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The Generic Distinction of Pied Woodpeckers

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The Generic Distinction of Pied Woodpeckers THE GENERIC DISTINCTION OF PIED WOODPECKERS M. RALPH BROWNING, 170 JacksonCreek Drive, Jacksonville,Oregon 97530 ABSTRACT: The ten speciesof New World four-toedwoodpeckers (scalaris, nuttallii, pubescens, villosus, stricklandi, arizonae, borealis, albolarvatus, lignarius,and m ixtusand the two borealthree-toed species (arcticus and tridactylus), currentlycombined in the genusPicoides, differ, in additionto the numberof toes,in modificationsof the skull,ribs, the belly of the pubo-ischio-femoralismuscle, head plumage,and behavior. I recommendthat the genericname Dryobates be reinstituted for the New World four-toedwoodpeckers. There are three generalmorphological groups of pied woodpeckers,a groupof nine four-toedspecies of the New World, a groupof 22 four-toed speciesof the Old World, and a groupof two three-toedspecies straddling bothregions. ! referto thesegroups of piedwoodpeckers beyond as the New World,Old World,and three-toedgroups. The three-toedspecies have long beenin the genusPicoides Lac•p•de, 1799, but the four-toedgroups have been combinedat the genericlevel in differentways. All four-toedpied woodpeckerswere long includedin the genusDryobates Boie, 1826, later changed to Dendrocopos Koch, 1816 an earlier name (Voous 1947, A.O.U. 1947, Peters 1948). Despite the differencein number of toes, Dendrocoposwas combined with Picoidesbecause of generalsimilarities in anatomy (Delacour 1951, Short 1971a), plumage and behavior (Short 1974a), and vocalizations(Winkler and Short 1978). The A.O.U (1976) followedthis mergerof the genera.On the basisof skeletalcharacters Rea (1983) was skepticalof the merger,but he did not providedetails. On the otherhand, Ouellet(1977), concludingthat the two generadiffer in external morphologyand some behaviors and vocalizations, separated the Old World four-toedwoodpeckers in Dendrocoposand three-toedand New World four-toedwoodpeckers in Picoides.The A.O.U. (1987, 1998), Sibleyand Monroe (1990), and most Europeanauthors (e.g., Hogstad 1978, Cramp 1985, Aulenand Lundberg1991) followedOuellet (1977). More recently, analysisof DNA sequencesof the genesfor cytochromeoxidase I, cyto- chromeb (Weibeland Moore 2002a), and l•-fibrinogenintron 7 (Weibeland Moore 2002b) suggestthat Picoides(sensu Short 1982) consistsof several groupsof species. In discussingthe characterson which he based his classificationof woodpeckers,Short (1982) commentedthat anatomicalstudies are unreli- able becauseof the lack of analysesof variousstructures' functions. Burr (1930), however,had founda positivecorrelation between certain modifica- tionsin the skulland habitsof woodpeckers.Spring (1965) concludedthat the lossof the halluxand a reducedangle of cranialkinesis are probably modificationsfor a woodpecker'speck to deliver a blow of maximum impact. Kirby (1980) concludedthat woodpeckerswith wider ribs are adaptedto more frequentand harderpecking than specieswith narrow ribs and that the pattern of increasingrib width follows that of increasing specializationin the skullin speciescharacterized by Burt (1930). WesternBirds 34:97-107, 2003 97 THE GENERIC DISTINCTION OF PIED WOODPECKERS Burt (1930) discussedas a modificationin the skullof woodpeckersthe meetingof the frontal bone (foreheadof the cranium)and the superior processesof the premaxillary(base of the maxilla),which forms a so-called cranio-facialangle. He qualitativelyranked the three-toedpied woodpeckers as havinga more acuteangle than the four-toedNew World species.The frontalbones of both groups,and of the distinct(Zusi and Marshall1970) genus$phyrapicu$, the sapsuckers,fold towardthe cranium(Burt 1930). Another modification in the skull is the relative distance between the lateralparts of the frontalbone that bulgeanteriorly and the anteriorcenter of the bone (Figure1). Here I summarizethis distanceand other measure- ments of the skull, with other anatomicaland behavioralinformation, to determinedifferences and similaritiesamong pied woodpeckers. METHODS I examinedskeletons including skulls and an associatedfemur of 17 of the 22 speciesof Old World four-toedpied woodpeckers,all speciesof New World four-toedpied woodpeckersexcept the CheckeredWoodpecker (Pi½oid½smixtus) of South America, both speciesof three-toed pied Figure 1. Diagrammaticdorsal view of a woodpeckerskull showing distances mea- sured.A, distancefrom the medialbase of the mostposterior part of the occiputto the anteriorlateral corners of the frontalbone; B, distance,measured medially, from the mostposterior part of the occiputto centerof the anteriormargin of the frontalbone. The craniofacialdistance formula, normalized for the sizeof the bird,is expressedby (A - B)/femur length(100).Two measurementsof widthsof the skull:W, maximum width;O, width at the orbits. 98 THE GENERIC DISTINCTION OF PIED WOODPECKERS woodpeckers(the Three-toed, P. tridactylus, and the Black-backed,P. arcticus),all speciesof Sphyrapicus,and three species of the Africangenus Dendropicos(sensu Short 1982) (Table 1). The last genuswas included becausePicoides and Dendropicos are probably closelyrelated (Short 1971b). On each skullI measuredthe distancealong the midlinebetween the mostanterior point of the frontalbone in itscenter and the mostanterior (fartheranterior) point of the frontalbone at its lateralcorners. I did thisby subtractingthe lengthof the cranium,measured medially from the baseof the occiputto the centerof the mostanterior part of the frontalbone (B in Figure1), fromthe distancefrom the medialbase of the occiputto the lateral cornersof the mostanterior part of the frontalbone (A in Figure1). To avoid possiblebias caused by the species'varying sizes (Kirby 1980, Risingand Somers1989), I dividedA - B by femurlength. Thus, the formula(A - B)/ femur(100)gives a valuethat, for brevity,I referto as facialdistance. The sexeswere combinedbecause the charactersaddressed in the studyare not sexuallydimorphic. I also comparedmaximum skull width (W in Figure 1) anddorsal skull width between the orbits(O in Figure1), dividedby lengthof femur.SYSTAT (Wilkonson 1989) wasused for the statisticalanalyses. For the speciesomitted from this analysis,no skeletonincluding an intactskull andat leastone femurwas available from the numerousmuseums surveyed. RESULTS AND DISCUSSION In the three-toedpied woodpeckers the facialdistance is greaterthan in the four-toedspecies. The averagefacial distance for boththree-toed species is at least2.5 greaterthan in any four-toedspecies, and individualvariation yieldsonly minimal overlap with onlyone of the four-toedspecies, the Old WorldLesser Spotted Woodpecker (Dendrocopos minor) (Tables1 and 2). The skullshapes of the two groupsare distinctvisually; a skullis readily identifiableas that of a three-toedor a four-toedwoodpecker without the facialdistance being measured (Figure 2). Facialdistances of New Worldand Old Worldfour-toed woodpeckers (Table 2), sapsuckers(mean 5.18, range 2.89-7.50, n = 8) and the availablespecies of Dendropicos(mean 3.63, range1.32-6.77, n = 4) are similarto one anotherand do not overlapthose of the three-toedwoodpeckers. The width of the skull(Figure 1) in all pied woodpeckersis similar.The widthof the skullof speciesof Dendropicos(0.84 to 0.95) andSphyrapicus (Figure3) is similarin both generaand narrowerthan in the pied species. The widthof the skullat the orbits(Figure 1) is similarin the four-toedpied woodpeckers.This variable is greater in the three-toed speciesand in Sphyrapicus,whereas in Dendropicosit is narrower than in the other woodpeckersmeasured. Principal-componentsanalysis, based on facial distance,widths, and lengthsof the skull(Figure 1) of the pied woodpeckers,revealed extensive overlapin canonicalfactors 1 and 2 of the Old and New Worldfour-toed species(Figure 4). The three-toedpied woodpeckersand Sphyrapicus, however, differ from each other, and both differ from four-toed pied woodpeckers(Figure 4). 99 THE GENERIC DISTINCTION OF PIED WOODPECKERS Table 1 Cranio-facial Distance of Adult Three- and Four-toed Pied Woodpeckersand Some RelatedSpecies Species n Range Mean+ SD• OldWorld four-toed woodpeckers (Dendrocopos) Sulawesi(temmincki) 1 3.30 Philippine(maculatus) 3 3.16-5.56 4.07 Brown-capped(nanus) 1 4.86 Pygmy(kizuki) 1 5.33 Gray-capped(½anicapillus) 2 3.82-4.69 4.23 LesserSpotted (minor) 5 7.60-9.43 8.50 Fulvous-breasted(macei) 1 3.45 Stripe•breasted(atratus) 2 1.40-2.90 2.10 Yellow-crowned(mahrattensis) 1 7.40 Darjeeling(darjellensis) 1 4.16 Rufous-bellied(hyperythrus) 1 5.80 MiddleSpotted (medius) 3 3.90-4.90 4.40 White-backed(leu½otos) 1 4.09 Himalayan(himalayensis) 1 4.16 Sind(assimilis) 1 5.82 Syrian(syriacus) 2 5.20-6.80 6.00 GreatSpotted (major) 5 3.89-5.33 4.80 New Worldfour-toed species (Dryobates) Striped(lignarius) 1 6.34 Ladder-backed(scalaris) 23 2.23-5.40 4.20 + 1.20 Nuttall's(nuttallii) 6 2.71-5.59 4.70 Downy(pubescens) 22 2.98-7.59 5.10 + 1.30 Red-cockaded(borealis) 10 3.41-6.53 4.60 + 1.81 Strickland's(stricklandi) b 9 2.55-6.01 4.90 Hairy (villosus) 24 2.25-7.05 5.30 + 1.20 White-headed(albolarvatus) 5 4.61-8.48 7.30 Three-toedspecies (Pi½oides) Three-toed(tridactylus) 10 8.60-13.83 11.10 + 1.58 Black-backed(arcticus) 8 10.08-13.25 11.14 Sapsuckers($phyrapicus) Yellow-bellied(varius) 3 7.32-2.89 4.45 Red-naped(nuchalis) 2 7.07-7.50 7.29 Red-breasted(tuber) 1 3.77 Williamson's(thyroideus) 2 3.38-6.37 4.88 Africanpied woodpeckers (Dendropi½os) Golden-crowned(xantholophus) 1 4.00 Bearded(namaquus) 1 1.32 Cardinal(fuscescens) 1 6.77 Gray (goertae) 1 2.42 aStandarddeviation (SD) calculatedfor n > 9. blncludesArizona Woodpecker
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