Cambroernids Temporal range: | |
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Herpetogaster | |
Eldonia | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Stem group: | Ambulacraria |
Clade: | † Cambroernida Caron, Conway Morris, & Shu, 2010 |
Subdivisions | |
The Cambroernida are a clade of unusual Paleozoic animals with coiled bodies and filamentous tentacles. They include a number of early to middle Paleozoic (Cambrian to Devonian) [1] 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," [2] later work with better-preserved fossils has strengthened the argument for Cambroernida as a monophyletic clade. [3]
Cambroernids encompass three particular types of enigmatic animals first appearing in the Cambrian: Herpetogaster (the type genus), Phlogites , and the Eldonioidea. They are united by a set of common features including at least one pair of bifurcated or divided oral tentacles, and a large stomach and narrower intestine enclosed together in a clockwise-coiled sac. [2]
Body coiling increased throughout this group's evolution. [4] Herpetogaster has a segmented and clockwise-curved body attached to the substrate via a narrow and partially mobile stolon (stalk). Phlogites was even more simple, with a thick immobile stolon leading up to a tentacle-bearing calyx (cup-shaped main body), with internal gut coiling. The eldoniids [2] [5] (also known as eldonioids [6] [7] [8] or eldonids [1] [7] ) were diverse and disc-shaped, commonly described as "medusiform", i.e. jellyfish-shaped. Though the lifestyle of eldoniids is still debated, it can be agreed that they had a large curved stomach and no stolon. [6] [9] [10] [7]
The lack of a post-anal tail in cambroernids suggests that, contrary to long-held assumptions, this feature was not present in the common ancestor of deuterostomes. This is congruent with the significant differences between the post-anal tails of chordates and hemichordates. This and other features of cambroernids suggest that post-anal tails, gill bars, and a U-shaped gut evolved multiple times in the deuterostomes through convergence. [11]
Segmentation, as seen in Herpetogaster, is a notable characteristic of chordates not seen in other ambulacrarians, indicating that it might be a trait of ancestral deuterostomes. [11]
Phylogenetic analysis offers strong support for Cambroernida as a clade of stem-group ambulacrarians. [4] The following cladogram is simpllified from Li et al. 2023; only a sampling of eldonioids were included in the analysis: [12]
Ambulacraria |
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A "?" preceding the name indicates the placement in the parent taxon is tentative:
Note that some authors continue to treat Stellostomites as a separate taxon. [14]
Previously, some cambroernids were compared to members of the broad invertebrate clade Lophotrochozoa. In particularl, they were allied with the lophophorates, a subset of lophotrochozoans bearing ciliated tentacles known as lophophores. [6] [15] However, this interpretation has more recently been considered unlikely relative to the deuterostome hypothesis for cambroernid origins. [2]
Hyoliths are animals with small conical shells, known from fossils from the Palaeozoic era. They are at least considered as being lophotrochozoan, and possibly being lophophorates, a group which includes the brachiopods, while others consider them as being basal lophotrochozoans, or even molluscs.
Vetulicolia is a group of bilaterian marine animals encompassing several extinct species from the Cambrian, and possibly Ediacaran, periods. As of 2023, the majority of workers favor placing Vetulicolians in the stem group of the Chordata, but some continue to favor a more crownward placement as a sister group to the Tunicata. It was initially erected as a monophyletic clade with the rank of phylum in 2001, with subsequent work supporting its monophyly. However, more recent research suggests that vetulicolians may be paraphyletic and form a basal evolutionary grade of stem chordates.
The Maotianshan Shales (帽天山页岩) are a series of Early Cambrian sedimentary deposits in the Chiungchussu Formation, famous for their Konservat Lagerstätten, deposits known for the exceptional preservation of fossilized organisms or traces. The Maotianshan Shales form one of some forty Cambrian fossil locations worldwide exhibiting exquisite preservation of rarely preserved, non-mineralized soft tissue, comparable to the fossils of the Burgess Shale of British Columbia, Canada. They take their name from Maotianshan Hill in Chengjiang County, Yunnan Province, China.
Nectocaris is a genus of squid-like animal of controversial affinities known from the Cambrian period. The initial fossils were described from the Burgess Shale of Canada. Other similar remains possibly referrable to the genus are known from the Emu Bay Shale of Australia and Chengjiang Biota of China.
Cathaymyrus is a genus of Early Cambrian chordate known from the Chengjiang biota in Yunnan Province, China. Both species have a long segmented body with no distinctive head. The segments resemble the v-shaped muscle blocks found in cephalochordates such as Amphioxus. A long linear impression runs along the "back" of the body looking something like a chordate notochord.
Dinomischusis an extinct genus of stalked filter-feeding animals within the Cambrian period, with specimens known from the Burgess Shale and the Maotianshan Shales. While long of uncertain affinities, recent studies have suggested it to be a stem-group ctenophore.
Didazoonidae is a vetulicolian family within the order Vetulicolata. It is charaterized by a relatively thin-walled, non-biomineralized body and a large, round anterior opening surrounded by an oral disc. It may be paraphyletic, even if the phylum Vetulicolia is monophyletic.
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 four phyla: Chordata, Echinodermata, Hemichordata, and the extinct Vetulicolia known from Cambrian fossils. The extinct clade Cambroernida is thought to be a member of Deuterostomia.
Eldonia is an extinct soft-bodied cambroernid animal of unknown affinity, best known from the Fossil Ridge outcrops of the Burgess Shale, particularly in the 'Great Eldonia layer' in the Walcott Quarry. In addition to the 550 collected by Walcott, 224 specimens of Eldonia are known from the Greater Phyllopod bed, where they comprise 0.43% of the community. Species also occur in the Chengjiang biota, Siberia, and in Upper Ordovician strata of Morocco.
Herpetogaster is an extinct cambroernid genus of animal from the Early Cambrian Chengjiang biota of China, Blang Formation of China, Pioche Formation of Nevada and Middle Cambrian Burgess Shale of Canada containing the species Herpetogaster collinsi and Herpetogaster haiyanensis.
Eldonioids are an extinct clade of enigmatic disc-shaped animals which lived in the early to middle Paleozoic. The terms "eldonioid" and "eldoniid" have been used somewhat informally and interchangeably, but technically refer to members of the class Eldonioidea and the family Eldoniidae, respectively. The lifestyle of eldonioids is still an unresolved question; some authors reconstruct eldonioids as free-floating planktonic predators similar to jellyfish, while others argue that they were passive detritivores, embedded within the seabed for much of their life.
Stellostomites is a discoidal animal known from the Cambrian Chengjiang biota. It is classified with the eldoniids, and considered a junior synonym of Eldonia by some authors.
Paropsonema is a discoidal animal known from the Cambrian Chengjiang biota and possibly from Devonian formations in upstate New York. It is classified with the eldonioids.
Rotadiscus is a genus of discoidal animal known from the Cambrian Chengjiang biota and classified with the eldonioids.
Phlogites is a member of the extinct ambulacrarian stem group Cambroernida, occupying an intermediate position between the basal Herpetogaster and the more derived Eldonioidea. It is known from the Lower Cambrian Haikou Chengjiang deposits of China.
Cotyledion tylodes is an extinct, stalked filter-feeder known from the Chengjiang lagerstatten. The living animal reached a couple of centimetres in height, and bore a loose scleritome of ovoid sclerites. Its interpretation has been controversial, but it is currently thought to be a member of the Entoprocta stem group.
Shankouclava is an extinct genus of tunicates. It is one of two candidates for the oldest member of this group, dating to 518 million years ago. It has been found in the Lower Cambrian Maotianshan Shale at Shankou village, Anning, near Kunming. Each of the eight specimens found and used for description were isolated, suggesting that the genus was solitary and not colonial.
Vetulocystidae is the only family of the taxon Vetulocystida, which is a group of extinct deuterostomes of uncertain phylogenetic position. Vetulocystidae is made up of the genera Vetulocystis, Dianchicystis and Thylacocercus.
Lenisambulatrix is a genus of extinct worm belonging to the group Lobopodia and known from the Lower Cambrian Maotianshan shale of China. It is represented by a single species L. humboldti. The incomplete fossil was discovered and described by Qiang Ou and Georg Mayer in 2018. Due to its missing parts, its relationship with other lobopodians is not clear. It shares many structural features with another Cambrian lobopodian Diania cactiformis, a fossil of which was found alongside it.
Cheungkongella is a fossil organism from the lower Cambrian Chengjiang lagerstatte, the affinity of which has been the subject of debate. It was announced as a "probable" tunicate while noting the lack of definitive Cambrian fossils from that group. However, this affinity was later disputed in a paper announcing the discovery of Shankouclava, also from Chengjiang, as the oldest known tunicate. Cheungkongella has been accepted as a distinct taxon and possible tunicate by multiple workers not involved in its discovery, but the dispute remains unresolved.