Profilicollis Major
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''Profilicollis'' is a
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 ...
of
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 ...
n
parasites 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 ...
of
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s. The status of the genus ''Profilicollis'' has been debated, and species placed in this genus were formerly included in the genus '' Polymorphus''. However, research on the morphology of the group and their use of hosts has concluded that ''Profilicollis'' and ''Polymorphus'' should be regarded as distinct genera, and species previously described as ''Polymorphus altmani'' are now referred to as ''Profilicollis altmani'' in taxonomic and biological literature. ''Profilicollis'' parasites infect decapod crustaceans, usually shore crabs, as intermediate hosts, and use many species of shorebirds as definitive (final) hosts.


Life cycle

This parasite first develops in the haemocoel or digestive gland of shore crabs, which are the intermediate host. The species of crabs that are parasitized differs between ''Profilicollis'' species. Mole crabs in the genus ''Emerita'' are parasitized throughout North and South America by ''Profilicollis altmani''. After infection of the crab, the parasite becomes a dormant cystacanth until the crab is eaten by a suitable bird, such as the
surf scoter The surf scoter (''Melanitta perspicillata'') is a large sea duck native to North America. Adult males are almost entirely black with characteristic white patches on the forehead and the nape and adult females are slightly smaller and browner. S ...
''Melanitta perspicillata'' or
herring gull Herring gull is a common name for several birds in the genus ''Larus'', all formerly treated as a single species. Three species are still combined in some taxonomies: * American herring gull (''Larus smithsonianus'') - North America * European h ...
, ''Larus argentatus'' (the final or definitive host). Once the parasite cystacanth has passed through the stomach of the bird, it develops into the adult worm and attaches to the intestines of the bird. Eggs produced by the parasite are released from the bird with bird feces and enter the ocean. Here the eggs are transported by the currents until they are ingested by a filter-feeding mole crab. The southern
sea otter The sea otter (''Enhydra lutris'') is a marine mammal native to the coasts of the northern and eastern Pacific Ocean, North Pacific Ocean. Adult sea otters typically weigh between , making them the heaviest members of ...
, ''Enhydra lutris nereis'' is an accidental host (or paratenic host) of ''Profilicollis altmani'' and ''P. major''. Between 1998 and 2001, between 13% and 16.2% of sea otter carcasses found along the coasts of central California were determined to be caused by acanthocephalan peritonitis, or inflammation of the intestines caused by acanthocephalan parasites. This is a large increase in deaths of sea otters due to these parasites; only a few decades ago, the proportion of sea otters dying because of these parasites was almost negligible. Most of the sea otters that were found to be killed by infections by ''Profilicollis altmani'' were either young juvenile or older female otters, and it is hypothesized that they feed on sand crabs because they are less skilled at foraging or because of a decrease in their preferred prey.


Behavioral manipulation

Like many other acanthocephalans, recent studies have shown that the presence of cystacanth of ''Profilicollis antarcticus'' causes behavioral alterations due to changes in the levels of hemolymph dopamine in its intermediate host, the crab '' Hemigrapsus crenulatus''. These changes in the biogenic amino levels can cause behavioral changes such as changes in response to external stimuli resulting from neurological damages (escape behavior), increased conspicuity resulting from modified pigmentation and/ or behavior and castration. Previous studies have also shown that crabs infected with cystacanths exhibit higher metabolic rates and activity levels than non-parasitized crabs. The free living conditions of cystacanths in the hemocoelomic cavity of the intermediate host maybe interpreted as the direct way of altering the dopamine metabolic pathway. The alteration of dopamine levels in the hemolymph system implies that there is a generalized effect of the parasite on the body of its intermediate host, rather than an effect on the nerve ganglion level. Therefore, the increase in activity and metabolic rates for this host parasite system could be a consequence of higher metabolic cost of maintenance for the host, linked to the synthesis of monoamines and/ or the physiological consequences of their bio-availability. This altered physiological condition is beneficial for the parasite, considering that their final goal is to increase transmission by increasing the probability of predation on the intermediate host (the crab) by its definitive host (the gull).


Species

* '' Profilicollis altmani'' (Perry, 1942) Van Cleave, 1947 Infects mole crabs as intermediate hosts in North and South America. Final hosts include scoters, ''
Melanitta The scoters are stocky seaducks in the genus ''Melanitta''. The drakes are mostly black and have swollen bills, the females are brown. They breed in the far north of Europe, Asia, and North America, and winter further south in temperate zones o ...
'' spp.; gulls, ''Chroicocephalus maculipennis'', ''Leucopheus pipixcan'', ''Larus modestus'', and ''L. dominicanus'';
lesser scaup The lesser scaup (''Aythya affinis'') is a small North American diving duck that migrates south as far as Central America in winter. It is colloquially known as the little bluebill or broadbill because of its distinctive blue bill. The origin of ...
, ''Marilla affnis'';
sanderling The sanderling (''Calidris alba'') is a small wading bird. The name derives from Old English ''sand-yrðling'', "sand-ploughman". The genus name is from Ancient Greek ''kalidris'' or ''skalidris'', a term used by Aristotle for some grey-colour ...
, ''Calidris alba'';
dunlin The dunlin (''Calidris alpina'') is a small wader in the genus '' Calidris''. The English name is a dialect form of "dunling", first recorded in 1531–1532. It derives from ''dun'', "dull brown", with the suffix ''-ling'', meaning a person or ...
, ''Calidris alpina''; whimbrel, ''Phaeopus hudsonicus'';
rhinoceros auklet The rhinoceros auklet (''Cerorhinca monocerata'') is a seabird and a close relative of the puffins. It is the only extant species of the genus ''Cerorhinca''. Given its close relationship with the puffins, the common name rhinoceros puffin has b ...
, ''Cerorhinca monocerata''; and
willet The willet (''Tringa semipalmata'') is a large shorebird in the family Scolopacidae. It is a relatively large and robust sandpiper and is the largest of the species called "shanks" in the genus ''Tringa''. Its closest relative is the lesser yell ...
, ''Catoptrophorus semipalmatus''. This species was moved to this genus by Van Cleave in 1947. This species was redescribed in great detail by Amin et al. in 2022 on the basis of specimens from Pacific mole crab ''
Emerita analoga ''Emerita analoga'', the Pacific sand crab, Pacific mole crab or coldwater mole crab, is a species of small, sand-burrowing Decapoda, decapod crustacean found living in the sand along the temperate western coasts of North and South America. It is ...
'' ( Hippidae) and Belcher's gull ''
Larus belcheri Belcher's gull (''Larus belcheri''), also known as the band-tailed gull, is a bird in the family Laridae found along the Pacific coast of South America. It formerly included the very similar Olrog's gull as a subspecies, but that bird occurs on t ...
'' (
Laridae Laridae is a family of seabirds in the order Charadriiformes that includes the gulls, terns (including white terns), noddies, and skimmers. It includes around 100 species arranged into 22 genera. They are an adaptable group of mostly aerial bird ...
), in Peru. * '' Profilicollis antarcticus'' (Zdzitowiecki, 1985) Infects shore crabs such as '' Hemigrapsus crenulatus'' and gulls such as '' Larus dominicanus'' in Chile and New Zealand. * '' Profilicollis arcticus'' (Van Cleave, 1920) * '' Profilicollis botulus'' (Van Cleave, 1916) Infects shore crabs such as '' Hemigrapsus oregonensis'' and the invasive green crab ''
Carcinus maenas ''Carcinus maenas'' is a common littoral crab. It is known by different names around the world. In the British Isles, it is generally referred to as the shore crab or green shore crab. In North America and South Africa, it bears the name Europe ...
'' as well as the lobster ''
Homarus americanus The American lobster (''Homarus americanus'') is a species of lobster found on the Atlantic coast of North America, chiefly from Labrador to New Jersey. It is also known as Atlantic lobster, Canadian lobster, true lobster, northern lobster, Cana ...
''. Its final hosts are shorebirds such as
common goldeneye The common goldeneye or simply goldeneye (''Bucephala clangula'') is a medium-sized sea duck of the genus ''Goldeneye (duck), Bucephala'', the goldeneyes. Its closest relative is the similar Barrow's goldeneye. The genus name is derived from th ...
, ''Bucephala clangula'';
Barrow's goldeneye Barrow's goldeneye (''Bucephala islandica'') is a medium-sized sea duck of the genus '' Bucephala'', the goldeneyes. This bird was named after Sir John Barrow. The genus name is derived from Ancient Greek ''boukephalos'', "bullheaded", from ''bo ...
, ''B. islandica'';
greater scaup The greater scaup (''Aythya marila''), just scaup in Europe or, colloquially, "bluebill" in North America, is a mid-sized diving duck, larger than the closely related lesser scaup and tufted duck. It spends the summer months breeding in Iceland ...
, ''Aythya marila'';
eider duck The eiders () are large Mergini, seaducks in the genus ''Somateria''. The three extant species all breed in the cooler latitudes of the Northern Hemisphere. The down feathers of eider ducks and some other ducks and geese are used to fill pillow ...
, ''Somateria mollissima''; and
long-tailed duck The long-tailed duck (''Clangula hyemalis'') or coween, is a medium-sized sea duck that breeds in the tundra and taiga regions of the arctic and winters along the northern coastlines of the Atlantic and Pacific Oceans. It is the only member of ...
, ''Clangula hyemalis''. * '' Profilicollis chasmagnathi'' (Holcman-Spector, Mane-Garzon and Dei-Cas, 1977) Infects estuarine crabs (described in Uruguay) and gulls, including ''
Larus atlanticus Olrog's gull (''Larus atlanticus'') is a species of gull found along the Atlantic coast of southern Brazil, Uruguay, and northern Argentina. It was formerly considered a subspecies of the very similar '' L. belcheri''. It is a large gull with a b ...
''. The species was moved to this genus by Golvan, 1994. * '' Profilicollis formosus'' Schmidt and Kuntz, 1967 Infects ''
Macrobrachium ''Macrobrachium'' is a genus of freshwater prawns or shrimps characterised by the extreme enlargement of the second pair of pereiopods, at least in the male. Species It contains these species: *'' Macrobrachium acanthochirus'' F. Villalobos, ...
'' and
domestic duck Domestic ducks (mainly mallards, ''Anas platyrhynchos domesticus'', with some Muscovy ducks, ''Cairina moschata domestica'') are ducks that have been domesticated and raised for meat and egg (food), eggs. A few are kept for show, or for thei ...
s in Taiwan. * '' Profilicollis major'' Lundström, 1942 Infects the Atlantic rock crab ''
Cancer irroratus Description ''This crab species occurs on the eastern coast of North America, Labrador Canada to Iceland to South Carolina. Rock crabs live over a large depth range, from well above the low tide line to as deep as . And live up to 8 years. Canc ...
'', although likely other species as well. This parasite has also found in the intestines of sea otters, although it appears to be much less pathogenic to the otters than ''Profilicollis altmani''. * '' Profilicollis novaezelandensis'' (Brockerhoff, 2002) Infects shore crabs such as '' Hemigrapsus crenulatus'', and mud crabs '' Macrophthalmus hirtipes'' and '' Helice crassa''. They then infect birds such as the
South Island pied oystercatcher The South Island oystercatcher or South Island pied oystercatcher (''Haematopus finschi'') is one of two common oystercatcher species found in New Zealand. Its name is often contracted to the acronym "SIPO" (rhyming with "typo"). The indigenous ...
''Haematopus ostralegus finschi'' and
bar-tailed godwit The bar-tailed godwit (''Limosa lapponica'') is a large and strongly migratory wader in the family Scolopacidae, which feeds on bristle-worms and shellfish on coastal mudflats and estuaries. It has distinctive red breeding plumage, long legs, ...
''Limosa lapponica'' in New Zealand. * '' Profilicollis rancoensis'' * '' Profilicollis sphaerocephalus'' (Bremser, 1819) Infects many grapsid crabs (including '' Brachynotus spinosus,'' ''
Cyclograpsus granulosus ''Cyclograpsus granulosus'', the purple mottled shore crab or smooth shore crab, is an Australasian crab species. Description ''Cyclograpsus granulosus'' grows to a carapace width of about , with a carapace mottled with red-brown or purple on ye ...
'', '' Paragrapsus gaimardii'', '' Paragrapsus laevis'' and '' Paragrapsus quadridentatus'') 1 species of portunid crab ('' Nectocarcinus integrifrons''). This species also infects gulls.


Proposed synonyms

* ''Profilicollis kenti'' and ''Profilicollis texensis'' were described as parasitizing mole crabs in North America, in addition to the species ''Profilicollis altmani''. Morphological examination first suggested that ''Profilicollis kenti'' and ''Profilicollis texensis'' were synonyms of ''Profilicollis altmani''. Later, DNA sequencing of acanthocephalans sampled form the mole crab ''Emerita analoga'' in the Pacific and ''E. talpoida'' in the Atlantic showed that only one species of acanthocephalan was present in ''Emerita'' crabs in North America. * ''Profilicollis bullocki'' was reported as parasitizing the
mole crab Hippoidea is a superfamily of decapod crustaceans known as mole crabs or sand crabs. Ecology Hippoids are adapted to burrowing into sandy beaches, a habit they share with raninid crabs, and the parallel evolution of the two groups is strikin ...
s in Chile and Peru. DNA sequences were used to test whether the species ''Profilicollis bullocki'' in South America was genetically distinct from ''Profilicollis altmani'' in North America. The parasites from the mole crab ''Emerita analoga'' in Chile and ''E. rathbunae'' in Panama were found to be highly similar to the acanthocephalans from mole crabs in North America, strongly suggesting they are the same species. Later research showed that acanthocephalan parasites from the Atlantic coast of South America (Uruguay) are also genetically similar, and belong to the same species.


References

{{Taxonbar, from=Q2095471 Polymorphidae Acanthocephala genera