Moniliformis Acomysi
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''Moniliformis'' is a genus of parasitic worms in the
Acanthocephala Acanthocephala ( Greek , ' 'thorn' + , ' 'head') is a group of parasitic worms known as acanthocephalans, thorny-headed worms, or spiny-headed worms, characterized by the presence of an eversible proboscis, armed with spines, which it uses t ...
phylum.


Taxonomy

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 In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
.


Description

Species of the genus ''Moniliformis'' are usually pseudosegmented and have a cylindrical proboscis with longitudinal rows of hooks that have posteriorly directed roots. ''Moniliformis'' species are further characterized by the presence of a simple, double-walled proboscis receptacle with the outer wall having spirally aligned muscle fibers, 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 The nephridium (: nephridia) is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Nephridia co ...
and males have eight
cement gland Cement glands are small organs found in Acanthocephala that are used to temporarily close the posterior end of the female after copulation. Cement glands are also mucus-secreting organs that can attach embryos or larvae to a solid substrate. The ...
s, each with a giant nucleus, which are used to temporarily close the posterior end of the female after copulation.


Species

The genus ''Moniliformis'' Travassos, 1915 contains nineteen 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. *''Moniliformis acomysi'' Ward and Nelson, 1967 ''M. acomysi'' was found infesting the
Cairo spiny mouse The Cairo spiny mouse (''Acomys cahirinus''), also known as the common spiny mouse, Egyptian spiny mouse, or Arabian spiny mouse, is a Nocturnality, nocturnal species of rodent in the family Muridae. It is found in Africa north of the Sahara De ...
(''Acomys cahirinus'') in
South Sinai South Sinai ( ') is the least populated governorate of Egypt. It is located in the east of the country, encompassing the southern half of the Sinai Peninsula. Saint Catherine's Monastery, an Eastern Orthodox Church monastery and UNESCO World Her ...
,
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
. The proboscis has eleven to sixteen longitudinal rows of hooks with six to ten hooks per row with the largest hooks being apical and range from 21 to 29 mm long in the male and 23 to 31 mm long in the female. There are eight cement glands in the male and the eggs of the female are oval. The species name ''acomysi'' derives from the host genus name ''Acomys''. *''Moniliformis aegyptiacus'' Meyer, 1932 ''M. aegyptiacus'' infests ''
Meriones sinaiticus Meriones may refer to: *Meriones (mythology) In Greek mythology, Meriones ( ; ) was the Cretan son of Molus and Melphis or Euippe. Molus was a half-brother of Idomeneus. Like other heroes of mythology, Meriones was said to be a descendant o ...
'' and the
four-toed jerboa The four-toed jerboa (''Scarturus tetradactylus'') is a rodent of the family Dipodidae and genus ''Scarturus'' that has four digits. Four-toed jerboas are native to Egypt and Libya. They live in coastal salt marshes and dry deserts. Physical a ...
(''Allactaga tetradactyla'' reported as ''Scarturus tetradactylus'') in
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
and the
North African hedgehog The North African hedgehog (''Atelerix algirus'') or Algerian hedgehog, is a mammal species in the family Erinaceidae native to Algeria, Libya, Malta, Morocco, Spain, and Tunisia. Little is known about this hedgehog, even though the most common b ...
(''Atelerix algirus'' reported as ''Erinaceus algiris'') in
Tunis Tunis (, ') is the capital city, capital and largest city of Tunisia. The greater metropolitan area of Tunis, often referred to as "Grand Tunis", has about 2,700,000 inhabitants. , it is the third-largest city in the Maghreb region (after Casabl ...
. The proboscis has 96 hooks arranged in twelve longitudinal rows with eight hooks per row. The eggs are oval. The species name ''aegyptiacus'' derives from locality of the host, Egypt. *''Moniliformis amini'' Martins, del Rosario Robles, and Navone 2017 ''M. amini'' was found in the small intestine parasitising the
olive grass mouse ''Abrothrix olivacea'', also known as the olive grass mouse or olive akodont,Musser and Carleton, 2005, p. 1090 is a species of rodent in the genus '' Abrothrix'' of family Cricetidae. It is found from northern Chile into southern Chile and ...
(''Abrothrix olivaceus'') in the
Santa Cruz Province, Argentina Santa Cruz Province (, , "Holy Cross") is a Provinces of Argentina, province of Argentina, located in the southern part of the country, in Patagonia. It borders Chubut Province to the north, and Chile to the west and south, with the Atlantic Ocean ...
. The species name ''amini'' was named after Omar Amin, a parasitologist specializing in Acanthocephala.
Sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
is pronounced with female over four times longer than the male. The proboscis has 12 to 14 longitudinal rows of hooks with 10 to 12 hooks per row with the largest hooks in the female being apical ranging from 22.21 to 24.47 μm, the middle being 15.50 μm long and the posterior being the smallest at 12 μm long. The largest hooks in the male are 24 μm. There are eight cement glands in the male, the eggs are oval. *''Moniliformis cestodiformis'' ( Linstow, 1904) ''M. cestodiformis'' has been found infesting hedgehogs of the genus ''
Erinaceus ''Erinaceus'' is a genus of hedgehog from the family of Erinaceidae. There are four main species of ''Erinaceus''. The range is all across Europe, throughout the Middle East, parts of Russia, and extending to northern China and Korea. The Europe ...
'' in
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. The proboscis is between 400 and 500 μm long and 200 μm wide with 16 to 18 rows of 7 or 8 hooks each and no apical pores. The largest hooks are found anteriorly and are 30 to 32 μm long with no dorsoventral differences. The eggs have 4 membranes and are 85 to 92 μm long and 49 to 51 μm wide giving them an elongation ratio of 1.73. *''Moniliformis clarki'' (Ward, 1917) ''M. clarki'' commonly infects squirrels, chipmunks, deer mice, and gophers in North America. If left untreated, it can kill its host. Ten
cotton-top tamarins The cotton-top tamarin (''Saguinus oedipus'') is a small New World monkey weighing less than . This New World monkey can live up to 24 years, but most of them die by 13 years. One of the smallest primates, the cotton-top tamarin is easily recog ...
(''Saguinus oedipus'') were infected at the
St. Louis Zoo The Saint Louis Zoo, officially known as the Saint Louis Zoological Park, is a zoo in Forest Park in St. Louis, Missouri. It is recognized as a leading zoo in animal management, research, conservation, and education. The zoo is accredited by th ...
with at least one being killed by the
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
. Eggs of what is thought to be ''M. clarki'' due to the heavy population of common rodent hosts were discovered while examining human
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name ...
specimens gathered from
Danger Cave Danger Cave is a North American Archaeology, archaeological site located in the Bonneville Basin of western Utah around the Great Salt Lakes region, that features Artifact (archaeology), artifacts of the Desert Culture from c. 9000 BC until c. 500 ...
in
Utah Utah is a landlocked state in the Mountain states, Mountain West subregion of the Western United States. It is one of the Four Corners states, sharing a border with Arizona, Colorado, and New Mexico. It also borders Wyoming to the northea ...
. It was theorized that the man had either eaten a raw insect or an under cooked rodent. The female is twice as long than the male. The proboscis has 12 or 13 longitudinal rows of hooks with six or seven hooks per row with the largest hooks being 23 to 28 mm long. The eggs have 4 membranes and are 75 μm long and have an elongation ratio of 2.42. *''Moniliformis convolutus'' Meyer, 1932 ''M. convolutus'' has been found infesting the
large flying fox The large flying fox (''Pteropus vampyrus'', formerly ''Pteropus giganteus''), also known as the greater flying fox, Malayan flying fox, Malaysian flying fox, large fruit bat, kalang, or kalong, is a southeast Asian species of megabat in the fam ...
(''Pteropus vampyrus'') in
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
. The proboscis is 500 μm long by 200 μm wide with 12 rows of 11 or 12 hooks each. The posterior hooks are rootless and the proboscis receptacle is 700 μm long. *''Moniliformis cryptosaudi'' Amin, Heckmann, Sharifdini, and Albayati, 2019 ''M. cryptosaudi'' is a
cryptic species In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
with ''M. saudi'' being morphologically identical, apart from the metal content of the hooks, but genetically distinct. The hooks consist of a
collagen Collagen () is the main structural protein in the extracellular matrix of the connective tissues of many animals. It is the most abundant protein in mammals, making up 25% to 35% of protein content. Amino acids are bound together to form a trip ...
material with very low levels of
phosphorus Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...
and
calcium Calcium is a chemical element; it has symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to it ...
which is in contrast to those of ''M. saudi'' which has high levels of these two metals. The trunk is between 20.50 ad 48.75 mm long by 0.72 to 1.05 mm wide in the male and 28.75 to 121.25 mm long by 0.65 to 2.25 mm wide in the much larger female. The anterior trunk tapers in a fan-shaped pattern and has a collar of vertical plates. The proboscis is between 343 and 416 μm long by 135 and 188 μm wide in the male and 300 to 416 μm long by 162 to 291 μm wide in the female. The proboscis is armed with 13 to 14 rows of hooks each containing 7 or 8 hooks with the largest being found anteriorly and measuring 25 to 35 μm long in the female and 16 to 25 μm long in the male. The ventral hooks longer than dorsal hooks. The proboscis receptacle is 624 to 1092 μm by 218 to 364 μm in the female and 551 to 936 μm by 200 to 291 μm in the male. The long lemnisci has 6 to 8 giant nuclei and measures 4.26 to 7.28 mm by 0.12 to 0.18 mm in the male and has 6 to 9 giant nuclei and measures 4.15 to 8.12 mm by 0.11 to 0.21 mm in the female. The short lemnisci has 3 or 4 giant nuclei and measures 3.74 to 5.72 mm by 0.09 to 0.15 mm in the male and has 5 or 6 giant nuclei and measures 2.18 to 7.25 mm by 0.11 to 0.21 mm in the female. The anterior testes are 1.65 to 3.67 mm by 0.40 to 0.62 mm in size and the posterior testes are 1.50 to 2.62 mm by 0.37 to 0.62 mm in size. In the male, the cement glands are 322 to 780 μm by 229 to 416 μm and Saefftigen's pouch measures 624 to 925 μm by 270 to 425 μm. In the female, The eggs are oval, measuring 50 to 104 μm by 25 to 36 μm. The species name ''cryptosaudi'' refers to this cryptic status with ''M. saudi''. It has been found in the intestine infesting the
long-eared Hedgehog The long-eared hedgehog (''Hemiechinus auritus'') is a species of hedgehog native to Central Asian countries and some countries of the Middle East. The long-eared hedgehog lives in burrows that it either makes or finds and is distinguished by it ...
(''Hemiechinus auritus'') near
Baqubah Baqubah (; BGN: Ba‘qūbah; also spelled Baquba and Baqouba) is the capital of Iraq's Diyala Governorate. The city is located some to the northeast of Baghdad, on the Diyala River. In 2003 it had an estimated population of some 280,000 people. ...
,
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. *''Moniliformis echinosorexi'' Deveaux, Schmidt and Krishnasamy, 1988 ''M. echinosorexi'' has been found infesting the moonrat (''Echinosorex gymnurus'') in
Malaysia Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
but may also infect hedgehogs. The worms have 12 to 15 rows of between 11 and 13 hooks that change from 34 to 36 μm long. The proboscis receptacle is 1.2 to 2.0 mm long. Males are between 95 and 98 mm long whereas females are longer ranging from 126 to 192 mm long. The proboscis is relatively large ranging from 690 to 1000 μm long with 12 to 15 rows of containing 11 and 13 hooks each. The posterior hooks do not have roots. The proboscis receptacle is large at 1.2 mm long in males and 2.0 mm long in females. The testes are relatively large being between 5.6 and 6.8 mm long. Lemnisci are similarly long at with three giant nuclei each. The species name ''echinosorexi'' derives from the genus of the host, ''Echinosorex''. *''Moniliformis gracilis'' (Rudolphi, 1819) ''M. gracilis'' is a parasite of the
European roller The European roller (''Coracias garrulus'') is the only member of the Coraciidae, roller family breeding in Europe. Its range extends into the Maghreb, West Asia and Central Asia. It winters in southern Africa, primarily in dry wooded savanna and ...
(''Coracias garrullus'') in
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
. The worms are very small being only 2.25 to 3.00 mm long. The proboscis has 10 rows each with 8 or 9 hooks with the largest hooks on the anterior portion. The ganglion at the base of the short receptacle is 360 μm long. *''Moniliformis ibunami'' *''Moniliformis kalahariensis'' Meyer, 1931 ''M. kalahariensis'' was found infesting the Southern African Hedgehog (''Atelerix frontalis'') in Mohlonong and at the
University of Limpopo The University of Limpopo () is a public university in the Limpopo Province, South Africa. It was formed on 1 January 2005, by merger of the University of the North and the Medical University of South Africa (MEDUNSA). These previous institution ...
in the
Limpopo Province Limpopo () is the northernmost province of South Africa. It is named after the Limpopo River, which forms the province's western and northern borders. The term Limpopo is derived from Rivombo (Livombo/Lebombo), a group of Tsonga settlers ...
,
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. and
Namaqua sandgrouse The Namaqua sandgrouse (''Pterocles namaqua''), is a species of ground-dwelling bird in the sandgrouse family. It is found in arid regions of south-western Africa. Taxonomy The Namaqua sandgrouse was formally described in 1789 by the German na ...
(''Pterocles namaqua'') in
Botswana Botswana, officially the Republic of Botswana, is a landlocked country in Southern Africa. Botswana is topographically flat, with approximately 70 percent of its territory part of the Kalahari Desert. It is bordered by South Africa to the sou ...
. The
intermediate host In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include ...
is the
German cockroach The German cockroach (''Blattella germanica''), colloquially known as the croton bug, is a species of small cockroach, typically about long. In color it varies from tan to almost black, and it has two dark, roughly parallel, streaks on the pro ...
(''Blattella germanica''). Juvelines are unsegmented whereas adults are pseudosegmented. There are 2 apical sensory pores on the proboscis. Males trunk is between 44.25 and 75.00 mm long by 1.62 and 1.75 mm wide whereas the females are larger being 51.25 to 81.25 mm long by 2.00 to 2.25 mm wide. In the male, proboscis 780 long by 384 wide and 763 to 863 μm long by 374 to 426 μm wide in the female. The proboscis has 15 or 16 usually straight rows of between 9 and 11 hooks. In the male, the rooted hooks measure 29 to 30 μm, 41 to 47 μm, 55 to 57 μm, 55 to 56 μm from the anterior and the spings spiniform hooks measure 67 to 75 μm, 75 to 77 μm, 65 to 75 μm, 60 to 62 μm, 57 to 60 μm, 35 to 40 μm from the anterior. Proboscis receptacle 1.52 mm long by 0.52 mm wide in the male and 1.75 to 1.80 mm long by 0.47 to 0.55 mm wide in the female. The lemnisci are 18.75 mm long by 0.17 mm wide male and 13.75 to 16.25 mm long by 0.24 to 0.27 mm wide female. The male reproductive system is located in the posterior part of trunk. The testes are elongate with the anterior testis measuring between 3.50 and 6.87 mm long by 0.75 and 1.25 mm wide and the posterior testis measuring between 4.05 and 5.55 mm long by 0.95 and 1.32 mm wide. Cement glands are located posterior of the testes in cluster of 8 and measure between 0.87 and 2.50 mm long by 0.50 and 1.07 mm wide with the anterior glands being the largest. Saefftigen's pouch is 1.62 to 1.75 mm long by 0.50 to 0.57 mm wide. In the juvenile, the reproductive system is underdeveloped, with ovoid testes and occasionally lacking cement glands. The female reproductive system measures 572 to 946 μm in total length but lacks vaginal muscles. The uterus is thick and double-walled measuring 343 to 624 μm long and well-developed ligaments hold the distal end of uterus to body wall. The uterine bell is between 208 and 343 μm long and contains many nuclei. The eggs are ovoid being 73 to 114 μm long by 31 to 62 μm wide and are notched at one pole. *''Moniliformis moniliformis'' (Bremser, 1811) ''M. moniliformis'' is a globally distributed parasite where the adult worms are usually found in intestines of
rodent Rodents (from Latin , 'to gnaw') are mammals of the Order (biology), order Rodentia ( ), which are characterized by a single pair of continuously growing incisors in each of the upper and Mandible, lower jaws. About 40% of all mammal specie ...
s or
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
s such as cats and dogs but can on rare occasions infest humans. The
intermediate hosts In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include a ...
are usually
beetle Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
s and
cockroach Cockroaches (or roaches) are insects belonging to the Order (biology), order Blattodea (Blattaria). About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known Pest (organism), pests. Modern cockro ...
es. The male averages 77 mm long and 1.66 mm wide whereas the female is much larger at 140 mm long and 1.24 mm wide. The
scolex Cestoda is a class of parasitic worms in the flatworm phylum (Platyhelminthes). Most of the species—and the best-known—are those in the subclass Eucestoda; they are ribbon-like worms as adults, commonly known as tapeworms. Their bodies consi ...
of this worm has a cylindrical proboscis and a multitude of curved hooks. The proboscis of both genders are 0.50 mm long and between 1.19 and 1.24 mm wide and the proboscis receptacle is between 0.89 and 1.07 mm long and 0.27 and 0.28 mm wide. The proboscis has 11 to 12 longitudinal rows of hooks with 9 to 14 hooks per row with the largest hooks range from 20 to 30 mm long in the male and 31 mm long in the female. Because of horizontal markings on the worm, there is the appearance of segmentation. The lemnisci are between 4.00 and 7.50 mm by 0.15 and 0.17 mm. Males have testes that are between 2.20 and 3.00 mm by 0.50 to 0.80 mm, copulatory bursas used to hold on to the female during copulation, and eight cement glands that occupy an area of between 0.6 and 1.26 mm by 0.40 to 0.60 mm. Females have floating ovaries within a ligament sac where fertilization of the eggs, which are oval in shape with a thick, clear outer coat measuring between 0.11 and 0.12 mm long by 0.04 and 0.06 mm wide, occurs. Another report states the eggs also have 4 membranes and are 75 μm long and have an elongation ratio of 2.42. In the life cycle of ''M. moniliformis'', the intermediate hosts ingest the eggs of the parasite. In the intermediate host, the acanthor, or the parasite in its first larval stage, morphs into the acanthella, the second larval stage. After 6 to 12 weeks in this stage, the acanthella becomes a cystacanth. The cystacanth, or infective acanthella, of ''M. moniliformis'' are cyst-shaped and encyst in the tissues of the intermediate hosts in contrast to most other acanthocephalans which have infective larvae that more closely resemble underdeveloped adult worms. The definitive hosts consume the cystacanths upon feeding on infested intermediate hosts. These cystacanths mature and mate in the small intestine in 8 to 12 weeks. After this time, the eggs are excreted with the feces, to be ingested yet again by another intermediate host and renew this cycle. In what is commonly known as "brain-jacking," the parasite induces a behavioural change in its intermediate host that increases the risk of predation for the host. It is thought that this behavioral change holds an evolutionary advantage for the parasite by increasing its chances of getting to its definitive host. When ''M. moniliformis'' infests its intermediate host, the cockroach species ''
Periplaneta americana The American cockroach (''Periplaneta americana'') is the largest species of common cockroach, and often considered a pest. In certain regions of the U.S. it is colloquially known as the waterbug, though it is not a true waterbug since it is not ...
'', it changes the cockroach's escape response. In one study, it was concluded that cockroaches infected by ''M. moniliformis'' took longer to respond to wind stimuli simulating the approach of a potential predator and displayed fewer escape responses implying that the parasite infection renders its intermediate host more vulnerable to predation by hindering its ability to detect and escape from its predator. It is thought that serotonin plays a role in upending the communication between giant interneurons and the thoracic interneurons and in turn hampers the escape response of the cockroach. In a similar study, the effects of parasitism on three ''
Periplaneta ''Periplaneta'' is a genus of cockroaches containing some of the well-known pest species with cosmopolitan distributions, such as: * ''Periplaneta americana'' (Linnaeus, 1758) – American cockroach * ''Periplaneta australasiae'' (Fabricius, 1775 ...
'' species found that '' Periplaneta australasiae'' uses substrates differently and moves around less when infected with ''M. moniliformis''. Another study concludes an increased vulnerability of infected ''Periplaneta americana'' due to increased
phototaxis Phototaxis is a kind of taxis, or locomotory movement, that occurs when a whole organism moves towards or away from a stimulus of light. This is advantageous for phototrophic organisms as they can orient themselves most efficiently to receive ...
, more time spent moving (due to slower movement) and movement in response to light (uninfected cockroaches hesitated before moving). Cases of moniliformiasis, a form of acanthocephaliasis, caused by human infestation by ''M. moniliformis'' have been reported in the United States, Iran, Iraq, and Nigeria, but human moniliformiasis is now rare as it requires consumption of raw infested beetles or cockroaches. Calandruccio provided the first description of the clinical manifestations of moniliformiasis and similar accounts are found in the few case studies since; many of the patients described were asymptomatic. When they showed symptoms, they normally experienced abdominal pain, diarrhea, dizziness, edema, and anorexia. In some patients, giddiness has also been reported. In rodents, moniliformiasis is fatal and manifests itself through hemorrhaging and gastrointestinal disturbance. *''Moniliformis monoechinus'' (Linstow, 1902) ''M. monoechinus'' was found infesting the
Giant anteater The giant anteater (''Myrmecophaga tridactyla'') is an Insectivore, insectivorous mammal native to Central America, Central and South America. It is the largest of the four living species of anteaters, which are classified with sloths in the or ...
(''Myrmecophaga tridactyla'') in South America. The proboscis is very long measuring 3.55 mm long by 0.71 mm wide with 12 rows of 40 hooks each. The largest hooks are 130 μm long and found anteriorly and are shorter, 30 μm long, on the posterior portion. *''Moniliformis necromysi'' Gomes, Costa, Gentile, Vilela and Maldonado, 2020 ''M. necromysi'' has been found infesting the
hairy-tailed bolo mouse The hairy-tailed bolo mouse or hairy-tailed akodont (''Necromys lasiurus'') is a South American rodent species of the family Cricetidae. It is found in Argentina, Bolivia, Brazil, and Paraguay. Description The hairy-tailed bolo mouse grows to a ...
(''Necromys lasiurus'') in the
cerrado The Cerrado () is a vast ecoregion of Tropics, tropical savanna in central Brazil, being present in the states of Goiás, Mato Grosso do Sul, Mato Grosso, Tocantins, Maranhão, Piauí, Bahia, Minas Gerais, São Paulo (state), São Paulo, Paraná ...
in the state of
Minas Gerais Minas Gerais () is one of the 27 federative units of Brazil, being the fourth largest state by area and the second largest in number of inhabitants with a population of 20,539,989 according to the 2022 Brazilian census, 2022 census. Located in ...
,
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
.
Phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analyses using the partial mitochondrial
cytochrome c oxidase subunit I Cytochrome c oxidase I (COX1) also known as mitochondrially encoded cytochrome c oxidase I (MT-CO1) is a protein that is encoded by the ''MT-CO1'' gene in eukaryotes. The gene is also called ''COX1'', ''CO1'', or ''COI''. Cytochrome c oxidase ...
gene and the nuclear large subunit ribosomal RNA gene demonstrated that this worm forms a well-supported monophyletic group within the genus ''Moniliformis''. The species name ''necromysi'' derives from the genus name of the host species ''Necromys''. *''Moniliformis saudi'' Amin, Heckmann, Mohammed, & Evans, 2016 ''M. saudi'' has been found parasitizing the
desert hedgehog The desert hedgehog (''Paraechinus aethiopicus'') is a species of mammal in the family Erinaceidae. Description The desert hedgehog is one of the smallest hedgehogs. It is long and weighs about . The spines on its back can be banded with col ...
(''Paraechinus aethiopicus'') in Saudi Arabia. The species was named after the country
Saudi Arabia Saudi Arabia, officially the Kingdom of Saudi Arabia (KSA), is a country in West Asia. Located in the centre of the Middle East, it covers the bulk of the Arabian Peninsula and has a land area of about , making it the List of Asian countries ...
in which it was discovered. ''M. saudi'' is morphologically identical to ''M. cryptosaudi'', apart from the metal content of the hooks and genetic analysis. The trunk is between 24.00 and 50.00 mm long by 0.67 and 1.15 mm wide in the male and 18.00 to 117.50 mm long by 0.60 to 1.82 mm wide in the much larger female. The anterior trunk tapers in a fan-shaped pattern and has a collar of vertical plates. The proboscis is between 315 and 520 μm long by 130 and 177 μm wide in the male and 354 to 468 μm long by 130 to 208 μm wide in the female. The proboscis is armed with 13 to 14 rows of hooks each containing 7 or 8 hooks with the largest being found anteriorly and measuring 25 to 31 μm long in the female and 25 to 27 μm long in the male. The ventral hooks longer than dorsal hooks. The proboscis receptacle is 718 to 1250 μm by 155 to 322 μm in the female and 603 to 880 μm by 190 to 300 μm in the male. The long lemnisci and measures 2.91 to 8.75 mm by 0.08 to 0.17 mm in the male and measures 3.30 to 12.50 mm by 0.11 to 0.22 mm in the female. The short lemnisci and measures 2.81 to 5.75 mm by 0.07 to 0.14 mm in the male and measures 3.02 to 11.25 mm by 0.10 to 0.17 mm in the female. There are has 7 to 10 giant nuclei in the lemnisci. The anterior testes are 1.25 to 3.50 mm by 0.27 to 0.82 mm in size and the posterior testes are 1.12 to 3.50 mm by 0.27 to 0.65 mm in size. In the male, the cement glands are 312 to 811 μm by 270 to 468 μm and Saefftigen's pouch measures 468 to 1400 μm by 177 to 450 μm. In the female, The eggs are oval, measuring 57 to 83 μm by 31 to 42 μm. *''Moniliformis siciliensis'' Meyer, 1932 ''M. siciliensis'' has been found infesting the
brown rat The brown rat (''Rattus norvegicus''), also known as the common rat, street rat, sewer rat, wharf rat, Hanover rat, Norway rat and Norwegian rat, is a widespread species of common rat. One of the largest Muroidea, muroids, it is a brown or grey ...
(''Rattus norvegicus'' reported as ''Mus decumanus''), and the
garden dormouse The garden dormouse (''Eliomys quercinus'') is a species of dormouse native to Europe. Characteristics The garden dormouse is gray or brown, with a whitish underside. It has black eye markings and large ears. Its hair is short, and its tail ha ...
(''Eliomys quercinus'') in
Sicily Sicily (Italian language, Italian and ), officially the Sicilian Region (), is an island in the central Mediterranean Sea, south of the Italian Peninsula in continental Europe and is one of the 20 regions of Italy, regions of Italy. With 4. ...
,
Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...
. The male is between 40 and 45 mm long by 1.0 and 1.50 mm wide whereas the female is much larger between 70 and 80 mm long by 1.0 and 1.50 mm wide. The proboscis is between 0.42 and 0.45 mm long by 0.17 to 0.19 mm wide in the male and female. The proboscis has 14 longitudinal rows of hooks with 8 hooks per row. The lemnisci are 10.0 by 0.17 mm. The eggs are oval being 0.08 mm long by 0.04 mm wide. *''Moniliformis spiralis'' Subrahmanian, 1927 ''M. spiralis'' been found infesting '' Nesokia bengalensis'' and the
black rat The black rat (''Rattus rattus''), also known as the roof rat, ship rat, or house rat, is a common long-tailed rodent of the stereotypical rat genus ''Rattus'', in the subfamily Murinae. It likely originated in the Indian subcontinent, but is n ...
(''Rattus rattus'') in
Rangoon Yangon, formerly romanized as Rangoon, is the capital of the Yangon Region and the largest city of Myanmar. Yangon was the List of capitals of Myanmar, capital of Myanmar until 2005 and served as such until 2006, when the State Peace and Dev ...
,
Burma Myanmar, officially the Republic of the Union of Myanmar; and also referred to as Burma (the official English name until 1989), is a country in northwest Southeast Asia. It is the largest country by area in Mainland Southeast Asia and ha ...
. It has also been found in
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
infesting a species of
Bandicota ''Bandicota'' is a genus of rodents from Asia known as the bandicoot rats. Their common name and genus name are derived from the Telugu language word ''pandikokku'' (పందికొక్కు). DNA studies have found the group to be a monop ...
and
Herpestes ''Herpestes'' is a genus within the mongoose family Herpestidae. Several species in the family are known as slender mongooses. It is the type genus of the family, and comprises 5-6 living species, each with several subspecies. Fossil remains of t ...
, and in bats in
Kyrgyzstan Kyrgyzstan, officially the Kyrgyz Republic, is a landlocked country in Central Asia lying in the Tian Shan and Pamir Mountains, Pamir mountain ranges. Bishkek is the Capital city, capital and List of cities in Kyrgyzstan, largest city. Kyrgyz ...
. The male is 16.94 mm long and 0.56 mm wide whereas the female is much larger at 40 mm long and 0.91 mm wide. The proboscis of the male is 0.43 mm long and 0.11 mm wide and in the female is between 0.35 and 0.47 mm long and 0.17 to 0.18 mm wide. The proboscis receptacle of the male is 0.64 to 0.85 mm long and 0.24 to 0.30 mm wide and in the female it is 0.98 mm long and 0.31 mm wide. The proboscis has 12 to 14 longitudinal rows of hooks with 12 to 14 hooks per row with the largest hooks being 40 mm long. The lemnisci are 0.81 by 0.25 mm. The anterior testes are 0.81 by 0.25 mm and the posterior testes are 0.85 by 0.28 mm. There are six to eight cement glands 0.53 by 0.21 mm in size. The eggs are oval being 0.06 mm long by 0.03 mm wide and the genital pore is terminal and 0.91 by 0.48 mm. *''Moniliformis tarsii'' Deveaux, Schmidt and Krishnasamy, 1988 ''M. tarsii'' was found infesting the small and large intestines of Horsfield's tarsier (''Tarsius bancanus'') in
Poring Poring is a small tourist resort in Sabah, Malaysia. Located 40 km south-east of the Kinabalu National Park Headquarters, in the district of Ranau, Poring is situated in lowland rainforest, contrasting with the montane and submontane rainfor ...
,
Sabah Sabah () is a States and federal territories of Malaysia, state of Malaysia located in northern Borneo, in the region of East Malaysia. Sabah has land borders with the Malaysian state of Sarawak to the southwest and Indonesia's North Kalima ...
,
Malaysia Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
. The male is 15 to 19 mm long and 800 to 900 μm wide whereas the female is larger being 21 to 41 mm long and 0.73 to 1.57 mm wide with a trunk lacking pseudosegmentation. The proboscis in the male is 490 μm long and 240 μm wide and in the female is 490 to 560 μm long and 230 to 320 μm wide. The proboscis is armed with 11 or 12 regularly alternating longitudinal rows of 6 or 7 hooks with the intermediate hooks being considerably larger than the others. Anterior hooks measure 26 to 37 μm, the next two hooks measure 41 to 55 μm, all three of which have powerful roots, whereas the remaining hooks four through seven measure 24 to 28 μm and have weak roots. The lemnsici are flat with three giant nuclei each and reach about 25–30% of the body length and are not bound posteriorly to the inner body wall. In the male, the anterior testes are 1.66 to 1.88 mm long and 330 μm wide, and the posterior testes are 1.5 mm long and 430 μm wide. The males also have eight cement glands each with a single giant nucleus located posterior to the testes. In the female, the uterine bell is 1.03 mm from posterior and eggs are very small and oval, being 47 to 55 μm long and 16 to 21 μm wide. The species name ''tarsii'' derives from the genus name of the host ''Tarsius''. *''Moniliformis travassosi'' Meyer, 1932 ''M. travassosi'' was found infesting the
brown rat The brown rat (''Rattus norvegicus''), also known as the common rat, street rat, sewer rat, wharf rat, Hanover rat, Norway rat and Norwegian rat, is a widespread species of common rat. One of the largest Muroidea, muroids, it is a brown or grey ...
(''Rattus norvegicus'') in
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
. The male measures between 60 and 80 mm long by 1.0 and 1.5 mm wide whereas the female is between 100 and 110 mm long and 1.5 mm wide. The proboscis has 14 longitudinal rows of hooks with 15 hooks per row with the largest hooks being 24 to 28 mm long. The lemnisci are 10.0 mm. In the male, the testes are 2.50 to 3.0 mm by 0.80 mm and there are eight cement glands. The eggs are oval being 0.12 to 0.12 mm long by 0.07 to 0.07 mm wide. It is named after
Lauro Travassos Lauro Pereira Travassos (2 July 1890 – 20 November 1970) was a Brazilian parasitologist. He collected helminth specimens from across South America and studied their life-histories. He built a Brazilian school of parasitology through his students ...
, a Brazilian helminthologist and entomologist who named the genus ''Moniliformis'' to which this species belongs. *''Moniliformis tupaia'' Chen, Yu, Ma, Zhao, Cao and Li, 2024


Hosts

The life cycle of an acanthocephalan consists of three stages beginning when an infective acanthor (development of an egg) is released from the intestines of the
definitive host In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include ...
and then ingested by an arthropod, the
intermediate host In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include ...
. The intermediate hosts of ''Moniliformis'' are usually insects. When the acanthor
molt In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is a process by which an animal casts off parts of its body to serve some beneficial purpose, either at ...
s, the second stage called the acanthella begins. This stage involves penetrating the wall of the
mesenteron The midgut is the portion of the human embryo from which almost all of the small intestine and approximately half of the large intestine develop. After it bends around the superior mesenteric artery, it is called the "midgut loop". It comprises t ...
or the intestine of the intermediate host and growing. The final stage is the infective cystacanth which is the
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
l or juvenile state of an Acanthocephalan, differing from the adult only in size and stage of sexual development. The cystacanths within the intermediate hosts are consumed by the definitive host, usually attaching to the walls of the intestines, and as adults they reproduce sexually in the intestines. The acanthor are passed in the
feces Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the ...
of the definitive host and the cycle repeats. There are no known paratenic hosts (hosts where parasites infest but do not undergo larval development or sexual reproduction) for ''Moniliformis''. ''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 A lesion is any damage or abnormal change in the tissue of an organism, usually caused by injury or diseases. The term ''Lesion'' is derived from the Latin meaning "injury". Lesions may occur in both plants and animals. Types There is no de ...
in the intestines, intestinal distension,
perforated ulcer A perforated ulcer is a condition in which an untreated ulcer has burned through the mucosal wall in a segment of the gastrointestinal tract (e.g., the stomach or colon) allowing gastric contents to leak into the abdominal cavity. Signs and symp ...
s,
enteritis Enteritis is inflammation of the small intestine. It is most commonly caused by food or drink contaminated with pathogenic microbes,Dugdale, David C., IIII, and George F Longretc"Enteritis" MedlinePlus Medical Encyclopedia, 18 October 2008. Acces ...
,
crypt A crypt (from Greek κρύπτη (kryptē) ''wikt:crypta#Latin, crypta'' "Burial vault (tomb), vault") is a stone chamber beneath the floor of a church or other building. It typically contains coffins, Sarcophagus, sarcophagi, or Relic, religiou ...
hypertrophy Hypertrophy is the increase in the volume of an organ or tissue due to the enlargement of its component cells. It is distinguished from hyperplasia, in which the cells remain approximately the same size but increase in number. Although hypertro ...
,
goblet cell Goblet cells are simple columnar epithelial cells that secrete gel-forming mucins, like mucin 2 in the lower gastrointestinal tract, and mucin 5AC in the respiratory tract. The goblet cells mainly use the merocrine method of secretion, secre ...
hyperplasia Hyperplasia (from ancient Greek ὑπέρ ''huper'' 'over' + πλάσις ''plasis'' 'formation'), or hypergenesis, is an enlargement of an organ or tissue caused by an increase in the amount of Tissue (biology), organic tissue that results from ...
, and occlusions of the intestinal tract in the
gray squirrel Gray squirrel or grey squirrel may refer to several species of squirrel indigenous to North America: *The eastern gray squirrel (''Sciurus carolinensis''), from the eastern United States and southeastern Canada; introduced into the United Kingdom, ...
(''Sciurus carolinensis''). There are multiple reports of human infection from Australia, Iran, Iraq, Nigeria, Zimbabwe, Saudi Arabia and United States, all from ''Moniliformis moniliformis''. File:Common spiny mouse.JPG, alt=A Cairo spiny mouse on dirt, The
Cairo spiny mouse The Cairo spiny mouse (''Acomys cahirinus''), also known as the common spiny mouse, Egyptian spiny mouse, or Arabian spiny mouse, is a Nocturnality, nocturnal species of rodent in the family Muridae. It is found in Africa north of the Sahara De ...
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 The four-toed jerboa (''Scarturus tetradactylus'') is a rodent of the family Dipodidae and genus ''Scarturus'' that has four digits. Four-toed jerboas are native to Egypt and Libya. They live in coastal salt marshes and dry deserts. Physical a ...
is a host of ''Moniliformis aegyptiacus'' File:Atelerix algirus.jpg, alt=A North African hedgehog curled up on hay, The
North African hedgehog The North African hedgehog (''Atelerix algirus'') or Algerian hedgehog, is a mammal species in the family Erinaceidae native to Algeria, Libya, Malta, Morocco, Spain, and Tunisia. Little is known about this hedgehog, even though the most common b ...
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 The cotton-top tamarin (''Saguinus oedipus'') is a small New World monkey weighing less than . This New World monkey can live up to 24 years, but most of them die by 13 years. One of the smallest primates, the cotton-top tamarin is easily recog ...
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 The large flying fox (''Pteropus vampyrus'', formerly ''Pteropus giganteus''), also known as the greater flying fox, Malayan flying fox, Malaysian flying fox, large fruit bat, kalang, or kalong, is a southeast Asian species of megabat in the fam ...
is a host of ''Moniliformis convolutus'' File:Hemiechinus auritus; Baikonur 09.jpg, alt=Long-eared hedgehog on a wood floor in front of a wood door, The
long-eared hedgehog The long-eared hedgehog (''Hemiechinus auritus'') is a species of hedgehog native to Central Asian countries and some countries of the Middle East. The long-eared hedgehog lives in burrows that it either makes or finds and is distinguished by it ...
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 The giant anteater (''Myrmecophaga tridactyla'') is an Insectivore, insectivorous mammal native to Central America, Central and South America. It is the largest of the four living species of anteaters, which are classified with sloths in the or ...
is a host of ''Moniliformis monoechinus'' File:Desert Hedgehog.JPG, alt=A Desert hedgehog under branches, The
desert hedgehog The desert hedgehog (''Paraechinus aethiopicus'') is a species of mammal in the family Erinaceidae. Description The desert hedgehog is one of the smallest hedgehogs. It is long and weighs about . The spines on its back can be banded with col ...
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 The brown rat (''Rattus norvegicus''), also known as the common rat, street rat, sewer rat, wharf rat, Hanover rat, Norway rat and Norwegian rat, is a widespread species of common rat. One of the largest Muroidea, muroids, it is a brown or grey ...
is a host of ''M. siciliensis'' and ''Moniliformis travassosi'' File:Eliomys quercinus01.jpg, alt=A Garden dormouse on a stone, The
garden dormouse The garden dormouse (''Eliomys quercinus'') is a species of dormouse native to Europe. Characteristics The garden dormouse is gray or brown, with a whitish underside. It has black eye markings and large ears. Its hair is short, and its tail ha ...
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 The black rat (''Rattus rattus''), also known as the roof rat, ship rat, or house rat, is a common long-tailed rodent of the stereotypical rat genus ''Rattus'', in the subfamily Murinae. It likely originated in the Indian subcontinent, but is n ...
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''


Notes


References

{{Taxonbar, from=Q2782820 Archiacanthocephala Acanthocephala genera