Ornithological Monographs No.41 Hindlimb Myology andEvolution ofthe OldWorldSuboscine Passerine Birds (Acanthisittidae, Pittidae, Philepittidae, Eurylaimidae) RobertJ Raikow HINDLIMB MYOLOGY EVOLUTION OLD AND OF THE WORLD SUBOSCINE PASSERINE BIRDS (ACANTHISITTIDAE, PITTIDAE, PHILEPITTIDAE, EURYLAIMIDAE) ORNITHOLOGICAL MONOGRAPHS This series,published by the American Ornithologists' Union, has been established for major papers too long for inclusion in the Union's journal, The Auk Publication has been made possiblethrough the generosityof the late Mrs Carll Tucker and the Marcia Brady Tucker Foundation, Inc Correspondenceconcerningmanuscriptsfor publication in the seriesshould be addressedto the Editor, Dr David W Johnston,Department of Biology, George MasonUniversity, Fairfax,Virginia22030 Copies of OrnithologicalMonographs may be ordered from the Assistant to the Treasurer of the AOU, Frank R Moore, Department of Biology, University of Southern Mississippi, Southern Station Box 5018, Hattiesburg, Mississippi 39406 (See price list on back and inside back covers.) OrnithologicalMonographs,No 41, viii + 81 pp Editor, David W Johnston Special Reviewers for this issue, Richard L Zusi, Division of Birds, National Museum of Natural History, Washington, D.C 20560; Andrew J Berger, 1349 Kainui Dr., Kailua, Hawaii 96734 Author, Robert J Raikow, Department of Biological Sciences,University of Pittsburgh,Pittsburgh,Pennsylvania 15260 and CarnegieMuseum of Natural History, Pittsburgh, Pennsylvania 15213 First received, 15 January 1985; final revision completed, 23 October 1985 Issued March 19, 1987 Price $12.50 prepaid ($9.50 to AOU members) Library of CongressCatalogue Card Number 87-70535 Printed by the Allen Press,Inc., Lawrence, Kansas 66044 Copyright ¸ by the American Ornithologists'Union, 1987 ISBN: 0-943610-51-6 HINDLIMB MYOLOGY EVOLUTION OLD AND OF THE WORLD SUBOSCINE PASSERINE BIRDS (ACANTHISITTIDAE, PITTIDAE, PHILEPITTIDAE, EURYLAIMIDAE) BY ROBERT J RAIKOW Department of Biological Sciences University of Pittsburgh Pittsburgh, Pennsylvania 15260 and Carnegie Museum of Natural History Pittsburgh, Pennsylvania 15213 ORNITHOLOGICAL MONOGRAPHS PUBLISHED THE AMERICAN BY ORNITHOLOGISTS' WASHINGTON, 1987 NO D.C UNION 41 TABLE LIST OF FIGURES LIST OF TABLES LIST OF APPENDICES INTRODUCTION OF CONTENTS vi viii viii Eurylaimidae Philepittidae Pittidae Acanthisittidae MATERIALS Dissection Data AND METHODS 7 Analysis MUSCLES OF THE THIGH M iliotibialis cranialis M iliotibialis lateralis M iliofibularis 10 M iliotrochantericus caudalis 11 Mm iliotrochantericus cranialis and iliotrochantericus medius 11 M iliofemoralis intemus 12 Mm femorotibialis extemus and femorotibialis medius 12 M femorotibialis intemus 13 M flexor cruris lateralis 13 M flexor cruris medialis M caudofemoralis 14 15 M ischiofemoralis 15 M obturatorius lateralis 15 M obturatorius medialis 16 16 M pubo-ischio-femoralis MUSCLES OF THE CRUS 17 M tibialis cranialis 17 17 M extensordigitomm longus 18 M fibularis longus M fibularis brevis 18 M M M M M M M M M 19 gastrocnemius 2O plantaris 21 flexor perforans et perforatus digiti II 21 flexor perforans et perforatus digiti III 21 flexor perforatus digiti II 22 flexor perforatusdigiti III 22 flexor perforamsdigiti IV 23 flexor hallucis longus flexor digitomm longus MUSCLES OF THE PES 26 M extensorhallucis longus 26 M flexor hallucis brevis 26 M abductor digiti IV 27 M lumbricalis DISCUSSION 27 OF MORPHOLOGICAL CHARACTERS 28 Myological Characters 28 Nonmyological Characters 34 PHYLOGENY OF THE METHODOLOGICAL OLD WORLD LIMITATIONS SUBOSCINES 34 37 Problems with Specimens 38 Individual Variation 38 Cladisfic Analysis 38 COMPARISON WITH CLASSIFICATION 41 CITED 43 LIST 39 41 LITERATURE Figure STUDIES 40 ACKNOWLEDGMENTS SUMMARY OTHER OF FIGURES Superficiallayer of lateral musclesof the thigh and crus in Eurylaimus ochromalus 48 Secondlayer of lateral musclesof the thigh and crusin Eurylaimus ochromalus 49 Third layer of muscleson the lateral surfaceof the crus in Eurylaimus ochromalus 50 Fourth layer of muscleson the lateral surfaceof the crus in Eurylaimus ochromalus 51 Deep muscleson the lateral surfaceof the thigh and crus in Eurylaimus ,ochromalus 52 Superficialmusculatureon the medial surfaceof the thigh and crus in Eurylaimus ochromalus 53 Muscles on the medial surfaceof the crus in Eurylaimus ochrom- alus 54 Deeper layer of muscleson the medial surfaceof the thigh and crus in Eurylaimus ochromalus 55 Structural variations in M iliotrochantericus cranialis and M iliotrochantericus medius 56 10 11 Variation in M extensor digitorum longus 57 Plantar surface of the tarsometatarsusshowing variation in the insertionof M gastrocnemius 57 12 Variation in M flexor perforatusdigiti IV 58 13 Musclesand tendonson the plantar surfaceof the tarsometatarsus in Eurylaimus ochromalus 59 14 Superficiallateral musculatureof the thigh and crus in Smithornis capensis 59 15 Superficialmusculatureon the lateral surfaceof the thigh in Calyptomena viridis 60 16 M iliotrochantericuscaudalisin Calyptomenawhiteheadi 61 17 Superficialmusculatureon the lateral surfaceof the thigh and crus in Pseudocalyptomena graueri 62 18 Superficial musculature on the medial surface of the thigh and crus in Pseudocalyptomena graueri 63 19 Superficialmusculatureon the lateral surfaceof the thigh and crus in Philepitta castanea 64 20 Superficial musculatureon the medial surfaceof the thigh and crus in Philepitta castanea 65 21 Superficialmusculatureon the lateral surfaceof the thigh and crus in Neodrepaniscoruscans 66 22 Superficial musculature on the medial surface of the thigh and crus in Neodrepaniscoruscans 67 23 Superficialmusculatureon the lateral surfaceof the thigh and crus in Pitta versicolor 68 24 Superficial musculatureon the medial surfaceof the thigh and 25 crus in Pitta versicolor 69 Variation in the insertions of M flexor cruris medialis and M 26 flexor cruris lateralis pars pelvica 70 Superficial muscles on the lateral surface of the knee in Pitta 27 brachyura 71 Superficialmusculatureon the lateral surfaceof the thigh and crus in Acanthisitta 28 chloris 72 Secondlayer of muscleson the lateral surfaceof the thigh and crus in Acanthisitta chloris 73 29 Third layer of muscleson the lateral surfaceof the thigh and crus in Acanthisitta 30 31 Acanthisitta 32 chloris 74 Deep musclesof the hip in Acanthisitta chloris 75 Superficialmuscleson the medial surfaceof the thigh and crus in chloris 76 Secondlayer of muscleson the medial surfaceof the thigh and crus in Acanthisitta 33 34 chloris 77 Deep caudomedial muscles of the crus in Acanthisitta chloris 78 Extremesof variation in M extensorhallucislongusand M flexor hallucis brevis 78 35 A cladogramhypothesizingthe phylogeneticrelationshipsof the 36 37 Old World suboscines 79 Strict consensus tree 80 A comparison of the results of three different studiesof passeriform relationships 81 vii LIST Table OF TABLES Character statesused in phylogeneticanalysis 29 Distribution of characterstatesamongtaxa analyzed 30 Suggestedclassification 41 LIST OF APPENDICES I List of Abbreviations in Figures 47 INTRODUCTION The purpose of this study is to increaseour understandingof the evolutionary history of the passefinebirds by adding to our knowledgeof their morphology, reconstructingtheir phylogeny, and using the information thus obtained to improve existing classifications.To this, I will provide detailed descriptionsof the hindlimb musculatureof the Old World suboscines(Acanthisittidae, Pittidae, Philepittidae, and Eurylaimidae), and will use this information to infer the phylogeneticrelationships among these taxa The relationshipsof birds are reasonablywell establishedat the specieslevel, but are progressivelylesssatisfactoryat higher levels The fact that a fairly stable classificationhas existedfor severaldecadesis misleadingif it suggests that the relationships among most avian families and orders are well understood: many ambiguitiesand disagreementsconcerningtheserelationshipsexist (Stresemann 1959; Cracraft 1972, 1981) An especiallyseriousproblem occurswith the largest order of birds, the Passefiformes.Although 25-30 ordersof living birds are generally recognized,the Passefiformesalone contain more than half of all species These are divided into about 70 families (Wetmore 1960), most of them in the suborderPasseres(oscinesor true songbirds).The relationshipsamong the Passefiformeshave long been poorly understood.Many families might prove to be monophyletic, but this possibility must be tested and not assumed.The relationshipsof the families to each other, and of the generawithin families, are very obscure.Some families have been erected for one genusor a few generasimply becausetheir relationshipsare not understood.To a considerableextent existing classifications reflectmore the proceduresand philosophyof traditionaltaxonomy than they the patternsof corroboratedphylogenetic hypotheses (Raikow 1985a) The basicproblem to be investigatedis the phylogenetichistory of the Passeriformes; this must be approached, for practical reasons, through subordinate problems of manageable size Previous studies in this program emphasized relationships between various oscine groups (Bentz 1979; Borecky 1977, 1978; Raikow 1973, 1976, 1977a,b, 1978, 1985d;Raikow et al., 1980; Ufik 1983), of nonpassefineorderssuspectedof closerelationshipto the Passefiformes(Berman and Raikow 1982; Maurer and Raikow 1981; Swierczewskiand Raikow 1981), and the demarcationof the Passefiformes asa monophyleticgroup(Raikow 1982) The present consideration of suboscinesthus forms a link between the separate approaches previously pursued A detailed history of passefineclassificationis beyondthe scopeof this work, but is thoroughlycoveredby Sibley(1970:23-31) and Ames (1971:127-129, 153164) Thus, only a brief review of the groups studied will be offered here The majorproblemsdealwith the arrangementof the suboscines within a phylogenetic framework.Are the Eurylaimidaethe most primitive passefinesas often suggested?Are they closelyrelated to the Cotingidae?Are the Acanthisittidaesuboscine or oscine? Are the Old World suboscines linked with the New World Tyrannoidea as traditional interpretationsof syfingealanatomy have suggested? Answerswill be offeredto suchquestionsin the form of a corroboratedphylogenetic hypothesis In this work I will consideras monophyletica group that appearsfrom its characteristics to includeall of the known descendants of a singlecommonances- 68 ORNITHOLOGICAL MONOGRAPHS NO 41 FTEP CF I½ IL FCRLP FCRM GL FPPD2 FPD4 FB FPPD FL FiG 23 Superficialmusculatureon the lateral surfaceof the thigh and crus in Pitta versicolor, showingthe acetabulargap in M iliotibialis lateralis (IL) Abbreviationsin Appendix I SUBOSCINE MYOLOGY AND EVOLUTION 69 ITCR IFI • CF ICR FTM OM TI FCR PIFCA GL GI TCR EDL FL GM FIG 24 Superficialmusculatureon the medial surfaceof the thighand crusin Pitta versicolor M gastrocnemius parsmedialis(GM) showsa primitive robustform with superficialand deepheadsof origin,differentfrom the derivedattenuateconditionin the PhilepittidaeandEurylaimidae.Thereis no Parssupramedialis M pubo-ischio-femoralis parscaudalis(PIFCA) showsa caudadenlargement of its area of origin, a derived condition.Abbreviationsin Appendix I 70 ORNITHOLOGICAL MONOG1L•PHS NO 41 FCRLP FCRM PL FCRLA EDL Fla 25 Variation in the insertionsof M flexor cruris medialis (FCRM) and M flexor cruris lateralisparspelvica(FCRLP).Left:condition in Pittabrachy'ura showing theprimitiveconditionof fusionof the two tendons.Right:derivedconditionexemplifiedby P versicolor showingthe derived separation of the two tendonsof insertion.Abbreviations in AppendixI SUBOSCINE MYOLOGY AND EVOLUTION 71 ILPR FTE / // / ILPO // •FPPD $ FL FiG 26 Superficialmuscleson the lateral surfaceof the knee in Pitta brachyura,illustrating a derived condition of pittasin which the distal part of the postacetabularbelly of M iliotibialis lateralis (ILPO) is dividedinto superficial(a) and deep(b) portionswith separateinsertions.Abbreviationsin Appendix I 72 ORNITHOLOGICAL MONOGRAPHS NO 41 ICR IL FCRLP GL FB FL FHL TCR FPD FPPD FIG 27 Superficialmusculatureon the lateral surfaceof the thigh and crusin Acanthisittachloris Note that in M iliotibialis lateralis (IL) the pre- and postacetabularportions are not separatedby an acetabulargap Abbreviations in Appendix I $UBOSCINE MYOLOGY AND EVOLUTION 73 ITCA IF ITCR ISF • FCRLP FT FPD FTED FPPD FHL FB TCR FPD FIO 28 Secondlayer of muscleson the lateral surfaceof the thigh and crusin Acanthisittachloris Abbreviations in Appendix I 74 ORNITHOLOGICAL MONOGRAPHS NO 41 ITCR ITM ISF FCRLP CF PIFCR FCRLA FCRM FPD FPD $ oo EDL FHL FDL FB FIG 29 Third layer of muscleson the lateral surfaceof the thigh and cms in Acanthisittachloris Abbreviationsin Appendix I SUBOSCINE MYOLOGY AND EVOLUTION 75 ITCA • ISF OLD :'.: • OLV OM Fxo 30 Deep musclesof the hip in Acanthisittachloris,showingthe enlargedconditionof M obturatoriuslateralisparsdorsalis(OLD) Abbreviationsin Appendix I 76 ORNITHOLOGICAL MONOGRAPHS NO 41 IF ISF ITCR IFI OLD FTM ICR OM PIFCR GL FTI PIFCA l'CR FPD5 FCRM GM GI FPD FIO 31 Superficialmuscleson the medialsurfaceof the thighand crusin Acanthisittachloris.M gastrocnemiuspars mediaIls (GM) has a primitive form rather than the attenuatederived form of the Philepittidaeand Eurylaimidae,and lacksPars supramedialis.Abbreviationsin Appendix I SUBOSCINE MYOLOGY AND EVOLUTION 77 IFI PIFCR PIFCA FPD $ FPD FTM FTI PL FDL TO R GI FORM EDL FtG 32 Secondlayerof muscleson the medial surfaceof the thighand crusin.4cantl•isittacldoris Abbreviationsin Appendix I 78 ORNITHOLOGICAL FPD,4 FPD3 MONOGRAPHS NO 41 '•,.;.'.:, i FDL PL FIG 33 Deep caudomedialmusclesof the crusin Acanthisittachloris.Abbreviationsin Appendix I FHB Calxpfomena Fro 34 Extremesof variation in the development of two intrinsic musclesof the pes, Mm extensor hallucislongus(EHL) and flexorhallucisbrevis(FHB) as illustratedby Acanthisittachlorisand Calyptornenaviridis.MTI is metatarsalI Abbreviationsin AppendixI SUBOSCINE MYOLOGY AND EVOLUTION 79 22,1 27.1 F]o 35 One of four similar and equallyparsimoniouscladogramsshowingthe relationshipsof the Old World suboscincsto each other and to the other passerincbirds They were generated by PAUPandhavea consistency indexof0.771,excluding autapomorphic character states Thenumbers designatethe characterstatesas in Tables and A minussignindicatesan evolutionaryreversal Derivedstatespostulated to havearisenindependently in twodifferentlineages areunderlined.Corydon was not dissected,and is includedtentativelyon the basisof bill shapealone;it was not includedin the computeranalysis 80 ORNITHOLOGICAL MONOGRAPHS Eurylaimus E NO 41 steeœi och•omalus Cymbirhynchus Seœilophus Psarisomus Smithoœnis I Calyptomena viridis C whiteheadi Pseudocalyptomena I Philepitta Neodrepanis Pitta versacolor P guajana P erythrogaster P brachyura Tyranni Furnarii Acanthisitta Oscines• inc Menurae FI0.36 Stfi• consensust•e •r •ursimilarcladograms, one of which isshowninFigu• Thisisthebasic phylogen•ichypo•esisofthepresent Mvesfigafion 35 SUBOSCINE MYOLOGY AND EVOLUTION 81 A Presentstudy B Sibley et al C Wetmore (1960) FIG 37 A comparisonof the results of three different studiesof passeriformrelationships.A Simplifieddadogramshowingresultsof the presentmorphologicalstudy.B Simplifiedcladogramof the resultsof the DNA hybridization studyof Sibleyet al (1982) and Sibleyand Ahlquist (1985b) C The hierarchicalstructureof the classificationof Wetmore (1960) A singlenomenclatureis used for easeof comparison.The taxon Furnarii correspondsto the Furnarioidea of Sibley et al (1982) and of Wetmore (1960), and to a corresponding but unnamedtaxon of Sibleyand Ahlquist(1985b) The taxon Tyranni correspondsto the Tyrannoidea of Sibley et al (1982), the Tyranni of Sibley and Ahlquist (1985b), and to the New World families of Wetmore's (1960) taxon Tyrannoidea ORNITHOLOGICAL No No No No No No MONOGRAPHS A Distributional Study oœthe BirdsoœBritish Honduras.StephenM Russell.1964.(Out of print) A ComparativeStudy oœSomeSocialCommunicationPatternsin the Pelecaniœorme• G F van Tets 1965.$2.50 The Birds oœKentucky R M Mengel 1965 $10.00 EvolutionoœSomeArctic Gulls (Larus):an ExperimentalStudyoœIsolatingMechanisms•Neal G Smith 1966 (Out of print) A ComparativeLi[e-historyStudy oœFour SpeciesoœWoodpeckers• Louisede Kiriline Lawrence.1967 (Out of print) Adaptations[or Locomotionand Feedingin the Anhinga and the Double-crestedCormorant.O T Owre 1967 $3.00 No No.& No No 10 No 11 No 12 No 13 No 14 No 15 No 16 No 17 A Distributional Survey oœthe Birds oœHonduras.B L Monroe, Jr 1968.$7.00 An Approachto the Study oœEcologicalRelationshipsamong GrasslandBirds.John A Wiens 1969 (Out of print) Mating Systems,Sexual Dimorphism, and the Role oœMale North American PassertheBirds in the Nesting Cycle.JaredVerner and Mary F Willson 1969.(Out of print) The BehavioroœSpottedAntbirds E O Willis 1972.$4.00 Behavior,Mimetic Son•s and SongDialects,and Relationshipsoœthe ParasiticIndigobirds(Vidua) oœ Aœrica R B Payne.1973.$6.00 Intra-ialand Variation in the MascareneWhite-eyeZosteropsborbontca.F B Gill 1973.$2.50 Evolutionary Trendsin the NeotropicalOvenbirdsand Woodhewer• A Feduccia.1973.$2.50 A Symposiumon the House Sparrow (Passerdomesticus)and EuropeanTree Sparrow (P montanus) in North America.S.C Kendeigh,Ed 1973.$3.00 FunctionalAnatomyand AdaptiveEvolutionoœthe FeedingApparatusin the HawaiianHoneycreeper GenusLoxops(Drepanididae).L P Richardsand W J Bock.1973.$5.00 The Red-tailed Tropicbirdon Kure Atoll R R Fleet 1974.$3.00 ComparativeBehavioroœthe American A vocet and the Black-neckedStilt (Recurvirostridae).R B Hamilton 1975 $4.00 No 18 No 19 Breeding Biology andBehavior oœ theOlds•uaw (Clangula hyemalis L.).R.M Alison 1975 $2.50 No 20 Soxual SizeDimorphism in HawksandOwlsoœ NorthAmerica N F.R.Snyderandj W Wiley.1976 No 21 Social Organization and Behavior of the Acorn Woodpeckerin Central Coastal Callœornla.M H No 22 MacRoberts and B R MacRoberts 1976 $4.00 Maintenance Behavior and Communication in the Brown Pelican R W Schreiber 1977 $3.50 No 23 No 24 SpeciesRelationshipsin the Arian GenusAimophila L L Wolf 1977.$7.00 Land Bird Communities oœGrand BahamaIsland: The Structure and Dynamics oœan Aviœauna.J T Bird PopulationsoœAspenForestsin WesternNorth America.J A.D Flack.1976.$4.00 $6.00 Emlen 1977 $5.00 No 25 Systematics oœSmallerAsianNight BirdsBasedon Voice.J T Marshall.1978.$4.00 No 26 No 27 Ecologyand EvolutionoœLek Mating Behaviorin the Long-tailedHermit Hummingbird.F G Stiles Ecologyand Behavior oœthe Prairie Warbler Dendroica discolor.V Nolan, Jr 1978 $15.00 and L L Wolf 1979 $4.50 No 28 The ForagingBehavioroœMountain Bluebirdswith Emphasison SoxualForagingDigference•H W Power 1980 $4.50 No 29 No 30 The Molt oœScrubJaysand BlueJaysin Florida G T Bancroftand G E Woolfenden.1982.$4.00 Avian Incubation:Egg Temperature,Nest Humidity, and BehavioralThermoregulationin a Hot Environment No 31 No 32 No 33 G S Grant 1982 $5.00 The Native Forest Birds oœGuam J M Jenkins 1983 $6.00 The Marine EcologyoœBirds in the Ross Sea, Antarctica D G Ainley, E F O'Connor, and R J Boekelheide.x + 97 pp 1984 $9.00 ($8.00) SoxualSolection,Lek and Arena Behavior,and SexualSize Dimorphismin Birds.R B Payne.viii + 52 pp 1984.$8.00 ($6.50) No 34 No 35 No 36 No 37 No 38 No 39 No 40 Pattern,Mechanism,and AdaptiveSigni[icanceof Territoriality in Herring Gulls (Larusargentatus) J Burger.xii + 92 pp 1984.$9.00($7.00) Ecogeographical Variation in Size and ProportionsoœSongSparrows(Melospizamelodia).J W Aldrich x + 134 pp 1984.$10.50($8.50) NeotropicalOrnithology.P A Buckleyet aL, Eds.xi + 1,041pp., colorplates.1985.$70.00 Arian Monogamy.P A Gowatyand D W Mock, Eds.vi + 12r pp 1985.$11.00($9.00) An Analysisof Physical,Physiological,and Optical AspectsoœArian Colorationwith Emphasison Wood-Warbler• E H Burtt,Jr x + 122 pp 1986.$15.00($12.50) The Lingual Apparatusoœthe AœricanGrey Parrot, PsittacuserithacusLinn• (Aves:Psittacidae): scriptionand TheoreticalMechanicalAnalysis.D G Hornberger.xii + 236pp 1986.$25.00($20.00) Patternsand EvolutionarySignil•canceoœGeographicVariation in the •chistaceaGroup oœthe Fox Sparrow(Passerellailiaca) R M Zink viii + 119 pp 1986.$15.00($12.50) Orderfrom:FrankR•Moore,Assistant to theTreasu.rer A.O.U.,Department of Biological Sciences, University of SouthernMississippi,Hattiesburg,Mississippi39406.Ordersmustbe prepaid,in U.S dollars;add percent(minimum$1.50)handlingandshippingcharge.Makechecks payableto AmericanOrnithologists' Union Pricesin parenthesesare for A.O.U members ... 5018, Hattiesburg, Mississippi 39406 (See price list on back and inside back covers.) OrnithologicalMonographs,No 41, viii + 81 pp Editor, David W Johnston Special Reviewers for this issue, Richard... Natural History Pittsburgh, Pennsylvania 15213 ORNITHOLOGICAL MONOGRAPHS PUBLISHED THE AMERICAN BY ORNITHOLOGISTS' WASHINGTON, 1987 NO D.C UNION 41 TABLE LIST OF FIGURES LIST OF TABLES LIST OF... its characteristics to includeall of the known descendants of a singlecommonances- ORNITHOLOGICAL MONOGRAPHS NO 41 tor This ancestoris hypothetical,but would be includedin the group if known Paraphyleticgroups,which