History

The Angora is said[by whom?] to have originated in Ankara (historically known as Angora), in present-day Turkey, and is known to have been brought to France in 1723. The Angora rabbit became a popular pet of the French royalty in the mid-18th century, and Angoras had spread to other parts of Europe by the end of that century. In the United States, garments made of Angora-rabbit wool have been popular ever since they first arrived in the early 20th century. However, only during World War II did domestic production expand to meet the demand for more than 120,000 pounds a year. This valuable, soft, silky, fiber aroused much interest, and quickly people became enamored with the production process.

Angora-rabbit wool

Angoras are bred mainly for their wool, which is silky and soft. At only 14–16 micrometres in diameter, it is similar to cashmere in fineness and softness to the touch. A healthy adult Angora's wool will grow approximately 3 centimetres (1 in) per month. Regular grooming is necessary to prevent the fibre from matting and felting on the rabbit, which causes discomfort and can lead to pain and even infection. Angora wool is harvested (plucked or shorn) every three to four months throughout the year. The coat needs to be monitored after 6 months of regrowth since it may tend to "die" and easily mat.

Angora wool may be gathered periodically by hand-plucking the hairs within the coat that are being naturally shed. Full harvesting is done by shearing the coat with clippers or small scissors, often while the rabbit sits atop a groomer's turntable. Shearing typically starts at the head, moving across the shoulders to the tail. The rabbit is then flipped and the underside is shorn from tail to chin. Between 340 and 510 g (12 and 18 oz) of wool may be harvested from a Giant Angora.

Health

Main article: Domestic rabbit § Health

Wool block

Main article: Domestic rabbit § Digestion

Because of the length and abundance of their hair, Angora rabbits are particularly susceptible to wool block, a potentially lethal blockage of the digestive tract. All rabbits ingest some of their wool when they groom themselves, but their digestive system is not able to pass that foreign matter. The length of Angora hairs compounds the risk of impaction, which can lead to death. Clipping or plucking an Angora's wool every 90–120 days is necessary to prevent wool block.[citation neede

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Wool mites

Main article: Rabbit § Rabbit diseases

Cheyletiella parasitovorax is a skin parasite commonly found in Angora rabbits. Signs of infestation are flaky skin patches and fur loss. Wool mites reduce fiber yields and the resulting skin flakes are detrimental to the fiber quality. Wool mites may be treated with ivermectin or with carbaryl powder.

Angora rabbit breeds

The iconic long coat of the Angora is the result of a rabbit gene referred to as l (i.e., lowercase "L"). This "Angora gene" is present in all Angora breeds. It has also sometimes been utilized in the development of other rabbit breeds or other breeds' new varieties. "Dwarf Wooly" breeds including American Fuzzy Lop, Lion Head and Jersey Woolie are now recognized in the U.S. by ARBA. Belgium and France have their own Dwarf Wooly breeds. There is also a rare Mini English Angora breed in New Zealand.

English Angora

Before 1939 there was one breed of "Angora Wooler". In 1939 ARBA reclassified 'Angora Wooler' as English Type and French Type. In 1944 ARBA officially separated Angora rabbits into two breeds: English Angora and French Angora.

Rabbits of the Angora breed are adorned with "fur", growths of wool on the ears and the entire face except above the nose, and front feet, along with their thick body, and wool. They are gentle in nature, but they are not recommended for those who do not groom their animals. Their wool is very dense and needs to be groomed twice a week.

This is the smallest Angora rabbit of the four ARBA-recognized breeds. This breed is more common as a pet because of the facial features that give it a puppy or teddy-bear look. If the texture of the wool is correct, the maintenance is relatively easy; if the texture of the rabbit is cottony, it requires a great deal of maintenance. Beginning spinners may find the wool challenging.

The English Angora can be bred to have broken colors—i.e., white with black spots—but this is not accepted by ARBA standards and would lead to a disqualification when showing the rabbit. When an English Angora rabbit is shown, the toenails should also be only one color, the ears could be folded over at the tips and the furnishings on the face may cover their eyes. The English Angora is the only one of the Angora breeds that has hair covering its eyes.

French Angora

This breed has a dense undercoat. If the texture is correct it requires less maintenance than other Angora breeds. Small ear tufts are allowed but not usually preferred by breeders. ARBA recognizes the same colors as with English Angora, plus ticked and wide band. They are shown at ARBA shows using the types 'white' and 'colored' (broken being a colored). As with other ARBA-shown rabbits, toenails should also be only one color.

The French Angora is one of the large Angora breeds at 7.5–10.5 pounds (3.4–4.8 kg), with a commercial body type. It differs from the English, Giant and German Angora in that it possesses a clean (hairless) face and front feet with only minor tufting on the rear legs. The color of a French Angora is determined by the color of its head, feet and tail (all the same color). This variety of angora fibre has smooth silky texture. Beginning spinners may find Angora wool a challenge. Desirable characteristics of the fibre include its texture, warmth, light weight and pure white color. It is used for sweaters, mittens, baby clothes and millinery.

German Angora

• Weight: 2.5–5.5 kg (5+1⁄2–12 lb).

• IAGARB-accepted varieties: Albino or Colored (but not bi-colored)

• Albino Color Group

• Albino

• Self Group

  • Black

  • Dilute Blacka/k/a Blue

  • Browna/k/a Chocolate

  • Dilute Browna/k/a Lilac

• Tortoiseshell Group

  • Tortoiseshell

  • Dilute Black Tortoiseshella/k/a Blue Tortoiseshell

  • Brown Tortoiseshella/k/a Chocolate Tortoiseshell

  • Dilute Brown Tortoiseshella/k/a Lilac Tortoiseshell

• Agouti Group

  • Agoutia/k/a Black Agouti, Chestnut Agouti, Wild Agouti

  • Dilute Black Agoutia/k/a Blue Agouti or Opal

  • Brown Agoutia/k/a Chocolate Agouti

  • Dilute Brown Agoutia/k/a Lilac Agouti or Lynx

• Wide Band Color Group

  • Yellowa/k/a Red or Fawn

• Chinchilla Color Group

  • Chinchilla

  • Dilute Black Chinchillaa/k/a Blue Chinchilla or Squirrel

  • Brown Chinchillaa/k/a Chocolate Chinchilla

  • Dilute Brown Chinchillaa/k/a Lilac Chinchilla

Though common, the German Angora is not currently recognized by ARBA. The International Association of German Angora Rabbit Breeders (IAGARB) maintains a breed standard for the German Angora.

Giant Angora

• Weight: Minimum 5.4 kg (12 lb)

• ARBA-recognized varieties: Ruby-eyed White

• The Giant Angora is the largest of the ARBA-recognized Angora breeds. It was originally developed to be an efficient commercial producer that could be sustained on 16–18% protein pellets plus hay, and live in the standard sized, all-wire cages.

  Because ARBA wouldn't allow German Angoras to be shown, their body type being considered too similar to other Angora breeds, Louise Walsh of Taunton, Massachusetts, created a new breed. She used German Angoras, French Lops and Flemish Giants to develop a completely different 'commercial' body type. ARBA officially recognized the Giant Angora in 1988. Its coat includes three types of wool: soft underwool, awn fluff and awn hair.

  The awn-type wool exists only in the Giant and German Angora breeds. The Giant Angora has furnishings on the face and ears. Many people confuse the German with the Giant Angora, but it is their body type that differs.

  The only color variety ARBA currently recognizes for the Giant Angora is the Ruby-eyed White (REW), a color that indicates the genetic absence of pigment (albino). The Giant Angora produces more wool than the French, Satin or English Angoras. The Giant Angora is the only 6-class animal in the Angora breed. It should have a commercial-type body with a very dense coat of wool. The head should be oval in appearance, that is broad across the forehead and slightly narrower at the muzzle. The Giant Angora should have forehead tufts (head trimmings) and cheek furnishings. The head trimmings should be noticeable, but does have lighter trimmings than bucks. The ears should be lightly fringed and well tasseled. The Giant Angora is also the only breed of angora that is shown only as a ruby-eyed white. A Black color variety of the Giant Angora is in development but has not been sanctioned by ARBA.

  The Giant Angora coat contains three fiber types for its texture. The underwool should be the most dominant over the other two types of hair. It should be medium-fine, soft and delicately waved and have a gentle shine. Beginning spinners may find Angora wool a challenge.

  The Awn Fluff has a guard hair tip and is a stronger, wavy wool. The Awn Fluff is found between the underwool and Awn Hair. The Awn Hair, also known as guard hair, is the third type of fiber. The Awn Hair is a strong straight hair that protrudes above the wool and must be present and evident.

  The classification of the Giant Angora is different from the other three breeds owing to it being a 6-class animal. The junior buck and junior doe must be under 6 months of age and have a minimum weight of 4 ¾ pounds. The intermediate buck and intermediate doe are 6–8 months of age. The senior buck and senior doe are 8 months of age or over. The senior buck must weigh at least 9 ½ pounds. The senior doe must weigh at least 10 pounds.

  When Giant Angoras are judged the majority of the points are based on the wool, which includes density, texture and length. The points for 'general type' include the body type, head, ears, eyes, feet, legs and tail.

  Like many other 'giant' breeds of rabbit, the Giant Angora grows slowly. A doe usually takes more than a year to reach maturity (size and weight). A buck can take up to 1.5 years to mature (size and weight).

  Satin Angora

*

• Weight: 3.0–4.5 kg (6+1⁄2–10 lb).

  • ARBA-recognized varieties: [Includes eight color groups. The color of a Satin Angora is determined by the uniform pigment on its head, feet, and tail.]

It is best known from the type species A. canadensis, found in the Stephen Formation (particularly the Burgess Shale) of British Columbia, Canada. The species A. daleyae is known from the somewhat older Emu Bay Shale of Australia. Other remains are known from China and the United States. Originally, several fossilized parts discovered separately (the mouth, frontal appendages and trunk) were thought to be three separate creatures, a misapprehension corrected by Harry B. Whittington and Derek Briggs in a 1985 journal article.

Like other radiodonts, Anomalocaris had swimming flaps running along its body, large compound eyes, and a single pair of segmented, "frontal appendages", which in Anomalocaris were used to grasp prey. Measuring up to 38 cm (1.25 ft) long excluding frontal appendages and tail fan, A. canadensis is one of the largest animals of the Cambrian, and thought to be one of the earliest examples of an apex predator, though others have been found in older Cambrian lagerstätten deposits.

Discovery and identification

From the start, Anomalocaris fossil was misidentified, followed by a series of misidentifications and taxonomic revisions. As Stephen Jay Gould, who popularised the Cambrian explosion in his 1989 book Wonderful Life, appropriately described:

[The story of Anomalocaris is] a tale of humor, error, struggle, frustration, and more error, culminating in an extraordinary resolution that brought together bits and pieces of three "phyla" in a singe reconstructed creature, the largest and fiercest of Cambrian organisms.

Anomalocaris fossils were first collected in 1886 by Richard G. McConnell of the Geological Survey of Canada (GSC). Having been informed of rich fossils at the Stephen Formation in British Columbia, McConnell climbed Mount Stephen on 13 September 1886. He found abundant trilobites, along with two unknown specimens. In August 1891, Henri-Marc Ami, Assistant Palaeontologist at GSC, collected many trilobites and brachiopod fossils, along with 48 more of the unknown specimens. The fifty specimens were examined and described in 1892 by GSC paleontologist Joseph Frederick Whiteaves. Whiteaves interpreted them as the abdomens of phyllocarid crustaceans, and gave the full scientific name Anomalocaris canadensis. He describe the crustacean characters:

Body or abdominal segments, which, in all the specimens collected, are abnormally flattened laterally, a little higher or deeper than long, broader above than below, the pair of ventral appendages proceeding from each, nearly equal in height or depth to the segment itself... The generic name Anomalocaris (from ανώμαλος, unlike,—καρίς, a shrimp, i.e., unlike other other shrimps) [the species name referring to Canada] is suggested by the unusual shape of the uropods or ventral appendages of the body segments and the relative position of the caudal spine.

In 1928, Danish paleontologist Kai Henriksen proposed that Tuzoia, a Burgess Shale arthropod which was known only from the carapace, represented the missing front half of Anomalocaris. The artists Elie Cheverlange and Charles R. Knight followed this interpretation in their depictions of Anomalocaris.

Unbeknownst to scientists at the time, the body parts of relatives of Anomalocaris had already been described but not recognized as such. The first fossilized mouth of such a kind of animal was discovered by Charles Doolittle Walcott, who mistook it for a jellyfish and placed it in the genus Peytoia. Walcott also discovered a frontal appendage but failed to realize the similarities to Whiteaves' discovery and instead identified it as feeding appendage or tail of the coexisted Sidneyia. In the same publication in which he named Peytoia, Walcott named Laggania, a taxon that he interpreted as a holothurian.

In 1966, the Geological Survey of Canada began a comprehensive revision of the Burgess Shale fossil record, led by Cambridge University paleontologist Harry B. Whittington. In the process of this revision, Whittington and his students Simon Conway Morris and Derek Briggs would discover the true nature of Anomalocaris and its relatives, but not without contributing to the history of misinterpretations first. In 1978, Conway Morris recognized that the mouthparts of Laggania were identical to Peytoia, but concluded that Laggania was a composite fossil made up of Peytoia and the sponge Corralio undulata. In 1979, Briggs recognized that the fossils of Anomalocaris were appendages, not abdomens, and proposed that they were the walking legs of a giant arthropod, and that the feeding appendage Walcott had assigned to Sidneyia was the feeding appendage of similar animal, referred to as "appendage F". Later, while clearing what he thought was an unrelated specimen, Harry B. Whittington removed a layer of covering stone to discover the unequivocally connected frontal appendage identical to Anomalocaris and mouthpart similar to Peytoia. Whittington linked the two species, but it took several more years for researchers to realize that the continuously juxtaposed Peytoia, Laggania and frontal appendages (Anomalocaris and "appendage F") actually represented a single group of enormous creatures. The two genera have now been placed into the order Radiodonta and are commonly known as radiodonts or anomalocaridids. Since Peytoia was named first, it is the accepted correct name for the entire animal. However, the original frontal appendage was from a larger species distinct from Peytoia and "Laggania" and therefore retains the name Anomalocaris.

In 2011 and 2020, compound eyes of Anomalocaris were recovered from a paleontological dig at Emu Bay on Kangaroo Island, Australia, proving that Anomalocaris was indeed an arthropod as had been suspected. The find also indicated that advanced arthropod eyes had evolved very early, before the evolution of jointed legs or hardened exoskeletons.

In 2021, "A." saron and "A." magnabasis were reassigned to the new genus Houcaris, in the family Tamisiocarididae. In the same year, "A." pennsylvanica was reassigned to the genus Lenisicaris. In 2022, specimen ELRC 20001 that was treated as an unnamed species of Anomalocaris or whole-body specimen of A. saron got a new genus, Innovatiocaris. In 2023, "A". kunmingensis was reassigned to the new genus Guanshancaris in the family Amplectobeluidae. Multiple phylogenetic analyses also suggested that "A". briggsi (tamisiocaridid) was not a species of Anomalocaris either, and it was reassigned to the genus Echidnacaris in the family Tamisiocarididae in 2023. In the same 2023 study, a new species of Anomalocaris, A. daleyae, was described based on remains found in the Emu Bay Shale in Australia.

Description

Size estimation of Anomalocaris.

For the time in which it lived, Anomalocaris was gigantic, A. canadensis is estimated to be up to 37.8 cm (1.24 ft) long excluding frontal appendages and tail fans. Previous estimation up to 1 m (3.3 ft) is unlikely based on the ratio of body parts (body length measured only about 2 times the length of frontal appendage in A. canadensis, respectively) and the size of largest frontal appendage (up to 18 centimetres (7.1 inches) in length when extended).

Anomalocaris propelled itself through the water by undulating the flexible flaps on the sides of its body. Each flap sloped below the one more posterior to it, and this overlapping allowed the lobes on each side of the body to act as a single "fin", maximizing the swimming efficiency. The construction of a remote-controlled model showed this mode of swimming to be intrinsically stable, implying that Anomalocaris would not have needed a complex brain to manage balance while swimming. The body was widest between the third and fifth lobe and narrowed towards the tail, with additional 3 pairs of small flaps on the constricted neck region. It is difficult to distinguish lobes near the tail, making an accurate count difficult. For the main trunk flaps, the type species A. canadensis had 13 pairs.

Anomalocaris had an unusual disk-like mouth known as oral cone. The oral cone was composed of several plates organized triradially. Three of the plates were quite large. Three to four medium sized plates could be found between each of the large plates, and several small plates between them. Most of the plates wrinkled and possess scale-like tubercles near the mouth opening. Such an oral cone is very different from those of a typical hurdiid radiodont like Peytoia and Hurdia, which is smooth and tetraradial. As a shared character across radiodonts, Anomalocaris also possessed three sclerites on the top and side of its head. The top one, known as a head shield, dorsal carapace or H-element, was shaped like an laterally-elongated oval, with a distinct rim on the outer edge. The remaining two lateral sclerites, known as P-elements, were also ovoid, but connected by a bar-like outgrowth. The P-elements were previously misinterpreted as two huge compound eyes.

Frontal appendages of Anomalocaris, with examples from multiple species.

Two large frontal appendage were positioned in front of the mouth, at the front of the head. Each frontal appendage of Anomalocaris usually possessed 14 podomeres (segmental units, at least 1 for shaft and 13 for distal articulated region), with each appendage being laterally-flattened (taller than wide). Most podomeres were tipped with a pair of endites (ventral spines). The endites themselves were both equipped with multiple auxiliary spines, which branches off from the anterior and posterior margin of the endites.

The tail was a large tail fan, composed of three pairs of large, lateral fin-shaped lobes and one terminal lobe-like tailpiece. Previous studies suggest the tail fan was used to propel it through Cambrian waters, while further hydrodynamic study rather suggest it was more adapted to provide steering function. The gills of the animal, in the form of long, thin, hair-like structures known as lanceolate blades, were arranged in rows forming setal blades. The setal blades were attached by their margin to the top side of the animal, two setal blades per body segment. A divide ran down the middle, separating the gills.

Based on fossilized eyes from the Emu Bay Shale, which belong to the species Anomalocaris daleyae, the stalked eyes of Anomalocaris were 30 times more powerful than those of trilobites, long thought to have had the most advanced eyes of any contemporary species. With one specimen having over 24,000 lenses in one eye, the resolution of the 3-centimetre-wide (1.2 in) eyes would have been rivalled only by that of the modern dragonfly, which has 28,000 lenses in each eye. Additionally, estimation of ecdysozoan opsins suggest that Anomalocaris may have had dichromatic color vision.

Paleobiology

Diet

Grasping movement of the frontal appendage of A. canadensis.

The interpretation of Anomalocaris as an active predator is widely accepted throughout the history of research, as its raptorial frontal appendages and mid-gut glands strongly suggest a predatory lifestyle. In the case of A. canadensis, its outstanding size amongst Burgess Shale fauna also make it one of the first apex predators known to exist.

However, the long-standing idea that Anomalocaris fed on hard-bodied animals, especially its ability to penetrate mineralized exoskeleton of trilobites, has been questioned, with many recent studies considering it more likely that Anomalocaris exclusively hunted soft-bodied prey. Some Cambrian trilobites have been found with round or W-shaped "bite" marks, which were identified as being the same shape as the mouthparts of Peytoia (previously misidentified as those of Anomalocaris). Stronger evidence that Anomalocaris ate trilobites comes from coprolite, which contain trilobite parts and are so large that the anomalocarids are the only known organism from that period large enough to have produced them. However, since Anomalocaris lacks any mineralized tissue, it seemed unlikely that it would be able to penetrate the hard, calcified shell of trilobites. Rather, the coprolites may have been produced by a different organisms, such as the trilobites of the genus Redlichia. Another suggested possibility was that Anomalocaris fed by grabbing one end of their prey in its oral cone while using its frontal appendages to quickly rock the other end of the animal back and forth. This produced stresses that exploited the weaknesses of arthropod cuticles, causing the prey's exoskeleton to rupture and allowing the predator to access its innards. This behaviour was originally thought to have provided an evolutionary pressure for trilobites to roll up, to avoid being flexed until they snapped.

Ecological reconstruction of Anomalocaris hunting Isoxys, after posture estimated in Bicknell et al. (2023)

The lack of wear on radiodont mouthparts suggests they did not come into regular contact with mineralized trilobite shells, and were possibly better suited to feeding on smaller, soft-bodied organisms by suction, since they would have experienced structural failure if they were used against the armour of trilobites. A. canadensis was suggested to have been capable of feeding on organisms with hard exoskeletons due to the short, robust spines on its frontal appendages. However, this conclusion is solely based on the comparison with the fragile frontal appendages of suspension feeding radiodonts (e.g. "A". briggsi and Houcaris spp.). The typical lack of damage to the endites on the frontal appendages of A. canadensis (with damage only present on a single specimen) suggests that they were not used to grasp hard-shelled prey. As opposed to Peytoia whose oral cone is more rectangular with short protruding spines, the oral cone of A. canadensis has a smaller and more irregular opening, not permitting strong biting motions, and indicating a suction-feeding behavior to suck in softer organisms. Three-dimensional modelling of various radiodont frontal appendages also suggest that A. canadensis is more capable to prey on smaller (2–5 cm in diameter), active, soft-bodied animals (e.g. vetulicolian; free-swimming arthropods like isoxyids and hymenocarines; Nectocaris).

Bicknell et al. (2023) examined the frontal appendages of Anomalocaris, suggesting it was an active nektonic apex predator. Postured with the frontal appendages outstretched, Anomalocaris would have been able to swim with maximized speed, similar to modern predatory water bugs. Its eyes would be suitable to hunt prey in well-lit waters. Anomalocaris would have hunted various free-swimming animals since there are a large diversity of nektonic and pelagic soft-bodied animals. It probably would have not hunted benthic animals like trilobites, considering the possibility of damaging the frontal appendages on the substrate while trying to grab prey from seafloor at speed. Instead, other animals such as other radiodonts (e.g. Hurdia, Cambroraster, Titanokorys, Stanleycaris) and artiopods (e.g. Sidneyia) would have been benthic predators in the Burgess Shale.

Paleoecology

Specimens of Anomalocaris have been found worldwide spanning from Cambrian Stage 3 to the Guzhangian. Aside from the Burgess Shale and Emu Bay Shale, fossils have been found in the Chengjiang Biota, Hongjingshao Formation, Balang Formation and the Kaili Formation of China, as well as the Eagar Formation and Weeks Formation in the United States.

Anomalocaris canadensis lived in the Burgess Shale in relatively great numbers. In the Burgess Shale, Anomalocaris is more common in the older sections, notably the Mount Stephen trilobite beds. However, in the younger sections, such as the Phyllopod bed, Anomalocaris could reach much greater sizes; roughly twice the size of its older, trilobite bed relatives. These rare giant specimens have previously been referred to a separate species, Anomalocaris gigantea; however, the validity of this species has been called into question, and is currently synonymized to A. canadensis.

Other unnamed species of Anomalocaris live in vastly different environments. For example, Anomalocaris cf. canadensis (JS-1880) lived in the Maotianshan Shales, a shallow tropical sea or even being delta in what is now modern China. Anomalocaris daleyae (Emu Bay Shale) lived in a comparable environment; the shallow, tropical waters of Cambrian Australia. The Maotianshan Shale and the Emu Bay Shale are very close in proximity, being separated by a small landmass, far from the Burgess Shale. These two locations also included "Anomalocaris" kunmingensis and "Anomalocaris" briggsi respectively, species that previously attributed but taxonomically unlikely to be a member of Anomalocaris nor even Anomalocarididae.

The water pipit in breeding plumage has greyish-brown upperparts, weakly streaked with darker brown, and pale pink-buff underparts fading to whitish on the lower belly. The head is grey with a broad white supercilium ("eyebrow"), and the outer tail feathers are white. In winter, the head is grey-brown, the supercilium is duller, the upperparts are more streaked, and the underparts are white, streaked lightly with brown on the breast and flanks. There are only minor differences among the three subspecies, the sexes are almost identical, and young birds resemble adults. The water pipit's song is delivered from a perch or in flight, and consists of four or five blocks, each consisting of about six repetitions of a different short note.

Water pipits construct a cup-like nest on the ground under vegetation or in cliff crevices and lay four to six speckled grey-ish white eggs, which hatch in about two weeks with a further 14–15 days to fledging. Although pipits occasionally catch insects in flight, they feed mainly on small invertebrates picked off the ground or vegetation, and also some plant material.

The water pipit may be hunted by birds of prey, infested by parasites such as fleas, or act as an involuntary host to the common cuckoo, but overall its population is large and stable, and it is therefore evaluated as a species of least concern by the International Union for Conservation of Nature (IUCN).

Taxonomy and systematics

The family Motacillidae consists of the wagtails, pipits and longclaws. The largest of the three groups is the pipits, genus Anthus, which are typically brown-plumaged terrestrial insectivores. Their similar appearances have led to taxonomic problems; the water pipit and the buff-bellied pipit were both formerly considered subspecies of the European rock pipit. Of these, the rock pipit is the more closely related to the water pipit, based on external and molecular characteristics. Other near relatives are the meadow, red-throated and rosy pipits.

The water pipit was first described by Carl Linnaeus in the tenth edition of his Systema Naturae in 1758 as Alauda spinoletta (characterised as A. rectricibus fuscis : extimis duabus oblique dimidiato-albis). The current genus Anthus was created for the pipits by German naturalist Johann Matthäus Bechstein in 1805. Anthus is the Latin name for a small bird of grasslands, and the specific spinoletta is a local dialect word for a pipit from the Florence area of Italy.

There are three recognised subspecies of the water pipit:

• Anthus spinoletta spinoletta, (Linnaeus, 1758), the nominate subspecies, breeds in the mountains of southern Europe from the Pyrenees eastwards to northwestern Turkey, and on Sardinia and Corsica.

• Anthus spinoletta coutellii, Audouin, 1828, breeds in the mountains of Turkey other than the northwest, the Caucasus, northern Iran and Turkmenistan.

• Anthus spinoletta blakistoni, R. Swinhoe, 1863, breeds in mountains across Asia from southern Russia east to central China.

A possible fourth race from the northwestern Caucasus, Anthus spinoletta caucasicus, cannot be reliably separated from A.s. coutellii. The latter form is itself a potential new species, based on genetic data, appearance and a characteristic flight call.

Description

The water pipit is 15–17 centimetres (5.9–6.7 in) long and weighs 18.7–23 grams (0.66–0.81 oz). The adult of the nominate race in spring plumage has greyish-brown upperparts, weakly streaked with darker brown, and pale pink-buff underparts fading to whitish on the lower belly. There may be some faint streaking on the breast and flanks. The head is grey with a broad white supercilium. The outer tail feathers are white, and the legs, bill and iris are dark brown or blackish. In non-breeding plumage, the head is grey-brown and the supercilium is less distinct. The upperparts are more streaked, and the underparts are white, marked lightly with brown on the breast and flanks.

The sexes are similar although the female has, on average, a greyer head. Young birds resembles the non-breeding adult, but are browner and more streaked above with prominent streaking on the underparts. A. s. coutellii is smaller than the nominate subspecies and the white of the outer tail feathers has a hint of grey. It is paler and more heavily streaked above, and in summer plumage the underparts' colour covers a larger area and has a rusty tint. A. s. blakistoni is large, pale and less strongly streaked.

The water pipit has a complete moult between July and September, although there is considerable individual variation in timing. There is a partial pre-breeding moult, mainly between January and March, but with much variability in timing. This moult typically involves replacing the head, body and some wing feathers, but the extent is again variable. Very occasionally, females may moult into what looks like non-breeding plumage, rather than the expected brighter garb. The chicks start to gain juvenile plumage as soon as a month after hatching, and most have completed the transition to near-adult appearance by September. The first pre-breeding moult is similar to that of the adult, but may be less complete or even absent.

The water pipit is closely related to the Eurasian rock pipit and the meadow pipit, and is rather similar to both in appearance. Compared to the meadow pipit, the water pipit is longer-winged and longer-tailed than its relative, and has much paler underparts. It has dark, rather than pinkish-red, legs. The water pipit in winter plumage is also confusable with the Eurasian rock pipit, but has a strong supercilium, greyer upperparts, and white, not grey, outer tail feathers; it is also typically much warier. The habitats used by European rock and water pipits are completely separate in the breeding season, and there is little overlap even when birds are not nesting. There is also little mixing with breeding meadow pipits, although since 1960 some overlapping territories have been found where the species coexist.

The European rock pipit's subspecies Anthus petrosus littoralis in summer plumage is particularly close in outward appearance to the water pipit. The rock pipit normally has a bluer tint to the head, streaking on the breast and flanks, and buff outer tail feathers, and the songs are also different. The species mostly occupy different habitat types even when they occur in the same general area.

Voice

The water pipit's song is delivered from a perch or in flight, and consists of four or five blocks, each consisting of about half a dozen repetitions of a different short note. In comparison, the European rock pipit's song is a sequence of about twenty tinkling cheepa notes followed by a rising series of thin gee calls, and finishing with a short trill. The call of the water pipit is a single or double sharp "dzip" or similar, slightly harsher than soft sip sip sip of the meadow pipit or the shrill pseep of the European rock pipit. The short, thin fist flight call is intermediate between the sip of the meadow pipit and the rock pipit's feest. The differences between the calls of the pipit species are very subtle, and not diagnostic in the absence of other evidence. The flight call of the subspecies A. s. coutellii is shorter and more buzzing than that of the nominate race.

Distribution and habitat

The breeding range of the water pipit is the mountains of southern Europe and Asia from Spain to central China, along with the Mediterranean islands of Sardinia and Corsica.

The water pipit is predominantly a mountain species in the breeding season, found in alpine pasture and high meadows with short grass and some bushes or rocks. It is typically found close to wetter areas and often on slopes. It breeds between 615–3,200 metres (2,020–10,500 ft) altitude, mostly 1,400–2,500 metres (4,600–8,200 ft). It migrates relatively short distances in autumn to lower ground, typically wintering on coastal wetlands, marshes, rice fields and similar habitats. Although most birds move to lowlands, some may remain at up to 2,000 metres (6,600 ft). Nominate A. s. spinoletta winters mainly in western and southern Europe and in northwestern Africa; in western Europe some birds show fidelity to the same wintering site, returning each year. Birds in Spain appear to move only lower down the mountains in which they breed. A.s. coutellii winters at lower altitudes near its breeding areas and also in the Arabian Peninsula and northeast Africa. A.s. blakistoni winters in Pakistan, northwest India and southern China.

Water pipits leave their breeding sites from mid-September, although the eastern subspecies may start moving south before then. The spring migration starts in February and March, with arrival on the breeding grounds in April and May. The water pipit has been recorded as a vagrant in Belarus, Gibraltar and Latvia, and on islands including the Canaries, Iceland, Malta and Svalbard.

Behaviour

The water pipit is a much less approachable bird as compared to the European rock pipit. It is warier than its relative and if approached it flies some distance before landing again, whereas the rock pipit typically travels only a short distance, close to the ground, before it alights.

Breeding

The water pipit is mainly monogamous, although both sexes may deviate from this occasionally. The male has a display flight in which he climbs to 10–30 metres (33–98 ft), flies in an arc and glides back down, singing throughout. The female constructs a cup nest from grass and leaves which is lined with finer plant material and animal hairs. The nest is hidden in vegetation on the ground, sometimes in a hollow. The normal clutch is four to six eggs laid from the end of April to early July. Eggs are greyish white with darker grey or brownish speckles mainly at the wider end, and they measure 21 by 16 millimetres (0.83 in × 0.63 in) and weigh 2.7 grams (0.095 oz) of which 5% is shel

l.

The eggs are incubated by the female for 14–15 days to hatching. Chicks are fed initially by the male, both parents sharing the duty after a few days when the female does not need to brood so often, and they fledge in a further 14–15 days. There may be two broods in a year.

In a Swiss study of the nominate subspecies, 76% of eggs hatched, and 58% of chicks fledged. Birds of the race A. s. blakistoni in the Tian Shan hatched 90% of their eggs, and hatchlings survived to fledging in 47% of the nests. In the latter study, early nests were more likely to fail because less plant cover made them more likely to be found by predators. Neither the average lifespan nor the maximum age of survival are known.

Feeding

The water pipit's feeding habitat is damp grassland, rather than the rocky coasts favoured by the Eurasian rock pipit. The water pipit feeds mainly on a wide range of invertebrates, including crickets and grasshoppers, beetles, snails, millipedes and spiders. Barkflies, true flies, caterpillars and homopterans can form a large part of the diet of fledglings. Birds close to snow fields take insects specialised for that habitat such as the springtails Isotoma saltans (the glacier flea) and I. nivalis, and the scorpion fly Boreus izyemalis.

Birds normally forage alone or in pairs; in bad weather, foraging is more frequent and involves longer flights, and may be concentrated around marmot burrow entrances. Prey items average 8.3 millimetres (0.33 in) in length and are mainly hunted on foot, although flying insects are occasionally caught in the air. Some plant material is taken, and one study on the border of Czechoslovakia and Poland found that 75% of the diet by volume consisted of algae, specifically Ulothrix zonata, despite large numbers of insects being available.

In areas with acidic soils, there is less calcium available, potentially leading to thinner egg shells. In such locations, pipits are more likely to select snails and similar prey with calcium-rich shells than is the case in limestone terrain.

Predators and parasites

The water pipit is hunted by birds of prey including the Eleonora's falcon, and eggs and young may be taken by terrestrial predators including stoats and snakes. As with other members of its genus, the water pipit is a host of the common cuckoo, a brood parasite. Eggs of cuckoos that specialise in parasitising pipits are similar in appearance to those of their hosts.

A new species of feather mite, Proctophyllodes schwerinensis, was discovered on the water pipit, which is also a host to the fleas Ceratophyllus borealis and Dasypsyllus gallinulae. Along with other Motacillidae species, the water pipit is a host of the protozoan parasite Haemoproteus anthi.

Status

Estimates of the European breeding population of the water pipit vary widely, but may be as high as two million pairs, which would suggest a global population of tens of millions of individuals spread over 3.7 millionsquare kilometres (1.4 millionsq mi). The range is discontinuous due to the mountain habitat this species uses, but the population is considered overall to be large and stable, and for this reason the water pipit is evaluated as a species of least concern by the IUCN.

Breeding densities (in pairs per ten ha) have been recorded as 2.4 in the Jura Mountains, 3.0–3.6 in the Alps and 4.5 in the Tatra Mountains of Poland.

It is common in well-wooded areas of the Indian Subcontinent and Southeast Asia. Most populations are resident. The species shows a preference for shady damp areas, and like many Geokichla and Zoothera thrushes, can be quite secretive.

The orange-headed thrush is omnivorous, eating a wide range of insects, earthworms and fruit. It nests in trees but does not form flocks.

The male of this small thrush has uniform grey upperparts, and an orange head and underparts. The females and young birds have browner upper parts.

Taxonomy

This species was first described by John Latham in 1790 as Turdus citrinus, the species name meaning "citrine" and referencing the colour of the head and underparts. It has about 12 subspecies. Rasmussen and Anderton (2005) suggest that this complex may consist of more than one species.

• G. c. citrina, the nominate subspecies breeds from northern India east along the Himalayas to eastern Bangladesh and possibly in western and northern Burma. It winters further south in India, Sri Lanka and Bangladesh.

• G. c. cyanota is mainly resident in Peninsular India south to Kerala. It has a white throat and face sides, with two black stripes running downwards from below the eyes. The spelling emendation cyanota is suggested by Rasmussen and Anderton.

• G. c. amadoni (not always recognized) found in northeastern part of peninsular India (Madhya Pradesh and Orissa) has brighter orange crown and longer wings than cyanota.

• G. c. innotata breeds through most of South-East Asia from southern Burma and southwestern China to northwest Thailand, central and southern Laos, Cambodia and southern Vietnam. It winter further south in southern Burma, and much of the rest of Thailand into Malaysia. It is very similar to the nominate but the male is brighter or deeper orange and lacks white tips to the median coverts; the female is duller on head and underparts, with an olive tinge to the grey of the mantle and back.

• G. c. melli breeds in southeastern China, and is partially migratory, regularly wintering in Hong Kong.

• G. c. courtoisi breeds in eastern-central China; its wintering range is unknown.

• G. c. aurimacula breeds in southern Vietnam, Hainan and possibly northern Laos. It resembles G. c. cyanota, but with a less defined head pattern. The face and neck-sides are whitish but flecked with orange or brownish and with weaker face stripes. The orange breast and flanks become paler orange on the belly and lower flanks.

• G. c. andamensis is resident in the Andaman Islands.

• G. c. albogularis is resident in the Nicobar Islands.

• G. c. gibsonhilli breeds from southern Burma to southern Thailand, and winters further south at lower levels in Peninsular Thailand, on islands in the Gulf of Thailand, and into Malaysia. It is similar to the nominate subspecies, but averages slightly brighter or deeper orange on head and upperparts and also has a slightly longer, heavier bill, and white tips to the median coverts.

• G. c. aurata is resident in the mountains of northern Borneo.

• G. c. rubecula is resident in Western Java.

• G. c. orientis is resident in Eastern Java and Bali and intergrades with G. c. rubecula in the west of its range. The separation of this form from the western Javan subspecies has been questioned.

Measurements

The following table summarises selected physical measurements for those subspecies for which data is available.

Distribution and habitat

The orange-headed thrush breeds in much of the Indian Subcontinent, including Bangladesh, India and Sri Lanka, and through Southeast Asia to Java and southern China. Its habitat is moist broadleaved evergreen woodlands, with a medium-density undergrowth of bushes and ferns, but it also utilises bamboo forests for secondary growth. G. c. cyanota also occurs in large gardens and orchards.

This species is often found in damp areas, near streams or in shady ravines. It occurs between 250 and 1830 metres (825– 6040 ft) in the Himalayas and up to about 1500 metres (5000 ft) in Malaysia, Thailand and Java. G. c. aurata is resident between 1000 and 1630 metres (3300–5400 ft) on Mt Kinabalu and Mt Trus Madi, northern Borneo. Some of the subspecies are completely or partially migratory; their wintering habitat is similar to the breeding forests, but more likely to be at lower altitudes.

Description

The orange-headed thrush is 205–235 milliimetres (8.1–9.25 in) long and weighs 47–60 grammes (1.7–2.1 oz). The adult male of the nominate subspecies of this small thrush has an entirely orange head and underparts, uniformly grey upperparts and wings, and white median and undertail coverts. It has a slate-coloured bill and the legs and feet have brown fronts and pink or yellowish rears.

The female resembles the male but has browner or more olive upperparts and warm brown wings, but some old females are almost identical to the male. The juvenile is dull brown with buff streaks on its back, and a rufous tone to the head and face; it has grey wings. The bill is brownish horn, and the legs and feet are brown.

This species' orange and grey plumage is very distinctive, and it is unlikely to be confused with any other species. Differences between the subspecies, as described above, can be quite striking, as with the strong head pattern on G. c. cyanota, but may be less obvious variations in plumage tone, or whether there is white on the folded wing. As with many other thrushes, all forms of this species shows a distinctive underwing pattern, with a strong white band.

Voice

Calls of the orange-headed thrush include a soft chuk or tchuk, a screeching teer-teer-teer, and a thin tsee or dzef given in flight. However, this bird is generally silent especially in winter. The song is a loud clear series of variably sweet lilting musical notes, recalling the quality of the common blackbird, but with the more repetitive structure of the song thrush. It also includes imitations of other birds like bulbuls, babblers and common tailorbird. It sings from a perch in a leafy tree, mostly early morning and late afternoon.

Behaviour

The orange-headed thrush is a shy, secretive bird usually occurring alone or in pairs, but is comparatively more easily seen than many other Zoothera thrushes, and several birds may congregate outside the breeding season at a good food source. It has a swift, silent flight, but when disturbed will often sit motionless until the threat has passed.

Breeding

The nest, built by both sexes, is a wide but shallow cup of twigs, bracken and rootlets lined with softer plant material like leaves, moss and conifer needles. It is constructed at a height of up to 4.5 metres (15 ft) in a small tree or bush, with mango trees and coffee bushes being preferred. Three or four, occasionally five, eggs are laid; they are cream or tinted with pale blue, grey or green, and have pale lilac blotches and reddish brown spots. They are incubated for 13–14 days to hatching, with another 12 days until the young birds leave the nest.

This species is a host of the pied cuckoo, Clamator jacobinus, a brood parasite which lay a single egg in the nest. Unlike the common cuckoo, neither the hen nor the hatched chick evict the host's eggs, but the host's young often die because they cannot compete successfully with the cuckoo for food. The chestnut-winged cuckoo, Clamator coromandus, and, very rarely, the common cuckoo, Cuculus canorus have also been claimed as parasites on this species.

Feeding

The orange-headed thrush feeds on the ground in dense undergrowth or other thick cover. It is most active at dawn and dusk, probing the leaf litter for insects and their larvae, spiders, other invertebrates and fruit. In Malaysia, wintering birds regularly feed on figs.

Status

The orange-headed thrush has an extensive range, estimated at 2,780,000 square kilometres (1,070,000 sq mi). The population size has not been quantified, but it is believed to be large due to its extensive range; it also reported as being locally common. The species is not believed to approach the thresholds for the global population decline criterion of the IUCN Red List (i.e., declining more than 30% in ten years or three generations), and is therefore evaluated as Least Concern.

It is very popular as cage-bird on Java, and numbers have severely declined in recent years owing to trapping for aviculture. Against the trend in Southeast Asia where loss or fragmentation of woodland poses a threat to forest birds, the orange-headed thrush has colonized Hong Kong, where it was first recorded in 1956, thanks to forest maturation.

Gallery

The ring ouzel is omnivorous, eating invertebrates, particularly insects and earthworms, some small vertebrates, and a wide range of fruit. Most animal prey is caught on the ground. During spring migration and the breeding season, invertebrates dominate the adult's diet and are also fed to the chicks. Later in the year, fruit becomes more important, particularly the common juniper.

With an extensive range and a large population, the ring ouzel is evaluated as least concern by the International Union for Conservation of Nature (IUCN). There are signs of decline in several countries; suspected causes including climate change, human disturbance, hunting and outdoor leisure activities. Loss of junipers may also be a factor in some areas. Natural hazards include predation by mammalian carnivores and birds of prey, and locally there may also be competition from other large thrushes such as the common blackbird, mistle thrush and fieldfare.

tymology

"Ouzel" is an old name for the common blackbird, the word being cognate with the German Amsel.  "Ouzel" may also be applied to a group of superficially similar but more distantly related birds, the dippers, the European representative of which is sometimes known as the water ouzel. "Ring Ouzel" was first used by John Ray in his 1674 Collection of English Words not Generally Used and became established with his 1678 book The Ornithology of Francis Willughby of Middleton in the County of Warwick. As with the English term, the scientific name also refers to the male's prominent white neck crescent, being derived from the Latin words turdus, "thrush", and torquatus, "collared". Old and local names for the ring ouzel include "fell blackbird", "hill blackbird", "moor blackbird", "rock ouzel" and "mountain blackbird".

Taxonomy

Male T. t. alpestris in Serbia

The ring ouzel was first described by Carl Linnaeus under its current scientific name in his 1758 10th edition of Systema Naturae. He noted earlier descriptions by Francis Willughby and Eleazar Albin, both of whom gave it the name Merula torquata.

There are about 85 species of medium to large thrushes in the genus Turdus. They are characterised by rounded heads, medium or longish pointed wings, and usually melodious songs.

A 2020 study of the genetics of Turdus suggested that the genus arose about 9.37 million years ago (Mya), expanding out of Africa around 7.2 Mya, and diverging into Palearctic and Oriental groups about 5.7 Mya. Further radiation from Africa to the Americas followed at about 5.3 Mya. Details of the study suggest that the ring ouzel, a member of the Eurasian group, may be more closely related to Naumann's and dusky thrushes than to the superficially more similar common blackbird.

Subspecies

Male T. t. amicorum in Artashavan, Armenia

The ring ouzel has three recognised subspecies: The northern ring ouzel, Turdus torquatus torquatus Linnaeus (1758) is the nominate subspecies (the subspecies that repeats the name of the species). It breeds across western and northern Europe from Ireland through Scandinavia to northwest Russia and winters in southern Europe and northwest Africa. The Alpine ring ouzel, T. t. alpestris Brehm, C L (1831) breeds in mountain ranges from Iberia through southern and central Europe to the Balkans, Greece and western Turkey, and also in North Africa. It winters in North Africa, southern Europe and southern Turkey. The Caucasian ring ouzel, T. t. amicorum Hartert E (1923) breeds in central and eastern Turkey east to Turkmenistan, and winters mainly in Iran and parts of Iraq. Analysis of mitochondrial DNA samples from across Europe suggests that this species had a much broader distribution after the Last Glacial Period that ended about 11,700 years ago than it does now.

Description

The ring ouzel is 23–24 centimetres (9.1–9.4 in) in length and weighs 90–138 grams (3.2–4.9 oz). The plumage of the male of the nominate race is entirely black except for a conspicuous white crescent on the breast, narrow greyish scaling on the upperparts and belly and pale edges to the wing feathers. The bill is yellow and the legs are greyish brown. The female resembles the male but is browner and with a duller breast band. Juveniles are like the female, but with a faint or non-existent breast crescent.

The pale breast marking makes adults of this species unmistakeable; first-winter males also sometimes show a pale crescent. Other young ouzels can be confused with the common blackbird, but always show a paler wing panel than that species.

Males of T. t. alpestris have broader white scalloping (repeated small curves) on their underparts than T. t. torquatus, giving a distinctly scaly appearance below. The wing panel is also paler than in the nominate subspecies. Females are much as the nominate race, but with broad white fringes on the chin and throat. Males of T. t. amicorum have the largest and whitest breast band of the three subspecies, and the broader white edges and tips of the wing feathers form a distinctive whitish panel in the wing. Females have narrow white fringes on their underparts. Adult ring ouzels undergo a complete moult after breeding from late June to early September, before their autumn migration. Juveniles have a partial moult between July and September, replacing their head, body and wing covert feathers.

Voice

The male ring ouzel sings from a low perch or occasionally in flight. The song consists of a repetition of 2–4 plaintive fluty notes, tri-ríí, tri-ríí, ti-ríí with pauses between repeats. The call is a loud tac-tac-tac, becoming harsher if the bird is alarmed. The contact call is a soft cherrr in flight. Males sing most frequently at dawn and sunset.

Distribution and habitat

The ring ouzel breeds discontinuously across western and northern Europe from north-west Ireland through Scandinavia to northwest Russia, and in mountains across central southern Europe from the Pyrenees through the Alps, the Balkans, Greece and Turkey east to Turkmenistan. In 2014, breeding was recorded on the Timan Ridge, Arkhangelsk Oblast, about 300 kilometres (190 mi) further east than previously known breeding sites in north Russia.

The species is migratory, birds leaving the breeding areas in September and October. Birds of the nominate subspecies winter in southern Spain and northwest Africa. Central European populations of T. t. alpestris move to higher elevations initially before moving south or southwest through the Swiss Alps; some two weeks later migrants of the nominate form pass through the same area to winter in the south of the breeding range or around the Mediterranean. Eastern alpestris ouzels migrate through the Balkans and Turkey. T. t. amoricus moves south to Egypt and neighbouring areas. The return migration is mainly in March and April, the males arriving some days before the females. Northern breeders arrive later, and in the mountains, some birds may ascend in stages as the snow melts. Many birds stop off at traditional well-grazed grassland locations in both spring and autumn.

The ring ouzel is extinct in Latvia and occurs only on migration in Denmark. It is a passage migrant in Syria and a vagrant to Iceland, Jordan, the Arabian Peninsula, Sudan, Kazakhstan, Mauritania, Svalbard and Jan Mayen. In the Atlantic, it is a regular winter visitor to the Canary Islands but a rarity in the Azores and Madeira.

In middle latitudes, the ring ouzel is a bird of continental mountains, but in the north of its range, it is found in coastal uplands. It can cope with wind and rain but avoids ice and snow. Nominate T. t. torquatus is usually found on open moorland with a few stunted trees above 250 metres (820 ft), and reaching 1,200 metres (3,900 ft) in Scotland and northern Europe. In Switzerland, ring ouzels breed on rugged upland slopes with heather, conifers, beech or hairy alpenrose at 1,100–1,300 metres (3,600–4,300 ft), although in Turkey birds are found from sea level to 1,500 metres (4,900 ft). In Armenia and the Caucasus, it occupies similar steep habitat with conifer stands, rhododendron thickets, and juniper scrub and shrub, from sea level to 2,000–3,000 metres (6,600–9,800 ft).

In northwest Africa ring ouzels winter in juniper forest at 1,800–2,200 metres (5,900–7,200 ft), often near rivers or ponds. On migration, ouzels may occur on coastal grassland and steep hillsides with short, unsown wild grass and sparse scrub.

Behaviour

The ring ouzel is territorial and normally seen alone or in pairs, although loose flocks may form on migration. When not breeding, several birds may be loosely associated in good feeding areas, such as a fruiting tree, often with other thrushes such as song thrushes or redwings. The ouzel's flight is direct, and birds often perch on rocks or heather clumps.

Breeding

Ring ouzels nest from mid-April to mid-July in the Alps and the British Isles, and from May to August in Scandinavia. Territories may be strung out along streams, 160–200 metres (520–660 ft) apart and the ranges may overlap, but this species does not form breeding colonies. The nest, built by the female, is a cup of leaves, dry grass and other plant material consolidated with mud. In the west of the range nests are almost always built on the ground, but T. t. alpestris may also nest in a small tree or scrub at an average height of 3.5 metres (11 ft).

The clutch is 3–6 pale blue or greenish-blue eggs flecked with reddish-brown. The eggs are 30 mm × 22 mm (1.18 in × 0.87 in) in size and weigh 7.4 grams (0.26 oz) of which 6% is shell. Incubation is almost always by the female, hatching typically occurring after 13 days. The altricial, downy chicks fledge in another 14 days. The young are dependent on their parents for about 12 days after fledging.

Adults breed after their first year and their average lifespan is two years, although nine years has been recorded. There may be two broods, especially in the south of the range, although triple-brooding is rare. This species is philopatric, returning to the same area to breed each year. Around 36% of juveniles survive their first year, while the annual survival rate for adults is 47% for males and 37% for females. The main causes of death in northwest Europe are predation (9%), accidental human-related incidents (10%), and hunting, mainly in France (77%).

Diet

The ring ouzel is omnivorous, eating a wide range of insects, earthworms, small amphibians and reptiles and fruit. Most animal prey is caught on the ground.

During spring migration and the breeding season, invertebrates dominate the diet, and include earthworms, beetles flies, ants, spiders and snails. Later in the year, fruit becomes more important, including bramble, strawberry, cherry, hawthorn, rowan and juniper. Where it is available common juniper makes up more than 90% of the ring ouzel's winter diet, with arthropods constituting most of the rest. As a result, the ring ouzel is an important vector for dispersing the juniper's seeds, and is key to the dispersal of the endemic Canary Islands juniper.

The young are mainly fed invertebrates, caterpillars and earthworms being major items where available. Although birds migrating in autumn use similar habitat to that used in spring, seasonal berries make up most of their diet, particularly elderberries, haws and, where available, juniper berries.

Predators and parasites

Predators of the ring ouzel include the tawny owl, long-eared owl, common buzzard, common kestrel and Eurasian sparrowhawk, least weasel and stoat. Most deaths are of young juveniles, and birds hatched early in the season are more likely to survive than later broods. A Scottish study showed that raptors were responsible for 59% of deaths and mammals for 27%. In Romania, eggs were taken by red squirrels and spotted nutcrackers. As with other Turdus thrushes, the ring ouzel is rarely a host of the common cuckoo, a brood parasite. If the thrush's nest cup is too deep for the cuckoo to evict the host's chicks, the young cuckoo cannot successfully compete for food with the fast-growing host species' chicks, and if the cuckoo does manage to expel its nest-mates, the parents are reluctant to feed it; either way, the young cuckoo will starve.

A study in the Carpathian Mountains found that a significant proportion of ring ouzels carried trombiculid mites, commonly known as chiggers. These mites commonly infect ground-feeding birds, and heavy infestations can cause birds to lose condition and stop feeding. The hard-bodied tick Ixodes festai commonly parasitises thrushes, including the ring ouzel. There is a record of this species carrying a *Haemoproteus *blood parasite.

Status and conservation

The ring ouzel has an extensive range, estimated at 9.17 million square kilometres (3.54 million sq mi), and a large population, estimated at 600,000–2 million individuals in Europe (which comprises 95% of the breeding range). The species is not believed to approach the thresholds for the population decline criteria of the IUCN Red List (i.e., declining more than 30% in ten years or three generations), and is therefore evaluated as least concern. The breeding population in Europe was estimated to be 299,000–598,000 pairs in 2019.

There are signs of decline in several countries. Its decline in Ireland in recent years has been striking, with regular breeding now confined to two counties. Suspected causes include climate change, human disturbance, hunting and outdoor leisure activities. Loss of junipers may be a factor in southern Spain and north-west Africa, as may upland forestation in the UK. There may also be competition from larger thrushes like the common blackbird, mistle thrush and fieldfare. A Scottish study suggested that sites at higher altitudes and with a good cover of heather were less likely to have been deserted by breeding ring ouzels than lower or more open locations.

In the Alps, the density of breeding pairs can reach 60–80 per square kilometre (160–210/sq mi) but is generally much lower with 37 per square kilometre (96/sq mi) in Haute-Savoie, 22 per square kilometre (57/sq mi) in the Jura Mountains, and 8 per square kilometre (21/sq mi) in more open habitats in Britain.