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.     

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.     

Behavioral and Physiological Responses to Emersion in Freshwater Bivalves

SYNOPSIS. Littoral lentic and shallow lotic freshwater habitatsare unpredictable in periodicity and duration of shore emersion.As a result, freshwater bivalves have evolved extensive capacitiesto withstand prolonged emersion. Valve movement behaviors allowemersed bivalves to control rate of water loss while maintainingat least partial aerial gas exchange; these behaviors are affectedby environmental variables such as temperature and relativehumidity. Aerial oxygen uptake is associated directly with valveventilatory behaviors and mantle edge exposure. Such behaviorsare often phasic, indicative of oxygen "debt" payment. Lackingeffective hemolymph buffer, respiratory acidosis during emersionis compensated by shell carbonate stores allowing hemolymphP_CO2 to rise to levels facilitating diffusion of CO₂ to theenvironment. During emersion, hemolymph calcium can increasefour fold while Na and Cl are tightly regulated. Ammonia productionceases in emersed bivalves. It resumes on reimmersion, indicativeof heavy reliance on non-protein catabolism during emersion.Some emersion adaptations of freshwater species appear to bemodifications of those displayed by intertidal and estuarinebivalves, while others appear independently evolved to allowsurvival of the extreme emersion periods associated with lifein shallow freshwaters.     

Evidence of predation in Early Cretaceous unionoid bivalves from freshwater sediments in the Cameros Basin,Spain

Here, we present evidence of possible vertebrate predation on freshwater bivalves from the Lower Cretaceous strata of the Cameros Basin (Spain). The described collection contains the largest number of vertebrate‐inflicted shell injuries in freshwater bivalve shells yet reported in the Mesozoic continental record. Several types of shell damage on fossil shells of Protopleurobema numantina (Bivalvia: Unionoida) are described and their respective modes of formation interpreted in the context of morphological attributes of the shell injuries and the inferred tooth morphology of predators that could have inflicted such injuries. Detailed study of these bite marks shows similarities with the well‐documented injuries in the shells of marine molluscs, namely ammonoids, that have likewise been attributed to reptilian predators. The most parsimonious interpretation suggests crocodiles as the vertebrates interacting with the bivalves in the Cameros Basin. □Barremian–Aptian; bite marks; freshwater bivalves; predation; reptile; Unionoida.     

Abbreviation of larval development and extension of brood care as key features of the evolution of freshwater Decapoda

The transition from marine to freshwater habitats is one of the major steps in the evolution of life. In the decapod crustaceans, four groups have colonized fresh water at different geological times since the Triassic, the freshwater shrimps, freshwater crayfish, freshwater crabs and freshwater anomurans. Some families have even colonized terrestrial habitats via the freshwater route or directly via the sea shore. Since none of these taxa has ever reinvaded its environment of origin the Decapoda appear particularly suitable to investigate life‐history adaptations to fresh water. Evolutionary comparison of marine, freshwater and terrestrial decapods suggests that the reduction of egg number, abbreviation of larval development, extension of brood care and lecithotrophy of the first posthatching life stages are key adaptations to fresh water. Marine decapods usually have high numbers of small eggs and develop through a prolonged planktonic larval cycle, whereas the production of small numbers of large eggs, direct development and extended brood care until the juvenile stage is the rule in freshwater crayfish, primary freshwater crabs and aeglid anomurans. The amphidromous freshwater shrimp and freshwater crab species and all terrestrial decapods that invaded land via the sea shore have retained ocean‐type planktonic development. Abbreviation of larval development and extension of brood care are interpreted as adaptations to the particularly strong variations of hydrodynamic parameters, physico‐chemical factors and phytoplankton availability in freshwater habitats. These life‐history changes increase fitness of the offspring and are obviously favoured by natural selection, explaining their multiple origins in fresh water. There is no evidence for their early evolution in the marine ancestors of the extant freshwater groups and a preadaptive role for the conquest of fresh water. The costs of the shift from relative r‐ to K‐strategy in freshwater decapods are traded‐off against fecundity, future reproduction and growth of females and perhaps against size of species but not against longevity of species. Direct development and extension of brood care is associated with the reduction of dispersal and gene flow among populations, which may explain the high degree of speciation and endemism in directly developing freshwater decapods. Direct development and extended brood care also favour the evolution of social systems, which in freshwater decapods range from simple subsocial organization to eusociality. Hermaphroditism and parthenogenesis, which have evolved in some terrestrial crayfish burrowers and invasive open water crayfish, respectively, may enable populations to adapt to restrictive or new environments by spatio‐temporal alteration of their socio‐ecological characteristics. Under conditions of rapid habitat loss, environmental pollution and global warming, the reduced dispersal ability of direct developers may turn into a severe disadvantage, posing a higher threat of extinction to freshwater crayfish, primary freshwater crabs, aeglids and landlocked freshwater shrimps as compared to amphidromous freshwater shrimps and secondary freshwater crabs.     

Physiological evolution in the branchiopods

The interactions of physiological and morphological evolution are discussed in relation to the branchiopod crustaceans. Many branchiopods are morphologically relatively primitive and evade competition from more advanced forms by physiological adaptations which enable them to survive in extreme environments. The most recently evolved group of branchiopods, the Cladocera, or water fleas, have specialisation of morphology and reproduction which have enabled them to return to less extreme freshwater environments. Most recently they have developed physiological adptations which enable them to exploit the marine and other saline environments.     

Morphology of the Bivalve Mollusks Crenomytilus grayanus and Mytilus coruscus in Relation to Their Spatial Distribution in the Upper Subtidal Zone

The morphology of the shell and byssus threads was studied in two closely related mussel species Crenomytilus grayanus and Mytilus coruscus. The two species differ significantly from each other in the shell shape and in the degrees of development and deformation of byssus threads. These differences, in turn, determine (either directly or indirectly) the differences in strength of the byssal attachment and are discussed in terms of their functional morphology with respect to the spatial distribution of the mussels in marine coastal zones.     

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.     

Aspects of the phylogeny of the marine Tubificidae

A tentative phylogeny of the oligochaete family Tubificidae, with emphasis on the marine representatives, is presented. The scheme is based on the morphology and arrangements of prostate glands and the setal patterns. The rhyacodriline, more or less diffuse prostates are regarded as a primitive stage in prostate evolution, preceded only by the aprostate condition assumed for the ancestor of the family. An early split of the subfamily Rhyacodrilinae supposedly led to (1) a marine branch, from which evolved the highly diverse, exclusively marine subfamilies Phallodrilinae and Limnodriloidinae, and (2) a freshwater branch, which later divided into the Telmatodrilinae, Tubificinae and Aulodrilinae. The marine subfamilies invariably lack hair setae, whereas about half of the species within the other, freshwater subfamilies possess such setae in their dorsal bundles. Some marine genera, such as Monopylephorus (Rhyacodrilinae), Tubificoides and Clitellio (both Tubificinae) are regarded as recent off-shoots from the main freshwater stock.The families Naididae and Opistocystidae are considered likely to have evolved from rhyacodriline Tubificidae, whereas Phreodrilidae, the fourth family within the suborder Tubificina, is regarded as a sister group to the Tubificidae.     

Zebra mussel adhesion: structure of the byssal adhesive apparatus in the freshwater mussel, Dreissena polymorpha

The freshwater zebra mussel (Dreissena polymorpha) owes a large part of its success as an invasive species to its ability to attach to a wide variety of substrates. As in marine mussels, this attachment is achieved by a proteinaceous byssus, a series of threads joined at a stem that connect the mussel to adhesive plaques secreted onto the substrate. Although the zebra mussel byssus is superficially similar to marine mussels, significant structural and compositional differences suggest that further investigation of the adhesion mechanisms in this freshwater species is warranted. Here we present an ultrastructural examination of the zebra mussel byssus, with emphasis on interfaces that are critical to its adhesive function. By examining the attached plaques, we show that adhesion is mediated by a uniform electron dense layer on the underside of the plaque. This layer is only 10-20 nm thick and makes direct and continuous contact with the substrate. The plaque itself is fibrous, and curiously can exhibit either a dense or porous morphology. In zebra mussels, a graded interface between the animal and the substrate mussels is achieved by interdigitation of uniform threads with the stem, in contrast to marine mussels, where the threads themselves are non-uniform. Our observations of several novel aspects of zebra mussel byssal ultrastructure may have important implications not only for preventing biofouling by the zebra mussel, but for the development of new bioadhesives as well.     

The Byssus and Byssus Glands in Chlamys islandica and Other Scallops (Lamellibranchia)

The foot has at its proximal end, a pouch opening by a duct onto the sole of the foot containing 25–40 thin lamellae. The primary byssus gland secretes into the spaces between these lamellae. A secondary byssus gland secretes into a groove which runs forwards from the duct opening for about two-thirds of the length of the foot. Polyphenoloxidase is produced by cells surrounding the distal end of this groove. The tip attachment gland secretes into the distal fissure at the tip of the foot. The sole of the foot and the neck of the duct are lined with mucus cells. Ribbons of byssus are produced by the primary byssus gland. These are spun out in the groove and attached directly to the substratum at the distal end of the groove in an area presmeared by the tip attachment gland. The ribbon is enveloped in the groove by a sheath produced by the secondary byssus gland. Subsequent ribbons are bound to the existing byssus by a new sheath. Both the tip attachment gland and the ribbons contain some phenolic protein which is probably tanned in the presence of polyphenoloxidase to form a bond with the substratum. The homologies between this system and that in Mytilidae is discussed.     

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.     

Functional anatomy of the tarsier foot

Tarsiers possess a very odd musculoskeletal foot anatomy that goes beyond their acknowledged specialized leaping adaptations. Tarsius has evolved a fundamentally different method of bone rotation to achieve an inverted foot position during grasping and has developed an unusual muscular system for holding onto vertical supports. Although galagos and tarsiers possess elongated foot bones as adaptations for leaping, galagos utilize many more types of movements, have specialized osteological surface for inversion, and have a more common type of muscle development in the foot and leg than tarsiers possess. Likewise, the Omomyidae, the ancestral lineage of Tarsius, exhibit a lack of morphological similarity with Tarsius in the known foot joints.     

Convergent evolution of shell shape in freshwater limpets: the African genus Burnupia

Convergent evolution of shell shape is a well known phenomenon in gastropods that has caused much confusion in taxonomic and systematic studies. A paradigm is the patelliform shell shape in several taxa of the order Basommatophora. Historically, most freshwater limpets were assigned to the family Ancylidae. Based on anatomical data, some taxa were subsequently moved to different families. However, there are still doubts about the monophyly of the remaining ancylids. This is also true for the African limpets and particularly for species of the genus Burnupia Walker, 1912. In the present paper, two independent molecular markers (COI, 18S rRNA) are used to (a) infer the position of Burnupia within the Basommatophora and (b) to test whether the shell shape of Burnupia evolved independently. Bayesian and maximum likelihood analyses of 12 genera of limnic Basommatophora indicate that Burnupia not only appears to be distinct from the Ancylidae, but also from all other representatives of the superfamily Planorboidea studied here. Based on a generally well-supported phylogeny and preliminary anatomical data, it is concluded that patelliform shell shapes evolved at least three times in the taxa studied here and that the shell shape of Burnupia represents another case of convergent evolution. Previously proposed functional and evolutionary scenarios for the evolution of patelliform shell and their relevance for freshwater limpets are discussed.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 577–586.     

A morphometric study on morphological plasticity of shell form in crevice-dwelling Pterioida (Bivalvia)

Relative warp analysis and eigenshape analysis were used to explore the patterns of variation of erratic shell shapes in and among six species of crevice-dwelling pterioid bivalves. The results of morphometric analyses revealed that a great deal of the variance of shell shape within each species can be reduced into principal components which display patterns of variation common to all species examined. In the species with striking variability, a few sets of principal components account for most of the variance of shell shape. On the other hand, in the species lacking considerable variability, several components contribute to form a given variety. Comparison of relative warp and eigenshape scores with centroid sizes indicates that the direction of ontogenetic shape change has been modified through evolution to produce adaptations to habitats such as crevices or the undersides of rocks.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 285–297.     

The mechanism of shell elevation in Haliotis (Mollusca: Gastropoda) and a consideration of the evolution of the hydrostatic skeleton in Mollusca

Early in molluscan evolution, the development of a conical shell with shell or pedal retractor muscles led to the need of a mechanism for the extension of the foot or the raising of the shell. The forces generated during pedal retraction and extension have been studied in Haliotis midae , an easily obtainable and conveniently large archaeogastropod. In the mantle cavity, cephalopedal venous sinus and ventricle pressure pulses were observed during pedal retraction elicited by the shadow withdrawal reflex, but were never present during extension. However, pressure pulses were recorded in the proximal region of the columellar (or shell) muscle, both during retraction and pedal extension. Sections of this region of the muscle show a three dimensional network of muscle fibres, consisting of retractor fibres passing down to the foot and circumferential and radial fibres. Contraction of the two latter sets of fibres would bring about extension of the retractors, without the use of a discrete hydrostatic skeleton, and appears to be the principal mechanism of pedal extension. Similar muscular structures, here termed the muscular antagonistic system, have been observed in the columellar muscle of other gastropods and in the cephalopod mantle. In contrast, this system has not been observed in the proximal region of the pedal retractors of bivalves or scaphopods, for the pedal haemocoel, which allows muscular antagonism in the manner of a classical hydrostatic skeleton, has developed in association with the burrowing habit. The significance of the muscular antagonistic system in molluscan evolution is discussed.     

Evolution of the molluscan body plan: the case of the anterior adductor muscle of bivalves

The separated shell plates with the rearranged musculature (adductor muscle) is a novelty for bivalves. Despite its importance in the bivalve bodyplan, the development of the anterior adductor muscle remains unresolved. In this study, we investigate the myogenesis of the bivalve species Septifer virgatus to reveal the developmental origin of the larval muscles in bivalves, focusing on the anterior adductor muscle. We observed that larval retractor muscles are differentiated from the ectomesoderm in bivalves, and that the anterior adductor muscles are derived from primordial larval retractor muscles via segregation of the myoblast during the veliger larval stage. Through the comparative study of myogenesis in bivalves and its related taxa, gastropods, we found that both species possess myoblasts that emerge bilaterally and later meet dorsally. We hypothesize that these myoblasts, which are a major component of the main larval retractor in limpets, are homologous to the anterior adductor muscle in bivalves. These observations imply that the anterior adductor muscle of bivalves evolved as a novel muscle by modifying the attachment sites of an existing muscle.     

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.     

Impact of habitat and life trait on character evolution of pallial eyes in Pectinidae (Mollusca: bivalvia)

Pectinidae, a large group of marine bivalves comprising more than 300 species worldwide, inhabit a diverse array of habitats, enabling an enormous radiation, and yielding many different life forms and adaptations. This apparent diversity led to the distinction of ecotypes based on shell morphology and lifestyle. Eyes in Pectinidae (Bivalvia, Pteriomorphia) have long sparked scientific interest and have been described for various species over the past two centuries. These eyes are morphologically and functionally highly complex. Despite this complexity, studies have focused mostly on functional aspects with only few examining the relationships associated with different environmental or evolutionary traits. Here, the pallial eye structure within the Pectinidae was examined using Masson Goldner Trichrom staining, and ancestral character estimation with BayesTraits was performed to reconstruct macro-evolutionary patterns. To evaluate the connection of substrate type and lifestyle to the evolution of eyes, we compared eyes within the major subgroups of Pectinidae while considering the different lifestyles and substrate types as well as different depth ranges. The results indicate a tendency towards a taxon-/clade-specific evolution in respect to characters such as the cornea and lens while depth specific adaptations occur mainly in the light sensitive compartments of the retina. Successive reduction of eyes seems to occur from shallow to deep water species and ends in a total reduction of all structures in deep sea species.     

Burrowing mechanisms and sculptures in Recent gastropods

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