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| The Burgess Shale |
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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. [1]
The fauna of the middle Cambrian has a cosmopolitan range. All assemblages preserving soft-part anatomy have a very similar fauna, even though they span almost every continent. [2] The wide distribution has been attributed to the advent of pelagic larvae. [2]
The fauna is composed of a range of soft-bodied organisms; creatures with hard, mineralised skeletons are rare, although trilobites are quite commonly found. The major soft-bodied groups are sponges, palaeoscolecid worms, lobopods, arthropods and anomalocaridids. [2] Assemblages are typically diverse, with the most famous localities each containing in the region of 150 described species. [2] The fauna of the Burgess Shale lived in the photic zone, as bottom-dwelling photosynthesisers are present in the assemblage. [3]
Sirius Passet is a lagerstätte in Greenland which was formed about 527 million years ago. Its most common fossils are arthropods, but there is only a handful of trilobite species. There are also very few species with hard parts: trilobites, hyoliths, sponges, brachiopods, and no echinoderms or molluscs. [4]
Halkieria has features associated with more than one living phylum, and is discussed below.
The strangest-looking animals from Sirius Passet are Pambdelurion and Kerygmachela . They are generally regarded as anomalocarids because they have long, soft, segmented bodies with a pair of broad fin-like flaps on most segments and a pair of segmented appendages at the rear. The outer parts of the top surfaces of the flaps have grooved areas which are thought to have acted as gills. Under each flap there is a short, fleshy leg. This arrangement suggests the animals are related to biramous arthropods. [5]
There are several Cambrian fossil sites in the Chengjiang county of China's Yunnan province. The most significant is the Maotianshan shale, a lagerstätte which preserves soft tissues very well. The Chengjiang fauna date to between 525 million and 520 million years ago, about the middle of the early Cambrian epoch, a few million years after Sirius Passet and at least 10 million years earlier than the Burgess Shale.
The Chengjiang sediments provide what are currently the oldest-known chordates, the phylum to which all vertebrates belong. The 8 chordate species include Myllokunmingia , possibly a very primitive agnathid and Haikouichthys , which may be related to lampreys. [6] Yunnanozoon may be the oldest-known hemichordate. [7]
Anomalocaris was a mainly soft-bodied swimming predator which was gigantic for its time (up to 70 cm = 2¼ feet long; some later species were 3 times as long); the soft, segmented body had a pair of broad fin-like flaps along each side, except that the last 3 segments had a pair of fans arranged in a V shape. Unlike Kerygmachela and Pambdelurion (see above), Anomalocaris apparently had no legs, and the grooved patches which are thought to have acted as gills were at the bases of the flaps, or even overlapping on to its back. The two eyes were on relatively long horizontal stalks; the mouth lay under the head and was a round-cornered square of plates which could not close completely; and in front of the mouth were two jointed appendages which were shaped like a shrimp's body, curved backwards and with short spines on the inside of the curve. Amplectobelua , also found at Chengjiang, was similar, smaller than Anomalocaris but considerably larger than most other Chengjiang animals. Both are thought to have been powerful predators.
Hallucigenia looks like a long-legged caterpillar with spines on its back, and almost certainly crawled on the seabed. [4]
Nearly half of the Chengjiang fossil species are arthropods, few of which had the hard, mineral-reinforced exoskeletons found in most later marine arthropods; only about 3% of the organisms known from Chengjiang have hard shells, and most of those are trilobites (although Misszhouia is a soft-bodied trilobite). Many other phyla are found there: Porifera (sponges) and Priapulida (burrowing "worms" which were ambush predators), Brachiopoda (these had bivalve-like shells, but fed by means of a lophophore, a fan-like filter which occupied about of half of the internal space), Chaetognatha (arrow worms), Cnidaria, Ctenophora (comb jellies), Echinodermata, Hyolitha (Lophophorata with small conical shells), [8] Nematomorpha, Phoronida (horseshoe worms), and Protista. [9]
The Burgess Shale was the first of the Cambrian lagerstätten to be discovered (by Walcott in 1909), and the re-analysis of the Burgess Shale by Whittington and others in the 1970s was the basis of Gould's book Wonderful Life , which was largely responsible for non-scientists' awareness of the Cambrian explosion. The fossils date from the mid Cambrian, about 515 million years ago and 10 million years later than the Chengjiang fauna.
The shelled fossils in the Burgess Shale are similar in proportions to other shelly fossil deposits; however, they are a minor component of the biota, accounting for only 14% of the Burgess Shale fossils. When organisms that were not preserved are entered into the equation, the shelly fossils probably represent about 2% of the animals that were alive at the time. [10]
Arthropods are the most abundant and diverse group of organisms in the Burgess Shale, followed closely by sponges. [11] Many Burgess Shale fossils are unusual and difficult to classify, for example:
But the "weird wonders", creatures that resembled nothing known in the 1970s, attracted the most publicity, for example:
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Other fauna include the Middle Cambrian Wheeler Shale Formation of Utah. [17]
Trace fossils are associated with many Burgess Shale-type deposits. [18] They are often associated with the innards of soft-bodied organisms, [19] and are particularly prevalent under the carapaces of bivalved arthropods. [20] Burrowing organisms seem to have used the high-sulfur decay fluids as a nutrient source when farming bacteria in the microenvironment under the carapaces, indicated by their repeated uses of individual burrows. [20]
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.
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, 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.
The Emu Bay Shale is a geological formation in Emu Bay, South Australia, containing a major Konservat-Lagerstätte. It is one of two in the world containing Redlichiidan trilobites. The Emu Bay Shale is dated as Cambrian Series 2, Stage 4, correlated with the upper Botomian Stage of the Lower Cambrian.
Anomalocaris is an extinct genus of radiodont, an order of early-diverging stem-group 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.
Waptia is an extinct genus of arthropod from the Middle Cambrian of North America. It grew to a length of 6.65 cm (3 in), and had a large bivalved carapace and a segmented body terminating into a pair of tail flaps. It was an active swimmer and likely a predator of soft-bodied prey. It is also one of the oldest animals with direct evidence of brood care. Waptia fieldensis is the only species classified under the genus Waptia, and is known from the Burgess Shale Lagerstätte of British Columbia, Canada. Specimens of Waptia are also known from the Spence Shale of Utah, United States.
Peytoia infercambriensis is a species of hurdiid radiodont in the genus Peytoia.
The Burgess Shale of British Columbia is famous for its exceptional preservation of mid-Cambrian organisms. Around 69 other sites have been discovered of a similar age, with soft tissues preserved in a similar, though not identical, fashion. Additional sites with a similar form of preservation are known from the Ediacaran and Ordovician periods.
Alalcomenaeus is one of the most widespread and longest-surviving arthropod genera of the Early and Middle Cambrian. Known from over 300 specimens in the Burgess Shale and the Chengjiang biota. It is a member of the family Leanchoiliidae in the group Megacheira.
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.
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 later surviving members include the subfamily Aegirocassisinae from the Early Ordovician of Morocco and the Early Devonian member Schinderhannes bartelsi from Germany.
The fossils of the Burgess Shale, like the Burgess Shale itself, are fossils that formed around 505 million years ago in the mid-Cambrian period. They were discovered in Canada in 1886, and Charles Doolittle Walcott collected over 65,000 specimens in a series of field trips up to the alpine site from 1909 to 1924. After a period of neglect from the 1930s to the early 1960s, new excavations and re-examinations of Walcott's collection continue to reveal new species, and statistical analysis suggests that additional discoveries will continue for the foreseeable future. Stephen Jay Gould's 1989 book Wonderful Life describes the history of discovery up to the early 1980s, although his analysis of the implications for evolution has been contested.
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.
Kiisortoqia soperi is an extinct species of arthropod from the Early Cambrian Sirius Passet Lagerstätte in Greenland. While it had a superficially trilobite-like bodyform, it also possessed large frontal appendages similar to those of radiodonts.
Squamacula is an extinct artiopodan arthropod from the Cambrian Series 2. The type species S. clypeata was described in 1997 from the Chengjiang biota of Yunnan, China. At the time of description there were only two known specimens of S. clypeata, but now there are at least six known specimens. In 2012 a second species S. buckorum was described from the Emu Bay Shale of Australia.
Retifacies abnormalis 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, and closely related to Pygmaclypeatus.
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.
Erratus is an extinct genus of marine arthropod from the Cambrian of China. Its type and only species is Erratus sperare. 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. Erratus on the other hand had not only flaps but also a set of primitive legs. 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.
Balhuticaris is a genus of extinct bivalved hymenocarine arthropod that lived in the Cambrian aged Burgess Shale in what is now British Columbia around 506 million years ago. This extremely multisegmented arthropod is the largest member of the group, and it was even one of the largest animals of the Cambrian, with individuals reaching lengths of 245 mm (9 in). Fossils of this animal suggests that gigantism occurred in more groups of Arthropoda than had been previously thought. It also presents the possibility that bivalved arthropods were very diverse, and filled in a lot of ecological niches.
