Family of birds (Picidae)
Woodpeckers are part of the bird family Picidae, which also includes the 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 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.
Woodpeckers include the tiny piculets, the smallest of which appears to be the bar-breasted piculet at in length and a weight of . Some of the largest woodpeckers can be more than in length. The largest surviving species is the great slaty woodpecker, which weighs on average and up to , and measures , but the extinct imperial woodpecker, at , and ivory-billed woodpecker, around and , were probably both larger.
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.
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.
Woodpeckers have strong bills that they use for drilling and drumming on trees, and long, sticky tongues for extracting food (insects and larvae). 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. Combined, this anatomy helps the beak absorb mechanical stress. 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.
Many of the foraging, breeding, and signaling behaviors of woodpeckers involve drumming and hammering using their bills. To prevent brain damage from the rapid and repeated powerful impacts, woodpeckers have a number of physical features that protect their brains. 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. 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.
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. During the millisecond before contact with wood, a thickened nictitating membrane closes, protecting the eye from flying debris. 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 (1000 g).
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.
Distribution, habitat, and movements.
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. The second piculet subfamily, the Sasiinae, contains the African piculet and two species in the genus "Sasia" that are found in Southeast Asia. The wrynecks (Jynginae) are found exclusively in the Old World, with the two species occurring in Europe, Asia, and Africa.
Most woodpeckers are sedentary, but a few examples of migratory species are known, such as the rufous-bellied woodpecker, yellow-bellied sapsucker, and Eurasian wryneck, which breeds in Europe and west Asia and migrates to the Sahel in Africa in the winter. 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. 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.
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. 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.
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, and the Andean flicker is another.
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.
Behavior.
Most woodpeckers live solitary 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 a termite colony or fruit-laden tree, driving away other conspecifics 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.
Group-living species tend to be communal group breeders. 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. 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.
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. The drumming is mainly a territorial call, equivalent to the song of a passerine. Woodpeckers choose a surface that resonates, such as a hollow tree, and may use man-made structures such as gutters and downpipes. 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 neighbors. Drumming