Woodpecker

Taxobox
name = Woodpeckers, piculets and wrynecks

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image_caption = Hispaniolan Woodpecker
regnum = Animalia
phylum = Chordata
classis = Aves
subclassis = Neornithes
infraclassis = Neognathae
superordo = Neoaves
ordo = Piciformes
subordo = Pici
familia = Picidae
familia_authority = Vigors, 1825
subdivision_ranks = Subfamilies
subdivision =
Jynginae - wrynecks

Nesoctitinae - Antillean Piculet

Picinae - woodpeckers

Picumninae - typical piculets

The woodpeckers, piculets and wrynecks are a family, Picidae, of near-passerine birds . Members of this family are found worldwide, except for Australia and New Zealand, Madagascar, and the extreme polar regions. Most species live in forests or woodland habitats, although a few species are known to live in desert areas.

The Picidae are just one of the eight living families in the order Piciformes. Members of the order Piciformes, such as the jacamars, puffbirds, barbets, toucans and honeyguides, have traditionally been thought to be very closely related to the woodpeckers, piculets and wrynecks. More recently, DNA sequence analyses have confirmed this view. [Johansson & Ericson (2003)]

There are about 200 species and about 30 genera in this family. Many species are threatened or endangered due to loss of habitat or habitat fragmentation. Two species of woodpeckers, the Ivory-billed Woodpecker and the Imperial Woodpecker, have been considered extinct for about 30 years (there has been some controversy recently whether these species still exist).

Description

The smallest woodpecker is the Bar-breasted Piculet, at 7 g and 8 cm (3.2 inches). The largest woodpecker was the Imperial Woodpecker, at an average of 58 cm (23 inches) and probably over 600 g (1.3 lb). The Ivory-billed Woodpecker is (or was) slightly smaller at 50 cm (20 inches) and a weight of 500 g (1.1 lb). If both the Ivory-billed and Imperial Woodpeckers are indeed extinct, the largest extant woodpecker is the Great Slaty Woodpecker of Southeast Asia, at about 50 cm (20 inches) and 450 g (1 lb). A number of species exhibit sexual dimorphism in size, bill length and weight. In the piculets it is often the females that are larger, amongt the woodpeckers that show sexual dimorphism it is usually the males that are larger.

Most species possess predominantly white, black and brown, green and red plumage, although many piculets show a certain amount of grey and olive green. In woodpeckers, many species exhibit patches of red and yellow on their heads and bellies, and these bright areas are important in signalling. The dark areas of plumage are often iridescent. Although the sexes of Picidae species tend to look alike, many woodpecker species have more prominent red or yellow head markings in males than in females.

Members of the family Picidae have strong bills for drilling and drumming on trees and long sticky tongues for extracting food.Winkler, 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. ISBN 8487334377] 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 shrp by the pecking action in birds that regularly use it on wood. 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 deep within a hole of a tree.

Many of the foraging, breeding and signalling behaviours of woodpeckers involve drumming and hammering using the bill.Gibson L. (2006) "Woodpecker pecking: how woodpeckers avoid brain injury" "Journal of Zoology" 270: 462–465 doi|10.1111/j.1469-7998.2006.00166.x] In order to prevent brain damage from the rapid and repeated decelerations woodpeckers have evolved a number of adaptations to protect the brain. These include small brain size, the orientation of the brain within the skull (which maximises the area of contact between the brain and the skull) and the short duration of contact. The millisecond before contact with wood a thickened nictitans membrane closes, protecting the eye from flying debris. [Schwab I (2002) "Cure for a headache" "British Journal of Ophthalmology" 86 : 843 doi|10.1136/bjo.86.8.843] The nostrils are also protected; they are often slit-like and have special feathers to cover them. Woodpeckers, piculets and wrynecks all possess zygodactyl feet. Zygodactyl feet consist of four toes, the first and the fourth facing frontward and the second and third facing back. This type of 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 the strong claws and feet woodpeckers have short strong legs, this is typical of birds that regularly forage on trunks. The tails of all woodpeckers except the piculets and wrybills are stiffened , and when perched on vertical surfaces the tail and feet work together to support the bird.

Distribution, habitat and movements

The woodpeckers have a mostly cosmopolitan distribution, although they are absent from Australasia, Madagascar and Antarctica. They are also absent from 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 woodpeckers. The Picumninae piculets have a pantropical distribution, with species in Southeast Asia, Africa and the Neotropics, with South America holding the majority of piculet species. The second piculet subfamily, Nesoctitinae, has a single species, the Antillean Piculet, which is restricted to the Caribbean island of Hispaniola. The wrynecks (Jynginae) have an exclusively Old World distribution, with the two species occurring in Europe, Asia and Africa.

Overall the 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, bamboo forests. Even grasslands and deserts have been colonised by various species. These habitats are more easily occupied where a small number of tree exist, or, in the case of desert species like the Gila Woodpecker, tall cactuses are available for nesting in. A number of species are adapted to spending a portion of their time feeding on the ground, and a very small minority of species 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.

Picidae species can either be sedentary or migratory. Many species are known to stay in the same area year around while others, such as the Eurasian Wryneck and the Yellow-bellied Sapsucker, travel great distances from their breeding grounds to their wintering ground.

Behaviour

Listen|filename=Woodpecker2.ogg|noicon|title=Woodpecker|description=A woodpecker pecking into a tree.|format=Ogg

The woodpeckers range from highly antisocial solitary species which are aggressive to other members of their species to group living species. Group living species tend to be communal group breeders. In addition to these species a number of species may join mixed-species feeding flocks with other insectivorous birds, although they tend to stay at the edges of these groups. Woodpeckers are diurnal, roosting at night inside holes. In most species the roost will become the nest during the breeding season.

Diet and feeding

The diet of these birds consists mainly of insects, such as ants and beetles, nuts, seeds, berries, some fruit and sap. Species may feed generally on all of these, or may specialize on one or two.

Breeding

All members of the family Picidae nest in cavities.Almost every species nests in tree cavities, although in deserts some species nest inside holes in cactus and a few species nest in holes dug into the earth. Woodpeckers and piculets will excavate their own nests, but wrynecks will not. The excavated nest is usually only lined from the wood chips produced as the hole was made. Many species of woodpeckers excavate one hole per breeding season, sometimes after multiple attempts. It takes around a month to finish the job. Abandoned holes are used by many other birds and mammals which are secondary cavity nesters. [Kotaka N & S Matsuoka (2002) " [http://www.jstage.jst.go.jp/article/osj/1/2/117/_pdf 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]

Members of Picidae are typically monogamous. 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 nest will usually consist of 2-5 round white eggs. Since these birds are cavity nesters their eggs do not need to be camouflaged and the white colour helps the parents to see them in dim light. The eggs are incubated for about 11-14 days before the chicks are born. It takes about 18-30 days before the young are ready to leave the nest.

ystematics and evolution

The phylogeny has been updated according to new knowledge about convergence patterns and evolutionary history. [Benz "et al." (2006), Moore "et al." (2006)] 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.

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 (10-15 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. [Grimaldi & Case (1995)]

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, Bahamas, has been described as "Bathoceleus hyphalus" and probably also is a woodpecker. [Cracraft & Morony (1969)]

List of genera

FAMILY: PICIDAE
* Basal
** Genus: "Palaeopicus" (Late Oligocene of France)
* "Incertae sedis"
** Picidae gen. et sp. indet. (Middle Miocene of New Mexico, USA)
** Picidae gen. et sp. indet. (Late Miocene of Gargano Peninsula, Italy)
* Subfamily: Jynginae - Wrynecks
** Genus: "Jynx" (2 species)
* Subfamily: Picumninae - Typical piculets
** Genus: "Picumnus" - American Piculets (c.27 species)
** Genus: "Verreauxia" - African Piculet (sometimes included in "Sasia")
** Genus: "Sasia" - Asian Piculets (2 species)
* Subfamily: Nesoctitinae
** Genus "Nesoctites" - Antillean Piculet
* Subfamily: Picinae - Woodpeckers
** "Incertae sedis"
*** Genus: "Palaeonerpes" (Ogalalla Early Pliocene of Hitchcock County, USA) - possibly dendropicine
*** Genus: "Pliopicus" (Early Pliocene of Kansas, USA) - possibly dendropicine
*** cf. "Colaptes" DMNH 1262 (Early Pliocene of Ainsworth, USA) - malarpicine?
** Tribe: Dendropicini
*** Genus: "Melanerpes" (some 22 species)
*** Genus: "Sphyrapicus" - sapsuckers (4 species)
*** Genus: "Xiphidiopicus" - Cuban Woodpecker (Placement in Dendropicini tentative)
*** Genus: "Dendropicos" (15 species)
*** Genus: "Dendrocopos" (21 species)
*** Genus: "Picoides" (presently 12 species; maybe only 3 belong here) - this genus is in need of revision [Moore "et al." (2006)] . See the genus article for more.
*** Genus: "Veniliornis" (14 species)
** Tribe: Malarpicini
*** Genus: "Campethera" (12 species)
*** Genus: "Geocolaptes" - Ground Woodpecker
*** Genus: "Dinopium" - malarpicine flamebacks (4 species)
*** Genus: "Meiglyptes" (3 species)
*** Genus: "Hemicircus" (2 species; placement in Malarpicini tentative)
*** Genus: "Micropternus" - Rufous Woodpecker (formerly in "Celeus")
** Tribe: Picini
*** Genus: "Picus" (c.15 species)
*** Genus: "Mulleripicus" (3 species)
*** Genus: "Dryocopus" (7 species)
*** Genus: "Celeus" (11 species)
*** Genus: "Piculus" (7 species)
*** Genus: "Colaptes" - flickers (about one dozen species)
** Tribe: Megapicini
*** Genus: "Campephilus" (11 species, 2 possibly recently extinct)
*** Genus: "Chrysocolaptes" - megapicine flamebacks (2 species)
*** Genus: "Reinwardtipicus" - Orange-backed Woodpecker
*** Genus: "Blythipicus" (2 species)
*** Genus: "Gecinulus" (2 species; placement in Megapicini tentative)
*** Genus: "Sapheopipo" - Okinawa Woodpecker (Placement in Megapicini tentative)

Footnotes

References

* (2006): Evolutionary history of woodpeckers and allies (Aves: Picidae): Placing key taxa on the phylogenetic tree. "Mol. Phylogenet. Evol." 40(2): 389–399. DOI|10.1016/j.ympev.2006.02.021 (HTML abstract)
* (1969): A new Pliocene woodpecker, with comments on the fossil Picidae. "American Museum Novitates" 2400: 1-8. [http://digitallibrary.amnh.org/dspace/bitstream/2246/2610/1/N2400.pdf PDF fulltext]
* (1995): A feather in amber from the Upper Cretaceous of New Jersey. "American Museum Novitates" 3126: 1-6. [http://digitallibrary.amnh.org/dspace/bitstream/2246/3571/1/N3126.pdf PDF fulltext]
* (2003): Molecular support for a sister group relationship between Pici and Galbulae (Piciformes sensu Wetmore 1960). "J. Avian Biol." 34(2): 185-197. doi|10.1034/j.1600-048X.2003.03103.x [http://www.nrm.se/download/18.4e32c81078a8d9249800021325/Johansson%2520&%2520Ericson%2520-%2520Piciformes%5B1%5D.pdf PDF fulltext]
* (1999): Oaks, acorns, and the geographical ecology of acorn woodpeckers. "J. Biogeogr." 26(1): 159-165. DOI|10.1046/j.1365-2699.1999.00256.x (HTML abstract)
* (2005): Foraging patterns of pileated woodpeckers in a managed Acadian forest: a resource selection function. "Can. J. Forest Res." 35(10): 2387-2393. doi|10.1139/x05-148 (HTML abstract)
* (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 (HTML abstract)
* (2006): Mitochondrial DNA phylogeny of the woodpecker genus "Veniliornis" (Picidae, Picinae) and related genera implies convergent evolution of plumage patterns. "Biol. J. Linn. Soc." 87(4): 611–624. doi|10.1111/j.1095-8312.2006.00586.x [http://bio.wayne.edu/profhtml/moore/PUBLICATIONS/MooreEtal2006Veniliornis.pdf PDF fulltext]
* (1998): A quantitative analysis of woodpecker drumming. "Condor" 100(2): 350-356. [http://elibrary.unm.edu/sora/Condor/files/DJVU/v100n02/P0350-P0356.djvu DjVu fulltext] [http://elibrary.unm.edu/sora/Condor/files/issues/v100n02/p0350-p0356.pdf PDF fulltext]
* (2004). How do woodpeckers extract grubs with their tongues? A study of the Guadeloupe woodpecker ("Melanerpes herminieri") in the French Indies. "Auk" 121: 509-514. DOI:10.1642/0004-8038(2004)121 [0509:HDWEGW] 2.0.CO;2 [http://www.bioone.org/perlserv/?request=get-abstract&doi=10.1642%2F0004-8038%282004%29121%5B0509%3AHDWEGW%5D2.0.CO%3B2 HTML abstract]
* (2005): A phylogenetic analysis of woodpeckers and their allies using 12S, Cyt "b", and COI nucleotide sequences (class Aves; order Piciformes). "Mol. Phylogenet. Evol." 36(2): 233-248. doi|10.1016/j.ympev.2005.03.015 [http://www.biosci.wayne.edu/profhtml/moore/PUBLICATIONS/Webb&Moore2005.pdf PDF fulltext]
* (2001): Clutch size relative to tree cavity size in northern flickers. "J. Avian Biol." 32(2): 167. DOI|10.1034/j.1600-048X.2001.320210.x (HTML abstract)
* (2000): Parental care and social mating system in the lesser spotted woodpecker "Dendrocopos minor". "J. Avian Biol." 31(4): 447. DOI|10.1034/j.1600-048X.2000.310003.x (HTML abstract)
* (1993): Incubation and fledging durations of woodpeckers. "Condor" 95(2): 282-287. [http://elibrary.unm.edu/sora/Condor/files/DJVU/v095n02/P0282-P0287.djvu DjVu fulltext] [http://elibrary.unm.edu/sora/Condor/files/issues/v095n02/p0282-p0287.pdf PDF fulltext]