Functional morphology and autecology of Pseudoptera (Bakevelliid bivalves,upper cretaceous of Portugal)

Pseudoptera sp. nov. is sinistrally twisted about the hinge axis, i.e., in the opposite direction to that of Hoernesia and the other twisted bakevelliids. Field observations confirm the semi-infaunal pleurothetic life habit predicted from the shell morphology. The twisted Bakevelliidae are unlikely to have evolved from soft-bottom forms. A transition from an epibyssate habit on solid substrates to an endobyssate, semi-infaunal life habit probably triggered the evolution of shell torsion in this family, as in the Arcidae and Mytilidae.     

Life habit and ontogeny of the unusual arcid bivalve Ambrogia mytiloides (Brocchi, 1814)

The unusual Plio-Pleistocene arcid Ambrogia mytiloides (Brocchi, 1814) has a large, elongate, smooth and streamlined shell. On the basis of these characters and the occurrence of moderate shell torsion, the mode of life of this species was formerly thought to be semi-infaunal endobyssate, obliquely oriented like the twisted arcid Trisidos. The discovery of shells in life position suggests that this species lived in a subvertical position. Rather than a recliner, this arcid was then a sticker, whose stability was provided by the byssus, which also was used to aid the bivalve in burrowing, and by its large size. The morphology of juvenile valves, smaller than 4 mm, suggests an epibyssate mode of life in its early growth stages. Ambrogia represents a remarkable pathway in the secondary return of arcids to soft bottoms: with this genus, they reached their deepest burrowing level. However, this strategy was not very successful, probably because of evolutionary constraints on the Arcoida.     

Epibiont habitation patterns and their implications for life habits and orientation among trigoniid bivalves

The bivalve superfamily Trigoniacea has persisted from the Late Paleozoic to the Recent. Late Jurassic and terminal Cretaceous mass extinctions decimated this once-dominant group in shallow marine facies; only a single genus with seven species survives today in the Austral Province. Trigoniacea retain a vestigial byssus and primitive but efficient schizodont dentition. They have been widely considered as infaunal bivalves, burrowing with a very large foot to shallow depths, with inhalant and exhalant apertures at or slightly below the sediment-water interface (SWI). Yet the Trigoniacea are poorly adapted for this life habit. The mantle in living species is unfused and non-siphonate, and some fossil Trigoniacea have permanent shell gapes over these apertures, enhancing the probability of sediment fouling of feeding and respiratory structures. Some living Neotrigonia , e.g., N. margaritacea , solve this problem by having a semi-infaunal life habit, with the inhalant and exhalant apertures elevated above the SWI and the zone of active sediment transport. Semi-infaunal species commonly have epibionts cohabiting the exposed posterior-posteroventral portion of the shell. Numerous well-preserved species of South American Mesozoic Trigoniacea have phototropically and geotropically oriented epibionts on co-attached valves, strongly suggesting a semi-infaunal life mode for at least some members of these taxa. These shell symbionts allow orientation of extinct trigoniid shells relative to the SWI during life, as well as analysis of their depth of burial. Careful analyses of the kinds, size classes, orientation, and dispersion of various epibionts on fossil Trigoniacea thus yield important new information on their life habits, and demonstrate that semi-infaunal life modes were far more common than previously supposed.     

Constructional morphology of strombid gastropods

The extreme diversity in shell shape of strombid gastropods is interpreted as the result of three independent factors: (1) The terminal growth pattern of the Strombidae allows the circumvention of geometric constraints on shell morphology found in gastropods with continuous or periodic growth patterns. (2) Shell morphology in the Strombidac is adaptive in epifaunal locomotion, burrowing. infaunal or semi-infaunal habits, and passive protection from predators. Specialization for one of these functions often conflicted with the others. thus bringing about a forced 'choice' among mutually exclusive morphological characters. (3) Conservatism in life habits and anatomy of the soft parts has allowed the multiple evolution of extreme shell morphologies, as well as the secondary return to relativcly unspecialized morphologies. □ Constructional morphology, functional morphology. growth. behaviour. evolution, locomotion, burrowing, predation, exoskeleton. shell. Mollusca. Gastropoda. Strombacea. Strombidae.     

ADAPTATIONS IN GRAMMYSIA OBLIQUA

The Grammysiidae are an important but little understood family of Paleozoic bivalves. Specimens of Grammysia obliqua (McCoy) from the Stonehouse Formation (Uppermost Silurian), Nova Scotia, permit interpretation of their functional morphology and autecology. Grammysia obliqua was an immobile, semi-infaunal, filter-feeding bivalve. It lived fixed by a byssus, with the shell inclined about 40° to the sediment-water interface. Though incapable of opening the shell, the animal maintained contact with the environment through byssal and dorsal-posterior gapes. This latter gape was in a peculiar position for a bivalve and required some anatomical adaptation, but it was functionally efficient for the animal's life attitude.     

Infaunal or semi-infaunal bellerophont gastropods: analysis of Euphemites and functionally related taxa

Burrowing and plowing gastropods may have existed prior to the Mesozoic, contrary to the conservative tendency of categorizing Paleozoic archaeogastropods as herbivorous hardground dwellers. We propose that Euphemites and other bellerophontiform molluscs such as Praematuratroph and Beyrichidiscus make functional and paleoecological sense only if reconstructed with an internal shell adapted to an infaunal mode of life. Attributes diagnostic of a probable infaunal mode of life in fossil bellerophontiform molluscs include: (1) lack of apertural flare; (2) restricted aperture; (3) low rate of whorl expansion; (4) semi-radial aperture; (5) extensive secondary shell deposits; (6) absence of extensive collabral ornament; (7) dorsolateral or umbilical furrows; (8) color patterns indicative of mantle extension and retraction capabilities; (9) absence of shell breakage and repair; and (10) associated lithologies indicative of penetrable substrates such as mudstones. Niche diversity in the bellerophontacean molluscs was probably much greater than previously suspected. This is entirely consistent with their long geologic range. Most bellerophontaceans, particularly the Late Paleozoic forms, were probably either 'nestling' or semi-infaunal with the apertural margins enwrapped by mantle or mantle and foot. As such they may have been particularly susceptible to predation during the postulated Mesozoic increase in number of infaunal predators- a factor which may have been contributory to the bellerophontacean demise.     

Torsion in Patella caerulea (Mollusca, Patellogastropoda): ontogenetic process, timing, and mechanisms

Abstract. Torsion is a process in gastropod ontogenesis where the visceral body portion rotates 180° relative to the head/foot region. We investigated this process in the limpet Patella caerulea by using light microscopy of living larvae, as well as scanning electron microscopy (SEM) of larvae fixed during the torsion process. The completion of the 180° twist takes considerably less time in larvae of Patella caerulea than previously described for other basal gastropod species. At a rearing temperature of 20–22°C, individuals complete ontogenetic torsion in ?2 h. Furthermore, the whole process is monophasic, i.e., carried out at a constant speed, without any evidence of distinct ‘fast” or ‘slow” phases. Both larval shell muscles—the main and the accessory larval retractor—are already fully contractile before the onset of torsion. During the torsion process both retractors perform cramp‐like contractions at ～30 s intervals, which are followed by hydraulic movements of the foot. However, retraction into the embryonic shell occurs only after torsion is completed. The formation of the larval operculum is entirely in‐dependent from ontogenetic torsion and starts before the onset of rotation, as does the mineralization of the embryonic shell. The reported variability regarding the timing (mono‐ versus biphasic; duration) of torsion in basal gastropod species precludes any attempt to interpret these data phylogenetically. The present findings indicate that the torsion process in Patella caerulea, and probably generally in basal gastropods, is primarily caused by contraction of the larval shell muscles in combination with hydraulic activities. In contrast, the adult shell musculature, which is independently formed after torsion is completed, does not contribute to ontogenetic torsion in any way. Thus, fossil data relying on muscle scars of adult shell muscles alone appear inappropriate to prove torted or untorted conditions in early Paleozoic univalved molluses. Therefore, we argue that paleontological studies dealing with gastropod phylogeny require data other than those based on fossilized attachment sites of adult shell muscles.     

Response of hermit crabs to sinistral shells

Shell rotating behavior of the hermit crabPagurus geminus was investigated. In preliminary observations, hermit crabs motivated to change shells rotated presented shells, filled with sand, in a way that dislodged the inside material. In order to determine if this behavior is stereotyped, or flexible and dependent on shell type, hermit crabs were tested with ordinary dextral shells ofLatirulus nagasakiensis and sinistral shells ofAntiplanes contraria. Sinistral shells are not normally encountered by hermit crabs. Their rotation of the dextral shell to the left was adequate for sand discharge. Sinistral shells were rotated in both directions. Analysis of recorded videotapes showed that variation in rotation direction could be attributed to variation in the position of the crab relative to the shell. When the crab faced the shell aperture from the inner lip, it rotated the sinistral shell to the right, and to opposite direction when it faced from the outer lip side. The crab always pushed the upper side of the horizontally laid shell, regardless of shell type or its own position.     

The effects of shell size and coil orientation on reproduction in female hermit crabs, Clibanarius vittatus

We investigated the effects of shell coil orientation and shell size on reproduction in field populations of the hermit crab, Clibanarius vittatus. Females were collected in the intertidal in Beaufort, NC. Shell parameters were measured and size (cephalothorax length) and reproductive status were determined for 70 females occupying Busycon shells. Crabs were categorized as berried (eggs on the pleopods), mature ovaries, or non-reproductive (no eggs). For berried females, the number of eggs was recorded. By offering a separate group of females access to empty shells, it was possible to calculate optimal shell size and the deficit in shell size for field-collected animals.Females that were berried were in shells closer to the optimal shell size than females with mature ovaries, both for shell weight and shell volume. And females with mature ovaries were in shells that were closer to the optimal size than females that were non-reproductive. For both categories of females without eggs on the pleopods, the majority of females were in shells that were too big (in weight and internal volume). While the percentage of berried females did not differ between dextral (Busycon carica) and sinistral (Busycon sinistrum) shells, the non-reproductive females had a much smaller deficit in volume in sinistral shells compared to dextral shells. For berried females, there was no relationship between the magnitude of their shell deficit and the number of eggs carried. Our results suggest that reproduction is inhibited when females occupy shells sufficiently greater than the optimal shell size.     

Linking taphonomy to community-level abundance: Insights into compositional fidelity of the Upper Triassic shell concentrations (Eastern Alps)

Although actualistic live/dead comparisons lead to robust estimates of fidelity of modern death assemblages, quantitative evaluation of fidelity of fossil assemblage remains uncertain. In this paper, effects of storm reworking on compositional fidelity of the Upper Triassic shell concentrations (Eastern Alps, Austria) are evaluated. An exploratory approach is based on comparison of reworked and non-reworked assemblages in ordination analyses. Non-reworked assemblages of one or more communities provide a baseline for evaluation of fidelity of reworked assemblages. In siliciclastic-rich intervals of the Kössen Formation, shell concentrations are represented by (1) packstones with small, shallow infaunal bivalves, (2) floatstones and pavements with large semi-infaunal bivalves, and (3) bioclastic marlstones. In carbonate-rich intervals, bioclastic floatstones with bivalves and brachiopods occur. Analyzing all shell concentrations, eight sample groups sharing similar species composition are discriminated. Limited effect of storm reworking on composition of shell concentrations is indicated by (1) a general persistence of six sample groups when only non-reworked assemblages are analyzed, (2) similarity in composition between reworked and non-reworked assemblages within sample groups, and (3) compositional segregation between non-reworked assemblages of distinctive sample groups, mostly without any reworked assemblages of intermediate composition.Depth-related variations in dead-shell production, shell destruction and body size governed preservation and distribution of the shell concentrations along onshore-offshore gradient in the Kössen Basin. First, at times when environmental conditions were unfavorable for shell producers, coupled with high background shell destruction rates, limestone beds formed during storm events were shell-poor. Second, less common shell concentrations in upper than in lower parts of siliciclastic intervals can be related to higher environmental stress in shallower habitats. Third, the difference between shell concentrations dominated by small and large bivalves is driven by between-habitat differences in body size and is not due to a differential sorting of small and large shells. Combining community analysis based on species abundances with taphonomic analysis can thus be helpful in tracking fidelity of fossil assemblages.     

Gonad pouches in a new clam-like monoplacophoran from the Silurian of Sweden

Beyrichidiscus fecundatus n. gen. n. sp. is described from the Upper Silurian of Gotland, Sweden. The minute, isostrophically coiled mollusc exhibits profound allometry and is considered to have been a monoplacophoran on the basis of functional interpretation of apertural structures. A clam-like, semi-infaunal mode of life is suggested. A pair of prominent lateral swellings in the adult shell are interpreted as reflecting the location of the gonads.     

Epizoan encrustation on Marsupites testudinarius – additional argument favoring an epifaunal mode of life of uintacrinoids?

Uintacrinoids (Uintacrinoidea Zittel) are among the best-known Late Cretaceous crinoids, but owing to their unusual morphology and widespread distribution their mode of life has become a subject of much discussion. Of several competing hypotheses, nektonic, pseudoplanktonic, hemipelagic, semi-infaunal and epibenthic lifestyles have been suggested. Recent study synthesizing and extending previous data has shown that these crinoids were epibenthic, holding their arms vertically for feeding. However, evidence supporting a rheophilic epibenthic model over an alternative rheophobic semi-infaunal model is still limited. Here we report epizoans, mostly represented by serpulids and bryozoans, encrusting thecal plates of Marsupites testudinarius from the Lägerdorf in Germany. Although a definitive interpretation whether recorded infestations occurred syn vivo or post mortem is not certain, it is remarkable that all epizoans (or their traces) are attached to the convex side (latera) of well-preserved isolated plates displaying no signs of reworking. Furthermore, a bryozoan colony crossing plate boundaries has been also found on the surface of a sub-articulated theca suggesting that it may have been colonised syn vivo. Recorded epibiotic associations, whether syn vivo or post-mortem, must have developed prior to burial of the specimens, above the surface of sea floor, and thus provide another argument against rheophobic semi-infaunal mode of life of uintacrinoids.     

Delayed prezygotic isolating mechanisms: evolution with a twist

Assortative mating characterizes the situation wherein reproducing individuals pair according to similarity. Usually, the impetus for this bias is attributed to some type of mate choice conferring benefits (e.g., increased fitness or genetic compatibility) and, thereby, promoting speciation and phenotypic evolution. We investigate, by computer simulation of an evolving deme-structured snail population, the ramifications ensuing from passive assortative mating wherein couples exhibiting opposite shell coil direction phenotypes experience a physical constraint on mating success: putative mating partners inhabiting stout dextral and sinistral shells are unable to exchange sperm. Because shell coil chirality genotype is encoded at a single locus by shell coil alleles that are inherited maternally, snails containing sinistral alleles can present the typical dextral phenotype. Consequently, the incidence of a sinistral allele in as few as one snail can be manifested as prezygotic reproductive isolation within a deme in a subsequent generation. However, because the efficacy of achieving this type of prezygotic reproductive isolation is affected by shell form, the likelihood and product of single-gene speciation should be determined by deme interaction (migration) and composition (morphological distribution). We test this hypothesis and show how stochastic migration interacts with passive assortative mating yielding morphologically induced prezygotic reproductive isolation to produce new species phenotypes. The results show that demes can achieve rapid macroscopic phenotypic transformation and indicate that sympatric speciation might be more plausible than naturalists recognize conventionally.     

Analysis of the shell torsion phenomenon in the Bivalvia

The curious phenomenon of shell torsion arose independently among members of two separate lineages within the Pteriomorpha (Bivalvia), the Arcidae and the Bakevelliidae. In torted bivalves the shell is twisted about the hinge axis normal to the antero-posterior direction, and the usually two-dimensional plane containing the commissure is deformed into a complex three-dimensional surface. The fabrication of shell torsion is particularly amenable to mathematical analysis, and by utilizing computer simulation it is possible to tort a normal bivalve by introducing 'errors' of a particular nature into the morphogenetic program. Analysis of the pattern of torsion within the Arcidae and Bakevelliidae, and the examination of modem untorted arcids, indicate that the interference of the byssus with the normal bivalve morphogenetic program played a definitive role in the origination of the torsion. Though quite different adaptive pathways were utilized by the two groups, the fabricational principle producing the torsion remained the same. The resultant torted morphologies observed in the Arcidae and Bakevelliidae can be understood in terms of the interaction between the adaptive significance of the torsion and the historical, or preexisting morphology of the animals involved.     

Larval muscle contraction fails to produce torsion in a trochoidean gastropod.

The causes and effects of ontogenetic torsion in gastropods have been debated intensely for more than a century (1-19). Occurring rapidly and very early in development, torsion figures prominently in shaping both the larval and adult body plans. We show that mechanical explanations of the ontogenetic event that invoke contraction of larval retractor muscles are inadequate to explain the observed consequences in some gastropods. The classic mechanical explanation of Crofts (4, 5) and subsequent refinements of her explanation have been based on species with rigid larval shell properties (18, 19) that cannot be extrapolated to all gastropods. We present visual evidence of the lack of rigidity of the uncalcified larval shell in a basal trochid gastropod, Margarites pupillus (Gould), and provide photographic confirmation of our prediction that larval retractor muscle contraction is insufficient to produce more than local deformation or dimpling at the site of muscle insertion. These findings do not refute muscular contraction as a primary cause of ontogenetic torsion in gastropods that calcify their larval shells prior to the onset of torsion, nor do they refute the monophyly of torsion. They do, however, suggest that torsion may be a loosely constrained developmental process with multiple pathways to the more constrained end result (20, 21).     

PREY CAPTURE, PREFERENCE AND CONSUMPTION BYLINATELLA CAUDATA (GASTROPODA: TONNOIDEA: RANELLIDAE) IN HONG KONG

On sheltered rocky beaches in Hong Kong, Linatella caudata preysupon the lower-littoral and sub-littoral fringe fauna. Ascidiantests are cut with the radula, while the proboscis is insertedbetween the parted shell valves of bivalves. The salivary glandssecrete sulphuric acid that is not used in prey penetration,as in the Cassidae, but is more likely used in digestion ordefence. In preference experiments, prey were chosen in the sequence:Barbatia virescens (Bivalvia: Arcidae) > Styela plicata (Ascidiacea)/Saccostreacucullata (Bivalvia: Ostreidae) > Lunella coronata (Gastropoda:Turbinidae) > Balanus amphitrite (Crustacea: Cirripedia).52% of all attacks were upon Barbatia Lunella and Balanus wereattacked rarely. Prey preferences by individual gastropods werealso demonstrated. Calibration plots of total weight against wet and dry tissueweight of prey (Barbatia virescens) and, finally, the predatorwere obtained and used in estimations of consumption. On average,an adult Linatella caudata (10-15g) consumed three B. virescens.week^–1 The mean weight of B. virescens flesh consumed.week^–1 ranged between 0. 208–0.412g dry weight prey.gdry weight predator^–1, i.e., a mean of 28.2% of the predator'body weight.week^–1 or 4%.day^–1. Such a figure accordswell with estimates of consumption obtained for other adultpredatory gastropods from Hong Kong. (Received 4 September 1989; accepted 5 December 1989)     

Fine Mapping of the Pond Snail Left-Right Asymmetry (Chirality) Locus Using RAD-Seq and Fibre-FISH

The left-right asymmetry of snails, including the direction of shell coiling, is determined by the delayed effect of a maternal gene on the chiral twist that takes place during early embryonic cell divisions. Yet, despite being a well-established classical problem, the identity of the gene and the means by which left-right asymmetry is established in snails remain unknown. We here demonstrate the power of new genomic approaches for identification of the chirality gene, “D”. First, heterozygous (Dd) pond snails Lymnaea stagnalis were self-fertilised or backcrossed, and the genotype of more than six thousand offspring inferred, either dextral (DD/Dd) or sinistral (dd). Then, twenty of the offspring were used for Restriction-site-Associated DNA Sequencing (RAD-Seq) to identify anonymous molecular markers that are linked to the chirality locus. A local genetic map was constructed by genotyping three flanking markers in over three thousand snails. The three markers lie either side of the chirality locus, with one very tightly linked (<0.1 cM). Finally, bacterial artificial chromosomes (BACs) were isolated that contained the three loci. Fluorescent in situ hybridization (FISH) of pachytene cells showed that the three BACs tightly cluster on the same bivalent chromosome. Fibre-FISH identified a region of greater that ∼0.4 Mb between two BAC clone markers that must contain D. This work therefore establishes the resources for molecular identification of the chirality gene and the variation that underpins sinistral and dextral coiling. More generally, the results also show that combining genomic technologies, such as RAD-Seq and high resolution FISH, is a robust approach for mapping key loci in non-model systems.     

Alternating torsions in a living '9 + 2' flagellum

Live spermatozoa of the Japanese quail were observed as they swam in highly viscous salines. Under these conditions, torsions of the flagellum were readily seen. The torsions had a characteristic magnitude (nominally 180 degrees) and pattern of incidence. As a cycle of bending propagated over it, each position on the flagellum experienced first a sinistral torsion and, later, a restoring dextral torsion. The two zones of torsion were each associated with bending; between them was a torsion-free zone that tended to be straight. The amount of interdoublet sliding needed to generate the torsions may be as little as 10 nm. These dynamic propagating torsions have been detected by following the angular displacements of individual (swollen) mitochondria lying adjacent to the axoneme. It is suggested that torque generation is a primary outcome when the unconstrained ''9 + 2'' axoneme is activated.     

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.     

Internal selection against the evolution of left-right reversal

Among metazoan species, left-right reversals in primary asymmetry have rarely gone to fixation. This suggests that a general mechanism suppresses the evolution of polarity reversal. Most metazoans appear externally symmetric and reproduce by external fertilization or copulation with genitalia located in the midline. Thus, reversal should generate little exogenous disadvantage when interacting with the external environment or in mating with the common wild-type. Accordingly, an endogenously caused fitness reduction may be responsible for the general absence of reversed species. However, how this selection operates is little understood. Phenotypic changes associated with reversal are usually inseparable from zygotic pleiotropy. By exploiting hermaphroditism and the maternal inheritance of left-right polarity, we generated dextral and sinistral snails that share the same zygotic genotype. Before hatching, these sinistrals developed lethal morphological anomalies more frequently than dextrals. Their shell shape at maturity differed from the mirror image of the dextral shell. These interchiral differences demonstrate pleiotropy in maternal effects of the polarity or linked genes. Variation in interchiral differences between parental crosses suggests the presence of epistatic variation in relative performance of sinistrals. Our results show that internal selection operates against polarity reversal, and we suggest that this is due to changes in blastomere configuration.