Fossil Embryos Research Articles

New ecdysozoan fossil embryos from the basal Cambrian of China

Invertebrate embryos have relatively low fossilization potential; however, when preserved, they can offer crucial insights into the evolutionary developmental biology of extinct animals. Here we report new exceptionally preserved fossil embryos from the early Fortunian (early Cambrian) of China, which are assigned to two new taxa, Saccus xixiangensis gen. et sp. nov. and Saccus necopinus gen. et sp. nov. All specimens are three-dimensionally phosphatized, akin to the Orsten-type preservation. The embryos are enclosed within a thin and smooth envelope ranging in diameter from 730 μm to 1 mm. The embryos have a bag-shaped body, a non-ciliated integument, radially arranged sclerites apically, and bilaterally arranged sclerites abapically, but lack any orifice, thus representing a developmental stage prior to the formation of a mouth or anus. The integument shows soft deformation, while the sclerites remain largely undeformed, suggesting that the sclerites are stiffer and likely cuticularized. The arrangement of the abapical sclerites imparts bilaterality, classifying these new embryos as bilaterians. Furthermore, the absence of cilia or cilium insertion sites and the presence of cuticularized sclerites suggest an ecdysozoan affinity. The relatively large embryo size indicates yolk-rich eggs, and the presence of cuticle implies that the embryos are in later embryonic stages, possibly close to hatching. In the absence of hatched specimens, it is inferred that these embryos undergo indirect development, hatching as lecithotrophic (yolk-feeding) larvae. In this case, the morphology of the juvenile and adult forms is uncertain due to metamorphosis during later growth. Alternatively, the embryos could undergo direct development, hatching as lecithotrophic juveniles, with both juveniles and adults potentially having a similar bag-shaped body without an introvert or paired limbs, closely reminiscent of Saccorhytus. This study expands our understanding of the morphological disparity and diversity of early Fortunian fossil embryos and ecdysozoans.

Asynchronized cell division in embryo-like fossils from the Ediacaran Zhenba microfossil assemblage.

Ediacaran embryo-like spherical fossils exhibit diverse cell adhesion patterns resembling partial cleavage-stage embryos of living animals. Two three-celled specimens characterized by a pair of small cells overlying a large cell have been recovered from the Ediacaran Zhenba microfossil assemblage. Their cell adhesion pattern is highly comparable to a phenomenon reported from the Weng'an biota that was interpreted as fossil embryos undergoing discoidal cleavage. However, our specimens contain fewer cells and thus probably represent developmental precursors of the Weng'an counterparts. Additionally, new material shows several anatomical features that are inconsistent with an embryo interpretation, including (1) an unusually large volume of "blastomeres,"(2) a putative nucleus preserved within the large "yolk cell,"and (3) completely separated cells. Collectively, the Zhenba embryo-like specimens permit a reconstruction of the consecutive developmental sequence from single-celled individuals to the three-celled individuals, leading us to interpret the newly found specimens as products of abnormal development of Ediacaran embryo-like organisms whose affinity remains unresolved.

Internal anatomy of a fossilized embryonic stage ofthe Cambrian-Ordovician scalidophoran Markuelia.

The Wangcun fossil Lagerstätte in Hunan, South China, has yielded hundreds of fossilized embryos of Markuelia hunanensis representing different developmental stages. Internal tissues have only rarely been observed, impeding further understanding of the soft tissue anatomy, phylogenetic affinity and evolutionary significance of Markuelia. In this study, we used synchrotron radiation X-ray tomographic microscopy (SRXTM) to study a new collection of fossil embryos from the Wangcun fossil Lagerstätte. We describe specimens exhibiting a spectrum of preservation states, the best of which preserves palisade structures underneath the cuticle of the head and tail, distinct from patterns of centripetal mineralization of the cuticle and centrifugal mineralization of hypha-like structures, seen elsewhere in this specimen and other fossils within the same assemblage. Our computed tomographic reconstruction of these mineralization phases preserves the gross morphology of (i) longitudinal structures associated with the tail spines, which we interpret as the proximal ends of longitudinal muscles, and (ii) a ring-shaped structure internal to the introvert, which we interpret as a ring-shaped brain, as anticipated of the cycloneuralian affinity of Markuelia. This is the first record of a fossilized nervous system in a scalidophoran, and the first instance in Orsten-style preservation, opening the potential for further such records within this widespread mode of high-fidelity three-dimensional preservation.

Interpretation of fossil embryos requires reasonable assessment of developmental age

AbstractDinosaur embryos cause a lot of excitement in the scientific literature and are often widely reported because of the general public's interest in dinosaur biology. Well-preserved, articulated oviraptorosaur embryos in eggs are usually interpreted as representing a stage of development close to hatching because of their large size and good level of skeletal ossification. Based on this evidence, a recent report suggested that the position of the one embryo's head was reminiscent of an avian-like hatching position. Here we explore how the developmental stage of well-preserved oviraptorosaur embryos can be estimated, rather than assumed. This will help in our understanding of their developmental biology and its evolutionary consequences. Using quantitative methods and comparison with modern crocodilian embryos, we show that all articulated oviraptorosaur embryos are small relative to the egg and most likely at a stage of development equivalent to around 50%–60% of the developmental period, that is, not even close to hatching. This conclusion is supported by the fact that many elements of the crocodilian skeleton are well ossified many weeks before hatching and the position of oviraptorosaur embryos’ heads was also comparable to a crocodilian embryo many days before hatching. Misunderstandings about the stage of the developmental biology of these well-preserved oviraptorosaur embryos hampers our understanding of the true nature of their reproductive biology. We urge a more conservative approach to their interpretation. This is important, because misunderstandings in the minds of the public about dinosaur biology are hard to counter once poorly evidenced ideas have been reported around the world.

An exquisitely preserved in-ovo theropod dinosaur embryo sheds light on avian-like prehatching postures

SummaryDespite the discovery of many dinosaur eggs and nests over the past 100 years, articulated in-ovo embryos are remarkably rare. Here we report an exceptionally preserved, articulated oviraptorid embryo inside an elongatoolithid egg, from the Late Cretaceous Hekou Formation of southern China. The head lies ventral to the body, with the feet on either side, and the back curled along the blunt pole of the egg, in a posture previously unrecognized in a non-avian dinosaur, but reminiscent of a late-stage modern bird embryo. Comparison to other late-stage oviraptorid embryos suggests that prehatch oviraptorids developed avian-like postures late in incubation, which in modern birds are related to coordinated embryonic movements associated with tucking — a behavior controlled by the central nervous system, critical for hatching success. We propose that such pre-hatching behavior, previously considered unique to birds, may have originated among non-avian theropods, which can be further investigated with additional discoveries of embryo fossils.

Synchrotron imagery of phosphatized eggs in Waptia cf. W. fieldensis from the middle Cambrian (Miaolingian, Wuliuan) Spence Shale of Utah

AbstractExceptionally preserved fossil eggs and embryos provide critical information regarding paleoembryogenesis, reproductive strategies, and the early ontogeny of early arthropods, but the rarity of preservation of both eggs and egg-bearing organisms in situ limits their use in detailed evolutionary developmental (evo-devo) studies. Burgess Shale-type deposits preserve rare instances of egg-bearing arthropods as carbonaceous compressions; however, the eggs are usually poorly preserved with no compelling evidence of embryos. We describe the first record of a brooding specimen of Waptia cf. W. fieldensis from the Spence Shale, a Cambrian (Wuliuan Stage) Burgess Shale-type deposit in northeastern Utah and southeastern Idaho. This is the first record of an egg-bearing arthropod from the Spence Shale and it exhibits two distinct modes of preservation among eggs within the single clutch: carbonization and phosphatization. Unlike the egg-bearing Burgess Shale specimens, many eggs of this Utah specimen are also preserved three-dimensionally. In addition, synchrotron radiation X-ray tomographic microscopy reveals internal distributions of mineral phases, along with potential remnants of the egg membrane and attachment structures, but, as in the Burgess Shale, no explicit traces of developing embryos. The distinct modes of preservation highlight the existence of diagenetic microenvironments within some eggs, but not in others during fossilization.

Characterization of the Multicellular Membrane‐Bearing Algae From the Kuanchuanpu Biota (Cambrian: Terreneuvian)

AbstractThe diversity of small shelly fossils (SSFs) demonstrates that multicellular organisms underwent large‐scale radiation at the beginning of the Cambrian, which is highlighted by the coexistence of various metazoans and the occurrence of their embryo fossils. However, little is known about early Cambrian eukaryotic multicellular algae, the primary producers that replaced oxygenic cyanobacteria and played a crucial ecological role in matter cycling and energy dynamics in marine ecosystems. In this study, hundreds of microscopic three‐dimensionally preserved multicellular aggregate fossils were obtained from the Kuanchuanpu Formation (Cambrian: Terreneuvian, ca. 535 Ma) in southern Shaanxi, South China, which consisted of several tightly packed multicellular clumps encapsulated in a thin organic membrane. Synchrotron tomography analysis further revealed that the cells of the whole aggregate, although partitioned into different clumps by a gelatinous membrane, were distinctly differentiated into an outer conical cell layer and an inner spherical‐cell layer, thus suggesting of a cortex‐medulla‐like differentiation. These characteristics resemble those of multicellular algae (e.g.,Wengania,Gremiphyca, andThallophyca) from the Weng'an biota (Ediacaran, South China) in morphology, size, and internal cell structure. Furthermore, a potential asexual life cycle for these membranous algae was proposed based on their morphological and structural characteristics.

An intermediate type of medusa from the early Cambrian Kuanchuanpu Formation, South China

AbstractThe tetraradial or pentaradial fossil embryos and related hatched individuals from the early Cambrian Kuanchuanpu Formation are of great interest for understanding the early evolution of medusozoans. The phylogenetic and evolutionary significance of their external and internal characters (e.g. manubrium, tentacles, septa and claustra) is still controversial. Here we describe a new pentamerous medusozoan, Hanagyroia orientalis gen. et sp. nov., characterized by five well‐developed perradial oral lips around a remarkably large manubrium, a conspicuous equatorial groove, and five short interradial pairs of extensile tentacles at the bell margin. Internally, five broad and stout interradial septa join horizontally to form the claustra. Hanagyroia orientalis lacks the frenula, apertural lappet and velarium seen in coeval microfossils and extant cubozoans. Although H. orientalis resembles extant coronate scyphozoans in its round medusa‐like bell margin and equatorial groove, cladistic analysis suggests close affinity with cubozoans. Hanagyroia may represent an intermediate morphological type between scyphozoans and cubozoans. The well‐developed oral lips and paired short strong tentacles of Hanagyroia suggest direct development.

Artificial Intelligence Identification of Multiple Microfossils from the Cambrian Kuanchuanpu Formation in Southern Shaanxi, China

AbstractThe Cambrian Kuanchuanpu Formation in southern Shaanxi, China is a critical window for the understanding of the Cambrian explosion, because of abundant and various exceptionally preserved metazoans and embryo fossils yielded. The efficiency of traditional sample manually selecting with microscopes is quite low and hinder the discoveries of new species, thus recognition and classification of microfossils by artificial intelligence (AI) is substantially in the request. In this paper, we develop a procedure for fossil area segmentation in common multi‐typed mixed photos by improved watershed algorithm. And for better fossil recognition, previous histogram of oriented grandient (HOG) algorithm is replaced by scale invariant feature transform (SIFT), which is feasible for the segmented images and increase the accuracy significantly. Thus, the scope of application of AI fossil recognition can be extended form single fossil image to multi‐typed mixed images and the reliability is also secured, as the result of our test presents a high (at least 84%) accuracy of fossil recognition.

Diverse radial symmetry among the Cambrian Fortunian fossil embryos from northern Sichuan and southern Shaanxi provinces, South China

The Cambrian Fortunian fossil embryos exhibit embryonic development of ancient animals and hence have important bearings on evolutionary developmental biology. They have radial symmetry, and may be early representatives of cnidarians. Here we report new material of three-dimensionally phosphatized fossil embryos from the Fortunian Kuanchuanpu Formation and coeval strata in northern Sichuan and southern Shaanxi provinces, South China. The new material includes previously reported fossil embryos assignable to Pseudooides prima with biradial symmetry or pseudo-hexaradial symmetry, Quadrapyrgites quadratacris with tetraradial symmetry, and Olivooides multisulcatus with pentaradial symmetry. Additionally, we recovered two new types of fossil embryos, i.e., Embryo I with hexaradial symmetry and Embryo II with octaradial symmetry, and they are tentatively suggested to represent new cnidarians. In contrast to the diverse radial symmetry of the Fortunian cnidarians, modern cnidarians exhibit stable tetraradial symmetry in medusozoans, biradial symmetry in anthozoans, and bilateral symmetry in siphonophores (Hydrozoa). The current study supports the view that the tetraradial symmetry of modern medusozoans is a surviving remnant of their Fortunian relatives.

The early Cambrian fossil embryo Pseudooides is a direct-developing cnidarian, not an early ecdysozoan.

Early Cambrian Pseudooides prima has been described from embryonic and post-embryonic stages of development, exhibiting long germ-band development. There has been some debate about the pattern of segmentation, but this interpretation, as among the earliest records of ecdysozoans, has been generally accepted. Here, we show that the ‘germ band’ of P. prima embryos separates along its mid axis during development, with the transverse furrows between the ‘somites’ unfolding into the polar aperture of the ten-sided theca of Hexaconularia sichuanensis, conventionally interpreted as a scyphozoan cnidarian; co-occurring post-embryonic remains of ecdysozoans are unrelated. We recognize H. sichuanensis as a junior synonym of P. prima as a consequence of identifying these two form-taxa as distinct developmental stages of the same organism. Direct development in P. prima parallels the co-occuring olivooids Olivooides, and Quadrapyrgites and Bayesian phylogenetic analysis of a novel phenotype dataset indicates that, despite differences in their tetra-, penta- and pseudo-hexa-radial symmetry, these hexangulaconulariids comprise a clade of scyphozoan medusozoans, with Arthrochites and conulariids, that all exhibit direct development from embryo to thecate polyp. The affinity of hexangulaconulariids and olivooids to extant scyphozoan medusozoans indicates that the prevalence of tetraradial symmetry and indirect development are a vestige of a broader spectrum of body-plan symmetries and developmental modes that was manifest in their early Phanerozoic counterparts.

Experimental precipitation of apatite pseudofossils resembling fossil embryos.

Certain phosphatic grains preserved in the rock record are interpreted as microfossils representing a diversity of microorganisms from bacteria to fossil embryos. In addition to bona fide primary biological features, phosphatic microfossils and fossil embryos commonly exhibit features that result from abiotic precipitation or diagenetic alteration. Distinguishing between abiotic and primary biological features can be difficult, and some features thought to represent biological tissue could instead be artifacts that are unrelated to the original morphology of a preserved organism. Here, we present experimentally generated, abiotically produced mineral precipitates that morphologically resemble biologically produced features, some of which may be observed in the rock record or noted in extant organisms, including embryos. These findings extend the diversity of biomorphic features known to result from abiotic precipitation.

Eric Davidson and deep time.

Eric Davidson had a deep and abiding interest in the role developmental mechanisms played in generating evolutionary patterns documented in deep time, from the origin of the euechinoids to the processes responsible for the morphological architectures of major animal clades. Although not an evolutionary biologist, Davidson's interests long preceded the current excitement over comparative evolutionary developmental biology. Here I discuss three aspects at the intersection between his research and evolutionary patterns in deep time: First, understanding the mechanisms of body plan formation, particularly those associated with the early diversification of major metazoan clades. Second, a critique of early claims about ancestral metazoans based on the discoveries of highly conserved genes across bilaterian animals. Third, Davidson's own involvement in paleontology through a collaborative study of the fossil embryos from the Ediacaran Doushantuo Formation in south China.

Revision of Eocene electric rays (Torpediniformes, Batomorphii) from the Bolca Konservat-Lagerstätte, Italy, reveals the first fossil embryo in situ in marine batoids and provides new insights into the origin of trophic novelties in coral reef fishes

The Eocene electric ray †Titanonarke Carvalho, 2010 from the Bolca Konservat-Lagerstätte, north-eastern Italy, is redescribed in detail based upon new material from recent excavations. This taxon exhibits a combination of features (large voids between the pectoral and the axial skeleton filled in life by electric organs, anteriorly directed fan-shaped antorbital cartilages, lack of dermal denticles, long prepelvic processes, and rounded basibranchial copula with a small caudal tab) that clearly supports its assignment to the order Torpediniformes. The analysis of new material also demonstrates that the previous apparent absence of typical narcinoid characters used to diagnose †Titanonarke was the result of taphonomic biases. †Titanonarke shares at least three synapomorphies (presence of a rostral fontanelle, low number of ribs, and rostral cartilage connected to the antorbital cartilage through lateral appendices) with the extant genera Benthobatis, Diplobatis, Discopyge and Narcine, with which it forms a clade (family Narcinidae) recognized herein as unquestionably monophyletic. Moreover, based upon a single specimen of †Titanonarke that exhibits a unique combination of morphometric and meristic features, a new species of Eocene numbfish, †T. megapterygia sp. nov., is recognized. The presence of several specimens representing different ontogenetic stages of at least two species of numbfishes suggests a close association of this taxon with shallow-water habitats corresponding to coral reefs as hypothesized for the Monte Postale palaeoenvironment. The occurrence of a fossilized marine batoid embryo is reported here for the first time. Moreover, the analysis of the gut contents suggests that the dietary adaptations of †Titanonarke can be related, at least in part, to an opportunistic strategy in the context of abundant larger foraminifera in the Monte Postale palaeobiotope, suggesting that this kind of feeding mode, known to occur in present-day reefs, already was realized 50 million years ago.http://zoobank.org/urn:lsid:zoobank.org:pub:EDD6E170-CA64-4FFB-8DD1-AED2D61D5504

New material of three-dimensionally phosphatized and microscopic cycloneuralians from the Cambrian Paibian Stage of South China

AbstractSome rare microscopic cycloneuralians are present in the Cambrian of South China, represented by Eopriapulites and Eokinorhynchus (both early Cambrian), fossil embryos of Markuelia (middle to late Cambrian), and palaeoscolecids (early to late Cambrian). Among them, palaeoscolecids are relatively diverse and abundant. Here, we describe new material of three-dimensionally phosphatized and microscopic cycloneuralians from the Paibian Stage of Wangcun Lagerstätte, western Hunan, South China. New material includes fossil embryos assignable to Markuelia sp., two other types of fossil embryos, and three species of palaeoscolecids, including Dispinoscolex decorus Duan, Dong, and Donoghue, 2012, Schistoscolex hunanensis Duan, Dong, and Donoghue, 2012, and Austroscolex sinensis new species. The palaeoscolecid fragments differ mainly in size and armor of the trunk annuli. Since Eokinorhynchus and Eopriapulites occurred the earliest among the Cambrian cycloneuralians, it is proposed here that: (1) cycloneuralians originated in the Cambrian Fortunian small shelly faunas rather than in the early Cambrian macrobenthos, (2) ancestral cycloneuralians may have simple trunk armor, and (3) Eopriapulites represents an ancestral cycloneuralian.

IDENTIFICATION AND MODES OF ACTION OF ENDOGENOUS BACTERIA IN TAPHONOMY OF EMBRYOS AND LARVAE

We use experimental taphonomy of embryos and larvae to determine the mechanisms by which endogenous bacteria stabilize rather than destroy soft tissues. Here, we show that bacteria can rapidly move from one dead organism to another through the surrounding medium—donor and recipient tissue need not be touching. In most cases tissue destruction results, but in some cases the bacteria generate stabilizing biofilms on recipient tissue that preserve its shape. We isolated, cultured, and identified phylogenetically diverse bacterial strains from the endogenous microbiotas of brine shrimp larvae ( Artemia sinica ) and sea urchin embryos ( Heliocidaris erythrogramma ) that had been killed anaerobically and then incubated aerobically to allow proliferation of endogenous bacteria in the dead tissue. We found that one Artemia -derived isolate, Marinobacter sp., is a potent generator of stabilizing biofilms and that other isolates can participate in biofilm formation or cause destruction. These results show the relative frequency of stabilizing bacteria in the endogenous microbiotas of newly dead organisms. Their ease of transmission reveals the potential for generation of a shared microbiology among groups of dead organisms, a possible contributor to uniform preservation in fossil assemblages such as Neoproterozoic and Cambrian fossil embryos.

Exceptional preservation of tiny embryos documents seed dormancy in early angiosperms.

The rapid diversification of angiosperms through the Early Cretaceous period, between about 130-100 million years ago, initiated fundamental changes in the composition of terrestrial vegetation and is increasingly well understood on the basis of a wealth of palaeobotanical discoveries over the past four decades and their integration with improved knowledge of living angiosperms. Prevailing hypotheses, based on evidence both from living and from fossil plants, emphasize that the earliest angiosperms were plants of small stature with rapid life cycles that exploited disturbed habitats in open, or perhaps understorey, conditions. However, direct palaeontogical data relevant to understanding the seed biology and germination ecology of Early Cretaceous angiosperms are sparse. Here we report the discovery of embryos and their associated nutrient storage tissues in exceptionally well-preserved angiosperm seeds from the Early Cretaceous. Synchrotron radiation X-ray tomographic microscopy of the fossil embryos from many taxa reveals that all were tiny at the time of dispersal. These results support hypotheses based on extant plants that tiny embryos and seed dormancy are basic for angiosperms as a whole. The minute size of the fossil embryos, and the modest nutrient storage tissues dictated by the overall small seed size, is also consistent with the interpretation that many early angiosperms were opportunistic, early successional colonizers of disturbance-prone habitats.

Macroevolutionary developmental biology: Embryos, fossils, and phylogenies.

The field of evolutionary developmental biology is broadly focused on identifying the genetic and developmental mechanisms underlying morphological diversity. Connecting the genotype with the phenotype means that evo-devo research often considers a wide range of evidence, from genetics and morphology to fossils. In this commentary, we provide an overview and framework for integrating fossil ontogenetic data with developmental data using phylogenetic comparative methods to test macroevolutionary hypotheses. We survey the vertebrate fossil record of preserved embryos and discuss how phylogenetic comparative methods can integrate data from developmental genetics and paleontology. Fossil embryos provide limited, yet critical, developmental data from deep time. They help constrain when developmental innovations first appeared during the history of life and also reveal the order in which related morphologies evolved. Phylogenetic comparative methods provide a powerful statistical approach that allows evo-devo researchers to infer the presence of nonpreserved developmental traits in fossil species and to detect discordant evolutionary patterns and processes across levels of biological organization.

Experimental Decay of Soft Tissues

The exceptionally preserved fossil record of soft tissues sheds light on a wide range of evolutionary episodes from across geological history. Understanding how soft tissues become hard fossils is not a trivial process. A powerful tool in this context is experimentally derived decay data. By studying decay in a laboratory setting and on a laboratory timescale, an understanding of the processes and patterns underlying soft-tissue preservation can be achieved. The considerations and problems particular to experimental decay are explored here in terms of experimental aims, design, variables, and utility. Aims in this context can relate to either reconstruction of the processes of soft-tissue preservation, or to elucidation of the patterns of morphological transformation and data loss occurring during decay. Experimental design is discussed in terms of hypotheses and relevant variables: i.e., the subject organism being decayed (phylogeny, ontogeny, and history), the environment of decay (biological, chemical, and physical) and the outputs (how to measure decay). Variables and practical considerations are illustrated with reference to previous experiments. The principles behind application of experimentally derived decay data to the fossil record are illustrated with three case studies: the interpretation of fossil color, feasibility of fossil embryos, and phylogenetic bias in chordate preservation. A rich array of possibilities for further decay experiments exists and it is hoped that the methodologies outlined herein will provide guidance and a conceptual framework for future studies.

The oldest known priapulid-like scalidophoran animal and its implications for the early evolution of cycloneuralians and ecdysozoans.

Morphological phylogenetic analyses suggest that scalidophorans (priapulids, loriciferans, and kinorhynchs) and nematoids (nematodes and nematomorphs) form the ecdysozoan clade Cycloneuralia, which is a sister group to panarthropods. It has been proposed that extant priapulids and Cambrian priapulid-like scalidophorans, because of their conserved evolution, have the potential to illuminate the ancestral morphology, ecology, and developmental biology of highly derived ecdysozoans such as nematods and arthropods. As such, Cambrian fossils, particularly Markuelia and possibly olivooids, can inform the early evolution of scalidophorans, cycloneuralians, and ecdysozoans. However, the scalidophoran Markuelia is known exclusively as embryo fossils, and the olivooids have been alternatively interpreted as cnidarians or cycloneuralians. Here, we describe a post-embryonic scalidophoran fossil Eopriapulites sphinx new genus and species, which represents the oldest known scalidophoran, from the early Cambrian Period (∼535 Ma) in South China. E. sphinx is similar to modern scalidophorans in having an introvert armed with hollow scalids, a collar with coronal scalids, and a pharynx with pharyngeal teeth, but its scalids and pharyngeal teeth are arranged in a hexaradial pattern. Phylogenetically resolved as a stem-group scalidophoran, E. sphinx shares a hexaradial pattern with the hexaradial arrangement of certain anatomical structures in kinorhynchs, loriciferans, nematoids, and Cambrian fossils such as Eolympia pediculata, which could also be a scalidophoran. Thus, the bodyplan of ancestral cycloneuralians may have had a component of hexaradial symmetry (i.e., some but not necessarily all anatomical parts are hexaradially arranged). If panarthropods are nested within paraphyletic cycloneuralians, as several molecular phylogenetic analyses suggest, the ancestral ecdysozoans may have been a legless worm possibly with a component of hexaradial symmetry.