Marrellomorpha

Marrellomorpha Temporal range: Middle Cambrian - Early Devonian, 508–390  Ma PreꞒ Ꞓ O S D C P T J K Pg N
Life restoration of Marrella from the Burgess Shale
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	† Marrellomorpha Beurlen, 1930
Subgroups
Acercostraca Marrellida

Marrellomorpha are an extinct group of arthropods known from the Cambrian to the Early Devonian. ^[1] 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.

Taxonomy

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 attached pairs of uniramous (single-branched) appendages, while Acercostraca generally have large ovoid carapaces that cover the entire upper half of the body, and five pairs of uniramous appendages attached to the head section. Both groups have unbranched antennae and a segmented trunk with biramous (divided into two branches) appendages. ^[2] Recent research has suggested the previously enigmatic Cambrian trilobite-like arthropods Skania and Primicaris belong to this group. ^{[3] [2]} Their phylogenetic position is uncertain, various studies have alternatively placed them in the Arachnomorpha as relatives of Artiopoda (trilobites and kin), as related to Mandibulata (the group containing crustaceans, insects and myriapods), or as stem group euarthropods. ^[2] The engimatic Cambrian arthropod Burgessia may be closely related to marrellomorphs. ^[4] 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. ^[2]

Internal taxonomy of Marrellomorpha after Moysiuk et al., 2022. ^[2]

• Acercostraca
  • Skania Burgess Shale, Canada, Cambrian (Miaolingian) ? Kaili Formation, China, Cambrian (Wuliuan)
  • Primicaris Burgess Shale, Canada, Cambrian (Miaolingian), Maotianshan Shales, China, Cambrian Stage 3
  • Vachonisiidae
    • Xylokorys Coalbrookdale Formation, England, Silurian (Wenlock)
    • Vachonisia Hunsrück Slate, Germany, Lower Devonian (Emsian)
    • Enosiaspis Fezouata Formation, Morocco, Early Ordovician (Floian)
• Marrellida
  • Marrella Balang Formation, China, Cambrian Stage 4 Kaili Formation, China, Cambrian (Wuliuan), Burgess Shale, Canada, Cambrian (Miaolingian)
  • Furca Letná Formation , Czech Republic, Upper Ordovician (Sandbian)
  • "Furca" mauretanica ( nomen nudum ) Fezouata Formation, Morocco, Early Ordovician (Floian)
  • "Mimetaster" florestaensis Floresta Formation Argentina, Lower Ordovician (Tremadocian)
  • Mimetasteridae
    • Tomlinsonus Kirkfield Formation, Canada, Late Ordovician (Katian)
    • Mimetaster Hunsrück Slate, Germany, Lower Devonian (Emsian)

Fragmentary taxa assigned to Marrellomorpha include Austromarrella from Cambrian Series 3 aged deposits in Australia, ^[5] and Dyrnwynia from the Ordovician (Darriwilian) aged Llanfallteg Formation of Wales, which in its original description was assigned to Marrelida. ^[6]

Phylogeny

After Legg, 2016. ^[7]

	Outgroups Marrellomorpha Marrellida Marrella splendens Mimetasteridae Furca bohemica Moroccan marrellid Mimetaster hexagonalis Acercostraca Skania fragilis Primicaris larvaformis "Skania" sundbergi Vachonisiidae Enosiaspis hrungnir Xylokorys chledophilia Vachonisia rogeri

After Moysiuk et al., 2022. ^[2]

Marrellomorpha Marrellida "Mimetaster" florestaensis Furca bohemica Mimetaster hexagonalis Tomlinsonus dimitrii Moroccan marrellid Marrella Palaeoisopus Haliestes Aquilonifer Acercostraca Vachonisia Xylokorys Enosiaspis Skania Primicaris Eoredlichia Mollisonia Waptia Tokummia Kylinxia

Taxa usually not considered marrellomorphs

Gallery

• 
  Ventral reconstruction of Xylokorys
• 
  Reconstruction of the head shield of Furca bohemica
• 
  Fossil of "Furca" mauretanica
• 
  Reconstruction of Mimetaster
• 
  Restoration of Vachonisia
• 
  Life restoration of Tomlinsonus
• 
  Life restoration of Primicaris
• 
  Diagram of Skania

References

1. Rak, Štĕpán; Ortega-Hernández, Javier; Legg, David A. (2011). "A revision of the Late Ordovician marrellomorph arthropod Furca bohemica from Czech Republic". Acta Palaeontologica Polonica. doi: 10.4202/app.2011.0038 .
2. Moysiuk, Joseph; Izquierdo-López, Alejandro; Kampouris, George E.; Caron, Jean-Bernard (July 2022). "A new marrellomorph arthropod from southern Ontario: a rare case of soft-tissue preservation on a Late Ordovician open marine shelf". Journal of Paleontology. 96 (4): 859–874. Bibcode:2022JPal...96..859M. doi:10.1017/jpa.2022.11. ISSN   0022-3360.
3. Legg, D.A. (2015-09-30). "The morphology and affinities of Skania fragilis (Arthropoda) from the middle Cambrian Burgess Shale". Bulletin of Geosciences: 509–518. doi: 10.3140/bull.geosci.1532 . ISSN   1802-8225.
4. 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. ISSN   0748-3007. PMID   34814371.
5. Haug, Joachim T.; Castellani, Christopher; Haug, Carolin; Waloszek, Dieter; Maas, Andreas (2012). "A Marrella-like arthropod from the Cambrian of Australia: A new link between "Orsten"-type and Burgess Shale assemblages". Acta Palaeontologica Polonica. 58 (3): 629–639. doi: 10.4202/app.2011.0120 .
6. Legg, David (2016). "A new marrellid arthropod from the Ordovician of Wales". Acta Palaeontologica Polonica. 61. doi: 10.4202/app.00246.2016 . ISSN   0567-7920.
7. Legg, David A. (2016-02-27). "An acercostracan marrellomorph (Euarthropoda) from the Lower Ordovician of Morocco". The Science of Nature. 103 (3): 21. Bibcode:2016SciNa.103...21L. doi:10.1007/s00114-016-1352-5. ISSN   1432-1904. PMID   26922777.