Acanthocheilonema
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''Acanthocheilonema'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
within the family
Onchocercidae The Onchocercidae are a family of nematodes in the superfamily Filarioidea. This family includes some of the most devastating human parasitic diseases, such as lymphatic filariasis, onchocerciasis, loiasis, and other filariases. Representativ ...
which comprises mainly tropical
parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has c ...
worms. Cobbold created the genus ''Acanthocheilonema'' with only one species, ''Acanthocheilonema dracunculoides'', which was collected from aardwolf (''Proteles cristatus'': Hyaenidae) in the region of South Africa in the nineteenth century. These parasites have a wide range of mammalian species as hosts, including members of
Carnivora Carnivora is a Clade, monophyletic order of Placentalia, placental mammals consisting of the most recent common ancestor of all felidae, cat-like and canidae, dog-like animals, and all descendants of that ancestor. Members of this group are f ...
, Macroscelidea,
Rodentia Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are nat ...
,
Pholidota Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (, from Ancient Greek ϕολιδωτός – "clad in scales"). The one extant family, the Manidae, has three genera: '' Manis'', ''Phataginus'', and ''Smutsia ...
,
Edentata Xenarthra (; from Ancient Greek ξένος, xénos, "foreign, alien" + ἄρθρον, árthron, "joint") is a major clade of placental mammals native to the Americas. There are 31 living species: the anteaters, tree sloths, and armadillos. Extin ...
, and
Marsupialia Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a pou ...
. Many species among several genera of filarioids exhibit a high degree of endemicity in studies done on mammalian species in Japan. However, no concrete evidence has confirmed any endemic species in the genus ''Acanthocheilonema''. There are about fifteen well characterized parasitic species found in genus ''Acanthocheilonema''. ''A. delicata'' n. sp. (2013), ''A. dracunculoides'' (Cobbold 1870), ''A. filaria'' (Kou, 1958), ''A. mansonbahri'' (Nelson, 1961), ''A. mephitis'' (Webster and Beauregard, 1964), ''A. odendhali'' (Perry, 1967), ''A. pachycephalum'' (Ortlepp, 1961), ''A. pricei'' (Vaz and Pereira, 1934), ''A. procyonis'' (Price, 1955), ''A. reconditum'' (Grassi, 1889), ''A. sabanicolae'' (Eberhard and Campo-Aasen, 1986), ''A. setariosa'' (Mönnig, 1926), ''A. spirocauda'' (Leidy, 1858), ''A. viteae'' (Krepkogorskaja, 1933), and ''A. weissi'' (Seurat, 1914) are the well characterized parasitic species found all around the world in variety of hosts. Some of the crucial parasites that affect a wide range of host species are discussed below.


''Acanthocheilonema delicata''

''
Acanthocheilonema delicata ''Acanthocheilonema'' is a genus within the family Onchocercidae which comprises mainly tropical parasitic worms. Cobbold created the genus ''Acanthocheilonema'' with only one species, ''Acanthocheilonema dracunculoides'', which was collected f ...
'' was discovered and characterized from an
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
badger species (''Meles anakuma'') in Japan. The researchers identified the DNA sequences of mitochondrial cytochrome c oxidase subunit 1 (cox1) gene in the Japanese badgers. Morphologically, filarioids analyzed were identified as thin, small and delicate and in females the length was longer (twice as in males). Anterior of the filarioids is slightly bulbous having two sets of four papilla (worms) and amphids. They possess distinct buccal cavity with buccal capsule consisting of a thick buccal ring. The esophagus is divided into short anterior muscular portion and long, posterior glandular portion. Caudal extremity in both males and females consists of three conical lappets. The location of the parasitic adult worms in the host species (''Meles anakuma'') was found to be the subcutaneous
connective tissue Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...
, whereas the
microfilaria ::''Microfilaria may also refer to an informal "collective group" genus name, proposed by Cobbold in 1882. While a convenient category for newly discovered microfilariae which can not be assigned to a known species because the adults are unknown, ...
was found in the skin. Data taken from the
Genbank The GenBank sequence database is an open access, annotated collection of all publicly available nucleotide sequences and their protein translations. It is produced and maintained by the National Center for Biotechnology Information (NCBI; a part ...
of fourteen species of genus ''Acanthocheilonema'' compared to the newly found species for possible similarities and distinctions shown in the table below:


''Acanthocheilonema dracunculoides''

''
Acanthocheilonema dracunculoides ''Acanthocheilonema'' is a genus within the family Onchocercidae which comprises mainly tropical parasitic worms. Cobbold created the genus ''Acanthocheilonema'' with only one species, ''Acanthocheilonema dracunculoides'', which was collected f ...
'' Cobbold, a species discovered in 1870, is a nematode parasitic worm particularly found in domestic dogs and other carnivores like aardwolf, spotted hyaena and red fox. These parasites can be located on various continents like Europe, Asia and Africa. it is known to be endemic in Morocco, Algeria, Tunisia, Mali, Niger, Democratic Republic of Congo, Sudan, Somalia, Kenya, Tanzania and South Africa. This parasite follows an indirect life cycle. In the
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
species, the male and female parasite tend to be biased to exist mainly in the peritoneal cavity. Males are typically shorter and are 15–32 mm long and 0.1–0.2 mm wide, whereas females are almost double, standing at 30–60 mm long and 0.1–0.3 mm wide in diameter. Sexually mature female parasites of ''A. dracunculoides'' are
viviparous Among animals, viviparity is development of the embryo inside the body of the parent. This is opposed to oviparity which is a reproductive mode in which females lay developing eggs that complete their development and hatch externally from the m ...
, they produce L1-stages, known as microfilariae, which eventually appear in the peripheral blood. Microfilariae are unsheathed and measure 185–276 μm (length) by 4.2–6 μm (width) Although the final host is the carnivores for ''A. dracunculoides'' there are some other intermediate hosts that are involved in the parasite life cycle. Louse fly ''Hippobosca longipennis'' and the hard tick ''
Rhipicephalus sanguineus ''Rhipicephalus sanguineus'', commonly called the brown dog tick, kennel tick, or pantropical dog tick, is a species of tick found worldwide, but more commonly in warmer climates. This species is unusual among ticks in that its entire lifecycle c ...
'', also known as dog tick, have been identified as intermediate hosts that helps the parasite to complete its life cycle. Although ''Acanthocheilonema dracunculoides'' has a reputation of not being an endemic parasite in the domestic dogs, recent evidence of the presence of the parasite in domestic animals show some signs of pathogenicity. As mostly these parasites are considered innocuous in the dogs, recent studies and evidence from Spain suggests that the parasite is not completely harmless and innocuous in the domestic dogs as believed earlier. The discovery of ''Acanthocheilonema'' parasites in the non-endemic species has triggered more research in this field. ''Dirofilaria immitis'', ''D. repens'', ''Acanthocheilonema dracunculoides'' and ''A. reconditum'' are the common species that have been known to be found in canine filarial nematodes. These nematodes present blood circulating microfilariae which are differentiated to the species level using the techniques like PCR and by acid phosphatase activity patterns. ''A. reconditum'' and ''A. dracunculoides'' live in the peritoneal cavity and adipose tissue of the host species and are less pathogenic, however ''D. immitis'' and ''D. repens'' are considered to be the emerging agents of parasitic zoonoses and are continuously expanding their ranges in the European. areas.


''Acanthocheilonema odendhali''

The filarial worm ''
Acanthocheilonema odendhali ''Acanthocheilonema'' is a genus within the family Onchocercidae which comprises mainly tropical parasitic worms. Cobbold created the genus ''Acanthocheilonema'' with only one species, ''Acanthocheilonema dracunculoides'', which was collected f ...
'' was first discovered in 1967 from the California
California sea lion The California sea lion (''Zalophus californianus'') is a coastal eared seal native to western North America. It is one of six species of sea lions. Its natural habitat ranges from southeast Alaska to central Mexico, including the Gulf of C ...
''
Zalophus californianus The California sea lion (''Zalophus californianus'') is a coastal eared seal native to western North America. It is one of six species of sea lions. Its natural habitat ranges from southeast Alaska to central Mexico, including the Gulf of Cali ...
''. This species was found inhabiting the intermuscular fascia and thoracic and the abdominal cavities. Initially this species was described in the ''Dipetalonema'' genus as ''Dipetalonema odendhali'', however due to later evidences it was transferred to the genus ''Acanthocheilonema''. ''A. odendhali'' was subsequently found in other marine species like Steller sea lion (''Eumetopias jubatus'') and northern fur seal (''Callorhinus ursinus''). Besides the ''A. odendhali'', other filarioidean species such as ''A. spirocauda'', commonly known as heart-worm has also been identified as parasitic in phocids. ''A. odendhali'' was primarily reported in the northern fur seals on the island of Pribilof Archipelago in 1967. Prior to that two scientists Mark C. Keyes and Eugene T. Lyons had discovered the
microfilariae ::''Microfilaria may also refer to an informal "collective group" genus name, proposed by Cobbold in 1882. While a convenient category for newly discovered microfilariae which can not be assigned to a known species because the adults are unknown, ...
in the blood of subadult male seals and adult filarioids and microfilariae in the samples collected from the
blubber Blubber is a thick layer of vascularized adipose tissue under the skin of all cetaceans, pinnipeds, penguins, and sirenians. Description Lipid-rich, collagen fiber-laced blubber comprises the hypodermis and covers the whole body, except for pa ...
, fur, and the skin from these male seals. In these studies, the prevalence and the intensity of these species were not examined. The life cycle of ''A. odendhali'' is still unknown. However, according to the National Marine Mammal Laboratory, Northwest and Alaska Fisheries Center (NWAFC) reports, some blood-sucking flies or insects that infest the wounds are potential intermediate hosts (vectors) in their life cycle. Pathogenicity of ''A. odendhali'' was however not studied in these experiments and research but this parasite is considered as non-pathogenic. In 2011 and 2012, a study that investigated
helminths Parasitic worms, also known as helminths, are large macroparasites; adults can generally be seen with the naked eye. Many are intestinal worms that are soil-transmitted and infect the gastrointestinal tract. Other parasitic worms such as schi ...
of northern fur seals on Saint Paul Island in Alaska discovered that ''A. odendhali'' was a parasite on the seals. Other research done by the same group investigated the intensity and prevalence of the parasite and discovered that prevalence of adult ''A. odendhali'' parasites in the fur seals was 18 percent; the intensity depends on the individual host species and the seven parasite specimens found in the parasite. Average intensity of infection was approximately 1.32 ±0.83 SD, whereas the abundance was found to be 0.24 SD.


''Acanthocheilonema reconditum''

''
Acanthocheilonema reconditum ''Acanthocheilonema'' is a genus within the family Onchocercidae which comprises mainly tropical parasitic worms. Cobbold created the genus ''Acanthocheilonema'' with only one species, ''Acanthocheilonema dracunculoides'', which was collected f ...
'' is a non-pathogenic canine
helminth Parasitic worms, also known as helminths, are large macroparasites; adults can generally be seen with the naked eye. Many are intestinal worms that are soil-transmitted and infect the gastrointestinal tract. Other parasitic worms such as schi ...
parasite of the specialized
connective tissue Connective tissue is one of the four primary types of animal tissue, along with epithelial tissue, muscle tissue, and nervous tissue. It develops from the mesenchyme derived from the mesoderm the middle embryonic germ layer. Connective tiss ...
. It is distributed in the same endemic areas as D. immitis, which is also known as heart-worm. Although it is endemic to some areas mentioned earlier, but the parasite is widely distributed as it has been identified in other species and around different continents. Dogs are frequently parasitized by the filarids. This transmission is usually carried out by the flea species and lice species. Flea vectors include species such as ''
Ctenocephalides canis The dog flea (''Ctenocephalides canis'') is a species of flea that lives as an ectoparasite on a wide variety of mammals, particularly the domestic dog and cat. It closely resembles the cat flea, ''Ctenocephalides felis'', which can live on a wid ...
'', ''
Pulex irritans The human flea (''Pulex irritans'') – once also called the house flea – is a cosmopolitan flea species that has, in spite of the common name, a wide host spectrum. It is one of six species in the genus '' Pulex''; the other five are all confi ...
'', ''Pulex simulans'', and ''
Echidnophaga gallinacea ''Echidnophaga gallinacea'', also known as the hen flea or sticktight flea, is part of the 2,500 known flea types in the Siphonaptera order. ''Echidnophaga gallinacea'' appear dark brown in colour and is a small flea measuring approximately 2 m ...
'', whereas the lice species include Linognathus setosus and Heterodoxus spiniger. These vectors are infected with the ''A. reconditum'' parasite with the microfilariae during the blood repast in a parasitized dog. These species act as intermediate hosts in completing the life cycle of the parasite.Otranto D, Torres FT, Brianti E et al. (2013) "Vector-borne helminths of dogs and humans in Europe". ''Parasites & Vectors'' 6:1–14 These microfilariae develop onto an infectious stage during the L3 (larvae stage 3) stage in the digestive tract of these vectors from where they can be inoculated into the new host. The third stage larvae could be found as soon as 7 days after the appearance of the vectors on the donor host species. (L3) Once it enters the new host species, the microfilariae develops into the adult stage. Most of these adult stage parasites are located in the
subcutaneous tissue The subcutaneous tissue (), also called the hypodermis, hypoderm (), subcutis, superficial fascia, is the lowermost layer of the integumentary system in vertebrates. The types of cells found in the layer are fibroblasts, adipose cells, and macr ...
. Although the infection by ''A. reconditum'' does not exhibit any clinical signs, a treatment to limit
dissemination To disseminate (from lat. ''disseminare'' "scattering seeds"), in the field of communication, is to broadcast a message to the public without direct feedback from the audience. Meaning Dissemination takes on the theory of the traditional view ...
is primarily recommended. Recent studies of ''A. reconditum'' occurrence in the southern regions of Italy has shown prevalence of this parasite as high as 13.3% with an annual incidence rate of 5.9%, which was observed in naturally exposed dogs. Recent experiments have enhanced scientific knowledge of the biology and ecology of this parasite. The full development of microfilariae occurs in the experimental infected cat flea ''Ctenocephalides felis felis'' in about 15 days. Localization and size of this parasite in this infected flea suggests the possibility that this might act as an intermediate host throughout the ingestion of the infected fleas rather than inoculation of the blood meal on dogs.Napoli, E., Gaglio, G., Falsone, L. et al. "New insights into the biology and ecology of ''Acanthocheilonema reconditum'' (spirurida: onchocercidae)". ''Parasites & Vectors'' 7, O29 (2014). https://doi.org/10.1186/1756-3305-7-S1-O29
Ivermectin Ivermectin (, '' EYE-vər-MEK-tin'') is an antiparasitic drug. After its discovery in 1975, its first uses were in veterinary medicine to prevent and treat heartworm and acariasis. Approved for human use in 1987, today it is used to treat inf ...
(0.25 mg/kg) is a drug that is preferred to treat infection by this parasite. As microfilariae of this parasite are mainly found in the blood, they are often mistaken for other parasites like ''
Dirofilaria immitis ''Dirofilaria immitis'', also known as heartworm or dog heartworm, is a parasitic roundworm that is a type of filarial worm, a small thread-like worm, that causes dirofilariasis. It is spread from host to host through the bites of mosquitoes ...
''. It is therefore crucial to identify and distinguish between them using modified Knott's procedure.


''Acanthocheilonema spirocauda''

'' Acanthocheilonema spirocauda'', also known as seal heartworm, is a filarial parasite that primarily infects the phocid seals, including ''Phoca vitulina'' (Harbor seal). The Heartworm was originally described as ''Filaria spirocauda'' by Leidy. Eventually ''Acanthocheilonema'' was raised to generic level and this species settled under this genus, and a researcher Anderson presented its current name.Anderson RC (1992) ''Nematode parasites of vertebrates. Their development and transmission'', 2nd edn. CABI, Wallingford ''A. spirocauda'' is found in the heart of the seals, and that is why is got its name 'Heartworm'. While there is no significant threat to the seal populations due to this parasite as of now, however the infection with this parasite can result in pathological consequences like in the case of ''D. immitis'', which includes the characteristics such as anorexia, fatigue, heart and lung complications, and potentially death. Researchers suggest that these parasites share strong evolutionary basis with its phocid hosts about some 45 million years ago It is believed to be transmitted via the seal louse ''(Echinophthirius horridus)''. The prevalence of the parasite in the host species is approximately 8.8%, and intensity ranges up to 44 specimens per heart. Additional researchers have investigated its additional potential hosts and global distribution patterns of this parasite. Studies discovered some additional hosts of this parasite that include ribbon seal ''P. fasciata'', harp seal ''P. groenlandica'', ringed seal ''P. hispida'', Baltic ringed seal ''P. h. botnica'', Ladoga seal ''P.h. ladogensis'', spotted seal ''P. largha'', hooded seal '' Cystophora cristata'', and bearded seal ''
Erignathus barbatus The bearded seal (''Erignathus barbatus''), also called the square flipper seal, is a medium-sized pinniped that is found in and near to the Arctic Ocean. It gets its generic name from two Greek words (''eri'' and ''gnathos'') that refer to its h ...
''.Westerling B, Stenman O, Rudbäck E (2005) "Pathology of seals from the Finnish coastal waters, Lake Saimaa and Lake Ladoga in the years 1982–2004". ''Proceedings from the Symposium on Biology and Management of Seals in the Baltic area'', 15–18 February 2005, Helsinki, pp 76–79 The primary sites of infection in the hosts are the right and left ventricles of the heart, pulmonary vein, pulmonary artery, and vena cava. These parasites are widely distributed around the globe. They can be found in the Northern Hemisphere and
Holarctic realm The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical region ...
. The present localities of ''A. spirocauda'' include the Danish North Sea (Agger Tange), Limfjord (Thy), Skagerrak (Lysekil) and Kattegat (Rungsted, Lille Vrøj), and the Baltic Sea (Lunds høje, Falster).


''Acanthocheilonema viteae''

''
Acanthocheilonema viteae ''Acanthocheilonema'' is a genus within the family Onchocercidae which comprises mainly tropical parasitic worms. Cobbold created the genus ''Acanthocheilonema'' with only one species, ''Acanthocheilonema dracunculoides'', which was collected fr ...
'' is a filarial nematode that infects the rodents species. ''A. Viteae''is also used as a model species in the research experiments in order to study Human filarial infections. The natural host for this parasite are
gerbils Gerbillinae is one of the subfamilies of the rodent family Muridae and includes the gerbils, jirds, and sand rats. Once known as desert rats, the subfamily includes about 110 species of African, Indian, and Asian rodents, including sand rats and ...
, whereas the experimental hosts include hamsters, jirds ('' Meriones unguiculatus'', also known as Mongolian gerbils), and ''
Mastomys ''Mastomys'' is a genus of rodent in the family Muridae endemic to Africa. It contains eight species: * Angolan multimammate mouse (''M. angolensis'') * Awash multimammate mouse or Awash mastomys (''M. awashensis'') * Southern multimammate mou ...
'' species. The vector that aids to finish its life cycle is a soft tick named ''Ornithodoros tartakovskyi'' (natural) and '' Ornithodoros moubata'' (experimental vector). The adults of ''A. Viteae'' resides in the deep
subcutaneous tissue The subcutaneous tissue (), also called the hypodermis, hypoderm (), subcutis, superficial fascia, is the lowermost layer of the integumentary system in vertebrates. The types of cells found in the layer are fibroblasts, adipose cells, and macr ...
, whereas the
microfilariae ::''Microfilaria may also refer to an informal "collective group" genus name, proposed by Cobbold in 1882. While a convenient category for newly discovered microfilariae which can not be assigned to a known species because the adults are unknown, ...
circulate in the blood. These worms are known to modulate the host immunological systems, however they do not prohibit the immunological pathways entirely. It renders them the increased survival. An excretory product called ES-62 is secreted by ''A. Viteae'' that is homologous to the one produced by the humans but not parasitic. ES-62 molecules tend to exhibit anti-inflammatory actions using phosphorycholine moiety to affect the intracellular pathways that are associated with the antigen receptors and TLR dependent cell responses. In nature, during the third larvae infected stage, ''A. Viteae'' parasites are transferred to their mammalian hosts with the aid of vector ''Ornithodoros tartakovskyi''. In the mammalian host, the L3 stage develops to mature adults and generates the microfilariae, which are again ingested by ticks. After reaching L3 parasites transfers to mammalian hosts via arthropods' mouth parts. Various stages of ''A. Viteae'' have two chitinase protein bands, one of 205 kDa and 68 Kda. Studies reveal that the 205 kDa band of stage L3 was an oligomer formed of 68kDa monomers. Chitin has been reported as a component of the eggshells of these nematodes Chitinase plays a vital role in the third larvae stage (L3) and female worms, as its inhibition during these stages leads to have increased mortality. There are no safe and efficient drugs to eliminate these infections so far but chitin is seen as a potential target for the development of
anthelmintic Anthelmintics or antihelminthics are a group of antiparasitic drugs that expel parasitic worms (helminths) and other internal parasites from the body by either stunning or killing them and without causing significant damage to the host. They may a ...
drugs and vaccines. Intracellular bacteria ''Wolbachia'' are prevalent among the different species of nematodes. However, ''Wolbachia'' is absent in ''Acanthocheilonema viteae''. Since ''A. Viteae'' lacks ''Wolbachia'', it is widely used as a negative control for experiments investigating the bacterium. Their role in the pathology of filarial infections and symbiotic nature with host makes it potential target to intervene in filarial infections in the humans.


References

{{Taxonbar, from=Q16747049 Spirurida Parasitic nematodes of mammals