Heteromorphus Temporal range: Cambrian Stage 3 | |
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Artist's reconstruction of Heteromorphus longicaudatus (with much smaller "Form A" above it) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade?: | † Vetulicolia |
Class: | † Banffozoa |
Order: | † Banffiata |
Family: | † Banffiidae |
Genus: | † Heteromorphus Luo and Hu in Luo et al. 1999 |
Type species | |
†Heteromorphus longicaudatus Luo and Hu in Luo et al. 1999 | |
Species | |
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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. [1] 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. [2]
Like Banffia, Heteromorphus has a two-part body with a notable constriction between the parts, and a crossover that effectively reverses the dorsal and ventral sides between the anterior and posterior sections. [3] The posterior portion is segmented, although the common presence of wrinkling makes counting the segments difficult. The anterior body shape ranges from torpedo-like to more rectangular, with a near-vertical anterior edge. [1]
Heteromorphus is separated from Banffia by the presence of a lateral groove, similar to that found in the Vetulicolida. [1] However, while a 2004 description claimed that four gill openings were present in the groove, later workers have not observed them. The lack of visible gills in banffiids has been speculated to indicate a burrowing lifestyle. [4] A partial "twist" has been described in Heteromorphus, compared to the well-developed torsion in Banffia. [5] While the initial description of H. confusus indicated an anus halfway along the ventral surface of the posterior section, [6] later descriptions confirmed the terminal location of the anus. [7]
Heteromorphus has been grouped with Banffia and Skeemella in the family Banffiidae. [8]
The following cladogram is simplified from Mussini et al. (2024), using the definition of Vetulicolidae from Li et al. (2018). It shows Heteromorphus as part of the earliest steps, corresponding to the class Banffozoa, in the evolutionary grade leading to extant chordates. [9]
"Banffozoa" "Vetulicolida" |
An earlier study in 2014 placed vetulicolians as the sister-group to tunicates, but was unable to resolve any relationships among vetulicolians as a group: [5]
Vetulicolia† | |
H. confusus was originally described as Banffia confusa in 1997, while H. longicaudatus was described and assigned to the new genus Heteromorphus in 1999. The discoverers of B. confusa referred to it as H. confusus in 2002, accepting its assignment to Heteromorphus and, according to a comprehensive 2007 review of vetulicolian phylogenetics, indicating that H. longicaudatus is its junior synonym. However, the same review also noted substantial variation among Heteromorphus specimens, and that more species (including a distinct H. longicaudatus) may exist. [1]
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.
Kunmingella is genus of Cambrian bradoriid from the Chengjiang biota, containing the single species K. douvillei. Kunmingella had 12 appendages, including a pair of antennae as well pairs of biramous limbs, including four anterior pairs of appendages bearing double rows of endites on their endopods, and a posterior 5 with only a single row of endites, as well as two terminal pairs of uniramous limbs. Eggs have been found preserved attached to the posteriormost three pairs of biramous limbs, suggesting it engaged in brood care. Around 50–80 eggs, each around 150–180 μm across were attached in total.
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.
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 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.
Banfiidae is a family of extinct banffozoan animals from North America and China. The family name is sometimes spelt Banffidae. It includes Banffia, Heteromorphus, and possibly Skeemella. The family may be paraphyletic. The family may be paraphyletic. 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.
Luolishania is an extinct genus of lobopodian panarthropod and known from the Lower Cambrian Chiungchussu Formation of the Chengjiang County, Yunnan Province, China. A monotypic genus, it contains one species Luolishania longicruris. It was discovered and described by Hou Xian-Guang and Chen Jun-Yuan in 1989. It is one of the superarmoured Cambrian lobopodians suspected to be either an intermediate form in the origin of velvet worms (Onychophora) or basal to at least Tardigrada and Arthropoda. It is the basis of the family name Luolishaniidae, which also include other related lobopods such as Acinocricus, Collinsium, Facivermis, and Ovatiovermis. Along with Microdictyon, it is the first lobopodian fossil discovered from China.