Hallucigeniidae | |
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Fossil of Hallucigenia from the Burgess Shale | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
(unranked): | Panarthropoda |
Phylum: | † "Lobopodia" |
Clade: | † Hallucishaniids |
Family: | † Hallucigeniidae Conway Morris, 1977 |
Genera | |
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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. [1] Classification of these lobopods and their relatives are still controversial, and the family consists of at least four genera.
The first fossil of hallucigeniid worm was discovered by an American palaeontologist Charles Doolittle Walcott from the Walcott Quarry that contains the Cambrian Burgess Shale. In 1911, Walcott gave the name Canadia sparsa as he believed that it was related to the polychaete worm (Annelida) Canadia spinosa, which he described simultaneously. [2] British palaeontologist Simon Conway Morris re-examined the specimen and concluded that it was not a Canadia species. He created a new genus Hallucigenia in 1977. [3] [4] With only a single species and fragmentary fossils available, the relationship of the worm with other animals was not obvious. The most prominent feature of the worm, its body projections were particularly difficult to understand as there were two distinct groups, the tube-like tentacles and thorn-like spines. Morris described the spines as the legs and the tentacles as feeding apparatus. [5] Two other species were later discovered from the Cambrian Maotianshan shales of China, H. fortis in 1995, [6] and H. hongmeia in 2012. [7]
In 1991, Lars Ramsköld (Uppsala University, Sweden) and Hou Xian-guang (Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences) described a new specimen, Microdictyon , from the lower Cambrian Maotianshan shales. With this relatively complete fossil, they assigned the animal to and reinterpreted Hallucigenia as a lobopodian, a legged worm-like taxon which were still thought to be exclusively related to onychophoran (velvet worm) at that time. [8] [9] They were also able to work out that by inverting the specimen upside down, the so-called tentacles were actually walking legs (called lobopods) and the spines were protective armours on the back. [5] [10] The reinterpretation was strengthened by the discovery of new species Cardiodictyon catenulum from the same Maotianshan shales, reported by Hou, Ramsköld and Jan Bergström in the same year. [11]
In 2012, a different lobopodian fossil was discovered from the Permian (about 296 million years old) sediments of the Mazon Creek in Illinois, US. Joachim T. Haug, Georg Mayer, Carolin Haug, Derek E.G. Briggs gave the name Carbotubulus waloszeki . [12] In 2018, Thanahita distos was described by Derek J. Siveter, Briggs, David J. Siveter, Mark D. Sutton and David Legg from the Coalbrookdale Formation (Herefordshire Lagerstätte) at the England–Wales border in UK. [13] Dated to about 430 million years old, it is the only known extinct lobopodian in Europe and the first Silurian lobopod known worldwide. [14] [15]
Hallucigeniid worms are elongated, soft-bodied animals characterised by several pairs of stumpy legs known as lobopods, for which they are included in the larger but informal group of animals, Lobopodia. Their body can be described in three parts: the head, neck, and the trunk. The head bears a pair of eyes. [1] [16] The neck can be prominent in some species such as T. distos in which it bears two pairs of small legs. [13] The trunk is the longest part of the body and contains several pairs of legs on the frontal (ventral) side and several pairs of spines on the back (dorsal) side. Each legs has terminal claws. C. catenulum, measuring 2.5 cm long, bears about 25 pairs of legs. [17]
T. distos, though incomplete, is the longest with 3 cm body length and bears at least nine pairs of legs. [18] Hallucigenia species are highly diverse in body sizes, H. fortis is only about 1 cm long, H. hongmeia is intermediate with about 3 cm in length, and H. sparsa being the longest measuring 5.5 cm. [7] [19]
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.
Aysheaia is an extinct genus of soft-bodied lobopodian, known from the Middle Cambrian Burgess Shale of British Columbia, Canada
Microdictyon is an extinct armoured worm-like panarthropod coated with net-like scleritic plates, known from the Early Cambrian Maotianshan shale of Yunnan China and other parts of the world. Microdictyon is part of the ill-defined taxon – Lobopodia – that includes several other odd worm-like animals that resembling worm with legs, such as Hallucigenia, Onychodictyon, Cardiodictyon, Luolishania, and Paucipodia. The isolated sclerites of Microdictyon are known from other Lower Cambrian deposits. Microdictyon sclerites appear to have moulted; one sclerite seems to have been preserved during ecdysis.
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.
Dinocaridida is a proposed fossil taxon of basal arthropods that flourished in the Cambrian period with occasional Ordovician and Devonian records. 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.
Parapeytoia is a genus of extinct arthropod that lived over 530 million years ago in the Maotianshan shales of prehistoric China. It was interpreted as an anomalocaridid (radiodont) with legs, but later studies reveal it was a megacheiran, a group of arthropods which are no longer thought to be closely related to the radiodonts.
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.
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.
Archaeopriapulida is a group of priapulid worms known from Cambrian lagerstätte. The group is closely related to, and very similar to, the modern Priapulids. It is unclear whether it is mono- or polyphyletic. Despite a remarkable morphological similarity to their modern cousins, they fall outside of the priapulid crown group, which is not unambiguously represented in the fossil record until the Carboniferous. In addition to well-preserved body fossils, remains of several archaeopriapulid taxa are known to have been preserved primarily as organic microfossils, such as isolated scalids and pharyngeal teeth. They are probably closely related or paraphyletic to the palaeoscolecids; the relationship between these basal worms is somewhat unresolved.
Cucumericrus decoratus is a species of putative radiodont known from a few poorly preserved specimens. Only fragments of trunk cuticle and corresponded appendages had been revealed, while important radiodont features such as frontal appendages are unknown in this species. The trunk cuticle possess irregular wrinkles and may had been soft in life. Each of the trunk appendage compose of a dorsal flap-like element and a ventral stubby leg with unknown distal region, structurally comparable to the trunk appendages of gill lobopodians and euarthropod biramous appendages. The legs have been interpreted as somewhere between annulated lobopod legs and segmented arthropod legs.
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.
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.
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.
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.
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.