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diff --git a/.gitattributes b/.gitattributes new file mode 100644 index 0000000..6833f05 --- /dev/null +++ b/.gitattributes @@ -0,0 +1,3 @@ +* text=auto +*.txt text +*.md text diff --git a/33558-8.txt b/33558-8.txt new file mode 100644 index 0000000..0a6732a --- /dev/null +++ b/33558-8.txt @@ -0,0 +1,1394 @@ +The Project Gutenberg EBook of Thoracic and Coracoid Arteries In Two +Families of Birds, Columbidae and Hirundinidae, by Marion Anne Jenkinson + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Thoracic and Coracoid Arteries In Two Families of Birds, Columbidae and Hirundinidae + +Author: Marion Anne Jenkinson + +Release Date: August 28, 2010 [EBook #33558] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK THORACIC AND CORACOID ARTERIES *** + + + + +Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci +and the Online Distributed Proofreading Team at +https://www.pgdp.net. + + + + + + + +UNIVERSITY OF KANSAS PUBLICATIONS + +MUSEUM OF NATURAL HISTORY + +Volume 12, No. 13, pp. 553-573, 7 figs. + +March, 2, 1964 + +Thoracic and Coracoid Arteries In Two Families of Birds, Columbidae and +Hirundinidae + +BY + +MARION ANNE JENKINSON + +UNIVERSITY OF KANSAS +LAWRENCE +1964 + +UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + +Editors: E. Raymond Hall, Chairman, Henry S. Fitch, +Theodore H. Eaton, Jr. + +Volume 12, No. 13, pp. 553-573, 7 figs. +Published March 2, 1964 + +UNIVERSITY OF KANSAS +Lawrence, Kansas + +PRINTED BY THE STATE PRINTER +TOPEKA, KANSAS +1964 + +[Transcriber's Note: Words surrounded by tildes, like ~this~ signifies +words in bold. Words surrounded by underscores, like _this_, signifies +words in italics.] + + + + +Thoracic and Coracoid Arteries In Two Families of Birds, Columbidae and +Hirundinidae + +BY + +MARION ANNE JENKINSON + + + + +CONTENTS + + PAGE + +INTRODUCTION 555 + +METHODS AND MATERIALS 556 + +MYOLOGY AND ANGIOLOGY: HIRUNDINIDAE 557 + Myology 557 + Angiology 558 + +MYOLOGY AND ANGIOLOGY: COLUMBIDAE 560 + Myology 560 + Angiology 560 + +SUMMARY OF ARTERIAL ARRANGEMENT 562 + +DISCUSSION AND CONCLUSIONS 562 + Individual Variation 562 + Intrafamilial Differences 563 + Interfamilial Differences 565 + +SUMMARY 567 + +LITERATURE CITED 573 + + + + +INTRODUCTION + + +Most descriptions of the circulatory system of birds, largely the work +of Glenny, have dealt with arteries of the neck and thorax in a wide +variety of species. As a result of his work, Glenny offered several +hypotheses concerning the phylogenetic, hence taxonomic, significance of +differences in some of these vessels. He also described six types of +thoracic arterial arrangements and stated that these categories might +represent various levels of evolution (Glenny, 1955:543-544). + +The families Columbidae (pigeons) and Hirundinidae (swallows) have two +nearly extreme arterial types described by Glenny, and are universally +acknowledged as monophyletic. Differences within the families, +therefore, can be considered as valid intrafamilial differences. I have +investigated the thoracic and coracoid arteries and their branches in +members of these two families to determine the degree of individual +variability of the vessels, and the possible causes of interspecific and +intrafamilial differences. + + + + +METHODS AND MATERIALS + + +All specimens studied are in The University of Kansas Museum of Natural +History. They were preserved in alcohol and their blood vessels were not +injected. Dissections were made with the aid of a binocular microscope +at magnifications of 10× and 20×. + +Following is a list of the species studied, the number of individuals of +each species dissected, and the catalogue numbers of the specimens. The +nomenclature and classification are those of the American +Ornithologists' Union's _Check-List of North American Birds_, fifth +edition (1957). + + Family Columbidae + + _Zenaidura macroura_ (Linnaeus), Mourning Dove 2: 40325, 40326. + _Zenaida asiatica_ (Linnaeus), White-winged Dove 1: 40328. + _Scardafella inca_ (Lesson), Inca Dove 5: 34894, 34896, 34902, 34906, + 34907. + _Columba livia_ Gmelin, Rock Dove (domestic pigeon) 1: 40321. + + Family Hirundinidae + + _Iridoprocne bicolor_ (Vieillot), Tree Swallow 1: 38101. + _Progne subis_ (Linnaeus), Purple Martin 5: 37711, 38794, 38796, + 38798, 38804. + _Stelgidopteryx ruficollis_ (Vieillot), Rough-winged Swallow 1: 38277. + _Riparia riparia_ (Linnaeus), Bank Swallow 2: 38784, 38785. + _Hirundo rustica_ (Linnaeus), Barn Swallow 1: 38839. + +The following descriptions are of _Progne subis_ and _Scardafella inca_. +Differences in the vascular system in other members of the families +represented by _P. subis_ and _S. inca_ are mentioned at the appropriate +places. The muscles briefly described for each of these two species are +those that are supplied by the thoracic or coracoid arteries or by +branches of the same, and muscles that, by their origin, location, or +insertion, seem to affect the course or origin of one of these arteries. + +The following sources have been particularly useful for the terminology +of muscles and of skeletal features: Ashley (1941), Beddard (1898), +Coues (1903), Howard (1929), Howell (1937), and Hudson and Lanzillotti +(1955). + +The names used for most arteries are those in common usage for +vertebrates. I have not used the terms "internal mammary" and +"intercostal" artery as substitutes for "thoracic" artery, except when +referring to the work of others. The vessel's homology with the internal +mammary artery of mammals has been denied (Glenny, 1955:541), and the +name "mammary" is certainly not useful descriptively in birds. The term +"intercostal" is less objectionable, except that such a name may call to +mind segmental vessels arising from the dorsal aorta. The term +"thoracic" seems best, as it is reasonably descriptive, and has been +used by Glenny in the majority of his descriptions covering a wide +variety of birds. The name "sternoclavicular" has been used by others as +a synonym for the "coracoid" artery. I have arbitrarily chosen to use +the latter. + + +ACKNOWLEDGMENTS + +I gratefully acknowledge many valuable suggestions in my research and +the preparation of this manuscript from Professors Theodore H. Eaton, A. +Byron Leonard, Richard F. Johnston, Robert M. Mengel, and E. Raymond +Hall. Mr. Abbot S. Gaunt and Miss Sandra Lovett assisted in collecting +specimens. Final drafts of the illustrations were prepared by Mr. Thomas +Swearingen. + + + + +MYOLOGY AND ANGIOLOGY: HIRUNDINIDAE + +Figs. 1, 2, 3, and 4 illustrate the following muscles and arteries +described for _Progne subis_. + + +Myology + +~_M. pectoralis thoracica_~, Fig. 1. The origin is from slightly less than +the posterior half of the sternum, from the ventral half of the keel, +almost the entire length of the posterolateral surface of the clavicle +and adjacent portion of the sterno-coraco-clavicular membrane, and +tendinously from the ventral thoracic ribs. This massive muscle covers +the entire ventral surface of the thorax and converges to insert on the +ventral side of the humerus on the pectoral surface. + +~_M. supracoracoideus_~, Fig. 1. The origin is from the dorsal portion of +the keel and medial portion of the sternum, and is bordered ventrally by +the origin of M. pectoralis thoracica, and laterally by _M. +coracobrachialis posterior_. The origin is also from the manubrium and +the anterolateral portion of the proximal half of the coracoid and to a +slight extent from the sterno-coraco-clavicular membrane adjacent to the +manubrium. This large pinnate muscle converges, passes through the +foramen triosseum, and inserts by a tendon on the external tuberosity of +the humerus, immediately proximal to the insertion of _M. pectoralis +thoracica_. + +~_M. coracobrachialis posterior_~, Figs. 1 and 3. The origin is from the +dorsolateral half of the coracoid, anterolateral portion of the sternum +(where the area of origin is bordered medially by _M. supracoracoideus_, +posteriorly by _M. pectoralis thoracica_, and laterally by _M. +sternocoracoideus_), and also to a slight extent from the area of +attachment of the thoracic ribs to the sternum. The muscle fibers +converge along the lateral edge of the coracoid and insert on the median +crest of the humerus immediately proximal to the pneumatic foramen. In +passing from the origin on the sternum to the insertion on the humerus, +the belly of the muscle bridges the angle formed by the costal process +of the sternum and the coracoid. + +~_M. sternocoracoideus_~, Figs. 2 and 3. The origin is from the entire +external surface of the costal process of the sternum, and to a small +extent from the extreme proximal ends of the thoracic ribs where they +articulate with the costal process. The muscle inserts on a triangular +area on the dorsomedial surface of the coracoid. Like _M. +coracobrachialis posterior_, this muscle bridges the angle formed by the +costal process and the coracoid. + +~_M. subcoracoideus_~ (ventral head), Figs. 2 and 3. The origin is from +the dorsomedial edge of the coracoid at its extreme proximal end, and to +a slight extent from the adjacent portion of the manubrium. The origin +is medial to the insertion of _M. sternocoracoideus_. The ventral head +passes anterodorsally along the medial edge of the coracoid and joins +the dorsal head (not here described). The combined muscle then inserts +by a tendon onto the internal tuberosity of the humerus. + +~_M. costi-sternalis_~, Figs. 1, 2, and 3. The origin is from the anterior +edge of the sternal portion of the first four thoracic ribs. This +triangular muscle narrows and inserts on the posterior edge of the apex +of the costal process. The portion arising from the first rib may share +slips with _M. sternocoracoideus_. + +~_M. costi-sternalis anterior_~, Figs. 1, 2, and 3. This muscle is +variously developed, and originates from a small area on the ventral end +of the vertebral portion of the last cervical rib. The insertion is on +the apex of the costal process, immediately anterior to the insertion of +_M. costi-sternalis_. + +~_Mm. intercostales externus_~, Fig. 1. These muscles extend +posteroventrally between the vertebral portions of successive thoracic +ribs, and between the last cervical and first thoracic ribs. In the more +posterior intercostal spaces these muscles are poorly developed, but +they become progressively better developed anteriorly, and are fully +represented in the most anterior intercostal spaces. + +~_Mm. intercostales internus_~, Fig. 3. These muscles resemble the +external intercostal muscles, but extend anteroventrally, with the +muscles being most fully developed posteriorly, and progressively less +so anteriorly. + +~_Costopulmonary muscles_~, Fig. 3. This diagonal series of muscle slips +from the thoracic ribs attaches to the aponeurosis covering the lungs. + + +Angiology + +Figs. 3 and 4 show all arteries discussed for this family. The numbers +following the names or descriptions of arteries in the text refer to +numbered arteries in one or both of these figures. + +The right and left innominate or brachiocephalic arteries arise from the +aortic trunk and give rise to the common carotid arteries (14). The +major vessel continuing across the thoracic cavity is the subclavian +artery. Classically the subclavian is considered as continuing into the +anterior appendage as the axillary artery. However, in the species +studied, the axillary artery can best be described as a branch from the +subclavian; the pectoral stem forms a more direct continuation of the +subclavian. In traversing the thoracic cavity, the subclavian gives rise +to the thoracic, coracoid, and axillary arteries, and leaves the +thoracic cavity as the pectoral trunk, dorsal to the area where _Mm. +coracobrachialis posterior_ and _sternocoracoideus_ span the angle +formed by the coracoid and costal process. + +The pectoral trunk bifurcates into two main pectoral arteries (9), which +penetrate _M. pectoralis thoracica_. Neither the axillary artery nor +these pectoral arteries were traced in my study. + +The coracoid artery (2) arises from the ventral face of the subclavian +(1), either opposite the base of, or medial to, the axillary artery +(10). The coracoid artery passes ventrad between the medial edge of the +coracoid and the ventral head of _M. subcoracoideus_, and an artery (7) +is given off to supply that muscle. The main vessel then penetrates _M. +supracoracoideus_ and bifurcates or ramifies into several vessels (12). + +Between the origin of the coracoid artery from the subclavian, and the +point where the coracoid artery passes the medial edge of the coracoid, +several branches are given off. These vessels are highly variable in +origin, as described below, and not all were always found. Along with +the coracoid artery, they are termed a "coracoid complex." + +The first artery (11) of this complex arises from any one of several +places: from the lateral face of the coracoid artery at its base; +independently from the subclavian immediately lateral to the origin of +the coracoid artery; and from the thoracic artery near its origin. This +vessel travels laterad, parallel to the subclavian, and penetrates _M. +coracobrachialis posterior_ at the same point that the pectoral artery +passes dorsal to that muscle. + +Another vessel (common stem of 4 and 5) of the coracoid complex in most +specimens arises from the anterior face of the coracoid artery and +branches into several vessels, some of which (5) supply _M. +subcoracoideus_, and some of which (4) feed _M. coracobrachialis +posterior_. The vessel occasionally shares a common stem with the main +vessel (11) to _M. coracobrachialis posterior_, and in some specimens +arises independently from the subclavian, immediately anterior to the +origin of the coracoid artery. The branch (4) to _M. coracobrachialis +posterior_ was also seen to arise independently from any of the +above-mentioned positions. + +Two remaining vessels (6 and 8) are often found as branches from the +coracoid artery. They were small and often were collapsed in the +individuals I dissected, but were most clearly seen in _Iridoprocne +bicolor_. The vessels occasionally had a common base, and in some +specimens only one vessel was found. The first artery (6) passes mediad +into _M. sternocoracoideus_, or continues across that muscle onto the +inner face of the sternum. The second vessel (8) also supplies _M. +sternocoracoideus_ or the inner surface of the sternum, and often a +large branch continues across the dorsal surface of the coracoid to _M. +coracobrachialis posterior_. Fig. 3 shows a composite of these vessels; +not all branches were seen in any one specimen. In the specimen of _I. +bicolor_ a foramen existed on the lateral edge of the coracoid where the +branch (of 8) to _M. coracobrachialis posterior_ passed. An examination +of skeletons of five to 10 individuals each of the five species for +which dissections were made, and of _Petrochelidon pyrrhonota_ (Cliff +Swallow) and _Tachycineta thalassina_ (Violet-green Swallow), in the +University of Kansas collection, showed that most coracoids of these +seven species of swallows had a small notch (as shown in Fig. 4) or a +complete foramen there. + +The thoracic artery (3) arises from the subclavian opposite the base of +the coracoid artery, or from the base of the coracoid artery. Of the +five specimens of _P. subis_ dissected, one individual had the former +arrangement on both sides, and one had the latter on both sides, whereas +in the remaining three the thoracic artery arose from the coracoid +artery on one side and from the subclavian on the other side. The +distance between these two possible sites of origin is slight. + +The thoracic artery usually passes ventral to _M. costi-sternalis +anterior_. Occasionally a small artery (13) could be traced from the +main trunk of the thoracic artery to that muscle. The main thoracic +artery bifurcates near the insertion of _M. costi-sternalis_, the +branches traveling posteriad on both sides of the muscle. On one side of +one specimen this artery bifurcated immediately after leaving the +subclavian, the dorsal trunk passing dorsal to _M. costi-sternalis +anterior_, and the ventral trunk ventral to the muscle. On the other +side of the same individual the artery passed dorsal to _M. +costi-sternalis anterior_, bifurcating at the normal point. + +From the ventral trunk of the thoracic artery a variable number of small +vessels arises to supply the costosternal articulations. The main +ventral trunk bifurcates into two branches, one of which passes onto the +inner face of the sternum, and one of which supplies the posterior two +intercostal spaces. + +The dorsal thoracic trunk supplies _M. costi-sternalis_, several dorsal +intercostal areas, and the costopulmonary muscles. Minor variations in +all of the smaller branches of the thoracic artery were common. + + + + +MYOLOGY AND ANGIOLOGY: COLUMBIDAE + +Figs. 5, 6, and 7 illustrate the following muscles and arteries +described for _Scardafella inca_. + + +Myology + +~_M. pectoralis thoracica~_, Fig. 5. The origin is from approximately the +ventral third of the keel, the lateral and anterior portion of the +clavicle and the adjacent sterno-coraco-clavicular membrane, and from +the lateral portion of the sternum and the fascia overlying the thoracic +ribs. This massive muscle covers the entire ventral surface of the +thorax, converges, and inserts on the pectoral surface on the ventral +side of the humerus. + +~_M. supracoracoideus~_, Fig. 5. The origin is from the dorsal two-thirds +of the keel and medial half of the sternum (where the origin is bordered +ventrally, posteriorly, and laterally by the origin of _M. pectoralis +thoracica_) and from the sterno-coraco-clavicular membrane adjacent to +the coracoid. This large pinnate muscle converges, passes through the +foramen triosseum, and inserts by means of a strong tendon on the dorsal +surface of the humerus on the deltoid ridge. + +~_M. coracobrachialis posterior_~, Fig. 5. The origin is from a prominent +lateral wing on the posterolateral portion of the coracoid, and from the +lateral surface of the proximal two-thirds of the coracoid. The +insertion is by means of a tendon on the internal tuberosity of the +humerus. Of the muscles described here, this one differs most strikingly +from the homologous muscle in _P. subis_. The difference can be seen by +comparing Figs. 1 and 5. + +~_M. sternocoracoideus_~, Figs. 5, 6, and 7. The origin is from the +external, and to a slight extent from the internal, surface of the +costal process. The insertion is on a posterolateral triangular area on +the dorsal surface of the coracoid. + +~_M. costi-sternalis_~, Figs. 5 and 6. The origin is from the anterior +edge of the sternal portion of the first three thoracic ribs. The muscle +converges and inserts on the apex of the costal process. + +~_M. subcoracoideus_~ (ventral head), Fig. 6. The origin is from the +manubrium and from approximately the posterior half of the coracoid and +on the medial and dorsal surface of that bone, and the medial side of +the sterno-coraco-clavicular membrane adjacent to the coracoid. The +ventral head passes anterodorsally to join with the dorsal head (not +here described), and the combined muscle inserts by a tendon on the +internal tuberosity of the humerus. + +~_Mm. intercostales externus_~, Fig. 5. These muscles extend +posteroventrally between successive thoracic ribs and between the last +cervical and first thoracic ribs. + +~_Mm. intercostales internus_~, Fig. 7. These muscles extend +anteroventrally between the last three thoracic ribs. + +~_Costopulmonary muscles_~, Fig. 7. This series of muscle slips from the +thoracic ribs attaches to the aponeurosis covering the lungs. + + +Angiology + +Figs. 5, 6, and 7 show all arteries discussed for this family. The +numbers following names or descriptions of arteries in the text refer to +numbered arteries in one of these figures. Insofar as possible, the +numbers used for these arteries are the same numbers used for the +homologous vessels in swallows. + +The right and left innominate arteries arise from the aortic trunk and +give rise to the common carotid (14) and subclavian (1) arteries. The +latter continues across the thoracic cavity, giving rise to the coracoid +(2) and axillary (10) arteries, and becoming the pectoral trunk. That +trunk swings posteriorly and leaves the thoracic cavity near the apex of +the costal process, as shown in Fig. 7. Where the trunk passes under _M. +sternocoracoideus_, the thoracic artery (3) is given off. + +The various branches of the coracoid artery, again referred to as a +"coracoid complex," are as follows: The first branch, from the posterior +face of the coracoid artery, is a relatively large vessel (6) here +termed the sternal artery; it passes mediad across _M. +sternocoracoideus_, sending off a branch (6a) to that muscle. The right +sternal artery continues posteriorly on the mid-line of the inner +surface of the sternum, and appears to send branches into the various +pneumatic foramina of the sternum, but these vessels are minute and +exceedingly difficult to trace accurately. The corresponding left vessel +is smaller and ramifies on the anteromedial surface of the sternum. +Variations found in these vessels were the following: In one specimen of +_S. inca_ the sternal artery had, on both sides, an independent origin +from the subclavian, lateral to the origin of the coracoid artery. In +_Zenaidura macroura_ both right and left sternal arteries were similar +to the left vessel described above, no median longitudinal vessel being +seen. In _Columba livia_ no vessel corresponding to the sternal artery +was seen. In _Zenaida asiatica_ these arteries penetrated _M. +sternocoracoideus_; no branch to the sternum was seen. + +A small complex of vessels (4 and 4a) arises from the lateral face of +the coracoid artery and feeds _M. coracobrachialis posterior_, and +occasionally _M. sternocoracoideus_. One branch (4a) passes under the +coracoid and travels along the lateral side of that bone, supplying +small branches to _M. coracobrachialis posterior_, and finally ramifying +on the head of the coracoid. In _C. livia_, _Zenaidura macroura_, and +_Zenaida asiatica_ this complex usually arises independently from the +subclavian, and in one case it arose from the axillary artery. + +Two other branches from the coracoid artery were regularly seen. The +first (8) passes across _M. sternocoracoideus_ and appears to supply the +area of the coracoid articulation with the sternum; the second (7) +supplies _M. subcoracoideus_ as the main vessel passes between that +muscle and the coracoid and penetrates _M. suparacoracoideus_. A small +notch on the medial side of the coracoid (shown in Figs. 6 and 7) often +marks the passage of the coracoid artery. + +All vessels of the coracoid complex are exceedingly variable, in number, +size, and site of origin. + +A prominent vessel (15) is given off from the posterior pectoral artery, +outside the thoracic cavity, passes ventrad, and sends two branches into +_M. supracoracoideus_. No corresponding artery was seen in the swallows +dissected. + +The thoracic artery (3), arising from the pectoral stem, +characteristically bifurcates at the anterior end of _M. +costi-sternalis_. The dorsal, and larger, branch passes posteriorly, +sends several small branches to _M. costi-sternalis_, and continues to +the most posterior rib. The ventral trunk bifurcates, one branch passing +along the edge of, and supplying, _M. costi-sternalis_, the other +branch passing onto the surface of the sternum. In some specimens two +such branches to the sternum were seen. + + + + +SUMMARY OF ARTERIAL ARRANGEMENT + + +In both families the vessels that are relatively constant in appearance +are: a subclavian giving rise to the carotid and axillary arteries, and +becoming the pectoral trunk; the thoracic artery arising variously, and +passing posteriorly to the rib cage; and the coracoid complex of +vessels. The coracoid complex includes the coracoid artery, the vessels +to _Mm. sternocoracoideus_ and _coracobrachialis posterior_, and the +sternal artery, which is variously present, and more extensive in some +species than in others. + + + + +DISCUSSION AND CONCLUSIONS + + +In the vessels studied individual variation is marked, but the arterial +arrangement within both families is relatively constant. Interfamilial +differences probably represent responses of the arteries to adaptive +structural differences of other systems of the body. + + +Individual Variation + +The term "individual variation" is used here to mean "continuous +non-sex-associated variation" (see Mayr, Linsley, and Usinger, 1953:93) +found between members of the same species or between the two sides of +the same individual. It is hazardous to define individual variation (and +also interspecific differences, as discussed later) in the origin of one +vessel by relating its location to other vessels, because these may +likewise vary in origin. But, by necessity, certain vessels that are +probably less variable (axillary, carotid, and pectoral arteries) have +been considered here as being constant in origin. If these three vessels +are accepted as reference points, individual variants, as well as +interspecific differences, can easily be described in the thoracic and +coracoid arteries and in their various branches. + +The thoracic artery in _P. subis_ arose either from the subclavian +artery, or from the coracoid artery. Likewise in other swallows, both of +these origins were found. In doves the thoracic artery arose +consistently from the pectoral stem, lateral to the origin of the +axillary artery. + +The coracoid artery in _P. subis_ and other swallows arose from the +subclavian artery, either opposite the base of the axillary artery, or +medial to that vessel. In all doves studied the coracoid artery arose +from the subclavian medial to the axillary artery. I observed much +individual variation in the branches of the coracoid artery (that is to +say, in the vessels of the coracoid complex). In _S. inca_ the sternal +artery arose either from the coracoid artery, or independently from the +subclavian. As mentioned earlier, in members of both families the +vessels to _Mm. coracobrachialis posterior_ and _subcoracoideus_ are +highly variable, arising in swallows from the coracoid artery or from +the subclavian artery, and in doves from either of these two sites or +from the axillary artery. The distribution of these arteries after their +origin is also diverse. + +Individual variation in the arteries of the thorax has been recorded +previously. Bhaduri, Biswas, and Das (1957:2) state that, in the +domestic pigeon, "the origin and course of various smaller arteries... +show noticeable variation," although they do not specifically state to +which vessels they are referring. Fisher (1955:287-288) found +variability in the Whooping Crane, _Grus americana_, of the axillary, +coracoid, thoracic, and pectoral arteries. In one specimen he found +these vessels arising on the right side from the subclavian, in the +sequence just listed, and on the left side all arose from the same +point. Berger (1956:439-440) strongly emphasized the variability of the +vascular system, calling it the most variable in the body. As he stated, +this high degree of individual variation seems to be due to the +embryological development of the system, wherein many of the adult +channels of circulation are derived from embryonic plexuses. + + +Intrafamilial Differences + +In spite of the rather extensive amount of individual variability in +some vessels, I found the over-all pattern of arteries to be relatively +constant within the family Columbidae and within the family +Hirundinidae. There are, nevertheless, several intrafamilial differences +needing some further discussion and clarification. + +Others have reported the occasional presence of more than one coracoid +artery on each side in some columbids, these arteries being described as +arising from various sites and being variously named. Bhaduri and Biswas +(1954) described the arterial situation in seven species of the family +Columbidae (_Columba livia_, _Streptopelia tranquebarica_, _S. +chinensis_, _S. senegalensis_, _Chalcophaps indica_, _Treron bicincta_, +and _T. phoenicoptera_) and stated (_op. cit._: 348) that "The +sternoclavicular [= coracoid] artery is similar in all the species, but +the domestic pigeon seems to be unique in that it has, in addition, a +small vessel, the accessory sternoclavicular." This artery was described +later, in the domestic pigeon, as follows (Bhaduri, Biswas, and Das, +1957:5): "A minute and insignificant vessel which has been termed the +_accessory sternoclavicular_ artery... is given off close to the origin +of the sternoclavicular. It passes anteroventrally to supply the +adjacent muscles." Glenny (1955:577) described the arterial pattern +characteristic of members of the family Columbidae (more than 30 species +studied by him) and stated that "three pairs of coracoid arteries are +found in _Otidiphaps nobilis_, normally one or two pairs may be found." +As suggested by Bhaduri and Biswas (1954:348), the "accessory" vessel +probably corresponds to a vessel previously described by Glenny (1940) +in _Streptopelia chinensis_ and referred to as the "coracoid minor." + +Bhaduri and Biswas (1954:348) have suggested that "the accessory +sternoclavicular artery occurring sporadically as it does in some +species of diverse groups may not have any phylogenetic value." + +In no case did I find more than one coracoid artery on a side. When one +of the highly variable arteries feeding _Mm. coracobrachialis posterior_ +and _sternocoracoideus_ (arteries 4 and 4a, Fig. 7) arises from the +subclavian or axillary artery instead of from the coracoid artery, that +vessel may have been interpreted by others as a second (accessory or +minor) coracoid artery. If so, this artery probably does not "occur +sporadically." Rather, its origin from the subclavian, axillary, or +thoracic artery may be sporadic, subject to individual variation, and it +may have been overlooked when it arose from the coracoid artery. + +Of the vessels described here, the only one that differed distinctly in +one species was the sternal artery. In _Scardafella inca_ the right +sternal vessel was long, extending down the mid-line of the inner +surface of the sternum, whereas in other columbids the right and left +arteries ramified on the anterior part of the inner surface of the +sternum, or were altogether lacking. I am unable to account for the +differential development of this artery in _S. inca_. + +In describing the arterial arrangement in the seven species of Indian +columbids named earlier, Bhaduri and Biswas (1954:348) state that all +species except _Treron phoenicoptera_ have two "internal mammary" +arteries on each side "showing variable sites of origin." These arteries +were later described (Bhaduri, Biswas, and Das, 1957:4-5) as "a slender +(_outer_) _internal mammary_ artery... to the outer wall of the thoracic +cavity" and "a slender (_inner_) _internal mammary_ artery... to supply +the inner wall of the chest cavity." From this description, the question +arises as to whether the "outer" one of these arteries should properly +be called an _external_ instead of _internal_ mammary artery. In any +case, I saw no specimen possessing two thoracic arteries on a side. + + +Interfamilial Differences + +As shown above, there is a high degree of individual variation in the +vessels being considered, while at the same time, few interspecific +differences were noted within the families. On the other hand, the +vascular arrangement of swallows consistently differed from that of +pigeons in the species studied. The differences are most easily +described by discussing the resulting change in the site of origin of +the thoracic artery. In swallows the thoracic artery arises between the +carotid and axillary arteries, either from the stem of the coracoid +artery or independently from the subclavian, but in pigeons the thoracic +artery arises from the pectoral stem, a site of attachment that is +relatively more lateral than in swallows. + +This difference, in my opinion, demonstrates well the topological +relationships of various systems of the body, here especially of the +skeletal, muscular, and vascular systems. The location of the thoracic +artery seems to be determined by the particular configuration of +skeletal and muscular elements, although even within the bounds set by +these elements, individual variation in the precise origin of the artery +is possible. In all swallows dissected _Mm. coracobrachialis posterior_ +and _sternocoracoideus_ bridge the angle formed by the costal process +and the coracoid. This arrangement makes it necessary for the subclavian +to leave the thoracic cavity dorsal to the costal process, although it +does pass immediately anterior to that process. The thoracic artery +arises from the vessel next to the apex of the costal process, hence +from the subclavian, between the axillary and carotid arteries. + +In pigeons, the wing of the coracoid extends farther laterally than does +the costal process, and the apex of the latter is displaced farther +posteriorly than it is in swallows. _M. coracobrachialis posterior_ does +not arise from the sternum, and only part of the costal process serves +as a point of origin for _M. sternocoracoideus_. Consequently, this +region differs from that of swallows; the area between the costal +process and coracoid is not entirely bridged by muscle, and the space +between the two skeletal elements is of a different shape and size. It +seems that these differences have resulted, in pigeons, in the +subclavian assuming a more anterior position with reference to the +costal process. The subclavian in these birds leads into the pectoral +artery, which runs posteriad, passing under _M. sternocoracoideus_ and +leaving the thoracic cavity approximately opposite the apex of the +costal process. The thoracic artery arises immediately opposite the apex +of the costal process from the main artery in the area, as it does in +swallows, except that in this case the adjacent artery from which it +arises is the pectoral stem. + +The thoracic area seems to be most "efficiently" arranged when the +thoracic artery arises _opposite the apex of the costal process, from +whatever main artery is closest to that site_. This arrangement existed +in all species studied. Considering the differences in skeletal and +muscular structures, between pigeons and swallows, it would be much more +remarkable if an alternative were the case, that is to say if the +thoracic artery _had the same site of attachment on the subclavian_ in +both groups. + +A comparison of these suggestions with statements made previously about +these arteries seems necessary. When Glenny (1955) summarized his +accumulative findings, concerning the main arteries in the region of the +heart, based on individuals representing more than 750 avian species of +27 orders and 120 families, he described five types of thoracic arteries +that were distinguished by differences in the site of their origin, and +one type in which there were two thoracic arteries on each side. His +statements regarding these differences were as follows (Glenny, +1955:543-544): + + "The thoracic, intercostal, or internal mammary artery of + birds... is found to arise at slightly different relative + positions--from a point at the base of the inferior pectoral + artery to a point near the base of the coracoid or + sternoclavicular artery, and in some instances both of these + vessels have a common root from the subclavian artery. Such + differences are found to be of common occurrence within + several orders of birds. In the Galliformes and the + Passeriformes there appears to be a graded series in the + sites of attachment of the thoracic artery from a lateral to + a medial position. As a result of these observations, + numerical values can be assigned to the site of attachment + of the intercostal or thoracic artery, and these values may + come to be used as an index in specific levels of + evolution.... + + "The medial migration of the thoracic artery appears to have + some phylogenetic significance as yet not understood." + +The six types of thoracic arteries described in Glenny's classification +were distinguished as follows (Glenny, 1955:544): + + "Type 1: attachment to the pectoral stem lateral to the + axillary. + + "Type 2: attachment to the subclavian between the axillary + and coracoid. + + "Type 3: attachment to the subclavian at the base of the + coracoid. + + "Type 4: attachment to the subclavian, but with a common + root for both the coracoid and thoracic. + + "Type 5: attachment to the subclavian medial to both the + axillary and coracoid. + + "Type 6: two separate thoracic arteries are present; the + primary thoracic is the same as type 1 above, while the + secondary thoracic is the same as type 3 or type 4 above." + +Possibly the thoracic artery has undergone migration but apparent +differences in its origin might well be due to differences in other +vessels of the thoracic area. Additionally, there seems to be no reason +to assume that the lateral position of the thoracic artery is the +primitive one, or that the medial is the derived position, as is implied +by the phrase "medial migration." Although the lateral site of +attachment (type 1) is predominant in the lower orders of birds, and the +medial attachment is found primarily in Passeriformes, a fact which may +indicate that type 1 is the more primitive, it must nevertheless be kept +in mind that a sequence of a single morphological character does not +necessarily represent the phylogenetic sequence of the character itself +(see Mayr, 1955:41). + +Also, a given arterial arrangement might be independently derived more +than once. If such has been the case, similarities in arterial +arrangements in different taxa would sometimes be "chance similarities," +that is to say, "resemblance in characteristics developed in separate +taxa by independent causes and without causal relationship involving the +similarity as such" (Simpson, 1961:79). + +The particular arrangement of the arteries of the thoracic area also +seems to be of limited value as a clue to taxonomic relationships. If +the origin of any artery is determined by skeletal and muscular +features, as I suggest, the artery perhaps ought not be considered as a +separate character, but as part of a "character complex" that varies as +a unit (see Mayr, Linsley, and Usinger, 1953:123). The skeleton offers a +potential fossil record for consideration. Changes in the skeleton and +muscles, great enough to affect the blood vessels, would probably be +detected more easily than would the resulting vascular changes. Also, I +did not find as much individual variation in the skeleton and muscles in +the area studied as I did in the vascular system. In other words, within +the bounds established by the skeletal and muscular features, the artery +still exhibited individual variation in exact origin. + + + + +SUMMARY + + +The origin, distribution, and individual variation of the thoracic and +coracoid arteries, and their branches, have been studied in four species +of the family Columbidae (pigeons) and in five species of the family +Hirundinidae (swallows). These arteries are described for _Scardafella +inca_ (Inca Dove) and _Progne subis_ (Purple Martin). Muscles that are +supplied by these vessels, and muscles the particular configuration of +which seems to effect the arrangement of the arteries have also been +described. Correlation of the arteries observed with those named and +described by other workers has been attempted. + +In most of the vessels studied there is a high degree of individual +variation, but few interspecific differences were noticed within either +family. Differences in the arteries of the thorax between the two +families are described by discussing the resulting different origins of +the thoracic artery. In swallows the thoracic artery arises from either +the subclavian artery or the coracoid artery, whereas in pigeons it +arises from the pectoral trunk. This difference in site of attachment +seems to be a result of differences between the two families in muscular +and skeletal elements of the thorax. + +The particular site of attachment of the thoracic artery is of limited +value as a taxonomic character. Several considerations influenced this +conclusion. (1) If the location of the artery is determined by skeletal +and muscular elements, these associated structures must be considered +taxonomically as a "character complex" (a set of characters varying as a +unit). (2) Even within the bounds established by the skeleton and +muscles, the artery displays a high degree of individual variation in +exact origin. (3) A given arterial arrangement could have been derived +independently many times. (4) Because differences are defined relative +to other likewise variable vessels, supposed similarities or differences +in the one artery may be artifacts of the system of description. + +My findings and interpretations do not support previous suggestions that +the thoracic artery has undergone a mediad migration, and that the +various sites of attachment of that vessel may come to represent various +levels of evolution. The primitive site of attachment of the vessel is +unknown, and it seems to me that it has not been sufficiently +demonstrated that the vessel has undergone any "migration." + +[Illustration: FIG. 1. _Progne subis._ Lateral view of left half of +thorax. _M. pectoralis thoracica_ (area of insertion indicated by dotted +line) has been removed. Muscles not described in text are not shown. (× +1.5.)] + +[Illustration: FIG. 2. _Progne subis._ Lateral view of left half of +thorax. Same view as shown in Fig. 1, but with _Mm. supracoracoideus_, +_coracobrachialis posterior_, and _intercostales externus_ removed. (× +1.5.)] + +[Illustration: FIG. 3. _Progne subis._ Medial view of left half of +thorax. Not all muscles shown. See Fig. 4 for identification of +arteries. (× 1.5.)] + +[Illustration: FIG. 4. _Progne subis._ Lateral view of left half of +thorax. (× 1.5.) + +(Applies also to Fig. 3.) + + 1. Subclavian artery. + 2. Coracoid artery. + 3. Thoracic artery. + 4. (Unnamed.) Supplies _M. coracobrachialis posterior_. + 5. (Unnamed.) Supplies _M. subcoracoideus_. + 6. (Unnamed.) Supplies _M. sternocoracoideus_ and sternum. + 7. (Unnamed.) Supplies _M. subcoracoideus_. + 8. (Unnamed.) Supplies _M. sternocoracoideus_, _M. coracobrachialis + posterior_, and sternum. + 9. Pectoral artery. + 10. Axilliary artery. + 11. (Unnamed.) Supplies _M. coracobrachialis posterior_. + 12. (Unnamed.) Supplies _M. supracoracoideus_. + 13. (Unnamed.) Supplies _M. costi-sternalis anterior_. + 14. Carotid artery.] + +[Illustration: FIG. 5. _Scardafella inca._ Lateral view of left half of +thorax. _M. pectoralis thoracica_ (area of insertion indicated by dotted +line) has been removed. Muscles not described in text are not shown. See +legend for Fig. 7 for identification of arteries. (× 1.)] + +[Illustration: FIG. 6. _Scardafella inca._ Lateral view of left half of +thorax. See legend for Fig. 7 for identification of arteries. (× 1.)] + +[Illustration: FIG. 7. _Scardafella inca._ Medial view of left half of +thorax. (× 1.) + +KEY + +(Applies also to Figs. 5 and 6.) Numerals not used are those used for +_Progne subis_ for which no homologous artery occurs in _Scardafella +inca_. + + 1. Subclavian artery. + 2. Coracoid artery. + 3. Thoracic artery. + 4. (Unnamed.) Supplies _Mm. coracobrachialis posterior_ and + _sternocoracoideus_. + 4a. (Unnamed.) Supplies _M. coracobrachialis posterior_. + 6. Sternal artery. (Shown as it appears on _right_ side. Left sternal + artery not so extensive.) + 6a. (Unnamed.) Supplies _M. sternocoracoideus_. + 7. (Unnamed.) Supplies _M. subcoracoideus_. + 8. (Unnamed.) Supplies coracoid-sternal articulation. + 9. Pectoral artery. + 10. Axillary artery. + 12. (Unnamed.) Supplies _M. supracoracoideus_. + 14. Carotid artery. + 15. (Unnamed.) Supplies _M. supracoracoideus_.] + + + + +LITERATURE CITED + + +AMERICAN ORNITHOLOGISTS' UNION + +1957. Check-List of North American birds. Baltimore, Maryland, Amer. +Ornith. Union, xiv + 691 pp. + + +ASHLEY, J. F. + +1941. A study of the structure of the humerus in the Corvidae. Condor, +43:184-195. + + +BEDDARD, F. E. + +1898. The structure and classification of birds. London, Longmans, +Green, & Co., xx + 548 pp. + + +BERGER, A. J. + +1956. Anatomical variation and avian anatomy. Condor, 58:433-441. + + +BHADURI, J. L., and BISWAS, B. + +1954. The main cervical and thoracic arteries of birds. Series 2. +Columbiformes, Columbidae, part 1. Anat. Anz., 100:337-350. + + +BHADURI, J. L., BISWAS, B., and DAS, S. K. + +1957. The arterial system of the domestic pigeon (_Columba livia_ +Gmelin). Anat. Anz., 104:1-14. + + +COUES, E. + +1903. Key to North American birds. Vol. I, Fifth edit. Boston, The Page +Company, xlii + 535 + [55] pp. + + +FISHER, H. I. + +1955. Major arteries near the heart in the Whooping Crane. Condor, +57:286-289. + + +GLENNY, F. H. + +1940. The main arteries in the region of the heart of three species of +doves. Bull. Fan Mem. Inst. Biol., Zool. ser., vol. 10, pt. 4, 271-278. +(Not seen.) + +1955. Modifications of pattern in the aortic arch system of birds and +their phylogenetic significance. Proc. U. S. Nat. Mus., 104:525-621. + + +HOWARD, H. + +1929. The avifauna of Emeryville Shellmound. Univ. Calif. Publs. Zool., +32:301-394. + + +HOWELL, A. B. + +1937. Morphogenesis of the shoulder architecture: Aves. Auk, 54:364-375. + + +HUDSON, G. E., and LANZILLOTTI, P. J. + +1955. Gross anatomy of the wing muscles in the family Corvidae. Amer. +Midl. Nat., 53:1-44. + + +MAYR, E. + +1955. Comments on some recent studies of song bird phylogeny. Wilson +Bull., 67:33-44. + + +MAYR, E., LINSLEY, E. G., and USINGER, R. L. + +1953. Methods and principles of systematic zoology. New York, +McGraw-Hill Book Co., x + 336 pp. + + +SIMPSON, G. G. + +1961. Principles of animal taxonomy. New York, Columbia Univ. Press, xiv ++ 247 pp. + + +_Transmitted June 24, 1963._ + + + + + +End of the Project Gutenberg EBook of Thoracic and Coracoid Arteries In Two +Families of Birds, Columbidae and Hirundinidae, by Marion Anne Jenkinson + +*** END OF THIS PROJECT GUTENBERG EBOOK THORACIC AND CORACOID ARTERIES *** + +***** This file should be named 33558-8.txt or 33558-8.zip ***** +This and all associated files of various formats will be found in: + https://www.gutenberg.org/3/3/5/5/33558/ + +Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci +and the Online Distributed Proofreading Team at +https://www.pgdp.net. + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Thoracic and Coracoid Arteries In Two Families of Birds, Columbidae and Hirundinidae + +Author: Marion Anne Jenkinson + +Release Date: August 28, 2010 [EBook #33558] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK THORACIC AND CORACOID ARTERIES *** + + + + +Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci +and the Online Distributed Proofreading Team at +https://www.pgdp.net. + + + + + + +</pre> + + + + +<p class="center"> +<span class="smcap">University of Kansas Publications</span> +<br /> +<span class="smcap">Museum of Natural History</span><br /> +<br /> +Volume 12, No. 13, pp. 553-573, 7 figs.<br /> +<br /> +March, 2, 1964<br /> +</p> + +<h1>Thoracic and Coracoid Arteries In Two Families of Birds, Columbidae and +Hirundinidae</h1> + +<h3>BY</h3> + +<h2>MARION ANNE JENKINSON</h2> + +<p class="center"> +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1964<br /> +<br /> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors: E. Raymond Hall, Chairman, Henry S. Fitch,<br /> +Theodore H. Eaton, Jr.<br /> +<br /> +Volume 12, No. 13, pp. 553-573, 7 figs.<br /> +Published March 2, 1964<br /> +<br /> +<span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /> +<br /> +PRINTED BY THE STATE PRINTER<br /> +TOPEKA, KANSAS<br /> +1964<br /> +</p> + + + +<hr style="width: 65%;" /><p><span class='pagenum'><a name="Page_555" id="Page_555">[Pg 555]</a></span></p> +<h2>Thoracic and Coracoid Arteries In Two Families of Birds, Columbidae and +Hirundinidae</h2> + +<h4>BY</h4> + +<h3>MARION ANNE JENKINSON</h3> + +<hr style="width: 65%;" /> + +<h2>CONTENTS</h2> + +<p> +<span class="tocnum">PAGE</span><br /> +<br /> +<span class="smcap">Introduction</span> <span class="tocnum"><a href='#Page_555'>555</a></span><br /> +<br /> +<span class="smcap">Methods and Materials</span> <span class="tocnum"><a href='#Page_556'>556</a></span><br /> +<br /> +<span class="smcap">Myology and Angiology: Hirundinidae</span> <span class="tocnum"><a href='#Page_557'>557</a></span><br /> +<span style="margin-left: 1em;">Myology <span class="tocnum"><a href='#Page_557'>557</a></span></span><br /> +<span style="margin-left: 1em;">Angiology <span class="tocnum"><a href='#Page_558'>558</a></span></span><br /> +<br /> +<span class="smcap">Myology and Angiology: Columbidae</span> <span class="tocnum"><a href='#Page_560'>560</a></span><br /> +<span style="margin-left: 1em;">Myology <span class="tocnum"><a href='#Page_560'>560</a></span></span><br /> +<span style="margin-left: 1em;">Angiology <span class="tocnum"><a href='#Page_560'>560</a></span></span><br /> +<br /> +<span class="smcap">Summary of Arterial Arrangement</span> <span class="tocnum"><a href='#Page_562'>562</a></span><br /> +<br /> +<span class="smcap">Discussion and Conclusions</span> <span class="tocnum"><a href='#Page_562'>562</a></span><br /> +<span style="margin-left: 1em;">Individual Variation <span class="tocnum"><a href='#Page_562'>562</a></span></span><br /> +<span style="margin-left: 1em;">Intrafamilial Differences <span class="tocnum"><a href='#Page_563'>563</a></span></span><br /> +<span style="margin-left: 1em;">Interfamilial Differences <span class="tocnum"><a href='#Page_565'>565</a></span></span><br /> +<br /> +<span class="smcap">Summary</span> <span class="tocnum"><a href='#Page_567'>567</a></span><br /> +<br /> +<span class="smcap">Literature Cited</span> <span class="tocnum"><a href='#Page_573'>573</a></span><br /> +</p> + + + +<hr style="width: 65%;" /> +<h2>INTRODUCTION</h2> + + +<p>Most descriptions of the circulatory system of birds, largely the work +of Glenny, have dealt with arteries of the neck and thorax in a wide +variety of species. As a result of his work, Glenny offered several +hypotheses concerning the phylogenetic, hence taxonomic, significance of +differences in some of these vessels. He also described six types of +thoracic arterial arrangements and stated that these categories might +represent various levels of evolution (Glenny, 1955:543-544).</p> + +<p>The families Columbidae (pigeons) and Hirundinidae (swallows) have two +nearly extreme arterial types described by Glenny, and are universally +acknowledged as monophyletic. Differences within the families, +therefore, can be considered as valid intrafamilial differences. I have +investigated the thoracic and coracoid arteries and their branches in +members of these two families to determine the degree of individual +variability of the vessels, and the possible causes of interspecific and +intrafamilial differences.</p> + + + +<hr style="width: 65%;" /><p><span class='pagenum'><a name="Page_556" id="Page_556">[Pg 556]</a></span></p> +<h2>METHODS AND MATERIALS</h2> + + +<p>All specimens studied are in The University of Kansas Museum of Natural +History. They were preserved in alcohol and their blood vessels were not +injected. Dissections were made with the aid of a binocular microscope +at magnifications of 10× and 20×.</p> + +<p>Following is a list of the species studied, the number of individuals of +each species dissected, and the catalogue numbers of the specimens. The +nomenclature and classification are those of the American +Ornithologists' Union's <i>Check-List of North American Birds</i>, fifth +edition (1957).</p> + +<div class="poem"><div class="stanza"> +<span class="i0">Family Columbidae<br /></span> +</div><div class="stanza"> +<span class="i0"><i>Zenaidura macroura</i> (Linnaeus), Mourning Dove 2: 40325, 40326.<br /></span> +<span class="i0"><i>Zenaida asiatica</i> (Linnaeus), White-winged Dove 1: 40328.<br /></span> +<span class="i0"><i>Scardafella inca</i> (Lesson), Inca Dove 5: 34894, 34896, 34902, 34906, 34907.<br /></span> +<span class="i0"><i>Columba livia</i> Gmelin, Rock Dove (domestic pigeon) 1: 40321.<br /></span> +</div><div class="stanza"> +<span class="i0">Family Hirundinidae<br /></span> +</div><div class="stanza"> +<span class="i0"><i>Iridoprocne bicolor</i> (Vieillot), Tree Swallow 1: 38101.<br /></span> +<span class="i0"><i>Progne subis</i> (Linnaeus), Purple Martin 5: 37711, 38794, 38796, 38798, 38804.<br /></span> +<span class="i0"><i>Stelgidopteryx ruficollis</i> (Vieillot), Rough-winged Swallow 1: 38277.<br /></span> +<span class="i0"><i>Riparia riparia</i> (Linnaeus), Bank Swallow 2: 38784, 38785.<br /></span> +<span class="i0"><i>Hirundo rustica</i> (Linnaeus), Barn Swallow 1: 38839.<br /></span> +</div></div> + +<p>The following descriptions are of <i>Progne subis</i> and <i>Scardafella inca</i>. +Differences in the vascular system in other members of the families +represented by <i>P. subis</i> and <i>S. inca</i> are mentioned at the appropriate +places. The muscles briefly described for each of these two species are +those that are supplied by the thoracic or coracoid arteries or by +branches of the same, and muscles that, by their origin, location, or +insertion, seem to affect the course or origin of one of these arteries.</p> + +<p>The following sources have been particularly useful for the terminology +of muscles and of skeletal features: Ashley (1941), Beddard (1898), +Coues (1903), Howard (1929), Howell (1937), and Hudson and Lanzillotti +(1955).</p> + +<p>The names used for most arteries are those in common usage for +vertebrates. I have not used the terms "internal mammary" and +"intercostal" artery as substitutes for "thoracic" artery, except when +referring to the work of others. The vessel's homology with the internal +mammary artery of mammals has been denied (Glenny, 1955:541), and the +name "mammary" is certainly not useful descriptively in birds. The term +"intercostal" is less objectionable, except that such a name may call to +mind segmental vessels arising from the dorsal aorta. The term +"thoracic" seems best, as it is reasonably descriptive, and has been +used by Glenny in the majority of his descriptions covering a wide +variety of birds. The name "sternoclavicular" has been used by others as +a synonym for the "coracoid" artery. I have arbitrarily chosen to use +the latter.</p> + + +<h3>ACKNOWLEDGMENTS</h3> + +<p>I gratefully acknowledge many valuable suggestions in my research and +the preparation of this manuscript from Professors Theodore H. Eaton, A. +Byron Leonard, Richard F. Johnston, Robert M. Mengel, and E. Raymond +Hall. Mr. Abbot S. Gaunt and Miss Sandra Lovett assisted in collecting +specimens. Final drafts of the illustrations were prepared by Mr. Thomas +Swearingen.</p> + + + +<hr style="width: 65%;" /><p><span class='pagenum'><a name="Page_557" id="Page_557">[Pg 557]</a></span></p> +<h2>MYOLOGY AND ANGIOLOGY: HIRUNDINIDAE</h2> + +<p>Figs. 1, 2, 3, and 4 illustrate the following muscles and arteries +described for <i>Progne subis</i>.</p> + + +<h3>Myology</h3> + +<p><b><i>M. pectoralis thoracica</i></b>, Fig. 1. The origin is from slightly less than +the posterior half of the sternum, from the ventral half of the keel, +almost the entire length of the posterolateral surface of the clavicle +and adjacent portion of the sterno-coraco-clavicular membrane, and +tendinously from the ventral thoracic ribs. This massive muscle covers +the entire ventral surface of the thorax and converges to insert on the +ventral side of the humerus on the pectoral surface.</p> + +<p><b><i>M. supracoracoideus</i></b>, Fig. 1. The origin is from the dorsal portion of +the keel and medial portion of the sternum, and is bordered ventrally by +the origin of M. pectoralis thoracica, and laterally by <i>M. +coracobrachialis posterior</i>. The origin is also from the manubrium and +the anterolateral portion of the proximal half of the coracoid and to a +slight extent from the sterno-coraco-clavicular membrane adjacent to the +manubrium. This large pinnate muscle converges, passes through the +foramen triosseum, and inserts by a tendon on the external tuberosity of +the humerus, immediately proximal to the insertion of <i>M. pectoralis +thoracica</i>.</p> + +<p><b><i>M. coracobrachialis posterior</i></b>, Figs. 1 and 3. The origin is from the +dorsolateral half of the coracoid, anterolateral portion of the sternum +(where the area of origin is bordered medially by <i>M. supracoracoideus</i>, +posteriorly by <i>M. pectoralis thoracica</i>, and laterally by <i>M. +sternocoracoideus</i>), and also to a slight extent from the area of +attachment of the thoracic ribs to the sternum. The muscle fibers +converge along the lateral edge of the coracoid and insert on the median +crest of the humerus immediately proximal to the pneumatic foramen. In +passing from the origin on the sternum to the insertion on the humerus, +the belly of the muscle bridges the angle formed by the costal process +of the sternum and the coracoid.</p> + +<p><b><i>M. sternocoracoideus</i></b>, Figs. 2 and 3. The origin is from the entire +external surface of the costal process of the sternum, and to a small +extent from the extreme proximal ends of the thoracic ribs where they +articulate with the costal process. The muscle inserts on a triangular +area on the dorsomedial surface of the coracoid. Like <i>M. +coracobrachialis posterior</i>, this muscle bridges the angle formed by the +costal process and the coracoid.</p> + +<p><b><i>M. subcoracoideus</i></b> (ventral head), Figs. 2 and 3. The origin is from +the dorsomedial edge of the coracoid at its extreme proximal end, and to +a slight extent from the adjacent portion of the manubrium. The origin +is medial to the insertion of <i>M. sternocoracoideus</i>. The ventral head +passes anterodorsally along the medial edge of the coracoid and joins +the dorsal head (not here described). The combined muscle then inserts +by a tendon onto the internal tuberosity of the humerus.</p> + +<p><b><i>M. costi-sternalis</i></b>, Figs. 1, 2, and 3. The origin is from the anterior +edge of the sternal portion of the first four thoracic ribs. This +triangular muscle narrows and inserts on the posterior edge of the apex +of the costal process. The portion arising from the first rib may share +slips with <i>M. sternocoracoideus</i>.<span class='pagenum'><a name="Page_558" id="Page_558">[Pg 558]</a></span></p> + +<p><b><i>M. costi-sternalis anterior</i></b>, Figs. 1, 2, and 3. This muscle is +variously developed, and originates from a small area on the ventral end +of the vertebral portion of the last cervical rib. The insertion is on +the apex of the costal process, immediately anterior to the insertion of +<i>M. costi-sternalis</i>.</p> + +<p><b><i>Mm. intercostales externus</i></b>, Fig. 1. These muscles extend +posteroventrally between the vertebral portions of successive thoracic +ribs, and between the last cervical and first thoracic ribs. In the more +posterior intercostal spaces these muscles are poorly developed, but +they become progressively better developed anteriorly, and are fully +represented in the most anterior intercostal spaces.</p> + +<p><b><i>Mm. intercostales internus</i></b>, Fig. 3. These muscles resemble the +external intercostal muscles, but extend anteroventrally, with the +muscles being most fully developed posteriorly, and progressively less +so anteriorly.</p> + +<p><b><i>Costopulmonary muscles</i></b>, Fig. 3. This diagonal series of muscle slips +from the thoracic ribs attaches to the aponeurosis covering the lungs.</p> + + +<h3>Angiology</h3> + +<p>Figs. 3 and 4 show all arteries discussed for this family. The numbers +following the names or descriptions of arteries in the text refer to +numbered arteries in one or both of these figures.</p> + +<p>The right and left innominate or brachiocephalic arteries arise from the +aortic trunk and give rise to the common carotid arteries (14). The +major vessel continuing across the thoracic cavity is the subclavian +artery. Classically the subclavian is considered as continuing into the +anterior appendage as the axillary artery. However, in the species +studied, the axillary artery can best be described as a branch from the +subclavian; the pectoral stem forms a more direct continuation of the +subclavian. In traversing the thoracic cavity, the subclavian gives rise +to the thoracic, coracoid, and axillary arteries, and leaves the +thoracic cavity as the pectoral trunk, dorsal to the area where <i>Mm. +coracobrachialis posterior</i> and <i>sternocoracoideus</i> span the angle +formed by the coracoid and costal process.</p> + +<p>The pectoral trunk bifurcates into two main pectoral arteries (9), which +penetrate <i>M. pectoralis thoracica</i>. Neither the axillary artery nor +these pectoral arteries were traced in my study.</p> + +<p>The coracoid artery (2) arises from the ventral face of the subclavian +(1), either opposite the base of, or medial to, the axillary artery +(10). The coracoid artery passes ventrad between the medial edge of the +coracoid and the ventral head of <i>M. subcoracoideus</i>, and an artery (7) +is given off to supply that muscle. The main vessel then penetrates <i>M. +supracoracoideus</i> and bifurcates or ramifies into several vessels (12).</p> + +<p>Between the origin of the coracoid artery from the subclavian, and the +point where the coracoid artery passes the medial edge of the coracoid, +several branches are given off. These vessels are highly variable in +origin, as described below, and not all were always found. Along with +the coracoid artery, they are termed a "coracoid complex."</p> + +<p>The first artery (11) of this complex arises from any one of several +places: from the lateral face of the coracoid artery at its base; +independently from the subclavian immediately lateral to the origin of +the coracoid artery; and from the thoracic artery near its origin. This +vessel travels laterad, parallel to the<span class='pagenum'><a name="Page_559" id="Page_559">[Pg 559]</a></span> subclavian, and penetrates <i>M. +coracobrachialis posterior</i> at the same point that the pectoral artery +passes dorsal to that muscle.</p> + +<p>Another vessel (common stem of 4 and 5) of the coracoid complex in most +specimens arises from the anterior face of the coracoid artery and +branches into several vessels, some of which (5) supply <i>M. +subcoracoideus</i>, and some of which (4) feed <i>M. coracobrachialis +posterior</i>. The vessel occasionally shares a common stem with the main +vessel (11) to <i>M. coracobrachialis posterior</i>, and in some specimens +arises independently from the subclavian, immediately anterior to the +origin of the coracoid artery. The branch (4) to <i>M. coracobrachialis +posterior</i> was also seen to arise independently from any of the +above-mentioned positions.</p> + +<p>Two remaining vessels (6 and 8) are often found as branches from the +coracoid artery. They were small and often were collapsed in the +individuals I dissected, but were most clearly seen in <i>Iridoprocne +bicolor</i>. The vessels occasionally had a common base, and in some +specimens only one vessel was found. The first artery (6) passes mediad +into <i>M. sternocoracoideus</i>, or continues across that muscle onto the +inner face of the sternum. The second vessel (8) also supplies <i>M. +sternocoracoideus</i> or the inner surface of the sternum, and often a +large branch continues across the dorsal surface of the coracoid to <i>M. +coracobrachialis posterior</i>. Fig. 3 shows a composite of these vessels; +not all branches were seen in any one specimen. In the specimen of <i>I. +bicolor</i> a foramen existed on the lateral edge of the coracoid where the +branch (of 8) to <i>M. coracobrachialis posterior</i> passed. An examination +of skeletons of five to 10 individuals each of the five species for +which dissections were made, and of <i>Petrochelidon pyrrhonota</i> (Cliff +Swallow) and <i>Tachycineta thalassina</i> (Violet-green Swallow), in the +University of Kansas collection, showed that most coracoids of these +seven species of swallows had a small notch (as shown in Fig. 4) or a +complete foramen there.</p> + +<p>The thoracic artery (3) arises from the subclavian opposite the base of +the coracoid artery, or from the base of the coracoid artery. Of the +five specimens of <i>P. subis</i> dissected, one individual had the former +arrangement on both sides, and one had the latter on both sides, whereas +in the remaining three the thoracic artery arose from the coracoid +artery on one side and from the subclavian on the other side. The +distance between these two possible sites of origin is slight.</p> + +<p>The thoracic artery usually passes ventral to <i>M. costi-sternalis +anterior</i>. Occasionally a small artery (13) could be traced from the +main trunk of the thoracic artery to that muscle. The main thoracic +artery bifurcates near the insertion of <i>M. costi-sternalis</i>, the +branches traveling posteriad on both sides of the muscle. On one side of +one specimen this artery bifurcated immediately after leaving the +subclavian, the dorsal trunk passing dorsal to <i>M. costi-sternalis +anterior</i>, and the ventral trunk ventral to the muscle. On the other +side of the same individual the artery passed dorsal to <i>M. +costi-sternalis anterior</i>, bifurcating at the normal point.</p> + +<p>From the ventral trunk of the thoracic artery a variable number of small +vessels arises to supply the costosternal articulations. The main +ventral trunk bifurcates into two branches, one of which passes onto the +inner face of the sternum, and one of which supplies the posterior two +intercostal spaces.</p> + +<p>The dorsal thoracic trunk supplies <i>M. costi-sternalis</i>, several dorsal +intercostal areas, and the costopulmonary muscles. Minor variations in +all of the smaller branches of the thoracic artery were common.</p> + + + +<hr style="width: 65%;" /><p><span class='pagenum'><a name="Page_560" id="Page_560">[Pg 560]</a></span></p> +<h2>MYOLOGY AND ANGIOLOGY: COLUMBIDAE</h2> + +<p>Figs. 5, 6, and 7 illustrate the following muscles and arteries +described for <i>Scardafella inca</i>.</p> + + +<h3>Myology</h3> + +<p><b><i>M. pectoralis thoracica</i></b>, Fig. 5. The origin is from approximately the +ventral third of the keel, the lateral and anterior portion of the +clavicle and the adjacent sterno-coraco-clavicular membrane, and from +the lateral portion of the sternum and the fascia overlying the thoracic +ribs. This massive muscle covers the entire ventral surface of the +thorax, converges, and inserts on the pectoral surface on the ventral +side of the humerus.</p> + +<p><b><i>M. supracoracoideus</i></b>, Fig. 5. The origin is from the dorsal two-thirds +of the keel and medial half of the sternum (where the origin is bordered +ventrally, posteriorly, and laterally by the origin of <i>M. pectoralis +thoracica</i>) and from the sterno-coraco-clavicular membrane adjacent to +the coracoid. This large pinnate muscle converges, passes through the +foramen triosseum, and inserts by means of a strong tendon on the dorsal +surface of the humerus on the deltoid ridge.</p> + +<p><b><i>M. coracobrachialis posterior</i></b>, Fig. 5. The origin is from a prominent +lateral wing on the posterolateral portion of the coracoid, and from the +lateral surface of the proximal two-thirds of the coracoid. The +insertion is by means of a tendon on the internal tuberosity of the +humerus. Of the muscles described here, this one differs most strikingly +from the homologous muscle in <i>P. subis</i>. The difference can be seen by +comparing Figs. 1 and 5.</p> + +<p><b><i>M. sternocoracoideus</i></b>, Figs. 5, 6, and 7. The origin is from the +external, and to a slight extent from the internal, surface of the +costal process. The insertion is on a posterolateral triangular area on +the dorsal surface of the coracoid.</p> + +<p><b><i>M. costi-sternalis</i></b>, Figs. 5 and 6. The origin is from the anterior +edge of the sternal portion of the first three thoracic ribs. The muscle +converges and inserts on the apex of the costal process.</p> + +<p><b><i>M. subcoracoideus</i></b> (ventral head), Fig. 6. The origin is from the +manubrium and from approximately the posterior half of the coracoid and +on the medial and dorsal surface of that bone, and the medial side of +the sterno-coraco-clavicular membrane adjacent to the coracoid. The +ventral head passes anterodorsally to join with the dorsal head (not +here described), and the combined muscle inserts by a tendon on the +internal tuberosity of the humerus.</p> + +<p><b><i>Mm. intercostales externus</i></b>, Fig. 5. These muscles extend +posteroventrally between successive thoracic ribs and between the last +cervical and first thoracic ribs.</p> + +<p><b><i>Mm. intercostales internus</i></b>, Fig. 7. These muscles extend +anteroventrally between the last three thoracic ribs.</p> + +<p><b><i>Costopulmonary muscles</i></b>, Fig. 7. This series of muscle slips from the +thoracic ribs attaches to the aponeurosis covering the lungs.</p> + + +<h3>Angiology</h3> + +<p>Figs. 5, 6, and 7 show all arteries discussed for this family. The +numbers following names or descriptions of arteries in the text refer to +numbered arteries<span class='pagenum'><a name="Page_561" id="Page_561">[Pg 561]</a></span> in one of these figures. Insofar as possible, the +numbers used for these arteries are the same numbers used for the +homologous vessels in swallows.</p> + +<p>The right and left innominate arteries arise from the aortic trunk and +give rise to the common carotid (14) and subclavian (1) arteries. The +latter continues across the thoracic cavity, giving rise to the coracoid +(2) and axillary (10) arteries, and becoming the pectoral trunk. That +trunk swings posteriorly and leaves the thoracic cavity near the apex of +the costal process, as shown in Fig. 7. Where the trunk passes under <i>M. +sternocoracoideus</i>, the thoracic artery (3) is given off.</p> + +<p>The various branches of the coracoid artery, again referred to as a +"coracoid complex," are as follows: The first branch, from the posterior +face of the coracoid artery, is a relatively large vessel (6) here +termed the sternal artery; it passes mediad across <i>M. +sternocoracoideus</i>, sending off a branch (6a) to that muscle. The right +sternal artery continues posteriorly on the mid-line of the inner +surface of the sternum, and appears to send branches into the various +pneumatic foramina of the sternum, but these vessels are minute and +exceedingly difficult to trace accurately. The corresponding left vessel +is smaller and ramifies on the anteromedial surface of the sternum. +Variations found in these vessels were the following: In one specimen of +<i>S. inca</i> the sternal artery had, on both sides, an independent origin +from the subclavian, lateral to the origin of the coracoid artery. In +<i>Zenaidura macroura</i> both right and left sternal arteries were similar +to the left vessel described above, no median longitudinal vessel being +seen. In <i>Columba livia</i> no vessel corresponding to the sternal artery +was seen. In <i>Zenaida asiatica</i> these arteries penetrated <i>M. +sternocoracoideus</i>; no branch to the sternum was seen.</p> + +<p>A small complex of vessels (4 and 4a) arises from the lateral face of +the coracoid artery and feeds <i>M. coracobrachialis posterior</i>, and +occasionally <i>M. sternocoracoideus</i>. One branch (4a) passes under the +coracoid and travels along the lateral side of that bone, supplying +small branches to <i>M. coracobrachialis posterior</i>, and finally ramifying +on the head of the coracoid. In <i>C. livia</i>, <i>Zenaidura macroura</i>, and +<i>Zenaida asiatica</i> this complex usually arises independently from the +subclavian, and in one case it arose from the axillary artery.</p> + +<p>Two other branches from the coracoid artery were regularly seen. The +first (8) passes across <i>M. sternocoracoideus</i> and appears to supply the +area of the coracoid articulation with the sternum; the second (7) +supplies <i>M. subcoracoideus</i> as the main vessel passes between that +muscle and the coracoid and penetrates <i>M. suparacoracoideus</i>. A small +notch on the medial side of the coracoid (shown in Figs. 6 and 7) often +marks the passage of the coracoid artery.</p> + +<p>All vessels of the coracoid complex are exceedingly variable, in number, +size, and site of origin.</p> + +<p>A prominent vessel (15) is given off from the posterior pectoral artery, +outside the thoracic cavity, passes ventrad, and sends two branches into +<i>M. supracoracoideus</i>. No corresponding artery was seen in the swallows +dissected.</p> + +<p>The thoracic artery (3), arising from the pectoral stem, +characteristically bifurcates at the anterior end of <i>M. +costi-sternalis</i>. The dorsal, and larger, branch passes posteriorly, +sends several small branches to <i>M. costi-sternalis</i>, and continues to +the most posterior rib. The ventral trunk bifurcates, one branch passing +along the edge of, and supplying, <i>M. costi-sternalis</i>, the other<span class='pagenum'><a name="Page_562" id="Page_562">[Pg 562]</a></span> +branch passing onto the surface of the sternum. In some specimens two +such branches to the sternum were seen.</p> + + + +<hr style="width: 65%;" /> +<h2>SUMMARY OF ARTERIAL ARRANGEMENT</h2> + + +<p>In both families the vessels that are relatively constant in appearance +are: a subclavian giving rise to the carotid and axillary arteries, and +becoming the pectoral trunk; the thoracic artery arising variously, and +passing posteriorly to the rib cage; and the coracoid complex of +vessels. The coracoid complex includes the coracoid artery, the vessels +to <i>Mm. sternocoracoideus</i> and <i>coracobrachialis posterior</i>, and the +sternal artery, which is variously present, and more extensive in some +species than in others.</p> + + + +<hr style="width: 65%;" /> +<h2>DISCUSSION AND CONCLUSIONS</h2> + + +<p>In the vessels studied individual variation is marked, but the arterial +arrangement within both families is relatively constant. Interfamilial +differences probably represent responses of the arteries to adaptive +structural differences of other systems of the body.</p> + + +<h3>Individual Variation</h3> + +<p>The term "individual variation" is used here to mean "continuous +non-sex-associated variation" (see Mayr, Linsley, and Usinger, 1953:93) +found between members of the same species or between the two sides of +the same individual. It is hazardous to define individual variation (and +also interspecific differences, as discussed later) in the origin of one +vessel by relating its location to other vessels, because these may +likewise vary in origin. But, by necessity, certain vessels that are +probably less variable (axillary, carotid, and pectoral arteries) have +been considered here as being constant in origin. If these three vessels +are accepted as reference points, individual variants, as well as +interspecific differences, can easily be described in the thoracic and +coracoid arteries and in their various branches.</p> + +<p>The thoracic artery in <i>P. subis</i> arose either from the subclavian +artery, or from the coracoid artery. Likewise in other swallows, both of +these origins were found. In doves the thoracic artery arose +consistently from the pectoral stem, lateral to the origin of the +axillary artery.</p> + +<p>The coracoid artery in <i>P. subis</i> and other swallows arose from the +subclavian artery, either opposite the base of the axillary artery, or +medial to that vessel. In all doves studied the coracoid artery arose +from the subclavian medial to the axillary artery. I observed much<span class='pagenum'><a name="Page_563" id="Page_563">[Pg 563]</a></span> +individual variation in the branches of the coracoid artery (that is to +say, in the vessels of the coracoid complex). In <i>S. inca</i> the sternal +artery arose either from the coracoid artery, or independently from the +subclavian. As mentioned earlier, in members of both families the +vessels to <i>Mm. coracobrachialis posterior</i> and <i>subcoracoideus</i> are +highly variable, arising in swallows from the coracoid artery or from +the subclavian artery, and in doves from either of these two sites or +from the axillary artery. The distribution of these arteries after their +origin is also diverse.</p> + +<p>Individual variation in the arteries of the thorax has been recorded +previously. Bhaduri, Biswas, and Das (1957:2) state that, in the +domestic pigeon, "the origin and course of various smaller arteries... +show noticeable variation," although they do not specifically state to +which vessels they are referring. Fisher (1955:287-288) found +variability in the Whooping Crane, <i>Grus americana</i>, of the axillary, +coracoid, thoracic, and pectoral arteries. In one specimen he found +these vessels arising on the right side from the subclavian, in the +sequence just listed, and on the left side all arose from the same +point. Berger (1956:439-440) strongly emphasized the variability of the +vascular system, calling it the most variable in the body. As he stated, +this high degree of individual variation seems to be due to the +embryological development of the system, wherein many of the adult +channels of circulation are derived from embryonic plexuses.</p> + + +<h3>Intrafamilial Differences</h3> + +<p>In spite of the rather extensive amount of individual variability in +some vessels, I found the over-all pattern of arteries to be relatively +constant within the family Columbidae and within the family +Hirundinidae. There are, nevertheless, several intrafamilial differences +needing some further discussion and clarification.</p> + +<p>Others have reported the occasional presence of more than one coracoid +artery on each side in some columbids, these arteries being described as +arising from various sites and being variously named. Bhaduri and Biswas +(1954) described the arterial situation in seven species of the family +Columbidae (<i>Columba livia</i>, <i>Streptopelia tranquebarica</i>, <i>S. +chinensis</i>, <i>S. senegalensis</i>, <i>Chalcophaps indica</i>, <i>Treron bicincta</i>, +and <i>T. phoenicoptera</i>) and stated (<i>op. cit.</i>: 348) that "The +sternoclavicular [= coracoid] artery is similar in all the species, but +the domestic pigeon seems to be unique in that it has, in addition, a +small vessel, the accessory sternoclavicular." This artery was described +later, in the domestic pigeon, as<span class='pagenum'><a name="Page_564" id="Page_564">[Pg 564]</a></span> follows (Bhaduri, Biswas, and Das, +1957:5): "A minute and insignificant vessel which has been termed the +<i>accessory sternoclavicular</i> artery... is given off close to the origin +of the sternoclavicular. It passes anteroventrally to supply the +adjacent muscles." Glenny (1955:577) described the arterial pattern +characteristic of members of the family Columbidae (more than 30 species +studied by him) and stated that "three pairs of coracoid arteries are +found in <i>Otidiphaps nobilis</i>, normally one or two pairs may be found." +As suggested by Bhaduri and Biswas (1954:348), the "accessory" vessel +probably corresponds to a vessel previously described by Glenny (1940) +in <i>Streptopelia chinensis</i> and referred to as the "coracoid minor."</p> + +<p>Bhaduri and Biswas (1954:348) have suggested that "the accessory +sternoclavicular artery occurring sporadically as it does in some +species of diverse groups may not have any phylogenetic value."</p> + +<p>In no case did I find more than one coracoid artery on a side. When one +of the highly variable arteries feeding <i>Mm. coracobrachialis posterior</i> +and <i>sternocoracoideus</i> (arteries 4 and 4a, Fig. 7) arises from the +subclavian or axillary artery instead of from the coracoid artery, that +vessel may have been interpreted by others as a second (accessory or +minor) coracoid artery. If so, this artery probably does not "occur +sporadically." Rather, its origin from the subclavian, axillary, or +thoracic artery may be sporadic, subject to individual variation, and it +may have been overlooked when it arose from the coracoid artery.</p> + +<p>Of the vessels described here, the only one that differed distinctly in +one species was the sternal artery. In <i>Scardafella inca</i> the right +sternal vessel was long, extending down the mid-line of the inner +surface of the sternum, whereas in other columbids the right and left +arteries ramified on the anterior part of the inner surface of the +sternum, or were altogether lacking. I am unable to account for the +differential development of this artery in <i>S. inca</i>.</p> + +<p>In describing the arterial arrangement in the seven species of Indian +columbids named earlier, Bhaduri and Biswas (1954:348) state that all +species except <i>Treron phoenicoptera</i> have two "internal mammary" +arteries on each side "showing variable sites of origin." These arteries +were later described (Bhaduri, Biswas, and Das, 1957:4-5) as "a slender +(<i>outer</i>) <i>internal mammary</i> artery... to the outer wall of the thoracic +cavity" and "a slender (<i>inner</i>) <i>internal mammary</i> artery... to supply +the inner wall of the chest cavity." From this description, the question +arises as to<span class='pagenum'><a name="Page_565" id="Page_565">[Pg 565]</a></span> whether the "outer" one of these arteries should properly +be called an <i>external</i> instead of <i>internal</i> mammary artery. In any +case, I saw no specimen possessing two thoracic arteries on a side.</p> + + +<h3>Interfamilial Differences</h3> + +<p>As shown above, there is a high degree of individual variation in the +vessels being considered, while at the same time, few interspecific +differences were noted within the families. On the other hand, the +vascular arrangement of swallows consistently differed from that of +pigeons in the species studied. The differences are most easily +described by discussing the resulting change in the site of origin of +the thoracic artery. In swallows the thoracic artery arises between the +carotid and axillary arteries, either from the stem of the coracoid +artery or independently from the subclavian, but in pigeons the thoracic +artery arises from the pectoral stem, a site of attachment that is +relatively more lateral than in swallows.</p> + +<p>This difference, in my opinion, demonstrates well the topological +relationships of various systems of the body, here especially of the +skeletal, muscular, and vascular systems. The location of the thoracic +artery seems to be determined by the particular configuration of +skeletal and muscular elements, although even within the bounds set by +these elements, individual variation in the precise origin of the artery +is possible. In all swallows dissected <i>Mm. coracobrachialis posterior</i> +and <i>sternocoracoideus</i> bridge the angle formed by the costal process +and the coracoid. This arrangement makes it necessary for the subclavian +to leave the thoracic cavity dorsal to the costal process, although it +does pass immediately anterior to that process. The thoracic artery +arises from the vessel next to the apex of the costal process, hence +from the subclavian, between the axillary and carotid arteries.</p> + +<p>In pigeons, the wing of the coracoid extends farther laterally than does +the costal process, and the apex of the latter is displaced farther +posteriorly than it is in swallows. <i>M. coracobrachialis posterior</i> does +not arise from the sternum, and only part of the costal process serves +as a point of origin for <i>M. sternocoracoideus</i>. Consequently, this +region differs from that of swallows; the area between the costal +process and coracoid is not entirely bridged by muscle, and the space +between the two skeletal elements is of a different shape and size. It +seems that these differences have resulted, in pigeons, in the +subclavian assuming a more anterior position with reference to the +costal process. The subclavian in these birds leads into the pectoral +artery, which runs posteriad, passing<span class='pagenum'><a name="Page_566" id="Page_566">[Pg 566]</a></span> under <i>M. sternocoracoideus</i> and +leaving the thoracic cavity approximately opposite the apex of the +costal process. The thoracic artery arises immediately opposite the apex +of the costal process from the main artery in the area, as it does in +swallows, except that in this case the adjacent artery from which it +arises is the pectoral stem.</p> + +<p>The thoracic area seems to be most "efficiently" arranged when the +thoracic artery arises <i>opposite the apex of the costal process, from +whatever main artery is closest to that site</i>. This arrangement existed +in all species studied. Considering the differences in skeletal and +muscular structures, between pigeons and swallows, it would be much more +remarkable if an alternative were the case, that is to say if the +thoracic artery <i>had the same site of attachment on the subclavian</i> in +both groups.</p> + +<p>A comparison of these suggestions with statements made previously about +these arteries seems necessary. When Glenny (1955) summarized his +accumulative findings, concerning the main arteries in the region of the +heart, based on individuals representing more than 750 avian species of +27 orders and 120 families, he described five types of thoracic arteries +that were distinguished by differences in the site of their origin, and +one type in which there were two thoracic arteries on each side. His +statements regarding these differences were as follows (Glenny, +1955:543-544):</p> + +<div class="blockquot"><p>"The thoracic, intercostal, or internal mammary artery of +birds... is found to arise at slightly different relative +positions—from a point at the base of the inferior pectoral +artery to a point near the base of the coracoid or +sternoclavicular artery, and in some instances both of these +vessels have a common root from the subclavian artery. Such +differences are found to be of common occurrence within +several orders of birds. In the Galliformes and the +Passeriformes there appears to be a graded series in the +sites of attachment of the thoracic artery from a lateral to +a medial position. As a result of these observations, +numerical values can be assigned to the site of attachment +of the intercostal or thoracic artery, and these values may +come to be used as an index in specific levels of +evolution....</p> + +<p>"The medial migration of the thoracic artery appears to have +some phylogenetic significance as yet not understood."</p></div> + +<p>The six types of thoracic arteries described in Glenny's classification +were distinguished as follows (Glenny, 1955:544):</p> + +<div class="blockquot"><p>"Type 1: attachment to the pectoral stem lateral to the +axillary.</p> + +<p>"Type 2: attachment to the subclavian between the axillary +and coracoid.</p> + +<p>"Type 3: attachment to the subclavian at the base of the +coracoid.</p> + +<p>"Type 4: attachment to the subclavian, but with a common +root for both the coracoid and thoracic.</p> + +<p>"Type 5: attachment to the subclavian medial to both the +axillary and coracoid.<span class='pagenum'><a name="Page_567" id="Page_567">[Pg 567]</a></span></p> + +<p>"Type 6: two separate thoracic arteries are present; the +primary thoracic is the same as type 1 above, while the +secondary thoracic is the same as type 3 or type 4 above."</p></div> + +<p>Possibly the thoracic artery has undergone migration but apparent +differences in its origin might well be due to differences in other +vessels of the thoracic area. Additionally, there seems to be no reason +to assume that the lateral position of the thoracic artery is the +primitive one, or that the medial is the derived position, as is implied +by the phrase "medial migration." Although the lateral site of +attachment (type 1) is predominant in the lower orders of birds, and the +medial attachment is found primarily in Passeriformes, a fact which may +indicate that type 1 is the more primitive, it must nevertheless be kept +in mind that a sequence of a single morphological character does not +necessarily represent the phylogenetic sequence of the character itself +(see Mayr, 1955:41).</p> + +<p>Also, a given arterial arrangement might be independently derived more +than once. If such has been the case, similarities in arterial +arrangements in different taxa would sometimes be "chance similarities," +that is to say, "resemblance in characteristics developed in separate +taxa by independent causes and without causal relationship involving the +similarity as such" (Simpson, 1961:79).</p> + +<p>The particular arrangement of the arteries of the thoracic area also +seems to be of limited value as a clue to taxonomic relationships. If +the origin of any artery is determined by skeletal and muscular +features, as I suggest, the artery perhaps ought not be considered as a +separate character, but as part of a "character complex" that varies as +a unit (see Mayr, Linsley, and Usinger, 1953:123). The skeleton offers a +potential fossil record for consideration. Changes in the skeleton and +muscles, great enough to affect the blood vessels, would probably be +detected more easily than would the resulting vascular changes. Also, I +did not find as much individual variation in the skeleton and muscles in +the area studied as I did in the vascular system. In other words, within +the bounds established by the skeletal and muscular features, the artery +still exhibited individual variation in exact origin.</p> + + + +<hr style="width: 65%;" /> +<h2>SUMMARY</h2> + + +<p>The origin, distribution, and individual variation of the thoracic and +coracoid arteries, and their branches, have been studied in four species +of the family Columbidae (pigeons) and in five species of the family +Hirundinidae (swallows). These arteries are described for <i>Scardafella +inca</i> (Inca Dove) and <i>Progne subis</i> (Purple Martin). Muscles that are +supplied by these vessels, and muscles the particular<span class='pagenum'><a name="Page_568" id="Page_568">[Pg 568]</a></span> configuration of +which seems to effect the arrangement of the arteries have also been +described. Correlation of the arteries observed with those named and +described by other workers has been attempted.</p> + +<p>In most of the vessels studied there is a high degree of individual +variation, but few interspecific differences were noticed within either +family. Differences in the arteries of the thorax between the two +families are described by discussing the resulting different origins of +the thoracic artery. In swallows the thoracic artery arises from either +the subclavian artery or the coracoid artery, whereas in pigeons it +arises from the pectoral trunk. This difference in site of attachment +seems to be a result of differences between the two families in muscular +and skeletal elements of the thorax.</p> + +<p>The particular site of attachment of the thoracic artery is of limited +value as a taxonomic character. Several considerations influenced this +conclusion. (1) If the location of the artery is determined by skeletal +and muscular elements, these associated structures must be considered +taxonomically as a "character complex" (a set of characters varying as a +unit). (2) Even within the bounds established by the skeleton and +muscles, the artery displays a high degree of individual variation in +exact origin. (3) A given arterial arrangement could have been derived +independently many times. (4) Because differences are defined relative +to other likewise variable vessels, supposed similarities or differences +in the one artery may be artifacts of the system of description.</p> + +<p>My findings and interpretations do not support previous suggestions that +the thoracic artery has undergone a mediad migration, and that the +various sites of attachment of that vessel may come to represent various +levels of evolution. The primitive site of attachment of the vessel is +unknown, and it seems to me that it has not been sufficiently +demonstrated that the vessel has undergone any "migration."<span class='pagenum'><a name="Page_569" id="Page_569">[Pg 569]</a></span></p> + +<div class="figcenter" style="width: 600px;"> +<img src="images/fig1.jpg" width="600" height="431" alt="Fig. 1. Progne subis. Lateral view of left half of +thorax. M. pectoralis thoracica (area of insertion indicated by dotted +line) has been removed. Muscles not described in text are not shown. (× +1.5.)" title="" /> +<span class="caption">Fig. 1. Progne subis. Lateral view of left half of +thorax. M. pectoralis thoracica (area of insertion indicated by dotted +line) has been removed. Muscles not described in text are not shown. (× +1.5.)</span> +</div> + +<div class="figcenter" style="width: 600px;"> +<img src="images/fig2.jpg" width="600" height="424" alt="Fig. 2. Progne subis. Lateral view of left half of +thorax. Same view as shown in Fig. 1, but with Mm. supracoracoideus, +coracobrachialis posterior, and intercostales externus removed. (× +1.5.)" title="" /> +<span class="caption">Fig. 2. Progne subis. Lateral view of left half of +thorax. Same view as shown in Fig. 1, but with Mm. supracoracoideus, +coracobrachialis posterior, and intercostales externus removed. (× +1.5.)</span> +</div><p><span class='pagenum'><a name="Page_570" id="Page_570">[Pg 570]</a></span></p> + +<div class="figcenter" style="width: 600px;"> +<img src="images/fig3.jpg" width="600" height="533" alt="Fig. 3. Progne subis. Medial view of left half of +thorax. Not all muscles shown. See Fig. 4 for identification of +arteries. (× 1.5.)" title="" /> +<span class="caption">Fig. 3. Progne subis. Medial view of left half of +thorax. Not all muscles shown. See Fig. 4 for identification of +arteries. (× 1.5.)</span> +</div> + +<div class="figcenter" style="width: 600px;"> +<img src="images/fig4.jpg" width="600" height="480" alt="Fig. 4. Progne subis. Lateral view of left half of +thorax. (× 1.5.)" title="" /> +<span class="caption">Fig. 4. Progne subis. Lateral view of left half of +thorax. (× 1.5.)</span> +</div> + +<h4>(Applies also to Fig. 3.)</h4> + +<div class="poem"><div class="stanza"> +<span class="i1">1. Subclavian artery.<br /></span> +<span class="i1">2. Coracoid artery.<br /></span> +<span class="i1">3. Thoracic artery.<br /></span> +<span class="i1">4. (Unnamed.) Supplies <i>M. coracobrachialis posterior</i>.<br /></span> +<span class="i1">5. (Unnamed.) Supplies <i>M. subcoracoideus</i>.<br /></span> +<span class="i1">6. (Unnamed.) Supplies <i>M. sternocoracoideus</i> and sternum.<br /></span> +<span class="i1">7. (Unnamed.) Supplies <i>M. subcoracoideus</i>.<br /></span> +<span class="i1">8. (Unnamed.) Supplies <i>M. sternocoracoideus</i>, <i>M. coracobrachialis posterior</i>, and sternum.<br /></span> +<span class="i1">9. Pectoral artery.<br /></span> +<span class="i0">10. Axilliary artery.<br /></span> +<span class="i0">11. (Unnamed.) Supplies <i>M. coracobrachialis posterior</i>.<br /></span> +<span class="i0">12. (Unnamed.) Supplies <i>M. supracoracoideus</i>.<br /></span> +<span class="i0">13. (Unnamed.) Supplies <i>M. costi-sternalis anterior</i>.<br /></span> +<span class="i0">14. Carotid artery.<br /></span> +</div></div> +<p><span class='pagenum'><a name="Page_571" id="Page_571">[Pg 571]</a></span></p> +<div class="figcenter" style="width: 600px;"> +<img src="images/fig5.jpg" width="600" height="438" alt="Fig. 5. Scardafella inca. Lateral view of left half of +thorax. M. pectoralis thoracica (area of insertion indicated by dotted +line) has been removed. Muscles not described in text are not shown. See +legend for Fig. 7 for identification of arteries. (× 1.)" title="" /> +<span class="caption">Fig. 5. Scardafella inca. Lateral view of left half of +thorax. M. pectoralis thoracica (area of insertion indicated by dotted +line) has been removed. Muscles not described in text are not shown. See +legend for Fig. 7 for identification of arteries. (× 1.)</span> +</div> + +<div class="figcenter" style="width: 600px;"> +<img src="images/fig6.jpg" width="600" height="383" alt="Fig. 6. Scardafella inca. Lateral view of left half of +thorax. See legend for Fig. 7 for identification of arteries. (× 1.)" title="" /> +<span class="caption">Fig. 6. Scardafella inca. Lateral view of left half of +thorax. See legend for Fig. 7 for identification of arteries. (× 1.)</span> +</div><p><span class='pagenum'><a name="Page_572" id="Page_572">[Pg 572]</a></span></p> + +<div class="figcenter" style="width: 600px;"> +<img src="images/fig7.jpg" width="600" height="379" alt="Fig. 7. Scardafella inca. Medial view of left half of +thorax. (× 1.)" title="" /> +<span class="caption">Fig. 7. Scardafella inca. Medial view of left half of +thorax. (× 1.)</span> +</div> + +<h4><span class="smcap">Key</span></h4> + +<p>(Applies also to Figs. 5 and 6.) Numerals not used are those used for +<i>Progne subis</i> for which no homologous artery occurs in <i>Scardafella +inca</i>.</p> + +<div class="poem"><div class="stanza"> +<span class="i1">1. Subclavian artery.<br /></span> +<span class="i1">2. Coracoid artery.<br /></span> +<span class="i1">3. Thoracic artery.<br /></span> +<span class="i1">4. (Unnamed.) Supplies <i>Mm. coracobrachialis posterior</i> and <i>sternocoracoideus</i>.<br /></span> +<span class="i0">4a. (Unnamed.) Supplies <i>M. coracobrachialis posterior</i>.<br /></span> +<span class="i1">6. Sternal artery. (Shown as it appears on <i>right</i> side. Left sternal artery not so extensive.)<br /></span> +<span class="i0">6a. (Unnamed.) Supplies <i>M. sternocoracoideus</i>.<br /></span> +<span class="i1">7. (Unnamed.) Supplies <i>M. subcoracoideus</i>.<br /></span> +<span class="i1">8. (Unnamed.) Supplies coracoid-sternal articulation.<br /></span> +<span class="i1">9. Pectoral artery.<br /></span> +<span class="i0">10. Axillary artery.<br /></span> +<span class="i0">12. (Unnamed.) Supplies <i>M. supracoracoideus</i>.<br /></span> +<span class="i0">14. Carotid artery.<br /></span> +<span class="i0">15. (Unnamed.) Supplies <i>M. supracoracoideus</i>.<br /></span> +</div></div> + + + +<hr style="width: 65%;" /><p><span class='pagenum'><a name="Page_573" id="Page_573">[Pg 573]</a></span></p> +<h2>LITERATURE CITED</h2> + + +<p><span class="smcap">American Ornithologists' Union</span></p> + +<p>1957. Check-List of North American birds. Baltimore, Maryland, Amer. +Ornith. Union, xiv + 691 pp.</p> + + +<p><span class="smcap">Ashley, J. F.</span></p> + +<p>1941. A study of the structure of the humerus in the Corvidae. Condor, +43:184-195.</p> + + +<p><span class="smcap">Beddard, F. E.</span></p> + +<p>1898. The structure and classification of birds. London, Longmans, +Green, & Co., xx + 548 pp.</p> + + +<p><span class="smcap">Berger, A. J.</span></p> + +<p>1956. Anatomical variation and avian anatomy. Condor, 58:433-441.</p> + + +<p><span class="smcap">Bhaduri, J. L.</span>, and <span class="smcap">Biswas, B.</span></p> + +<p>1954. The main cervical and thoracic arteries of birds. Series 2. +Columbiformes, Columbidae, part 1. Anat. Anz., 100:337-350.</p> + + +<p><span class="smcap">Bhaduri, J. L.</span>, <span class="smcap">Biswas, B.</span>, and <span class="smcap">Das, S. K.</span></p> + +<p>1957. The arterial system of the domestic pigeon (<i>Columba livia</i> +Gmelin). Anat. Anz., 104:1-14.</p> + + +<p><span class="smcap">Coues, E.</span></p> + +<p>1903. Key to North American birds. Vol. I, Fifth edit. Boston, The Page +Company, xlii + 535 + [55] pp.</p> + + +<p><span class="smcap">Fisher, H. I.</span></p> + +<p>1955. Major arteries near the heart in the Whooping Crane. Condor, +57:286-289.</p> + + +<p><span class="smcap">Glenny, F. H.</span></p> + +<p>1940. The main arteries in the region of the heart of three species of +doves. Bull. Fan Mem. Inst. Biol., Zool. ser., vol. 10, pt. 4, 271-278. +(Not seen.)</p> + +<p>1955. Modifications of pattern in the aortic arch system of birds and +their phylogenetic significance. Proc. U. S. Nat. Mus., 104:525-621.</p> + + +<p><span class="smcap">Howard, H.</span></p> + +<p>1929. The avifauna of Emeryville Shellmound. Univ. Calif. Publs. Zool., +32:301-394.</p> + + +<p><span class="smcap">Howell, A. B.</span></p> + +<p>1937. Morphogenesis of the shoulder architecture: Aves. Auk, 54:364-375.</p> + + +<p><span class="smcap">Hudson, G. E.</span>, and <span class="smcap">Lanzillotti, P. J.</span></p> + +<p>1955. Gross anatomy of the wing muscles in the family Corvidae. Amer. +Midl. Nat., 53:1-44.</p> + + +<p><span class="smcap">Mayr, E.</span></p> + +<p>1955. Comments on some recent studies of song bird phylogeny. Wilson +Bull., 67:33-44.</p> + + +<p><span class="smcap">Mayr, E.</span>, <span class="smcap">Linsley, E. G.</span>, and <span class="smcap">Usinger, R. L.</span></p> + +<p>1953. Methods and principles of systematic zoology. New York, +McGraw-Hill Book Co., x + 336 pp.</p> + + +<p><span class="smcap">Simpson, G. G.</span></p> + +<p>1961. Principles of animal taxonomy. New York, Columbia Univ. Press, xiv ++ 247 pp.</p> + + +<h4><i>Transmitted June 24, 1963.</i></h4> + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of Thoracic and Coracoid Arteries In Two +Families of Birds, Columbidae and Hirundinidae, by Marion Anne Jenkinson + +*** END OF THIS PROJECT GUTENBERG EBOOK THORACIC AND CORACOID ARTERIES *** + +***** This file should be named 33558-h.htm or 33558-h.zip ***** +This and all associated files of various formats will be found in: + https://www.gutenberg.org/3/3/5/5/33558/ + +Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci +and the Online Distributed Proofreading Team at +https://www.pgdp.net. + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: Thoracic and Coracoid Arteries In Two Families of Birds, Columbidae and Hirundinidae + +Author: Marion Anne Jenkinson + +Release Date: August 28, 2010 [EBook #33558] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK THORACIC AND CORACOID ARTERIES *** + + + + +Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci +and the Online Distributed Proofreading Team at +https://www.pgdp.net. + + + + + + + +UNIVERSITY OF KANSAS PUBLICATIONS + +MUSEUM OF NATURAL HISTORY + +Volume 12, No. 13, pp. 553-573, 7 figs. + +March, 2, 1964 + +Thoracic and Coracoid Arteries In Two Families of Birds, Columbidae and +Hirundinidae + +BY + +MARION ANNE JENKINSON + +UNIVERSITY OF KANSAS +LAWRENCE +1964 + +UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + +Editors: E. Raymond Hall, Chairman, Henry S. Fitch, +Theodore H. Eaton, Jr. + +Volume 12, No. 13, pp. 553-573, 7 figs. +Published March 2, 1964 + +UNIVERSITY OF KANSAS +Lawrence, Kansas + +PRINTED BY THE STATE PRINTER +TOPEKA, KANSAS +1964 + +[Transcriber's Note: Words surrounded by tildes, like ~this~ signifies +words in bold. Words surrounded by underscores, like _this_, signifies +words in italics.] + + + + +Thoracic and Coracoid Arteries In Two Families of Birds, Columbidae and +Hirundinidae + +BY + +MARION ANNE JENKINSON + + + + +CONTENTS + + PAGE + +INTRODUCTION 555 + +METHODS AND MATERIALS 556 + +MYOLOGY AND ANGIOLOGY: HIRUNDINIDAE 557 + Myology 557 + Angiology 558 + +MYOLOGY AND ANGIOLOGY: COLUMBIDAE 560 + Myology 560 + Angiology 560 + +SUMMARY OF ARTERIAL ARRANGEMENT 562 + +DISCUSSION AND CONCLUSIONS 562 + Individual Variation 562 + Intrafamilial Differences 563 + Interfamilial Differences 565 + +SUMMARY 567 + +LITERATURE CITED 573 + + + + +INTRODUCTION + + +Most descriptions of the circulatory system of birds, largely the work +of Glenny, have dealt with arteries of the neck and thorax in a wide +variety of species. As a result of his work, Glenny offered several +hypotheses concerning the phylogenetic, hence taxonomic, significance of +differences in some of these vessels. He also described six types of +thoracic arterial arrangements and stated that these categories might +represent various levels of evolution (Glenny, 1955:543-544). + +The families Columbidae (pigeons) and Hirundinidae (swallows) have two +nearly extreme arterial types described by Glenny, and are universally +acknowledged as monophyletic. Differences within the families, +therefore, can be considered as valid intrafamilial differences. I have +investigated the thoracic and coracoid arteries and their branches in +members of these two families to determine the degree of individual +variability of the vessels, and the possible causes of interspecific and +intrafamilial differences. + + + + +METHODS AND MATERIALS + + +All specimens studied are in The University of Kansas Museum of Natural +History. They were preserved in alcohol and their blood vessels were not +injected. Dissections were made with the aid of a binocular microscope +at magnifications of 10x and 20x. + +Following is a list of the species studied, the number of individuals of +each species dissected, and the catalogue numbers of the specimens. The +nomenclature and classification are those of the American +Ornithologists' Union's _Check-List of North American Birds_, fifth +edition (1957). + + Family Columbidae + + _Zenaidura macroura_ (Linnaeus), Mourning Dove 2: 40325, 40326. + _Zenaida asiatica_ (Linnaeus), White-winged Dove 1: 40328. + _Scardafella inca_ (Lesson), Inca Dove 5: 34894, 34896, 34902, 34906, + 34907. + _Columba livia_ Gmelin, Rock Dove (domestic pigeon) 1: 40321. + + Family Hirundinidae + + _Iridoprocne bicolor_ (Vieillot), Tree Swallow 1: 38101. + _Progne subis_ (Linnaeus), Purple Martin 5: 37711, 38794, 38796, + 38798, 38804. + _Stelgidopteryx ruficollis_ (Vieillot), Rough-winged Swallow 1: 38277. + _Riparia riparia_ (Linnaeus), Bank Swallow 2: 38784, 38785. + _Hirundo rustica_ (Linnaeus), Barn Swallow 1: 38839. + +The following descriptions are of _Progne subis_ and _Scardafella inca_. +Differences in the vascular system in other members of the families +represented by _P. subis_ and _S. inca_ are mentioned at the appropriate +places. The muscles briefly described for each of these two species are +those that are supplied by the thoracic or coracoid arteries or by +branches of the same, and muscles that, by their origin, location, or +insertion, seem to affect the course or origin of one of these arteries. + +The following sources have been particularly useful for the terminology +of muscles and of skeletal features: Ashley (1941), Beddard (1898), +Coues (1903), Howard (1929), Howell (1937), and Hudson and Lanzillotti +(1955). + +The names used for most arteries are those in common usage for +vertebrates. I have not used the terms "internal mammary" and +"intercostal" artery as substitutes for "thoracic" artery, except when +referring to the work of others. The vessel's homology with the internal +mammary artery of mammals has been denied (Glenny, 1955:541), and the +name "mammary" is certainly not useful descriptively in birds. The term +"intercostal" is less objectionable, except that such a name may call to +mind segmental vessels arising from the dorsal aorta. The term +"thoracic" seems best, as it is reasonably descriptive, and has been +used by Glenny in the majority of his descriptions covering a wide +variety of birds. The name "sternoclavicular" has been used by others as +a synonym for the "coracoid" artery. I have arbitrarily chosen to use +the latter. + + +ACKNOWLEDGMENTS + +I gratefully acknowledge many valuable suggestions in my research and +the preparation of this manuscript from Professors Theodore H. Eaton, A. +Byron Leonard, Richard F. Johnston, Robert M. Mengel, and E. Raymond +Hall. Mr. Abbot S. Gaunt and Miss Sandra Lovett assisted in collecting +specimens. Final drafts of the illustrations were prepared by Mr. Thomas +Swearingen. + + + + +MYOLOGY AND ANGIOLOGY: HIRUNDINIDAE + +Figs. 1, 2, 3, and 4 illustrate the following muscles and arteries +described for _Progne subis_. + + +Myology + +~_M. pectoralis thoracica_~, Fig. 1. The origin is from slightly less than +the posterior half of the sternum, from the ventral half of the keel, +almost the entire length of the posterolateral surface of the clavicle +and adjacent portion of the sterno-coraco-clavicular membrane, and +tendinously from the ventral thoracic ribs. This massive muscle covers +the entire ventral surface of the thorax and converges to insert on the +ventral side of the humerus on the pectoral surface. + +~_M. supracoracoideus_~, Fig. 1. The origin is from the dorsal portion of +the keel and medial portion of the sternum, and is bordered ventrally by +the origin of M. pectoralis thoracica, and laterally by _M. +coracobrachialis posterior_. The origin is also from the manubrium and +the anterolateral portion of the proximal half of the coracoid and to a +slight extent from the sterno-coraco-clavicular membrane adjacent to the +manubrium. This large pinnate muscle converges, passes through the +foramen triosseum, and inserts by a tendon on the external tuberosity of +the humerus, immediately proximal to the insertion of _M. pectoralis +thoracica_. + +~_M. coracobrachialis posterior_~, Figs. 1 and 3. The origin is from the +dorsolateral half of the coracoid, anterolateral portion of the sternum +(where the area of origin is bordered medially by _M. supracoracoideus_, +posteriorly by _M. pectoralis thoracica_, and laterally by _M. +sternocoracoideus_), and also to a slight extent from the area of +attachment of the thoracic ribs to the sternum. The muscle fibers +converge along the lateral edge of the coracoid and insert on the median +crest of the humerus immediately proximal to the pneumatic foramen. In +passing from the origin on the sternum to the insertion on the humerus, +the belly of the muscle bridges the angle formed by the costal process +of the sternum and the coracoid. + +~_M. sternocoracoideus_~, Figs. 2 and 3. The origin is from the entire +external surface of the costal process of the sternum, and to a small +extent from the extreme proximal ends of the thoracic ribs where they +articulate with the costal process. The muscle inserts on a triangular +area on the dorsomedial surface of the coracoid. Like _M. +coracobrachialis posterior_, this muscle bridges the angle formed by the +costal process and the coracoid. + +~_M. subcoracoideus_~ (ventral head), Figs. 2 and 3. The origin is from +the dorsomedial edge of the coracoid at its extreme proximal end, and to +a slight extent from the adjacent portion of the manubrium. The origin +is medial to the insertion of _M. sternocoracoideus_. The ventral head +passes anterodorsally along the medial edge of the coracoid and joins +the dorsal head (not here described). The combined muscle then inserts +by a tendon onto the internal tuberosity of the humerus. + +~_M. costi-sternalis_~, Figs. 1, 2, and 3. The origin is from the anterior +edge of the sternal portion of the first four thoracic ribs. This +triangular muscle narrows and inserts on the posterior edge of the apex +of the costal process. The portion arising from the first rib may share +slips with _M. sternocoracoideus_. + +~_M. costi-sternalis anterior_~, Figs. 1, 2, and 3. This muscle is +variously developed, and originates from a small area on the ventral end +of the vertebral portion of the last cervical rib. The insertion is on +the apex of the costal process, immediately anterior to the insertion of +_M. costi-sternalis_. + +~_Mm. intercostales externus_~, Fig. 1. These muscles extend +posteroventrally between the vertebral portions of successive thoracic +ribs, and between the last cervical and first thoracic ribs. In the more +posterior intercostal spaces these muscles are poorly developed, but +they become progressively better developed anteriorly, and are fully +represented in the most anterior intercostal spaces. + +~_Mm. intercostales internus_~, Fig. 3. These muscles resemble the +external intercostal muscles, but extend anteroventrally, with the +muscles being most fully developed posteriorly, and progressively less +so anteriorly. + +~_Costopulmonary muscles_~, Fig. 3. This diagonal series of muscle slips +from the thoracic ribs attaches to the aponeurosis covering the lungs. + + +Angiology + +Figs. 3 and 4 show all arteries discussed for this family. The numbers +following the names or descriptions of arteries in the text refer to +numbered arteries in one or both of these figures. + +The right and left innominate or brachiocephalic arteries arise from the +aortic trunk and give rise to the common carotid arteries (14). The +major vessel continuing across the thoracic cavity is the subclavian +artery. Classically the subclavian is considered as continuing into the +anterior appendage as the axillary artery. However, in the species +studied, the axillary artery can best be described as a branch from the +subclavian; the pectoral stem forms a more direct continuation of the +subclavian. In traversing the thoracic cavity, the subclavian gives rise +to the thoracic, coracoid, and axillary arteries, and leaves the +thoracic cavity as the pectoral trunk, dorsal to the area where _Mm. +coracobrachialis posterior_ and _sternocoracoideus_ span the angle +formed by the coracoid and costal process. + +The pectoral trunk bifurcates into two main pectoral arteries (9), which +penetrate _M. pectoralis thoracica_. Neither the axillary artery nor +these pectoral arteries were traced in my study. + +The coracoid artery (2) arises from the ventral face of the subclavian +(1), either opposite the base of, or medial to, the axillary artery +(10). The coracoid artery passes ventrad between the medial edge of the +coracoid and the ventral head of _M. subcoracoideus_, and an artery (7) +is given off to supply that muscle. The main vessel then penetrates _M. +supracoracoideus_ and bifurcates or ramifies into several vessels (12). + +Between the origin of the coracoid artery from the subclavian, and the +point where the coracoid artery passes the medial edge of the coracoid, +several branches are given off. These vessels are highly variable in +origin, as described below, and not all were always found. Along with +the coracoid artery, they are termed a "coracoid complex." + +The first artery (11) of this complex arises from any one of several +places: from the lateral face of the coracoid artery at its base; +independently from the subclavian immediately lateral to the origin of +the coracoid artery; and from the thoracic artery near its origin. This +vessel travels laterad, parallel to the subclavian, and penetrates _M. +coracobrachialis posterior_ at the same point that the pectoral artery +passes dorsal to that muscle. + +Another vessel (common stem of 4 and 5) of the coracoid complex in most +specimens arises from the anterior face of the coracoid artery and +branches into several vessels, some of which (5) supply _M. +subcoracoideus_, and some of which (4) feed _M. coracobrachialis +posterior_. The vessel occasionally shares a common stem with the main +vessel (11) to _M. coracobrachialis posterior_, and in some specimens +arises independently from the subclavian, immediately anterior to the +origin of the coracoid artery. The branch (4) to _M. coracobrachialis +posterior_ was also seen to arise independently from any of the +above-mentioned positions. + +Two remaining vessels (6 and 8) are often found as branches from the +coracoid artery. They were small and often were collapsed in the +individuals I dissected, but were most clearly seen in _Iridoprocne +bicolor_. The vessels occasionally had a common base, and in some +specimens only one vessel was found. The first artery (6) passes mediad +into _M. sternocoracoideus_, or continues across that muscle onto the +inner face of the sternum. The second vessel (8) also supplies _M. +sternocoracoideus_ or the inner surface of the sternum, and often a +large branch continues across the dorsal surface of the coracoid to _M. +coracobrachialis posterior_. Fig. 3 shows a composite of these vessels; +not all branches were seen in any one specimen. In the specimen of _I. +bicolor_ a foramen existed on the lateral edge of the coracoid where the +branch (of 8) to _M. coracobrachialis posterior_ passed. An examination +of skeletons of five to 10 individuals each of the five species for +which dissections were made, and of _Petrochelidon pyrrhonota_ (Cliff +Swallow) and _Tachycineta thalassina_ (Violet-green Swallow), in the +University of Kansas collection, showed that most coracoids of these +seven species of swallows had a small notch (as shown in Fig. 4) or a +complete foramen there. + +The thoracic artery (3) arises from the subclavian opposite the base of +the coracoid artery, or from the base of the coracoid artery. Of the +five specimens of _P. subis_ dissected, one individual had the former +arrangement on both sides, and one had the latter on both sides, whereas +in the remaining three the thoracic artery arose from the coracoid +artery on one side and from the subclavian on the other side. The +distance between these two possible sites of origin is slight. + +The thoracic artery usually passes ventral to _M. costi-sternalis +anterior_. Occasionally a small artery (13) could be traced from the +main trunk of the thoracic artery to that muscle. The main thoracic +artery bifurcates near the insertion of _M. costi-sternalis_, the +branches traveling posteriad on both sides of the muscle. On one side of +one specimen this artery bifurcated immediately after leaving the +subclavian, the dorsal trunk passing dorsal to _M. costi-sternalis +anterior_, and the ventral trunk ventral to the muscle. On the other +side of the same individual the artery passed dorsal to _M. +costi-sternalis anterior_, bifurcating at the normal point. + +From the ventral trunk of the thoracic artery a variable number of small +vessels arises to supply the costosternal articulations. The main +ventral trunk bifurcates into two branches, one of which passes onto the +inner face of the sternum, and one of which supplies the posterior two +intercostal spaces. + +The dorsal thoracic trunk supplies _M. costi-sternalis_, several dorsal +intercostal areas, and the costopulmonary muscles. Minor variations in +all of the smaller branches of the thoracic artery were common. + + + + +MYOLOGY AND ANGIOLOGY: COLUMBIDAE + +Figs. 5, 6, and 7 illustrate the following muscles and arteries +described for _Scardafella inca_. + + +Myology + +~_M. pectoralis thoracica~_, Fig. 5. The origin is from approximately the +ventral third of the keel, the lateral and anterior portion of the +clavicle and the adjacent sterno-coraco-clavicular membrane, and from +the lateral portion of the sternum and the fascia overlying the thoracic +ribs. This massive muscle covers the entire ventral surface of the +thorax, converges, and inserts on the pectoral surface on the ventral +side of the humerus. + +~_M. supracoracoideus~_, Fig. 5. The origin is from the dorsal two-thirds +of the keel and medial half of the sternum (where the origin is bordered +ventrally, posteriorly, and laterally by the origin of _M. pectoralis +thoracica_) and from the sterno-coraco-clavicular membrane adjacent to +the coracoid. This large pinnate muscle converges, passes through the +foramen triosseum, and inserts by means of a strong tendon on the dorsal +surface of the humerus on the deltoid ridge. + +~_M. coracobrachialis posterior_~, Fig. 5. The origin is from a prominent +lateral wing on the posterolateral portion of the coracoid, and from the +lateral surface of the proximal two-thirds of the coracoid. The +insertion is by means of a tendon on the internal tuberosity of the +humerus. Of the muscles described here, this one differs most strikingly +from the homologous muscle in _P. subis_. The difference can be seen by +comparing Figs. 1 and 5. + +~_M. sternocoracoideus_~, Figs. 5, 6, and 7. The origin is from the +external, and to a slight extent from the internal, surface of the +costal process. The insertion is on a posterolateral triangular area on +the dorsal surface of the coracoid. + +~_M. costi-sternalis_~, Figs. 5 and 6. The origin is from the anterior +edge of the sternal portion of the first three thoracic ribs. The muscle +converges and inserts on the apex of the costal process. + +~_M. subcoracoideus_~ (ventral head), Fig. 6. The origin is from the +manubrium and from approximately the posterior half of the coracoid and +on the medial and dorsal surface of that bone, and the medial side of +the sterno-coraco-clavicular membrane adjacent to the coracoid. The +ventral head passes anterodorsally to join with the dorsal head (not +here described), and the combined muscle inserts by a tendon on the +internal tuberosity of the humerus. + +~_Mm. intercostales externus_~, Fig. 5. These muscles extend +posteroventrally between successive thoracic ribs and between the last +cervical and first thoracic ribs. + +~_Mm. intercostales internus_~, Fig. 7. These muscles extend +anteroventrally between the last three thoracic ribs. + +~_Costopulmonary muscles_~, Fig. 7. This series of muscle slips from the +thoracic ribs attaches to the aponeurosis covering the lungs. + + +Angiology + +Figs. 5, 6, and 7 show all arteries discussed for this family. The +numbers following names or descriptions of arteries in the text refer to +numbered arteries in one of these figures. Insofar as possible, the +numbers used for these arteries are the same numbers used for the +homologous vessels in swallows. + +The right and left innominate arteries arise from the aortic trunk and +give rise to the common carotid (14) and subclavian (1) arteries. The +latter continues across the thoracic cavity, giving rise to the coracoid +(2) and axillary (10) arteries, and becoming the pectoral trunk. That +trunk swings posteriorly and leaves the thoracic cavity near the apex of +the costal process, as shown in Fig. 7. Where the trunk passes under _M. +sternocoracoideus_, the thoracic artery (3) is given off. + +The various branches of the coracoid artery, again referred to as a +"coracoid complex," are as follows: The first branch, from the posterior +face of the coracoid artery, is a relatively large vessel (6) here +termed the sternal artery; it passes mediad across _M. +sternocoracoideus_, sending off a branch (6a) to that muscle. The right +sternal artery continues posteriorly on the mid-line of the inner +surface of the sternum, and appears to send branches into the various +pneumatic foramina of the sternum, but these vessels are minute and +exceedingly difficult to trace accurately. The corresponding left vessel +is smaller and ramifies on the anteromedial surface of the sternum. +Variations found in these vessels were the following: In one specimen of +_S. inca_ the sternal artery had, on both sides, an independent origin +from the subclavian, lateral to the origin of the coracoid artery. In +_Zenaidura macroura_ both right and left sternal arteries were similar +to the left vessel described above, no median longitudinal vessel being +seen. In _Columba livia_ no vessel corresponding to the sternal artery +was seen. In _Zenaida asiatica_ these arteries penetrated _M. +sternocoracoideus_; no branch to the sternum was seen. + +A small complex of vessels (4 and 4a) arises from the lateral face of +the coracoid artery and feeds _M. coracobrachialis posterior_, and +occasionally _M. sternocoracoideus_. One branch (4a) passes under the +coracoid and travels along the lateral side of that bone, supplying +small branches to _M. coracobrachialis posterior_, and finally ramifying +on the head of the coracoid. In _C. livia_, _Zenaidura macroura_, and +_Zenaida asiatica_ this complex usually arises independently from the +subclavian, and in one case it arose from the axillary artery. + +Two other branches from the coracoid artery were regularly seen. The +first (8) passes across _M. sternocoracoideus_ and appears to supply the +area of the coracoid articulation with the sternum; the second (7) +supplies _M. subcoracoideus_ as the main vessel passes between that +muscle and the coracoid and penetrates _M. suparacoracoideus_. A small +notch on the medial side of the coracoid (shown in Figs. 6 and 7) often +marks the passage of the coracoid artery. + +All vessels of the coracoid complex are exceedingly variable, in number, +size, and site of origin. + +A prominent vessel (15) is given off from the posterior pectoral artery, +outside the thoracic cavity, passes ventrad, and sends two branches into +_M. supracoracoideus_. No corresponding artery was seen in the swallows +dissected. + +The thoracic artery (3), arising from the pectoral stem, +characteristically bifurcates at the anterior end of _M. +costi-sternalis_. The dorsal, and larger, branch passes posteriorly, +sends several small branches to _M. costi-sternalis_, and continues to +the most posterior rib. The ventral trunk bifurcates, one branch passing +along the edge of, and supplying, _M. costi-sternalis_, the other +branch passing onto the surface of the sternum. In some specimens two +such branches to the sternum were seen. + + + + +SUMMARY OF ARTERIAL ARRANGEMENT + + +In both families the vessels that are relatively constant in appearance +are: a subclavian giving rise to the carotid and axillary arteries, and +becoming the pectoral trunk; the thoracic artery arising variously, and +passing posteriorly to the rib cage; and the coracoid complex of +vessels. The coracoid complex includes the coracoid artery, the vessels +to _Mm. sternocoracoideus_ and _coracobrachialis posterior_, and the +sternal artery, which is variously present, and more extensive in some +species than in others. + + + + +DISCUSSION AND CONCLUSIONS + + +In the vessels studied individual variation is marked, but the arterial +arrangement within both families is relatively constant. Interfamilial +differences probably represent responses of the arteries to adaptive +structural differences of other systems of the body. + + +Individual Variation + +The term "individual variation" is used here to mean "continuous +non-sex-associated variation" (see Mayr, Linsley, and Usinger, 1953:93) +found between members of the same species or between the two sides of +the same individual. It is hazardous to define individual variation (and +also interspecific differences, as discussed later) in the origin of one +vessel by relating its location to other vessels, because these may +likewise vary in origin. But, by necessity, certain vessels that are +probably less variable (axillary, carotid, and pectoral arteries) have +been considered here as being constant in origin. If these three vessels +are accepted as reference points, individual variants, as well as +interspecific differences, can easily be described in the thoracic and +coracoid arteries and in their various branches. + +The thoracic artery in _P. subis_ arose either from the subclavian +artery, or from the coracoid artery. Likewise in other swallows, both of +these origins were found. In doves the thoracic artery arose +consistently from the pectoral stem, lateral to the origin of the +axillary artery. + +The coracoid artery in _P. subis_ and other swallows arose from the +subclavian artery, either opposite the base of the axillary artery, or +medial to that vessel. In all doves studied the coracoid artery arose +from the subclavian medial to the axillary artery. I observed much +individual variation in the branches of the coracoid artery (that is to +say, in the vessels of the coracoid complex). In _S. inca_ the sternal +artery arose either from the coracoid artery, or independently from the +subclavian. As mentioned earlier, in members of both families the +vessels to _Mm. coracobrachialis posterior_ and _subcoracoideus_ are +highly variable, arising in swallows from the coracoid artery or from +the subclavian artery, and in doves from either of these two sites or +from the axillary artery. The distribution of these arteries after their +origin is also diverse. + +Individual variation in the arteries of the thorax has been recorded +previously. Bhaduri, Biswas, and Das (1957:2) state that, in the +domestic pigeon, "the origin and course of various smaller arteries... +show noticeable variation," although they do not specifically state to +which vessels they are referring. Fisher (1955:287-288) found +variability in the Whooping Crane, _Grus americana_, of the axillary, +coracoid, thoracic, and pectoral arteries. In one specimen he found +these vessels arising on the right side from the subclavian, in the +sequence just listed, and on the left side all arose from the same +point. Berger (1956:439-440) strongly emphasized the variability of the +vascular system, calling it the most variable in the body. As he stated, +this high degree of individual variation seems to be due to the +embryological development of the system, wherein many of the adult +channels of circulation are derived from embryonic plexuses. + + +Intrafamilial Differences + +In spite of the rather extensive amount of individual variability in +some vessels, I found the over-all pattern of arteries to be relatively +constant within the family Columbidae and within the family +Hirundinidae. There are, nevertheless, several intrafamilial differences +needing some further discussion and clarification. + +Others have reported the occasional presence of more than one coracoid +artery on each side in some columbids, these arteries being described as +arising from various sites and being variously named. Bhaduri and Biswas +(1954) described the arterial situation in seven species of the family +Columbidae (_Columba livia_, _Streptopelia tranquebarica_, _S. +chinensis_, _S. senegalensis_, _Chalcophaps indica_, _Treron bicincta_, +and _T. phoenicoptera_) and stated (_op. cit._: 348) that "The +sternoclavicular [= coracoid] artery is similar in all the species, but +the domestic pigeon seems to be unique in that it has, in addition, a +small vessel, the accessory sternoclavicular." This artery was described +later, in the domestic pigeon, as follows (Bhaduri, Biswas, and Das, +1957:5): "A minute and insignificant vessel which has been termed the +_accessory sternoclavicular_ artery... is given off close to the origin +of the sternoclavicular. It passes anteroventrally to supply the +adjacent muscles." Glenny (1955:577) described the arterial pattern +characteristic of members of the family Columbidae (more than 30 species +studied by him) and stated that "three pairs of coracoid arteries are +found in _Otidiphaps nobilis_, normally one or two pairs may be found." +As suggested by Bhaduri and Biswas (1954:348), the "accessory" vessel +probably corresponds to a vessel previously described by Glenny (1940) +in _Streptopelia chinensis_ and referred to as the "coracoid minor." + +Bhaduri and Biswas (1954:348) have suggested that "the accessory +sternoclavicular artery occurring sporadically as it does in some +species of diverse groups may not have any phylogenetic value." + +In no case did I find more than one coracoid artery on a side. When one +of the highly variable arteries feeding _Mm. coracobrachialis posterior_ +and _sternocoracoideus_ (arteries 4 and 4a, Fig. 7) arises from the +subclavian or axillary artery instead of from the coracoid artery, that +vessel may have been interpreted by others as a second (accessory or +minor) coracoid artery. If so, this artery probably does not "occur +sporadically." Rather, its origin from the subclavian, axillary, or +thoracic artery may be sporadic, subject to individual variation, and it +may have been overlooked when it arose from the coracoid artery. + +Of the vessels described here, the only one that differed distinctly in +one species was the sternal artery. In _Scardafella inca_ the right +sternal vessel was long, extending down the mid-line of the inner +surface of the sternum, whereas in other columbids the right and left +arteries ramified on the anterior part of the inner surface of the +sternum, or were altogether lacking. I am unable to account for the +differential development of this artery in _S. inca_. + +In describing the arterial arrangement in the seven species of Indian +columbids named earlier, Bhaduri and Biswas (1954:348) state that all +species except _Treron phoenicoptera_ have two "internal mammary" +arteries on each side "showing variable sites of origin." These arteries +were later described (Bhaduri, Biswas, and Das, 1957:4-5) as "a slender +(_outer_) _internal mammary_ artery... to the outer wall of the thoracic +cavity" and "a slender (_inner_) _internal mammary_ artery... to supply +the inner wall of the chest cavity." From this description, the question +arises as to whether the "outer" one of these arteries should properly +be called an _external_ instead of _internal_ mammary artery. In any +case, I saw no specimen possessing two thoracic arteries on a side. + + +Interfamilial Differences + +As shown above, there is a high degree of individual variation in the +vessels being considered, while at the same time, few interspecific +differences were noted within the families. On the other hand, the +vascular arrangement of swallows consistently differed from that of +pigeons in the species studied. The differences are most easily +described by discussing the resulting change in the site of origin of +the thoracic artery. In swallows the thoracic artery arises between the +carotid and axillary arteries, either from the stem of the coracoid +artery or independently from the subclavian, but in pigeons the thoracic +artery arises from the pectoral stem, a site of attachment that is +relatively more lateral than in swallows. + +This difference, in my opinion, demonstrates well the topological +relationships of various systems of the body, here especially of the +skeletal, muscular, and vascular systems. The location of the thoracic +artery seems to be determined by the particular configuration of +skeletal and muscular elements, although even within the bounds set by +these elements, individual variation in the precise origin of the artery +is possible. In all swallows dissected _Mm. coracobrachialis posterior_ +and _sternocoracoideus_ bridge the angle formed by the costal process +and the coracoid. This arrangement makes it necessary for the subclavian +to leave the thoracic cavity dorsal to the costal process, although it +does pass immediately anterior to that process. The thoracic artery +arises from the vessel next to the apex of the costal process, hence +from the subclavian, between the axillary and carotid arteries. + +In pigeons, the wing of the coracoid extends farther laterally than does +the costal process, and the apex of the latter is displaced farther +posteriorly than it is in swallows. _M. coracobrachialis posterior_ does +not arise from the sternum, and only part of the costal process serves +as a point of origin for _M. sternocoracoideus_. Consequently, this +region differs from that of swallows; the area between the costal +process and coracoid is not entirely bridged by muscle, and the space +between the two skeletal elements is of a different shape and size. It +seems that these differences have resulted, in pigeons, in the +subclavian assuming a more anterior position with reference to the +costal process. The subclavian in these birds leads into the pectoral +artery, which runs posteriad, passing under _M. sternocoracoideus_ and +leaving the thoracic cavity approximately opposite the apex of the +costal process. The thoracic artery arises immediately opposite the apex +of the costal process from the main artery in the area, as it does in +swallows, except that in this case the adjacent artery from which it +arises is the pectoral stem. + +The thoracic area seems to be most "efficiently" arranged when the +thoracic artery arises _opposite the apex of the costal process, from +whatever main artery is closest to that site_. This arrangement existed +in all species studied. Considering the differences in skeletal and +muscular structures, between pigeons and swallows, it would be much more +remarkable if an alternative were the case, that is to say if the +thoracic artery _had the same site of attachment on the subclavian_ in +both groups. + +A comparison of these suggestions with statements made previously about +these arteries seems necessary. When Glenny (1955) summarized his +accumulative findings, concerning the main arteries in the region of the +heart, based on individuals representing more than 750 avian species of +27 orders and 120 families, he described five types of thoracic arteries +that were distinguished by differences in the site of their origin, and +one type in which there were two thoracic arteries on each side. His +statements regarding these differences were as follows (Glenny, +1955:543-544): + + "The thoracic, intercostal, or internal mammary artery of + birds... is found to arise at slightly different relative + positions--from a point at the base of the inferior pectoral + artery to a point near the base of the coracoid or + sternoclavicular artery, and in some instances both of these + vessels have a common root from the subclavian artery. Such + differences are found to be of common occurrence within + several orders of birds. In the Galliformes and the + Passeriformes there appears to be a graded series in the + sites of attachment of the thoracic artery from a lateral to + a medial position. As a result of these observations, + numerical values can be assigned to the site of attachment + of the intercostal or thoracic artery, and these values may + come to be used as an index in specific levels of + evolution.... + + "The medial migration of the thoracic artery appears to have + some phylogenetic significance as yet not understood." + +The six types of thoracic arteries described in Glenny's classification +were distinguished as follows (Glenny, 1955:544): + + "Type 1: attachment to the pectoral stem lateral to the + axillary. + + "Type 2: attachment to the subclavian between the axillary + and coracoid. + + "Type 3: attachment to the subclavian at the base of the + coracoid. + + "Type 4: attachment to the subclavian, but with a common + root for both the coracoid and thoracic. + + "Type 5: attachment to the subclavian medial to both the + axillary and coracoid. + + "Type 6: two separate thoracic arteries are present; the + primary thoracic is the same as type 1 above, while the + secondary thoracic is the same as type 3 or type 4 above." + +Possibly the thoracic artery has undergone migration but apparent +differences in its origin might well be due to differences in other +vessels of the thoracic area. Additionally, there seems to be no reason +to assume that the lateral position of the thoracic artery is the +primitive one, or that the medial is the derived position, as is implied +by the phrase "medial migration." Although the lateral site of +attachment (type 1) is predominant in the lower orders of birds, and the +medial attachment is found primarily in Passeriformes, a fact which may +indicate that type 1 is the more primitive, it must nevertheless be kept +in mind that a sequence of a single morphological character does not +necessarily represent the phylogenetic sequence of the character itself +(see Mayr, 1955:41). + +Also, a given arterial arrangement might be independently derived more +than once. If such has been the case, similarities in arterial +arrangements in different taxa would sometimes be "chance similarities," +that is to say, "resemblance in characteristics developed in separate +taxa by independent causes and without causal relationship involving the +similarity as such" (Simpson, 1961:79). + +The particular arrangement of the arteries of the thoracic area also +seems to be of limited value as a clue to taxonomic relationships. If +the origin of any artery is determined by skeletal and muscular +features, as I suggest, the artery perhaps ought not be considered as a +separate character, but as part of a "character complex" that varies as +a unit (see Mayr, Linsley, and Usinger, 1953:123). The skeleton offers a +potential fossil record for consideration. Changes in the skeleton and +muscles, great enough to affect the blood vessels, would probably be +detected more easily than would the resulting vascular changes. Also, I +did not find as much individual variation in the skeleton and muscles in +the area studied as I did in the vascular system. In other words, within +the bounds established by the skeletal and muscular features, the artery +still exhibited individual variation in exact origin. + + + + +SUMMARY + + +The origin, distribution, and individual variation of the thoracic and +coracoid arteries, and their branches, have been studied in four species +of the family Columbidae (pigeons) and in five species of the family +Hirundinidae (swallows). These arteries are described for _Scardafella +inca_ (Inca Dove) and _Progne subis_ (Purple Martin). Muscles that are +supplied by these vessels, and muscles the particular configuration of +which seems to effect the arrangement of the arteries have also been +described. Correlation of the arteries observed with those named and +described by other workers has been attempted. + +In most of the vessels studied there is a high degree of individual +variation, but few interspecific differences were noticed within either +family. Differences in the arteries of the thorax between the two +families are described by discussing the resulting different origins of +the thoracic artery. In swallows the thoracic artery arises from either +the subclavian artery or the coracoid artery, whereas in pigeons it +arises from the pectoral trunk. This difference in site of attachment +seems to be a result of differences between the two families in muscular +and skeletal elements of the thorax. + +The particular site of attachment of the thoracic artery is of limited +value as a taxonomic character. Several considerations influenced this +conclusion. (1) If the location of the artery is determined by skeletal +and muscular elements, these associated structures must be considered +taxonomically as a "character complex" (a set of characters varying as a +unit). (2) Even within the bounds established by the skeleton and +muscles, the artery displays a high degree of individual variation in +exact origin. (3) A given arterial arrangement could have been derived +independently many times. (4) Because differences are defined relative +to other likewise variable vessels, supposed similarities or differences +in the one artery may be artifacts of the system of description. + +My findings and interpretations do not support previous suggestions that +the thoracic artery has undergone a mediad migration, and that the +various sites of attachment of that vessel may come to represent various +levels of evolution. The primitive site of attachment of the vessel is +unknown, and it seems to me that it has not been sufficiently +demonstrated that the vessel has undergone any "migration." + +[Illustration: FIG. 1. _Progne subis._ Lateral view of left half of +thorax. _M. pectoralis thoracica_ (area of insertion indicated by dotted +line) has been removed. Muscles not described in text are not shown. (x +1.5.)] + +[Illustration: FIG. 2. _Progne subis._ Lateral view of left half of +thorax. Same view as shown in Fig. 1, but with _Mm. supracoracoideus_, +_coracobrachialis posterior_, and _intercostales externus_ removed. (x +1.5.)] + +[Illustration: FIG. 3. _Progne subis._ Medial view of left half of +thorax. Not all muscles shown. See Fig. 4 for identification of +arteries. (x 1.5.)] + +[Illustration: FIG. 4. _Progne subis._ Lateral view of left half of +thorax. (x 1.5.) + +(Applies also to Fig. 3.) + + 1. Subclavian artery. + 2. Coracoid artery. + 3. Thoracic artery. + 4. (Unnamed.) Supplies _M. coracobrachialis posterior_. + 5. (Unnamed.) Supplies _M. subcoracoideus_. + 6. (Unnamed.) Supplies _M. sternocoracoideus_ and sternum. + 7. (Unnamed.) Supplies _M. subcoracoideus_. + 8. (Unnamed.) Supplies _M. sternocoracoideus_, _M. coracobrachialis + posterior_, and sternum. + 9. Pectoral artery. + 10. Axilliary artery. + 11. (Unnamed.) Supplies _M. coracobrachialis posterior_. + 12. (Unnamed.) Supplies _M. supracoracoideus_. + 13. (Unnamed.) Supplies _M. costi-sternalis anterior_. + 14. Carotid artery.] + +[Illustration: FIG. 5. _Scardafella inca._ Lateral view of left half of +thorax. _M. pectoralis thoracica_ (area of insertion indicated by dotted +line) has been removed. Muscles not described in text are not shown. See +legend for Fig. 7 for identification of arteries. (x 1.)] + +[Illustration: FIG. 6. _Scardafella inca._ Lateral view of left half of +thorax. See legend for Fig. 7 for identification of arteries. (x 1.)] + +[Illustration: FIG. 7. _Scardafella inca._ Medial view of left half of +thorax. (x 1.) + +KEY + +(Applies also to Figs. 5 and 6.) Numerals not used are those used for +_Progne subis_ for which no homologous artery occurs in _Scardafella +inca_. + + 1. Subclavian artery. + 2. Coracoid artery. + 3. Thoracic artery. + 4. (Unnamed.) Supplies _Mm. coracobrachialis posterior_ and + _sternocoracoideus_. + 4a. (Unnamed.) Supplies _M. coracobrachialis posterior_. + 6. Sternal artery. (Shown as it appears on _right_ side. Left sternal + artery not so extensive.) + 6a. (Unnamed.) Supplies _M. sternocoracoideus_. + 7. (Unnamed.) Supplies _M. subcoracoideus_. + 8. (Unnamed.) Supplies coracoid-sternal articulation. + 9. Pectoral artery. + 10. Axillary artery. + 12. (Unnamed.) Supplies _M. supracoracoideus_. + 14. Carotid artery. + 15. (Unnamed.) Supplies _M. supracoracoideus_.] + + + + +LITERATURE CITED + + +AMERICAN ORNITHOLOGISTS' UNION + +1957. Check-List of North American birds. Baltimore, Maryland, Amer. +Ornith. Union, xiv + 691 pp. + + +ASHLEY, J. F. + +1941. A study of the structure of the humerus in the Corvidae. Condor, +43:184-195. + + +BEDDARD, F. E. + +1898. The structure and classification of birds. London, Longmans, +Green, & Co., xx + 548 pp. + + +BERGER, A. J. + +1956. Anatomical variation and avian anatomy. Condor, 58:433-441. + + +BHADURI, J. L., and BISWAS, B. + +1954. The main cervical and thoracic arteries of birds. Series 2. +Columbiformes, Columbidae, part 1. Anat. Anz., 100:337-350. + + +BHADURI, J. L., BISWAS, B., and DAS, S. K. + +1957. The arterial system of the domestic pigeon (_Columba livia_ +Gmelin). Anat. Anz., 104:1-14. + + +COUES, E. + +1903. Key to North American birds. Vol. I, Fifth edit. Boston, The Page +Company, xlii + 535 + [55] pp. + + +FISHER, H. I. + +1955. Major arteries near the heart in the Whooping Crane. Condor, +57:286-289. + + +GLENNY, F. H. + +1940. The main arteries in the region of the heart of three species of +doves. Bull. Fan Mem. Inst. Biol., Zool. ser., vol. 10, pt. 4, 271-278. +(Not seen.) + +1955. Modifications of pattern in the aortic arch system of birds and +their phylogenetic significance. Proc. U. S. Nat. Mus., 104:525-621. + + +HOWARD, H. + +1929. The avifauna of Emeryville Shellmound. Univ. Calif. Publs. Zool., +32:301-394. + + +HOWELL, A. B. + +1937. Morphogenesis of the shoulder architecture: Aves. Auk, 54:364-375. + + +HUDSON, G. E., and LANZILLOTTI, P. J. + +1955. Gross anatomy of the wing muscles in the family Corvidae. Amer. +Midl. Nat., 53:1-44. + + +MAYR, E. + +1955. Comments on some recent studies of song bird phylogeny. Wilson +Bull., 67:33-44. + + +MAYR, E., LINSLEY, E. G., and USINGER, R. L. + +1953. Methods and principles of systematic zoology. New York, +McGraw-Hill Book Co., x + 336 pp. + + +SIMPSON, G. G. + +1961. Principles of animal taxonomy. New York, Columbia Univ. Press, xiv ++ 247 pp. + + +_Transmitted June 24, 1963._ + + + + + +End of the Project Gutenberg EBook of Thoracic and Coracoid Arteries In Two +Families of Birds, Columbidae and Hirundinidae, by Marion Anne Jenkinson + +*** END OF THIS PROJECT GUTENBERG EBOOK THORACIC AND CORACOID ARTERIES *** + +***** This file should be named 33558.txt or 33558.zip ***** +This and all associated files of various formats will be found in: + https://www.gutenberg.org/3/3/5/5/33558/ + +Produced by Chris Curnow, Joseph Cooper, Josephine Paolucci +and the Online Distributed Proofreading Team at +https://www.pgdp.net. + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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