Soft‐tissue imprints in fossil and Recent cephalopod septa and septum formation

Several soft‐tissue imprints and attachment sites have been discovered on the inside of the shell wall and on the apertural side of the septum of various fossil and Recent ectocochleate cephalopods. In addition to the scars of the cephalic retractors, steinkerns of the body chambers of bactritoids and some ammonoids from the Moroccan and the German Emsian (Early Devonian) display various kinds of striations; some of these striations are restricted to the mural part of the septum, some start at the suture and terminate at the anterior limit of the annular elevation. Several of these features were also discovered in specimens of Mesozoic and Recent nautilids. These structures are here interpreted as imprints of muscle fibre bundles of the posterior and especially the septal mantle, blood vessels as well as the septal furrow. Most of these structures were not found in ammonoids younger than Middle Devonian. We suggest that newly formed, not yet mineralized (or only slightly), septa were more tightly stayed between the more numerous lobes and saddles in more strongly folded septa of more derived ammonoids and that the higher tension in these septa did not permit soft‐parts to leave imprints on the organic preseptum. It is conceivable that this permitted more derived ammonoids to replace the chamber liquid faster by gas and consequently, new chambers could be used earlier than in other ectocochleate cephalopods, perhaps this process began even prior to mineralization. This would have allowed faster growth rates in derived ammonoids.     

Structure of shell muscles in the Cambrian gastropod genus Bemella (Gastropoda: Archaeobranchia: Helcionellidae)

Zones with peculiar microornamentation interpreted as muscle scars were found on the internal molds of the Early Cambrian gastropod genus Bemella Missarzhevsky, 1969 (family Helcionellidae). Shell muscles of helcionellids are reconstructed based on the topographic pattern of muscle scars, i.e., the pedal, cephalic, and mantle retractors are recognized. The reconstruction proposed here of the shell musculature corroborates affinity between ancient gastropod and helcionelloid mollusks.     

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.     

Shell Structure And Inferred Growth, Functions And Affinities Of The Sclerites Of The Problematic Micrina

The stratiform laminae of Micrina sclerites originally consisted of rheomorphic successions of monolayers of micrometric–sized, apatitic tablets, presumably interleaved with chitin and glycosaminoglycans (GAGs). Paired laminae enclose slot–like chambers swelling into lobes distally that originally contained GAGs and deposits of spherulitic and prismatic apatite. The laminae are pervaded by apatitic tubes, apparently secreted by microvillous setoblasts and containing, at the surface, chitinous setae. Internal markings suggest that the triangular (sellate) sclerite supported a pair of muscles and the planospiral (mitral) sclerite, a medial muscle and gonadal sacs flanked by a pair of crescentic muscle bases. Both sclerites were secreted by a mantle with a circumferential fold. The sellate and mitral sclerites are homologized with the anterior and posterior shells of Halkieria and could have become the dorsal and ventral valves of the ancestral brachiopod by a sequence of transformations. These include: the folding of the halkieriid body axis; accelerated mixoperipheral growth of the anterior (dorsal) shell to enclose, with the posterior (ventral) shell, a mantle cavity lined with modified ciliated epithelium of the foot; reduction of sclerite–secreting epithelium to the locus of the brachiopod pedicle epithelium; and the anterior (dorsal) spread of gonadal lamellae.     

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.     

Modern insights on gastropod development: Reevaluation of the evolution of a novel body plan

More than a century of speculation about the evolutionary originof the contorted gastropod body plan has been inspired by adultanatomy and by long-standing developmental observations. Theresult has been a concept of gastropod torsion that I call the"rotation hypothesis." Under the rotation hypothesis, gastropodsoriginated when all components of the visceropallium (shell,mantle, mantle cavity with contained structures, and viscera)rotated by 180° relative to the head and foot. This evolutionaryrotation is echoed during early development of patellogastropodsand vetigastropods and occurs to some extent during developmentof more derived clades. However, comparative developmental dataon ontogenetic torsion are minimal and I argue that the rotationhypothesis is a tautological argument. More recent studies onrepresentatives from 3 major clades of gastropods suggest thatthe highly conserved aspect of gastropod development is notsynchronous rotation of all components of the visceropalliumrelative to the head and foot but rather a state of anatomicalorganization in which the developing mantle cavity is on theright but the shell coil is posterior (endogastric orientation).This conserved state of developmental anatomy has inspired analternative hypothesis for the evolutionary origin of the gastropodbody plan, the "asymmetry hypothesis." Under the asymmetry hypothesis,the gastropod mantle cavity originated from 1 side only of abilateral set of mantle cavities. The asymmetry hypothesis doesnot require a saltation event to explain the origin of gastropods,nor does it require that the ancient molluscan precursor ofgastropods carried the shell coil over the head (exogastricorientation).     

An electromyographic analysis of the biting mechanism of the lemon shark,Negaprion Brevirostris: Functional and evolutionary implications

The kinetics of the head and function of select jaw muscles were studied during biting behavior in the lemon shark, Negaprion brevirostris. High speed cinematography and electromyography of seven cranial muscles were recorded during bites elicited by a probe to the oral cavity. In weak bites mandible depression was followed by mandible elevation and jaw closure without cranial elevation. In strong bites cranial elevation always preceded lower jaw depression, lower jaw elevation, and cranial depression. The average duration of the strong bites was rapid (176 msec), considering the size of the animal relative to other fishes. Different electromyographic patterns distinguished the two forms of bite, primarily in activity of the epaxial muscles, which effect cranial elevation. A composite reconstruction of the activity of seven cranial muscles during biting revealed that epaxial muscle activity and consequently cranial elevation preceded all other muscle activity. Mandible depression was primarily effected by contraction of the common coracoarcual and coracomandibularis, with assistance by the coracohyoideus. Simultaneous activity of the levator hyomandibulae is believed to increase the width of the orobranchial chamber. The adductor mandibulae dorsal was the primary jaw adductor assisted by the adductor mandibulae ventral. This biomechanically conservative mechanism for jaw opening in aquatic vertebrates is conserved, with the exception of the coracomandibularis, which is homologous to prehyoid muscles of salamanders.     

How to recognize in situ fossil cephalopods: evidence from experiments with modern Nautilus

Field and flume experiments with modern Nautilus pompilius establish two prerequisites to recognize in situ preservation of fossil cephalopod shells (soft parts were within body chamber in situ at the time of fossilization): occurrence of the upper jaw within the body chamber and the position of jaws within the body chamber. Morphology of shells and jaws in modern and fossil nautiloids is so similar that these prerequisites can be applied for fossil nautiloids and provide implications for ammonoids. The upper jaws of Nautilus start to move at a water velocity of > 0.2 m/s, when the shells are reoriented with the aperture downstream; jaws are therefore unlikely to be secondarily deposited near the shell aperture by bottom currents. The lower jaws, moved at the velocity of > 0.1 m/s, can be deposited around the shell aperture by weak current (0.1–0.2 m/s in velocity), but never enter the inside of body chamber. Neither jaw is likely to be separately and selectively displaced from the inside of the body chamber through scavenging of the soft parts by burrowing infaunal animals. An upper jaw preserved inside the body chamber, together with a lower jaw, is thus a reliable indicator of in situ preservation; a sole lower jaw preserved around the shell aperture is likely to be secondarily deposited. Sedimentary structures inferring rapid burial events and jaw size are useful as additional evidence. Smaller jaws were more likely to be displaced from the body chamber by scavenging by infaunal animals after in situ burial, so that smaller jaws preserved within the body chamber suggest less scavenging. These findings are crucial to interpreting the taphonomic history and palaeo-ecology of fossil cephalopods.     

The biology and functional morphology of Arca noae (Bivalvia: Arcidae) from the Adriatic Sea, Croatia, with a discussion on the evolution of the bivalve mantle margin

In the Croatian Adriatic, Arca noae occurs from the low intertidal to a depth of 60 m; it can live for > 15 years and is either solitary or forms byssally attached clumps with Modiolus barbatus. The shell is anteriorly foreshortened and posteriorly elongate. The major inhalant flow is from the posterior although a remnant anterior stream is retained. There are no anterior but huge posterior byssal retractor muscles and both anterior and posterior pedal retractors. The ctenidia are of Type B(1a) and the ctenidial–labial palp junction is Category 3. The ctenidia collect, filter and undertake the primary sorting of potential food in the inhalant water. The labial palps are small with simple re‐sorting tracks on the ridges of their inner surfaces. The ciliary currents of the mantle cavity appear largely concerned with the rejection of particulate material. The mantle margin comprises an outer and an (either) inner or middle fold. The outer fold is divided into outer and inner components that secrete the shell and are photo‐sensory, respectively. The latter bears a large number of pallial eyes, especially posteriorly. The inner/middle mantle fold of A. noae, possibly representative of simpler, more primitive conditions, may have differentiated into distinct folds in other recent representatives of the Bivalvia.     

Notes on the phylogeny of the nautiloidea

Critical notes on the monograph “Phylogeny of the Nautiloidea” by J. Dzik (1984), especially with respect to cephalopod material from the Lower Palaeozoic of Bohemia are presented. Oncocerid type of muscle scars in the Lower Devonian genusPtenoceras as well as succession of growth stages of the shell in Silurian genusSphooceras is figured.     

Chamber growth in ammonites inferred from colour markings and naturally etched surfaces of Cretaceous vascoceratids from Nigeria

Cretaceous Vascoceras and Jurassic Lytoceras show colour markings and etched surfaces representing original organic membranes between the septa. The main difference between the formation of ammonite and Nautilus chambers involved the continuous secretion of a gelatinous cameral liquid to support the ammonite mantle when it moved forward. The gel containing cyclically secreted membranes. here named pseudosepta, resembled the intra-cameral structures of the cuttlebone in Sepia. Pseudoscpta are attached to the shell wall in pseudosutures (Pseudoloben) which are particularly visible in the saddles of the septal suture and tend to mimic them. Their shape suggests reconstruction of posterior mantle shape during translocation. Drag-bands (Schleppstreifen) are spiral markings formed by the overlapping pseudosepta along the axial traces of the foliole folds. The chamber of ammonites was formed by a locally muscular mantle in a tripartite cycle: (1) the mantle initially remained attached to the saddles of the completed septal suture while muscular tissue within the umbilical lobes was contracted and rapidly reattached to the side of the lateral saddles; (2) the whole mantle subsequently crept forward by secreting a gelatinous matrix which contained telescoped membranes, with an adhesive function on pseudolobc flanks; (3) the mantle almost ceased to move within the sites of future lobules, but expanded and crept on before forming the mural and 'gutter' ridges of the septum. □ Ammonites, chamber growth, vascoceratids, LYTOCERAS, Nigeria.     

Siphonariid development: Quintessential euthyneuran larva with a mantle fold innovation (Gastropoda; Panpulmonata)

Recent phylogenetic revisions of euthyneuran gastropods (“opisthobranchs” and “pulmonates”) suggest that clades with a planktotrophic larva, the ancestral life history for euthyneurans, are more widely distributed along the trunk of the euthyneuran tree than previously realized. There is some indication that the planktotrophic larva of euthyneurans has distinctive features, but information to date has come mainly from traditional “opisthobranch” groups. Much less is known about planktotrophic “pulmonate” larvae. If planktotrophic larvae of “pulmonates” share unique traits with those of “opisthobranchs,” then a distinctive euthyneuran larval-type has been the developmental starting template for a spectacular amount of evolved morphological and ecological disparity among adult euthyneurans. We studied development of a siphonariid by preparing sections of larval and postmetamorphic stages for histological and ultrastructural analysis, together with 3D reconstructions and data from immunolabeling of the larval apical sensory organ. We also sought a developmental explanation for the unusual arrangement of shell-attached, dorso-ventral muscles relative to the mantle cavity of adult siphonariids. Adult siphonariids (“false limpets”) have a patelliform shell but their C-shaped shell muscle partially embraces a central mantle cavity, which is different from the arrangement of these components in patellogastropods (“true limpets”). It is not obvious how shell muscles extending into the foot become placed anterior to the mantle cavity during siphonariid development from a veliger larva. We found that planktotrophic larvae of Siphonaria denticulata are extremely similar to previously described, planktotrophic “opisthobranch” larvae. To emphasize this point, we update a list of distinctive characteristics of planktotrophic euthyneuran larvae, which can anchor future studies on the impressive evolvability of this larval-type. We also describe how premetamorphic and postmetamorphic morphogenesis of larval mantle fold tissue creates the unusual arrangement of shell-muscles and mantle cavity in siphonariids. This result adds to the known postmetamorphic evolutionary innovations involving mantle fold tissue among euthyneurans.     

Reconstruction of cranial and hyobranchial muscles in the triassic temnospondyl Gerrothorax provides evidence for akinetic suction feeding

The cranial and hyobranchial muscles of the Triassic temnospondyl Gerrothorax have been reconstructed based on direct evidence (spatial limitations, ossified muscle insertion sites on skull, mandible, and hyobranchium) and on phylogenetic reasoning (with extant basal actinopterygians and caudates as bracketing taxa). The skeletal and soft‐anatomical data allow the reconstruction of the feeding strike of this bottom‐dwelling, aquatic temnospondyl. The orientation of the muscle scars on the postglenoid area of the mandible indicates that the depressor mandibulae was indeed used for lowering the mandible and not to raise the skull as supposed previously and implies that the skull including the mandible must have been lifted off the ground during prey capture. It can thus be assumed that Gerrothorax raised the head toward the prey with the jaws still closed. Analogous to the bracketing taxa, subsequent mouth opening was caused by action of the strong epaxial muscles (further elevation of the head) and the depressor mandibulae and rectus cervicis (lowering of the mandible). During mouth opening, the action of the rectus cervicis muscle also rotated the hyobranchial apparatus ventrally and caudally, thus expanding the buccal cavity and causing the inflow of water with the prey through the mouth opening. The strongly developed depressor mandibulae and rectus cervicis, and the well ossified, large quadrate‐articular joint suggest that this action occurred rapidly and that powerful suction was generated. Also, the jaw adductors were well developed and enabled a rapid mouth closure. In contrast to extant caudate larvae and most extant actinopterygians (teleosts), no cranial kinesis was possible in the Gerrothorax skull, and therefore suction feeding was not as elaborate as in these extant forms. This reconstruction may guide future studies of feeding in extinct aquatic tetrapods with ossified hyobranchial apparatus. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

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.     

A paradox survival – report of a repaired syn vivo perforation in a nautiloid phragmocone

The nautiloid Trocholites depressus (Eichwald, 1840) from the Lasnamägi regional stage (Darrivillian, Middle Ordovician) of Vaike Pakri Island (North-West Estonia) is the only known ectocochleate cephalopod that survived and healed a perforation of the phragmocone. Two chambers of the specimen were broken during its lifetime. The injury is located on the venter of the conch directly above the peristomal opening of the body chamber. It is reconstructed that the peristomal mantle tissue carried out an initial sealing of the injured chambers. The complete calcified sealing and compensation of the irregular shell surface started late with the overgrowth of the septa of the preceding whorl. The position and diameter of the siphuncle were not disturbed by the regeneration showing that these characters allowed a low phenotypic variability. Despite the trauma in the midlife growth history of this specimen, it appears to have reached maturity.     

Comparative morphology of changeable skin papillae in octopus and cuttlefish

A major component of cephalopod adaptive camouflage behavior has rarely been studied: their ability to change the three‐dimensionality of their skin by morphing their malleable dermal papillae. Recent work has established that simple, conical papillae in cuttlefish (Sepia officinalis) function as muscular hydrostats; that is, the muscles that extend a papilla also provide its structural support. We used brightfield and scanning electron microscopy to investigate and compare the functional morphology of nine types of papillae of different shapes, sizes and complexity in six species: S. officinalis small dorsal papillae, Octopus vulgaris small dorsal and ventral eye papillae, Macrotritopus defilippi dorsal eye papillae, Abdopus aculeatus major mantle papillae, O. bimaculoides arm, minor mantle, and dorsal eye papillae, and S. apama face ridge papillae. Most papillae have two sets of muscles responsible for extension: circular dermal erector muscles arranged in a concentric pattern to lift the papilla away from the body surface and horizontal dermal erector muscles to pull the papilla's perimeter toward its core and determine shape. A third set of muscles, retractors, appears to be responsible for pulling a papilla's apex down toward the body surface while stretching out its base. Connective tissue infiltrated with mucopolysaccharides assists with structural support. S. apama face ridge papillae are different: the contraction of erector muscles perpendicular to the ridge causes overlying tissues to buckle. In this case, mucopolysaccharide‐rich connective tissue provides structural support. These six species possess changeable papillae that are diverse in size and shape, yet with one exception they share somewhat similar functional morphologies. Future research on papilla morphology, biomechanics and neural control in the many unexamined species of octopus and cuttlefish may uncover new principles of actuation in soft, flexible tissue. J. Morphol. 275:371–390, 2014. © 2013 Wiley Periodicals, Inc.     

Irregular calcareous sheets preserved in a conch of the Cretaceous ammonoid Hypophylloceras from Hokkaido,Japan

The mantle tissue is essential for understanding the diverse ecology and shell morphology of ammonoid cephalopods. Here, we report on irregular calcareous sheets in a well-preserved shell of a Late Cretaceous phylloceratid ammonoid Hypophylloceras subramosum from Hokkaido, Japan, and their significance for repairing the conch through the mantle inside the body chamber. The sheets are composed of nacreous layers arranged parallel to the irregularly distorted outer whorl surface. The nacreous sheets formed earlier are unevenly distributed and attached to the outer shell wall locally, whereas the last formed sheet covers a wide area of the outer shell wall. The absence of any interruption of ribbing around the irregular area suggests that these sheets were secreted inside the body chamber from the inner mantle. Gross morphological and X-ray computed tomography observations revealed that the spacing of septal formation was not affected by this event. The complex structure of the irregular sheets suggests a highly flexible mantle inside the body chamber.     

Development of the pedicle in the articulate brachiopod Terebratalia transversa (Brachiopoda,Terebratulida)

Summary The development of the pedicle in the articulate brachiopod Terebratalia transversa has been examined by electron microscopy. The posterior half of the free-swimming larva comprises a non-ciliated pedicle lobe that contains the primordium of the juvenile pedicle at its distal end. During settlement at five to six days post-fertilization, the pedicle lobe secretes a sticky sheet that attaches the larva to the substratum. As metamorphosis proceeds, the epithelium in the posterior half of the pedicle lobe produces a thin overlying cuticle, and the pedicle primordium develops into a stalk-like anchoring organ. The juvenile pedicle protrudes through the gape that occurs between the posterior margins of the shell valves. A cup-like canopy, called the pedicle capsule, lines the posterior end of the shell and surrounds the newly formed pedicle. The core of the juvenile pedicle is filled with a solid mass of connective tissue. Numerous tonofibrils occur in the pedicle epithelium, and the overlying cuticle consists of amorphous material covered by a thin granular fringe. By one year post-metamorphosis, a body cavity develops anterior to the pedicle. Two pairs of adjustor muscles extend from the posterior end of the shell and traverse the cavity to insert in the pedicle. The connective tissue core of the pedicle in sub-adult specimens lacks muscle cells but contains numerous fibroblasts and collagen fibers. Three regions are recognizable in the connective tissue compartment of the adult pedicle: a subepithelial layer of non-fibrous connective tissue, a central fibrous zone, and a proximal mass of tissue that resembles cartilage.List of abbreviations as adhesive sheet - bc body cavity - bv brachial valve of shell - cf collagen fibrils - ct connective tissue - cu cuticle - di diductor muscle - ec epithelial cell - f fibroblast - fz fibrous zone - g gut - gc granular cell - gd gastric diverticulum - ht hinge tooth - ia interarea of pedicle valve - icl inner cuticular layer - lo lophophore - lu lumen of gut - m mesenchyme - ma mantle - ml mantle lobe - ocl outer cuticular layer - p periostracum - pc pedicle capsule - pce pedicle capsule epithelium - pcl pedicle collar of shell - pcn pedicle connectives - pd pedicle - pe pedicle epithelium - pl pedicle lobe - pv pedicle valve of shell - pzc proximal zone of cartilage-like tissue - s substratum - sel subepithelial layer - t tendon - tf tonofibril - vam ventral adjustor muscle