Pentastomida

(′pen·tə′stäm·ə·də)

(invertebrate zoology) A class of bloodsucking parasitic arthropods; the adult is vermiform, and there are two pairs of hooklike, retractile claws on the cephalothorax.

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Animal Classification: Pentastomida

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(Tongue worms)

Phylum: Arthropoda

Subphylum: Crustacea

Class: Pentastomida

Number of families: 8

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Parasitic tongue worms inhabit the respiratory systems of terrestrial vertebrates

Evolution and systematics

Pentastomida once was classified as a minor phylum, a fact reflected in most modern textbooks of parasitology. The classification is being changed, however. The evolutionary history of tongue worms is unique among parasites. The fossil record apparently extends to the late Cambrian period (500 million years ago [mya]), exceeding that of the next oldest parasites, certain copepods, by some 370 million years. Tiny fossil tongue worms have been etched from ancient fine-grained, deep-water limestone when the rock has been dissolved with dilute acid. In most respects these 0.04 to 0.08 in long (1–2 mm) larvae are indistinguishable from modern tongue worms. Therein lies a conundrum, because extant adult tongue worms are parasitic in terrestrial vertebrates, which do not appear in the fossil record until the late Devonian, approximately 350 mya. Nonetheless, fossil agnathan fishes are known from the Upper Cambrian, and it is noteworthy that all of the fossil tongue worms discovered so far have been collected from limestone containing diverse conodont fauna. Conodonts are primitive fish-like chordates. Although a marine ancestry for tongue worms seems assured, the exact nature of the ancestral host may never be known. In addition to having anterior pairs of claws, some of these ancient tongue worms have two pairs of vestigial nonsegmented trunk limbs. These structures are lacking in other fossil specimens and in all modern forms. A compounding problem is that the larvae of modern tongue worms are highly modified for tissue migration. Clawed limbs, together with other limb-like outgrowths evident in early larval development within the egg, have so far proved impossible to homologize with euarthropod head and trunk appendages.

A relation between tongue worms and branchiuran crustaceans (class Maxillopoda, subclass Branchiura) is supported by results of studies of sperm of other species and by comparison of ribosomal 18S recombinant RNA sequences in the two groups. In numerous extant parasitic maxillopods representative of several subclasses, the degree of cephalization together with the development of the trunk and anchoring devices is highly variable, to the extent that many adults bear no resemblance to their free-living counterparts. Always, however, the first larval instar, the nauplius, is easily recognized. The highly modified first instar of tongue worms, the so-called primary larva, is very different from its putative naupliar fore-bear because it has evolved to penetrate and traverse tissues in (mostly) terrestrial hosts. These larvae hatch with only two limb-bearing head somites and three trunk somites. Subsequent growth of the trunk is by a form of pseudometamerism, without the addition of further somites. The development of some copepods, relatives of branchiurans, also fits this pattern.

The present, still unresolved debate hinges heavily on the relative merits of the fossil evidence versus that of ribosomal RNA and sperm morphology. In this entry, tongue worms are considered a class of crustaceans. The class Pentastomida contains eight families and approximately 110 species assorted between two orders—the primitive Cephalobaenida and the advanced Porocephalida.

Cephalobaenida

Cephalobaenidae. Three genera. Cephalobaena, one species found in snakes; Raillietiella, more than 35 species found in amphibians, snakes, lizards, amphisbaenans, and birds; Rileyiella, one species found in mammals.
Reighardiidae. One genus. Reighardia, two species found in marine birds.

Porocephalida

Sebekidae. Seven genera. Alofia, five species; Leiperia, two species; Selfia, one species; Agema, one species, all found in crocodiles; Sebekia, 12 species, found in crocodiles and chelonians (one species); Pelonia, one species; Diesingia, one species, found in chelonians.

Subtriquetridae. One genus. Subtriquetra, three species, found in crocodilians.
Sambonidae. Four genera. Sambonia, four species; Elenia, two species, found in monitor lizards; Waddycephalus, 10 species; Parasambonia, two species, found in snakes.
Porocephalidae. Two genera. Porocephalus, eight species; Kiricephalus, five species, found in snakes.
Armilliferidae. Three genera. Armillifer, seven species; Cubirea, two species; Gigliolella, one species, found in snakes.
Linguatulidae. One genus. Linguatula, more than six species, found in mammals.

Physical characteristics

All tongue worms inhabit the respiratory systems of terrestrial vertebrates. As a consequence, all aspects of their structure and function have adapted to life in this unusual habitat. All are aerodynamic, possessing an elongated, worm-like, mostly cylindrical body, which is rounded both anteriorly and posteriorly. The body is differentiated into an anterior head region, bearing a small ventral mouth flanked by two pairs of retractile hooks, and a long posterior trunk, which carries numerous raised annuli that are not true segments. Sexual dimorphism is pronounced, females are invariably larger than males. Females may be as small as 0.06 × 0.01 in (1.5 × 0.3 mm) (Rileyiella) to 4.7 × 0.4 in (12 × 1 cm) (Armillifer), but males are much shorter and proportionately more slender. The chitinous cuticle is thin, flexible, and translucent, so that in living specimens the body organs, suspended in an extensive fluid-filled haemocoel, are clearly visible. Peristaltic contractions of the body wall musculature affect locomotion, which is comparable with that of soft-skinned dipteran maggots. The cuticle is shed periodically during growth, and simple metamorphosis occurs (developing nymphs resemble adults). Numerous, exceedingly fine, chitinlined ducts erupt over the entire surface of the cuticle, each connected to an extensive system of subparietal gland cells that abound in the haemocoel immediately beneath the tegumental epidermis. Secretion from these glands, composed largely of membranous secretory droplets, is critical to prolonged survival of these parasites in the delicate environment of the lung. Large numbers of distinctive, flask-shaped cells, analogous to those involved in ion transport in other invertebrates, are embedded in the cuticle. Additional gland systems, located mainly in the head (both orders), but also flanking the intestine in porocephalids, discharge copious enzymic secretions over the head and into hook pits. There are no respiratory or excretory systems.

The sucking mouth leads into a short esophagus, which is separated by a valve from a simple undifferentiated tubular intestine. The intestine terminates at a short rectum. The simple nervous system forms initially as separate ganglia that fuse progressively during development to varying degrees within the two orders. In Porocephalida, all ganglia fuse to form a compact "brain." In cephalobaenids only the most anterior ganglia do so. A variety of small sense organs, often visible as distinct papillae, are arranged over the head, but most are concentrated on the ventral surface around the mouth.

The reproductive system differs between the two orders. The uterus of cephalobaenids is saccate, and the vagina opens anteriorly close to the junction of the head and trunk. In porocephalids, the uterus is elongate and tubular. Because it is many times longer than the body, the uterus is irregularly coiled to occupy most of the available haemocoel. The vagina opens near the anus. In both orders, the dorsal ovary leads into a paired oviduct, which passes around the gut. Close to the junction between the oviduct and the uterus are paired spermathecae, which are responsible for long-term storage of sperm.

The lower reproductive tract of males comprises paired, elongate penises—basically thin, coiled tubes of chitin—close to elaborate chitinous spicules called dilators. Dilators may be extruded through the anterior genital pore by muscles and thereby carry the tip of the penis either to the entrance of the spermathecal ducts in cephalobaenids or into the vagina/uterus in porocephalids. Peristalsis within the uterus of porocephalids pulls the ornamented heads of the paired penises toward the spermathecal ducts. The testis is dorsal, and the paired vasa deferentia empty into a seminal vesicle, which functions as a sperm storage depot before intromission.

Distribution

Most tongue worms, approximately 96%, live in tetrapod definitive hosts that are widespread in the tropics and subtropics. Only a few species are found outside these regions. Porocephalus crotali is unique in that it is cold-adapted, infecting North American rattlesnakes, which hibernate each winter. Reighardia sternae is a cosmopolitan species in gulls and terns, which are widespread in both hemispheres. Another cosmopolitan species, Linguatula serrata, lives in the nasal sinuses of canines (dogs, foxes, wolves), whereas the reindeer host of Linguatula arctica, inhabits the arctic tundra.

Habitat

Most tongue worms live in the lungs of their definitive hosts, although two genera, Leiperia and Subtriquetra, inhabit the trachea and nasal sinuses respectively. At least one member of the genus Elenia infests the throats of monitor lizards. All Linguatula species live in the nasal sinuses of mammals, and Reighardia species are located in the body cavity and air sacs of their avian hosts. An intermediate host is usual in the life cycle of tongue worms. In these cases infective larvae encyst in the tissues of arthropods or vertebrates, depending on the species.

Behavior

Because tongue worms are endoparasites of the respiratory tracts of tetrapods, what little is known about behavior has been inferred mostly from findings at autopsy. In the case of direct life cycles, eggs containing primary larvae gain entry to the definitive host through the alimentary tract as contaminants of food or water. When there are intermediate hosts in the life cycle, larvae acquired by the same means escape from the egg but invade the viscera, where they molt several times to form infective nymphs. The nymphs excyst when intermediate hosts are eaten, and larvae penetrate the stomach or intestinal wall of the final host. This stage is followed by a period of growth in the body cavity before larvae penetrate the lung through the pleura. It is possible to culture in vitro the lung-stage worms of certain species in a blood-based medium under sterile conditions. For example, developing nymphs of Porocephalus crotali, normally resident in the lung of their rattlesnake host, ingest ad libitum the medium in which they are suspended and molt normally through several instars to the adult stage. Thus lung-dwelling species appear not to require specific cues for successful development.

Feeding ecology and diet

With the exception of species belonging to the genus Linguatula, which browse on cells and mucus lining the nasal sinuses, all tongue worms feed on blood. In most cases the blood is pumped from capillaries lining the respiratory surface of the lung by the sucking action of an oral papilla. In some genera (Alofia, Leiperia, Elenia, Waddycephalus, Kiricephalus, and Cuberia) the head of females is separated from the trunk by a distinct neck, which is permanently encapsulated by inflammatory tissue and thereby anchored to the lung wall. In the lung females may also feed on inflammatory cells, as is known to occur during the development of Porocephalus nymphs in rodent intermediate hosts.

Reproductive biology

As far as is known, all tongue worm females become sexually mature precociously. Copulation occurs when the uterus is undeveloped and when males and females are of similar size. Copulation is a lengthy and complex process, entailing docking of the paired penises in the narrow spermathecal duct and concomitant transfer of millions of filiform sperm. Stored sperm fertilize ova released continuously from the ovaries of mature females. Fertilized eggs mature as they descend the uterus of porocephalids, and gravid females of Armillifer and Linguatula species may contain millions of eggs. In contrast, the eggs of cephalobaenids are stored temporarily in a saccate uterus until 30–50% become infectious (i.e., they contain a fully mature primary larva). Then egg deposition begins. The vagina is equipped with a sieve-like mechanism that retains small, undeveloped eggs but allows mature eggs to escape. Thus in both orders, continuous egg production is usual. Eggs shed into lungs are wafted out by the hosts' ciliation and swallowed.

Conservation status

No species are listed by the IUCN.

Significance to humans

The eggs of five tongue worm species are infective to humans, and in four of these species (Armillifer grandis, A. armillatus, A. moniliformis, and A. agkistrodontis), humans are merely an accidental intermediate host. Nearly always, nymphal infections are acquired when eggs in undercooked meat, derived from tongue worm–infected snakes, are consumed. The epidemiology of the remaining species, Linguatula serrata from the nasal sinuses of dogs, is complicated, because both eggs and infective larvae can become established in humans. Ingested eggs hatch to produce infective nymphs. In contrast, if ingested in contaminated offal from sheep or goats, nymphs attempt to migrate from the stomach to the nasal passages, producing acute symptoms of nasopharyngeal linguatulosis.

Resources

Books:

Kabata, Z. Parasitic Copepoda of British Fishes. London: Ray Society, 1979.

Mehlhorn, H. Parasitology in Focus. Berlin: Springer Verlag, 1988.

Riley, J. "Pentastomids." In Reproductive Biology of Invertebrates. Vol. VI, edited by K. G. Adiyodi and R. G. Adiyodi. Oxford: IBH Publishing, 1994.

Periodicals:

Almeida, W. O., and M. L. Christoffersson. "A Cladistic Approach to Relationships in Pentastomida." Journal of Parasitology 85 (1999): 695–704.

Böckeler, W. "Embryogenese und ZNS-Differenzierung bei Reighardia sternae, Lichtund electronenmikroskopishe Untersuchungen zur Tagmosis und systematischen Stellung der Pentastomiden." Zoologische Jahrbucher (Anatomie) 11 (1984): 297–342.

Buckle, A. C., J. Riley, and G. F. Hill. "The in vitro Development of the Pentastomid Porocephalus crotali from the Infective Instar to the Adult Stage." Parasitology 115 (1997): 503–512.

Martin, J. W., and G. E. Davis. "An Updated Classification of the Recent Crustacea." Science Series, Natural History Museum of Los Angeles County 39 (2001): 1–124.

Riley, J. "The Biology of Pentastomids." Advances in Parasitology 25 (1986): 46–128.

Riley, J., and R. J. Henderson. "Pentastomids and the Tetrapod Lung." Parasitology 119, supplement (1999): S89–105.

Storch, V., and B.G.M. Jamieson. "Further Spermatological Evidence for Including the Pentastomida (Tongue Worms) in the Crustacea." International Journal of Parasitology 22 (1992): 95–108.

Walossek, D., and K. J. Müller. "Pentastomid Parasites from the Lower Palaeozoic of Sweden." Transactions of the Royal Society of Edinburgh, Earth Sciences 85 (1994): 1–37.

Wingstrand K. G. "Comparative Spermatology of a Pentastomid Raillietiella hemidactyli and a Branchiuran Crustacean Argulus foliaceus with a Discussion of Pentastomid Relationships. Biologiske Skrifter 19 (1972): 1–72.

[Article by: John Riley, PhD]

Sci-Tech Encyclopedia: Pentastomida

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A class of bloodsucking arthropods, parasitic in the respiratory organs of vertebrates, that frequently are referred to as the Linguatulida or tongue worms. The adults are vermiform, with a short cephalothorax and an elongate, annulate abdomen that may be cylindrical or flattened.

The class is divided into two orders: the Cephalobaenida, a more primitive group, and the Porocephalida, a more specialized one. The first has six-legged larvae and the other, four-legged larvae. The mitelike form of the larvae, with short stumpy legs, demonstrates relationship to the arthropods. Characteristic arthropod features include the presence of (1) jointed appendages in the larvae; (2) stigmata or breathing pores in the body wall; (3) specialized reproductive organs, especially those of the male; and (4) ecdysis or molting of larvae and nymphs. More than 50 species have been described.

Human infection occurs frequently in Africa and the Orient, where humans are an accidental intermediate host of the nymphal form. The liver is a common site of infection, and large numbers of larvae may produce serious and even fatal effects. See also Arthropoda; Cephalobaenida; Porocephalida.

Columbia Encyclopedia: Pentastomida

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Pentastomida (pĕn'təstō'mə), tongue worms, small phylum of fewer than 100 species of parasites living in the upper respiratory passages of reptiles, and occasionally of birds and mammals. They are closely related to arthropods, but zoologists disagree as to their exact placement in that phylum. Recent evidence indicates a possible origin from parasitic crustaceans. The tapering wormlike body, varying in length from ¹/₂ in. to 5 in. (1.3-13 cm), is unsegmented. The front of the body bears five short appendages; one bears the mouth, the other four attach to host tissues. Internal organs are much reduced, except the reproductive organs, which occupy most of the internal space. Eggs are released by the mature pentastomids and hatch when eaten by a suitable intermediate host. When the intermediate host, in turn, is eaten, the larvae migrate to the respiratory passages of the final host, where they take up permanent residence and mature.

Word Tutor: Pentastomida

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IN BRIEF: n. - Tongue worms.

Wikipedia: Pentastomida

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Tongue worms

Linguatula taenioides, a pentastomid parasite of sheep
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Subphylum:	Crustacea
Class:	Maxillopoda
Subclass:	Pentastomida Diesing, 1836
Orders
Cephalobaenida Porocephalida

The Pentastomida are an enigmatic group of parasitic invertebrates commonly known as tongue worms due to the resemblance of the species of the genus Linguatula to a vertebrate tongue.

There are about 130 extant species of pentastomids; all are obligate parasites with correspondingly degenerate anatomy. Adult tongue worms vary from about 1 cm to 14 cm 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.

Alternative names for the Pentastomida include Pentastoma (strictly a genus name), Linguatulida, and Acanthotheca.

1 Biology
- 1.1 Life cycle
2 Human infestation
- 2.1 Linguatuliasis
- 2.2 Pentastomiasis
3 Affinities
- 3.1 Crustaceans?
- 3.2 Stem-arthropods?
4 Fossil record
5 Classification
6 See also
7 References

Biology

Historically significant accounts of tongue worm biology and systematics include early work by J. A. 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].

Life cycle

Pentastomids live in the upper respiratory tract of reptiles, birds, and mammals, where they lay eggs. 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. The first 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 pentastomid reaches the main host when the intermediate host is eaten by the main host.

Human infestation

Tongue worms occasionally parasitise humans ^[10]. While there is a report of Sebakia inducing dermatitis ^[11], the two genera responsible for internal human infestation are Linguatula and Armillifer. Medical terms sometimes used are linguatuliasis, pentastomiasis or porocephalosis, usually depending on the genus involved; although the latter term is misleading as there is no record of Porocephalus affecting humans and 'porocephalosis' probably refers to Armillifer. A modern account of human pentastomid infestation can be found in the study of Dennis Tappe & Dietrich Büttner^[12].

Linguatuliasis

Human infestation by Linguatula was historically more commonplace than is sometimes realised. Human liver autopsies in Berlin from the early part of the 20th century revealed an infection rate of nearly 12%^[13]. The usual final host for Linguatula serrata is a carnivore, like a dog or jackal, and the species is sometimes known as the dog tongueworm for this reason^[14]. Humans can become infected in two ways: either as an intermediate host (visceral linguatuliasis) or as an accidental final host (nasopharyngeal linguatuliasis).

In visceral linguatuliasis Linguatula eggs are sneezed or defecated out by the primary host. Normally they would be eaten by a herbivorous mammal (the intermediate host), including various domestic animals^[15]. The eggs hatch in the intestines and the resulting larvae burrow their way into the visceral cavity of the body. Here they form cysts or granulomas, typically in the liver or the lymph nodes. This can also occur in humans if the eggs are accidentally swallowed, although the victim may not be aware of the infestation and misdiagnosis as liver disease can even occur^[16]. Rarely, the larvae of both Lingulatula and Armillifer can enter the eye accidentally^[17]

In nasopharyngeal linguatuliasis it is the encysted larvae which are consumed, usually via raw or poorly-cooked meat. Once in the stomach, the larvae become liberated from their cysts within a couple of hours and then crawl up the oesophagus and establish themselves in the nose, pharynx or lungs — holding themselves in place with the hooks flanking the mouth ^[18]. Presence of the parasite can induce headaches, coughing and nasal discharge which obviously helps to spread the infection. Nasopharyngeal linguatuliasis appears to be quite prevalent throughout the Middle East where it is often known as the Halzoun syndrome after religious festivals in which infected raw meat is consumed^[19]. In Sudan nasopharyngeal linguatuliasis is known as Marrara syndrome; whereby Marrara is a popular local dish prepared from raw offal. It has been suggested that up to 20% of the population in some parts of Sudan may be affected by this syndrome at some stage of their lives ^[20].

Pentastomiasis

Human infestation by Armillifer is, by contrast, associated with contact to snakes. It is thus prevalent in parts of Africa ^[21] and Asia ^[22] where eating snake meat is common. In Africa it has also been associated with groups who use the snake as a totem ^[23]. Unlike linguatuliasis, humans are only even an accidental intermediate host for Armillifer, i.e. the larvae establish themselves in the visceral organs causing human visceral pentastomiasis, but adults do not occur in the human respiratory system. After a while the larvae die within the host and sometimes calcify, leaving characteristic crescent-shaped structures seen in X-ray^[24]. In extreme cases a heavy parasite burden can have serious medical consequences ^[25] and can even be fatal^[26].

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^[27]; 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 ^[28]. 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^[29] who compared them to parasitic copepods. The modern form of this hypothesis dates from Karl Georg Wingstrand's study of sperm morphology^[30] which recognised similarities in sperm structure between tongue worms and fish lice (Branchiura); a group of maxillipod crustaceans which live as parasites on fish and occasionally amphibians. John Riley and colleagues^[31] also offered a detailed justification for the inclusion of the tongue worms among the crustaceans. The fish louse model received significant further support from the molecular work of Lawrence G. Abele and colleagues^[32]. A number of subsequent molecular phylogenies have corroborated these results^[33]^[34] and the name Ichthyostraca has been proposed for a (Pentastomida + Branchiura) clade^[35]. Thus a number of important standard works^[36] and databases on crustaceans now include the pentastomids as members of this group.

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 exteremely ancient Cambrian origins of these animals and interprets tongue worms as stem-group arthropods^[37]. A recent morphological analysis^[38] recovered Pentastomida outside the arthropods, as sister group to a clade including nematodes, priapulids and similar ecdyzoan 'worm' groups. Adding fossils, they suggested an extinxt animal called Facivermes 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^[39] and the Cambrian/Ordovician boundary of Canada^[40] 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 in placing Pentastomida in the class Maxillopoda within the subphylum Crustacea. The species list below is derived from Heymons (1935) and Self (1969), as well as the more recent literature.

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

Subclass Pentastomida Diesing, 1836

- fossil stem-group taxa
  - †Aengapentastomum Walossek, Repetski & Maas, 2006
    - †Aengapentastomum andresi Walossek, Repetski & Maas, 2006 — Cambrian 'Orsten' of Sweden; host unknown
  - †Boeckelericambria Walossek & Müller, 1994
    - †Boeckelericambria pelturae Walossek & Müller, 1994 — Cambrian 'Orsten' of Sweden; host unknown
  - †Haffnericambria Walossek & Müller, 1994
    - †Haffnericambria trolmeniensis Walossek & Müller, 1994 — Cambrian 'Orsten' of Sweden; host unknown
  - †Heymonsicambria Walossek & Müller, 1994
    - †Heymonsicambria gossmannae Walossek & Müller, 1994 — Cambrian 'Orsten' of Sweden; host unknown
    - †Heymonsicambria kinnekullensis Walossek & Müller, 1994 — Cambrian 'Orsten' of Sweden; host unknown
    - †Heymonsicambria repetski Walossek & Müller, 1994 — Cambrian 'Orsten' of Sweden; host unknown
    - †Heymonsicambria scandia Walossek & Müller, 1994 — Cambrian 'Orsten' of Sweden; host unknown
    - †Heymonsicambria taylori Walossek, Repetski & Müller, 1994 — Cambrian/Ordovician of Canada; host unknown

Taxon Eupentastomida Waloszek, Repetski & Maas, 2006
Order Cephalobaenida Heymons, 1935
- Family Cephalobaenidae Fain, 1961
  - Cephalobaena Heymons, 1922
    - Cephalobaena freitarsi (Motta & da Silva Gomes, 1968)
      - = Travassostulida acutiacanthus Gomes & da Silva Motta, 1968
    - Cephalobaena giglioli (Hett, 1924) — South America; primary host: amphisbaenians
    - Cephalobaena tetrapoda Heymons, 1922 — South America; primary host: snakes
      - = Cepahlobaena recurvocauda da Silva Motta, 1963
  - Mahafaliella Gretillat, Brygoo & Domergue, 1962
    - Mahafaliella tetrapoda Gretillat, Brygoo & Domergue, 1962 — Madagascar; primary host: Madagascar boa
    - Mahafaliella venteli da Silva Motta, 1965
  - Raillietiella Sambon, 1910
    - = Gretillatia da Silva Motta, 1965
    - Raillietiella aegypti Ali, Riley & Self, 1982 — Egypt; primary host: lizards
    - Raillietiella affinis Bovien, 1927
    - Raillietiella agocai Tubangui & Masilungan, 1936 — Philippines; primary host: cobra
    - Raillietiella ampanihyensis Gretillat, Brygoo & Domergue, 1962 — Madagascar: primary host: snakes
    - Raillietiella amphiboluri Mahon, 1954 — Australia; primary host: bearded lizard
    - Railietiella bicaudata Heymons & Vitzthum, 1935
    - Raillietiella bufonis Ali, Riley & Self, 1982 — Puerto Rico; primary host: toads
    - Raillietiella boulengeri (Vaney & Sambon, 1910)— Africa; primary host: snakes
    - Raillietiella boulengeri var. spiralis Heymons, 1939
    - Raillietiella cartagenensis Ali, Riley & Self, 1985 — Colombia; primary host: lizards
    - Raillietiella chamaeleonis Gretillat & Brygoo, 1959
    - Raillietiella chautedeni Fain, 1960
    - Raillietiella congolensis Fain, 1960
    - Raillietiella crotali Ali, Riley & Self, 1984 — Mexico — primary host: rattlesnakes
    - Raillietiella frenatus Ali, Riley & Self, 1980 — Malaysia; primary host: geckos
    - Raillietiella furcocerca (Diesing, 1835)
      - = Raillietella gomesi da Silva Motta, 1963
    - Raillietiella gehyrae Bovien, 1927 — Indonesia
    - Raillietiella gowrii Rajula & Rajendran, 1970
    - Raillietiella hebithamata Self & Kuntz, 1960
    - Raillietiella hemidactyli Hett, 1934 — SE Asia
    - Raillietiella indica Gedoelst, 1921 — India; primary host: toads
    - Raillietiella kochi Heymons, 1926
      - = Raillietiella shipleyi Heymons, 1926
    - Raillietiella mabuiae Heymons, 1922 —— south-west Africa
    - Raillietiella maculatus Rao & Hiregauder, 1959 — India
    - Raillietiella mediterranea (Hett, 1915)
    - Raillietiella morenoi Abreu-Acosta, Foronda Rodriguez, Valladares & Casanova, 2006 — Canary Islands; primary host: lizards
    - Raillietiella orientalis (Hett, 1915) — Asia; primary host: snakes
    - Railletiella spiralis Hett, 1923
    - Raillietiella trachea Riley, Oaks & Gilbert, 2003 — Pakistan; primary host: white backed vulture; autoinfection?
  - Rileyella Spratt, 2003 NB: Homonym of a fly genus!
    - Rileyella petauri Spratt, 2003
- Family Reighardiidae Heymons, 1935
  - Hispania Martínez et al., 2004
    - Hispana vulturis Martínez et al., 2004 — Spain; primary host: black vulture
  - Reighardia Ward, 1899
    - Reighardia lomviae Dyck, 1975 — northern Europe; primary host; guillemot
    - Reighardia sternae Diesing, 1864 — widespread in the northern Hemisphere particularly; primary host: seabirds; no intermediate host (autoinfection)
Order Porocephalida Heymons, 1935
- Superfamily Lingulatoidea Heymons, 1935
- Family Linguatulidae Heymons, 1935
  - Linguatula Frölich, 1789
    - Linguatula dingophila Johnson, 1910 — Australia; primary host: dingos
    - Lingulatula nuttali Sambon, 1922 — East Africa; primary host: lion; intermediate host antilopes
    - Lingulatula recurvata Diesing, 1850 — South America; primary host: jaguar; intermediate host: mammals, fish
    - Linguatula serrata Frölich, 1789 — cosmopolitan; primary host: dogs, wolves, etc; intermediate host: mammals
      - = Taenia rhinaris Meyer, 1789
      - = Taenia capraea Abildgaard, 1789
      - = Polystoma serratum Goeze, 1803
      - = Linguatula denticulata Rudolphi, 1805
      - = Polystoma taenoides Rudolphi, 1809
      - = Prinoderma rhinarium Rudolphi, 1810
      - = Prinoderma lanceolata Cuvier, 1817
      - = Pentatsoma emarginatum Rudolphi, 1819
      - = Pentastoma fera Creplin, 1829
- Family Subtriquetridae Fain, 1961
  - Subtriquetra Sambon, 1922
    - Subtriquetra megacephala (Baird, 1853) — India; primary host: crocodiles; intermediate host: fish
    - Subtriquetra rileyi Junker, Boomker & Booyse, 1998 — Africa; primary host: nile crocodiles?; intermediate host: fish
    - Subtriquetra shipleyi Hett, 1924 — India; primary host: crocodile; intermediate host: fish?
    - Subtriquetra subtriquetra (Diesing, 1835) — South America; primary host: crocodilians; intermediate host: fish
- Superfamily Porocephaloidea Fain, 1961
- Family Porocephalidae Fain, 1961
  - = Armilliferidae Fain, 1961
  - Armillifer Sambon, 1922
    - = Ligamifer Heymons, 1932
    - Armillifer agkistrodontis Self & Kuntz, 1966
    - Armillifer armillatus (Wyman, 1845
      - = Linguatula constrictor Pruner, 1847
      - = Linguatula diesingii van Beneden, 1849
      - = Pentastomum euryzonum Diesing, 1850
      - = Nematoideum hominis (viscerum) Diesing, 1851
      - = Pentastomum polyzonum Harley, 1857
      - = Pentastoma leonis Wedl, 1863
      - = Pentastomum protelis Hoyle, ????
    - Armillifer grandis Hett, 1915
    - Armillifer mazzai (Sambon, 1932)
    - Armillifer moniliformis (Diesing, 1835)
      - = Pentastoma wedlii Cobbald, 1866
      - = Pentastoma aonycis Macalister, 1877
    - Armillifer moniliformis var. heymonsi (Sambon, 1922)
  - Cuberia Kishida, 1928
    - Cuberia armulata (Baird, 1853)
      - = Pentastoma multicinctum Harley, 1857
    - Cuberia pomeroyi (Woodland, 1921)
  - Gigliolella Chaubard & Choquet, 1954
    - Gigliolella brumpti (Giglioli, 1922) — Madagascar; primary host: snakes
  - Kiricephalus Sambon, 1922
    - Kiricephalus clelii Riley & Self, 1980 — West Indies; primary host: pesudoboa
    - Kiricephalus coarctatus (Diesing, 1850)-
      - = Porocephalus seurati Neveu-Lemaire, 1900
      - = Porocephalus herpetodryados Shipley, 1905
      - = Porocephalus globicephalus Hett, 1915
    - Kiricephalus constrictor Riley & Self, 1980 — Central and South America?; primary host boa constrictors
    - Kiricephalus gabonensis Riley & Self, 1980 — West Africa´; primary host colubrid snakes
    - Kiricephalus pattoni (Stephens, 1908) — Asia, Australia; primary host: snakes; intermediate host: amphibians, lizards, snakes
    - Kiricephalus tortus (Shipley, 1898)
  - Porocephalus Humboldt, 1811
    - Porocephalus basilicus Riley & Self, 1979 — Mexico; primary host: rattlesnakes; intermediate host: small mammals?
    - Porocephalus benoiti Fain, 1960 — West Africa; primary/intermediate host?: cobras
    - Porocephalus clavatus (Wyman, 1845) — Central and South America; primary host: constrictors; intermediate host: small mammals
      - = Pentastomum didelphidis virginianae Leidy, 1850
      - = Pentastoma imperatoris Macalister, 1875
    - Porocephalus crotali Humboldt, 1811 — Americas; primary host: rattlesnakes and moccasins; intermediate host: small mammals
      - = Polystoma proboscideum Rudolphi, 1812
      - = Pentastoma subcylindricum Diesing, 1835
    - Porocephalus dominicana Riley & Walters, 1980 — Dominican Republic; primary host: Dominican boas; intermediate host: small mammals?
    - Porocephalus stilesi Sambon, 1910 — Central and South America; primary host: pit vipers; intermediate host: small mammala
    - Porocephalus subulifer (Leuckart, 1860) — Africa; primary host: snake-eating snakes; intermediate host: other snakes
      - = Porocephalus cercopitheci Breinl & Hindle, 1909
      - = Porocephalus bouvieri Vaney & Sambon, 1910
    - Porocephalus tortugensis Riley & Self, 1979 — Tortga Island, Mexico; primary host rattlesnakes; intermediate host: small mammals?
- Family Sebekiidae Fain, 1961
  - = Diesingidae Fain, 1961
  - Agema Riley, Hill & Huchzermeyer, 1997
    - Agema silvaepalustris Riley, Hill & Huchzermeyer, 1997
  - Alofia Giglioli, 1922
    - Alofia ginae Giglioli, 1922
    - Alofia merki Giglioli, 1922
    - Alofia parva Riley & Huchzermeyer, 1995
    - Alofia platycephala (Lohrmann, 1889)
  - Diesingia Sambon, 1922
    - Diesingia kachugensis Sambon, 1922
    - Diesingia megastoma (Diesing, 1835) — South America; primary host: terrapins
  - Elenia Heymons, 1932
    - Elenia australis Heymons, 1932
    - Elenia travassosi Heymons
  - Leiperia Sambon, 1922
    - Leiperia australis Riley & Huchzermeyer, 1996
    - Leiperia cincinnalis Sambon, 1922 — Africa; primary host: crocodiles
  - Pelonia Junker & Boomker, 2002
    - Pelonia africana Junker & Boomker, 2002 — South Africa; primary host: terrapins
  - Sambonia Noc & Giglioli, 1922
    - Sambonia lohrmanni (Sambon, 1910) — Africa; primary host: monitor lizards
    - Sambonia parapodum (Self & Kuntz, 1966)
    - Sambonia solomenensis (Self & Kuntz, 1957)
    - Sambonia varani (Self & Kuntz, 1957)
  - Sebekia Sambon, 1922
    - Sebekia cesarisi Giglioli, 1922
    - Sebekia divestei Giglioli, 1922
    - Sebekia johnstoni Riley, Spratt & Winch, 1990
    - Sebekia mississippiensis Overstreet, Self & Vliet, 1985 — southern USA; primary host: alligators; intermediate host: fish
    - Sebakia multiannulata Riley, Spratt & Winch, 1990
    - Sebakia novaguineae Riley, Spratt & Winch, 1990
    - Sebekia oxycephala (Diesing, 1835)
    - Sebekia purdieae Riley, Spratt & Winch, 1990
    - Sebekia samboni Travassos, 1924
    - Sebekia wedli Giglioli, 1922
  - Selfia Riley, 1994
    - Selfia porosus Riley, 1994
  - Waddycephalus Sambon, 1922
    - Waddycephalus teretiusculus (Baird, 1862)
    - Waddycephalus vitensis Heymons, 1932

nomina dubia
- Alofia adriatica Hirst, 1922
- Alofia indica von Linstow, 1906
- Elenia lialisi Heymons, 1939 — immature Waddycephalus specimens from Indonesia
- Pentastoma gracile Diesing, 1835 — immature specimens from South America
- Pentastomum crocidurae Parona, 1889 — immature specimens from Burma
- Pentastomum najae sputatricis Leuckart, 1860 — immature specimens
- Porocephalus siamensis Koch, 1906 — immature specimens from Thailand
- Raillietiella geckonis Diesing, 1850 — original description inadequate, host uncertain
- Sebekia acuminata Travassos, 1924
- Sebekia jubini Vaney & Sambon, 1910
- Sebekia samboni Travassos, 1924

References

^ Frölich, J. A. 1789. Beschreibung einiger neuer Eingeweidewürmer. Der Naturforscher, 24: 101–162.
^ Humboldt, A. von 1811. Sur un ver intestin trouvé dans les poumons du serpent à sonnettes, de Cumana. In Voyage de Humboldt et Bonpand 2. Ptie. F. Schoell et G. Dufour, Paris: 298–304.
^ Rudolphi, K. A. 1819. Entozoorum Synopsis. Augustus Rücker, Berlin.
^ Diesing, K. M. 1835. Versuch einer Monographie der Gattung Pentastoma. Annalen des Wiener Museums der Naturgeschichte, 1: 1–32.
^ Leuckart, R. 1860. Bau und Entwicklungsgeschichte der Pentastomen nach Untersuchungen besonders von Pent. taenoides und P. denticulatum. C. F. Winter’sche Verlagshandlung, Leipzig, vi + 160 pp.
^ Sambon, L. W. 1922. A synopsis of the family Linguatulidae. Journal of Tropical Medicine and Hygiene, 12: 188–206, 391–428.
^ Heymons, R. 1935. Pentastomida. In Dr. H. G. Bronns Klassen und Ordnungen des Tierreichs. Fünfter Band. IV Abteilung, 1. Buch. Akademische Verlagsgesellschaft m.b.H., Leipzig, 1–268 pp.
^ Riley, J. 1986. The biology of pentastomids. Advances in Parasitology, 25: 45–128.
^ Self, J. T. 1969. Biological relationships of the Pentastomida: a bibliography on the Pentastomida. Experimental Parasitology, 21: 63–119.
^ Fain, A. 1975. The Pentastomida parasitic in man. Annales de la Société belge de médecine tropicale, 55: 59–64.
^ Mairena H., Solano, M. & Venegas, W. 1989. Human dermatitis caused by a nymph of Sebekia. The American Journal of Tropical Medicine and Hygiene, 41: 352–354.
^ Tappe D, Büttner DW 2009. Diagnosis of Human Visceral Pentastomiasis. PLoS Neglected Tropical Diseases, 5(2): e320. doi:10.1371/journal.pntd.0000320 [1]
^ Koch, M. 1906. Zur Kenntnis des Parasitismus der Pentastomen. Verhandlungen der Deutschen Gesellschaft für Pathologie, 10: 265–279.
^ Heymons, R. 1942. Der Nasenwurm des Hundes (Linguatula serrata Froelich), seine Wirte und Beziehungen zur europäischen Tierwelt, seine Herkunft und praktische Bedeutung auf Grund unserer bisherigen Kenntnisse. Zeitschrift für Parasitenkunde, 12: 607–638.
^ Ravindran, R., Lakshmanan, B., Ravishankar, C. & Subramanian, H. 2008. Prevalance of Linguatula serrata in domestic ruminants in South India. Southeast Asian Journal of Tropical Medicine and Public Health, 39: 808–812.
^ Machado, M. A. C., Makdissi, F. F., Canedo, L. F., Martino. R. B., Crescentini, F., Chieffi, P. P., Bachella, T. & Machado, M. C. C. 2006. Unusual case of pentastomiasis mimicking liver tumor. Journal of Gasteroenterology and Hepatology, 21: 1218–1220.
^ Lazo, R. F., Hidalgo, E., Lazo, J. E., Bermeo, A., Llaguno, M., Murillo, J. & Teixeria, V. P. A. 1999. Ocular linguatuliasis in Ecuador: case report and morphometric study of the larva of Linguatula serrata. American Journal of Tropical Medicine and Hygiene, 60: 405–409.
^ Tappe, D., Winzer, R., Büttner, D. W., Ströbel, P., Stich, A., Klinker, H., Frosch, M. 2006. Linguatuliasis in Germany. Emerging Infectious Diseases, 12: 1034–1036 [2].
^ Siavashi, M. R., Assmat, M. & Vatankhah, A. 2002. Nasopharyngeal pentastomiasis (Halzoun): report of 3 cases. Iranian Journal of Medical Science, 27: 191–192.
^ Yagi, H., El Bahari, S. Mohamed, H. A., Sid Ahmed, E. R., Mustafa, B., Mahmoud, M., Saad, M. B. A., Sulaiman, S. M. & El Hassan, A. M. 1996. The Marrara syndrome: a hypersensitivity reaction of the upper respiratory tract and buccopharyngeal mucosa to nymphs of Linguatula serrata. Acta Tropica, 62: 127–134.
^ Du Plessis, V., Birnie, A. J., Eloff, I., Reuter, H. & Andronikou, S. 2007. Pentastomiasis (Armillifer armillatus infestation). South African Medical Journal, 97: 928–930.
^ Yao, M. H., Wu, F., Tang, L. F. 2008. Human pentastomiasis in China: case report and literature review. Journal of Parasitology, 94: 1295–1298.
^ Dakubo, J. C. B., Naaeder, S. B., Kumodji, R. 2008. Totemism and the transmission of human pentastomiasis. Ghana Medical Journal, 42: 165–168.
^ Mapp, E. M., Pollack, M. D. & Goldman, L. H. 1976. Roentgen diagnosis of Armillifer armillatus infestation (Porocephalosis) in man. Journal of the National Medical Association, 68: 198–200.
^ Pan, C.-m., Tang, H.-f., Qui, M.-h. & Xiong, Q.-x. 2005. Heavy infestation with Armillifer moniliformis: a case report. Chinese Medical Journal, 118: 262–264.
^ Yapo Ette, H., Fanton, L., Adou Bryn, K. D., Botti, K., Koffi, K. & Malicier, D. 2003. Human pentastomiasis discovered post-mortem. Forensic Science International, 137: 52–54.
^ Schubart, T. D. 1853. Ueber die Entwicklung des Pentastoma taenioides. Zeitschrift für Wissenschaftliche Zoologie, 4: 117-118
^ Osche, G. 1959. “Arthropodencharaktere” bei einem Pentastomiden Embryo (Reighhardia sernae). Zoologischer Anzeiger, 163: 169–178.
^ Beneden, P. J. van 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.
^ Wingstrand, K. G. 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.
^ Riley, J., Banaja, A. A. & James, J. L. 1978 The phylogenetic relationship of the Pentastomida: the case for their inclusion within the Crustacea. International Journal of Parasitology, 8: 245–254
^ Abele, L. G., Kim, W. & Felgenhauer, B. E. 1989. Molecular evidence for inclusion of the Phylum Pentastomida in the Crustacea. Molecular Biology and Evolution, 6: 685–691.
^ Lavrov, D. V., Brown W. M. & Boore, J. L. 2004. Phylogenetic position of the Pentastomida and (pan)crustacean relationships. Proceedings of the Royal Society of London B, 271: 537–544.
^ Møller, O. S., Olesen, J., Avenant-Oldewage, A., Thomsen, P. F. & Glenner, H. 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 and Development, 37: 333–346.
^ Zrzavý, J. 2001. The interrelationships of metazoan parasites: a review of phylum- and higher-level hypotheses from recent morphological and molecular phylogenetic analyses. Folia Parasitologica, 48: 81–103.
^ Martin, J. W. & Davis, G. E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series, 39: 1–124.
^ Waloszek, D., Repetski, J. E. & Maas, A. 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, 93: 163–176.
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^ Walossek, D. & Müller, K. J. 1994. Pentastomid parasites from the Lower Palaeozoic of Sweden. Transactions of the Royal Society of Edinburgh: Earth Sciences, 85: 1–37.
^ Walossek, D., Repetski, J. E., Müller, K. J. 1994. An exceptionally preserved parasitic arthropod, Heymonsicambria taylori n.sp. (Arthropoda increate sedis: Pentastomida), from Cambrian – Ordovician boundary beds of Newfoundland, Canada. Canadian Journal of Earth Sciences, 31: 1664–1671.

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