Luolishaniidae Temporal range: | |
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Collinsovermis (1), Luolishania (= Miraluolishania , 2), Ovatiovermis (3) and Facivermis (4) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
(unranked): | Panarthropoda |
Phylum: | † "Lobopodia" |
Clade: | † Hallucishaniids |
Family: | † Luolishaniidae |
Genera | |
The Luolishaniidae [5] or Luolishaniida [Note 1] are a group of Cambrian and Ordovician [6] lobopodians with anterior 5 or 6 pairs of setiferous lobopods. [1] [3] [2] Most luolishaniids also have posterior lobopods each with a hooked claws, and thorn-shaped sclerites arranged as three or more per trunk segment. [4] [1] [2] The type genus is based on Luolishania longicruris Hou and Chen, 1989, from the Chengjiang Lagerstatte, South China. [5] They are presumed to have been benthic suspension or filter feeders. [4] [1] [3] [2]
New specimens of the previously enigmatic Facivermis show that it was a sessile tube-dweller, and part of this group. [3]
A 2023 analysis concluded, on the basis of numerous morphological similarities, that luolishaniids might be the closest known relatives of tardigrades. [7]
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. However, other genera like Kerygmachela and Pambdelurion are often referred to as “gilled lobopodians”.
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.
Dinocaridida is a proposed fossil taxon of basal arthropods, which flourished during the Cambrian period and survived up to Early Devonian. Characterized by a pair of frontal appendages and series of body flaps, the name of Dinocaridids refers to the suggested role of some of these members as the largest marine predators of their time. Dinocaridids are occasionally referred to as the 'AOPK group' by some literatures, as the group compose of Radiodonta, Opabiniidae, and the "gilled lobopodians" Pambdelurion and Kerygmachelidae. It is most likely paraphyletic, with Kerygmachelidae and Pambdelurion more basal than the clade compose of Opabiniidae, Radiodonta and other arthropods.
Pambdelurion is an extinct genus of panarthropod from the Cambrian aged Sirius Passet site in northern Greenland. Like the morphologically similar Kerygmachela from the same locality, Pambdelurion is thought to be closely related to arthropods, combining characteristics of "lobopodians" with those of primitive arthropods.
Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and were used for a variety of functions. Radiodonts were among the earliest large predators, but they also included sediment sifters and filter feeders. Some of the most famous species of radiodonts are the Cambrian taxa Anomalocaris canadensis, Hurdia victoria, Peytoia nathorsti, Titanokorys gainesi, Cambroraster falcatus and Amplectobelua symbrachiata. The later surviving members include the subfamily Aegirocassisinae from the Early Ordovician of Morocco and the Early Devonian member Schinderhannes bartelsi from Germany.
Miraluolishania is an extinct lobopodian known from Chengjiang County in China. It is remarkable for the possession of lensed pit-eyes. The only species, Miraluolishania haikouensis, was described from the Maotianshan Shales at Haikou by Jianni Liu and Degan Shu in 2004. In 2009, a team of palaeontologists at the Yunnan University, led by Xiaoya Ma reported the discovery of 42 other specimens from Haikou. With the help of Swiss palaeontologist Jan Bergström, Ma and Hou came to the conclusion that all the specimens were the same species as Luolishania; another lobopod discovered from the Chengjian in 1989. Chengjian is 40 kms from Haikou and the fossil fauna are different. A reassessment by Liu and Shu's team at the Northwest University in 2008 established that Luolishania and Miraluolishania are distinct animals.
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.
Omnidens, meaning "all-tooth", is an extinct genus of large Cambrian animal known only from a series of large mouth apparatus and sclerotized talon-like structures, originally mistaken as the mouthparts of anomalocaridids. When first named, it was interpreted as a giant priapulid, but is now considered a panarthropod. Its mouth apparatus closely resembles that of the smaller gilled lobopodian Pambdelurion, indicating it is likely to have been a close relative of that species, potentially even synonymous. With a maximum estimated body length of 1.5 metres (4.9 ft), Omnidens is suggested to have been the largest known free-living Cambrian organism. Omnidens fossils are found in the Maotianshan Shales.
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.
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 6 fine anterior appendages. Its body was covered in hair-like papillae and its fine anterior appendages were lined with setae. 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.
Houcaris is a possibly paraphyletic radiodont genus, tentatively assigned to either Amplectobeluidae, Anomalocarididae or Tamisiocarididae, known from Cambrian Series 2 of China and the United States. The type species is Houcaris saron which was originally described as a species of the related genus Anomalocaris. Other possible species include H. magnabasis and H. consimilis. The genus Houcaris was established for the two species in 2021 and honors Hou Xianguang, who had discovered and named the type species Anomalocaris saron in 1995 along with his colleagues Jan Bergström and Per E. Ahlberg.
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. A similar lobopodian is known from the Emu Bay Shale, however it remains unnamed.
Acinocricus is a genus of extinct panarthropod 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.
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. It was discovered and described by Joachim T. Haug, Georg Mayer, Carolin Haug, and Derek E.G. Briggs in 2012. With an age of about 300 million years, it is the first long-legged lobopodian discovered after the period of Cambrian explosion.
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.