Woodpeckers are part of thebirdfamilyPicidae, which also includes thepiculets,wrynecks andsapsuckers.[1] Members of this family are found worldwide, except forAustralia,New Guinea,New Zealand,Madagascar and the extreme polar regions. Most species live inforests orwoodland habitats, although a few species are known that live in treeless areas, such as rocky hillsides and deserts, and theGila woodpecker specialises in exploiting cacti.
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, andcarrion. 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.
Woodpeckers include the tinypiculets, the smallest of which appears to be thebar-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 thegreat 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 extinctimperial woodpecker, at 55 to 61 cm (22 to 24 in), andivory-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]
Theplumage 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 forcamouflage; others are boldly patterned in black, white, and red, and many have a crest or tufted feathers on their crowns. Woodpeckers tend to besexually dimorphic, but differences between the sexes are generally small; exceptions to this areWilliamson's sapsucker and theorange-backed woodpecker, which differ markedly. The plumage ismoulted fully once a year apart from thewrynecks, which have an additional partial moult before breeding.[8]
Woodpeckers, piculets, and wrynecks all possess characteristiczygodactyl 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 theblack-backed woodpecker and theAmerican andEurasian 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 strongbills 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 theirmorphology is very similar. The bill's chisel-like tip is kept sharp by thepecking action in birds that regularly use it on wood. The beak consists of three layers; an outer sheath calledrhamphotheca, made of scales formed fromkeratin proteins, an inner layer of bone which has a large cavity and mineralisedcollagen 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 barbs, 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]
Many of the foraging, breeding, and signaling behaviors of woodpeckers involve drumming and hammering using their bills.[12] To preventbrain 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, narrowsubdural space, littlecerebrospinal 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 elongatedhyoid 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 ofstrain 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 thickenednictitating membrane closes, protecting the eye from flying debris.[15] These membranes also prevent theretina 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 highdecelerations on the order of 10,000 m/s2 (33,000 ft/s2) (1000g).[12]
Some large woodpeckers such asDryocopus 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 genusMelanerpes have distinctive, rowing wing-strokes while the piculets engage in short bursts of rapid direct flight.[16]
Use ofcacti for breeding and roosting holes allows some woodpeckers to live in treeless deserts, such as theladder-backed woodpecker, which uses cacti for nesting.
Woodpeckers have a mostlycosmopolitan distribution, although they are absent fromAustralasia, Madagascar, andAntarctica. They are also absent from some of the world's oceanicislands, although many insular species are found oncontinental islands. The true woodpeckers, subfamilyPicinae, are distributed across the entire range of the family. The Picumninae piculets have a pantropical distribution, with species inSoutheast Asia, Africa, and theNeotropics, with the greatest diversity being in South America.[17] The second piculet subfamily, the Sasiinae, contains theAfrican piculet and two species in the genusSasia that are found in Southeast Asia.[18] The wrynecks (Jynginae) are found exclusively in the Old World, with the two species occurring in Europe, Asia, and Africa.[17]
Most woodpeckers are sedentary, but a few examples ofmigratory species are known, such as therufous-bellied woodpecker,yellow-bellied sapsucker,[17] andEurasian wryneck, which breeds in Europe and west Asia and migrates to theSahel in Africa in the winter.[19] More northerly populations ofLewis's woodpecker,northern flicker, Williamson's sapsucker,red-breasted sapsucker, andred-naped sapsucker all move southwards in the fall in North America.[17] 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 thegrey-capped pygmy woodpecker, which moves to lowlands from hills during winter. The woodpeckers that do migrate, do so during the day.[4]
Overall, woodpeckers are arboreal birds of woodedhabitats. They reach their greatestdiversity intropical rainforests, but occur in almost all suitable habitats, including woodlands,savannahs,scrublands, andbamboo forests. Evengrasslands anddeserts 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, tallcacti are available for nesting.[20] Some are specialists and are associated withconiferous ordeciduous woodlands, or even, like theacorn woodpecker, with individual tree genera (oaks in this case). Other species are generalists and are able to adapt to forest clearance by exploitingsecondary growth, plantations,orchards, and parks. In general, forest-dwelling species need rotting or dead wood on which to forage.[21]
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. Theground woodpecker is one such species, inhabiting the rocky and grassy hills ofSouth Africa,[22] and theAndean flicker is another.[21]
Most woodpeckers livesolitary lives, but their behavior 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 atermite colony or fruit-laden tree, driving away otherconspecifics and returning frequently until the resource is exhausted.Aggressive behaviors include bill pointing and jabbing, head shaking, wing flicking, chasing, drumming, and vocalizations. Ritual actions do not usually result in contact, and birds may "freeze" for a while before they resume their dispute. The colored patches may be flouted, and in some instances, these antagonistic behaviors resemble courtship rituals.[24]
Group-living species tend to be communal group breeders.[24] In addition to these species, a number of species may joinmixed-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.[25] 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; thegrey-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 theMagellanic woodpecker and acorn woodpecker are cooperative roosters.[24]
The sound of a woodpecker (Picidae) tapping, with a few background sounds as well. Species/location unknown, presumably from the continental United States.
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.[26][27] The drumming is mainly a territorial call, equivalent to the song of apasserine.[28] Woodpeckers choose a surface that resonates, such as a hollow tree, and may use man-made structures such as gutters and downpipes.[29] Drumming serves for the mutual recognition of conspecifics and plays a part incourtship rituals. Individual birds are thought to be able to distinguish the drumming of their mates and those of their neighbors.[30] Drumming can be reliably used to distinguish between multiple species in a region, even if those species are phenotypically similar. Cadence (or the mean number of drum beats per second) is heavily conserved within species.[31] Comparative analyses within species between distant geographic populations have shown that cadence is heavily conserved across species' respective ranges, indicating that there likely are not 'dialects' as seen in passerine song.[32] 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.[33] A 2023 study revealed a strong association between extractive foraging and relative brain size across the FamilyPicidae, indicating that a larger brain does not necessarily result in more powerful drumming abilities, but is implicated in foraging behaviors, as the act of sensing and retrieving wood-boring larvae from woody substrates likely requires an increase in sensory and motor control capabilities.[34]
Woodpeckers do not have such a wide range ofsongs and calls as dopasserine 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 incommunication 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 2.5kHz 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.[28] 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.[35] 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 alertsornithologists to the presence of the birds, as they are easily overlooked.[36]
Most woodpecker species feed oninsects and otherinvertebrates 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.[24]Crustaceans,molluscs, and carrion may be eaten by some species, including the great spotted woodpecker, and bird feeders are visited forsuet and domestic scraps.[37]
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 thebuff-spotted woodpecker feeds on and nests intermite mounds. Other species, such as the wrynecks and the Andean flicker, feed wholly or partly on the ground.[24]
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% ofemerald ash borer larvae from individual ash trees.[38]
The ability to excavate allows woodpeckers to obtaintree sap, an important source of food for some species. Most famously, the sapsuckers (genusSphyrapicus) feed in this fashion, but the technique is not restricted to these, and others such as theacorn woodpecker andwhite-headed woodpecker also feed on sap. The technique was once thought to be restricted to the New World, but Old World species, such as theArabian woodpecker and great spotted woodpecker, also feed in this way.[4]
All members of the family Picidaenest 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, thegilded flicker andladder-backed woodpecker excavate holes in cactus, and the Andean flicker and ground woodpecker dig holes in earth banks. Thecampo flicker sometimes chooses termite mounds, the rufous woodpecker prefers to use ants' nests in trees and thebamboo woodpecker specialises in bamboos.[39] Woodpeckers also excavate nest holes in residential and commercial structures and wooden utility poles.[38]
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.[40] 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.[41]
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, thered-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.[42]
Members of Picidae are typically monogamous, with a few species breeding cooperatively and somepolygamy reported in a few others.[43]Polyandry, where a female raises two broods with two separate males, has also been reported in theWest Indian woodpecker.[44] Another unusual social system is that of the acorn woodpecker, which is apolygynandrous 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.[45]
A pair works together to help build the nest,incubate the eggs, and raise theiraltricial 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 fullyfledged 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 theHispaniolan 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 arebrood parasites of woodpeckers.[40]
Black-rumped goldenback (Dinopium benghalense) in Guwahati, India
The Picidae are just one of nine living families in the orderPiciformes. Other members of this group, such as thejacamars,puffbirds,barbets,toucans, andhoneyguides, have traditionally been thought to be closely related to the woodpecker family (true woodpeckers,piculets,wrynecks, andsapsuckers). The clade Pici (woodpeckers, barbets, toucans, and honeyguides) is well supported and shares azygodactyl foot with the Galbuli (puffbirds and jacamars). More recently, severalDNA sequence analyses have confirmed thatPici andGalbuli are sister groups.[46]
The name Picidae for the family was introduced by English zoologistWilliam Elford Leach in a guide to the contents of theBritish Museum published in 1819.[50][51] The phylogeny has been updated according to new knowledge aboutconvergence patterns and evolutionary history.[52][53] Most notably, the relationship of the Picinae genera has been largely clarified, and theAntillean piculet was found to be a surviving offshoot of protowoodpeckers. Genetic analysis supports themonophyly 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 asparaphyletic.[52] Morphological and behavioural characters, in addition to DNA evidence, highlights genusHemicircus as the sister group of all remaining true woodpeckers, besides a sister-group relationship between the true woodpecker tribes Dendropicini and Malarpicini.[54]
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 LateOligocene, about 25million 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 theEarly 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 fossilamber from theDominican Republic, dated to about 25 Mya, however, seems to indicate that the Nesoctitinae were already a distinct lineage by then.[55]
Stepwise adaptations for drilling, tapping, and climbing head first on vertical surfaces have been suggested.[54] 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 reinforcedrhamphotheca, frontal overhang, andprocessus dorsalispterygoidei) evolved in the ancestral lineage of piculets and true woodpeckers. Additional adaptations for drilling and tapping (enlargedcondylus lateralis of thequadrate and fused lower mandible) have evolved in the ancestral lineage of true woodpeckers (Hemicircus excepting). The innerrectrix pairs became stiffened, and thepygostyle lamina was enlarged in the ancestral lineage of true woodpeckers (Hemicircus included), which facilitated climbing head first up tree limbs. GenusHemicircus excepting, the tail feathers were further transformed for specialized support, the pygostyle disc became greatly enlarged, and theectropodactyl toe arrangement evolved. These latter characters may have facilitated enormous increases in body size in some lineages.[54]
Prehistoric representatives of the extant Picidae genera are treated in the genus articles. An enigmatic form based on acoracoid, found inPliocene deposits ofNew Providence inthe Bahamas, has been described asBathoceleus hyphalus and probably also is a woodpecker.[56]
The followingcladogram is based on the comprehensivemolecular phylogenetic study of the woodpeckers published in 2017 together with the list of bird species maintained byFrank Gill,Pamela Rasmussen and David Donsker on behalf of theInternational Ornithological Committee (IOC). TheCuban green woodpecker in themonotypic genusXiphidiopicus was not included in the study.[57][49] 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 wassister to Picinae (as shown below), the other method found that Sasiinae was sister to a clade containing both Picumninae and Picinae.[57]
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.[59] 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.[60]
Woodpeckers also drum on various reverberatory structures on buildings such as gutters, downspouts, chimneys, vents, and aluminium sheeting.[61] Drumming is a less-forceful type of pecking that serves to establish territory and attract mates.[60] Houses withshingles 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 oftongue-and-groove boarding. The birds may also drill holes in houses as they forage for insect larvae and pupae hidden behind the woodwork.[61]
Woodpeckers sometimes cause problems when they raid fruit crops, but their foraging activities are mostly beneficial as they control forest insect pests such as thewoodboring beetles that create galleries behind the bark and can kill trees. They also eat ants, which may be tending sap-sucking pests such asmealybugs, as is the case with the rufous woodpecker in coffee plantations in India.[59] Woodpeckers can serve asindicator 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.[59]
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; ablack 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.[62] The design of protective helmets is another field being influenced by the study of woodpeckers.[62]
One of the accounts of thefounding of Rome, preserved in the work known asOrigo Gentis Romanae (unknown), refers to a legend of a woodpecker bringing food to the boysRomulus and Remus during the time they were abandoned in the wild, thus enabling them to survive and play their part in history.
Woody Woodpecker is an animated character that appeared in theatrical short films produced between 1940 and 1972.
The Pokémon Pikipek was introduced in the seventh generation gamesPokémon Sun and Moon. In addition to being a visual homage to apileated woodpecker, entries in the game'sPokédex encyclopedia describes the smallFlying-type as analogous to its real-world counterpart.[63] Its later forms (called "evolutions" in the series) Trumbeak and Toucannon resemble ahoneyguide andtoucan, respectively, perhaps as a tongue-in-cheek reference to the phylogenetic relationship woodpeckers share with thesePiciformes families.
Theivory-billed woodpecker is classified as critically endangered by theIUCN,[64] and some authorities believe it may already be extinct.
In a global survey of the risk ofextinction faced by the various bird families, woodpeckers were the only bird family to have significantly fewer species at risk than would be expected.[65]
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 fewconservation projects directed primarily at woodpeckers, but they benefit whenever their habitat is conserved.[59] Thered-cockaded woodpecker has been the focus of much conservation effort in the southeastern United States, with artificial cavities being constructed in thelongleaf pines they favour as nesting sites.[66]
Two species of woodpeckers in the Americas are on the verge of extinction: theivory-billed woodpecker, which is classified as critically endangered; and theimperial woodpecker, which is classified as extinct in the wild. Some authorities believe them to be extinct, though possible but disputed ongoing sightings of ivory-billed woodpeckers have been made in the United States[67] and a small population may survive in Cuba.[64] A critically endangered species is theOkinawa 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.[68]
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 andocciput but not in other regions.[69] Along with the longhyoid bone “safety belt” the woodpecker has uneven beak lengths which drastically reduce strains when compared to equal length.[69][70] 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.[71] Others dispute shock-absorption in the head (which reduces the force of pecking) but instead point to adaptations within the brain itself.[72]
Tau protein accumulation is associated withchronic 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.[73] 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.[73] 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.[74]
Straight-line trajectory was theorized to be the reason why woodpeckers do not injure themselves, sincecentripetal forces were the cause ofconcussion, but they do not always peck in straight lines, so they produce and resist centripetal forces.[69] 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.[75] The cranial bone has a high bone mineral density with plate-like structures that are thick with high numbers oftrabeculae 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 isimpulse 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.[76]
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 ofcarbon 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.[77] 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.
^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.
^abDunning, John B. Jr., ed. (2008).CRC Handbook of Avian Body Masses (2nd ed.). CRC Press.ISBN978-1-4200-6444-5.
^abcdefWinkler, Hans & Christie, David A. (2002), "Family Picidae (Woodpeckers)"in del Hoyo, J.; Elliot, A. & Sargatal, J. (editors). (2002).Handbook of the Birds of the World. Volume 7: Jacamars to Woodpeckers. Lynx Edicions.ISBN978-84-87334-37-5
^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.S2CID83528456.
^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.S2CID226025386.
^abcGill, Frank; Donsker, David;Rasmussen, Pamela, eds. (February 2025)."Woodpeckers".IOC World Bird List Version 15.1. International Ornithologists' Union. Retrieved11 March 2025.
^Leach, William Elford (1819)."Eleventh Room".Synopsis of the Contents of the British Museum (15th ed.). London: British Museum. pp. 63–68 [66]. The name of the author is not specified in the document, Leach was the Keeper of Zoology at the time.
^abBenz 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.Bibcode:2006MolPE..40..389B.doi:10.1016/j.ympev.2006.02.021.PMID16635580.
^Fuchs J, Ohlson J, Ericson P, Pasquet E (2006). "Molecular phylogeny and biogeographic history of the piculets (Piciformes: Picumninae)".Journal of Avian Biology.37 (5):487–496.doi:10.1111/j.0908-8857.2006.03768.x.
^Purvis, Andy; Rambaut, Andrew (1995). "Comparative analysis by independent contrasts (CAIC): An Apple Macintosh application for analysing comparative data".Computer Applications in the Biosciences.11 (3):247–251.doi:10.1093/bioinformatics/11.3.247.PMID7583692.
^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.JSTOR3809497.
^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.PMID33439040.S2CID231596453.
^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.
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.Bibcode:2015MolPE..88...28F.doi:10.1016/j.ympev.2015.03.016.PMID25818851.
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.
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.Bibcode:2005MolPE..36..233W.doi:10.1016/j.ympev.2005.03.015.PMID15869887.
_female.jpg)
_female.JPG)
_female.JPG)
