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Picinae
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| Woodpeckers | |
|---|---|
| Hispaniolan woodpecker | |
| Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Class: | Aves |
| Order: | Piciformes |
| Family: | Picidae |
| Subfamily: | Picinae Bonaparte 1838 |
| Genera | |
|
Several, see text | |
Picinae containing the true woodpeckers is one of four subfamilies that make up the woodpecker family Picidae. True woodpeckers are found over much of the world, but do not occur in Madagascar or Australasia.
Woodpeckers gained their English name because of the habit of some species of tapping and pecking noisily on tree trunks with their beaks and heads. This is both a means of communication to signal possession of territory to their rivals, and a method of locating and accessing insect larvae found under the bark or in long winding tunnels in the tree or upright log.

Physiology and behaviour
[edit]Some woodpeckers and wrynecks in the order Piciformes have zygodactyl feet, with two toes pointing forward, and two backward. These feet, though adapted for clinging to a vertical surface, can be used for grasping or perching. Several species have only three toes. The woodpecker's long tongue, in many cases as long as the woodpecker itself, can be darted forward to capture insects. The tongue is not attached to the woodpecker's head in the same way as it is in most birds, but instead it curls back up around its skull, which allows it to be so long.
The woodpecker first locates a tunnel by tapping on the trunk with its head. Once a tunnel is found, the woodpecker chisels out wood until it makes an opening into the tunnel. Then it worms its tongue into the tunnel to try to locate the grub. The tongue of the woodpecker is long and ends in a barb. With its tongue the woodpecker skewers the grub and draws it out of the trunk.
Woodpeckers also use their beaks to create larger holes for their nests which are 15–45 cm (6–18 inches) below the opening. These nests are lined only with wood chips and hold 2–8 white eggs. Because the nests are out of sight, they are not visible to predators and eggs do not need to be camouflaged. Cavities created by woodpeckers are also reused as nests by other birds, such as grackles, starlings, some ducks and owls, and mammals, such as tree squirrels.
Several adaptations combine to protect the woodpecker's brain from the substantial pounding that the pecking behaviour causes: it has a relatively thick skull with relatively spongy bone to cushion the brain; there is very little cerebrospinal fluid in its small subarachnoid space; the bird contracts mandibular muscles just before impact, thus transmitting the impact past the brain and allowing its whole body to help absorb the shock; its relatively small brain is less prone to concussion than other animals'.[1]
Some species have modified joints between bones in the skull and upper jaw, as well as muscles which contract to absorb the shock of the hammering. Strong neck and tail-feather muscles, and a chisel-like bill are other hammering adaptations which are seen in most species. Other species of woodpecker, such as the Flicker, uses its long tongue primarily to grab prey from the ground or from under loose bark. It has few shock-absorbing adaptations, and prefers to feed on the ground or to chip away at rotting wood and bark, habits observed in birds outside of the woodpecker family. A "continuum" in skull structures, from little- to highly specialized for pounding is seen in different genera (groups of related species) of woodpeckers alive today. In his classic "Birds of America," John James Audubon describes the slight gradations in hyoid horn length found in different species of living woodpeckers. The slack of tongue is kept under the loose skin behind its neck. The tiny bones divide into essentially two tongues, coming back together before entering the beak.
Systematics
[edit]The systematics of woodpeckers is quite convoluted. Based on an assumption of unrealistically low convergence in details of plumage and behavior, 5 subfamilies were distinguished. However, it has turned out that similar plumage patterns and modes of life are not reliable to determine higher phylogenetic relationships in woodpeckers, and thus only 3 subfamilies should be accepted.
For example, the genera Dryocopus (Eurasia and Americas) and Campephilus (Americas) of large woodpeckers were believed to form a distinct group. However, they are quite unrelated and instead close, respectively, to the genera, Mulleripicus and Chrysocolaptes, of Southeast Asia. In addition, the genus allocation of many species, e.g. the rufous woodpecker, has turned out to be in error, and some taxa with unclear relationships could be placed into the phylogeny.[2][3]
In 1975 John Morony and colleagues in their Reference List of the Birds of the World divided the true woodpeckers into six tribes: Melanerpini, Campetherini, Colaptini, Campephilini, Picini, Meiglyptini.[4] This classification was used in 1982 by Lesley Short in his Woodpeckers of the World.[5] The introduction of molecular methods led to a substantial reorganization of the phylogeny making the earlier groups obsolete. In 2005 David Webb and William Moore divided the main woodpecker genera into three tribes: Megapicini, Malarpicini and Dendropicini.[6] These names were rapidly accepted by other ornithologists but in 2013 Edward Dickinson and Leslie Christidis in the fourth edition of the Howard and Moore Complete Checklist of the Birds of the World rejected these new names on the grounds that the earlier names had precedence. Webb and Moore's Malarpicini, Megapicini and Dendropicini were replaced by Pici, Campephilini and Melanerpini. Dickinson and Christidis also introduced Nesoctitini for the Antillean piculets and Hemicircini for the two species in Hemicircus giving a total of five tribes.[7]
Genera
[edit]
| |||||||||||||||||||||
| Relationships between the genera based on a phylogenetic study by Sabir Shakya and colleagues published in 2017.[8] |
The world bird list maintained by Frank Gill, Pamela Rasmussen and David Donsker on behalf of the International Ornithologists' Union recognises 208 species of true woodpecker which are split up into 33 genera.[9] The division into tribes is based on the 2017 study by Sabir Shakya and colleagues.[8]
Tribe Nesoctitini
- Nesoctites – monotypic: Antillean piculet (Caribbean)
Tribe Hemicircini
- Hemicircus – 2 species (Indomalaya)
Tribe Picini
- Micropternus – monotypic: rufous woodpecker (Indomalaya)
- Meiglyptes – 4 species (Indomalaya)
- Gecinulus – 3 species (Indomalaya)
- Dinopium – 5 species, flamebacks (Indomalaya)
- Picus – 14 species (Indomalaya and Palearctic)
- Chrysophlegma – 3 species (Indomalaya)
- Pardipicus – 2 species (Africa)
- Geocolaptes – monotypic: ground woodpecker (Africa)
- Campethera – 11 species (Africa)
- Mulleripicus – 4 species (Indomalaya)
- Dryocopus – 6 species (Eurasia and Americas)
- Celeus – 12 species (Neotropic)
- Piculus – 7 species (Neotropic)
- Colaptes – 15 species (Americas)
Tribe Campephilini
- Campephilus – 12 species (Americas)
- Blythipicus – 2 species (Indomalaya)
- Reinwardtipicus – monotypic: orange-backed woodpecker (Indomalaya)
- Chrysocolaptes – 9 species, flamebacks (Indomalaya)
Tribe Melanerpini (pied woodpeckers)
- Sphyrapicus – 4 species, sapsuckers (Americas)
- Melanerpes – 23 species (Americas)
- Picoides – 3 species (Holarctic)
- Yungipicus – 7 species (Asia)
- Leiopicus – monotypic: yellow-crowned woodpecker (Indomalaya)
- Dendrocoptes – 3 species (Eurasia)
- Chloropicus – 3 species (Africa)
- Dendropicos – 12 species (Africa)
- Dendrocopos – 12 species (Eurasia)
- Dryobates – 6 species (Americas and Eurasia)[a]
- Leuconotopicus – 6 species (Americas)
- Veniliornis – 14 species (Neotropic)
- Xiphidiopicus – monotypic: Cuban green woodpecker (Caribbean)[b]
Unassigned fossil forms
[edit]- Genus Palaeonerpes (Ogallala 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) - possibly malarpicine
Notes
[edit]- ^ In the eBird/Clements Checklist of Birds of the World which is maintained by the Cornell Lab of Ornithology, the genus Dryobates is expanded to include the species in Leuconotopicus and Veniliornis.[10]
- ^ The Cuban green woodpecker (Xiphidiopicus percussus) was not sampled by Shakya and colleagues in their 2017 study.[8] An earlier study in which DNA sequences of mitochondrial cytochrome-b were compared indicated that the Cuban woodpecker was most closely related to species in the genus Melanerpes.[11]
References
[edit]- ^ Cure for a headache, Ivan R. Schwab, British Journal of Ophthalmology. 86(8):843, August 2002.
- ^ Benz, B.W.; Robbins, M.B.; Peterson, A.T. (2006). "Evolutionary history of woodpeckers and allies (Aves: Picidae): Placing key taxa on the phylogenetic tree". Molecular Phylogenetics and Evolution. 40 (2): 389–399. Bibcode:2006MolPE..40..389B. doi:10.1016/j.ympev.2006.02.021. PMID 16635580.
- ^ Moore, W.S.; Weibel, A.C.; Aguis, A. (2006). "DNA phylogeny of the woodpecker genus Veniliornis (Picidae, Picinae) and related genera implies convergent evolution of plumage patterns". Biological Journal of the Linnean Society. 87 (4): 611–624. doi:10.1111/j.1095-8312.2006.00586.x.
- ^ Morony, John J.; Bock, Walter J.; Farrand, John (1975). Reference List of the Birds of the World. New York: Department of Ornithology, American Museum of Natural History. pp. 66–68. hdl:2246/6700.
- ^ Short, Lester L. (1982). Woodpeckers of the World. Monograph Series 4. Greenville, Delaware: Delaware Museum of Natural History. p. 36. ISBN 0-913176-05-2.
- ^ Webb, David Matthew; Moore, William S. (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): 233–248. Bibcode:2005MolPE..36..233W. doi:10.1016/j.ympev.2005.03.015. PMID 15869887.
- ^ Dickinson, E.C.; Remsen, J.V. Jr., eds. (2013). The Howard & Moore Complete Checklist of the Birds of the World. Vol. 1: Non-passerines (4th ed.). Eastbourne, UK: Aves Press. pp. 298, 307, 309. ISBN 978-0-9568611-0-8.
- ^ a b c 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. Bibcode:2017MolPE.116..182S. doi:10.1016/j.ympev.2017.09.005. PMID 28890006.
- ^ Gill, Frank; Donsker, David; Rasmussen, Pamela, eds. (February 2025). "Woodpeckers". IOC World Bird List Version 15.1. International Ornithologists' Union. Retrieved 23 May 2025.
- ^ Clements, J.F.; Schulenberg, T.S.; Iliff, M.J.; Billerman, S.M.; Fredericks, T.A.; Sullivan, B.L.; Wood, C.L. (2019). "The eBird/Clements Checklist of Birds of the World: v2019". Retrieved 18 May 2020.
- ^ Overton, Lowell C.; Rhoads, Douglas D. (2006). "Molecular phylogenetic relationships of Xiphidiopicus percussus, Melanerpes, and Sphyrapicus (Aves: Picidae) based on cytochrome b sequence". Molecular Phylogenetics and Evolution. 41 (2): 288–294. Bibcode:2006MolPE..41..288O. doi:10.1016/j.ympev.2006.05.014. PMID 16814571.
External links
[edit]- Woodpecker videos on the Internet Bird Collection
- Anatomy and Evolution of the Woodpecker's Tongue
Picinae
View on GrokipediaTaxonomy and Phylogeny
Classification and Definition
Picinae is the largest subfamily within the family Picidae, commonly known as the "true woodpeckers," encompassing approximately 176–200 species distributed across 28–30 genera.[3] It excludes the basal subfamilies Jynginae (wrynecks, 2 species in 1 genus) and Picumninae (piculets, ~29 species in 3 genera), which differ in morphology and behavior, such as the lack of rigid tail support in wrynecks and the smaller size and weaker bills of piculets.[3] The subfamily is characterized by its monophyletic nature, as confirmed by molecular analyses, and represents the core group of woodpeckers adapted for excavating and climbing on trees.[4] The taxonomic framework of Picinae traces back to early 19th-century classifications, with the family Picidae first established by William Elford Leach in 1820 based on morphological similarities among woodpecking birds.[5] Initial subfamily divisions relied on plumage, bill shape, and foot structure, but these were often limited by convergent evolution, leading to unstable groupings in works like those of Short (1982).[3] Modern revisions, driven by molecular phylogenetics, have refined the classification; for instance, Shakya et al. (2017) provided a comprehensive tree using mitochondrial and nuclear DNA from 158 species, resolving major clades like the tribes Picini, Campephilini, and Melanerpini within Picinae.[4] Similarly, Fuchs et al. (2015) restructured the pied woodpecker assemblage (Dendropicini) using multi-locus data, highlighting paraphyly in genera like Dendrocopos and supporting transfers to align with evolutionary relationships. Key diagnostic traits of Picinae include zygodactyl feet (with toes II and III forward, I and IV backward) for enhanced grip on vertical surfaces, stiffened tail feathers that act as a prop during climbing and foraging, and nestlings hatched without natal down feathers, relying instead on sparse contour feathers that develop rapidly.[6][7] These features distinguish true woodpeckers from the softer-tailed wrynecks and less specialized piculets, enabling their specialized arboreal lifestyle.[3] Recent taxonomic updates as of 2025 in the eBird/Clements checklist reflect ongoing refinements, including the 2025 splits within Leuconotopicus (elevating the Hairy Woodpecker complex to three species) and the earlier recognition of Dryobates as a distinct genus split from the former Picoides, with approximately 12 species in Dryobates based on phylogenetic evidence of non-monophyly in the original grouping.[8][4] These changes, informed by molecular data, ensure the classification better mirrors evolutionary history while maintaining stability for conservation and field identification.[9]Evolutionary History and Fossil Record
The subfamily Picinae, comprising the true woodpeckers, originated through divergence from other Picidae subfamilies during the late Eocene to early Oligocene, approximately 35–40 million years ago, as indicated by molecular clock estimates for the broader family's diversification.[10] Phylogenetic analyses using multi-locus data place the crown-group age of Picinae around 25–30 Ma, consistent with the timing of early divergences within the clade. These estimates derive from Bayesian relaxed-clock models calibrated with fossil and biogeographic constraints, revealing an Old World origin for Picinae, likely in Asia, with subsequent dispersals to other regions. Major diversification events occurred during the Miocene, driven by climatic changes and habitat expansion. In the New World, radiation of lineages such as Melanerpes (e.g., red-headed woodpecker group) and Dryocopus (e.g., pileated woodpecker) coincided with the spread of forests across North and South America around 20–10 Ma. Similarly, in the Old World, the expansion of Picus (e.g., green woodpecker lineage) reflects adaptation to Eurasian woodlands during the same period, with diversification rate shifts detected early in Picinae evolution using Bayesian analysis of macroevolutionary dynamics. Basal clades, including Indomalayan genera like Micropternus (rufous woodpecker), highlight early branching in tropical Asia, supported by mitochondrial and nuclear DNA phylogenies. The fossil record of Picinae is sparse and fragmentary, limiting precise phylogenetic placement, with approximately 15 described genera from Miocene to Pleistocene deposits worldwide.[11] The earliest piciform fossils potentially allied to Pici (including Picidae) date to the late Eocene, but definitive Picinae records begin in the Oligocene, such as an isolated tarsometatarsus from Germany (~25 Ma) resembling modern woodpeckers in proportions.[12] Key Miocene examples include Piculoides saulcetensis from France (~22.5 Ma), a stem-group representative possibly linking Picumninae and Picinae based on tarsometatarsal morphology, and unassigned forms like those from the Caribbean Miocene, underscoring incomplete preservation challenges.[11] Convergent evolution of traits like drumming behaviors and specialized tongue structures appears across lineages, as evidenced by morphological comparisons, rather than shared ancestry in basal groups.List of Genera
The subfamily Picinae, comprising the true woodpeckers, includes 28 recognized genera according to the IOC World Bird List version 15.1 (2025), encompassing approximately 200 species distributed across all major biogeographic realms except Antarctica and Australasia (beyond introduced populations). Recent taxonomic updates, such as the 2025 eBird revision splitting the Hairy Woodpecker complex in Leuconotopicus, have refined species-level boundaries within existing genera, increasing the total species count. The following table lists all current genera, with species counts, primary geographic ranges, and brief distinguishing morphological or ecological traits that aid in identification, based on phylogenetic and distributional syntheses.| Genus | Species Count | Geographic Range | Distinguishing Traits |
|---|---|---|---|
| Micropternus | 1 | Indomalaya (Southeast Asia) | Rufous plumage; arboreal but mimics ant nests for camouflage; small size (17-20 cm). |
| Meiglyptes | 5 | Indomalaya (Southeast Asia) | Streaked underparts; short-tailed; inhabits lowland forests; subtle barring on wings. |
| Gecinulus | 3 | Indomalaya (Southeast Asia) | Buff-rumped; pale supercilium; ground-foraging tendencies in bamboo undergrowth. |
| Dinopium | 6 | Asia (Indian subcontinent to Southeast Asia) | Golden-backed in males; ladder-like wing markings; medium-sized (20-25 cm). |
| Chrysocolaptes | 11 | Asia to Australasia (Philippines, Indonesia) | Crimson-rumped; large crest; powerful bill for excavating in dense forests. |
| Picus | 15 | Eurasia and Africa (widespread) | Mottled plumage; long bill; undulating flight; often in open woodlands. |
| Mulleripicus | 3 | Asia (Philippines, Indonesia) | Black-and-white with red crown; large (30-35 cm); island endemics with robust build. |
| Hemicircus | 2 | Asia (Southeast Asia) | Heart-spotted; short crest; inhabits mangroves and coastal forests. |
| Dendropicos | 7 | Sub-Saharan Africa | Small pied woodpeckers; spotted underparts; adaptable to savannas and gardens. |
| Campethera | 12 | Sub-Saharan Africa | Striped back; ground-probing bill; terrestrial foraging in grasslands. |
| Geocolaptes | 1 | Southern Africa | Ground Woodpecker; long legs for walking; sandy-brown plumage for camouflage on earth banks. |
| Nesoctites | 1 | Caribbean (Hispaniola) | Antillean; olive-backed; unique island radiation with weak flight. |
| Melanerpes | 24 | Americas (New World, widespread) | Black-and-white with red head; cavity-nesters in open habitats; vocal mimics. |
| Sphyrapicus | 4 | North America | Sapsuckers; pale yellow bellies; sap-drilling specialists with vertical foraging posture. |
| Xiphidiopicus | 1 | Cuba (Caribbean) | Cuban Green; greenish plumage; slender bill for probing in humid forests. |
| Piculus | 10 | Neotropics (Central/South America) | Green-backed; yellow underparts; ladder-marked wings; canopy dwellers. |
| Celeus | 12 | Neotropics (Central/South America) | Golden-olive; crested or bushy head; large (30-35 cm); fruit-supplemented diet. |
| Dryocopus | 8 | Worldwide (Holarctic, Neotropics) | Black-backed; massive bill; powerful excavators; includes Pileated Woodpecker. |
| Campephilus | 7 | Americas (Neotropics, North America) | Ivory-billed; red crests; double-taps in display; large (35-50 cm) forest giants. |
| Colaptes | 21 | Americas to Eurasia (introduced) | Flickers; ground-foragers; spotted underparts; migratory in some species. |
| Chrysoptilus | 6 | South America (Andes, Patagonia) | Checkered; small to medium; pied patterns; high-elevation specialists. |
| Veniliornis | 16 | Neotropics (South America) | Spotted; ladder-backed; diverse habitats from lowlands to montane forests. |
| Dryobates | 12 | Worldwide (especially Holarctic, Neotropics) | Small piculets to woodpeckers; includes Downy; post-2014 split from Dendrocopos/Picoides. |
| Leuconotopicus | 5 | North America | Ladder-backed; arid-adapted; white underparts with black streaks; 2014 split from Picoides, with 2025 further splits. |
| Dendrocopos | 14 | Eurasia and Africa | Spotted; small; Eurasian focus post-2014 split; includes Great Spotted. |
| Sapheopipo | 1 | Asia (Taiwan) | Taiwan; blue-gray plumage; highland endemic with weak bill. |
| Reinwardtipicus | 1 | Asia (Sulawesi) | Sulawesi; black-and-crimson; island specialist with robust form. |
| Blythipicus | 3 | Asia (Himalayas to Southeast Asia) | Rufous-winged; long bill; montane forest dwellers. |
