Banffia Temporal range: Middle Cambrian | |
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Fossil of B. constricta from the Burgess Shale | |
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Fossil of B. episoma from the Spence Shale | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | † Vetulicolia |
Class: | † Heteromorphida |
Order: | † Banffiata |
Family: | † Banffidae |
Genus: | † Banffia Walcott, 1911 |
Type species | |
†Banffia constricta Walcott, 1911 | |
Species | |
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Banffia is a genus of animals described from Middle Cambrian fossils. The genus commemorates Banff, Alberta, near where the first fossil specimens were discovered. Its placement in higher taxa is controversial. It is considered to be a member of the enigmatic phylum Vetulicolia.
Banffia constricta is known from hundreds of fossils found in the Burgess Shales. It is up to 10 cm in length, and divided equally into anterior and posterior parts. The entire body is twisted in a clockwise spiral, as seen from the front. This is believed to be a secondary adaptation from an initial bilateral condition for a burrowing lifestyle. The anterior section is covered by two carapace-like un-mineralized shells that are fused together. A crown-like structure formed of three concentric circular features surrounds the mouth. An antenna-form structure just posterior to the mouth may be a sensory organ. The posterior section is composed of 40 to 50 segments. The gut is straight, and the anus is at the terminal tip of the posterior section. The gut appears to have a series of diverticula or pouches. [1] A possible circulatory system is visible in the fossils. [2] B. constricta and its relative Skeemella were probably filter or deposit feeders.
There is no agreement on the classification of Banffia. B. constricta was assigned to the annelids by Walcott in 1911. [3] As of 2006, different proposals would place Banffia in Vetulicolia, [1] [4] Arthropoda, [2] or Urochordata. While the body plan (equal anterior and posterior sections with segmentation) resembles that of the Vetulicolians, it is argued that the absence of gills and an endostyle, and the presence of gut diverticula makes Banffia unlikely to be a member of the deuterostomes. However, an apparent complete lack of appendages (aside from the antenna-like structures) makes B. constricta's placement within Arthropoda equally unlikely. More recent finds of similarly shaped animals show structures thought to be a notochord, making Banffia and their relatives chordates, possibly the sister group to tunicates. [5]
The species Banffia confusa, known from fossils from the Chengjiang shales, was originally assigned to this genus, [6] though, recent research shows that this species is not closely related to B. constricta, having been renamed as its junior synonym, Heteromorphus longicaudatus, and placed in a different class altogether, "Heteromorphida." [7] Later, Aldridge, et al. resurrected B. confusa as Heteromorphus confusus, noting a variety of anatomical differences among the various specimens of Heteromorphus, suggesting that there may be more species recognized with more fossils discovered. [8]
Vetulicolia is a phylum of 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. The vetulicolian body comprises two parts: a voluminous anterior forebody, tipped with an anteriorly positioned mouth and lined with a row of five round to oval-shaped features on each lateral side, which have been interpreted as gills ; and a posterior section that primitively comprises seven segments and functions as a tail. All vetulicolians lack preserved appendages of any kind, having no legs, feelers or even eyes. The area where the anterior and posterior parts join is constricted.
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.
Opabinia regalis is an extinct, stem group arthropod found in the Middle Cambrian Burgess Shale Lagerstätte of British Columbia. Opabinia was a soft-bodied animal, measuring up to 7 cm in body length, and its segmented trunk had flaps along the sides and a fan-shaped tail. The head shows unusual features: five eyes, a mouth under the head and facing backwards, and a clawed proboscis that probably passed food to the mouth. Opabinia probably lived on the seafloor, using the proboscis to seek out small, soft food. Fewer than twenty good specimens have been described; 3 specimens of Opabinia are known from the Greater Phyllopod bed, where they constitute less than 0.1% of the community.
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.
Yunnanozoon lividum is an extinct species of possible vertebrate or chordate from the Lower Cambrian, Chengjiang biota of Yunnan province, China. It is thought of as a deuterostome suspected of being either a hemichordate or chordate. In 2022, a study reanalyzed fossils of Yunnanozoon and found it to be one of the earliest members of the vertebrate family tree.
Pikaia gracilens is an extinct species of primitive chordate animal known from the Middle Cambrian Burgess Shale of British Columbia. Described in 1911 by Charles Doolittle Walcott as an annelid, and in 1979 by Harry B. Whittington and Simon Conway Morris as a chordate, it became "one of the most famous early chordate fossils," or "famously known as the earliest described Cambrian chordate". It is estimated to have lived during the latter period of the Cambrian explosion. Since its initial discovery, more than a hundred specimens have been recovered.
Vetulicola is an extinct genus of marine animal discovered from the Cambrian of China. It is the eponymous member of the enigmatic phylum Vetulicolia, which is of uncertain affinities but may belong to the deuterostomes. The name was derived from Vetulicola cuneata, the first species described by Hou Xian-guang in 1987 from the Lower Cambrian Chiungchussu Formation in Chengjiang, China.
Pomatrum is an extinct vetulicolian, the senior synonym of Xidazoon; the latter taxon was described by Shu, et al. (1999) based on fossils found in the Qiongzhusi (Chiungchussu) Formation, Yu'anshan Member, Lower Cambrian, Haikou, (Kunming), about 50 km west of Chengjiang, China. It has been likened to the chordate Pipiscius.
Leanchoilia is a megacheiran arthropod known from Cambrian deposits of the Burgess Shale in Canada and the Chengjiang biota of China.
Naraoia is a genus of small to average size marine arthropods within the family Naraoiidae, that lived from the early Cambrian to the late Silurian period. The species are characterized by a large alimentary system and sideways oriented antennas.
Misszhouia is a genus of small to average sized marine trilobite-like arthropods within the Naraoiidae family, that lived during the early Cambrian period. The species are M. longicaudata, from the Maotianshan Shales, described in 1985, and M. canadensis, from the Burgess Shale and described in 2018, although later species may belongs to genus Naraoia instead.
A number of assemblages bear fossil assemblages similar in character to that of the Burgess Shale. While many are also preserved in a similar fashion to the Burgess Shale, the term "Burgess Shale-type fauna" covers assemblages based on taxonomic criteria only.
Haikoucaris is a genus of megacheiran arthropod that contains the single species Haikoucaris ercaiensis. It was discovered in the Cambrian Chengjiang biota of China.
Primicaris is genus of Cambrian arthropod from the Chengjiang biota of China and the Burgess Shale of Canada. It contains a single described species, P. larvaformis.
Vetulicola cuneata is a species of extinct animal from the Early Cambrian Chengjiang biota of China. It was described by Hou Xian-guang in 1987 from the Lower Cambrian Chiungchussu Formation, and became the first animal under an eponymous phylum Vetulicolia.
Ooedigera peeli is an extinct vetulicolian from the Early Cambrian of North Greenland. The front body was flattened horizontally, oval-shaped, likely bearing a reticulated or anastomosing pattern, and had 5 evenly-spaced gill pouches along the midline. The tail was also bulbous and flattened horizontally, but was divided into 7 plates connected by flexible membranes, allowing movement. Ooedigera likely swam by moving side-to-side like a fish. It may have lived in an oxygen minimum zone alongside several predators in an ecosystem based on chemosynthetic microbial mats, and was possibly a deposit or filter feeder living near the seafloor.
Skeemella clavula is an elongate animal from what is now the Middle Cambrian Wheeler Shale and Marjum lagerstätte of Utah. It has been classified with the vetulicolians.
Vetulicola rectangulata is a species of extinct animal from the Early Cambrian of the Chengjiang biota of China. Regarded as a deuterostome, it has characteristic rectangular anterior body on which the posterior tail region is attached. It was described by Luo Huilin and Hu Shi-xue in 1999.
Nesonektris aldridgei is an extinct deuterostome chordate from the Late Botomian-aged Emu Bay Shale Lagerstätte in Kangaroo Island, Australia. So far, it is the fourth described vetulicolian that is not restricted to the Maotianshan Shales.
The Cambrian chordates are an extinct group of animals belonging to the phylum Chordata that lived during the Cambrian, between 485 and 538 million years ago. The first Cambrian chordate known is Pikaia gracilens, a lancelet-like animal from the Burgess Shale in British Columbia, Canada. The discoverer, Charles Doolittle Walcott, described it as a kind of worm (annelid) in 1911, but it was later identified as a chordate. Subsequent discoveries of other Cambrian fossils from the Burgess Shale in 1991, and from the Chengjiang biota of China in 1991, which were later found to be of chordates, several Cambrian chordates are known, with some fossils considered as putative chordates.