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舌形動物類：修订间差异

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{| border="1" cellspacing="0" align="right" cellpadding="2"
{{Taxobox
! align="center" bgcolor=pink|'''扁形动物门'''
| name = 舌形動物<br />Pentastomida
|-
| fossil_range = {{fossil_range|寒武紀第三期|現代}}
! align="center" bgcolor=pink|'''[[科学分类]]'''
| image = Linguatula.jpg
|-
| image_width = 220px
|
| image_caption = Adult female ''[[Linguatula serrata]]''
{| align="center"
| regnum = [[動物界]] Animalia
|[[域 (生物)|域]]：||[[真核生物|真核生物域]]
| phylum = [[節肢動物門]] Arthropoda
|-
| subphylum = [[甲殼亞門]] Crustacea
|[[界 (生物)|界]]：||[[动物|动物界]]
| classis = [[顎足綱]] Maxillopoda
|-
| subclassis = '''舌形動物亞綱 Pentastomida'''
|[[门 (生物)|门]]：||'''Pentastoma'''
| subclassis_authority = [[:en:K. M. Diesing|Diesing]], 1836
|}
| subdivision_ranks = [[目 (生物)|目]]
|-
| subdivision =
! align="center" bgcolor=pink|'''[[纲]]'''
*[[Cephalobaenida]]
|-
*[[Porocephalida]]
|[[舌形动物纲]]<br>
*[[Raillietiellida]]
|}
*[[Reighardiida]]
}}


'''舌形动物门(Pentastoma)'''是动物界的一个门，又称'''五口动物门'''，'''舌形动物门'''一类动物介于[[环节动物]]和[[节肢动物]]之间的[[寄生]]类动物，体软，扁而长，无色，透明，无足。身長從數毫米到十數厘米，体表分近百个清晰的节段，内部却并不分节。虫体前端口部突出，呈椭圆形，周围有钩2对,可伸缩，用以附着在寄主组织上。该门动物没有[[呼吸系统]]、[[排泄系统]]和[[循环系统]]，雌雄异体。
'''舌形动物類'''（[[學名]]：{{lang|la|'''Pentastomida'''}}），又称'''五口动物類'''，是[[动物界]]的一个[[分類學|分類]]。這一種謎樣的動物屬於[[無脊椎動物]]，介乎於[[环节动物]]和[[节肢动物]]之间，均為[[寄生動物|寄生性]]動物。其特徵是体软、扁而长，无色，透明，无足。身長從數毫米到十數厘米，体表分近百个清晰的节段，内部却并不分节。虫体前端口部突出，呈椭圆形，周围有钩2对,可伸缩，用以附着在寄主组织上。该门动物没有[[呼吸系统]]、[[排泄系统]]和[[循环系统]]，雌雄异体。


本物種的名稱是因為從其部分物種的外型酷似脊椎動物的舌頭，因而為名；而其學名及另一名稱「五口動物」的來源，則是因為其口器的五個鈎，好像有五個口一樣，因而為名。這五個「鈎」，其實只有一個為口器，其餘四個只是顎足。過往由於對本分類物種的分類不明，單單從外形來判斷，所以將其歸類到門一層。後來到了2001年，本物種生物跟其他物種的關係得到確認，所以被降至歸入成為[[顎足綱]]的一員，是為'''舌形動物亞綱'''。

現時本分類有物種約130個，皆為被動式寄生物，其部分解剖結果為退化性。成年的舌蟲長約{{convert|1|to|14|cm|1}} ，多數於宿主的呼吸系統黏付着。

==生物==
Historically significant accounts of tongue worm biology and systematics include early work by [[Josef Aloys Frölich]],<ref>{{cite journal |author=J. A. Frölich |year=1789 |title= Beschreibung einiger neuer Eingeweidewürmer |journal=[[Der Naturforscher]] |volume=24 |pages=101–162}}</ref> [[Alexander von Humboldt]],<ref>{{cite book |author=A. von Humboldt |year=1811 |chapter=Sur un ver intestin trouvé dans les poumons du serpent à sonnettes, de Cumana |title=Voyage de Humboldt et Bonpand 2 |publisher=Ptie. F. Schoell et G. Dufour, Paris |pages=298–304}}</ref> [[Karl Asmund Rudolphi]],<ref>{{cite book |author=K. A. Rudolphi |year=1819 |title= Entozoorum Synopsis |publisher=Augustus Rücker Berlin}}</ref> [[Karl Moriz Diesing]]<ref>{{cite journal |author=K. M. Diesing |year=1835 |title= Versuch einer Monographie der Gattung ''Pentastoma'' |journal=[[Annalen des Wiener Museums der Naturgeschichte]] |volume=1 |pages=1–32}}</ref> and [[Rudolph Leuckart]].<ref>{{cite journal |author=R. Leuckart |year=1860 |title= Bau und Entwicklungsgeschichte der Pentastomen nach Untersuchungen besonders von ''Pent. taenoides'' und ''P. denticulatum'' |publisher=C. F. Winter'sche Verlagshandlung, Leipzig |pages=vi + 160}}</ref>

Other important summaries have been published by Louis Westenra Sambon,<ref>{{cite journal |author=L. W. Sambon |year=1922 |title= A synopsis of the family Linguatulidae |journal=[[Journal of Tropical Medicine and Hygiene]] |volume=12 |pages=188–206, 391–428}}</ref> [[Richard Heymons]]<ref name="Heymons">{{cite book |author=R. Heymons |year=1935 |chapter= Pentastomida |editor=H. G. Bronns |title=Klassen und Ordnungen des Tierreichs. Fünfter Band. IV Abteilung, 1. Buch |publisher=Akademische Verlagsgesellschaft m.b.H., Leipzig |pages=1–268 pp}}</ref> and John Riley,<ref>{{cite journal |author=J. Riley |year=1986 |title= The biology of pentastomids |journal=[[Advances in Parasitology]] |volume=25 |pages=45–128 |doi=10.1016/S0065-308X(08)60342-5}}</ref> and a review of their evolutionary relationships with a bibliography up to 1969 was published by J. T. Self.<ref name="Self">{{cite journal |author=J. T. Self |year=1969 |title= Biological relationships of the Pentastomida: a bibliography on the Pentastomida |journal=[[Experimental Parasitology]] |volume=21 |pages=63–119 |doi=10.1016/0014-4894(69)90222-7}}</ref>

===Anatomy===
Pentastomids are worm-like animals ranging from {{convert|2|to|13|cm|in}} in length. The anterior end of the body bears five protuberances, four of which are clawed legs, while the fifth bears the mouth. The body is segmented and covered in a [[chitin]]ous cuticle. The digestive tract is simple and tubular, since the animal feeds entirely on blood, although the mouth is somewhat modified as a muscular pump.<ref name=IZ>{{cite book |author= Barnes, Robert D. |year=1982 |title= Invertebrate Zoology |publisher= Holt-Saunders International |location= Philadelphia, PA|pages= 880–881|isbn= 0-03-056747-5}}</ref>

The nervous system is similar to that of other arthropods, including a ventral nerve cord with [[ganglion|ganglia]] in each segment. Although the body contains a [[haemocoel]], there are no circulatory, respiratory, or excretory organs.<ref name=IZ/>

===Life cycle===
Pentastomids live in the upper respiratory tract of reptiles, birds, and mammals, where they lay eggs. They are [[gonochorism|gonochoric]] (having 2 sexes), and employ [[internal fertilisation]]. The eggs are either coughed out by the host or leave the host body through the digestive system. The eggs are then ingested by an intermediate host, which is commonly either a fish or a small herbivorous mammal.<ref name=IZ/>

The larva hatches in the intermediate host and breaks through the wall of the intestine. It then forms a cyst in the intermediate host's body. The larva is initially rounded in form, with four or six short legs, but [[ecdysis|moults]] several times to achieve the adult form. The pentastomid reaches the main host when the intermediate host is eaten by the main host, and crawls into the respiratory tract from the [[oesophagus]].<ref name=IZ/>

==Human infestation==
[[File:Ocular-Pentastomiasis-in-the-Democratic-Republic-of-the-Congo-pntd.0003041.s001.ogv|thumb|Extraction of an ''[[Armillifer grandis]]'' nymph from a human eye.]]
Tongue worms occasionally parasitise humans.<ref>{{cite journal |author=A. Fain |year=1975 |title= The Pentastomida parasitic in man |journal=[[Annales de la Société belge de médecine tropicale]] |volume=55 |pages=59–64}}</ref> While there is a report of ''[[Sebakia]]'' inducing dermatitis,<ref>{{cite journal |author=H. Solano Mairena & W. Venegas |year=1989 |title= Human dermatitis caused by a nymph of ''Sebekia'' |journal=[[American Journal of Tropical Medicine and Hygiene]] |volume=41 |pages=352–354 |url=http://www.ajtmh.org/cgi/content/abstract/41/3/352 |pmid=2802021 |issue=3}}</ref> the two genera responsible for most internal human infestation are ''[[Linguatula]]'' and ''[[Armillifer]]''. Visceral pentastomiasis can be caused by ''[[Linguatula serrata]]'', ''[[Armillifer armillatus]]'', ''[[Armillifer moniliformis]]'', ''[[Armillifer grandis]]'', and ''[[Porocephalus crotali]]''.<ref name="pmid19238218">{{cite journal |author=Dennis Tappe & Dietrich W. Büttner |editor1-last=Bethony |editor1-first=Jeffrey M. |title=Diagnosis of human visceral pentastomiasis |journal=[[PLoS Neglected Tropical Diseases]] |volume=3 |issue=2 |pages=e320 |year=2009 |pmid=19238218 |pmc=2643528 |doi=10.1371/journal.pntd.0000320 |url=http://dx.plos.org/10.1371/journal.pntd.0000320}}</ref>

The terms associated with infections can vary:
* ''[[Linguatula]]'' disease can be called [[linguatuliasis]] or [[linguatulosis]].
* ''[[Porocephalus]]'' disease can be called [[porocephaliasis]] or [[porocephalosis]].
* ''[[Armillifer]]'' disease can also be called [[porocephalosis]].<ref name="pmid933188">{{cite journal |author=Esmond M. Mapp, Howard M. Pollack & Louis H. Goldman |title=Roentgen diagnosis of ''Armillifer armillatus'' infestation (porocephalosis) in man |journal=[[Journal of the National Medical Association]] |volume=68 |issue=3 |pages=198–200, 191 |date=May 1976 |pmid=933188 |pmc=2609651 |doi= |url=}}</ref> (An alternate name for ''Armillifer moniliformis'' is ''Porocephalus moniliformis''.)<ref name="PalmerReeder2001">{{cite book|author1=Philip E. S. Palmer|author2=Maurice Merrick Reeder|title=Imaging of tropical diseases: with epidemiological, pathological, and clinical correlation|url=http://books.google.com/?id=b3_RMZl464cC&pg=PA389|accessdate=19 April 2010|year=2001|publisher=Birkhäuser|isbn=978-3-540-62471-4|pages=389–}}</ref>
* "Pentastomiasis" can refer to any infection of Pentastomida.

''Porocephalus'' and ''Armillifer'' (which are all cylindrical and all inhabit snakes) have much more in common with each other than they do with ''Linguatula'' (which is flat and inhabits dogs and wolves.)

==Affinities==
The affinities of tongue worms have long proved controversial. Historically, they were initially compared to various groups of parasitic worms. Once the arthropod-like nature of their cuticle was recognised, similarities were drawn with mites,<ref>{{cite journal |author=T. D. Schubart |year=1853 |title= Ueber die Entwicklung des ''Pentastoma taenioides'' |journal=[[Zeitschrift für Wissenschaftliche Zoologie]] |volume=4 |pages=117–118}}</ref> particularly gall mites ([[Eriophyidae]]). Although gall mites are much smaller than tongue worms they also have a long, segmented body and only two pairs of legs. Later work drew comparisons with millipedes and centipedes ([[Myriapoda]]), with velvet worms ([[Onychophora]]) and water bears ([[Tardigrada]]). Some authors interpreted tongue worms as essentially intermediate between [[annelids]] and [[arthropods]], while others suggested that they deserved a [[phylum]] of their own. Tongue worms grow by [[ecdysis|moulting]], which suggests they belong to [[Ecdysozoa]], while other work has identified the arthropod-like nature of their larvae,<ref>{{cite journal |author=G. Osche |year=1959 |title= "Arthropodencharaktere" bei einem Pentastomiden Embryo (''Reighhardia sernae'') |journal=[[Zoologischer Anzeiger]] |volume=163 |pages=169–178}}</ref> In general, there are two current alternative interpretations: pentastomids are highly modified and parasitic crustaceans, probably related to fish lice, or they are an ancient group of stem-arthropods, close to the origins of Arthropoda.

===Crustaceans===
The possibility that tongue worms are [[crustacean]]s can be traced back to the work of [[Pierre-Joseph Van Beneden]],<ref>{{cite journal |author=P. J. van Beneden |year=1849 |title= Recherches sur l’organisation et le développement des Lingatules (Pentastoma Rud.), suivies de la description d’une espèce nouvelle provenant d’un Mandrill |journal=[[Annales des Sciences Naturelles]] Zoologie Series |volume=3 |issue=11 |pages=313–348}}</ref> who compared them to parasitic [[copepod]]s. The modern form of this hypothesis dates from Karl Georg Wingstrand's study of [[sperm]] morphology,<ref>{{cite journal |author=K. G. Wingstrand |year=1972 |title= Comparative spermatology of a pentastomid, ''Raillietiella hemidactyli'', and a branchiuran crustacean, ''Argulus foliaceus'', with a discussion of pentastomid relationships |journal=[[Det Kongelige Danske Videnskabernes Selskab Biologiske Skrifter]] |volume=19 |issue=4 |pages=1–72}}</ref> which recognised similarities in sperm structure between tongue worms and fish lice ([[Argulidae]]) – a group of [[maxillipod]] crustaceans which live as parasites on fish and occasionally amphibians. John Riley and colleagues also offered a detailed justification for the inclusion of the tongue worms among the crustaceans.<ref>{{cite journal |author=J. Riley, A. A. Banaja & J. L. James |year=1978 |title=The phylogenetic relationships of the Pentastomida: the case for their inclusion within the Crustacea |journal=[[International Journal of Parasitology]] |volume=8 |issue=4 |pages=245–254 |doi=10.1016/0020-7519(78)90087-5}}</ref> The fish louse model received significant further support from the molecular work of Lawrence G. Abele and colleagues.<ref>{{cite journal |author=L. G. Abele, W. Kim & B. E. Felgenhauer |year=1989 |title= Molecular evidence for inclusion of the Phylum Pentastomida in the Crustacea |journal=[[Molecular Biology and Evolution]] |volume=6 |pages=685–691 |url=http://mbe.oxfordjournals.org/cgi/reprint/6/6/685.pdf |issue=6}}</ref> A number of subsequent molecular phylogenies have corroborated these results,<ref>{{cite journal |author=D. V. Lavrov, W. M. Brown & J. L. Boore |year=2004 |title=Phylogenetic position of the Pentastomida and (pan)crustacean relationships |journal=[[Proceedings of the Royal Society of London B]] |volume=271 |pages=537–544 |url=http://www.eeob.iastate.edu/faculty/LavrovD/publications/PDF_files/pentastomida.pdf |doi=10.1098/rspb.2003.2631 |pmid=15129965 |issue=1538 |pmc=1691615}}</ref><ref>{{cite journal |author=O. S. Møller, J. Olesen, A. Avenant-Oldewage, P. F. Thomsen & H. Glenner |year=2008 |title=First maxillae suction discs in Branchiura (Crustacea): development and evolution in light of the first molecular phylogeny of Branchiura, Pentastomida, and other "Maxillopoda" |journal=[[Arthropod Structure & Development]] |volume=37 |pages=333–346 |doi=10.1016/j.asd.2007.12.002 |pmid=18394959 |issue=4}}</ref><ref name="Oakley2013">{{cite journal |author=Todd H. Oakley, Joanna M. Wolfe, Annie R. Lindgren & Alexander K. Zaharoff |year=2013 |title=Phylotranscriptomics to bring the understudied into the fold: monophyletic Ostracoda, fossil placement, and Pancrustacean phylogeny |journal=[[Molecular Biology and Evolution]] |volume=30 |issue=1 |pages=215–233 |pmid=22977117 |doi=10.1093/molbev/mss216}}</ref> and the name '''Ichthyostraca''' has been proposed for a (Pentastomida + Branchiura) clade.<ref>{{cite journal |author=J. Zrzavý |year=2001 |title= The interrelationships of metazoan parasites: a review of phylum- and higher-level hypotheses from recent morphological and molecular phylogenetic analyses |journal=[[Folia Parasitologica]] |volume=48 |pages=81–103 |url=http://faculty.uml.edu/rhochberg/hochberglab/Courses/Parasite/PDF%20Papers/Parasitology%20Papers/Interrelationships%20of%20Metazoan%20Parasites.pdf}}</ref> Thus a number of important standard works and databases on crustaceans now include the pentastomids as members of this group.<ref name="M&D">{{cite book |url=http://atiniui.nhm.org/pdfs/3839/3839.pdf |title=An Updated Classification of the Recent Crustacea |author=J. W. Martin & G. E. Davis |year=2001 |pages=132 pp |publisher=[[Natural History Museum of Los Angeles County]]}}</ref>

===Stem-arthropods===
Critics of the Ichthyostraca hypothesis have pointed out that even parasitic crustaceans can still be recognised as crustaceans based on their larvae; but that tongue worms and their larvae do not express typical characters for Crustacea or even [[Euarthropoda]]. An alternative model notes the extremely ancient Cambrian origins of these animals and interprets tongue worms as stem-group arthropods.<ref>{{cite journal |author=Dieter Waloszek, John E. Repetski & Andreas Maas |year=2006 |title= A new Late Cambrian pentastomid and a review of the relationships of this parasitic group |journal=[[Transactions of the Royal Society of Edinburgh]]: Earth Sciences |volume=96 |issue=2 |pages=163–176 |doi=10.1017/S0263593300001280}}</ref> A recent morphological analysis recovered Pentastomida outside the arthropods, as sister group to a clade including [[nematode]]s, [[priapulids]] and similar ecdyzoan 'worm' groups.<ref>{{cite journal |author=W. O. Almeida, M. L. Christoffersen, D. S. Amorim & E. C. C. Eloy |year=2008 |title=Morphological support for the phylogenetic positioning of Pentastomida and related fossils |journal=[[Revista Biotemas]] |volume=21 |pages=81–90 |url=http://www.biotemas.ufsc.br/volumes/pdf/volume213/p81a90.pdf |doi=10.5007/2175-7925.2008v21n3p81}}</ref> Adding fossils, they suggested an extinct animal called ''[[Facivermis]]'' could be closely related to tongue worms. However it should be stressed that these authors did not explicitly test pentastomid/crustacean relationships.

==Fossil record==
Exceptionally preserved, three-dimensional and [[Phosphatic fossilization|phosphatised fossils]] from the Upper [[Cambrian]] [[Orsten]] of [[Sweden]]<ref>{{cite journal |author=D. Walossek & K. J. Müller |year=1994 |title= Pentastomid parasites from the Lower Palaeozoic of Sweden |journal=[[Transactions of the Royal Society of Edinburgh]]: Earth Sciences |volume=85 |pages=1–37 |doi=10.1017/s0263593300006295}}</ref> and the Cambrian/[[Ordovician]] boundary of [[Canada]]<ref>{{cite journal |author=Dieter Walossek, John E. Repetski & Klaus J. Müller |year=1994 |title= An exceptionally preserved parasitic arthropod, ''Heymonsicambria taylori'' n. sp. (Arthropoda increate sedis: Pentastomida) from Cambrian – Ordovician boundary beds of Newfoundland |journal=[[Canadian Journal of Earth Sciences]] |volume=31 |issue=11 |pages=1664–1671 |doi=10.1139/e94-149}}</ref> have been identified as pentastomids. Four fossil genera have been identified so far: ''Aengapentastomum'', ''Bockelericambria'', ''Haffnericambria'' and ''Heymonsicambria''. These fossils suggest that pentastomids evolved very early and raise questions about whether these animals were parasites at this time, and if so, on which hosts. [[Conodont]]s have sometimes been mentioned as possible hosts in this context.

==Classification==
This article follows Martin and Davis<ref name="M&D"/> in placing Pentastomida in the class [[Maxillopoda]] within the subphylum [[crustacean|Crustacea]]. The subclass contains the following extant orders, superfamilies, families, genera and species:<ref>{{cite WoRMS |author=Geoff Boxshall |year=2013 |title=Pentastomida |id=22602 |accessdate=May 13, 2013}}</ref>
[[File:Armillifer armillatus.jpg|thumb|''Armillifer armillatus'' Wyman, 1848, a 4 cm individual collected from the respiratory system of a python, ''[[Python sebae]]''. Specimen deposited in the [[Humboldt Museum|Museum für Naturkunde Berlin]]]]
[[File:Armillifer.jpg|thumb|right|Female (right) and male (left) ''[[Armillifer]] sp.'']]
*'''[[Cephalobaenida]]''' <small>Heymons, 1935</small>
:*'''[[Cephalobaenidae]]''' <small>Heymons, 1922</small>
:::*''[[Cephalobaena]]'' <small>Heymons, 1922</small>
::::*''[[Cephalobaena tetrapoda]]'' <small>Heymons, 1922</small>
*'''[[Porocephalida]]''' <small></small>
:*'''[[Linguatuloidea]]''' <small></small>
::*'''[[Linguatulidae]]''' <small>Haldeman, 1851</small>
:::*''[[Linguatula]]'' <small>Frölich, 1789</small>
::::*''[[Linguatula arctica]]'' <small>Riley, Haugerud & Nilssen, 1987</small>
::::*''[[Linguatula multiannulata]]'' <small>Haffner & Rack ''in'' Haffner, Rack & Sachs, 1969</small>
::::*''[[Linguatula recurvata]]'' <small>(Diesing, 1850)</small>
::::*''[[Linguatula serrata]]'' <small>Frölich, 1789</small>
:::*''[[Neolinguatula]]'' <small>Haffner & Rack ''in'' Haffner, Rack & Sachs, 1969</small>
::::*''[[Neolinguatula nuttalli]]'' <small>(Sambon, 1922)</small>
::*'''[[Subtriquetridae]]''' <small>Fain, 1961</small>
:::*''[[Subtriquetra]]'' <small>Sambon, 1922</small>
::::*''[[Subtriquetra megacephalum]]'' <small>(Baird, 1853)</small>
::::*''[[Subtriquetra rileyi]]'' <small>Junker, Boomker & Booyse, 1998</small>
::::*''[[Subtriquetra shipleyi]]'' <small>Hett, 1924</small>
::::*''[[Subtriquetra subtriquetra]]'' <small>(Diesing, 1836)</small>
:*'''[[Porocephaloidea]]''' <small></small>
::*'''[[Porocephalidae]]''' <small>Sambon, 1922</small>
:::*''[[Armillifer]]'' <small>Sambon, 1922</small>
::::*''[[Armillifer aborealis]]'' <small>Riley & Self, 1981</small>
::::*''[[Armillifer agkistrodontis]]'' <small>Self & Kuntz, 1966</small>
::::*''[[Armillifer armillatus]]'' <small>(Wyman, 1845)</small>
::::*''[[Armillifer australis]]'' <small>Heymons, 1935</small>
::::*''[[Armillifer grandis]]'' <small>(Hett, 1915)</small>
::::*''[[Armillifer mazzai]]'' <small>(Sambon, 1922)</small>
::::*''[[Armillifer moniliformis]]'' <small>(Diesing, 1836)</small>
::::*''[[Armillifer yoshidai]]'' <small>Kishida, 1928</small>
:::*''[[Cubirea]]'' <small>Kishida, 1928</small>
::::*''[[Cubirea annulata]]'' <small>(Baird, 1853)</small>
::::*''[[Cubirea pomeroyi]]'' <small>(Woodland, 1921)</small>
:::*''[[Elenia]]'' <small>Heymons, 1932</small>
::::*''[[Elenia australis]]'' <small>Heymons, 1932</small>
:::*''[[Gigliolella]]'' <small>[[Alain Chabaud|Chabaud]] & Choquet, 1954</small>
::::*''[[Gigliolella brumpti]]'' <small>(Giglioli, 1922)</small>
:::*''[[Kiricephalus]]'' <small>Sambon, 1922</small>
::::*''[[Kiricephalus clelii]]'' <small>Riley & Self, 1980</small>
::::*''[[Kiricephalus coarctatus]]'' <small>(Diesing, 1850)</small>
::::*''[[Kiricephalus constrictor]]'' <small>Riley & Self, 1980</small>
::::*''[[Kiricephalus gabonensis]]'' <small>Riley & Self, 1980</small>
::::*''[[Kiricephalus pattoni]]'' <small>(Stephens, 1908)</small>
::::*''[[Kiricephalus tortus]]'' <small>(Shipley, 1898)</small>
:::*''[[Parasambonia]]'' <small>Stunkard & Gandal, 1968</small>
::::*''[[Parasambonia bridgesi]]'' <small>Stunkard & Gandal, 1968</small>
::::*''[[Parasambonia minor]]'' <small>Riley & Self, 1982</small>
:::*''[[Porocephalus]]'' <small>Humboldt, 1812</small>
::::*''[[Porocephalus basiliscus]]'' <small>Riley & Self, 1979</small>
::::*''[[Porocephalus benoiti]]'' <small>Fain, 1960</small>
::::*''[[Porocephalus bifurcatus]]''
::::*''[[Porocephalus clavatus]]'' <small>(Wyman, 1845)</small>
::::*''[[Porocephalus crotali]]'' <small>Humboldt, 1812</small>
::::*''[[Porocephalus dominicana]]'' <small>Riley & Walters, 1980</small>
::::*''[[Porocephalus stilesi]]'' <small>Sambon ''in'' Vaney & Sambon, 1910</small>
::::*''[[Porocephalus subuliferum]]'' <small>(Leuckart, 1860)</small>
::::*''[[Porocephalus taiwana]]'' <small>Qiu, Ma, Fan & Lu, 2005</small>
::::*''[[Porocephalus tortugensis]]'' <small>Riley & Self, 1979</small>
:::*''[[Waddycephalus]]'' <small>Sambon, 1922</small>
::::*''[[Waddycephalus calligaster]]'' <small>Riley & Self, 1981</small>
::::*''[[Waddycephalus komodoensis]]'' <small>Riley & Self, 1981</small>
::::*''[[Waddycephalus longicauda]]'' <small>Riley & Self, 1981</small>
::::*''[[Waddycephalus porphyriacus]]'' <small>Riley & Self, 1981</small>
::::*''[[Waddycephalus punctulatus]]'' <small>Riley & Self, 1981</small>
::::*''[[Waddycephalus radiata]]'' <small>Riley & Self, 1981</small>
::::*''[[Waddycephalus scutata]]'' <small>Riley & Self, 1981</small>
::::*''[[Waddycephalus superbus]]'' <small>Riley & Self, 1981</small>
::::*''[[Waddycephalus teretiusculus]]'' <small>(Baird, 1862)</small>
::::*''[[Waddycephalus vitiensis]]'' <small>Heymons, 1932</small>
::*'''[[Sebekidae]]''' <small>Sambon, 1922</small>
:::*''[[Alofia]]'' <small>Giglioli ''in'' Sambon, 1922</small>
::::*''[[Alofia ginae]]'' <small>Giglioli ''in'' Sambon, 1922</small>
::::*''[[Alofia indica]]'' <small>(von Linstow, 1906)</small>
::::*''[[Alofia merki]]'' <small>Giglioli 'in'' Sambon, 1922</small>
::::*''[[Alofia nilotici]]'' <small>Riley & Huchzermeyer, 1995</small>
::::*''[[Alofia parva]]'' <small>Riley & Huchzermeyer, 1995</small>
::::*''[[Alofia platycephalum]]'' <small>(Lohrmann, 1889)</small>
::::*''[[Alofia simpsoni]]'' <small>Riley, 1994</small>
::::*''[[Alofia travassosi]]'' <small>(Heymons, 1932)</small>
:::*''[[Diesingia]]'' <small>Sambon, 1922</small>
::::*''[[Diesingia kachugensis]]'' <small>(Shipley, 1910)</small>
::::*''[[Diesingia megastomum]]'' <small>(Diesing, 1836)</small>
:::*''[[Selfia]]'' <small>Riley, 1994</small>
::::*''[[Selfia porosus]]'' <small>Riley, 1994</small>
:::*''[[Leiperia]]'' <small>Sambon, 1922</small>
::::*''[[Leiperia australiensis]]'' <small>Riley & Huchzermeyer, 1996</small>
::::*''[[Leiperia cincinnalis]]'' <small>(Sambon ''in'' Vaney & Sambon, 1910)</small>
::::*''[[Leiperia gracilis]]'' <small>(Diesing, 1836)</small>
:::*''[[Sambonia]]'' <small>Noc & Giglioli, 1922</small>
::::*''[[Sambonia clavata]]'' <small>(Lohrmann, 1889)</small>
::::*''[[Sambonia parapodum]]'' <small>Self & Kuntz, 1966</small>
::::*''[[Sambonia solomenensis]]'' <small>(Self & Kuntz, 1957)</small>
::::*''[[Sambonia varani]]'' <small>(Self & Kuntz, 1957)</small>
::::*''[[Sambonia wardi]]'' <small>(Sambon ''in'' Vaney & Sambon, 1910)</small>
:::*''[[Agema (genus)|Agema]]'' <small>Riley ''et al.'', 1997</small>
::::*''[[Agema silvaepalustris]]'' <small>Riley ''et al.'', 1997</small>
:::*''[[Pelonia]]'' <small>Junker & Boomker, 2002</small>
::::*''[[Pelonia africana]]'' <small>Junker & Boomker, 2002</small>
:::*''[[Sebekia]]'' <small>Sambon, 1922</small>
::::*''[[Sebekia cesarisi]]'' <small>Giglioli ''in'' Sambon, 1922</small>
::::*''[[Sebekia divestei]]'' <small>Giglioli ''in'' Sambon, 1922</small>
::::*''[[Sebekia johnstoni]]'' <small>Riley, Spratt & Winch, 1990</small>
::::*''[[Sebekia microhamus]]'' <small>Self & Rego, 1985</small>
::::*''[[Sebekia minor]]'' <small>(Wedl, 1861)</small>
::::*''[[Sebekia mississippiensis]]'' <small>Overstreet, Self & Vliet, 1985</small>
::::*''[[Sebekia multiannulata]]'' <small>Riley, Spratt & Winch, 1990</small>
::::*''[[Sebekia novaeguineae]]'' <small>Riley, Spratt & Winch, 1990</small>
::::*''[[Sebekia okavangoensis]]'' <small>Riley & Huchzermeyer, 1995</small>
::::*''[[Sebekia oxycephalum]]'' <small>(Diesing, 1836)</small>
::::*''[[Sebekia purdieae]]'' <small>Riley, Spratt & Winch, 1990</small>
::::*''[[Sebekia trinitatis]]'' <small>Riley, Spratt & Winch, 1990</small>
*'''[[Raillietiellida]]''' <small>Almeida & Christoffersen, 1999</small>
::*'''[[Raillietiellidae]]''' <small>Sambon, 1922</small>
:::*''[[Raillietiella]]'' <small>Sambon, 1910</small>
::::*''[[Raillietiella aegypti]]'' <small>Ali, Riley & Self, 1982</small>
::::*''[[Raillietiella affinis]]'' <small>Bovien, 1927</small>
::::*''[[Raillietiella agcoi]]'' <small>Tubangui & Masiluñgan, 1936</small>
::::*''[[Raillietiella ampanihyensis]]'' <small>Gretillat, Brygoo & Domergue, 1962</small>
::::*''[[Raillietiella amphiboluri]]'' <small>Mahon, 1954</small>
::::*''[[Raillietiella belohaensis]]'' <small>McAllister, Riley, Freed & Freed, 1993</small>
::::*''[[Raillietiella bicaudata]]'' <small>Heymons & Vitzthum, 1935</small>
::::*''[[Raillietiella boulengeri]]'' <small>(Vaney & Sambon, 1910)</small>
::::*''[[Raillietiella bufonis]]'' <small>Ali, Riley & Self, 1982</small>
::::*''[[Raillietiella cartagenensis]]'' <small>Ali, Riley & Self, 1985</small>
::::*''[[Raillietiella chamaeleonis]]'' <small>Gretillat & Brygoo, 1959</small>
::::*''[[Raillietiella colubrilineati]]'' <small>(Leuckart, 1860)</small>
::::*''[[Raillietiella congolensis]]'' <small>Fain, 1961</small>
::::*''[[Raillietiella crotali]]'' <small>Ali, Riley & Self, 1984</small>
::::*''[[Raillietiella freitasi]]'' <small>(Motta & Gomes, 1968)</small>
::::*''[[Raillietiella furcocercum]]'' <small>(Diesing, 1836)</small>
::::*''[[Raillietiella gehyrae]]'' <small>Bovien, 1927</small>
::::*''[[Raillietiella gigliolii]]'' <small>Hett, 1924</small>
::::*''[[Raillietiella gowrii]]'' <small>Rajalu & Rajendran, 1970</small>
::::*''[[Raillietiella hebitihamata]]'' <small>Self & Kuntz, 1960</small>
::::*''[[Raillietiella hemidactyli]]'' <small>Hett, 1934</small>
::::*''[[Raillietiella indica]]'' <small>Gedoelst, 1921</small>
::::*''[[Raillietiella kochi]]'' <small>Heymons, 1926</small>
::::*''[[Raillietiella mabuiae]]'' <small>Heymons, 1922</small>
::::*''[[Raillietiella maculatus]]'' <small>Rao & Hiregaudar, 1962</small>
::::*''[[Raillietiella maculilabris]]'' <small>Ali, Riley & Self, 1984</small>
::::*''[[Raillietiella madagascariensis]]'' <small>McAllister ''et al.'', 1993</small>
::::*''[[Raillietiella mediterranea]]'' <small>(Hett, 1915)</small>
::::*''[[Raillietiella monarchus]]'' <small>Ali, Riley & Self, 1984</small>
::::*''[[Raillietiella morenoi]]'' <small>Abreu-Acosta ''et al.'', 2006</small>
::::*''[[Raillietiella mottae]]'' <small>Almeida, Freire & Lopes, 2008</small>
::::*''[[Raillietiella namibiensis]]'' <small>Riley & Heideman 1998</small>
::::*''[[Raillietiella orientalis]]'' <small>(Hett, 1915)</small>
::::*''[[Raillietiella piscator]]'' <small>Nair, 1967</small>
::::*''[[Raillietiella rileyi]]'' <small>Krishnasamy ''et al.'', 1995</small>
::::*''[[Raillietiella schoutedeni]]'' <small>Fain, 1960</small>
::::*''[[Raillietiella scincoides]]'' <small>Ali, Riley & Self, 1984</small>
::::*''[[Raillietiella spiralis]]'' <small>Hett, 1924</small>
::::*''[[Raillietiella teagueselfi]]'' <small>Riley, McAllister & Freed, 1988</small>
::::*''[[Raillietiella tetrapoda]]'' <small>(Gretillat, Brygoo & Domergue, 1962)</small>
::::*''[[Raillietiella trachea]]'' <small>Riley, Oaks & Gilbert, 2003</small>
::::*''[[Raillietiella venteli]]'' <small>(Motta, 1965)</small>
:::*''[[Yelirella]]'' <small>Spratt, 2010</small>
::::*''[[Yelirella petauri]]'' <small>(Spratt, 2010)</small>
*'''[[Reighardiida]]''' <small>Almeida & Christoffersen, 1999</small>
::*'''[[Reighardiidae]]''' <small>Heymons, 1926</small>
:::*''[[Hispania (genus)|Hispania]]'' <small>Martínez ''et al.'', 2004</small>
::::*''[[Hispania vulturis]]'' <small>Martínez ''et al.'', 2004</small>
:::*''[[Reighardia]]'' <small>Ward, 1899</small>
::::*''[[Reighardia lomviae]]'' <small>Dyck, 1975</small>
::::*''[[Reighardia sternae]]'' <small>(Diesing, 1864)</small>

== References ==
{{reflist|32em}}

{{Arthropods}}
{{Arthropod infestations}}

[[Category:Maxillopoda]]
[[Category:Parasitic crustaceans]]
== 分類 ==
== 分類 ==
最新的證據顯示，五口動物實為退化了的寄生性節肢動物。
最新的證據顯示，五口動物實為退化了的寄生性節肢動物。

2014年8月28日 (四) 19:03的版本

舌形動物
Pentastomida
化石時期： 寒武紀第三期至現代
表达式错误：预期外的<运算符。

表达式错误：预期外的<运算符。

Adult female Linguatula serrata
Adult female Linguatula serrata
科學分類
界： 動物界 Animalia
門： 節肢動物門 Arthropoda
亞門： 甲殼亞門 Crustacea
綱： 顎足綱 Maxillopoda
亞綱： 舌形動物亞綱 Pentastomida
Diesing, 1836
目

舌形动物類（學名：Pentastomida），又称五口动物類，是动物界的一个分類。這一種謎樣的動物屬於無脊椎動物，介乎於环节动物和节肢动物之间，均為寄生性動物。其特徵是体软、扁而长，无色，透明，无足。身長從數毫米到十數厘米，体表分近百个清晰的节段，内部却并不分节。虫体前端口部突出，呈椭圆形，周围有钩2对,可伸缩，用以附着在寄主组织上。该门动物没有呼吸系统、排泄系统和循环系统，雌雄异体。

本物種的名稱是因為從其部分物種的外型酷似脊椎動物的舌頭，因而為名；而其學名及另一名稱「五口動物」的來源，則是因為其口器的五個鈎，好像有五個口一樣，因而為名。這五個「鈎」，其實只有一個為口器，其餘四個只是顎足。過往由於對本分類物種的分類不明，單單從外形來判斷，所以將其歸類到門一層。後來到了2001年，本物種生物跟其他物種的關係得到確認，所以被降至歸入成為顎足綱的一員，是為舌形動物亞綱。

現時本分類有物種約130個，皆為被動式寄生物，其部分解剖結果為退化性。成年的舌蟲長約1至14厘米（0.4至5.5英寸） ，多數於宿主的呼吸系統黏付着。

生物

Historically significant accounts of tongue worm biology and systematics include early work by Josef Aloys Frölich,[1] Alexander von Humboldt,[2] Karl Asmund Rudolphi,[3] Karl Moriz Diesing[4] and Rudolph Leuckart.[5]

Other important summaries have been published by Louis Westenra Sambon,[6] Richard Heymons[7] and John Riley,[8] and a review of their evolutionary relationships with a bibliography up to 1969 was published by J. T. Self.[9]

Anatomy

Pentastomids are worm-like animals ranging from 2至13厘米（0.79至5.12英寸） in length. The anterior end of the body bears five protuberances, four of which are clawed legs, while the fifth bears the mouth. The body is segmented and covered in a chitinous cuticle. The digestive tract is simple and tubular, since the animal feeds entirely on blood, although the mouth is somewhat modified as a muscular pump.[10]

The nervous system is similar to that of other arthropods, including a ventral nerve cord with ganglia in each segment. Although the body contains a haemocoel, there are no circulatory, respiratory, or excretory organs.[10]

Life cycle

Pentastomids live in the upper respiratory tract of reptiles, birds, and mammals, where they lay eggs. They are gonochoric (having 2 sexes), and employ internal fertilisation. The eggs are either coughed out by the host or leave the host body through the digestive system. The eggs are then ingested by an intermediate host, which is commonly either a fish or a small herbivorous mammal.[10]

The larva hatches in the intermediate host and breaks through the wall of the intestine. It then forms a cyst in the intermediate host's body. The larva is initially rounded in form, with four or six short legs, but moults several times to achieve the adult form. The pentastomid reaches the main host when the intermediate host is eaten by the main host, and crawls into the respiratory tract from the oesophagus.[10]

Human infestation

Extraction of an Armillifer grandis nymph from a human eye.

Tongue worms occasionally parasitise humans.[11] While there is a report of Sebakia inducing dermatitis,[12] the two genera responsible for most internal human infestation are Linguatula and Armillifer. Visceral pentastomiasis can be caused by Linguatula serrata, Armillifer armillatus, Armillifer moniliformis, Armillifer grandis, and Porocephalus crotali.[13]

The terms associated with infections can vary:

Porocephalus and Armillifer (which are all cylindrical and all inhabit snakes) have much more in common with each other than they do with Linguatula (which is flat and inhabits dogs and wolves.)

Affinities

The affinities of tongue worms have long proved controversial. Historically, they were initially compared to various groups of parasitic worms. Once the arthropod-like nature of their cuticle was recognised, similarities were drawn with mites,[16] particularly gall mites (Eriophyidae). Although gall mites are much smaller than tongue worms they also have a long, segmented body and only two pairs of legs. Later work drew comparisons with millipedes and centipedes (Myriapoda), with velvet worms (Onychophora) and water bears (Tardigrada). Some authors interpreted tongue worms as essentially intermediate between annelids and arthropods, while others suggested that they deserved a phylum of their own. Tongue worms grow by moulting, which suggests they belong to Ecdysozoa, while other work has identified the arthropod-like nature of their larvae,[17] In general, there are two current alternative interpretations: pentastomids are highly modified and parasitic crustaceans, probably related to fish lice, or they are an ancient group of stem-arthropods, close to the origins of Arthropoda.

Crustaceans

The possibility that tongue worms are crustaceans can be traced back to the work of Pierre-Joseph Van Beneden,[18] who compared them to parasitic copepods. The modern form of this hypothesis dates from Karl Georg Wingstrand's study of sperm morphology,[19] which recognised similarities in sperm structure between tongue worms and fish lice (Argulidae) – a group of maxillipod crustaceans which live as parasites on fish and occasionally amphibians. John Riley and colleagues also offered a detailed justification for the inclusion of the tongue worms among the crustaceans.[20] The fish louse model received significant further support from the molecular work of Lawrence G. Abele and colleagues.[21] A number of subsequent molecular phylogenies have corroborated these results,[22][23][24] and the name Ichthyostraca has been proposed for a (Pentastomida + Branchiura) clade.[25] Thus a number of important standard works and databases on crustaceans now include the pentastomids as members of this group.[26]

Stem-arthropods

Critics of the Ichthyostraca hypothesis have pointed out that even parasitic crustaceans can still be recognised as crustaceans based on their larvae; but that tongue worms and their larvae do not express typical characters for Crustacea or even Euarthropoda. An alternative model notes the extremely ancient Cambrian origins of these animals and interprets tongue worms as stem-group arthropods.[27] A recent morphological analysis recovered Pentastomida outside the arthropods, as sister group to a clade including nematodes, priapulids and similar ecdyzoan 'worm' groups.[28] Adding fossils, they suggested an extinct animal called Facivermis could be closely related to tongue worms. However it should be stressed that these authors did not explicitly test pentastomid/crustacean relationships.

Fossil record

Exceptionally preserved, three-dimensional and phosphatised fossils from the Upper Cambrian Orsten of Sweden[29] and the Cambrian/Ordovician boundary of Canada[30] have been identified as pentastomids. Four fossil genera have been identified so far: Aengapentastomum, Bockelericambria, Haffnericambria and Heymonsicambria. These fossils suggest that pentastomids evolved very early and raise questions about whether these animals were parasites at this time, and if so, on which hosts. Conodonts have sometimes been mentioned as possible hosts in this context.

Classification

This article follows Martin and Davis[26] in placing Pentastomida in the class Maxillopoda within the subphylum Crustacea. The subclass contains the following extant orders, superfamilies, families, genera and species:[31]

Armillifer armillatus Wyman, 1848, a 4 cm individual collected from the respiratory system of a python, Python sebae. Specimen deposited in the Museum für Naturkunde Berlin
Female (right) and male (left) Armillifer sp.
  • Alofia Giglioli in Sambon, 1922

References

  1. ^ J. A. Frölich. Beschreibung einiger neuer Eingeweidewürmer. Der Naturforscher. 1789, 24: 101–162. 
  2. ^ A. von Humboldt. Sur un ver intestin trouvé dans les poumons du serpent à sonnettes, de Cumana. Voyage de Humboldt et Bonpand 2. Ptie. F. Schoell et G. Dufour, Paris. 1811: 298–304. 
  3. ^ K. A. Rudolphi. Entozoorum Synopsis. Augustus Rücker Berlin. 1819. 
  4. ^ K. M. Diesing. Versuch einer Monographie der Gattung Pentastoma. Annalen des Wiener Museums der Naturgeschichte. 1835, 1: 1–32. 
  5. ^ R. Leuckart. Bau und Entwicklungsgeschichte der Pentastomen nach Untersuchungen besonders von Pent. taenoides und P. denticulatum. C. F. Winter'sche Verlagshandlung, Leipzig: vi + 160. 1860. 
  6. ^ L. W. Sambon. A synopsis of the family Linguatulidae. Journal of Tropical Medicine and Hygiene. 1922, 12: 188–206, 391–428. 
  7. ^ R. Heymons. Pentastomida. H. G. Bronns (编). Klassen und Ordnungen des Tierreichs. Fünfter Band. IV Abteilung, 1. Buch. Akademische Verlagsgesellschaft m.b.H., Leipzig. 1935: 1–268 pp. 
  8. ^ J. Riley. The biology of pentastomids. Advances in Parasitology. 1986, 25: 45–128. doi:10.1016/S0065-308X(08)60342-5. 
  9. ^ J. T. Self. Biological relationships of the Pentastomida: a bibliography on the Pentastomida. Experimental Parasitology. 1969, 21: 63–119. doi:10.1016/0014-4894(69)90222-7. 
  10. ^ 10.0 10.1 10.2 10.3 Barnes, Robert D. Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. 1982: 880–881. ISBN 0-03-056747-5. 
  11. ^ A. Fain. The Pentastomida parasitic in man. Annales de la Société belge de médecine tropicale. 1975, 55: 59–64. 
  12. ^ H. Solano Mairena & W. Venegas. Human dermatitis caused by a nymph of Sebekia. American Journal of Tropical Medicine and Hygiene. 1989, 41 (3): 352–354. PMID 2802021. 
  13. ^ Dennis Tappe & Dietrich W. Büttner. Bethony, Jeffrey M. , 编. Diagnosis of human visceral pentastomiasis. PLoS Neglected Tropical Diseases. 2009, 3 (2): e320. PMC 2643528 可免费查阅. PMID 19238218. doi:10.1371/journal.pntd.0000320. 
  14. ^ Esmond M. Mapp, Howard M. Pollack & Louis H. Goldman. Roentgen diagnosis of Armillifer armillatus infestation (porocephalosis) in man. Journal of the National Medical Association. May 1976, 68 (3): 198–200, 191. PMC 2609651 可免费查阅. PMID 933188. 
  15. ^ Philip E. S. Palmer; Maurice Merrick Reeder. Imaging of tropical diseases: with epidemiological, pathological, and clinical correlation. Birkhäuser. 2001: 389– [19 April 2010]. ISBN 978-3-540-62471-4. 
  16. ^ T. D. Schubart. Ueber die Entwicklung des Pentastoma taenioides. Zeitschrift für Wissenschaftliche Zoologie. 1853, 4: 117–118. 
  17. ^ G. Osche. "Arthropodencharaktere" bei einem Pentastomiden Embryo (Reighhardia sernae). Zoologischer Anzeiger. 1959, 163: 169–178. 
  18. ^ P. J. van Beneden. Recherches sur l’organisation et le développement des Lingatules (Pentastoma Rud.), suivies de la description d’une espèce nouvelle provenant d’un Mandrill. Annales des Sciences Naturelles Zoologie Series. 1849, 3 (11): 313–348. 
  19. ^ K. G. Wingstrand. Comparative spermatology of a pentastomid, Raillietiella hemidactyli, and a branchiuran crustacean, Argulus foliaceus, with a discussion of pentastomid relationships. Det Kongelige Danske Videnskabernes Selskab Biologiske Skrifter. 1972, 19 (4): 1–72. 
  20. ^ J. Riley, A. A. Banaja & J. L. James. The phylogenetic relationships of the Pentastomida: the case for their inclusion within the Crustacea. International Journal of Parasitology. 1978, 8 (4): 245–254. doi:10.1016/0020-7519(78)90087-5. 
  21. ^ L. G. Abele, W. Kim & B. E. Felgenhauer. Molecular evidence for inclusion of the Phylum Pentastomida in the Crustacea (PDF). Molecular Biology and Evolution. 1989, 6 (6): 685–691. 
  22. ^ D. V. Lavrov, W. M. Brown & J. L. Boore. Phylogenetic position of the Pentastomida and (pan)crustacean relationships (PDF). Proceedings of the Royal Society of London B. 2004, 271 (1538): 537–544. PMC 1691615 可免费查阅. PMID 15129965. doi:10.1098/rspb.2003.2631. 
  23. ^ O. S. Møller, J. Olesen, A. Avenant-Oldewage, P. F. Thomsen & H. Glenner. First maxillae suction discs in Branchiura (Crustacea): development and evolution in light of the first molecular phylogeny of Branchiura, Pentastomida, and other "Maxillopoda". Arthropod Structure & Development. 2008, 37 (4): 333–346. PMID 18394959. doi:10.1016/j.asd.2007.12.002. 
  24. ^ Todd H. Oakley, Joanna M. Wolfe, Annie R. Lindgren & Alexander K. Zaharoff. Phylotranscriptomics to bring the understudied into the fold: monophyletic Ostracoda, fossil placement, and Pancrustacean phylogeny. Molecular Biology and Evolution. 2013, 30 (1): 215–233. PMID 22977117. doi:10.1093/molbev/mss216. 
  25. ^ J. Zrzavý. The interrelationships of metazoan parasites: a review of phylum- and higher-level hypotheses from recent morphological and molecular phylogenetic analyses (PDF). Folia Parasitologica. 2001, 48: 81–103. 
  26. ^ 26.0 26.1 J. W. Martin & G. E. Davis. An Updated Classification of the Recent Crustacea (PDF). Natural History Museum of Los Angeles County. 2001: 132 pp. 
  27. ^ Dieter Waloszek, John E. Repetski & Andreas Maas. A new Late Cambrian pentastomid and a review of the relationships of this parasitic group. Transactions of the Royal Society of Edinburgh: Earth Sciences. 2006, 96 (2): 163–176. doi:10.1017/S0263593300001280. 
  28. ^ W. O. Almeida, M. L. Christoffersen, D. S. Amorim & E. C. C. Eloy. Morphological support for the phylogenetic positioning of Pentastomida and related fossils (PDF). Revista Biotemas. 2008, 21: 81–90. doi:10.5007/2175-7925.2008v21n3p81. 
  29. ^ D. Walossek & K. J. Müller. Pentastomid parasites from the Lower Palaeozoic of Sweden. Transactions of the Royal Society of Edinburgh: Earth Sciences. 1994, 85: 1–37. doi:10.1017/s0263593300006295. 
  30. ^ Dieter Walossek, John E. Repetski & Klaus J. Müller. An exceptionally preserved parasitic arthropod, Heymonsicambria taylori n. sp. (Arthropoda increate sedis: Pentastomida) from Cambrian – Ordovician boundary beds of Newfoundland. Canadian Journal of Earth Sciences. 1994, 31 (11): 1664–1671. doi:10.1139/e94-149. 
  31. ^ Geoff Boxshall. Pentastomida. WoRMS. 2013 [May 13, 2013]. 
按亞門分類的現存节肢动物
螯肢亞門
真螯肢類
頭胸足類
平背板類（英语：Planaterga）
十三腹節類
有顎類
多足亞門
泛甲殼動物
(甲殼亞門 +
六足亞門)
寡甲總綱（英语：Oligostraca）
魚介形綱
Altocrustacea
多甲總綱
Allotriocarida
Labiocarida
六足亞門
由节肢动物感染引起的疾病（B85–B89，132–134）
昆虫纲
蛛形纲
甲壳亚门
描述
疾病
治療

分類

最新的證據顯示，五口動物實為退化了的寄生性節肢動物。

動物界現存各門
動
物
界
真後生
動物亞界
副同源异形基因动物
（浮浪幼蟲樣動物）
两侧对称动物
腎管動物
  • （见下 ↓）
 
原口动物
蛻
皮
動
物
有棘動物（英语：Scalidophora）
線蟲-鎧甲-
泛節肢動物
分支
線蟲動物
鎧甲-泛節肢
動物分支
泛节肢动物
序足動物
螺
旋
動
物
有顎動物（英语：Gnathifera (clade)）
微顎-合皮
動物分支
合皮動物（英语：Syndermata）
扁
蟲
冠
輪
動
物（英语：Platytrochozoa）
吮食-中生
動物分支
吮食动物
中生动物
冠
輪
動
物
软体-隐担轮
动物分支
隱擔輪動物（英语：Kryptotrochozoa）
觸手冠動物
苔蘚動物
帚腕动物（英语：Brachiozoa）
後口動物
步带动物
一些門下的
主要子類群
检索自“https://zh.wikipedia.org/w/index.php?title=舌形動物類&oldid=32475991”