Wikipedia

Stork

For other uses, seeStork (disambiguation).

Storksare large, long-legged, long-neckedwading birdswith long, stoutbills.They belong to thefamilyCiconiidae,and make up the orderCiconiiformes/sɪˈkoʊni.ɪfɔːrmiːz/.Ciconiiformes previously included a number of other families, such asheronsandibises,but those families have been moved to other orders.[2]

Stork
Temporal range:Early Oligoceneto present30–0Ma
White stork(Ciconia ciconia)
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Clade: Aequornithes
Order: Ciconiiformes
Bonaparte,1854[1]
Family: Ciconiidae
J. E. Gray,1840[1]
Genera

Storks dwell in many regions and tend to live in drier habitats than the closely relatedherons,spoonbillsandibises;they also lack thepowder downthat those groups use to clean offfishslime. Bill-clattering is an important mode ofcommunicationat the nest. Many species aremigratory.Most storks eatfrogs,fish,insects,earthworms,smallbirdsand smallmammals.There are 20 livingspeciesof storks in sixgenera.

Various terms are used to refer to groups of storks,[3]two frequently used ones being amusterof storks and aphalanxof storks.

Storks tend to usesoaring,gliding flight, which conserves energy. Soaring requiresthermalair currents.Ottomar Anschütz's famous 1884 album of photographs of storks inspired the design ofOtto Lilienthal's experimentalglidersof the late nineteenth century. Storks are heavy, with widewingspans:themarabou stork,with a wingspan of 3.2 m (10 ft 6 in) and weight up to 8 kg (18 lb), joins theAndean condorin having the widest wingspan of all living land birds.

Theirnestsare often very large and may be used for many years. Some nests have been known to grow to over 2 metres (6 ft 7 in) in diameter and about 3 metres (9.8 ft) in depth. All storks were once thought to bemonogamous,but this is only partially true. While storks are generally socially monogamous, some species exhibit regular extra-pair breeding.[4]

Popular conceptions of storks' fidelity,serial monogamy,and doting parental care contribute to their prominence inmythologyand culture, especially in western folklore as the deliverers of newborn humans.[5]

All 20 stork species have been assessed by the IUCN and carry a confident Red List status. However, the assessment for several species were based on incorrect assumptions and a general absence of sound information on stork habits.[6]

Etymology

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The word "stork" was first used in its current sense by at least the 12th century inMiddle English.[7]It is derived from theOld Englishword "storc", which itself comes from the hypothesisedProto-Germanic*storkand ultimately theProto-Indo-European*sr̥ǵos.[7][8]The name refers to the rigid posture of storks, a meaning reflected in the related wordstark,which is derived from the Old English "stearc".[9]Several species of storks are known by other common names. Thejabiruis named after theTupí-Guaraniwords meaning "that which has" and "swollen", referring to its thickset neck.[10]Themarabou storkis named after the Arabic word for holy man,murābiṭ,due to the perceived holy nature of the species.[11]Theadjutantsare named afterthe military rank,referring to their stiff, military-like gait.[12]

Systematics

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See also:List of storks

A DNA study found that the familiesArdeidae,Balaenicipitidae,Scopidaeand theThreskiornithidaebelong to thePelecaniformes.This would makeCiconiidaethe only group.[13][14]

Storks were distinct and possibly widespread by theOligocene.Like most families of aquatic birds, storks seem to have arisen in thePalaeogene,maybe 40–50million years ago(mya). For the fossil record of living genera, documented since theMiddle Miocene(about 15 mya) at least in some cases, see the genus articles.

No species or subspecies of stork is known to have goneextinctin historic times. A systematic literature review uncovered nearly 1,000 papers on storks, but showed most stork species to lack scientific understanding suggesting that many species should be classified as Data Deficient on the IUCN Red List. ACiconiabone found in arock shelteron the island ofRéunionwas probably of a bird taken there as food by early settlers; no known account mentions the presence of storks on theMascarene Islands.

Phylogeny

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The following phylogeny is recognized by the International Ornithological Congress, partially based on de Sousaet al(2023):[15][16]

African openbill(Anastomus lamelligerus)

Asian openbill(Anastomus oscitans)

Marabou stork(Leptoptilos crumenifer)

Lesser adjutant(Leptoptilos javanicus)

Greater adjutant(Leptoptilos dubius)

Wood stork(Mycteria americana)

Yellow-billed stork(Mycteria ibis)

Painted stork(Mycteria leucocephala)

Milky stork (Mycteria cinerea)

Jabiru(Jabiru mycteria)

Saddle-billed stork(Ephippiorhynchus senegalensis)

Black-necked stork(Ephippiorhynchus asiaticus)

Abdim's stork(Ciconia abdimii)

Asian woolly-necked stork(Ciconia episcopus)

Storm's stork(Ciconia stormi)

Black stork(Ciconia nigra)

Maguari stork(Ciconia maguari)

African woolly-necked stork(Ciconia microscelis)

White stork(Ciconia ciconia)

Oriental stork(Ciconia boyciana)

Fossil storks

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  • GenusPalaeoephippiorhynchus(fossil:Early Oligocene ofFayyum,Egypt)
  • GenusGrallavis(fossil:Early Miocene of Saint-Gérand-le-Puy, France, and Djebel Zelten, Libya) – may be same asProciconia
  • Ciconiidae gen. et sp. indet. (Ituzaingó Late Miocene of Paraná, Argentina)[note 1][17][18]
  • Ciconiidae gen. et sp. indet. (Puerto Madryn Late Miocene of Punta Buenos Aires, Argentina)[note 2][18]
  • GenusProciconia(fossil:Late Pleistocene of Brazil) – may belong to modern genusJabiruorCiconia
  • GenusPelargosteon(fossil:Early Pleistocene of Romania)
  • Ciconiidae gen. et sp. indet. – formerlyAquilavus/Cygnus bilinicus(fossil:Early Miocene of Břešťany, Czech Republic)
  • cf.Leptoptilosgen. et sp. indet. – formerlyL. siwalicensis(fossil:Late Miocene? – Late Pliocene of Siwalik, India)[19]
  • Ciconiidae gen. et sp. indet. (fossil:Late Pleistocene of San Josecito Cavern, Mexico)[20]
  • Ciconia nana(fossil:Pleistocene of Darling Downs, Queensland, Australia, andCiconia louisebolesaefossil:Olig-Miocene of Riversleigh WHA, Queensland, Australia[21]

The fossil generaEociconia(Middle Eocene of China) andCiconiopsis(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,Brazil) might be of the livingwood storkM. americana;it is at most ofLate Pleistoceneage, a few 10,000s of years.[22]

European white stork in a nest in Bisag,Croatia

Morphology

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Mycteriastorks, like this yellow-billed stork, have sensitive bills that allow them to hunt by touch

Storks range in size from the marabou, which stands 152 cm (60 in) tall and can weigh8.9 kg (19+1⁄2lb), to the Abdim's stork, which is only 75 cm (30 in) high and weighs only1.3 kg (2+3⁄4lb). Their shape is superficially similar to the herons, with long legs and necks, but they are more heavy-set. There is somesexual 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 theirisof the two species in the genusEphippiorhynchus.[23]

Thebillsof 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 theCiconiastorks are the least specialized. Larger are the massive and slightly upturned bills of theEphippiorhynchusand 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 defense against other scavengers, as well as for taking other prey.[23]The long, ibis-like downcurved bills of theMycteriastorks have sensitive tips that allow them to detect prey by touch (tactilocation) where cloudy conditions would not allow them to see it.[24]The most specialised bills of any storks are those of the two openbills (Anastomus), which as their name suggests, is open in the middle when their bill is closed. These bills have evolved to help openbills feed on their primary prey item, aquatic snails.[25]

Although it is sometimes reported that storks lacksyrinxesand are mute,[26]they do have syrinxes,[27]and are capable of making some sounds, although they do not do so often.[23][28]The syrinxes of storks are "variably degenerate" however,[27]and the syringeal membranes of some species are found between tracheal rings or cartilage, an unusual arrangement shared with theovenbirds.[29]

Distribution and habitat

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Lesser adjutantswill forage in marine habitats, unlike most storks
Maraboustork atEtosha National ParkinNamibia
See also:List of Ciconiiformes by population

Storks have a nearlycosmopolitan distribution,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 storkandjabiru,which is the tallest flying bird of the Americas. Two species,whiteandblack 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.[23]

Storks are more diverse and common in the tropics, and the species that live in temperate climates for the most partmigrateto avoid the worst of winter. They are fairly diverse in their habitat requirements. Some species, particularly theMycteria"wood storks" andAnastomusopenbills, 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 themarabouandAbdim's storkwill 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.[23][30]Some species like the woolly-necked storks and lesser adjutant storks have adapted to changing crops of tropical agricultural landscapes that enables them to remain resident despite the transformations brought about by seasonal crops.[31][32][33][34]In South Africa, the woolly-necked storks have adapted to artificial feeding and now largely nest on trees in gardens with swimming pools.[35]

Less typical habitats include the dense temperate forests used by Europeanblack storks,or the rainforest habitat sought byStorm's storkin South East Asia. They generally avoid marine habitats, with the exception of thelesser adjutant,milky storkandwood 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, for foraging and breeding.[30][36][37][38]In the absence of persecution several stork species breed close to people, and species such as the marabou, greater adjutant, and white stork feed at landfill sites.[23][39]

Migration and movements

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Abdim's storks are regular intra-African migrants

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.[23]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 inpasserinemigrants.[40]Migrating black storks are split between those that make stopovers on the migration between Europe and their wintering grounds in Africa, and those that do not.[41]

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.[42]Many species that are not 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 130 km (80 mi) from their breeding colony.[23]

Behaviour

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Feeding and diet

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African openbill foraging in shallow water

Storks are carnivorous predators, taking a range of reptiles, small mammals, insects, fish, amphibians and other small invertebrates. Storks usually hunt for animals in shallow water. Any plant material consumed is usually by accident.Mycteriastorks 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 inSumatra,milky storks feed onmudskippers,probing the burrow with the bill and even the whole head into the mud.[citation needed]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.[43][23]

Openbills are specialists in freshwater molluscs, particularly apple snails. They feed in small groups,[30]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.[23]

The other genera of storks are more generalised.Ciconiastorks are very generalised in their diets, and some species including Abdim's stork and marabous will feed in large flocks on swarms of locusts and at wildfires.[23][44]This is why white storks and Abdim's storks are known as "grasshopper birds".Ephippiorhynchusare carnivorous though have a very diverse diet when living on human modified habitats such as agricultural landscapes.[45]The foraging method used by the generalists is to stalk or walk across grassland or shallow water, watching for prey.[23]

Breeding

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Storks range from being solitary breeders through loose breeding associations to fullycolonial.The jabiru,Ephippiorhynchusstorks and several species ofCiconiaare entirely solitary when breeding.[38][46][47]In contrast theMycteriastorks, Abdim's stork, openbills andLeptoptilosstorks breed in colonies which can range from a couple of pairs to thousands.[31][36][37]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.[23]Storks use trees in a variety of habitats to breed including forests, cities, farmlands, and large wetlands.[32][35][36][48]

In culture

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Storks feature in several ofAesop's Fables,such asThe Fox and the Stork

Many ancient mythologies feature stories and legends involving storks. InAncient Egypt,saddle-billed storks were seen as being amongst the most powerful animals and were used to represent theba,the Ancient Egyptian conception of the soul, during theOld Kingdom.[49]Bennu,anEgyptian deitythat was later the inspiration for thephoenix,may also have been inspired by a stork, although it was more likely an ibis or heron.[50][51]

GreekandRoman mythologyportrays storks as models of parental devotion. The 3rd century Roman writerAelian,citing the authority ofAlexander of Myndus,noted in hisDe natura animalium(book 3, chapter 23) that aged storks flew away to oceanic islands where they were transformed into humans as a reward for their piety towards their parents.[52]Storks were also thought to care for their aged parents, feeding them and even transporting them, and children's books depicted them as a model of filial values. A Greek law calledPelargonia,from theAncient Greekwordpelargosfor stork, required citizens to take care of their aged parents.[citation needed]The Greeks also held that killing a stork could be punished with death.[53]

Storks feature in several of Aesop's Fables, most notably inThe Farmer and the Stork,The Fox and the Stork,andThe Frogs Who Desired a King.The first fable involves a stork who is caught with a group of cranes who are eating grain in a farmer's field, with the moral that those who associate with wicked people can be held accountable for their crimes.[54]The Fox and the Stork involves a fox who invites a stork for dinner and provides soup in a dish that the stork cannot drink from, and is in turn invited for dinner by the stork and given food in a narrow jug which he cannot access. It cautions readers to follow the principle ofdo no harm.[55]The third fable involves a group of frogs that are dissatisfied with the king that Zeus has given them, an inanimate log, and who are then punished with a newKing Stork(awater-snakein some versions) who eats the frogs.[56][57]King Storkhas subsequently entered the English language as a term for a particularly tyrannical ruler.[58]

Associations with fertility

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According to European folklore, the white stork is responsible for bringing babies to new parents. The legend is very ancient, but was popularised by an 1839Hans Christian Andersenstory called "The Storks".[59]German folklore held that storks found babies in caves or marshes and brought them to households in a basket on their backs or held in their beaks. These caves containedadebarsteineor "stork stones". The babies would then be given to the mother or dropped down the chimney. Households would notify when they wanted children by placing sweets for the stork on the window sill.[60]Subsequently, the folklore has spread around the world to thePhilippinesand countries inSouth America.[60]Birthmarks on the back of the head of newborn babies,nevus flammeus nuchae,are sometimes referred to as stork-bite.[61]InSlavic mythology and pagan religion,storks were thought to carry unbornsoulsfromVyrajtoEarthin spring and summer.[62]This belief still persists in the modern folk culture of many Slavic countries, in the simplified child story that "storks bring children into the world".[63][64]

Psychoanalyst Marvin Margolis suggests the enduring nature of the stork fable of the newborn is linked to its addressing a psychological need, in that it allays the discomfort of discussing sex and procreation with children. Birds have long been associated with the maternal symbols from pagan goddesses such asJunoto theHoly Ghost,and the stork may have been chosen for its white plumage (depicting purity), size, and flight at high altitude (likened to flying between Earth and Heaven).[60]

There were negative aspects to stork folklore as well; a Polish folktale relates how God made the stork's plumage white, while the Devil gave it black wings, imbuing it with both good and evil impulses. They were also associated with handicapped or stillborn babies in Germany, explained as the stork having dropped the baby en route to the household, or as revenge or punishment for past wrongdoing. A mother who was confined to bed around the time of childbirth was said to have been "bitten" by the stork. In Denmark, storks were said to toss a nestling off the nest and then an egg in successive years.[60]In medieval England, storks were also associated with adultery, possibly inspired by their courtship rituals. Their preening and posture saw them linked with the attribute of self-conceit.[65]Children ofAfrican Americanslaveswere sometimes told that white babies were brought by storks, while black babies were born frombuzzardeggs.[66]

As food

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Storks have never been a particularly common food, but occasionally featured in medievalbanquets.[67]They may also have been eaten in Ancient Egypt.[68]

Footnotes

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  1. ^Tarsometatarsusfragments somewhat similar toMycteria
  2. ^SpecimenMEF1363: Incomplete skeleton of a large stork somewhat similar toJabirubut apparently moreplesiomorphic

References

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  1. ^abBrands, Sheila (14 August 2008)."Systema Naturae 2000 / Classification, Class Aves".Project: The Taxonomicon.Retrieved11 June2012.
  2. ^"Anastomus lamelligerus subsp. lamelligerus".www.gbif.org.Archived fromthe originalon 2018-08-03.Retrieved2018-03-22.
  3. ^About the Wood Stork: Denizens of the WetlandsArchived2011-07-27 at theWayback Machine,Accessed on 13.12.2010
  4. ^Turjeman, Sondra F.; Centeno-Cuadros, Alejandro; Eggers, Ute; Rotics, Shay; Blas, Julio; Fielder, Wolfgang; Kaatz, Michael; Jeltsch, Florian; Wikelski, Martin; Ran, Nathan (2016)."Extra-pair paternity in the socially monogamous white stork (Ciconia ciconia) is fairly common and independent of local density".Scientific Reports.6:27976.Bibcode:2016NatSR...627976T.doi:10.1038/srep27976.PMC4916429.PMID27328982.
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  19. ^SpecimensBMNH39741 (holotype, leftproximaltarsometatarsus) and BMNH 39734 (rightdistaltibiotarsus). Similar toEphippiorhynchusandLeptotilos,may be from a small female ofLeptotilos falconeri,fromL. dubius,or from another species:Louchart, Antoine; Vignaud, Patrick; Likius, Andossa; Brunet, Michel & White, Tim D. (2005)."A large extinct marabou stork in African Pliocene hominid sites, and a review of the fossil species ofLeptoptilos"(PDF).Acta Palaeontologica Polonica.50(3): 549–563. Archived fromthe original(PDF)on 2008-02-28.Retrieved2008-01-15.
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