Pentastomida

The Pentastomida are an enigmatic group of parasitic arthropods commonly known as tongue worms due to the resemblance of the species of the genus Linguatula to a vertebrate tongue; molecular studies point to them being highly derived crustaceans.[1]

About 130 species of pentastomids are known; all are obligate parasites with correspondingly degenerate anatomy. Adult tongue worms vary from about 1 to 14 cm (0.4 to 5.5 in) in length and parasitize the respiratory tracts of vertebrates. They have five anterior appendages. One is the mouth; the others are two pairs of hooks, which they use to attach to the host. This arrangement led to their scientific name, meaning "five openings", but although the appendages are similar in some species, only one is a mouth.

## Contents

* 1 Taxonomy
* 1.1 Affinities
* 1.2 Crustaceans
* 1.3 Stem-arthropods
* 1.4 Fossil record
* 1.5 Classification
* 2 Description
* 3 Behaviour and ecology
* 4 Human infestation
* 5 References
* 6 External links

## Taxonomy

Historically significant accounts of tongue worm biology and systematics include early work by Josef Aloys Frölich,[2] Alexander von Humboldt,[3] Karl Asmund Rudolphi,[4] Karl Moriz Diesing[5] and Rudolph Leuckart.[6]

Other important summaries have been published by Louis Westenra Sambon,[7] Richard Heymons[8] and John Riley,[9] and a review of their evolutionary relationships with a bibliography up to 1969 was published by J. T. Self.[10]

### 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 recognized, similarities were drawn with mites,[11] 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{{citation needed|date=July 2021} ntially 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.[12] In general, the two current alternative interpretations are: 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 discovery that tongue worms are crustaceans can be traced back to the work of Pierre-Joseph Van Beneden,[13] who compared them to parasitic copepods. The modern form of this hypothesis dates from Karl Georg Wingstrand's study of sperm morphology,[14] which recognized similarities in sperm structure between tongue worms and fish lice (Argulidae) – a group of maxillopod 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.[15] The fish louse model received significant further support from the molecular work of Lawrence G. Abele and colleagues.[16] A number of subsequent molecular phylogenies have corroborated these results,[17][18][19] and the name Ichthyostraca has been proposed for a (Pentastomida + Branchiura) clade.[20] Thus a number of important standard works and databases on crustaceans now include the pentastomids as members of this group.[21]

### Stem-arthropods

Critics of the Ichthyostraca classification have pointed out that even parasitic crustaceans can still be recognized 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.[22] A 2008 morphological analysis recovered Pentastomida outside the arthropods, as sister group to a clade including nematodes, priapulids and similar ecdysozoan 'worm' groups.[23] 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 and do not take the molecular information into account.

### Fossil record

Exceptionally preserved, three-dimensional and phosphatised fossils from the Upper Cambrian Orsten of Sweden[24] and the Cambrian/Ordovician boundary of Canada[25] have been identified as pentastomids. Also one from the Wuluian (middle Cambrian) of Greenland.[26] Five fossil genera have been identified from the Cambrian so far: Aengapentastomum, Boeckelericambria, Dietericambria, 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 (primitive fish) have sometimes been mentioned as possible hosts in this context.[25] A fifth genus, Invavita, is from Silurian-aged marine strata of England: fossil specimens of Invavita are found firmly attached to their ostracod hosts of the species Nymphatelina gravida.[27][28] It possessed a head, a worm-like body, and two pairs of limbs.[29]

### Classification

There are four extant orders recognized in the subclass Pentastomida:

* Cephalobaenida
* Porocephalida
* Raillietiellida
* Reighardiida

## Description

Pentastomids are worm-like animals ranging from 1 to 14 centimetres (0.39 to 5.51 in) in length. The female is larger than the male. 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, except from genus Linguatula which lives in the nasal cavity of carnivorous mammals where they feed mainly on mucus and dead cells,[30][31] although the mouth is somewhat modified as a muscular pump.[32]

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, no circulatory, respiratory, or excretory organs are present.[32]

## Behaviour and ecology

Armillifer armillatus from a python

Pentastomids live in the upper respiratory tract of reptiles, birds, and mammals, where they lay eggs. They are gonochoric (having two 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.[32]

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. At least one species, Subtriquetra subtriquetra, has a free-living larva.[33] There is both indirect development with nymphal stages and direct development. 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.[32][34]

## Human infestation

File:Ocular-Pentastomiasis-in-the-Democratic-Republic-of-the-Congo-pntd.0003041.s001.ogv

Tongue worms occasionally parasitise humans.[35] While a report exists of Sebekia inducing dermatitis,[36][37] 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.[38]

Armillifer armillatus Wyman, 1848, a 4 cm individual collected from the respiratory system of a python, Python sebae. Specimen deposited in the Natural History Museum of Berlin. Female (right) and male (left) Armillifer sp.

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.[39] (An alternate name for Armillifer moniliformis is Porocephalus moniliformis.)[40]
* "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).

## References

1. ↑ Lavrov, Dennis V.; Brown, Wesley M.; Boore, Jeffrey L. (7 March 2004). "Phylogenetic position of the Pentastomida and (pan)crustacean relationships". Proceedings of the Royal Society of London. Series B: Biological Sciences 271 (1538): 537–544. doi:10.1098/rspb.2003.2631. PMID 15129965.
2. ↑ J. A. Frölich (1789). "Beschreibung einiger neuer Eingeweidewürmer". Der Naturforscher 24: 101–162.
3. ↑ A. von Humboldt (1811). "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. pp. 298–304.
4. ↑ K. A. Rudolphi (1819). Entozoorum Synopsis. Augustus Rücker Berlin.
5. ↑ K. M. Diesing (1835). "Versuch einer Monographie der Gattung Pentastoma". Annalen des Wiener Museums der Naturgeschichte 1: 1–32.
6. ↑ R. Leuckart (1860) (in German). Bau und Entwicklungsgeschichte der Pentastomen nach Untersuchungen besonders von Pent. taenoides und P. denticulatum. C. F. Winter'sche Verlagshandlung, Leipzig. pp. vi + 160.
7. ↑ L. W. Sambon (1922). "A synopsis of the family Linguatulidae". Journal of Tropical Medicine and Hygiene 12: 188–206, 391–428.
8. ↑ R. Heymons (1935). "Pentastomida". in H. G. Bronns. Klassen und Ordnungen des Tierreichs. Fünfter Band. IV Abteilung, 1. Buch. Leipzig: Akademische Verlagsgesellschaft m.b.H.. pp. 1–268 pp.
9. ↑ J. Riley (1986). "The biology of pentastomids". Advances in Parasitology 25: 45–128. doi:10.1016/S0065-308X(08)60342-5. ISBN 9780120317257. PMID 3535437.
10. ↑ J. T. Self (1969). "Biological relationships of the Pentastomida: a bibliography on the Pentastomida". Experimental Parasitology 24 (1): 63–119. doi:10.1016/0014-4894(69)90222-7. PMID 4887218.
11. ↑ T. D. Schubart (1853). "Ueber die Entwicklung des Pentastoma taenioides". Zeitschrift für Wissenschaftliche Zoologie 4: 117–118.
12. ↑ G. Osche (1959). ""Arthropodencharaktere" bei einem Pentastomiden Embryo (Reighhardia sernae)". Zoologischer Anzeiger 163: 169–178.
13. ↑ P. J. van Beneden (1849). "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 3 (11): 313–348.
14. ↑ K. G. Wingstrand (1972). "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 19 (4): 1–72.
15. ↑ J. Riley, A. A. Banaja & J. L. James (1978). "The phylogenetic relationships of the Pentastomida: the case for their inclusion within the Crustacea". International Journal for Parasitology 8 (4): 245–254. doi:10.1016/0020-7519(78)90087-5.
16. ↑ L. G. Abele, W. Kim & B. E. Felgenhauer (1989). "Molecular evidence for inclusion of the Phylum Pentastomida in the Crustacea". Molecular Biology and Evolution 6 (6): 685–691. http://mbe.oxfordjournals.org/cgi/reprint/6/6/685.pdf.
17. ↑ D. V. Lavrov, W. M. Brown & J. L. Boore (2004). "Phylogenetic position of the Pentastomida and (pan)crustacean relationships". Proceedings of the Royal Society of London B 271 (1538): 537–544. doi:10.1098/rspb.2003.2631. PMID 15129965. PMC 1691615. http://www.eeob.iastate.edu/faculty/LavrovD/publications/PDF_files/pentastomida.pdf. Retrieved 2009-12-20.
18. ↑ O. S. Møller; J. Olesen; A. Avenant-Oldewage; P. F. Thomsen; H. Glenner (2008). "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 37 (4): 333–346. doi:10.1016/j.asd.2007.12.002. PMID 18394959. Bibcode: 2008ArtSD..37..333M.
19. ↑ Todd H. Oakley; Joanna M. Wolfe; Annie R. Lindgren; Alexander K. Zaharoff (2013). "Phylotranscriptomics to bring the understudied into the fold: monophyletic Ostracoda, fossil placement, and Pancrustacean phylogeny". Molecular Biology and Evolution 30 (1): 215–233. doi:10.1093/molbev/mss216. PMID 22977117. http://pdxscholar.library.pdx.edu/cgi/viewcontent.cgi?article=1050&context=bio_fac.
20. ↑ J. Zrzavý (2001). "The interrelationships of metazoan parasites: a review of phylum- and higher-level hypotheses from recent morphological and molecular phylogenetic analyses". Folia Parasitologica 48 (2): 81–103. doi:10.14411/fp.2001.013. PMID 11437135.
21. ↑ J. W. Martin; G. E. Davis (2001). An Updated Classification of the Recent Crustacea. Natural History Museum of Los Angeles County. pp. 132 pp. http://atiniui.nhm.org/pdfs/3839/3839.pdf. Retrieved 2009-12-20.
22. ↑ Dieter Waloszek, John E. Repetski & Andreas Maas (2006). "A new Late Cambrian pentastomid and a review of the relationships of this parasitic group". Transactions of the Royal Society of Edinburgh: Earth Sciences 96 (2): 163–176. doi:10.1017/S0263593300001280.
23. ↑ W. O. Almeida; M. L. Christoffersen; D. S. Amorim; E. C. C. Eloy (2008). "Morphological support for the phylogenetic positioning of Pentastomida and related fossils". Biotemas 21 (3): 81–90. doi:10.5007/2175-7925.2008v21n3p81.
24. ↑ D. Walossek; K. J. Müller (1994). "Pentastomid parasites from the Lower Palaeozoic of Sweden". Transactions of the Royal Society of Edinburgh: Earth Sciences 85: 1–37. doi:10.1017/s0263593300006295.
25. ↑ 25.0 25.1 Dieter Walossek, John E. Repetski & Klaus J. Müller (1994). "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 31 (11): 1664–1671. doi:10.1139/e94-149.
26. ↑ Peel, John S. (2022). "The oldest tongue worm: A stem-group pentastomid arthropod from the early middle Cambrian (Wuliuan Stage) of North Greenland (Laurentia)". GFF 144 (2): 97–105. doi:10.1080/11035897.2022.2064543. Bibcode: 2022GFF...144...97P.
27. ↑ Gill, Victoria (22 May 2015). "A 425-million-year-old parasite found attached to host". BBC Online. https://www.bbc.co.uk/news/science-environment-32829628.
28. ↑ Siveter, David J.; Briggs, Derek E.G.; Siveter, Derek J.; Sutton, Mark D. (2015). "A 425-Million-Year-Old Silurian Pentastomid Parasitic on Ostracods". Current Biology 25 (12): 1632–1637. doi:10.1016/j.cub.2015.04.035. PMID 26004764. Bibcode: 2015CBio...25.1632S.
29. ↑ "Requiem for an ancient tongue worm". 21 May 2015. http://news.yale.edu/2015/05/21/requiem-ancient-tongue-worm.
30. ↑ Mehlhorn, Heinz (21 May 2001). Encyclopedic Reference of Parasitology: Biology, Structure, Function. Springer. ISBN 978-3-540-66819-0. https://books.google.com/books?id=9uGQnKdxhyUC&dq="if+they+are+living+in+the+nasopharyngeal+cavity+of+mammals"&pg=PA474.
31. ↑ Barton, Diane P.; Gherman, Calin Mircea; Zhu, Xiaocheng; Shamsi, Shokoofeh (2022). "Characterization of tongue worms, Linguatula SPP. (Pentastomida) in Romania, with the first record of an unknown adult Linguatula from roe deer (Capreolus capreolus Linnaeus)". Parasitology Research 121 (8): 2379–2388. doi:10.1007/s00436-022-07566-9. PMID 35689112.
32. ↑ 32.0 32.1 32.2 32.3 Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 880–881. ISBN 0-03-056747-5.
33. ↑ Williams Jr, E. H.; Bunkley-Williams, L. (2019). "Life Cycle and Life History Strategies of Parasitic Crustacea". Parasitic Crustacea. Zoological Monographs. 3. pp. 179–266. doi:10.1007/978-3-030-17385-2_5. ISBN 978-3-030-17383-8.
34. ↑ Martin, Joel W.; Olesen, Jørgen; Høeg, Jens T. (July 2014). Atlas of Crustacean Larvae. JHU Press. ISBN 978-1-4214-1198-9. https://books.google.com/books?id=PPnZAwAAQBAJ&dq=pentastomida+1+species+has+a+free-living+larva&pg=PA135.
35. ↑ A. Fain (1975). "The Pentastomida parasitic in man". Annales de la Société Belge de Médecine Tropicale 55 (1): 59–64. PMID 1231664.
36. ↑ H. Solano Mairena; W. Venegas (1989). "Human dermatitis caused by a nymph of Sebekia". American Journal of Tropical Medicine and Hygiene 41 (3): 352–354. doi:10.4269/ajtmh.1989.41.352. PMID 2802021.
37. ↑ Correct spelling: Sebakia → Sebekia, See "Sebekia Sambon, 1922". WoRMS. World Register of Marine Species. http://www.marinespecies.org/aphia.php?p=taxdetails&id=384727.
38. ↑ Dennis Tappe & Dietrich W. Büttner (2009). Bethony, Jeffrey M.. ed. "Diagnosis of human visceral pentastomiasis". PLOS Neglected Tropical Diseases 3 (2): e320. doi:10.1371/journal.pntd.0000320. PMID 19238218.
39. ↑ Esmond M. Mapp, Howard M. Pollack & Louis H. Goldman (May 1976). "Roentgen diagnosis of Armillifer armillatus infestation (porocephalosis) in man". Journal of the National Medical Association 68 (3): 198–200, 191. PMID 933188.
40. ↑ Philip E. S. Palmer; Maurice Merrick Reeder (2001). Imaging of tropical diseases: with epidemiological, pathological, and clinical correlation. Birkhäuser. pp. 389–. ISBN 978-3-540-62471-4. https://books.google.com/books?id=b3_RMZl464cC&pg=PA389. Retrieved 19 April 2010.

## External links

* v
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Extant Arthropoda classes by subphylum

* Kingdom Animalia
* Subkingdom Eumetazoa
* (unranked) Bilateria
* (unranked) Protostomia
* Superphylum Ecdysozoa

Chelicerata|

* Pycnogonida (sea spiders)

Euchelicerata|

* Xiphosura (horseshoe crabs)
* Arachnida (spiders, scorpions, ticks, mites)

Myriapoda|

* Chilopoda (centipedes)
* Diplopoda (millipedes)
* Pauropoda (pauropods)
* Symphyla (symphylans or garden centipedes)

Pancrustacea
(Crustacea +
Hexapoda)|

| Oligostraca|

* Ostracoda (seed shrimps)
* Mystacocarida¹

Ichthyostraca|

* Pentastomida (tongue worms)¹
* Branchiura (fish lice)¹

Altocrustacea|

| Multicrustacea|

* Malacostraca (woodlice, shrimps, crayfish, lobsters, crabs)
* Hexanauplia (copepods¹, barnacles and relatives and tantulocaridans¹)

Allotriocarida|

* Cephalocarida (horseshoe shrimps)
* Branchiopoda (fairy, tadpole, clam shrimps, water fleas)

Labiocarida|

* Remipedia

Hexapoda|

* Protura (coneheads)²
* Collembola (springtails)²
* Diplura (two-pronged bristletails)²
* Insecta (insects)

* ¹Maxillopoda
* ²Entognatha
* italic are paraphyletic groups
* Sources: Edgecombe et al. (2014), Petrunina (2012) for pancrustaceans.

Template:Arthropod infestations

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Extant Animal phyla

Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta

Animal
Fungi)

| Animalia|

* Porifera (sponges)

Diploblasts
(Eumetazoa)|

* Ctenophora (comb jellies)

ParaHoxozoa
(Planulozoa)|

| P+C|

* Placozoa (Trichoplax)
* Cnidaria (jellyfish and relatives)

Bilateria
(Triploblasts)|

* Xenacoelomorpha
* Xenoturbellida (Xenoturbella)
* Acoelomorpha
* acoels
* nemertodermatids

Nephrozoa|

* (see below↓)

Nephrozoa

| Deuterostomia|

* Chordata
* lancelets
* tunicates
* Vertebrata/Craniata

Ambulacraria|

* Echinodermata (starfish and relatives)
* Hemichordata
* acorn worms
* pterobranchs

Protostomia|

| Ecdysozoa|

| Scalidophora|

* Kinorhyncha (mud dragons)
* Priapulida (penis worms)

|

N+L+P|

| Nematoida|

* Nematoda (roundworms)
* Nematomorpha (horsehair worms)

|

L+P|

* Loricifera (corset animals)

Panarthropoda|

* Onychophora (velvet worms)

Tactopoda|

* Arthropoda (arthropods)
* Tardigrada (waterbears)

Spiralia|

| Gnathifera|

* Chaetognatha (arrow worms)
* Gnathostomulida (jaw worms)

M+S|

* Micrognathozoa (Limnognathia)
* Syndermata
* Rotifera (wheel animals)
* Acanthocephala (thorny-headed worms)

Platytrochozoa|

| R+M|

| Mesozoa|

* Orthonectida
* Dicyemida or Rhombozoa
* Monoblastozoa (Salinella)†

|

Rouphozoa|

* Platyhelminthes (flatworms)
* Gastrotricha (hairybacks)

Lophotrochozoa|

* Cycliophora (Symbion)
* Annelida (ringed worms)
* Sipuncula ‡

M+K|

* Mollusca (molluscs)

Kryptotrochozoa|

* Nemertea (ribbon worms)

Lophophorata|

| Bryozoa s.l.|

* Entoprocta or Kamptozoa
* Ectoprocta (moss animals)

|

Brachiozoa|

* Brachiopoda (lamp shells)
* Phoronida (horseshoe worms)

* * *

| Major groups
within phyla|

* Sponges
* Hexactinellid
* Demosponge
* Calcareous
* Homoscleromorpha
* Cnidarians
* Anthozoa inc. corals
* Hexacorallia
* Octocorallia
* Medusozoa inc. jellyfish
* Myxozoa
* Vertebrates
* Jawless fish
* Cartilaginous fish
* Bony fish
* Amphibians
* Reptiles
* Birds
* Mammals
* Echinoderms
* Sea lilies
* Asterozoa inc. starfish
* Echinozoa
* Nematodes
* Chromadorea
* Enoplea
* Secernentea
* Arthropods
* Chelicerates/Arachnids
* Myriapods
* Crustaceans
* Hexapods/Insects
* Platyhelminths
* Turbellaria
* Trematoda
* Monogenea
* Cestoda
* Bryozoans
* Phylactolaemata
* Stenolaemata
* Gymnolaemata
* Annelids
* Polychaetes
* Clitellata
* Echiura
* Sipuncula
* Molluscs
* Gastropods
* Cephalopods
* Bivalves
* Chitons
* Tusk shells

|

Phyla with ≥1000 extant species bolded
Potentially dubious phyla †
Disputed phyla ‡
See also
Diploblasts

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