02Winkler_Layout 2 30.11.15 15:51 Seite 7 Phylogeny, biogeography and systematics Hans W INKLER Abstract : After a historical oVerVieW of Woodpecker sYstematics, a short accoUnt of the sYstematic relationships Within the Pici - formes and Picidae is giVen. The honeYgUides (Indicatoridae) are the sister groUp of the Woodpeckers (Picidae). FUrther members of the order are the barbets (Megalaimidae, LYbiidae, Capitonidae), toUcans (Ramphastidae), pUffbirds (BUcconidae) and jaca - mars (GalbUlidae). Three sUbfamilies are cUrrentlY recogniZed, With the Picinae containing the greatest nUmber of species in fiVe tribes. The Picidae do not occUr in the AUstralo-PapUan region. SoUth East Asia, Northern SoUth America and eqUatorial Africa form the diVersitY hotspots of the familY. Woodpecker plUmage characteristics haVe mislead sYstematists repeatedlY. MolecUlar methods haVe UncoVered manY cases of parallel eVolUtion and begin to help to Understand plUmage Variation dUe to habitat Vari - ables and social mimicrY. The neW scoring sYstem Used in the checklist bY del HOYO et al. (2014a) does not accoUnt for these facts. The species list presented here thUs does not complY With all the changes in this neW checklist. Key words : Piciformes, Picidae, phYlogeographY, taXonomY, species delimitation, HBW checklist. A brief history of woodpecker ‘Raptatores’ or birds of preY, ‘Rasores’ or scratching classifications from Aristotle to the birds, ‘CUrsores’ or rUnners, ‘Grallatores’ or Waders, and ‘Natatores’ or sWimming birds. He recogniZed fiVe fam - 20 th century ilies Within the ‘Scansores’: ‘Psittacini’ (parrots), ‘Ser - rati’ (birds With serrated bills, e.g. toUcans or trogons), The beginnings ‘Amphiboli’ (those With an opposable toe, e.g. pUffbirds BRUCE (2003) gaVe a splendid introdUction to the and cUckoos), ‘SagittilingUes’ (birds With arroW shaped fascinating historY of bird classification that also con - tongUes) and the ‘SYndactYli’ (birds With stitched tains a feW references to Woodpeckers. SIBLEY & together toes, kingfishers, jacamars). The WrYnecks and AHLQUIST (1972) and MOORE & M IGLIA 2009 dis - Woodpeckers Were part of the ‘arroW-tongUes’. ClearlY, cUssed the historical deVelopment of the sYstematics of morphological featUres Were important for establishing Woodpeckers and allies in greater detail. similarities and hence classification. SqUabbles aboUt th Aristotle classified all passerines (With the eXcep - priorities and plagiarism characteriZe the earlY 19 cen - tion of sWalloWs and martins) together With Woodpeck - tUrY as Well as attempts to reflect ‘diVine order’ in the ers and other smaller birds, bUt also recogniZed their classification of organisms ( BRUCE 2003). The Wood - special attribUtes, sUch as the hard bill. Modern sYstem - peckers Were groUped together With the passeriform atics began With the Works of Carl LinnaeUs or Linné. birds in a groUp named ‘SYlVicolae’, forest birds, bY Three species of Woodpeckers appear 1740 in the 2 nd VIEILLOT (1816). William SWAINSON (1837) listed edition (‘ Picus niger’ , ‘ P. viridis’ , ‘ P. varius’ ) and Were them after the parrots and before the treecreepers, and presented manY figUres on foot and bill characters. placed betWeen the cUckoos (‘ Cuculus ’ Which also Within the familY Picidae, he recogniZed the Picinae or inclUded the WrYneck!) and treecreepers ( Certhia ) and trUe Woodpeckers, and the BUccoinae into Which he the nUthatch ( Sitta ). Toe arrangement, bill shape and classified the WrYnecks and picUlets, and Bucco and climbing behaVior seem to haVe been the criteria for Oxyrynchus (noW an obsolete name referring to the this placement. He maintained the close relationship Sharpbill Oxyruncus in the SUboscines). It Was GLOGER betWeen Woodpeckers and cUckoos eVen in the 10 th edi - (1834) Who made it clear that Woodpeckers and passer - tion (1758) bY pUtting the latter in the ‘Picae’ Which ine birds shoUld be pUt into different orders becaUse of also contained parrots and hUmmingbirds. ‘ Picus ’ fol - the pronoUnced anatomical differences especiallY in loWed ‘ Jynx ’ (WrYneck) in the seqUence of species. Denisia 36 , the sYrinX, and ‘ Picus’ and Jynx appear betWeen cUckoos zugleich Kataloge des oberösterreichischen ILLIGER (1811) listed seVen orders of birds, ‘Scan - and the Hoopoe ( Upupa epops ), With ‘ Picus’ being Landesmuseums Neue Serie 164 (2015): sores’ or climbing birds, ‘AmbUlatores’ or Walking birds, diVided into tree-liVing and ant-eating Woodpeckers. 7-35 02Winkler_Layout 2 30.11.15 15:51 Seite 8 BY the second half of the 19 th centUrY knoWledge on Woodpeckers, the Middle Spotted Woodpecker far aWaY the anatomY of birds had increased sUbstantiallY, and from the Great Spotted Woodpecker and close to the collectors aroUnd the World sent specimens to the big YelloW-croWned Woodpecker and Dendropicos . HAR - national mUseUms. In a list based on the holdings of the GITT also seems to haVe recogniZed the closer link bird collection of the British MUseUm bY GRAY (1869) betWeen Chrysocolaptes and Campephilus , as Well as jacamars, hoopoes and rollers are still foUnd Within the betWeen Dryocopus (Mulleripicus ) and American repre - Passeres. The Woodpeckers, hoWeVer Were placed bY this sentatiVes of Dryocopus . There Were, of coUrse, also taX - aUthor far aWaY from those groUps in the ‘Scansores’ onomic decisions that one cannot reconcile With the With the parrots and cUckoos, and also the barbets and modern sYstem. toUcans ( GRAY 1870). EXtensiVe anatomical Work The sYstem of birds Used in the 20 th centUrY had seV - spUrred seVeral aUthors to prodUce neW classifications. eral „fathers“ ( BRUCE 2003) of Which one Was Hans GARROD (1874) recogniZed the similarities betWeen the Friedrich GADOW Who presented a detailed accoUnt of Woodpeckers and barbets, bUt had other coraciform the anatomY of birds (1891) and conseqUentlY a sYstem birds, sUch as the pUffbirds, bee-eaters and jacamars still (1893) that Was VerY inflUential With respect to the Within the Passeriformes. HoWeVer, FÜBRINGER (1888) higher classification of birds and inclUded an oVerVieW pUt together the honeYeaters (Indicatoridae), toUcans of the preVioUs attempts on a higher classification of (Ramphastidae) and the Woodpeckers in the Pici. On birds. The Woodpeckers Were pUt into the Coraciiformes the other side of the Atlantic, Elliot COUES , one of the and Pici Which comprised the families GalbUlidae, great American ornithologists of the 19th centUrY, had Capitonidae, Ramphastidae, and Picidae With the sUb - stUdied the anatomY of Woodpeckers qUite thoroUghlY families JYnginae and Picinae. He deViated slightlY from (see BOCK , this VolUme), placed them betWeen cUckoos FÜBRINGER (1888) in the placement of the honeYgUides and parrots, bUt did not consider other relationships Which he considered a sUbfamilY of the barbets. becaUse of his focUs on North American birds ( COUES 1884) . He listed the Pileated Woodpecker right after TWo inflUential projects on the classification of birds the IVorY-billed Woodpecker, bUt did recogniZe the con - shoUld be brieflY mentioned here. One Was the VerY siderable differences in toe proportions and arrange - inflUential list of higher taXa (doWn to the familY leVel) ment betWeen the genera these species represent. He bY F. AleXander WETMORE (e.g. 1930, 1960), and the also groUped the pied Woodpeckers together (inclUding other the checklist bY James L. PETERS Which respected Picoides ), folloWed bY Melanerpes and Sphyrapicus , and all taXonomic leVels doWn to the sUbspecies and serVed finallY Colaptes . as an important reference in the bird collections aroUnd At the tUrn of the 20 th centUrY Richard B SHARPE the World. With regard to the higher classification, Undertook to catalogUe the then alreadY enormoUs bird these lists Were deriVed from GADOW ’S. In WETMORE collection of the British MUseUm, both as aUthor and (1930, 1960) the Woodpeckers and honeYgUides had a editor. The CatalogUe treats the Woodpeckers as part of similar position as in GADOW (1893) . The Peters-list the ‘Picariae’, not in VolUme 19 that deals With the from 1948 (reprinted 1964) split the order Piciformes ‘Scansores and CoccYges’, and hence the honeYgUides, into the sUborders GalbUlae and Pici, With the latter barbets, cUckoos and tUracos, and reserVed VolUme 18 containing onlY one familY, the Picidae. It treated the solelY for the treatment of the Woodpecker familY as part honeYgUides as a familY that together With the barbet of the Scansores bY HARGITT (1890) . This VolUme lists familY (WithoUt the toUcans) comprised the sUperfamilY all the 50 genera recogniZed at that time and Which all Capitonoidea. Three sUbfamilies, the JYnginae (WrY - Were represented at the British MUseUm. Then, the col - necks), PicUmninae (picUlets) and Picinae (trUe Wood - lection held almost 8000 Woodpecker specimens of 385 peckers), constitUted the Woodpecker familY. Like in species and sUbspecies. Using modern names, the list HARGITT (1890) , the Picinae started With Geocolaptes begins With the African GroUnd Woodpecker, immedi - and Colaptes . HoWeVer, fernandinae Which Was pUt into atelY folloWed bY American Colaptes (not inclUding a monotYpic genUs ( Nesoceleus ), Was alreadY part of the CUban fernandinae positioned betWeen Dinopium and groUp of species that todaY comprise the genera Colaptes Celeus in this arrangement of species). Then a miXtUre and Piculus . The pUZZling insUlar species Sapheopipo of EUrasian Picus , Chrysophlegma , one species of noguchii (OkinaWa) and Xiphidiopicus percussu s (CUba) Dinopium