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Storks are large, long-legged, long-necked wading birds with long, stout bills. They belong to the family called Ciconiidae, and make up the order Ciconiiformes . Ciconiiformes previously included a number of other families, such as
heron The herons are long-legged, long-necked, freshwater and coastal birds in the family Ardeidae, with 72 recognised species, some of which are referred to as egrets or bitterns rather than herons. Members of the genera ''Botaurus'' and ''Ixobrychus ...
s and ibises, but those families have been moved to other orders. Storks dwell in many regions and tend to live in drier habitats than the closely related
heron The herons are long-legged, long-necked, freshwater and coastal birds in the family Ardeidae, with 72 recognised species, some of which are referred to as egrets or bitterns rather than herons. Members of the genera ''Botaurus'' and ''Ixobrychus ...
s,
spoonbill Spoonbills are a genus, ''Platalea'', of large, long-legged wading birds. The spoonbills have a global distribution, being found on every continent except Antarctica. The genus name ''Platalea'' derives from Ancient Greek and means "broad", refe ...
s and ibises; they also lack the powder down that those groups use to clean off fish slime. Bill-clattering is an important mode of communication at the nest. Many species are migratory. Most storks eat frogs, fish, insects, earthworms, small birds and small
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s. There are 19 living species of storks in six
genera 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 nomenclat ...
. Various terms are used to refer to groups of storks, two frequently used ones being a ''muster'' of storks and a ''phalanx'' of storks. Storks tend to use soaring, gliding flight, which conserves energy. Soaring requires thermal air currents. Ottomar Anschütz's famous 1884 album of photographs of storks inspired the design of Otto Lilienthal's experimental gliders of the late nineteenth century. Storks are heavy, with wide wingspans: the
marabou stork The marabou stork (''Leptoptilos crumenifer'') is a large wading bird in the stork family Ciconiidae native to sub-Saharan Africa. It breeds in both wet and arid habitats, often near human habitation, especially landfill sites. It is someti ...
, with a wingspan of and weight up to , joins the Andean condor in having the widest wingspan of all living land birds. Their nests are often very large and may be used for many years. Some nests have been known to grow to over in diameter and about in depth. All storks were once thought to be
monogamous Monogamy ( ) is a form of Dyad (sociology), dyadic Intimate relationship, relationship in which an individual has only one Significant other, partner during their lifetime. Alternately, only one partner at any one time (Monogamy#Serial monogamy, ...
, but this is only partially true. Some species may change mates after migrations, and may migrate without a mate. Storks' large size, serial monogamy, and faithfulness to an established nesting site contribute to their prominence in mythology and culture.


Morphology

Storks range in size from the marabou, which stands tall and can weigh , to the Abdim's stork, which is only high and weighs only . Their shape is superficially similar to the herons, with long legs and necks, but they are more heavy-set. There is some sexual dimorphism (differences between males and females) in size, with males being up to 15% bigger than females in some species (for example the saddle-billed stork), but almost no difference in appearance. The only difference is in the colour of the iris of the two species in the genus '' Ephippiorhynchus''. The bills of storks are large to very large, and vary considerably between the genera. The shape of the bills is linked to the diet of the different species. The large bills of the ''Ciconia'' storks are the least specialised. Larger are the massive and slightly upturned bills of the ''Ephippiorhynchus'' and the jabiru. These have evolved to hunt for fish in shallow water. Larger still are the massive daggers of the two adjutants and marabou (''Leptoptilos''), which are used to feed on carrion and in defence against other scavengers, as well as for taking other prey. The long, ibis-like downcurved bills of the ''Mycteria'' storks have sensitive tips that allow them to detect prey by touch (tactilocation) where cloudy conditions would not allow them to see it. The most specialised bills of any storks are those of the two openbills (''Anastomus.''), which as their name suggested is open in the middle when their bill is closed. These bills have evolved to help openbills feed on their only prey item, aquatic snails. Although it is sometimes reported that storks lack syrinxes and are mute, they do have syrinxes, and are capable of making some sounds, although they do not do so often. The syrinxes of storks are "variably degenerate" however, and the syringeal membranes of some species are found between tracheal rings or cartilage, an unusual arrangement shared with the
ovenbirds Ovenbirds or furnariids are a large family of small suboscine passerine birds found from Mexico and Central to southern South America. They form the family Furnariidae. This is a large family containing around 315 species and 70 genera. The oven ...
.


Distribution and habitat

Storks have a nearly
cosmopolitan distribution In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the world in appropriate habitats. Such a taxon, usually a species, is said to exhibit cosmopolitanism or cosmopolitism. The ext ...
, being absent from the poles, most of North America and large parts of Australia. The centres of stork diversity are in tropical Asia and sub-Saharan Africa, with eight and six breeding species respectively. Just three species are present in the New World: wood stork, maguari stork and jabiru, which is the tallest flying bird of the Americas. Two species, white and black stork, reach Europe and western temperate Asia, while one species, Oriental stork, reaches temperate areas of eastern Asia, and one species, black-necked stork, is found in Australasia.del Hoyo, J. Elliott, A. & Sargatal, J. (1992). ''Handbook of the Birds of the World. Volume 1: Ostrich to Ducks. Lynx Edicions''. . Storks are more diverse and common in the tropics, and the species that live in temperate climates for the most part
migrate Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
to avoid the worst of winter. They are fairly diverse in their habitat requirements. Some species, particularly the ''
Mycteria ''Mycteria'' is a genus of large tropical storks with representatives in the Americas, east Africa and southern and southeastern Asia. Two species have "ibis" in their scientific or old common names, but they are not related to these birds and ...
'' "wood storks" and '' Anastomus'' openbills, are highly dependent on water and aquatic prey, but many other species are far less dependent on this habitat type, although they will frequently make use of it. Species like the marabou and Abdim's stork will frequently be found foraging in open grasslands of savannah. Preferred habitats include flooded grasslands, light woodland, marshes and paddyfields, wet meadows, river backwaters and ponds. Many species will select shallow pools, particularly when lakes or rivers are drying out, as they concentrate prey and make it harder for prey to escape, or when monsoonal rainfall increases water depth of larger waterbodies. Some species like the woolly-necked storks and lesser adjutant storks have adapted to changing crops of tropical agricultural landscapes showcasing behavioral plasticity that enables them to remain resident despite the transformations brought about by seasonal crops. In South Africa, the woolly-necked storks have adapted to artificial feeding and now largely nest on trees in gardens with swimming pools. Less typical habitats include the dense temperate forests used by European black storks, or the rainforest habitat sought by Storm's stork in South East Asia. They generally avoid marine habitats, with the exception of the lesser adjutant, milky stork and wood stork, all of which forage in mangroves, lagoons and estuarine mudflats. A number of species, especially woolly-necked storks, black-necked storks, Asian openbills and lesser adjutant Storks in south Asia, have adapted to highly modified human habitats, either for foraging or breeding or both. In the absence of persecution several species breed close to people, and species such as the marabou, greater adjutant, and white stork will feed at landfill sites.


Migration and movements

Storks vary in their tendency towards migration. Temperate species like the white stork, black stork and Oriental stork undertake long annual migrations in the winter. The routes taken by these species have developed to avoid long distance travel across water, and from Europe this usually means flying across the Straits of Gibraltar or east across the Bosphorus and through Israel and the Sinai. Studies of young birds denied the chance to travel with others of their species have shown that these routes are at least partially learnt, rather than being innate as they are in passerine migrants. Migrating black storks are split between those that make stopovers on the migration between Europe and their wintering grounds in Africa, and those that don't. The Abdim's stork is another migrant, albeit one that migrates within the tropics. It breeds in northern Africa, from Senegal to the Red Sea, during the wet season, and then migrates to Southern Africa. Many species that aren't regular migrants will still make smaller movements if circumstances require it; others may migrate over part of their range. This can also include regular commutes from nesting sites to feeding areas. Wood storks have been observed feeding from their breeding colony.


Behaviour


Feeding and diet

Storks are carnivorous predators, taking a range of reptiles, small mammals, insects, fish, amphibians and other small invertebrates. Any plant material consumed is usually by accident. ''Mycteria'' storks are specialists in feeding on aquatic vertebrates, particularly when prey is concentrated by lowering water levels or flooding into shallows. On marine mudflats and mangrove swamps in
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
, milky storks feed on
mudskipper Mudskippers are any of the 23 extant species of amphibious fish from the subfamily Oxudercinae of the goby family Oxudercidae. They are known for their unusual body shapes, preferences for semiaquatic habitats, limited terrestrial locomotion and ...
s, probing the burrow with the bill and even the whole head into the mud. The characteristic feeding method involves standing or walking in shallow water and holding the bill submerged in the water. When contact is made with prey the bill reflexively snaps shut in 25 milliseconds, one of the fastest reactions known in any vertebrate. The reaction is able to distinguish between prey items and inanimate objects like branches, although the exact mechanism is unknown. Openbills are specialists in freshwater molluscs, particularly apple snails. They feed in small groups, and sometimes African openbills ride on the backs of hippos while foraging. Having caught a snail it will return to land or at least to the shallows to eat it. The fine tip of the bill of the openbills is used to open the snail, and the saliva has a narcotic effect, which causes the snail to relax and simplifies the process of extraction. The other genera of storks are more generalised. ''Ciconia'' storks are very generalised in their diets, although Abdim's stork is something of a specialist in feeding in large flocks on swarms of locusts and at wildfires, although other storks will opportunistically feed in this way if the opportunity arises. This is why white storks and Abdim's storks are known as "grasshopper birds". ''Ephippiorhynchus'' are carnivorous though have a very diverse diet when living on human modified habitats such as agricultural landscapes. The foraging method used by the generalists is to stalk or walk across grassland or shallow water, watching for prey.


Breeding

Storks range from being solitary breeders through loose breeding associations to fully
colonial Colonial or The Colonial may refer to: * Colonial, of, relating to, or characteristic of a colony or colony (biology) Architecture * American colonial architecture * French Colonial * Spanish Colonial architecture Automobiles * Colonial (1920 a ...
. The jabiru, ''Ephippiorhynchus'' storks and several species of ''Ciconia'' are entirely solitary when breeding. In contrast the ''Mycteria'' storks, Abdim's stork, openbills and ''Leptoptilos'' storks all breed in colonies which can range from a couple of pairs to thousands. Many of these species breed in colonies with other waterbirds, which can include other species of storks, herons and egrets, pelicans, cormorants and ibises. White storks, Oriental storks and Maguari storks are all loosely colonial, and may breed in nests that are within visual range of others of the same species, but have little to do with one another. They also may nest solitarily, and the reasons why they choose to nest together or apart are not understood.


Systematics

A DNA study found that the families Ardeidae,
Balaenicipitidae Balaenicipitidae is a family of birds in the order Pelecaniformes, although it was traditionally placed in Ciconiiformes. The shoebill is the sole extant species and its closest relative is the hamerkop (''Scopus umbretta''), which belongs to ano ...
,
Scopidae ''Scopus'' is a genus of wading birds containing the hamerkop ''(Scopus umbretta)'' and its extinct Pliocene relative, ''Scopus xenopus''. This genus is the sole representative of the family Scopidae. Taxonomy Hamerkops were traditionally includ ...
and the Threskiornithidae belong to the Pelecaniformes. This would make Ciconiidae the only group. Storks were distinct and possibly widespread by the
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
. Like most families of aquatic birds, storks seem to have arisen in the Palaeogene, maybe 40–50 million years ago (mya). For the fossil record of living genera, documented since the
Middle Miocene The Middle Miocene is a sub-epoch of the Miocene Epoch made up of two stages: the Langhian and Serravallian stages. The Middle Miocene is preceded by the Early Miocene. The sub-epoch lasted from 15.97 ± 0.05 Ma to 11.608 ± 0.005 Ma (million y ...
(about 15 mya) at least in some cases, see the genus articles. Though some storks are highly threatened, no species or subspecies are known to have gone
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
in historic times. A ''Ciconia'' bone found in a
rock shelter A rock shelter (also rockhouse, crepuscular cave, bluff shelter, or abri) is a shallow cave-like opening at the base of a bluff or cliff. In contrast to solutional caves (karst), which are often many miles long, rock shelters are almost alway ...
on the island of
Réunion Réunion (; french: La Réunion, ; previously ''Île Bourbon''; rcf, label= Reunionese Creole, La Rényon) is an island in the Indian Ocean that is an overseas department and region of France. It is located approximately east of the island ...
was probably of a bird taken there as food by early settlers; no known account mentions the presence of storks on the Mascarene Islands.


Extant storks


Fossil storks

*Genus ''
Palaeoephippiorhynchus Palaeoephippiorhynchus is an extinct genus of large stork from the Oligocene of Africa: its closest living relative is the Saddle-billed stork The saddle-billed stork or saddlebill (''Ephippiorhynchus senegalensis'') is a large wading bird in t ...
'' ( fossil: Early Oligocene of Fayyum, Egypt) *Genus ''Grallavis'' ( fossil: Early Miocene of Saint-Gérand-le-Puy, France, and Djebel Zelten, Libya) – may be same as ''Prociconia'' *Ciconiidae gen. et sp. indet. (Ituzaingó Late Miocene of Paraná, Argentina)
Tarsometatarsus The tarsometatarsus is a bone that is only found in the lower leg of birds and some non-avian dinosaurs. It is formed from the fusion of several bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) and meta ...
fragments somewhat similar to ''Mycteria''
Noriega, Jorge Ignacio & Cladera, Gerardo (2005). ''First Record of Leptoptilini (Ciconiiformes: Ciconiidae) in the Neogene of South America''. Abstracts of Sixth International Meeting of the Society of Avian Paleontology and Evolution: 47
PDF fulltext
*Ciconiidae gen. et sp. indet. (Puerto Madryn Late Miocene of Punta Buenos Aires, Argentina)Specimen MEF 1363: Incomplete skeleton of a large stork somewhat similar to ''Jabiru'' but apparently more plesiomorphic *Genus ''Prociconia'' ( fossil: Late Pleistocene of Brazil) – may belong to modern genus ''Jabiru'' or ''Ciconia'' *Genus ''Pelargosteon'' ( fossil: Early Pleistocene of Romania) *Ciconiidae gen. et sp. indet. – formerly ''Aquilavus/Cygnus bilinicus'' ( fossil: Early Miocene of Břešťany, Czech Republic) *cf. ''Leptoptilos'' gen. et sp. indet. – formerly ''L. siwalicensis'' ( fossil: Late Miocene? – Late Pliocene of Siwalik, India)Specimens BMNH 39741 (holotype, left proximal
tarsometatarsus The tarsometatarsus is a bone that is only found in the lower leg of birds and some non-avian dinosaurs. It is formed from the fusion of several bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) and meta ...
) and BMNH 39734 (right
distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
tibiotarsus). Similar to ''Ephippiorhynchus'' and ''Leptotilos'', may be from a small female of ''Leptotilos falconeri'', from ''L. dubius'', or from another species:
*Ciconiidae gen. et sp. indet. ( fossil: Late Pleistocene of San Josecito Cavern, Mexico)
Distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
radius of a mid-sized ''Ciconia'' or smallish ''Mycteria'':
The fossil genera ''Eociconia'' (Middle Eocene of China) and ''Ciconiopsis'' (Deseado Early Oligocene of Patagonia, Argentina) are often tentatively placed with this family. A "ciconiiform" fossil fragment from the Touro Passo Formation found at Arroio Touro Passo (
Rio Grande do Sul Rio Grande do Sul (, , ; "Great River of the South") is a Federative units of Brazil, state in the South Region, Brazil, southern region of Brazil. It is the Federative_units_of_Brazil#List, fifth-most-populous state and the List of Brazilian st ...
, Brazil) might be of the living wood stork ''M. americana''; it is at most of
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...
age, a few 10,000s of years.Schmaltz Hsou, Annie (2007). ''O estado atual do registro fóssil de répteis e aves no Pleistoceno do Estado do Rio Grande do Sul, Brasil'' The current state of the fossil record of Pleistocene reptiles and birds of Rio Grande do Sul" Talk held on 2007-JUN-20 at ''Quaternário do RS: integrando conhecimento'', Canoas, Rio Grande do Sul, Brazil
PDF abstract


Etymology

The Modern English word can be traced back to Proto-Germanic ''*sturkaz''. Nearly every Germanic language has a descendant of this proto-language word to indicate the ( white) stork. Related names also occur in Latvian, ''stārķis'', and some Slavic languages, e.g. ''štorklja'' in Slovenian and “щъркел” htŭrkelin Bulgarian, originating as Germanic loanwords. According to the ''New Shorter Oxford English Dictionary'', the Germanic root is probably related to the modern English "stark", in reference to the stiff or rigid posture of a European species, the white stork. A non-Germanic word linked to it may be Greek ''torgos'' ("vulture"). In some West Germanic languages cognate words of a different etymology exist, e.g. '' ooievaar'' in Dutch. They originate from ''*uda-faro'', ''uda'' being related to ''water'' meaning something like ''swamp'' or ''moist area'' and ''faro'' being related to ''fare''; so ''*uda-faro'' is something like ''he who walks in the swamp''. In later times this name got reanalysed as ''*ōdaboro'', ''ōda'' "fortune, wealth" + ''boro'' "bearer" meaning ''he who brings wealth'' adding to the myth of storks as maintainers of welfare and bringers of children. In
Estonian Estonian may refer to: * Something of, from, or related to Estonia, a country in the Baltic region in northern Europe * Estonians, people from Estonia, or of Estonian descent * Estonian language * Estonian cuisine * Estonian culture See also

...
, "stork" is ''toonekurg'', which is derived from ''toonela'' ( underworld in Estonian folklore) + ''kurg'' ( crane). At the times storks were named, the now-rare black stork was probably the more common species.


In fiction

Storks have many stories surrounding them, like in Aesop's (6th century BCE) fables The Farmer and the Stork and The Fox and the Stork. The first fable begins with a farmer plowing his fields, sowing his seeds and spreading his nets. These nets catch several
cranes Crane or cranes may refer to: Common meanings * Crane (bird), a large, long-necked bird * Crane (machine), industrial machinery for lifting ** Crane (rail), a crane suited for use on railroads People and fictional characters * Crane (surname), ...
who hopped behind him picking up the seed. Along with the cranes tangled in his net, the farmer discovered a stork with a broken leg. The stork begged the farmer to spare his life, arguing that he was not a crane, but a stork. He pointed to his feathers and told the farmer that they didn't resemble a crane's feathers in the least. The farmer laughed at the stork and said, "I have taken you with these robbers, the cranes, and you must die in their company." A common euphemism is that storks deliver babies to their mothers instead of mothers
giving birth Childbirth, also known as labour and delivery, is the ending of pregnancy where one or more babies exits the internal environment of the mother via vaginal delivery or caesarean section. In 2019, there were about 140.11 million births globally ...
. Storks play a prominent role in Hans Christian Andersen's lengthy fairy tale, "The Marsh King's Daughter."


Footnotes


References


External links

* *
Scott MacDonald, "The Stork"
emblematic uses

Image documentation
Stork videos
on the Internet Bird Collection

Image documentation
Storks vocalizing
{{Authority control Extant Rupelian first appearances Taxa named by John Edward Gray fr:Cigogne