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Moniliformidae
Adult Moniliformis moniliformis
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Acanthocephala
Class:	Archiacanthocephala
Order:	Moniliformida Schmidt, 1972
Family:	Moniliformidae Van Cleave, 1924

Moniliformidae is a family of parasitic spiny-headed (or thorny-headed) worms. It is the only family in the Moniliformida order and contains three genera: Australiformis containing a single species, Moniliformis containing eighteen species and Promoniliformis containing a single species. Genetic analysis have determined that the clade is monophyletic despite being distributed globally. These worms primarily parasitize mammals, including humans in the case of Moniliformis moniliformis, and occasionally birds by attaching themselves into the intestinal wall using their hook-covered proboscis. The intermediate hosts are mostly cockroaches. The distinguishing features of this order among archiacanthocephalans is the presence of a cylindrical proboscis with long rows of hooks with posteriorly directed roots and proboscis retractor muscles that pierce both the posterior and ventral end or just posterior end of the receptacle. Infestation with Monoliformida species can cause moniliformiasis, an intestinal condition characterized as causing lesions, intestinal distension, perforated ulcers, enteritis, gastritis, crypt hypertrophy, goblet cell hyperplasia, and blockages.

Taxonomy and description

Species of the family Moniliformidae are usually pseudosegmented and have a cylindrical proboscis with longitudinal rows of hooks that have posteriorly directed roots. Moniliformidae are further characterized by the presence of a simple, double-walled proboscis receptacle with the outer wall having spirally aligned muscle fibers (with the exception of Australiformis), brain at posterior end of receptacle, and dorsal and ventral lacunar canals. The proboscis retractor muscles pierce both the posterior and ventral end or just posterior end of the receptacle. The cerebral ganglion is in the mid to posterior region, and the lemnisci are long and flat and not bound to the body wall. These worms also lack protonephridia and males have eight cement glands, each with a giant nucleus, which are used to temporarily close the posterior end of the female after copulation. Genetic analysis has been conducted on four species: Moniliformis moniliformis, M. saudi, M. cryptosaudi and M. kalahariensis. Based on these results, Moniliformidae has been determined to be monophyletic.

Genera

There are three genera and twenty living species in the order Moniliformida.

Australiformis

Australiformis is a monotypic genus that infest marsupials in Australia and New Guinea. It was described by Schmidt and Edmonds in 1989. Its body consists of a proboscis armed with hooks which it uses to pierce and hold the gut wall of its host, and a long trunk. It contains a single species, Australiformis semoni. This genus resembles species in the genus Moniliformis but is characterized by a lack of spiral muscles in the outer wall of the proboscis receptacle. The proboscis is armed with 12 rows of 13 to 15 hooks which are used to attach themselves to the small or large intestines of the host. The female worms range from 95 to 197 millimetres, long virtually all of which is the trunk, and 1.75 to 3.5 millimetres wide. There is pronounced sexual dimorphism in this species as females are around twice the size of the males whose trunks range from 46 to 80 millimetres long and 2 millimetres wide. Infestation of marsupials by A. semoni may cause debilitating inflammation of the stomach (gastritis) with granulomatous ulcers.

Moniliformis

The genus Moniliformis Travassos, 1915 contains eighteen species. Description of the genus is the same as the family Moniliformidae with the exception of possessing spiral muscles in the outer wall of the proboscis receptacle and a single distinct kind of proboscis hooks.

Promoniliformis

The genus Promoniliformis Dollfus and Golvan, 1963 is characterized by possessing two distinct kinds of proboscis hooks. There is only one species in this genus. It contains a single species P. ovocristatus.

Hosts

Moniliformidae species are found in the intestines parasitizing mammals and occasionally birds. Intermediate hosts are mostly cockroaches but also other insect groups. Infestation can cause moniliformiasis, which is characterized as lesions in the intestines, intestinal distension, perforated ulcers, enteritis, crypt hypertrophy, goblet cell hyperplasia, and occlusions of the intestinal tract in the gray squirrel (Sciurus carolinensis).

• Hosts for Moniliformidae species
• The Southern brown bandicoot is a host for Australiformis semoni
• The Cairo spiny mouse is a host of Moniliformis acomysi
• The four-toed jerboa is a host of Moniliformis aegyptiacus
• The North African hedgehog is a host of Moniliformis aegyptiacus
• The cotton-top tamarins is a host of Moniliformis clarki
• The large flying fox is a host of Moniliformis convolutus
• The long-eared hedgehog is a host of Moniliformis cryptosaudi
• The moonrat is a host of Moniliformis echinosorexi
• The giant anteater is a host of Moniliformis monoechinus
• The desert hedgehog is a host of Moniliformis saudi
• The brown rat is a host of M. siciliensis and Moniliformis travassosi
• The garden dormouse is a host of M. siciliensis
• The black rat is a host of Moniliformis spiralis
• The Horsfield's tarsier is a host of Moniliformis tarsii
• Promoniliformis ovocristatus is a parasite of the tailless tenrec
• Promoniliformis ovocristatus is a parasite of the lesser hedgehog tenrec

Notes

1. A binomial authority in parentheses indicates that the species was originally described in a genus other than the present genus.

References

1. Van Cleave, H. J. (1924). "A Critical Study of the Acanthocephala Described and Identified by Joseph Leidy". Proceedings of the Academy of Natural Sciences of Philadelphia. 76: 279–334. JSTOR 4063926.
2. ^ Schmidt, Gerald D.; Edmonds, Stanley J. (1989). "Australiformis semoni (Linstow, 1898) n. Gen., n. Comb. (Acanthocephala: Moniliformidae) from Marsupials of Australia and New Guinea". The Journal of Parasitology. 75 (2): 215–7. doi:10.2307/3282769. JSTOR 3282769. PMID 2926590.
3. ^ Kükenthal, W (2014). Gastrotricha and Gnathifera. Göttingen, Germany: Walter de Gruyter. p. 322. ISBN 978-3110274271.
4. ^ Amin, O. M. (1987). "Key to the families and subfamilies of Acanthocephala, with the erection of a new class (Polyacanthocephala) and a new order (Polyacanthorhynchida)". Journal of Parasitology. 73 (6): 1216–1219. doi:10.2307/3282307. JSTOR 3282307. PMID 3437357.
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6. ^ Amin, Omar M.; Heckmann, Richard A.; Osama, Mohammed; Evans, R. Paul (2016). "Morphological and molecular descriptions of Moniliformis saudi sp. n. (Acanthocephala: Moniliformidae) from the desert hedgehog, Paraechinus aethiopicus (Ehrenberg) in Saudi Arabia, with a key to species and notes on histopathology". Folia Parasitologica. 63. doi:10.14411/fp.2016.014. ISSN 0015-5683. PMID 27189420.
7. ^ Amin, Omar M.; Heckmann, Richard A.; Sharifdini, Meysam; Albayati, Nagham Yaseen (2019). "Moniliformis cryptosaudi n. sp. (Acanthocephala: Moniliformidae) from the Long-eared Hedgehog Hemiechinus auritus (Gmelin) (Erinaceidae) in Iraq; A Case of Incipient Cryptic Speciation Related to M. saudi in Saudi Arabia". Acta Parasitologica. 64 (1): 195–204. doi:10.2478/s11686-018-00021-9. PMID 30666546. S2CID 58572640.
8. Nascimento Gomes, Ana Paula; Cesário, Clarice Silva; Olifiers, Natalie; de Cassia Bianchi, Rita; Maldonado, Arnaldo; Vilela, Roberto do Val (December 2019). "New morphological and genetic data of Gigantorhynchus echinodiscus (Diesing, 1851) (Acanthocephala: Archiacanthocephala) in the giant anteater Myrmecophaga tridactyla Linnaeus, 1758 (Pilosa: Myrmecophagidae)". International Journal for Parasitology: Parasites and Wildlife. 10: 281–288. Bibcode:2019IJPPW..10..281N. doi:10.1016/j.ijppaw.2019.09.008. PMC 6906829. PMID 31867208.
9. Amin, O.M.; Sharifdini, M.; Heckmann, R.A.; Zarean, M. (2020). "New perspectives on Nephridiacanthus major (Acanthocephala: Oligacanthorhynchidae) collected from hedgehogs in Iran". Journal of Helminthology. 94: e133. doi:10.1017/S0022149X20000073. PMID 32114988. S2CID 211725160.
10. Amin, Omar M. (September 19, 2013). "Classification of the Acanthocephala". Folia Parasitologica. 60 (4): 273–305. doi:10.14411/fp.2013.031. PMID 24261131.
11. "Moniliformida Schmidt, 1972". Integrated Taxonomic Information System (ITIS). November 23, 2019. Retrieved January 30, 2020.
12. Lenhaus, Cornelius; Obendorf, David; Wright, Frank H. (1990). "Veterinary aspects of Perameles gunnii biology with special reference to species conservation". In Clark, Tim W.; Seebeck, John H. (eds.). Management and conservation of small populations. Chicago Zoological Society. pp. 89–108. ISBN 0-913934-16-X. Retrieved March 23, 2020.
13. ^ Ward, Helen L.; Nelson, Diane R. (1967). "Acanthocephala of the Genus Moniliformis from Rodents of Egypt with the Description of a New Species from the Egyptian Spiny Mouse (Acomys cahirinus)". The Journal of Parasitology. 53 (1): 150–156. doi:10.2307/3276638. JSTOR 3276638. PMID 6066757.
14. Dollfus, Robert-Ph.; Golvan, Yves-J. (1963). "Sur un singulier Métacanthocéphale parasite d'insectivores (Tenrecinae) de Madagascar et des Comores". Annales de Parasitologie Humaine et Comparée. 38 (5): 793–806. doi:10.1051/parasite/1963385793.
15. Singleton, Jeurel; Richardson, Dennis J.; Lockhart, J. Mitchell (January 1993). "Severe Moniliformiasis (Acanthocephala: Moniliformidae) in a Gray Squirrel, Sciurus carolinensis, from Arkansas, USA". Journal of Wildlife Diseases. 29 (1): 165–168. doi:10.7589/0090-3558-29.1.165. PMID 8445783. S2CID 10220897.

• Acanthocephala families
• Archiacanthocephala