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Arthropoda
Fossil range: CambrianRecent
Marrella
Marrella, one of the puzzling arthropods from the Burgess Shale.
Scientific classification

Superphylum

Ecdysozoa

Phylum

Arthropoda
Latreille, 1829

Subphyla and Classes




An arthropod is an invertebrate that has an exoskeleton (external skeleton), a segmented body, and jointed attachments called appendages.

Evolution Edit

Last common ancestorEdit

The last common ancestor of all arthropods is reconstructed as a modular organism with each module covered by its own sclerite (armor plate) and bearing a pair of biramous limbs.[1] Whether the ancestral limb was uniramous or biramous is far from a settled debate, though. This Ur-arthropod had a ventral mouth, pre-oral antennae and dorsal eyes at the front of the body. It was a non-discriminatory sediment feeder, processing whatever sediment came its way for food.[1]

Fossil recordEdit

It has been proposed that the Ediacaran animals Parvancorina and Spriggina, from 550 to 543 million years ago, were arthropods.[2][3][4] Small arthropods with bivalve-like shells have been found in Early Cambrian fossil beds in China.[5] The earliest Cambrian trilobite fossils are about 530 million years old, but the class was already quite diverse and worldwide, suggesting that they had been around for quite some time.[6] Re-examination in the 1970s of the Burgess Shale fossils from about 505 million years ago identified many arthropods, some of which could not be assigned to any of the well-known groups, and thus intensified the debate about the Cambrian explosion.[7][8][9] A fossil of Marrella from the Burgess Shale has provided the earliest clear evidence of molting.[10]

The earliest fossil crustaceans date from about 513 million years ago in the Cambrian,[11] and fossil shrimp from about 500 million years ago apparently formed a tight-knit procession across the seabed.[12] Crustacean fossils are common from the Ordovician period onwards.[13] They have remained almost entirely aquatic, possibly because they never developed excretory systems that conserve water.

Arthropods provide the earliest identifiable fossils of land animals, from about 419 million years ago in the Late Silurian, and terrestrial tracks from about 450 million years ago appear to have been made by arthropods.[14] Arthropods were well pre-adapted to colonize land, because their existing jointed exoskeletons provided protection against desiccation, support against gravity and a means of locomotion that was not dependent on water.[15] Around the same time the aquatic, scorpion-like eurypterids became the largest ever arthropods, some as long as 2.5 meters (8.2 ft).[16]

The oldest known arachnid is the trigonotarbid Palaeotarbus jerami, from about 420 million years ago in the Silurian period.[17]Attercopus fimbriunguis, from 386 million years ago in the Devonian period, bears the earliest known silk-producing spigots, but its lack of spinnerets means it was not one of the true spiders,[18] which first appear in the Late Carboniferous over 299 million years ago.[19] The Jurassic and Cretaceous periods provide a large number of fossil spiders, including representatives of many modern families.[20] Fossils of aquatic scorpions with gills appear in the Silurian and Devonian periods, and the earliest fossil of an air-breathing scorpion with book lungs dates from the Early Carboniferous period.[21]

The oldest definitive insect fossil is the Devonian Rhyniognatha hirsti, dated at 396 to 407 million years ago, but its mandibles are of a type found only in winged insects, which suggests that the earliest insects appeared in the Silurian period.[22] The Mazon Creek lagerstätten from the Late Carboniferous, about 300 million years ago, include about 200 species, some gigantic by modern standards, and indicate that insects had occupied their main modern ecological niches as herbivores, detritivores and insectivores. Social termites and ants first appear in the Early Cretaceous, and advanced social bees have been found in Late Cretaceous rocks but did not become abundant until the Mid Cenozoic.[23]

Evolutionary family treeEdit



onychophorans,
including Aysheaia and Peripatus





armored lobopods,
including Hallucigenia and Microdictyon




anomalocarid-like taxa,
including modern tardigrades as
well as extinct animals like
Kerygmachela and Opabinia




Anomalocaris



arthropods,
including living groups and
extinct forms such as trilobites







Simplified summary of Budd's "broad-scale" cladogram (1996)[24]


From the late 1950s to the late 1970s, Sidnie Manton and others argued that arthropods are polyphyletic, in other words, they do not share a common ancestor that was itself an arthropod. Instead, they proposed that three separate groups of "arthropods" evolved separately from common worm-like ancestors: the chelicerates, including spiders and scorpions; the crustaceans; and the uniramia, consisting of onychophorans, myriapods and hexapods. These arguments usually bypassed trilobites, as the evolutionary relationships of this class were unclear. Proponents of polyphyly argued the following: that the similarities between these groups are the results of convergent evolution, as natural consequences of having rigid, segmented exoskeletons; that the three groups use different chemical means of hardening the cuticle; that there were significant differences in the construction of their compound eyes; that it is hard to see how such different configurations of segments and appendages in the head could have evolved from the same ancestor; and that crustaceans have biramous limbs with separate gill and leg branches, while the other two groups have uniramous limbs in which the single branch serves as a leg.[25]

Further analysis and discoveries in the 1990s reversed this view, and led to acceptance that arthropods are monophyletic, in other words they do share a common ancestor that was itself an arthropod.[26][27] For example Graham Budd's analyses of Kerygmachela in 1993 and of Opabinia in 1996 convinced him that these animals were similar to onychophorans and to various Early Cambrian "lobopods", and he presented an "evolutionary family tree" that showed these as "aunts" and "cousins" of all arthropods.[24][28] These changes made the scope of the term "arthropod" unclear, and Claus Nielsen proposed that the wider group should be labelled "Panarthropoda" ("all the arthropods") while the animals with jointed limbs and hardened cuticles should be called "Euarthropoda" ("true arthropods").[29]

A contrary view was presented in 2003, when Jan Bergström and Xian-Guang Hou argued that, if arthropods were a "sister-group" to any of the anomalocarids, they must have lost and then re-evolved features that were well-developed in the anomalocarids. The earliest known arthropods ate mud in order to extract food particles from it, and possessed variable numbers of segments with unspecialized appendages that functioned as both gills and legs. Anomalocarids were, by the standards of the time, huge and sophisticated predators with specialized mouths and grasping appendages, fixed numbers of segments some of which were specialized, tail fins, and gills that were very different from those of arthropods. This reasoning implies that Parapeytoia, which has legs and a backward-pointing mouth like that of the earliest arthropods, is a more credible closest relative of arthropods than is Anomalocaris.[32] In 2006, they suggested that arthropods were more closely related to lobopods and tardigrades than to anomalocarids.[33]

Higher up the "family tree", the Annelida have traditionally been considered the closest relatives of the Panarthropoda, since both groups have segmented bodies, and the combination of these groups was labelled Articulata. There had been competing proposals that arthropods were closely related to other groups such as nematodes, priapulids and tardigrades, but these remained minority views because it was difficult to specify in detail the relationships between these groups.

In the 1990s, molecular phylogenetics analyses that compared sequences of RNA and DNA produced a coherent scheme showing arthropods as members of a superphylum labelled Ecdysozoa ("animals that molt"), which contained nematodes, priapulids and tardigrades but excluded annelids. This was backed up by studies of the anatomy and development of these animals, which showed that many of the features that supported the Articulata hypothesis showed significant differences between annelids and the earliest Panarthropods in their details, and some were hardly present all in arthropods. This hypothesis groups annelids with molluscs and brachiopods in another superphylum, Lophotrochozoa.

If the Ecdysozoa hypothesis is correct, then segmentation of arthropods and annelids either has evolved convergently or has been inherited from a much older ancestor, and has been subsequently lost in several other lineages, such as the non-arthropod members of the Ecdysozoa.[34][30]

ReferencesEdit

  1. ^ a b Bergström; Hou (2005), "Early Palaeozoic non-lamellipedian arthropods", Crustacea and Arthropod Relationships, Boca Raton: Taylor & Francis, ISBN 0-8493-3498-5 
  2. ^ Glaessner, M. F. (1958), "New Fossils from the Base of the Cambrian in South Australia" (PDF), Transactions of the Royal Society of South Australia 81: 185–188, Archived from the original on 2007-09-29, http://web.archive.org/web/20070929094012/http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_V081/TRSSA_V081_p185p188.pdf, retrieved on 2008-10-21 
  3. ^ doi:10.1080/08912960500508689
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  4. ^ McMenamin, M.A.S (2003), "Spriggina is a trilobitoid ecdysozoan" (abstract), Abstracts with Programs (Geological Society of America) 35 (6): 105, http://gsa.confex.com/gsa/2003AM/finalprogram/abstract_62056.htm 
  5. ^ Yuan, X.; Xiao, S., Parsley, R.L., Zhou, C., Chen, Z. and Hu, J. (April 2002). "Towering sponges in an Early Cambrian Lagerstätte: Disparity between nonbilaterian and bilaterian epifaunal tierers at the Neoproterozoic-Cambrian transition". Geology 30 (4): 363–366. doi:10.1130/0091-7613(2002)030<0363:TSIAEC>2.0.CO;2. 
  6. ^ Lieberman, BS (01 Mar 1999), "Testing the Darwinian Legacy of the Cambrian Radiation Using Trilobite Phylogeny and Biogeography" (abstract), Journal of Paleontology 73 (2): 176, http://jpaleontol.geoscienceworld.org/cgi/content/abstract/73/2/176 
  7. ^ Whittington, H. B. (1979). Early arthropods, their appendages and relationships. In M. R. House (Ed.), The origin of major invertebrate groups (pp. 253–268). The Systematics Association Special Volume, 12. London: Academic Press.
  8. ^ Whittington, H.B.; Geological Survey of Canada (1985), The Burgess Shale, Yale University Press, ISBN 0660119013 
  9. ^ Gould, S.J. (1989), Wonderful Life: The Burgess Shale and the Nature of History, W.W. Norton & Company, ISBN 0393027058 
  10. ^ García-Bellido, D.C.; Collins, D.H. (May 2004). "Moulting arthropod caught in the act". Nature 429: 40. doi:10.1038/429040a. 
  11. ^ Budd, G.E., Butterfield, N.J., and Jensen, S. (December 2001), "Crustaceans and the “Cambrian Explosion″", Science 294 (5549): 2047, doi:10.1126/science.294.5549.2047a 
  12. ^ Callaway, E. (9 October 2008), Fossilised shrimp show earliest group behaviour, New Scientist, http://www.newscientist.com/channel/life/dn14903-fossilised-shrimp-show-earliest-group-behaviour.html?feedId=online-news_rss20, retrieved on 2008-10-21 
  13. ^ Zhang, X-G., Siveter, D.J., Waloszek, D., and Maas, A. (October 2007), "An epipodite-bearing crown-group crustacean from the Lower Cambrian", Nature 449 (7162): 595–598, doi:10.1038/nature06138 
  14. ^ Pisani, D., Laura L Poling, L.L., Lyons-Weiler M., and Hedges, S.B. (2004), "The colonization of land by animals: molecular phylogeny and divergence times among arthropods", BMC Biology 2: 1, doi:10.1186/1741-7007-2-1 
  15. ^ Cowen, R. (2000). History of Life (3rd ed.). Blackwell Science. p. p. 126. ISBN 0632044446. 
  16. ^ Braddy, S.J., Markus Poschmann, M., and Tetlie, O.E. (2008), "Giant claw reveals the largest ever arthropod", Biology Letters 4: 106–109, doi:10.1098/rsbl.2007.0491 
  17. ^ Dunlop, J.A. (September 1996), "A trigonotarbid arachnid from the Upper Silurian of Shropshire" (PDF), Palaeontology 39 (3): 605–614, http://palaeontology.palass-pubs.org/pdf/Vol%2039/Pages%20605-614.pdf, retrieved on 2008-10-12  The fossil was originally named Eotarbus but was renamed when it was realized that a Carboniferous arachnid had already been named Eotarbus: Dunlop, J.A., "A replacement name for the trigonotarbid arachnid Eotarbus Dunlop", Palaeontology 42 (1): 191 
  18. ^ Selden, P.A., and Shear, W.A. (July 2008), "Fossil evidence for the origin of spider spinnerets" (PDF), Nature Precedings, http://precedings.nature.com/documents/2088/version/1/files/npre20082088-1.pdf, retrieved on 2008-10-12 
  19. ^ Selden, P.A. (February 1996), "Fossil mesothele spiders", Nature 379: 498–499, doi:10.1038/379498b0 
  20. ^ Vollrath, F., and Selden, P.A. (December 2007), "The Role of Behavior in the Evolution of Spiders, Silks, and Webs" (PDF), Annual Review of Ecology, Evolution, and Systematics 38: 819–846, doi:10.1146/annurev.ecolsys.37.091305.110221, Archived from the original on 2008-12-09, http://web.archive.org/20081209102852/homepage.mac.com/paulselden/Sites/Website/ARES.pdf, retrieved on 2008-10-12 
  21. ^ Jeram, A.J. (January 1990), "Book-lungs in a Lower Carboniferous scorpion", Nature 343: 360–361, doi:10.1038/343360a0 
  22. ^ Engel, M.S., and Grimaldi, D.A. (February 2004), "New light shed on the oldest insect", Nature 427: 627–630, doi:10.1038/nature02291 
  23. ^ Labandeira, C., and Eble, G.J. (2000), "The Fossil Record of Insect Diversity and Disparity", in Anderson, J., Thackeray, F., van Wyk, B., and de Wit, M. (PDF), Gondwana Alive: Biodiversity and the Evolving Biosphere, Witwatersrand University Press, http://www.santafe.edu/research/publications/workingpapers/00-08-044.pdf, retrieved on 2008-09-07 
  24. ^ a b Budd, G.E. (1996), "The morphology of Opabinia regalis and the reconstruction of the arthropod stem-group", Lethaia 29 (1): 1–14, doi:10.1111/j.1502-3931.1996.tb01831.x 
  25. ^ Gillott, C. (1995), Entomology, Springer, pp. 17–19, ISBN 0306449676 
  26. ^ Adrain, J. (15 March 1999), Book Review: Arthropod Fossils and Phylogeny, edited by Gregory D. Edgecomb, Palaeontologia Electronica, http://palaeo-electronica.org/1999_1/books/arthropo.htm, retrieved on 2008-09-28  The book is G.D., ed. (1998), Arthropod Fossils and Phylogeny, Columbia University Press, pp. 347 
  27. ^ Chen, J.-y. (1995), "Head segmentation in Early Cambrian Fuxianhuia: implications for arthropod evolution", Science 268 (5215): 1339–1343, doi:10.1126/science.268.5215.1339, http://www.sciencemag.org/cgi/content/abstract/268/5215/1339, retrieved on 2008-09-28 
  28. ^ Budd, G.E. (1993), "A Cambrian gilled lobopod from Greenland", Nature 364: 709–711, doi:10.1038/364709a0 
  29. ^ Nielsen, C. (2001), Animal Evolution: Interrelationships of the Living Phyla (2nd ed.), Oxford University Press, pp. 194–196, ISBN 978-0-19-850681-2, http://books.google.co.uk/books?id=UmCg6c0HkqMC&pg=PA194&lpg=PA194&dq=nielsen+panarthropoda+euarthropoda&source=web&ots=NGNQl10IJk&sig=g3xEdLrpt6WVOsuxeuFt1HNOw-0&hl=en&sa=X&oi=book_result&resnum=7&ct=result#PPA19,M1, retrieved on 2008-09-28 
  30. ^ a b Telford, M.J., Bourlat, S.J., Economou, A., Papillon, D., and Rota-Stabelli, O. (January 2008), "The evolution of the Ecdysozoa", Philospohical Transactions of the Royal Society: Biology 363: 1529–1537, doi:10.1098/rstb.2007.2243, http://journals.royalsociety.org/content/rh11255r948k040r/, retrieved on 2008-09-29 
  31. ^ Vaccari, N.E., Edgecombe, G.D., and Escudero, C. (2004), "Cambrian origins and affinities of an enigmatic fossil group of arthropods", Nature 430: 554–557, doi:10.1038/nature02705 
  32. ^ Bergström, J., and Hou, X-G. (2003), "Arthropod origins" (PDF), Bulletin of Geosciences 78 (4): 323–334, http://www.geology.cz/bulletin/contents/2003/vol79no4/323_bergstrom.pdf, retrieved on 2008-10-22 
  33. ^ Hou, X-G., Bergström, J., and Jie, Y. (2006), "Distinguishing anomalocaridids from arthropods and priapulids", Geological Journal 41 (3–4): 259–269, doi:10.1002/gj.1050 
  34. ^ Schmidt-Rhaesa, A., Bartolomaeus, T., Lemburg, C., Ehlers, U., and Garey, J.R. (January 1999), "The position of the Arthropoda in the phylogenetic system", Journal of Morphology 238 (3): 263–285, doi:0.1002/(SICI)1097-4687(199812)238:3<263::AID-JMOR1>3.0.CO;2-L, http://www3.interscience.wiley.com/journal/5005954/abstract, retrieved on 2008-09-29 


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