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The stork also carries several types of internal parasites, including ''[[Toxoplasma gondii]]''<ref>{{cite book |title=Foodborne Parasites |first=Ynes R. |last=Ortega |year=2006 |location=New York |publisher=Springer |page=121 |isbn=0387300686}}</ref> and intestinal parasites of the genus ''[[Giardia]]''.<ref>{{cite journal |title=Giardiasis in a White Stork in The Netherlands |first1=F. F. |last1=Franssen |first2=J |last2=Hooimeijer |first3=B. |last3=Blankenstein |first4=Dirk J. |last4=Houwers |journal=Journal of Wildlife Diseases |volume=36 |issue=4|pages=764–66 |year=2000 |format=abstract |url= |pmid=11085441}}</ref> A study of 120 White Stork carcasses from Saxonia-Anhalt and Brandenburg in Germany yielded eight species of [[trematoda|trematode]] (fluke), four [[cestoda|cestode]] (tapeworm) species, and at least three species of [[nematoda|nematode]].<ref>{{cite journal|last=Schuster, R.; Schaffer, T.; Shimalov, V.|year=2002|title=[The helminth fauna of indigenous white storks (Ciconia ciconia)].|journal=Berl Munch Tierarztl Wochenschr.|volume=115|pages=435–39|pmid=12481650|language=German}}</ref> One species of fluke, ''[[Chaunocephalus ferox]]'', caused lesions in the wall of the small intestine in a number of birds admitted to two rehabilitation centers in central Spain, and was associated with reduced weight. It is a recognised pathogen and cause of morbidity in the [[Asian Openbill]] (''Anastomus oscitans'').<ref>{{cite journal|last=Höfle, Ursula; Krone, Oliver; Blanco, Juan Manuel; Pizarro, Manuel|year=2003|title=''Chaunocephalus ferox'' in Free-Living White Storks in Central Spain|journal=Avian Diseases|volume=47|issue=2|pages=506–12|doi=10.1637/0005-2086(2003)047[0506:CFIFWS]2.0.CO;2}}</ref>
The stork also carries several types of internal parasites, including ''[[Toxoplasma gondii]]''<ref>{{cite book |title=Foodborne Parasites |first=Ynes R. |last=Ortega |year=2006 |location=New York |publisher=Springer |page=121 |isbn=0387300686}}</ref> and intestinal parasites of the genus ''[[Giardia]]''.<ref>{{cite journal |title=Giardiasis in a White Stork in The Netherlands |first1=F. F. |last1=Franssen |first2=J |last2=Hooimeijer |first3=B. |last3=Blankenstein |first4=Dirk J. |last4=Houwers |journal=Journal of Wildlife Diseases |volume=36 |issue=4|pages=764–66 |year=2000 |format=abstract |url= |pmid=11085441}}</ref> A study of 120 White Stork carcasses from Saxonia-Anhalt and Brandenburg in Germany yielded eight species of [[trematoda|trematode]] (fluke), four [[cestoda|cestode]] (tapeworm) species, and at least three species of [[nematoda|nematode]].<ref>{{cite journal|last=Schuster, R.; Schaffer, T.; Shimalov, V.|year=2002|title=[The helminth fauna of indigenous white storks (Ciconia ciconia)].|journal=Berl Munch Tierarztl Wochenschr.|volume=115|pages=435–39|pmid=12481650|language=German}}</ref> One species of fluke, ''[[Chaunocephalus ferox]]'', caused lesions in the wall of the small intestine in a number of birds admitted to two rehabilitation centers in central Spain, and was associated with reduced weight. It is a recognised pathogen and cause of morbidity in the [[Asian Openbill]] (''Anastomus oscitans'').<ref>{{cite journal|last=Höfle, Ursula; Krone, Oliver; Blanco, Juan Manuel; Pizarro, Manuel|year=2003|title=''Chaunocephalus ferox'' in Free-Living White Storks in Central Spain|journal=Avian Diseases|volume=47|issue=2|pages=506–12|doi=10.1637/0005-2086(2003)047[0506:CFIFWS]2.0.CO;2}}</ref>


On August 26, 1998, a flock of about 1,200 migrating White Storks that had been blown off course on their southward journey landed in [[Eilat]], in southern Israel. The flock was stressed as it had resorted to flapping flight to return to its migratory route, and a number of birds died. A virulent strain of [[West Nile virus]] was isolated from the brains of eleven dead juveniles. Other White Storks subsequently tested in Israel have shown anti-WNV antibodies.<ref>{{cite journal | title=Introduction of West Nile virus in the Middle East by Migrating White Storks|author=Malkinson, Mertyn; Banet, Caroline; Weisman, Yoram; Pokamunski, Shimon; King, Roni; Drouet, Marie-Thérèse; Deubel, Vincent | journal=Emerging Infectious Diseases|year=2002|volume=8|issue=4|pages=392-97|url=http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2730252/|accessdate=15 February 2011}}</ref> In 2008 three juvenile White Storks from a Polish wildlife refuge yielded seropositive results indicating exposure to the virus, but the context or existence of the virus in that country is unclear.<ref>{{cite journal |url= |journal=Viral Immunology |volume=21 |issue=2 |pages=247–54 |first1=Zdenek |last1=Hubálek |first2=Elżbieta |last2=Wegner |year= 2008 |first3=Jiří et al |last4= Tryjanowski |last3=Halouzka |first4= Piotr |last5= Jerzak |first5= Leszek |last6= Šikutová |first6= Silvie |last7= Rudolf |first7= Ivo |last8= Kruszewicz |first8= Andrzej G. |last9= Jaworski |first9= Zbigniew |title=Serologic survey of potential vertebrate hosts for West Nile Virus in Poland |format= |doi=10.1089/vim.2007.0111}}</ref> Migrating birds appear to be important in spread of the virus,<ref>{{cite journal|last=Malkinson, Mertyn; Banet, Caroline|year=2002|title=The role of birds in the ecology of West Nile virus in Europe and Africa|journal=Current Topics in Microbiology and Immunology|volume=267|pages=309–22|pmid=12082995}}</ref> the ecology of which remains poorly known.<ref name="Zeller2004">{{cite journal|last=Zeller|first=Hervé G.|coauthors=Schuffenecker, Isabelle|year=2004|title=West Nile Virus: An Overview of Its Spread in Europe and the Mediterranean Basin in Contrast to Its Spread in the Americas|journal=European Journal of Clinical Microbiology & Infectious Diseases|volume=23|issue=3|pages=147–56|issn=0934-9723|doi=10.1007/s10096-003-1085-1}}</ref>
[[West Nile virus]] (WNV) is mainly a bird infection that is transmitted between birds by [[Mosquito]]s.<ref>{{cite journal |author=Hayes EB, Komar N, Nasci RS, Montgomery SP, O'Leary DR, Campbell GL |title=Epidemiology and transmission dynamics of West Nile virus disease |journal=Emerging Infect. Dis. |volume=11 |issue=8 |pages=1167–73 |year=2005 |pmid=16102302 |url=http://www.cdc.gov/ncidod/EID/vol11no08/05-0289a.htm}}</ref> Migrating birds appear to be important in spread of the virus,<ref>{{cite journal|last=Malkinson, Mertyn; Banet, Caroline|year=2002|title=The role of birds in the ecology of West Nile virus in Europe and Africa|journal=Current Topics in Microbiology and Immunology|volume=267|pages=309–22|pmid=12082995}}</ref> the ecology of which remains poorly known.<ref name="Zeller2004">{{cite journal|last=Zeller|first=Hervé G.|coauthors=Schuffenecker, Isabelle|year=2004|title=West Nile Virus: An Overview of Its Spread in Europe and the Mediterranean Basin in Contrast to Its Spread in the Americas|journal=European Journal of Clinical Microbiology & Infectious Diseases|volume=23|issue=3|pages=147–56|issn=0934-9723|doi=10.1007/s10096-003-1085-1}}</ref>
On August 26, 1998, a flock of about 1,200 migrating White Storks that had been blown off course on their southward journey landed in [[Eilat]], in southern Israel. The flock was stressed as it had resorted to flapping flight to return to its migratory route, and a number of birds died. A virulent strain of [[West Nile virus]] was isolated from the brains of eleven dead juveniles. Other White Storks subsequently tested in Israel have shown anti-WNV antibodies.<ref>{{cite journal | title=Introduction of West Nile virus in the Middle East by Migrating White Storks|author=Malkinson, Mertyn; Banet, Caroline; Weisman, Yoram; Pokamunski, Shimon; King, Roni; Drouet, Marie-Thérèse; Deubel, Vincent | journal=Emerging Infectious Diseases|year=2002|volume=8|issue=4|pages=392-97|url=http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2730252/|accessdate=15 February 2011}}</ref> In 2008 three juvenile White Storks from a Polish wildlife refuge yielded seropositive results indicating exposure to the virus, but the context or existence of the virus in that country is unclear.<ref>{{cite journal |url= |journal=Viral Immunology |volume=21 |issue=2 |pages=247–54 |first1=Zdenek |last1=Hubálek |first2=Elżbieta |last2=Wegner |year= 2008 |first3=Jiří et al |last4= Tryjanowski |last3=Halouzka |first4= Piotr |last5= Jerzak |first5= Leszek |last6= Šikutová |first6= Silvie |last7= Rudolf |first7= Ivo |last8= Kruszewicz |first8= Andrzej G. |last9= Jaworski |first9= Zbigniew |title=Serologic survey of potential vertebrate hosts for West Nile Virus in Poland |format= |doi=10.1089/vim.2007.0111}}</ref>


== Conservation ==
== Conservation ==

Revision as of 22:31, 15 February 2011

White Stork
Adult at Karlsruhe Zoo, Germany
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Family:
Genus:
Species:
C. ciconia
Binomial name
Ciconia ciconia
(Linnaeus, 1758)
Approximate ranges and routes

   Breeding range
   Winter range

  Migration routes
Synonyms

Ardea ciconia L.

The White Stork (Ciconia ciconia) is a large wading bird in the stork family Ciconiidae. Its plumage is mainly white, with black on its wings. Adults have long red legs and a long pointed red beak. Its two subspecies, which differ slightly in size, breed in the warmer parts of Europe (north to Estonia), northwestern Africa, southwestern Asia (east to southern Kazakhstan), and southern Africa. It is a long-distance migrant, wintering in Africa from tropical Sub-Saharan Africa to as far south as South Africa, or on the Indian subcontinent.

A carnivore, the White Stork eats a wide range of animal prey, including insects, crayfish, fish, amphibians, reptiles, small mammals and small birds. It takes most of its food from the ground, among low vegetation, and from shallow water. It is a monogamous breeder, but does not pair for life. Both members of the pair build a large stick nest, which may be used for several years. The female lays one clutch of eggs per year usually consisting four eggs, which hatch asynchronously 33–34 days later. Both parents take turns incubating the eggs and both feed the young. The young leave the nest 58–64 days after hatching, and they continue to be fed by the parents for a further 7–20 days.

The White Stork benefited from human activities during the Middle Ages as woodland was cleared; however, a decline in population began in the 19th Century due to changes in farming methods. Continuing threats to the species include the drainage of wetlands, collisions with overhead power lines, use of persistent pesticides, and illegal hunting on migration routes and wintering grounds. It has few natural predators, but may harbour several types of parasite. This conspicuous bird has given rise to many legends across its range, of which the best known is the story of babies being brought by storks.

Taxonomy and evolution

The White Stork was one of the many species originally described by Linnaeus in his 18th century work, Systema Naturae,[2] where it was given the binomial name of Ardea ciconia. It was moved to (and was designated the type species of) the new genus Ciconia by French zoologist Mathurin Jacques Brisson two years later,[3][4] Both the genus and the species names, ciconia, are the Latin word for "stork".[5] The word stork is derived from the Old English word storc, related to the Old High German storah, "stork", and similar words in many other European languages, all of which are descended from the Teutonic sturko-z.[6]

There are two subspecies of the White Stork:

  • C. c. ciconia, the nominate subspecies described by Linnaeus in 1758, breeds from Europe to northwest Africa and westernmost Asia, and in southern Africa, and winters mainly in Africa south of the Sahara Desert.[7] Some birds however winter in India.[8]
  • C. c. asiatica, which was described by Russian naturalist Nikolai Severtzov in 1873, breeds in Turkestan and winters from Iran to India. It is slightly larger than the nominate subspecies.[7][9]

The stork family contains six genera in three major groups: the open-billed and wood storks (Mycteria and Anastomus), the giant storks (Ephippiorhynchus, Jabiru and Leptoptilos), and the "typical storks", Ciconia. The typical storks include the White Stork and six other extant species,[10] which are characterised by straight pointed beaks and mainly black and white plumage.[11] The White Stork's closest relatives are the larger black-billed Oriental White Stork (Ciconia boyciana) of East Asia, which was formerly classified as a subspecies of the White Stork,[7] and the Maguari Stork (C. maguari) of South America. Close evolutionary relationships within the Ciconia genus are suggested by behavioural similarities and biochemically through analysis of both mitochondrial cytochrome b gene sequences and DNA-DNA hybridization.[12] A Ciconia fossil representing the distal end of a right humerus has been recovered from Miocene beds of Rusinga Island, Lake Victoria, Kenya.[13] The 24–6 mya fossil could have originated from either a White Stork or a Black Stork (C. nigra), which are species of about the same size with very similar bone structures. Middle Miocene beds of Maboko Island have yielded further remains.[13]

Description

White bird with black flight feathers, long legs and a long neck soars against a hazy blue sky.
Flying in Germany. Storks fly with their necks outstretched.

The White Stork is a large bird. It has a length from the tip of its beak to the tip of its tail of 100–115 cm (39–45 in), a wingspan of 195–215 cm (77–85 in),[14] and a weight of 2.3–4.4 kg (5.1–9.7 lb). Like all storks, it has long legs, a long neck, and a long straight pointed beak.[10] Its plumage is mainly white with some black in the wings; the primaries, secondaries, primary coverts, greater upperwing coverts, scapulars, and alula are black.[15] The breast feathers are long and shaggy forming a ruff which is used in some courtship displays.[15] The irises are dull brown or grey, and the peri-orbital skin is black. The adult has a bright red beak and red legs.[7] The sexes are identical in appearance, except that males are larger than females on average.[7] As with other storks, the wings of the White Stork are long and broad, allowing the bird to soar.[16] In flapping flight its wingbeats are slow and regular. It flies with its neck outstretched forward and with its long legs outstretched beyond the end of its short tail. It walks at a slow and steady pace with its neck outstretched and vertical. In contrast, it often tucks its head between its shoulders when resting.[17] Moulting has not been extensively studied, but appears to takes place over the course of the year, with the primary flight feathers replaced over the breeding season.[15]

Head, neck and upper body of a white stork with a long beak with is reddish at the base fading to black at the tip
An older juvenile at Vogelpark Avifauna, Netherlands. Beaks turn red starting at the base.

Upon hatching, the young White Stork is partially covered with short sparse whitish down feathers. This early down is replaced about a week later with a denser coat of woolly white down. By its third week, the young bird has acquired its black scapulars and flight feathers. On hatching the chick has pinkish legs, which turn to greyish-black as the nestling ages. Its beak is black with a brownish tip.[15] By the time it fledges, the juvenile bird's plumage is similar to that of the adult, though its black feathers are often tinged with brown, and its beak and legs are a duller brownish red or orange. Its beak is typically orange or red with a darker tip.[17] The bills gain the adult red colour the following summer, although the black tips persist in some individuals. Young storks adopt adult plumage by their second summer.[18] The oldest known wild White Stork lived for 39 years after being ringed in Switzerland,[19] while captive birds have lived for more than 35 years.[7]

Similar species

In its range the White Stork is distinctive when seen on the ground, but when seen at a distance in flight it can be confused with several other species with similar underwing patterns, such as the Yellow-billed Stork, Great White Pelican, and Egyptian Vulture.[17] The Yellow-billed Stork is identified by its black tail and a longer slightly curved yellow beak.[20] The Great White Pelican has short legs which do not extend beyond its tail, and it flies with its neck retracted keeping it head near to its stocky body, which gives it a different flight profile.[21] Pelicans also behave differently, soaring in orderly, synchronised flocks rather than in disorganised groups of individuals as the stork does.[22] The Egyptian Vulture is much smaller, with a long wedge-shaped tail, shorter legs and a small yellow-tinged head on a short neck.[23] The Common Crane, which can also look black and white in strong light, shows longer legs and a longer neck in flight.[24]

Distribution and habitat

Several black and white birds with long red legs and long red beaks walk in a green grassy area.
A flock foraging in Turkey. White Stork avoid areas overgrown with tall grass and shrub.

The nominate race of the White Stork has a widely distributed although disjunct summer range across Europe, clustered in the Iberian peninsula and North Africa in the west, and much of eastern and central Europe, with 25% of the world's population concentrated in Poland,[25] as well as parts of western Asia. The asiatica population of about 1500 birds is restricted to a region in central Asia between the Aral Sea and Xinjiang in western China.[18][26] The Xinjiang population is believed to have become extinct around 1980.[27] Migration routes extend the range of this species into many parts of Africa and India. Some populations adhere to the eastern migration route, which passes across Israel into eastern and central Africa.[28][29] A few records of breeding from South Africa have been known since 1933 at Calitzdorp, and about 10 birds have been known to breed since the 1990s around Bredasdorp.[30] A small population of storks winters in India and are thought to belong mostly to the C. c. asiatica population[9] as flocks of up to 200 birds have been observed on spring migration in the early 1900s through the Kurram Valley.[31] However, birds ringed in Germany have been recovered in western (Bikaner) and southern (Tirunelveli) India.[8][32] An atypical specimen with red orbital skin, a feature of the Oriental White Stork, has been recorded[33] and further study of the Indian population has been called for.[9] North of the breeding range, it is a passage migrant or vagrant in Finland, Great Britain, Iceland, Ireland, Norway, and Sweden, and west to the Azores and Madeira. In recent years, the range has expanded into western Russia.[34][35]

The White Stork's preferred feeding grounds are grassy meadows, farmland and shallow wetlands. It avoids areas overgrown with tall grass and shrub.[36] In the Chernobyl area, stork populations declined after the nuclear accident as farmland changed into tall grass shrub.[37] In parts of Poland, poor natural foraging grounds have forced birds to seek food at rubbish dumps since 1999.[38] White Storks have also been reported foraging in rubbish dumps in the Middle East, North Africa, and South Africa.[39]

The White Stork breeds in greater numbers in areas with open grasslands, particularly grassy areas which are wet or periodically flooded, and less in areas with taller vegetation cover such as forest and shrubland.[40] They make use of grasslands, wetlands, and farmland on the winter grounds in Africa.[30] White Storks were probably aided by human activities during the Middle Ages as woodland was cleared and new pastures and farmland was created, and they were found across much of Europe, breeding even as far north as Sweden. In 1416, a pair nested atop St Giles's Church in Edinburgh, Scotland and are otherwise only known to be rare visitors on the British Isles.[41] A decline in populations began in the 19th Century due to industrialisation and changes in agricultural methods. White Storks no longer nest in many countries, and the current strongholds of the western population are in Spain, Ukraine, and Poland. In the Iberian Peninsula, populations are concentrated in the southwest, and have also declined due to agricultural practices.[40] A study published in 2005 found that the Podhale region in the uplands of southern Poland has seen an influx of White Stork, which first bred in there in 1931 and have nested at progressively higher altitudes since, reaching 890 m (3000 ft) in 1999. The authors proposed that this was related to climate warming, and the influx of other animals and plants to higher altitudes.[42] White Storks arriving in Poznań province in western Poland in spring to breed did so some 10 days earlier in the latter twenty years of the twentieth century than at the end of the nineteenth century.[43]

Migration

A blue sky with many tiny silhouettes of distant flying birds
A flock in migration over Israel. Migrating White Storks use the uplift of air thermals to reduce the effort of long-distance flying.

Systematic research into migration began with German ornithologist Johannes Thienemann who commenced banding studies in 1906 at the Rossitten Bird Observatory, on the Curonian Spit in what was then East Prussia. Although not many storks passed through Rossitten itself, the observatory coordinated the large-scale banding of the species throughout Germany and elsewhere in Europe. Between 1906 and the Second World War about 100,000, mainly juvenile, storks were banded, with over 2,000 long-distance recoveries of birds wearing Rossitten bands reported between 1908 and 1954.[44]

Routes

White storks fly south from their summer breeding grounds in Europe in August and September, heading for Africa.[36] There, the storks spend the winter in savanna from Kenya and Uganda south to the Cape Province of South Africa.[45] In these areas they congregate in large flocks which may exceed a thousand individuals.[17] Some diverge westwards into western Sudan and Chad, and may reach Nigeria.[28] In spring, the birds return north; they are recorded from Sudan and Egypt from February to April.[46] They arrive back in Europe around late March and April,[36] after an average journey of 49 days. By comparison, the autumn journey is completed in about 26 days. Tailwinds and scarcity of food and water en route (birds fly faster over regions lacking resources) increase average speed.[29]

To bypass a long sea crossing over the Mediterranean, birds from central Europe either follow an eastern migration route by crossing the Bosphorus to Turkey, traversing the Levant, then avoiding the Sahara Desert by following the Nile valley southwards, or follow a western route over the Strait of Gibraltar.[47] These migration corridors maximise help from the thermals and thus save energy.[48][49] The eastern route is by far the more important with 530,000 storks using it annually, making the species the second commonest migrant there (after the Honey Buzzard). The flocks of raptors, storks and Great White Pelicans can stretch for 200 km (125 mi).[50] The eastern route is twice as long as the western, but storks take the same time to reach the wintering grounds by either route.[51]

Juvenile storks set of on their first southward migration in an inherited direction but, if displaced from that bearing by weather conditions, they are unable to compensate, and may end up in a new wintering location. Adults can compensate for strong winds and adjust their direction to finish at their normal winter sites, because they are familiar with the location. For the same reason, all spring migrants, even those from displaced wintering locations, can find their way back to the traditional breeding sites.[52] An experiment with young White Storks raised in captivity in Kaliningrad and released in the absence of wild storks to show them the way showed that they appeared to have an instinct to fly south, although the scatter in direction was large.[53]

Energetics

White Storks rely on the uplift of air thermals to soar and glide the long distances of their annual migrations between Europe and Sub-Saharan Africa. For many storks the shortest route would take them over the Mediterranean Sea; however, air thermals only form over land, so storks take a detour to fly over land and avoid long flights over water that would require prolonged energetic wing flapping.[54] It has been estimated that flapping flight metabolises 23 times more body fat than soaring flight per distance travelled.[55] Thus, groups of storks spiral upwards on rising warm air until they emerge at the top, up to 1200–1500 m (4000–5000 ft) above the ground (though one record from Western Sudan observed an altitude of 3300 m (10,800 ft)).[46]

Long flights over water may occasionally be undertaken. A young bird ringed at the nest in Denmark subsequently arrived in mainland England, where it spent some days before moving on. It was seen flying over St Mary's, Isles of Scilly, and arrived in a poor condition in Madeira three days later. That island is 500 km (320 mi) from Africa, and twice as far from the European mainland.[56] Migration in the middle east may be hampered by the khamsin, winds bringing gusty overcast days unsuitable for flying. In these situations, flocks of White Storks sit out the adverse weather on the ground, standing and facing into the wind.[46]

Behaviour

An adult in Tsavo East National Park, Kenya. The lower parts of its legs are a whitish colour due to being covered with its droppings—an example of thermoregulation by urohidrosis

The White Stork is a gregarious bird, flocks of thousands of individuals have been recorded on migration routes and at wintering areas in Africa. Non-breeding birds gather in groups of 40 or 50 during the breeding season.[17] Breeding pairs of White Storks may gather in small groups to hunt, and colony nesting has been recorded in some areas.[57] However, groups among White Stork colonies vary widely in size and the social structure is loosely defined. Young storks are often restricted to peripheral nests, while older storks attain higher breeding success while occupying the better quality nests toward the centers of breeding colonies.[58] Social structure and group cohesion is maintained by altruistic behaviours such as allopreening. White Storks exhibit this behaviour exclusively at the nest site. Standing birds preen the heads of sitting birds, sometimes these are parents grooming juveniles, and sometimes juveniles preen each other.[59] Unlike most storks, it never adopts a spread-winged posture, though it is known to droop its wings (holding them away from its body with the primary feathers pointing downwards) when its plumage is wet.[60] A White Stork's droppings, containing faeces and urine, sometimes go onto its own legs, making them appear white.[20] The resulting evaporation provides cooling and is termed urohidrosis.[61] Birds that have been ringed can sometimes be affected by the accumulation of droppings leading to constriction and trauma to the legs.[62] The White Stork has also been noted for tool use by squeezing moss in the beak to drip water into the mouths of their chicks.[63]

Communication

The adult White Stork's main sound is noisy bill-clattering, which has been likened to distant machine gun fire. The bird makes these sounds by rapidly opening and closing its beak so that a knocking sound is made each time its beak closes. The clattering is amplified by its throat pouch, which acts as a resonator. Used in a variety of social interactions, bill-clattering generally grows louder the longer it lasts, and takes on distinctive rhythms depending on the situation—for example, slower during copulation and briefer when given as an alarm call. The only vocal sound adult birds generate is a weak barely audible hiss; however, young birds can generate a harsh hiss, various cheeping sounds, and a cat-like mew they use to beg for food. Like the adults, young also clatter their beaks.[64] The up-down display is used for a number of interactions with other members of the species. Here a White Stork quickly throws its head backwards so that its crown rests on its back before slowly bringing its head and neck forwards again, and this is repeated several times. The display is used as a greeting between birds, post coitus, and also as a threat display. Breeding pairs are territorial over the summer, and use this display, as well as crouching forward with the tails cocked and wings extended.[65]

Feeding

The White Stork consumes a wide variety of animal prey. They prefer to forage in meadows that are within roughly 5 km (3 mi) of their nest and sites where the vegetation is shorter so that their prey is more accessible.[36] Its diet varies depending on season, locality and prey availability. Common food items include insects (primarily beetles, grasshoppers, locusts and crickets), earthworms, reptiles, amphibians (particularly frogs) and small mammals such as voles, moles, and shrews. Less commonly, it also eats bird eggs and young birds, fish, molluscs, crustaceans and scorpions. It hunts principally during the day, swallowing small prey whole, but killing and breaking apart larger prey before swallowing it.[57] Birds returning to Latvia during spring have been shown to locate their prey, moor frogs (Rana arvalis), by homing in on the mating calls produced by aggregations of male frogs.[66] The red colour of the beak is derived from carotenoids in the diet. In parts of Spain, studies have shown that their colour is based on astaxanthin obtained from an introduced species of crayfish (Procambarus clarkii) and the bright red beak colours show up even in nestlings, in contrast to the duller beaks of young White Storks elsewhere.[67]

The diet of non-breeding birds is similar to that of breeding birds, but food items are more often taken from dry areas.[68] Storks wintering in western India have been observed to follow blackbuck to capture insects disturbed by them.[69] Wintering storks in India sometimes forage along with Woolly-necked Stork (Ciconia episcopus).[70] Food piracy has been recorded in India with a rodent captured by Western Marsh Harrier appropriated by a White Stork while Montagu's Harrier is known to harass storks foraging for voles in some parts of Poland.[71][72]

Breeding

At least eight tall, black and white birds, in three nests on the roof of a building.
Nests on a belfry in Spain. White Storks often form small nesting colonies

The White Stork breeds in open farmland areas with access to marshy wetlands, building a large stick nest in trees, on buildings, or on purpose built man-made platforms.[73] Each nest is 1–2 m (3.3–6.6 ft) in depth, 0.8–1.5 m (2.6–4.9 ft) in diameter, and 60–250 kg (130–550 lb) in weight.[74] Nests are built in loose colonies.[75] Not persecuted as it is viewed as a good omen, it often nests close to human habitation; in southern Europe, nests can be seen on churches and other buildings. The nest is typically used year after year especially by older males. The males arrive earlier in the season and choose the nests. Nest change is often related to a change in the pairing and these are more common in the younger birds.[76][77] A succession of pairs have been observed occupying a nest for a few days before moving on, the reason for which is unclear.[78]

Other bird species often nest within the storks' large nest; regular occupants are House Sparrows, Tree Sparrows, and Common Starlings; less common residents include Eurasian Kestrels, Little Owls, European Rollers, White Wagtails, Black Redstarts, Eurasian Jackdaws, and Spanish Sparrows.[79] Paired birds greet by engaging in up-down and head-shaking crouch displays, and clattering the beak while throwing back the head.[7] Pairs copulate frequently throughout the month before eggs are laid. High frequency pair copulation is usually associated with sperm competition and high frequency of extra-pair copulation; however, extra-pair copulation is infrequent in White Storks.[80]

A white oval-shaped egg, tinged with yellow and faint blackish lines
Egg: average size 72.6 × 51.9 mm (2.86 × 2.04 in)

A White Stork pair raises a single brood a year. The female typically lays four eggs, though clutches of 1–7 have been recorded.[64] The eggs are white, but often look dirty or yellowish due to a glutinous covering. They measure 72.58 mm × 51.86 mm (2.857 in × 2.042 in),[81] and weigh 96–129 g (3.4–4.6 oz),[64] of which 10.76 g (0.380 oz) is shell.[81] Incubation begins as soon as the first egg is laid, so the brood hatches asynchronously, beginning 33 to 34 days later. The first hatchling typically has a competitive edge over the others. While stronger chicks are not aggressive towards weaker siblings, as is the case in some species, particularly weak chicks are sometimes killed by their parents.[82][83] The temperature and weather around the time of hatching in spring is important; cool temperatures and wet weather increase chick mortality and reduce breeding success rates.[40] Somewhat unexpectedly, studies have found that later-hatching chicks which successfully reach adulthood produce more chicks than do their earlier-hatching nestmates.[84] The body weight of the chicks increases rapidly in the first few weeks and reaches a plateau of about 3.4 kg (7.5 lb) in 45 days. The length of the beak increases linearly for about 50 days.[85] Young birds are fed with earthworms and insects, which are regurgitated by the parents onto the floor of the nest. Older chicks reach into the mouths of parents to obtain food.[86] Chicks fledge 58–64 days after hatching.[18] White Stork parents have been known to kill one of their young, generally the smallest, in times of food shortage to reduce brood size and hence increase the chance of survival of the remaining nestlings. Stork nestlings do not attack each other, and their parents' method of feeding them (disgorging large amounts of food at once) means that stronger siblings cannot outcompete weaker ones for food directly, hence parental infanticide is an efficient way of reducing brood size. Despite this, this behaviour has not commonly been observed.[87]

Parasites and diseases

White Stork nests are habitats for an array of small arthropods, particularly over the warmer months after the birds arrive to breed. Nesting over successive years, the storks bring more material to line their nests and layers of organic material accumulate within them. Not only do their bodies tend to regulate temperatures within the nest, but excrement, food remains and feather and skin fragments provide nourishment for a large and diverse population of free-living mesostigmatic mites. A survey of twelve nests found 13,352 individuals of 34 species, the most common being Macrocheles merdarius, M. robustulus, Uroobovella pyriformis and Trichouropoda orbicularis, which together represented almost 85% of all the specimens collected. These feed on the eggs and larvae of insects and on nematodes, which are abundant in the nest litter. These mites are dispersed by coprophilous beetles, often of the family Scarabaeidae, or on dung brought by the storks during nest construction. Parasitic mites do not occur, perhaps being controlled by the predatory species. The overall impact of the mite population is unclear, the mites may have a role in suppressive harmful organisms (and hence be beneficial), or they may themselves have an adverse effect on nestlings.[88][89]

The birds themselves host species belonging to more than four genera of feather mites.[90] These mites, including Freyana pelargica, Pterolichus ciconiae and Xoloptes didactylus live on fungi growing on the feathers.[91] The fungi found on the plumage may feed on the keratin of the outer feathers or on feather oil.[92] Chewing lice such as Colpocephalum zebra tend to be found on the wings, and Neophilopterus incompletus elsewhere on the body.[93]

The stork also carries several types of internal parasites, including Toxoplasma gondii[94] and intestinal parasites of the genus Giardia.[95] A study of 120 White Stork carcasses from Saxonia-Anhalt and Brandenburg in Germany yielded eight species of trematode (fluke), four cestode (tapeworm) species, and at least three species of nematode.[96] One species of fluke, Chaunocephalus ferox, caused lesions in the wall of the small intestine in a number of birds admitted to two rehabilitation centers in central Spain, and was associated with reduced weight. It is a recognised pathogen and cause of morbidity in the Asian Openbill (Anastomus oscitans).[97]

West Nile virus (WNV) is mainly a bird infection that is transmitted between birds by Mosquitos.[98] Migrating birds appear to be important in spread of the virus,[99] the ecology of which remains poorly known.[100] On August 26, 1998, a flock of about 1,200 migrating White Storks that had been blown off course on their southward journey landed in Eilat, in southern Israel. The flock was stressed as it had resorted to flapping flight to return to its migratory route, and a number of birds died. A virulent strain of West Nile virus was isolated from the brains of eleven dead juveniles. Other White Storks subsequently tested in Israel have shown anti-WNV antibodies.[101] In 2008 three juvenile White Storks from a Polish wildlife refuge yielded seropositive results indicating exposure to the virus, but the context or existence of the virus in that country is unclear.[102]

Conservation

The White Stork benefitted from human activities during the Middle Ages as woodland was cleared and new pastures and farmland was created, but a decline in population began in the 19th Century due to industrialisation and changes in agricultural methods. The last wild stork in Belgium was seen in 1895, in Sweden in 1955, in Switzerland in 1950 and in Holland in 1991. However, storks have since been reintroduced to many regions.[103] Threats to the species include the drainage of wetlands, collisions with overhead power lines, use of persistent pesticides (such as DDT) to combat locusts in Africa, and largely illegal hunting on migration routes and wintering grounds.[7] The White Stork is one of the species to which the Agreement on the Conservation of African-Eurasian Migratory Waterbirds (AEWA) applies.[104] Parties to the Agreement are required to engage in a wide range of conservation strategies described in a detailed action plan. The plan is intended to address key issues such as species and habitat conservation, management of human activities, research, education, and implementation.[105]

Three long-legged, long-billed black and white birds stand on a huge pile of sticks atop an artificial platform on a pole
Three young birds on their stick nest built on a man-made platform in Poland

A large population of White Storks breeds in central and eastern Europe. In a 2004 census, there were 52,500 pairs in Poland, 12,000–18,000 pairs in Ukraine, 10,500–13,000 pairs in Belarus, 10,000 pairs in Lithuania (the highest known density of this species in the world), and 8,500 pairs in Latvia. In Estonia the population has also been increasing, numbering about 4000 pairs in 2004. There were around 5,000 pairs in Romania and an estimated 4,818 breeding pairs in Bulgaria.[106] In Germany, 3,000 of the total 3,400 pairs are in the former East Germany. In southwestern Asia, Turkey has the highest population, with 15,000–35,000 pairs. Apart from Spain (14,000 pairs) and Portugal (ca. 10,000 pairs in 2008), populations in western Europe are generally much less stable for instance the once sizable Danish population declined to just five pairs in 1995.

In the 1980s, the White Stork population had fallen to fewer than nine pairs in the entire upper Rhine River valley, an area closely identified with the White Stork for centuries. Conservation efforts successfully increased the population of birds to 270 pairs largely due to the actions of the Association for the Protection and Reintroduction of Storks in Alsace and Lorraine.[107] The re-introduction of zoo-reared birds has halted further declines in Italy (30 pairs), the Netherlands (9–12 pairs), and Switzerland (120–160 pairs). There were 601 pairs breeding in Armenia and around 700 pairs in the Netherlands in 2008,[108] and few pairs also breed in South Africa, typically recent colonists from within the normal wintering population.[7][18] In Poland, electric poles have been modified with a platform at the top to prevent the stork's large nest form disrupting the electricity supply, and sometimes nests are moved from an electric pole to a man-made platform.[74] Introductions of zoo-reared birds in the Netherlands has been followed up by feeding and nest-building programs by volunteers.[108] Similar reintroduction programs are taking place in Sweden,[109] and Switzerland,[110] where 175 pairs were recorded breeding in 2000.[111] Long term viability of the population in Switzerland is unclear as breeding success rates are low, and supplementary feeding does not appear to be of benefit.[110]

Cultural associations

"Der Klapperstorch" (The Stork), a painting by Carl Spitzweg (1808–1885)

Due to its large size, predation on vermin, and nesting behaviour close to human settlements and on rooftops, the White Stork has an imposing presence that has had an impact on human culture and folklore.[56] Greek and Roman mythology portray storks as models of parental devotion, and it was believed that they did not die of old age, but flew to islands and took the appearance of humans. The bird is featured in two of Aesop's Fables: The Fox and the Stork and The Farmer and the Stork.[112] In Ancient Egypt, the stork was associated with, and was the hieroglyph for, the Ba, or "soul".[113] Followers of Islam also revered storks because they too made an annual pilgrimage to Mecca on their migration.[114] In ancient Greece, killing a stork could be punished with death.[115]

Supposed filial virtues of the stork (1831)

An ancient Greek belief was that storks cared for their aged parents, feeding them and even transporting them. In Greece children's books depicted the stork as a model of filial values. A Greek law called Pelargonia, from the Greek word pelargos for stork, required citizens to take care of their aged parents.[7] Another idea was that storks were only found in countries having a republican form of government.[116]

Storks have little fear of humans in places where they are not disturbed, and often nest on buildings in Europe. In Germany, the presence of a nest on a house was believed to protect against fires. they were also protected in the belief that their souls were human.[117] According to northern European legend, the stork is responsible for bringing babies to new parents. The legend is very ancient, but was popularised by a nineteenth century Hans Christian Andersen story called The Storks.[114]

A long term study that showed a spurious correlation between the numbers of stork nests and human births is widely used in the teaching of basic statistics as an example to highlight that correlation does not necessarily indicate causation.[118][119] The child-bringing myth has appeared in different forms in history. Children of African American slaves were sometimes told that white babies where brought by storks, while black babies were born from buzzard eggs.[120][121] Some of the earliest understanding on bird migration were initiated by an interest in White Storks, particularly due to examples, known in German as Pfeilstorch ("arrow storks"), that were found in Europe with African arrows embedded in their bodies. A well known example of such a stork found in the summer of 1822 in the German town of Klutz in Mecklenburg was made into a mounted taxidermy specimen, complete with the ornate African arrow, that is now in the University of Rostock.[122]

In Slavic mythology and religion, it was believed that in spring and summer storks carry unborn souls from Iriy to Earth.[123] This belief still persist in the modern folk culture of many Slavic countries, in the simplified child story that "storks bring children into the world".[124] Storks were seen by the Slavs as bringing luck, and killing one would bring misfortune.[125] Poles, Lithuanians and Ukrainians believe that storks bring harmony to a family on whose property they nest.[126] The White Stork is the national bird of Lithuania,[127] and it was a Polish mascot at the Expo 2000 Fair in Hanover.[128] Polish poet Cyprian Kamil Norwid mentioned storks in his poem Moja piosnka (II) ("My Song (II)"):[129]

For the land where it's a great travesty
To harm a stork's nest in a pear tree,
For storks serve us all…

I am homesick, Lord!...

— translated by Walter Whipple

The White Stork is a popular motif on postage stamps, and it is featured on more than 120 stamps issued by more than 60 stamp-issuing entities.[130]

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Works cited

  • Cramp, Stanley, ed. (1977). Handbook of the Birds of Europe the Middle East and North Africa, the Birds of the Western Palearctic, Volume 1: Ostrich to Ducks. Oxford: Oxford University Press. ISBN 0-19-857358-8.
  • Elliott, Andrew (1992). "Family Ciconiidae (Storks)". Handbook of the Birds of the World Volume 1: Ostrich to Ducks. Barcelona: Lynx Edicions. ISBN 8487334105. {{cite book}}: Unknown parameter |editors= ignored (|editor= suggested) (help)
  • Haverschmidt, François (1949). The Life of the White Stork. Leiden: E J Brill.
  • Newton, Ian (2010). Collins New Naturalist Library (113) - Bird Migration. London: Collins. ISBN 0007307322.
  • Svensson, Lars; Grant, Peter J (1999). Collins Bird Guide. London: HarperCollins. ISBN 0-00-219728-6.