Yanjiahella Biscarpa
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''Yanjiahella biscarpa'' is an extinct species of
Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and th ...
and
Early Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
deuterostome which may represent the earliest stem group
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea ...
. This species is known from the
Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and th ...
to
Fortunian The Fortunian age marks the beginning of the Phanerozoic Eon, the Paleozoic Era, and the Cambrian Period. It is the first of the two stages of the Terreneuvian series. Its base is defined as the first appearance of the trace fossil ''Treptichnus p ...
Yanjiahe Formation The Yanjiahe Formation is a Cambrian fossiliferous geologic formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics ( lithology) that distinguishes it from adjacent bodies of ...
(~541.0–534.6 Ma) in Hubei province, China and was first described by Guo et al. who had difficulty in assigning a taxonomy to the animal due to the shared nature of its features between the
hemichordate 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 ...
s and echinoderms.


Etymology

The ''Yanjiahella'' genus takes its name from the
Yanjiahe Formation The Yanjiahe Formation is a Cambrian fossiliferous geologic formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics ( lithology) that distinguishes it from adjacent bodies of ...
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. 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''. 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''.


Description

''Yanjiahella biscarpa'' is described from 35 specimens. ''Yanjiahella biscarpa'' was a small,
bilateral Bilateral may refer to any concept including two sides, in particular: * Bilateria, bilateral animals *Bilateralism, the political and cultural relations between two states *Bilateral, occurring on both sides of an organism ( Anatomical terms of ...
deuterostome Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some exampl ...
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. 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.


Taxonomic classification

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. tentatively placed the Yanjiahella genus, including ''Y. biscarpa'' in the echinoderm family, and later investigation by Topper et al. placed ''Y. biscarpa'' in order echinodermata based on the nature of the theca and
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis which indicated ''Y. biscarpa'' was a basal echinoderm. Topper et al. proposed that ''Y. biscarpa'' was a basal member of the echinoderm family sharing traits with the last common ambicularian 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.


Dispute over classification

Zamora et al. dispute the placement of ''Y. biscarpa'' with the echinoderms based on the lack of features typifying echinoderms: a
stereom 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 b ...
, 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. 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 group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
s 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. was conducted and no evidence of stereom was observed. Topper et al. pointed to the lack of observed stereom structures in other Cambrian echinoderm fossils due to
taphonomic Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientist Ivan Efremov t ...
factors, and thus the lack of a stereom observed in ''Y. biscarpa'' is not evidence of absence.


Significance for echinoderm evolution

''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. ''Yanjiahella biscarpa'' provides further evidence that echinoderms evolved their familiar pentaradial symmetry after their emergence as a distinct group in the early Cambrian.


References

{{Taxonbar, from= Q105686771 Prehistoric deuterostomes Prehistoric echinoderms Fortunian genus first appearances Cambrian animals of Asia Cambrian China Paleontology in Hubei Fossil taxa described in 2019 Ediacaran life Cambrian genus extinctions