Cheloniellida

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Cheloniellida
Temporal range: Upper Ordovician–Early Devonian
20200415 Cheloniellon calmani.png
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
(unranked):† Artiopoda
(unranked):† Vicissicaudata
(unranked):† Cheloniellida
Broili, 1932
Genera

Cheloniellida is a taxon (usually referred to as an order [1] [2] ) of extinct Paleozoic arthropods. As of 2018, [2] 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. [2] It was later accepted as a member of Vicissicaudata within Artiopoda. [3] [4] [5]

Contents

Morphology

Ventral structures of the anterior region of Cheloniellon calmani, showing differentiation between appendages. 20200615 Cheloniellon calmani anterior ventral.png
Ventral structures of the anterior region of Cheloniellon calmani, showing differentiation between appendages.

The flattened, ovoid body of cheloniellid comprises an eye-bearing cephalon (head) and segmented trunk region, dorsally divided by a series of tergites (dorsal exoskeleton). The cephalon could be divided into procephalon and gnathocephalon. [2] Compared to other members of Artiopoda, the head shield (dorsal exoskeleton of cephalon) of cheloniellid is relatively reduced. [2] [3] The trunk is wider than the cephalon and is made up of 8-13 tergites. [2] The pleural (lateral) tips of first few tergites are directed anterolaterally, becoming increasingly posterolaterally directed rearward, giving the segmental boundaries between tergites a radiated appearance. [3] [2] The last trunk segment, also known as postabdomen, [4] is tiny and laterally encompassed by the pleural regions of previous tergite. [2]

Based on available materials, the cephalon comprises a pair of antennae and 5 pairs of uniramus appendages, with the posterior 4 pairs bore gnathobases. [6] There are evidences that the non-gnathobasic second cephalic appendages are specialized or even raptorial in some species. [6] [2] Each of the trunk segments (except the last one) has a pair of biramous appendages each consisting of a leg-like endopod and a shorter exopod. [2] The last trunk segment has a pair of spine/whip-like appendages referred as furcae [3] or cerci, [6] some species bore a medial spine between it which may or may not be a telson. [2]

Classification

Diagram of various cheloniellids and related taxa; (C) Tardisia broedeae, (D) Cheloniellon calmani, (E) Triopus draboviensis, (F) Paraduslia talimaae. Diagram of Tardisia broedeae and various Cheloniellids.png
Diagram of various cheloniellids and related taxa; (C) Tardisia broedeae , (D) Cheloniellon calmani, (E) Triopus draboviensis , (F) Paraduslia talimaae .

Phylogenetic position

The specialized second appendages (Pap, green) and gnathobasic appendages (Gap, cyan) of Cheloniellon had been compared to chelicerate prosomal appendages by some studies. 20200615 Cheloniellon calmani anterior ventral abbreviation.png
The specialized second appendages (Pap, green) and gnathobasic appendages (Gap, cyan) of Cheloniellon had been compared to chelicerate prosomal appendages by some studies.

While Boudreaux (1979) regarded Cheloniellida as a class, [8] further studies usually treat Cheloniellida as an order. [1] [2] Cheloniellida has a controversial phylogenetic position within arthropod higher classifications, with studies mainly around 20th century suggested it as a relative/member of either Crustacea, Trilobita, Chelicerata or Aglaspidida. [2] Some species even had been misidentified as polyplacophoran mollusks (chiton) when being first described. [9] Originally, the iconic cheloniellid Cheloniellon was believed to be a crustacean similar to trilobites. [10] [11] Subsequent authors suggests that it occupied a position intermediate between trilobitomorphs and chelicerates, [6] [7] while some also interpreted it as a sister group of crustaceans [12] or chelicerates [13] [14] as well. The suggested close relationship between cheloniellids and chelicerates was inferred by the gnathobasic appendages similar to those of merostomes (e.g. Xiphosurans and Eurypterids), and the hypothesis that the chelicerates arose from trilobitomorphs through the loss of deutocerebral antennae (i.e. first antennae) and specialization of tritocerebral appendages into chelicerae (comparable to the cheloniellid antennae and specialized 2nd appendages), [6] [7] a scenario which is not supported by gene expression, [15] [16] [17] [18] [19] neuroanatomical [20] [21] and developmental [22] evidences (suggests that chelicerae are in fact deutocerebral).

Artiopoda
Summarized phylogeny of Artiopoda with focus on Cheloniellida, based on Lerosey-Aubril et al. (2017). [4]

As of 21st century, Cheloniellida was mostly found to form a clade with Aglaspidida and Xenopoda (e.g. Sidneyia and Emeraldella ). [23] [3] [24] [25] [4] The clade was formally named Vicissicaudata in 2013, [3] united by a differentiated terminal trunk area (postabdomen) that bears a pair of non-leg-like appendages. [3] [4] Numerous phylogenetic analyses also retrieved Vicissicaudata within Artiopoda, [4] a diverse arthropod taxon comprising trilobites and similar fossil taxa that may [26] [24] or may not [3] [25] [4] be closely related to chelicerates.

Included genera and species

The unambiguous members of cheloniellids shown as follows:

In 2018, a new species (informally named " Latromirus " in an unpublished thesis from 2016 [27] ) was described in a preprint by Wendruff et al. [2] According to Braddy & Dunlop 2021, the enigmatic Parioscorpio may have cheloniellid affinities, [28] however that interpretation is denied by Van Roy et al. 2022, only remaining specimen UWGM 2439 (holotype specimen of "Latromirus") as possible cheloniellid. [29]

Related Research Articles

<span class="mw-page-title-main">Chelicerata</span> Subphylum of arthropods

The subphylum Chelicerata constitutes one of the major subdivisions of the phylum Arthropoda. Chelicerates include the sea spiders, horseshoe crabs, and arachnids, as well as a number of extinct lineages, such as the eurypterids and chasmataspidids.

<span class="mw-page-title-main">Sea spider</span> Class of marine arthropods

Sea spiders are marine arthropods of the class Pycnogonida, hence they are also called pycnogonids. The class include the only now-living order Pantopoda, alongside a few fossil species which could trace back to the early or mid Paleozoic. They are cosmopolitan, found in oceans around the world. The over 1,300 known species have leg spans ranging from 1 mm (0.04 in) to over 70 cm (2.3 ft). Most are toward the smaller end of this range in relatively shallow depths; however, they can grow to be quite large in Antarctic and deep waters.

<span class="mw-page-title-main">Xiphosura</span> Order of marine chelicerates

Xiphosura is an order of arthropods related to arachnids. They are more commonly known as horseshoe crabs. They first appeared in the Hirnantian. Currently, there are only four living species. Xiphosura contains one suborder, Xiphosurida, and several stem-genera.

<span class="mw-page-title-main">Arthropod head problem</span> Dispute concerning the evolution of arthropods

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, and crustaceans, as well as 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.

<span class="mw-page-title-main">Chasmataspidida</span> Order of arthropods

Chasmataspidids, sometime referred to as chasmataspids, are a group of extinct chelicerate arthropods that form the order Chasmataspidida. Chasmataspidids are probably related to horseshoe crabs (Xiphosura) and/or sea scorpions (Eurypterida), with more recent studies suggest that they form a clade (Dekatriata) with Eurypterida and Arachnida. Chasmataspidids are known sporadically in the fossil record through to the mid-Devonian, with possible evidence suggesting that they were also present during the late Cambrian. Chasmataspidids are most easily recognised by having an opisthosoma divided into a wide forepart (preabdomen) and a narrow hind part (postabdomen) each comprising 4 and 9 segments respectively. There is some debate about whether they form a natural group.

<i>Urokodia</i> Extinct genus of artiopod

Urokodia is an extinct genus of arthropod from the early Cambrian. The only known species is Urokodia aequalis from the Maotianshan Shales of China based on some 15 specimens.

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

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 Cambrian to the upper Ordovician.

<i>Weinbergina</i> Extinct genus of chelicerate

Weinbergina is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Fossils of the single and type species, W. opitzi, have been discovered in deposits of the Devonian period in the Hunsrück Slate, Germany.

<i>Schinderhannes bartelsi</i> Extinct species of radiodont

Schinderhannes bartelsi is a species of hurdiid radiodont (anomalocaridid), known from one specimen from the Lower Devonian Hunsrück Slates. Its discovery was astonishing because the latest definitive radiodonts were known only from the Early Ordovician, at least 66 million years earlier than this taxon.

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

Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and were used for a variety of functions. Radiodonts were among the earliest large predators, 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 gainesi, 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.

<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. Some authors have proposed that they may be closely related to sea spiders (Pycnogonida) within Chelicerata though the cladistical support for such a relationship is relatively weak.

<span class="mw-page-title-main">Synziphosurina</span> Group of arthropods

Synziphosurina is a paraphyletic group of chelicerate arthropods previously thought to be basal horseshoe crabs (Xiphosura). It was later identified as a grade composed of various basal euchelicerates, eventually excluded from the monophyletic Xiphosura sensu stricto and only regarded as horseshoe crabs under a broader sense. Synziphosurines survived at least since early Ordovician to early Carboniferous in ages, with most species are known from the in-between Silurian strata.

<i>Mollisonia</i> Extinct genus of Cambrian Arthropod

Mollisonia is an extinct genus of Cambrian arthropod. 4 species had been described from North America and China. Studies suggest it is a basal member of Chelicerata, a group which includes horseshoe crabs and arachnids.

<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 abundance of findings owing to their mineralized exoskeletons, are by far the best recorded, diverse, and long lived members of the clade. Other members, which lack mineralised exoskeletons, are known mostly from Cambrian deposits.

<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>Camanchia</i> Extinct genus of arthropods

Camanchia is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Camanchia was regarded as part of the clade Prosomapoda. Fossils of the single and type species, C. grovensis, have been discovered in deposits of the Silurian period in Iowa, in the United States. Alongside Venustulus, Camanchia is one of the only Silurian synziphosurine with fossil showing evidence of appendages.

<i>Palaeoisopus</i> Extinct genus of sea spiders

Palaeoisopus is a genus of fossil pycnogonid. The only known species is Palaeoisopus problematicus from the Lower Devonian Hunsrück Slate of Germany. It is characterized by several features unusual for a pycnogonid, such as swimming legs of different sizes and a long, segmented abdomen.

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

Cheloniellon is a monotypic genus of cheloniellid arthropod, known only by one species, Cheloniellon calmani, discovered from the Lower Devonian Hunsrück Slate of Germany.

<span class="mw-page-title-main">Deuteropoda</span> Clade of arthropods

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.

<i>Bundenbachiellus</i> Extinct arthropod genus

Bundenbachiellus is an extinct genus of arthropod described from the Lower Devonian Hunsrück Slate of Germany. This genus is known from only one species, B. giganteus. Alongside its possible relative Enalikter from Silurian, it is possible that genus is late-living example of Megacheira, "great-appendage arthropod".

References

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  2. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 Wendruff, Andrew James, et al. "New cheloniellid arthropod with large raptorial appendages from the Silurian of Wisconsin, USA." BioRxiv (2018): 407379.
  3. 1 2 3 4 5 6 7 8 Ortega-Hernández, Javier; Legg, David A.; Braddy, Simon J. (February 2013). "The phylogeny of aglaspidid arthropods and the internal relationships within Artiopoda". Cladistics. 29 (1): 15–45. doi:10.1111/j.1096-0031.2012.00413.x. PMID   34814371. S2CID   85744103.
  4. 1 2 3 4 5 6 7 Lerosey-Aubril, Rudy; Zhu, Xuejian; Ortega-Hernández, Javier (2017-09-11). "The Vicissicaudata revisited – insights from a new aglaspidid arthropod with caudal appendages from the Furongian of China". Scientific Reports. 7 (1): 11117. Bibcode:2017NatSR...711117L. doi: 10.1038/s41598-017-11610-5 . ISSN   2045-2322. PMC   5593897 . PMID   28894246.
  5. ↑ Du, Kun-sheng; Ortega-Hernández, Javier; Yang, Jie; Zhang, Xi-guang (2019). "A soft-bodied euarthropod from the early Cambrian Xiaoshiba Lagerstätte of China supports a new clade of basal artiopodans with dorsal ecdysial sutures". Cladistics. 35 (3): 269–281. doi: 10.1111/cla.12344 . ISSN   0748-3007. PMID   34622993. S2CID   89985331.
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  8. ↑ Boudreaux H. B., 1979. Significance of intersegmental tendon system in arthropod phylogeny and monophyletic classification of Arthropoda. pp. 551-586. In.: Gupta A.P. [Ed.]. Arthropod Phylogeny. Van Nostrand Reinhold Company, N.Y., 1-762
  9. ↑ Chlupác I. The enigmatic arthropod Duslia from the Ordovician of Czechoslovakia. Palaeontol. 1988; 31:611–620.
  10. ↑ BROILI, F. (1932): Ein neuer Crustacee aus dem rheinischen Unterdevon. -- Sitzungsber. bayer. Akad. Wiss.: 27--38; Miinchen.
  11. ↑ BROILI, F. (1933): Ein zweites Exemplar von Cheloniellon. -- Sitzungsber. bayer. Akad. Wiss.: 11--32; Miinchen.
  12. ↑ Delle Cave, L. and A.M. Simonetta (1991) Early Palaeozoic arthropods and problems of arthropod phylogeny; with some notes on taxa of doubtful affinities. In The Early Evolution of Metazoa and the Significance of Problematic Taxa, eds. A. M. Simonetta and S. Conway Morris. Cambridge University Press, Cambridge, England. Vol. 189–244.
  13. ↑ Wills, M. A., Briggs, D. E. G., Fortey, R. A. & Wilkinson, M. 1995. The significance of fossils in understanding arthropod evolution. Verhandlungen der deutschen zoologischen Gesselschaft 88, 203–15.
  14. ↑ Dunlop, J. A.; Selden, P. A. (1998), Fortey, R. A.; Thomas, R. H. (eds.), "The early history and phylogeny of the chelicerates", Arthropod Relationships, The Systematics Association Special Volume Series, Dordrecht: Springer Netherlands, pp. 221–235, doi:10.1007/978-94-011-4904-4_17, ISBN   978-94-011-4904-4
  15. ↑ Telford, Maximilian J.; Thomas, Richard H. (1998-09-01). "Expression of homeobox genes shows chelicerate arthropods retain their deutocerebral segment". Proceedings of the National Academy of Sciences. 95 (18): 10671–10675. Bibcode:1998PNAS...9510671T. doi: 10.1073/pnas.95.18.10671 . ISSN   0027-8424. PMC   27953 . PMID   9724762.
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  17. ↑ Jager, Muriel; Murienne, Jérôme; Clabaut, Céline; Deutsch, Jean; Guyader, Hervé Le; Manuel, Michaël (2006). "Homology of arthropod anterior appendages revealed by Hox gene expression in a sea spider". Nature. 441 (7092): 506–508. Bibcode:2006Natur.441..506J. doi:10.1038/nature04591. ISSN   1476-4687. PMID   16724066. S2CID   4307398.
  18. ↑ Manuel, Michaël; Jager, Muriel; Murienne, Jérôme; Clabaut, Céline; Guyader, Hervé Le (2006-07-01). "Hox genes in sea spiders (Pycnogonida) and the homology of arthropod head segments". Development Genes and Evolution. 216 (7): 481–491. doi:10.1007/s00427-006-0095-2. ISSN   1432-041X. PMID   16820954. S2CID   833103.
  19. ↑ Brenneis, Georg; Ungerer, Petra; Scholtz, Gerhard (2008). "The chelifores of sea spiders (Arthropoda, Pycnogonida) are the appendages of the deutocerebral segment". Evolution & Development. 10 (6): 717–724. doi:10.1111/j.1525-142X.2008.00285.x. ISSN   1525-142X. PMID   19021742. S2CID   6048195.
  20. ↑ Mittmann, Beate; Scholtz, Gerhard (2003-02-01). "Development of the nervous system in the "head" of Limulus polyphemus (Chelicerata: Xiphosura): morphological evidence for a correspondence between the segments of the chelicerae and of the (first) antennae of Mandibulata". Development Genes and Evolution. 213 (1): 9–17. doi:10.1007/s00427-002-0285-5. ISSN   1432-041X. PMID   12590348. S2CID   13101102.
  21. ↑ Harzsch, Steffen; Wildt, Miriam; Battelle, Barbara; Waloszek, Dieter (2005-07-01). "Immunohistochemical localization of neurotransmitters in the nervous system of larval Limulus polyphemus (Chelicerata, Xiphosura): evidence for a conserved protocerebral architecture in Euarthropoda". Arthropod Structure & Development. Arthropod Brain Morphology and Evolution. 34 (3): 327–342. doi:10.1016/j.asd.2005.01.006. ISSN   1467-8039.
  22. ↑ Mittmann, Beate; Scholtz, Gerhard (2003-02-01). "Development of the nervous system in the "head" of Limulus polyphemus (Chelicerata: Xiphosura): morphological evidence for a correspondence between the segments of the chelicerae and of the (first) antennae of Mandibulata". Development Genes and Evolution. 213 (1): 9–17. doi:10.1007/s00427-002-0285-5. ISSN   1432-041X. PMID   12590348. S2CID   13101102.
  23. ↑ Edgecombe, Gregory D.; García-Bellido, Diego C.; Paterson, John R. (2011). "A New Leanchoiliid Megacheiran Arthropod from the Lower Cambrian Emu Bay Shale, South Australia". Acta Palaeontologica Polonica. 56 (2): 385–400. doi: 10.4202/app.2010.0080 . hdl: 10261/61352 . ISSN   0567-7920.
  24. 1 2 Legg, David A. (2014). "Sanctacaris uncata: the oldest chelicerate (Arthropoda)". Naturwissenschaften. 101 (12): 1065–1073. Bibcode:2014NW....101.1065L. doi:10.1007/s00114-014-1245-4. ISSN   0028-1042. PMID   25296691. S2CID   15290784.
  25. 1 2 Edgecombe, Gregory D.; Paterson, John R.; García-Bellido, Diego C. (2017). "A new aglaspidid-like euarthropod from the lower Cambrian Emu Bay Shale of South Australia". Geological Magazine. 154 (1): 87–95. Bibcode:2017GeoM..154...87E. doi:10.1017/S0016756815001053. ISSN   0016-7568. S2CID   133058010.
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  27. ↑ Wendruff, Andrew J. (2016). Paleobiology and Taphonomy of Exceptionally Preserved Organisms from the Brandon Bridge Formation (Silurian), Wisconsin, USA (Thesis). The Ohio State University.
  28. ↑ Braddy, S.J.; Dunlop, J.A. (2021). "A sting in the tale of Parioscorpio venator from the Silurian of Wisconsin: is it a cheloniellid arthropod?". Lethaia. 54 (1): 1–7. doi:10.1111/let.12457. S2CID   245285654.
  29. ↑ Roy, Peter Van; Rak, Štěpán; Budil, Petr; Fatka, Oldřich (2022-06-13). "Redescription of the cheloniellid euarthropod Triopus draboviensis from the Upper Ordovician of Bohemia, with comments on the affinities of Parioscorpio venator". Geological Magazine. 159 (9): 1471–1489. doi:10.1017/S0016756822000292. hdl:1854/LU-8756253. ISSN   0016-7568. S2CID   249652930.
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