White stork

The white stork (Ciconia ciconia) is a large bird in the stork family, Ciconiidae. Its plumage is mainly white, with black on the bird's wings. Adults have long red legs and long pointed red beaks, and measure on average 100–115 cm (39–45 in) from beak tip to end of tail, with a 155–215 cm (61–85 in) wingspan. The two subspecies, which differ slightly in size, breed in Europe (north to Finland), northwestern Africa, southwestern Asia (east to southern Kazakhstan) and southern Africa. The white stork 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. When migrating between Europe and Africa, it avoids crossing the Mediterranean Sea and detours via the Levant in the east or the Strait of Gibraltar in the west, because the air thermals on which it depends for soaring do not form over water.

White stork

Least Concern  (IUCN 3.1)[1]
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Ciconiiformes
Family: Ciconiidae
Genus: Ciconia
Species:
C. ciconia
Binomial name
Ciconia ciconia
Approximate ranges and routes

   Breeding range
   Winter range

  Migration routes
Synonyms
  • Ardea ciconia Linnaeus, 1758

A carnivore, the white stork eats a wide range of animal prey, including insects, 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. Each year the female can lay one clutch of usually four eggs, which hatch asynchronously 33–34 days after being laid. Both parents take turns incubating the eggs and both feed the young. The young leave the nest 58–64 days after hatching, and continue to be fed by the parents for a further 7–20 days.

The white stork has been rated as least concern by the International Union for Conservation of Nature (IUCN). It benefited from human activities during the Middle Ages as woodland was cleared, but changes in farming methods and industrialisation saw it decline and disappear from parts of Europe in the 19th and early 20th centuries. Conservation and reintroduction programs across Europe have resulted in the white stork resuming breeding in the Netherlands, Belgium, Switzerland, Sweden and the United Kingdom. It has few natural predators, but may harbour several types of parasite; the plumage is home to chewing lice and feather mites, while the large nests maintain a diverse range of mesostigmatic mites. This conspicuous species 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

English naturalist Francis Willughby wrote about the white stork in the 17th century, having seen a drawing sent to him by his friend and natural history enthusiast Sir Thomas Brown of Norwich. He named it Ciconia alba. They noted they were occasional vagrants to England, blown there by storms.[2][lower-alpha 1] It was one of the many bird species originally described by Carl Linnaeus in his landmark 1758 10th edition of Systema Naturae,[4] where it was given the binomial name of Ardea ciconia. It was reclassified to (and was designated the type species of) the new genus Ciconia by French zoologist Mathurin Jacques Brisson in 1760.[5][6] Both the genus and specific epithet, cĭcōnia, are the Latin word for "stork",[7] originally recorded in the works of Horace and Ovid.[8] The Latin word survives in most Romance languages (cicogna, cigüeña, cegonha and the like). The word stork is derived from the Old English word storc, and appeared in the 10th-century works the Erfurt Glossary, where the word is equated with Ciconia, and Aelfric's Homilies. The word is related to the Old High German storah, "stork", and similar words in many other European languages, all of which are descended from the Germanic sturko-z.[9]

Skeleton

There are two subspecies:

  • 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,[10] though some birds winter in India.[11]
  • C. c. asiatica, 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.[10][12]

The stork family contains six genera in three broad 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,[13] which are characterised by straight pointed beaks and mainly black and white plumage.[14] Its closest relatives are the larger, black-billed Oriental stork (Ciconia boyciana) of East Asia, which was formerly classified as a subspecies of the white stork,[10] and the maguari stork (C. maguari) of South America. Close evolutionary relationships within Ciconia are suggested by behavioural similarities and, biochemically, through analysis of both mitochondrial cytochrome b gene sequences and DNA-DNA hybridization.[15]

A Ciconia fossil representing the distal end of a right humerus has been recovered from Miocene beds of Rusinga Island, Lake Victoria, Kenya.[16] The 24–6 million year old 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. The Middle Miocene beds of Maboko Island have yielded further remains.[16]

Description
A juvenile feeding on an insect

The white stork is a large bird. It has a length of 100–115 cm (39–45 in),[lower-alpha 2][17] and a standing height of 100–125 cm (39–49 in). The wingspan is 155–215 cm (61–85 in) and its weight is 2.3–4.5 kg (5.1–9.9 lb).[18][19] Like all storks, it has long legs, a long neck and a long straight pointed beak.[13] The sexes are identical in appearance, except that males are larger than females on average.[10] The plumage is mainly white with black flight feathers and wing coverts; the black is caused by the pigment melanin.[20] The breast feathers are long and shaggy forming a ruff which is used in some courtship displays.[21] The irises are dull brown or grey, and the peri-orbital skin is black. The adult has a bright red beak and red legs,[10] the colouration of which is derived from carotenoids in the diet. In parts of Spain, studies have shown that the pigment 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.[22]

In flight. White storks fly with their necks outstretched.

As with other storks, the wings are long and broad enabling the bird to soar.[23] In flapping flight its wingbeats are slow and regular. It flies with its neck stretched forward and with its long legs extended well beyond the end of its short tail. It walks at a slow and steady pace with its neck upstretched. In contrast, it often hunches its head between its shoulders when resting.[24] Moulting has not been extensively studied, but appears to take place throughout the year, with the primary flight feathers replaced over the breeding season.[21]

An older juvenile at Vogelpark Avifauna, Netherlands. Beaks turn red starting at the base.

Upon hatching, the young white stork is partly 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 three weeks, the young bird acquires black scapulars and flight feathers. On hatching the chick has pinkish legs, which turn to greyish-black as it ages. Its beak is black with a brownish tip.[21] 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. The beak is typically orange or red with a darker tip.[24] The bills gain the adults' red colour the following summer, although the black tips persist in some individuals. Young storks adopt adult plumage by their second summer.[25]

Similar species

Within its range the white stork is distinctive when seen on the ground. The winter range of C. c. asiatica overlaps that of the Asian openbill, which has similar plumage but a different bill shape. When seen at a distance in flight, the white stork can be confused with several other species with similar underwing patterns, such as the yellow-billed stork, great white pelican and Egyptian vulture.[24] The yellow-billed stork is identified by its black tail and a longer, slightly curved, yellow beak. The white stork also tends to be larger than the yellow-billed stork.[26] The great white pelican has short legs which do not extend beyond its tail, and it flies with its neck retracted, keeping its head near to its stocky body, giving it a different flight profile.[27] Pelicans also behave differently, soaring in orderly, synchronised flocks rather than in disorganised groups of individuals as the white stork does.[28] The Egyptian vulture is much smaller, with a long wedge-shaped tail, shorter legs and a small yellow-tinged head on a short neck.[29] The common crane, which can also look black and white in strong light, shows longer legs and a longer neck in flight.[30]

Distribution and habitat
A flock foraging in Turkey. White storks avoid areas overgrown with tall grass and shrub.

The nominate race of the white stork has a wide 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,[31] as well as parts of western Asia. The asiatica population of about 1450 birds is restricted to a region in central Asia between the Aral Sea and Xinjiang in western China.[32][33] The Xinjiang population is believed to have become extinct around 1980.[34] 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.[35][36]

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.[37] A small population of white storks winters in India and is thought to derive principally from the C. c. asiatica population[12] as flocks of up to 200 birds have been observed on spring migration in the early 1900s through the Kurram Valley.[38] However, birds ringed in Germany have been recovered in western (Bikaner) and southern (Tirunelveli) India.[11][39] An atypical specimen with red orbital skin, a feature of the Oriental white stork, has been recorded[40] and further study of the Indian population is required.[12] North of the breeding range, it is a passage migrant or vagrant in Finland, Iceland, Ireland, Norway and Sweden, and west to the Azores and Madeira. In recent years, the range has expanded into western Russia.[41][42]

The white stork's preferred feeding grounds are grassy meadows, farmland and shallow wetlands. It avoids areas overgrown with tall grass and shrubs.[43] In the Chernobyl area of northern Ukraine, white stork populations declined after the 1986 nuclear accident there as farmland was succeeded by tall grass shrubs.[44] In parts of Poland, poor natural foraging grounds have forced birds to seek food at rubbish dumps since 1999.[45] White storks have also been reported foraging in rubbish dumps in the Middle East, North Africa and South Africa.[46]

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.[47] They make use of grasslands, wetlands, and farmland on the wintering grounds in Africa.[37] White storks were probably aided by human activities during the Middle Ages as woodland was cleared and new pastures and farmland were created, and they were found across much of Europe, breeding as far north as Sweden. The population in Sweden is thought to have established in the 16th century after forests were cut down for agriculture. About 5000 pairs were estimated to breed in the 18th century which declined subsequently. The first accurate census in 1917 found 25 pairs and the last pair failed to breed around 1955.[48] The white stork has been a rare visitor to the British Isles, with about 20 birds seen in Britain every year, and prior to 2020 there were no records of nesting since a pair nested atop St Giles High Kirk in Edinburgh, Scotland in 1416.[49] In 2020, a pair bred in the United Kingdom for the first time in over 600 years,[50] as part of an re-introduction initiative called the White Stork Project.[51]

A decline in population 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 Portugal, Spain, Ukraine and Poland. In the Iberian Peninsula, populations are concentrated in the southwest, and have also declined due to agricultural practices.[47] A study published in 2005 found that the Podhale region in the uplands of southern Poland had seen an influx of white storks, which first bred 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.[52] White storks arriving in Poznań province (Greater Poland Voivodeship) in western Poland in spring to breed did so some 10 days earlier in the last twenty years of the 20th century than at the end of the 19th century.[53]

Migration
In 1822, the Rostocker Pfeilstorch provided early evidence of long-distance stork migration
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 of the white stork began with German ornithologist Johannes Thienemann who commenced bird ringing 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 ringing of the species throughout Germany and elsewhere in Europe. Between 1906 and the Second World War about 100,000, mainly juvenile, white storks were ringed, with over 2,000 long-distance recoveries of birds wearing Rossitten rings reported between 1908 and 1954.[54]

Routes

White storks fly south from their summer breeding grounds in Europe in August and September, heading for Africa.[43] There, they spend the winter in savannah from Kenya and Uganda south to the Cape Province of South Africa.[55] In these areas they congregate in large flocks which may exceed a thousand individuals.[24] Some diverge westwards into western Sudan and Chad, and may reach Nigeria.[35] In spring, the birds return north; they are recorded from Sudan and Egypt from February to April.[56] They arrive back in Europe around late March and April,[43] 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.[36]

Eastern route: storks resting in Rahat (spring 2017).

To avoid 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 bypassing the Sahara Desert by following the Nile valley southwards, or follow a western route over the Strait of Gibraltar.[57] These migration corridors maximise help from the thermals and thus save energy.[58][59] The eastern route is by far the more important with 530,000 white storks using it annually, making the species the second commonest migrant there (after the European honey buzzard). The flocks of migrating raptors, white storks and great white pelicans can stretch for 200 km (125 mi).[60] The eastern route is twice as long as the western, but storks take the same time to reach the wintering grounds by either.[61]

Juvenile white storks set off 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.[62] An experiment with young birds raised in captivity in Kaliningrad and released in the absence of wild storks to show them the way revealed that they appeared to have an instinct to fly south, although the scatter in direction was large.[63]

Energetics
Otto Lilienthal studied the flight of storks in 1889 while designing his glider. Sketches by his brother Gustav Lilienthal.

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, the shortest route would take them over the Mediterranean Sea; however, since air thermals do not form over water, they generally detour over land to avoid the trans-Mediterranean flights that would require prolonged energetic wing flapping.[64] It has been estimated that flapping flight metabolises 23 times more body fat than soaring flight per distance travelled.[65] Thus, flocks spiral upwards on rising warm air until they emerge at the top, up to 1,200–1,500 m (3,900–4,900 ft) above the ground (though one record from Western Sudan observed an altitude of 3,300 m (10,800 ft)).[56]

Long flights over water may occasionally be undertaken. A young white stork ringed at the nest in Denmark subsequently appeared in England, where it spent some days before moving on. It was later 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.[66] Migration through 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.[56]

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.[24] The smaller dark-plumaged Abdim's stork is often encountered with white stork flocks in southern Africa.[67] Breeding pairs of white stork may gather in small groups to hunt, and colony nesting has been recorded in some areas.[68] However, groups among white stork colonies vary widely in size and the social structure is loosely defined; young breeding storks are often restricted to peripheral nests, while older storks attain higher breeding success while occupying the better quality nests toward the centres of breeding colonies.[69] 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.[70] 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.[71]

A white stork's droppings, containing faeces and uric acid, are sometimes directed onto its own legs, making them appear white.[26] The resulting evaporation provides cooling and is termed urohidrosis.[72] Birds that have been ringed can sometimes be affected by the accumulation of droppings around the ring leading to constriction and leg trauma.[73] The white stork has also been noted for tool use by squeezing moss in the beak to drip water into the mouths of its chicks.[74]

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.[75] The up-down display is used for a number of interactions with other members of the species. Here a 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.[76]

Breeding and lifespan
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.[77] 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.[78] Nests are built in loose colonies.[69] 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. Larger nests are associated with greater numbers of young successfully fledged, and appear to be sought after.[79] Nest change is often related to a change in the pairing and failure to raise young the previous year, and younger birds are more likely to change nesting sites.[80] Although a pair may be found to occupy a nest, partners may change several times during the early stages and breeding activities begin only after a stable pairing is achieved.[81]

Mating

Several bird species often nest within the large nests of the white stork. 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.[82] Paired birds greet by engaging in up-down and head-shaking crouch displays, and clattering the beak while throwing back the head.[10] 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.[83]

A white stork pair raises a single brood a year. The female typically lays four eggs, though clutches of one to seven have been recorded.[75] The eggs are white, but often look dirty or yellowish due to a glutinous covering. They typically measure 73 mm × 52 mm (2.9 in × 2.0 in),[84] and weigh 96–129 g (3.4–4.6 oz),[75] of which about 11 g (0.39 oz) is shell.[84] 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, weak or small chicks are sometimes killed by their parents.[85][86] This behaviour occurs in times of food shortage to reduce brood size and hence increase the chance of survival of the remaining nestlings. White 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.[85]

Egg

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.[47] Somewhat unexpectedly, studies have found that later-hatching chicks which successfully reach adulthood produce more chicks than do their earlier-hatching nestmates.[87] 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.[88] 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.[89] Chicks fledge 58 to 64 days after hatching.[90]

White storks generally begin breeding when about four years old, although the age of first breeding has been recorded as early as two years and as late as seven years.[21] The oldest known wild white stork lived for 39 years after being ringed in Switzerland,[91] while captive birds have lived for more than 35 years.[10]

Feeding

White storks consume 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.[43] Their diet varies according to season, locality and prey availability. Common food items include insects (primarily beetles, grasshoppers, locusts and crickets), earthworms, reptiles, amphibians, particularly frog species such as the edible frog (Pelophylax kl. esculentus) and common frog (Rana temporaria) and small mammals such as voles, moles and shrews. Less commonly, they also eat bird eggs and young birds, fish, molluscs, crustaceans and scorpions. They hunt mainly during the day, swallowing small prey whole, but killing and breaking apart larger prey before swallowing.[68] Rubber bands are mistaken for earthworms and consumed, occasionally resulting in fatal blockage of the digestive tract.[92]

White stork picking a dead young rabbit

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.[93]

The diet of non-breeding birds is similar to that of breeding birds, but food items are more often taken from dry areas.[94] White storks wintering in western India have been observed to follow blackbuck to capture insects disturbed by them.[95] Wintering white storks in India sometimes forage along with the woolly-necked stork (Ciconia episcopus).[96] Food piracy has been recorded in India with a rodent captured by a western marsh harrier appropriated by a white stork, while Montagu's harrier is known to harass white storks foraging for voles in some parts of Poland.[97][98] White storks can exploit landfill sites for food during the breeding season, migration period and winter.[99]

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 effect of the mite population is unclear, the mites may have a role in suppressing harmful organisms (and hence be beneficial), or they may themselves have an adverse effect on nestlings.[100][101]

The birds themselves host species belonging to more than four genera of feather mites.[102] These mites, including Freyanopterolichus pelargicus[103][104] and Pelargolichus didactylus[104] live on fungi growing on the feathers. The fungi found on the plumage may feed on the keratin of the outer feathers or on feather oil.[105] Chewing lice such as Colpocephalum zebra tend to be found on the wings, and Neophilopterus incompletus elsewhere on the body.[106]

The white stork also carries several types of internal parasites, including Toxoplasma gondii[107] and intestinal parasites of the genus Giardia.[108] A study of 120 white stork carcasses from Saxony-Anhalt and Brandenburg in Germany yielded eight species of trematode (fluke), four cestode (tapeworm) species, and at least three species of nematode.[109] One species of fluke, Chaunocephalus ferox, caused lesions in the wall of the small intestine in a number of birds admitted to two rehabilitation centres in central Spain, and was associated with reduced weight. It is a recognised pathogen and cause of morbidity in the Asian openbill (Anastomus oscitans).[110] More recently, the thorough study performed by J. Sitko and P. Heneberg in the Czech Republic in 1962–2013 suggested that the central European white storks host 11 helminth species. Chaunocephalus ferox, Tylodelphys excavata and Dictymetra discoidea were reported to be the dominant ones. The other species found included Cathaemasia hians, Echinochasmus spinulosus, Echinostoma revolutum, Echinostoma sudanense, Duboisia syriaca, Apharyngostrigea cornu, Capillaria sp. and Dictymetra discoidea. Juvenile white storks were shown to host less species, but the intensity of infection was higher in the juveniles than in the adult storks.[111]

West Nile virus (WNV) is mainly a bird infection that is transmitted between birds by mosquitos.[112] Migrating birds appear to be important in spread of the virus,[113] the ecology of which remains poorly known.[114] On 26 August 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.[115] 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 Poland is unclear.[116]

Conservation
Results of the 2004/05 white stork census in Europe (numbers of breeding pairs)

The white stork's decline due to industrialisation and agricultural changes (principally the draining of wetlands and conversion of meadows to crops such as maize) began in the 19th century: the last wild individual in Belgium was seen in 1895, in Sweden in 1955, in Switzerland in 1950 and in the Netherlands in 1991. However, the species has since been reintroduced to many regions.[117] It has been rated as least concern by the IUCN since 1994, after being evaluated as near threatened in 1988.[1] The white stork is one of the species to which the Agreement on the Conservation of African-Eurasian Migratory Waterbirds (AEWA) applies.[118] 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.[119] Threats include the continued loss 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.[10]

A man-made nest platform in Poland built as a conservation measure and to prevent storks disrupting electricity supplies through nesting on pylons. Three young white storks are on the top of the nest and two Eurasian tree sparrows are perching on the side of the nest.

A large population of white storks breeds in central (Poland, Ukraine and Germany) and southern Europe (Spain and Turkey). In a 2004/05 census, there were 52,500 pairs in Poland, 30,000 pairs in Ukraine, 20,000 pairs in Belarus, 13,000 pairs in Lithuania (the highest known density of this species in the world), 10,700 pairs in Latvia, and 10,200 in Russia. There were around 5,500 pairs in Romania, 5,300 in Hungary and an estimated 4,956 breeding pairs in Bulgaria.[120] In Germany, the majority of the total 4,482 pairs were in the eastern region, especially in the states of Brandenburg and Mecklenburg-Vorpommern (1296 and 863 pairs in 2008 respectively).[121] Apart from Spain and Portugal (33,217 and 7,684 pairs in 2004/05 respectively), populations are generally much less stable. The Danish population declined to just three pairs in 2005. In the eastern Mediterranean region Turkey has a sizeable population of 6,195 pairs, and Greece 2,139 pairs. In Western Europe the white stork remains a rare bird despite conservation efforts. In 2004 France had only 973 pairs, and the Netherlands 528 pairs.[120] In Armenia the population of the white stork slightly increased in the period between 2005 and 2015, and by last data reached 652 pairs.[122][123][124]

In the early 1980s, the 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 there to 270 pairs (in 2008), largely due to the actions of the Association for the Protection and Reintroduction of Storks in Alsace and Lorraine.[125] The reintroduction of zoo-reared birds has halted further declines in Italy, the Netherlands, and Switzerland. There were 601 pairs breeding in Armenia and around 700 pairs in the Netherlands in 2008,[126] and few pairs also breed in South Africa, typically recent colonists from within the normal wintering population.[10] In Poland, electric poles have been modified with a platform at the top to prevent the white stork's large nest from disrupting the electricity supply, and sometimes nests are moved from an electric pole to a man-made platform.[78] Introductions of zoo-reared birds in the Netherlands has been followed up by feeding and nest-building programs by volunteers.[126] Similar reintroduction programs are taking place in Sweden,[127] and Switzerland,[128] where 175 pairs were recorded breeding in 2000.[129] 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.[128] However, as of 2017, 470 adults and 757 young ones were recorded in Switzerland.[130]

In August 2019, 24 juveniles were released at the Knepp Estate in West Sussex, and others at a site near Tunbridge Wells and at the Wintershall Estate, near Godalming, as part of a project to reintroduce the white stork as a breeding species in South East England,[131] for the first time since 1416.[132] In 2020, the program was successful with the birth of five baby storks.[50]

Cultural associations
Bociany (Storks), a painting by Józef Chełmoński (1849–1914)

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 influenced human culture and folklore.[66] The Hebrew word for the white stork is chasidah (חסידה), meaning "merciful" or "kind".[133] Greek and Roman mythology portray storks as models of parental devotion. The 3rd century Roman writer Aelian citing the authority of Alexander of Myndus noted in his De 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.[134] The bird is featured in at least three of Aesop's Fables: The Fox and the Stork, The Farmer and the Stork, and The Frogs Who Desired a King. 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 called Pelargonia, from the Ancient Greek word pelargos for stork, required citizens to take care of their aged parents.[10] The Greeks also held that killing a stork could be punished with death.[135] Storks were allegedly protected in Ancient Thessaly as they hunted snakes, and widely held to be Virgil's "white bird".[136] Roman writers noted the white stork's arrival in spring, which alerted farmers to plant their vines.[137]

Followers of Islam revered storks because they made an annual pilgrimage to Mecca on their migration.[138] Some of the earliest understanding on bird migration were initiated by an interest in white storks; Pfeilstörche ("arrow storks") 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 Klütz in Mecklenburg was made into a mounted taxidermy specimen, complete with the ornate African arrow, that is now in the University of Rostock.[139]

Supposed filial virtues of the stork[140]

Storks have little fear of humans if 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 because of the belief that their souls were human.[141] German and Dutch households would encourage storks to nest on houses, sometimes by constructing purpose-built high platforms, to bring good luck.[133] Poles, Lithuanians, Ukrainians and Belarusians believe that storks bring harmony to a family on whose property they nest.[142]

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.[143] It is the national bird of Lithuania, Belarus[144] and Poland, and it was a Polish mascot at the Expo 2000 Fair in Hanover.[145] Storks nesting in Polish villages such as Zywkowo have made them tourist attractions, drawing 2000–5000 visitors a year in 2014.[146] In the 19th century, storks were also thought to only live in countries having a republican form of government.[147] Polish poet Cyprian Kamil Norwid mentioned storks in his poem Moja piosnka (II) ("My Song (II)"):[148]

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

In 1942 Heinrich Himmler sought to use storks to carry Nazi propaganda leaflets so as to win support from the Boers in South Africa. The idea for this "Storchbein-Propaganda" plan was a secret that was transmitted by Walter Schellenberg to be examined by the German ornithologist Ernst Schüz at the Rossiten bird observatory, who pointed out that the probability of finding marked storks in Africa was less than one percent, requiring a 1000 birds to transmit 10 leaflets successfully. The plan was then dropped.[149]

Storks and delivery of babies
Der Klapperstorch (The Stork), a painting by Carl Spitzweg (1808–1885)

According to European folklore, the stork is responsible for bringing babies to new parents. The legend is very ancient, but was popularised by a 19th-century Hans Christian Andersen story called "The Storks".[138] 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 contained adebarsteine or "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.[133] From there the folklore has spread around the world to the Philippines and countries in South America.[133] Birthmarks on the back of the head of newborn baby, nevus flammeus nuchae, are sometimes referred to as stork-bite.[150]

Stork bringing baby - Colmar, Alsace

In Slavic mythology and religion, storks were thought to carry unborn souls from Vyraj to Earth in spring and summer.[151] 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".[152] Storks were seen by Early Slavs as bringing luck, and killing one would bring misfortune.[153] 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.[154][155] The child-bringing myth has appeared in different forms in history. Children of African American slaves were sometimes told that white babies were brought by storks, while black babies were born from buzzard eggs.[156]

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 as Juno to the Holy Ghost, and the stork may have been chosen for its white plumage (depicting purity), size (in that it is big enough to carry an infant), and flight at high altitude (likened to flying between Earth and Heaven).[133] The fable and its relation to the internal world of the child have been discussed by Sigmund Freud[133] and Carl Jung.[157] In fact, Jung recalled being told the story himself upon the birth of his own sister.[158] The traditional link with the newborn continues with their use in advertising for such products as nappies and baby announcements.[133]

There were negative aspects to stork folklore as well; a Polish folk tale 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.[133] In Denmark, storks were said to toss a nestling off the nest and then an egg in successive years.[133] 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.[159]

Notes
  1. The universally accepted starting point of modern taxonomy for animals is set at 1758, with the publishing of Linnaeus' 10th edition of Systema Naturae, although scientists had been coining names in the previous century.[3]
  2. By convention, length is measured from the tip of the bill to the tip of the tail on a dead bird (or skin) laid on its back.

Citations
  1. BirdLife International (2016). "Ciconia ciconia". IUCN Red List of Threatened Species. 2016: e.T22697691A86248677. Retrieved 19 May 2020.
  2. Willughby, Francis (1681). The Ornithology of Francis Willughby of Middleton in the County of Warwick, esq. ... London, United Kingdom: A.C. for John Martyn. p. 286.
  3. Polaszek, Andrew (2010). Systema Naturae 250 - The Linnaean Ark. Boca Raton, Florida: CRC Press. p. 34. ISBN 9781420095029.
  4. Linnaeus, Carl (1758). Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis (in Latin). Vol. I (10th revised ed.). Holmiae: (Laurentii Salvii). p. 142 – via The Internet Archive.
  5. Brisson, Mathurin Jacques (1760). Ornithologie, ou, Méthode Contenant la Division des Oiseaux en Ordres, Sections, Genres, Especes & leurs Variétés (in French and Latin). Paris: Jean-Baptiste Bauche. Vol. 1, p. 48, Vol. 5, p. 361.
  6. Boles, Walter E. (2005). "A review of the Australian fossil storks of the genus Ciconia (Aves : Ciconiidae), with the description of a new species" (PDF). Records of the Australian Museum. 57 (2): 165–78. doi:10.3853/j.0067-1975.57.2005.1440.
  7. Lewis, Charlton Thomas; Kingery, Hugh Macmaster (1918). An Elementary Latin Dictionary. New York: American Book Company. p. 126. ISBN 978-0-19-910205-1.
  8. Simpson, D.P. (1979). Cassell's Latin Dictionary (5th ed.). London: Cassell Ltd. p. 103. ISBN 978-0-304-52257-6.
  9. Simpson, John; Weiner, Edmund, eds. (1989). "Stork". Oxford English Dictionary (2nd ed.). Oxford, UK: Clarendon Press. p. 1915. ISBN 978-0-19-861186-8.
  10. Elliott 1992, pp. 460-1.
  11. Ali, Salim; Ripley, S. Dillon (2001). Handbook of the Birds of India and Pakistan (paperback). Vol. 1 (2nd ed.). India: Oxford University Press. pp. 99–101. ISBN 978-0-19-565934-4.
  12. Rasmussen, Pamela C.; Anderton, John C. (2005). Birds of South Asia: The Ripley Guide. Vol. 2. Washington, DC: Smithsonian Institution and Barcelona: Lynx edicions. p. 63. ISBN 978-84-87334-66-5.
  13. Elliott 1992, p. 437.
  14. Kahl, M. Philip (1987). "An overview of the storks of the world". Colonial Waterbirds. 10 (2): 131–34. doi:10.2307/1521251. JSTOR 1521251.
  15. Slikas, Beth (1997). "Phylogeny of the avian family Ciconiidae (storks) based on cytochrome b Sequences and DNA–DNA hybridization distances". Molecular Phylogenetics and Evolution. 8 (3): 275–300. doi:10.1006/mpev.1997.0431. PMID 9417889.
  16. Dykes, Gareth J.; Alexander, Cyril Walker (2008). "New records of fossil 'waterbirds' from the Miocene of Kenya". American Museum Novitates (3610). hdl:2246/5906.
  17. Cramp 1977, p. 3.
  18. Mead, C.; Ogilvie, M. (2007). The Atlas of Bird Migrations: Tracing the Great Journeys of the World's Birds. Cape Town: Struik. pp. 88–89. ISBN 978-1-77007-499-6.
  19. Hancock, James; Kushan, James A. (1992). "White stork". Storks, Ibises and Spoonbills of the World. Princeton University Press. ISBN 978-0-12-322730-0.
  20. Grande, Juan Manuel; Negro, Juan José; María Torres, José (2004). "The evolution of bird plumage colouration; A role for feather-degrading bacteria?" (PDF). Ardeola. 51 (2): 375–83. doi:10.1007/s00114-008-0462-0. PMID 18853129.
  21. Cramp 1977, p. 335.
  22. Negro, Juan José; Garrido-Fernandez, Juan (2000). "Astaxanthin is the major carotenoid in tissues of White Storks (Ciconia ciconia) feeding on introduced Crayfish (Procambarus clarkii)". Comparative Biochemistry and Physiology B. 126 (3): 347–52. doi:10.1016/S0305-0491(00)00180-2. PMID 11007176.
  23. Elliott 1992, p. 438.
  24. Cramp 1977, p. 328.
  25. Van den Bossche 2002, p. 11.
  26. Sinclair, Ian; Davidson, Ian (2006). Southern African Birds: A Photographic Guide. Cape Town, RSA: Struik. p. 34. ISBN 978-1-77007-244-2.
  27. Cramp 1977, p. 228.
  28. Svensson & Grant 1999, p. 23.
  29. Svensson & Grant 1999, p. 74.
  30. Svensson & Grant 1999, p. 34.
  31. Chernetsov, Nikita; Chromik, Wiesław; Dolata, Pawel T.; Profus, Piotr; et al. (2006). "Sex-related natal dispersal of White Storks (Ciconia ciconia) in Poland: How far and where to?" (PDF). The Auk. 123 (4): 1103–9. doi:10.1642/0004-8038(2006)123[1103:SNDOWS]2.0.CO;2 – via The Russian Academy of Sciences: Zoological Institute: Biological Station Rybachy.
  32. Van den Bossche 2002, p. 10.
  33. Scully, John (1876). "A contribution to the ornithology of eastern Turkestan". Stray Feathers. 4: 41–205 – via The Internet Archive.
  34. Ma, Ming; Dai, Cai (2002). "The Fate of the White Stork (Ciconia ciconia asiatica) in Xinjiang, China". Abstract Volume. 23rd International Ornithological Congress, Beijing, August 11–17, 2002. p. 352.
  35. Berthold, Peter; Van Den Bossche, Willem; Fiedler, Wolfgang; Kaatz, Christoph; et al. (2001). "Detection of a new important staging and wintering area of the White Stork Ciconia ciconia by satellite tracking" (PDF). Ibis. 143 (4): 450–5. CiteSeerX 10.1.1.579.7034. doi:10.1111/j.1474-919X.2001.tb04946.x – via Tel Aviv University.
  36. Shamoun-Baranes, Judy; Baharad, Anat; Alpert, Pinhas; Berthold, Peter; et al. (2003). "The effect of wind, season and latitude on the migration speed of White Storks Ciconia ciconia, along the eastern migration route" (PDF). Journal of Avian Biology. 34: 97–104. doi:10.1034/j.1600-048X.2003.03079.x – via Tel Aviv University.
  37. Allan, David G. (1997). "White Stork" (PDF). In Harrison, J.A.; Allan, D.G.; Underhill, L.G.; Herremans, M.; Tree, A.J. Parker, V; Brown, C.J. (eds.). The Atlas of Southern African Birds. Vol. 1: Non-passerines. BirdLife South Africa. pp. 82–3. ISBN 978-0-620-20729-4 – via South African Bird Atlas Project 2.
  38. Whitehead, Charles Hughes Tempest (1911). "On the birds of Kohat and the Kurram Valley. Northern India. Part 3". Journal of the Bombay Natural History Society. 20 (4): 954–80.
  39. Prater, Stanley Henry (1931). "The migration of the White Stork Ciconia ciconia". Journal of the Bombay Natural History Society. 35 (2): 459.
  40. Jadeja, Vijayraj; Vyas, Raju; Parasharya, B.M. (1998). "Record of Ciconia ciconia asiaticus from Indian Territory". Journal of the Bombay Natural History Society. 95 (1): 113.
  41. Snow, David William; Perrin, C.M.; Gillmor, Robert; et al., eds. (1998). The Birds of the Western Palearctic. Vol. 1: Non-Passerines (Concise ed.). Oxford University Press. pp. 141–3. ISBN 978-0-19-854099-1.
  42. Thomsen, Kai-Michael; Hötker, Hermann (2006). "The Sixth International White Stork Census: 2004–2005" (PDF). In Boere, G.C.; Galbraith, C.A.; Stroud, D.A. (eds.). Waterbirds Around the World. Edinburgh: The Stationery Office. pp. 493–5. ISBN 978-0-11-497333-9.
  43. Johst, Karin; Brandl, Rolan; Pfeifer, Robert (2001). "Foraging in a patchy and dynamic landscape: Human land use and the White Stork". Ecological Applications. 11: 60–9. doi:10.1890/1051-0761(2001)011[0060:FIAPAD]2.0.CO;2. ISSN 1051-0761.
  44. Samusenko, Irina (2004). "Some aspects of White Stork Ciconia ciconia population dynamics in the region of Chernobyl's accident" (PDF). Bird Census News. 13 (2000): 157–60. Archived from the original (PDF) on 2011-06-27. Retrieved 2010-12-03.
  45. Kruszyk, Robert; Ciach, Michał (2010). "White Storks, Ciconia ciconia, forage on rubbish dumps in Poland — A novel behaviour in population" (PDF). European Journal of Wildlife Research. 56 (1): 83–7. doi:10.1007/s10344-009-0313-0.
  46. Ciach, Michał; Kruszyk, Robert (2010). "Foraging of White Storks Ciconia ciconia on rubbish dumps on non-breeding grounds". Waterbirds. 33 (1): 101–4. doi:10.1675/063.033.0112.
  47. Carrascal, Luis María; Bautista, Luis Miguel; Lázaro, Encarnación (1993). "Geographical variation in the density of the White Stork Ciconia ciconia in Spain: Influence of habitat structure and climate". Biological Conservation. 65 (1): 83–7. doi:10.1016/0006-3207(93)90200-K.
  48. Cavallin, Berith (1997). "Storklandskapet". Skånes Natur (in Swedish). 84: 1–160.
  49. Gurney, John Henry (1921). Early Annals of Ornithology. London: H F & G Witherby. p. 80 – via The Internet Archive.
  50. Elassar, Alaa (16 May 2020). "Wild white storks hatched in the UK for the first time in centuries". CNN. Retrieved 18 May 2020.
  51. "Wild white storks hatch in UK for first time in hundreds of years". The Guardian. 16 May 2020. Retrieved 18 May 2020.
  52. Tryjanowski, Piotr; Sparks, Tim H.; Profus, Piotr (2005). "Uphill shifts in the distribution of the White Stork Ciconia ciconia in southern Poland: The importance of nest quality". Diversity and Distributions. 11 (3): 219–23. doi:10.1111/j.1366-9516.2005.00140.x.
  53. Ptaszyk, J.; Kosicki, J.; Sparks, T.H.; Tryjanowski, P. (2003). "Changes in the timing and pattern of arrival of the White Stork (Ciconia ciconia) in western Poland". Journal für Ornithologie. 144 (3): 323–9. doi:10.1046/j.1439-0361.2003.03011.x.
  54. Sproll, Alexandra; Fiedler, Wolfgang (2001). "Digging in old data: Migration and causes of death in White Storks (Ciconia ciconia) according to ringing recovery data of the Vogelwarte Rossitten (Eastern Prussia) before the second World War". Euring Newsletter. European Union for Bird Ringing. 3. Archived from the original on 2011-07-22.
  55. Cramp 1977, p. 331.
  56. Reed, C.A.; Lovejoy, T.E. (1969). "The migration of the White Stork in Egypt and adjacent areas" (PDF). The Condor. 71 (2): 146–54. doi:10.2307/1366076. JSTOR 1366076 – via University of New Mexico: Searchable Ornithological Research Archive.
  57. Leshem, Yossi; Yom-Tov, Yoram (1998). "Routes of migrating soaring birds" (PDF). Ibis. 140: 41–52. doi:10.1111/j.1474-919X.1998.tb04539.x – via Tel Aviv University.
  58. Alexander, R. McNeill (1992). Exploring Biomechanics, Animals in Motion. New York: Scientific American Library. ISBN 978-0-7167-5035-2.
  59. Liechti, Felix; Ehrich, Dorothee; Bruderer, Bruno (1996). "Flight behaviour of White Storks Ciconia ciconia on their migration over southern Israel" (PDF). Ardea. 84: 3–13.
  60. Newton 2010, pp. 74–75.
  61. Newton 2010, p. 144.
  62. Newton 2010, p. 229.
  63. Chernetsov, Nikita; Berthold, Peter; Querner, Ulrich (2004). "Migratory orientation of first-year White Storks (Ciconia ciconia): Inherited information and social interactions". Journal of Experimental Biology. 207 (6): 937–43. doi:10.1242/jeb.00853. PMID 14766952.
  64. Meyburg, Bernd-U.; Matthes, Joachim; Meyburg, Christiane (2002). "Satellite-tracked Lesser Spotted Eagle avoids crossing water at the Gulf of Suez" (PDF). British Birds. 95: 372–6 – via Raptor Research.
  65. Spaar, Reto; Bruderer, Bruno (1996). "Soaring migration of Steppe Eagles Aquila nipalensis in southern Israel: Flight behaviour under various wind and thermal conditions" (PDF). Journal of Avian Biology. 27 (4): 289–301. doi:10.2307/3677260. JSTOR 3677260.
  66. Cocker, Mark; Mabey, Richard (2005). Birds Britannica. London: Chatto & Windus. p. 58. ISBN 978-0-7011-6907-7.
  67. Sinclair, Ian; Hockey, Phil; Arlott, Norman (2005). The Larger Illustrated Guide to Birds of Southern Africa. Cape Town, RSA: Struik. p. 34. ISBN 978-1-77007-243-5.
  68. Cramp 1977, p. 332.
  69. Vergara, Pablo; Aguirre, José I. (2006). "Age and breeding success related to nest position in a White Stork Ciconia ciconia colony". Acta Oecologica. 30 (3): 414–8. Bibcode:2006AcO....30..414V. doi:10.1016/j.actao.2006.05.008 – via ResearchGate.
  70. Harrison, Colin James Oliver (1965). "Allopreening as agonistic behaviour". Behaviour. 24 (3–4): 161–209. doi:10.1163/156853965X00011. JSTOR 4533105.
  71. Kahl, M. Philip (1984). "Spread-wing postures and their possible functions in Ciconiidae" (PDF). The Auk. 88 (4): 715–22. doi:10.2307/4083833. JSTOR 4086376 – via University of New Mexico: Searchable Ornithological Research Archive.
  72. Elphick, Chris; Dunning, John B. Jr; Sibley, David, eds. (2001). The Sibley Guide to Bird Life and Behaviour. London: Christopher Helm. p. 575. ISBN 978-0-7136-6250-4.
  73. Herholdt, J.J. (1987). "Recovery of a ring-maimed White Stork". Safring News. 16: 82.
  74. Lefebvre, Louis; Nicolakakis, Nektaria; Boire, Dennis (2002). "Tools and brains in birds" (PDF). Behaviour. 139 (7): 939–73. doi:10.1163/156853902320387918 – via McGill University.
  75. Cramp 1977, p. 334.
  76. Cramp 1977, p. 333.
  77. Tryjanowski, Piotr; Sparks, Tim H.; Jerzak, Leszek, eds. (2006). The White Stork in Poland. Poznań, Poland: Bogucki Wydaw. ISBN 978-83-60247-35-8.
  78. Tryjanowski, Piotr; Kosicki, Jakub Z.; Kuzniak, Stanisław; Sparks, Tim H. (2009). "Long-term changes and breeding success in relation to nesting structures used by the White Stork, Ciconia ciconia" (PDF). Annales Zoologici Fennici. 46: 34–8. doi:10.5735/086.046.0104.
  79. Vergara, Pablo; Gordo, O.; Aguirre, José I. (2010). "Nest size, nest building behaviour and breeding success in a species with nest reuse: The White Stork Ciconia ciconia" (PDF). Annales Zoologici Fennici. 47 (3): 184–94. doi:10.5735/086.047.0303.
  80. Vergara, Pablo; Aguirre, José I.; Fargallo, Juan A.; Dávila, José A. (2006). "Nest-site fidelity and breeding success in White Stork Ciconia ciconia". Ibis. 148 (4): 672–7. doi:10.1111/j.1474-919X.2006.00565.x.
  81. Wuczyński, Andrzej (2005). "The turnover of White Storks Ciconia ciconia on nests during spring migration". Acta Ornithologica. 40 (1): 83–5. doi:10.3161/068.040.0104.
  82. Haverschmidt, François (1949). The Life of the White Stork. Leiden: E. J. Brill. pp. 33–4. OCLC 1576336 – via Google Books.
  83. Tortosa, Francisco S.; Redondo, Tomas (1992). "Frequent copulations despite low sperm competition in White Storks (Ciconia ciconia)". Behaviour. 121 (3&4): 288–315. doi:10.1163/156853992X00408. hdl:10261/60750. JSTOR 4535031.
  84. van Pelt Lechner, A. A. (1911). Oologia Neerlandica : Eggs of Birds Breeding in the Netherlands. Vol. II. The Hague, Netherlands: Nijhof. p. 118 – via The Internet Archive.
  85. Zielinski, Piotr (2002). "Brood reduction and parental infanticide – are the White Stork Ciconia ciconia and the Black Stork C. nigra exceptional?" (PDF). Acta Ornithologica. 37 (2): 113–9. doi:10.3161/068.037.0207. Archived from the original (PDF) on 2012-03-15 – via University of Łódź.
  86. Tortosa, Francisco S.; Redondo, Tomas (1992). "Motives for parental infanticide in White Storks Ciconia ciconia". Ornis Scandinavica. 23 (2): 185–9. doi:10.2307/3676447. JSTOR 3676447.
  87. Aguirre, José I.; Vergara, Pablo (2007). "Younger, weaker White Stork (Ciconia ciconia) nestlings become the best breeders" (PDF). Evolutionary Ecology Research. 9: 355–64.
  88. Tsachalidis, Efstathios P.; Liordos, Vasilios; Goutner, Vassilis (2005). "Growth of White Stork Ciconia ciconia nestlings" (PDF). Ardea. 93 (1): 133–7 – via Aristotle University of Thessaloniki.
  89. Mužinic, Jasmina; Rašajski, Javor (1992). "On food and feeding habits in the White Stork, Ciconia c. ciconia, in the central Balkans". Ökologie der Vögel (Ecology of Birds). 14: 211–23.
  90. Van den Bossche 2002, p. 8.
  91. "EURING list of longevity records for European birds". EURING. 2010. Retrieved 2 December 2010.
  92. Henry, PY; Wey, GR; Balança, G (2011). "Rubber band ingestion by a rubbish dump dweller, the White Stork (Ciconia ciconia)". Waterbirds. 34 (4): 504–8. doi:10.1675/063.034.0414.
  93. Igaune, Kristine; Krams, Indrikis; Krama, Tatjana; Bobkova, Jadviga (2008). "White Storks Ciconia ciconia eavesdrop on mating calls of Moor Frogs Rana arvalis" (PDF). Journal of Avian Biology. 39 (2): 229–32. doi:10.1111/j.2008.0908-8857.04180.x. Archived from the original (PDF) on 2011-07-22. Retrieved 2010-12-10 – via Institute of Systematic Biology: Laboratory of Animal Ecology and Evolution.
  94. Antczak, Marcin; Konwerski, Szymon; Grobelny, Seweryn; Tryjanowski, Piotr (2002). "The food composition of immature and non-breeding White Storks in Poland". Waterbirds. 25 (4): 424–8. doi:10.1675/1524-4695(2002)025[0424:TFCOIA]2.0.CO;2.
  95. Parasharya, Bhavbhuti M.; Vyas, Raju (1998). "Foraging association of White Stork Ciconia ciconia with Blackbuck Antilope cervicapra". Journal of the Bombay Natural History Society. 95 (1): 112.
  96. Pande, Satish; Sant, Niranjan; Bhate, Rohan; Ponkshe, Aditya; et al. (2007). "Recent records of wintering white Ciconia ciconia and black C. nigra storks and flocking behaviour of White-necked Storks C. episcopus in Maharashtra and Karnataka states, India". Indian Birds. 3 (1): 28–32. ISSN 0973-1407 – via The Internet Archive.
  97. Akhtar, S. Asad; Tiwari, J.K. (1993). "Food piracy by a White Stork Ciconia ciconia (Linn.)". Journal of the Bombay Natural History Society. 90 (1): 90–91.
  98. Kitowski, Ignacy (2007). "Why do Montagu's Harriers disturb foraging sessions of White Storks in South-East Poland" (PDF). Berkut. 16 (1): 110–8. Archived from the original (PDF) on 2011-08-11.
  99. Arizaga, J.; Resano-Mayor, J.; Villanúa, D.; Alonso, D.; Barbarin, J.M.; Herrero, A.; Lekuona, J.M.; Rodriguez, R. (2018). "Importance of artificial stopover sites through avian migration flyways: a landfill‐based assessment with the White Stork Ciconia ciconia". Ibis. 160 (3): 542–553. doi:10.1111/ibi.12566.
  100. Błoszyk, Jerzy; Gwiazdowicz, Dariusz J.; Bajerlein, Daria; Halliday, Robert Bruce (2005). "Nests of the White Stork Ciconia ciconia (L.) as a habitat for Mesostigmatic Mites (Acari, Mesostigmata)" (PDF). Acta Parasitologica. 50 (2): 171–75.
  101. Bajerlein, Daria; Błoszyk, Jerzy; Gwiazdowicz, Dariusz J.; Ptaszyk, Jerzy; et al. (2006). "Community structure and dispersal of mites (Acari, Mesostigmata) in nests of the White Stork (Ciconia ciconia)". Biologia. 61 (5): 525–30. doi:10.2478/s11756-006-0086-9.
  102. Rothschild, Miriam; Clay, Theresa (1953). Fleas, Flukes and Cuckoos. A Study of Bird Parasites. London, UK: Collins. p. 152 – via The Internet Archive.
  103. Trouessart, Édouard Louis; Mégnin, Jean Pierre (1885). "Monographie du genre Freyana et description des espèces nouvelles du Musée d'Angers". Bulletin de la Société d'Études Scientifiques d'Angers (in French). 14: 29–45 – via The Internet Archive.
  104. "Acari Collection & Systematics". Royal Museum for Central Africa. Retrieved 10 March 2012.
  105. Pugh, Geoffrey John Frederick (1972). "The contamination of birds' feathers by Fungi". Ibis. 114 (2): 172–7. doi:10.1111/j.1474-919X.1972.tb02602.x.
  106. Fryderyk, Sławomira; Izdebska, Joanna N. (2009). "Chewing lice (Insecta, Phthiraptera) of the White Stork (Ciconia ciconia L.) in Poland" (PDF). Annales UMCS, Biologia. 64 (2): 83–8. doi:10.2478/v10067-010-0017-6.
  107. Ortega, Ynes R. (2006). Foodborne Parasites. New York: Springer. p. 121. ISBN 978-0-387-30068-9.
  108. Franssen, F.F.; Hooimeijer, J.; Blankenstein, B.; Houwers, Dirk J. (2000). "Giardiasis in a White Stork in The Netherlands". Journal of Wildlife Diseases. 36 (4): 764–6. doi:10.7589/0090-3558-36.4.764. PMID 11085441.
  109. Schuster, Rolf; Schaffer, Thoralf; Shimalov, Vladimir (2002). "[The helminth fauna of indigenous White Storks (Ciconia ciconia)]". Berliner und Munchener Tierarztliche Wochenschrift (in German). 115 (11–12): 435–9. PMID 12481650.
  110. Höfle, U.; Krone, O.; Blanco, J.M.; Pizarro, M. (2003). "Chaunocephalus ferox in free-living White Storks in central Spain". Avian Diseases. 47 (2): 506–12. doi:10.1637/0005-2086(2003)047[0506:CFIFWS]2.0.CO;2. PMID 12887215.
  111. Sitko, J.; Heneberg, P. (2015). "Composition, structure and pattern of helminth assemblages associated with central European storks (Ciconiidae)". Parasitology International. 64 (2): 130–4. doi:10.1016/j.parint.2014.11.004. PMID 25462710.
  112. Hayes, Edward B.; Komar, Nicholas; Nasci, Roger S.; Montgomery, Susan P.; et al. (2005). "Epidemiology and transmission dynamics of West Nile Virus Disease". Emerging Infectious Diseases. 11 (8): 1167–73. doi:10.3201/eid1108.050289a. PMC 3320478. PMID 16102302.
  113. Malkinson, M.; Banet, C. (2002). The role of birds in the ecology of West Nile Virus in Europe and Africa. Current Topics in Microbiology and Immunology. 267. pp. 309–22. doi:10.1007/978-3-642-59403-8_15. ISBN 978-3-642-63966-1. PMID 12082995.
  114. Zeller, Hervé G.; Schuffenecker, Isabelle (2004). "West Nile Virus: An overview of its spread in Europe and the Mediterranean Basin in contrast to its spread in the Americas". European Journal of Clinical Microbiology & Infectious Diseases. 23 (3): 147–56. doi:10.1007/s10096-003-1085-1. PMID 14986160.
  115. Malkinson, Mertyn; Banet, Caroline; Weisman, Yoram; Pokamunski, Shimon; et al. (2002). "Introduction of West Nile virus in the Middle East by migrating White Storks". Emerging Infectious Diseases. 8 (4): 392–7. doi:10.3201/eid0804.010217. PMC 2730252. PMID 11971773.
  116. Hubálek, Zdenek; Wegner, Elżbieta; Halouzka, Jiří; Tryjanowski, Piotr; et al. (2008). "Serologic survey of potential vertebrate hosts for West Nile Virus in Poland". Viral Immunology. 21 (2): 247–54. doi:10.1089/vim.2007.0111. PMID 18433332.
  117. Berthold, Peter; Fiedler, Wolfgang; Querner, Ulrich (2000). "White Stork (Ciconia ciconia) migration studies: Basic research devoted to conservation measures". Global Environment Research. 2: 133–41.
  118. "Annex 2: Waterbird Species to Which the Agreement Applies" (PDF). Agreement on the Conservation of African-Eurasian Migratory Waterbirds (AEWA). UNEP/ AEWA Secretariat. Archived from the original (PDF) on 28 July 2011. Retrieved 9 December 2010.
  119. "Introduction". African-Eurasian Waterbird Agreement. UNEP/ AEWA Secretariat. Archived from the original on 11 February 2012. Retrieved 9 December 2010.
  120. "Preliminary Results of the VI International White Stork Census 2004/05" (PDF). Naturschutzbund Deutschland (NABU). Archived from the original (PDF) on 12 August 2011.
  121. "National data". Naturschutzbund Deutschland (NABU). Retrieved 14 January 2012.
  122. Aghababyan K. 2011. White Storks (Ciconia ciconia L.): Population tendencies in Armenia. // Proceedings of the International Conference “Biological Diversity and Conservation Problems of the Fauna of the Caucasus”. September 26–29, Yerevan, Asoghik: 9-13.
  123. Aghababyan K., Kochinyan M., and Stepanyan L. (2013). White Storks (Ciconia ciconia L.) in Armenia: population, trend, and relationships to humans. // in Hötker, H. & K.-M. Thomsen (ed): White Stork on the top? - Results of the VIth International White Stork Census 2004/05. NABU (Naturschutzbund Deutschland e.V.), Berlin.
  124. White Stork in Armenia. 2016. Armenian Bird Census Council. Retrieved from http://www.abcc-am.org/white-stork.html
  125. Moore, Molly (22 June 2008). "French region saves iconic White Storks from brink: population hits 270 pairs after dwindling to nine". The Washington Post. p. A10.
  126. Altenburg, Jouke (2010). "Restoring the Dutch White Stork Population: A Charismatic Species Brought Back by Targeted Conservation Work" (PDF). BirdLife International. Vogelbescherming Netherlands (BirdLife Netherlands). Retrieved 30 January 2011.
  127. "The Swedish White Stork Reintroduction Program". Valkommen till Storkprojektet!. Scanian Ornithological Society (SkOF). 17 December 2010. Archived from the original on 25 December 2012. Retrieved 8 February 2011.
  128. Moritzi, Martin; Maumary, Lionel; Schmid, David; Steiner, Isabelle; et al. (2001). "Time budget, habitat use and breeding success of White Storks (Ciconia ciconia) under variable foraging conditions during the breeding season in Switzerland" (PDF). Ardea. 89 (3): 457–70.
  129. Schaub, Michael; Pradela, Roger; Lebretona, Jean-Dominique (2004). "Is the reintroduced White Stork (Ciconia ciconia) population in Switzerland self-sustainable?" (PDF). Biological Conservation. 119 (1): 105–14. doi:10.1016/j.biocon.2003.11.002.
  130. "Switzerland sees record stork numbers". SWI swissinfo.ch.
  131. "White Storks released in Surrey and Sussex". BirdGuides. 19 August 2019. Retrieved 19 August 2019.
  132. Crisp, Wil (7 July 2018). "White storks to breed in Britain for the first time in 600 years". The Telegraph.
  133. Margolis, Marvin; Parker, Philip (1972). "The stork fable−some psychodynamic considerations". Journal of the American Psychoanalytic Association. 20 (3): 494–511. doi:10.1177/000306517202000304. PMID 4116100.
  134. Schofield, A.F., ed. (1958). Aelian on the characteristics of animals. Volume I. Books I-V. Harvard University Press. p. 185.
  135. Dolata, Pawel T. (2006). "The White Stork Ciconia ciconia Protection in Poland by Fradition, Customs, Law, and Active Efforts". In Tryjanowski, Piotr; Sparks, Tim H.; Jerzak, L. (eds.). The White Stork in Poland: Studies in Biology, Ecology and Conservation. Poznań, Poland: Bogucki Wydawnictwo Naukowe. ISBN 978-83-60247-35-8.
  136. Royds, Thomas Fletcher (1914). The Beasts, Birds, and Bees of Virgil; A Naturalist's Handbook to the Georgics. Oxford University Press – via The Internet Archive.
  137. Thomas, Richard F. (1988). "Vergil's "White Bird" and the Alexandrian reference (G. 2. 319-20)" (PDF). Classical Philology. 83 (3): 214–7. doi:10.1086/367107.
  138. Sax, Boria (2001). The Mythical Zoo. Oxford, UK: ABC-CLIO. pp. 153–4. ISBN 978-1-57607-612-5.
  139. Kinzelbach, Ragnar (2003). "Der Pfeilstorch in der Zoologischen Sammlung der Universität Rostock" (PDF). Der Sprössling: Fachschaftsrat Biologie der Universität Rostock (in German) (3): 9–10. Archived from the original (PDF) on 2014-11-25.
  140. Unknown (1831). Descriptive Scenes for Children. Boston: N.S. and S.G. Simpkins. p. 3. OCLC 31373438 – via The Internet Archive.
  141. Hardwick, Charles (1872). Traditions, Superstition and Folk-lore. Manchester, UK: A. Ireland and Co. pp. 238, 247–8 – via The Internet Archive.
  142. "Stork". Encyclopedia of Ukraine. Canadian Institute of Ukrainian Studies. Retrieved 5 December 2010.
  143. Scharning, Kjell. "White Stork". Theme Birds on Stamps. Retrieved 5 December 2010.
  144. "Lithuania". Understanding the European Union: Member States. Centre d'Information sur les Institutions Européennes (CIIE). Archived from the original on 20 July 2011. Retrieved 5 December 2010.
  145. "Kingdom of Storks". PAIIZ Newsletter. Polish Information and Foreign Investment Agency (PAIIZ). 22 March 2007. Retrieved 18 March 2011.
  146. Czajkowski, Mikołaj; Giergiczny, Marek; Kronenberg, Jakub; Tryjanowski, Piotr (2014). "The economic recreational value of a white stork nesting colony: A case of 'stork village' in Poland". Tourism Management. 40: 352–360. doi:10.1016/j.tourman.2013.07.009.
  147. Hulme, F. Edward (1895). Natural History Lore and Legend. London: Bernard Quaritch. p. 260 – via The Internet Archive. The extraordinary idea that storks were found only in countries having a republican form of government held its ground for a considerable time, though it would appear as though nothing could have been simpler than its prompt disproof.
  148. Norwid, Cyprian Kamil. "My Song (II)". Polish Literature. LDS Mission Network. Archived from the original on 17 May 2011. Retrieved 23 December 2010.
  149. "Flugblätter für Afrika". Der Spiegel (in German). 10 January 1994. pp. 53–54. Retrieved June 30, 2018.
  150. Jacobs, Alvin H.; Walton, Robert G. (1976). "The incidence of birthmarks in the neonate". Pediatrics. 58 (2): 218–22. PMID 951136.
  151. Gieysztor, Aleksander (1982). Mitologia Słowian (in Polish). Warsaw: Wydawnictwa Artystyczne i Filmowe. ISBN 978-83-221-0152-0.
  152. Jakubiec, Z. (2009). "Dlaczego bocian przynosił dzieci?". Bocianopedia (in Polish). Retrieved 10 March 2011.
  153. Szczepanowicz, Barbara (2005). "Ptaki Ziemi Świętej: Bocian, czapla, ibis" [Birds in the Holy Land: Stork, heron, ibis]. Ziemia Święta (in Polish) (rok XI 1(41)). Retrieved 8 February 2011.
  154. Matthews, Robert (2000). "Storks deliver babies" (PDF). Teaching Statistics. 22 (2): 36–8. doi:10.1111/1467-9639.00013.
  155. Didelez, Vanessa (2007). "Statistical Causality" (PDF). In Østreng, Willy (ed.). Consilience: Interdisciplinary Communications 2005/2996. Centre for Advanced Study, Oslo. pp. 114–20. ISBN 978-82-996367-4-2.
  156. Bay, Mia (2000). The White Image in the Black Mind: African-American Ideas about White People, 1830–1925. New York: Oxford University Press. p. 120. ISBN 978-0-19-513279-3.
  157. Jung, Carl (1910). "The Association Method – Lecture III: Experiences concerning the psychic life of the child". American Journal of Psychology. 31 (3): 219–69. doi:10.2307/1413002. JSTOR 1422691 – via York University.
  158. Pickles, Penny (2011). "Jung the Man". The Society of Analytical Psychology: Jungian Analysis and Psychotherapy. London: The Society of Analytical Psychology. Archived from the original on 3 November 2011. Retrieved 2 March 2011.
  159. de Vries, Ad (1976). Dictionary of Symbols and Imagery. Amsterdam: North-Holland Publishing Company. p. 445. ISBN 978-0-7204-8021-4.

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 University Press. ISBN 978-0-19-857358-6.
  • Elliott, Andrew (1992). "Family Ciconiidae (Storks)". In del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi (eds.). Handbook of the Birds of the World. Volume 1: Ostrich to Ducks. Barcelona: Lynx Edicions. ISBN 978-84-87334-10-8.
  • Newton, Ian (2010). Bird Migration. Collins New Naturalist Library no. 113. London: Collins. ISBN 978-0-00-730732-6.
  • Svensson, Lars; Grant, Peter J. (1999). Collins Bird Guide. London: HarperCollins. ISBN 978-0-00-219728-1.
  • Van den Bossche, Willem (2002). Eastern European White Stork Populations: Migration Studies and Elaboration of Conservation Measures (PDF). (In collaboration with: Berthold, Peter; Kaatz, Michael; Nowak, Eugeniusz; Querner, Ulrich). Bonn: Bundesamt für Naturschutz (BfN)/German Federal Agency for Nature Conservation.
gollark: Or [EVERY OTHER PROXY IN EXISTENCE].
gollark: Can you access them through `archive.org` maybe?
gollark: Also, why do I have three IPv6 addresses listed under `ip addr`...?
gollark: Wico: Same in the US! And in the UK we have massive government spying too!
gollark: HTTPS won't obscure URLs.
This article is issued from Wikipedia. The text is licensed under Creative Commons - Attribution - Sharealike. Additional terms may apply for the media files.