephemeral small pond. Because of the advantages as a model animal, enantiomorphs of L. stagnalis can be a unique system to study aspects of chirality in diverse biological disciplines.     

Coiling dimorphism within a genetic type of the planktonic foraminifer Globorotalia truncatulinoides

The coiling direction—sinistral (left-coiled) or dextral (right-coiled)—of planktonic foraminiferal shells is a classical proxy used to assess past environmental changes and to understand their evolutionary patterns. Globorotalia truncatulinoides is composed of five different genetic types (I to V), each with a specific biogeographic distribution. So far, type II is the only type within which both coiling types have been frequently found. Here we examine the coiling direction as a dimorphic variation in a single genetic type, and show that the two forms with different coiling directions differ in ecology. The studied 140 left- and 137 right-coiled specimens from eight depth layers at four stations in the Sargasso Sea all belonged to Type II, based on the phylogenetic analyses of the ITS (internal transcribed spacer) region of ribosomal DNA. The distributions of left- and right-coiled Type II (Type II-L and II-R) differed strongly: Type II-L dominated between 400 and 200 m depth in the central water mass, whereas Type II-R was common in the top 200 m of the subtropical gyre. The left- and right-coiled forms are thus associated with different water masses. However, the vertical distribution along a temperature gradient indicates that their coiling direction is not determined by temperature, and might have some genetic basis. Our molecular phylogenetic analyses showed that the two forms probably share the same gene pool through their dispersals between different water masses. Moreover, the lineage of G. truncatulinoides is young (< 3 Ma), which might not be long enough to evolve into two lineages with opposite coiling direction. We discuss the practical and theoretical usefulness of a simple genetic model for coil dimorphism, in order to build testable hypotheses regarding the evolutionary history of planktonic foraminifera at the population level.     

THE CONSEQUENCES OF BEING DIFFERENT: SINISTRAL COILING IN CERION

The overwhelming predominance of dextral coiling in gastropods is an outstanding and puzzling phenomenon. A few sinistral specimens (left coiling individuals) have been found in many dextral species. Only six sinistral shells have ever been found in Cerion; we base this analysis on the five available shells. We ask whether reversed symmetry is a simple either-or switch without further consequences for shell form, or whether sinistrality engenders associated effects, making left-coiling shells unlike their dextral deme-mates in other ways. All five sinistral shells differ in features of size and coiling late in growth, leading to relatively small apertures and a slight twist in the axis of coiling. We detect and measure this effect as follows: in multivariate morphospace, sinistrals occupy peripheral positions among their dextral deme-mates; in univariate analysis, sinistrals are consistently different for a set of characters involving covariance patterns never before seen in a decade of studies on ontogenetic and age-standardized variation in dextrals; a bootstrap procedure does not recover similar patterns in randomly constituted samples of dextrals matching the true sinistral distribution; direct x-ray measures of the coiling axis detect its slight twist in sinistrals. We discuss the implications of these unsuspected associations for the issues of developmental constraint upon the evolution of morphology.     

Silk Weave and Silk Glands in Aquatic Instars of Two Species of Helicopsyche von Siebold, 1856 (Trichoptera,Helicopsychidae)

Under SEM the silk weave in the snail-like cases of Helicopsyche crispata and H. shuttleworthi, the two species present in Italy consist of several types of meshes. The silk which connects the sand grains of the external wall is made up of multi-layered threads forming irregular meshes. The sand grains of the vertical pillars in the wall of the centr al columella are held together by very loosely woven silk and are supported by thick silken threads. The pupal silken membrane consists of concentric threads and the pupal case is attached to the substrate by a disordered mass of silken threads. The two glands which secrete the silk are long, double folded tubes.     

Multiple reversals of chirality in the land snail genus Albinaria (Gastropoda,Clausiliidae)

Reversed chirality has frequently evolved in snails, although the vast majority coils dextrally. However, there are often sinistral species within a dextral genus or almost exclusively sinistral families, such as the Clausiliidae. Some populations of the predominantly sinistral clausiliid genus Albinaria, in the southern Greek mainland, coil dextrally. The origin, evolution and distribution of the dextral Albinaria are puzzling, and as there is no reliable phylogenetic reconstruction for this speciose genus, it remains unclear how many times a shift in chirality has really occurred. In this study, our aim was to elucidate the evolutionary pathways of dextrality in Albinaria. We undertook a molecular phylogenetic analysis of two mtDNA (16S and COI) and one nDNA marker (ITS1) and included dextral and sinistral representatives found in syntopy or not. Both mtDNA and nDNA tree topologies imply that dextrals did not evolve as a monophyletic lineage. Instead, dextral lineages have evolved from sinistral ancestors multiple times independently. The fragmented population structure in Albinaria facilitates genetic drift and contributes to fixation of the opposite chirality and overcoming of the mating disadvantage of left–right reversal. Stochastic phenomena and biogeographical barriers have trapped those reversals in a limited geographical area.     

Dextral and sinistral <Emphasis Type="Italic">Amphidromus inversus</Emphasis> (Gastropoda: Pulmonata: Camaenidae) produce dextral sperm

Coiling direction in pulmonate gastropods is determined by a single gene via a maternal effect, which causes cytoskeletal dynamics in the early embryo of dextral gastropods to be the mirror image of the same in sinistral ones. We note that pulmonate gastropod spermatids also go through a helical twisting during their maturation. Moreover, we suspect that the coiling direction of the helical elements of the spermatozoa may affect their behaviour in the female reproductive tract, giving rise to the possibility that sperm chirality plays a role in the maintenance of whole-body chiral dimorphism in the tropical arboreal gastropod Amphidromus inversus (Müller, 1774). For these reasons, we investigated whether there is a relationship between a gastropod’s body chirality and the chirality of the spermatozoa it produces. We found that spermatozoa in A. inversus are always dextrally coiled, regardless of the coiling direction of the animal itself. However, a partial review of the literature on sperm morphology in the Pulmonata revealed that chiral dimorphism does exist in certain species, apparently without any relationship with the coiling direction of the body. Though our study shows that body and sperm chirality follows independent developmental pathways, it gives rise to several questions that may be relevant to the understanding of the chirality of spermatid ultrastructure and spermatozoan motility and sexual selection.     

Phylogenetic reconstruction and shell evolution of the Diplommatinidae (Gastropoda: Caenogastropoda)

The fascinating and often unlikely shell shapes in the terrestrial micromollusc family Diplommatinidae (Gastropoda: Caenogastropoda) provide a particularly attractive set of multiple morphological traits to investigate evolutionary patterns of shape variation. Here, a molecular phylogenetic reconstruction, based on five genes and 2700 bp, was undertaken for this family, integrated with ancestral state reconstruction and phylogenetic PCA of discrete and quantitative traits, respectively. We found strong support for the Diplommatininae as a monophyletic group, separating the Cochlostomatidae into a separate family. Five main clades appear within the Diplommatininae, corresponding with both coiling direction and biogeographic patterns. A Belau clade (A) with highly diverse (but always sinistral) morphology comprised Hungerfordia, Palaina, and some Diplommatina. Arinia (dextral) and Opisthostoma (sinistroid) are sister groups in clade B. Clade C and D solely contain sinistral Diplommatina that are robust and little ornamented (clade C) or slender and sculptured (clade D). Clade E is dextral but biogeographically diverse with species from all sampled regions save the Caroline Islands. Adelopoma, Diplommatina, Palaina, and Hungerfordia require revision to allow taxonomy to reflect phylogeny, whereas Opisthostoma is clearly monophyletic. Ancestral state reconstruction suggests a sinistral origin for the Diplommatinidae, with three reversals to dextrality.     

新疆晚奥陶世晚期管状体及其古植物学物意义

本文系统描述了产自我国新疆塔里木盆地晚奥陶世晚期的管状体,包括2属8种,其中一个新种。通过对比研究,本管状体组合与Burgess和Edwards(1991）的下组合相当。根据本组合的研究,对Burgess和Edwards的组合进行了补充,其组合由不同类型的光壁管、低分异度的内壁加厚管组成、未见外壁加厚管和丝状体,组合的时代从中奥陶世到早志留世（晚Llandovery)。根据植物学的对比研究,管状体主要产自Nematophytales Lang.     

Sex chromatin and sex chromosome systems in nonditrysian Lepidoptera (Insecta)

Eleven representatives of the superorder Amphiesmenoptera (Trichoptera + Lepidoptera) were examined for sex chromatin status. Three species represent stenopsychoid, limnephiloid and leptoceroid branches of the Trichoptera; eight species belong to the primitive, so-called nonditrysian Lepidoptera and represent the infra-orders Zeugloptera, Dacnonypha, Exoporia, Incurvariina, Nepticulina and Tischeriina. The female-specific sex chromatin body was found in the interphase somatic nuclei of Tischeria ekebladella (Bjerkander 1795) (Lepidoptera, Tischeriina). The sex chromatin was absent in all investigated Trichoptera species as well as in all representatives of the nonditrysian Lepidoptera except Tischeria ekebladella . The sex chromosome mechanism of Limnephilus lunatus Curtis 1834 (Trichoptera, Limnephilidae) is Z/ZZ. The sex chromosome mechanism of Tischeria ekebladella (Lepidoptera, Tischeriina) is ZW/ZZ including the W chromosome as the largest element in the chromosome set. The data obtained support the hypothesis that the Z/ZZ sex chromosome system, the female heterogamety and the absence of the sex chromatin body in interphase nuclei are ancestral traits in the superorder Amphiesmenoptera. These ancestral characters are probably kept constant in all the Trichoptera and in the most primitive Lepidoptera. The W sex chromosome and the sex chromatin evolved later in the nonditrysian grade of the Lepidoptera. It is proposed that the sex chromatin is a synapomorphy of Tischeriina and Ditrysia.     

Molecular phylogeny of the land snail genus Alopia (Gastropoda: Clausiliidae) reveals multiple inversions of chirality

Whereas the vast majority of gastropods possess dextral shell and body organization, members of the Clausiliidae family are almost exclusively sinistral. Within this group a unique feature of the alpine genus Alopia is the comparable representation of sinistral and dextral taxa, and the existence of enantiomorph taxon pairs that appear to differ only in their chirality. We carried out a molecular phylogenetic study, using mitochondrial cytochrome c oxidase subunit I (COI) gene sequences, in order to find out whether chiral inversions are more frequent in this genus than in other genera of land snails. Our results revealed multiple independent inversions in the evolutionary history of Alopia and a close genetic relationship between members of the enantiomorph pairs. The inferred COI phylogeny also provided valuable clues for the taxonomic division and zoogeographical evaluation of Alopia species. The high number of inverse forms indicates unstable fixation of the coiling direction. This deficiency and the availability of enantiomorph pairs may make Alopia species attractive experimental models for genetic studies aimed at elucidating the molecular basis of chiral stability. © 2013 The Linnean Society of London     

Shell chirality in Cambrian gastropods and sinistral members of the genus <Emphasis Type="Italic">Aldanella</Emphasis> Vostokova, 1962

Shell chirality among Cambrian gastropods is discussed. It is demonstrated that the earliest members of the class include chiral aberrations with abnormal opposite coiling of the shell. It is assumed that, in Cambrian gastropods, speciation could have occurred by mutation in the locus determining the chirality, as is proposed for extant gastropods. In contrast to modern gastropods, the existence of chiral morphs within single species has not been recorded in Cambrian mollusks, whereas the presence of chiral twin species is possible. The systematic position of sinistral representatives of the genus Aldanella Vostokova, 1962 is considered. Aldanella golubevi sp. nov. with sinistral shell is described from the base of the Tommotian Stage of the Anabar Region. Aberrant sinistral specimens of the normally dextral species Aldanella utchurica Missarzhevsky in Rozanov et al., 1969 and Pelagiella adunca Missarzhevsky in Rozanov et al., 1969 are figured.     

Morphological evidence of correlational selection and ecological segregation between dextral and sinistral forms in a polymorphic flatfish, Platichthys stellatus

Phenotypic polymorphisms in natural systems are often maintained by ecological selection, but only if niche segregation between morphs exists. Polymorphism for eyed-side direction is rare among the approximately 700 species of flatfish (Pleuronectiformes), and the evolutionary mechanisms that maintain it are unknown. Platichthys stellatus (starry flounder) is a polymorphic pleuronectid flatfish exhibiting large, clinal variation in proportion of left-eyed (sinistral) morphs, from 50% in California to 100% in Japan. Here I examined multiple traits related to swimming and foraging performance between sinistral and dextral morphs of P. stellatus from 12 sites to investigate if the two morphs differ in ways that may affect function and ecology. Direction of body asymmetry was correlated with several other characters: on an average, dextral morphs had longer, wider caudal peduncles, shorter snouts and fewer gill rakers than sinistral morphs. Although the differences were small in magnitude, they were consistent in direction across samples, implying that dextral and sinistral starry flounder may be targeting different prey types. Morphological differences between morphs were greatest in samples where the chances of competitive interactions between them were the greatest. These results suggest that the two morphs are not ecologically identical, may represent a rare example of divergent selection maintaining polymorphism of asymmetric forms, and that correlational selection between body asymmetry and other characters may be driven by competitive interactions between sinistral and dextral flatfish. This study is one of very few that demonstrates the ecological significance of direction in a species with polymorphic asymmetric forms.     

Evolution of a dextral lineage by left–right reversal in Cristataria (Gastropoda,Pulmonata, Clausiliidae)

It has long been debated whether mirror image‐like similarity in shell morphology between enantiomorphic pairs of dextral and sinistral taxa represents their sister relationship, or each of them is closer related to other congeners with the same coiling direction. The obligate rock‐dwelling genus Cristataria Vest, 1867 of the eastern Mediterranean region belongs to the Alopiinae subfamily of door snails (Clausiliidae). Cristataria and a few other genera of this subfamily include enantiomorphic pairs that are conchologically very similar to each other. Dextral C. colbeauiana (Pfeiffer, 1861) and its sinistral counterpart of such an enantiomorphic pair occur nearby one another in southern Turkey. However, the latter has been classified either as the sinistral subspecies C. colbeauiana inversa Szekeres, 1998 or as a form of sinistral C. leprevieri (Pallary, 1922). To examine the phylogenetic relationship of this enantiomorphic pair, we carried out molecular phylogenetic analysis of all the Turkish and two other Cristataria taxa based on both mitochondrial and nuclear DNA markers. Our results show that dextral C. colbeauiana and its sinistral counterpart are closest related to one another. This supports the classification of this enantiomorphic pair as dextral C. colbeauiana colbeauiana and sinistral C. colbeauiana inversa. Our results also reveal that these taxa and C. intersita Németh & Szekeres, 1995, sharing a characteristic collar behind the aperture of the shell, represent a monophyletic lineage. By contrast, the Cristataria species of non‐collared shells belong to another clade.     

High gene flow despite opposite chirality in hybrid zones between enantiomorphic door snails

We studied differentiation and geneflow patterns between enantiomorphic door‐snail species in two hybrid zones in the Bucegi Mountains (Romania) to investigate the effects of intrinsic barriers (complications in copulation) and extrinsic selection by environmental factors. A mitochondrial gene tree confirmed the historical separation of the examined populations into the dextral Alopia livida and the sinistral Alopia straminicollis in accordance with the morphological classification, but also indicated gene flow between the species. By contrast, a network based on amplified fragment length polymorphisms (AFLP) markers revealed local groups of populations as units independent of their species affiliation. Admixture analyses based on AFLP data showed that the genomes of most individuals in the hybrid zones are composed of parts of the genomes of both parental taxa. The introgression patterns of a notable fraction of the examined markers deviated from neutral introgression. However, the patterns of most non‐neutral markers were not concordant between the two hybrid zones. There was also no concordance between non‐neutral markers in the two genomic clines and markers that were correlated with environmental variables or markers that were correlated with the proportion of dextral individuals in the populations. Neither extrinsic selection by environmental factors nor intrinsic barriers resulting from positive frequency‐dependent selection of the prevailing coiling direction were sufficient to maintain the distinctness of A. straminicollis and A. livida. Despite being historically separated units, we conclude that these taxa now merge where they come into contact.     

Momentary maximizing and optimal foraging theories of performance on concurrent VR schedules

Optimal foraging theory proposes that animals obtain the highest rate of reinforcers for the least effort and momentary maximizing theory proposes that animals make the response that at that instant is most likely to be reinforced. While each theory may account for matching on concurrent schedules, the data supporting each theory are weak. Two experiments assessed these theories by considering concurrent choice as consisting of two pairs of stay and switch schedules. Symmetrical arrangements, which are equivalent to standard concurrent schedules, maintained behavior described by the generalized matching law. Weighted arrangements, in which the programmed rate of earning reinforcers was always greater at one alternative, maintained behavior that was biased towards the weighted alternative, yet the bias was less than that predicted by optimal foraging theory. Asymmetrical arrangements, in which the stay and switch schedules operating at an alternative are the same, maintained behavior that favored one alternative, even though momentary maximizing predicted indifference. The generalized matching law poorly described each rat's pooled data from all conditions but these data were described by an equation based on the stay and switch reinforcers earned per-visit and included elements of optimal foraging and momentary maximizing theories of choice.     

Drosophila tracheal morphogenesis: intricate cellular solutions to basic plumbing problems

The cellular architecture of tubular organs suggests striking similarities in the mechanisms of tubulogenesis between species. The formation of the Drosophila respiratory organ (trachea) highlights the basic principles of branch patterning and tube growth that generate a highly elaborate but stereotyped epithelial tubular network. Oriented cell migration, changes in cell shape, selective growth of the apical cell membrane and intracellular lumen formation are essential events in this process. These morphogenetic processes build four structurally distinct classes of tubes that facilitate optimal airflow and gas exchange with target tissues. The molecular players in these plots include attractant and repellent signals, differentiation factors that cause a high diversity of cell fates within the epithelium, and determinants of tube formation and dimensions.     

Female genital configuration in the classification of Lepidoptera

In Coleoptera, Neuroptera, and Megaloptera, and the panorpoid orders Diptera, Trichoptera, and Mecoptera the common oviduct is ventral to, or opens into, the vagina or genital chamber. In Lepidoptera, in the superfamilies Micropterigoidea, Eriocranioidea, Incurvarioidea, and Nepticuloidea, the common oviduct enters the copulatory chamber ventrally; in Mnesarchaeidae, Hepialoidea, and all Ditrysia auct. the common oviduct is dorsal to the copulatory chamber, and the vagina (that region posterior to the entry of the spermatheca) opens separately from the genital ostium. Lepidoptera are unique in the complex and various arrangements of the ectodermal elements in the female genitalia.

The arrangements of, and connections between rectal and genital structures in representatives of Megaloptera, Neuroptera, and the panorpoid orders are re‐examined for comparison with the systems found in Zeugloptera, Dacnonypha, Monotrysia, and Ditrysia auct.

The Zeugloptera are here included in Lepidoptera because they have a circumcloacal chamber. Other female zeuglopteran genital structures intergrade with those of Dacnonypha, here restricted to Eriocraniidae, Agathiphagidae, Lophocoronidae, and the divergent Acanthopteroctetes. Dacnonypha have a less specialised spermathecal and vaginal structure than do most of the Monotrysia, here restricted to Incurvarioidea and Nepticuloidea (including Tischeriidae); many Monotrysia lack a cloaca, whereas it is always present in Dacnonypha.

There is no basis for retaining either Mnesarchaeidae or Hepialoidea in Monotrysia auct., as the dorsal common oviduct and the two genital openings indicate that these groups are ditrysian. They are here regarded as exoporian Ditrysia, a group characterised by the lack of a free, tubular ductus seminalis. A fixed gutter or channel between ostium and ovipore characterises the Hepialoidea, and the absence of this channel (ostium and ovipore opposable within an external genital pouch) characterises Mnesarchaeidae.

The endoporian Ditrysia all have a free, tubular ductus seminalis; where a cloaca is present it is incomplete, i.e., combines ovipore and rectum but never copulatory structures, in contrast to the complete cloaca found in Zeugloptera, Dacnonypha, and many Monotrysia. The endoporian Ditrysia comprise all other superfamilies, i.e., about 97% of species of Lepidoptera.