Tube construction in the watering pot shell Brechites vaginiferus (Bivalvia; Anomalodesmata; Clavagelloidea)

The bizarre watering pot shells of the clavagellid bivalve Brechites comprise a calcareous tube encrusted frequently with sand grains and other debris, the anterior end of which terminates in a convex perforated plate (the ‘watering pot’). It has not proved easy to understand how such extreme morphologies are produced. Previously published models have proposed that the tube and ‘watering pot’ are formed separately, outside the periostracum, and fuse later. Here we present the results of a detailed study of the structure and repair of the tubes of Brechites vaginiferus which suggest that these models are not correct. Critical observations include the fact that the external surface of the tube and ‘watering pot’ are covered by a thin organic film, on to the inner surface of which the highly organized aragonite crystals are secreted. There is no evidence of a suture between the tube and the ‘watering pot’ or that the periostracum of the juvenile shell passes through the wall of the tube. Live individuals of B. vaginiferus are able to repair substantial holes in the tube or ‘watering pot’ by laying down a new organic film followed by subsequent calcareous layers. Brechites vaginiferus displays Type C mantle fusion, with the result that the whole animal is encased by a continuous ring of mantle and periostracum, thereby making it possible to secrete a continuous ‘ring’ of shell material. On the basis of these observations we suggest that watering pot shells are not extra‐periostracal but are the product of simple modification of ‘normal’ shell‐secreting mechanisms.     

Reconstructing the Anomalodesmata (Mollusca: Bivalvia): morphology and molecules

The extant anomalodesmatan bivalves have always proved rather enigmatic and difficult to interpret, both in terms of their relationships to other bivalve taxa and the interrelationships of the constituent families. These difficulties stem from their diverse and often highly specialized life habits which have resulted in a wide array of disparate morphologies, and also from the fact that many are extremely rare. Classifications based on morphological characters have been dogged by fears that convergent and parallel evolution has masked phylogenetic signals. Molecular surveys of members of 12 of the 15 constituent families, using the 18S rRNA gene, have revealed that anomalodesmatans are robustly monophyletic and lie within the basal heterodonts. The Anomalodesmata should no longer be regarded as a subclass, but as a part of the Heterodonta. Here we present an enhanced analysis of 32 anomalodesmatan species (representatives of 12 families). Our results, subjected to Maximum Parsimony, Maximum Likelihood and Bayesian analyses, challenge our understanding of the internal relationships within the Anomalodesmata. In particular they indicate the need for a re-distribution of the families traditionally placed in the Thracioidea and Pandoroidea into a 'thraciid' lineage (Thraciidae + Cleidothaeridae + Myochamidae) and a 'lyonsiid' lineage (polyphyletic Lyonsiidae + Clavagellidae + Laternulidae + Pandoridae). The endolithic Clavagella and endobenthic Brechites and Penicillus form a robust clade. The hypothesis that the carnivorous septibranchs are monophyletic can, thus far, be neither supported nor rejected. Mapping critical morphological characters onto our molecular results provides evidence of multiple loss of some characters (e.g. prismato-nacreous shell microstructure and shell spicules) and also multiple gain of others (e.g. chondrophores).  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 395–420.     

Arenophilic mantle glands in the Laternulidae (Bivalvia: Anomalodesmata) and their evolutionary significance

The mantle margins of several anomalodesmatans bear multicellular arenophilic glands, the mucoid secretions of which attach sand grains and other foreign particles to the outer surface of the periostracum. These glands have been recorded for many of the anomalodesmatan families and are used as a key morphological character in recent attempts to unravel the evolutionary relationships within the Anomalodesmata. The glands occur in Laternula elliptica, L. truncata, L. boschasina and L. marilina, discharging from the top of muscular papillae at the distal tip of the siphons. The secretions are laid down as threads organized in longitudinal lines along the length of the periostracum that covers the siphonal walls. This is the first record of arenophilic mantle glands in members of the Laternulidae, a finding that not only broadens our current knowledge of the family's morphology, but assists in the reconstruction of anomalodesmatan evolutionary history.     

Spermatozoa of the Anomalodesmata (Bivalvia, Mollusca) with special reference to relationships within the group

Sperm ultrastructure in the marine bivalve order Anomalodesmata is considered in the light of new information for Australian Myochama anomioides and Cuspidaria latesulcata . In M. anomioides , the acrosomal complex lies posterior to the nucleus, in contact with the asymmetrical midpiece mitochondria – an unusual configuration reported from most of the studied Anomalodesmata. Spermatozoa of M. anomioides resemble those of Myadora brevis (both in family Myochamidae). Myochamid spermatozoa are considerably less modified than those of the Lyonsiidae or Laternulidae, and may prove a basal type within the non-septibranch anomalodesmatans. In contrast, C. latesulcata differs from other examined anomalodesmatans in having an anterior acrosomal complex and radial midpiece mitochondria (classic aquasperm features). Sperm data for the Anomalodesmata are limited but congruent with the most recent phylogenetic analyses that recognize distinct 'thraciid' and 'lyonsiid' clades. Results for C. latesulcata suggest septibranch origins before the development of a posteriorly positioned acrosomal complex and mitochondrial asymmetry.     

Discovery and anatomy of the arenophilic system of cuspidariid clams (Bivalvia: Anomalodesmata)

The arenophilic system of anomalodesmatan clams comprises multicellular glands, muscular papillae, and radial lines of adhesive secretion, which glue sand grains and other extraneous particles to the external surface of the shell. The presence and morphology of these organs have been established in an increasing number of taxa, but arenophilic glands were not previously known to occur in the deep‐sea, carnivorous family Cuspidariidae. Dissections, histological sections and ultrastructural examination of members of the cuspidariid genera Cuspidaria and Cardiomya revealed a functional arenophilic system positioned around the siphonal apparatus. Glands are contained in muscular papillae at the posterior tip of the siphonal sheath, from where they deposit their adhesive secretion onto the shell rostrum. Discovery of an arenophilic system in cuspidariid clams adds support to the idea that the glands are a synapomorphy of Anomalodesmata, secondarily absent in only one of the major component branches (the clade joining Thraciidae, Cleidothaeridae, and Myochamidae). Moreover, similarity between the arenophilic papillae we observed on the siphonal sheath of Cuspidaria, Cardiomya, and Myonera, and literature reports on the crown of tentacles of Multitentacula is puzzling and highlights the need for further studies of the latter genus. J. Morphol. 275:9–16, 2014. © 2013 Wiley Periodicals, Inc.     

Molecular phylogeny of Anomalodesmata (Mollusca: Bivalvia) inferred from 18S rRNA sequences

The origin of the anomalodesmatan bivalves and the relationships of the constituent families are far from being settled. Phylogenetic uncertainties result from the morphological heterogeneity of the Anomalodesmata and from parallel/convergent evolution of several character complexes due to similar life habits. Here, we assess these problems with 26 near-complete anomalodesmatan 18S rRNA sequences from 12 out of 15 families and a selection of heteroconch outgroup taxa. The robustly monophyletic Anomalodesmata share insertions in the V2 and V4 expansion regions. Both parsimony and maximum-likelihood analyses confirm their position among the basal heterodonts rooting between Carditidae and Lucinidae or, together with the latter, between Carditidae and the remaining Heterodonta. There is no support for monophyletic Myoida, nor for a close relationship of Anomalodesmata with any myoid taxon. At the base of the Anomalodesmata is an unstable cluster of long-branch species belonging to the Poromyidae, Verticordiidae, Lyonsiellidae and Thraciidae. The remaining Anomalodesmata split consistently but with varying branch support into three major clades: the Cuspidariidae excluding Myonera ; a 'thraciid' clade consisting of (Euciroidae, ( Myonera ( Thracia, Cleidothaerus , Myochamidae))); and a 'lyonsiid' clade with Laternulidae, Pandoridae, diphyletic Lyonsiidae due to a robust clade of Lyonsia norwegica and the clavagellid Brechites vaginiferus . Tests of various alternative topologies showed that all are significantly longer but optimal likelihood trees with monophyletic carnivorous taxa and/or Thraciidae are not significantly less likely. These results differ greatly from previous morphological studies. Palaeontological data and homology decisions for selected characters are evaluated in the light of the molecular trees.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 229–246.     

The fine structure of the pallial eyes of Laternula truncata (Bivalvia: Anomalodesmata: Pandoracea)

The structure of the pallial eyes of Laternula truncata (Lamarck 1818) has been studied using the light and electron microscopes. The eye is complex and can be- considered to be- the most advanced yet described for a bivalve mollusc. The cornea consists of modified flattened epithelial cells with an external border of microvilli. The cornea covers a large, circular, multinucleate lens. The lens comprises (1) centrally located translucent lens cells, (2) laterally located supporting cells from which cell processes interdigitate with processes from the lens cells. The retina is two layered and inverted. The proximal and distal retinae are made up of concentrically arranged laminae derived from the membranes of ciliary basal bodies. The cilia comprise a base and feet, but no root system and have a 9+0 arrangement of filaments.
The pigment cup or tapetum is bounded by a sclerotic coat and is three layered, each layer possessing characteristic pigment granules. From the base of the eye arises a large optic nerve.
The eye possesses an eye appendage, the epithelium of which is invaginated on its internal border to form a groove within which are found some 28 cilia. The cilia, it is thought, make contact with the microvilli of the epithelium when the appendage is touched. Such an action serves to protect the delicate eye from damage.
The structure of the eye is compared with that of other molluscs, particularly members of the Bivalvia.     

Structure and formation of the adventitious tube of the Japanese watering-pot shell Stirpulina ramosa (Bivalvia, Anomalodesmata, Clavagellidae) and a comparison with that of the Penicillidae

Abstract. Stirpulina ramosa is the only extant endobenthic representative of the Clavagellidae and is restricted to the waters of Japan. A single intact adventitious tube of this species has been obtained and its structure is described. The right valve is 16 mm long and located within the adventitious tube. It has an opisthodetic ligament located on resilifers. There are anterior and posterior adductor muscle scars, a thick pallial line, and pallial and pedal gape (right valve only) sinuses. The left shell valve is but 9 mm long and is united into the fabric of the adventitious tube via the intermediary of a shelly saddle. Internally, only the anterior adductor muscle scar and a small element of the pallial line scar are identifiable on the left valve. The posterior adductor and the rest of the pallial line scar (including a pallial sinus) are, remarkably, located on the adventitious tube beyond the shell valve margin. The adventitious tube of S. ramosa is formed in a manner wholly dissimilar from that of Brechites vaginiferus (Penicillidae). In B. vaginiferus, the tube is secreted as a single entity from the general outer mantle surface, including the siphons, covering the body. As a consequence, both shell valves are incorporated into the structure of the tube and the watering pot is bilaterally symmetrical. In S. ramosa, the tube and watering pot are secreted from the mantle margin and surface surrounding and extending from the left shell valve, so that only the left valve is incorporated into its structure. A dorsally derived mantle element is progressively extended over to the right side of the body, meeting a ventrally derived counterpart that passes beneath it, forming a pleat in the calcareous structure of the right side of the tube that they secrete. This pleat extends into the complex of watering‐pot tubules and forms the pedal gape. The watering pot is thus Ω shaped. The ventrally derived mantle element forms a sinusoidal crest on the right‐hand base of the watering pot, creating a pedal gape sinus scar on the right valve. The Clavagellidae radiated widely in the Mesozoic, leaving behind a rich fossil record for Stirpulina. Only S. ramosa, however, has survived until the present. In contrast, the Cenozoic Penicillidae has a poor fossil record, but there is a rich variety of extant endobenthic watering‐pot shells. It has been argued hitherto that the two families represent a remarkable example of convergent evolution. In view of the success of the Penicillidae and thus the endobenthic, tube‐dwelling lifestyle, however, it is hard to understand why Stirpulina has largely died out—even S. ramosa being known by but one or two specimens. A study of the anatomy of S. ramosa might one day answer this question.     

Teranota n. gen. (Bivalvia; Anomalodesmata) aus dem Mittel-Devon (Givetium) des Rheinischen Schiefergebirges

From the Rhenish Middle Devonian (Givetian) a new genus of posteriorly extremely elongated, endobenthonic bivalves preserved in life orientation is described. The phylogenetic relationships within the superorder Anomalodesmata are discussed.     

The ecology and functional morphology of Trigonothracia jinxingae (Bivalvia: Anomalodesmata: Thracioidea) from Xiamen, China

All anomalodesmatans are 'rare' but Trigonothrucia jinxingae is relatively common in Xiamen Harbour, Fujian Province, China. This is because the species has a life span of approximately one year and is a simultaneous hermaphrodite, probably with either a short or absent planktonic larval stage. That is, success results from rapid maturation, self-fertilization, direct development and within-habitat recruitment over an extended period in early summer.
Trigonothracia jinxingae is interesting in another way, however. The Thraciidae is the Mesozoic stem group of the Thracioidea which also contains the more modern (Caenozoic) Laternulidae and Periplomatidae. Features of the anatomy of T. jinxingae , such as the method of hydraulically moving the foot by the pumping of blood into a capacious pallial haemocoel, and the structure of the stomach, are reminiscent of the earliest (Palaeozoic) anomalodesmaans, i.e. the Pholadomyoidea, represented today by Pholadomya candida. The thraciid Asthenothaerus sp. (Pelseneer, 1911) even has, like P. cundida , an opisthopodium on its visceral mass. P. candida , however, fed on sub-surface deposits using its foot. T. jinxingae is also a deposit feeder, but on surface deposits using the inhalant siphon.
Modern periplomatids resemble thraciids in their separate siphons, but both representatives of this family and the Laternulidae are suspension feeders with extensive sorting areas on the wall of the stomach to process such material. The Thraciidae thus form a link between the oldest, pedal feeding, pholadomyoidean anomalodesmatan and the most advanced, suspension feeding, laternulids and periplomatids.