Erratus Temporal range: | |
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Schematic reconstruction in dorsal and ventral views | |
Life restoration | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Genus: | † Erratus Fu et al. 2022 |
Type species | |
Erratus sperare Fu et al. 2022 |
Erratus [lower-alpha 1] is an extinct genus of marine arthropod from the Cambrian of China. Its type and only species is Erratus sperare. [lower-alpha 2] Erratus is likely one of the most basal known arthropods, and its discovery has helped scientists understand the early evolution of arthropod trunk appendages. Some of the stem-arthropods like radiodonts did not have legs, instead they had flap like appendages that helped them swim. [1] Erratus on the other hand had not only flaps but also a set of primitive legs. [2] It also supported the theory that the gills of aquatic arthropods probably evolved into the wings and lungs of terrestrial arthropods later in the Paleozoic. [3]
Fossils of Erratus have been found in the Chengjiang Lagerstätte of China, [4] dating to around 520 million years ago. [5] Erratus was a small, nektonic organism with a bivalved carapace that probably ate by deposit feeding or filter feeding.
The holotype consists of specimens XDBZ101 and XDBZ102. [2] The paratype specimen, NWUS92-310, had previously been referred to Isoxys auritus . [6] [2] Both specimens were found in the Helinpu Formation in Yunnan, China. Erratus sperare was named by Fu and colleagues in 2022. The genus name is Latin for "roaming", in reference to the inferred nektonic habit of the organism, and the specific name is Latin for "to hope". The holotype specimens consist of a part and counterpart of an individual, and an isolated carapace. The paratype consists of a carapace with an anterior spine, as well as seven anterior body fragments. [2]
The relationship between the components of the biramous limb of many euarthropod groups, which consists of an exopod and an endopod (appendages), and the lopopods (stubby legs) and flaps of lobopodians and radiodonts has long been controversial. Erratus appears to show one of the earliest steps in the evolution of the endopod, with an unsclerotized endopod fused to a flap that is attached to the body wall, rather than being a branch of the appendage like an exopod. Fu et al. 2022 describes the species as having "unique trunk appendages formed of lateral anomalocaridid-type flaps and ventral subconical endopods". The fossils indicate that the species represent "an intermediate stage of biramous limb evolution". [2]
Known specimens of Erratus measure 37–72 millimetres (1.5–2.8 in) in length. The arthropod had a bivalved carapace that covered the top, which possesses an anterior spine but rounded posteriorly. A pair of lateral eyes located near the anterior spine of the carapace. The eyes appear to have no lenses, and the stalks appear to be as long as the eyes. On the bottom of the arthropod were 11 pairs of wide body flaps with gill-like wrinklings. The flaps decreased in size towards the bottom of the arthropod. As in gilled lobopodians and radiodonts, the flaps were connected to the body wall. The anterior 7 pairs of body flaps each possesses a medially-arranged endopod (inner branch). Each endopod was a simple, 7-segmented leg, lacking any evidences of endites (internal projections) and claws. The posterior 4 pairs of body flaps lacking endopods, but the wrinklings are more prominent. Ventral head structures between the eyes and body flaps are yet to be discovered, it is also uncertain if the trunk-end possesses a telson. [2]
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Phylogenetic position of Erratus within the arthropod stem group, after Fu et al. 2022 [2] |
A phylogenetic analysis conducted by Fu et al. 2022 found that Erratus was a basal arthropod, branched before isoxyids ( Isoxys and Surusicaris ) and other euarthropods (hymenocarines, fuxianhuiids, megacheirans, trilobites and so on), but more derived than other stem-arthropods like gilled lobopodians ( Pambdelurion and Kerygmachela ), Opabinia and radiodonts ( Anomalocaris , Hurdia and so on). [2]
Erratus was a member of the Chengjiang Biota, which dates to the Cambrian period, 520 million years ago. [2] During this time the area was a tropical region with sea level changes and tectonic activity. Most of the fauna were primarily benthic, and were probably buried via turbidity currents. Brachiopods, ctenophores, phoronids, lobopodians, worms, primitive chordates, and other arthropods are represented in the biota. [7] [8] The site is known for its incredible preservation of fossils similar to that of the younger Burgess Shale. [9]
Sidneyia is an extinct arthropod known from fossils found from the Early to the Mid Cambrian of China and the Mid Cambrian Burgess Shale of British Columbia, Canada.
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.
Canadaspis is an extinct genus of bivalved Cambrian arthropod, known from North America and China. They are thought to have been benthic feeders that moved mainly by walking and possibly used its biramous appendages to stir mud in search of food. They have been placed within the Hymenocarina, which includes other bivalved Cambrian arthropods.
Amplectobelua is an extinct genus of late Early Cambrian amplectobeluid radiodont, a group of stem arthropods that mostly lived as free-swimming predators during the first half of the Paleozoic Era.
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.
Chuandianella ovata is an extinct bivalved arthropod that lived during Cambrian Stage 3 of the Early Cambrian. It is the only species classified under the genus Chuandianella. Its fossils were recovered from the Chengjiang Biota in Yunnan, China.
Haikoucaris is a genus of megacheiran arthropod that contains the single species Haikoucaris ercaiensis. It was discovered in the Cambrian Chengjiang biota of China.
Isoxys is a genus of extinct bivalved Cambrian arthropod; the various species of which are thought to have been freely swimming predators. It had a pair of large spherical eyes, and two large frontal appendages used to grasp prey.
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.
Occacaris oviformis is an extinct nektonic predatory arthropod from the Lower Cambrian Maotianshan shale Lagerstätte. It bears a superficial resemblance to the Cambrian arthropod, Canadaspis, though, was much smaller, and had a pair of "great appendages", with which it may have grasped prey. It was originally considered to belong to Megacheira, however it is questioned in later study.
Pectocaris is an extinct genus of bivalved arthropods from the Cambrian Maotianshan Shales, Yunnan Province of China. There are currently three known species within the genus.
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.
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.
Aegirocassis is an extinct genus of giant radiodont arthropod belonging to the family Hurdiidae that lived 480 million years ago during the early Ordovician in the Fezouata Formation of Morocco. It is known by a single species, Aegirocassis benmoulai. Van Roy initiated scientific study of the fossil, the earliest known of a "giant" filter-feeder discovered to date. Aegirocassis is considered to have evolved from early predatory radiodonts. This animal is characterized by its long, forward facing head sclerite, and the endites on its frontal appendages that bore copious amounts of baleen-like auxiliary spines. This animal evolving filter-feeding traits was most likely a result of the Great Ordovician Biodiversification Event, when environmental changes caused a diversification of plankton, which in turn allowed for the evolution of new suspension feeding lifeforms. Alongside the closely related Pseudoangustidontus, an unnamed hurdiid from Wales, the middle Ordovician dinocaridid Mieridduryn, and the Devonian hurdiid Schinderhannes this radiodont is one of the few known dinocaridids known from post-Cambrian rocks.
Tokummia is a genus of fossil hymenocarine arthropod, known only by one species, Tokummiakatalepsis, from the middle Cambrian Burgess Shale as found in a quarry in Marble Canyon in Canada.
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.
Deuteropoda is a proposed clade of arthropods whose members are distinguished from more basal stem-group arthropods like radiodonts by an anatomical reorganization of the head region, namely the appearance of a differentiated first appendage pair, a multisegmented head, a hypostome/labrum complex, and by bearing pairs of segmented biramous limbs.
Fengzhengia is an extinct genus of arthropod known from a single species, Fengzhengia mamingae from the Cambrian aged Chengjiang Biota of Yunnan, China. It is thought to be a basal arthropod, as one of the most basal members of Deuteropoda. Like other basal deuteropods, Fengzhengia has an upward curling pair of "frontal appendages" with at least 6 segments which bear endites, contrasting with the downward curving pair of frontal appendages possessed by radiodonts. The head has a pair of stalked eyes. The trunk had 15 tergites the first nine of which had upward facing spines, with the trunk terminating with a tail fan. The trunk seemingly had pairs of biramous limbs, with paddle-shaped exopods. It is thought to have been nektobenthic, and either a scavenger or a predator. A close relationship with the genus Kylinxia has been suggested.
Jugatacaris is an extinct genus of bivalved arthropod known from a single species, Jugatacaris agilis found in the Cambrian Stage 3 aged Chengjiang biota of Yunnan, China. The carapace is around 28 to 37 millimetres in length, with a pronounced ridge at the top of the carapace separating the two valves, which formed a fin-like structure raised above the carapace. The head has a pair of stalked eyes, as well as a dumbbell shaped medial eye between them. The head also bore a pair of mandibles as well as at least one and possibly two pairs of antennules. The trunk had up to 65 segments, each with biramous appendages. The appendages had thin endopods with 30 podomeres, each bearing a spiny endite, with the endopods ending with a terminal claw. The appendages also had overlapping flap-like exopods, which are elongated, being at maximum eight times as long as they are wide, which on their posterior edge are covered with setae. The trunk ended with a forked tail. It was likely an actively swimming filter feeder, using its constantly beating appendages to sift food from the water column, which was then passed forward along the U-shaped food groove between the appendage pairs towards the mouth. While initially placed as a crustaceanomorph, later studies considered to be a member of Hymenocarina, which contains numerous other similar bivalved Cambrian arthropods.
Isoxyids are members of the order Isoxyida and the family Isoxyidae, a group of basal arthropods that existed during the Cambrian period. It contains two genera, Isoxys, with 20 species found worldwide, and Surusicaris known from a single species found in the Burgess Shale of Canada. They are distinguished by their bivalved carapaces and pair of upward curving grasping frontal appendages.