Apororhynchus silesiacus
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''Apororhynchus'' is a genus of small parasitic spiny-headed (or thorny-headed) worms. It is the only
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
in the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Apororhynchidae, which in turn is the only member of the order Apororhynchida. A lack of features commonly found in the phylum
Acanthocephala Acanthocephala (Greek , ', thorn + , ', head) is a phylum 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 to p ...
(primarily musculature) suggests an evolutionary branching from the other three orders of class
Archiacanthocephala Archiacanthocephala is a class within the phylum of Acanthocephala. They are parasitic worms that attach themselves to the intestinal wall of terrestrial vertebrates, including humans. They are characterised by the body wall and the lemnisci (wh ...
; however no genetic analysis has been completed to determine the evolutionary relationship between species. The distinguishing features of this order among
archiacanthocephala Archiacanthocephala is a class within the phylum of Acanthocephala. They are parasitic worms that attach themselves to the intestinal wall of terrestrial vertebrates, including humans. They are characterised by the body wall and the lemnisci (wh ...
ns is a highly enlarged
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
which contain small hooks. The musculature around the proboscis (the proboscis receptacle and receptacle protrusor) is also structured differently in this order. This genus contains six species that are distributed globally, being collected sporadically in Hawaii, Europe, North America, South America, and Asia. These worms exclusively parasitize birds by attaching themselves around the cloaca using their hook-covered
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
. The bird
hosts A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places *Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman *Michel Host ( ...
are of different orders, including
owl Owls are birds from the order Strigiformes (), which includes over 200 species of mostly solitary and nocturnal birds of prey typified by an upright stance, a large, broad head, binocular vision, binaural hearing, sharp talons, and feathers a ...
s,
wader 245px, A flock of Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflat ...
s, and
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
s.
Infestation Infestation is the state of being invaded or overrun by pests or parasites. It can also refer to the actual organisms living on or within a host. Terminology In general, the term "infestation" refers to parasitic diseases caused by animals s ...
by an ''Apororhynchus'' species may cause
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. Access ...
and
anemia Anemia or anaemia (British English) is a blood disorder in which the blood has a reduced ability to carry oxygen due to a lower than normal number of red blood cells, or a reduction in the amount of hemoglobin. When anemia comes on slowly, t ...
.


Taxonomy

The first species in this order to be described was ''Apororhynchus hemignathi'' which was originally named ''Arhynchus hemignathi'' by
Arthur Shipley Sir Arthur Everett Shipley GBE FRS (10 March 1861 – 22 September 1927) was an English zoologist and Vice-Chancellor of the University of Cambridge. Biography Shipley was born in Walton-on-Thames, Surrey on 10 March 1861. He was brought up in ...
in 1896. The name ''Arhynchus'' was chosen based on the characteristic absence of a proboscis in this species of Acanthocephala. It was later renamed ''Apororhynchus'' (along with the family name Apororhynchidae) by Shipley in 1899 due to the name ''Arhynchus'' having been used by Dujean in 1834 for a beetle. Although ''Apororhynchus'' has not been included in
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses thus far due to insufficiency of morphological data, the lack of features such as an absence of a muscle plate, a midventral
longitudinal Longitudinal is a geometric term of location which may refer to: * Longitude ** Line of longitude, also called a meridian * Longitudinal engine, an internal combustion engine in which the crankshaft is oriented along the long axis of the vehicl ...
muscle,
lateral Lateral is a geometric term of location which may refer to: Healthcare *Lateral (anatomy), an anatomical direction * Lateral cricoarytenoid muscle * Lateral release (surgery), a surgical procedure on the side of a kneecap Phonetics *Lateral co ...
receptacle flexors, and an apical sensory organ when compared to the other three orders of class
Archiacanthocephala Archiacanthocephala is a class within the phylum of Acanthocephala. They are parasitic worms that attach themselves to the intestinal wall of terrestrial vertebrates, including humans. They are characterised by the body wall and the lemnisci (wh ...
indicate it is an early offshoot ( basal).


Description

The genus ''Apororhynchus'' consists of
ectoparasitic 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 ha ...
worms that attach themselves beneath the skin and around the anus of birds. The distinguishing features of this order among
acanthocephala Acanthocephala (Greek , ', thorn + , ', head) is a phylum 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 to p ...
ns are a highly enlarged
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...
with limited
motility Motility is the ability of an organism to move independently, using metabolic energy. Definitions Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...
and a reduced size of the hooks (or spines). ''Apororhynchus'' species have short conical trunks and a reduced or absent neck. The proboscis is large and globular with numerous deeply set spirally arranged rootless hooks usually not reaching the surface, or with no hooks. They contain sets of muscles that are common to all Acanthocephala including a proboscis receptacle, a receptacle-surrounding muscle called a receptacle protrusor,
retinacula A retinaculum (plural ''retinacula'') is a band of thickened deep fascia around tendons that holds them in place. It is not part of any muscle. Its function is mostly to stabilize a tendon. The term retinaculum is New Latin, derived from the Latin ...
(connective tissue that stabilizes tendons), a neck retractor, proboscis and receptacle retractors, circular and longitudinal musculature under the
metasoma The metasoma is the posterior part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the mesosoma. In insects, it contains most of the digestive tract, respiratory system, and circul ...
l (trunk) tegument, and a single muscular layer beneath the proboscis wall. Two regions of musculature are considerably different in ''Apororhynchus'' compared to the other acanthocephalan orders: the proboscis receptacle and receptacle protrusor are both reorganized in ''Apororhynchus'' with the muscles subdivided into strands extending from the cerebral ganglion, or nerve bundle, to the proboscis wall. These two muscles suspend the cerebral ganglion but are not involved in the eversion of the proboscis. Additional anatomical features that can be used to distinguish this genus among other acanthocephalans include a cerebral ganglion located under the anterior wall of the proboscis, long and tubular lemnisci (bundles of sensory nerve fibers) that run along a central canal, the lack of any
protonephridia The nephridium (plural ''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. Neph ...
(an organ which functions as a kidney), and the presence of eight pear-shaped cement glands used to temporarily close the posterior end of the female after copulation.


Species

There are six species in the genus ''Apororhynchus''. A seventh species, ''Apororhynchus bivoluerus'' Das, 1950 (also called ''A. bivolucrus'') from an
Egyptian vulture The Egyptian vulture (''Neophron percnopterus''), also called the white scavenger vulture or pharaoh's chicken, is a small Old World vulture and the only member of the genus ''Neophron''. It is widely distributed from the Iberian Peninsula and ...
(''Neophron percnopterus'') from India was considered to be a strigeid trematode by Yamaguti (1963). * ''Apororhynchus aculeatus'' Meyer, 1931 ''A. aculeatus'' has been found in Santos, Brazil, parasitizing a
New World oriole New World orioles are a group of birds in the genus ''Icterus'' of the blackbird family. Unrelated to Old World orioles of the family Oriolidae, they are strikingly similar in size, diet, behavior, and strongly contrasting plumage. As a resu ...
. The parasite was discovered in 1931, in the Berlin Museum, taken from the digestive tube of a bird named at that time as "''Oriolus cristatus''", which was likely a
crested oropendola The crested oropendola (''Psarocolius decumanus''), also known as the Suriname crested oropendola or the cornbird, is a New World tropical icterid bird. It is a resident breeder in lowland South America east of the Andes, from Panama and Colombia ...
(''Psarocolius decumanus''). ''A. aculeatus'' was the second parasite to be discovered in its genus, and the specimen used to describe the species was female. Numerous fine hooks on the bulb-shaped proboscis, as well as the different host and location, distinguish it from ''A. hemignathi''. * ''Apororhynchus amphistomi'' Byrd and Denton, 1949 The parasite was discovered in the summer of 1947 infesting a
Canada warbler The Canada warbler (''Cardellina canadensis'') is a small boreal songbird of the New World warbler family (Parulidae). It summers in Canada and northeastern United States and winters in northern South America. Taxonomy In 1760 the French zoolo ...
(''Cardellina canadensis'' formerly ''Wilsonia canadensis'') in Mountain Lake, Virginia, and a
Northern parula The northern parula (''Setophaga americana'') is a small New World warbler. It breeds in eastern North America from southern Canada to Florida. Description The northern parula is one of the smaller North American migratory warblers, often being ...
(''Setophaga americana'' formerly ''Compsothlyphis americana'') in
Augusta, Georgia Augusta ( ), officially Augusta–Richmond County, is a consolidated city-county on the central eastern border of the U.S. state of Georgia. The city lies across the Savannah River from South Carolina at the head of its navigable portion. Georgi ...
, thus making it the third species in its genus to be discovered, and the first species in its genus to be discovered in North America. It was found in the digestive tract, just inside the vent. The species name ''amphistomi'' derives from a superficial resemblance to the amphistomate
trematodes Trematoda is a class of flatworms known as flukes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is usually a snail. The definitive host ...
(flukes with stoma on opposite sides). On the proboscis, there are approximately 20 very fine hooks found on each of 40 rows. The two lemnisci are longer than the body length and are folded in the body cavity. Females are 2.13 mm long by 0.83 mm in maximum width and the proboscis is 0.36 mm long by 0.44 mm in maximum width. The male is smaller being 1.43 mm long by 0.53 mm in maximum width and the proboscis is 0.44 mm long by 0.74 mm in maximum width. * ''Apororhynchus chauhani'' Sen, 1975 ''A. chauhani'' is the only ''Apororhynchus'' species described from India. It was discovered in Srishailam, Andhra Pradesh, in 1975. ''A. chauhani'' was named after Birendra Singh Chauhan, a member of the Zoological Society of India. It parasitizes the spotted owlet (''Athene brama'') and has been found in the intestine. The body is 4.70 mm by 1.70 mm long with a proboscis that is 1.11 mm by 1.68 mm in length and immature eggs are around 0.015 mm to 0.035 mm in diameter. The hooks on the proboscis are finger shaped and numerous, especially in the posterior. The rest of the hooks on the anterior side are irregularly directed and sparse. The proboscis sheath is absent. The nerve ganglion is large and located near the anterior proboscis. The lemnisci are very long and unequal. * ''Apororhynchus hemignathi'' ( Shipley, 1896) ''A. hemignathi'' was the first species of ''Apororhynchus'' to be described with the creation of the genus and family by Arthur Shipley in 1896 due to its uniqueness among already described Acanthocephala. It has been found in
Kaua'i Kauai, () anglicized as Kauai ( ), is geologically the second-oldest of the main Hawaiian Islands (after Niʻihau). With an area of 562.3 square miles (1,456.4 km2), it is the fourth-largest of these islands and the List of islands of th ...
,
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only state ...
, parasitizing the now extinct Kauaʻi ʻakialoa (''Akialoa stejnegeri''). ''A. hemignathi'' was named after the genus of the Kauaʻi ʻakialoa, which was ''Hemignathus'' at the time of the description. Specimens range from 2.5 mm to 3.5 mm long, distended 1 mm to 1.5 mm longer. It has two to four nuclei in the lemnisci. It is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
for the genus. * ''Apororhynchus paulonucleatus'' Khokhlova and Cimbaluk, 1966 This parasite has been found in the
black-winged pratincole The black-winged pratincole (''Glareola nordmanni'') is a wader in the pratincole bird family, Glareolidae. The genus name is a diminutive of Latin ''glarea'', "gravel", referring to a typical nesting habitat for pratincoles. The species nam ...
(''Glareola nordmanni'') at Malye Chany Lake, in
Russia Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It is the largest country in the world, with its internationally recognised territory covering , and encompassing one-eig ...
, and in the colon and cloaca of the Eastern yellow wagtail (''Motacilla tschutschensis'') in Chukotka and
Kamchatka The Kamchatka Peninsula (russian: полуостров Камчатка, Poluostrov Kamchatka, ) is a peninsula in the Russian Far East, with an area of about . The Pacific Ocean and the Sea of Okhotsk make up the peninsula's eastern and west ...
(including the
Karaginsky Island Karaginsky Island or Karaginskiy Island (russian: Карагинский остров) is an island in the Karaginsky Gulf of the Bering Sea. The -wide strait between the Kamchatka Peninsula and this island is called Litke Strait. Karaginsky Isl ...
), also in
Russia Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It is the largest country in the world, with its internationally recognised territory covering , and encompassing one-eig ...
. It was described in 1966. The proboscis is large compared to the body and spherical. It is armed with 10–12 spiral rows of hooks with 14–15 hooks in each row. The hook has a thin blade with a curved tip and root thickened with broadened base. In the body, there are 10–16 giant nuclei with a diameter of 0.050–0.077 mm. There is a very short neck (0.153 mm long) between the proboscis and the body with attached ribbon-like lemnisci longer than their own body length. Females are 3.7 mm long by 0.92 mm in maximum width and the proboscis is 1.30 mm long by 1.53 mm in maximum width. The male is smaller being 3.21 mm long by 0.766 mm in maximum width and the proboscis is 0.796 mm long by 0.995 mm in maximum width. The eggs were oval shaped with three concentric shells around 0.074–0.080 mm long and 0.040–0.043 mm wide. * ''Apororhynchus silesiacus'' Okulewicz and Maruszewski, 1980 ''A. silesiacus'' was found in the cloaca of the
European robin The European robin (''Erithacus rubecula''), known simply as the robin or robin redbreast in Great Britain & Ireland, is a small insectivorous passerine bird that belongs to the chat subfamily of the Old World flycatcher family. About in len ...
(''Erithacus rubecula''), the
thrush nightingale The thrush nightingale (''Luscinia luscinia''), also known as the sprosser, is a small passerine bird that was formerly classed as a member of the thrush family Turdidae, but is now more generally considered to be an Old World flycatcher, Musci ...
(''Luscinia luscinia'') and the
common nightingale The common nightingale, rufous nightingale or simply nightingale (''Luscinia megarhynchos''), is a small passerine bird best known for its powerful and beautiful song. It was formerly classed as a member of the thrush family Turdidae, but is no ...
(''Luscinia megarhynchos'') in Wroclaw, Poland, and of the European robin in Alsóperepuszta, Hungary. Described in 1980, it is the most recently classified species of the ''Apororhynchus''. ''A. silesiacus'' is named after
Silesia Silesia (, also , ) is a historical region of Central Europe that lies mostly within Poland, with small parts in the Czech Republic and Germany. Its area is approximately , and the population is estimated at around 8,000,000. Silesia is split ...
, a region in Poland where the parasite was found. Specimens were between 3.21 and 3.51 mm in total length, with a maximum width of 0.80 to 1.05 mm at middle and the eggs were around 0.07 mm long and 0.035 mm wide. It has a proboscis wider than the anterior part of the trunk, with about 40 spiral rows of hooks, each row bearing 14 to 16 hooks. There are 28 to 31 giant nuclei in the body wall and 9 to 12 in the lemnisci. The testes are almost parallel. The parasite has been found infesting juvenile European robins, indicating that the infestation occurred in the nesting area.


Hosts

''Apororhynchus'' species exclusively parasitize avian hosts of different orders including
owl Owls are birds from the order Strigiformes (), which includes over 200 species of mostly solitary and nocturnal birds of prey typified by an upright stance, a large, broad head, binocular vision, binaural hearing, sharp talons, and feathers a ...
s,
wader 245px, A flock of Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflat ...
s, and
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
s. The parasite attaches to the cloaca and in some cases the intestinal wall using a hook-covered proboscis. Infestation can cause
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. Access ...
and
anemia Anemia or anaemia (British English) is a blood disorder in which the blood has a reduced ability to carry oxygen due to a lower than normal number of red blood cells, or a reduction in the amount of hemoglobin. When anemia comes on slowly, t ...
in
Hawaiian honeycreeper Hawaiian honeycreepers are a group of small, passerine birds endemic to Hawaii. They are closely related to the rosefinches in the genus ''Carpodacus'', but many species have evolved features unlike those present in any other finch. Their great ...
s. File:Crested oropendola (Psarocolius decumanus insularis).jpg, alt=Crested oropendola on a branch, The
crested oropendola The crested oropendola (''Psarocolius decumanus''), also known as the Suriname crested oropendola or the cornbird, is a New World tropical icterid bird. It is a resident breeder in lowland South America east of the Andes, from Panama and Colombia ...
is one of the hosts of ''A. aculeatus'' File:Northern Parula by Dan Pancamo.jpg, alt=Northern parula on a branch, The
northern parula The northern parula (''Setophaga americana'') is a small New World warbler. It breeds in eastern North America from southern Canada to Florida. Description The northern parula is one of the smaller North American migratory warblers, often being ...
is one of the hosts of ''A. amphistomi'' File:Spotted owlet (Athene brama indica) (cropped).jpg, alt=A spotted owlet on a branch, The spotted owlet is a host of ''A. chauhani'' File:Akialoa stejnegeri, Bishop Museum, Honolulu.JPG, alt=A display of a stuffed Kauaʻi ʻakialoa bird, The now extinct Kauaʻi ʻakialoa was a host of ''A. hemignathi'' File:Tyrkyshka.jpg, alt=Black-winged pratincole in flight, The
black-winged pratincole The black-winged pratincole (''Glareola nordmanni'') is a wader in the pratincole bird family, Glareolidae. The genus name is a diminutive of Latin ''glarea'', "gravel", referring to a typical nesting habitat for pratincoles. The species nam ...
is a host of ''A. paulonucleatus'' File:Luscinia megarhynchos - 01 (cropped).jpg, alt=Common nightingale on a branch, The
common nightingale The common nightingale, rufous nightingale or simply nightingale (''Luscinia megarhynchos''), is a small passerine bird best known for its powerful and beautiful song. It was formerly classed as a member of the thrush family Turdidae, but is no ...
is one of the hosts of ''A. silesiacus''


Notes


References


External links

* {{Featured article Archiacanthocephala