Squamacula

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Squamacula
Temporal range: Cambrian Series 2
S. clypeata.png
Reconstruction of Squamacula clypeata
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
(unranked): Artiopoda
Genus: Squamacula
Hou and Bergström, 1997
Type species
Squamacula clypeata
Hou and Bergström, 1997 [1]
Species
  • Squamacula clypeataHou and Bergström, 1997
  • Squamacula buckorumPaterson et al. 2012

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. [2] In 2012 a second species S. buckorum was described from the Emu Bay Shale of Australia. [3]

Contents

Etymology

The genus Squamacula is derived from the Latin word squama, meaning scale, and the diminutive suffix -culus, indicating that the animal is relatively small. The species clypeata is derived from the Latin word clypeatus, meaning shield-shaped. It was named this in reference to its shield-like outline. [1]

Description

Squamacula clypeata is flattened (dorsoventrally). It has 11 segments in total: the cephalon (the head), nine thoracic tergites (each of which covers a somite), and one pygidium. [4] It has a doublure, a piece of exoskeleton that covers part of the underside of the animal. Its doublure is, on average, about twice as long as the length of its cephalon. It has been hypothesized that this large doublure could have aided the animal in digging through sediment to find food, as it was thought to have been both a predator of small animals and a scavenger. [4]

Squamacula clypeata has one pair of antennae attached to its cephalon, but no other appendages or auditory or visual structures. [4] No mouth is present in the specimens, but the gut is present, so S. clypeata must have had a mouth, which is thought to have been between the cephalon and doublure on the underside. [4]

Squamacula clypeata has one pair of biramous appendages (an appendage which branches into two) per thoracic tergite. [4] The endopod (inside branch) has seven segments, the first six of which are roughly even in size and shape, with small spines on the underside, and the last of which terminates with a claw and appears to have been used for walking and eating, as the claw could be useful in grabbing hold of and tearing up food. [4] The exopod (outside branch) was not segmented, but flap-like, with setae (bristles) near the tip, which may have been used for both locomotion and respiration. [4]

S. buckorum differs from S. clypeata in that it has 11 thoracic tergites, which appear to exhibit less overlap that those of S. clypeata, though the differences between the two species are relatively minor. The species also preserves elements of the gut tract not preserved in S. clypeata , which show that it was narrow annulated tube running along the midline of the animal, with three or possibly four pairs of midgut glands within the cephalic shield, with other gland pairs extending along the trunk. [3]

Taxonomy

In phylogenetic analyses, Squamacula has consistently been placed as the earliest diverging and most primitive member of Artiopoda, a clade that includes trilobites and a variety of morphologically similar soft-bodied forms. [5]

Cladogram after Jiao et al. 2021. [5]

Artiopoda 

Squamacula clypeata

Protosutura  

Zhiwenia coronata

Australimicola spriggi

Acanthomeridion serratum

Acanthomeridion anacanthus

Bailongia longicaudata

Trilobitomorpha  

Retifacies abnormalis

Kwanyinaspis maotiashanensis

Trilobita

Olenoides serratus

Eoredlichia intermedia

Conciliterga

Haifengella corona

Kuamaia lata

Helmetia expansa

Tegopelte gigas

Skioldia aldna

Saperion glumaceum

Xandarellida

Cindarella eucalla

Luohuilinella deletres

Luohuilinella rarus

Sinoburius lunaris

Xandarella spectaculum

Phytophilaspis pergamena

Nektaspida

Buenaspis forteyi

Tariccoia arrusensis

Soomaspis splendida

Liwia convexa

Naraoia spinosa

Naraoia compacta

Misszhouia longicaudata

Vicissicaudata  

Sidneyia inexpectans

Emeraldella brocki

Cheloniellida

Cheloniellon calmani

Triopus draboviensis

Duslia insignis

Kodymirus vagans

Eozetetes gemmelli

Aglaspidida

Beckwithia typa

Chlupacaris dubia

Quasimodaspis brentsae

Tremaglaspis unite

Tremaglaspis vanroyi

Brachyaglaspis singularis

Cyclopites vulgaris

Australaglaspis stonyensis

Uarthrus instabilis

Flobertia kochi

Aglaspella granulifera

Gogglops ensifer

Glypharthrus simplex

Aglaspis spinifer

Chraspedops modesta

Glypharthrus thomasi

Glypharthrus trispinicaudatus

Glypharthrus magnoculus

A close relationship has been proposed with the genus Thulaspis from the Sirius Passet site of Greenland. [6]

Related Research Articles

<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.

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

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.

<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.

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.

<i>Acanthomeridion</i> Extinct genus of Cambrian Era Arthropod

Acanthomeridion is an extinct arthropod found in the Chengjiang fauna deposits of China. In 1997, it was placed in its own, monotypic family, Acanthomeridiidae. It is known from eight specimens, all found in China.

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

Plenocaris plena is a Cambrian arthropod with a bivalved carapace, and is known from the Burgess shale and Chengjiang. Originally described as a species of Yohoia by Walcott in 1912, it was placed into its own genus in 1974.

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

Kuamaia is an extinct genus of artiopodan in the phylum Arthropoda. Fossils of the type species K. lata were discovered in the Chengjiang biota. The other species in the genus,K. muricata has also been identified there, but neither species has been found elsewhere.

<span class="mw-page-title-main">Kunmingella</span>

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.

<i>Pygmaclypeatus</i> Extinct genus of arthropod

Pygmaclypeatus is a genus of trilobite-like arthropod from the Cambrian aged Chengjiang biota of southern China. The carapace is flat and broad, and slightly shorter than it is wide, reaching a maximum width of 17.5 millimetres (0.69 in) and length of 14 millimetres (0.55 in). The headshield makes up about 25% of the total length, and has attached a pair of antennae, as well as four pairs of biramous limbs. The trunk has 6 tergites, each associated with a pair of biramous limbs along with a terminal pygidium associated with four pairs of biramous appendages and a segmented short tailspine. The well developed paddle-like exopodites on the trunk and pygidium limbs along with its small size suggests that it was an effective swimmer with a strong power stroke, and that it probably had a nektobenthic mode of life, swimming close to the ocean floor. Given its delicate spinose endites on the limbs it likely only consumed soft food and organic particles. It is considered to be closely related to Retifacies from the same deposit with shared characters including a segmented tailspine. It has been placed as a member of Artiopoda, possibly along with Retifacies the earliest diverging lineage of the Trilobitomorpha.

<i>Miraluolishania</i> Extinct genus of lobopodians

Miraluolishania is an extinct lobopodian known from Chengjiang County in China. It is remarkable for the possession of lensed pit-eyes. The only species, Miraluolishania haikouensis, was described from the Maotianshan Shales at Haikou by Jianni Liu and Degan Shu in 2004. In 2009, a team of palaeontologists at the Yunnan University, led by Xiaoya Ma reported the discovery of 42 other specimens from Haikou. With the help of Swiss palaeontologist Jan Bergström, Ma and Hou came to the conclusion that all the specimens were the same species as Luolishania; another lobopod discovered from the Chengjian in 1989. Chengjian is 40 kms from Haikou and the fossil fauna are different. A reassessment by Liu and Shu's team at the Northwest University in 2008 established that Luolishania and Miraluolishania are distinct animals.

<i>Phytophilaspis</i> Extinct genus of trilobites

Phytophilaspis is a phosphatized genus of trilobite-like arthropod with eyes, found in association with algal remains. It dwelt in well-lit, shallow waters.

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

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.

<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">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>Luohuilinella</i> Genus of arthropods (fossil)

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<i>Retifacies</i> Species of arthropod

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.

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

Fuxianhuiida is an extinct clade of arthropods from the Cambrian of China. All currently known species are from Cambrian Series 2 aged deposits in Yunnan Province, including the Chengjiang biota. They are generally considered to be close to the base of Euarthropoda, but have also been considered to be early mandibulates. Many specimens are known with exceptional soft tissue preservation, including preserved guts and neural tissue, which given their basal phylogenetic position makes them important in understanding the evolution of arthropoda as a whole. They reach a size of up to 15 cm, and are interpreted as benthic predators and scavengers. The fuxianhuiid exoskeleton is unmineralised, and the number of tergites ranges from 15 to over 40. The cephalon is covered by a head shield and contains stalked eyes connected by the anterior sclerite, antennae, a butterfly shaped hyposome and a posterior facing mouth. Fuxianhuiids possess specialized post-antennal appendages with serrated edges used for food processing. The presence of gnathobases in members of Chengjiangocardidae suggests that they were capable of durophagy. In most fuxianhuiids, the thorax tergites narrow posteriorly, terminating in either a swimming paddle or paired flukes with a tailspine. In members of Fuxianhuiidae the thorax is divided into two sections, the anterior wide opisthothorax and the posterior narrow limbless tail-like abdomen.

<i>Cheloniellon</i>

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

Luolishania is an extinct genus of lobopodian panarthropod and known from the Lower Cambrian Chiungchussu Formation of the Chengjiang County, Yunnan Province, China. A monotypic genus, it contains one species Luolishania longicruris. It was discovered and described by Hou Xian-Guang and Chen Jun-Yuan in 1989. It is one of the superarmoured Cambrian lobopodians suspected to be either an intermediate form in the origin of velvet worms (Onychophora) or basal to at least Tardigrada and Arthropoda. It is the basis of the family name Luolishaniidae, which also include other related lobopods such as Acinocricus, Collinsium, Facivermis, and Ovatiovermis. Along with Microdictyon, it is the first lobopodian fossil discovered from China.

References

  1. 1 2 Hou, Xianguang; Bergström, Jan (1997). Arthropods of the Lower Cambrian Chengjiang fauna, southwest China. Bergström. Oslo: Scandinavian University Press. ISBN   8200376931. OCLC   38305908.[ page needed ]
  2. The Cambrian fossils of Chengjiang, China : the flowering of early animal life. Hou, Xianguang. (Second ed.). Chichester, West Sussex. 8 March 2017. ISBN   9781118896310. OCLC   970396735.{{cite book}}: CS1 maint: location missing publisher (link) CS1 maint: others (link)
  3. 1 2 Paterson, John R.; García-Bellido, Diego C.; Edgecombe, Gregory D. (March 2012). "New artiopodan arthropods from the early Cambrian Emu Bay Shale Konservat-Lagerstätte of South Australia". Journal of Paleontology. 86 (2): 340–357. Bibcode:2012JPal...86..340P. doi:10.1666/11-077.1. ISSN   0022-3360. S2CID   87520649.
  4. 1 2 3 4 5 6 7 Zhang, Xingliang; Han, Jian; Zhang, Zhifei; Liu, Huqin; Shu, Degan (2004). "Redescription of the Chengjiang arthropod Squamacula clypeata Hou and Bergström, from the Lower Cambrian, south-west China". Palaeontology. 47 (3): 605–617. Bibcode:2004Palgy..47..605Z. doi: 10.1111/j.0031-0239.2004.00363.x . ISSN   1475-4983.
  5. 1 2 Jiao, De-Guang; Du, Kun-Sheng; Zhang, Xi-Guang; Yang, Jie; Eggink, Daniel (May 2022). "A new small soft-bodied non-trilobite artiopod from the Cambrian Stage 4 Guanshan Biota". Geological Magazine. 159 (5): 730–734. Bibcode:2022GeoM..159..730J. doi: 10.1017/S0016756821001254 . ISSN   0016-7568.
  6. Berks, Harry O.; Lunde Nielsen, Morten; Flannery‐Sutherland, Joseph; Thorshøj Nielsen, Arne; Park, Tae‐Yoon S.; Vinther, Jakob (May 2023). "A possibly deep branching artiopodan arthropod from the lower Cambrian Sirius Passet Lagerstätte (North Greenland)". Papers in Palaeontology. 9 (3). doi: 10.1002/spp2.1495 . ISSN   2056-2799.