Mechanical adaptation of brachiopod shells via hydration-induced structural changes.

The function-optimized properties of biominerals arise from the hierarchical organization of primary building blocks. Alteration of properties in response to environmental stresses generally involves time-intensive processes of resorption and reprecipitation of mineral in the underlying organic scaffold. Here, we report that the load-bearing shells of the brachiopod Discinisca tenuis are an exception to this process. These shells can dynamically modulate their mechanical properties in response to a change in environment, switching from hard and stiff when dry to malleable when hydrated within minutes. Using ptychographic X-ray tomography, electron microscopy and spectroscopy, we describe their hierarchical structure and composition as a function of hydration to understand the structural motifs that generate this adaptability. Key is a complementary set of structural modifications, starting with the swelling of an organic matrix on the micron level via nanocrystal reorganization and ending in an intercalation process on the molecular level in response to hydration.

Warm afterglow from the Toarcian Oceanic Anoxic Event drives the success of deep-adapted brachiopods.

Many aspects of the supposed hyperthermal Toarcian Oceanic Anoxic Event (T-OAE, Early Jurassic, c. 182 Ma) are well understood but a lack of robust palaeotemperature data severely limits reconstruction of the processes that drove the T-OAE and associated environmental and biotic changes. New oxygen isotope data from calcite shells of the benthic fauna suggest that bottom water temperatures in the western Tethys were elevated by c. 3.5 °C through the entire T-OAE. Modelling supports the idea that widespread marine anoxia was induced by a greenhouse-driven weathering pulse, and is compatible with the OAE duration being extended by limitation of the global silicate weathering flux. In the western Tethys Ocean, the later part of the T-OAE is characterized by abundant occurrences of the brachiopod Soaresirhynchia, which exhibits characteristics of slow-growing, deep sea brachiopods. The unlikely success of Soaresirhynchia in a hyperthermal event is attributed here to low metabolic rate, which put it at an advantage over other species from shallow epicontinental environments with higher metabolic demand.

Cephalodiscus planitectus sp. nov. (Hemichordata: Pterobranchia) from Sagami Bay, Japan.

We describe here a new pterobranch, Cephalodiscus planitectus sp. nov. This pterobranch was collected from rocky slopes, at 100-300 m depth, off Jogashima Island, Sagami Bay, Japan. The tubaria of this new species have unique morphological features that differentiate it from known species. The tubaria are usually isolated from one another and have a completely flat and smooth surface that is devoid of erect features and projecting spines. Each has a simple, non-branched tubular cavity that is usually inhabited by a mature animal and its asexually budding offspring. The zooids have three pairs of tentaculated arms. A single bud is produced on the dorsal side of the stalk in adult zooids. In one instance, a live embryo was observed rotating at the bottom of a tubarium. Molecular phylogenetic analysis showed that C. planitectus is a sister group to all other Cephalodiscus species analyzed to date.

Morphological variation suggests that chitinozoans may be fossils of individual microorganisms rather than metazoan eggs.

Chitinozoans are organic-walled microfossils widely recorded in Ordovician to Devonian (ca 485-359 Mya) marine sediments and extensively used in high-resolution biostratigraphy. Their biological affinity remains unknown, but most commonly, they are interpreted as eggs of marine metazoans. Here, we provide new insights into their palaeobiology from three lines of inquiry. We examine morphological variation of a new, well-preserved Late Ordovician species, Hercochitina violana; analyse a compiled dataset of measurements on 378 species representing all known chitinozoan genera; and compare these data with the size variation of eggs of both extinct and extant aquatic invertebrates. The results indicate that the magnitude of size variation within chitinozoan species is larger than observed in fossil and modern eggs. Additionally, delicate morphological structures of chitinozoans, such as prosome and complex ornamentation, are inconsistent with the egg hypothesis. Distinct and continuous morphological variation in H. violana is more plausibly interpreted as an ontogenetic series of individual microorganisms, rather than as eggs.

Fibers on the Fly: Multiscale Mechanisms of Fiber Formation in the Capture Slime of Velvet Worms.

Many organisms have evolved a capacity to form biopolymeric fibers outside their bodies for functions such as defense, prey capture, attachment, and protection. In particular, the adhesive capture slime of onychophorans (velvet worms) is remarkable for its ability to rapidly form stiff fibers through mechanical drawing. Notably, fibers that are formed ex vivo from extracted slime can be dissolved in water and new fibers can be drawn from the solution, indicating that fiber formation is encoded in the biomolecules that comprise the slime. This review highlights recent findings on the biochemical and physicochemical principles guiding this circular process in the Australian onychophoran Euperipatoides rowelli. A multiscale cross-disciplinary approach utilizing techniques from biology, biochemistry, physical chemistry, and materials science has revealed that the slime is a concentrated emulsion of nanodroplets comprised primarily of proteins, stabilized via electrostatic interactions, possibly in a coacervate phase. Upon mechanical agitation, droplets coalesce, leading to spontaneous self-assembly and fibrillation of proteins-a completely reversible process. Recent investigations highlight the importance of subtle transitions in protein structure and charge balance. These findings have clear relevance for better understanding this adaptive prey capture behavior and providing inspiration toward sustainable polymer processing.

Pseudocnidae of archinemerteans (Nemertea, Palaeonemertea) and their implications for nemertean systematics.

The structure of pseudocnidae of 16 species of Palaeonemertea clade Archinemertea (= Cephalotrichida s.l.) was investigated with confocal laser, scanning, and transmission electron microscopy (TEM). All species of the genus Cephalothrix possess two kinds of pseudocnidae, large and small. Only one type of pseudocnida is present in Balionemertes and Cephalotrichella. TEM revealed variation in the ultrastructure of large and small pseudocnidae of four species of Cephalothrix. Pseudocnidae of Balionemertes, Cephalotrichella, and Cephalothrix differ in substructure: in Balionemertes and Cephalotrichella the medulla is located in the basal half of the pseudocnidae with а precore layer situated in the apical half, whereas in Cephalothrix spp. and other palaeonemerteans the medulla surrounds a precore layer. Our results confirm the division of archinemerteans into Cephalotrichidae (with genus Cephalothrix) and Cephalotrichellidae (with genera Cephalotrichella and Balionemertes). The synapomorphy of Cephalotrichidae is pseudocnida dimorphism and the synapomorphies of Cephalotrichellidae are the position of the pseudocnidae on epithelial ridges and the distinct organization of pseudocnida layers, specifically the relative position of the medulla and precore layers. The pseudocnida lateral process, one or more of which is present in most species observed, is a probable synapomorphy of the clade Archinemertea. This is the first application of pseudocnida features to distinguish super-generic nemertean taxa and the results suggest that pseudocnidae provide a useful source of characters for nemertean systematics.

A comparative ultrastructure study of storage cells in the eutardigrade Richtersius coronifer in the hydrated state and after desiccation and heating stress.

Tardigrades represent an invertebrate phylum with no circulatory or respiratory system. Their body cavity is filled with free storage cells of the coelomocyte-type, which are responsible for important physiological functions. We report a study comparing the ultrastructure of storage cells in anhydrobiotic and hydrated specimens of the eutardigrade Richtersius coronifer. We also analysed the effect of temperature stress on storage cell structure. Firstly, we verified two types of ultrastructurally different storage cells, which differ in cellular organelle complexity, amount and content of reserve material and connection to oogenetic stage. Type I cells were found to differ ultrastructurally depending on the oogenetic stage of the animal. The main function of these cells is energy storage. Storage cells of Type I were also observed in the single male that was found among the analysed specimens. The second cell type, Type II, found only in females, represents young undifferentiated cells, possibly stem cells. The two types of cells also differ with respect to the presence of nucleolar vacuoles, which are related to oogenetic stages and to changes in nucleolic activity during oogenesis. Secondly, this study revealed that storage cells are not ultrastructurally affected by six months of desiccation or by heating following this desiccation period. However, heating of the desiccated animals (tuns) tended to reduce animal survival, indicating that long-term desiccation makes these animals more vulnerable to heat stress. We confirmed the degradative pathways during the rehydration process after desiccation and heat stress. Our study is the first to document two ultrastructurally different types of storage cells in tardigrades and reveals new perspectives for further studies of tardigrade storage cells.

Ground plan of the larval nervous system in phoronids: Evidence from larvae of viviparous phoronid.

Nervous system organization differs greatly in larvae and adults of many species, but has nevertheless been traditionally used for phylogenetic studies. In phoronids, the organization of the larval nervous system depends on the type of development. With the goal of understanding the ground plan of the nervous system in phoronid larvae, the development and organization of the larval nervous system were studied in a viviparous phoronid species. The ground plan of the phoronid larval nervous system includes an apical organ, a continuous nerve tract under the preoral and postoral ciliated bands, and two lateral nerves extending between the apical organ and the nerve tract. A bilobed larva with such an organization of the nervous system is suggested to be the primary larva of the taxonomic group Brachiozoa, which includes the phyla Brachiopoda and Phoronida. The ground plan of the nervous system of phoronid larvae is similar to that of the early larvae of annelids and of some deuterostomians. The protostome- and deuterostome-like features, which are characteristic of many organ systems in phoronids, were probably inherited by phoronids from the last common bilaterian ancestor. The information provided here on the ground plan of the larval nervous system should be useful for future analyses of phoronid phylogeny and evolution.

The first data on the innervation of the lophophore in the rhynchonelliform brachiopod Hemithiris psittacea: what is the ground pattern of the lophophore in lophophorates?

BACKGROUND: The nervous system in brachiopods has seldom been studied with modern methods. An understanding of lophophore innervation in adult brachiopods is useful for comparing the innervation of the same lophophore type among different brachiopods and can also help answer questions about the monophyly of the lophophorates. Although some brachiopods are studied with modern methods, rhynchonelliform brachiopods still require investigation. The current study used transmission electron microscopy, immunocytochemistry, and confocal laser scanning microscopy to investigate the nerve system of the lophophore and tentacles in the rhynchonelliform Hemithiris psittacea. RESULTS: Four longitudinal nerves pass along each brachium of the lophophore: the main, accessory, second accessory, and lower. The main brachial nerve extends at the base of the dorsal side of the brachial fold and gives rise to the cross nerves, passing through the extracellular matrix to the tentacles. Cross nerves skirt the accessory brachial nerve, branch, and penetrate into adjacent outer and inner tentacles, where they are referred to as the frontal tentacular nerves. The second accessory nerve passes along the base of the inner tentacles. This nerve consists of Ʊ-like parts, which repetitively skirt the frontal and lateral sides of the inner tentacle and the frontal sides of the outer tentacles. The second accessory nerve gives rise to the latero-frontal nerves of the inner and outer tentacles. The abfrontal nerves of the inner tentacles also originate from the second accessory nerve, whereas the abfrontal nerves of the outer tentacles originate from the lower brachial nerve. The lower brachial nerve extends along the outer side of the lophophore brachia and gives rise to the intertentacular nerves, which form a T-like branch and penetrate the adjacent outer tentacles where they are referred to as abfrontal nerves. The paired outer radial nerves start from the lower brachial nerve, extend into the second accessory nerve, and give rise to the lateroabfrontal tentacular nerves of the outer tentacles. CONCLUSIONS: The innervation of the lophophore in the rhynchonelliform Hemithiris psittacea differs from that in the inarticulate Lingula anatina in several ways. The accessory brachial nerve does not participate in the innervation of the tentacles in H. psittacea as it does in L. anatina. The second accessory nerve is present in H. psittacea but not in L. anatina. There are six tentacular nerves in the outer tentacles of H. psittacea but only four in all other brachiopods studied to date. The reduced contribution of the accessory brachial nerve to tentacle innervation may reflect the general pattern of reduction of the inner lophophoral nerve in both phoronids and brachiopods. Bryozoan lophophores, in contrast, have a weakened outer nerve and a strengthened inner nerve. Our results suggest that the ancestral lophophore of all lophophorates had a simple shape but many nerve elements.

Ultrastructure of the coelom in the brachiopod Lingula anatina.

The organization of the coelomic system and the ultrastructure of the coelomic lining are used in phylogenetic analysis to establish the relationships between major taxa. Investigation of the anatomy and ultrastructure of the coelomic system in brachiopods, which are poorly studied, can provide answers to fundamental questions about the evolution of the coelom in coelomic bilaterians. In the current study, the organization of the coelom of the lophophore in the brachiopod Lingula anatina was investigated using semithin sectioning, 3D reconstruction, and transmission electron microscopy. The lophophore of L. anatina contains two main compartments: the preoral coelom and the lophophoral coelom. The lining of the preoral coelom consists of ciliated cells. The lophophoral coelom is subdivided into paired coelomic sacs: the large and small sinuses (= canals). The lining of the lophophoral coelom varies in structure and includes monociliate myoepithelium, alternating epithelial and myoepithelial cells, specialized peritoneum and muscle cells, and podocyte-like cells. Connections between cells of the coelomic lining are provided by adherens junctions, tight-like junctions, septate junctions, adhesive junctions, and direct cytoplasmic bridges. The structure of the coelomic lining varies greatly in both of the main stems of the Bilateria, that is, in the Protostomia and Deuterostomia. Because of this great variety, the structure of the coelomic lining cannot by itself be used in phylogenetic analysis. At the same time, the ciliated myoepithelium can be considered as the ancestral type of coelomic lining. The many different kinds of junctions between cells of the coelomic lining may help coordinate the functioning of epithelial cells and muscle cells.

Earliest Onychophoran in Amber Reveals Gondwanan Migration Patterns.

The anomalous occurrence of supposedly Gondwanan taxa in Laurasian-derived regions remains an intriguing chapter of paleobiogeographical history. Representatives of Peripatidae, a major subgroup of velvet worms (Onychophora), show a disjointed distribution in the neotropics, tropical Africa, and Southeast Asia, the latter being the only landmass previously associated with Laurasia [1, 2]. The arrival of these animals in Southeast Asia is explained by two alternative, albeit not mutually exclusive, hypotheses: an early migration via Europe before continental drift (Eurogondwana hypothesis) or transportation via insular India during the Cretaceous and Paleogene ("out-of-India" hypothesis) [3-6]. The latter hypothesis is based on a single extant species of Peripatidae, Typhloperipatus williamsoni, in India. †Cretoperipatus burmiticus from Myanmar is the oldest fossil onychophoran found in amber [7], dating to sometime between the two proposed scenarios, and hence crucial for clarifying how Gondwanan lineages of these low-vagility animals reached Southeast Asia (see also Supplemental Information). Based on the anatomical reconstruction of †C. burmiticus using synchrotron radiation-based X-ray microtomography (SRµCT) and comparisons with extant taxa, we resolved this fossil species within Onychophora, particularly within Peripatidae, with T. williamsoni as its closest extant relative. This suggests that an early Eurogondwanan migration of peripatids was the most likely event, as Burmese amber is too old to be compatible with the out-of-India hypothesis. Moreover, peripatids probably colonized India only recently from Myanmar, refuting the putative Gondwanan relict status of Indian onychophorans. Finally, preservation artifacts identified in the novel amber material might have a major impact on studies of onychophoran stem and/or crown groups.

Distribution of tetrodotoxin in the ribbon worm Lineus alborostratus (Takakura, 1898) (nemertea): Immunoelectron and immunofluorescence studies.

Transmission electron and confocal laser scanning (CLSM) microscopies with monoclonal anti-tetrodotoxin antibodies were used to locate tetrodotoxin (TTX) in tissues and gland cells of the ribbon worm Lineus alborostratus. CLSM studies have shown that the toxin is primarily localized in the cutis (special subepidermal layer) of the body wall and in the glandular epithelium of the proboscis. Immunoelectron micrographs have shown that only subepidermal bacillary gland cells type I in cutis and pseudocnidae-containing and mucoid gland cells manifested TTX-gold labeling. TTX was associated with the nuclear envelope, endoplasmic reticulum membrane, and secretory granules of TTX-positive gland cells. These studies indicate that ТТХ is brought into the cytoplasm of the glandular cells of the cutis and proboscis epithelium, where it is associated with membrane-enclosed organelles involved in protein secretion and then concentrated in glandular granules.

Larval development of Brachiopod Coptothyris grayi (Davidson, 1852) (Brachiopoda, Rhynchonelliformea).

The larval development of the Brachiopod Coptothyris grayi (Davidson, 1852) from the Sea of Japan is described for the first time. Ciliated blastula proved to represent the first free-swimming stage. The blastopore is initially formed as a rounded hole stretching later along the anteroposterior axis. The larva is first divided into two lobes (the apical lobe and the trunk); the mantle lobe is formed later as two lateral folds. Two pairs of seta bundles appear in the late stage larvae. The apical larval lobe in brachiopods is supposed to match the pre-oral lobe and anterior part of the trunk with tentacles in phoronids.

Malacobdella arrokeana: Parasite or Commensal of the Giant Clam Panopea abbreviata?

We examined trophic relationship between the nemertean Malacobdella arrokeana and its host, the edible geoduck Panopea abbreviata by studying the diets of both species by direct (stomach contents) and indirect methods (stable-isotope analysis of C and N). In addition to these methods, the feeding behavior of M. arrokeana within the host and the morphology of its feeding organs were examined. The feeding behavior of M. arrokeana did not exhibit parasitic characteristics, and the proboscis morphology indicates it is unable to injure host tissues. Analysis of stomach contents revealed a diet consisting mainly of microalgae and diatoms. Panopea abbreviata and M. arrokeana shared similar trophic levels, presenting no differences in the spread of the isotopic niches and high overlap (SEAB overlapped 63%). Consistent with this, our results showed no differences in δ(15)N or δ(13)C values between the two species. The combination of direct and indirect approaches revealed that M. arrokeana has a diet similar to that of its host, confirming a commensal relationship.

Structure of the Proboscis Endothelium in Nemertea.

We studied the ultrastructure of the proboscis endothelium of 14 nemertean species. In all nemerteans examined, the endothelium is organized as a pseudostratified myoepithelium consisting of two types of cells resting on the basal extracellular matrix: apically situated supportive cells and subapical myocytes covered by cytoplasmic sheets of the supportive cells. Myocytes form the inner circular musculature of the proboscis; the endothelium in the bulb of monostiliferous nemerteans lacks myocytes. The endothelium of the studied species differs in the number of rows of muscle fibres (one vs. several rows), the number of myofibrils in myocytes (one vs. two to five), the number of processes of myocytes covered by one supportive cell (one vs. two to 23), and in the number of processes in supportive cells (one vs. two to five). In some of the species, rudimentary cilia of supportive cells were revealed by using cLSM and an antibody against tubulin. The data obtained indicate that the proboscis endothelium in nemerteans is in fact a coelothelium, but recognition of the ancestral state of the coelomic lining in Nemertea is problematic, as the rhynchocoel peritoneum lacks myocytes.

Ediacaran skeletal metazoan interpreted as a lophophorate.

While many skeletal biomineralized genera are described from Ediacaran (635-541 million years ago, Ma) strata, none have been suggested to have an affinity above the Porifera-Cnidaria metazoan grade. Here, we reinterpret the widespread terminal Ediacaran (approx. 550-541 Ma) sessile goblet-shaped Namacalathus as a triploblastic eumetazoan. Namacalathus has a stalked cup with radially symmetrical cross section, multiple lateral lumens and a central opening. We show that the skeleton of Namacalathus is composed of a calcareous foliated ultrastructure displaying regular concordant columnar inflections, with a possible inner organic-rich layer. These features point to an accretionary growth style of the skeleton and an affinity with the Lophotrochozoa, more specifically within the Lophophorata (Brachiopoda and Bryozoa). Additionally, we present evidence for asexual reproduction as expressed by regular budding in a bilateral pattern. The interpretation of Namacalathus as an Ediacaran total group lophophorate is consistent with an early radiation of the Lophophorata, as known early Cambrian representatives were sessile, mostly stalked forms, and in addition, the oldest known calcareous Brachiopoda (early Cambrian Obolellida) and Bryozoa (Ordovician Stenolaemata) possessed foliated ultrastructures.

A low-cost technique to manufacture a container to process meiofauna for scanning electron microscopy.

An easy and low-cost method to elaborate a container to dehydrate nematodes and other meiofauna in order to process them for scanning electron microscopy (SEM) is presented. Illustrations of its elaboration, step by step, are included. In addition, a brief methodology to process meiofauna, especially nematodes and kinorhynchs, and illustrations are provided. With this methodology it is possible to easily introduce the specimens, to lock them in a closed chamber allowing the infiltration of fluids and gases (ethanol, acetone, carbon dioxide) but avoiding losing the specimens. After using this meiofauna basket for SEM the results are efficient. Examples of nematode and kinorhynch SEM pictures obtained using this methodology are also included.

An Integrated Morphological and Molecular Approach to the Description and Systematisation of a Novel Genus and Species of Macrodasyida (Gastrotricha).

BACKGROUND: Gastrotricha systematics is in a state of flux mainly due to the conflicts between cladistic studies base on molecular markers and the classical systematisation based on morphological traits. In sandy samples from Thailand, we found numerous macrodasyidan gastrotrichs belonging to an undescribed species of difficult taxonomic affiliation. The abundance and original nature of the specimens prompted us to undertake a deep survey of both morphological and molecular traits aiming at a reliable systematisation of the new taxon. METHODOLOGY/PRINCIPAL FINDINGS: Using several microscopical techniques we investigated the external and internal anatomy, including the muscular and nervous systems of the new species. Additional specimens were used to obtain the 18S rRNA gene sequence; molecular data was analysed cladistically in conjunction with data from additional species belonging to the near complete Macrodasyida taxonomic spectrum. Specimens are vermiform, up to 806 µm in total length, and show a well-defined head equipped with peculiar leaf-like sensorial organs and a single-lobed posterior end. The adhesive apparatus includes anterior, ventrolateral, dorsal and posterior tubes. Pharynx is about 1/4 of the total length and shows pores at its posterior 3/4. Adult specimens exhibit maturing eggs and a bulky, muscular caudal organ, but do not show sperm nor the frontal organ. Musculature and nervous system organisation resemble the usual macrodasyidan plan; however, the somatic circular muscles of the intestinal region surround all other muscular components and a third FMRFamide-IR commissure ventral to the pharyngo-intestinal junction appear to be an autoapomorphic traits of the new species. CONCLUSIONS/SIGNIFICANCE: While the anatomical characteristics of the Asian specimens appear so unique to grant the establishment of a new taxon, for which the name Thaidasys tongiorgii gen. et sp. nov. is proposed, the result of phylogenetic analyses based on the 18S rRNA gene unites the new genus with the family Macrodasyidae.

Critical appraisal of tubular putative eumetazoans from the Ediacaran Weng'an Doushantuo biota.

Molecular clock analyses estimate that crown-group animals began diversifying hundreds of millions of years before the start of the Cambrian period. However, the fossil record has not yielded unequivocal evidence for animals during this interval. Some of the most promising candidates for Precambrian animals occur in the Weng'an biota of South China, including a suite of tubular fossils assigned to Sinocyclocyclicus, Ramitubus, Crassitubus and Quadratitubus, that have been interpreted as soft-bodied eumetazoans comparable to tabulate corals. Here, we present new insights into the anatomy, original composition and phylogenetic affinities of these taxa based on data from synchrotron radiation X-ray tomographic microscopy, ptychographic nanotomography, scanning electron microscopy and electron probe microanalysis. The patterns of deformation observed suggest that the cross walls of Sinocyclocyclicus and Quadratitubus were more rigid than those of Ramitubus and Crassitubus. Ramitubus and Crassitubus specimens preserve enigmatic cellular clusters at terminal positions in the tubes. Specimens of Sinocyclocyclicus and Ramitubus have biological features that might be cellular tissue or subcellular structures filling the spaces between the cross walls. These observations are incompatible with a cnidarian interpretation, in which the spaces between cross walls are abandoned parts of the former living positions of the polyp. The affinity of the Weng'an tubular fossils may lie within the algae.

Hallucigenia's head and the pharyngeal armature of early ecdysozoans.

The molecularly defined clade Ecdysozoa comprises the panarthropods (Euarthropoda, Onychophora and Tardigrada) and the cycloneuralian worms (Nematoda, Nematomorpha, Priapulida, Loricifera and Kinorhyncha). These disparate phyla are united by their means of moulting, but otherwise share few morphological characters--none of which has a meaningful fossilization potential. As such, the early evolutionary history of the group as a whole is largely uncharted. Here we redescribe the 508-million-year-old stem-group onychophoran Hallucigenia sparsa from the mid-Cambrian Burgess Shale. We document an elongate head with a pair of simple eyes, a terminal buccal chamber containing a radial array of sclerotized elements, and a differentiated foregut that is lined with acicular teeth. The radial elements and pharyngeal teeth resemble the sclerotized circumoral elements and pharyngeal teeth expressed in tardigrades, stem-group euarthropods and cycloneuralian worms. Phylogenetic results indicate that equivalent structures characterized the ancestral panarthropod and, seemingly, the ancestral ecdysozoan, demonstrating the deep homology of panarthropod and cycloneuralian mouthparts, and providing an anatomical synapomorphy for the ecdysozoan supergroup.