Isoxys

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Isoxys
Temporal range: Atdabanian to Paibian
Artistic reconstruction of Isoxys curvirostratus.jpg
Isoxys curvirostratus from the Chengjiang Biota
Isoxys acutangulus 2.jpg
Fossil of Isoxys acutangulus from the Burgess Shale
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Order:† Isoxyida
Family:† Isoxyidae
Genus:† Isoxys
Walcott, 1890
Species

See text

Isoxys (meaning "equal surfaces") is a genus of extinct bivalved Cambrian arthropod; the various species of which are thought to have been freely swimming predators. [1] It had a pair of large spherical eyes (which are the most commonly preserved feature of the soft-bodied anatomy), [2] and two large frontal appendages used to grasp prey. [1]

Contents

Description

Diagram of I. curvirostratus showing frontal (b) and biramous appendages (c,d) (c) and (d) represent first four limb pairs and remaining posterior limbs, respectively Figure 3. Morphological reconstruction and phylogenetic position of Isoxys curvirostratus (cropped).jpg
Diagram of I. curvirostratus showing frontal (b) and biramous appendages (c,d) (c) and (d) represent first four limb pairs and remaining posterior limbs, respectively

Species of Isoxys have roughly semicircular bivalved carapaces, which vary in morphology between species. The front and rear edges of the carapaces bear forward and posterior facing spines, respectively which in some species are greatly elongated. [3] The carapaces of Isoxys are typically 1.1 to 3.3 centimetres (0.43 to 1.30 in) in length, excluding the spines, though some species are known to reach over 6 centimetres (2.4 in). In long-spined species when including spine length, some specimens exceed 10 centimetres (3.9 in). [4] The opening angle of the carapace was close to vertical, giving it a narrow profile when viewed from above. [3]

The head had a pair of large spherical stalked eyes, as well as a pair of upward-curling frontal appendages, which have a varying number of podomeres/segments, depending on the species. Most podomeres on the frontal appendages have upward-facing endite spines, with the number and placement of spines varying between species. The last podomere of the frontal appendage is a curved terminal claw. The trunk lacks clear segmentation (arthrodization). Along the body are pairs of biramous appendages, the counts of which differ between species (Isoxys curvirostratus has 14, while Isoxys auritus has 11). In Isoxys curvirostratus the endopods have well defined segments/podomeres. The first four biramous limb pairs of Isoxys curvirostratus were shorter than the remaining pairs, with their endopods having borne well developed endite (inward facing) spines and ended with a curved subchela (claw), with these features absent in the endopods of the more posterior limbs. The exopods of Isoxys curvirostratus had thick paddle-shaped lamellae which projected perpendicular to the limb axis. [5] Isoxys volucris from Greenland had paddle-shaped exopods suggested to have been fringed with setae. [6] The end of the trunk has paired telson flaps. [5]

Ecology

Species of Isoxys are thought to have been actively swimming predators, using its frontal appendages to capture soft-bodied prey, with the frontalmost pairs of biramous limbs aiding in food processing. [5] The various Isoxys species are thought to have occupied a variety of niches, from swimming just above the seafloor (nektobenthic) to open ocean swimmers (pelagic). [3] Swimming was likely accomplished by rhythmic movement of the legs. [5] Eyes of different specimens appear to have been adapted to different light intensities; one specimen of I. auritus was either crepuscular in shallow water, or lived in waters around 140 m below the sea surface; whereas another was morphologically adapted to a diurnal light intensity in shallow waters. [7] Isoxys species with elongated carapace spines are likely to have engaged in vertical migration up and down the water column, like many modern marine invertebrates. [3] Specimens of Isoxys minor have been found with eggs adhered to the inner surface of the carapace, indicating they engaged in brood care. The brood size was large, with approximately 300 eggs, each 0.5 millimetres (3⁄128 in) across per (presumably female) individual. Egg bearing individuals were only around half the maximum size, suggesting that individuals continued to grow beyond sexual maturity. [8]

Taxonomy

Specimen of Isoxys longissimus Isoxys longissimus.jpg
Specimen of Isoxys longissimus

Isoxys is thought to be one of the basalmost known stem-group arthropods, showing a combination of traits characteristic of more primitive stem-group arthropods like radiodonts, like lacking an arthrodized (sclerotized and jointed) trunk exoskeleton, with those of modern arthropods, like possessing sclerotized and jointed (arthropodized) biramous limbs. It is one of two genera within the family Isoxyidae, alongside Surusicaris . [5] A close relationship to the bivalved arthropod Tuzoia had historically been proposed based on the similarities of some aspects of their carapaces, [9] but preserved soft tissues of Tuzoia described in 2022 suggest that they are not closely related. [10]

Species

20 species of Isoxys have been described, [3] which have a global distribution, having been found in North America, Siberia, Australia, China and Europe, spanning from Cambrian Series 2 into the Miaolingian. [11]

Indeterminate species are also known from the Spence Shale of Utah, dating to the Miaolingian, as well as the Kaili Biota in Guizhou, China, which also dates to the Miaolingian. [11]

See also

Related Research Articles

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<i>Waptia</i> Cambrian arthropod

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

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

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<i>Chuandianella</i> Extinct genus of Cambrian animals

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

Clypecaris is genus of bivalved Cambrian arthropod known from the Chengjiang biota of Yunnan, China. The genus was initially described for the type species C. pteroidea by Hou, 1999. A second species C. serrata was described by Yang et al. in 2016. The species are primarily distinguished by the presence of a serrated edge on the front of the carapace of C. serrata. C. serrata is noted for the modification of an anterior pair of limbs into spined grasping appendages, indicating a predatory lifestyle. It is unknown whether a similar structure was present in C. pteroidea.Clypecaris is considered to likely be a member or a close relative of Hymenocarina, and is closely related to Perspicaris. As well as to Ercaicunia.

<i>Ercaicunia</i>

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<span class="mw-page-title-main">Pectocaris</span> Extinct genus of crustaceans

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<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

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<i>Retifacies</i> Species of arthropod

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<i>Erratus</i> Extinct genus of Cambrian arthropod

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<i>Laminacaris</i> Genus of extinct arthropods

Laminacaris is a genus of extinct stem-group arthropods (Radiodonta) that lived during the Cambrian period. It is monotypic with a single species Laminacaris chimera, the fossil of which was described from the Chengjiang biota of China in 2018. Around the same time, two specimens that were similar or of the same species were discovered at the Kinzers Formation in Pennsylvania, USA. The first specimens from China were three frontal appendages, without the other body parts.

<i>Jugatacaris</i> Extinct genus of bivalved arthropod

Jugatacaris is an extinct genus of bivalved arthropod known from a single species, Jugatacaris agilis found in the Cambrian Stage 3 aged Chengjiang biota of Yunnan, China. The carapace is around 28 to 37 millimetres in length, with a pronounced ridge at the top of the carapace separating the two valves, which formed a fin-like structure raised above the carapace. The head has a pair of stalked eyes, as well as a dumbbell shaped medial eye between them. The head also bore a pair of mandibles as well as at least one and possibly two pairs of antennules. The trunk had up to 65 segments, each with biramous appendages. The appendages had thin endopods with 30 podomeres, each bearing a spiny endite, with the endopods ending with a terminal claw. The appendages also had overlapping flap-like exopods, which are elongated, being at maximum eight times as long as they are wide, which on their posterior edge are covered with setae. The trunk ended with a forked tail. It was likely an actively swimming filter feeder, using its constantly beating appendages to sift food from the water column, which was then passed forward along the U-shaped food groove between the appendage pairs towards the mouth. While initially placed as a crustaceanomorph, later studies considered to be a member of Hymenocarina, which contains numerous other similar bivalved Cambrian arthropods.

<i>Surusicaris</i>

Surusicaris is an extinct genus of bivalved arthropod, known from the Cambrian Burgess Shale of British Columbia, Canada. It is considered to be closely related to Isoxys, and like it has spined grasping frontal appendages.

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

Isoxyids are members of the order Isoxyida and the family Isoxyidae, a group of basal arthropods that existed during the Cambrian period.

References

  1. 1 2 Vannier, J.; Garcia-Bellido, C.; Hu, X.; Chen, L. (Jul 2009). "Arthropod visual predators in the early pelagic ecosystem: evidence from the Burgess Shale and Chengjiang biotas". Proceedings: Biological Sciences. 276 (1667): 2567–2574. doi:10.1098/rspb.2009.0361. ISSN   0962-8452. PMC   2686666 . PMID   19403536.
  2. ↑ García-Bellido, D.; Vannier, J.; Collins, D. (2009). "Soft-part preservation in two species of the arthropod Isoxys from the middle Cambrian Burgess Shale of British Columbia, Canada" (PDF). Acta Palaeontologica Polonica. 54 (4): 699. doi: 10.4202/app.2009.0024 . S2CID   53613716.
  3. 1 2 3 4 5 Pates, Stephen; Daley, Allison C.; Legg, David A.; Rahman, Imran A. (2021-06-30). "Vertically migrating Isoxys and the early Cambrian biological pump". Proceedings of the Royal Society B: Biological Sciences. 288 (1953): 20210464. doi:10.1098/rspb.2021.0464. PMC   8220267 . PMID   34157876.
  4. ↑ García-Bellido, Diego C.; Paterson, John R.; Edgecombe, Gregory D.; Jago, James B.; Gehling, James G.; Lee, Michael S. Y. (November 2009). "The bivalved arthropods Isoxys and Tuzoia with soft-part preservation from the Lower Cambrian Emu Bay Shale Lagerstätte (Kangaroo Island, Australia): Isoxys and Tuzoia From Emu Bay Shale (Australia)". Palaeontology. 52 (6): 1221–1241. doi: 10.1111/j.1475-4983.2009.00914.x . S2CID   129374179.
  5. 1 2 3 4 5 Zhang, Caixia; Liu, Yu; Ortega-Hernández, Javier; Wolfe, Joanna; Jin, Changfei; Mai, Huijuan; Hou, Xian-guang; Guo, Jin; Zhai, Dayou (19 April 2023). "Three-dimensional morphology of the biramous appendages in Isoxys from the early Cambrian of South China, and its implications for early euarthropod evolution". Proceedings of the Royal Society B: Biological Sciences. 290 (1997). doi:10.1098/rspb.2023.0335. PMC   10113025 . PMID   37072042.
  6. ↑ Stein, Martin; Peel, John S.; Siveter, David J.; Williams, Mark (2009-09-07). "Isoxys (Arthropoda) with preserved soft anatomy from the Sirius Passet Lagerstätte, lower Cambrian of North Greenland: Isoxys with soft anatomy from Greenland". Lethaia. 43 (2): 258–265. doi:10.1111/j.1502-3931.2009.00189.x.
  7. ↑ Schoenemann, B.; Clarkson, E. N. K. (2010). "Eyes and vision in the Chengjiang arthropod Isoxys indicating adaptation to habitat". Lethaia. 44 (2): no. doi:10.1111/j.1502-3931.2010.00239.x.
  8. ↑ Ma, Jiaxin; Pates, Stephen; Wu, Yu; Lin, Weiliang; Liu, Cong; Wu, Yuheng; Zhang, Mingjing; Fu, Dongjing (2023-05-25). "Ontogeny and brooding strategy of the early Cambrian arthropod Isoxys minor from the Qingjiang biota". Frontiers in Ecology and Evolution. 11. doi: 10.3389/fevo.2023.1174564 . ISSN   2296-701X.
  9. ↑ Ma, Jiaxin; Lin, Weiliang; Liu, Cong; Sun, Ao; Wu, Yu; Wu, Yuheng; Fu, Dongjing (January 2022). "A new bivalved arthropod from the Cambrian (Stage 3) Qingjiang biota expands the palaeogeographical distribution and increases the diversity of Tuzoiidae". Journal of the Geological Society. 179 (1): jgs2020–229. Bibcode:2022JGSoc.179..229M. doi:10.1144/jgs2020-229. ISSN   0016-7649. S2CID   236289449.
  10. ↑ Izquierdo-López, Alejandro; Caron, Jean-Bernard (December 2022). "The problematic Cambrian arthropod Tuzoia and the origin of mandibulates revisited". Royal Society Open Science. 9 (12): 220933. Bibcode:2022RSOS....920933I. doi:10.1098/rsos.220933. ISSN   2054-5703. PMC   9727825 . PMID   36483757.
  11. 1 2 Liu, S.; Peng, J.; Wen, R.; Liang, B. (2018-06-30). "New data for Isoxys of the Balang Fauna (Cambrian Stage 4), South China". Bulletin of Geosciences: 147–162. doi: 10.3140/bull.geosci.1673 . ISSN   1802-8225. S2CID   134294843.
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