The Auk 113(1):94-104, 1996

WING SHAPE IN INSECTIVOROUS INHABITING NEW GUINEA AND AUSTRALIAN RAIN FORESTS AND EUCALYPT FOREST/EUCALYPT WOODLANDS

ALLEN KEAST Departmentof Biology,Queen's University, Kingston, Ontario K7L 3N6, Canada

ABSTRACT.--Basedon museum skins of 47 speciesin 17 genera, I evaluated the sup- positionsfor membersof varioustaxonomic groups in New Guineaand Australianrain forest comparedto open eucalypt forest/eucalyptwoodland that in rain forest comparedto eucalyptforest/eucalypt woodland counterparts in eachgroup: (1) outer primary 9 tendsto be proportionatelyshorter; (2) secondaryI is proportionally longer; and (3) the longest primary is more proximally placed relative to the body in its series.The postulationsare largely confirmed. The results are presagedby the findings of earlier writers that operatingin confinedspaces should have broad wings with rounded tips, with low wing loading conferringhigh manoeuvrability,and that distanceflyers shouldhave wings with long wingspansand pointed tips for economyof effort. ReceivedI April 1993, accepted4 September1994.

WHEN MEASURING SKINS of New Guinea and chord.These features reduce drag and conserve Australian tropical rain-forest insectivorous power. passerines, and eucalypt forest/woodland Laterstudents of bird flight (Pennycuick1969, counterparts,I found regularly occurring dif- Norberg 1979,1981, 1990,Norberg and Raynet ferencesin wing form. The former tended to 1987, Raynet 1988, Thollesson and Norberg have more rounded wings, with the outer pri- 1991) have explored wing design in the more mary 9 relatively short, while the longest pri- meaningful framework of wingspan, wing mary in the serieswas placedmore proximally loading, and aspectratio. They have demon- than the othersin its seriesrelative to the body strated that slow-flying speciesoperating in in the rain-forestspecies. Examples were found confinedspaces should have broad wings with in foliage-gleaningwarblers (Gerygone),whis- roundedtips and a low wing loading to confer tlers (Pachycephala),ground-feeding robins, and high manoeuvrability. Long-distance-flying severalother groups,suggesting that a general migrants,by contrast,should have a long wing- habitat adaptation was operating, as has been spanand high-aspect-ratiowings with pointed shown for several body structuresby Winkler tips to reduce profile power and confer econ- and Leisler(1992). In this paper,I quantitatively omy of effort (see detailed review in Bairlein exploredthis observationand postulatedcon- 1992). Various factors are likely to influence nection in a diverse set of ecomorphological wing shapein birds, including: (1) habitat; (2) types of birds. feeding zone utilized (e.g. thicketscompared to An early paper by Savile (1957) documented exposedbranches); (3) way of life (being aerial the occurrenceof two alternative wing types in feedervs. undergrowth skulker);(4) relative de- small birds--"elliptical" wings found in birds velopment of migratory habits;and (5) innate that lived in confinedspaces, and "high-speed" phylogeneticconstraints. wings characteristicof speciesthat remain in All of the species I have evaluated can be the air for long periods. In the former group allocated readily to a precise habitat category the wing is curved to increasenegative pressure usingdata for New Guinea birds from Rand and and, hence,lift above,with slotting of the tips Gilliard (1967), Diamond (1972), Beehler et al. servingto minimize turbulenceand ensureuni- (1986), and for Australian birds from Keast form pressureover the entire surface.The result (1961), Hall (1974), Pizzey (1980), and Blakers is high lift, as well as good control and man- et al. (1984). Quantitative dataon feeding zones oeuvrability. By contrast,the high-speedwing and feeding habitsof eachspecies are not avail- has a lower camber,a slender tapering tip, a able, but general information on these charac- sweepbackof the leading edge, and a short teristics are known (see above references). The 94 January1996] WingShape in InsectivorusPasserines 95 need for the former is less critical if the taxo- The hypotheseswere evaluatedby comparingthe nomicgroups are considered indi¾idually, since groupedrain-forest, and groupedeucalypt forest/eu- in general,members of eachshare common eco- calyptwoodland membersof each taxonomicgroup morphologiesand ways of life. Seasonalmove- usingthe Mann-Whitney U-test(0.05 level of signif- icance; Zar 1984). ments are not a major factor influencing wing The vegetation featuresof the contrastinghabitat shapein thesebird species.Only three (the Aus- typesare summarizedin Table 2 (Specht1981, Webb tralianPachycephala rufiventris, Gerygone olivacea, and Tracey 1981).Tropical rain foresthas high struc- and G. fusca)are partial migrants;all other spe- tural diversity and compactcover-providing vegeta- cies are residents. It is valid, therefore, to ex- tion. The Australianeucalypt forests are classified into: plore habitat as the major variable influencing (a) tall open forest;and (b) woodland and dry forest. wing form. In bothof the latter,the treesand branchesare widely In this paper, I test the hypothesesthat con- spaced.Vines, epiphytes, and thicketsare lacking.In generic and counterpart species inhabiting woodland,the treesare more widely spacedthan in tropical rain forest and thickets, compared to tall forest.The realleeand mangrovehabitats, to which Pachycephalainornata and P. lanioidesare confined, have those of eucalypt forest/eucalypt woodland, the general structural attributesof woodland (Table should have: (1) a relatively short primary 9 to 2). give a more rounded tip to wing; (2) a conspic- uous,if small, primary 10 (seeAverill 1925);(3) secondariesthat are relatively long to maximize RESULTS lift in the proximal part of the wing; and (4) the longestprimary in the primary seriesplaced The basicmeasurements of the 47 speciesare more proximal than the others to the body. The presentedin the Appendix. The Appendix and degreeto which a featureis developed,of course, Table 1 give the longest primary or primaries will vary with closenessof its link to specific (on average)for eachspecies, and the ratiosof structural features of the alternative habitats. primary 9, primary 10, and secondary1 to lon- gestprimary. Standarddeviations are inappro- priate for ratios (Atchley et al. 1976). In Table MATERIALS AND METHODS 1, the habitat and feeding zone of each species alsoare given. Figures1 and 2 showwing shape The studycovers 47 speciesin 17genera (Appendix, Table 1), and includes: foliage-gleaning warblers for a seriesof contrastingspecies in diagram- (Gerygone);robinlike birds that are mostly ground matic form. The wings of tree-creepersand nut- feeders; whistlers (Pachycephala)that are foliage hatches, which do not differ between habitats, gleanersand foliagepluckers; aerial-sallying flycatch- are not illustrated. ers (Rhipidura);and representativesof four groupsof Gerygone (Acanthizidae).--Inthis group of specialistspecies (Table t). These are the: thicket- small-bodiedfoliage-gleaning warblers (habits dwelling whipbirds (Androphobus,Psophodes, Sphen- documented in Keast and Recher [unpubl. ostorna);trunk- and bark-feedingshrike-tits (Eulaces- manuscript]),primary 6 is marginally the lon- torna,Falcunculus); trunk- and branch-feedingtree- gest primary. In the Australian rain-forest G. creepersand nuthatches(Clirnacteris, Daphoenositta); mouki, the more inner primary 5 averagesslight- and ground-feedingmagpie-larks (Grallina). Measurementswere made of the wings of six adult ly longer; in G. olivaceaand G. fusca,both of males of each speciesfrom museum skins. Starting which are partial migrants found in eucalypt with the outermost primary (i.e. primary t0), the woodlands it is the more distal primary 7 or length of eachfeather was measured by insertingfine primary 6 that usually is the longest (see Fig. dividers between adjacent feathers from point of 1). Primary9 is relatively longest(83.4% of lon- emergenceof each from the skin and measuring to gest primary) in G. olivacea;in G. fusca,it is the feather tip. The primaries were measuredto pri- 88.9%,and in G. ruficollis80.4%. Although the mary 5 to contour the outer half of the wing. The first latter is a rain-forestinhabitant, it occupiesthe two secondaries(i.e. the distal two in secondaryse- upper stratumand outer edge (pers.obs. made ries) were similarly measured.The length of longest at Tari, Western Highlands). Primary 10 shows primary in eachindividual wasdetermined. Ratios of little variation. The secondaries are shortest rel- the lengthsof the primary t0 and primary 9 to longest primary provided a measure of relative degree of ative to the longest primary in G. fuscaand G. pointednessof the wing. Ratiosof secondariest and olivacea(Fig. 1). Thus, in Gerygone,trends are in 2 to the longestprimary provided an index of breadth the direction hypothesized. of the proximal sectionof the wing. Robins(Petroicidae).--In the 10 species (six 96 ALLENKEAST [Auk,Vol. 113 January1996] WingShape inInsectivorus Passerines 97 98 ALLENKEAST [Auk, Vol. 113

TABLE2. Basicstructural features of major New Guinea and Australianbird habitats(derived from Specht [1981] and Webb and Tracey [1981]).

Tropicalrain forest.--Dense evergreen closed forest with upper stratumcover of over 70%;several dense intergradingsublayers that include vines, epiphytes,and tree-ferns;often densethickets along edge;com- monly with bare floor with rotting logs, ferns, leaves,mesophyll and notophyll. Eucalypttall openforest.--Open evergreen forest with upperstratum cover 50-70%; height variable; trees spaced;open branches;medium to denselow-tree or tall-shrubsubstratum; variably denselow-shrub cov- er. Eucalyptwoodland and dry forest.--Evergreen; trees widely spaced;foliage projective cover of upperstra- tum cover,30-70% in open forestand 10-30%in woodland;height 10-30 m; medium-dense;small-tree and tall-shrub layer sparse;leaf size notophyll; low shrubssparse; understory grassy layer. Mallee openshrub.--Evergreen; spare and open, stuntedsmall-tree and tall-shrubstratum; medium dense; foliage projectivecover 30-70%; ground coverincomplete and of sparsehummock grass clumps. Mangroves.--Evergreenestuarine formation varying structurallyfrom closedforest to woodland and tall shrubland;branches sparse; substratum absent; intertidal.

genera)of robins evaluated,primaries 6 and 5 robins the greatest ratio of primary 9 to the characteristicallyare the longest, but in the longestprimary exceptfor the Peneothellocy- ground-feedingAustralian woodland anus, which is characteristic of rain forest. Pe- goodenoviiand Melanodryascucullata, primary 7 troicagoodenovil, cucullata and Pe- and 6 are the longest(Appendix, Fig. 1). These troica multicolor, as well as the Australian rain- speciesand the Australian dry-forest-dwelling forest P. rosea,have relatively the shortestpri- Petroicamulticolor have relative to this group of mary 10. The first secondaryis relatively the

GERYGONE- WARBLERS ROBINS Geq/gone Gerygone Gerygone Peneothello Eopsaltr• Melanod•as rnouki ruficollis oliva cea sigillatus australis cucullata WING P-10 133 1 42 [ 29 P-9 P-8

P-7 P-6 P-5

S-1

S-2

40 80 80 lOO 40 60 80 100 40 80 80 lOO 40 60 80 lOO 40 60 80 lOO 40 80 80 lOO % % % % % %

WHISTLERS RHIPIDURAFLYCATCHERS Pachycephala Pachycephala Pachycephala Rhipidura Rhipidura Rhipidura rufinucha olivacea WING rufiventris brachyryncha fuliginosa leucophrys P-10 144 28 ß 40 P-9 •1•71 P-8

P-7 P-6 P-5

S-1 m 77 S-2

40 80 80 100 40 80 80 100 40 60 80 100 40 60 80 100 40 60 80 100 % % % % % Percentlength of longestprimary (*) Percentlength of longestprimary (*) Fig. 1. Wing shapebased on lengthsof individual primary and secondaryfeathers for representative speciesof Gerygonewarblers, robins, Pachycephala, and Rhipidurafantails. Longest primary (on average)for eachspecies indicated by asterisk.Figures beside primary 10,primary 9, and secondary1 are percentagesof longestprimary in series.Data basedon averagesof six adult malesfor eachspecies. January1996] WingShape in InsectivorusPassetines 99 shortest in the first three species and in the others,the longestis primary 6. Andophobushas Australian eucalypt-forestEopsaltria australis. relatively the shortestprimary 9. Sphenostoma Thus,for the robinlike birds,the hypotheses has the longestprimary 10 and shortestsec- are largely supported,although in relative size ondary 1. Wing shapein the three is relatively of primary 10 in Petroicarosea groups with those similar (Fig. 2). of the other Australianspecies. Feeding data Shrike-tits().--Feeding in the that might suggestwhy Peneothellocyanus has eucalypt-forestFalcunculus is largely from the a long primary 9 are lacking. branchesand hangingbark (Recherand Holmes Pachycephala(Pachycephalidae).--In the 10 1985, Holmes and Recher 1986). The rain-forest membersof this ,which variouslyfeed Eulacestomafeeds in vines and thickets, com- byfoliage gleaning and plucking prey from the monly hanging its head downwards (Beehler leaves(see data on P. rufiventrisand A. pectoralis et al. 1986).Eulacestoma has the shorterprimary in Recherand Holmes 1985, Recher et al. 1985), 9 and longer secondary1 (Fig. 2). or largely by the former method (my observa- Tree-creepersand nuthatches.--Tree-creepers tions on P. rufinucha),the longest primary is (Climacteris,Climacteridae) and nuthatches (Da- characteristicallyprimary 6 or primary 5. The phoenositta,Neosittiae) are unrelated trunk and exception is the partly-migratory Australian- branchfeeders of the upper stratum.There was woodlandP. rufiventris,where the outerprimary little differencein wing form in the two pairs 7 is longest.Primary 9 is proportionatelylong of New Guinea and Australian counterparts in this speciesand in the Australian mangrove- considered here. However, in the Australian- and mallee-inhabitingP. lanioidesand P. inor- woodland Climacterispicumnus, primary 7 is nata.It is alsolong in the Australian rain-forest/ marginally the longestprimary, the primary 9 eucalypt-forestP. pectoralis,and its close New to longest primary ratio averages94% (90-97), Guinea rain-forest relatives, P. sororand P. aurea. and ratio of secondary1 to the longestprimary Primary 10 tends to be shortestin P. rufiventris averages79%. and P. pectoralis.The secondariesare shortestin Magpie-larks(Grallinidae).--Comparison of the P. rufiventrisand P. inornata,and longest in the wingsof the New Guineamountain-stream in- thicket-dwelling P. rufinuchaand P. olivacea habiting Grallinabruijni and larger-bodiedAus- (habitat data in Beehleret al. [1986],plus pers. tralian open-countryG. cyanoleucashow that the obs.). former has a more rounded wing and longer Pachycephalarufiventris, the only true open- secondaries(Fig. 2). country dweller, standsapart from the others. Statisticalanalysis.--Are the seeming differ- The contrastingwing shaperelative to thoseof encesbetween the counterpartsin rain forest P. rufinuchaand P. olivaceaare shownin Figure1. and eucalypt forest/eucalypt woodland statis- Rhipidura (Rhipiduridae).--These insectivo- tically significant?Table 3 comparesthem. Ra- rousfantail flycatchersfeed mainly by sallying tios of primary 9 to the longest primary are into the air for prey. The feeding habitsof the significantlydifferent in six of the eight groups various speciesare closely similar (pers. obs., (exceptionsare whipbirds and tree-creepers/ Diamond 1972, Cameron 1985, Beehler et al nuthatches).Ratios of secondary1 to the longest 1986). primary are significantly different in seven Primaries6 and 5 are the longest.In the unique groups(but not in tree-creepersand nuthatch- large-bodiedopen-terrain ground-feedingR. es). In each case, the differences lie in the di- leucophrys,the ratio of the primary 9 to longest rectionhypothesized. There is someintragroup primary ratio is 81.3%,compared to 64.9 to 71.7% variation.Differences in body sizesbeing eval- in the others.Rhipidura leucophrys also has a long uated must, of course, be borne in mind when primary 10 and somewhat short outer second- weighing the validity of the statistics.The hy- aries.Rhipidura rufiventris has a shortprimary 10 pothesisthat the diminutive primary 10 should and the shortestsecondary in the series. be smallerin more activelyflying birds(Averill Whipbirds(Cinclosomatidae).--The relictual 1925) is not confirmed. New Guinea Androphobusand Australian Pso- phodeslive in densethickets near the ground, DISCUSSION while the Australian interior Sphenostomain- habits desert shrubbery.The longest primary The predictionsare supportedfor mostof the in Androphobusis, marginally, primary 5; in the taxonomicgroups. Wing shapesof rain-forest 100 ALLENKEAST [Auk, Vol. 113

WHIPBIRDS Androphobus Psophodes Sphenostoma viridis olivaceus occidentalis WING P-10 41 1 43 / 5o P-9 68 P-8 P-7 P-6 P-5

S-1 S-2

40 60 80 lOO 40 60 80 lOO 40 60 80 lOO % % % SHRIKE - TITS MAGPIE- LARKS Eulacestoma Falcunculus Grallina Grallina frontalis WING nigropectus brui/ni cyanoleuca P-10 1 42 BIB145 P-9 P-8 P-7 P-6 P-5

S-1 • 88 • 72 S-2

40 60 80 lOO 40 60 80 100 40 60 80 100 40 60 80 100 % % % % Percentlength of longestprimary (*) Fig. 2. Wing shapebased on lengthsof individualprimary and secondaryfeathers for representative speciesof whipbirds(Androphobus, Psophodes, Sphenostoma), shrike-tits (Eulacestoma and Falcunculus),and mag- pie-larks(Grallina). Longest primary (on average)for eachspecies indicated by asterisk.Figures beside primary 10, primary 9, and secondary1 are percentagesof longestprimary in series.Data basedon averagesof six adult males for each species. forms compared to congenersinhabiting eu- shapein this group.Note, for example,that the calypt forest/eucalyptwoodland tend to be dif- relative length of primary 9 in the woodland ferent. However, New Guinea and Australian R. rufiventrisis similar to that of the rain-forest counterpartsin the trunk-and-branch-feeding species. Climacterisand Daphoenosittaare no different. Although I have identified and shown the All Daphoenosittahave long pointed wings, longestprimary for eachspecies, it often differs which might be associatedwith high mobility from adjacentones by as little as 1 to 2 mm and life in the upper branches. The forms of (Appendix). Often in a particular individual, Climacterismeasured represent a conspecificpair two or three feathers are nearly equally long. (C. leucophaea).Wing form in Rhipidura,except When averagesare struck,one emergesas the in the large-bodiedR. leucophrys,shows little longest.For presentpurposes minor differences variation; most are rain-forest dwellers. Possi- in lengths of individual primates serve mainly bly, the needs of aerial feeding dictateswing to identify trends.Differences in relativelengths January1996] WingShape in InsectivorusPasserines 101

T^nLœ3. Resultsof Mann-WhitneyU-tests for differencesin proportionatelengths relative to longestprimary of primary 10, primary 9, and secondary1. Comparisonsof speciesin eight taxonomicgroups in tropical rain forest(RF) vs. open eucalyptforest (EF) and eucalyptwoodland (EW). U representsranking done from low to high, U' the reverse.

No. species compareda Primary 10 Primary 9 Secondary! RF EF/EW U U' P U U' P U U' P Gerygonewarblers (foliage gleaners) 4 2 !28.0 !60.0 ns 27.5 260.5 *** 3.5 284.5 *** Robins (ground gleanersand aerial snatchers) 7 3 85.0 67!.0 *** !3!.0 625.0 *** !6!.5 594.5 *** Pachycephala(test 1; foliage gleaners) 9 ! !!.0 3!3.0 *** 29.0 295.0 *** !5.5 308.5 *** Pachycephala(test 2; foliage gleaners) 7 2 5!.0 453.0 *** 82.0 422.0 *** !25.0 379.0 ** Rhipidura fantails (aerial sallyers) 8 2 273.5 302.5 ns !!2.0 392.0 ** 68.5 4!!.5 *** Whipbirds (thicket dwellers) 2 ! 0.0 72.0 *** 17.0 55.0 ns !4.0 58.0 * Shrike-tits (trunk and bark feeders) ! ! !6.0 20.0 ns 0.0 36.0 ** 0.0 36.0 ** Tree-creepersand nuthatches(trunk and branch feeders) 2 2 44.5 99.5 ns 66.5 77.5 ns 57.5 86.5 ns Magpie-larks (ground feeders) ! ! 7.0 29.0 ns 0.0 36.0 ** 0.0 36.0 **

*, P < 0.05; **, P < 0.01; ***, P < 0.001; ns, P > 0.05. ' Numberof speciesfrom rain forest(RF) compared with numberfrom eucalyptforest/eucalypt woodland (EF/EW). Species compared indicated in Table 1. For Pachycephalatests 1 and 2, see Table 2.

of primary 9 and secondary1 are more clearcut. It would be fruitful to extend my study to Wing features should be viewed as part of a incorporate data on wingspan, wing loading, composite. This is obvious when contrasting and aspectratio. Suchinformation would great- wing types are diagrammed(Figs. 1 and 2). ly improve understandingand interpretation. The significanceof differencesin relative Unfortunately,many of the speciescovered here length of the diminutive primary 10 remainsto are relatively inaccessibleand, sincethese mea- be resolved.In an early study, Averill (1925) surementscan only be made on live material, found that the length of this feather, which is the repeat study may have to be based on a commonlyregarded as "vestigial," actually var- smaller subsetof species. ied inversely with the length of the wing. It The avifaunas of New Guinea and Australia was short,if not absent,in migratorymembers representtwin, but quite differently adapted, of the Vireonidae and Turdinae, but showed no radiations from a common stock. The bulk of regressionin the Troglodytidae,few speciesof New Guinea birds inhabitat rain forest and which are migrants. He found that it was not montane forest; most Australian speciesare reduced in the large-bodied crows and ravens found in open eucalyptforest and eucalypt and, hence, he suggestedthat in these species, woodland (Keast 1981). The features docu- it must function in flight. A much more com- mented here presumablyhave evolved repeat- prehensive study than the present one would edly in the two faunas. be necessaryto explore the role of the tenth primary in Australo-Papuanbirds. It is reduced ACKNOWLEDGMENTS in size in only one of the three migratory spe- The study was carried out while the writer held a cies,Pachycephala rufiventris. CanadianNatural Sciencesand EngineeringResearch 102 ALLENKEAST [Auk, Vol. 113

Council Grants.I thank this body for its support.Spec- and tail of three coniferous forest tits, the Gold- imens were kindly made available by the American crest, and the Treecreeper in relation to loco- Museumof Natural History, New York, and I would motor pattern and feeding stationselection. Phi- also like to thank this body. The calculationsand los. Trans. R. Soc. Lond. 287:131-165. diagramswere generatedby Laura Pearce,who also NORBERG,U. M. 1981. Allometry of bat wings and ran the statistical tests. legs and comparisonwith bird wings. Philos. Trans. R. Soc. London. Biol. Ser. 292:369-398. NORBERG,U.M. 1990. Vertebrate flight. Springer- LITERATURE CITED Verlag, New York. NORSERG,U. M., ANDJ. M. V. RAYNER.1987. Ecolog- ATCHELY, W. R., C. T. GASKINS,AND D. ANDERSON. ical morphology and flight in bats (Mammalia: 1976. Statisticproperties of ratios. 1. Empirical Chiroptera), lung adaptations, flight perfor- results.Syst. Zool. 25:137-148. mance,foraging strategy, and echolocation.Phi- AvFaULL,C. K. 1925. The outer primary in relation los. Trans. R. Soc. London. Biol. Ser. 316:335-427. to migration in the ten-primariedoscines. Auk PENNYCUICK,C.J. 1969. The mechanics of bird mi- 42:353-358. gration. Ibis 96:565-585. BAIRLEIN,F. 1992. Morphology-habitatrelationships PIzzEY,G. 1980. A field guide to the birds of Aus- in migratingsongbirds. Pages 356-369 in Ecology tralia. Princeton Univ. Press, Princeton, New Jer- and conservationof Neotropical migrant land- sey. birds (J. M. Hagan III and D. W. Johnston,Eds.). RAND, A. L., AND E. T. GILLIARD. 1967. Handbook of SmithsonianInstitution Press,Washington, D.C. New Guinea birds. Natural History Press,New BEEHLER,B. M., T. K. PRATT,AND D. A. ZIMMERMAN. York. 1986. Birds of New Guinea. Princeton Univ. Press, RAYNER,J. M.V. 1988. Form and function in avian Princeton, New Jersey. flight. Curr. Ornithol. 5:1-66. BtArmRs,M., S. J. J. F. DAVIES,AND P.M. REILLY. 1984. RECHER,H. F., AND R. T. HOLMES. 1985. Foraging The atlas of Australian birds. Melbourne Univ. ecologyand seasonalpatterns of abundancein a Press, Melbourne, Australia. forestavifauna. Pages 79-96 in Birdsof eucalypt CAMERON,E. 1985. Habitat use and foraging behav- forests and woodlands: Ecology, conservation, iour of three fantails (Rhipidura:Pachycephali- management(A. Keast,H. F. Recher, H. Ford, dae). Pages177-191 in Birds of eucalypt forests and D. Saunders, Eds.). Surrey Beatty & Sons, and woodlands:Ecology, conservation, manage- Chipping Norton, New South Wales,Australia. ment (A. Keast, H. F. Recher, H. Ford, and D. RECHER,H. F., R. T. HOLMES, M. SCHULZ, AND R. KA- Saunders,Eds.). Surrey Beatty& Sons,Chipping VANAGI-I.1985. Foraging patterns of breeding Norton, New South Wales, Australia. birds in eucalypt forest and woodland of south DIAMOND,J.M. 1972. Avifauna of the EasternHigh- eastern Australia. Aust. J. Ecol. 10:399-419. lands of New Guinea. Publ. Nuttall Ornithol. Club SAVILLE,D. B. O. 1957. Adaptive evolution in the 12. avian wing. Evolution 11:212-224. HALL, B. P. (Ed.). 1974. Birds of the Harold Hall SPECHT,R.L. 1981. Major vegetationformations in Expedition1962-1970. A reporton the collections Australia.Pages 163-298 in Ecologicalbiogeog- made for the British Museum (Natural History). raphy of Australia (A. Keast,Ed.). Dr. W. Junk, Trustees of the British Museum (Natural His- The Hague. tory), London. THOLLESSON,M., AND U. M. NORBERG. 1991. Mo- HOLMES,R. T., AND H. F. RECHER.1986. Search tactics ments of inertia of bat wings and body. J. Exp. of insectivorousbirds foraging in an Australian Biol. 158:19-35. eucalyptforest. Auk 103:515-530. WENS,L. J., AND J. G. TRACEY. 1981. Australian rain- KEAST,A. 1961. Bird speciationon the Australian forests:Patterns and change. Pages 605-694 in continent. Bull. Mus. Comp. Zool. 123:307-495. Ecologicalbiogeography of Australia (A. Keast, KEAST,A. 1981. The evolutionarybiogeography of Ed.). Dr. W. Junk, The Hague. Australianbirds. Pages1585-1635 in Ecological WINKLER, H., AND B. LFaSLER.1992. On the ecomor- biogeographyof Australia3 (A. Keast,Ed.). Dr. phologyof migrants.Ibis 134 (Suppl. 1):21-28. W. Junk, The Hague. ZAR,J.H. 1984. Biostatisticalanalysis.Prentice-Hall, NORBERG,U.M. 1979. Morphologyofthewings, legs New Jersey. January1996] WingShape in InsectivorusPasserines 103

+1 +1 +1 +1 +1 +1 +1 +1 +1 +1

+1+1+1+1+1+1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1+1+1+{+1+1+1+1 +1+{

• +1+1+1+1+1+1 •

• +l+l+l+l+l+l•

+1+1+1+1+1+1 +l +1 +1 +1 +1 +1 +1 +1 +1 +1 +l+l+l+l+l+l+l+l+l+l

vzvgg 104 ALLENKEAST [Auk, Vol. 113

+1 +1

•+1+1+1+1

+1+1+1+1

• +1+1+1

•+l+l+l+l

+1+1+1 +1+1 +l+l+l+l

z zzz•L•-<,•<• z<< z< z