Author Institution: Department of Zoology and Entomology, The Ohio State University, Columbus 1

Glenny, Fred H.

The Ohio Journal of Science

Author Institution: Department of Anatomy, The Ohio State University, Columbus 1

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A SYSTEMATIC STUDY OP THE MAIN ARTERIES IN THEREGION OF THE HEART. AVES XIII.ClCONIIFORMES, PART 21FRED H. GLENNYDepartment of Anatomy, The Ohio State University, Columbus 10Further studies on the arrangement-pattern of the main arteries in the neckand thorax of 19 species of Ciconiiformes reveals the presence of a basic ordinalarrangement of these vessels, and further substantiates the view of Glenny (1945)that there are notable steps in the evolution of the carotids from a bicarotid tounicarotid condition. As has been shown by Garrod (1873) and Glenny (1945),union of the carotid (trunk) arteries results in the conjugate condition. Subsequentreduction of the basal portion of either side (usually the left side in the Ciconii-formes) may lead to complete loss of the reduced vessel.It may be noted that there is not any notable uniformity in occurrence of thesereductions with respect to generic status at the present time. As a result, bicarotidand conjugate carotids may occur in different species of a single genus. Further-more, these variations may occur in the several families and subfamilies sincethey are not restricted to any single grouping of genera.The characteristic ordinal arrangement-pattern for the Ciconiiformes is asfollows; bicarotidinae-normales (with variations); ligamentum aortae present;ligamentum botalli reduced or lacking; branches of subclavian artery: coracoidor sterno-clavicular, axillary, internal mammary or intercostal, two pectorals;branches of the common carotid: ductus shawi and tracheal, thyroid, superficialcervical (comes nervi vagi), vertebral, internal carotid (trunk).The following species have been studied, and differences in the arrangement-pattern noted. Unless otherwise noted, the following species present the patterncharacteristic of the order.ARDEIDAE:Ardea herodias treganzai Court: conjugate carotids, ligamentum botalli present but reduced.Ardea herodias herodias (Linnaeus): bicarotid, ligamentum botalli present.Butorides virescens virescens (Linnaeus): conjugate carotids, ligamentum botalli lacking.Butorides sundevalli Reichenow: bicarotid, ligamentum botalli lacking.Florida caerulea (Linnaeus): bicarotid, no ligamentum botalli.Ardeola speciosa (Horsfield): conjugate carotids, left side reduced to about x/i that of theright side, no ligamentum botalli.Bubulcus ibis coromandus (Boddaert): bicarotid, ligamentum botalli much reduced andpartial.Bubulcus ibis ibis (Linnaeus): same as above.Dichromanassa rufescens rufescens (Gmelin): same as above.Casmerodius albus albus (Linnaeus): same as above.Demiegretta sacra (Gmelin): same as above, small ventro-lateral superficial cervicals alsoarise from base common to thyroids.Leucophoyx thula brewsteri (Thayer & Bangs): conjugate carotids, ligamentum botalliabsent.Hydranassa tricolor ruficollis (Gosse): bicarotid, ligamentum reduced or fused to radixaortae.Contribution of Blue Sea Lake Biological Laboratory, Messines, Quebec.THE OHIO JOURNAL OF SCIENCE 53(6): 347, November, 1953.348 FRED H. GLENNY Vol. L I UNyctanassa violacea pauper (Sclater & Salvin): bicarotid, ligamentum botalli reducedto a white linea botalli on the face of the right radix aortae.Ixobrychus minutus minutus (Linnaeus): bicarotid, ligamentum botalli present.Ixobrychus sinensis bryani (Seale): bicarotid, ligamentum botalli absent.Botaurus lentiginosus (Montagu): conjugate carotids, ligamentum botalli lacking.COCHLEARIIDAE:Cochlearius cochlaerius zeledoni (Ridgway): bicarotid, linea botalli present.THRESKIORNITHIDAE:Guara alba (Linnaeus): bicarotid, ligamentum botalli short.The writer wishes to express his gratitude to Dr. Herbert Friedmann, Curator,Division of Birds, U. S. National Museum, and to Mr. J. W. Cowland, Curator,Sudan Government Museum, Natural History, Wad Medani (Sudan), for theirassistance in making available the materials for this study.REFERENCESGarrod, A. H. 1873. On the carotid arteries of birds. London, Proc. Zool. Soc, 457-472.Glenny, F. H. 1945. A systematic study of the main arteries in the region of the heart—AVES XIII. Ciconiiformes, Part 1. Amer. Mid. Nat., 33: 449-454.

A Systematic Study of the Main Arteries in the Region of the Heart. Aves XIII : Ciconiiformes, Part 2

A SYSTEMATIC STUDY OF THE MAIN ARTERIES IN THEREGION OF THE HEART—AVES XXCAPRIMULGIFORMES, PART I1FRED H. GLENNYDepartment of Anatomy, The Ohio State University, Columbus 10Studies on 12 species of Caprimulgiformes, shows the basic arterial arrangement-pattern in the neck and thorax to be that referred to as bicarotid-normales. Thelaevo-carotid variation was observed, however, in representatives of two familieswithin the order. Furthermore, an evolutionary tendency sometimes observedin a few other orders of birds was noted in the different families of this order. Thischaracteristic tendency is the rather singular lateral migration of the sterno-clavicular or coracoid arteries from a "normal position on the subclavian betweenthe common carotid and axillary arteries to one on the pectoral stem just lateralto the axillary and medial to the internal mammary or intercostal artery.The following observations characterize the materials studied.STEATORNITHIDAE:Steatornis caripensis Humboldt (USNM #18308): bicarotid; both ligamentum aortae andligamentum botalli present; coracoid arises from subclavian medial to origin of axillary; ductusshawi sends branches to oesophagus and trachea as well as posteriorly to bronchi and connectivetissues of the thorax; common carotid then sends off a short thyroid artery before dividing toform a superficial cervical (ascending oesophageal on the right side), vertebral, and internalcarotid (trunk) artery.Garrod (1873) observed that both carotids are present in Steatornis caripensis.PODARGIDAE:Laevo-carotidinae; ligamentum aortae present; ligamentum botalli reduced to a linea orentirely lacking; coracoid opposite or just lateral to axillary; right common carotid reducedgiving rise to an ascending oesophageal artery in addition to the normal superficial cervicaland finally the vertebral artery; left carotid divides to form the superficial cervical, vertebral,and internal carotid (trunk) arteries; ductus shawi and thyroid artery arise as branches ofboth left and right carotids.Podargus strigoides strigoides (Latham) (USNM #19361): the coracoids are opposite theaxillaries.Batrachostomus hodgsoni indochinae Stresemann (USNM #321751): the coracoids are justlateral to the axillaries.NYCTIBIIDAE:Nyctibius griseus (Gmelin) (USNM #346233): laevo-carotidinae; ligamentum aortae present;coracoid lateral to axillary; ductus shawi prominent; common carotid divides into superficialcervical and vertebral arteries (right side) and superficial cervical, vertebral and internalcarotid (trunk) arteries after giving off the thyroid artery.CAPRIMULGIDAE:Bicarotid normales; ligamentum aortae present; ligamentum botalli present and reducedor lacking; coracoids medial or lateral to axillary; ductus shawi and thyroid artery arise asbranches of carotid before it divides to form the superficial cervical, vertebral, and internalcarotid (trunk) arteries.Contribution of Blue Sea Lake Biological Laboratory, Messines, Quebec.THE OHIO JOURNAL OF SCIENCE 53(6): 356, November, 1953.No. 6 MAIN ARTERIES NEAR THE HEART 357Coracoid artery medial to axillary artery:Chordeiles virginianus = Chordeiles minor minor (Forster), Podager nacunda nacunda(Vieillot), Nyctidromus albicollis albicollis (Gmelin), Scotornis fossii fossii (Hartlaub), andHydropsalis brasiliana furcifera (Vieillot).Coracoid artery opposite or just lateral to axillary artery: Chordeiles minor vicinus Riley.Coracoid artery lateral to axillary artery:Nyctiprogne latifascia (USNM #319819) and Caprimulgus carolinensis Gmelin.Garrod (1873) pointed out that Chordeiles acutipennis texensis Lawrence and Caprimulguseuropaeus Linnaeus both present two carotid arteries which enter the hypapophysial canal.It may be concluded, from the above observations, that the members of thisorder and especially the families within the order are undergoing noticable ana-tomical evolution especially with respect to the arrangement of arteries in theneck and thorax. Such major differences as are herein noted may have sometaxonomic significance.The writer is indebted to Dr. Herbert Friedmann, U. S. National Museum,for making available most of the materials used in this study.REFERENCEGarrod, A. H. 1873. On the carotid arteries in birds. Proc. Zoo. Soc, London, pp. 457-472.

A Systematic Study of the Main Arteries in the Region of the Heart Aves XX : Caprimulgiformes, Part 1

A SYSTEMATIC STUDY OF THE MAIN ARTERIES IN THEREGION OF THE HEART. AVES XVI.CHARADRIIFORMES, PART 2lFRED H. GLENNYDepartment of Zoology and Entomology, The Ohio State University, Columbus 10Recent continued studies on the arteries in the neck and thorax of seven speciesof Charadriiformes were made possible by the loan of alcoholic specimens from theanatomical collections of the Chicago Natural History Museum and the U. S.National Museum.The writer wishes to express his gratitude to Mr. Dwight D. Davis, Dr. HerbertFriedmann, and Dr. Alexander Wetmore for their help in making these materialsavailable for this study.Except where otherwise indicated, single specimens were dissected and diagramsof the arterial arrangement-patterns prepared.MATERIALSSuborder CHARADRIIFamily Charadriidae ;Belonopterus chilensis lampronotus (Wagler) (CNHM #106097).Suborder LARIFamily LaridaeRissa tridactyla pollicaris Ridgway (USNM #81523).Suborder ALCAEFamily AlcidaeUria aalge inornate Salomonsen (USNM #81379).Synthliborhamphus antiquus (Gmelin) (USNM Nos. 81608, 81707, 18317 to18321 incl.)Synthliborhampus wumizusume (Temminck) (USNM #87266 & 87267).Aethia cristatella (Pallajs) (USNM #18313).Cydorrhynchus psittacula (Pallas) (USNM #18314 & 18315).OBSERVATIONSThe arterial arrangement-pattern of Belonopterus chilensis lampronotus isalmost identical with that of Charadrius vociferus vociferus (Oxyechus v. vociferus)(Glenny, 1947) except that the origin of the sterno-tracheal artery could be deter-mined in Belonopterus where it arises as a branch of the coracoid major artery;no vestigeal right ligamentum botalli could be accurately located; origin of theleft vertebral artery was separate from, but close to the base of, the superficialcervical artery (comes nervi vagi) which gives rise to an accessory oesophagealartery.Rissa tridactyla pollicaris is similar to Larus argentatus (Glenny, 1947) in basicarrangement, but differs in the following respects: the right ligamentum botalliis short, the proximal portion becoming completely atrophied; the sterno-trachealartery arises as a branch of the coracoid major artery; the superficial cervicalarteries (comes nervi vagi) arise separately (not in common with the vertebrals)from the common carotid arteries.Uria aalge inornata follows the basic ordinal pattern of the Charadriiformes(Glenny, 1948). The superficial cervical and vertebral arteries arise separatelyContributions from the Blue Sea Lake Biological Laboratory, Messines, P. Q., Canada.THE OHIO JOURNAL OF SCIENCE 52(6): 314, November, 1952.No. 6 ARTERIES NEAR HEART 315from the common carotid arteries. An accessory oesophageal artery arises as abranch of the left common carotid. The ligamentum aortae is present, althoughthe right ligamentum botalli appears to be entirely lacking.Except for the accessory oesophageal artery, which appears to be absent, thearrangement-pattern of Synthliborhamphus wumizusume is like that of Uria aalgeinornata (above).Basically, Synthliborhamphus antiquus is like that of ,5". wumizusume andUria aalge inornata, except that in one type (bicarotidinae) (USNM #18317 &18318) an accessory superficial cervical artery was observed to arise in commonwith the left vertebral artery (Glenny, 1952). The ligamentum botalli appearedto be completely lacking in this species. A second type of arrangement (laevo-carotidinae) (fig. 2, Glenny, 1952) was observed in five specimens (USNM #s81608, 81707, 18319-18321). In these specimens, the basic ordinal pattern—asseen in .5. wumizusume—was characteristic except for the very singular fact thatthe right internal carotid trunk was lacking. The embryonic connection of theright third aortic arch with the anterior dorsal radix of the right side had becomelost, and the ventral portion of this arch became functionally modified as an acces-sory vertebral artery—passing dorsally and anteriorly just anterior to the normalvertebral artery (in the region of the brachial plexus). Such a functional modifica-tion differs somewhat from that, of this same structure, in Tockus alboterminatusalboterminatus (Buttikofer) (Lophoceros melanoleucos alboterminatus) (Glenny,1943) wherein the vessel in question comes to serve the ventro-cervical musclesat the base of the hypapophysial canal.Cyclorrhynchus psittacula differs from the Alcidae included in this study inthat the vertebral and superficial cervical arteries have a common origin from thecommon carotid artery, an accessory oesophageal artery arises as a branch of theleft common carotid, a coracoid minor artery is found in addition to and justlateral to the coracoid major artery, only the proximal portion of the rightligamentum botalli could be found as it maintains its proximal connection with thepulmonary artery, and finally the left radix aortae was found to be patent andfunctional (several small twigs were observed branching from the radix in muchthe same manner as was reported in Ceryle alcyon) (Glenny, 1939, 1943) with aterminal ligamentous portion attached to the left pulmonary artery.The arterial arrangement-pattern in Aethia cristatella is very similar to that ofCyclorrhynchus psittacula except that the common origin of the superficial cervicaland vertebral arteries is not so pronounced or may be separate, and the ligamentumbotalli is lacking. Both the coracoid major and minor arteries were present.DISCUSSIONWhile the Charadriiformes are fundamentally bicarotid as pointed out byGarrod (1873), Beddard (1898), and Glenny (1947, 1948), it may be observedthat this condition may be altered in cases such as has been observed inSynthliborhamphus antiquus.Bremer (1928) has shown that the normal loss of the left fourth aortic archin the chick is due chiefly to mechanical forces. It may well be reasoned thatsome of the functional modifications, along with the atrophy or complete loss ofcertain of the embryonic vessels or their derivatives may be due in large part tosome, as yet unexplained, mechanical force or forces. These may be inherentcharacters which have become reasonably well established within a strain whichmight develop under conditions of "breeding-site isolation" such as may occurin the Aleutians and other islands along the coast of Alaska. Such may be thecase in the instance of Synthliborhamphus antiquus as reported in this paper.In general, the Alcidae show a high degree of similarity in their arterialarrangement-patterns. Furthermore, while some differences are found in the316 FRED H. GLENNY Vol. LI Iseveral suborders and families, the basic ordinal arrangement holds, except invery special instances.A trophic gradients appear to function at different levels in different species,and the level of atrophy of any vessel is not to be regarded as characteristic ofeither a suborder or family, but rather more as a specific characteristic.REFERENCESBeddard, F. E. 1898. The Structure and Classification of Birds. Longmans, Green & Co.,London.Bremer, J. L. 1928. Experiments on the aortic arches in the chick. Anat. Rec, 37: 225-254.Garrod, A. H. 1873. On the carotid arteries of birds. Proc. Zoo. Soc. London, pp. 457-472.Glenny, F. H. 1939. An anomalous artery in the kingfisher (Ceryle alcyon). Ohio Jour.Sci., 39: 94-96.Glenny, F. H. (series). A systematic study of the main arteries in the region of the heart.. 1943. Aves VII. Coraciiformes, Part 1. Trans. R.S.C., Sec. V, 37: 35-53.. 1947. Aves XVI. Charadriiformes, Part 1. Ohio Jour. Sci., 47: 152-154.. 1948. Aves XVI. Charadriiformes, Part 2. Ohio Jour. Sci., 48: 194-198.Glenny, F. H. 1952. The problem of specific and subspecific status and morphologic deviationin the Ancient Murrelet Synthliborhatnphus antiquus (Gmelin). Ohio Jour. Sci., 52: 221-223.

A Systematic Study of the Main Arteries in the Region of the Heart. Aves XVI. Charadriiformes, Part 2

A SYSTEMATIC STUDY OF THE MAIN ARTERIES IN THEREGION OF THE HEART—AVES XIXAPODIFORMES, PART I1FRED H. GLENNYDepartment of Anatomy, The Ohio State University, Columbus 10Recent studies on 39 species of Apodiformes, with respect to the arterial arrange-ment-pattern in the neck and thorax tend to substantiate the observations ofGarrod (1873) on members of this order of birds. Garrod showed that whilethe left carotid alone persists as the chief cephalic supply in most of the Apodiformesa few species present the bicarotid condition commonly found in most ordersof birds.. 1 Ft*.FIGURES 1-3. Diagrammatic representation of the main arteries in the neck and thoraxof: 1. Nephoecetes and Streptoprocne. 2. Right side: Aeronautes. 3. Right side: Apus.KEY TO ABBREVIATIONS1. aortic root 10.2. innominate artery 11.3. right systemic arch 12.4. right radix aortae 12a.5. ligamentum aortae 13.6. subclavian artery 14.7. coracoid or sterno-clavicular 15.8. axillary 16.9. intercostal or internal mammarypectoraliscommon carotidvertebralaccessory vertebralsuperficial cervical (comes nervi vagi)internal carotid (trunk)ductus shawitracheal.In the Apodiformes, the coracoid or sterno-clavicular artery arises as the firstbranch of the subclavian which then gives rise to the pectoral branch as the sub-clavian continues as the axillary artery. The first branch of the pectoral stemis the intercostal or internal mammary artery. The common carotid then sendsContribution of Blue Sea Lake Biological Laboratory, Messines, Quebec.THE OHIO JOURNAL OF SCIENCE 53(6): 367, November, 1953.368 FRED H. GLENN Y Vol. LI 11off a thyroid artery and further divides to form the superficial cervical (comesnervi vagi), vertebral, and carotid trunk artery. In most Apodiformes, the leftcarotid alone enters the hypapophysial canal. In one notable instance, the basalportion of the right internal carotid becomes modified as an accessory vertebralartery (fig. 2, 12a). In a few other species, especially of the Apodidae, the bicarotidcondition persists in the adult. In the Trochilidae, two coracoid or sterno-clavi-cular arteries were found to arise from the face of the subclavian before the axillaryartery passed anterolaterally to the wing. The ligamentum aortae is present,but unless otherwise noted, the ligamentum botalli is lacking.APODIDAE:Collocalia inexpectata bartschi Mearns: laevo-carotidinae.Streptoprocne zonaris pallidifrons (Hartert): bicarotid.Chaetura pelagica (Linnaeus): laevo-carotidinae; ascending oesophageal artery arises fromcommon carotid after origin of vertebral on the right side.Nephoecetes niger niger (Gmelin): bicarotid.Apus andecolus andecolus (d'Orbigny & Lafresnaye): laevo-carotidinae.Aeronautes saxatalis saxatalis (Woodhouse) 2 specimens: two right vertebral arteries;laevo-carotidinae.Tachornis phoenicobia phoenicobia Gosse: laevo-carotidinae.HEMIPROCNIDAE:Hemiprocne comata major (Hartert): laevo-carotidinae.TROCHILIDAE:No major differences in arterial pattern are found in the species listed below. Onlyapparent differences due to reduction in size of the system and length of vessels may be noted,such "apparent" differences are not real differences and are therefore not mentioned herein.The following species show the same arrangement-pattern for arteries in the neck and thorax.Glaucis hirsuta hirsuta (Gmelin).Phaethornis sp. (U. S. National Museum #225016).Florisuga mellivora mellivora (Linnaeus).Anthracothorax dominicus (Linnaeus).Eulampis jugularis (Linnaeus).Sericotes holosericeus holosericeus (Linnaeus).Chrysolampis mosquitus (Linnaeus).Orthorhynchus cristatus exilis (Gmelin).Lophornis sp. (U. S. National Museum #81493).Chlorostilbon canivetti caribaeus Lawrence.Chlorostilbon ricordii ricordii (Gervais)Cynanthus latirostris latirostris Swainson.Thalurania furcata colombica (Bourcier).Lepidopyga goudoti goudoti (Bourcier).Hylocharis xantusii (Lawrence).Hylocharis chrysura (Shaw).Trochilus polytmus polytmus Linnaeus.Leucippus fallax fallax (Bourcier).Amazilia tobaci (U. S. N. M. #346366) zoo specimen.Amazilia tzacatl tzacatl (De la Llave).Chalybura buffonii aeneicauda Lawrence.Oreotrochilus leucopleurus Gould.Sephanoides fernandensis fernandensis (King).Sephanoides sephanoides (Lesson).Sappho sparganura sapho (Lesson).Heliomasterfurcifer (Shaw).Philodice evelynae evelynae (Bourcier).No. 6 MAIN ARTERIES NEAR HEART 369Archilochus colubris (Linnaeus).Mellisuga minima vieillot (Shaw).Stellula calliope calliope (Gould).Selasphorus platycercus platycercus (Swainson).Garrod (1873) pointed out that the left carotid alone is present in the following species:Patagona gigas (Vieillot).Chlorolampis osberti (Probably = Chlorostilbon canivetti osberti).Cypselus apus = A pus apus apus (Linnaeus).Cypselus alpinus (Possibly = Apus melba).Chetura vauxi = Chaetura vauxi (J. K. Townsend).Chetura spinicauda = Chaetura s. spinicauda (Temminck).Chetura caudacuta = Hirund-apus caudacutus (Latham).Dendrochelidon coronata (Possibly = Hemiprocne longipennis coronata), while both carotidsare present in:Cypseloides fumigatus (Streubel).In the Trochilidae there is a very high degree of uniformity in the arterial arrangement -pattern, while in the Apodidae there is a strong tendency for the laevo-carotid condition butwith some evidence that this order of birds had their origin perhaps somewhat more recentlyfrom a bicarotid stock of birds.The writer wishes to express his gratitude to Dr. Herbert Friedmann for making availablethe materials used in this study.REFERENCEGarrod, A. H. 1873. On the carotid arteries in birds. London, Proc. Zool. Soc, 457-472.

A Systematic Study of the Main Arteries in the Region of the Heart Aves XIX : Apodiformes, Part 1

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A Systematic Study of the Main Arteries in the Region of the Heart. Aves XVI. Charadriiformes, Part 2

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