Earliest ontogeny of the Silurian orthotetide brachiopod Coolinia and its significance for interpreting strophomenate phylogeny

Well‐preserved juvenile specimens of the orthotetide brachiopod Coolinia pecten (Linnaeus, 1758 ) from the Silurian of Gotland, Sweden, demonstrate evidence of a planktotrophic larval habit. Larval shell morphology indicates the absence of a pedicle sheath: this character is otherwise typical of derived billingsellides, strophomenides and productides, which form the conventional strophomenide clade. The presence of a rudimentary colleplax structure in the larval shell of Colinia suggests instead a phylogenetic link to chiliate brachiopods and the enigmatic genus Salanygolina. This relationship suggests an early divergence of rhynchonellate and strophomenate brachiopods.     

The protegulum and juvenile shell of a Recent articulate brachiopod: patterns of growth and chemical composition

Patterns of shell formation and the chemical composition of the shell deposited during early post-larval life were investigated in laboratory-reared cultures of the Recent articulate brachiopod Terebraralia transversa (Sowerby). A non-hinged protegulum averaging 148 pm in length is secreted by the mantle within a day after larval metamorphosis. The inner surface of the protegulum exhibits finely granular, non-fibrous material. A rudimentary periostracum constitutes the outer layer of the primordial shell. and concentrically arranged growth lines are lacking. By four days post-metamorphosis, a brephic type of juvenile shell develops from periodic additions of shell material to the anterior and lateral edges of the protegulum. Imbricated secondary fibers occur throughout the inner layer of the newly formed juvenile shell, and a rudimentary hinge apparatus is present posteriorly. The external surface of the shell exhibits concentric growth lines anterior to the caudally situated protegulum, and unbranched punctae begin to form in the subperiostracal region of the shell. At 23 days post-metamorphosis, the shell weighs an average of 1.7 μg and measures 318 μm in length. Electron microprobe analyses reveal that the protegulum is calcified. Minor amounts of sulfur, magnesium, iron, chlorine, aluminum, and silicon are also present in protegula and juvenile shells. Based on electron diffraction data, the mineral phase of juvenile shells consists of calcite, and protegula also appear to contain calcite.     

Shell microstructure of <Emphasis Type="Italic">Discinisca suborbicularis</Emphasis> sp. nov. (Brachiopoda,Lingulata) from the Upper Jurassic of Western Siberia

A new species of the phosphatic brachiopod Discinisca suborbicularis (family Discinidae) from the Upper Jurassic of Western Siberia (latitudinal Priob’e) is established. The shell microstructure under protegulum and brephic shell and the microstructure of adult shell (that corresponds to three developmental stages) are described in detail. The shell microstructure of new species considerably differs from Paleozoic discinids and is similar to Recent discinids. D. suborbicularis differs from recent discinids in the presence of protegulum and thicker organophosphatic primary layer.     

Protegulum and brephic shell of the earliest organophosphatic brachiopods

Early development stages imprinted on the shells of Cambrian brachiopods from the class Linguliformea (orders Paterinida, Lingulida, Acrotretida) were studied with scanning electron microscope based on a large collection from the Siberian Platform. Some specimens of all three orders preserved protegulum (embryonic shell); their brephic (juvenile) shells, also were studied. Many of them might lack larval development stage as it is known for the recent representatives of the family Lingulidae. But unlike recent lingulids, the surface of the juvenile dorsal valves of all studied linguliformeans bore two (rarely three) pairs of gentle elevations accommodating bundles of setae, which sometime preserve as groups of fine imprints along their margins. Recent Lingula and Glottidia lack setae in this stage. On the other hand, recent classes Craniformea and Rhynchonelliformea have setae, but they develop in the larval stage. Most of brachiopod groups have the setae in the adult stage but these setae have different origin, are short, located on the lateral and anterior ends of the shell, and grow from the marginal cells of the mantle.     

Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny

Popov, L.E., Bassett, M.G., Holmer, L.E., Skovsted, C.B. & Zuykov, M.A. 2010: Earliest ontogeny of Early Palaeozoic Craniiformea: implications for brachiopod phylogeny. Lethaia, Vol. 43, pp. 323–333. Well preserved specimens of the Early Palaeozoic craniiform brachiopods Orthisocrania and Craniops retain clear evidence of a lecithotrophic larval stage, indicating the loss of planktotrophy early in their phylogeny. The size of the earliest mineralized dorsal shell was <100 μm across, and the well preserved shell structure in these fossil craniiforms allows their earliest ontogeny to be compared directly with that of living Novocrania, in which the first mineralized dorsal shell (metamorphic shell) is secreted only after settlement of the lecithotrophic larvae. Immediately outside this earliest shell (early post‐metamorphic or brephic shell) and in the rest of the dorsal valve the primary layer in both fossil and living craniiforms has characteristic radially arranged laths, which are invariably lacking in the earliest dorsal shell. The ventral valve of the fossil specimens commonly preserves traces of an early attachment scar (cicatrix), which is equal in size to the dorsal metamorphic shell, and the brephic post‐metamorphic ventral valve also has a primary shell with radially arranged laths. However, a primary shell with radial laths is completely lacking in the ventral valve of living Novocrania, indicating that heterochrony may have been involved in the origin of the encrusting mode of life in living craniids; the entire ventral valve of Recent craniids (with the possible exception of Neoancistrocrania) may correspond to the earliest attachment scar of some fossil taxa such as Orthisocrania. It is also probable that the unique absence of an inner mantle lobe as well as the absence of lobate cells in Novocrania could be the result of heterochronic changes. The dorsal valve of both fossil and living craniiforms has a marked outer growth ring, around 500 μm across, marking the transition to the adult, and a significant change in regime of shell secretion. The earliest craniiform attachment is considered to be homologous to the unique attachment structures described recently in polytoechioids (e.g. Antigonambonites) and other members of the strophomenate clade. However, unlike the craniiforms, polytoechioids and strophomenates all have planktotrophic larvae, and planktotrophy is most probably a plesiomorphic character for all Brachiopoda. □Brachiopoda, Craniiformea, Early Palaeozoic, ontogeny, phylogeny.     

Chemico-structure of the organophosphatic shells of siphonotretide brachiopods

The organophosphatic shell of siphonotretide brachiopods is stratiform with orthodoxly secreted primary and secondary layers. The dominant apatitic constituents of the secondary layer are prismatic laths and rods arranged in monolayers (occasionally in cross-bladed successions), normally recrystallized as platy laminae. Sporadically distributed, interlaminar, lenticular chambers, containing apatitic meshes of laths and aggregates of plates and spherulites, probably represent degraded, localized exudations of glycosaminoglycans (GAGs) with dispersed apatite.
The shells of Helmersenia and Gorchakovia are perforated by canals with external depressions (antechambers) that possibly contained chitinous tubercles in vivo . The immature shell of Siphonotreta and most other siphonotretids is similarly perforated and pitted; but the mature part bears recumbent, rheomorphic, hollow spines that grew forward out of pits. Internally, spines pierce the shell as independent structures to terminate as pillars in GAGs chambers. Spines and pillars were probably secreted by collectives of specialized cells (acanthoblasts) within the mantle.
The shell of the oldest siphonotretide, Schizambon , is imperforate but the ventral valve has a pedicle foramen that lies forward of the posterior margin of the juvenile valve. This relationship characterizes all siphonotretides, suggesting that the pedicle, in vivo , originated within the ventral outer epithelium and not from the posterior body wall as in lingulides.     

The ontogeny of shell secretion in Terebratalia transversa (Brachiopoda, Articulata). II. Formation of the protegulum and juvenile shell

The fine structure of the shell and underlying mantle in young juveniles of the articulate brachiopod Terebratalia transversa has been examined by electron microscopy. The first shell produced by the mantle consists of a nonhinged protegulum that lacks concentric growth lines. The protegulum is secreted within a day after larval metamorphosis and typically measures 140-150 micron long. A thin organic periostracum constitutes the outer layer of the protegulum, and finely granular shell material occurs beneath the periostracum. Protegula resist digestion in sodium hypochlorite and are refractory to sectioning, suggesting that the subperiostracal portion of the primordial shell is mineralized. The juvenile shell at 4 days postmetamorphosis possesses incomplete sockets and rudimentary teeth that consist of nonfibrous material. The secondary layer occuring in the inner part of the juvenile shell contains imbricated fibers, whereas the outer portion of the shell comprises a bipartite periostracum and an underlying primary layer of nonfibrous shell. Deposition of the periostracum takes place within a slot that is situated between the so-called lobate and vesicular cells of the outer mantle lobe. Vesicular cells deposit the basal layer of the periostracum, while lobate cells contribute materials to the overlying periostracal superstructure. Cells with numerous tonofibrils and hemidesmosomes differentiate in the outer mantle epithelium at sites of muscle attachments, and unbranched punctae that surround mantle caeca develop throughout the subperiostracal portion of the shell. Three weeks after metamorphosis, the juvenile shell averages about 320 micron in length and is similar in ultrastructure to the shells secreted by adult articulates.     

ONTOGENETIC NICHE SHIFT IN THE BRACHIOPOD TEREBRATALIA TRANSVERSA: RELATIONSHIP BETWEEN THE LOSS OF ROTATION ABILITY AND ALLOMETRIC GROWTH

Abstract: Many articulated brachiopods experience marked life habit variations during ontogeny because they experience their fluid environment at successively higher Reynolds numbers, and they can change the configuration of their inhalant and exhalant flows as body size increases. We show that the extant brachiopod Terebratalia transversa undergoes a substantial ontogenetic change in reorientation governed by rotation around the pedicle. T. transversa′s reorientation angle (maximum ability to rotate on the pedicle) decreases during ontogeny, from 180 degrees in juveniles to 10–20 degrees in individuals exceeding 5 mm, to complete cessation of rotation in individuals larger than 10 mm. Rotation ability is substantially reduced after T. transversa achieves the adult lophophore configuration and preferred orientation with respect to ambient water currents at a length of 2.5–5 mm. We hypothesize that the rotation angle of T. transversa is determined mainly by the position of ventral and dorsal points of attachment of dorsal pedicle muscles relative to the pedicle. T. transversa shows a close correlation between the ontogenetic change in reorientation angle and ontogeny of morphological traits that are related to points of attachment of dorsal pedicle muscles, although other morphological features can also limit rotation in the adult stage. The major morphological change in cardinalia shape and the observed reduction of rotation affect individuals 2.5–10 mm in length. The position of ventral insertions of dorsal pedicle muscles remains constant, but contraction of dorsal pedicle muscles is functionally handicapped because dorsal insertions shift away from the valve midline, rise above the dorsal valve floor, and become limited by a wide cardinal process early in ontogeny (<5 mm). The rate of increase of cardinal process width and of distance between dorsal pedicle muscle scars substantially decreases in the subadult stage (5–10 mm), and most of the cardinalia shell traits grow nearly isometrically in the adult stage (>10 mm). T. transversa attains smaller shell length in crevices than on exposed substrates. The proportion of small‐sized individuals and population density is lower on exposed substrates than in crevices, indicating higher juvenile mortality on substrates prone to grazing and physical disturbance. The loss of reorientation ability can be a consequence of morphological changes that strengthen substrate attachment and maximize protection against biotic or physical disturbance (1) by minimizing torques around the pedicle axis and/or (2) by shifting energy investments into attachment strength at the expense of the cost involved in reorientation.     

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     

The transition from planktotrophy to lecithotrophy in larvae of Lower Palaeozoic Rhynchonelliform brachiopods

Criteria are established for defining the presence of protegula formed on embryonic or larval mantle in representative genera of Lower Palaeozoic Obolellata, Strophomenata and Rhynchonellata. Width was used to define protegular type. Taxa with only an embryonic protegulum are inferred to have had lecithotrophic larvae while taxa with a larval protegulum or an embryonic protegulum surrounded by a larval protegulum are inferred to have had planktotrophic larvae. All or most of the taxa examined in the Obolellata, the Strophomenata and the orders Protorothida and Orthida in the Rhynchonellata had planktotrophic larvae. In the Pentamerida a minority of genera had only a larval, or an embryonic and a larval protegulum while a majority had protegular widths indicating lecithotrophy. In the orders Rhynchonellida, Atrypida, Athyrida and Spiriferida derived from the Pentamerida (with the exception of one species in the Atrypida) a number of the genera had protegular widths indicating lecithotrophy. It is suggested that the onset of lecithotrophy in the Pentamerida was associated with a developmental innovation in which the mantle lobe of the larva was reflected over the apical lobe during the process of metamorphosis. This evolutionary innovation probably occurred during the late Cambrian or early Ordovician and was subsequently inherited during the process of cladogenesis.     

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.     

Changes in the timing of mantle formation and larval life history traits in linguliform and craniiform brachiopods

The distribution of embryonic and larval mantles is documented in linguliform and craniiform brachiopods. Criteria are presented for identifying these mantle types. The mantle type is related to planktotrophic and lecithotrophic larval life history patterns. In the Linguliformea and Craniiformea, all Lower Palaeozoic families with adequate preservation had larval mantles, indicating the presence of a planktotrophic larva. Heterochronic changes in the time of mantle origin, from the larval to the embryonic stage of development, has occurred several times. In the Lingulidae this change appears to have taken place at about the time the family originated in the Devonian and has been retained in extant genera. The family Discinidae has also retained a planktotrophic larval stage from the Lower Palaeozic to the present. The extant genus Crania in the Craniidae has a short-lived lecithotrophic larva that lacks a mantle. Through the Lower Jurassic, this family had planktotrophic larvae with a larval shell. During the Upper Jurassic, genera with a lecithotrophic larva that lacked a larval shell began to appear; however, the last genera in this family with a planktotrophic larva and a larval shell did not become extinct until the Tertiary.     

Earliest ontogeny of Middle Ordovician rhynchonelliform brachiopods (Clitambonitoidea and Polytoechioidea): implications for brachiopod phylogeny

New data on the earliest ontogeny of Mid-Ordovician Baltoscandian clitambonitoid ( Apomatella , Neumania and Oslogonites ) and polytoechioid ( Antigonambonites and Raunites ) brachiopods reveal significant differences in the life history of the taxa belonging to these two superfamilies. The Polytoechioidea and probably other members of the Billingsellida had planktotrophic larvae, in which the dorsal and ventral mantle lobes formed separately and without reversion. The 'pedicle sheath' in Antigonambonites is secreted by a section of modified ventral mantle and thus this 'pedicle' is not homologous within the pedicle of rhynchonellate brachiopods. It is likely that polytoechioids and other members of the strophomenate clade had the same type of ontogeny and mode of attachment. In contrast, the ontogeny and mode of attachment of clitambonitoids are similar to that of recent rhynchonellates: their mantle lobes and both valves formed simultaneously, and the pedicle most likely formed from the larval pedicle lobe. Evidence for the lecithotrophic nature of clitambonitoid larva is discussed. This confirms that the Clitambonitoidea, unlike the Polytoechioidea, represents an ingroup within the Rhynchonellata.     

The food and feeding habits of five freshwater and brackish-water fish species in Nigeria

The food and feeding habits of five economically important fresh-and brackish-water fishes, Channa obscura, Chrysichthys nigrodigitatus, Heterotis niloticus, Synodontis nigrita and Trachinotus maxillosus, were investigated. A number of techniques were used to carry out gut content analysis, including the Hynes 'point' system based on volume estimations expressed as relative percentages. Juveniles of C. nigrodigitatus were omnivorous, consuming 32% gastropods, 30% nematodes, 14% diatoms and 8% crustaceans, while adults were planktotrophic, consuming 23% diatoms, 33% Chlorophyceae and 22% crustaceans. Synodontis nigrita juveniles fed almost exclusively (91%) on nematodes, while adults were predominantly planktotrophic, their diet comprising 50% diatoms and 50% crustaceans. Trachinotus maxillosus was exclusively benthotrophic, consuming 99.5% gastropods and 0.46% nematodes. Heterotis niloticus was planktotrophic at the adult stage and consumed 72% crustaceans, 12% gastropods, 3% fish and about 90% planktotrophic at the juvenile phase. Chrysichthys obscura was purely benthotrophic at the juvenile stage, feeding 100% on nematodes, but fed mainly (89%) on fish at the adult stage.     

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.     

First records of brachiopods of the family Discinidae (Class Lingulata) from the Upper Jurassic of West Siberia

The organophosphatic brachiopods of the superfamily Discinoidea, family Discinidae are described from the Upper Jurassic of West Siberia. The protegulum (embryonic shell) preserved in adult shells is for the first time discovered in Mesozoic discinids. The shell microstructure is studied. A new species, Discinisca undata Smirnova, sp. nov., is established.     

Dark bands on pyritic internal moulds of the Early Jurassic ammonites Oxynoticeras and Cheltonia from Gloucestershire,England: interpretation and significance to ammonite growth analysis

Abstract: Thin, radiating, darker bands occur on pyritic internal moulds of the Early Jurassic ammonites Oxynoticeras and Cheltonia from Bishop’s Cleeve, Gloucestershire. They closely resemble true colour patterns preserved in Early Jurassic Calliphylloceras from Kutch, India, and false colour patterns reported in Carboniferous and Triassic ammonoids. Up to five dark bands occur within the body chamber, suggesting that they do not represent serially repeated anatomical structures, but the same feature repeatedly formed during growth. Dark bands are interpreted as traces of black bands deposited on the inside of the shell at the aperture during pauses in growth. The angles between dark bands and between septa correlate strongly in Cheltonia, suggesting that pauses in growth coincided with septal secretion during the chamber formation cycle. There are, however, no other indications that growth was episodic in either genus.     

Larval ecology and morphology in fossil gastropods

The shell of marine gastropods conserves and reflects early ontogeny, including embryonic and larval stages, to a high degree when compared with other marine invertebrates. Planktotrophic larval development is indicated by a small embryonic shell (size is also related to systematic placement) with little yolk followed by a multiwhorled shell formed by a free‐swimming veliger larva. Basal gastropod clades (e.g. Vetigastropoda) lack planktotrophic larval development. The great majority of Late Palaeozoic and Mesozoic ‘derived’ marine gastropods (Neritimorpha, Caenogastropoda and Heterobranchia) with known protoconch had planktotrophic larval development. Dimensions of internal moulds of protoconchs suggest that planktotrophic larval development was largely absent in the Cambrian and evolved at the Cambrian–Ordovician transition, mainly due to increasing benthic predation. The evolution of planktotrophic larval development offered advantages and opportunities such as more effective dispersal, enhanced gene flow between populations and prevention of inbreeding. Early gastropod larval shells were openly coiled and weakly sculptured. During the Mid‐ and Late Palaeozoic, modern tightly coiled larval shells (commonly with strong sculpture) evolved due to increasing predation pressure in the plankton. The presence of numerous Late Palaeozoic and Triassic gastropod species with planktotrophic larval development suggests sufficient primary production although direct evidence for phytoplankton is scarce in this period. Contrary to previous suggestions, it seems unlikely that the end‐Permian mass extinction selected against species with planktotrophic larval development. The molluscan classes with highest species diversity (Gastropoda and Bivalvia) are those which may have planktotrophic larval development. Extremely high diversity in such groups as Caenogastropoda or eulamellibranch bivalves is the result of high phylogenetic activity and is associated with the presence of planktotrophic veliger larvae in many members of these groups, although causality has not been shown yet. A new gastropod species and genus, Anachronistella peterwagneri, is described from the Late Triassic Cassian Formation; it is the first known Triassic gastropod with an openly coiled larval shell.     

Post‐metamorphic allometry in the earliest acrotretoid brachiopods from the lower Cambrian (Series 2) of South China,and its implications

The earliest growth of post‐metamorphic (post‐larval) shells in two species of Eohadrotreta is described from the Cambrian Shuijingtuo Formation of South China. Two different growth patterns can be observed by quantifying developmental variations in size and shape of successive stages of post‐metamorphic shell growth (including the pedicle foramen forming stage, pedicle foramen enclosing stage and intertrough increasing stage) of Eohadrotreta zhenbaensis and Eohadrotreta ? zhujiahensis . The pedicle foramen is never enclosed within the metamorphic shell of E. zhenbaensis , while the enclosed pedicle foramen of E .? zhujiahensis is located directly outside the metamorphic shell after the pedicle foramen enclosing stage. A strongly allometric growth pattern of E. zhenbaensis is demonstrated by the early enclosure of the pedicle foramen; an accelerated lengthening of the ventral intertrough is associated with the development of a more complex dorsal median septum during the intertrough increasing stage. By contrast, E .? zhujiahensis demonstrates possible paedomorphic development by delayed enclosure of pedicle foramen and an associated decreased lengthening of ventral intertrough during the intertrough increasing stage. This ontogenetic developmental sequence represents the marginal accretionary formation and growth of the pedicle foramen, which resembles that of linguloid brachiopods. Furthermore, the developmental process of the pedicle foramen of Eohadrotreta seems to recapitulate the likely evolutionary transition from the Botsfordiidae, with open delthyrium, to the Acrotheloidea, with an enclosed foramen. This study provides a unique opportunity to obtain a complete understanding of the ontogenetic development of the earliest acrotretoids, and casts new light on the phylogeny of lingulate brachiopods.     

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.