(ISBN: 0-943610-49-4) THE LINGUAL APPARATUS THE AFRICAN OF GREY PARROT, PSITTA CUS ERITHA CUS LINNl• (AVES: PSITTACIDAE): DESCRIPTION AND MECHANICAL THEORETICAL ANALYSIS BY DOMINIQUE G HOMBERGER Department of Zoology and Physiology Louisiana State University Baton Rouge, Louisiana 70803 ORNITHOLOGICAL MONOGRAPHS PUBLISHED THE AMERICAN BY ORNITHOLOGISTS' WASHINGTON, 1986 NO D.C UNION 39 THE LINGUAL APPARATUS OF THE AFRICAN GREY PARROT, PSITTA CUS ERITHA CUS LINN]• (AVES: PSITTACIDAE): DESCRIPTION AND MECHANICAL THEORETICAL ANALYSIS ORNITHOLOGICAL MONOGRAPHS This series,publishedby 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 Mason University, Fairfax, VA 22030 Copies of Ornithological Monographs 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 39, xi + 233 pp Editor of AOU Monographs, Mercedes S Foster and David W Johnston Special Reviewers for this issue, Richard L Zusi, Division of Birds, National Museum of Natural History, Washington, D.C.; Kathleen K Smith, Department of Anatomy, Duke University Medical Center, Durham, N.C Author, Dominique G Hornberger, Department of Zoology and Physiology, Louisiana State University, Baton Rouge, Louisiana 70803 First received, 31 January 1983; accepted, 25 July 1983; final revision completed, 12 September 1984 Issued June 12, 1986 Price $25.00 prepaid ($20.00 to AOU members) Library of CongressCatalogueCard Number 86-71332 Printed by the Allen Press,Inc., Lawrence, Kansas 66044 Copyright ¸ by the American Ornithologists'Union, 1986 ISBN: 0-943610-49-4 THE LINGUAL APPARATUS THE AFRICAN OF GREY PARROT, PSITTA C US ERI THA C US LINNI• (AVES: PSITTACIDAE): DESCRIPTION AND MECHANICAL THEORETICAL ANALYSIS BY DOMINIQUE G HOMBERGER Department of Zoology and Physiology Louisiana State University Baton Rouge, Louisiana 70803 ORNITHOLOGICAL MONOGRAPHS PUBLISHED THE AMERICAN BY ORNITHOLOGISTS' WASHINGTON, 1986 NO D.C UNION 39 TABLE LIST OF FIGURES LIST OF TABLES LIST viii x OF APPENDICES INTRODUCTION MATERIAL AND OF CONTENTS xi METHODS ANATOMY INTRODUCTION SICeLœTON Introduction Hyoid Skeleton Os epibranchiale Os ceratobranchiale Os basihyale, Os urohyale, and Nodulus Os paraglossale 11 Discussion 14 Articulations of the Hyoid Skeleton: Structure and Function 14 Articulatio epi-ceratobranchialis 14 Articulatio cerato-basihyalis 15 Articulatio paraglosso-basihyalis 17 Discussion 19 Mandible 20 Larynx and Trachea 20 LXGnMœNTS ^D FnSCX^ (SYNDSMOLOGắ) 20 Introduction 20 Intraosseal Ligaments 21 Articular Ligaments 22 Linkage Ligaments 23 Fasciae 23 Discussion 24 MUSCULATURe 25 Introduction 25 Extrinsic Lingual Musculature 27 M serpihyoideus 27 M stylohyoideus 29 M branchiomandibularis 31 M mylohyoideus 34 M genioglossus 37 M tracheohyoideus 38 Intrinsic Lingual Musculature 42 M ceratohyoideus 42 M ceratoglossus 44 M hypoglossusobliquus 51 M supraglossus 55 M hypoglossusanterior 60 M hypoglossustransversus 63 M mesoglossus 64 BURSAE SYNOvlALES 66 Introduction 66 Description and Discussion 67 Bursa synovialis urohyalis 67 Bursa synovialis paraglossalis 67 THE TONGUE; GENERALSHAPE,SURFACESTRUCTURES,AND RELATIONSHIPS TO UNDERLYrNG AND SURROUNDING STRUCTURES 67 CAVERNOUS VASCULARTISSUE(CORPORACAVERNOSA) 71 Introduction 71 Description and Discussion 72 Corpus cavernosum apicale 72 Corpus cavernosum laterale 72 SALIVARY GLANDS 72 Introduction 72 Description and Discussion 73 Glandula lingualis 73 Glandula sublingualis 75 Glandula FUNCTIONAL mandibularis ANALYSIS INTRODUCTION 75 76 76 Methods of Functional Analysis 76 Reductionistic approach 76 Holistic approach 77 Synthetic approach 78 Physical and PhysiologicalProperties of Tissues 82 Supporting and connective tissues 82 Muscle 83 Epithelial structures 87 Hydraulic structures 89 Biomechanical Principles 91 Constructional basis of movements 91 Guidance and regulation of movements 91 Actions and interactions of muscles 94 Synchronizationof movements 96 Construction of a Theoretical Mechanical Model 96 Description of the Theoretical Mechanical Model 97 MOVEMENTS OF THE HYOID RELATIVE TO THE MANDIBLE 99 Back-and-Forth Movement of the Hyoid 99 Retraction of the hyoid 99 Protraction of the hyoid 101 Up-and-Down Movement of the Hyoid 102 Lifting of the retracted, lowered hyoid 103 Protraction of the hyoid without simultaneouslifting 104 Lowering of the hyoid 105 Side-to-SideMovement of the Hyoid 106 Functions of intrinsic lingual muscles 107 Functions of extrinsic lingual muscles 110 Discussion MOVEMENTS 112 OF THE EPIBRANCHIALE RELATIVE TO THE CERATO- m•ANCHI^LE 113 Movements of the Epibranchiale During Protraction of the Hyoid 113 Movements of the Epibranchiale During Retraction of the Hyoid 114 MOVEMENTS OF THE PARAGLOSSALE RELATIVE TO THE BASIHYALE 114 Up-and-Down Movement of the Paraglossale 114 Extension of the paraglossalewith stationary hyoid 115 Flexion of the paraglossalewith stationary hyoid 116 Movements of the paraglossale during retraction of the hyoid 117 Movements of the paraglossale during protraction of the hyoid 119 Discussion 120 Side-to-Side Movement of the Paraglossale 120 MOVEMENTS AT THE TIP OF THE TONGUE 121 Movements of the Epithelial Surfaceof the Tip of the Tongue 122 Depression of the central area of the dorsal epithelial surface 123 Flattening of the dorsal epithelial surface 124 Inflation of the Corpora cavernosa 126 Deflation of the Corpora cavernosa 128 Movements of the Epithelial Surfaceof the Tip of the Tongue During Movements of the Paraglossale 129 Depression of the dorsal epithelial surfaceduring flexion of the paraglossale 129 Flattening of the dorsalepithelial surfaceduring flexion of the paraglossale 130 Depressionof the dorsal epithelial surfaceduring extension of the paraglossale 132 Flattening of the dorsalepithelial surfaceduring extensionof the paraglossale 133 Discussion 134 Movements of the Epithelial Surfaceof the Tongue'sTip During Movements of the Hyoid 134 Movements of the epithelial surfaceand paraglossaleduring retraction of the hyoid 135 Movements of the epithelial surfaceand paraglossaleduring protraction of the hyoid 136 Discussion 138 CONCI•USIONS 138 ACKNOWLEDGMENTS 144 vii SUMMARIES 144 ENGLISH: SUMMARY 144 GERMAN: ZUSAMMENFASSUNG 147 LITERATURE CITED 149 LIST OF FIGURES The bony elementsof the articulatedhyoid skeletonin protracted position 170 Lateral views of the bony elements of the articulated hyoid skeletonand of the basihyaleand Nodulus 171 Left epibranchiale 172 Right ceratobranchiale 173 Left ceratobranchiale 174 Apical end of the right ceratobranchiale(lateral and dorsalviews) 175 Apical end of the right ceratobranchiale(dorso-medialand medial views) 176 Apical end of the basihyale,and right articular facet of the Art cerato-basihyalison the basihyale 177 Ventral views of the bony portion of the articulated paraglossalia178 10 Dorsal view of the bony portion of the articulated paraglossalia, and medial view of the left paraglossale 179 11 Mandible with rhamphotheca,originsofextrinsiclingualmuscles, and insertion of M ethmomandibularis 180 12 Hyoid skeletonand attachmentsitesof lingual muscles 181 13 Ventral views of the hyoid skeletonwith attachment sitesof lin- gual muscles 182 14 Articular facetsof the hyoid skeletonshowingtrajectoriesof the movements of the complementaryarticular facets 183 15 In situ views of the superficialjaw and lingual musculature 184 16 Ventral views of the lingual apparatusafter separationfrom the mandible 17 18 19 20 21 22 23 24 25 26 27 28 185 Ventral views of the central portion of the lingual apparatus 186 Ventral view of the central portion of the lingual apparatus 187 Ventral views of the central portion of the lingual apparatus 188 Lateral view of the antero-centralportion of the lingualapparatus 189 Superficial lingual musculature and epithelial structures of the lingual apparatus 190 Enlargeddorsalview of anterior half of the tongue 191 Ventral views of the lingual apparatus 192 Dorsal views of the lingual apparatus,excludingthe tongue'stip 193 Dorsal views of the lingual apparatus 194 Dorsal views of central portions of the lingual apparatus 195 Ventro-lateral view of the apical three-quartersof the lingual apparatus, and enlargedventral view of the tongue'stip 196 Ventral views of central portions of the lingual apparatus 197 viii 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 Lateral views of the apical portion of the lingual apparatus 198 Ventral views of the tongue 199 Ventral views of the M supraglossus-M.mesoglossussystemof the lingual tip and of the tongue'stip 200 Tip of the tongue 201 Medial and dorsal views of the ceratobranchialeand epibranchiale, with attached muscles 202 Apical view of a cross-sectionthrough the lingual tip 203 Cross-sectionsthrough the lingual tip 204 Apical views of cross-sectionsthrough the lingual tip 205 Apical views of cross-sectionsthrough the lingual tip 206 Apical views of cross-sectionsthrough the lingual tip 207 Apical views of cross-sectionsthrough the lingual tip 208 Apical views of cross-sectionsthrough the lingual tip 209 Apical views of cross-sectionsthrough the lingual tip 210 Apical views of cross-sectionsthrough the lingual tip 211 Longitudinal, slightlyparasagittalsectionthroughthe tongue'stip 212 Longitudinal sectionsthrough the anterior portion of the lingual apparatus 213 Longitudinal sectionsthrough the anterior portion of the lingual apparatus 214 Retraction of the protracted hyoid 215 Protraction of the retracted hyoid 216 Lateral views of the up-and-down movement of the hyoid with immobilized paraglossale 217 Lateral views of the up-and-down movement of the hyoid 218 Dorsal views of the sinistral movement of the hyoid with immobilized paraglossale 219 Lateral views of the left half of the hyoid with immobilized paraglossaleduring the sinistral movement 220 Lateral views of the right half of the hyoid with immobilized paraglossaleduring the sinistral movement 221 Movements of the paraglossalewith the hyoid immobilized in intermediate position between protracted and retracted position 222 Lateral views of movements of the paraglossaleduring retraction of the protracted hyoid 223 Lateral views of movements of the paraglossaleduring protraction of the retracted hyoid 224 Dorsal views of lateral movements of the paraglossalewith the hyoid immobilized in intermediate position 225 Midsagittal views of movements of the epithelial surface of the tongue's tip 226 Dorsal views of movements of the epithelial surfaceof the tongue's tip 227 59 Movements of the tongue's tip brought about by the Corpora cavernosa 228 ix LINGUAL APPARATUS OF PSITTACUS 221 A B • • Mcgm • Mbma Mbmp F•G 52 •te• •ews of •e fi•t haw of •e hyoid •th •mob•ed p•a•oss•e d•g •e s•s• movementin PsRtac• eritha•; hyoidat inte•e•ate s•ge be•n promoted•d •ted position.A: Action of M •rato•ossus m•is B: Actionsof the M branchio•bM•s •d M cemtohyoideus (M mylohyoideuspost not sho•; seeFig 50B) (For me•ng of •o• seeFig 46.) C2• •d C• muscleattachmentson •ratob•cM•e (C: centerof Mse•ion of Mb• med., Ca •u• end of o•n of Mch, C4 apic• end of ofi•n of Mch, C• •u• end of ofi•n of M•), E•E: insetohs of Mbm on epibrancM•e (E• •nter of Mse•on of •mp, E: •nter of in•on of Mb• •t.), M4-M• ofi•ns of Mbm on •ble (M• •nter of ofi•n of Mb•, M• dor• end of Mbmp), MD: ven• e•e of m•ble • jaw musclesover wMch •e •mp pasts, • N•Mus •d •nnecfive tissueas in•on sitefor Mch, P3centerof M•on of M• on p•a•os•e -• o'• //// • B2{ Mcgp Mcgs FIO 53 Movementsof the paraglossale with hyoid immobilized in intermediatepositionbetween protractedand retractedpositionin Psittacuserithacus.A-B: Extensionand protractionof the flexed paraglossale (A: Lateralview; B: Dorsalview) C: Lateral view of flexionand retractionof the extended paraglossale (For meaningof arrowsseeFig 46.) B=-B2originsofM hypoglossus obliquusmed.on basihyale(B• caudo-medial end of origin,B2apicalend of origin),Cs-Csoriginsof intrinsiclingual muscles on ceratobranchiale (Cscaudalendof originofM ceratoglossus lat., Cscaudalendof origin of M ceratoglossus med.,C?centerof originof M ceratoglossus prof.,Cscaudalendof originof M hypoglossus obliquuslat.),G3-G4insertions of Mho ontendinous envelope of Gl lingualis (G3caudal endof insertionof Mhom, G4caudalendof insertionof Mhol), L centerof originofM ceratoglossus supf.onLig.nodulo-ceratobranchiale, P•-P4insertions of intrinsiclingualmuscles on paraglossale (P3 centerof insertionof Meg, Psapicalend of insertionof 1VIhom) LINGUAL APPARATUS OF PSITTACUS 223 A • Mhom • Mcgm xX x Mcgs Mcgp FIG 54 Lateral views of movementsof the paraglossaleduring retractionof the protractedhyoid in Psittacu•erithacu, A: Extensionand protractionof the flexedparaglossale B: Flexion and retraction of the extendedparaglossale C: Immobilizationof the extendedparaglossale throughsimultaneous contractionsof the M ceratoglossus and M hypoglossusobliquus(Mhom not shown).(For meaning of arrowsseeFig 46.) B•-B2 originsof Mhom on basihyale(B• caudo-medialend of origin,B2apical endof origin),Cs-C, originsof intrinsiclingualmuscleson ceratobranchiale (Cscaudalendof origin of Mcgl, C6caudalend of originof Mcgm, C?centerof originof Mcgp,Cgcaudalend of originof Mho0, G3 G4insertions of Mho on tendinous envelopeof GI lingualis(G3caudalendof insertion of Mhom, G4 caudalend of insertionof Mhol), L centerof origin of Mcgs on Lig nodulo-ceratobranchiale, P3-P4insertions of intrinsiclingualmuscles on paraglossale (P3centerof insertionof Mcg, P4 apical end of insertionof Mhom) 224 ORNITHOLOGICAL MONOGRAPHS NO 39 Mhol Mhom ,,• Mcgp Mcgm / • • Mcgl, •,,.•/"' • •.- ""' - Megs - •.• *•'"'•' •'/G4 ,/// /// FIG 55 Lateralviewsof movementsof the paxaglossalc duringprotractionof the retractedhyoid in Psittacuserithacus.A: Extensionandprotractionof the flexedparaglossale B: Flexionandretraction of theextended paxaglossale C: Iramobilizationof thepaxaglossale throughsimultaneous contractions of the M hypoglossus obliquus(Mhom not shown)and M ceratoglossus (For meaningof arrowssee Fig 46.) B,-B2originsof Mhom on basihyale (B, caudo-medial endof origin,B2apicalendof origin), C,-C, originsof intrinsiclingualmuscleson ceratobranchiale (C5caudalend of originof Mcgl, C6 caudalendof originof Mcgm,C7centerof originof Mcgp,C, caudalendof originof Mhol), insertionsof Mho on tendinousenvelopeof GI lingualis(Ga caudalend of insertionof Mhom, G4 caudal end of insertionof Mhol), L centerof origin of Megs on Lig nodulo-ceratobranchiale, P3-P4 insertionsof intrinsiclingualmuscleson paxaglossale (Pacenterof insertionof Mcg, P4apicalend of insertion of Mhom) LINGUAL APPARATUS OF PSITTACUS 225 Mhol Mhom -• Mcgp Mcgl Mcgm B _ -: z_- - _ F•a 56 Dorsal views of lateral movementsof the paraglossale with the hyoid immobilized in intermediate position in Psittacuserithacus.A: Turning of the straight paraglossaleto the right B: Straighteningof the laterally turned paraglossale.(For meaning of arrows seeFig 46.) Bi-B2 origins of Mhom on basihyale (B• caudo-medial end of origin, B2 apical end of origin), C•-C8 origins of intrinsiclingualmuscleson ceratobranchiale (C5caudalend of origin of Mcgl, C6caudalend of origin of Mcgm, C7centerof origin of Mcgp, Cs caudalend.of origin of Mhol), G3 G4insertionsof Mho on tendinousenvelopeof GI lingualis(G3 caudal end of insertion of Mhom, G4 caudal end of insertion of Mhol), L center of origin of Mcgs on Lig nodulo-ceratobranchiale,P3-P4 insertions of intrinsic lingualmuscleson paraglossale (P3centerof insertdon of Meg, P• apicalendof insertionof Mhom) 226 ORNITHOLOGICAL MONOGRAPHS NO 39 A Mmg ""'-'"' 'ß Msg Mhga SPG Lipga P5 FIG 57 Midsagittalviews of movementsof the epithelial surfaceof the tongue'stip in Psittacus erithacus.A: Depressionof central area of the dorsalepithelialsurfacethroughcontractionof M hypoglossus anteriorand M mesoglossus B: Flatteningof the dorsalepithelialsurfacethroughcontractionofM supraglossus (For meaningof arrowsseeFig.46.) A•-A3 attachments of intrinsiclingual muscleson aponeurosis ofMhga (A• insertionof tendinousportionE of Msg,A2 caudalendof origin of Mmgp, A3 apicalend of originof Mmgp), B4 origin of Mhga on basihyale,EP•-EP• insertionsof intrinsiclingualmuscleson epithelialsurfaceof tongue'stip (EP• caudalend of insertionof Mmgp, EP2apical end of insertionof Mmgp, EP3centralinsertionof Mmga, EP4apical end of insertionof tendinousportionC ofMsg, EP5insertionof tendinousportionE of Msg),NL• centraloriginof Mmga on lingualnail, P6apicaledgeof Synchonclrosis paraglossalis overwhichthe tendonof Mhgapasses, P? directionof caudo-dorsalend of insertionof Mhga on paraglossale, T2-T3 originsof tendinous portionsof Msgfromcentraltendon(T2originoftenclinous portionC, T• originoftenclinous portionE) LINGUAL APPARATUS OF P$1TTACU$ 227 A Mmgp+Mmga' Msg ApMhga• B N b-•G.58 Dorsalviewsof movements of theepithelial surface of thetongue's tip in Psittacus erithacus A: Depression of centralareaof thedorsalepithelial surface through contraction of M hypoglossus ant.andM mesoglossus (leftside: superposition ofskeletal elements andepithelial surface structures of thetongue's tip;rightside:Ming,agoneurosis of Mhga,tendinous fibersoriginating caudo-laterally fromtheagoneurosis oftheMhgatoinsert ondorsal border oflingual nail,andcaudal contourof G1.lingualis) B: Flattening of thedorsalepithelial surface throughcontraction of M supraglossus (leftside: superposition ofskeletal elements andepithelial surface structures ofthetongue's tip;rightside: tendinous portions ofMsgandcaudal contour ofG1.lingualis) (Formeaning ofarrows seeFig.46.)A•-A•origins of muscles andtendinous fibers onagoneurosis of Mhga(A:caudal end oforigin ofMmgp, A3apical endoforigin ofMmgp, A• origin oftendinous fibers originating caudolaterally fromtheaponeurosis ofMhgatoattach ondorsal border oflingual nail),EP•andEP3-EP8 insertions ofmuscles andtendinous fibers onepithelial surface ofthetongue's tip(EP•caudal endof insertion ofMmgp,EP•central insertion of Mmga,EP4insertion of tendinous portion C of Msgon •entralareaof thetongue's tip, EPsinsertion of tendinous portionE of Msgoncentral areaof the tongue's tip,EP6insertion of maintendon of portion D of Msg,EP7insertion of tendinous portion D ofMsgoncentral areaofthetongue's tip,EP8direction ofinsertion oftendinous portion C ofMsg onV-shaped mound), G•insertion oftendinous portion A ofMsgonG1.lingualis, NLz-NL3insertions of tendinous fibers of intrinsic muscles onlingual nailandlateralpartof V-shaped mound (NIn insertion oftendinous portion BofMsg,NL3insertion oftendinous fibers originating caudo-laterally fromagoneurosis ofMhga), P•insertion oftendinous portion CofMsgonparaglossale, P6apical edge of Synchondrosis paraglossalis overwhichthetendonof Mhgapasses, T•-Tnorigins of tendinous portions ofMsgfromcentral tendon (T•origin oftendinous portion A, T2origin oftendinous portions C andD, T• originof tendinous portionE, T4originof tendinous portionB) 228 ORNITHOLOGICAL MONOGRAPHS NO 39 + ApMhga I ApMhga-I-TMhga• SPG / NL l C PG• D l•o, 59 Movements of the tongue's tip brought about by the Corpora cavernosa in Psittacus erithacus.A-B: Cross-sections throughthe tongue'stip, adaptedfrom Fig 37A (A: Broadeningand flatteningof deflatedtip of tonguethroughinflation of the Corpuscavernosumlat.; B: Narrowing of inflated tip of tongueand relaxation of central area through deflation ofC c lat.) C-D: Dorsal views of the tongue'stip Left sides:epithelialsurfacestructures;right sides:superpositionof skeletalelements, caudal outline of G1 lingualisand outline of anterior part of the tongue'stip (C: Inflation of the Corporacavernosalateraliaet apicale;D: Deflation ofCc cc latt et ap.) (For meaningof arrowssee Fig 46.) LINGUAL APPARATUS OF PSITTACUS 229 A B I'"' / F•o 60 Late•a]viewsoœthe œu•ctions oœthe M hypoglossus arttenordur•_gmoYementsoœthe paraglossale in Psittacuserithacus.A-B: Contractionof Mhga during flexion of the paraglossale(A: Initial stagewith paraglossaleextended and aponeurosisof Mhga relaxed; B: Final stagewith paraglossaleflexedand aponeurosisof Mhga taut) C-D: Contractionof Mhga during extensionof paraglossale(C: Initial stagewith paraglossale flexedand aponeurosis of Mhga relaxed;D: Final stagewith paraglossale extendedand aponeurosisof Mhga taut) (For meaningof arrowsseeFig 46.) B4 origill of Mhga on basihyale,P, apicaledgeof Synchondrosis paraglossalis over which the tendonof Mhga passes,P7 caudo-dorsalend of insertionof Mhga on paraglossale 230 ORNITHOLOGICAL MONOGRAPHS NO 39 : ! ! Msg Mhgo c9! F]o 61 Lateral viewsof the contractionof the M supraglossus duringretractionof the hyoid and flexion of the paraglossalein Psittacu$erithacu&A: Initial stagewith protractedhyoid, extended paraglossale, taut aponeurosisofM hypoglossus ant., and uncompressed G1 lingualis.B: Final stage with retractedhyoid, flexedparaglossale, relaxedaponeurosisof Mhga, and compressed G1 lingualis (For meaningof arrows seeFig 46.) A• insertion of tendinousportion E of Msg on aponeurosisof Mhga, B•-B4 originsof intrinsic lingual muscleson basihyale(B• caudal end of origin of Msg bas., B4 origin of Mhga), C9 caudal end of origin of Msgc on ceratobranchiale,Gs insertion of tendinous portion A of Msg on GI lingualis, Ps insertion of tendinous portion C of Msg on paraglossale,P6 apical edgeof Synchondrosis paraglossalis over which the tendon of Mhga passes,T•-Ta originsof tendinousportionsof Msg from centraltendon(T• origin oftenclinousportion A, T2 origin oftenclinous portion C, Ta origin of tendinousportion E) LINGUAL APPARATUS OF PSITTACUS 231 Fla 62 Lateral viewsof the contractionof the M supraglossus duringretractionof the hyoid and extensionof the paraglossale in Psittacuserithacus.A: Initial stagewith protractedhyoid, flexed paraglossale, taut aponeurosisof M hypoglossus ant., and compressedG1 lingualis.B: Final stage with retractedhyoid, extendedparaglossale, relaxedaponeurosisof Mhga, and uncompressed G1 lingualis.(For meaningof arrowsseeFig 46.) A• insertionof tendinousportionf of Msgon aponeurosis of M]aga,B3-B4originsof intrinsiclingualmuscles on basihyale (B3caudalendof originof Msgb,Bn origin of Mhga), C9 caudalend of origin of Msgc on ceratobranchiale, Gs insertionof tendinous portion A of Msg on G1 lingualis,Ps insertion of tendinousportion C of Msg on paraglossale,P6 apicaledgeof Synchondrosis paraglossalis over which the tendonof Mhga passes,T,-T3 originsof tendinousportionsof Msg from centraltendon(T• originof tendinousportionA, T2 originof tendinous portion C, T3 origin of tendinousportion f.) 232 ORNITHOLOGICAL MONOGRAPHS NO 39 )% A Msg Mhga i/•G5 • • B '" /' ?._ p \ ',?% /./"/ FI• 63 •ter• •ews of ae •ntmcfion of the M supra½ossus d•g pro•aaion of the hyoid •d flexion of •e •a•os•e in Psitta• erith• A: Iffitial stage• retraced hyoid, extended •ossale, rout a•neurosis of M h•o•ossus •t., •d •mpressed G1 H•is B: Fi• stage • proWa•edhyoid,flexed•a•os•le, relied a•ne•osis ofMh•, •d compressed GI •is (For me•i• of •ows s• Fig 46.) A• in•on of ten•nous •on E of Msg on a•ne•osis of •, B•-Bn o•s of •ffinsic lin• muscleson bas•yale (B• mu• end of od• of Msgb, Bn odin of Mh•), C• mu• end of o•n of Msgc on •ratobmc•e, G• •se•on of ten•nous •on A of Msg on GI •g•Hs, P• insetion of ten•nous •on C of Msg on pa•ossale, P• api• e•e of Synchon•osis •os•is over w•ch the tendon of Mh• •sses, T•-T• od•s of ten•nous •ons ofMsg •om •n• tendon(T• o•n of•n•nous •on A, T2 od• often•nous po•on C, T• odin of ten•nous •nion E) LINGUAL APPARATUS OF PSITTACUS /G (' •,5 233 T3 ! T• ; ! / ! ! ! ! A ! c9: / Msg Mhga Fro 64 Lateral views of the contraction of the M supraglossus during protraction of the hyoid and extensionof the paraglossale in Psittacuserithacus.A: Initial stagewith retractedhyoid, flexed paxaglossale, taut aponeurosisof M hypoglossus ant., and compressedG1 lingualis.B: Final stage with protracted hyoid, extended paxaglossale, relaxed aponeurosisof Mhga, and uncompressedG1 lingualis.(Formeaningof arrowsseeFig.46.) A• insertionoftendinousportionE ofMsgonaponeurosis of Mhga, B3-B4originsof intrinsiclingual muscleson basihyale(B3caudalend of origin of Msgb, B4 origin of Mhga), C9 caudal end of origin of Msgc on ceratobranchiale,G• insertion of tendinous portionA of Msg on paraglossale, P6apicaledgeof Synchondrosis paxaglossalis over whichthe tendon of Mhga passes,T•-T3 originsof tendinousportionsof Msg from centraltendon(T• originof tendinous portion A, T2 origin of tendinousportion C, T3 origin of tendinousportion E) ORNITHOLOGICAL MONOGRAPHS No A Distributional StudyoftheBirdsof BritishHonduras, byStephen M Russell.1964 $7.00 ($5.50 to AOU members) No A Comparative Studyof SomeSocialCommunication Patternsin the Pelecaniformes, by GerardFrederick vanTets.1965.$3.50($2.50to AOU members) No TheBirdsof Kentucky, by RobertM Mengel.1965.$15.00($12.50to AOU members) No A Comparative Life-history Studyof FourSpecies of Woodpeckers, by Louisede Kiriline Lawrence.1967.$6.00 ($4.50 to AOU members) No Adaptations for Locomotion andFeeding in theAnhingaandtheDouble-crested Cormorant, by OscarT Owre.1967.$6.00($4.50to AOU members) No A DistributionalSurveyof the Birdsof Honduras,by Burt L Monroe, Jr 1968 $14.00 ($11.00 to AOU members) No MatingSystems, SexualDimorphism, andthe Roleof Male North AmericanPasserineBirds in the NestingCycle,by JaredVerner and MaryF Willson.1969.$4.00($3.00to AOU members) No 10 The Behaviorof SpottedAntbirds,by EdwinO Willis 1972.$9.00 ($7.50 to AOU members) No 11 Behavior, MimeticSongsandSongDialects,andRelationships of the Parasitic Indigobirds (Vidua)ofAfrica,byRobert B.Payne 1973.$12.50 ($10.00 to AOU members) No 12 Intra-isiand Variationin theMascarene White-eyeZosterops borbonica, by FrankB Gill 1973.$3.50($2.50to AOU members) No 13 Evolutionary Trendsin theNeotropical Ovenbirds andWoodhewers, by Alan Fcduccia.1973 $3.50 ($2.50 to AOU members) No 14 A Symposium ontheHouseSparrow (Passer domesticus) andEuropean TreeSparrow (P montanus) in NorthAmerica, byS.Charles Kendeigh 1973 $6.00 ($4.50 to AOU members) No 15 FunctionalAnatomyand AdaptiveEvolutionof the FeedingApparatus in the HawaiianHoneycreeper GenusLoxops(Drepanididae), by Lawrenee P Richardsand Walter J Bock 1973 $9.00 ($7.50 to AOU members) No 16 No 17 The Red-tailedTropicbirdon Kure Atoll, by RobertR Fleet 1974 $5.50 ($4.50 to AOU members) Comparative Behavior of the American Avocetandthe Black-necked Stilt(Recurvirostridae), byRobertBruce Hamilton.1975.$7.50($6.00 to AOU members) No 18 No 19 No 20 No 21 Breeding Biology andBehavior oftheOldsquaw (Clangula byemalls L.), by RobertM Alison.1975.$3.50($2.50to AOU members) BirdPopulations of AspenForests in WesternNorthAmerica, byJ A DouglasFlack.1976.$7.50($6.00to AOU members) SexualSizeDimorphism in HawksandOwlsofNorthAmerica,byNoel F R.Snyder andJames W.Wiley.1976.$7.00($6.00toAOUmembers) SocialOrganization andBehavior of theAcornWoodpecker in Central CoastalCalifornia,by MichaelH MacRobertsand BarbaraR MacRoberts 1976 $7.50 ($6.00 to AOU members) (Continued on insideback cover) No.22.Maintenance Behavior andCommunication intheBrown Pelican, by Ralph W.Schreiber 1977.Price $6.50($5.00toAOUmembers) No 23 Species Relationships intheArianGenus ,'limopM!a, byLarryL.Wolf 1977.Price$12.00($10.50toAOU members) No 24 LandBirdCommunities of GrandBahama Island:TheStructure and Dynamics ofanAvifauna, byJohnT Emlen 1977.Price$9.00($8.00 to AOU members) No 25 Systematics ofSmaller Asian NightBirdsBased onVoice, byJoeT Marshall 1978.Price$7.00($6.00toAOUmembers) No.26 Ecology andBehavior ofthePrairie Warbler Dendroica discolor, byVal Nolan, Jr 1978 Price $29.50 No 27 Ecology andEvolution ofLekMating Behavior intheLong-tailed HermitHummingbird, byF GaryStiles andLarryL Wolf.viii q-78pp., 26textfigures 1979.Price$8.50($7.50toAOUmembers) No 28 TheForaging Behavior ofMountain Bluebirds withEmphasis onSexual Foraging Differences, byHarryW.Power x q-72pp.,color frontispiece, 12textfigures 1980.Price$8.50($7.50toAOUmembers) ' No 29 TheMolt of ScrubJaysandBlueJaysin Florida,byG Thomas BancroftandGlenE Woolfenden 1982.Price$8.00($6.50toAOU members) No 30 ArianIncubation: EggTemperature, NestHumidity, andBehavioral Thermoregulation in a Hot Environment, by GilbertS Grant 1982 Price$9.00($7.00to AOU members) No 31 TheNativeForest BirdsofGuam, byJ.MarkJenkins 1983.Price$9.00 ($7.00 to AOU members) No: 32.TheMarine Ecology ofBirds intheRoss Sea,Antarctica, byDavidG Ainley,EdmundF O'Connor, andRobertJ Boekelheide 1984.Price $9.00 ($8.00 to AOU members) No 33 SexualSelection, Lek andArenaBehavior, andSexualSizeDimorphismin Birds,byRobertB.Payne.1984.Price$8.00($6.50toAOU members) No 34 Pattern, Mechanism, andAdaptive Significance ofTerritoriality inHerringGulls (Larus argentatus), byJoanna Burger 1984 Price $9.00($7.00 to AOU members) No 35.Ecogeographic Variation inSizeandProportions ofSong Sparrows (Me!osplza melodia), byJohnW.Aldrich.1984.Price$10.50($8.50toAOU members) No 36 Neotropical Ornithology, P.A Bucldey, Mercedes S.Foster, Eugene S Morton,RobertS.Ridgely, andFrancine G Bucldey (eds.).1985.Price $70.00 No 37 ArianMonogamy, byPatricia A Gowaty andDouglas W.Mock.1985 Price$11.00($9.00to AOU members) No 38 An Analysis of Physical, Physiological, andOpticalAspects of Avian Coloration withEmphasis onWood-warblers, byEdward H Burtt,Jr 1986.Price$15.00($12.50to AOU members) Likeallother AOUpublications, Ornithological Monographs areshipped prepaid Make checks payable to "TheAmerican Ornithologists' Union."Fortheconvenience ofthose whowishto maintain complete setsof Ornithological Monographs andto receive new numbers immediately uponissue, standing orders areencouraged Order from: Frank R.Moore, Assistant totheTreasurer AOU,Department ofBiology, University ofSouthern Mississippi, Southern Station Box5018, Hattiesburg, Mississippi 39406 ... Station Box 5018, Hattiesburg, Mississippi 394 06 (See price list on back and inside back covers.) OrnithologicalMonographs,No 39, xi + 233 pp Editor of AOU Monographs, Mercedes S Foster and David... State University Baton Rouge, Louisiana 70803 ORNITHOLOGICAL MONOGRAPHS PUBLISHED THE AMERICAN BY ORNITHOLOGISTS' WASHINGTON, 1986 NO D.C UNION 39 TABLE LIST OF FIGURES LIST OF TABLES LIST ... functional interpretationswere basedon intuitive considerations.Functional analyses,such ORNITHOLOGICAL MONOGRAPHS NO 39 as these, for the most part proved to be inaccurate or even incorrect becausethe