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 (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
in the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Apororhynchidae, which in turn is the only member of the
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
Apororhynchida. A lack of features commonly found in the phylum
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 ...
(primarily musculature) suggests an evolutionary branching from the other three orders of class Archiacanthocephala; however no genetic analysis has been completed to determine the evolutionary relationship between species. The distinguishing features of this order among archiacanthocephalans 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 arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...
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 A cloaca ( ), : cloacae ( or ), or vent, is the rear orifice that serves as the only opening for the digestive (rectum), reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles, birds, cartilagin ...
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 arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...
. The bird hosts 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 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 mudflats in order to foraging, ...
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 generally have an anisodactyl arrangement of their ...
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 su ...
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. Acces ...
and
anemia Anemia (also spelt anaemia in British English) is a blood disorder in which the blood has a reduced ability to carry oxygen. This can be due to a lower than normal number of red blood cells, a reduction in the amount of hemoglobin availabl ...
.


Taxonomy

The first species in this order to be described was ''Apororhynchus hemignathi'' which was originally named ''Arhynchus hemignathi'' by Arthur Shipley 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. The
National Center for Biotechnology Information The National Center for Biotechnology Information (NCBI) is part of the National Library of Medicine (NLM), a branch of the National Institutes of Health (NIH). It is approved and funded by the government of the United States. The NCBI is lo ...
does not indicate that any
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 ...
analysis has been published on ''Apororhynchida'' that would confirm its position as a unique order in the class Archiacanthocephala. The lack of morphological features such as an absence of a muscle plate, a midventral longitudinal muscle,
lateral Lateral is a geometric term of location which may also refer to: Biology and healthcare * Lateral (anatomy), a term of location meaning "towards the side" * Lateral cricoarytenoid muscle, an intrinsic muscle of the larynx * Lateral release ( ...
receptacle flexors, and an apical sensory organ when compared to the other three orders of class Archiacanthocephala 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 (at least some of the time) on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The en ...
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 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 ...
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 arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...
with limited
motility Motility is the ability of an organism to move independently using metabolism, metabolic energy. This biological concept encompasses movement at various levels, from whole organisms to cells and subcellular components. Motility is observed in ...
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 (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 (biology), 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 sy ...
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 A ganglion (: ganglia) is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system, this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system, there are ...
, 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 (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 bivolucrus'' Das, 1950 (also called ''A. bivolucrus'') from an Egyptian vulture (''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 Icterid, blackbird family. Although they are not closely related to Old World orioles of the family Oriolidae, they are strikingly similar in size, diet, behavior, and strong ...
. 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 (''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 (''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 is migratory and breeds in eastern North America from southern Canada to Florida. Taxonomy The northern parula was Species description, formally described in 1758 by ...
(''Setophaga americana'' formerly ''Compsothlyphis americana'') in
Augusta, Georgia Augusta is a city on the central eastern border of the U.S. state of Georgia (U.S. state), Georgia. The city lies directly across the Savannah River from North Augusta, South Carolina at the head of its navigable portion. Augusta, the third mos ...
, 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 trematodes, and commonly 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 a moll ...
(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
Srisailam Srisailam is a census town in Nandyal district of the Indian state of Andhra Pradesh. It is the mandal headquarters of Srisailam mandal in Atmakur revenue division, Nandyal district, Atmakur revenue division. It is located about from the distri ...
, 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 ( or ), is one of the main Hawaiian Islands. It has an area of 562.3 square miles (1,456.4 km2), making it the fourth-largest of the islands and the 21st-largest island in the United States. Kauai lies 73 mi ...
,
Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
, 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 International_Code_of_Zoological_Nomenclature, 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 spe ...
for the genus. * ''Apororhynchus paulonucleatus'' Khokhlova and Cimbaluk, 1966 This parasite has been found in the black-winged pratincole (''Glareola nordmanni'') at Malye Chany Lake, in
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
, and in the colon and cloaca of the Eastern yellow wagtail (''Motacilla tschutschensis'') in Chukotka and
Kamchatka The Kamchatka Peninsula (, ) 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 western coastlines, respectively. Immediately offshore along the Pacific ...
(including the Karaginsky Island), also in
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
. 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 In geometry, two or more objects are said to be ''concentric'' when they share the same center. Any pair of (possibly unalike) objects with well-defined centers can be concentric, including circles, spheres, regular polygons, regular polyh ...
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 A cloaca ( ), : cloacae ( or ), or vent, is the rear orifice that serves as the only opening for the digestive (rectum), reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles, birds, cartilagin ...
of the
European robin The European robin (''Erithacus rubecula''), known simply as the robin or robin redbreast in the British Isles, is a small insectivorous passerine bird that belongs to the Old World flycatcher family Muscicapidae. It is found across Europe, ea ...
(''Erithacus rubecula''), the thrush nightingale (''Luscinia luscinia'') and the
common nightingale The common nightingale, rufous nightingale or simply nightingale (''Luscinia megarhynchos''), is a small passerine bird which is best known for its powerful and beautiful Bird vocalization, song. It was formerly classed as a member of the Thrush ...
(''Luscinia megarhynchos'') in
Wrocław Wrocław is a city in southwestern Poland, and the capital of the Lower Silesian Voivodeship. It is the largest city and historical capital of the region of Silesia. It lies on the banks of the Oder River in the Silesian Lowlands of Central Eu ...
, 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 (see names #Etymology, below) is a historical region of Central Europe that lies mostly within Poland, with small parts in the Czech Silesia, Czech Republic and Germany. Its area is approximately , and the population is estimated at 8, ...
, 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

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 ''Apororhynchus'' are not known. 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 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 ''Apororhynchus''. ''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 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 mudflats in order to foraging, ...
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 generally have an anisodactyl arrangement of their ...
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. Acces ...
and
anemia Anemia (also spelt anaemia in British English) is a blood disorder in which the blood has a reduced ability to carry oxygen. This can be due to a lower than normal number of red blood cells, a reduction in the amount of hemoglobin availabl ...
in
Hawaiian honeycreeper Hawaiian honeycreepers are a group of small birds endemic to Hawaii. They are members of the finch family Fringillidae, closely related to the rosefinches (''Carpodacus''), but many species have evolved features unlike those present in any othe ...
s. There are no reported cases of any ''Apororhynchus'' species infesting humans in the English language medical literature. File:Crested oropendola (Psarocolius decumanus insularis).jpg, alt=Crested oropendola on a branch, The crested oropendola 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 is migratory and breeds in eastern North America from southern Canada to Florida. Taxonomy The northern parula was Species description, formally described in 1758 by ...
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 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 which is best known for its powerful and beautiful Bird vocalization, song. It was formerly classed as a member of the Thrush ...
is one of the hosts of ''A. silesiacus''


Notes


References


External links

* {{Featured article Archiacanthocephala