A Systematic Study of the Main Arteries in the Region of the Heart, Aves Vi

Vol.6o'] GLœNNY,A Study o! Arteriesin theRegion o! theHeart 235

A SYSTEMATIC STUDY OF THE MAIN ARTERIES IN THE REGION OF THE HEART, AVES VI. TROGONIFORMES, PART 1. BY FRED H. GLENNY •

INTRODUCTION IN severalrecent papers,the writer has presentedthe important featuresand arrangementsof the main arteriesin the neck and thorax of several families and orders of birds. In addition to the functional vesselsof the adult, the writer hasincluded notations on the persistence of certain vestigesof embryonicvessels. During thesestudies, the writer has observedthat within orders and even familiesof birds there is a certain constancyin the arrangement-patternof the main arteriesin the regionof the neckand thorax of birds. In the presentpaper, the writer presentsthe basicarrangement-patternof the arteriesas found in severalspecies of trogons. Materialsfor this studywere made availablethrough the co6opera- tion of Dr. AlexanderWetmore, AssistantSecretary, Smithsonian In- stitution, and Dr. Herbert Friedmann, Curator, Division of Birds, United States National Museum.

Singlespecimens of eightspecies of trogonswere dissected and the arrangementof the arteriescompared. Spedmensof Chrysotrogon caligatus (Gould), Priotelus temnurus (Temminck), Pyrotrogon erythrocephalus(Gould), Temnotrogonroseigaster (Vieillot), Curu- cujus (Trogon) massena(Gould), Trogon melanocephalusGould, Trogonstrigilatus Linn& and Trogonurusvariegatus behni (Gould) were included in this study. The information thus obtained is set forth in the following observations.

OUSERVATIONS The basicarrangement of the main arteriesin the neckand thorax is essentiallythe samein eachof the speciesstudied. The left and right innominatearteries arise from the aortic root and passanteriorly and laterallyto divide into the commoncarotid and subclavianarteries. The right systemicarch alonepersists as the functional 4th aortic arch which joins the right radix aortae. The subclavianarteries each receive the coracoldmajor, axillary, inter-

a On active servicewith the Royal Canadian Army Medical Corps. 236 Studyo[ Arteriesin theRegion o[ theHeart IL'Auk April costal, and two pectoral arteries in order. The intercostal artery arisesfrom the subclavianartery at the baseof the posteriorpectoral artery. The left commoncarotid artery givesrise to the ductusshawl (a tracheo-oesophagealartery), thyroid, internal carotid,superficial cervical, and vertebral arteries. The ductus shawl sends off branches to the syrinx,trachea, and oesophagusanteriorly and to the oesophagus and accessorytissues posteriorly. The left internal carotid artery aloneenters the hypapophysialcanal and passesanteriorly along the median-ventralline of the neck. The superficialcervical artery gives rise to the vertebral and basi-cervicalarteries before passinganteriorly to supplythe musculatureof the neck. The vertebralartery passes dorsally,in the regionof the brachialplexus, and anteriorly. The basi-cervicalartery passesposteriorly to the musculatureof the shoulder. The thyroidartery arises at the point of bifurcationof the commoncarotid artery. The right commoncarotid artery givesrise to the basi-cervical, thyroid, ascending-oesophageal,superficial cervical, vertebral, and ductus shawi arteries. The right basi-cervicalartery suppliesthe musculatureof the right shoulder. The right ductusshawl supplies the right sideof the tracheaand syrinxas well as the baseof the oesophagus.The ascending-oesophagealartery is comparableto the left internal carotidand servesto supplythe right face of the oeso- phagusas well as the muscleson the right sideof the neck. The superficialcervical of the right sidemay or may not be present,but when presentgenerally arises as a small branchof the ascending- oesophagealartery. The left ligamenturnaortae is presentand entire in Chrysotrogon caligatus,Priotelus temnurus, Pyrotrogon erythrocephalus, and Tro- gonurusvariegatus behni; and in Temnotrogonroseigaster, Curucujus massena,Trogon melanocephalus,and Trogon strigilatusthe distal portionof the radix presentsat leasta shortlumen while the anterior portion persistsas the ligamenturnaortae. The right ligamenturnbotalli is minute and tendsto fusewith the right radix aortae in Chrysotrogoncaligatus, Pyrotrogon erythrocephalus, Temnotrogonroseigaster, Curucujus massena, and Trogon strigilatus. In Trogonurusvariegatus behni the embryonicductus arteriosusremains only as a small ligamentous'button' and marks the distal attachmentof the ductusto the right radix, while the liga- menturn botalli is very prominent in Trogon melanocephalusand absent(probably entirely fused to the radix) in Priotelustemnurus. Vol.60] GLENNY,A Study O[ Arteries in theRegion of theHeart 237

TEXT-FIG.l.--Diagrammatic representation of the main arteriesin the neck and thorax of Trogon melanocephalus.Ventral view. a.o., ascending-oesophagealartery. l.r.a, ligamenturnaortae. a.r., aortic root. pec, pectoral arteries. a.s.c.,superficial cervical artery. p.s.c.,basi-cervical artery. ax, axillary artery. put., pulmonary artery. c.c., common carotid artery. r.l.b., right ligamentum botalli. cor, coracoldmajor artery. r.r.a., right radix aortae. d.a., dorsal aorta. r.s.a., right systemic(4th aortic) arch. d.i.c., vertebral artery. sub, subclavianartery. d.s., right ductus shawl t, thyroid artery. e.c., internal carotid artery. t.o., left ductus shawl im, intercostal artery. (tracheo-oesophagealartery). in, innominate artery. tr, tracheal branches of ductus shawl l.l.b., left ligamenturnbotalli. 238 GLENNY,Zl Study O! Arteries in theRegion o! theHeart Li'Auk April

DISCUSSION From the above evidences,we may concludethat the speciesoœ trogonsrepresented in this study show a characteristicrelationship basedon the arrangement-patternoœ the main arteriesin the neckand thorax. Variation in the degreeoœ persistence of the left radix aortae and the right ductusbotalli as ligamentousvestiges appears to be a specificor an individual matter and doesnot appear to have any genericvalue. Although there are definite pattern-variationsin the Coraciiformes, Anseriformes,Ciconiiformes, and othergroups oœ birds, this is not the casein the trogonsin sofar as they couldbe representedin this study. It may be observedthat the left internal carotidartery alone enters the hypapophysialcanal, whereasin someof the Coraciiformes,An- $eriformes,Strigiformes, etc., both left and right internal carotidsare present and enter the canal. In the opinion oœthe writer, the trogonspresent a significantly characteristicpattern oœthe arteriesin the neck and thorax region.

1. Eight speciesof trogonswere dissectedand drawingsof the arrangement-patternof the arteries in the neck and thorax prepared. 2. A singlebasic arrangement-pattern was found to be characteristic for the family. 3. Only slight variationsin persistenceof the ligamentousvestiges of the left radix aortaeand right ductusbotalli were recorded. 4. The left internal carotidartery alone enteredthe hypapophysial canal. ]•EFEREN CES BEDDARD,F. E. 1898. The Structure and Classificationof Birds. Longmans, Green and Co., London. BH^DURI,J. L. 1939. A caseof persistenceof the ductus caroticusin the Indian Blue Rock- pigeon Columbalivia intermedia (Strick.). Anat. Anz., 88: 178-182. G•,Dow, HANs 1891. Bronn's'Klassen und Ordnungendes Thief-Reichs', Leipzig, SechsterBand. Vierte Abtheilung. V6gel: 768-780. GARROD,A. H. 1873. On the Carotid Arteries of Birds. Proc. Zoo1. Soc. London: 457-472. GLENNy, F. H. 1939. An anomalousartery in the Kingfisher (Ccryle alcyon). Ohio ,Jour.Sci., 39 (2): 94-96. 1940. A systematicstudy of the main arteriesin the regionof the heart--AvesI. Anat. Rec., 76 (4): 371-380. Vol.0o' I E•)wAm)s,Hearing Ranges o[ FourSpecies o[ Birds 239 •94• -•

1941. Presenceof the ligamenturnbotalli in the GoldenEagle, Red-tailed Hawk, and the commonPigeon. Ohio Jour. Sci.,41 (1): 46-49. 1942. A systematicstudy of the main arteriesin the regionof the heart--AvesIII. Ohio Jour. Sci.,42 (2): 84-90. 1942. Main Arteriesin the Regionof the Neck and Thorax of the Australian Gassowary.Gan. Jour. Res., 20 (12): g6g-g67. 1948. A systematicstudy of the mainarteries in the regionof the heart--AvesIV. [Jour. Anat. (in press)]. HOCHSTETTER•F. 1890. t3ber den Ursprungder Arteria subclaviader V6gel. Morph. Jahrb., 16: 484. Bird ResearchLaboratory Summit County Wildlife ResearchStation Akron, Ohio

HEARING RANGES OF FOUR SPECIES OF BIRDS

BY ERNEST P. EDWARDS IN continuingthe experimentson the hearingof birds which the late Albert R. Brand and Prof. P. P. Kelloggbegan at Cornell Uni- versityin 1958,observations have recently been made on the frequency rangeof hearingof four otherspecies of wild birdsin captivity. The apparatusused in the earlier experimentswas placed at the author's disposalfor the presentexperiments, and hasmade up a major part of the equipmentused. This seriesof experimentswas begun in March, 1941,and is still in progress.The objectof the work is to determinethe frequency rangesof soundsto whichdifferent birds will respond. Observations have been completedon six captiveindividuals representing four specieswhose hearing ranges had not theretoforebeen investigated. Theseare asfollows: one Canvas-back(Nyroca valisineria); one Great Horned Owl (Bubo virginianus);three Prairie Horned Larks (Oto- corisalpestris praticola); one SnowBunting (Plectrophenaxnivalis). An electricshock was usedin teachingthe birds to give a definite responsewhen the soundstimulus was applied. Within onesecond aftera pure tonewas sounded close to the bird'scage, the bird was givenan electricshock, and it soonlearned that whenthe tonewas sounded,a shockwas imminent. Therefore, after a number of trials (usually30 to 60) thebird beganto respondto the soundas it did to the shock--byjumping or fluttering. A beat-frequencyoscillator was usedto producesounds with no overtones.Tones with frequenciesfrom 20 to 17,000vibrations per