Does variation in morphology correspond with variation in habitat use in freshwater gastropods?

We tested if variations (i.e., breadth) in morphology and habitat use vary predictably among six aquatic gastropod species that were collected across Indiana and Illinois, USA. We predicted that interspecific morphological variation would positively covary with variation in habitat use among species. We used geometric morphometrics (Procrustes technique and relative warp analysis) to quantify morphology and multivariate analyses (PCA) to quantify habitat. Increased morphological breadth did not vary predictably with increased habitat breadth. However, we found that life history traits correspond with patterns in morphological and habitat breadth for these six aquatic gastropods. Pulmonate gastropods (use lungs for respiration) that lack an operculum cover exhibited decreased morphological breadth compared to coenogastropods (use gills for respiration). This pattern may ultimately be a function of behavioral adaptations in freshwater gastropods. Gastropods that are capable of breathing air or using other behavioral modifications such as burrowing to escape predators may not require high morphological breadth. Conversely, selection may favor higher morphological breadth in gastropods with gills that also do not move out of the water column to escape predators.     

Locomotion and shell-righting behaviour in adult and juvenile Aporrhais occidentalis (Gastropoda: Strombacea)

Differences in behaviour between adult and juvenile Aporrhais occidentalis are related to age-dependent changes in shell morphology and habitat. Juvenile snails right their shells by pulling with the propodium, and do not ‘kick’ at the substratum until the expanded outer lip of the adult shell has formed. Locomotion of juveniles is by arhythmical crawling, with ‘leaping’ taking place only during escape from predators. Adult snails leap during both normal and escape locomotion. Data are presented that demonstrate the importance of the outer lip in stabilizing the shell during leaping locomotion. Escape reactions of A. occidentalis are similar to, but slower than, those reported in the literature for species of strombids. Most of the basic locomotory and shell-righting behaviour patterns seen in the highly specialized Strombidae are observable in the more primitive A. occidentalis. However, the use of the operculum in locomotion and shell righting, although characteristic of the Strombidae, is not found in A. occidentalis. The behaviour of Aporrhais is discussed in relation to evolution within the superfamily Strombacea.     

Larval development of hyolithids

The larval development of hyolithids is inferred to have been closely similar to that of primitive gastropods. A trochophore-like larva produced a swollen initial conch and a convex operculum. The initial conch was either subsphaerical and smooth or fusiform with a pointed apex and growth lines. This is taken to indicate an initial development within egg covers or free-living, respectively. A pelagic veliger phase followed. preceded by hatching in the first case. Veliger shells are similar to adult ones. Metamorphosis (transformation of veliger into benthic adult animal) in both hyolithids and gastropods was accompanied by an increase in mortality.     

Comparative morphology of the amphibian opercularis system: I. General design features and functional interpretation

The morphology of the opercularis system of anuran and caudate amphibians suggests that it acts to produce motion of the operculum that in turn produces fluid motion within the inner ear. The operculum and opercularis muscle form a lever system, with a narrow connection between the operculum and otic capsule acting as a fulcrum about which the operculum moves in response to forces applied via the muscle. The opercula of many species possess a muscular process on which the muscle inserts, thereby increasing the moment arm through which the muscle acts. The tonicity of the opercularis muscle allows tensile forces produced by substrate vibration or other mechanical energy applied to the forelimb to be effectively transmitted to the operculum; the elasticity of the connective tissue holding the operculum in place should act to return the operculum to its original position. The opercularis systems of frogs and non-plethodontid salamanders are similar structurally and functionally; that of plethodontid salamanders is structurally distinct but also functions as a lever system. Fluid motion produced by opercular motion could stimulate various end organs of the inner ear; the saccule, lagena, and amphibian papilla are in close approximation and wave energy could directly affect their otoconial or tectorial structures. In those anurans with a tympanic ear, the stapedial footplate and operculum articulate, but this articulation allows both to move independently. The stapes-tympanum complex and opercularis system therefore appear to be independent functional systems, and it is unlikely that the opercularis system modulates middle ear responsiveness. The general design of the opercularis system is consistent with a function in reception of substrate vibrations.     

Chitin in gastropod operculi

The operculi of 114 species of gastropods representing 38 families with wide ranging morphological, ecological and geographical differences have been examined for the presence of chitin using a quantitative enzymatic method along with qualitative tests. All the species having calcified operculi so far analysed (28 species) were relatively rich in chitin and neighbouring species had a similar composition. No chitin was found in the non-calcified, corneous operculi (86 species). The level of calcification was the unique character found to be correlated to the presence of chitin in the operculum; no other morphological parameters showed any reciprocity. Within the calcified operculi, chitin is restricted to the mineralized matrix. The tanned corneous sheet seems completely devoid of chitin. The possibility of a direct relation between calcification processes and the presence of chithin in the mineralizing matrix of some molluscan hard structures is discussed.     

Usefulness of the opercular nucleus for inferring early development in neritimorph gastropods

The early ontogeny of gastropods (i.e., planktotrophic vs. nonplanktotrophic) may be inferable from the morphology of the protoconch in adult shells. The protoconch consists of both embryonic and larval shells in species with planktotrophic development; the embryonic shell forms in the intracapsular period and the succeeding larval shell gradually develops during the larval period. In nonplanktotrophic species, on the other hand, there is no additional growth of the larval shell and the protoconch consists exclusively of a relatively large embryonic shell formed prior to hatching. This "shell apex theory" has been applied to many species of shell-bearing gastropods, but biotic and abiotic erosion of the apex often prevents detailed examination of the protoconch and subsequent inferences about ontogeny. I examined the gastropod operculum to test its utility for predicting developmental mode, drawing on the Neritimorpha as model taxa. Most aquatic members of Neritimorpha were found to bear an operculum with a clearly demarcated nucleus; SEM observations reveal four types of nuclei, which correspond to different types of protoconch morphologies and observed ontogenies for the study species. The nucleus is secreted before metamorphosis, fits into the shell aperture of the larva, and reflects early ontogeny as morphology, as does the protoconch. Moreover, the apparently organic (rather than calcareous) composition of the nucleus makes it nearly invulnerable to erosion and very advantageous, compared to the protoconch, in this ecologically diverse group, whose habitats range from freshwater streams and mangrove swamps to rocky shores and deep-sea hydrothermal vents. The measurements of the nucleus are also valuable for taxonomic purposes, especially in the species identification of veliger larvae and juvenile snails. On the other hand, the opercular nuclei of the Caenogastropoda and Heterobranchia are often eroded away in adult individuals; even if present, the morphology of the nuclei does not seem to clearly reflect early ontogeny in those groups.     

Righting kinematics in beetles (Insecta: Coleoptera)

Twenty modes of stereotyped righting motions were observed in 116 representative species of coleoptera. Methods included cine and stereocine recording with further frame by frame analysis, stereogrammetry, inverse kinematic reconstruction of joint angles, stroboscopic photography, recording of electromyograms, 3D measurements of the articulations, etc. The basic mode consists of a search phase, ending up with grasping the substrate, and a righting, overturning phase. Leg coordination within the search cycle differs from the walking cycle with respect to phasing of certain muscle groups. Search movements of all legs appear chaotic, but the tendency to move in antiphase is still present in adjacent ipsilateral and contralateral leg pairs. The system of leg coordination might be split: legs of one side might search, while contralateral legs walk, or fore and middle legs walk while hind legs search. Elaborated types of righting include somersaults with the aid of contralateral or diagonal legs, pitch on elytra, jumps with previous energy storage with the aid of unbending between thoracic segments (well-known for Elateridae), or quick folding of elytra (originally described in Histeridae). Righting in beetles is compared with righting modes known in locusts and cockroaches. Search in a righting beetle is directed dorsad, while a walking insect searches for the ground downwards. Main righting modes were schematized for possible application to robotics.     

A new species of Haplosporidium Caullery & Mesnil, 1899 in the marine false limpet Siphonaria lessonii (Gastropoda: Siphonariidae) from Patagonia

A new species of Haplosporidium Caullery & Mesnil, 1899 parasitising the pulmonate gastropod Siphonaria lessonii Blainville in Patagonia, Argentina, is described based on morphological (scanning and transmission electron microscopy) and sequence (small subunit ribosomal RNA gene) data. Different stages of sporulation were observed as infections disseminated in the digestive gland. Haplosporidium patagon n. sp. is characterised by oval or slightly subquadrate spores with an operculum that is ornamented with numerous short digitiform projections of regular height, perpendicular to and covering its outer surface. The operculum diameter is slightly larger than the apical diameter of the spore. Neither the immature nor mature spores showed any kind of projections of the exosporoplasm or of the spore wall. Regarding phylogenetic affinities, the new species was recovered as sister to an undescribed species of Haplosporidium Caullery & Mesnil, 1899 from the polychaete family Syllidae Grube from Japanese waters. The morphological characters (ornamentation of the operculum, spore wall structure, shape and size of spores, and the lack of spore wall projections) corroborate it as an as yet undescribed species of Haplosporidium and the first for the phylum in marine gastropods of South America. Siphonaria lessonii is the only known host to date.     

Developmental role of dpp in the gastropod shell plate and co-option of the dpp signaling pathway in the evolution of the operculum

The operculum is a novel structure in gastropod molluscs. Because the operculum shows notable similarities to the shell plate, we asked whether there were an evolutionary link between these two secretory organs. We found that some of the genes involved in shell-field development are expressed in the operculum, such as dpp and grainyhead, whereas engrailed and Hox1 are not. Specific knockdown of dpp by injection of double-stranded RNA (dsRNA) resulted in malformation of the shell plate. The shell plate was smaller due to failure of activation of cell proliferation in the shell-field margin. The expressions of grainyhead and chitin synthase 1 in the shell field margin were suppressed by dpp-dsRNA. However, matrix secretion was not completely abolished, and the expressions of ferritin, engrailed or Hox1 were not affected by dpp-dsRNA, indicating that dpp is partly involved in the developmental pathway for shell matrix secretion. We also present evidence that dpp performs a key role in operculum development. Indeed, dpp-dsRNA impaired matrix secretion in the operculum as well as expression of grainyhead. Based on these observations that dpp is important for development of both the shell plate and operculum, we conclude that co-option of dpp to the posterior part of the foot contributed to the innovation of the operculum in gastropods.     

Seed Micromorphology and Its Taxonomic Significance of Begonia (Begoniaceae) in China and Vietnam

The seed morphology of 47 species in Begonia in China and Vietnam were examined under Light Microscope (LM) and Scanning Electron Microscope (SEM), respectively. Seed varied from ellipsoidal to obovoid, sometimes broadly obovoid shape with large variation in size. The shape of operculum could be classified into three types: short claviform, broadly nipple shaped and obtuse. There are four types of epidermal ornamentations: straight striae, undulated striae, rounded punctum and herringbone shaped punctum. The principal component analysis and hierarchical cluster analysis revealed that the classification of Begonia based on the seed micromorphology was incompletely consistent with the traditional classification on the section level based on the placenta; the ornamentation of seeds showed stability in different populations of a species; the operculum shape shows a considerable difference among morphologically similar species. Seed morphological characters may relate to the dispersal patterns which showed an adaptation to the living environment.     

Connecting species richness, abundance and body size in deep-sea gastropods

Aim This paper examines species richness, abundance, and body size in deep‐sea gastropods and how they vary over depth, which is a strong correlate of nutrient input. Previous studies have documented the empirical relationships among these properties in terrestrial and coastal ecosystems, but a full understanding of how these patterns arise has yet to be obtained. Examining the relationships among macroecological variables is a logical progression in deep‐sea ecology, where patterns of body size, diversity, and abundance have been quantified separately but not linked together. Location 196–5042 m depth in the western North Atlantic. Method Individuals analysed represent all Vetigastropoda and Caenogastropoda (Class Gastropoda) with intact shells, excluding Ptenoglossa, collected by the Woods Hole Benthic Sampling Program (3424 individuals representing 80 species). Biovolume was measured for every individual separately (i.e. allowing the same species to occupy multiple size classes) and divided into log₂ body size bins. Analyses were conducted for all gastropods together and separated into orders and depth regions (representing different nutrient inputs). A kernel smoothing technique, Kolmogorov‐Smirnov test of fit, and OLS and RMA were used to characterize the patterns. Results Overall, the relationship between the number of individuals and species is right skewed. There is also a positive linear relationship between the number of individuals and the number of species, which is independent of body size. Variation among these relationships is seen among the three depth regions. At depths inferred to correspond with intermediate nutrient input levels, species are accumulated faster given the number of individuals and shift from a right‐skewed to a log‐normal distribution. Conclusion A strong link between body size, abundance, and species richness appears to be ubiquitous over a variety of taxa and environments, including the deep sea. However, the nature of these relationships is affected by the productivity regime and scale at which they are examined.     

Operculum regeneration following failed predation in the Silurian gastropod Oriostoma

Regeneration of the calcareous rigiclaudent operculum following severe, non‐lethal, predatory attacks is described in two specimens of the characteristic Silurian gastropod Oriostoma from the English Midlands. This is the first record of the regeneration of opercula in Palaeozoic gastropods and of opercula conjoined with shells that have been repaired after significant, failed, durophagous attacks. In one specimen, a series of repaired injuries culminates in a repaired aperture closed by the operculum. In the second specimen, new shell growth has failed to recover the full extent of the original broken whorl, and the new aperture and operculum, though well developed, are smaller than the originals. Both opercula are unusually thin centrally when compared to other, strongly domed, Oriostoma opercula.     

Anatomy and systematic affinity of Stanleya neritinoides (Smith, 1880), an enigmatic member of the thalassoid gastropod species flock in Lake Tanganyika, East Africa (Cerithioidea, Paludomidae)

The radiation of gastropods in Lake Tanganyika is an ideal system for testing competing hypotheses of species flock formation. Yet, much of the basic biology of these species remains unknown. In an ongoing effort to understand the evolution of Tanganyikan gastropods, we here describe Stanleya neritinoides. Alcohol‐preserved material of the soft parts is rare, consequently, the systematic position of the species, and a repeated suggested affinity to Tanganyicia rufofilosa, have been based primarily on features of the shell. However, features of the radula and operculum are unique and do not suggest an affinity to any other Tanganyikan species. Thus, S. neritinoides has remained a particularly poorly known and enigmatic member of the species flock. This investigation confirmed that several aspects of internal and external anatomy are shared between S. neritinoides, T. rufofilosa, and other Tanganyikan gastropods, but that S. neritinoides is unique in features of the radula and seminal receptacle. Moreover, S. neritinoides differs from T. rufofilosa in features of the foregut, midgut, hindgut, kidney, nervous system, reproductive system and reproductive strategy. These new data are inconsistent with an interpretation of identity of Stanleya and Tanganyicia. In addition, given the pervasive differences between the two, a sister‐group relationship between the two is unlikely. More precise systematic placement of S. neritinoides awaits the establishment of a phylogenetic framework for all Tanganyikan gastropods.     

Avoidance and escape responses of the sub-antarctic limpet Nacella edgari (Powell) (Mollusca: Gastropoda) to the sea star Anasterias perrieri (Smith) (Echinodermata: Asteroidea)

Summary The sub-antarctic limpet Nacella edgari (Powell) (Patellidae) has avoidance and escape responses to the sea star Anasterias perrieri (Smith) (Asteriidae) in a kelp forest and in the laboratory. Nacella edgari was preyed upon in the field by A. perrieri, but frequency of capture was low, suggesting the effectiveness of defensive behaviors. Both avoidance and escape responses by the limpet consisted of extension and waving of the pallial tentacles, mushrooming and rotation of the shell and flight. However, escape responses were characterized by greatly intensified rotation of the shell through a horizontal plane. This study extends the evaluation of defensive responses in gastropods to a species which occurs in the southern Indian Ocean.     

Motor activity and trajectory control during escape jumping in the locust <Emphasis Type="Italic">Locusta migratoria</Emphasis>

We investigated the escape jumps that locusts produce in response to approaching objects. Hindleg muscular activity during an escape jump is similar to that during a defensive kick. Locusts can direct their escape jumps up to 50° either side of the direction of their long axis at the time of hindleg flexion, allowing them to consistently jump away from the side towards which an object is approaching. Variation in jump trajectory is achieved by rolling and yawing movements of the body that are controlled by the fore- and mesothoracic legs. During hindleg flexion, a locust flexes the foreleg ipsilateral to its eventual jump trajectory and then extends the contralateral foreleg. These foreleg movements continue throughout co-contraction of the hindleg tibial muscles, pivoting the locust’s long axis towards its eventual jump trajectory. However, there are no bilateral differences in the motor programs of the left and right hindlegs that correlate with jump trajectory. Foreleg movements enable a locust to control its jump trajectory independent of the hindleg motor program, allowing a decision on jump trajectory to be made after the hindlegs have been cocked in preparation for a jump.     

Effects of quantitative and qualitative variation in phenolic compounds on feeding in three species of marine invertebrate herbivores

The deterrent effects of brown algal phenolic compounds and the terrestrial polyphenolic tannic acid on feeding by three species of invertebrate herbivores from central California, including the gastropods Tegula funebralis (Adams) and Tegula brunnea (Phillipi) and the echinoid Strongylocentrotus purpuratus (Stimpson) were examined. Algal phenolics used were the monomeric phenolic phloroglucinol and polyphloroglucinols from Fucus vesiculosus (Linnaeus), Halidrys siliquosa (Linnaeus) Lyngbye and Eisenia arborea Areschoug. All of the polyphenolics deterred feeding by all three herbivores at concentrations of 5 mg · ml⁻¹ in agar disks. Concentrations of 2 mg · ml⁻¹ also generally deterred feeding by the gastropods (these levels were not tested against S. purpuratus). Relative amounts of deterrence by different compounds were similar, especially for the gastropods. Phloroglucinol deterred feeding by the echinoids, but not by T. funebralis. Responses of the echinoids were otherwise similar to the gastropods, but more variable. I also demonstrated deterrence of S. purpuratus by tannic acid using the “tanned” kelp technique of Steinberg (1985). Reactivity of the different phenolic compounds in the Folin-Denis procedure, a common colorimetric assay used to estimate levels of phenolics in plant tissue, was similar. This suggests that measuring phenolic levels in brown algae by this technique will not be greatly confounded by the occurrence of different kinds of phenolic molecules in different brown algae. This result, in combination with the similarity of the deterrent effects of the compounds used in this study, increases the validity of previous studies in the northeastern Pacific Ocean which correlate algal phenolic levels and diets or feeding preferences of invertebrate herbivores. For plants and herbivores in this region, this assay is a reasonable measure of a biologically meaningful phenomenon — levels of phenolic deterrents in the algae.     

Polymorphisms of octopine dehydrogenase (Odh) in mollusks and implications for the neutralism-selectionism hypothesis

Octopine dehydrogenase (Odh) was examined in several species of bivalves and gastropods and complemented with bibliographic data, to assess the controversy between neutralism and selectionism in explaining the maintenance of genetic variation in natural populations. This debate was the center of the molecular evolution and population genetic research in the 1970s and 1980s, but waned thereafter, without resolution. Although DNA data have been produced, implications are not understood. We examined the polymorphims of Odh in several species of bivalves and gastropods, and the kinetic properties (apparent Km) of the different isozymes in the scallop Euvola ziczac that indicates an apparent case of overdominance of the heterozygous individuals. The question "which of the two hypothesis is correct" has shifted with time to "how much influence did each factor have in the maintenance of genetic variation".     

Development of asymmetry in the caenogastropods Amphissa columbiana and Euspira lewisii

Abstract. The asymmetry displayed by the body plan of gastropods has been directly or indirectly attributed to an evolutionary process called torsion. Torsion is defined as a rotation of 180° between the cephalopodium (head and foot) and visceropallium (visceral organs, mantle, mantle cavity, and shell). During development, the displacement of anatomical components occurs during a process called "ontogenetic torsion." Although ontogenetic torsion is central to theories of gastropod evolution, surprisingly few studies have documented actual tissue movements during the development of asymmetry in gastropods. We investigated the development of the mantle cavity and pleurovisceral nerve connective (visceral nerve loop) in the caenogastropods Amphissa columbiana and Euspira lewisii , because displacements of both of these structures are interpreted as major consequences of torsion. Scanning electron micrographs, histological sections, and immunofluorescence images showed that the developing vis-ceropallium twists by 90° relative to the cephalopodium, the mantle cavity initially forms on the right side, and displacements of the visceral nerve loop become evident on the left side before the right side. A developmental stage in which the mantle cavity is confined to the right side has also been reported in members of the Vetigastropoda and Heterobranchia. We suggest that further comparative studies should test the hypothesis that early development throughout the Gastropoda converges on an embryonic organization in which the mantle cavity and anus are located laterally, despite clade-specific differences in developmental patterns both before and after this stage.     

Internal morphology of two species of Lagenochitina (Chitinozoa)

Internal and external structures of Lagenochitina boja Bockelie 1980 and L. esthonica Eisenack 1955 have been examined with a scanning electron microscope. The material derives from limestone samples from the Arenig to ?Lower Llanvirn succession of NE Spitsbergen. The species possesses both an operculum and a prosoma (Bockelie, 1978). The operculum, in the shape of a thin flat disc, closes the aperture of the test. A cylindroconical structure is enclosed in the test aborally of the operculum. the cylindrical part of this structure, the prosoma, is transversed by numerous thin disc-shaped laminae. The sequence of thin laminae is terminated in a thicker lamina, here termed the subex. The prosoma wall is extended aborally forming a conical structure, the rica. The prosomai of individual specimens show variation in length, microstructures and the attachment to the test. The internal structures of Lagenochitina species are compared with similar structures known from other chitinozoan species.