Carbotubulus Temporal range: | |
---|---|
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
(unranked): | Panarthropoda |
Phylum: | † "Lobopodia" |
Clade: | † Hallucishaniids |
Family: | † Hallucigeniidae |
Genus: | † Carbotubulus Haug et al., 2012 [1] |
Species: | †C. waloszeki |
Binomial name | |
†Carbotubulus waloszeki Haug et al., 2012 | |
Carbotubulus is a genus of extinct worm belonging to the group Lobopodia and known from the Carboniferous Carbondale Formation of the Mazon Creek area in Illinois, US. A monotypic genus, it contains one species Carbotubulus waloszeki. [1] It was discovered and described by Joachim T. Haug, Georg Mayer, Carolin Haug, and Derek E.G. Briggs in 2012. [2] With an age of about 300 million years, it is the first long-legged lobopodian discovered after the period of Cambrian explosion. [3]
Carbotubulus, represented by a single fossil, was discovered by Joachim T. Haug (University of Greifswald, Germany), Georg Mayer (Leipzig University), Carolin Haug (Yale University, US), and Derek E.G. Briggs (Yale University) from the Carbondale Formation (Francis Creek Shale Member) of the Mazon Creek area in Illinois, US. It was recovered from the location named Pit 11 along with a fossil of an extinct velvet worm Helenodora inopinata (also mentioned as Ilyodes inopinata). [4] The original fossil is kept at the Royal Ontario Museum in Toronto. [1]
The name is derived from Latin words, carbo, referring to the geological age Carboniferous; and tubulus, meaning "small pipe", a description for the pipe-like legs. The specific name is after the German zoologist Dieter Waloszek, one of the leaders in the study of arthropod evolution. [1]
Carbotubulus is a soft-bodied worm with stumpy legs called lobopods. It has nine pairs of lobopods that are tube-like and elongated. Unlike other lobopodians, its head is relatively large and cylindrical in shape, occupying about one-third of the body length. [1] In other related hallucigeniid worms, the head is usually rounded and tiny such that it is often difficult to describe. [5] [6] [7] Living around 300 million years ago, it supports the fact that Cambrian-type worms survived for over 200 million years after extinction events during the Middle Cambrian (between 510 and 502 million years ago) by which most Cambrian animals disappeared. [8] [9] Majority of the long-legged lobopodians such as Hallucigenia , Paucipodia and Orstenotubulus are known only during the Cambrian explosion. [10]
When Carbotubulus was first described, its systematic position was not clear and was loosely assigned to the phylum Arthropoda. Discovery of Cambrian lobopod Ovatiovermis cribratus from the Burgess Shale in 2017 led to a reanalysis of lobopodian classification, and Carbotubulus was assigned to a group Panarthropoda, specifically belonging to the family Hallucigeniidae along with the various species of Hallucigenia and Cardiodictyon catenulum. [11] This classification is still controversial, especially after the discovery of the second post-Cambrian (Silurian) long-legged lobopodian, Thanahita distos , from the Herefordshire Lagerstätte at the England–Wales border in UK in 2018. The new interpretation suggest that Carbotubulus and Cardiodictyon may lie outside the hallucigeniid family. [12]
Lobopodians are members of the informal group Lobopodia, or the formally erected phylum Lobopoda Cavalier-Smith (1998). They are panarthropods with stubby legs called lobopods, a term which may also be used as a common name of this group as well. While the definition of lobopodians may differ between literatures, it usually refers to a group of soft-bodied, marine worm-like fossil panarthropods such as Aysheaia and Hallucigenia.
Hallucigenia is a genus of lobopodian, known from Cambrian aged fossils in Burgess Shale-type deposits in Canada and China, and from isolated spines around the world. The generic name reflects the type species' unusual appearance and eccentric history of study; when it was erected as a genus, H. sparsa was reconstructed as an enigmatic animal upside down and back to front. Lobopodians are a grade of Paleozoic panarthropods from which the velvet worms, water bears, and arthropods arose.
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.
Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada and Onychophora. Panarthropods also include extinct marine legged worms known as lobopodians ("Lobopodia"), a paraphyletic group where the last common ancestor and basal members (stem-group) of each extant panarthropod phylum are thought to have risen. However the term "Lobopodia" is sometimes expanded to include tardigrades and onychophorans as well.
Parapeytoia is a genus of Cambrian arthropod. The type and only described species is Parapeytoia yunnanensis, lived over 518 million years ago in the Maotianshan shales of Yunnan, China. Unidentified fossils from the same genus also had been discovered from the nearby Wulongqing Formation.
Helenodora is an extinct basal onychophoran or lobopodian genus known from the Carboniferous Carbondale Formation of Illinois. The only known species described is H. inopinata. The ecology of this animal is not well known, but it is thought that it may have lived on land and/or underwater.
Megacheira is an extinct class of predatory arthropods defined by their possession of spined "great appendages". Their taxonomic position is controversial, with studies either considering them stem-group euarthropods, or stem-group chelicerates. The homology of the great appendages to the cephalic appendages of other arthropods is also controversial. Uncontested members of the group were present in marine environments worldwide from the lower to middle Cambrian.
Haikoucaris is a genus of megacheiran arthropod that contains the single species Haikoucaris ercaiensis. It was discovered in the Cambrian Chengjiang biota of China.
Cucumericrus ("cucumber-leg") is an extinct genus of stem-arthropod. The type and only species is Cucumericrus decoratus, with fossils discovered from the Maotianshan Shales of Yunnan, China.
Diania is an extinct genus of lobopodian panarthropod found in the Lower Cambrian Maotianshan shale of China, represented by a single species - D. cactiformis. Known during its investigation by the nickname "walking cactus", this organism belongs to a group known as the armoured lobopodians, and has a simple worm-like body with robust, spiny legs. Initially, the legs were thought to have a jointed exoskeleton and Diania was suggested to be evolutionarily close to early arthropods, but many later studies have rejected this interpretation.
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.
The Luolishaniidae or Luolishaniida are a group of Cambrian and Ordovician lobopodians with anterior 5 or 6 pairs of setiferous lobopods. Most luolishaniids also have posterior lobopods each with a hooked claws, and thorn-shaped sclerites arranged as three or more per trunk segment. The type genus is based on Luolishania longicruris Hou and Chen, 1989, from the Chengjiang Lagerstatte, South China. They are presumed to have been benthic suspension or filter feeders.
Collinsium was a genus of lobopodian from the Early Cambrian. It is represented by a single fossil species, Collinsium ciliosum, found in the Xiaoshiba Lagerstätte of China. Similar to the later Hallucigenia it was a small worm-like creature with spikes along its back and feeding tentacles near its head. Unlike Hallucigenia, Collinsium had 9 walking appendages and 4 fine anterior appendages. Its body was covered in hair-like Papillae, and its fine anterior appendages were lined with Setae likely used for filter feeding. The generic name honors paleontologist Desmond Collins.
Onychodictyon is a genus of extinct lobopodian known from the Lower Cambrian Chengjiang Maotianshan Shales in the Yunnan Province in China. It was characterized by a stout body covered by fleshy papillae and pairs of sclerotized plates with spines, representing part of the diverse "armoured lobopodians" alongside similar forms such as Microdictyon and Hallucigenia.
Collinsovermis is a genus of extinct panarthropod belonging to the group Lobopodia and known from the middle Cambrian Burgess Shale in British Columbia, Canada. It is monotypic having only one species, Collinsovermis monstruosus. After its initial discovery in 1983, Desmond H. Collins popularised it as a unique animal and was subsequently dubbed "Collins' monster" for its unusual super armoured body. The formal scientific description and name were given in 2020.
Acinocricus is a genus of extinct worm belonging to the group Lobopodia and known from the middle Cambrian Spence Shale of Utah, United States. As a monotypic genus, it has one species Acinocricus stichus. The only lobopodian discovered from the Spence Shale, it was described by Simon Conway Morris and Richard A. Robison in 1988. Owing to the original fragmentary fossils discovered since 1982, it was initially classified as an alga, but later realised to be an animal belonging to Cambrian fauna.
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.
Thanahita is a genus of extinct lobopodian and known from the middle Silurian Herefordshire Lagerstätte at the England–Wales border in UK. It is monotypic and contains one species, Thanahita distos. Discovered in 2018, it is estimated to have lived around 430 million years ago and is the only known extinct lobopodian in Europe, and the first Silurian lobopodian known worldwide.
Hallucigeniidae is a family of extinct worms belonging to the group Lobopodia that originated during the Cambrian explosion. It is based on the species Hallucigenia sparsa, the fossil of which was discovered by Charles Doolittle Walcott in 1911 from the Burgess Shale of British Columbia. The name Hallucigenia was created by Simon Conway Morris in 1977, from which the family was erected after discoveries of other hallucigeniid worms from other parts of the world. Classification of these lobopods and their relatives are still controversial, and the family consists of at least four genera.
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.