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Wikipedia

Woodpecker

Woodpeckers are part of the bird family Picidae, which also includes the piculets, wrynecks, and sapsuckers.[1] Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar, and the extreme polar regions. Most species live in forests or woodland habitats, although a few species are known that live in treeless areas, such as rocky hillsides and deserts, and the Gila woodpecker specialises in exploiting cacti.

Woodpecker
Temporal range: 26–0 Ma Late Oligocene to present
Pileated woodpecker

Tapping sound of a woodpecker 

Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Piciformes
Infraorder: Picides
Family: Picidae
Leach, 1820
Subfamilies

Members of this family are chiefly known for their characteristic behaviour. They mostly forage for insect prey on the trunks and branches of trees, and often communicate by drumming with their beaks, producing a reverberatory sound that can be heard at some distance. Some species vary their diet with fruits, birds' eggs, small animals, tree sap, human scraps, and carrion. They usually nest and roost in holes that they excavate in tree trunks, and their abandoned holes are of importance to other cavity-nesting birds. They sometimes come into conflict with humans when they make holes in buildings or feed on fruit crops, but perform a useful service by their removal of insect pests on trees.

The Picidae are one of nine living families in the order Piciformes, the others being barbets (comprising three families), toucans, toucan-barbets, and honeyguides, which (along with woodpeckers) comprise the clade Pici, and the jacamars and puffbirds in the clade Galbuli. DNA sequencing has confirmed the sister relationships of these two groups. The family Picidae includes about 240 species arranged in 35 genera. Almost 20 species are threatened with extinction due to loss of habitat or habitat fragmentation, with one, the Bermuda flicker, being extinct and a further two possibly being so.

General characteristics

 
A black-rumped flameback using its tail for support

Woodpeckers include the tiny piculets, the smallest of which appears to be the bar-breasted piculet at 7.5 cm (3.0 in) in length and a weight of 8.9 g (0.31 oz).[2][3] Some of the largest woodpeckers can be more than 50 cm (20 in) in length. The largest surviving species is the great slaty woodpecker, which weighs 430 g (15 oz) on average and up to 563 g (19.9 oz), and measures 45 to 55 cm (18 to 22 in), but the extinct imperial woodpecker, at 55 to 61 cm (22 to 24 in), and ivory-billed woodpecker, around 48 to 53 cm (19 to 21 in) and 516 g (18.2 oz), were probably both larger.[4][3][5][6][7]

The plumage of woodpeckers varies from drab to conspicuous. The colours of many species are based on olive and brown and some are pied, suggesting a need for camouflage; others are boldly patterned in black, white, and red, and many have a crest or tufted feathers on their crowns. Woodpeckers tend to be sexually dimorphic, but differences between the sexes are generally small; exceptions to this are Williamson's sapsucker and the orange-backed woodpecker, which differ markedly. The plumage is moulted fully once a year apart from the wrynecks, which have an additional partial moult before breeding.[8]

Woodpeckers, piculets, and wrynecks all possess characteristic zygodactyl feet, consisting of four toes, the first (hallux) and the fourth facing backward and the second and third facing forward. This foot arrangement is good for grasping the limbs and trunks of trees. Members of this family can walk vertically up tree trunks, which is beneficial for activities such as foraging for food or nest excavation. In addition to their strong claws and feet, woodpeckers have short, strong legs. This is typical of birds that regularly forage on trunks. Exceptions are the black-backed woodpecker and the American and Eurasian three-toed woodpeckers, which have only three toes on each foot. The tails of all woodpeckers, except the piculets and wrynecks, are stiffened, and when the bird perches on a vertical surface, the tail and feet work together to support it.[4]

Woodpeckers have strong bills that they use for drilling and drumming on trees, and long, sticky tongues for extracting food (insects and larvae).[4] Woodpecker bills are typically longer, sharper, and stronger than the bills of piculets and wrynecks, but their morphology is very similar. The bill's chisel-like tip is kept sharp by the pecking action in birds that regularly use it on wood. The beak consists of three layers; an outer sheath called rhamphotheca, made of scales formed from keratin proteins, an inner layer of bone which has a large cavity and mineralised collagen fibers, and a middle layer made of porous bone which connects the two other layers. Furthermore, the tongue bone (or hyoid bone) of the woodpecker is very long, and winds around the skull through a special cavity, thereby cushioning the brain.[9] Combined, this anatomy helps the beak absorb mechanical stress.[10] Species of woodpecker and flicker that use their bills in soil or for probing as opposed to regular hammering tend to have longer and more decurved bills. Due to their smaller bill size, many piculets and wrynecks forage in decaying wood more often than woodpeckers. Their long, sticky tongues, which possess bristles, aid these birds in grabbing and extracting insects from deep within a hole in a tree. The tongue was reported to be used to spear grubs, but more detailed studies published in 2004 have shown that the tongue instead wraps around the prey before being pulled out.[11]

 
Diagram showing the hyoid bone of Dendrocopos major

Many of the foraging, breeding, and signaling behaviors of woodpeckers involve drumming and hammering using their bills.[12] To prevent brain damage from the rapid and repeated powerful impacts, woodpeckers have a number of physical features that protect their brains.[13] These include a relatively small and smooth brain, narrow subdural space, little cerebrospinal fluid surrounding it to prevent it from moving back and forth inside the skull during pecking, the orientation of the brain within the skull (which maximises the contact area between the brain and the skull) and the short duration of contact. The skull consists of strong but compressible, sponge-like bone, which is most concentrated in the forehead and the back of the skull.[13] Another anatomical adaptation of woodpeckers is the enormously elongated hyoid bone which subdivides, passes on either side of the spinal column and wraps around the brain case, before ending in the right nostril cavity. It plays the role of safety-belt.[9]

Computer simulations have shown that 99.7% of the energy generated in pecking is stored in the form of strain energy, which is distributed throughout the bird's body, with only a small remaining fraction of the energy going into the brain. The pecking also causes the woodpecker's skull to heat up, which is part of the reason why they often peck in short bursts with brief breaks in between, giving the head some time to cool.[14] During the millisecond before contact with wood, a thickened nictitating membrane closes, protecting the eye from flying debris.[15] These membranes also prevent the retina from tearing. Their nostrils are also protected; they are often slit-like and have special feathers to cover them. Woodpeckers are capable of repeated pecking on a tree at high decelerations on the order of 10,000 m/s2 (33,000 ft/s2) (1000 g).[16]

Some large woodpeckers such as Dryocopus have a fast, direct form of flight, but the majority of species have a typical undulating flight pattern consisting of a series of rapid flaps followed by a swooping glide. Many birds in the genus Melanerpes have distinctive, rowing wing-strokes while the piculets engage in short bursts of rapid direct flight.[17]

Distribution, habitat, and movements

 
Use of cacti for breeding and roosting holes allows some woodpeckers to live in treeless deserts, such as the ladder-backed woodpecker, which uses cacti for nesting.

Global distribution

Woodpeckers have a mostly cosmopolitan distribution, although they are absent from Australasia, Madagascar, and Antarctica. They are also absent from some of the world's oceanic islands, although many insular species are found on continental islands. The true woodpeckers, subfamily Picinae, are distributed across the entire range of the family. The Picumninae piculets have a pantropical distribution, with species in Southeast Asia, Africa, and the Neotropics, with the greatest diversity being in South America.[18] The second piculet subfamily, the Sasiinae, contains the African piculet and two species in the genus Sasia that are found in Southeast Asia.[19] The wrynecks (Jynginae) are found exclusively in the Old World, with the two species occurring in Europe, Asia, and Africa.[18]

Most woodpeckers are sedentary, but a few examples of migratory species are known, such as the rufous-bellied woodpecker, yellow-bellied sapsucker,[18] and Eurasian wryneck, which breeds in Europe and west Asia and migrates to the Sahel in Africa in the winter.[20] More northerly populations of Lewis's woodpecker, northern flicker, Williamson's sapsucker, red-breasted sapsucker, and red-naped sapsucker all move southwards in the fall in North America.[18] Most woodpecker movements can be described as dispersive, such as when young birds seek territories after fledging, or eruptive, to escape harsh weather conditions. Several species are altitudinal migrants, for example the grey-capped pygmy woodpecker, which moves to lowlands from hills during winter. The woodpeckers that do migrate, do so during the day.[4]

Habitat requirements

Overall, woodpeckers are arboreal birds of wooded habitats. They reach their greatest diversity in tropical rainforests, but occur in almost all suitable habitats, including woodlands, savannahs, scrublands, and bamboo forests. Even grasslands and deserts have been colonised by various species. These habitats are more easily occupied where a small number of trees exist, or in the case of desert species like the Gila woodpecker, tall cacti are available for nesting.[21] Some are specialists and are associated with coniferous or deciduous woodlands, or even, like the acorn woodpecker, with individual tree genera (oaks in this case). Other species are generalists and are able to adapt to forest clearance by exploiting secondary growth, plantations, orchards, and parks. In general, forest-dwelling species need rotting or dead wood on which to forage.[22]

Several species are adapted to spending a portion of their time feeding on the ground, and a very small minority have abandoned trees entirely and nest in holes in the ground. The ground woodpecker is one such species, inhabiting the rocky and grassy hills of South Africa,[23] and the Andean flicker is another.[22]

The Swiss Ornithological Institute has set up a monitoring program to record breeding populations of woodland birds. This has shown that deadwood is an important habitat requirement for the black woodpecker, great spotted woodpecker, middle spotted woodpecker, lesser spotted woodpecker, European green woodpecker, and Eurasian three-toed woodpecker. Populations of all these species increased by varying amounts from 1990 to 2008. During this period, the amount of deadwood in the forest increased and the range of the white-backed woodpecker enlarged as it extended eastwards. With the exception of the green and middle-spotted woodpeckers, the increase in the amount of deadwood is likely to be the major factor explaining the population increase of these species.[24]

Behaviour

Most woodpeckers live solitary lives, but their behaviour ranges from highly antisocial species that are aggressive towards their own kind, to species that live in groups. Solitary species defend such feeding resources as a termite colony or fruit-laden tree, driving away other conspecifics and returning frequently until the resource is exhausted. Aggressive behaviours include bill pointing and jabbing, head shaking, wing flicking, chasing, drumming, and vocalisations. Ritual actions do not usually result in contact, and birds may "freeze" for a while before they resume their dispute. The coloured patches may be flouted, and in some instances, these antagonistic behaviours resemble courtship rituals.[25]

Group-living species tend to be communal group breeders.[25] In addition to these species, a number of species may join mixed-species foraging flocks with other insectivorous birds, although they tend to stay at the edges of these groups. Joining these flocks allows woodpeckers to decrease their anti-predator vigilance and increase their feeding rate.[26] Woodpeckers are diurnal, roosting at night inside holes and crevices. In many species the roost will become the nest-site during the breeding season, but in some species they have separate functions; the grey-and-buff woodpecker makes several shallow holes for roosting which are quite distinct from its nesting site. Most birds roost alone and will oust intruders from their chosen site, but the Magellanic woodpecker and acorn woodpecker are cooperative roosters.[25]

Drumming

Drumming is a form of nonvocal communication used by most species of woodpeckers, and involves the bill being repeatedly struck on a hard surface with great rapidity. After a pause, the drum roll is repeated, with each species having a pattern that is unique in the number of beats in the roll, the length of the roll, the length of the gap between rolls, and the cadence.[27][28] The drumming is mainly a territorial call, equivalent to the song of a passerine.[29] Woodpeckers choose a surface that resonates, such as a hollow tree, and may use man-made structures such as gutters and downpipes.[30] Drumming serves for the mutual recognition of conspecifics and plays a part in courtship rituals. Individual birds are thought to be able to distinguish the drumming of their mates and those of their neighbours.[31] Drumming in woodpeckers is controlled by a set of nuclei in the forebrain that closely resemble the brain regions that underlie song learning and production in many songbirds.[32]

Calls

Woodpeckers do not have such a wide range of songs and calls as do passerine birds, and the sounds they make tend to be simpler in structure. Calls produced include brief, high-pitched notes, trills, rattles, twittering, whistling, chattering, nasal churrs, screams, and wails. These calls are used by both sexes in communication and are related to the circumstances of the occasion; these include courtship, territorial disputes, and alarm calls. Each species has its own range of calls, which tend to be in the 1.0 to 2.5 kHz range for efficient transmission through forested environments. Mated couples may exchange muted, low-pitched calls, and nestlings often issue noisy begging calls from inside their nest cavity.[29] The wrynecks have a more musical song, and in some areas, the song of the newly arrived Eurasian wryneck is considered to be the harbinger of spring.[33] The piculets either have a song consisting of a long, descending trill, or a descending series of two to six (sometimes more) individual notes, and this song alerts ornithologists to the presence of the birds, as they are easily overlooked.[34]

Diet and feeding

 
Holes bored by feeding woodpeckers

Most woodpecker species feed on insects and other invertebrates living under bark and in wood, but overall, the family is characterized by its dietary flexibility, with many species being both highly omnivorous and opportunistic. The diet includes ants, termites, beetles and their larvae, caterpillars, spiders, other arthropods, bird eggs, nestlings, small rodents, lizards, fruit, nuts, and sap. Many insects and their grubs are taken from living and dead trees by excavation. The bird may hear sounds from inside the timber indicating where creating a hole would be productive.[25] Crustaceans, molluscs, and carrion may be eaten by some species, including the great spotted woodpecker, and bird feeders are visited for suet and domestic scraps.[35]

Other means are also used to garner prey. Some species, such as the red-naped sapsucker, sally into the air to catch flying insects, and many species probe into crevices and under bark, or glean prey from leaves and twigs. The rufous woodpecker specialises in attacking the nests of arboreal ants, and the buff-spotted woodpecker feeds on and nests in termite mounds. Other species, such as the wrynecks and the Andean flicker, feed wholly or partly on the ground.[25]

Ecologically, woodpeckers help to keep trees healthy by keeping them from suffering mass infestations. The family is noted for its ability to acquire wood-boring grubs from the trunks and branches, whether the timber is alive or dead. Having hammered a hole into the wood, the prey is extracted by use of a long, barbed tongue. Woodpeckers consume beetles that burrow into trees, removing as many as 85% of emerald ash borer larvae from individual ash trees.[36]

The ability to excavate allows woodpeckers to obtain tree sap, an important source of food for some species. Most famously, the sapsuckers (genus Sphyrapicus) feed in this fashion, but the technique is not restricted to these, and others such as the acorn woodpecker and white-headed woodpecker also feed on sap. The technique was once thought to be restricted to the New World, but Old World species, such as the Arabian woodpecker and great spotted woodpecker, also feed in this way.[4]

Breeding

 
A male black woodpecker attending its chicks

All members of the family Picidae nest in cavities, nearly always in the trunks and branches of trees, well away from the foliage. Where possible, an area of rotten wood surrounded by sound timber is used. Where trees are in short supply, the gilded flicker and ladder-backed woodpecker excavate holes in cactus, and the Andean flicker and ground woodpecker dig holes in earth banks. The campo flicker sometimes chooses termite mounds, the rufous woodpecker prefers to use ants' nests in trees and the bamboo woodpecker specialises in bamboos.[37] Woodpeckers also excavate nest holes in residential and commercial structures and wooden utility poles.[36]

Woodpeckers and piculets excavate their own nests, but wrynecks do not, and need to find pre-existing cavities. A typical nest has a round entrance hole that just fits the bird, leading to an enlarged vertical chamber below. No nesting material is used, apart from some wood chips produced during the excavation; other wood chips are liberally scattered on the ground, thus providing visual evidence of the site of the nest.[38] Many species of woodpeckers excavate one hole per breeding season, sometimes after multiple attempts. It takes around a month to finish the job and abandoned holes are used by other birds and mammals that are cavity nesters unable to excavate their own holes.[39]

Cavities are in great demand for nesting by other cavity nesters, so woodpeckers face competition for the nesting sites they excavate from the moment the hole becomes usable. This may come from other species of woodpecker, or other cavity-nesting birds such as swallows and starlings. Woodpeckers may aggressively harass potential competitors, and also use other strategies to reduce the chance of being usurped from their nesting sites; for example, the red-crowned woodpecker digs its nest in the underside of a small branch, which reduces the chance that a larger species will take it over and expand it.[40]

Members of Picidae are typically monogamous, with a few species breeding cooperatively and some polygamy reported in a few others.[41] Polyandry, where a female raises two broods with two separate males, has also been reported in the West Indian woodpecker.[42] Another unusual social system is that of the acorn woodpecker, which is a polygynandrous cooperative breeder where groups of up to 12 individuals breed and help to raise the young.[4] Young birds from previous years may stay behind to help raise the group's young, and studies have found reproductive success for the group goes up with group size, but individual success goes down. Birds may be forced to remain in groups due to a lack of habitat to which to disperse.[43]

 
Woodpecker[which?] feeding its chick

A pair works together to help build the nest, incubate the eggs, and raise their altricial young. In most species, though, the male does most of the nest excavation and takes the night shift while incubating the eggs. A clutch usually consists of two to five round, white eggs. Since these birds are cavity nesters, their eggs do not need to be camouflaged and the white color helps the parents to see them in dim light. The eggs are incubated for about 11–14 days before they hatch. About 18–30 days are then needed before the chicks are fully fledged and ready to leave the nest. In most species, soon after this, the young are left to fend for themselves, exceptions being the various social species, and the Hispaniolan woodpecker, where adults continue to feed their young for several months. In general, cavity nesting is a successful strategy and a higher proportion of young is reared than is the case with birds that nest in the open. In Africa, several species of honeyguide are brood parasites of woodpeckers.[38]

Systematics and evolutionary history

 
Lesser goldenback (Dinopium benghalense) in Guwahati, India

The Picidae are just one of nine living families in the order Piciformes. Other members of this group, such as the jacamars, puffbirds, barbets, toucans, and honeyguides, have traditionally been thought to be closely related to the woodpecker family (true woodpeckers, piculets, wrynecks, and sapsuckers). The clade Pici (woodpeckers, barbets, toucans, and honeyguides) is well supported and shares a zygodactyl foot with the Galbuli (puffbirds and jacamars). More recently, several DNA sequence analyses have confirmed that Pici and Galbuli are sister groups.[44]

The name Picidae for the family was introduced by English zoologist William Elford Leach in a guide to the contents of the British Museum published in 1820.[45][46] The phylogeny has been updated according to new knowledge about convergence patterns and evolutionary history.[47][48] Most notably, the relationship of the Picinae genera has been largely clarified, and the Antillean piculet was found to be a surviving offshoot of protowoodpeckers. Genetic analysis supports the monophyly of the Picidae, which seem to have originated in the Old World, but the geographic origins of the Picinae is unclear. The Picumninae are returned as paraphyletic.[47] Morphological and behavioural characters, in addition to DNA evidence, highlights genus Hemicircus as the sister group of all remaining true woodpeckers, besides a sister-group relationship between the true woodpecker tribes Dendropicini and Malarpicini.[49]

The evolutionary history of this group is not well documented, but the known fossils allow some preliminary conclusions; the earliest known modern picids were piculet-like forms of the Late Oligocene, about 25 million years ago (Mya). By that time, however, the group was already present in the Americas and Europe, and they actually may have evolved much earlier, maybe as early as the Early Eocene (50 Mya). The modern subfamilies appear to be rather young by comparison; until the mid-Miocene (10–15 Mya), all picids seem to have been small or mid-sized birds similar to a mixture between a piculet and a wryneck. A feather enclosed in fossil amber from the Dominican Republic, dated to about 25 Mya, however, seems to indicate that the Nesoctitinae were already a distinct lineage by then.[50]

Stepwise adaptations for drilling, tapping, and climbing head first on vertical surfaces have been suggested.[49] The last common ancestor of woodpeckers (Picidae) was incapable of climbing up tree trunks or excavating nest cavities by drilling with its beak. The first adaptations for drilling (including reinforced rhamphotheca, frontal overhang, and processus dorsalis pterygoidei) evolved in the ancestral lineage of piculets and true woodpeckers. Additional adaptations for drilling and tapping (enlarged condylus lateralis of the quadrate and fused lower mandible) have evolved in the ancestral lineage of true woodpeckers (Hemicircus excepting). The inner rectrix pairs became stiffened, and the pygostyle lamina was enlarged in the ancestral lineage of true woodpeckers (Hemicircus included), which facilitated climbing head first up tree limbs. Genus Hemicircus excepting, the tail feathers were further transformed for specialized support, the pygostyle disc became greatly enlarged, and the ectropodactyl toe arrangement evolved. These latter characters may have facilitated enormous increases in body size in some lineages.[49]

Prehistoric representatives of the extant Picidae genera are treated in the genus articles. An enigmatic form based on a coracoid, found in Pliocene deposits of New Providence in the Bahamas, has been described as Bathoceleus hyphalus and probably also is a woodpecker.[51]

The following cladogram is based on the comprehensive molecular phylogenetic study of the woodpeckers published in 2017 together with the list of bird species maintained by Frank Gill, Pamela Rasmussen and David Donsker on behalf of the International Ornithological Committee (IOC). The Cuban green woodpecker in the monotypic genus Xiphidiopicus was not included in the study.[52][53] The relative positions of Picumninae, Sasiinae and Picinae in the cladogram are uncertain. In the 2017 study the results depended upon which of two different statistical procedures were used to analyse the DNA sequence data. One method found that Sasiinae was sister to Picinae (as shown below), the other method found that Sasiinae was sister to a clade containing both Picumninae and Picinae.[52]

Picidae
Jynginae

Jynx – 2 species (wrynecks)

Picumninae

Picumnus – 26 species (piculets)

Sasiinae

Verreauxia – African piculet

Sasia – 2 species (piculets)

Picinae
Nesoctitini

Nesoctites – Antillean piculet

Hemicirini

Hemicircus – 2 species

Picini

Micropternus – rufous woodpecker

Meiglyptes – 3 species

Gecinulus – 3 species

Dinopium – 5 species (flamebacks)

Picus – 14 species

Chrysophlegma – 3 species

Pardipicus – 2 species

Geocolaptes – ground woodpecker

Campethera – 11 species

Mulleripicus – 4 species

Dryocopus – 6 species

Celeus – 13 species

Piculus – 7 species

Colaptes – 14 species

Campephilini

Campephilus – 11 species

Blythipicus – 2 species

Reinwardtipicus – orange-backed woodpecker

Chrysocolaptes – 9 species

Melanerpini

Sphyrapicus – 4 species (sapsuckers)

Melanerpes – 24 species

Picoides – 3 species

Yungipicus – 7 species

Dendrocoptes – 4 species (including Leiopicus)

Chloropicus – 3 species

Dendropicos – 12 species

Dendrocopos – 12 species

Dryobates – 5 species

Leuconotopicus – 6 species

Veniliornis – 14 species


List of genera

 
Ochre-collared piculet
(Picumnus temminckii)
 
Red-crowned woodpecker
(Melanerpes rubricapillus rubricapillus)
female, Tobago

The woodpecker family Picidae contains 37 genera.[53] For more detail, see list of woodpecker species.

 
Cuban green woodpecker
(Xiphidiopicus percussus)
female, Cuba
 
Campo flicker
Colaptes campestris
female, Brazil

Family: Picidae

Relationship with humans

In general, humans consider woodpeckers in a favourable light; they are viewed as interesting birds and fascinating to watch as they drum or forage, but their activities are not universally appreciated.[55] Many woodpecker species are known to excavate holes in buildings, fencing, and utility poles, creating health and/or safety issues for affected structures. Such activity is very difficult to discourage and can be costly to repair.[56]

Woodpeckers also drum on various reverberatory structures on buildings such as gutters, downspouts, chimneys, vents, and aluminium sheeting.[57] Drumming is a less-forceful type of pecking that serves to establish territory and attract mates.[56] Houses with shingles or wooden boarding are also attractive as possible nesting or roosting sites, especially when close to large trees or woodland. Several exploratory holes may be made, especially at the junctions of vertical boards or at the corners of tongue-and-groove boarding. The birds may also drill holes in houses as they forage for insect larvae and pupae hidden behind the woodwork.[57]

Woodpeckers sometimes cause problems when they raid fruit crops, but their foraging activities are mostly beneficial as they control forest insect pests such as the woodboring beetles that create galleries behind the bark and can kill trees. They also eat ants, which may be tending sap-sucking pests such as mealybugs, as is the case with the rufous woodpecker in coffee plantations in India.[55] Woodpeckers can serve as indicator species, demonstrating the quality of the habitat. Their hole-making abilities make their presence in an area an important part of the ecosystem, because these cavities are used for breeding and roosting by many bird species that are unable to excavate their own holes, as well as being used by various mammals and invertebrates.[55]

The spongy bones of the woodpecker's skull and the flexibility of its beak, both of which provide protection for the brain when drumming, have provided inspiration to engineers; a black box needs to survive intact when a plane falls from the sky, and modelling the black box with regard to a woodpecker's anatomy has increased the resistance of this device to damage 60-fold.[58] The design of protective helmets is another field being influenced by the study of woodpeckers.[58]

One of the accounts of the founding of Rome, preserved in the work known as Origo Gentis Romanae, refers to a legend of a woodpecker bringing food to the boys Romulus and Remus during the time they were abandoned in the wild, thus enabling them to survive and play their part in history.

Status and conservation

 
The ivory-billed woodpecker is classified as critically endangered by the IUCN,[59] and some authorities believe it may already be extinct.

In a global survey of the risk of extinction faced by the various bird families, woodpeckers were the only bird family to have significantly fewer species at risk than would be expected.[60] Nevertheless, several woodpeckers are under threat as their habitats are destroyed. Being woodland birds, deforestation and clearance of land for agriculture and other purposes can reduce populations dramatically. Some species adapt to living in plantations and secondary growth, or to open countryside with forest remnants and scattered trees, but some do not. A few species have even flourished when they have adapted to man-made habitats. There are few conservation projects directed primarily at woodpeckers, but they benefit whenever their habitat is conserved.[55] The red-cockaded woodpecker has been the focus of much conservation effort in the southeastern United States, with artificial cavities being constructed in the longleaf pines they favour as nesting sites.[61]

Two species of woodpeckers in the Americas, the ivory-billed woodpecker is critically endangered and the imperial woodpecker is classified as extinct in the wild, with some authorities believing them extinct, though possible but disputed ongoing sightings of ivory-billed woodpeckers have been made in the United States[62] and a small population may survive in Cuba.[59] A critically endangered species is the Okinawa woodpecker from Japan, with a single declining population of a few hundred birds. It is threatened by deforestation, golf course, dam, and helipad construction, road building, and agricultural development.[63]

Brain impact research

Anatomy

Woodpeckers possess many sophisticated shock-absorption mechanisms that help protect them from head injury. Micro-CT scans show that plate-like spongy bones are in the skull with an uneven distribution, highly accumulated in the forehead and occiput but not in other regions.[64]  Along with the long hyoid bone “safety belt” the woodpecker has uneven beak lengths which drastically reduce strains when compared to equal length.[64][65] Models have shown that pecking force is changed to strain energy and stored into the body at around 99% absorption while 1% is in the head. The head also has many factors that reduce strain to the brain and small portions of energy are dissipated into the form of heat, therefore the pecks are always intermittent.[66]

Tau protein accumulation is associated with chronic traumatic encephalopathy (CTE), and thus has been studied in sports where athletes suffer repeated concussions. Tau is important as it helps hold together and stabilize brain neurons. Woodpeckers' brains share similarities to humans with CTE showing most build-up in the frontal and temporal lobes of the brain.[67] It is not yet known whether these accumulations are pathological or the result of behavioral changes. More research is being done on the subject and the woodpecker is a suitable animal model to study.[67] The orientation of the brain within the skull increases the area of contact when pecking to reduce stress on the brain, and their small size helps, given the acceleration speeds.[68]

Mechanical properties

Straight-line trajectory was theorized to be the reason why woodpeckers do not injure themselves, since centripetal forces were the cause of concussion, but they do not always peck in straight lines, so they produce and resist centripetal forces.[64] Laboratory tests show that the woodpeckers' cranial bone produces a significantly higher Young's modulus and ultimate strength scores compared to other birds its size.[69] The cranial bone has a high bone mineral density with plate-like structures that are thick with high numbers of trabeculae that are spaced closely together which all may lead to lower deformation while pecking.

The jaw apparatus was studied, looking into its cushioning effects. When comparing the same impact to the beak and to the forehead, the forehead experiences an impact force 1.72 times that of the beak, due to the contact time being 3.25 ms in the forehead and 4.9 ms in the beak. This is impulse momentum where impulse is the integral of force over time. The quadrate bone and joints play an important role in extending impact time, which decreases impact load to brain tissue.[70]

Bio-inspired ideas

Beams

Bio-inspired honeycomb sandwich beams are inspired by the woodpecker's skull design; this beam's goal is to withstand continuous impacts without the need of replacement. The BHSB is composed of carbon fiber-reinforced plastic (CFRP), this is to mimic the high-strength beak. Next is a rubber layer core for the hyoid bone for absorbing and spreading impact, a second core layer of aluminum honeycomb that is porous and light like the woodpecker's spongey bone for impact cushioning. The final layer is the same as the first a CFRP to act as the skull bone.[71] Bio-inspired honeycomb sandwich beams when compared to conventional beams reduced area damage by 50–80% and carried 40 to 5% of the level of stresses in the bottom layer while having an impact-resistance efficiency 1.65 to 16.22 times higher.

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Cited sources

Further reading

  • Dufort MJ (January 2016). "An augmented supermatrix phylogeny of the avian family Picidae reveals uncertainty deep in the family tree". Molecular Phylogenetics and Evolution. 94 (Pt A): 313–26. doi:10.1016/j.ympev.2015.08.025. PMID 26416706.
  • Fuchs J, Pons JM (July 2015). "A new classification of the Pied Woodpeckers assemblage (Dendropicini, Picidae) based on a comprehensive multi-locus phylogeny". Molecular Phylogenetics and Evolution. 88: 28–37. doi:10.1016/j.ympev.2015.03.016. PMID 25818851.
  • Fuchs J, Pons JM, Bowie RC (March 2017). "Biogeography and diversification dynamics of the African woodpeckers". Molecular Phylogenetics and Evolution. 108: 88–100. doi:10.1016/j.ympev.2017.01.007. PMID 28089840.
  • Gorman G (2004). Woodpeckers of Europe: a study of European Picidae. Chalfont St Peter, Bucks.: Bruce Coleman. ISBN 978-1-872842-05-9.
  • Gorman G (2011). The Black Woodpecker: a monograph on Dryocopus martius (1st ed.). Lynx. ISBN 978-84-96553-79-8.
  • Gorman G (2020). The green woodpecker : a monograph on Picus viridis. Great Britain: Amazon/Picus Press. ISBN 9798676711870.</ref>
  • Koenig WD, Haydock J (1999). "Oaks, acorns, and the geographical ecology of acorn woodpeckers". Journal of Biogeography. 26 (1): 159–165. doi:10.1046/j.1365-2699.1999.00256.x. S2CID 5068060.
  • Lemaitre J, Villard MA (2005). "Foraging patterns of pileated woodpeckers in a managed Acadian forest: a resource selection function". Canadian Journal of Forest Research. 35 (10): 2387–2393. doi:10.1139/x05-148.
  • Michalek KG, Winkler H (2001). "Parental care and parentage in monogamous great spotted woodpeckers (Picoides major) and middle spotted woodpeckers (Picoides medius)". Behaviour. 138 (10): 1259–1285. doi:10.1163/15685390152822210.
  • Shakya SB, Fuchs J, Pons JM, Sheldon FH (November 2017). "Tapping the woodpecker tree for evolutionary insight". Molecular Phylogenetics and Evolution. 116: 182–191. doi:10.1016/j.ympev.2017.09.005. PMID 28890006.
  • Stark RD, Dodenhoff DJ, Johnson EV (1998). "A quantitative analysis of woodpecker drumming" (PDF). Condor. 100 (2): 350–356. doi:10.2307/1370276. JSTOR 1370276.
  • Webb DM, Moore WS (August 2005). "A phylogenetic analysis of woodpeckers and their allies using 12S, Cyt b, and COI nucleotide sequences (class Aves; order Piciformes)". Molecular Phylogenetics and Evolution. 36 (2): 233–48. doi:10.1016/j.ympev.2005.03.015. PMID 15869887.
  • Wiebe KL, Swift TL (2001). "Clutch size relative to tree cavity size in northern flickers". Journal of Avian Biology. 32 (2): 167–173. doi:10.1034/j.1600-048X.2001.320210.x.
  • Yom-Tov Y, Ar A (1993). "Incubation and fledging durations of woodpeckers" (PDF). Condor. 95 (2): 282–287. doi:10.2307/1369350. JSTOR 1369350.

External links

  • Woodpecker videos, photos & sounds on the Internet Bird Collection
  • "Woodpecker" . The American Cyclopædia. 1879.
  • "Woodpecker" . The New Student's Reference Work . 1914.

woodpecker, other, uses, disambiguation, part, bird, family, picidae, which, also, includes, piculets, wrynecks, sapsuckers, members, this, family, found, worldwide, except, australia, guinea, zealand, madagascar, extreme, polar, regions, most, species, live, . For other uses see Woodpecker disambiguation Woodpeckers are part of the bird family Picidae which also includes the piculets wrynecks and sapsuckers 1 Members of this family are found worldwide except for Australia New Guinea New Zealand Madagascar and the extreme polar regions Most species live in forests or woodland habitats although a few species are known that live in treeless areas such as rocky hillsides and deserts and the Gila woodpecker specialises in exploiting cacti WoodpeckerTemporal range 26 0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Late Oligocene to presentPileated woodpeckerTapping sound of a woodpecker help info Scientific classificationKingdom AnimaliaPhylum ChordataClass AvesOrder PiciformesInfraorder PicidesFamily PicidaeLeach 1820SubfamiliesJynginae wrynecks Picinae true woodpeckers Picumninae piculets Sasiinae piculetsMembers of this family are chiefly known for their characteristic behaviour They mostly forage for insect prey on the trunks and branches of trees and often communicate by drumming with their beaks producing a reverberatory sound that can be heard at some distance Some species vary their diet with fruits birds eggs small animals tree sap human scraps and carrion They usually nest and roost in holes that they excavate in tree trunks and their abandoned holes are of importance to other cavity nesting birds They sometimes come into conflict with humans when they make holes in buildings or feed on fruit crops but perform a useful service by their removal of insect pests on trees The Picidae are one of nine living families in the order Piciformes the others being barbets comprising three families toucans toucan barbets and honeyguides which along with woodpeckers comprise the clade Pici and the jacamars and puffbirds in the clade Galbuli DNA sequencing has confirmed the sister relationships of these two groups The family Picidae includes about 240 species arranged in 35 genera Almost 20 species are threatened with extinction due to loss of habitat or habitat fragmentation with one the Bermuda flicker being extinct and a further two possibly being so Contents 1 General characteristics 2 Distribution habitat and movements 2 1 Global distribution 2 2 Habitat requirements 3 Behaviour 3 1 Drumming 3 2 Calls 3 3 Diet and feeding 3 4 Breeding 4 Systematics and evolutionary history 4 1 List of genera 5 Relationship with humans 5 1 Status and conservation 6 Brain impact research 6 1 Anatomy 6 2 Mechanical properties 7 Bio inspired ideas 7 1 Beams 8 References 9 Cited sources 10 Further reading 11 External linksGeneral characteristics Edit A black rumped flameback using its tail for support Woodpeckers include the tiny piculets the smallest of which appears to be the bar breasted piculet at 7 5 cm 3 0 in in length and a weight of 8 9 g 0 31 oz 2 3 Some of the largest woodpeckers can be more than 50 cm 20 in in length The largest surviving species is the great slaty woodpecker which weighs 430 g 15 oz on average and up to 563 g 19 9 oz and measures 45 to 55 cm 18 to 22 in but the extinct imperial woodpecker at 55 to 61 cm 22 to 24 in and ivory billed woodpecker around 48 to 53 cm 19 to 21 in and 516 g 18 2 oz were probably both larger 4 3 5 6 7 The plumage of woodpeckers varies from drab to conspicuous The colours of many species are based on olive and brown and some are pied suggesting a need for camouflage others are boldly patterned in black white and red and many have a crest or tufted feathers on their crowns Woodpeckers tend to be sexually dimorphic but differences between the sexes are generally small exceptions to this are Williamson s sapsucker and the orange backed woodpecker which differ markedly The plumage is moulted fully once a year apart from the wrynecks which have an additional partial moult before breeding 8 Woodpeckers piculets and wrynecks all possess characteristic zygodactyl feet consisting of four toes the first hallux and the fourth facing backward and the second and third facing forward This foot arrangement is good for grasping the limbs and trunks of trees Members of this family can walk vertically up tree trunks which is beneficial for activities such as foraging for food or nest excavation In addition to their strong claws and feet woodpeckers have short strong legs This is typical of birds that regularly forage on trunks Exceptions are the black backed woodpecker and the American and Eurasian three toed woodpeckers which have only three toes on each foot The tails of all woodpeckers except the piculets and wrynecks are stiffened and when the bird perches on a vertical surface the tail and feet work together to support it 4 Woodpeckers have strong bills that they use for drilling and drumming on trees and long sticky tongues for extracting food insects and larvae 4 Woodpecker bills are typically longer sharper and stronger than the bills of piculets and wrynecks but their morphology is very similar The bill s chisel like tip is kept sharp by the pecking action in birds that regularly use it on wood The beak consists of three layers an outer sheath called rhamphotheca made of scales formed from keratin proteins an inner layer of bone which has a large cavity and mineralised collagen fibers and a middle layer made of porous bone which connects the two other layers Furthermore the tongue bone or hyoid bone of the woodpecker is very long and winds around the skull through a special cavity thereby cushioning the brain 9 Combined this anatomy helps the beak absorb mechanical stress 10 Species of woodpecker and flicker that use their bills in soil or for probing as opposed to regular hammering tend to have longer and more decurved bills Due to their smaller bill size many piculets and wrynecks forage in decaying wood more often than woodpeckers Their long sticky tongues which possess bristles aid these birds in grabbing and extracting insects from deep within a hole in a tree The tongue was reported to be used to spear grubs but more detailed studies published in 2004 have shown that the tongue instead wraps around the prey before being pulled out 11 Diagram showing the hyoid bone of Dendrocopos major Many of the foraging breeding and signaling behaviors of woodpeckers involve drumming and hammering using their bills 12 To prevent brain damage from the rapid and repeated powerful impacts woodpeckers have a number of physical features that protect their brains 13 These include a relatively small and smooth brain narrow subdural space little cerebrospinal fluid surrounding it to prevent it from moving back and forth inside the skull during pecking the orientation of the brain within the skull which maximises the contact area between the brain and the skull and the short duration of contact The skull consists of strong but compressible sponge like bone which is most concentrated in the forehead and the back of the skull 13 Another anatomical adaptation of woodpeckers is the enormously elongated hyoid bone which subdivides passes on either side of the spinal column and wraps around the brain case before ending in the right nostril cavity It plays the role of safety belt 9 Computer simulations have shown that 99 7 of the energy generated in pecking is stored in the form of strain energy which is distributed throughout the bird s body with only a small remaining fraction of the energy going into the brain The pecking also causes the woodpecker s skull to heat up which is part of the reason why they often peck in short bursts with brief breaks in between giving the head some time to cool 14 During the millisecond before contact with wood a thickened nictitating membrane closes protecting the eye from flying debris 15 These membranes also prevent the retina from tearing Their nostrils are also protected they are often slit like and have special feathers to cover them Woodpeckers are capable of repeated pecking on a tree at high decelerations on the order of 10 000 m s2 33 000 ft s2 1000 g 16 Some large woodpeckers such as Dryocopus have a fast direct form of flight but the majority of species have a typical undulating flight pattern consisting of a series of rapid flaps followed by a swooping glide Many birds in the genus Melanerpes have distinctive rowing wing strokes while the piculets engage in short bursts of rapid direct flight 17 Distribution habitat and movements Edit Use of cacti for breeding and roosting holes allows some woodpeckers to live in treeless deserts such as the ladder backed woodpecker which uses cacti for nesting See also List of Piciformes by population Global distribution Edit Woodpeckers have a mostly cosmopolitan distribution although they are absent from Australasia Madagascar and Antarctica They are also absent from some of the world s oceanic islands although many insular species are found on continental islands The true woodpeckers subfamily Picinae are distributed across the entire range of the family The Picumninae piculets have a pantropical distribution with species in Southeast Asia Africa and the Neotropics with the greatest diversity being in South America 18 The second piculet subfamily the Sasiinae contains the African piculet and two species in the genus Sasia that are found in Southeast Asia 19 The wrynecks Jynginae are found exclusively in the Old World with the two species occurring in Europe Asia and Africa 18 Most woodpeckers are sedentary but a few examples of migratory species are known such as the rufous bellied woodpecker yellow bellied sapsucker 18 and Eurasian wryneck which breeds in Europe and west Asia and migrates to the Sahel in Africa in the winter 20 More northerly populations of Lewis s woodpecker northern flicker Williamson s sapsucker red breasted sapsucker and red naped sapsucker all move southwards in the fall in North America 18 Most woodpecker movements can be described as dispersive such as when young birds seek territories after fledging or eruptive to escape harsh weather conditions Several species are altitudinal migrants for example the grey capped pygmy woodpecker which moves to lowlands from hills during winter The woodpeckers that do migrate do so during the day 4 Habitat requirements Edit Overall woodpeckers are arboreal birds of wooded habitats They reach their greatest diversity in tropical rainforests but occur in almost all suitable habitats including woodlands savannahs scrublands and bamboo forests Even grasslands and deserts have been colonised by various species These habitats are more easily occupied where a small number of trees exist or in the case of desert species like the Gila woodpecker tall cacti are available for nesting 21 Some are specialists and are associated with coniferous or deciduous woodlands or even like the acorn woodpecker with individual tree genera oaks in this case Other species are generalists and are able to adapt to forest clearance by exploiting secondary growth plantations orchards and parks In general forest dwelling species need rotting or dead wood on which to forage 22 Several species are adapted to spending a portion of their time feeding on the ground and a very small minority have abandoned trees entirely and nest in holes in the ground The ground woodpecker is one such species inhabiting the rocky and grassy hills of South Africa 23 and the Andean flicker is another 22 The Swiss Ornithological Institute has set up a monitoring program to record breeding populations of woodland birds This has shown that deadwood is an important habitat requirement for the black woodpecker great spotted woodpecker middle spotted woodpecker lesser spotted woodpecker European green woodpecker and Eurasian three toed woodpecker Populations of all these species increased by varying amounts from 1990 to 2008 During this period the amount of deadwood in the forest increased and the range of the white backed woodpecker enlarged as it extended eastwards With the exception of the green and middle spotted woodpeckers the increase in the amount of deadwood is likely to be the major factor explaining the population increase of these species 24 Behaviour Edit Woodpecker source source A woodpecker pecking into a tree Problems playing this file See media help Most woodpeckers live solitary lives but their behaviour ranges from highly antisocial species that are aggressive towards their own kind to species that live in groups Solitary species defend such feeding resources as a termite colony or fruit laden tree driving away other conspecifics and returning frequently until the resource is exhausted Aggressive behaviours include bill pointing and jabbing head shaking wing flicking chasing drumming and vocalisations Ritual actions do not usually result in contact and birds may freeze for a while before they resume their dispute The coloured patches may be flouted and in some instances these antagonistic behaviours resemble courtship rituals 25 Group living species tend to be communal group breeders 25 In addition to these species a number of species may join mixed species foraging flocks with other insectivorous birds although they tend to stay at the edges of these groups Joining these flocks allows woodpeckers to decrease their anti predator vigilance and increase their feeding rate 26 Woodpeckers are diurnal roosting at night inside holes and crevices In many species the roost will become the nest site during the breeding season but in some species they have separate functions the grey and buff woodpecker makes several shallow holes for roosting which are quite distinct from its nesting site Most birds roost alone and will oust intruders from their chosen site but the Magellanic woodpecker and acorn woodpecker are cooperative roosters 25 Drumming Edit Drumming is a form of nonvocal communication used by most species of woodpeckers and involves the bill being repeatedly struck on a hard surface with great rapidity After a pause the drum roll is repeated with each species having a pattern that is unique in the number of beats in the roll the length of the roll the length of the gap between rolls and the cadence 27 28 The drumming is mainly a territorial call equivalent to the song of a passerine 29 Woodpeckers choose a surface that resonates such as a hollow tree and may use man made structures such as gutters and downpipes 30 Drumming serves for the mutual recognition of conspecifics and plays a part in courtship rituals Individual birds are thought to be able to distinguish the drumming of their mates and those of their neighbours 31 Drumming in woodpeckers is controlled by a set of nuclei in the forebrain that closely resemble the brain regions that underlie song learning and production in many songbirds 32 Calls Edit Woodpeckers do not have such a wide range of songs and calls as do passerine birds and the sounds they make tend to be simpler in structure Calls produced include brief high pitched notes trills rattles twittering whistling chattering nasal churrs screams and wails These calls are used by both sexes in communication and are related to the circumstances of the occasion these include courtship territorial disputes and alarm calls Each species has its own range of calls which tend to be in the 1 0 to 2 5 kHz range for efficient transmission through forested environments Mated couples may exchange muted low pitched calls and nestlings often issue noisy begging calls from inside their nest cavity 29 The wrynecks have a more musical song and in some areas the song of the newly arrived Eurasian wryneck is considered to be the harbinger of spring 33 The piculets either have a song consisting of a long descending trill or a descending series of two to six sometimes more individual notes and this song alerts ornithologists to the presence of the birds as they are easily overlooked 34 Diet and feeding Edit Holes bored by feeding woodpeckers Most woodpecker species feed on insects and other invertebrates living under bark and in wood but overall the family is characterized by its dietary flexibility with many species being both highly omnivorous and opportunistic The diet includes ants termites beetles and their larvae caterpillars spiders other arthropods bird eggs nestlings small rodents lizards fruit nuts and sap Many insects and their grubs are taken from living and dead trees by excavation The bird may hear sounds from inside the timber indicating where creating a hole would be productive 25 Crustaceans molluscs and carrion may be eaten by some species including the great spotted woodpecker and bird feeders are visited for suet and domestic scraps 35 Other means are also used to garner prey Some species such as the red naped sapsucker sally into the air to catch flying insects and many species probe into crevices and under bark or glean prey from leaves and twigs The rufous woodpecker specialises in attacking the nests of arboreal ants and the buff spotted woodpecker feeds on and nests in termite mounds Other species such as the wrynecks and the Andean flicker feed wholly or partly on the ground 25 Ecologically woodpeckers help to keep trees healthy by keeping them from suffering mass infestations The family is noted for its ability to acquire wood boring grubs from the trunks and branches whether the timber is alive or dead Having hammered a hole into the wood the prey is extracted by use of a long barbed tongue Woodpeckers consume beetles that burrow into trees removing as many as 85 of emerald ash borer larvae from individual ash trees 36 The ability to excavate allows woodpeckers to obtain tree sap an important source of food for some species Most famously the sapsuckers genus Sphyrapicus feed in this fashion but the technique is not restricted to these and others such as the acorn woodpecker and white headed woodpecker also feed on sap The technique was once thought to be restricted to the New World but Old World species such as the Arabian woodpecker and great spotted woodpecker also feed in this way 4 Breeding Edit A male black woodpecker attending its chicks All members of the family Picidae nest in cavities nearly always in the trunks and branches of trees well away from the foliage Where possible an area of rotten wood surrounded by sound timber is used Where trees are in short supply the gilded flicker and ladder backed woodpecker excavate holes in cactus and the Andean flicker and ground woodpecker dig holes in earth banks The campo flicker sometimes chooses termite mounds the rufous woodpecker prefers to use ants nests in trees and the bamboo woodpecker specialises in bamboos 37 Woodpeckers also excavate nest holes in residential and commercial structures and wooden utility poles 36 Woodpeckers and piculets excavate their own nests but wrynecks do not and need to find pre existing cavities A typical nest has a round entrance hole that just fits the bird leading to an enlarged vertical chamber below No nesting material is used apart from some wood chips produced during the excavation other wood chips are liberally scattered on the ground thus providing visual evidence of the site of the nest 38 Many species of woodpeckers excavate one hole per breeding season sometimes after multiple attempts It takes around a month to finish the job and abandoned holes are used by other birds and mammals that are cavity nesters unable to excavate their own holes 39 Cavities are in great demand for nesting by other cavity nesters so woodpeckers face competition for the nesting sites they excavate from the moment the hole becomes usable This may come from other species of woodpecker or other cavity nesting birds such as swallows and starlings Woodpeckers may aggressively harass potential competitors and also use other strategies to reduce the chance of being usurped from their nesting sites for example the red crowned woodpecker digs its nest in the underside of a small branch which reduces the chance that a larger species will take it over and expand it 40 Members of Picidae are typically monogamous with a few species breeding cooperatively and some polygamy reported in a few others 41 Polyandry where a female raises two broods with two separate males has also been reported in the West Indian woodpecker 42 Another unusual social system is that of the acorn woodpecker which is a polygynandrous cooperative breeder where groups of up to 12 individuals breed and help to raise the young 4 Young birds from previous years may stay behind to help raise the group s young and studies have found reproductive success for the group goes up with group size but individual success goes down Birds may be forced to remain in groups due to a lack of habitat to which to disperse 43 Woodpecker which feeding its chick A pair works together to help build the nest incubate the eggs and raise their altricial young In most species though the male does most of the nest excavation and takes the night shift while incubating the eggs A clutch usually consists of two to five round white eggs Since these birds are cavity nesters their eggs do not need to be camouflaged and the white color helps the parents to see them in dim light The eggs are incubated for about 11 14 days before they hatch About 18 30 days are then needed before the chicks are fully fledged and ready to leave the nest In most species soon after this the young are left to fend for themselves exceptions being the various social species and the Hispaniolan woodpecker where adults continue to feed their young for several months In general cavity nesting is a successful strategy and a higher proportion of young is reared than is the case with birds that nest in the open In Africa several species of honeyguide are brood parasites of woodpeckers 38 Systematics and evolutionary history Edit Lesser goldenback Dinopium benghalense in Guwahati India The Picidae are just one of nine living families in the order Piciformes Other members of this group such as the jacamars puffbirds barbets toucans and honeyguides have traditionally been thought to be closely related to the woodpecker family true woodpeckers piculets wrynecks and sapsuckers The clade Pici woodpeckers barbets toucans and honeyguides is well supported and shares a zygodactyl foot with the Galbuli puffbirds and jacamars More recently several DNA sequence analyses have confirmed that Pici and Galbuli are sister groups 44 The name Picidae for the family was introduced by English zoologist William Elford Leach in a guide to the contents of the British Museum published in 1820 45 46 The phylogeny has been updated according to new knowledge about convergence patterns and evolutionary history 47 48 Most notably the relationship of the Picinae genera has been largely clarified and the Antillean piculet was found to be a surviving offshoot of protowoodpeckers Genetic analysis supports the monophyly of the Picidae which seem to have originated in the Old World but the geographic origins of the Picinae is unclear The Picumninae are returned as paraphyletic 47 Morphological and behavioural characters in addition to DNA evidence highlights genus Hemicircus as the sister group of all remaining true woodpeckers besides a sister group relationship between the true woodpecker tribes Dendropicini and Malarpicini 49 The evolutionary history of this group is not well documented but the known fossils allow some preliminary conclusions the earliest known modern picids were piculet like forms of the Late Oligocene about 25 million years ago Mya By that time however the group was already present in the Americas and Europe and they actually may have evolved much earlier maybe as early as the Early Eocene 50 Mya The modern subfamilies appear to be rather young by comparison until the mid Miocene 10 15 Mya all picids seem to have been small or mid sized birds similar to a mixture between a piculet and a wryneck A feather enclosed in fossil amber from the Dominican Republic dated to about 25 Mya however seems to indicate that the Nesoctitinae were already a distinct lineage by then 50 Stepwise adaptations for drilling tapping and climbing head first on vertical surfaces have been suggested 49 The last common ancestor of woodpeckers Picidae was incapable of climbing up tree trunks or excavating nest cavities by drilling with its beak The first adaptations for drilling including reinforced rhamphotheca frontal overhang and processus dorsalis pterygoidei evolved in the ancestral lineage of piculets and true woodpeckers Additional adaptations for drilling and tapping enlarged condylus lateralis of the quadrate and fused lower mandible have evolved in the ancestral lineage of true woodpeckers Hemicircus excepting The inner rectrix pairs became stiffened and the pygostyle lamina was enlarged in the ancestral lineage of true woodpeckers Hemicircus included which facilitated climbing head first up tree limbs Genus Hemicircus excepting the tail feathers were further transformed for specialized support the pygostyle disc became greatly enlarged and the ectropodactyl toe arrangement evolved These latter characters may have facilitated enormous increases in body size in some lineages 49 Prehistoric representatives of the extant Picidae genera are treated in the genus articles An enigmatic form based on a coracoid found in Pliocene deposits of New Providence in the Bahamas has been described as Bathoceleus hyphalus and probably also is a woodpecker 51 The following cladogram is based on the comprehensive molecular phylogenetic study of the woodpeckers published in 2017 together with the list of bird species maintained by Frank Gill Pamela Rasmussen and David Donsker on behalf of the International Ornithological Committee IOC The Cuban green woodpecker in the monotypic genus Xiphidiopicus was not included in the study 52 53 The relative positions of Picumninae Sasiinae and Picinae in the cladogram are uncertain In the 2017 study the results depended upon which of two different statistical procedures were used to analyse the DNA sequence data One method found that Sasiinae was sister to Picinae as shown below the other method found that Sasiinae was sister to a clade containing both Picumninae and Picinae 52 Picidae Jynginae Jynx 2 species wrynecks Picumninae Picumnus 26 species piculets Sasiinae Verreauxia African piculetSasia 2 species piculets Picinae Nesoctitini Nesoctites Antillean piculetHemicirini Hemicircus 2 speciesPicini Micropternus rufous woodpeckerMeiglyptes 3 speciesGecinulus 3 speciesDinopium 5 species flamebacks Picus 14 speciesChrysophlegma 3 speciesPardipicus 2 speciesGeocolaptes ground woodpeckerCampethera 11 speciesMulleripicus 4 speciesDryocopus 6 speciesCeleus 13 speciesPiculus 7 speciesColaptes 14 speciesCampephilini Campephilus 11 speciesBlythipicus 2 speciesReinwardtipicus orange backed woodpeckerChrysocolaptes 9 speciesMelanerpini Sphyrapicus 4 species sapsuckers Melanerpes 24 speciesPicoides 3 speciesYungipicus 7 speciesDendrocoptes 4 species including Leiopicus Chloropicus 3 speciesDendropicos 12 speciesDendrocopos 12 speciesDryobates 5 speciesLeuconotopicus 6 speciesVeniliornis 14 species List of genera Edit Ochre collared piculet Picumnus temminckii Red crowned woodpecker Melanerpes rubricapillus rubricapillus female Tobago The woodpecker family Picidae contains 37 genera 53 For more detail see list of woodpecker species Cuban green woodpecker Xiphidiopicus percussus female Cuba Campo flickerColaptes campestrisfemale Brazil Family Picidae Subfamily Jynginae wrynecks Jynx 2 species Subfamily Picumninae piculets 54 Picumnus piculets 26 species Subfamily Sasiinae 19 Verreauxia African piculet Sasia Asian piculets 2 species Subfamily Picinae true woodpeckers Tribe Nesoctitini Nesoctites monotypic Antillean piculet Tribe Hemicircini Hemicircus 2 species Tribe Picini Micropternus monotypic rufous woodpecker Meiglyptes 3 species Gecinulus 3 species Dinopium 5 species flamebacks Picus 14 species Chrysophlegma 3 species Pardipicus 2 species Geocolaptes monotypic ground woodpecker Campethera 11 species Mulleripicus 4 species Dryocopus 6 species Celeus 13 species Piculus 7 species Colaptes 14 species Tribe Campephilini Campephilus 11 species Blythipicus 2 species Reinwardtipicus monotypic orange backed woodpecker Chrysocolaptes 9 species flamebacks Tribe Melanerpini Sphyrapicus 4 species sapsuckers Melanerpes 24 species Picoides 3 species Yungipicus 7 species Leiopicus monotypic yellow crowned woodpecker Dendrocoptes 3 species Chloropicus 3 species Dendropicos 12 species Dendrocopos 12 species Dryobates 5 species Leuconotopicus 6 species Veniliornis 14 species Xiphidiopicus monotypic Cuban green woodpecker Incertae sedis fossils Genus Palaeopicus Late Oligocene of France Picidae gen et sp indet Middle Miocene of New Mexico US Picidae gen et sp indet Late Miocene of Gargano Peninsula Italy Genus Palaeonerpes Ogallala Early Pliocene of Hitchcock County US possibly dendropicine Genus Pliopicus Early Pliocene of Kansas US possibly dendropicine cf Colaptes DMNH 1262 Early Pliocene of Ainsworth US malarpicine Relationship with humans EditIn general humans consider woodpeckers in a favourable light they are viewed as interesting birds and fascinating to watch as they drum or forage but their activities are not universally appreciated 55 Many woodpecker species are known to excavate holes in buildings fencing and utility poles creating health and or safety issues for affected structures Such activity is very difficult to discourage and can be costly to repair 56 Woodpeckers also drum on various reverberatory structures on buildings such as gutters downspouts chimneys vents and aluminium sheeting 57 Drumming is a less forceful type of pecking that serves to establish territory and attract mates 56 Houses with shingles or wooden boarding are also attractive as possible nesting or roosting sites especially when close to large trees or woodland Several exploratory holes may be made especially at the junctions of vertical boards or at the corners of tongue and groove boarding The birds may also drill holes in houses as they forage for insect larvae and pupae hidden behind the woodwork 57 Woodpeckers sometimes cause problems when they raid fruit crops but their foraging activities are mostly beneficial as they control forest insect pests such as the woodboring beetles that create galleries behind the bark and can kill trees They also eat ants which may be tending sap sucking pests such as mealybugs as is the case with the rufous woodpecker in coffee plantations in India 55 Woodpeckers can serve as indicator species demonstrating the quality of the habitat Their hole making abilities make their presence in an area an important part of the ecosystem because these cavities are used for breeding and roosting by many bird species that are unable to excavate their own holes as well as being used by various mammals and invertebrates 55 The spongy bones of the woodpecker s skull and the flexibility of its beak both of which provide protection for the brain when drumming have provided inspiration to engineers a black box needs to survive intact when a plane falls from the sky and modelling the black box with regard to a woodpecker s anatomy has increased the resistance of this device to damage 60 fold 58 The design of protective helmets is another field being influenced by the study of woodpeckers 58 One of the accounts of the founding of Rome preserved in the work known as Origo Gentis Romanae refers to a legend of a woodpecker bringing food to the boys Romulus and Remus during the time they were abandoned in the wild thus enabling them to survive and play their part in history Status and conservation Edit The ivory billed woodpecker is classified as critically endangered by the IUCN 59 and some authorities believe it may already be extinct In a global survey of the risk of extinction faced by the various bird families woodpeckers were the only bird family to have significantly fewer species at risk than would be expected 60 Nevertheless several woodpeckers are under threat as their habitats are destroyed Being woodland birds deforestation and clearance of land for agriculture and other purposes can reduce populations dramatically Some species adapt to living in plantations and secondary growth or to open countryside with forest remnants and scattered trees but some do not A few species have even flourished when they have adapted to man made habitats There are few conservation projects directed primarily at woodpeckers but they benefit whenever their habitat is conserved 55 The red cockaded woodpecker has been the focus of much conservation effort in the southeastern United States with artificial cavities being constructed in the longleaf pines they favour as nesting sites 61 Two species of woodpeckers in the Americas the ivory billed woodpecker is critically endangered and the imperial woodpecker is classified as extinct in the wild with some authorities believing them extinct though possible but disputed ongoing sightings of ivory billed woodpeckers have been made in the United States 62 and a small population may survive in Cuba 59 A critically endangered species is the Okinawa woodpecker from Japan with a single declining population of a few hundred birds It is threatened by deforestation golf course dam and helipad construction road building and agricultural development 63 Brain impact research EditAnatomy Edit Woodpeckers possess many sophisticated shock absorption mechanisms that help protect them from head injury Micro CT scans show that plate like spongy bones are in the skull with an uneven distribution highly accumulated in the forehead and occiput but not in other regions 64 Along with the long hyoid bone safety belt the woodpecker has uneven beak lengths which drastically reduce strains when compared to equal length 64 65 Models have shown that pecking force is changed to strain energy and stored into the body at around 99 absorption while 1 is in the head The head also has many factors that reduce strain to the brain and small portions of energy are dissipated into the form of heat therefore the pecks are always intermittent 66 Tau protein accumulation is associated with chronic traumatic encephalopathy CTE and thus has been studied in sports where athletes suffer repeated concussions Tau is important as it helps hold together and stabilize brain neurons Woodpeckers brains share similarities to humans with CTE showing most build up in the frontal and temporal lobes of the brain 67 It is not yet known whether these accumulations are pathological or the result of behavioral changes More research is being done on the subject and the woodpecker is a suitable animal model to study 67 The orientation of the brain within the skull increases the area of contact when pecking to reduce stress on the brain and their small size helps given the acceleration speeds 68 Mechanical properties Edit Straight line trajectory was theorized to be the reason why woodpeckers do not injure themselves since centripetal forces were the cause of concussion but they do not always peck in straight lines so they produce and resist centripetal forces 64 Laboratory tests show that the woodpeckers cranial bone produces a significantly higher Young s modulus and ultimate strength scores compared to other birds its size 69 The cranial bone has a high bone mineral density with plate like structures that are thick with high numbers of trabeculae that are spaced closely together which all may lead to lower deformation while pecking The jaw apparatus was studied looking into its cushioning effects When comparing the same impact to the beak and to the forehead the forehead experiences an impact force 1 72 times that of the beak due to the contact time being 3 25 ms in the forehead and 4 9 ms in the beak This is impulse momentum where impulse is the integral of force over time The quadrate bone and joints play an important role in extending impact time which decreases impact load to brain tissue 70 Bio inspired ideas EditBeams Edit Bio inspired honeycomb sandwich beams are inspired by the woodpecker s skull design this beam s goal is to withstand continuous impacts without the need of replacement The BHSB is composed of carbon fiber reinforced plastic CFRP this is to mimic the high strength beak Next is a rubber layer core for the hyoid bone for absorbing and spreading impact a second core layer of aluminum honeycomb that is porous and light like the woodpecker s spongey bone for impact cushioning The final layer is the same as the first a CFRP to act as the skull bone 71 Bio inspired honeycomb sandwich beams when compared to conventional beams reduced area damage by 50 80 and carried 40 to 5 of the level of stresses in the bottom layer while having an impact resistance efficiency 1 65 to 16 22 times higher References Edit Bouglouan Nicole Family Picidae Woodpeckers Piculets Wrynecks oiseaux birds com Retrieved 2 December 2022 Winkler H Christie DA Bonan A 2020 Bar breasted Piculet Picumnus aurifrons In del Hoyo J Elliott A Sargatal J Christie DA de Juana E eds Birds of the World Ithaca NY USA Cornell Lab of Ornithology a b Dunning John B Jr ed 2008 CRC Handbook of Avian Body Masses 2nd ed CRC Press ISBN 978 1 4200 6444 5 a b c d e f Winkler Hans amp Christie David A 2002 Family Picidae Woodpeckers in del Hoyo J Elliot A amp Sargatal J editors 2002 Handbook of the Birds of the World Volume 7 Jacamars to Woodpeckers Lynx Edicions ISBN 978 84 87334 37 5 Jackson JA 2020 Ivory billed Woodpecker Campephilus principalis In Poole AF Gill FB eds Birds of the World Ithaca NY USA Cornell Lab of Ornithology Howell SN Webb S 1995 A guide to the birds of Mexico and northern Central America Oxford University Press Styring Alison R Hussin Mohamed Zakaria bin 2004 Foraging ecology of woodpeckers in lowland Malaysian rain forests Journal of Tropical Ecology 20 5 487 494 doi 10 1017 S0266467404001579 S2CID 83528456 Gorman 2014 pp 22 23 a b Wang L Cheung JT Pu F Li D Zhang M Fan Y 2011 Why do woodpeckers resist head impact injury a biomechanical investigation PLOS ONE 6 10 e26490 Bibcode 2011PLoSO 626490W doi 10 1371 journal pone 0026490 PMC 3202538 PMID 22046293 Helmenstine T 8 May 2014 Woodpecker Beak Shock Absorbers Science Notes Retrieved 24 July 2017 Villard P Cuisin J 2004 How do woodpeckers extract grubs with their tongues A study of the Guadeloupe woodpecker Melanerpes herminieri in the French Indies Auk 121 2 509 514 doi 10 1642 0004 8038 2004 121 0509 HDWEGW 2 0 CO 2 S2CID 86781719 Gibson L 2006 Woodpecker pecking how woodpeckers avoid brain injury PDF Journal of Zoology 270 3 462 465 doi 10 1111 j 1469 7998 2006 00166 x hdl 1721 1 70094 a b Puiu T 23 March 2017 Why woodpeckers don t get headaches ZME Science Retrieved 24 July 2017 Gammon K 25 August 2014 Woodpecker Bodies Cushion Collision Impact On Bird Brains Inside Science Retrieved 24 July 2017 May PR Fuster JM Haber J Hirschman A June 1979 Woodpecker drilling behavior An endorsement of the rotational theory of impact brain injury Archives of Neurology 36 6 370 3 doi 10 1136 bjo 86 8 843 PMC 1771249 PMID 454236 Gibson LJ 2006 Woodpecker pecking how woodpeckers avoid brain injury Journal of Zoology 270 3 462 465 doi 10 1111 j 1469 7998 2006 00166 x hdl 1721 1 70094 Gorman 2014 p 27 a b c d Gorman 2014 p 15 a b Sangster G Gaudin J Fuchs J 2022 A new subfamily taxon for Sasia and Verreauxia Picidae Bulletin of the British Ornithologists Club 142 4 478 479 doi 10 25226 bboc v142i4 2022 a6 S2CID 254367038 Reichlin TS Schaub M Menz MH Mermod M Portner P Arlettaz R Jenni L 2008 Migration patterns of Hoopoe Upupa epops and Wryneck Jynx torquilla an analysis of European ring recoveries PDF Journal of Ornithology 150 2 393 400 doi 10 1007 s10336 008 0361 3 S2CID 43360238 Korol J Hutto R 1984 Factors Affecting Nest Site Location in Gila Woodpeckers PDF Condor 86 1 73 78 doi 10 2307 1367350 JSTOR 1367350 a b Gorman 2014 p 18 Short L 1971 The evolution of terrestrial woodpeckers American Museum Novitates 2467 hdl 2246 2675 Mollet P Zbinden N Schmid H 2009 An increase in the population of woodpeckers and other bird species thanks to an increase in the quantities of deadwood FAO Retrieved 28 March 2017 a b c d e Gorman 2014 pp 19 20 Kimberly S 1984 Information Exploitation By Downy Woodpeckers in Mixed Species Flocks Behaviour 91 4 294 311 doi 10 1163 156853984X00128 Miles MC Schuppe ER Ligon RM Fuxjager MJ 2018 Macroevolutionary patterning of woodpecker drums reveals how sexual selection elaborates signals under constraint Proceedings of the Royal Society B Biological Sciences 285 1873 doi 10 1098 rspb 2017 2628 PMC 5832706 PMID 29467264 Miles MC Schuppe ER Fuxjager MJ 2020 Selection for Rhythm as a Trigger for Recursive Evolution in the Elaborate Display System of Woodpeckers The American Naturalist 195 5 772 787 doi 10 1086 707748 PMID 32364790 S2CID 212917887 a b Gorman 2014 p 28 Williams Jr EH 2005 The Nature Handbook A Guide to Observing the Great Outdoors Oxford University Press p 118 ISBN 978 0 19 972075 0 Sarkar A 2003 Fundamentals Of Animals Behaviour Discovery Publishing House p 264 ISBN 978 81 7141 742 1 Schuppe ER Cantin L Chakraborty M Biegler MT Jarvis ER Chen CC 2022 Forebrain nuclei linked to woodpecker territorial drum displays mirror those that enable vocal learning in songbirds PLOS Biology 20 9 e3001751 doi 10 1371 journal pbio 3001751 PMC 9488818 PMID 36125990 Noel T 1841 Rymes and Roundelayes Smith p 144 Hilty SL 2002 Birds of Venezuela Princeton University Press p 464 ISBN 978 1 4008 3409 9 Winkler H Christie DA Kirwan GM 2020 del Hoyo J Elliott A Sargatal J Christie DA de Juana E eds Great Spotted Woodpecker Dendrocopos major version 1 0 Birds of the World Ithaca NY USA Cornell Lab of Ornithology doi 10 2173 bow grswoo 01 S2CID 226025386 a b Graham R 24 July 2014 Resilient Woodpeckers hard to knock or stop Birds News Archived from the original on 4 April 2016 Retrieved 24 March 2016 Gorman 2014 p 20 a b Gorman 2014 p 22 Kotaka N Matsuoka S 2002 Secondary users of Great Spotted Woodpecker Dendrocopos major nest cavities in urban and suburban forests in Sapporo City northern Japan Ornithological Science 1 2 117 122 doi 10 2326 osj 1 117 Short LL 1979 Burdens of the Picid Hole Excavating Habit PDF Wilson Bulletin 91 1 16 28 Wiktander U Olsson O Nilsson SG 2000 Parental care and social mating system in the Lesser Spotted Woodpecker Dendrocopos minor Journal of Avian Biology 31 4 447 456 doi 10 1034 j 1600 048X 2000 310003 x Willimont LA Jackson JA Jackson BJ 1991 Classical polyandry in the West Indian woodpecker on Abaco Bahamas PDF Wilson Bulletin 103 124 125 Koenig WD 1981 Reproductive success group size and the evolution of cooperative breeding in the acorn woodpecker The American Naturalist 117 4 421 443 doi 10 1086 283726 JSTOR 2460453 S2CID 85399703 Johansson US Ericson GP 2003 Molecular support for a sister group relationship between Pici and Galbulae Piciformes sensu Wetmore 1960 PDF Journal of Avian Biology 34 2 185 197 doi 10 1034 j 1600 048X 2003 03103 x Leach WE 1820 Eleventh Room Synopsis of the Contents of the British Museum Vol 17 17th ed London British Museum p 68 Although the name of the author is not specified in the document Leach was the keeper of zoology at the time Bock WJ 1994 History and Nomenclature of Avian Family Group Names Bulletin of the American Museum of Natural History Vol Number 222 New York American Museum of Natural History pp 146 192 hdl 2246 830 a b Benz BW Robbins MB Peterson AT August 2006 Evolutionary history of woodpeckers and allies Aves Picidae placing key taxa on the phylogenetic tree Molecular Phylogenetics and Evolution 40 2 389 99 doi 10 1016 j ympev 2006 02 021 PMID 16635580 Moore WS Weibel AC Agius A 2006 Mitochondrial DNA phylogeny of the woodpecker genus Veniliornis Picidae Picinae and related genera implies convergent evolution of plumage patterns PDF Biological Journal of the Linnean Society 87 4 611 624 doi 10 1111 j 1095 8312 2006 00586 x a b c Manegold A Topfer T 2013 The systematic position of Hemicircus and the stepwise evolution of adaptations for drilling tapping and climbing up in true woodpeckers Picinae Picidae Journal of Zoological Systematics and Evolutionary Research 51 1 72 82 doi 10 1111 jzs 12000 Grimaldi DA Case GR 1995 A feather in amber from the Upper Cretaceous of New Jersey PDF American Museum Novitates 3126 1 6 Cracraft J Morony Jr JJ 1969 A new Pliocene woodpecker with comments on the fossil Picidae PDF American Museum Novitates 2400 1 8 a b Shakya S B Fuchs J Pons J M Sheldon F H 2017 Tapping the woodpecker tree for evolutionary insight Molecular Phylogenetics and Evolution 116 182 191 doi 10 1016 j ympev 2017 09 005 PMID 28890006 a b Gill Frank Donsker David Rasmussen Pamela eds January 2023 Woodpeckers IOC World Bird List Version 13 1 International Ornithologists Union Retrieved 19 February 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1688257 PMID 7583692 Copeyon CK Walters JR Carter III JH 1991 Induction of Red Cockaded Woodpecker Group Formation by Artificial Cavity Construction The Journal of Wildlife Management 55 4 549 556 doi 10 2307 3809497 JSTOR 3809497 The search for the ivory billed woodpecker Big Woods Conservation Partnership Retrieved 2017 03 26 BirdLife International 2018 Dendrocopos noguchii IUCN Red List of Threatened Species 2018 e T22681531A125513230 doi 10 2305 IUCN UK 2018 2 RLTS T22681531A125513230 en Retrieved 12 November 2021 a b c Wang L Cheung JT Pu F Li D Zhang M Fan Y 2011 10 26 Briffa M ed Why do woodpeckers resist head impact injury a biomechanical investigation PLOS ONE 6 10 e26490 Bibcode 2011PLoSO 626490W doi 10 1371 journal pone 0026490 PMC 3202538 PMID 22046293 May PR Newman P Fuster JM Hirschman A February 1976 Woodpeckers and Head Injury The Lancet 307 7957 454 455 doi 10 1016 s0140 6736 76 91477 x PMID 55721 S2CID 28685873 How the woodpecker avoids brain injury despite high speed impacts via optimal anti shock body structure phys org 2014 08 11 Retrieved 2021 04 16 a b Farah G Siwek D Cummings P 2018 02 02 Tau accumulations in the brains of woodpeckers PLOS ONE 13 2 e0191526 Bibcode 2018PLoSO 1391526F doi 10 1371 journal pone 0191526 PMC 5796688 PMID 29394252 Gibson LJ November 2006 Woodpecker pecking how woodpeckers avoid brain injury Journal of Zoology 270 3 462 465 doi 10 1111 j 1469 7998 2006 00166 x hdl 1721 1 70094 Wang L Zhang H Fan Y November 2011 Comparative study of the mechanical properties micro structure and composition of the cranial and beak bones of the great spotted woodpecker and the lark bird Science China Life Sciences 54 11 1036 41 doi 10 1007 s11427 011 4242 2 PMID 22173310 S2CID 25697639 Xu P Ni Y Lu S Liu S Zhou X Fan Y January 2021 The cushioning function of woodpecker s jaw apparatus during the pecking process Computer Methods in Biomechanics and Biomedical Engineering 24 5 527 537 doi 10 1080 10255842 2020 1838489 PMID 33439040 S2CID 231596453 Abo Sabah SH Kueh AB Al Fasih MY April 2018 Bio inspired vs conventional sandwich beams A low velocity repeated impact behavior exploration Construction and Building Materials 169 193 204 doi 10 1016 j conbuildmat 2018 02 201 Cited sources EditGorman G 2014 Woodpeckers of the World A Photographic Guide Firefly Books ISBN 978 1 77085 309 6 Further reading EditDufort MJ January 2016 An augmented supermatrix phylogeny of the avian family Picidae reveals uncertainty deep in the family tree Molecular Phylogenetics and Evolution 94 Pt A 313 26 doi 10 1016 j ympev 2015 08 025 PMID 26416706 Fuchs J Pons JM July 2015 A new classification of the Pied Woodpeckers assemblage Dendropicini Picidae based on a comprehensive multi locus phylogeny Molecular Phylogenetics and Evolution 88 28 37 doi 10 1016 j ympev 2015 03 016 PMID 25818851 Fuchs J Pons JM Bowie RC March 2017 Biogeography and diversification dynamics of the African woodpeckers Molecular Phylogenetics and Evolution 108 88 100 doi 10 1016 j ympev 2017 01 007 PMID 28089840 Gorman G 2004 Woodpeckers of Europe a study of European Picidae Chalfont St Peter Bucks Bruce Coleman ISBN 978 1 872842 05 9 Gorman G 2011 The Black Woodpecker a monograph on Dryocopus martius 1st ed Lynx ISBN 978 84 96553 79 8 Gorman G 2020 The green woodpecker a monograph on Picus viridis Great Britain Amazon Picus Press ISBN 9798676711870 lt ref gt Koenig WD Haydock J 1999 Oaks acorns and the geographical ecology of acorn woodpeckers Journal of Biogeography 26 1 159 165 doi 10 1046 j 1365 2699 1999 00256 x S2CID 5068060 Lemaitre J Villard MA 2005 Foraging patterns of pileated woodpeckers in a managed Acadian forest a resource selection function Canadian Journal of Forest Research 35 10 2387 2393 doi 10 1139 x05 148 Michalek KG Winkler H 2001 Parental care and parentage in monogamous great spotted woodpeckers Picoides major and middle spotted woodpeckers Picoides medius Behaviour 138 10 1259 1285 doi 10 1163 15685390152822210 Shakya SB Fuchs J Pons JM Sheldon FH November 2017 Tapping the woodpecker tree for evolutionary insight Molecular Phylogenetics and Evolution 116 182 191 doi 10 1016 j ympev 2017 09 005 PMID 28890006 Stark RD Dodenhoff DJ Johnson EV 1998 A quantitative analysis of woodpecker drumming PDF Condor 100 2 350 356 doi 10 2307 1370276 JSTOR 1370276 Webb DM Moore WS August 2005 A phylogenetic analysis of woodpeckers and their allies using 12S Cyt b and COI nucleotide sequences class Aves order Piciformes Molecular Phylogenetics and Evolution 36 2 233 48 doi 10 1016 j ympev 2005 03 015 PMID 15869887 Wiebe KL Swift TL 2001 Clutch size relative to tree cavity size in northern flickers Journal of Avian Biology 32 2 167 173 doi 10 1034 j 1600 048X 2001 320210 x Yom Tov Y Ar A 1993 Incubation and fledging durations of woodpeckers PDF Condor 95 2 282 287 doi 10 2307 1369350 JSTOR 1369350 External links Edit Wikimedia Commons has media related to Picidae Wikispecies has information related to Picidae Woodpecker videos photos amp sounds on the Internet Bird Collection Woodpecker The American Cyclopaedia 1879 Woodpecker The New Student s Reference Work 1914 Retrieved from https en wikipedia org w index php title Woodpecker amp oldid 1147797651, wikipedia, wiki, book, books, library,

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