Fuxianhuia Temporal range: Lower Cambrian | |
---|---|
Fuxianhuia protensa | |
Fuxianhuia xiaoshibaensis | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Order: | † Fuxianhuiida |
Family: | † Fuxianhuiidae |
Genus: | † Fuxianhuia Hou, 1987 |
Type species | |
Fuxianhuia protensa Hou, 1987 | |
Other species | |
Fuxianhuia xiaoshibaensisYang et al, 2013 |
Fuxianhuia is a genus of Lower Cambrian fossil arthropod known from the Chengjiang fauna in China. Its purportedly primitive features have led to its playing a pivotal role in discussions about the euarthropod stem group. Nevertheless, despite being known from many specimens, disputes about its morphology, in particular its head appendages, have made it one of the most controversial of the Chengjiang taxa, and it has been discussed extensively in the context of the arthropod head problem.
The genus is named after Fuxian Lake (Fuxian Hu), where it was unearthed. Its specific name protensa refers to its extended trunk. [1]
Complete Fuxianhuia specimens are approximately 4 centimetres long. The anterior of Fuxianhuia is encased in an oval sclerite, from which two stalked eyes emerge. Inserting directly behind this sclerite, on the head shield proper, are two stout antennae. When the head of Fuxianhuia was originally described, two additional head appendages, the "sub-chelate" pair were also described. [2] These are geniculate, backward-pointing appendages that lie in a highly stereotypical position (i.e., their position does not vary much from one specimen to another). Partly because of this, and partly because of their rather indistinct morphology, their status as appendages has been questioned. Indeed, on the grounds that these structures seem to lie between two cuticular layers, Waloszek and colleagues have suggested that they are not appendages at all, but rather gut diverticula; [3] a reassignment that has however not been universally accepted. [4] Ventrally, a large plate has been interpreted as a hypostome.
The head shield overlaps a tapering series of 12–17 trunk tergites, which lead into a set of limb-bearing segments comprising the thorax. The limbs are simple in form, consisting of a smooth oval exopod and a stout, annulated endopod. There is no one-to-one correspondence between the thoracic tergites and the limbs, but, rather, there appear to be two or three limbs per tergite.
Behind the thorax is a narrower abdominal region, consisting of 14 tergites, that bears no appendages. The abdomen is terminated by a telson-like spine.
In 2012, a Fuxianhuia fossil was described with exceptional preservation of brain and optic lobes. [5] The shape and complexity corresponds roughly to that of a modern malacostracan brain. [5] In general, the Fuxianhuia brain shows the same tripartite morphology of Malacostraca, Chilopoda and Insecta, indicating that such an organization could be precedent to the divergence between these clades. [5]
In 2014 a fossil was described [6] that preserved in exquisite, unequaled detail the tubular heart and blood vessels, which represent the oldest cardiovascular system yet identified. "The rich vascularization in the head... suggests that the brain of this species required a good supply of oxygen for its performance," said University of Arizona neuroscientist Nicholas Strausfeld, one of the researchers. [7]
Fuxianhuia was first described from incomplete material, [8] and its true nature did not become apparent until the head and limbs were discovered. Its articulated head region, lack of tergite-segment correspondence and undifferentiated limbs have all been taken to indicate a very basal position in the arthropods, [2] [9] [10] even though an early cladistic analysis suggested, rather, that it was a stem-group chelicerate. [11] The presence of a distinct anterior sclerite bearing the eyes has been taken to suggest that a distinct acron once existed in front of the euarthropod head. [2]
Fuxianhuia is not a unique arthropod, it is now recognised to belong to the clade Fuxianhuiida including other similar forms from the Cambrian of China, within Fuxianhuiida it is most closely related to Guangweicaris, together forming the family Fuxianhuiidae. A cladogram is given below.
From. [12]
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The lobopodians, members of the informal group Lobopodia, or the formally erected phylum Lobopoda Cavalier-Smith (1998), 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, worm-like fossil panarthropods such as Aysheaia and Hallucigenia.
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 comes from Greek, "deinos" and "caris", referring 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, Pambdelurion and Kerygmachela. It is most likely paraphyletic, with Kerygmachela and Pambdelurion more basal than the clade compose of Opabiniidae, Radiodonta and other arthropods.
Anomalocaris is an extinct genus of radiodont, an order of early-diverging stem-group arthropods. The first fossils of Anomalocaris were discovered in the Ogygopsis Shale of the Stephen Formation in British Columbia, Canada by Joseph Frederick Whiteaves, with more examples found by Charles Doolittle Walcott in the Burgess Shale unit of the Stephen Formation. Other closely related fossils have been found in the older Emu Bay Shale of Australia, as well as possibly elsewhere. Originally several fossilized parts discovered separately were thought to be three separate creatures, a misapprehension corrected by Harry B. Whittington and Derek Briggs in a 1985 journal article. With a body length close to 40 centimetres, A. canadensis is thought to be one of the earliest examples of an apex predator, though others have been found in older Cambrian lagerstätten deposits.
Cheloniellida is a taxon of extinct Paleozoic arthropods. As of 2018, 7 monotypic genera of cheloniellids had been formally described, whose fossils are found in marine strata ranging from Ordovician to Devonian in age. Cheloniellida has a controversial phylogenetic position, with previous studies associated it as either a member or relative of various fossil and extant arthropod taxa. It was later accepted as a member of Vicissicaudata within Artiopoda.
Anomalocarididae is an extinct family of Cambrian radiodonts, a group of stem-group arthropods.
Parapeytoia is a prehistoric 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.
The (pan)arthropod head problem is a long-standing zoological dispute concerning the segmental composition of the heads of the various arthropod groups, and how they are evolutionarily related to each other. While the dispute has historically centered on the exact make-up of the insect head, it has been widened to include other living arthropods such as chelicerates, myriapods, crustaceans; and fossil forms, such as the many arthropods known from exceptionally preserved Cambrian faunas. While the topic has classically been based on insect embryology, in recent years a great deal of developmental molecular data has become available. Dozens of more or less distinct solutions to the problem, dating back to at least 1897, have been published, including several in the 2000s.
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.
Schinderhannes bartelsi is a species of hurdiid radiodont (anomalocaridid) known from one specimen from the lower Devonian Hunsrück Slates. Its discovery was astonishing because previously, radiodonts were known only from exceptionally well-preserved fossil beds (Lagerstätten) from the Cambrian, 100 million years earlier.
Haikoucaris is a genus of megacheiran arthropod that contains the single species Haikoucaris ercaiensis. It was discovered in the Cambrian Chengjiang biota of China.
Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. They may be referred to as radiodonts, radiodontans, radiodontids, anomalocarids, or anomalocaridids, although the last two originally refer to the family Anomalocarididae, which previously included all species of this order but is now restricted to only a few species. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and used for a variety of functions. Radiodonts included the earliest large predators known, 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 gainessii, Cambroraster falcatus and Amplectobelua symbrachiata, the Ordovician Aegirocassis benmoulai and the Devonian Schinderhannes bartelsi.
Cardiodictyon is a genus of lobopodian known from 518 millions years old Chengjiang Lagerstätte. 525 millions years old partial fossil is also reported. It has ~25 pairs of legs, each associated with a pair of dorsal plates.
Amplectobeluidae is a clade of Cambrian radiodonts.
Lyrarapax is a radiodont genus of the family Amplectobeluidae that lived in the early Cambrian period 520 million years ago. Its neural tissue indicates that the radiodont frontal appendage is protocerebral, resolving parts of the arthropod head problem and showing that the frontal appendage is homologous to the antennae of Onychophorans and labrum of euarthropods. Its fossilized remains were found in Yunnan in southwestern China. A second species was described in 2016, differing principally in the morphology of its frontal appendages.
Chengjiangocaris is an extinct genus of fuxianhuiid arthropod known from the Cambrian of South China. It contains two species, C. longiformis which was described in 1991. C. kunmingensis was described in 2013 by Javier Ortega-Hernández and colleagues. One specimen of C. kunmingensis shows detailed evidence of a nervous system. The nervous system of the chengjiangocaris is and has always been very complex to understand but a recent discovery of the ladder like ventral nerval cords and segmental ganglia of the related chengjiangocaris together with the brain provide the most comprehensive reconstruction of any lower Cambrian arthropod.
Retifacies is an extinct arthropod, that lived in the lower Cambrian. Its fossil remains have been found in the Maotianshan Shales of Yunnan, China. It is a member of the Artiopoda.
Hymenocarina is an order of extinct arthropods known from the Cambrian. They possess bivalved carapaces, typically with exposed posteriors. Members of the group are morphologically diverse and had a variety of ecologies, including as filter feeders and as predators. Recent research has generally considered them to be stem or crown group members of Mandibulata, due the presence of mandibles in at least some species.
Fuxianhuiida is an extinct clade of arthropods from the Cambrian of China. All currently known species are from Cambrian Series 2 aged deposits in Yunnan Province, including the Chengjiang biota. They are generally considered to be close to the base of Euarthropoda, but have also been considered to be early mandibulates. Many specimens are known with exceptional soft tissue preservation, including preserved guts and neural tissue, which given their basal phylogenetic position makes them important in understanding the evolution of arthropoda as a whole. They reach a size of up to 15 cm, and are interpreted as benthic predators and scavengers. The fuxianhuiid exoskeleton is unmineralised, and the number of tergites ranges from 15 to over 40. The cephalon is covered by a head shield and contains stalked eyes connected by the anterior sclerite, antennae, a butterfly shaped hyposome and a posterior facing mouth. Fuxianhuiids possess specialized post-antennal appendages with serrated edges used for food processing. The presence of gnathobases in members of Chengjiangocardidae suggests that they were capable of durophagy. In most fuxianhuiids, the thorax tergites narrow posteriorly, terminating in either a swimming paddle or paired flukes with a tailspine. In members of Fuxianhuiidae the thorax is divided into two sections, the anterior wide opisthothorax and the posterior narrow limbless tail-like abdomen.
Kylinxia is a genus of extinct arthropod described in 2020. It was described from six specimens discovered in Yu'anshan Formation in southern China. The specimens are assigned to one species Kylinxia zhangi. Dated to 518 million years, the fossils falls under the Cambrian period. Announcing the discovery on 4 November 2020 at a press conference, Zeng Han of the Nanjing Institute of Geology and Paleontology, said that the animal "bridges the evolutionary gap from Anomalocaris to true arthropods and forms a key ‘missing link’ in the origin of arthropods," which was "predicted by Darwin’s evolutionary theory." The same day the formal description was published in Nature.
Luolishania is an extinct genus of lobopodian worm 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.
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