Nektaspida

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Nektaspida
Temporal range: Atdabanian–Silurian
Naraoia compacta fossil.jpg
Naraoia compacta fossils with preserved appendages
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Trilobitomorpha
Order: Nektaspida
Raymond, 1920
Families

Nektaspida (also called Naraoiida, Nektaspia and Nectaspida) is an extinct order of non-mineralised artiopodan arthropods. They are known from the mid-Cambrian to the upper Silurian. Originally classified as trilobites, which they superficially resemble, they are now placed as close relatives as members of the Trilobitomorpha within Artiopoda. The order is divided into three major families; Emucarididae, Liwiidae, and Naraoiidae.

Contents

Naming history and taxonomic placement

The order was originally proposed by Raymond in 1920 as Nektaspia. [1] Størmer corrected it to Nectaspida for the 1959 Treatise on Invertebrate Paleontology to conform with the names of the other trilobite orders. Whittington described it in 1985 with the spelling Nektaspida; the revised 1997 Treatise by Raymond and Fortey uses this spelling, as do other modern works. [2] Whittington (1985) placed the order in the Trilobita. Cotton & Braddy (2000) place it in a new "Trilobite clade" containing the Trilobita, recognizing the close affinities of the nektaspids to trilobites. However this necessitates the inclusion of genera that look very little like trilobites., [2] it was formerly placed in the stem-group to the chelicerata subdivision of the Arthropoda phylum. [3] However, it currently considered part of Artiopoda, the clade that contains trilobites and their close relatives. [4]

Anatomy

Digestive systems of various naraoiid species Nektaspida guts.jpg
Digestive systems of various naraoiid species

The group is united by several morphological characters, including reduced or absent lateral eyes, a hypostome with a natant attachment, extensive articulation overlap between unfused trunk tergites, and fused pleurae that do not form lateral spines. [5] Preserved soft tissue of the group indicates that the cephalic (head) shield of nektaspids and liwiids contained branched digestive glands. [6] The Naraoiidae have their exoskeletons mostly composed of only two major shields, which have a single articulation point. [7]

Ecology

The naraoiids have been interpreted as benthic organisms that were opportunistic scavengers and predators of soft-bodied prey, with the spine-like endites of the limbs possibly allowing soft-bodied prey to be shredded before ingestion. The differences in gut morphology between some species of naraoiids suggests that some species only intermittently fed, while others regularly fed. [8]

Evolutionary history

The group first appeared and was most diverse in the Cambrian, especially during the Cambrian Series 2. Emucarididae are only known from the Cambrian Series 2 of East Gondwana (now Australia) and the then nearby South China. Naraoiidae were diverse in low latitudes during Cambrian Series 2 and the following Miaolingian in Laurentia and in South China. Liwiidae first appeared in Baltica during the Cambrian, and are absent from major Cambrian deposits elsewhere during this period, but are widespread in the southern high-latitude margin of Gondwana during the Ordovician. [6] The youngest known member of the group is the Laurentian naraoiid Naraoia bertensis from the Bertie Formation in Ontario, Canada, dating to the Upper Silurian. [9]

Related Research Articles

<span class="mw-page-title-main">Agnostida</span> Extinct order of arthropods

Agnostida are an order of extinct arthropods which have classically been seen as a group of highly modified trilobites, though some recent research has doubted this placement. Regardless, they appear to be close relatives as part of the Artiopoda. They are present in the Lower Cambrian fossil record along with trilobites from the Redlichiida, Corynexochida, and Ptychopariida orders, and were highly diverse throughout the Cambrian. Agnostidan diversity severely declined during the Cambrian-Ordovician transition, and the last agnostidans went extinct in the Late Ordovician.

<span class="mw-page-title-main">Naraoiidae</span> Family of arthropods (fossil)

Naraoiidae is a family, of extinct, soft-shelled trilobite-like arthropods, that belongs to the order Nektaspida. Species included in the Naraoiidae are known from the second half of the Lower Cambrian to the end of the Upper Silurian. The total number of collection sites is limited and distributed over a vast period of time: Maotianshan Shale and Balang Formation (China), Burgess Shale and Bertie Formation (Canada), the Šárka Formation, Emu Bay Shale (Australia), Idaho and Utah (USA). This is probably due to the rare occurrence of the right circumstances for soft tissue preservation, needed for these non-calcified exoskeletons.

<span class="mw-page-title-main">Trilobite</span> Class of extinct, Paleozoic arthropods

Trilobites are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period and they flourished throughout the lower Paleozoic before slipping into a long decline, when, during the Devonian, all trilobite orders except the Proetida died out. The last trilobites disappeared in the mass extinction at the end of the Permian about 251.9 million years ago. Trilobites were among the most successful of all early animals, existing in oceans for almost 270 million years, with over 22,000 species having been described.

<span class="mw-page-title-main">Emu Bay Shale</span> Geological formation in South Australia

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.

<span class="mw-page-title-main">Cheloniellida</span> Order of arthropods (fossil)

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.

<i>Naraoia</i> Extinct genus of arthropods

Naraoia is a genus of small to average size marine arthropods within the family Naraoiidae, that lived from the early Cambrian to the late Silurian period. The species are characterized by a large alimentary system and sideways oriented antennas.

<i>Misszhouia</i>

Misszhouia is a genus of small to average sized marine trilobite-like arthropods within the Naraoiidae family, that lived during the early Cambrian period. The species are M. longicaudata, from the Maotianshan Shales, described in 1985, and M. canadensis, from the Burgess Shale and described in 2018, although later species may belongs to genus Naraoia instead.

<span class="mw-page-title-main">Radiodonta</span> Extinct order of Cambrian arthropods

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.

<span class="mw-page-title-main">Marrellomorpha</span> Extinct class of arthropods

Marrellomorpha are an extinct group of arthropods known from the Cambrian to the Early Devonian. They lacked mineralised hard parts, so are only known from areas of exceptional preservation, limiting their fossil distribution. The best known member is Marrella, with thousands of specimens found in the Cambrian aged Burgess Shale of Canada. The group is divided up into two major orders, Marrellida and Acercostraca. Marrellida is recognised by the possession of head shields with two or three pairs of elongate spine-like projections, and three pairs of uniramous appendages on the cephalon, while Acercostraca generally have large ovoid carapaces that cover the entire upper half of the body, and five pairs of uniramous cephalic appendages. Both groups have unbranched antennules and a segmented trunk with biramous appendages. Recent research has suggested the previously enigmatic Cambrian trilobite-like arthropods Skania and Primicaris belong to this group. Their phylogenetic position is uncertain, various studies have alternatively placed them in the Arachnomorpha as relatives of Artiopoda, as related to Mandibulata, or as stem group euarthropods.

<i>Buenaspis</i> Small Cambrian arthropod

Buenaspis is a genus of small nektaspid arthropod, that lived during the early Cambrian period. Fossil remains of Buenaspis were collected from the Lower Cambrian Sirius Passet Lagerstätte of North Greenland. Buenaspis looks like a soft eyeless trilobite. It has a headshield slightly larger than the tailshield (pygidium), and in between them six thoracic body segments (somites). The genus is monotypic, its sole species being Buenaspis forteyi.

<span class="mw-page-title-main">Emucarididae</span> Extinct family of arthropods

Emucarididae is an extinct family of soft-shelled trilobite-like arthropods (nektaspids) from the Lower Cambrian of South Australia and South China. It contains only two genera – Emucaris and Kangacaris. Two species were described in 2010 from specimens recovered from Emu Bay Shale Lagerstätte, one species in 2012 from the Maotianshan Shales. It is classified under the order Nektaspida, and is a sister-group to the families Liwiidae and Naraoiidae.

<i>Tariccoia</i>

Tariccoia is a genus of nektaspid arthropods belonging to the family Liwiidae, known from fossils found in Ordovician strata in Sardinia and Morocco. It is between 2.5 centimetres (0.98 in) and 6 centimetres (2.4 in) long. It has a headshield wider than the tailshield (pygidium), and in between them three thoracic body segments (somites).

<span class="mw-page-title-main">Artiopoda</span> Extinct group of arthropods

The Artiopoda is a grouping of extinct arthropods that includes trilobites and their close relatives. It was erected by Hou and Bergström in 1997 to encompass a wide diversity of arthropods that would traditionally have been assigned to the Trilobitomorpha. Trilobites, in part due to their mineralising exoskeletons, are by far the most diverse and long lived members of the clade, with most records of other members, which lack mineralised exoskeletons, being from Cambrian deposits.

<i>Squamacula</i> Extinct genus of arthropods

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.

<i>Parioscorpio</i> Extinct genus of enigmatic arthropod

Parioscorpio is an extinct genus of arthropod containing the species P. venator known from the Silurian-aged Waukesha Biota of the Brandon Bridge Formation near Waukesha, Wisconsin. This animal has gone through a confusing taxonomic history, being called an arachnid, crustacean, and an artiopodan arthropod at various points. This animal is one of the more famous fossil finds from Wisconsin, due to the media coverage it received based on its original description in 2020 as a basal scorpion.

<i>Arthroaspis</i> Extinct genus of arthropods

Arthroaspis is an extinct genus of arthropod known from the Cambrian aged Sirius Passet Lagerstatte in Greenland. It is relatively large in size for Cambrian arthropods, attaining a length of up to 215 mm. It is a common component of the Passet fauna, being located at multiple localities within the formation. It possessed 14 tergites. In the describing paper, it was recovered as a member of a non-monophyletic Artiopoda. It has subsequently been considered a potential close relative of nektaspids.

<span class="mw-page-title-main">Hymenocarina</span> Extinct order of arthropods

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.

2021 in arthropod paleontology is a list of new arthropod fossil taxa, including arachnids, crustaceans, insects, trilobites, and other arthropods that were announced or described, as well as other significant arthropod paleontological discoveries and events which occurred in 2021.

<i>Acheronauta</i> Possible extinct mandibulate arthropod

Acheronauta is a genus of extinct vermiform arthropod that lived in the early Silurian Waukesha biota fossil site in southeast Wisconsin. This arthropod was first discovered alongside the biota in 1985, but was not fully described until October 2022. This creature was recognized and described as a possible early mandibulate. This description is very important as much of the fauna of the biota remain undescribed, and its discovery has allowed for paleontologists to get a better grasp of the diversity of the arthropod fauna at the site. Multiple phylogenetic analyses were performed, and it seems that this arthropod forms a previously undiscovered clade with the Devonian stem-arthropod Captopodus, and the somewhat enigmatic group Thylacocephala.

2023 in arthropod paleontology is a list of new arthropod fossil taxa, including arachnids, crustaceans, trilobites, and other arthropods that were announced or described, as well as other significant arthropod paleontological discoveries and events which occurred in 2023.

References

  1. Raymond, P.E. (1920). "The appendages, anatomy, and relationships of trilobites". Memoirs of the Connecticut Academy of Arts and Sciences. 7: 1–169.
  2. 1 2 Budd, G. E. (1999). "A nektaspid arthropod from the Early Cambrian Sirius Passet fauna, with a description of retrodeformation based on functional morphology". Palaeontology. 42 (1): 99–122. Bibcode:1999Palgy..42...99B. doi: 10.1111/1475-4983.00064 . S2CID   128566745.
  3. Budd, G. E.; Jensen, S. (2000). "A critical reappraisal of the fossil record of the bilaterian phyla". Biological Reviews of the Cambridge Philosophical Society. 75 (2): 253–95. doi:10.1111/j.1469-185X.1999.tb00046.x. PMID   10881389. S2CID   39772232.
  4. Pérez-Peris, Francesc; Laibl, Lukáš; Lustri, Lorenzo; Gueriau, Pierre; Antcliffe, Jonathan B.; Enright, Orla G. Bath; Daley, Allison C. (March 2021). "A new nektaspid euarthropod from the Lower Ordovician strata of Morocco". Geological Magazine. 158 (3): 509–517. Bibcode:2021GeoM..158..509P. doi:10.1017/S001675682000062X. ISSN   0016-7568. S2CID   221859513.
  5. Mayers, Benjamin; Aria, Cédric; Caron, Jean‐Bernard (January 2019). Zhang, Xi‐Guang (ed.). "Three new naraoiid species from the Burgess Shale, with a morphometric and phylogenetic reinvestigation of Naraoiidae". Palaeontology. 62 (1): 19–50. Bibcode:2019Palgy..62...19M. doi:10.1111/pala.12383. ISSN   0031-0239. S2CID   135331371.
  6. 1 2 Pérez-Peris, Francesc; Laibl, Lukáš; Lustri, Lorenzo; Gueriau, Pierre; Antcliffe, Jonathan B; Bath Enright, Orla G; Daley, Allison C (2020-07-10). "A new nektaspid euarthropod from the Lower Ordovician strata of Morocco". Geological Magazine. 158 (3): 509–517. doi:10.1017/s001675682000062x. ISSN   0016-7568. S2CID   221859513.
  7. Bond, Andrew D.; Edgecombe, Gregory D. (January 2021). Zhang, Xi‐Guang (ed.). "Phylogenetic response of naraoiid arthropods to early–middle Cambrian environmental change". Palaeontology. 64 (1): 161–177. Bibcode:2021Palgy..64..161B. doi: 10.1111/pala.12516 . ISSN   0031-0239. S2CID   229454194.
  8. VANNIER, JEAN; CHEN, JUN-YUAN (2007-01-02). "Digestive system and feeding mode in Cambrian naraoiid arthropods". Lethaia. 35 (2): 107–120. doi:10.1111/j.1502-3931.2002.tb00072.x. ISSN   0024-1164.
  9. Caron, Jean-Bernard; Rudkin, David M.; Milliken, Stuart (November 2004). "A new Late Silurian (Pridolian) naraoiid (Euarthropoda: Nektaspida) from the Bertie Formation of southern Ontario, Canada—delayed fallout from the Cambrian explosion". Journal of Paleontology. 78 (6): 1138–1145. doi:10.1666/0022-3360(2004)078<1138:ANLSPN>2.0.CO;2. ISSN   0022-3360. S2CID   131595361.