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The limpkin (''Aramus guarauna''), also called carrao, courlan, and crying bird, is a large wading bird related to
rails Rail or rails may refer to: Rail transport *Rail transport and related matters *Rail (rail transport) or railway lines, the running surface of a railway Arts and media Film * ''Rails'' (film), a 1929 Italian film by Mario Camerini * ''Rail'' ( ...
and cranes, and the only extant species 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 ...
Aramidae. It is found mostly in wetlands in warm parts of the
Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...
, from
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
to northern Argentina. It feeds on molluscs, with the diet dominated by apple snails of the genus '' Pomacea''. Its name derives from its seeming limp when it walks.


Taxonomy and systematics

The limpkin is placed in the family Aramidae, which is in turn placed within the crane and
rail Rail or rails may refer to: Rail transport *Rail transport and related matters *Rail (rail transport) or railway lines, the running surface of a railway Arts and media Film * ''Rails'' (film), a 1929 Italian film by Mario Camerini * ''Rail'' ( ...
order
Gruiformes The Gruiformes are an order containing a considerable number of living and extinct bird families, with a widespread geographical diversity. Gruiform means "crane-like". Traditionally, a number of wading and terrestrial bird families that did ...
. The limpkin had been suggested to be close to the
ibis The ibises () (collective plural ibis; classical plurals ibides and ibes) are a group of long-legged wading birds in the family Threskiornithidae, that inhabit wetlands, forests and plains. "Ibis" derives from the Latin and Ancient Greek word ...
and spoonbill family
Threskiornithidae The family Threskiornithidae includes 36 species of large wading birds. The family has been traditionally classified into two subfamilies, the ibises and the spoonbills; however recent genetic studies have cast doubt on this arrangement, and ha ...
, based upon shared bird lice. The Sibley–Ahlquist taxonomy of birds, based upon DNA–DNA hybridization, suggested that the limpkin's closest relatives were the Heliornithidae finfoots, and Sibley and Monroe even placed the species in that family in 1990. More recent studies have found little support for this relationship. More recent DNA studies have confirmed a close relationship with particularly the cranes, with the limpkin remaining as a family close to the cranes and the two being sister taxa to the
trumpeter The trumpet is a brass instrument commonly used in classical and jazz ensembles. The trumpet group ranges from the piccolo trumpet—with the highest register in the brass family—to the bass trumpet, pitched one octave below the standard B ...
s. Although the limpkin is the only extant species in the family today, several
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s of extinct Aramidae are known from across the Americas. The earliest known species, '' Aramus paludigrus'', is dated to the middle
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
, while the oldest supposed members of the family, '' Aminornis'' and '' Loncornis'', have been found in early Oligocene deposits in Argentina, although whether these are indeed related is not certain; in fact, ''Loncornis'' seems to be a misidentified mammal bone. Another Oligocene fossil from Europe, ''
Parvigrus ''Parvigrus'' is an extinct bird genus, with the single species ''Parvigrus pohli''. It is considered a family, the Parvigruidae. The remains of ''Parvigrus pohli'' have been described from fossils found in Vachères in France, from rocks from ...
pohli'' (family Parvigruidae), has been described as a mosaic of the features shared by the limpkins and the cranes. It shares many morphological features with the cranes and limpkins, but also was much smaller than either group, and was more rail-like in its proportions. In the paper describing the fossil,
Gerald Mayr Gerald Mayr is a German palaeontologist who is Curator of Ornithology at the Senckenberg Research Institute in Frankfurt am Main, Hesse. He has published extensively on fossil birds, especially the Paleogene avifauna of Europe. He is an expert on ...
suggested that it was similar to the stem species of the grues (the cranes and limpkins), and that the limpkins evolved massively long bills as a result of the specialisation to feeding on snails. In contrast, the cranes evolved into long-legged forms to walk and probe on open grasslands.


Subspecies

Between 1856 and 1934, the limpkin was treated as two species, one in South America (''Aramus guarauna'') and the other found in Central America, the Caribbean, and Florida (''Aramus pictus''). Today, it is treated as a single species with four subspecies. Along with the nominate subspecies ''A. g. guarauna'', ''A. g. dolosus'', ''A. g. elucus'' (both J. L. Peters, 1925), and ''A. g. pictus'' ( F. A. A. Meyer, 1794) are recognized. The difference between the subspecies are related to slight differences in size and plumage. * ''Aramus guarauna guarauna'' -
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...
(except the arid west coast, the
Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...
and extreme south) * ''Aramus guarauna pictus'' - Florida, Georgia,
The Bahamas The Bahamas (), officially the Commonwealth of The Bahamas, is an island country within the Lucayan Archipelago of the West Indies in the North Atlantic. It takes up 97% of the Lucayan Archipelago's land area and is home to 88% of the ar ...
,
Cuba Cuba ( , ), officially the Republic of Cuba ( es, República de Cuba, links=no ), is an island country comprising the island of Cuba, as well as Isla de la Juventud and several minor archipelagos. Cuba is located where the northern Caribbea ...
and
Jamaica Jamaica (; ) is an island country situated in the Caribbean Sea. Spanning in area, it is the third-largest island of the Greater Antilles and the Caribbean (after Cuba and Hispaniola). Jamaica lies about south of Cuba, and west of His ...
* ''Aramus guarauna elucus'' - Hispaniola and (formerly)
Puerto Rico Puerto Rico (; abbreviated PR; tnq, Boriken, ''Borinquen''), officially the Commonwealth of Puerto Rico ( es, link=yes, Estado Libre Asociado de Puerto Rico, lit=Free Associated State of Puerto Rico), is a Caribbean island and unincorporated ...
* ''Aramus guarauna dolosus'' - Southwestern
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
to
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a transcontinental country spanning the southern part of North America and the northern part of South America. It is bordered by Co ...


Description

The limpkin is a somewhat large bird, long, with a wingspan of . Body mass ranges from , averaging . The males are slightly larger than the females in size, but no difference in plumage is seen. Its plumage is drab—dark brown with an olive luster above. The feathers of the head, neck, wing coverts, and much of the back and underparts (except the rear) are marked with white, making the body look streaked and the head and neck light gray. It has long, dark-gray legs and a long neck. Its bill is long, heavy, and downcurved, yellowish bill with a darker tip. The bill is slightly open near but not at the end to give it a tweezers-like action in removing snails from their shells, and in many individuals the tip curves slightly to the right, like the apple snails' shells. The white markings are slightly less conspicuous in first-year birds. Its wings are broad and rounded and its tail is short. It is often confused with the immature American white ibis. This bird is easier to hear than see. Its common vocalization is a loud wild wail or scream with some rattling quality, represented as "''kwEEEeeer'' or ''klAAAar''." This call is most often given at night and at dawn and dusk. Other calls include "wooden clicking", clucks, and in alarm, a "piercing ''bihk, bihk...''". Limpkin-2.jpg, Limpkin performing a wing-stretch Aramus guarauna-takes off.jpg, Taking off


Distribution and habitat

The limpkin occurs from peninsular Florida (and the
Okefenokee Swamp The Okefenokee Swamp is a shallow, 438,000-acre (177,000 ha), peat-filled wetland straddling the Georgia–Florida line in the United States. A majority of the swamp is protected by the Okefenokee National Wildlife Refuge and the Okefenokee ...
in southern
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
) and southern Mexico through the Caribbean and Central America to northern
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
. In South America, it occurs widely east of the
Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...
; west of them its range extends only to the Equator. It inhabits freshwater
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found a ...
es and swamps, often with tall reeds, as well as
mangrove A mangrove is a shrub or tree that grows in coastal saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evolution in severa ...
s. In the Caribbean, it also inhabits dry brushland. In Mexico and northern Central America, it occurs at altitudes up to . In Florida, the distribution of apple snails is the best predictor of where limpkins can be found. The limpkin undertakes some localized migrations, although the extent of these is not fully understood. In some parts in the northern part of the range, females (and a few males) leave the breeding areas at the end of summer, returning at the end of winter. In Brazil, birds breeding in some seasonal marshes leave during the dry season and return again with the rains. Birds may also migrate between Florida and Cuba, as several limpkins on the Florida Keys and
Dry Tortugas Dry Tortugas National Park is a national park located about west of Key West in the Gulf of Mexico. The park preserves Fort Jefferson and the seven Dry Tortugas islands, the westernmost and most isolated of the Florida Keys. The archipelago's c ...
have been reported, but these records may also represent
vagrants Vagrancy is the condition of homelessness without regular employment or income. Vagrants (also known as bums, vagabonds, rogues, tramps or drifters) usually live in poverty and support themselves by begging, scavenging, petty theft, tempora ...
or postbreeding dispersal. One study in Florida using wing tags found limpkins dispersed up to away from the breeding site. This tendency may explain vagrant limpkins seen in other parts of the United States and at sea near the Bahamas.


Behavior and ecology

Limpkins are active during the day, but also forage at night. Where they are not persecuted, they are also very tame and approachable. Even so, they are usually found near cover. They are not aggressive for the most part, being unconcerned by other species and rarely fighting with members of their own species. Because of their long toes, they can stand on floating water plants. They also swim well, both as adults or as newly hatched chicks, but they seldom do so. They fly strongly, the neck projecting forward and the legs backward, the wings beating shallowly and stiffly, with a jerky upstroke, above the horizontal most of the time.


Feeding

Limpkins forage primarily in shallow water and on floating vegetation such as
water hyacinth ''Pontederia crassipes'' (formerly ''Eichhornia crassipes''), commonly known as common water hyacinth is an aquatic plant native to South America, naturalized throughout the world, and often invasive outside its native range.water lettuce. When wading, they seldom go deeper than having half the body underwater, and never are submerged up to the back. They walk slowly with a gait described as "slightly undulating" and "giving the impression of lameness or limping", "high-stepping", or "strolling", looking for food if the water is clear or probing with the bill. They do not associate with other birds in
mixed-species feeding flock A mixed-species feeding flock, also termed a mixed-species foraging flock, mixed hunting party or informally bird wave, is a flock of usually insectivorous birds of different species that join each other and move together while foraging. These are ...
s, as do some other wading birds, but may forage in small groups with others of their species. The diet of the limpkin is dominated by apple snails (
Ampullariidae Ampullariidae, commonly known as the apple snails, is a family of large freshwater snails, aquatic gastropod mollusks with a gill and an operculum. These snails simultaneously have a gill and a lung as functional respiratory structures ...
) of the genus '' Pomacea''. The availability of this one
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
has a significant effect on the local distribution of the limpkin.
Freshwater mussel Freshwater bivalves are one kind of freshwater mollusc, along with freshwater snails. They are bivalves that live in fresh water as opposed to salt water, which is the main habitat type for bivalves. The majority of species of bivalve molluscs ...
s, including '' Anodonta cowperiana'', '' Villosa vibex'', '' Elliptio strigosus'', ''E. jayensis'', and '' Uniomerus obesus'', as well as other kinds of snails, are a secondary food sources. Less important prey items are
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
s,
frog A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-frog" ''Triadobatrachus'' is ...
s, lizards,
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
s (such as crayfish) and
worm Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limbs, and no eyes (though not always). Worms vary in size from microscopic to over in length for marine polychaete wo ...
s, as well as
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s.https://sta.uwi.edu/fst/lifesciences/sites/default/files/lifesciences/images/Aramus%20guarana%20-%20Limpkin.pdf These prey items may be important in periods of drought or flooding when birds may be pushed into less than optimal foraging areas. In one site in Florida,
moon snails Naticidae, common name moon snails or necklace shells, is a family of medium to large-sized predatory sea snails, marine gastropod molluscs in the clade Littorinimorpha. The shells of the species in this family are mostly globular in shape. Nat ...
and mussels were the most important prey items. Two studies, both in Florida, have looked at the percentage composition of the diet of limpkins. One, looking at stomach contents, found 70% ''Pomacea'' apple snails, 3% '' Campeloma'', and 27% unidentified mollusc, probably ''Pomacea''. When a limpkin finds an apple snail, it carries it to land or very shallow water and places it in mud, the opening facing up. It deftly removes the operculum or "lid" and extracts the snail, seldom breaking the shell. The extraction takes 10 to 20 seconds. The orange-yellow yolk gland of female snails is usually shaken loose and not eaten. It often leaves piles of empty shells at favored spots.


Reproduction and breeding

Males have exclusive
territories A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...
, which can vary in size from . In large, uniform swamps, nesting territories can often be clumped together, in the form of large
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
. These are vigorously defended, with males flying to the territory edges to challenge intruders and passing limpkins being chased out of the territory. Territorial displays between males at boundaries include ritualized charging and wing-flapping. Females may also participate in territorial defense, but usually only against other females or juveniles. Territories may be maintained year-round or abandoned temporarily during the nonbreeding season, usually due to lack of food. Limpkins may be either monogamous, with females joining a male's territory, or serially polyandrous, with two or more females joining a male. With the monogamous pairs, banding studies have shown that a small number of pairs reform the following year (four out of 18 pairs). Nests may be built in a wide variety of places – on the ground, in dense floating vegetation, in bushes, or at any height in trees. They are bulky structures of rushes, sticks, or other materials. Nest building is undertaken by the male initially, which constructs the nest in his territory prior to pair-bond formation. Unpaired females visit a number of territories before settling on a male with which to breed. Males may initially challenge and fight off prospective mates, and may not accept first-year females as mates. Pair-bond formation may take a few weeks. Courtship feeding is part of the bonding process, where males catch and process a snail and then feed it to the female. The clutch consists of three to eight eggs, with five to seven being typical and averaging 5.5, which measure . The egg color is highly variable. Their background color ranges from gray-white through buff to deep olive, and they are marked with light-brown and sometimes purplish-gray blotches and speckles. The eggs are laid daily until the clutch is complete, and incubation is usually delayed until the clutch is completed. Both parents incubate the eggs during the day, but only the female incubates at night. The shift length is variable, but the male incubates for longer during the day. The male remains territorial during incubation, and leaves the clutch to chase off intruders; if this happens, the female returns quickly to the eggs. The incubation period is about 27 days, and all the eggs hatch within 24 hours of each other. The young hatch covered with down, capable of walking, running, and swimming. They follow their parents to a platform of aquatic vegetation, where they are brooded. They are fed by both parents; they reach adult size at 7 weeks and leave their parents at about 16 weeks.


Ecology

Limpkins are reported to be attacked and eaten by
American alligator The American alligator (''Alligator mississippiensis''), sometimes referred to colloquially as a gator or common alligator, is a large crocodilian reptile native to the Southeastern United States. It is one of the two extant species in the gen ...
s. Also, adults with serious foot and leg injuries have been reported, suggesting they may have been attacked by turtles while standing on floating vegetation. Their nests are apparently preyed upon by snakes,
raccoon The raccoon ( or , ''Procyon lotor''), sometimes called the common raccoon to distinguish it from other species, is a mammal native to North America. It is the largest of the procyonid family, having a body length of , and a body weight of ...
s, crows, and muskrats. Foraging adults may in times of drought be victims of
kleptoparasitism Kleptoparasitism (etymologically, parasitism by theft) is a form of feeding in which one animal deliberately takes food from another. The strategy is evolutionarily stable when stealing is less costly than direct feeding, which can mean when f ...
by
snail kite The snail kite (''Rostrhamus sociabilis'') is a bird of prey within the family Accipitridae, which also includes the eagles, hawks, and Old World vultures. Its relative, the slender-billed kite, is now again placed in ''Helicolestes'', making ...
s, and the attempted theft of apple snails caught by limpkins has also been observed in
boat-tailed grackle The boat-tailed grackle (''Quiscalus major'') is a passerine bird of the family Icteridae found as a permanent resident on the coasts of the Southeastern United States. Habitat The boat-tailed grackle is found in coastal saltwater marshes and, i ...
s. Limpkins in Florida were examined for parasites, which included
trematode 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 h ...
s, nematodes, and biting lice. Two biting lice species were found, '' Laemobothrion cubense'' and ''
Rallicola ''Rallicola'' is a genus of chewing louse. It is an ectoparasite of rails and other birds. It was named by Thomas Harvey Johnston and Launcelot Harrison in 1911. There are two subgenera aside from the nominotypical subgenus: ''Aptericola'', w ...
funebris''. The trematode '' Prionosoma serratum'' was found in the intestines of some birds; this species may enter the bird after first infecting apple snails (this has been shown to be the route of infection for a closely related trematode to infect snail kites). Nematodes '' Amidostomum acutum'' and '' Strongyloides'' spp. are also ingested and live in the gut.


Relationship with humans

Many of the limpkin's names across its range are onomatopoeic and reflect the bird's call; for example, '' carau ''in Argentina, ''carrao'' in Venezuela, and ''guareáo ''in Cuba. The species also has a range of common names that refer to its call, for example lamenting bird, or to its supposed gait, crippled bird. The limpkin does not feature much in folklore, although in the Amazon people believe that when the limpkin starts to call, the river will not rise any more. Its call has been used for jungle sound effects in
Tarzan Tarzan (John Clayton II, Viscount Greystoke) is a fictional character, an archetypal feral child raised in the African jungle by the Mangani great apes; he later experiences civilization, only to reject it and return to the wild as a heroic adv ...
films and for the
hippogriff The hippogriff, or sometimes spelled hippogryph ( el, Ἱππόγρυπας), is a legendary creature with the front half of an eagle and the hind half of a horse. It was invented by Ludovico Ariosto in his ''Orlando Furioso'', at the beginning o ...
in the film ''
Harry Potter and the Prisoner of Azkaban ''Harry Potter and the Prisoner of Azkaban'' is a fantasy novel written by British author J. K. Rowling and is the third in the '' Harry Potter'' series. The book follows Harry Potter, a young wizard, in his third year at Hogwarts School of ...
''.


References


External links

*
Limpkin Bird Sound
at Florida Museum of Natural History * * * * {{Taxonbar, from=Q725276 Gruiformes Wading birds Native birds of the Southeastern United States Birds of the Americas Birds of Hispaniola Birds of the Dominican Republic Birds of Haiti Birds of the Guianas Birds described in 1766 Taxa named by Carl Linnaeus