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Woodpeckers are part of the family Picidae, a group of near-passerine birds that also consist

of piculets, wrynecks, and sapsuckers. 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.
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
beak, producing a reverberatory sound that can be heard at some distance. Some species vary their
diet with fruits, bird's eggs and small animals, and some suck sap. They mostly 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 man 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 just one of eight living families in the order Piciformes, the others
being barbets, toucans, and honeyguides in 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 probably being so.


1General characteristics
2Distribution, habitat and movements
o 2.1Global distribution
o 2.2Habitat requirements
o 3.1Drumming
o 3.2Calls
o 3.3Diet and feeding
o 3.4Breeding
4Systematics and evolution
o 4.1List of genera
5Relationship with humans
o 5.1Status and conservation
o 7.1Bibliography
8Further reading
9External links

General characteristics[edit]
A black-rumped flameback using its tail for support

Woodpeckers range from tiny piculets measuring no more than 7 cm (2.8 in) in length and weighing
7 g (0.25 oz) to large woodpeckers which can be more than 50 cm (20 in) in length. The largest
surviving species is the great slaty woodpecker, which weighs 360563 g (12.719.9 oz), but the
probably extinct imperial woodpecker and ivory-billed woodpecker were both larger.[1]
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 the crown.
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.[2]
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 a tree trunk, 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.[1]
Woodpeckers have strong bills for drilling and drumming on trees, and long sticky tongues for
extracting food.[1] Woodpecker bills are typically longer, sharper and stronger than the bills of piculets
and wrynecks; however, 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 collagenfibers, and a middle layer made of porous bone
which connects the two other layers. Combined, this anatomy helps the beak absorb mechanical
stress.[3] 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 will forage in decaying wood more often than woodpeckers. The long sticky
tongues, which possess bristles, aid these birds in grabbing and extracting insects from deep within
a hole in a tree. It has been reported that the tongue was 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.[4]
Many of the foraging, breeding and signaling behaviors of woodpeckers involve drumming and
hammering using the bill.[5] To prevent brain damage from the rapid and repeated impacts,
woodpeckers have a number of physical features evolved to protect the brain.[6]These include a
relatively small and smooth brain, narrow subdural space, little cerebrospinal fluid (CSF) 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.[6] Computer simulations have shown that 99.7
percent of the energy generated in pecking was stored in the form of strain energy, which was
distributed throughout the bird's body, with only a small remaining fraction of the energy going into
the brain. The pecking also heats up the woodpecker's skull, 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.[7] During
the millisecond before contact with wood, a thickened nictitating membrane closes, protecting the
eye from flying debris.[8] These membranes also prevent the retina from tearing. The 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).[9]
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 Melanerpes genus have distinctive, rowing wing-strokes while the
piculets engage in short bursts of rapid direct flight.[10]

Distribution, habitat and movements[edit]

Use of cacti for breeding and roosting holes allows some woodpeckers to live in treeless deserts, like
the ladder-backed woodpeckerwhich 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. The second piculet subfamily, Nesoctitinae, has a single
member, the Antillean piculet, which is restricted to the Caribbean island of Hispaniola. The
wrynecks (Jynginae) are found exclusively in the Old World, with the two species occurring
in Europe, Asia, and Africa.[11]
The majority of woodpeckers are sedentary but there are a few examples of migratory species such
as the rufous-bellied woodpecker and yellow-bellied sapsucker,[11] and the Eurasian wryneck breeds
in Europe and west Asia and migrates to the Sahel in Africa in the winter.[12] 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.[13] 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 woodpecker, which moves to lowlands from hills during the
winter months. The woodpeckers that do migrate do so during the day.[1]

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.[14] Some are specialists and are associated with coniferous or deciduous woodland 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.[15]
A number of 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,[16] and
the Andean flicker is another.[17]
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 in the period 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.[18]



A woodpecker pecking
into a tree

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The majority of woodpeckers live solitary lives, but the spectrum of behaviour ranges from highly
antisocial species that are aggressive towards their own kind, to species that live in groups. Solitary
species will defend such feeding resources as a termitecolony 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.[19]
Group-living species tend to be communal group breeders.[19] 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.[20] 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.[19]

Drumming is a form of non-vocal communication used by most species of woodpecker and involves
the bill being repeatedly struck on a hard surface with great rapidity. After a pause, the drum roll is
repeated, each species having a pattern which 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. The drumming is mainly a territorial
call, equivalent to the song of a passerine, with male birds drumming more frequently than
females.[21] Woodpeckers choose a surface that resonates, such as a hollow tree, and may use man-
made structures such as gutters and downpipes.[22] 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 that of their neighbours.[23]

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 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.[21] 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.[24] 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.[25]

Diet and feeding[edit]

Holes bored by feeding woodpeckers

The majority of woodpecker species live up to their name and 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 it will be
productive to create a hole.[19]
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.[19]
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 percent of emerald ash borer larvae from individual ash trees.[26]
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. It was once thought that the technique was restricted to the New
World, but Old World species, such as the Arabian woodpecker and great spotted woodpecker, also
feed in this way.[1]

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 woodpeckerexcavate
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.[27] Woodpeckers also excavate nest holes in
residential and commercial structures as well as wooden utility poles.[26]
Woodpeckers and piculets will excavate their own nests, but wrynecks will 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 providing
visual evidence of the site of the nest.[28]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.[29]
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 like swallows and starlings. Woodpeckers
may aggressively harass potential competitors, and also use other strategies to reduce the chance
of being usurped from their nesting site; 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.[30]
Members of Picidae are typically monogamous, with a few species breeding cooperatively and
some polygamy reported in a few species.[31] Polyandry, where a female raises two broods with two
separate males, has also been reported in the West Indian woodpecker.[32] 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.[1] 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 disperse to.[33]
A pair will work together to help build the nest, incubate the eggs and raise their altricial young.
However, in most species the male does most of the nest excavation and takes the night shift while
incubating the eggs. A clutch will usually consist 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 1114 days before they hatch. It then
takes about 1830 days 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 are
reared than is the case with birds that nest in the open. In Africa, several species of honeyguide
are brood parasites of woodpeckers.[34]

Systematics and evolution[edit]

The Picidae are just one of eight 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 (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.[35]
The family Picidae was introduced by the English zoologist William Elford Leach in a guide to the
contents of the British Museumpublished in 1820.[36][37] The phylogeny has been updated according to
new knowledge about convergence patterns and evolutionary history.[38][39] Most notably, the
relationship of the picine genera has been largely clarified, and it was determined that the Antillean
piculet is a surviving offshoot of proto-woodpeckers. Genetic analysis supports the monophyly of
Picidae, which seems to have originated in the Old World, but the geographic origins of the Picinae
is unclear. The Picumninae is returned as paraphyletic.[38]
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 it is hypothesized that they actually 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 (1015 mya), all picids seem to have been small or mid-sized
birds similar to a mixture between a piculet and a wryneck. On the other hand, there exists a feather
enclosed in fossil amber from the Dominican Republic, dated to about 25 mya, which seems to
indicate that the Nesoctitinae were already a distinct lineage by then.[40]
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.[41]