Yanjiahella Temporal range: Fortunian ~ | |
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Fossil Y. biscarpa from the Ediacaran and Fortunian of China. Scale bar=5mm | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Echinodermata (?) |
Genus: | † Yanjiahella |
Species: | †Y. biscarpa |
Binomial name | |
†Yanjiahella biscarpa (Guo et al., 2012) | |
Yanjiahella biscarpa is an extinct species of Early Cambrian deuterostome which may represent the earliest stem group echinoderm. [1] [2]
This species is known from the Fortunian Yanjiahe Formation (~541.0–534.6 Ma) in Hubei province, China and was first described by Guo et al. [3] who had difficulty in assigning a taxonomy to the animal due to the shared nature of its features between the hemichordates and echinoderms.
The Yanjiahella genus takes its name from the Yanjiahe Formation combined with the Latin diminutive "-ella" in reference to the small size of the species with the species name biscarpa deriving from the Latin "bis" meaning two, and Greek "carpa" meaning arms. [3]
Guo et al. originally divided the Yanjiahella genus into three species based on the number of feeding appendages, Y. ancarpa (meaning "no armed") Y. monocarpa (meaning "single armed") and Y. biscarpa. [3] However, analysis by Topper et al. showed identical morphological features between the three initial species, and that the varying number of these appendages was due to their lack of preservation in some specimens and joined all species identified by Guo et al. as Y. biscarpa. [1]
Yanjiahella biscarpa is described from 35 specimens. Yanjiahella biscarpa was a small, bilateral deuterostome animal typically 20-50 mm in length with a stem, theca and two feeding appendages. The stem is divided into two parts, distal and proximal. The proximal stem displays transverse ridges and a medial ridge interpreted as a linear digestive tract, with the anus interpreted as located above the junction between the proximal and distal stem. The proximal stem was likely lightly mineralized, aiding in support for the theca. The distal part of the stem is typically curved with lineations indicative of musculature and interpreted as a holdfast used to anchor the animal either erect or prostrate in the sediment. [1] [3]
The theca of Y.biscarpa was covered in small ovoid to polygonal plates typically 1-2 mm in length arranged in an uneven distribution. The feeding appendages emerge from opposite sides of the theca, and extend 15-20 mm with one specimen possessing arms 43 mm in length. The nature of the feeding appendages is ambiguous, with no conclusive observations supporting a muscular, collagenous or vascular make up. The mouth is interpreted as being located between the feeding appendages on the body though it has not been observed in any specimen. [3]
The classification of Y. biscarpa is difficult due to sharing features with the sister phylum hemichordates, most notably the muscled stalk and linear digestive tract. However, the presence of the plated body structure has led to its placement with other echinoderms. Guo et al. [3] tentatively placed the Yanjiahella genus, including Y. biscarpa in the echinoderm family, and later investigation by Topper et al. [1] placed Y. biscarpa in order echinodermata based on the nature of the theca and phylogenetic analysis which indicated Y. biscarpa was a basal echinoderm. Topper et al. [1] proposed that Y. biscarpa was a basal member of the echinoderm family sharing traits with the last common ambulacrarian ancestor. Later early echinoderms such as the ctenocystoids lost the muscular stalk of these basal echinoderms, while maintaining bilateral form before evolving the familiar pentaradial symmetry of later echinoderms, indicating that the bilateralism was a common trait among early echinoderms. [1] [4]
Zamora et al. [2] dispute the placement of Y. biscarpa with the echinoderms based on the lack of features typifying echinoderms: a stereom, pentaradial symmetry, and a water vascular system. These authors additionally conducted an independent phylogenetic analysis, which placed Y. biscarpa with the hemichordates. Topper et al. [4] responded to these criticisms by acknowledging the difficulty of interpreting early Cambrian fossils, while noting that the common basal stem animals of the Cambrian often lacked the typical features of their crown groups early in their evolutionary stages, and thus interpreting the fossil based these features is unwarranted. Additional analysis of specimens to identify stereom structures by Topper et al. [4] was conducted and no evidence of stereom was observed. Topper et al. [4] pointed to the lack of observed stereom structures in other Cambrian echinoderm fossils due to taphonomic factors, and thus the lack of a stereom observed in Y. biscarpa is not evidence of absence.
Yanjiahella biscarpa appears to represent the most basal form of the echinoderms predating other echinoderm groups by several million years, and represents one of the earliest deuterostome animals. The similar features to its ambicularian ancestors is likely to help further elucidate the divergence of echinoderms and their early evolutionary history. [1] Yanjiahella biscarpa provides further evidence that echinoderms evolved their familiar pentaradial symmetry after their emergence as a distinct group in the early Cambrian.
A chordate is a deuterostomic animal belonging to the phylum Chordata. All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics (synapomorphies) that distinguish them from other taxa. These five synapomorphies are a notochord, a hollow dorsal nerve cord, an endostyle or thyroid, pharyngeal slits, and a post-anal tail. The name "chordate" comes from the first of these synapomorphies, the notochord, which plays a significant role in chordate body plan structuring and movements. Chordates are also bilaterally symmetric, have a coelom, possess a closed circulatory system, and exhibit metameric segmentation.
Hemichordata is a phylum which consists of triploblastic, enterocoelomate, and bilaterally symmetrical marine deuterostome animals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and include two main classes: Enteropneusta, and Pterobranchia. A third class, Planctosphaeroidea, is known only from the larva of a single species, Planctosphaera pelagica. The class Graptolithina, formerly considered extinct, is now placed within the pterobranchs, represented by a single living genus Rhabdopleura.
Vetulicolia is a phylum of bilaterian animals encompassing several extinct species belonging to the Cambrian period. The phylum was created by Degan Shu and his research team in 2001, and named after Vetulicola cuneata, the first species of the phylum described in 1987.
Graptolites are a group of colonial animals, members of the subclass Graptolithina within the class Pterobranchia. These filter-feeding organisms are known chiefly from fossils found from the Middle Cambrian through the Lower Carboniferous (Mississippian). A possible early graptolite, Chaunograptus, is known from the Middle Cambrian. Recent analyses have favored the idea that the living pterobranch Rhabdopleura represents an extant graptolite which diverged from the rest of the group in the Cambrian.
Anomalocaris is an extinct genus of radiodont, an order of early-diverging stem-group arthropods.
The stylophorans are an extinct, possibly polyphyletic group allied to the Paleozoic Era echinoderms, comprising the prehistoric cornutes and mitrates. It is synonymous with the subphylum Calcichordata. Their unusual appearances have led to a variety of very different reconstructions of their anatomy, how they lived, and their relationships to other organisms.
Ambulacraria, or Coelomopora, is a clade of invertebrate phyla that includes echinoderms and hemichordates; a member of this group is called an ambulacrarian. Phylogenetic analysis suggests the echinoderms and hemichordates separated around 533 million years ago. The Ambulacraria are part of the deuterostomes, a clade that also includes the many Chordata, and the few extinct species belonging to the Vetulicolia.
Deuterostomes are bilaterian animals of the superphylum Deuterostomia, typically characterized by their anus forming before the mouth during embryonic development. Deuterostomia is further divided into 4 phyla: Chordata, Echinodermata, Hemichordata, and the extinct Vetulicolia known from Cambrian fossils. The extinct clade Cambroernida is also thought to be a member of Deuterostomia.
Schinderhannes bartelsi is a species of hurdiid radiodont (anomalocaridid), known from one specimen from the Lower Devonian Hunsrück Slates. Its discovery was astonishing because the latest definitive radiodonts were known only from the Early Ordovician, at least 66 million years earlier than this taxon.
Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. They may be referred to as radiodonts, radiodontans, radiodontids, anomalocarids, or anomalocaridids, although the last two originally refer to the family Anomalocarididae, which previously included all species of this order but is now restricted to only a few species. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and used for a variety of functions. Radiodonts included the earliest large predators known, but they also included sediment sifters and filter feeders. Some of the most famous species of radiodonts are the Cambrian taxa Anomalocaris canadensis, Hurdia victoria, Peytoia nathorsti, Titanokorys gainessii, Cambroraster falcatus and Amplectobelua symbrachiata. The later surviving members include the subfamily Aegirocassisinae from the Early Ordovician of Morocco and the Early Devonian member Schinderhannes bartelsi from Germany.
Homalozoa is an obsolete extinct subphylum of Paleozoic era echinoderms, prehistoric marine invertebrates. They are also referred to as carpoids.
Pelmatozoa was once a clade of Phylum Echinodermata. It included stalked and sedentary echinoderms. The main class of Pelmatozoa were the Crinoidea which includes sea lily and feather star.
The calcichordate hypothesis holds that each separate lineage of chordate evolved from its own lineage of mitrate, and thus the echinoderms and the chordates are sister groups, with the hemichordates as an out-group.
The cambroernids are a clade of unusual Paleozoic animals with coiled bodies and filamentous tentacles. They include a number of early to middle Paleozoic genera noted as "bizarre" or "orphan" taxa, meaning that their affinities with other animals, living or extinct, have long been uncertain. While initially defined as an "informal stem group," later work with better-preserved fossils has strengthened the argument for Cambroernida as a monophyletic clade.
Stereom is a calcium carbonate material that makes up the internal skeletons found in all echinoderms, both living and fossilized forms. It is a sponge-like porous structure which, in a sea urchin may be 50% by volume living cells, and the rest being a matrix of calcite crystals. The size of openings in stereom varies in different species and in different places within the same organism. When an echinoderm becomes a fossil, microscopic examination is used to reveal the structure and such examination is often an important tool to classify the fossil as an echinoderm or related creature.
The Yanjiahe Formation is an Ediacaran to Cambrian fossiliferous geologic formation found in South China.
Cincta is an extinct class of echinoderms that lived only in the Middle Cambrian epoch. Homostelea is a junior synonym. The classification of cinctans is controversial, but they are probably part of the echinoderm stem group.
Ctenocystoidea is an extinct clade of echinoderms, which lived during the Cambrian and Ordovician periods. Unlike other echinoderms, ctenocystoids had bilateral symmetry, or were only very slightly asymmetrical. They are believed to be one of the earliest-diverging branches of echinoderms, with their bilateral symmetry a trait shared with other deuterostomes. Ctenocystoids were once classified in the taxon Homalozoa, also known as Carpoidea, alongside cinctans, solutes, and stylophorans. Homalozoa is now recognized as a polyphyletic group of echinoderms without radial symmetry. Ctenocystoids were geographically widespread during the Middle Cambrian, with one species surviving into the Late Ordovician.
Laminacaris is a genus of extinct stem-group arthropods (Radiodonta) that lived during the Cambrian period. It is monotypic with a single species Laminacaris chimera, the fossil of which was described from the Chengjiang biota of China in 2018. Around the same time, two specimens that were similar or of the same species were discovered at the Kinzers Formation in Pennsylvania, USA. The first specimens from China were three frontal appendages, without the other body parts.
Gyaltsenglossus is a monospecific hemichordate known from the Burgess Shale of Canada that is notable for advancing the understanding of the early evolution of the phylum Hemichordata. Its discovery has been hailed as a "breakthrough" due to featuring both Pterobranch-like feeding tentacles and an Enteropneust-like proboscis-tipped elongate body, thus uniting the two morphologically disparate orders of the Hemichordata. A phylogenetic analysis recovered Gyaltsenglossus senis as the first known stem group hemichordate.