Banffiidae Temporal range: about Cambrian Stage 3—Drumian | |
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Fossils and reconstructions of banffiids (with much smaller non-banffiid "Form A" above H. confusus) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade?: | † Vetulicolia |
Class: | † Banffozoa |
Order: | † Banffiata |
Family: | † Banffiidae Caron, 2006 |
Type genus | |
† Banffia Walcott 1911 | |
Genera | |
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Banfiidae is a family of extinct banffozoan animals from North America and China. The family name is sometimes spelt Banffidae. [1] [2] It includes Banffia , Heteromorphus , and possibly Skeemella . [2] The family may be paraphyletic. The family may be paraphyletic. [3] A Heteromorphus-like dwarf "Form A" is allied with this group at the class level, but has not been formally described or assigned to this family. [4]
Banffiids have a bipartite body with a smooth anterior part that may or may not have a lateral groove, and a posterior part with many segments. Lateral pouches have not been seen, although some questions remain regarding their possible presence in Skeemella. The entire body is twisted dextrally (from an anterior view). [1] Gut diverticula may be present, but the structures interpreted as such could alternatively be a circulatory system. [5] The anus is terminal, [6] despite initial reports to the contrary in Heteromorphus. [7]
Banffiids have been found across multiple paleocontienents, with Heteromporphus found in the Chengjiang lagerstatte of South China while Banffia has been found in the Burgess Shale and Spence Shale of western Laurentia. [2] Skeemella has been found in the Pierson Cove and Marjum Formations, both above the Wheeler Shale and also in western Laurentia. [8]
The family Banffiidae and the class Banffozoa were erected in a paper published in 2006, [9] in order to contain Banffia constricta in a redescription of that species, with Heteromorphus only mentioned as a possible home outside of Banffiidae for Banffia confusa . [10] A comprehensive phylogenetic revision in 2007 expanded Banffozoa to encompass the proposed class Heteromorphida (which had excluded Banffia), and placed Heteromorphus in the Banffiidae. It tentatively placed Skeemella outside of Banffiidae and the newly-erected order Banffiata, but under Banffozoa. [1] "Form A", which had been included in the Heteromorphida, [4] was not mentioned or reassigned. [1] Two later sources have placed Skemella in Banffiidae while continuing to note doubts as to whether it is actually a vetulicolian. [2] [8]
Banffozoans have been recognized as "rather different from vetulicolians proper," [11] and proposed to be more closely related to protostomes. [12] However, multiple sources treat Banffozoa as part of Vetulicolia. [1] [2] [8]
A 2024 study has found the Banffiidae to form the earliest part of the paraphyletic stem-chordate evolutionary grade, as shown in this simplified cladogram: [13]
"Banffiidae" "Banffiidae" ? |
An earlier study in 2014 placed vetulicolians as the sister-group of tunicates, but was unable to resolve any relationships among vetulicolians as a group: [14]
Vetulicolia† | |
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.
Yunnanozoon lividum is an extinct species of bilaterian animal from the Lower Cambrian Chengjiang biota of Yunnan province, China. Its affinities have been long the subject of controversy.
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.
Didazoon haoae is an extinct species of vetulicolid vetulicolian described by Shu, et al. based on fossils found in the Qiongzhusi (Chiungchussu) Formation, Yu'anshan Member, Lower Cambrian, in the Dabanqiao area (Kunming), about 60 km northwest of 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.
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, with it mostly being considered to be a member of the enigmatic phylum Vetulicolia.
Vetulicolidae is a vetulicolian family from the Cambrian Stage 3 Maotianshan Shale and Sirius Passet Lagerstätte that consists of Vetulicola, Beidazoon, and Ooedigera. It is distinguished from the Didazoonidae by a harder body wall and the lack of an oral disc.
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.
Banffozoa is an extinct class of bilaterians. Most workers place it in the Vetulicolia, but the protostome-like features of some members have motivated ongoing debate. Banffozoa consists of the order Banffiata as well as a dwarf "Form A" that has not been formally described or named. Skeemella has been placed incertae sedis in this class, but has more recently been placed with the Banffiidae. Banffozoa may be paraphyletic even if Vetulicolia is monophyletic.
Yuyuanozoon magnificissimi, from the Cambrian Stage 3 Chengjiang lagerstatte, is the largest known vetulicolian, with specimens up to 20 cm in length compared to 5–14 cm for other vetulicolian species.
The palaeoscolecids are a group of extinct ecdysozoan worms resembling armoured priapulids. They are known from the Lower Cambrian to the lower Ludfordian ; they are mainly found as disarticulated sclerites, but are also preserved in many of the Cambrian lagerstätten. They take their name from the typifying genus Palaeoscolex. Other genera include Cricocosmia from the Lower Cambrian Chengjiang biota. Their taxonomic affinities within Ecdysozoa have been the subject of debate.
Vetulicola cuneata is a species of extinct marine 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 is a genus of elongate animal from the Middle Cambrian Wheeler Shale and Marjum lagerstätte of Utah. It has been classified with the banffozoan vetulicolians.
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.
Nesonektris aldridgei is an extinct vetulicolian 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.
Beidazoon venustum is a marine deuterostome from the group Vetulicolia. It originates from the lower Cambrian Chengjiang biota of Yunnan Province, China, and was discovered in 2005. It is known as the smallest described vetulicolian, and for its surface being covered in many small nodes.
Shenzianyuloma is an extinct genus of vetulicolian represented by a single species, Shenzianyuloma yunnanense, from the Maotianshan Shale during Stage 3 of the Cambrian period. It is notable for having a compact body shape akin to that of an angelfish. It's exact phylogenetic position is unclear, and it was not included in a 2024 phylogenetic analysis of vetulicolians.
Heteromorphus is an extinct genus of banffiid from the lower Cambrian Chengjiang lagerstatte. It contains one broadly accepted species, Heteromorphus confusus, as well as a proposed junior synonym, Heteromorphus longicaudatus that may prove to be a separate species as additional specimens are examined. A much smaller species labeled "Form A" is allied with Heteromorphus at the class level but has not been formally described or assigned to Heteromorphus itself.