Moniliformidae

{{short description|Family of worms}} {{Automatic taxobox | taxon = Moniliformidae | image = Moniliformis moniliformis.jpg | image_caption = Adult ''Moniliformis moniliformis'' | parent_authority = Schmidt, 1972 | authority = Van Cleave, 1924 <ref name="VanCleave1924">{{Cite journal |jstor = 4063926|title = A Critical Study of the Acanthocephala Described and Identified by Joseph Leidy|last1 = Van Cleave|first1 = H. J.|journal = Proceedings of the Academy of Natural Sciences of Philadelphia|volume = 76|pages = 279–334|year = 1924}}</ref> }}

'''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.<ref name="Schmidt1989">{{Cite journal |doi = 10.2307/3282769|pmid = 2926590|jstor = 3282769|title = ''Australiformis semoni'' (Linstow, 1898) n. Gen., n. Comb. (Acanthocephala: Moniliformidae) from Marsupials of Australia and New Guinea|journal = The Journal of Parasitology|volume = 75|issue = 2|pages = 215–7|year = 1989|last1 = Schmidt|first1 = Gerald D.|last2 = Edmonds|first2 = Stanley J.}}</ref><ref name="Kükenthal">{{cite book |last=Kükenthal|first=W |date=2014 |title=Gastrotricha and Gnathifera |url=https://books.google.com/books?id=hwCTBgAAQBAJ&q=apororhynchus&pg=PP11 |location=Göttingen, Germany |publisher=Walter de Gruyter |page=322 |isbn=978-3110274271}}</ref> The proboscis retractor muscles pierce both the posterior and ventral end or just posterior end of the receptacle.<ref name = "Amin1987">{{cite journal |last1=Amin |first1=O. M.|date=1987 |title=Key to the families and subfamilies of Acanthocephala, with the erection of a new class (Polyacanthocephala) and a new order (Polyacanthorhynchida) |journal=Journal of Parasitology |volume=73 |issue=6 |pages=1216–1219 |doi=10.2307/3282307 |jstor=3282307 |pmid=3437357 }}</ref> 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.<ref name="Schmidt1989"/><ref name="Kükenthal"/><ref name="Bush2001">{{cite book | last1=Bush | first1=Albert O.| last2=Fernández| first2=Jacqueline C.| last3=Esch| first3=Gerald W.| last4=Seed| first4=J. Richard | title=Parasitism : the diversity and ecology of animal parasites | publisher=Cambridge University Press | location=Cambridge, UK New York, NY | year=2001 | isbn=0-521-66278-8 | oclc=44131774 | page=203}}</ref> Genetic analysis has been conducted on four species: ''Moniliformis moniliformis'', ''M. saudi'', ''M. cryptosaudi'' and ''M. kalahariensis''.<ref name="AminHeckmann2016">{{cite journal|last1=Amin|first1=Omar M.|last2=Heckmann|first2=Richard A.|last3=Osama|first3=Mohammed|last4=Evans|first4=R. Paul|title=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|journal=Folia Parasitologica|volume=63|year=2016|issn=0015-5683|doi=10.14411/fp.2016.014|pmid=27189420|doi-access=free}}</ref><ref name="Amin2019">{{Cite journal | doi=10.2478/s11686-018-00021-9| pmid=30666546| title=''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| journal=Acta Parasitologica| volume=64| issue=1| pages=195–204| year=2019| last1=Amin| first1=Omar M.| last2=Heckmann| first2=Richard A.| last3=Sharifdini| first3=Meysam| last4=Albayati| first4=Nagham Yaseen |s2cid=58572640 | doi-access=free}}</ref> Based on these results, Moniliformidae has been determined to be monophyletic.<ref name="AminHeckmann2016"/><ref name="Amin2019"/>

{{cladogram|align=center|title=Archiacanthocephala |caption=Phylogenetic reconstruction for select species in the class Archiacanthocephala<ref name="Gomes 2019">{{cite journal |last1=Nascimento Gomes |first1=Ana Paula |last2=Cesário |first2=Clarice Silva |last3=Olifiers |first3=Natalie |last4=de Cassia Bianchi |first4=Rita |last5=Maldonado |first5=Arnaldo |last6=Vilela |first6=Roberto do Val |title=New morphological and genetic data of Gigantorhynchus echinodiscus (Diesing, 1851) (Acanthocephala: Archiacanthocephala) in the giant anteater Myrmecophaga tridactyla Linnaeus, 1758 (Pilosa: Myrmecophagidae) |journal=International Journal for Parasitology: Parasites and Wildlife |date=December 2019 |volume=10 |pages=281–288 |doi=10.1016/j.ijppaw.2019.09.008|pmid=31867208 |pmc=6906829 |bibcode=2019IJPPW..10..281N }}</ref><ref name="Amin2020">{{Cite journal |doi = 10.1017/S0022149X20000073|title = New perspectives on Nephridiacanthus major (Acanthocephala: Oligacanthorhynchidae) collected from hedgehogs in Iran|year = 2020|last1 = Amin|first1 = O.M.|last2 = Sharifdini|first2 = M.|last3 = Heckmann|first3 = R.A.|last4 = Zarean|first4 = M.|journal = Journal of Helminthology|volume = 94|pages = e133|pmid = 32114988|s2cid = 211725160}}</ref> |cladogram=

{{clade |label1=Archiacanthocephala |1={{clade |label1=Oligacanthorhynchidae |1= {{clade |1=''Macracanthorhynchus ingens'' |2=''Oncicola venezuelensis'' |3=''Oligacanthorhynchus tortuosa'' |4=''Nephridiacanthus major''}} |2={{clade |1=''Moniliformis moniliformis'' |label1='''Moniliformidae''' |label2=Gigantorhynchida |2={{clade |1=''Mediorhynchus'' sp. |2=''Gigantorhynchus echinodiscus'' }} }} }} }} }}

== Genera == There are three genera and twenty living species in the order Moniliformida.<ref name="AminHeckmann2016"/><ref name="Amin2013">{{cite journal |last1=Amin |first1=Omar M. |title=Classification of the Acanthocephala |journal=Folia Parasitologica |date=September 19, 2013 |volume=60 |issue=4 |pages=273–305 |doi=10.14411/fp.2013.031 |pmid=24261131 |doi-access=free }}</ref><ref name="ITIS">{{cite web |url=https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=196572|title=Moniliformida Schmidt, 1972|date=November 23, 2019 |website=Integrated Taxonomic Information System (ITIS) |access-date=January 30, 2020}}</ref>{{#tag:ref|A binomial authority in parentheses indicates that the species was originally described in a genus other than the present genus.|group=lower-alpha}}

===''Australiformis''=== {{main|Australiformis}} ''Australiformis'' is a monotypic genus that infest marsupials in Australia and New Guinea. It was described by Schmidt and Edmonds in 1989.<ref name="Schmidt1989"/> 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.<ref name="Schmidt1989" /> Infestation of marsupials by ''A. semoni'' may cause debilitating inflammation of the stomach (gastritis) with granulomatous ulcers.<ref name="Lenhaus1990">{{Cite book |last1=Lenhaus |first1=Cornelius |title=Management and conservation of small populations |last2=Obendorf |first2=David |last3=Wright |first3=Frank H. |date=1990 |publisher=Chicago Zoological Society |isbn=0-913934-16-X |editor-last=Clark |editor-first=Tim W. |pages=89–108 |chapter=Veterinary aspects of ''Perameles gunnii'' biology with special reference to species conservation |access-date=March 23, 2020 |editor-last2=Seebeck |editor-first2=John H. |chapter-url=https://www.researchgate.net/publication/258341847}}</ref>

===''Moniliformis''=== {{main|Moniliformis}} The genus ''Moniliformis'' <small>Travassos, 1915</small> 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<ref name="Schmidt1989"/> and a single distinct kind of proboscis hooks.<ref name="Ward1967">{{Cite journal |doi = 10.2307/3276638|jstor = 3276638|title = Acanthocephala of the Genus Moniliformis from Rodents of Egypt with the Description of a New Species from the Egyptian Spiny Mouse (''Acomys cahirinus'')|journal = The Journal of Parasitology|volume = 53|issue = 1|pages = 150–156|year = 1967|last1 = Ward|first1 = Helen L.|last2 = Nelson|first2 = Diane R.|pmid = 6066757}}</ref>

===''Promoniliformis''=== {{Main|Promoniliformis}} The genus ''Promoniliformis'' <small>Dollfus and Golvan, 1963</small><ref name="Dollfus1963">{{cite journal |last1=Dollfus |first1=Robert-Ph. |last2=Golvan |first2=Yves-J. |title=Sur un singulier Métacanthocéphale parasite d'insectivores (Tenrecinae) de Madagascar et des Comores |journal=Annales de Parasitologie Humaine et Comparée |date=1963 |volume=38 |issue=5 |pages=793–806 |doi=10.1051/parasite/1963385793 |doi-access=free }}</ref> is characterized by possessing two distinct kinds of proboscis hooks.<ref name="Ward1967"/> 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.<ref name = "Amin1987"/> Intermediate hosts are mostly cockroaches but also other insect groups.<ref name="AminHeckmann2016"/> 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'').<ref name="Singleton1993">{{cite journal |last1=Singleton |first1=Jeurel |last2=Richardson |first2=Dennis J. |last3=Lockhart |first3=J. Mitchell |title=Severe Moniliformiasis (Acanthocephala: Moniliformidae) in a Gray Squirrel, Sciurus carolinensis, from Arkansas, USA |journal=Journal of Wildlife Diseases |date=January 1993 |volume=29 |issue=1 |pages=165–168 |doi=10.7589/0090-3558-29.1.165 |pmid=8445783 |s2cid=10220897 }}</ref>

<gallery mode="packed" widths="180" caption="Hosts for ''Moniliformidae'' species"> File:Southern Brown Bandicoot (Isoodon obesulus) 2, Vic, jjron, 09.01.2013.jpg|alt=Southern Brown Bandicoot on the ground in a forest|The Southern brown bandicoot is a host for ''Australiformis semoni'' File:Common spiny mouse.JPG|alt=A Cairo spiny mouse on dirt|The Cairo spiny mouse is a host of ''Moniliformis acomysi'' File:Four-toes-jerboa.jpg|alt=A Four-toed jerboa on a kitchen table with people in the background|The four-toed jerboa is a host of ''Moniliformis aegyptiacus'' File:Atelerix algirus.jpg|alt=A North African hedgehog curled up on hay|The North African hedgehog is a host of ''Moniliformis aegyptiacus'' File:Lisztaffe - Cottontop Tamarin - Saguinus oedipus.jpg|alt=A cotton-top tamarin on a tree stump in the jungle|The cotton-top tamarins is a host of ''Moniliformis clarki'' File:Wilhelma Kalong-Flughund Pteropus vampyrus 0513.jpg|alt=A large flying fox hanging upsidedown on a branch|The large flying fox is a host of ''Moniliformis convolutus'' File:Hemiechinus auritus; Baikonur 09.jpg|alt=Long-eared hedgehog on a wood floor infront of a wood door|The long-eared hedgehog is a host of ''Moniliformis cryptosaudi'' File:Haarigel (Echinosorex gymnura).jpg|alt=A stuffed moonrat on display|The moonrat is a host of ''Moniliformis echinosorexi'' File:Myresluger2.jpg|alt=A giant anteater|The giant anteater is a host of ''Moniliformis monoechinus'' File:Desert Hedgehog.JPG|alt=A Desert hedgehog under branches|The desert hedgehog is a host of ''Moniliformis saudi'' File:Rattus norvegicus -Fairlands Valley Park, Stevenage, England-8.jpg|alt=A brown rat on the ground|The brown rat is a host of ''M. siciliensis'' and ''Moniliformis travassosi'' File:Eliomys quercinus01.jpg|alt=A Garden dormouse on a stone|The garden dormouse is a host of ''M. siciliensis'' File:CSIRO ScienceImage 10564 The black rat Rattus rattus.jpg|alt=A black rat on the ground|The black rat is a host of ''Moniliformis spiralis'' File:Bornean Tarsier (Cephalopachus bancanus borneanus) (8067063777).jpg|alt=Close-up of the face of Horsfield's tarsier on a branch|The Horsfield's tarsier is a host of ''Moniliformis tarsii'' File:Tanrek.jpg|alt=A Tailless tenrec sitting|''Promoniliformis ovocristatus'' is a parasite of the tailless tenrec File:Kleiner-igeltanrek-a.jpg|alt=A Lesser hedgehog tenrec in front of rocks|''Promoniliformis ovocristatus'' is a parasite of the lesser hedgehog tenrec </gallery>

==Notes== {{notelist}}

==References== {{reflist}}

{{Taxonbar|from=Q3449713}}

Category:Acanthocephala families Category:Archiacanthocephala