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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/33914-8.txt b/33914-8.txt new file mode 100644 index 0000000..e7f5c2a --- /dev/null +++ b/33914-8.txt @@ -0,0 +1,3344 @@ +The Project Gutenberg EBook of Myology and Serology of the Avian Family +Fringillidae, by William B. Stallcup + +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: Myology and Serology of the Avian Family Fringillidae + A Taxonomic Study + +Author: William B. Stallcup + +Release Date: October 19, 2010 [EBook #33914] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK MYOLOGY AND SEROLOGY OF THE *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + + + + + + + ================================================================== + UNIVERSITY OF KANSAS PUBLICATIONS + MUSEUM OF NATURAL HISTORY + + + Volume 8, No. 2, pp. 157-211, figures 1-23, 4 tables + + ---------------------- November 15, 1954 ---------------------- + + + Myology and Serology + of the Avian Family Fringillidae, + A Taxonomic Study + + BY + WILLIAM B. STALLCUP + + + UNIVERSITY OF KANSAS + LAWRENCE + 1954 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, A. Byron Leonard, + Robert W. Wilson + + + Volume 8, No. 2, pp. 157-211, figures 1-23, 4 tables + Published November 15, 1954 + + + + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + + + + PRINTED BY + FERD VOILAND, JR., STATE PRINTER + TOPEKA, KANSAS + 1954 + [Union Label] + 25-4632 + + + + + Myology and Serology + of the Avian Family Fringillidae, + A Taxonomic Study + + BY + WILLIAM B. STALLCUP + + + + +CONTENTS + + + PAGE + + INTRODUCTION 160 + + MYOLOGY OF THE PELVIC APPENDAGE 162 + General Statement 162 + Materials and Methods 163 + Description of Muscles 164 + Discussion of Myological Investigations 175 + + COMPARATIVE SEROLOGY 185 + General Statement 185 + Preparation of Antigens 186 + Preparation of Antisera 188 + Methods of Serological Testing 188 + Experimental Data 190 + Discussion of Serological Investigations 190 + + CONCLUSIONS 201 + + SUMMARY 208 + + LITERATURE CITED 210 + + + + +INTRODUCTION + + +The relationships of many groups of birds within the Order +Passeriformes are poorly understood. Most ornithologists agree that +some of the passerine families of current classifications are +artificial groups. These artificial groupings are the result of early +work which gave chief attention to readily adaptive external +structures. The size and shape of the bill, for example, have been +over-emphasized in the past as taxonomic characters. It is now +recognized that the bill is a highly adaptive structure and that it +frequently shows convergence and parallelism. + +Since studies of external morphology have failed in some cases to +provide a clear understanding of the relationships of passerine birds, +it seems appropriate that attention be given to other morphological +features, to physiological features, and to life history studies in an +attempt to find other clues to relationships at the family and +subfamily levels. + +This paper reports the results of a study of the relationships of some +birds of the Family Fringillidae and is based on the comparative +myology of the pelvic appendage and on the comparative serology of +saline-soluble proteins. Where necessary for comparative purposes, +birds from other families have been included in these investigations. + +It has long been recognized that the Fringillidae include dissimilar +groups. Recent work by Beecher (1951b, 1953) on the musculature of the +jaw and by Tordoff (1954) primarily on the structure of the bony +palate has emphasized the artificial nature of the assemblage although +these authors disagree regarding major divisions within it (see +below). + +The Fringillidae have been distinguished from other families of +nine-primaried oscines by only one character--a heavy and conical bill +(for crushing seeds). Bills of this form have been developed +independently in several other, unrelated, groups; as Tordoff (1954:7) +has pointed out, _Molothrus_ of the Family Icteridae, _Psittorostra_ +of the Family Drepaniidae, and most members of the Family Ploceidae +have bills as heavy and conical as those of the fringillids. The +ploceids are distinguished from the fringillids by a single external +character: a fairly well-developed tenth primary whereas in +fringillids the tenth primary is absent or vestigial. Tordoff +(1954:20) points out, however, that this distinction is of limited +value since in other passerine families the tenth primary may be +present in some species of a genus and absent in others. The Genus +_Vireo_ is an example. Furthermore, at least one ploceid +(_Philetairus_) has a small, vestigial tenth primary, whereas some +fringillids (_Emberizoides_, for example) possess a tenth primary +which is rather large and ventrally placed (Chapin, 1917:253-254). +Thus, it is obvious that studies based on other features are necessary +in order to attain a better understanding of the relationships of the +birds involved. + +Sushkin's studies (1924, 1925) of the structure of the bony and horny +palates have served as a basis for the division of the Fringillidae +into as many as five subfamilies (Hellmayr, 1938:v): Richmondeninae, +Geospizinae, Fringillinae, Carduelinae, and Emberizinae. + +Beecher (1951b:280) points out that "the richmondenine finches arise +so uninterruptedly out of the tanagers that ornithologists have had +to draw the dividing line between the two groups arbitrarily." His +study of pattern of jaw-musculature substantiates this. He states +further that the cardueline finches arise without disjunction +from the tanagers. He suggests, therefore, that the two groups of +"tanager-finches" be made subfamilies of the Thraupidae and that a +third subfamily be maintained for the more typical tanagers. He states +that the emberizine finches are of different origin, arising from the +wood warblers (1953:307). Beecher (1951a:431; 1953:309) includes the +Dickcissel, _Spiza americana_, in the Family Icteridae, chiefly on the +basis of jaw muscle-pattern and the horny palate. + +Tordoff (1954:10-11) presents evidence that the occurrence of +palato-maxillary bones in nine-primaried birds indicates relationship +among the forms possessing them. He points out that all fringillids +except the Carduelinae possess palato-maxillaries that are either free +or more or less fused to the prepalatine bar. He points out also that +in all carduelines, the prepalatine bar is flared at its juncture with +the premaxilla, and that the mediopalatine processes are fused across +the midline; noncardueline fringillids lack these characteristics. In +addition to the above he cites differences between the carduelines and +the "other" fringillids in the appendicular skeletons, in geographic +distribution, in patterns of migration, and in habits. Tordoff +concludes, therefore, that the carduelines are not fringillids but +ploceids, their closest affinities being with the ploceid Subfamily +Estrildinae. On the basis of palatal structure, the Fringillinae and +Geospizinae are combined with the Emberizinae, the name Fringillinae +being maintained for the subfamily. The tanagers merge with the +Richmondeninae on the one hand and with the Fringillinae on the other. +On this basis, Tordoff (1954:32) suggests that the Family Fringillidae +be divided into subfamilies as follows: Richmondeninae, Thraupinae, +and Fringillinae. The carduelines are placed as the Subfamily +Carduelinae in the Family Ploceidae. + +From the foregoing, it is apparent that the two most recent lines of +research have given rise to conflicting theories regarding +relationships within the Family Fringillidae. The purpose of my +investigation, therefore, has been to gather information, from other +fields, which might clarify the relationships of these birds. + +Since the muscle pattern of the leg in the Order Passeriformes is +thought to be one of long standing and slow change, any variation +which consistently distinguishes one group of species from another +could be significant. With the hope that such variation might be +found, a study of the comparative myology of the legs was undertaken. + +The usefulness of comparative serology as a means of determining +relationship has been demonstrated in many investigations. Its use in +this instance was undertaken for several reasons: comparative serology +has its basis in biochemical systems which seem to evolve slowly; its +methods are objective; and its use has, heretofore, resulted in the +accumulation of data which seem compatible, in most instances, with +data obtained from other sources. + +I acknowledge with pleasure the guidance received in this study from +Prof. Harrison B. Tordoff of the University of Kansas. I am indebted +also to Prof. Charles A. Leone without whose direction and assistance +the serological investigations would not have been possible; to +Professors E. Raymond Hall and A. Byron Leonard whose suggestions and +criticisms have been most helpful in the preparation of this paper; +and to T. D. Burleigh of the U. S. Fish and Wildlife Service for gifts +of several specimens used in this work. Assistance with certain parts +of the study were received from a contract (NR163014) between the +Office of Naval Research of the United States Navy and the University +of Kansas. + + + + +MYOLOGY OF THE PELVIC APPENDAGE + + +General Statement + +In an excellent paper in which the muscles of the pelvic appendage of +birds are carefully and accurately described, Hudson (1937) reviewed +briefly the more important literature pertaining to the musculature of +the leg which had been published to that date. A review of such +information here, therefore, seems unnecessary. + +Myological formulae suggested by Garrod (1873, 1874) have been +extensively used by taxonomists as aids in characterizing the orders +of birds. Relatively few investigations, however, involving the +comparative myology of the leg have been undertaken at family and +subfamily levels. The works of Fisher (1946), Hudson (1948), and +Berger (1952) are notable exceptions. + +The terminology for the muscles used in this paper follows that of +Hudson (1937), except that I have followed Berger (1952) in Latinizing +all names. Homologies are not given since these are reviewed by +Hudson. Osteological terms are from Howard (1929). + + +Materials and Methods + +Specimens were preserved in a solution of one part formalin to eight +parts of water. Thorough injection of all tissues was necessary for +satisfactory preservation. Most of the down and contour feathers were +removed to allow the preservative to reach the skin. + +In preparing specimens for study, the legs and pelvic girdle were +removed and washed in running water for several hours to remove much +of the formalin. They were then transferred to a mixture of 50 per +cent alcohol and a small amount of glycerine. + +All specimens were dissected with the aid of a low power binocular +microscope. Where possible, several specimens of each species were +examined for individual differences. Such differences were found to be +slight, involving mainly size and shape of the muscles. The size is +dependent partly on the age of the bird, muscles from older birds +being larger and better developed. The shape of a muscle (whether long +and slender or short and thick) is due in part to the position in +which the leg was preserved; that is to say, a muscle may be extended +in one bird and contracted in another. For these reasons, descriptions +and comparisons are based mainly on the origin and insertion of a +muscle and on its position in relation to adjoining muscles. + +Birds dissected in this study are listed below (in the order of the A. +O. U. Check-List): + + SPECIES + + _Vireo olivaceus_ (Linnaeus) _Leucosticte tephrocotis_ + _Seiurus motacilla_ (Vieillot) (Swainson) + _Passer domesticus_ (Linnaeus) _Spinus tristis_ (Linnaeus) + _Estrilda amandava_ (Linnaeus) _Loxia curvirostra_ Linnaeus + _Poephila guttata_ (Reichenbach) _Chlorura chlorura_ (Audubon) + _Icterus galbula_ (Linnaeus) _Pipilo erythrophthalmus_ + _Molothrus ater_ (Boddaert) (Linnaeus) + _Piranga rubra_ (Linnaeus) _Calamospiza melanocorys_ + _Richmondena cardinalis_ (Linnaeus) Stejneger + _Guiraca caerulea_ (Linnaeus) _Chondestes grammacus_ (Say) + _Passerina cyanea_ (Linnaeus) _Junco hyemalis_ (Linnaeus) + _Spiza americana_ (Gmelin) _Spizella arborea_ (Wilson) + _Hesperiphona vespertina_ (Cooper) _Zonotrichia querula_ (Nuttall) + _Carpodacus purpureus_ (Gmelin) _Passerella iliaca_ (Merrem) + _Pinicola enucleator_ (Linnaeus) _Calcarius lapponicus_ (Linnaeus) + + +Description of Muscles + +The descriptions which follow are those of the muscles in the leg of +the Red-eyed Towhee, _Pipilo erythrophthalmus_. Differences between +species, where present, are noted for each muscle. The term thigh is +used to refer to the proximal segment of the leg; the term crus is +used for that segment of the leg immediately distal to the thigh. + + +_+Musculus iliotrochantericus posticus+_ (Fig. 2).--The origin of this +muscle is fleshy from the entire concave lateral surface of the ilium +anterior to the acetabulum. The fibers converge posteriorly, and the +muscle inserts by a short, broad tendon on the lateral surface of the +femur immediately distal to the trochanter. It is the largest muscle +which passes from the ilium to the femur. + +Action.--Moves femur forward and rotates it anteriorly. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus iliotrochantericus anticus+_ (Fig. 3).--Covered laterally +by the _m. iliotrochantericus posticus_, this slender muscle +has a fleshy origin from the anteroventral edge of the ilium +between the origins of the _m. sartorius_ anteriorly and the _m. +iliotrochantericus medius_ posteriorly. The _m. iliotrochantericus +anticus_ is directed caudoventrally and inserts by a broad, flat +tendon on the anterolateral surface of the femur between the heads of +the _m. femorotibialis externus_ and _m. femorotibialis medius_ and +just distal to the insertion of the _m. iliotrochantericus medius_. + +Action.--Moves femur forward and rotates it anteriorly. + +Comparison.--No significant differences noted among the species studied. + + +_+Musculus iliotrochantericus medius+_ (Fig. 3).--Smallest of the +three _iliotrochantericus_ muscles, this bandlike muscle has a fleshy +origin from the ventral edge of the ilium just posterior to the origin +of the _m. iliotrochantericus anticus_. The fibers are directed +caudoventrally, and the insertion is tendinous on the anterolateral +surface of the femur between the insertion of the other two +_iliotrochantericus_ muscles. + +Action.--Moves femur forward and rotates it anteriorly. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus iliacus+_ (Figs. 4, 5).--Arising from a fleshy origin on +the ventral edge of the ilium just posterior to the origin of the _m. +iliotrochantericus medius_, this small slender muscle passes +posteroventrally to its fleshy insertion on the posteromedial surface +of the femur just proximal to the origin of the _m. femorotibialis +internus_. + +Action.--Moves femur forward and rotates it posteriorly. + +Comparison.--No significant differences among the species studied. + + +_+Musculus sartorius+_ (Figs. 1, 4).--A long, straplike muscle, the +_sartorius_ forms the anterior edge of the thigh. The origin is +fleshy, half from the anterior edge of the ilium and from the median +dorsal ridge of this bone and half from the posterior one or two free +dorsal vertebrae. The insertion is fleshy along a narrow line on the +anteromedial edge of the head of the tibia and on the medial region of +the patellar tendon. + +Action.--Moves thigh forward and upward and extends shank. + +Comparison.--In _Loxia_ and _Spinus_, only one-third of the origin is +from the last free dorsal vertebra. In _Hesperiphona_, _Carpodacus_, +_Pinicola_, and _Leucosticte_, only one-fifth of the origin is from +this vertebra. + + +_+Musculus iliotibialis+_ (Fig. 1).--Broad and triangular, this muscle +covers most of the deeper muscles of the lateral aspect of the thigh. +The middle region is fused with the underlying _femorotibialis_ +muscles. In the distal half of this muscle there are three distinct +parts; the anterior and posterior edges are fleshy and the central +part is aponeurotic. The origin is from a narrow line along the iliac +crests--from the origin of the _m. sartorius_, anteriorly, to the +origin of the _m. semitendinosus_ posteriorly. The origin is +aponeurotic in the preacetabular region but fleshy in the +postacetabular region. The distal part of the muscle is aponeurotic +and joins with the _femorotibialis_ muscles in the formation of the +patellar tendon. This tendon incloses the patella and inserts on a +line along the proximal edges of the cnemial crests of the +tibiotarsus. + +Action.--Extends crus. + +Comparison.--In _Vireo_ the central aponeurotic portion of this muscle +is absent. + + +_+Musculus femorotibialis externus+_ (Fig. 2).--Covering the lateral +and anterolateral surfaces of the femur, this large muscle has a +fleshy origin from the lateral edge of the proximal three-fourths of +the femur. The origin separates the insertion of the _m. +iliotrochantericus anticus_ from that of the _m. ischiofemoralis_ and, +in turn, is separated from the origin of the _m. femorotibialis +medius_ by the insertions of the _m. iliotrochantericus anticus_ and +_m. iliotrochantericus medius_. Approximately midway of the length of +the femur this muscle fuses anteromesially with the _m. femorotibialis +medius_. Distally, the _m. femorotibialis externus_ contributes to the +formation of the patellar tendon which inserts on a line along the +proximal edges of the cnemial crests of the tibiotarsus. + +Action.--Extends crus. + +Comparison.--No significant differences noted among the species studied. + + +_+Musculus femorotibialis medius+_ (Figs. 2, 4).--The origin of this +muscle, which lies along the anterior edge of the femur, is fleshy +from the entire length of the femur proximal to the level of +attachment of the proximal arm of the biceps loop. Laterally this +muscle is completely fused for most of its length with the _m. +femorotibialis externus_ and contributes to the formation of the +patellar tendon, which inserts on a line along the proximal edges of +the cnemial crests of the tibiotarsus. Many of the fibers, +nevertheless, insert on the proximal edge of the patella. + +Action.--Extends crus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus femorotibialis internus+_ (Fig. 4).--One of the most +superficial muscles lying on the medial surface of the thigh, this +muscle is divided, especially near the distal end, into two parts, +lateral and medial. The origin of the lateral part is fleshy from a +line on the medial surface of the femur; the origin begins proximally +at a point near the insertion of the _m. iliacus_. The medial, bulkier +part of the muscle has a fleshy origin on the medial surface of the +lower one-third of the femur. The two parts fuse to some extent above +the points of insertion and insert on the medial edge of the head of +the tibia. + +Action.--Rotates tibia anteriorly. + +Comparison.--Two parts of this muscle variously fused; otherwise, no +significant differences in the species studied. + + +_+Musculus piriformis+_ (Fig. 3).--This muscle is represented by the +_pars caudifemoralis_ only, the _pars iliofemoralis_ being absent in +passerine birds as far as is known. The _pars caudifemoralis_ is flat, +somewhat spindle-shaped, and passes anteroventrally from the pygostyle +to the femur. The origin is tendinous from the anteroventral edge of +the pygostyle, and the insertion is semitendinous on the +posterolateral surface of the shaft of the femur about one-fourth its +length from the proximal end. + +Action.--Moves femur posteriorly and rotates it in this direction; +moves tail laterally and depresses it. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus semitendinosus+_ (Figs. 2, 3, 5).--The origin from the +extreme posterior edge of the posterior iliac crest of the ilium is +fleshy and is aponeurotic from the last vertebra of the synsacrum and +the transverse processes of several caudal vertebrae. The straplike +belly passes along the posterolateral margin of the thigh. Immediately +posterior to the knee, the muscle is divided transversely by a +ligament. That portion passing anteriorly from the ligament is the _m. +accessorius semitendinosi_ (here considered a part of the _m. +semitendinosus_) and is discussed below. The ligament continues +distally in two parts; one part inserts on the medial surface of the +_pars media_ of the _m. gastrocnemius_ and the other part fuses with +the tendon of insertion of the _m. semimembranosus_. + +The _m. accessorius semitendinosi_ extends anteriorly from the above +mentioned ligament to a fleshy insertion on the posterolateral surface +of the femur immediately proximal to the condyles. + +Action.--Moves femur posteriorly, flexes the crus and aids in +extending the tarsometatarsus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus semimembranosus+_ (Figs. 3, 4, 5).--This straplike muscle +passes along the posteromedial surface of the thigh. The origin is +semitendinous along a line on the ischium, from a point dorsal to the +middle of the ischiopubic fenestra to the posterior end of the +ischium, and from a small area of the abdominal musculature posterior +to the ischium. The insertion is by means of a broad, thin tendon on a +ridge on the medial surface of the tibia immediately distal to the +head of this bone. The tendon of insertion passes between the head of +the _pars media_ and _pars interna_ of the _m. gastrocnemius_ and is +fused with the tendon of the _m. semitendinosus_. + +Action.--Flexes crus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus biceps femoris+_ (Fig. 2).--Long, thin, and somewhat +triangular, this muscle lies on the lateral side of the thigh just +underneath the _m. iliotibialis_. Its origin is from a line along the +anterior and posterior iliac crests underneath the origin of the _m. +iliotibialis_. Anterior to the acetabulum the origin is aponeurotic, +and the edge of this aponeurosis passes over the proximal end of the +femur. The origin posterior to the acetabulum is fleshy. The most +anterior point of origin is difficult to ascertain but it lies near +the center of the anterior iliac crest. The most posterior point of +origin is immediately dorsal to the posterior end of the ilioischiatic +fenestra. Behind the knee the fibers of this muscle converge to form +the strong tendon of insertion which passes through the biceps loop, +under the tendon of origin of the _m. flexor perforatus digiti II_, +and inserts on a small tubercle on the posterolateral edge of the +fibula at the point of the tibia-fibula fusion. + +The biceps loop is tendinous and the distal end attaches to a +protuberance on the posterolateral edge of the femur at the proximal +edge of the external condyle. The proximal end attaches to the +anterolateral edge of the femur immediately proximal to the distal end +of the loop, which extends posterior to the femur. The distal arm of +this loop is connected with the tendon of origin of the _m. flexor +perforatus digiti II_ by a strong tendon. + +Action.--Flexes crus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus ischiofemoralis+_ (Fig. 3).--Short and thick, this muscle +arises directly from the lateral surface of the ischium between the +posterior iliac crest and the ischiopubic fenestra. The area of origin +extends to the posterior edge of the ischium. The insertion is +tendinous on the lateral surface of the trochanter opposite the +insertion of the _m. iliotrochantericus medius_. + +Action.--Moves femur posteriorly and rotates it in this direction. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus obturator internus+_ (Figs. 4, 7).--Lying on the inside of +the pelvis and covering the medial surface of the ischiopubic +fenestra, is this flat, pinnate, leaf-shaped muscle. The origin is +fleshy and is from the ischium and pubis around the edges of this +fenestra; none of the fibers arises from the membrane stretched across +the fenestra. Anteriorly the fibers converge and form a strong tendon +that passes through the obturator foramen and inserts on the +posterolateral surface of the trochanter of the femur. + +Action.--Rotates femur posteriorly. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus obturator externus+_ (Fig. 7).--Short and fleshy, this +muscle consists of two parts which are not easily separable but which +may be traced throughout its length. The parts are more nearly +distinct at the origin. The dorsal part arises directly from the +ischium along the dorsal edge of the obturator foramen. The larger +ventral part arises directly from the anterior and ventral edges of +the obturator foramen. The fibers of the dorsal part pass anteriorly, +cover the tendon of the _m. obturator internus_ laterally, and insert +on the trochanter around the point of insertion of the latter muscle. +The fibers of the ventral part pass parallel with the tendon of the +_m. obturator internus_ and insert on the trochanter immediately +distal and posterior to the tendon of the latter muscle. + +Action.--Rotates femur posteriorly. + +Comparison.--In _Passer_, _Estrilda_, _Poephila_, _Hesperiphona_, +_Carpodacus_, _Pinicola_, _Leucosticte_, _Spinus_ and _Loxia_, this +muscle is undivided and, in its position, origin, and insertion, +resembles the ventral part of the bipartite muscle described above. +The origin is from the anterior and ventral edges of the obturator +foramen and the insertion is on the trochanter of the femur +immediately distal and posterior to the insertion of the _m. obturator +internus_. In all other genera examined, the muscle is bipartite. In +_Chlorura_ the dorsal part is larger and better developed than it is +in the other genera. + + +_+Musculus adductor longus et brevis+_ (Figs. 3, 4, 5).--Consisting of +two distinct, straplike parts, this large muscle lies on the medial +surface of the thigh, posterior to the femur. + +The _pars anticus_ has a semitendinous origin on a line that extends +posteriorly from the posteroventral edge of the obturator foramen to a +point half way across the membrane that covers the ischiopubic +fenestra. The insertion is fleshy along the posterior surface of the +femur from the level of the insertion of the _m. piriformis_ distally +to the medial surface of the internal condyle. + +The _pars posticus_ originates by a broad, flat tendon on a line +across the posterior half of the membrane that covers the ischiopubic +fenestra. The insertion is at the point of origin of the _pars media_ +of the _m. gastrocnemius_ on the posteromedial surface of the proximal +end of the internal condyle of the femur. There is a broad tendinous +connection with the proximal end of the _pars media_ of the _m. +gastrocnemius_. The anterior edge of the _pars posticus_ is overlapped +medially by the posterior edge of the _pars anticus_. + +Action.--Flexes thigh; may flex crus also and may extend +tarsometatarsus. + +Comparison.--In _Vireo olivaceous_, the origin of this muscle does not +extend the length of the ischiopubic fenestra. The origin, +furthermore, is along the dorsal edge of the ischiopubic fenestra and +not from the membrane covering the fenestra. Finally, in this species, +the origin of the _pars posticus_ is fleshy. + + +_+Musculus tibialis anticus+_ (Figs. 2, 5).--Lying along the anterior +edge of the crus, a part of this muscle is covered by the _m. peroneus +longus_. The origin is by two distinct heads, each of which is +pinnate. The anterior head arises directly from the edges of the outer +and inner cnemial crests. The posterior head arises by a short, strong +tendon from a small pit on the anterodistal edge of the external +condyle of the femur. This tendon and the proximal end of the muscle +pass between the head of the fibula and the outer cnemial crest. The +two heads of the muscle fuse at a place slightly more than one-half of +the distance down the crus. At the distal end of the crus this muscle +gives rise to a strong tendon which passes under a fibrous loop +immediately proximal to the external condyle in company with the _m. +extensor digitorum longus_ and which passes between the condyles of +the tibia and inserts on a tubercle on the anteromedial edge of the +proximal end of the tarsometatarsus. + +Action.--Flexes tarsometatarsus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus extensor digitorum longus+_ (Figs. 3, 5, 8).--Slender and +pinnate, this muscle lies along the anteromedial surface of the tibia. +The origin is fleshy from most of the region between the cnemial +crests and from a line along the anterior surface of the proximal +fourth of the tibia. Approximately two-thirds of the distance down the +crus the muscle gives rise to the tendon of insertion which passes +through the fibrous loop near the distal end of the tibia in company +with the _m. tibialis anticus_. The tendon then passes along beneath +the supratendinal bridge at the distal end of the tibia, traverses the +anterior intercondylar fossa, and passes beneath a bony bridge on the +anteromedial surface of the proximal end of the tarsometatarsus. The +tendon continues along the anterior surface of the tarsometatarsus to +a point immediately above the bases of the toes and there gives rise +to three branches, one to the anterior surface of each foretoe. The +insertions of each branch are on the anterior surfaces of the +phalanges as shown in Fig. 8. + +Action.--Extends foretoes. + +Comparison.--This muscle is weakly developed in _Leucosticte_ and +_Calvarius_; the belly is slender and extends only half way down the +crus before giving rise to the tendon of insertion. The functional +significance of this variation is difficult to understand. The +convergence in muscle pattern shown by these two genera, however, is +in all probability the result of similarities in behavior patterns. +These birds perch less frequently than do the other birds studied. +Thus, the toes are neither flexed nor extended as often; the smaller +size of the _m. extensor digitorum longus_ may have resulted in part +from this lessened activity. Except for the variations just noted, +there are no significant differences among the species studied; even +the rather complex patterns of insertion are identical. + + +_+Musculus peroneus longus+_ (Fig. 1).--Relatively thin and straplike, +this muscle lies on the anterolateral surface of the crus and is +intimately attached to the underlying muscles. The part of the origin +from the proximal edges of the inner and outer cnemial crests is +semitendinous but the part of the origin from the lateral edge of the +shaft of the fibula is tendinous. Approximately two-thirds the +distance down the crus the muscle gives rise to the tendon of +insertion. Immediately above the external condyle of the tibiotarsus +this tendon divides. The posterior branch inserts on the proximal end +of the lateral edge of the tibial cartilage. The anterior branch +passes over the lateral surface of the external condyle to the +posterior surface of the tarsometatarsus and there unites with the +tendon of the _m. flexor perforatus digiti III_. + +Action.--Extends tarsometatarsus and flexes third digit. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus peroneus brevis+_ (Figs. 2, 3).--Lying along the +anterolateral surface of the tibia, this slender, pinnate muscle +arises from a fleshy origin along this surface and along the anterior +surface of the fibula from a point immediately proximal to the +insertion of the _m. biceps femoris_ to a point approximately +two-thirds of the way down the crus. Near the distal end of the tibia +the muscle gives rise to the tendon of insertion that passes through a +groove on the anterolateral edge of the tibia just above the external +condyle. Here the tendon is held in place by a broad fibrous loop and +passes under the anterior branch of the tendon of insertion of the _m. +peroneus longus_ and inserts on a prominence on the lateral edge of +the proximal end of the tarsometatarsus. + +Action.--Extends tarsometatarsus and may abduct it slightly. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus gastrocnemius+_ (Figs. 1, 4).--The largest muscle of the +pelvic appendage, it covers superficially all of the posterior +surface, most of the medial surface, and half of the lateral surface +of the crus. The muscle originates by three distinct heads. + +The _pars externa_ covers the posterolateral surface of the crus, is +intermediate in size between the other two heads, and arises by a +short, strong tendon from a small bony protuberance on the +posterolateral side of the distal end of the femur immediately +proximal to the fibular condyle. The tendon is intimately connected +with the distal arm of the loop for the _m. biceps femoris_. + +The _pars media_ is the smallest of the three heads and lies on the +medial surface of the crus. The head of the _pars media_ is separated +from the _pars interna_ by the tendon of insertion of the _m. +semimembranosus_ and originates by a short, strong tendon from the +posteromedial surface of the proximal end of the internal condyle of +the femur. The proximal portion of the _pars media_ has tendinous +connections with the tendon of the _m. semitendinosus_ and with the +_pars posticus_ of the _m. adductor longus et brevis_. + +The _pars interna_ is the largest of the three heads and covers most +of the medial surface of the crus. This head in its proximal portion +is distinctly divided into anterior and posterior parts, the former +overlapping the latter medially. The origin of the posterior part is +fleshy from the anterior half of the tibial head. Some of the fibers +of the anterior part arise directly from the inner cnemial crest while +its remaining fibers arise from the patellar tendon (Fig. 1) and form +a band that extends around the anterior surface of the knee, covering +the insertion of the _m. sartorius_. + +Approximately half way down the crus, the three heads give rise to the +tendon of insertion, the _tendo achillis_, which passes over and is +tightly bound to the posterior surface of the tibial cartilage. The +insertion is tendinous on the posterior surface of the hypotarsus and +along the posterolateral ridge of the tarsometatarsus. This tendon +seems to be continuous with a fascia which forms a sheath around the +posterior surface of the tarsometatarsus holding the other tendons of +this region firmly in the posterior sulcus. + +Action.--Extends tarsometatarsus. + +Comparison.--Study of the _pars externa_ and _pars media_ reveals no +significant differences among the species dissected. The _pars +interna_, however, is subject to some variation which is described +below. + + _Pars interna_ bipartite + + _Vireo_ _Chlorura_ + _Seiurus_ _Pipilo_ + _Icterus_ _Calamospiza_ + _Molothrus_ _Chondestes_ + _Piranga_ _Junco_ + _Richmondena_ _Spizella_ + _Guiraca_ _Zonotrichia_ + _Passerina_ _Passerella_ + _Spiza_ _Calcarius_ + +The two parts of the _m. gastrocnemius_ are most distinct in _Vireo_. +_Icterus_, _Molothrus_, _Richmondena_, _Guiraca_, and _Passerina_ lack +the fibrous band that passes around the front of the knee. In _Spiza_ +this band of fibers is smaller than in the other species. + + _Pars interna_ undivided + + _Passer_ _Pinicola_ + _Estrilda_ _Leucosticte_ + _Poephila_ _Spinus_ + _Hesperiphona_ _Loxia_ + _Carpodacus_ + +In _Leucosticte_, although the _pars interna_ is undivided, there is a +band of fibers which extends around the front of the knee (see +discussion, p. 183). + + +_+Musculus plantaris+_ (Fig. 5).--Small and slender, this muscle lies +on the posteromedial surface of the crus, beneath the _pars interna_ +of the _m. gastrocnemius_ and originates by fleshy fibers from the +posteromedial surface of the proximal end of the tibia immediately +distal to the internal articular surface. The belly extends +approximately one-sixth of the way down the crus and gives rise to a +long, slender tendon that inserts on the proximomedial edge of the +tibial cartilage. + +Action.--Extends tarsometatarsus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus flexor perforatus digiti II+_ (Figs. 3, 9).--This is a +slender muscle which lies on the lateral side of the crus beneath the +_pars externa_ of the _m. gastrocnemius_ and is intimately connected +anteromedially with the _m. flexor digitorum longus_ and +posteromedially with the _m. flexor hallucis longus_. The origin is by +a strong tendon from the lateral surface of the external condyle of +the femur at the point of origin of the _m. flexor perforans et +perforatus digiti II_. This tendon serves also as the origin of the +anterior head of the _m. flexor hallucis longus_. The tendon connects +also by a broad tendinous band with the distal arm of the loop for the +_m. biceps femoris_ and by a similar band with the lateral edge of the +fibula immediately distal to the head. The tendon of insertion passes +distally, perforates the tibial cartilage near its lateral edge, +traverses the middle medial canal of the hypotarsus (Fig. 6), and +passes distally to the foot. At the distal end of the tarsometatarsus +the tendon is held against the medial surface of the first metatarsal +by a straplike sheath. The tendon then passes over a sesamoid bone +between the first metatarsal and the base of the second digit and is +bound to this bone by a sheath. The tendon inserts mainly along the +posteromedial edge of the proximal end of the first phalanx of the +second digit, although the termination is sheathlike and covers the +entire posterior surface of this phalanx. This sheathlike termination +is perforated by the tendons of the _m. flexor perforans et perforatus +digiti II_ and the branch of the _m. flexor digitorum longus_ that +inserts on the second digit. + +Action.--Flexes second digit. + +Comparison.--In _Vireo_ this muscle is larger and more deeply situated +than it is in the other species examined and has no connection with +the _m. flexor hallucis longus_. + + +_+Musculus flexor perforatus digiti III+_ (Fig. 5).--Long and +flattened, this muscle lies on the posteromedial side of the crus +beneath the _m. gastrocnemius_. The belly is tightly fused laterally +with the belly of the _m. flexor hallucis longus_ and posteriorly with +the belly of the _m. flexor perforatus digiti IV_. The origin is by a +long, strong tendon from a small tubercle just medial to, and at the +proximal end of, the external condyle of the femur. Below the middle +of the crus this muscle terminates in a strong tendon which perforates +the tibial cartilage near its lateral edge. In this region the tendon +is sheathlike and wrapped around the tendon of the _m. flexor +perforatus digiti IV_. These two tendons together pass through the +posterolateral canal of the hypotarsus (Fig. 6). Immediately distal to +the hypotarsus the two tendons separate, and the tendon of the _m. +flexor perforatus digiti III_ receives a branch of the tendon of the +_m. peroneus longus_. The tendon passes distally over the surface of +the second trochlea, and its insertion is sheathlike on the posterior +surface of the first phalanx, and on the proximal end of the second. +In the area of insertion this tendon is perforated by that of the _m. +flexor perforans et perforatus digiti III_ and by that of the _m. +flexor digitorum longus_ to the third digit. + +Action.--Flexes digit III. + +Comparison.--In _Passer_, _Estrilda_, _Poephila_, _Hesperiphona_, +_Carpodacus_, _Pinicola_, _Leucosticte_, _Spinus_, and _Loxia_ the +edges of the sheathlike tendon are thickened at the points of +insertion, so that the tendon appears to have two branches which +insert along the posterolateral edges of the first phalanx and are +connected medially by a fascia. + + +_+Musculus flexor perforatus digiti IV+_ (Fig. 3).--Extending along +the posterior edge of the crus, this slender muscle lies beneath the +_m. gastrocnemius_. The belly is fused with those of the _m. flexor +hallucis longus_ and _m. flexor perforatus digiti III_. Its origin is +fleshy from the intercondyloid region of the distal end of the femur +and has a few fibers arising from the tendon of origin of the _m. +flexor perforatus digiti III_. Near the distal end of the crus the +muscle gives rise to the strong tendon of insertion which perforates +the tibial cartilage near its lateral edge and in this region is +ensheathed by the tendon of the _m. flexor perforatus digiti III_. The +two tendons pass together through the posterolateral canal of the +hypotarsus (Fig. 6). The tendon continues distally along the +tarsometatarsus and the posterior surface of digit IV. The tendon +bifurcates at approximately the middle of the first phalanx. A short +lateral branch inserts on the posterolateral edge of the proximal end +of the second phalanx. The long medial branch is perforated by a +branch of the _m. flexor digitorum longus_; the distal end is +flattened, has thickened edges, and inserts over the posterior +surfaces of the distal end of the second phalanx, and over the +proximal end of the third phalanx. + +Action.--Flexes digit IV. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus flexor perforans et perforatus digiti II+_ (Figs. 2, +9).--Small and spindle-shaped, this muscle lies on the posterolateral +side of the crus immediately beneath the _pars externa_ of the _m. +gastrocnemius_. The origin is fleshy and arises in company with the +_m. flexor perforans et perforatus digiti III_ from a point on the +posterolateral surface of the distal end of the femur between the +point of origin of the _pars externa_ of the _m. gastrocnemius_ and +the fibular condyle. The belly extends approximately one-fourth of the +way down the crus and gives rise to the tendon of insertion which +passes distally and superficially through the posterior edge of the +tibial cartilage. The tendon traverses the posteromedial canal of the +hypotarsus (Fig. 6) and continues along the posterior surface of the +tarsometatarsus. Between the first metatarsal and the base of the +second digit the tendon is enclosed by the medial surface of a +sesamoid bone. This tendon then perforates that of the _m. flexor +perforatus digiti II_ at the level of the first phalanx and in turn is +perforated by the tendon of the _m. flexor digitorum longus_ at the +proximal end of the second phalanx. The insertion is on the posterior +surface of the second phalanx. + +Action.--Flexes digit II. + +Comparison.--In _Passer_, _Estrilda_, _Poephila_, _Hesperiphona_, +_Carpodacus_, _Pinicola_, _Leucosticte_, _Spinus_, and _Loxia_ the +proximal portion of this muscle is more intimately connected with the +posterior edge of the _m. flexor perforans et perforatus digiti III_ +than it is in the other species examined. + + +_+Musculus flexor perforans et perforatus digiti III+_ (Fig. 2).--Long +and pinnate, this muscle lies on the lateral surface of the crus +beneath the _m. peroneus longus_ and _pars externa_ of the _m. +gastrocnemius_. There are two distinct heads. The origin of the +anterior head is fleshy from the proximal edge of the outer cnemial +crest and from the internal edge of the distal end of the patellar +tendon. The posterior head arises by a tendon from the femur in +company with the _m. flexor perforans et perforatus digiti II_, is +connected also with the tendon of origin of the _m. flexor perforatus +digiti II_, and is loosely attached to the head of the fibula. Fibers +from the belly of the muscle attach throughout its length to the +lateral edge of the fibula, and the muscle is tightly fused also with +adjacent muscles. The tendon of insertion is formed approximately +one-half the way down the crus. The tendon perforates the posterior +surface of the tibial cartilage and passes through the posteromedial +canal of the hypotarsus (Fig. 6). At the base of the third digit the +tendon ensheathes that of the _m. flexor digitorum longus_ and the two +together perforate the tendon of the _m. flexor perforatus digiti +III_. Immediately distal to this perforation the tendon of the _m. +flexor perforans et perforatus digiti III_ ceases to ensheath that of +the _m. flexor digitorum longus_. The latter passes beneath that of +the former. Near the distal end of the second phalanx the tendon of +the _m. flexor digitorum longus_ perforates that of the _m. flexor +perforans et perforatus digiti III_. The latter inserts on the +posterior surface of the distal end of the second phalanx and the +proximal end of the third. + +Action.--Flexes digit III. + +Comparison.--In _Passer_, _Estrilda_, and _Poephila_, and in all the +cardueline finches examined the proximal portion of this muscle is +more intimately connected with the anterior edge of the _m. flexor +perforans et perforatus digiti II_ than it is in the other species +examined. + + +_+Musculus flexor digitorum longus+_ (Figs. 3, 5).--This strong, +pinnate muscle is deeply situated along the posterior surfaces of the +tibia and fibula. There are two distinct heads of origin. The lateral +head arises by means of fleshy fibers from the posterior edge of the +head of the fibula. The medial head arises by means of fleshy fibers +from the region under the ledgelike external and internal articular +surfaces of the proximal end of the tibia. Neither head has any +connection with the femur in contrast to the condition, described by +Hudson (1937: 46-47) in the crow, _Corvus brachyrhynchos_, and in the +raven, _Corvus corax_. Near the point of insertion of the _m. biceps +femoris_ the two heads fuse. The common belly is attached by fleshy +fibers to the posterior surface of the tibia and fibula for two-thirds +of the distance down the crus. Near the distal end of the crus the +muscle terminates in a strong tendon which passes deeply through the +tibial cartilage and traverses the anteromedial canal of the +hypotarsus (Fig. 6). About midway down the tarsometatarsus this tendon +becomes ossified. Immediately above the bases of the toes it gives +rise to three branches, one to the posterior surface of each of the +foretoes. These branches perforate the other flexor muscles of the +toes as described in the accounts of those muscles and insert as +follows: The branch to digit II inserts on the base of the ungual +phalanx and by a stout, tendinous slip on the distal end of the second +phalanx (Fig. 9). The branch to digit III inserts on the base of the +distal end of the third phalanx and a stronger slip to the distal end +of the second or proximal end of the third. The branch to digit IV +inserts on the base of the ungual phalanx, with one tendinous slip to +the distal end of the third phalanx and another to the distal end of +the fourth. + +Action.--Flexes foretoes. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus flexor hallucis longus+_ (Fig. 3).--Situated immediately +posterior to the _m. flexor digitorum longus_, the belly of this +large, pinnate muscle is intimately connected anteriorly to that of +the _m. flexor perforatus digiti II_. The _m. flexor hallucis longus_ +arises by two heads which are separated by the tendon of insertion of +the _m. biceps femoris_. The smaller anterior head arises from the +same tendon as does the _m. flexor perforatus digiti II_. The larger +posterior head arises by means of fleshy fibers from the +intercondyloid region of the posterior surface of the femur along with +the _m. flexor perforatus digiti III_ and _IV_. The two heads join +just distal to the point of insertion of the _m. biceps femoris_. +There is no trace of a tendinous band connecting the two heads as +there is in the crow and in the raven (Hudson, 1937:49). Near the +distal end of the shank the muscle gives rise to a strong tendon which +perforates the tibial cartilage along its lateral edge and passes +through the anterolateral canal of the hypotarsus (Fig. 6). The tendon +crosses over to the medial surface of the tarsometatarsus, passes +distally, and perforates the sheathlike tendon of the _m. flexor +hallucis brevis_ between the first metatarsal and the trochlea for +digit II. The tendon continues along the posterior surface of the +hallux and has a double insertion; the main tendon attaches to the +base of the ungual phalanx and a smaller branch inserts on the distal +end of the proximal phalanx. + +Action.--Flexes hallux. + +Comparison.--In _Vireo_ this muscle has only the posterior head of +origin and is not connected with the _m. flexor perforatus digiti II_. +The muscle is proportionately smaller and weaker than in any of the +other species studied. + + +_+Musculus extensor hallucis longus+_ (Fig. 4).--One of the smallest +muscles of the leg, the origin is fleshy from the anteromedial edge of +the proximal end of the tarsometatarsus. The belly is long and slender +and terminates distally in a slender tendon which passes distally +along the posterior surfaces of the first metatarsal and the first +digit. The insertion is on the base of the ungual phalanx. Near the +distal end of the proximal phalanx, the tendon passes between two +thick bands of fibro-elastic tissue which insert also on the ungual +phalanx. These bands of tissue function as automatic extensors of the +claw. + +Action.--Extends hallux; action must be slight. + +Comparison.--In _Vireo_ this muscle is proportionately larger and +better developed than it is in any of the other species examined. + + +_+Musculus flexor hallucis brevis+_ (Fig. 4).--This minute muscle has +a fleshy origin from the medial surface of the hypotarsus. The short +belly terminates in a weak, slender tendon which passes down the +posteromedial surface of the tarsometatarsus and into the space +between the first metatarsal and the trochlea for digit II. In this +region the tendon envelops the tendon of the _m. flexor hallucis +longus_ and inserts on the distal end of the first metatarsal and on +the proximal end of the first phalanx of the first digit. + +Action.--Flexes hallux; action must be slight. + +Comparison.--The small size of this muscle makes it exceedingly +difficult to study. The muscle is larger in _Vireo_ than in any of the +other species examined. This may be correlated with the smaller size +of the _m. flexor hallucis longus_ in this species. The muscle does +not seem to be so well developed in the cardueline finches as it is in +the other species. + + +_+Musculus abductor digiti IV+_ (Fig. 2).--Extremely small, delicate +and difficult to demonstrate, this muscle arises in a fleshy origin +immediately from underneath the posterior edge of the external cotyla +of the tarsometatarsus. The tendon of insertion is long and slender +and inserts along the lateral edge of the first phalanx of digit IV. + +Action.--Abducts digit IV. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus lumbricalis.+_--Semitendinous throughout its length, this +muscle arises from the ossified tendon of the _m. flexor digitorum +longus_ at a point immediately proximal to the branching of this +tendon. The insertion is on the joint pulleys and capsules at the base +of the third and fourth digits. + +Action.--Hudson (1937:57) states that: "Meckel (_vide_ Gadow--1891, p. +204) considered this muscle as serving to draw the joint pulley behind +in order to protect it from pinching during the bending of the toes. +It perhaps also tends to flex the third and fourth digits." + +Comparison.--No significant differences noted among the species +studied. + + +Discussion of the Myological Investigations + +Simpson (1944:12) and others have emphasized that different parts of +organisms evolve at different rates. Beecher (1951b:275) in stating +that "... the hind limb is very similar in muscle pattern throughout +the Order Passeriformes and seems to have become relatively static +after attaining a high level of general efficiency ..." implies that +the muscle pattern of the leg must be one of long standing and slow +change. This concept was emphasized by Hudson (1937) who found but +little variation in muscle pattern among members of the several +families of passerine birds. The concept is further confirmed by the +present investigation. The intricate patterns of origin and of +insertion seem to remain almost the same throughout the order in spite +of adaptive radiation which has occurred. + +Two major differences in patterns of leg-musculature, however, were +found among the species studied, and these differences are significant +since they are consistent between subfamilies. The muscles involved +are the _m. obturator externus_ and the _pars interna_ of the _m. +gastrocnemius_. + +The _m. obturator externus_ is bipartite, consisting of dorsal and +ventral parts, in the passerine species studied by Hudson (1937) and +in all of the species examined by me except the ploceids and the +cardueline finches. In the ploceids and cardueline finches this muscle +is undivided and resembles in its position, origin, and insertion only +the ventral portion of the muscle found in the other birds studied. It +is difficult to imagine what advantage or disadvantage might be +associated with the bipartite or with the undivided condition. The +action of this muscle is to rotate the femur (right femur clockwise, +left femur counterclockwise), and certainly the greater mass of the +bipartite muscle could lend greater strength to such action. The +possible significance of this is discussed below. + + List of Abbreviations Used in Figures + + Abd. dig. IV _M. abductor digiti IV_ + Acc. _M. accessorius semitendinosi_ + Add. long. _M. adductor longus et brevis_ + Anterolat. can. Anterolateral canal of hypotarsus + Anteromed. can. Anteromedial canal of hypotarsus + Bic. fem. _M. biceps femoris_ + Bic. loop Loop for _m. biceps femoris_ + Ext. cot. External cotyla + Ext. dig. l. _M. extensor digitorum longus_ + Ext. hal. l. _M. extensor hallucis longus_ + Fem. tib. ext. _M. femorotibialis externus_ + Fem. tib. int. _M. femorotibialis internus_ + Fem. tib. med. _M. femorotibialis medius_ + F. dig. l. _M. flexor digitorum longus_ + F. hal. brev. _M. flexor hallucis brevis_ + F. hal. l. _M. flexor hallucis longus_ + F. p. et p. d. II _M. flexor perforans et perforatus digiti II_ + F. p. et p. d. III _M. flexor perforans et perforatus digiti III_ + F. per. d. II _M. flexor perforatus digiti II_ + F. per. d. III _M. flexor perforatus digiti III_ + F. per. d. IV _M. flexor perforatus digiti IV_ + Gas. _M. gastrocnemius_ + Iliacus _M. iliacus_ + Il. tib. _M. iliotibialis_ + Il. troc. ant. _M. iliotrochantericus anticus_ + Il. troc. med. _M. iliotrochantericus medius_ + Il. troc. post. _M. iliotrochantericus posticus_ + Int. cot. Internal cotyla + Isch. fem. _M. ischiofemoralis_ + Midmed. can. Midmedial canal of hypotarsus + Obt. ext. _M. obturator externus_ + Obt. int. _M. obturator internus_ + P. ant. _Pars anticus_ + P. ext. _Pars externa_ + P. int. _Pars interna_ + P. med. _Pars media_ + P. post. _Pars posticus_ + Per. brev. _M. peroneus brevis_ + Per. long. _M. peroneus longus_ + Pirif. _M. piriformis_ + Plan. _M. plantaris_ + Posterolat. can. Posterolateral canal of hypotarsus + Posteromed. can. Posteromedial canal of hypotarsus + Sar. _M. sartorius_ + Semim. _M. semimembranosus_ + Semit. _M. semitendinosus_ + Tib. ant. _M. tibialis anticus_ + Tib. cart. Tibial cartilage + + [Illustration: FIG. 1. _Pipilo erythrophthalmus._ Lateral view of + the superficial muscles of the left leg, × 1.5.] + + [Illustration: FIG. 2. _Pipilo erythrophthalmus._ Lateral view of + the left leg showing a deeper set of muscles. The superficial + muscles _iliotibialis_, _sartorius_, _gastrocnemius_ and + _peroneus longus_ have been removed, × 1.5.] + + [Illustration: FIG. 3. _Pipilo erythrophthalmus._ Lateral view of + the left leg showing the still deeper muscles. In addition to + those listed for figure 2, the following muscles have been + wholly or partly removed: _iliotrochantericus posticus_, + _femorotibialis externus_, _femorotibialis medius_, + _biceps femoris_, _semitendinosus_, _tibialis anticus_, + _flexor perforans et perforatus digiti II_, and _flexor + perforans et perforatus digiti III_, × 1.5.] + + [Illustration: FIG. 4. _Pipilo erythrophthalmus._ Medial view of + the superficial muscles of the left leg, × 1.5.] + + [Illustration: FIG. 5. _Pipilo erythrophthalmus._ Medial view of + the left leg showing a deeper set of muscles than those seen + in figure 4. The following superficial muscles have been + removed: _iliotibialis_, _sartorius_, _femorotibialis internus_, + _obturator internus_, _adductor longus (pars posticus)_, + _gastrocnemius_, and _peroneus longus_, × 1.5.] + + [Illustration: FIG. 6. _Pipilo erythrophthalmus._ Proximal end of + left tarsometatarsus and the hypotarsus, × 4.] + + [Illustration: FIG. 7. _Pipilo erythrophthalmus._ Lateral view of + proximal end of left femur and a portion of the pelvis, × 3.5.] + + [Illustration: FIG. 8. _Pipilo erythrophthalmus._ Upper surfaces + of the phalanges of the foretoes of the left foot showing + insertions of the _M. extensor digitorum longus_, × 3.] + + [Illustration: FIG. 9. _Pipilo erythrophthalmus._ Medial view of + the second digit of the left foot, showing insertions of the + flexor muscles, × 3.] + +The division of the _pars interna_ of the _m. gastrocnemius_ into +anterior and posterior parts has not been reported by previous authors +yet the division is quite distinct in those birds in which it occurs. +Hudson (1937:36) points out that in some non-passerine birds the _pars +interna_ is double, but that in these species the _m. semimembranosus_ +inserts between the two parts. This is not the condition in those +species studied by me. Only the ploceids and the cardueline finches in +the present investigation fail to show such a division. The undivided +muscle in these birds resembles, in its origin and position, the +posterior portion of the muscle found in those species showing the +bipartite condition. The greater mass of the bipartite muscle probably +makes possible a stronger extension of the tarsometatarsus. + +Thus, the divided or undivided conditions of the _m. obturator +externus_ and the _pars interna_ of the _m. gastrocnemius_ seem to be +correlated with the degrees of strength of certain movements of the +leg. It is conceivable that these differences in structure are +correlated with the manner in which food is obtained, the birds having +the bipartite muscles being those which spend the most time on the +ground searching and scratching for seeds and other sorts of food. +Yet, in _Leucosticte_, a cardueline, and in _Calcarius_, an +emberizine, whose foraging habits are rather similar, the structure is +unlike. _Leucosticte_ does resemble the emberizines and also _Piranga_ +and _Spzia_ in the extension of a band of muscle fibers from the _pars +interna_ of the _m. gastrocnemius_ around the front of the knee. A +band of muscle fibers of this sort strengthens the knee joint and +gives still more strength to the _pars interna_. This condition has +been reported in a number of birds by Hudson (1937) and is, in all +probability, an adaptation for greater strength of certain leg +movements. The development of this band in _Leucosticte_ seems to +parallel that in the other birds studied and does not indicate +relationship, since in _Leucosticte_ this band arises from the +undivided muscle which (as stated above) resembles only the posterior +portion of the bipartite muscle described for the other birds. In the +latter, the muscular band arises from the anterior part of the muscle. + +Minor differences in muscle pattern, like those already mentioned, are +consistent also between subfamilies, but correlation of these minor +differences with function is difficult. There is the implication, +however, that in all the groups except the carduelines and ploceids, +the emphasis is on greater strength and mobility of the leg. In the +carduelines that were studied the origin of the _m. sartorius_ does +not extend so far craniad as in the other species. In the latter, at +least half of the origin is from the last one or two free dorsal +vertebrae; in the carduelines no more than one third of the origin is +anterior to the ilium. It is conceivable that the more craniad the +origin, the stronger the forward movement of the thigh would be. + +In _Passer_, _Estrilda_ and _Poephila_, and in all the cardueline +finches examined, the bellies of the _m. flexor perforans et +perforatus digiti II_ and the _m. flexor perforans et perforatus +digiti III_ are more intimately connected than they are in the other +species studied. Thus, the amount of independent action of these +muscles in _Passer_, in the estrildines, and in the carduelines +probably is reduced. + +In _Passer_, the estrildines, and the carduelines the edges of the +sheathlike tendon of insertion of the _m. perforatus digiti III_ are +thickened; as a result the insertion appears superficially to be +double but closer examination reveals that there is a fascia stretched +between the thickened edges. In the other species examined, the +insertion is sheathlike throughout and there are no thick areas. I +cannot explain this on the basis of function. The difference, however, +is obvious and constant. + +Aside from the differences noted above, there were variations of +muscle pattern that seem to be significant only in _Vireo olivaceus_. +In this species the central, aponeurotic portion of the _m. +iliotibialis_ is absent. The origin of the _m. adductor longus et +brevis_ is from the dorsal edge of the ischiopubic fenestra and not +from the membrane covering this fenestra. The origin of the _pars +posticus_ of this muscle, furthermore, is fleshy and not tendinous as +it is in the other species. The _m. flexor perforatus digiti II_ is +larger and more deeply situated in _Vireo_ and has, furthermore, no +connection with the _m. flexor hallucis longus_. The latter muscle is +smaller and weaker than in any of the other species and has only one +(the posterior) head of origin. The _m. flexor hallucis brevis_, on +the contrary, is larger than in the other birds, compensating, +probably, for the small _m. flexor hallucis longus_. In those +differences, however, which separate the carduelines and ploceids from +the other birds studied, _Vireo_ resembles, in every instance, the +richmondenines, emberizines, tanagers, warblers, and blackbirds. + +On the basis of differences in leg-musculature the species which are +now included in the Family Fringillidae may be separated into two +groups. One group includes the richmondenines and the emberizines; the +other, the carduelines. The muscle patterns of the legs of the birds +of the first group are indistinguishable from those of _Seiurus_, +_Icterus_, _Molothrus_, and _Piranga_, and except for the differences +noted are similar to those in _Vireo_. The carduelines, on the other +hand, are similar in every point of leg-musculature to the ploceids +which were studied. Thus, the heterogeneity of the Family +Fringillidae, as now recognized, is emphasized by differences in the +muscle patterns of the leg. + + + + +COMPARATIVE SEROLOGY + + +General Statement + +The application of serological techniques to the problems of animal +relationships has been attempted with varying degrees of success over +a period of approximately fifty years. Few of the earlier studies were +of a quantitative nature, but within the past decade, satisfactory +quantitative serological techniques have been developed whereby +taxonomic relationships may be estimated. The usefulness of +comparative serology in taxonomy has been demonstrated in +investigations of many groups wherein results obtained have, in most +instances, been compatible with the results obtained by more +conventional methods, such as comparative morphology. As Boyden +(1942:141) stated, "comparative serology ... is no simple guide to +animal relationship." However, the objectiveness of its methods, the +fact that it has its basis in the comparisons of biochemical systems +which seem to be relatively slow to change in response to external +environmental influences, and the fact that the results are of +quantitative nature favor, where possible, the inclusion of data from +comparative serology along with that from more conventional sources +when an attempt is made to determine the relationships of groups of +animals. + +The application of serological methods in ornithology has not been +extensive. Irwin and Cole (1936) and Cumley and Irwin (1941, 1944) +used two species of doves and their hybrids and demonstrated that a +distinction between the red cells of these birds could be made by use +of immunological methods involving the agglutinin reaction. McGibbon +(1945) was able to distinguish the red cells of interspecific hybrids +in ducks by similar methods. Irwin (1953) used similar techniques in +his study of the evolutionary patterns of some antigenic substances of +the blood cells of birds of the Family Columbidae. Sasaki (1928) +demonstrated the usefulness of the precipitin technique in +distinguishing species of ducks and their hybrids. This technique +was used successfully also by DeFalco (1942) and by Martin and +Leone (1952). Working with groups of known relationships, these +investigators showed that the "accepted" systematic positions of +certain birds were confirmed by serological procedures. The precipitin +reaction, however, has never been applied to actual problems in avian +taxonomy prior to the present study. + + +Preparation of Antigens + +Although most previous work in comparative serology in which +precipitin tests were used has involved the use of whole sera as +antigens, Martin and Leone (1952) indicated that tissue extracts are +satisfactory as antigens and that serological differentiation can be +obtained with these extracts and the antisera to them. I decided, +therefore, to use such extracts in these investigations, since the +small sizes of the birds to be tested made it impracticable to obtain +enough whole sera. + +Most of the birds used were obtained by shooting, but a few were +trapped and the exotic species were purchased alive from a pet dealer. +When a bird was killed, the entire digestive tract was carefully +removed to prevent the escape of digestive enzymes into the tissues +and to prevent putrefaction by action of intestinal bacteria. As soon +as possible (and within three hours in every instance) the bird was +skinned, the head, wings, and legs were removed, and the body was +frozen. Each specimen, consisting of trunk, heart, lungs, and kidneys, +was wrapped separately and carefully in aluminum foil to prevent +dehydration of the tissues. The specimens were kept frozen until the +time when the extracts were made. + +When an extract was to be prepared, the specimen was allowed to thaw +but not to become warm. In the cold room with the temperature of all +equipment and reagents at 2°C., the specimen was placed in a Waring +blender with 0.9 per cent aqueous solution of NaCl buffered with M/150 +K_{2}HPO_{4} and M/150 Na_{2}HPO_{4} to a pH of 7.0. The amount of +reagent used was 75 ml. of saline for each gram of tissue to be +extracted. The tissues were minced in the blender, allowed to stand at +2°C. for 72 hours, and the tissue residues removed by centrifugation +in a refrigerated centrifuge. Formalin was added to a portion of the +supernatant in the amount necessary to make the final dilution 0.4 per +cent. This formolization was found to be necessary to inhibit the +action of autolytic enzymes over the period of time required to +complete the investigations. The effects of formolization on the +antigenicity and reactivity of proteins are discussed later. It was +necessary to sterilize and clarify the "native" (unformolized) +extracts; this was done by filtration through a Seitz filter. These +"native" substances were used only in the early stages of the +investigation (see below). The filtrate was bottled and stored at 2°C. +In the early stages of this investigation clarification of the +formolized extract was accomplished by the same sort of filtration. It +was determined, however, that centrifugation in a refrigerated +centrifuge at high speeds (17,000g) served the same purpose and was +quicker. The formolized extracts were bottled and also stored at 2°C. +(although refrigerated storage of the formolized extracts does not +seem necessary). For each extract the amount of protein present was +determined colorimetrically by the method of Greenberg (1929) with a +Leitz Photrometer. + +Species for which extracts were prepared and the protein values of the +extracts are listed in Table 1. Extracts of some species were used +throughout most of the experiment; extracts of others were used only +when needed for purposes of comparison. + + TABLE 1.--Species from Which Extracts Were Prepared and Injection + Schedules for Extracts Against Which Antisera Were Produced + + ==========================+==========+================================= + | Protein, | + SPECIES | gms. per | Injection schedules for + | 100 ml. | production of antisera + --------------------------+----------+--------------------------------- + _Myiarchus crinitus_ | 0.65 | Series 1: Intravenous, 0.5, 1.0, + (Linnaeus) | | 2.0, and 4.0 ml. + --------------------------+----------+--------------------------------- + _Passer domesticus_ | 1.40 | Series 1: Subcutaneous, 0.5, + | | 1.0, 2.0, and 4.0 ml. + --------------------------+----------+--------------------------------- + _Estrilda amandava_ | 0.45 | [A]Series 1: Intravenous, 0.5, + | | 1.0, 2.0, and 4.0 ml. + | | + | | [A]Series 2: Subcutaneous, 0.5, + | | 1.0, and 2.0 ml. + | | + | | Intraperitoneal, 8.0 ml. + --------------------------+----------+--------------------------------- + _Poephila guttata_ | 0.56 | [A]Same as for _Estrilda_. + --------------------------+----------+--------------------------------- + _Molothrus ater_ | 0.65 | Series 1: Intravenous and + | | subcutaneous, respectively, 0.5 + | | and 0.5 ml., 1.0 and 1.0 ml., + | | 3.0 and 1.0 ml., 5.0 and 3.0 ml. + | | + | | Series 2: Subcutaneous, 0.5, + | | 1.0, 2.0 and 4.0 ml. + --------------------------+----------+--------------------------------- + _Piranga rubra_ | 0.50 | Same as for _Molothrus_. + --------------------------+----------+--------------------------------- + _Richmondena cardinalis_ | 0.70 | [A]Same as for _Estrilda_. + --------------------------+----------+--------------------------------- + _Richmondena cardinalis_ | 0.60 | Same as for _Spinus_. + --------------------------+----------+--------------------------------- + _Passerina cyanea_ | 0.45 | Antiserum not prepared. + --------------------------+----------+--------------------------------- + _Spiza americana_ | 0.70 | Same as for _Molothrus_. + --------------------------+----------+--------------------------------- + _Carpodacus purpureus_ | 0.50 | Antiserum not prepared. + --------------------------+----------+--------------------------------- + _Spinus tristis_ | 0.49 | Series 1: Intravenous, 0.5, 1.0, + | | 2.0, and 4.0 ml. + | | + | | Series 2: Intravenous, 0.5, 1.0, + | | 2.0, and 4.0 ml. + | | + | | Series 3: Subcutaneous, 0.5, + | | 1.0, 2.0, and 4.0 ml. + --------------------------+----------+--------------------------------- + _Pipilo erythrophthalmus_ | 0.92 | Antiserum not prepared. + --------------------------+----------+--------------------------------- + _Junco hyemalis_ | 0.56 | Same as for _Spinus_. + --------------------------+----------+--------------------------------- + _Spizella arborea_ | 0.48 | Same as for _Spinus_. + --------------------------+----------+--------------------------------- + _Zonotrichia querula_ | 0.48 | Same as for _Spinus_. + --------------------------+----------+--------------------------------- + _Zonotrichia albicollis_ | 0.92 | Antiserum not prepared. + (Gmelin) | | + --------------------------+----------+--------------------------------- + + [A] Antiserum prepared against formolized antigen. + + +Preparation of Antisera + +All antisera were produced in rabbits (laboratory stock of +_Oryctolagus cuniculus_). Three methods of injection of antigen were +used in various combinations: intravenous, subcutaneous, and +intraperitoneal. Injection schedules used in the production of each +antiserum are listed in Table 1. Both formolized and "native" antigens +were used. Each rabbit received one or more series of four injections, +each injection being administered on alternate days and doubling in +amount: 0.5 ml., 1.0 ml., 2.0 ml., and 4.0 ml. In all but two +instances more than one series of injections was necessary to produce +a useful antiserum. More than two series, however, resulted in little +or no improvement of the reactivity of the antiserum. + +The injection-series were separated by intervals of eight days. On the +eighth day after the last injection of each series, 10 ml. of blood +were withdrawn from the main artery of the ear of the rabbit, and the +antiserum was used in a homologous precipitin test to determine its +usefulness. If the antiserum contained sufficient amounts of +antibodies to conduct the projected tests, the rabbit was completely +exsanguinated by cardiac puncture, by using an 18-gauge needle and a +50 ml. syringe. The whole blood was placed in clean test tubes and +allowed to clot. It was allowed to stand at 2°C. for 12 to 18 hours so +that most of the serum would be expressed from the clot. The serum was +then decanted, centrifuged to remove all blood cells, sterilized in a +Seitz filter, bottled in sterile vials, and stored at 2°C. until used. + + +Methods of Serological Testing + +The precipitin reaction is the most successful of the serological +techniques thus far devised for systematic comparisons. The reaction +occurs because antigenic substances introduced into the body of an +animal cause the formation of antibodies which precipitate antigens +when the two are mixed. The antisera which are produced show +quantitative specificities in their actions; therefore, when an +antiserum containing precipitins is mixed with each of several +antigens, the reaction involving the homologous antigen (that used in +the production of the antiserum) is greater than those reactions +involving the heterologous antigens (antigens other than those used in +the production of the antiserum). Furthermore, the magnitudes of the +reactions between the antiserum and the heterologous antigens vary +according to the degrees of similarity of these antigens to the +homologous one. + +The method of precipitin testing follows that outlined by Leone +(1949). The Libby (1938) Photronreflectometer was used to measure the +turbidities developed by the interaction of antigen and antiserum. +With this instrument parallel rays of light are passed through the +turbid systems being measured. Light rays are reflected from the +suspended particles to the sensitive plate of a photoelectric cell; +this generates a current of electricity which causes a deflection on a +galvanometer. The deflection is proportional to the amount of +turbidity developed and readings may be taken directly from the scale +of the instrument. + +The reaction-cells of the photronreflectometer are designed to operate +with a volume of 2 ml.; therefore, this volume was used in all +testing. In every series of tests the amount of antiserum was held +constant and the amount of antigen was varied. The volume for each +antigen dilution was always 1.7 ml., and to this was added 0.3 ml. of +antiserum to make up a volume of 2 ml. + + TABLE 2.--Percentage values obtained from analyses of precipitin + reactions. Numerals represent relative amounts of reaction between + antigens and antisera. Homologous reactions are arbitrarily valued + as 100 per cent, and heterologous reactions are expressed + accordingly. _Comparisons are meaningful only if made within each + horizontal row of values._ + + Table headings: + Col A: _Estrilda amandava_ + Col B: _Poephila guttata_ + Col C: _Piranga rubra_ + Col D: _Richmondena cardinalis_ + Col E: _Spiza americana_ + Col F: _Spinus tristis_ + Col G: _Junco hyemalis_ + Col H: _Zonotrichia querula_ + + ========================+============================================== + | ANTISERA + ANTIGENS +-----+-----+-----+-----+-----+-----+-----+---- + | A | B | C | D | E | F | G | H + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Passer domesticus_ | 75 | 74 | 73 | 66 | 81 | 72 | ... | 81 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Estrilda amandava_ | 100 | 88 | 75 | ... | 79 | 72 | 53 | ... + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Poephila guttata_ | 95 | 100 | 77 | 67 | 87 | 81 | ... | ... + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Molothrus ater_ | 66 | 54 | 69 | 65 | 86 | 75 | 69 | 75 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Piranga rubra_ | ... | ... | 100 | ... | ... | ... | ... | 89 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Richmondena cardinalis_| 75 | 80 | 91 | 100 | 98 | 65 | 88 | 91 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Spiza americana_ | 65 | 68 | ... | 71 | 100 | 64 | 67 | 80 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Carpodacus purpureus_ | 70 | 71 | 71 | 61 | 89 | 93 | 53 | 70 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Spinus tristis_ | 72 | 74 | 73 | 60 | 89 | 100 | 60 | ... + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Junco hyemalis_ | 64 | 56 | 74 | 65 | 87 | 68 | 100 | ... + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Zonotrichia querula_ | 65 | 71 | ... | 67 | 89 | 75 | ... | 100 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + +Antigens were diluted with 0.9 per cent phosphate-buffered saline +solution. Tests were run in standard Kolmer test-tube racks, each test +consisting of 12 tubes. Each dilution was made on the basis of the +known protein concentration of the antigen. The first tube contained +an initial dilution of 1 part protein in 250 parts saline and each +successive tube contained a protein dilution one-half the +concentration of the preceding tube, ranging up to 1:512,000. Saline +controls, antiserum controls, and antigen controls were maintained +with each test to determine the turbidities inherent in these +solutions. These control-turbidities were deducted from the total +turbidity developed in each reaction-tube, the resultant turbidity +then being considered as that which was caused by the interaction of +antigens and antibodies. The turbidities were allowed to develop over +a 24-hour period. In the early stages of this investigation the +reactions were allowed to take place at 2°C. in order to inhibit +bacterial growth. + +Later tests were carried out at room temperatures, and bacterial +growth was prevented by the addition to each tube of 'Merthiolate' in +a final dilution of 1:10,000. + + +Experimental Data + +Corrected values for the turbidities obtained were plotted with the +turbidity values on the ordinate and the antigen dilutions on the +abscissa. The homologous reaction was the standard of reference for +all other test reactions with the same antiserum. By summing the +plotted turbidity readings, numerical values are obtained which are +indices serving to characterize the curves. Such values were converted +to percentage values, that of the homologous reaction being considered +100 per cent. These values, plus the curves, provide the data by means +of which the proteins of the birds may be compared. Plots +representative of the precipitin curves are presented in Figs. 10 to +21. For convenience each plot represents only several of the 10 curves +obtained with each antiserum. + +A summary of the serological relationships of the birds involved in +the precipitin tests is presented in Table 2, in which percentage +values are presented. Since the techniques involved in testing were +greatly improved as the investigation proceeded, the summary is based +solely on those tests run in the later stages of the investigation. +For reasons which will become apparent in later discussion, it should +be emphasized that in Table 2 comparisons may be made only within each +horizontal row of values. + + +Discussion of the Serological Investigations + +One of the problems met early in this investigation was instability of +the proteins in the extracts that were prepared. Extracts in which no +attempt was made to inactivate the enzymes present proved +unsatisfactory. It was necessary to maintain the temperature of the +"native" antigens at 2°C, and all work with such antigens had to be +performed at this temperature. This arrangement was inconvenient; +furthermore, inactivation of the enzymes was not complete even at this +low temperature, and some denaturation of the proteins took place as +evidenced by the gradual appearance of insoluble precipitates in the +stored vials. + +The preservatives, 'Merthiolate' and formalin, were used in an attempt +to inhibit the autolytic action of the enzymes present. Formalin, when +added to make a final dilution of 0.4 per cent, proved to be the more +satisfactory of the two preservatives and was used throughout most of +the work. Formalin caused slight denaturation of some of the proteins, +but this effect was complete within a few hours, after which any +denatured material was removed by filtration or centrifugation. The +proteins remaining in solution were stable over the period necessary +to complete the investigations. + +The addition of formalin reduces the reactivity of the extracts when +they are tested with antisera prepared against "native" antigens and +causes changes in the nature of the precipitin curves. This effect has +been pointed out by Horsfall (1934) and by Leone (1953) in their work +on the effects of formaldehyde on serum proteins. Their data indicate, +however, that even though changes in the immunological characteristics +of proteins are brought about by formolization, the proteins retain +enough of their specific chemical characteristics to allow consistent +differentiation of species by immunological methods. In the tests +which I performed, the relative positions of the precipitin curves, +whether native or formolized extracts were involved, remained +unchanged (Figs. 10, 11). _All data used in interpretation of the +serological relationships were obtained from tests in which formolized +antigens of equivalent age were used._ + +Only three antisera were produced against formolized antigens, all +others being produced against "native" extracts. The formolized +antigens seemed to have a greater antigenicity, in most instances, +than did those which were unformolized, and precipitin reactions +involving antisera produced against formolized antigens developed +higher turbidities. The antisera produced against formolized antigens +were equal to but no better than those prepared against "native" +extracts in separating the birds tested (Figs. 12, 13). + +The rabbit is a variable to be considered in serological tests. Two +rabbits exposed to the same antigen, under the same conditions, may +produce antisera which differ greatly in their capacities to +distinguish different antigens. It is logical to assume, therefore, +that two rabbits exposed to different antigens may produce antisera +which also differ in this respect. This explains the unequal values of +reciprocal tests shown in Table 2. Thus, in the test involving the +antiserum to the extracts of _Richmondena_, a value of 71 per cent was +obtained for _Spiza_ antigen, whereas in the test involving +anti-_Spiza_ serum, a value of 98 per cent was obtained for +_Richmondena_ antigen. In Table 2, therefore, comparisons may be made +only among values for the proteins of birds tested with the same +antiserum. + +Since the amount of any one antiserum is limited, there is, of +necessity, a limit as to the number of birds used in a series of +serological tests. Therefore, although the results reveal the actual +serological relationships of the individual species, interpretation of +the relationships of the taxonomic groups must be undertaken with the +realization that such an interpretation is based on tests involving +relatively few species of each group. It is reasonable to assume, +however, that a species which has been placed in a group on the basis +of resemblances other than serological resemblance would show greater +serological correspondence to other members of that group than it +would to members of other groups. Specifically, in the Fringillidae +and their allies, there seems to be little reason to doubt that +genera, and even subfamilies, are natural groups. This is illustrated +in tests involving closely related genera: _Richmondena_ and _Spiza_ +(Figs. 14, 15, 18), _Estrilda_ and _Poephila_ (Fig. 21), _Spinus_ and +_Carpodacus_ (Figs. 12, 17, 19, 20). In each of these tests the pairs +of genera mentioned show greater serological correspondence to each +other than they do to other kinds involved. This point is illustrated +further by a test (not illustrated) involving _Zonotrichia querula_ +(the homologous antigen) and _Zonotrichia albicollis_. Although this +test was one of an earlier series in which difficulties were +encountered (the data, therefore, were not used), it is of interest +that the two species were almost indistinguishable serologically. + +The serological homogeneity of passeriform birds is emphasized by the +fact that the value of every heterologous reaction was more than 50 +per cent of the value of the homologous reaction, except in the test +involving the anti-_Richmondena_ serum and _Myiarchus_ (Fig. 13) in +which the value of the heterologous reaction was 45 per cent. Because +most ornithologists consider these genera to be only distantly related +(they are in different suborders within the Order Passeriformes), the +relatively high value of the heterologous reaction emphasizes the +close serological correspondence of passerine birds and indicates that +small consistent serological differences among these birds are +actually significant. The possibility that some of the serological +correspondence is due to the "homologizing" effect of formalin on +proteins should not be excluded. I think, however, that this effect is +not entirely responsible for the close correspondence observed here. + +An additional point to consider in interpretation of the serological +tests is that the techniques used tend to separate sharply species +that are closely related whereas species that are distantly related +are not so easily separated. In other words, comparative serological +studies with the photronreflectometer tend to minimize the differences +between distant relatives and to exaggerate the differences between +close relatives. + +In analyzing the serological relationships of the species used in this +study, it becomes obvious that two or more series of tests must be +considered before the birds can be placed in relation to each other. +For example, the data presented in Fig. 14 indicate that _Spiza_ and +_Molothrus_ show approximately the same degree of serological +correspondence to _Richmondena_. This does not imply necessarily that +_Spiza_ and _Molothrus_ are closely related. If Fig. 15 is examined, +it can be determined that _Richmondena_ shows much greater serological +correspondence to _Spiza_ than does _Molothrus_. Thus, an analysis of +both figures serves to clarify the true serological relationships of +the three genera. By reference to other series of tests involving +these three birds a more exact determination of their relationships +may be obtained. + +To illustrate this point by a hypothetical example, two species might +seem equidistant, serologically, from a third species. Additional +testing should indicate if the first two species are equidistant in +the same direction (therefore, by implication, close relatives) or in +opposite directions (therefore, distant relatives). A single test +supplies only two dimensions of a three dimensional arrangement. + +It is impossible to interpret and to picture the serological data +satisfactorily in two dimensions; therefore, a three-dimensional model +(Figs. 22, 23) was constructed to summarize the serological +relationships of the birds involved. Each of the eleven kinds used +consistently throughout the investigation is represented in the model. +By use of the percentage values (Table 2), each bird was located in +relation to the other birds. Where possible, averages of reciprocal +tests (Table 3) were used in determining distances between the +elements of the model. In this way seven of the birds were accurately +located in relation to each other. Lacking reciprocal tests, the +positions of the other birds were determined by the values of single +tests (Table 4). Although these birds were placed with less certainty, +at least four points of reference were used in locating each species. +At least one serological test is represented by each connecting bar in +the model. The lengths of the bars connecting any two elements were +determined as follows: a percentage value (Table 3 and Table 4) +representing the degree of serological correspondence between two +birds was subtracted from 100 per cent; the remainder was multiplied +by a factor of five to increase the size of the model and the product +was expressed in millimeters; a bar of proper length connects the two +elements involved. + +From the model it is observed that, _Molothrus_ and _Passer_ excluded, +the birds fall into two distinct groups: one includes _Piranga_, +_Richmondena_, _Spiza_, _Junco_, and _Zonotrichia_; the other includes +_Estrilda_, _Poephila_, _Carpodacus_, and _Spinus_. + + TABLE 3.--Reciprocal Values Used to Determine Distances Between + Elements of the Model; Each Value Represents the Average of + Serological Tests Between the Species Involved + + Table Headings: + Col A: _Estrilda amandava_ + Col B: _Poephila guttata_ + Col C: _Richmondena cardinalis_ + Col D: _Spiza americana_ + Col E: _Spinus tristis_ + Col F: _Junco hyemalis_ + Col G: _Zonotrichia querula_ + + ==========================+====+====+====+====+====+====+====+ + | A | B | C | D | E | F | G | + --------------------------+----+----+----+----+----+----+----+ + _Estrilda amandava_ | .. | 92 | .. | 72 | 72 | 59 | .. | + --------------------------+----+----+----+----+----+----+----+ + _Poephila guttata_ | 92 | .. | 74 | 78 | 78 | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + _Richmondena cardinalis_ | .. | 74 | .. | 85 | 63 | 77 | 79 | + --------------------------+----+----+----+----+----+----+----+ + _Spiza americana_ | 72 | 78 | 85 | .. | 77 | 77 | 85 | + --------------------------+----+----+----+----+----+----+----+ + _Spinus tristis_ | 72 | 78 | 63 | 77 | .. | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + _Junco hyemalis_ | .. | .. | 77 | 77 | .. | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + _Zonotrichia querula_ | .. | .. | 79 | 85 | .. | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + + TABLE 4.--Single Values Used to Determine Distances Between Elements + of the Model; Each Value Represents a Single Test Between the + Species Involved + + Table headings: + Col A: _Estrilda amandava_ + Col B: _Poephila guttata_ + Col C: _Piranga rubra_ + Col D: _Richmondena cardinalis_ + Col E: _Spinus tristis_ + Col F: _Junco hyemalis_ + Col G: _Zonotrichia querula_ + + ==========================+====+====+====+====+====+====+====+ + | A | B | C | D | E | F | G | + --------------------------+----+----+----+----+----+----+----+ + _Passer domesticus_ | .. | 74 | 73 | .. | 72 | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + _Molothrus ater_ | .. | 54 | .. | 65 | .. | 69 | 75 | + --------------------------+----+----+----+----+----+----+----+ + _Piranga rubra_ | .. | 77 | .. | 91 | 73 | 74 | .. | + --------------------------+----+----+----+----+----+----+----+ + _Carpodacus purpureus_ | 70 | 71 | .. | 61 | 93 | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + + [Illustration: FIGS. 10-13. Graphs of precipitin reactions + illustrating effects of formalin on antigenicity and reactivity + of the extracts. For further information, see text, pp. 190-193. + + FIG. 10. Reactions of unformolized antigens of _Richmondena_, + _Zonotrichia_, and _Molothrus_ with anti-_Richmondena_ serum. + FIG. 11. Reactions of formolized antigens of _Richmondena_, + _Zonotrichia_, and _Molothrus_ with anti-_Richmondena_ serum. + FIG. 12. Reactions of anti-_Richmondena_ serum prepared against + native antigen with antigens of _Richmondena_, _Zonotrichia_, + _Carpodacus_, and _Spinus_. + FIG. 13. Reactions of anti-_Richmondena_ serum prepared against + formolized antigen with antigens of _Richmondena_, _Zonotrichia_, + _Poephila_, _Spinus_, and _Myiarchus_.] + + [Illustration: FIGS. 14-17. Graphs of precipitin reactions + illustrating serological relationships. For further explanation, + see text, pp. 190-193. + + FIG. 14. Serological relationships of _Richmondena_, _Spiza_, and + _Molothrus_. + FIG. 15. Serological relationships of _Richmondena_, _Spiza_, and + _Molothrus_. + FIG. 16. Serological relationships of _Carpodacus_ with the + richmondenine-emberizine-thraupid assemblage. + FIG. 17. Serological relationships of _Carpodacus_ and _Spinus_ with + _Richmondena_ and _Junco_.] + + [Illustration: FIGS. 18-21. Graphs of precipitin reactions + illustrating serological relationships. For further explanation, + see text, pp. 190-193. + + FIG. 18. Serological relationships of _Spinus_ and _Poephila_ with + the richmondenines. + FIG. 19. Serological relationships of _Carpodacus_ and _Spinus_ + with _Richmondena_ and _Piranga_. + FIG. 20. Serological relationships of _Poephila_ and Richmondena + with the carduelines. + FIG. 21. Serological relationships of _Richmondena_ and _Spinus_ + with the estrildines.] + + [Illustration: FIG. 22. Two views of a model illustrating + serological relationships among fringillid and related birds. + For further explanation, see text, pp. 193-194. + + Genera Pi . . . . _Piranga_ + C . . . . _Carpodacus_ Po . . . . _Poephila_ + E . . . . _Estrilda_ R . . . . _Richmondena_ + J . . . . _Junco_ Sn . . . . _Spinus_ + M . . . . _Molothrus_ Sz . . . . _Spiza_ + Pa . . . . _Passer_ Z . . . . _Zonotrichia_] + + [Illustration: FIG. 23. Two additional views of the model shown in + fig. 22 illustrating serological relationships among fringillid + and related birds. For further explanation, see text, + pp. 193-194. + + Genera Pi . . . . _Piranga_ + C . . . . _Carpodacus_ Po . . . . _Poephila_ + E . . . . _Estrilda_ R . . . . _Richmondena_ + J . . . . _Junco_ Sn . . . . _Spinus_ + M . . . . _Molothrus_ Sz . . . . _Spiza_ + Pa . . . . _Passer_ Z . . . . _Zonotrichia_] + +Within the richmondenine-emberizine-thraupid assemblage, _Junco_ +and _Zonotrichia_ constitute a sub-group apart from the others. +_Piranga_ and _Richmondena_ show close serological correspondence. +The present taxonomic position of _Spiza_ in the Richmondeninae, +which has been questioned by Beecher (1951a:431; 1953:309), is +corroborated at least insofar as the serological evidence is +concerned. Certainly, serological correspondence of _Spiza_ with the +richmondenine-emberizine-thraupid assemblage is greater than with any +other group of birds tested. + +It is obvious that the serological affinities of the carduelines do +not lie with the richmondenines, emberizines, or thraupids. The +carduelines show greater serological correspondence with the +estrildines than they do with any of the other groups tested. Further +serological investigation involving other species, however, is +necessary before the nearest relatives of the carduelines can be +determined with certainty. + +The two estrildines tested (_Estrilda_ and _Poephila_) show close +serological relationship. Their nearest relatives, serologically, seem +to be the carduelines. The classification (Wetmore, 1951) that places +_Passer_ in the same family with the estrildines is not upheld by the +serological data available. _Passer_ is not, serologically, closely +related to any of the birds tested. It is of interest that Beecher +(1953:303-305), on the basis of jaw musculature, places _Passer_ and +the estrildines in separate families (Ploceidae and Estrildidae, +respectively). + +_Molothrus_ shows greater serological correspondence to the +richmondenine-emberizine-thraupid assemblage than to any of the other +birds tested. It is definitely set apart from this group, however, and +its position, serologically, is compatible with that based on evidence +from other sources. + +There seems to be but little argument among ornithologists that +icterids, fringillids, and ploceids constitute families which are +distinct from one another. If, then, the serological differences +between _Molothrus_ (Icteridae) and _Richmondena_ (Fringillidae), +between _Molothrus_ and _Zonotrichia_ (Fringillidae), and between +_Richmondena_ and _Poephila_ (Ploceidae) are indicative of family +differences, there are four families represented by the birds +involved. _Molothrus_ represents one family; _Piranga_, _Richmondena_, +_Spiza_, _Junco_, and _Zonotrichia_, a second; _Estrilda_, _Poephila_, +_Carpodacus_, and _Spinus_, a third; and _Passer_, a fourth. + + + + +CONCLUSIONS + + +The heterogeneity of the Family Fringillidae has been emphasized by +many authors. The relationships of the species now included in this +Family have been the subject of much discussion and constitute an +important problem in avian systematics. + +Sushkin's studies (1924, 1925) of features of the horny and bony +palates have served as a basis for the present division of the Family +into subfamilies. Recently, Beecher (1951a, 1951b, 1953) and Tordoff +(1954) have used these features and others which they thought to be of +value in an attempt to clarify the relationships of the species +involved. + +Beecher's work (1951a, 1951b, 1953) on jaw-musculature is a valuable +contribution to our knowledge of the anatomy of passerine birds. His +myological studies were so thorough and his presentation so detailed +that students who disagree with his interpretations can draw their own +conclusions. Beecher (1951b:276) points out that there are two basic +types of skeletal muscle--those with parallel fibers and those with +pinnately arranged fibers. The muscles with pinnate fibers seem to be +more efficient, each muscle having a greater functional cross section +for its bulk than does one with parallel fibers. He assumes that +muscles with parallel fibers are more primitive, phylogenetically, +than are those with fibers arranged pinnately. Since his study of the +jaw muscles of the Icteridae (1951a) revealed that patterns of +jaw-musculature within this Family remain constant regardless of the +methods used in procuring food, he assumes that such patterns may be +used as indicators of relationship throughout the entire oscinine +group. These two assumptions, then, serve as the basis for his +hypothesis concerning relationship and phylogeny within this +assemblage. Beecher (1951b:278-280; 1953:310-312) maintains that +within the Family Thraupidae there are two main lines which lead with +almost no disjunction to the Carduelinae and Richmondeninae. The +thraupid-richmondenine line involves a shift in the nature of the _m. +adductor mandibulae externus superficialis_, which becomes more +pinnate in the richmondenines. This results in greater crushing power. +The thraupid-cardueline line involves a shift in emphasis from the the +_m. adductor mandibulae externus medialis_ to the _m. pseudotemporalis +superficialis_ and the forward advance of the insertion of the latter. +This, also, promotes greater crushing ability. He states that features +of the horny palate and of the plumage provide further evidence of +close relationship of these groups. He includes, therefore, the +Thraupinae, the Carduelinae, and the Pyrrhuloxiinae (=Richmondeninae) +in the Family Thraupidae. Beecher (1953:307) indicates that the +patterns of jaw-musculature of the Parulinae (wood warblers) and +Emberizinae (buntings) are similar and suggests that the buntings had +their origin from the wood warblers. He includes these subfamilies, +therefore, in the Family Parulidae. + +Beecher's reasoning may be criticized on several points. It may be, as +he suggests, that muscles with parallel fibers evolved earlier, +phylogenetically, than did muscles with pinnate fibers, but he does +not give adequate consideration, it seems to me, to the possibility +that parallel fibers may also have evolved secondarily from pinnate +fibers. Since Beecher (1951a) found that patterns of jaw-musculature +within the Family Icteridae were conservative, he is reluctant to +admit the possibility of convergence among any of the other families. +Differences in patterns of jaw-musculature are, however, functional +adaptations and like the bill, which is also associated with +food-getting may be subject to rapid evolutionary change. Finally, in +attempting to classify the oscines, he has relied almost entirely on a +single character--the pattern of jaw-musculature. + +Tordoff's attempts (1954) to clarify the relationships of the +fringillids and related species are based chiefly on features of the +bony palate. He assumes that since palato-maxillaries seem to be +absent in the majority of passerine birds, their occurrence in certain +nine-primaried oscine groups indicates relationship among these +groups. He points out that these bones, when present, are important +areas of origin of the _m. pterygoideus_ which functions in depression +of the upper jaw and in elevation of the lower jaw. He assumes, +therefore, that palato-maxillaries were evolved to provide for a more +effective action of the _m. pterygoideus_. The need for such action +could be associated with a seed-eating habit. All richmondenines and +emberizines possess palato-maxillary bones either free or fused to the +prepalatine bar, but there is no trace of these bones in the +carduelines. Carduelines, furthermore, possess prepalatine bars that +are characteristically flared anteriorly. This condition does not +exist in the richmondenines or in the emberizines. + +Tordoff points out, also, that the irregular, erratic migrations of +the New World Carduelinae are unlike the more regular migrations of +the richmondenines and emberizines. The carduelines, furthermore, are +more arboreal in their habits than are these other groups and exhibit +a decided lack of nest sanitation during the later stages of nesting, +a situation which contrasts with that found in the Richmondeninae and +Emberizinae. He suggests, therefore, that the carduelines are not so +closely related to the richmondenines and the emberizines as +previously has been thought. + +Since there are only two cardueline genera, _Loximitris_ and +_Hesperiphona_, endemic to the New World and at least 10 genera with +many species endemic to the Old World, Tordoff (1954:15) suggests an +Old World origin for the carduelines. He strengthens his argument for +this hypothesis by pointing out that in features of the bony palate +and in habits the carduelines resemble the estrildines of the Family +Ploceidae. + +Tordoff (1954:29-30) states that the tanagers not only merge with the +richmondenines but also grade imperceptibly into the emberizines. He +includes, therefore, the Richmondeninae, Emberizinae, and Thraupinae +in the Family Fringillidae. He suggests that the carduelines are +ploceids, closely related to the Subfamily Estrildinae, on the basis +of structure of the bony palate, geographic distribution, social +behavior, and habits such as nest-fouling and nest-building. + +Tordoff, like Beecher, has based his interpretations chiefly on one +feature--structure of the bony palate. Since this feature also is +associated with food-getting, the possibilities of convergence of +distantly related species with similar habits and divergence of +closely related species with different habits may not be excluded. + +The hazard of unrecognized adaptive convergence cannot, of course, be +excluded from most fields of taxonomic research, but some features of +morphology and biochemistry are notably more conservative than others +and undergo slower evolutionary change. Such features are often of +utmost importance in distinguishing the higher taxonomic categories. + +Most ornithologists are aware that, within the Order Passeriformes, +patterns of musculature in the leg have evolved at a slow rate and +exhibit little variation within the Order. Differences which do occur, +therefore, probably are significant, especially those that are +consistent between groups of species. As I have pointed out earlier +(p. 184), there are no significant differences in leg-musculature +between the Richmondeninae, Emberizinae, and Thraupidae. Indeed, it is +difficult to define these groups on the basis of leg-musculature. If +these groups are of common origin, the lack of distinct boundaries +between them is not surprising. A muscular band which extends from the +_pars interna_ of the _m. gastrocnemius_ around the front of the knee +is present in every emberizine species that I studied and in the Genus +_Piranga_. With the exception of _Spiza_ none of the richmondenines +possesses this band. + +The significant differences in leg-musculature which have been +discussed above (pp. 183-184) distinguish the carduelines from the New +World finches and tanagers. Even the cardueline _Leucosticte_ and the +emberizine _Calcarius_, which resemble one another in general +adaptations and in several myological features of the leg (p. 183), +agree in significant features of the musculature with the respective +groups to which they belong. The carduelines agree in the major +features of leg-musculature with the ploceids which I studied. + +The use of serological techniques in taxonomic work has two main +advantages. The biochemical systems involved in such investigations +seem to be relatively slow to change in response to external +environmental influences, and the quantitative nature of the results +obtained makes possible objective measurement of resemblances among +species. + +I have pointed out (p. 200) that the carduelines are excluded, +serologically, from the distinct assemblage formed by the +richmondenines, emberizines, and tanagers. Actually, the carduelines +show less serological resemblance to this assemblage than do the +estrildines, and most ornithologists agree that the Estrildinae are +not at all closely related to the Richmondeninae, Emberizinae, and +Thraupidae. _Molothrus_, representing a family (Icteridae) recognized +as distinct from the Family Fringillidae, also more closely resembles +the fringillid assemblage, serologically, than do the carduelines. +Although the Carduelinae constitute a distinct group serologically, +they show greater serological resemblance to the estrildines of the +Family Ploceidae than to any of the other species tested. At least the +carduelines and the estrildines form a group as compact as the +subfamilies of the Fringillidae. Thus, the serological data correlate +well with those obtained from the study of the leg-musculature. + +Present systems of classification include the subfamilies Passerinae +and Estrildinae in the Family Ploceidae. _Passer_, however, is less +closely related to the estrildines serologically than are the +carduelines, and is less closely related to the estrildines than +_Molothrus_, an icterid, is to the fringillids. This raises a question +as to the homogeneity of the Family Ploceidae as presently recognized +by most ornithologists. If the Passerinae and the Estrildinae are +placed in a single family, the serological divergence among members of +this group is certainly greater than it is in the Family Fringillidae. +Additionally, Beecher (1953:303-304) found that the estrildines +possess a pattern of jaw-musculature different from those in other +ploceids. + +The combined evidence from jaw-musculature and serology has caused me +to conclude that the estrildines should be excluded from the Family +Ploceidae (see below). + +In an attempt to clarify the relationships of the Fringillidae and +allied groups, I here review briefly the evidence which has been +presented. From his studies of jaw-musculature (1951a, 1951b, +1953) Beecher concludes that the Pyrrhuloxinae (=Richmondeninae), +the Carduelinae, and the Thraupinae are closely related. +He places these groups in the Family Thraupidae. He excludes the +Emberizinae from this group and places them with the wood warblers +in the Family Parulidae. He suggests that the estrildines constitute +a family (Estrildidae) separate from the Family Ploceidae. + +From his studies of certain features of the bony palate Tordoff +(1954:25-26, 32) concludes that the richmondenines, the emberizines, +and the tanagers have a common origin and places these groups in the +Family Fringillidae. He excludes the carduelines from this assemblage, +suggests that they are closely related to the estrildines, and +includes them as the Subfamily Carduelinae in the Family Ploceidae. + +In this paper I have presented data obtained from the study of certain +features of morphology and biochemistry which I think are less subject +to the influence of environmental factors than those features studied +by recent workers. It is significant that the data obtained by use of +serological techniques and those obtained from the study of +leg-musculature point to the same conclusions. On the basis of these +data I have drawn several conclusions concerning the relationships of +the groups which I studied. + +The richmondenines, emberizines, and tanagers are closely related and +should be included in a single family, Fringillidae. The Carduelinae +and the Estrildinae are closely related subfamilies. Although most +recent classifications place the Estrildinae and Passerinae in the +Family Ploceidae, the serological evidence indicates that these groups +are not closely related. Beecher (1953:303-304) drew the same +conclusion from his study of jaw-musculature (see above). I suggest, +therefore, that the Carduelinae and the Estrildinae be placed in a +family separate from the Ploceidae and that the name Carduelidae +(rather than Estrildidae) be used for this group. At present, neither +is an accepted family name. Because _Carduelis_ Brisson 1760 is an +older name than _Estrilda_ Swainson 1827 and because _Carduelis_ seems +to be a centrally located genus in the family, I have chosen the +former (although the International Rules of Zoological Nomenclature do +not specify that priority must apply in forming family names). + +I have been unable to study any of the species included in the +subfamilies Fringillinae (not Fringillinae of Tordoff, see 1954:23-24, +and below) and Geospizinae of recent classifications; thus these +groups have not been discussed above. Beecher (1953:307-308) includes +_Fringilla_ in the Subfamily Carduelinae; he includes the geospizines +in a separate family, Geospizidae, and states that they are derived +from the emberizines. Tordoff (1954:23-24) found that in features of +the bony palate _Fringilla_ and the geospizines resemble the +emberizines and, on this basis, includes them in the Subfamily +Fringillinae. + +The Dickcissel, _Spiza americana_, possesses certain features which +merit special discussion. Beecher (1951a:431; 1953:309), on the basis +of jaw-musculature, considers it an icterid. To be sure _Spiza_ is in +many ways an aberrant member of the group to which it is now assigned +(Subfamily Richmondeninae). _Spiza_, serologically, is closely related +to all species of the richmondenine-emberizine-thraupid assemblage. +Within this assemblage its nearest relatives are the richmondenines. +_Spiza_ differs from the other richmondenines studied and resembles +the emberizines and tanagers in the possession of the muscular band +which extends from the _pars interna_ of the _m. gastrocnemius_ around +the front of the knee. This band, in _Spiza_, is smaller, however, +than in any of the other species. No icterid dissected possesses such +a structure. Tordoff (1954:29) states that _Spiza_ is typically +richmondenine in palatal structure and makes the suggestion, with +which I agree, that _Spiza_ is a richmondenine and may be closely +related to the ancestral stock which gave rise to the fringillid +assemblage. The serological position of _Spiza_, approximately +equidistant from the other fringillids (Figs. 22, 23), and the +presence of the small muscular band around the front of the knee +constitute evidence supporting the central position of _Spiza_. + +After consideration of evidence from the studies of external +morphology, ethology, myology, osteology, and serology, I propose here +an arrangement of the groups which I have studied and submit for +comparison the arrangements (of these groups) proposed by Beecher and +Tordoff. The names of subfamilies that I have been unable to study are +included in my classification and are placed in brackets. + + ------------------------+----------------------+----------------------- + | Proposed by Tordoff | Proposed by Beecher + Here proposed: | (1954) on the basis | (1953) on the basis + | of the bony palate: | of jaw-musculature: + ========================+======================+======================= + FAMILY PLOCEIDAE | FAMILY PLOCEIDAE | FAMILY PLOCEIDAE + | | + [Subf. Bubalornithinae] |Subf. Bubalornithinae | + | | + Subfamily Passerinae: |Subfamily Passerinae | Subfamily Passerinae + distinguished from the | | + Estrildinae by patterns | | + of jaw-musculature | | + (Beecher, 1953:303-304) | | + and on the basis of | | + comparative serology of | | + saline-soluble proteins.| | + | | + [Subfamily Ploceinae] |Subfamily Ploceinae | Subfamily Ploceinae + | | + [Subfamily Viduinae] |Subfamily Viduinae | Subfamily Viduinae + | | + FAMILY CARDUELIDAE | | + | | + Subfamily Estrildinae: |Subfamily Estrildinae | FAMILY ESTRILDIDAE + similar to the | | + Carduelinae in features | | + of the bony palate and | | + habits (Tordoff, 1954: | | + 18-22) and in patterns | | + of leg-musculature and | | + comparative serology | | + of saline-soluble | | + proteins. | | + | | + Subfamily Carduelinae: |Subfamily Carduelinae | [In Thraupidae below] + distinguished from the | | + Fringillidae by features| | + of the palate, | | + geographic distribution,| | + migration patterns, and | | + habits (Tordoff, 1954: | | + 14-18) and by patterns | | + of leg-musculature and | | + comparative serology | | + of saline-soluble | | + proteins. | | + | | + FAMILY FRINGILLIDAE: all| FAMILY FRINGILLIDAE | FAMILY PARULIDAE + members of this family | | Subfamily Parulinae + show similarities in | | Subfamily Emberizinae + features of the bony | | + palate (Tordoff, 1954: | | + 22-23), patterns of | | + leg-musculature, and | | + in comparative serology | | + of saline-soluble | | + proteins. | | FAMILY THRAUPIDAE + | | + Subf. Richmondeninae |Subf. Richmondeninae | Subfamily + | | Pyrrhuloxiinae + | | + Subfamily Thraupinae |Subfamily Thraupinae | Subfamily Thraupinae + | | + Subfamily Emberizinae |Subfamily Fringillinae| [In Parulidae above] + |(including Emberizinae| + [Subfamily Fringillinae]| and Geospizinae) | Subfamily Carduelinae + | | + [Subfamily Geospizinae] | | + ------------------------+----------------------+----------------------- + + + + +SUMMARY + + +It has long been recognized that the Family Fringillidae includes some +dissimilar groups. Specifically, the relationships of the subfamilies +Richmondeninae, Emberizinae, and Carduelinae of the Family +Fringillidae are poorly understood. Data from two recent studies, one +on patterns of jaw-musculature and the other on features of the bony +palate, emphasize the dissimilarity of these subfamilies but have +given rise to conflicting concepts of the relationships of subfamilies +within the Family. + +This paper reports the results of studies involving morphological and +biochemical features that I consider less sensitive to external +environmental factors than are features which have been studied +previously. Patterns of leg-musculature were chosen for study because +earlier work showed that muscle patterns in the legs of passerine +birds are highly stable and vary but little. Variations, therefore, +which are consistent in separating groups of species should be +significant. Serological techniques were used because the biochemical +systems involved seem to be relatively slow to change in response to +environmental influences and because the data obtained may be used in +a highly objective manner to measure resemblance among species. + +Individual differences in the patterns of leg-musculature were found +to be slight and involved mainly the sizes and shapes of muscles. For +this reason variations involving origin, insertion, or relative +position of a muscle, were judged significant. In leg-musculature the +Richmondeninae, the Emberizinae, and the Thraupidae resemble one +another closely. Several differences in muscle pattern were found, +however, which distinguish these groups from the Carduelinae. The +leg-musculature of the carduelines closely resembles that of the +Ploceidae. + +Serological techniques involved the extraction of saline-soluble +proteins from the tissues of the species to be studied. These extracts +were carefully processed and were used as antigens. Formolization of +the antigens was necessary as a means of preventing denaturation of +the proteins by enzymatic activity. Antisera were produced in rabbits. +The method of testing involved turbidimetric analysis of the +precipitin reaction. Utilizing the values for the precipitin tests a +model was constructed which showed the relationships of the eleven +species used in these tests. From a study of the model and the data +used in its construction, it was determined that the Richmondeninae, +Emberizinae, and Thraupidae constitute an assemblage distinct from the +other species studied. The Carduelinae are excluded from the +assemblage and serologically are most closely related to the +Estrildinae. The estrildines, serologically, do not closely resemble +_Passer_, Subfamily Passerinae, although recent classifications place +these two subfamilies in the Family Ploceidae. + +Upon consideration of all evidence now available--from external +morphology, ethology, myology, osteology, and serology--several +hypotheses regarding the relationships of the groups studied are set +forth. The richmondenines, emberizines, and tanagers are closely +related subfamilies and are here included in the Family Fringillidae. +The Estrildinae and Carduelinae are closely related subfamilies, but +neither group is closely related to the Passerinae. The estrildines +and carduelines, therefore, are placed in a separate family, the +Carduelidae. In some ways, _Spiza_ is an aberrant member of the +Subfamily Richmondeninae but should be retained in that subfamily. It +is suggested that _Spiza_ is a primitive richmondenine closely related +to the ancestral fringillid stock. + + + + +LITERATURE CITED + + +AMERICAN ORNITHOLOGISTS' UNION + + 1931. Check-list of North American birds. Fourth edition. + Lancaster, Pa., xix + 526 pp. + + +BEECHER, W. J. + + 1951a. Adaptations for food-getting in the American blackbirds. + Auk, 68:411-440, 11 figs. + + 1951b. Convergence in the Coerebidae. Wilson Bull., 63:274-287, + 5 figs. + + 1953. A phylogeny of the oscines. Auk, 70:270-333, 18 figs. + + +BERGER, A. J. + + 1952. The comparative functional morphology of the pelvic + appendage in three genera of Cuculidae. + Amer. Mid. Nat., 47:513-605, 29 pls. + + +BOYDEN, A. + + 1942. Systematic serology: a critical appreciation. + Physiol. Zool., 15:109-145, 12 figs. + + +CHAPIN, J. P. + + 1917. The classification of the weaver-birds. Bull. Amer. Mus. + Nat. Hist., 37:243-280, 10 pls., 9 figs. + + +CUMLEY, R. W., and IRWIN, M. R. + + 1941. Pictorial representation of the antigenic differences + between two dove species. Jour. Hered., 32:178-182, + frontispiece, 2 figs. + + 1941. Interaction of antigens in dove hybrids. Ibid., 429-434, + 3 figs. + + 1944. The correlation between antigenic composition and geographic + range in the Old and New World of some species of _Columba_. + Amer. Nat., 78:238-256, 1 fig. + + +DEFALCO, R. J. + + 1942. A serological study of some avian relationships. + Biol. Bull., 83:205-218. + + +FISHER, H. I. + + 1946. Adaptations and comparative anatomy of the locomotor + apparatus of New World vultures. Amer. Mid. Nat., + 35:545-727, 13 pls., 28 figs. + + +GADOW, H., and SELENKA, E. + + 1891. Vögel, vol. I, Anatomischer Theil. In Bronn's Klassen und + Ordnungen des Thier-Reichs, Sechster Band, Vierte Abtheilung. + Leipzig, 1008 pp., 59 pls. + + +GARROD, A. H. + + 1873. On certain muscles in the thigh of birds and their value in + classification. Proc. Zool. Soc. London, Part I:626-644, + 6 figs. + + 1874. On certain muscles in the thigh of birds and their value in + classification. Ibid., Part II:111-123. + + +GREENBERG, D. M. + + 1929. The colorimetric determination of serum proteins. + J. Biol. Chem., 82:545-550. + + +HELLMAYR, C. E. + + 1935. Catalogue of birds of the Americas. Field Mus. Nat. Hist., + Zool. ser. 13, pt. 8, vi + 541 pp. + + 1936. Catalogue of birds of the Americas. Ibid., 13, pt. 9, + v + 458 pp. + + 1937. Catalogue of birds of the Americas. Ibid., 13, pt. 10, + v + 228 pp. + + 1938. Catalogue of birds of the Americas. Ibid., 13, pt. 11, + vi + 662 pp. + + +HOWARD, H. + + 1929. The avifauna of the Emeryville shellmound. Univ. California + Publ. Zool., 32:301-394, 3 pls., 54 figs. + + +HUDSON, G. E. + + 1937. Studies on the muscles of the pelvic appendage in birds. + Amer. Mid. Nat., 18:1-108, 26 pls. + + +IRWIN, M. R. + + 1953. Evolutionary patterns of antigenic substances of the blood + corpuscles in Columbidae. Evol., 7:31-50. + + +IRWIN, M. R., and COLE, L. J. + + 1936. Immunogenetic studies of species and of species hybrids in + doves, and the separation of species-specific substances in + the backcross. Jour. Exp. Zool., 73:85-108, 1 fig. + + +LEONE, C. A. + + 1949. Comparative serology of some brachyuran crustacea and + studies in hemocyanin correspondence. Biol. Bull., + 97:273-286, 3 figs. + + 1953. Some effects of formalin on the serological activity of + crustacean and mammalian sera. Jour. Immun., 70:386-392, + 2 figs. + + +LIBBY, R. L. + + 1938. The photronreflectometer--an instrument for the measurement + of turbid systems. Jour. Immun., 34:71-73, 1 fig. + + +MARTIN, E. P., and LEONE, C. A. + + 1952. Serological relationships among domestic fowl as shown by + comparisons of protein preparations from corresponding organ + systems. Trans. Kansas Acad. Sci., 55:439-444, 1 fig. + + +MCGIBBON, W. H. + + 1945. Further division of contrasting antigens in species hybrids + in ducks. Genetics, 30:252-265. + + +SASAKI, K. + + 1928. Serological examination of the blood-relationship between + wild and domestic ducks. Jour. Dept. Agri., Kyushu Imp. + Univ., 2:117-132. + + +SIMPSON, G. G. + + 1944. Tempo and mode in evolution. Columbia Univ. Press, New York, + xviii + 237 pp., 36 figs. + + +SUSHKIN, P. P. + + 1924. [On the Fringillidae and allied groups.] Bull. British + Ornith. Club, 45:36-39. + + 1925. The evening grosbeak (Hesperiphona), the only American genus + of a Palaearctic group. Auk, 42:256-261, 2 figs. + + +TORDOFF, H. B. + + 1954. A systematic study of the avian family Fringillidae, based + on the structure of the skull. Univ. Michigan Mus. Zool. + Misc. Publ. No. 81:1-42, 77 figs. + + +WETMORE, A. + + 1951. A revised classification for the birds of the world. + Smithsonian Misc. Coll., 117(4):1-22. + + +_Transmitted June 8, 1954._ + + +25-4632 + + + + +UNIVERSITY OF KANSAS PUBLICATIONS + +MUSEUM OF NATURAL HISTORY + + +Institutional libraries interested in publications exchange may obtain +this series by addressing the Exchange Librarian, University of Kansas +Library, Lawrence, Kansas. Copies for individuals, persons working in +a particular field of study, may be obtained by addressing instead the +Museum of Natural History, University of Kansas, Lawrence, Kansas. +There is no provision for sale of this series by the University +Library which meets institutional requests, or by the Museum of +Natural History which meets the requests of individuals. However, +when individuals request copies from the Museum, 25 cents should +be included, for each separate number that is 100 pages or more +in length, for the purpose of defraying the costs of wrapping and +mailing. + + * An asterisk designates those numbers of which the Museum's supply + (not the Library's supply) is exhausted. Numbers published to date, + in this series, are as follows: + + Vol. 1. 1. The pocket gophers (Genus Thomomys) of Utah. By Stephen D. + Durrant. Pp. 1-82, 1 figure in text; August 15, 1946. + + 2. The systematic status of Eumeces pluvialis Cope, and + noteworthy records of other amphibians and reptiles from + Kansas and Oklahoma. By Hobart M. Smith. Pp. 85-89. + August 15, 1946. + + 3. The tadpoles of Bufo cognatus Say. By Hobart M. Smith. + Pp. 93-96, 1 figure in text. August 15, 1946. + + 4. Hybridization between two species of garter snakes. + By Hobart M. Smith. Pp. 97-100. August 15, 1946. + + 5. Selected records of reptiles and amphibians from Kansas. + By John Breukelman and Hobart M. Smith. Pp. 101-112. + August 15, 1946. + + 6. Kyphosis and other variations in soft-shelled turtles. + By Hobart M. Smith. Pp. 117-124, 3 figures in text. + July 7, 1947. + + *7. Natural history of the prairie vole (Mammalian Genus + Microtus). By E. W. Jameson, Jr. Pp. 125-151, 4 figures in + text. October 6, 1947. + + 8. The postnatal development of two broods of great horned + owls (Bubo virginianus). By Donald F. Hoffmeister and + Henry W. Setzer. Pp. 157-173, 5 figures in text. + October 6, 1947. + + 9. Additions to the list of the birds of Louisiana. By George + H. Lowery, Jr. Pp. 177-192. November 7, 1947. + + 10. A check-list of the birds of Idaho. By M. Dale Arvey. + Pp. 193-216. November 29, 1947. + + 11. Subspeciation in pocket gophers of Kansas. By Bernardo + Villa R. and E. Raymond Hall. Pp. 217-236, 2 figures in + text. November 29, 1947. + + 12. A new bat (Genus Myotis) from Mexico. By Walter W. + Dalquest and E. Raymond Hall. Pp. 237-244, 6 figures in + text. December 10, 1947. + + 13. Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. + By Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, + 1 figure in text. December 10, 1947. + + 14. A new pocket gopher (Thomomys) and a new spiny pocket + mouse (Liomys) from Michoacán, México. By E. Raymond Hall + and Bernardo Villa R. Pp. 249-256, 6 figures in text. + July 26, 1948. + + 15. A new hylid frog from eastern Mexico. By Edward H. Taylor. + Pp. 257-264, 1 figure in text. August 16, 1948. + + 16. A new extinct emydid turtle from the Lower Pliocene of + Oklahoma. By Edwin C. Galbreath. Pp. 265-280, 1 plate. + August 16, 1948. + + 17. Pliocene and Pleistocene records of fossil turtles from + western Kansas and Oklahoma. By Edwin C. Galbreath. + Pp. 281-284. August 16, 1948. + + 18. A new species of heteromyid rodent from the Middle + Oligocene of northeastern Colorado with remarks on the + skull. By Edwin C. Galbreath. Pp. 285-300, 2 plates. + August 16, 1948. + + 19. Speciation in the Brazilian spiny rats (Genus Proechimys, + Family Echimyidae). By João Moojen. Pp. 301-406, + 140 figures in text. December 10, 1948. + + 20. Three new beavers from Utah. By Stephen D. Durrant and + Harold S. Crane. Pp. 407-417, 7 figures in text. + December 24, 1948. + + 21. Two new meadow mice from Michoacán, Mexico. By E. Raymond + Hall. Pp. 423-427, 6 figures in text. December 24, 1948. + + 22. An annotated check list of the mammals of Michoacán, + Mexico. By E. Raymond Hall and Bernardo Villa R. + Pp. 431-472, 2 plates, 1 figure in text. December 27, 1949. + + 23. Subspeciation in the kangaroo rat, Dipodomys ordii. + By Henry W. Setzer. Pp. 473-573, 27 figures in text, + 7 tables. December 27, 1949. + + 24. Geographic range of the hooded skunk, Mephitis macroura, + with description of a new subspecies from Mexico. + By E. Raymond Hall and Walter W. Dalquest. Pp. 575-580, + 1 figure in text. January 20, 1950. + + 25. Pipistrellus cinnamomeus Miller 1902 referred to the Genus + Myotis. By E. Raymond Hall and Walter W. Dalquest. + Pp. 581-590, 5 figures in text. January 20, 1950. + + 26. A synopsis of the American bats of the Genus Pipistrellus. + By E. Raymond Hall and Walter W. Dalquest. Pp. 591-602, + 1 figure in text. January 20, 1950. + + Index. Pp. 605-638. + + *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. + Pp. 1-444, 140 figures in text. April 9, 1948. + + Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and + distribution. By Rollin H. Baker. Pp. 1-359, 16 figures + in text. June 12, 1951. + + *2. A quantitative study of the nocturnal migration of birds. + By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. + June 29, 1951. + + 3. Phylogeny of the waxwings and allied birds. By M. Dale + Arvey. Pp. 473-530, 49 figures in text, 13 tables. + October 10, 1951. + + 4. Birds from the state of Veracruz, Mexico. By George H. + Lowery, Jr., and Walter W. Dalquest. Pp. 531-649, + 7 figures in text, 2 tables. October 10, 1951. + + Index. Pp. 651-681. + + *Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, + 41 plates, 31 figures in text. December 27, 1951. + + Vol. 5. 1. Preliminary survey of a Paleocene faunule from the Angels + Peak area, New Mexico. By Robert W. Wilson. Pp. 1-11, + 1 figure in text. February 24, 1951. + + 2. Two new moles (Genus Scalopus) from Mexico and Texas. + By Rollin H. Baker. Pp. 17-24. February 28, 1951. + + 3. Two new pocket gophers from Wyoming and Colorado. + By E. Raymond Hall and H. Gordon Montague. Pp. 25-32. + February 28, 1951. + + 4. Mammals obtained by Dr. Curt von Wedel from the barrier + beach of Tamaulipas, Mexico. By E. Raymond Hall. + Pp. 33-47, 1 figure in text. October 1, 1951. + + 5. Comments on the taxonomy and geographic distribution of + some North American rabbits. By E. Raymond Hall and Keith + R. Kelson. Pp. 49-58. October 1, 1951. + + 6. Two new subspecies of Thomomys bottae from New Mexico and + Colorado. By Keith R. Kelson. Pp. 59-71, 1 figure in text. + October 1, 1951. + + 7. A new subspecies of Microtus montanus from Montana and + comments on Microtus canicaudus Miller. By E. Raymond Hall + and Keith R. Kelson. Pp. 73-79. October 1, 1951. + + 8. A new pocket gopher (Genus Thomomys) from eastern Colorado. + By E. Raymond Hall. Pp. 81-85. October 1, 1951. + + 9. Mammals taken along the Alaskan Highway. By Rollin H. + Baker. Pp. 87-117, 1 figure in text. November 28, 1951. + + *10. A synopsis of the North American Lagomorpha. By E. Raymond + Hall. Pp. 119-202, 68 figures in text. December 15, 1951. + + 11. A new pocket mouse (Genus Perognathus) from Kansas. + By E. Lendell Cockrum. Pp. 203-206. December 15, 1951. + + 12. Mammals from Tamaulipas, Mexico. By Rollin H. Baker. + Pp. 207-218. December 15, 1951. + + 13. A new pocket gopher (Genus Thomomys) from Wyoming and + Colorado. By E. Raymond Hall. Pp. 219-222. + December 15, 1951. + + 14. A new name for the Mexican red bat. By E. Raymond Hall. + Pp. 223-226. December 15, 1951. + + 15. Taxonomic notes on Mexican bats of the Genus Rhogeëssa. + By E. Raymond Hall. Pp. 227-232. April 10, 1952. + + 16. Comments on the taxonomy and geographic distribution of + some North American woodrats (Genus Neotoma). By Keith R. + Kelson. Pp. 233-242. April 10, 1952. + + 17. The subspecies of the Mexican red-bellied squirrel, + Sciurus aureogaster. By Keith R. Kelson. Pp. 243-250, + 1 figure in text. April 10, 1952. + + 18. Geographic range of Peromyscus melanophrys, with + description of new subspecies. By Rollin H. Baker. + Pp. 251-258, 1 figure in text. May 10, 1952. + + 19. A new chipmunk (Genus Eutamias) from the Black Hills. + By John A. White. Pp. 259-262. April 10, 1952. + + 20. A new piñon mouse (Peromyscus truei) from Durango, Mexico. + By Robert B. Finley, Jr. Pp. 263-267. May 23, 1952. + + 21. An annotated checklist of Nebraskan bats. By Olin L. Webb + and J. Knox Jones, Jr. Pp. 269-279. May 31, 1952. + + 22. Geographic variation in red-backed mice (Genus + Clethrionomys) of the southern Rocky Mountain region. + By E. Lendell Cockrum and Kenneth L. Fitch. Pp. 281-292, + 1 figure in text. November 15, 1952. + + 23. Comments on the taxonomy and geographic distribution of + North American microtines. By E. Raymond Hall and + E. Lendell Cockrum. Pp. 293-312. November 17, 1952. + + 24. The subspecific status of two Central American sloths. + By E. Raymond Hall and Keith R. Kelson. Pp. 313-317. + November 21, 1952. + + 25. Comments on the taxonomy and geographic distribution of + some North American marsupials, insectivores, and + carnivores. By E. Raymond Hall and Keith R. Kelson. + Pp. 319-341. December 5, 1952. + + 26. Comments on the taxonomy and geographic distribution of + some North American rodents. By E. Raymond Hall and Keith + R. Kelson. Pp. 343-371. December 15, 1952. + + 27. A synopsis or the North American microtine rodents. + By E. Raymond Hall and E. Lendell Cockrum. Pp. 373-498, + 149 figures in text. January 13, 1953. + + 28. The pocket gophers (Genus Thomomys) of Coahuila, Mexico. + By Rollin H. Baker. Pp. 499-514, 1 figure in text. + June 1, 1953. + + 29. Geographic distribution of the pocket mouse, Perognathus + fasciatus. By J. Knox Jones, Jr. Pp. 515-526, 7 figures in + text. August 1, 1953. + + 30. A new subspecies of wood rat (Neotoma mexicana) from + Colorado. By Robert B. Finley, Jr. Pp. 527-534, 2 figures + in text. August 15, 1953. + + 31. Four new pocket gophers of the genus Cratogeomys from + Jalisco, Mexico. By Robert J. Russell. Pp. 535-542. + October 15, 1953. + + 32. Genera and subgenera of chipmunks. By John A. White. + Pp. 543-561, 12 figures in text. December 1, 1953. + + 33. Taxonomy of the chipmunks, Eutamias quadrivittatus and + Eutamias umbrinus. By John A. White. Pp. 563-582, + 6 figures in text. December 1, 1953. + + 34. Geographic distribution and taxonomy of the chipmunks of + Wyoming. By John A. White. Pp. 584-610, 3 figures in text. + December 1, 1953. + + 35. The baculum of the chipmunks of western North America. + By John A. White. Pp. 611-631, 19 figures in text. + December 1, 1953. + + 36. Pleistocene Soricidae from San Josecito Cave, Nuevo Leon, + Mexico. By James S. Findley. Pp. 633-639. December 1, 1953. + + 37. Seventeen species of bats recorded from Barro Colorado + Island, Panama Canal Zone. By E. Raymond Hall and William + B. Jackson. Pp. 641-646. December 1, 1953. + + Index. Pp. 647-676. + + *Vol. 6. (Complete) Mammals of Utah, _taxonomy and distribution_. + By Stephen D. Durrant. Pp. 1-549, 91 figures in text, + 30 tables. August 10, 1952. + + Vol. 7. *1. Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, + 73 figures in text, 37 tables. August 25, 1952. + + 2. Ecology of the opossum on a natural area in northeastern + Kansas. By Henry S. Fitch and Lewis L. Sandidge. + Pp. 305-338, 5 figures in text. August 24, 1953. + + 3. The silky pocket mice (Perognathus flavus) of Mexico. + By Rollin H. Baker. Pp. 339-347, 1 figure in text. + February 15, 1954. + + 4. North American jumping mice (Genus Zapus). By Philip H. + Krutzsch. Pp. 349-472, 47 figures in text, 4 tables. + April 21, 1954. + + 5. Mammals from Southeastern Alaska. By Rollin H. Baker and + James S. Findley. Pp. 473-477. April 21, 1954. + + 6. Distribution of Some Nebraskan Mammals. By J. Knox Jones. + Pp. 479-487. April 21, 1954. + + 7. Subspeciation in the montane meadow mouse, Microtus + montanus, in Wyoming and Colorado. By Sydney Anderson. + Pp. 489-506, 2 figures in text. July 23, 1954. + + 8. A new subspecies of bat (Myotis velifer) from Southeastern + California and Arizona. By Terry A. Vaughn. Pp. 507-512. + July 23, 1954. + + 9. Mammals of the San Gabriel Mountains of California. + By Terry A. Vaughn. Pp. 513-582, 1 figure in text, + 12 tables. November 15, 1954. + + More numbers will appear in volume 7. + + Vol. 8. 1. Life History and Ecology of the Five-Lined Skink, Eumeces + fasciatus. By Henry S. Fitch. Pp. 1-156, 26 figures in + text. September 1, 1954. + + 2. Myology and Serology of the Avian Family Fringillidae, + a Taxonomic Study. By William B. Stallcup. Pp. 157-211, + 23 figures in text, 4 tables. November 15, 1954. + + More numbers will appear in volume 8. + + + + + * * * * * + + + Transcriber's Notes + + The text presented is essentially that in the original printed + document with the exception of some minor punctuation changes and + the typographical correction detailed below. Some of the tables + split between paragraphs in the original and they were moved and + the paragraphs restored into one. The captions for Figures 10-13 + and 14-17 were reformatted to enhance readability. + + + Empasis Notation + + _Text_ - Italics + + +Text+ - Bold + + + Typographical Corrections + + Page 187, Table 1 Item 5: Intavenous => Intravenous + + + * * * * * + + + + + +End of the Project Gutenberg EBook of Myology and Serology of the Avian +Family Fringillidae, by William B. 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Stallcup. + </title> + <style type="text/css"> + + body {margin-left: 10%; margin-right: 10%;} + p {text-align: justify; text-indent: 1.5em;} + sub {font-size:70%; vertical-align: -4px;} + hr {width: 95%; color: #000; clear: both;} + .hr30 {width:30%;} + table {margin-left: auto; padding:4px; margin-right: auto; border-collapse: collapse;} + .bb {border-bottom: solid #000 1px;} + .bl {border-left: solid #000 1px;} + .bt {border-top: solid #000 1px;} + .pagenum {position: absolute; left: 92%; text-indent:0; font-size: 0.75em; text-align: right; color: #b0b0b0;} + .reference {margin-left: 5.5em; text-indent: -3em;} + .vtop {vertical-align: top;} + .center {text-align: center;} + .justify {text-align: justify;} + .text_lf {text-align: left;} + .text_rt {text-align: right;} + .smaller {font-size: 0.75em;} + .smcap {font-variant: small-caps;} + .cover {background: #d0d0d0;} + .caption1 {font-weight: bold; font-size:1.75em; text-align: center;} + .caption2h {font-weight: bold; font-size:1.65em; text-align: center;} + .caption2 {font-weight: bold; font-size:1.50em; text-align: center;} + .caption3 {font-weight: bold; font-size:1.15em; text-align: center;} + .caption3nc {font-size:1.15em;} + .caption4 {font-weight: bold; font-size:0.85em; text-align: center;} + .trans_notes {background:#d0d0d0; padding: 12px; border:solid black 1px;} + .ind2em {margin-left: 2em; text-indent:-2em;} + .footnote {margin-left: 10%; margin-right: 10%; font-size: 0.9em;} + .footnote .label {position: absolute; right: 84%; text-align: right;} + .fnanchor {vertical-align: super; font-size: .8em; text-decoration: none;} + + </style> + </head> +<body> + + +<pre> + +The Project Gutenberg EBook of Myology and Serology of the Avian Family +Fringillidae, by William B. Stallcup + +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: Myology and Serology of the Avian Family Fringillidae + A Taxonomic Study + +Author: William B. Stallcup + +Release Date: October 19, 2010 [EBook #33914] + +Language: English + +Character set encoding: ISO-8859-1 + +*** START OF THIS PROJECT GUTENBERG EBOOK MYOLOGY AND SEROLOGY OF THE *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + +</pre> + + + + +<div class="trans_notes"> +<a name="typos"></a> +<div class="caption2">Transcriber's Notes</div> + +<p>Except for the typographical correction noted below and a few minor changes +(missing/extra punctuation) which may have been made but not noted here, the +text is the same as presented in the original publication. Some text has +been rearranged to restore paragraphs that were split by tables or images. +Most of the illustrations have notation to denote the scale compared to the +original specimen (example: × 3). Due to the variation in monitor resolution +and geometry, the scale is most likely not correct; but is provided as a guide.</p> + +<div class="caption2">Typographical Corrections</div> + +<p>Page 187, Table 1 Item 5 : Intavenous => Intravenous</p> +<p> </p> +</div> + +<p><span class="pagenum">[Cover]</span></p> +<div class="cover"> +<p> </p> +<img src="images/bar_double.png" width="100%" height="15" border="0" alt="double bar"> +<div class="caption2 smcap">University of Kansas Publications<br /> +Museum of Natural History</div> +<hr class="hr30"><br /> +<div class="caption3">Volume 8, No. 2, pp. 157-211, figures 1-23, 4 tables</div><br /> +<div class="center"><img src="images/bar_single.png" width="28%" height="15" title="bar" alt="bar" /> <span class="caption3">November 15, 1954</span> <img src="images/bar_single.png" width="28%" height="15" title="bar" alt="bar" /></div> +<p> </p> +<p> </p> +<p> </p> +<div class="caption1"> +Myology and Serology<br /> +of the Avian Family Fringillidae,<br /> +A Taxonomic Study +</div> +<p> </p> +<p> </p> + +<div class="caption3"> +BY<br /> +<p> </p> +WILLIAM B. STALLCUP<br /> +</div> +<p> </p> +<p> </p> +<p> </p> +<p> </p> +<p> </p> +<p> </p> + +<div class="caption2"> +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1954 +</div> +<p> </p> +<p> </p> +</div> +<p> </p> +<p> </p> + + +<p><span class="pagenum"><a name="Page_157" id="Page_157">[Pg 157]</a></span></p> + +<p> </p> +<img src="images/bar_double.png" width="100%" height="15" border="0" alt="double bar"> +<div class="caption2 smcap">University of Kansas Publications<br /> +Museum of Natural History</div> +<hr class="hr30"><br /> +<div class="caption3">Volume 8, No. 2, pp. 157-211, figures 1-23, 4 tables</div><br /> +<div class="center"><img src="images/bar_single.png" width="28%" height="15" title="bar" alt="bar" /> <span class="caption3">November 15, 1954</span> <img src="images/bar_single.png" width="28%" height="15" title="bar" alt="bar" /></div> +<p> </p> +<p> </p> +<p> </p> +<div class="caption1"> +Myology and Serology<br /> +of the Avian Family Fringillidae,<br /> +A Taxonomic Study +</div> +<p> </p> +<p> </p> + +<div class="caption3"> +BY<br /> +<p> </p> +WILLIAM B. STALLCUP<br /> +</div> +<p> </p> +<p> </p> +<p> </p> +<p> </p> +<p> </p> +<p> </p> + +<div class="caption2"> +<span class="smcap">University of Kansas</span><br /> +<span class="smcap">Lawrence</span><br /> +1954 +</div> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_158" id="Page_158">[Pg 158]</a></span></p> + +<div class="center"> +<div class="caption3"> +<span class="smcap">University of Kansas Publications, Museum of Natural History</span><br /> +<br /> +Editors: E. Raymond Hall, Chairman, A. Byron Leonard,<br /> +Robert W. Wilson<br /> +</div> +<p> </p> +<p> </p> + +<div class="caption3"> +Volume 8, No. 2, pp. 157-211, figures 1-23, 4 tables<br /> +<br /> +Published November 15, 1954<br /> +</div> +<p> </p> +<p> </p> + +<div class="caption3"> +<span class="smcap">University of Kansas</span><br /> +Lawrence, Kansas<br /> +</div> +<p> </p> +<p> </p> +<div class="caption4"> +PRINTED BY<br /> +FERD VOILAND, JR., STATE PRINTER<br /> +TOPEKA, KANSAS<br /> +1954<br /> +<br /> +<img src="images/union_label.png" width="71" height="26" title="" alt="Union Label" /><br /> +<br /> +25-4632<br /> +</div> +</div> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_159" id="Page_159">[Pg 159]</a></span></p> + +<div class="caption2">Myology and Serology<br /> +of the Avian Family Fringillidae,<br /> +a Taxonomic Study</div> +<p> </p> + +<div class="caption4"> +BY<br /><br /> +WILLIAM B. STALLCUP<br /> +</div> +<p> </p> +<p> </p> + +<a name="toc" id="toc"></a> +<div class="caption3nc"> +<div class="smcap center">Contents</div> +<p> </p> + +<table width="100%" summary="TOC List"> +<tr><td colspan=2 class="text_rt smaller">PAGE</td></tr> +<tr><td class="smcap"><a href="#Introduction">Introduction</a></td><td class="text_rt">160</td></tr> +<tr><td class="smcap"><a href="#Myology_of_the_Pelvic_Appendage">Myology of the Pelvic Appendage</a></td><td class="text_rt">162</td></tr> +<tr><td> <a href="#General_Statement_1">General Statement</a></td><td class="text_rt">162</td></tr> +<tr><td> <a href="#Materials_and_Methods">Materials and Methods</a></td><td class="text_rt">163</td></tr> +<tr><td> <a href="#Description_of_Muscles">Description of Muscles</a></td><td class="text_rt">164</td></tr> +<tr><td> <a href="#Discussion_of_the_Myological_Investigations">Discussion of Myological Investigations</a></td><td class="text_rt">175</td></tr> +<tr><td class="smcap"><a href="#Comparative_Serology">Comparative Serology</a></td><td class="text_rt">185</td></tr> +<tr><td> <a href="#General_Statement_2">General Statement</a></td><td class="text_rt">185</td></tr> +<tr><td> <a href="#Preparation_of_Antigens">Preparation of Antigens</a></td><td class="text_rt">186</td></tr> +<tr><td> <a href="#Preparation_of_Antisera">Preparation of Antisera</a></td><td class="text_rt">188</td></tr> +<tr><td> <a href="#Methods_of_Serological_Testing">Methods of Serological Testing</a></td><td class="text_rt">188</td></tr> +<tr><td> <a href="#Experimental_Data">Experimental Data</a></td><td class="text_rt">190</td></tr> +<tr><td> <a href="#Discussion_of_the_Serological_Investigations">Discussion of Serological Investigations</a></td><td class="text_rt">190</td></tr> +<tr><td class="smcap"><a href="#Conclusions">Conclusions</a></td><td class="text_rt">201</td></tr> +<tr><td class="smcap"><a href="#Summary">Summary</a></td><td class="text_rt">208</td></tr> +<tr><td class="smcap"><a href="#Literature_Cited">Literature Cited</a></td><td class="text_rt">210</td></tr> +</table> +</div> +<p> </p> +<p> </p> + +<a name="Introduction"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span><br /> +<p><span class="pagenum"><a name="Page_160" id="Page_160">[Pg 160]</a></span></p> +<div class="caption2h smcap">Introduction</div> + +<p>The relationships of many groups of birds within the Order +Passeriformes are poorly understood. Most ornithologists agree +that some of the passerine families of current classifications are +artificial groups. These artificial groupings are the result of early +work which gave chief attention to readily adaptive external structures. +The size and shape of the bill, for example, have been +over-emphasized in the past as taxonomic characters. It is now +recognized that the bill is a highly adaptive structure and that it +frequently shows convergence and parallelism.</p> + +<p>Since studies of external morphology have failed in some cases +to provide a clear understanding of the relationships of passerine +birds, it seems appropriate that attention be given to other morphological +features, to physiological features, and to life history studies +in an attempt to find other clues to relationships at the family and +subfamily levels.</p> + +<p>This paper reports the results of a study of the relationships of +some birds of the Family Fringillidae and is based on the comparative +myology of the pelvic appendage and on the comparative +serology of saline-soluble proteins. Where necessary for comparative +purposes, birds from other families have been included in these +investigations.</p> + +<p>It has long been recognized that the Fringillidae include dissimilar +groups. Recent work by Beecher (1951b, 1953) on the +musculature of the jaw and by Tordoff (1954) primarily on the +structure of the bony palate has emphasized the artificial nature +of the assemblage although these authors disagree regarding major +divisions within it (see below).</p> + +<p>The Fringillidae have been distinguished from other families of +nine-primaried oscines by only one character—a heavy and conical +bill (for crushing seeds). Bills of this form have been developed +independently in several other, unrelated, groups; as Tordoff +(1954:7) has pointed out, <i>Molothrus</i> of the Family Icteridae, +<i>Psittorostra</i> of the Family Drepaniidae, and most members of the +Family Ploceidae have bills as heavy and conical as those of the +fringillids. The ploceids are distinguished from the fringillids by +a single external character: a fairly well-developed tenth primary +whereas in fringillids the tenth primary is absent or vestigial. Tordoff +(1954:20) points out, however, that this distinction is of limited +value since in other passerine families the tenth primary may be +present in some species of a genus and absent in others. The Genus +<span class="pagenum"><a name="Page_161" id="Page_161">[Pg 161]</a></span> +<i>Vireo</i> is an example. Furthermore, at least one ploceid (<i>Philetairus</i>) +has a small, vestigial tenth primary, whereas some fringillids +(<i>Emberizoides</i>, for example) possess a tenth primary which is +rather large and ventrally placed (Chapin, 1917:253-254). Thus, +it is obvious that studies based on other features are necessary in +order to attain a better understanding of the relationships of the +birds involved.</p> + +<p>Sushkin's studies (1924, 1925) of the structure of the bony and +horny palates have served as a basis for the division of the Fringillidae +into as many as five subfamilies (Hellmayr, 1938:v): Richmondeninae, +Geospizinae, Fringillinae, Carduelinae, and Emberizinae.</p> + +<p>Beecher (1951b:280) points out that "the richmondenine finches +arise so uninterruptedly out of the tanagers that ornithologists have +had to draw the dividing line between the two groups arbitrarily." +His study of pattern of jaw-musculature substantiates this. He +states further that the cardueline finches arise without disjunction +from the tanagers. He suggests, therefore, that the two groups of +"tanager-finches" be made subfamilies of the Thraupidae and that +a third subfamily be maintained for the more typical tanagers. He +states that the emberizine finches are of different origin, arising from +the wood warblers (1953:307). Beecher (1951a:431; 1953:309) +includes the Dickcissel, <i>Spiza americana</i>, in the Family Icteridae, +chiefly on the basis of jaw muscle-pattern and the horny palate.</p> + +<p>Tordoff (1954:10-11) presents evidence that the occurrence of +palato-maxillary bones in nine-primaried birds indicates relationship +among the forms possessing them. He points out that all fringillids +except the Carduelinae possess palato-maxillaries that are either +free or more or less fused to the prepalatine bar. He points out also +that in all carduelines, the prepalatine bar is flared at its juncture +with the premaxilla, and that the mediopalatine processes are fused +across the midline; noncardueline fringillids lack these characteristics. +In addition to the above he cites differences between the +carduelines and the "other" fringillids in the appendicular skeletons, +in geographic distribution, in patterns of migration, and in habits. +Tordoff concludes, therefore, that the carduelines are not fringillids +but ploceids, their closest affinities being with the ploceid Subfamily +Estrildinae. On the basis of palatal structure, the Fringillinae and +Geospizinae are combined with the Emberizinae, the name Fringillinae +being maintained for the subfamily. The tanagers merge with +the Richmondeninae on the one hand and with the Fringillinae on +the other. On this basis, Tordoff (1954:32) suggests that the Family +<span class="pagenum"><a name="Page_162" id="Page_162">[Pg 162]</a></span> +Fringillidae be divided into subfamilies as follows: Richmondeninae, +Thraupinae, and Fringillinae. The carduelines are placed as +the Subfamily Carduelinae in the Family Ploceidae.</p> + +<p>From the foregoing, it is apparent that the two most recent lines +of research have given rise to conflicting theories regarding relationships +within the Family Fringillidae. The purpose of my investigation, +therefore, has been to gather information, from other fields, +which might clarify the relationships of these birds.</p> + +<p>Since the muscle pattern of the leg in the Order Passeriformes is +thought to be one of long standing and slow change, any variation +which consistently distinguishes one group of species from another +could be significant. With the hope that such variation might be +found, a study of the comparative myology of the legs was undertaken.</p> + +<p>The usefulness of comparative serology as a means of determining +relationship has been demonstrated in many investigations. Its use +in this instance was undertaken for several reasons: comparative +serology has its basis in biochemical systems which seem to evolve +slowly; its methods are objective; and its use has, heretofore, resulted +in the accumulation of data which seem compatible, in most +instances, with data obtained from other sources.</p> + +<p>I acknowledge with pleasure the guidance received in this study +from Prof. Harrison B. Tordoff of the University of Kansas. I am +indebted also to Prof. Charles A. Leone without whose direction +and assistance the serological investigations would not have been +possible; to Professors E. Raymond Hall and A. Byron Leonard +whose suggestions and criticisms have been most helpful in the +preparation of this paper; and to T. D. Burleigh of the U. S. Fish +and Wildlife Service for gifts of several specimens used in this work. +Assistance with certain parts of the study were received from a contract +(NR163014) between the Office of Naval Research of the +United States Navy and the University of Kansas.</p> +<p> </p> +<p> </p> + +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span> +<a name="Myology_of_the_Pelvic_Appendage" id="Myology_of_the_Pelvic_Appendage"></a> +<div class="caption2h smcap">Myology of the Pelvic Appendage</div> +<p> </p> +<a name="General_Statement_1"></a> +<div class="caption2">General Statement</div> + +<p>In an excellent paper in which the muscles of the pelvic appendage +of birds are carefully and accurately described, Hudson (1937) +reviewed briefly the more important literature pertaining to the +musculature of the leg which had been published to that date. A +review of such information here, therefore, seems unnecessary.</p> + +<p>Myological formulae suggested by Garrod (1873, 1874) have +<span class="pagenum"><a name="Page_163" id="Page_163">[Pg 163]</a></span> +been extensively used by taxonomists as aids in characterizing the +orders of birds. Relatively few investigations, however, involving +the comparative myology of the leg have been undertaken at family +and subfamily levels. The works of Fisher (1946), Hudson (1948), +and Berger (1952) are notable exceptions.</p> + +<p>The terminology for the muscles used in this paper follows that +of Hudson (1937), except that I have followed Berger (1952) in +Latinizing all names. Homologies are not given since these are +reviewed by Hudson. Osteological terms are from Howard (1929).</p> + +<a name="Materials_and_Methods"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span> +<div class="caption2">Materials and Methods</div> + +<p>Specimens were preserved in a solution of one part formalin to eight parts +of water. Thorough injection of all tissues was necessary for satisfactory preservation. +Most of the down and contour feathers were removed to allow the +preservative to reach the skin.</p> + +<p>In preparing specimens for study, the legs and pelvic girdle were removed +and washed in running water for several hours to remove much of the formalin. +They were then transferred to a mixture of 50 per cent alcohol and a small +amount of glycerine.</p> + +<p>All specimens were dissected with the aid of a low power binocular microscope. +Where possible, several specimens of each species were examined for +individual differences. Such differences were found to be slight, involving +mainly size and shape of the muscles. The size is dependent partly on the +age of the bird, muscles from older birds being larger and better developed. +The shape of a muscle (whether long and slender or short and thick) is due in +part to the position in which the leg was preserved; that is to say, a muscle +may be extended in one bird and contracted in another. For these reasons, +descriptions and comparisons are based mainly on the origin and insertion of a +muscle and on its position in relation to adjoining muscles.</p> + +<p>Birds dissected in this study are listed below (in the order of the A. O. U. +Check-List):</p> + +<div class="center"> +<span class="caption3">SPECIES</span><br /> + +<table width="80%" summary="Bird Checklist"> +<tr><td> +<i>Vireo olivaceus</i> (Linnaeus)<br /> +<i>Seiurus motacilla</i> (Vieillot)<br /> +<i>Passer domesticus</i> (Linnaeus)<br /> +<i>Estrilda amandava</i> (Linnaeus)<br /> +<i>Poephila guttata</i> (Reichenbach)<br /> +<i>Icterus galbula</i> (Linnaeus)<br /> +<i>Molothrus ater</i> (Boddaert)<br /> +<i>Piranga rubra</i> (Linnaeus)<br /> +<i>Richmondena cardinalis</i> (Linnaeus)<br /> +<i>Guiraca caerulea</i> (Linnaeus)<br /> +<i>Passerina cyanea</i> (Linnaeus)<br /> +<i>Spiza americana</i> (Gmelin)<br /> +<i>Hesperiphona vespertina</i> (Cooper)<br /> +<i>Carpodacus purpureus</i> (Gmelin)<br /> +</td><td> </td><td class="vtop"> +<i>Pinicola enucleator</i> (Linnaeus)<br /> +<i>Leucosticte tephrocotis</i> (Swainson)<br /> +<i>Spinus tristis</i> (Linnaeus)<br /> +<i>Loxia curvirostra</i> Linnaeus<br /> +<i>Chlorura chlorura</i> (Audubon)<br /> +<i>Pipilo erythrophthalmus</i> (Linnaeus)<br /> +<i>Calamospiza melanocorys</i> Stejneger<br /> +<i>Chondestes grammacus</i> (Say)<br /> +<i>Junco hyemalis</i> (Linnaeus)<br /> +<i>Spizella arborea</i> (Wilson)<br /> +<i>Zonotrichia querula</i> (Nuttall)<br /> +<i>Passerella iliaca</i> (Merrem)<br /> +<i>Calcarius lapponicus</i> (Linnaeus)<br /> +</td></tr> +</table> +</div> +<p> </p> +<p> </p> + +<a name="Description_of_Muscles"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span><br /> +<p><span class="pagenum"><a name="Page_164" id="Page_164">[Pg 164]</a></span></p> +<div class="caption2">Description of Muscles</div> + +<p>The descriptions which follow are those of the muscles in the leg of the +Red-eyed Towhee, <i>Pipilo erythrophthalmus</i>. Differences between species, +where present, are noted for each muscle. The term thigh is used to refer to +the proximal segment of the leg; the term crus is used for that segment of the +leg immediately distal to the thigh.</p> +<p> </p> + +<p><i><b>Musculus iliotrochantericus posticus</b></i> (Fig.<a href="#Fig_2"> 2</a>).—The origin of this muscle is +fleshy from the entire concave lateral surface of the ilium anterior to the acetabulum. +The fibers converge posteriorly, and the muscle inserts by a short, +broad tendon on the lateral surface of the femur immediately distal to the +trochanter. It is the largest muscle which passes from the ilium to the femur.</p> + +<p>Action.—Moves femur forward and rotates it anteriorly.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus iliotrochantericus anticus</b></i> (Fig.<a href="#Fig_3"> 3</a>).—Covered laterally by the <i>m. +iliotrochantericus posticus</i>, this slender muscle has a fleshy origin from the +anteroventral edge of the ilium between the origins of the <i>m. sartorius</i> anteriorly +and the <i>m. iliotrochantericus medius</i> posteriorly. The <i>m. iliotrochantericus +anticus</i> is directed caudoventrally and inserts by a broad, flat tendon on the +anterolateral surface of the femur between the heads of the <i>m. femorotibialis +externus</i> and <i>m. femorotibialis medius</i> and just distal to the insertion of the <i>m. +iliotrochantericus medius</i>.</p> + +<p>Action.—Moves femur forward and rotates it anteriorly.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus iliotrochantericus medius</b></i> (Fig.<a href="#Fig_3"> 3</a>).—Smallest of the three <i>iliotrochantericus</i> +muscles, this bandlike muscle has a fleshy origin from the ventral +edge of the ilium just posterior to the origin of the <i>m. iliotrochantericus anticus</i>. +The fibers are directed caudoventrally, and the insertion is tendinous on the +anterolateral surface of the femur between the insertion of the other two <i>iliotrochantericus</i> +muscles.</p> + +<p>Action.—Moves femur forward and rotates it anteriorly.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus iliacus</b></i> (Figs.<a href="#Fig_4"> 4</a>,<a href="#Fig_5"> 5</a>).—Arising from a fleshy origin on the ventral +edge of the ilium just posterior to the origin of the <i>m. iliotrochantericus medius</i>, +this small slender muscle passes posteroventrally to its fleshy insertion on the +posteromedial surface of the femur just proximal to the origin of the <i>m. femorotibialis +internus</i>.</p> + +<p>Action.—Moves femur forward and rotates it posteriorly.</p> + +<p>Comparison.—No significant differences among the species studied.</p> +<p> </p> + +<p><i><b>Musculus sartorius</b></i> (Figs.<a href="#Fig_1"> 1</a>,<a href="#Fig_4"> 4</a>).—A long, straplike muscle, the <i>sartorius</i> +forms the anterior edge of the thigh. The origin is fleshy, half from the +anterior edge of the ilium and from the median dorsal ridge of this bone and +half from the posterior one or two free dorsal vertebrae. The insertion is +fleshy along a narrow line on the anteromedial edge of the head of the tibia and +on the medial region of the patellar tendon.</p> + +<p>Action.—Moves thigh forward and upward and extends shank.</p> + +<p>Comparison.—In <i>Loxia</i> and <i>Spinus</i>, only one-third of the origin is from the +last free dorsal vertebra. In <i>Hesperiphona</i>, <i>Carpodacus</i>, <i>Pinicola</i>, and <i>Leucosticte</i>, +only one-fifth of the origin is from this vertebra.</p> +<p> </p> + +<p><span class="pagenum"><a name="Page_165" id="Page_165">[Pg 165]</a></span> +<i><b>Musculus iliotibialis</b></i> (Fig.<a href="#Fig_1"> 1</a>).—Broad and triangular, this muscle covers +most of the deeper muscles of the lateral aspect of the thigh. The middle +region is fused with the underlying <i>femorotibialis</i> muscles. In the distal half +of this muscle there are three distinct parts; the anterior and posterior edges +are fleshy and the central part is aponeurotic. The origin is from a narrow line +along the iliac crests—from the origin of the <i>m. sartorius</i>, anteriorly, to the +origin of the <i>m. semitendinosus</i> posteriorly. The origin is aponeurotic in the +preacetabular region but fleshy in the postacetabular region. The distal part +of the muscle is aponeurotic and joins with the <i>femorotibialis</i> muscles in the +formation of the patellar tendon. This tendon incloses the patella and inserts +on a line along the proximal edges of the cnemial crests of the tibiotarsus.</p> + +<p>Action.—Extends crus.</p> + +<p>Comparison.—In <i>Vireo</i> the central aponeurotic portion of this muscle is +absent.</p> +<p> </p> + +<p><i><b>Musculus femorotibialis externus</b></i> (Fig.<a href="#Fig_2"> 2</a>).—Covering the lateral and anterolateral +surfaces of the femur, this large muscle has a fleshy origin from the +lateral edge of the proximal three-fourths of the femur. The origin separates +the insertion of the <i>m. iliotrochantericus anticus</i> from that of the <i>m. ischiofemoralis</i> +and, in turn, is separated from the origin of the <i>m. femorotibialis +medius</i> by the insertions of the <i>m. iliotrochantericus anticus</i> and <i>m. iliotrochantericus +medius</i>. Approximately midway of the length of the femur this +muscle fuses anteromesially with the <i>m. femorotibialis medius</i>. Distally, the +<i>m. femorotibialis externus</i> contributes to the formation of the patellar tendon +which inserts on a line along the proximal edges of the cnemial crests of the +tibiotarsus.</p> + +<p>Action.—Extends crus.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus femorotibialis medius</b></i> (Figs.<a href="#Fig_2"> 2</a>,<a href="#Fig_4"> 4</a>).—The origin of this muscle, +which lies along the anterior edge of the femur, is fleshy from the entire length +of the femur proximal to the level of attachment of the proximal arm of the +biceps loop. Laterally this muscle is completely fused for most of its length +with the <i>m. femorotibialis externus</i> and contributes to the formation of the +patellar tendon, which inserts on a line along the proximal edges of the cnemial +crests of the tibiotarsus. Many of the fibers, nevertheless, insert on the proximal +edge of the patella.</p> + +<p>Action.—Extends crus.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus femorotibialis internus</b></i> (Fig.<a href="#Fig_4"> 4</a>).—One of the most superficial +muscles lying on the medial surface of the thigh, this muscle is divided, +especially near the distal end, into two parts, lateral and medial. The origin of +the lateral part is fleshy from a line on the medial surface of the femur; the +origin begins proximally at a point near the insertion of the <i>m. iliacus</i>. The +medial, bulkier part of the muscle has a fleshy origin on the medial surface of +the lower one-third of the femur. The two parts fuse to some extent above the +points of insertion and insert on the medial edge of the head of the tibia.</p> + +<p>Action.—Rotates tibia anteriorly.</p> + +<p>Comparison.—Two parts of this muscle variously fused; otherwise, no significant +differences in the species studied.</p> +<p> </p> + +<p><i><b>Musculus piriformis</b></i> (Fig.<a href="#Fig_3"> 3</a>).—This muscle is represented by the <i>pars caudifemoralis</i> +<span class="pagenum"><a name="Page_166" id="Page_166">[Pg 166]</a></span> +only, the <i>pars iliofemoralis</i> being absent in passerine birds as far as +is known. The <i>pars caudifemoralis</i> is flat, somewhat spindle-shaped, and passes +anteroventrally from the pygostyle to the femur. The origin is tendinous from +the anteroventral edge of the pygostyle, and the insertion is semitendinous on +the posterolateral surface of the shaft of the femur about one-fourth its length +from the proximal end.</p> + +<p>Action.—Moves femur posteriorly and rotates it in this direction; moves tail +laterally and depresses it.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus semitendinosus</b></i> (Figs.<a href="#Fig_2"> 2</a>,<a href="#Fig_3"> 3</a>,<a href="#Fig_5"> 5</a>).—The origin from the extreme posterior +edge of the posterior iliac crest of the ilium is fleshy and is aponeurotic +from the last vertebra of the synsacrum and the transverse processes of several +caudal vertebrae. The straplike belly passes along the posterolateral margin +of the thigh. Immediately posterior to the knee, the muscle is divided transversely +by a ligament. That portion passing anteriorly from the ligament is +the <i>m. accessorius semitendinosi</i> (here considered a part of the <i>m. semitendinosus</i>) +and is discussed below. The ligament continues distally in two parts; +one part inserts on the medial surface of the <i>pars media</i> of the <i>m. gastrocnemius</i> +and the other part fuses with the tendon of insertion of the <i>m. semimembranosus</i>.</p> + +<p>The <i>m. accessorius semitendinosi</i> extends anteriorly from the above mentioned +ligament to a fleshy insertion on the posterolateral surface of the femur +immediately proximal to the condyles.</p> + +<p>Action.—Moves femur posteriorly, flexes the crus and aids in extending the +tarsometatarsus.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus semimembranosus</b></i> (Figs.<a href="#Fig_3"> 3</a>,<a href="#Fig_4"> 4</a>,<a href="#Fig_5"> 5</a>).—This straplike muscle passes +along the posteromedial surface of the thigh. The origin is semitendinous along +a line on the ischium, from a point dorsal to the middle of the ischiopubic +fenestra to the posterior end of the ischium, and from a small area of the +abdominal musculature posterior to the ischium. The insertion is by means of +a broad, thin tendon on a ridge on the medial surface of the tibia immediately +distal to the head of this bone. The tendon of insertion passes between the +head of the <i>pars media</i> and <i>pars interna</i> of the <i>m. gastrocnemius</i> and is fused +with the tendon of the <i>m. semitendinosus</i>.</p> + +<p>Action.—Flexes crus.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus biceps femoris</b></i> (Fig.<a href="#Fig_2"> 2</a>).—Long, thin, and somewhat triangular, +this muscle lies on the lateral side of the thigh just underneath the <i>m. iliotibialis</i>. +Its origin is from a line along the anterior and posterior iliac crests underneath +the origin of the <i>m. iliotibialis</i>. Anterior to the acetabulum the origin is aponeurotic, +and the edge of this aponeurosis passes over the proximal end of the +femur. The origin posterior to the acetabulum is fleshy. The most anterior +point of origin is difficult to ascertain but it lies near the center of the anterior +iliac crest. The most posterior point of origin is immediately dorsal to the +posterior end of the ilioischiatic fenestra. Behind the knee the fibers of this +muscle converge to form the strong tendon of insertion which passes through +the biceps loop, under the tendon of origin of the <i>m. flexor perforatus digiti II</i>, +<span class="pagenum"><a name="Page_167" id="Page_167">[Pg 167]</a></span> +and inserts on a small tubercle on the posterolateral edge of the fibula at the +point of the tibia-fibula fusion.</p> + +<p>The biceps loop is tendinous and the distal end attaches to a protuberance +on the posterolateral edge of the femur at the proximal edge of the external +condyle. The proximal end attaches to the anterolateral edge of the femur immediately +proximal to the distal end of the loop, which extends posterior to the +femur. The distal arm of this loop is connected with the tendon of origin of +the <i>m. flexor perforatus digiti II</i> by a strong tendon.</p> + +<p>Action.—Flexes crus.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus ischiofemoralis</b></i> (Fig.<a href="#Fig_3"> 3</a>).—Short and thick, this muscle arises directly +from the lateral surface of the ischium between the posterior iliac crest +and the ischiopubic fenestra. The area of origin extends to the posterior edge +of the ischium. The insertion is tendinous on the lateral surface of the trochanter +opposite the insertion of the <i>m. iliotrochantericus medius</i>.</p> + +<p>Action.—Moves femur posteriorly and rotates it in this direction.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> + +<p><i><b>Musculus obturator internus</b></i> (Figs.<a href="#Fig_4"> 4</a>,<a href="#Fig_7"> 7</a>).—Lying on the inside of the pelvis +and covering the medial surface of the ischiopubic fenestra, is this flat, pinnate, +leaf-shaped muscle. The origin is fleshy and is from the ischium and pubis +around the edges of this fenestra; none of the fibers arises from the membrane +stretched across the fenestra. Anteriorly the fibers converge and form a strong +tendon that passes through the obturator foramen and inserts on the posterolateral +surface of the trochanter of the femur.</p> + +<p>Action.—Rotates femur posteriorly.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus obturator externus</b></i> (Fig.<a href="#Fig_7"> 7</a>).—Short and fleshy, this muscle consists +of two parts which are not easily separable but which may be traced throughout +its length. The parts are more nearly distinct at the origin. The dorsal +part arises directly from the ischium along the dorsal edge of the obturator +foramen. The larger ventral part arises directly from the anterior and ventral +edges of the obturator foramen. The fibers of the dorsal part pass anteriorly, +cover the tendon of the <i>m. obturator internus</i> laterally, and insert on the trochanter +around the point of insertion of the latter muscle. The fibers of the +ventral part pass parallel with the tendon of the <i>m. obturator internus</i> and insert +on the trochanter immediately distal and posterior to the tendon of the latter +muscle.</p> + +<p>Action.—Rotates femur posteriorly.</p> + +<p>Comparison.—In <i>Passer</i>, <i>Estrilda</i>, <i>Poephila</i>, <i>Hesperiphona</i>, <i>Carpodacus</i>, <i>Pinicola</i>, +<i>Leucosticte</i>, <i>Spinus</i> and <i>Loxia</i>, this muscle is undivided and, in its position, +origin, and insertion, resembles the ventral part of the bipartite muscle +described above. The origin is from the anterior and ventral edges of the +obturator foramen and the insertion is on the trochanter of the femur immediately +distal and posterior to the insertion of the <i>m. obturator internus</i>. In all +other genera examined, the muscle is bipartite. In <i>Chlorura</i> the dorsal part is +larger and better developed than it is in the other genera.</p> +<p> </p> + +<p><i><b>Musculus adductor longus et brevis</b></i> (Figs.<a href="#Fig_3"> 3</a>,<a href="#Fig_4"> 4</a>, <a href="#Fig_5">5</a>).—Consisting of two distinct, +straplike parts, this large muscle lies on the medial surface of the thigh, +posterior to the femur.</p> + +<p><span class="pagenum"><a name="Page_168" id="Page_168">[Pg 168]</a></span> +The <i>pars anticus</i> has a semitendinous origin on a line that extends posteriorly +from the posteroventral edge of the obturator foramen to a point half way across +the membrane that covers the ischiopubic fenestra. The insertion is fleshy +along the posterior surface of the femur from the level of the insertion of the +<i>m. piriformis</i> distally to the medial surface of the internal condyle.</p> + +<p>The <i>pars posticus</i> originates by a broad, flat tendon on a line across the +posterior half of the membrane that covers the ischiopubic fenestra. The insertion +is at the point of origin of the <i>pars media</i> of the <i>m. gastrocnemius</i> on +the posteromedial surface of the proximal end of the internal condyle of the +femur. There is a broad tendinous connection with the proximal end of the +<i>pars media</i> of the <i>m. gastrocnemius</i>. The anterior edge of the <i>pars posticus</i> is +overlapped medially by the posterior edge of the <i>pars anticus</i>.</p> + +<p>Action.—Flexes thigh; may flex crus also and may extend tarsometatarsus.</p> + +<p>Comparison.—In <i>Vireo olivaceous</i>, the origin of this muscle does not extend +the length of the ischiopubic fenestra. The origin, furthermore, is along the +dorsal edge of the ischiopubic fenestra and not from the membrane covering +the fenestra. Finally, in this species, the origin of the <i>pars posticus</i> is fleshy.</p> +<p> </p> + +<p><i><b>Musculus tibialis anticus</b></i> (Figs.<a href="#Fig_2"> 2</a>,<a href="#Fig_5"> 5</a>).—Lying along the anterior edge of the +crus, a part of this muscle is covered by the <i>m. peroneus longus</i>. The origin is +by two distinct heads, each of which is pinnate. The anterior head arises +directly from the edges of the outer and inner cnemial crests. The posterior +head arises by a short, strong tendon from a small pit on the anterodistal edge +of the external condyle of the femur. This tendon and the proximal end of +the muscle pass between the head of the fibula and the outer cnemial crest. +The two heads of the muscle fuse at a place slightly more than one-half of the +distance down the crus. At the distal end of the crus this muscle gives rise to +a strong tendon which passes under a fibrous loop immediately proximal to +the external condyle in company with the <i>m. extensor digitorum longus</i> and +which passes between the condyles of the tibia and inserts on a tubercle on the +anteromedial edge of the proximal end of the tarsometatarsus.</p> + +<p>Action.—Flexes tarsometatarsus.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus extensor digitorum longus</b></i> (Figs.<a href="#Fig_3"> 3</a>,<a href="#Fig_5"> 5</a>, <a href="#Fig_8">8</a>).—Slender and pinnate, +this muscle lies along the anteromedial surface of the tibia. The origin is fleshy +from most of the region between the cnemial crests and from a line along the +anterior surface of the proximal fourth of the tibia. Approximately two-thirds +of the distance down the crus the muscle gives rise to the tendon of insertion +which passes through the fibrous loop near the distal end of the tibia in company +with the <i>m. tibialis anticus</i>. The tendon then passes along beneath the +supratendinal bridge at the distal end of the tibia, traverses the anterior intercondylar +fossa, and passes beneath a bony bridge on the anteromedial surface +of the proximal end of the tarsometatarsus. The tendon continues along the +anterior surface of the tarsometatarsus to a point immediately above the bases +of the toes and there gives rise to three branches, one to the anterior surface of +each foretoe. The insertions of each branch are on the anterior surfaces of the +phalanges as shown in Fig.<a href="#Fig_8"> 8</a>.</p> + +<p>Action.—Extends foretoes.</p> + +<p>Comparison.—This muscle is weakly developed in <i>Leucosticte</i> and <i>Calvarius</i>; +the belly is slender and extends only half way down the crus before giving rise +<span class="pagenum"><a name="Page_169" id="Page_169">[Pg 169]</a></span> +to the tendon of insertion. The functional significance of this variation is difficult +to understand. The convergence in muscle pattern shown by these two +genera, however, is in all probability the result of similarities in behavior patterns. +These birds perch less frequently than do the other birds studied. Thus, +the toes are neither flexed nor extended as often; the smaller size of the <i>m. +extensor digitorum longus</i> may have resulted in part from this lessened activity. +Except for the variations just noted, there are no significant differences among +the species studied; even the rather complex patterns of insertion are identical.</p> +<p> </p> + +<p><i><b>Musculus peroneus longus</b></i> (Fig.<a href="#Fig_1"> 1</a>).—Relatively thin and straplike, this +muscle lies on the anterolateral surface of the crus and is intimately attached +to the underlying muscles. The part of the origin from the proximal edges of +the inner and outer cnemial crests is semitendinous but the part of the origin +from the lateral edge of the shaft of the fibula is tendinous. Approximately +two-thirds the distance down the crus the muscle gives rise to the tendon of +insertion. Immediately above the external condyle of the tibiotarsus this tendon +divides. The posterior branch inserts on the proximal end of the lateral edge +of the tibial cartilage. The anterior branch passes over the lateral surface of +the external condyle to the posterior surface of the tarsometatarsus and there +unites with the tendon of the <i>m. flexor perforatus digiti III</i>.</p> + +<p>Action.—Extends tarsometatarsus and flexes third digit.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus peroneus brevis</b></i> (Figs.<a href="#Fig_2"> 2</a>,<a href="#Fig_3"> 3</a>).—Lying along the anterolateral surface +of the tibia, this slender, pinnate muscle arises from a fleshy origin along +this surface and along the anterior surface of the fibula from a point immediately +proximal to the insertion of the <i>m. biceps femoris</i> to a point approximately +two-thirds of the way down the crus. Near the distal end of the tibia +the muscle gives rise to the tendon of insertion that passes through a groove on +the anterolateral edge of the tibia just above the external condyle. Here the +tendon is held in place by a broad fibrous loop and passes under the anterior +branch of the tendon of insertion of the <i>m. peroneus longus</i> and inserts on a +prominence on the lateral edge of the proximal end of the tarsometatarsus.</p> + +<p>Action.—Extends tarsometatarsus and may abduct it slightly.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus gastrocnemius</b></i> (Figs.<a href="#Fig_1"> 1</a>,<a href="#Fig_4"> 4</a>).—The largest muscle of the pelvic appendage, +it covers superficially all of the posterior surface, most of the medial +surface, and half of the lateral surface of the crus. The muscle originates by +three distinct heads.</p> + +<p>The <i>pars externa</i> covers the posterolateral surface of the crus, is intermediate +in size between the other two heads, and arises by a short, strong tendon from +a small bony protuberance on the posterolateral side of the distal end of the +femur immediately proximal to the fibular condyle. The tendon is intimately +connected with the distal arm of the loop for the <i>m. biceps femoris</i>.</p> + +<p>The <i>pars media</i> is the smallest of the three heads and lies on the medial surface +of the crus. The head of the <i>pars media</i> is separated from the <i>pars +interna</i> by the tendon of insertion of the <i>m. semimembranosus</i> and originates +by a short, strong tendon from the posteromedial surface of the proximal end +of the internal condyle of the femur. The proximal portion of the <i>pars media</i> +has tendinous connections with the tendon of the <i>m. semitendinosus</i> and with +the <i>pars posticus</i> of the <i>m. adductor longus et brevis</i>.</p> + +<p><span class="pagenum"><a name="Page_170" id="Page_170">[Pg 170]</a></span> +The <i>pars interna</i> is the largest of the three heads and covers most of the +medial surface of the crus. This head in its proximal portion is distinctly +divided into anterior and posterior parts, the former overlapping the latter +medially. The origin of the posterior part is fleshy from the anterior half of +the tibial head. Some of the fibers of the anterior part arise directly from the +inner cnemial crest while its remaining fibers arise from the patellar tendon +(Fig.<a href="#Fig_1"> 1</a>) and form a band that extends around the anterior surface of the knee, +covering the insertion of the <i>m. sartorius</i>.</p> + +<p>Approximately half way down the crus, the three heads give rise to the +tendon of insertion, the <i>tendo achillis</i>, which passes over and is tightly bound +to the posterior surface of the tibial cartilage. The insertion is tendinous on +the posterior surface of the hypotarsus and along the posterolateral ridge of +the tarsometatarsus. This tendon seems to be continuous with a fascia which +forms a sheath around the posterior surface of the tarsometatarsus holding the +other tendons of this region firmly in the posterior sulcus.</p> + +<p>Action.—Extends tarsometatarsus.</p> + +<p>Comparison.—Study of the <i>pars externa</i> and <i>pars media</i> reveals no significant +differences among the species dissected. The <i>pars interna</i>, however, is +subject to some variation which is described below.</p> + +<div class="center"> +<span class="caption3"><i>Pars interna</i> bipartite</span><br /> + +<table width="50%" summary="pars interna bipartite"> +<tr><td> + <i>Vireo</i><br /> + <i>Seiurus</i><br /> + <i>Icterus</i><br /> + <i>Molothrus</i><br /> + <i>Piranga</i><br /> + <i>Richmondena</i><br /> + <i>Guiraca</i><br /> + <i>Passerina</i><br /> + <i>Spiza</i><br /> +</td><td> </td><td> + <i>Chlorura</i><br /> + <i>Pipilo</i><br /> + <i>Calamospiza</i><br /> + <i>Chondestes</i><br /> + <i>Junco</i><br /> + <i>Spizella</i><br /> + <i>Zonotrichia</i><br /> + <i>Passerella</i><br /> + <i>Calcarius</i><br /> +</td></tr> +</table> +</div> +<p> </p> + +<p>The two parts of the <i>m. gastrocnemius</i> are most distinct in <i>Vireo</i>. <i>Icterus</i>, +<i>Molothrus</i>, <i>Richmondena</i>, <i>Guiraca</i>, and <i>Passerina</i> lack the fibrous band that +passes around the front of the knee. In <i>Spiza</i> this band of fibers is smaller +than in the other species.</p> + +<div class="center"> +<span class="caption3"><i>Pars interna</i> undivided</span><br /> + +<table width="50%" summary="pars interna bipartite"> +<tr><td> + <i>Passer</i><br /> + <i>Estrilda</i><br /> + <i>Poephila</i><br /> + <i>Hesperiphona</i><br /> + <i>Carpodacus</i><br /> +</td><td> </td><td> + <i>Pinicola</i><br /> + <i>Leucosticte</i><br /> + <i>Spinus</i><br /> + <i>Loxia</i><br /> +</td></tr> +</table> +</div> +<p> </p> + +<p>In <i>Leucosticte</i>, although the <i>pars interna</i> is undivided, there is a band of +fibers which extends around the front of the knee (see discussion, p.<a href="#Page_183"> 183</a>).</p> +<p> </p> + +<p><i><b>Musculus plantaris</b></i> (Fig.<a href="#Fig_5"> 5</a>).—Small and slender, this muscle lies on the +posteromedial surface of the crus, beneath the <i>pars interna</i> of the <i>m. gastrocnemius</i> +and originates by fleshy fibers from the posteromedial surface of the +proximal end of the tibia immediately distal to the internal articular surface. +The belly extends approximately one-sixth of the way down the crus and gives +rise to a long, slender tendon that inserts on the proximomedial edge of the +tibial cartilage.<span class="pagenum"><a name="Page_171" id="Page_171">[Pg 171]</a></span></p> + +<p>Action.—Extends tarsometatarsus.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus flexor perforatus digiti II</b></i> (Figs.<a href="#Fig_3"> 3</a>,<a href="#Fig_9"> 9</a>).—This is a slender muscle +which lies on the lateral side of the crus beneath the <i>pars externa</i> of the <i>m. +gastrocnemius</i> and is intimately connected anteromedially with the <i>m. flexor +digitorum longus</i> and posteromedially with the <i>m. flexor hallucis longus</i>. The +origin is by a strong tendon from the lateral surface of the external condyle of +the femur at the point of origin of the <i>m. flexor perforans et perforatus digiti II</i>. +This tendon serves also as the origin of the anterior head of the <i>m. flexor +hallucis longus</i>. The tendon connects also by a broad tendinous band with the +distal arm of the loop for the <i>m. biceps femoris</i> and by a similar band with the +lateral edge of the fibula immediately distal to the head. The tendon of insertion +passes distally, perforates the tibial cartilage near its lateral edge, traverses +the middle medial canal of the hypotarsus (Fig.<a href="#Fig_6"> 6</a>), and passes distally +to the foot. At the distal end of the tarsometatarsus the tendon is held against +the medial surface of the first metatarsal by a straplike sheath. The tendon +then passes over a sesamoid bone between the first metatarsal and the base of +the second digit and is bound to this bone by a sheath. The tendon inserts +mainly along the posteromedial edge of the proximal end of the first phalanx +of the second digit, although the termination is sheathlike and covers the entire +posterior surface of this phalanx. This sheathlike termination is perforated by +the tendons of the <i>m. flexor perforans et perforatus digiti II</i> and the branch of +the <i>m. flexor digitorum longus</i> that inserts on the second digit.</p> + +<p>Action.—Flexes second digit.</p> + +<p>Comparison.—In <i>Vireo</i> this muscle is larger and more deeply situated than +it is in the other species examined and has no connection with the <i>m. flexor +hallucis longus</i>.</p> +<p> </p> + +<p><i><b>Musculus flexor perforatus digiti III</b></i> (Fig.<a href="#Fig_5"> 5</a>).—Long and flattened, this +muscle lies on the posteromedial side of the crus beneath the <i>m. gastrocnemius</i>. +The belly is tightly fused laterally with the belly of the <i>m. flexor hallucis longus</i> +and posteriorly with the belly of the <i>m. flexor perforatus digiti IV</i>. The origin +is by a long, strong tendon from a small tubercle just medial to, and at the +proximal end of, the external condyle of the femur. Below the middle of the +crus this muscle terminates in a strong tendon which perforates the tibial +cartilage near its lateral edge. In this region the tendon is sheathlike and +wrapped around the tendon of the <i>m. flexor perforatus digiti IV</i>. These two +tendons together pass through the posterolateral canal of the hypotarsus (Fig.<a href="#Fig_6"> +6</a>). Immediately distal to the hypotarsus the two tendons separate, and the +tendon of the <i>m. flexor perforatus digiti III</i> receives a branch of the tendon of +the <i>m. peroneus longus</i>. The tendon passes distally over the surface of the +second trochlea, and its insertion is sheathlike on the posterior surface of the +first phalanx, and on the proximal end of the second. In the area of insertion +this tendon is perforated by that of the <i>m. flexor perforans et perforatus digiti +III</i> and by that of the <i>m. flexor digitorum longus</i> to the third digit.</p> + +<p>Action.—Flexes digit III.</p> + +<p>Comparison.—In <i>Passer</i>, <i>Estrilda</i>, <i>Poephila</i>, <i>Hesperiphona</i>, <i>Carpodacus</i>, +<i>Pinicola</i>, <i>Leucosticte</i>, <i>Spinus</i>, and <i>Loxia</i> the edges of the sheathlike tendon are +thickened at the points of insertion, so that the tendon appears to have two +branches which insert along the posterolateral edges of the first phalanx and are +connected medially by a fascia.</p> +<p> </p> + +<p><span class="pagenum"><a name="Page_172" id="Page_172">[Pg 172]</a></span> +<i><b>Musculus flexor perforatus digiti IV</b></i> (Fig.<a href="#Fig_3"> 3</a>).—Extending along the posterior +edge of the crus, this slender muscle lies beneath the <i>m. gastrocnemius</i>. +The belly is fused with those of the <i>m. flexor hallucis longus</i> and <i>m. flexor perforatus +digiti III</i>. Its origin is fleshy from the intercondyloid region of the distal +end of the femur and has a few fibers arising from the tendon of origin of the +<i>m. flexor perforatus digiti III</i>. Near the distal end of the crus the muscle gives +rise to the strong tendon of insertion which perforates the tibial cartilage near +its lateral edge and in this region is ensheathed by the tendon of the <i>m. flexor +perforatus digiti III</i>. The two tendons pass together through the posterolateral +canal of the hypotarsus (Fig.<a href="#Fig_6"> 6</a>). The tendon continues distally along the +tarsometatarsus and the posterior surface of digit IV. The tendon bifurcates +at approximately the middle of the first phalanx. A short lateral branch inserts +on the posterolateral edge of the proximal end of the second phalanx. The +long medial branch is perforated by a branch of the <i>m. flexor digitorum longus</i>; +the distal end is flattened, has thickened edges, and inserts over the posterior +surfaces of the distal end of the second phalanx, and over the proximal end of +the third phalanx.</p> + +<p>Action.—Flexes digit IV.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus flexor perforans et perforatus digiti II</b></i> (Figs.<a href="#Fig_2"> 2</a>,<a href="#Fig_9"> 9</a>).—Small and +spindle-shaped, this muscle lies on the posterolateral side of the crus immediately +beneath the <i>pars externa</i> of the <i>m. gastrocnemius</i>. The origin is fleshy +and arises in company with the <i>m. flexor perforans et perforatus digiti III</i> from +a point on the posterolateral surface of the distal end of the femur between the +point of origin of the <i>pars externa</i> of the <i>m. gastrocnemius</i> and the fibular +condyle. The belly extends approximately one-fourth of the way down the +crus and gives rise to the tendon of insertion which passes distally and superficially +through the posterior edge of the tibial cartilage. The tendon traverses +the posteromedial canal of the hypotarsus (Fig.<a href="#Fig_6"> 6</a>) and continues along the +posterior surface of the tarsometatarsus. Between the first metatarsal and the +base of the second digit the tendon is enclosed by the medial surface of a +sesamoid bone. This tendon then perforates that of the <i>m. flexor perforatus +digiti II</i> at the level of the first phalanx and in turn is perforated by the tendon +of the <i>m. flexor digitorum longus</i> at the proximal end of the second phalanx. +The insertion is on the posterior surface of the second phalanx.</p> + +<p>Action.—Flexes digit II.</p> + +<p>Comparison.—In <i>Passer</i>, <i>Estrilda</i>, <i>Poephila</i>, <i>Hesperiphona</i>, <i>Carpodacus</i>, +<i>Pinicola</i>, <i>Leucosticte</i>, <i>Spinus</i>, and <i>Loxia</i> the proximal portion of this muscle is +more intimately connected with the posterior edge of the <i>m. flexor perforans et +perforatus digiti III</i> than it is in the other species examined.</p> +<p> </p> + +<p><i><b>Musculus flexor perforans et perforatus digiti III</b></i> (Fig.<a href="#Fig_2"> 2</a>).—Long and pinnate, +this muscle lies on the lateral surface of the crus beneath the <i>m. peroneus +longus</i> and <i>pars externa</i> of the <i>m. gastrocnemius</i>. There are two distinct heads. +The origin of the anterior head is fleshy from the proximal edge of the outer +cnemial crest and from the internal edge of the distal end of the patellar tendon. +The posterior head arises by a tendon from the femur in company with the <i>m. +flexor perforans et perforatus digiti II</i>, is connected also with the tendon of +origin of the <i>m. flexor perforatus digiti II</i>, and is loosely attached to the head +of the fibula. Fibers from the belly of the muscle attach throughout its length +<span class="pagenum"><a name="Page_173" id="Page_173">[Pg 173]</a></span> +to the lateral edge of the fibula, and the muscle is tightly fused also with +adjacent muscles. The tendon of insertion is formed approximately one-half the +way down the crus. The tendon perforates the posterior surface of the tibial +cartilage and passes through the posteromedial canal of the hypotarsus (Fig.<a href="#Fig_6"> +6</a>). At the base of the third digit the tendon ensheathes that of the <i>m. flexor +digitorum longus</i> and the two together perforate the tendon of the <i>m. flexor +perforatus digiti III</i>. Immediately distal to this perforation the tendon of the +<i>m. flexor perforans et perforatus digiti III</i> ceases to ensheath that of the <i>m. +flexor digitorum longus</i>. The latter passes beneath that of the former. Near +the distal end of the second phalanx the tendon of the <i>m. flexor digitorum +longus</i> perforates that of the <i>m. flexor perforans et perforatus digiti III</i>. The +latter inserts on the posterior surface of the distal end of the second phalanx and +the proximal end of the third.</p> + +<p>Action.—Flexes digit III.</p> + +<p>Comparison.—In <i>Passer</i>, <i>Estrilda</i>, and <i>Poephila</i>, and in all the cardueline +finches examined the proximal portion of this muscle is more intimately connected +with the anterior edge of the <i>m. flexor perforans et perforatus digiti II</i> +than it is in the other species examined.</p> +<p> </p> + +<p><i><b>Musculus flexor digitorum longus</b></i> (Figs.<a href="#Fig_3"> 3</a>,<a href="#Fig_5"> 5</a>).—This strong, pinnate muscle +is deeply situated along the posterior surfaces of the tibia and fibula. There +are two distinct heads of origin. The lateral head arises by means of fleshy +fibers from the posterior edge of the head of the fibula. The medial head arises +by means of fleshy fibers from the region under the ledgelike external and internal +articular surfaces of the proximal end of the tibia. Neither head has any +connection with the femur in contrast to the condition, described by Hudson +(1937: 46-47) in the crow, <i>Corvus brachyrhynchos</i>, and in the raven, <i>Corvus +corax</i>. Near the point of insertion of the <i>m. biceps femoris</i> the two heads fuse. +The common belly is attached by fleshy fibers to the posterior surface of the +tibia and fibula for two-thirds of the distance down the crus. Near the distal +end of the crus the muscle terminates in a strong tendon which passes deeply +through the tibial cartilage and traverses the anteromedial canal of the hypotarsus +(Fig.<a href="#Fig_6"> 6</a>). About midway down the tarsometatarsus this tendon becomes +ossified. Immediately above the bases of the toes it gives rise to three branches, +one to the posterior surface of each of the foretoes. These branches perforate +the other flexor muscles of the toes as described in the accounts of those muscles +and insert as follows: The branch to digit II inserts on the base of the ungual +phalanx and by a stout, tendinous slip on the distal end of the second phalanx +(Fig.<a href="#Fig_9"> 9</a>). The branch to digit III inserts on the base of the distal end of the +third phalanx and a stronger slip to the distal end of the second or proximal end +of the third. The branch to digit IV inserts on the base of the ungual phalanx, +with one tendinous slip to the distal end of the third phalanx and another to +the distal end of the fourth.</p> + +<p>Action.—Flexes foretoes.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> + +<p><i><b>Musculus flexor hallucis longus</b></i> (Fig.<a href="#Fig_3"> 3</a>).—Situated immediately posterior to +the <i>m. flexor digitorum longus</i>, the belly of this large, pinnate muscle is intimately +connected anteriorly to that of the <i>m. flexor perforatus digiti II</i>. The +<i>m. flexor hallucis longus</i> arises by two heads which are separated by the tendon +of insertion of the <i>m. biceps femoris</i>. The smaller anterior head arises from +<span class="pagenum"><a name="Page_174" id="Page_174">[Pg 174]</a></span> +the same tendon as does the <i>m. flexor perforatus digiti II</i>. The larger posterior +head arises by means of fleshy fibers from the intercondyloid region of the posterior +surface of the femur along with the <i>m. flexor perforatus digiti III</i> and <i>IV</i>. +The two heads join just distal to the point of insertion of the <i>m. biceps femoris</i>. +There is no trace of a tendinous band connecting the two heads as there is in +the crow and in the raven (Hudson, 1937:49). Near the distal end of the +shank the muscle gives rise to a strong tendon which perforates the tibial +cartilage along its lateral edge and passes through the anterolateral canal of +the hypotarsus (Fig.<a href="#Fig_6"> 6</a>). The tendon crosses over to the medial surface of the +tarsometatarsus, passes distally, and perforates the sheathlike tendon of the <i>m. +flexor hallucis brevis</i> between the first metatarsal and the trochlea for digit II. +The tendon continues along the posterior surface of the hallux and has a +double insertion; the main tendon attaches to the base of the ungual phalanx +and a smaller branch inserts on the distal end of the proximal phalanx.</p> + +<p>Action.—Flexes hallux.</p> + +<p>Comparison.—In <i>Vireo</i> this muscle has only the posterior head of origin and +is not connected with the <i>m. flexor perforatus digiti II</i>. The muscle is proportionately +smaller and weaker than in any of the other species studied.</p> +<p> </p> + +<p><i><b>Musculus extensor hallucis longus</b></i> (Fig.<a href="#Fig_4"> 4</a>).—One of the smallest muscles of +the leg, the origin is fleshy from the anteromedial edge of the proximal end of +the tarsometatarsus. The belly is long and slender and terminates distally in +a slender tendon which passes distally along the posterior surfaces of the first +metatarsal and the first digit. The insertion is on the base of the ungual +phalanx. Near the distal end of the proximal phalanx, the tendon passes between +two thick bands of fibro-elastic tissue which insert also on the ungual +phalanx. These bands of tissue function as automatic extensors of the claw.</p> + +<p>Action.—Extends hallux; action must be slight.</p> + +<p>Comparison.—In <i>Vireo</i> this muscle is proportionately larger and better developed +than it is in any of the other species examined.</p> +<p> </p> + +<p><i><b>Musculus flexor hallucis brevis</b></i> (Fig.<a href="#Fig_4"> 4</a>).—This minute muscle has a fleshy +origin from the medial surface of the hypotarsus. The short belly terminates +in a weak, slender tendon which passes down the posteromedial surface of the +tarsometatarsus and into the space between the first metatarsal and the trochlea +for digit II. In this region the tendon envelops the tendon of the <i>m. flexor +hallucis longus</i> and inserts on the distal end of the first metatarsal and on the +proximal end of the first phalanx of the first digit.</p> + +<p>Action.—Flexes hallux; action must be slight.</p> + +<p>Comparison.—The small size of this muscle makes it exceedingly difficult to +study. The muscle is larger in <i>Vireo</i> than in any of the other species examined. +This may be correlated with the smaller size of the <i>m. flexor hallucis longus</i> in +this species. The muscle does not seem to be so well developed in the cardueline +finches as it is in the other species.</p> +<p> </p> + +<p><i><b>Musculus abductor digiti IV</b></i> (Fig.<a href="#Fig_2"> 2</a>).—Extremely small, delicate and difficult +to demonstrate, this muscle arises in a fleshy origin immediately from +underneath the posterior edge of the external cotyla of the tarsometatarsus. The +tendon of insertion is long and slender and inserts along the lateral edge of the +first phalanx of digit IV.</p> + +<p><span class="pagenum"><a name="Page_175" id="Page_175">[Pg 175]</a></span> +Action.—Abducts digit IV.</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> + +<p><i><b>Musculus lumbricalis.</b></i>—Semitendinous throughout its length, this muscle +arises from the ossified tendon of the <i>m. flexor digitorum longus</i> at a point immediately +proximal to the branching of this tendon. The insertion is on the +joint pulleys and capsules at the base of the third and fourth digits.</p> + +<p>Action.—Hudson (1937:57) states that: "Meckel (<i>vide</i> Gadow—1891, p. +204) considered this muscle as serving to draw the joint pulley behind in order +to protect it from pinching during the bending of the toes. It perhaps also +tends to flex the third and fourth digits."</p> + +<p>Comparison.—No significant differences noted among the species studied.</p> +<p> </p> +<p> </p> + +<a name="Discussion_of_the_Myological_Investigations"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span> +<div class="caption2">Discussion of the Myological Investigations</div> + +<p>Simpson (1944:12) and others have emphasized that different +parts of organisms evolve at different rates. Beecher (1951b:275) +in stating that "... the hind limb is very similar in muscle +pattern throughout the Order Passeriformes and seems to have become +relatively static after attaining a high level of general efficiency +..." implies that the muscle pattern of the leg must be one of +long standing and slow change. This concept was emphasized by +Hudson (1937) who found but little variation in muscle pattern +among members of the several families of passerine birds. The concept +is further confirmed by the present investigation. The intricate +patterns of origin and of insertion seem to remain almost the same +throughout the order in spite of adaptive radiation which has occurred.</p> + +<p>Two major differences in patterns of leg-musculature, however, +were found among the species studied, and these differences are +significant since they are consistent between subfamilies. The +muscles involved are the <i>m. obturator externus</i> and the <i>pars interna</i> +of the <i>m. gastrocnemius</i>.</p> + +<p>The <i>m. obturator externus</i> is bipartite, consisting of dorsal and +ventral parts, in the passerine species studied by Hudson (1937) and +in all of the species examined by me except the ploceids and the +cardueline finches. In the ploceids and cardueline finches this +muscle is undivided and resembles in its position, origin, and insertion +only the ventral portion of the muscle found in the other birds +studied. It is difficult to imagine what advantage or disadvantage +might be associated with the bipartite or with the undivided condition. +The action of this muscle is to rotate the femur (right femur +clockwise, left femur counterclockwise), and certainly the greater +mass of the bipartite muscle could lend greater strength to such +action. The possible significance of this is discussed below.</p> + + + +<p><span class="pagenum"><a name="Page_176" id="Page_176">[Pg 176]</a></span></p> + +<div class="caption3 center">List of Abbreviations Used in Figures</div> + +<br /><br /> +Abd. dig. IV <i>M. abductor digiti IV</i><br /> +Acc. <i>M. accessorius semitendinosi</i><br /> +Add. long. <i>M. adductor longus et brevis</i><br /> +Anterolat. can. Anterolateral canal of hypotarsus<br /> +Anteromed. can. Anteromedial canal of hypotarsus<br /> +Bic. fem. <i>M. biceps femoris</i><br /> +Bic. loop Loop for <i>m. biceps femoris</i><br /> +Ext. cot. External cotyla<br /> +Ext. dig. l. <i>M. extensor digitorum longus</i><br /> +Ext. hal. l. <i>M. extensor hallucis longus</i><br /> +Fem. tib. ext. <i>M. femorotibialis externus</i><br /> +Fem. tib. int. <i>M. femorotibialis internus</i><br /> +Fem. tib. med. <i>M. femorotibialis medius</i><br /> +F. dig. l. <i>M. flexor digitorum longus</i><br /> +F. hal. brev. <i>M. flexor hallucis brevis</i><br /> +F. hal. l. <i>M. flexor hallucis longus</i><br /> +F. p. et p. d. II <i>M. flexor perforans et perforatus digiti II</i><br /> +F. p. et p. d. III <i>M. flexor perforans et perforatus digiti III</i><br /> +F. per. d. II <i>M. flexor perforatus digiti II</i><br /> +F. per. d. III <i>M. flexor perforatus digiti III</i><br /> +F. per. d. IV <i>M. flexor perforatus digiti IV</i><br /> +Gas. <i>M. gastrocnemius</i><br /> +Iliacus <i>M. iliacus</i><br /> +Il. tib. <i>M. iliotibialis</i><br /> +Il. troc. ant. <i>M. iliotrochantericus anticus</i><br /> +Il. troc. med. <i>M. iliotrochantericus medius</i><br /> +Il. troc. post. <i>M. iliotrochantericus posticus</i><br /> +Int. cot. Internal cotyla<br /> +Isch. fem. <i>M. ischiofemoralis</i><br /> +Midmed. can. Midmedial canal of hypotarsus<br /> +Obt. ext. <i>M. obturator externus</i><br /> +Obt. int. <i>M. obturator internus</i><br /> +P. ant. <i>Pars anticus</i><br /> +P. ext. <i>Pars externa</i><br /> +P. int. <i>Pars interna</i><br /> +P. med. <i>Pars media</i><br /> +P. post. <i>Pars posticus</i><br /> +Per. brev. <i>M. peroneus brevis</i><br /> +Per. long. <i>M. peroneus longus</i><br /> +Pirif. <i>M. piriformis</i><br /> +Plan. <i>M. plantaris</i><br /> +Posterolat. can. Posterolateral canal of hypotarsus<br /> +Posteromed. can. Posteromedial canal of hypotarsus<br /> +Sar. <i>M. sartorius</i><br /> +Semim. <i>M. semimembranosus</i><br /> +Semit. <i>M. semitendinosus</i><br /> +Tib. ant. <i>M. tibialis anticus</i><br /> +Tib. cart. Tibial cartilage<br /> +<p> </p> +<p> </p> + +<a name="Fig_1"></a> +<p><span class="pagenum"><a name="Page_177" id="Page_177">[Pg 177]</a></span></p> +<div class="center"> + <img src="images/fig_1.png" width="378" height="596" title="Superficial Leg Muscles" alt="Superficial Leg Muscles" /><br /><br /> + <p><span class="smcap">Fig. 1.</span> <i>Pipilo erythrophthalmus.</i> Lateral view of the superficial muscles of the left leg, × 1.5.</p> +</div> +<p> </p> +<p> </p> + +<a name="Fig_2"></a> +<p><span class="pagenum"><a name="Page_178" id="Page_178">[Pg 178]</a></span></p> +<div class="center"> + <img src="images/fig_2.png" width="417" height="600" title="Deeper Leg Muscles" alt="Deeper Leg Muscles" /><br /><br /> + <p><span class="smcap">Fig. 2.</span> <i>Pipilo erythrophthalmus.</i> Lateral view of the left leg showing a deeper set of muscles. The superficial muscles <i>iliotibialis</i>, <i>sartorius</i>, <i>gastrocnemius</i> and <i>peroneus longus</i> have been removed, × 1.5.</p> +</div> +<p> </p> +<p> </p> + + +<a name="Fig_3"></a> +<p><span class="pagenum"><a name="Page_179" id="Page_179">[Pg 179]</a></span></p> +<div class="center"> + <img src="images/fig_3.png" width="404" height="600" title="Still Deeper Leg Muscles" alt="Still Deeper Leg Muscles" /><br /><br /> + <p><span class="smcap">Fig. 3.</span> <i>Pipilo erythrophthalmus.</i> Lateral view of the left leg showing the still deeper muscles. In addition to those listed for figure<a href="#Fig_2"> 2</a>, the following muscles have been wholly or partly removed: <i>iliotrochantericus posticus</i>, <i>femorotibialis externus</i>, <i>femorotibialis medius</i>, <i>biceps femoris</i>, <i>semitendinosus</i>, <i>tibialis anticus</i>, <i>flexor perforans et perforatus digiti II</i>, and <i>flexor perforans et perforatus digiti III</i>, × 1.5.</p> +</div> +<p> </p> +<p> </p> + +<a name="Fig_4"></a> +<p><span class="pagenum"><a name="Page_180" id="Page_180">[Pg 180]</a></span></p> +<div class="center"> + <img src="images/fig_4.png" width="418" height="600" title="Medial View Superficial Leg Muscles" alt="Medial View Superficial Leg Muscles" /><br /><br /> + <p><span class="smcap">Fig. 4.</span> <i>Pipilo erythrophthalmus.</i> Medial view of the superficial muscles of the left leg, × 1.5.</p> +</div> +<p> </p> +<p> </p> + +<a name="Fig_5"></a> +<p><span class="pagenum"><a name="Page_181" id="Page_181">[Pg 181]</a></span></p> +<div class="center"> + <img src="images/fig_5.png" width="397" height="600" title="Medial View Deeper Leg Muscles" alt="Medial View Deeper Leg Muscles" /><br /><br /> + <p><span class="smcap">Fig. 5.</span> <i>Pipilo erythrophthalmus.</i> Medial view of the left leg showing a deeper set of muscles than those seen in figure<a href="#Fig_4"> 4</a>. The following superficial muscles have been removed: <i>iliotibialis</i>, <i>sartorius</i>, <i>femorotibialis internus</i>, <i>obturator internus</i>, <i>adductor longus (pars posticus)</i>, <i>gastrocnemius</i>, and <i>peroneus longus</i>, × 1.5.</p> +</div> +<p> </p> +<p> </p> + +<a name="Fig_6"></a> +<a name="Fig_7"></a> +<a name="Fig_8"></a> +<a name="Fig_9"></a> +<p><span class="pagenum"><a name="Page_182" id="Page_182">[Pg 182]</a></span></p> +<table width="100%" summary="Figs. 6-9"> +<tr><td><img src="images/fig_6.png" width="285" height="85" title="Fig. 6" alt="Fig. 6" /><br /><div class="smaller center">Figure 6</div></td><td width="76%"> </td></tr> +<tr><td rowspan=2 class="center"><p> </p><img src="images/fig_8.png" width="206" height="474" title="Fig. 8" alt="Fig. 8" /><br /><span class="smaller">Figure 8</span></td> +<td class="center"><img src="images/fig_7.png" width="225" height="136" title="Fig. 7" alt="Fig. 7" /><br /><span class="smaller">Figure 7</span></td></tr> +<tr><td class="center"><img src="images/fig_9.png" width="181" height="313" title="Fig. 9" alt="Fig. 9" /><br /><span class="smaller">Figure 9</span></td></tr> +</table> +<p> </p> +<p> </p> + +<p><span class="smcap">Fig. 6.</span> <i>Pipilo erythrophthalmus.</i> Proximal end of left tarsometatarsus and the hypotarsus, × 4.</p> + +<p><span class="smcap">Fig. 7.</span> <i>Pipilo erythrophthalmus.</i> Lateral view of proximal end of left femur and a portion of the pelvis, × 3.5.</p> + +<p><span class="smcap">Fig. 8.</span> <i>Pipilo erythrophthalmus.</i> Upper surfaces of the phalanges of the foretoes of the left foot showing insertions of the <i>M. extensor digitorum longus</i>, × 3.</p> + +<p><span class="smcap">Fig. 9.</span> <i>Pipilo erythrophthalmus.</i> Medial view of the second digit of the left foot, showing insertions of the flexor muscles, × 3.</p> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_183" id="Page_183">[Pg 183]</a></span> +The division of the <i>pars interna</i> of the <i>m. gastrocnemius</i> into +anterior and posterior parts has not been reported by previous +authors yet the division is quite distinct in those birds in which it +occurs. Hudson (1937:36) points out that in some non-passerine +birds the <i>pars interna</i> is double, but that in these species the <i>m. +semimembranosus</i> inserts between the two parts. This is not the +condition in those species studied by me. Only the ploceids and the +cardueline finches in the present investigation fail to show such a +division. The undivided muscle in these birds resembles, in its +origin and position, the posterior portion of the muscle found in +those species showing the bipartite condition. The greater mass +of the bipartite muscle probably makes possible a stronger extension +of the tarsometatarsus.</p> + +<p>Thus, the divided or undivided conditions of the <i>m. obturator +externus</i> and the <i>pars interna</i> of the <i>m. gastrocnemius</i> seem to be +correlated with the degrees of strength of certain movements of the +leg. It is conceivable that these differences in structure are correlated +with the manner in which food is obtained, the birds having +the bipartite muscles being those which spend the most time on the +ground searching and scratching for seeds and other sorts of food. +Yet, in <i>Leucosticte</i>, a cardueline, and in <i>Calcarius</i>, an emberizine, +whose foraging habits are rather similar, the structure is unlike. +<i>Leucosticte</i> does resemble the emberizines and also <i>Piranga</i> and +<i>Spzia</i> in the extension of a band of muscle fibers from the <i>pars +interna</i> of the <i>m. gastrocnemius</i> around the front of the knee. A +band of muscle fibers of this sort strengthens the knee joint and +gives still more strength to the <i>pars interna</i>. This condition has +been reported in a number of birds by Hudson (1937) and is, in all +probability, an adaptation for greater strength of certain leg movements. +The development of this band in <i>Leucosticte</i> seems to +parallel that in the other birds studied and does not indicate relationship, +since in <i>Leucosticte</i> this band arises from the undivided +muscle which (as stated above) resembles only the posterior portion +of the bipartite muscle described for the other birds. In the latter, +the muscular band arises from the anterior part of the muscle.</p> + +<p>Minor differences in muscle pattern, like those already mentioned, +are consistent also between subfamilies, but correlation of these +minor differences with function is difficult. There is the implication, +however, that in all the groups except the carduelines and +ploceids, the emphasis is on greater strength and mobility of the leg. +In the carduelines that were studied the origin of the <i>m. sartorius</i> +<span class="pagenum"><a name="Page_184" id="Page_184">[Pg 184]</a></span> +does not extend so far craniad as in the other species. In the latter, +at least half of the origin is from the last one or two free dorsal +vertebrae; in the carduelines no more than one third of the origin is +anterior to the ilium. It is conceivable that the more craniad the +origin, the stronger the forward movement of the thigh would be.</p> + +<p>In <i>Passer</i>, <i>Estrilda</i> and <i>Poephila</i>, and in all the cardueline finches +examined, the bellies of the <i>m. flexor perforans et perforatus digiti +II</i> and the <i>m. flexor perforans et perforatus digiti III</i> are more intimately +connected than they are in the other species studied. Thus, +the amount of independent action of these muscles in <i>Passer</i>, in +the estrildines, and in the carduelines probably is reduced.</p> + +<p>In <i>Passer</i>, the estrildines, and the carduelines the edges of the +sheathlike tendon of insertion of the <i>m. perforatus digiti III</i> are +thickened; as a result the insertion appears superficially to be double +but closer examination reveals that there is a fascia stretched between +the thickened edges. In the other species examined, the +insertion is sheathlike throughout and there are no thick areas. I +cannot explain this on the basis of function. The difference, however, +is obvious and constant.</p> + +<p>Aside from the differences noted above, there were variations of +muscle pattern that seem to be significant only in <i>Vireo olivaceus</i>. +In this species the central, aponeurotic portion of the <i>m. iliotibialis</i> +is absent. The origin of the <i>m. adductor longus et brevis</i> is from +the dorsal edge of the ischiopubic fenestra and not from the membrane +covering this fenestra. The origin of the <i>pars posticus</i> of this +muscle, furthermore, is fleshy and not tendinous as it is in the other +species. The <i>m. flexor perforatus digiti II</i> is larger and more deeply +situated in <i>Vireo</i> and has, furthermore, no connection with the <i>m. +flexor hallucis longus</i>. The latter muscle is smaller and weaker than +in any of the other species and has only one (the posterior) head +of origin. The <i>m. flexor hallucis brevis</i>, on the contrary, is larger +than in the other birds, compensating, probably, for the small <i>m. +flexor hallucis longus</i>. In those differences, however, which separate +the carduelines and ploceids from the other birds studied, <i>Vireo</i> +resembles, in every instance, the richmondenines, emberizines, tanagers, +warblers, and blackbirds.</p> + +<p>On the basis of differences in leg-musculature the species which +are now included in the Family Fringillidae may be separated into +two groups. One group includes the richmondenines and the emberizines; +the other, the carduelines. The muscle patterns of the +legs of the birds of the first group are indistinguishable from those +of <i>Seiurus</i>, <i>Icterus</i>, <i>Molothrus</i>, and <i>Piranga</i>, and except for the differences +<span class="pagenum"><a name="Page_185" id="Page_185">[Pg 185]</a></span> +noted are similar to those in <i>Vireo</i>. The carduelines, on +the other hand, are similar in every point of leg-musculature to the +ploceids which were studied. Thus, the heterogeneity of the Family +Fringillidae, as now recognized, is emphasized by differences in the +muscle patterns of the leg.</p> +<p> </p> +<p> </p> + +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span> +<a name="Comparative_Serology" id="Comparative_Serology"></a> +<div class="caption2h smcap">Comparative Serology</div> +<p> </p> + +<a name="General_Statement_2"></a> +<div class="caption2">General Statement</div> + +<p>The application of serological techniques to the problems of +animal relationships has been attempted with varying degrees of +success over a period of approximately fifty years. Few of the +earlier studies were of a quantitative nature, but within the past +decade, satisfactory quantitative serological techniques have been +developed whereby taxonomic relationships may be estimated. The +usefulness of comparative serology in taxonomy has been demonstrated +in investigations of many groups wherein results obtained +have, in most instances, been compatible with the results obtained +by more conventional methods, such as comparative morphology. +As Boyden (1942:141) stated, "comparative serology ... is no +simple guide to animal relationship." However, the objectiveness +of its methods, the fact that it has its basis in the comparisons of +biochemical systems which seem to be relatively slow to change in +response to external environmental influences, and the fact that the +results are of quantitative nature favor, where possible, the inclusion +of data from comparative serology along with that from more +conventional sources when an attempt is made to determine the +relationships of groups of animals.</p> + +<p>The application of serological methods in ornithology has not +been extensive. Irwin and Cole (1936) and Cumley and Irwin +(1941, 1944) used two species of doves and their hybrids and +demonstrated that a distinction between the red cells of these birds +could be made by use of immunological methods involving the agglutinin +reaction. McGibbon (1945) was able to distinguish the +red cells of interspecific hybrids in ducks by similar methods. Irwin +(1953) used similar techniques in his study of the evolutionary +patterns of some antigenic substances of the blood cells of birds of +the Family Columbidae. Sasaki (1928) demonstrated the usefulness +of the precipitin technique in distinguishing species of ducks +and their hybrids. This technique was used successfully also by +DeFalco (1942) and by Martin and Leone (1952). Working with +groups of known relationships, these investigators showed that the +"accepted" systematic positions of certain birds were confirmed by +<span class="pagenum"><a name="Page_186" id="Page_186">[Pg 186]</a></span> +serological procedures. The precipitin reaction, however, has never +been applied to actual problems in avian taxonomy prior to the +present study.</p> +<p> </p> + +<a name="Preparation_of_Antigens"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span> +<div class="caption2">Preparation of Antigens</div> + +<p>Although most previous work in comparative serology in which precipitin +tests were used has involved the use of whole sera as antigens, Martin and +Leone (1952) indicated that tissue extracts are satisfactory as antigens and +that serological differentiation can be obtained with these extracts and the +antisera to them. I decided, therefore, to use such extracts in these investigations, +since the small sizes of the birds to be tested made it impracticable to +obtain enough whole sera.</p> + +<p>Most of the birds used were obtained by shooting, but a few were trapped +and the exotic species were purchased alive from a pet dealer. When a bird +was killed, the entire digestive tract was carefully removed to prevent the +escape of digestive enzymes into the tissues and to prevent putrefaction by +action of intestinal bacteria. As soon as possible (and within three hours in +every instance) the bird was skinned, the head, wings, and legs were removed, +and the body was frozen. Each specimen, consisting of trunk, heart, lungs, +and kidneys, was wrapped separately and carefully in aluminum foil to prevent +dehydration of the tissues. The specimens were kept frozen until the time +when the extracts were made.</p> + +<p>When an extract was to be prepared, the specimen was allowed to thaw but +not to become warm. In the cold room with the temperature of all equipment +and reagents at 2°C., the specimen was placed in a Waring blender with 0.9 +per cent aqueous solution of NaCl buffered with M/150 K<sub>2</sub>HPO<sub>4</sub> and M/150 +Na<sub>2</sub>HPO<sub>4</sub> to a pH of 7.0. The amount of reagent used was 75 ml. of saline for +each gram of tissue to be extracted. The tissues were minced in the blender, +allowed to stand at 2°C. for 72 hours, and the tissue residues removed by +centrifugation in a refrigerated centrifuge. Formalin was added to a portion of +the supernatant in the amount necessary to make the final dilution 0.4 per cent. +This formolization was found to be necessary to inhibit the action of autolytic +enzymes over the period of time required to complete the investigations. The +effects of formolization on the antigenicity and reactivity of proteins are discussed +later. It was necessary to sterilize and clarify the "native" (unformolized) +extracts; this was done by filtration through a Seitz filter. These "native" +substances were used only in the early stages of the investigation (see below). +The filtrate was bottled and stored at 2°C. In the early stages of this investigation +clarification of the formolized extract was accomplished by the same +sort of filtration. It was determined, however, that centrifugation in a refrigerated +centrifuge at high speeds (17,000g) served the same purpose and +was quicker. The formolized extracts were bottled and also stored at 2°C. +(although refrigerated storage of the formolized extracts does not seem necessary). +For each extract the amount of protein present was determined colorimetrically +by the method of Greenberg (1929) with a Leitz Photrometer.</p> + +<p>Species for which extracts were prepared and the protein values of the +extracts are listed in Table<a href="#Table_1"> 1</a>. Extracts of some species were used throughout +most of the experiment; extracts of others were used only when needed for +purposes of comparison.</p> +<p> </p> +<p> </p> + +<a name="Table_1"></a> +<p><span class="pagenum"><a name="Page_187" id="Page_187">[Pg 187]</a></span></p> + +<p class="smcap">Table 1.—Species from Which Extracts Were Prepared and Injection +Schedules for Extracts Against Which Antisera Were Produced</p> + +<table width="100%" cellpadding=4 summary="Species and Extract Injection Data"> +<tr><td colspan=3 class="bt white"><img src="images/bar_single.png" width="1" height="5" title="bar" alt="bar" /></td></tr> +<tr><td class="bt bb"><span class="smcap">Species</span></td><td class="bt bl bb">Protein, gms. per 100 ml</td><td class="bt bl bb">Injection schedules for<br />production of antisera</td></tr> +<tr><td class="bb"><i>Myiarchus crinitus</i> (Linnaeus)</td><td class="bl bb center">0.65</td><td class="bl bb">Series 1: Intravenous, 0.5, 1.0, 2.0, and 4.0 ml.</td></tr> +<tr><td class="bb"><i>Passer domesticus</i></td><td class="bl bb center">1.40</td><td class="bl bb"><div class="ind2em">Series 1: Subcutaneous, 0.5, 1.0, 2.0, and 4.0 ml.</div></td></tr> +<tr><td class="bb"><i>Estrilda amandava</i></td><td class="bl bb center">0.45</td><td class="bl bb"><div class="ind2em"><a name="FNanchor_A_1" id="FNanchor_A_1"></a><a href="#Footnote_A_1" class="fnanchor">[A]</a>Series 1: Intravenous, 0.5, 1.0, 2.0, and 4.0 ml.</div><div class="ind2em"><a href="#Footnote_A_1" class="fnanchor">[A]</a>Series 2: Subcutaneous, 0.5, 1.0, and 2.0 ml.</div><div class="ind2em">Intraperitoneal, 8.0 ml.</div></td></tr> +<tr><td class="bb"><i>Poephila guttata</i></td><td class="bb bl center">0.56</td><td class="bb bl"><a href="#Footnote_A_1" class="fnanchor">[A]</a>Same as for <i>Estrilda</i>.</td></tr> +<tr><td class="bb"><i>Molothrus ater</i></td><td class="bl bb center">0.65</td><td class="bl bb"><div class="ind2em">Series 1: <a href="#typos">Intravenous</a> and subcutaneous, respectively, 0.5 and 0.5 ml., 1.0 and 1.0 ml., 3.0 and 1.0 ml., 5.0 and 3.0 ml.</div><div class="ind2em">Series 2: Subcutaneous, 0.5, 1.0, 2.0 and 4.0 ml.</div></td></tr> +<tr><td class="bb"><i>Piranga rubra</i></td><td class="bb bl center">0.50</td><td class="bb bl">Same as for <i>Molothrus</i>.</td></tr> +<tr><td class="bb"><i>Richmondena cardinalis</i></td><td class="bb bl center">0.70</td><td class="bb bl"><a href="#Footnote_A_1" class="fnanchor">[A]</a>Same as for <i>Estrilda</i>.</td></tr> +<tr><td class="bb"><i>Richmondena cardinalis</i></td><td class="bb bl center">0.60</td><td class="bb bl">Same as for <i>Spinus</i>.</td></tr> +<tr><td class="bb"><i>Passerina cyanea</i></td><td class="bb bl center">0.45</td><td class="bb bl">Antiserum not prepared.</td></tr> +<tr><td class="bb"><i>Spiza americana</i></td><td class="bb bl center">0.70</td><td class="bb bl">Same as for <i>Molothrus</i>.</td></tr> +<tr><td class="bb"><i>Carpodacus purpureus</i></td><td class="bb bl center">0.50</td><td class="bb bl">Antiserum not prepared.</td></tr> +<tr><td class="bb"><i>Spinus tristis</i></td><td class="bl bb center">0.49</td><td class="bl bb"><div class="ind2em">Series 1: Intravenous, 0.5, 1.0, 2.0, and 4.0 ml.</div><div class="ind2em">Series 2: Intravenous, 0.5, 1.0, 2.0, and 4.0 ml.</div><div class="ind2em">Series 3: Subcutaneous, 0.5, 1.0, 2.0, and 4.0 ml.</div></td></tr> +<tr><td class="bb"><i>Pipilo erythrophthalmus</i></td><td class="bb bl center">0.92</td><td class="bb bl">Antiserum not prepared.</td></tr> +<tr><td class="bb"><i>Junco hyemalis</i></td><td class="bb bl center">0.56</td><td class="bb bl">Same as for <i>Spinus</i>.</td></tr> +<tr><td class="bb"><i>Spizella arborea</i></td><td class="bb bl center">0.48</td><td class="bb bl">Same as for <i>Spinus</i>.</td></tr> +<tr><td class="bb"><i>Zonotrichia querula</i></td><td class="bb bl center">0.48</td><td class="bb bl">Same as for <i>Spinus</i>.</td></tr> +<tr><td class="bb"><i>Zonotrichia albicollis (Gmelin)</i></td><td class="bl bb center">0.92</td><td class="bl bb">Antiserum not prepared.</td></tr> +</table> + +<div class="footnote"><a name="Footnote_A_1" id="Footnote_A_1"></a><p><a href="#FNanchor_A_1"><span class="label">[A]</span></a> Antiserum prepared against formolized antigen.</p></div> + +<p> </p> +<p> </p> + +<a name="Preparation_of_Antisera"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span><br> +<p><span class="pagenum"><a name="Page_188" id="Page_188">[Pg 188]</a></span></p> +<div class="caption2">Preparation of Antisera</div> + +<p>All antisera were produced in rabbits (laboratory stock of <i>Oryctolagus +cuniculus</i>). Three methods of injection of antigen were used in various combinations: +intravenous, subcutaneous, and intraperitoneal. Injection schedules +used in the production of each antiserum are listed in Table<a href="#Table_1"> 1</a>. Both formolized +and "native" antigens were used. Each rabbit received one or more series +of four injections, each injection being administered on alternate days and doubling +in amount: 0.5 ml., 1.0 ml., 2.0 ml., and 4.0 ml. In all but two instances +more than one series of injections was necessary to produce a useful antiserum. +More than two series, however, resulted in little or no improvement of the +reactivity of the antiserum.</p> + +<p>The injection-series were separated by intervals of eight days. On the eighth +day after the last injection of each series, 10 ml. of blood were withdrawn from +the main artery of the ear of the rabbit, and the antiserum was used in a +homologous precipitin test to determine its usefulness. If the antiserum contained +sufficient amounts of antibodies to conduct the projected tests, the rabbit +was completely exsanguinated by cardiac puncture, by using an 18-gauge needle +and a 50 ml. syringe. The whole blood was placed in clean test tubes and +allowed to clot. It was allowed to stand at 2°C. for 12 to 18 hours so that +most of the serum would be expressed from the clot. The serum was then +decanted, centrifuged to remove all blood cells, sterilized in a Seitz filter, +bottled in sterile vials, and stored at 2°C. until used.</p> +<p> </p> + +<a name="Methods_of_Serological_Testing"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span><br> +<div class="caption2">Methods of Serological Testing</div> + +<p>The precipitin reaction is the most successful of the serological techniques +thus far devised for systematic comparisons. The reaction occurs because +antigenic substances introduced into the body of an animal cause the formation +of antibodies which precipitate antigens when the two are mixed. The antisera +which are produced show quantitative specificities in their actions; therefore, +when an antiserum containing precipitins is mixed with each of several antigens, +the reaction involving the homologous antigen (that used in the production of +the antiserum) is greater than those reactions involving the heterologous antigens +(antigens other than those used in the production of the antiserum). +Furthermore, the magnitudes of the reactions between the antiserum and the +heterologous antigens vary according to the degrees of similarity of these +antigens to the homologous one.</p> + +<p>The method of precipitin testing follows that outlined by Leone (1949). The +Libby (1938) Photronreflectometer was used to measure the turbidities developed +by the interaction of antigen and antiserum. With this instrument +parallel rays of light are passed through the turbid systems being measured. +Light rays are reflected from the suspended particles to the sensitive plate of a +photoelectric cell; this generates a current of electricity which causes a deflection +on a galvanometer. The deflection is proportional to the amount of turbidity +developed and readings may be taken directly from the scale of the instrument.</p> + +<p>The reaction-cells of the photronreflectometer are designed to operate with +a volume of 2 ml.; therefore, this volume was used in all testing. In every +series of tests the amount of antiserum was held constant and the amount of +antigen was varied. The volume for each antigen dilution was always 1.7 ml., +and to this was added 0.3 ml. of antiserum to make up a volume of 2 ml.</p> +<p> </p> +<p> </p> + +<a name="Table_2"></a> +<p><span class="pagenum"><a name="Page_189" id="Page_189">[Pg 189]</a></span></p> +<p><span class="smcap">Table 2.</span>—Percentage values obtained from analyses of precipitin reactions. +Numerals represent relative amounts of reaction between antigens and antisera. +Homologous reactions are arbitrarily valued as 100 per cent, and heterologous +reactions are expressed accordingly. <i>Comparisons are meaningful only if made +within each horizontal row of values.</i></p> + +<table width="100%" cellpadding=4 class="center" summary="Species and Extract Injection Data"> +<tr><td colspan=9 class="bt"><img src="images/bar_single.png" width="1" height="5" title="bar" alt="bar" /></td></tr> +<tr><td rowspan=2 class="bt bb smcap">Antigens</td><td colspan=8 class="bt bl bb">ANTISERA</td></tr> +<tr><td class="bl bb vbot"><img src="images/t2n1.png" width="23" height="138" title="A" alt="A " /></td><td class="bl bb vbot"> <img src="images/t2n2.png" width="20" height="125" title="B" alt="B " /></td><td class="bl bb vbot"><img src="images/t2n3.png" width="18" height="111" title="C" alt="C " /></td><td class="bl bb vbot"><img src="images/t2n4.png" width="20" height="170" title="D" alt="D " /></td><td class="bl bb vbot"><img src="images/t2n5.png" width="19" height="120" title="E" alt="E " /></td><td class="bl bb vbot"><img src="images/t2n6.png" width="19" height="98" title="F" alt="F " /></td><td class="bl bb vbot"><img src="images/t2n7.png" width="19" height="112" title="G" alt="G " /></td><td class="bl bb vbot"><img src="images/t2n8.png" width="20" height="142" title="H" alt="H " /></td></tr> +<tr><td class="text_lf bb"><i>Passer domesticus</i></td><td class="bl bb">75</td><td class="bl bb">74</td><td class="bl bb">73</td><td class="bl bb">66</td><td class="bl bb">81</td><td class="bl bb">72</td><td class="bl bb">...</td><td class="bl bb">81</td></tr> +<tr><td class="text_lf bb"><i>Estrilda amandava</i></td><td class="bl bb">100</td><td class="bl bb">88</td><td class="bl bb">75</td><td class="bl bb">...</td><td class="bl bb">79</td><td class="bl bb">72</td><td class="bl bb">53</td><td class="bl bb"> ...</td></tr> +<tr><td class="text_lf bb"><i>Poephila guttata</i></td><td class="bl bb">95</td><td class="bl bb">100</td><td class="bl bb">77</td><td class="bl bb">67</td><td class="bl bb">87</td><td class="bl bb">81</td><td class="bl bb">...</td><td class="bl bb"> ...</td></tr> +<tr><td class="text_lf bb"><i>Molothrus ater</i></td><td class="bl bb">66</td><td class="bl bb">54</td><td class="bl bb">69</td><td class="bl bb">65</td><td class="bl bb">86</td><td class="bl bb">75</td><td class="bl bb">69</td><td class="bl bb">75</td></tr> +<tr><td class="text_lf bb"><i>Piranga rubra</i></td><td class="bl bb">...</td><td class="bl bb">...</td><td class="bl bb">100</td><td class="bl bb">...</td><td class="bl bb">...</td><td class="bl bb">...</td><td class="bl bb">...</td><td class="bl bb">89</td></tr> +<tr><td class="text_lf bb"><i>Richmondena cardinalis</i> </td><td class="bl bb">75</td><td class="bl bb">80</td><td class="bl bb">91</td><td class="bl bb">100</td><td class="bl bb">98</td><td class="bl bb">65</td><td class="bl bb">88</td><td class="bl bb">91</td></tr> +<tr><td class="text_lf bb"><i>Spiza americana</i></td><td class="bl bb">65</td><td class="bl bb">68</td><td class="bl bb">...</td><td class="bl bb">71</td><td class="bl bb">100</td><td class="bl bb">64</td><td class="bl bb">67</td><td class="bl bb">80</td></tr> +<tr><td class="text_lf bb"><i>Carpodacus purpureus</i></td><td class="bl bb">70</td><td class="bl bb">71</td><td class="bl bb">71</td><td class="bl bb">61</td><td class="bl bb">89</td><td class="bl bb">93</td><td class="bl bb">53</td><td class="bl bb">70</td></tr> +<tr><td class="text_lf bb"><i>Spinus tristis</i></td><td class="bl bb">72</td><td class="bl bb">74</td><td class="bl bb">73</td><td class="bl bb">60</td><td class="bl bb">89</td><td class="bl bb">100</td><td class="bl bb">60</td><td class="bl bb">...</td></tr> +<tr><td class="text_lf bb"><i>Junco hyemalis</i></td><td class="bl bb">64</td><td class="bl bb">56</td><td class="bl bb">74</td><td class="bl bb">65</td><td class="bl bb">87</td><td class="bl bb">68</td><td class="bl bb">100</td><td class="bl bb">...</td></tr> +<tr><td class="text_lf bb"><i>Zonotrichia querula</i></td><td class="bl bb">65</td><td class="bl bb">71</td><td class="bl bb">...</td><td class="bl bb">67</td><td class="bl bb">89</td><td class="bl bb">75</td><td class="bl bb">...</td><td class="bl bb">100</td></tr> +</table> +<p> </p> +<p> </p> + +<p>Antigens were diluted with 0.9 per cent phosphate-buffered saline solution. +Tests were run in standard Kolmer test-tube racks, each test consisting of 12 +tubes. Each dilution was made on the basis of the known protein concentration +of the antigen. The first tube contained an initial dilution of 1 part protein +in 250 parts saline and each successive tube contained a protein dilution one-half +the concentration of the preceding tube, ranging up to 1:512,000. Saline +controls, antiserum controls, and antigen controls were maintained with each +test to determine the turbidities inherent in these solutions. These control-turbidities +were deducted from the total turbidity developed in each reaction-tube, +the resultant turbidity then being considered as that which was caused +by the interaction of antigens and antibodies. The turbidities were allowed to +develop over a 24-hour period. In the early stages of this investigation the +reactions were allowed to take place at 2°C. in order to inhibit bacterial growth. +<span class="pagenum"><a name="Page_190" id="Page_190">[Pg 190]</a></span> +Later tests were carried out at room temperatures, and bacterial growth was +prevented by the addition to each tube of 'Merthiolate' in a final dilution of +1:10,000.</p> +<p> </p> + +<a name="Experimental_Data"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span> +<div class="caption2">Experimental Data</div> + +<p>Corrected values for the turbidities obtained were plotted with the turbidity +values on the ordinate and the antigen dilutions on the abscissa. The homologous +reaction was the standard of reference for all other test reactions with the +same antiserum. By summing the plotted turbidity readings, numerical values +are obtained which are indices serving to characterize the curves. Such values +were converted to percentage values, that of the homologous reaction being +considered 100 per cent. These values, plus the curves, provide the data by +means of which the proteins of the birds may be compared. Plots representative +of the precipitin curves are presented in Figs.<a href="#Fig_10"> 10</a> to<a href="#Fig_21"> 21</a>. For convenience +each plot represents only several of the 10 curves obtained with each antiserum.</p> + +<p>A summary of the serological relationships of the birds involved in the +precipitin tests is presented in Table<a href="#Table_2"> 2</a>, in which percentage values are presented. +Since the techniques involved in testing were greatly improved as the +investigation proceeded, the summary is based solely on those tests run in the +later stages of the investigation. For reasons which will become apparent in +later discussion, it should be emphasized that in Table<a href="#Table_2"> 2</a> comparisons may be +made only within each horizontal row of values.</p> +<p> </p> + +<a name="Discussion_of_the_Serological_Investigations"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span><br> +<div class="caption2">Discussion of the Serological Investigations</div> + +<p>One of the problems met early in this investigation was instability +of the proteins in the extracts that were prepared. Extracts in which +no attempt was made to inactivate the enzymes present proved unsatisfactory. +It was necessary to maintain the temperature of the +"native" antigens at 2°C, and all work with such antigens had to be +performed at this temperature. This arrangement was inconvenient; +furthermore, inactivation of the enzymes was not complete even at +this low temperature, and some denaturation of the proteins took +place as evidenced by the gradual appearance of insoluble precipitates +in the stored vials.</p> + +<p>The preservatives, 'Merthiolate' and formalin, were used in an +attempt to inhibit the autolytic action of the enzymes present. +Formalin, when added to make a final dilution of 0.4 per cent, +proved to be the more satisfactory of the two preservatives and was +used throughout most of the work. Formalin caused slight denaturation +of some of the proteins, but this effect was complete +within a few hours, after which any denatured material was removed +by filtration or centrifugation. The proteins remaining in +solution were stable over the period necessary to complete the investigations.</p> + +<p>The addition of formalin reduces the reactivity of the extracts +when they are tested with antisera prepared against "native" antigens +<span class="pagenum"><a name="Page_191" id="Page_191">[Pg 191]</a></span> +and causes changes in the nature of the precipitin curves. This +effect has been pointed out by Horsfall (1934) and by Leone (1953) +in their work on the effects of formaldehyde on serum proteins. +Their data indicate, however, that even though changes in the +immunological characteristics of proteins are brought about by formolization, +the proteins retain enough of their specific chemical characteristics +to allow consistent differentiation of species by immunological +methods. In the tests which I performed, the relative positions +of the precipitin curves, whether native or formolized extracts +were involved, remained unchanged (Figs.<a href="#Fig_10"> 10</a>,<a href="#Fig_11"> 11</a>). <i>All data used +in interpretation of the serological relationships were obtained from +tests in which formolized antigens of equivalent age were used.</i></p> + +<p>Only three antisera were produced against formolized antigens, +all others being produced against "native" extracts. The formolized +antigens seemed to have a greater antigenicity, in most instances, +than did those which were unformolized, and precipitin reactions +involving antisera produced against formolized antigens developed +higher turbidities. The antisera produced against formolized antigens +were equal to but no better than those prepared against "native" +extracts in separating the birds tested (Figs.<a href="#Fig_12"> 12</a>,<a href="#Fig_13"> 13</a>).</p> + +<p>The rabbit is a variable to be considered in serological tests. Two +rabbits exposed to the same antigen, under the same conditions, may +produce antisera which differ greatly in their capacities to distinguish +different antigens. It is logical to assume, therefore, that +two rabbits exposed to different antigens may produce antisera +which also differ in this respect. This explains the unequal values +of reciprocal tests shown in Table<a href="#Table_2"> 2</a>. Thus, in the test involving +the antiserum to the extracts of <i>Richmondena</i>, a value of 71 per cent +was obtained for <i>Spiza</i> antigen, whereas in the test involving anti-<i>Spiza</i> +serum, a value of 98 per cent was obtained for <i>Richmondena</i> +antigen. In Table<a href="#Table_2"> 2</a>, therefore, comparisons may be made only +among values for the proteins of birds tested with the same antiserum.</p> + +<p>Since the amount of any one antiserum is limited, there is, of necessity, +a limit as to the number of birds used in a series of serological +tests. Therefore, although the results reveal the actual serological +relationships of the individual species, interpretation of the relationships +of the taxonomic groups must be undertaken with the realization +that such an interpretation is based on tests involving relatively +few species of each group. It is reasonable to assume, however, +that a species which has been placed in a group on the basis of resemblances +other than serological resemblance would show greater +<span class="pagenum"><a name="Page_192" id="Page_192">[Pg 192]</a></span> +serological correspondence to other members of that group than it +would to members of other groups. Specifically, in the Fringillidae +and their allies, there seems to be little reason to doubt that genera, +and even subfamilies, are natural groups. This is illustrated in tests +involving closely related genera: <i>Richmondena</i> and <i>Spiza</i> (Figs. +<a href="#Fig_14"> 14</a>,<a href="#Fig_15"> 15</a>,<a href="#Fig_18"> 18</a>), <i>Estrilda</i> and <i>Poephila</i> (Fig.<a href="#Fig_21"> 21</a>), <i>Spinus</i> and <i>Carpodacus</i> +(Figs.<a href="#Fig_12"> 12</a>,<a href="#Fig_17"> 17</a>,<a href="#Fig_19"> 19</a>,<a href="#Fig_20"> 20</a>). In each of these tests the pairs of genera +mentioned show greater serological correspondence to each other +than they do to other kinds involved. This point is illustrated further +by a test (not illustrated) involving <i>Zonotrichia querula</i> (the +homologous antigen) and <i>Zonotrichia albicollis</i>. Although this test +was one of an earlier series in which difficulties were encountered +(the data, therefore, were not used), it is of interest that the two +species were almost indistinguishable serologically.</p> + +<p>The serological homogeneity of passeriform birds is emphasized +by the fact that the value of every heterologous reaction was more +than 50 per cent of the value of the homologous reaction, except in +the test involving the anti-<i>Richmondena</i> serum and <i>Myiarchus</i> (Fig.<a href="#Fig_13"> +13</a>) in which the value of the heterologous reaction was 45 per cent. +Because most ornithologists consider these genera to be only distantly +related (they are in different suborders within the Order +Passeriformes), the relatively high value of the heterologous reaction +emphasizes the close serological correspondence of passerine +birds and indicates that small consistent serological differences +among these birds are actually significant. The possibility that +some of the serological correspondence is due to the "homologizing" +effect of formalin on proteins should not be excluded. I think, however, +that this effect is not entirely responsible for the close correspondence +observed here.</p> + +<p>An additional point to consider in interpretation of the serological +tests is that the techniques used tend to separate sharply species that +are closely related whereas species that are distantly related are not +so easily separated. In other words, comparative serological studies +with the photronreflectometer tend to minimize the differences between +distant relatives and to exaggerate the differences between +close relatives.</p> + +<p>In analyzing the serological relationships of the species used in +this study, it becomes obvious that two or more series of tests must +be considered before the birds can be placed in relation to each +other. For example, the data presented in Fig.<a href="#Fig_14"> 14</a> indicate that +<i>Spiza</i> and <i>Molothrus</i> show approximately the same degree of serological +correspondence to <i>Richmondena</i>. This does not imply necessarily +<span class="pagenum"><a name="Page_193" id="Page_193">[Pg 193]</a></span> +that <i>Spiza</i> and <i>Molothrus</i> are closely related. If Fig.<a href="#Fig_15"> 15</a> is +examined, it can be determined that <i>Richmondena</i> shows much +greater serological correspondence to <i>Spiza</i> than does <i>Molothrus</i>. +Thus, an analysis of both figures serves to clarify the true serological +relationships of the three genera. By reference to other series of +tests involving these three birds a more exact determination of their +relationships may be obtained.</p> + +<p>To illustrate this point by a hypothetical example, two species +might seem equidistant, serologically, from a third species. Additional +testing should indicate if the first two species are equidistant +in the same direction (therefore, by implication, close relatives) or +in opposite directions (therefore, distant relatives). A single test +supplies only two dimensions of a three dimensional arrangement.</p> + +<p>It is impossible to interpret and to picture the serological data +satisfactorily in two dimensions; therefore, a three-dimensional +model (Figs.<a href="#Fig_22"> 22</a>,<a href="#Fig_23"> 23</a>) was constructed to summarize the serological +relationships of the birds involved. Each of the eleven kinds used +consistently throughout the investigation is represented in the model. +By use of the percentage values (Table<a href="#Table_2"> 2</a>), each bird was located +in relation to the other birds. Where possible, averages of reciprocal +tests (Table<a href="#Table_3"> 3</a>) were used in determining distances between the +elements of the model. In this way seven of the birds were accurately +located in relation to each other. Lacking reciprocal tests, +the positions of the other birds were determined by the values of +single tests (Table<a href="#Table_4"> 4</a>). Although these birds were placed with less +certainty, at least four points of reference were used in locating each +species. At least one serological test is represented by each connecting +bar in the model. The lengths of the bars connecting any +two elements were determined as follows: a percentage value +(Table<a href="#Table_3"> 3</a> and Table<a href="#Table_4"> 4</a>) representing the degree of serological correspondence +between two birds was subtracted from 100 per cent; +the remainder was multiplied by a factor of five to increase the size +of the model and the product was expressed in millimeters; a bar +of proper length connects the two elements involved.</p> + +<p>From the model it is observed that, <i>Molothrus</i> and <i>Passer</i> excluded, +the birds fall into two distinct groups: one includes <i>Piranga</i>, +<i>Richmondena</i>, <i>Spiza</i>, <i>Junco</i>, and <i>Zonotrichia</i>; the other includes +<i>Estrilda</i>, <i>Poephila</i>, <i>Carpodacus</i>, and <i>Spinus</i>. +<p> </p> +<p> </p> + +<a name="Table_3"></a> +<p><span class="pagenum"><a name="Page_194" id="Page_194">[Pg 194]</a></span></p> + +<p class="smcap">Table 3.—Reciprocal Values Used to Determine Distances Between +Elements of the Model; Each Value Represents the Average of Serological +Tests Between the Species Involved</p> + +<table width="100%" cellpadding=4 class="center" summary="Species and Extract Injection Data"> +<tr><td colspan=9 class="bt"><img src="images/bar_single.png" width="1" height="5" title="bar" alt="bar" /></td></tr> +<tr><td class="bt bb"> </td><td class="bt bl bb vbot"><img src="images/t2n1.png" width="23" height="138" title="A" alt="A " /></td><td class="bt bl bb vbot"> <img src="images/t2n2.png" width="20" height="125" title="B" alt="B " /></td><td class="bt bl bb vbot"><img src="images/t2n4.png" width="20" height="170" title="D" alt="D " /></td><td class="bt bl bb vbot"><img src="images/t2n5.png" width="19" height="120" title="E" alt="E " /></td><td class="bt bl bb vbot"><img src="images/t2n6.png" width="19" height="98" title="F" alt="F " /></td><td class="bt bl bb vbot"><img src="images/t2n7.png" width="19" height="112" title="G" alt="G " /></td><td class="bt bl bb vbot"><img src="images/t2n8.png" width="20" height="142" title="H" alt="H " /></td></tr> +<tr><td class="text_lf bb"><i>Estrilda amandava</i></td><td class="bl bb">..</td><td class="bl bb">92</td><td class="bl bb">..</td><td class="bl bb">72</td><td class="bl bb">72</td><td class="bl bb">59</td><td class="bl bb">..</td></tr> +<tr><td class="text_lf bb"><i>Poephila guttata</i></td><td class="bl bb">92</td><td class="bl bb">..</td><td class="bl bb">74</td><td class="bl bb">78</td><td class="bl bb">78</td><td class="bl bb">..</td><td class="bl bb">..</td></tr> +<tr><td class="text_lf bb"><i>Richmondena cardinalis</i></td><td class="bl bb">..</td><td class="bl bb">74</td><td class="bl bb">..</td><td class="bl bb">85</td><td class="bl bb">63</td><td class="bl bb">77</td><td class="bl bb">79</td></tr> +<tr><td class="text_lf bb"><i>Spiza americana</i></td><td class="bl bb">72</td><td class="bl bb">78</td><td class="bl bb">85</td><td class="bl bb">..</td><td class="bl bb">77</td><td class="bl bb">77</td><td class="bl bb">85</td></tr> +<tr><td class="text_lf bb"><i>Spinus tristis</i></td><td class="bl bb">72</td><td class="bl bb">78</td><td class="bl bb">63</td><td class="bl bb">77</td><td class="bl bb">..</td><td class="bl bb">..</td><td class="bl bb">..</td></tr> +<tr><td class="text_lf bb"><i>Junco hyemalis</i></td><td class="bl bb">..</td><td class="bl bb">..</td><td class="bl bb">77</td><td class="bl bb">77</td><td class="bl bb">..</td><td class="bl bb">..</td><td class="bl bb">..</td></tr> +<tr><td class="text_lf bb"><i>Zonotrichia querula</i></td><td class="bl bb">..</td><td class="bl bb">..</td><td class="bl bb">79</td><td class="bl bb">85</td><td class="bl bb">..</td><td class="bl bb">..</td><td class="bl bb">..</td></tr> +</table> +<p> </p> +<p> </p> + +<a name="Table_4"></a> +<div class="smcap">Table 4.—Single Values Used to Determine Distances Between Elements +of the Model; Each Value Represents a Single Test Between the +Species Involved</div> + +<table width="100%" cellpadding=4 class="center" summary="Species and Extract Injection Data"> +<tr><td colspan=9 class="bt"><img src="images/bar_single.png" width="1" height="5" title="bar" alt="bar" /></td></tr> +<tr><td class="bt bb"> </td><td class="bt bl bb vbot"><img src="images/t2n1.png" width="23" height="138" title="A" alt="A" /></td><td class="bt bl bb vbot"><img src="images/t2n2.png" width="20" height="125" title="B" alt="B" /></td><td class="bt bl bb vbot"><img src="images/t2n3.png" width="18" height="111" title="C" alt="C" /></td><td class="bt bl bb vbot"><img src="images/t2n4.png" width="20" height="170" title="D" alt="D" /></td><td class="bt bl bb vbot"><img src="images/t2n6.png" width="19" height="98" title="F" alt="F" /></td><td class="bt bl bb vbot"><img src="images/t2n7.png" width="19" height="112" title="G" alt="G" /></td><td class="bt bl bb vbot"><img src="images/t2n8.png" width="20" height="142" title="H" alt="H" /></td></tr> +<tr><td class="text_lf bb"><i>Passer domesticus</i></td><td class="bl bb">..</td><td class="bl bb">74</td><td class="bl bb">73</td><td class="bl bb">..</td><td class="bl bb">72</td><td class="bl bb">..</td><td class="bl bb">..</td></tr> +<tr><td class="text_lf bb"><i>Molothrus ater</i></td><td class="bl bb">..</td><td class="bl bb">54</td><td class="bl bb">..</td><td class="bl bb">65</td><td class="bl bb">..</td><td class="bl bb">69</td><td class="bl bb">75</td></tr> +<tr><td class="text_lf bb"><i>Piranga rubra</i></td><td class="bl bb">..</td><td class="bl bb">77</td><td class="bl bb">..</td><td class="bl bb">91</td><td class="bl bb">73</td><td class="bl bb">74</td><td class="bl bb">..</td></tr> +<tr><td class="text_lf bb"><i>Carpodacus purpureus</i></td><td class="bl bb">70</td><td class="bl bb">71</td><td class="bl bb">..</td><td class="bl bb">61</td><td class="bl bb">93</td><td class="bl bb">..</td><td class="bl bb">..</td></tr> +</table> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_195" id="Page_195">[Pg 195]</a></span></p> + +<div class="center"> + <table summary="Graphs"> + <tr> + <td><a name="Fig_10"></a><img src="images/fig_10.png" width="228" height="296" title="" alt="" /></td> + <td><a name="Fig_11"></a><img src="images/fig_11.png" width="223" height="291" title="" alt="" /></td> + </tr> + <tr> + <td><a name="Fig_12"></a><img src="images/fig_12.png" width="228" height="298" title="" alt="" /></td> + <td><a name="Fig_13"></a><img src="images/fig_13.png" width="228" height="294" title="" alt="" /></td> + </tr> + <tr> + <td colspan=2 class="center"><img src="images/key_10_13.png" width="320" height="47" title="key" alt="key" /></td> + </tr> + </table> +</div> + +<p><span class="smcap">Figs. 10-13.</span> Graphs of precipitin reactions illustrating effects of formalin on antigenicity and reactivity of the extracts. For further information, see text, pp.<a href="#Page_190"> 190-193</a>.</p> + +<p><span class="smcap">Fig. 10.</span> Reactions of unformolized antigens of <i>Richmondena</i>, <i>Zonotrichia</i>, and <i>Molothrus</i> with anti-<i>Richmondena</i> serum. <span class="smcap">Fig. 11.</span> Reactions of formolized antigens of <i>Richmondena</i>, <i>Zonotrichia</i>, and <i>Molothrus</i> with anti-<i>Richmondena</i> serum. <span class="smcap">Fig. 12.</span> Reactions of anti-<i>Richmondena</i> serum prepared against native antigen with antigens of <i>Richmondena</i>, <i>Zonotrichia</i>, <i>Carpodacus</i>, and <i>Spinus</i>. <span class="smcap">Fig. 13.</span> Reactions of anti-<i>Richmondena</i> serum prepared against formolized antigen with antigens of <i>Richmondena</i>, <i>Zonotrichia</i>, <i>Poephila</i>, <i>Spinus</i>, and <i>Myiarchus</i>.</p> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_196" id="Page_196">[Pg 196]</a></span></p> + +<div class="center"> + <table summary="Graphs"> + <tr> + <td><a name="Fig_14"></a><img src="images/fig_14.png" width="229" height="298" title="" alt="" /></td> + <td><a name="Fig_15"></a><img src="images/fig_15.png" width="221" height="296" title="" alt="" /></td> + </tr> + <tr> + <td><img src="images/fig_16.png" width="218" height="290" title="" alt="" /></td> + <td><a name="Fig_17"></a><img src="images/fig_17.png" width="214" height="296" title="" alt="" /></td> + </tr> + <tr> + <td colspan=2 class="center"><img src="images/key_14_17.png" width="317" height="49" title="" alt="key" /></td> + </tr> + </table> +</div> + +<p><span class="smcap">Figs. 14-17.</span> Graphs of precipitin reactions illustrating serological relationships. For further explanation, see text, pp.<a href="#Page_190"> 190-193</a>.</p> + +<p><span class="smcap">Fig. 14.</span> Serological relationships of <i>Richmondena</i>, <i>Spiza</i>, and <i>Molothrus</i>. <span class="smcap">Fig. 15.</span> Serological relationships of <i>Richmondena</i>, <i>Spiza</i>, and <i>Molothrus</i>. <span class="smcap">Fig. 16.</span> Serological relationships of <i>Carpodacus</i> with the richmondenine-emberizine-thraupid assemblage. <span class="smcap">Fig. 17.</span> Serological relationships of <i>Carpodacus</i> and <i>Spinus</i> with <i>Richmondena</i> and <i>Junco</i>.</p> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_197" id="Page_197">[Pg 197]</a></span></p> + +<div class="center"> + <table summary="Graphs"> + <tr> + <td><a name="Fig_18"></a><img src="images/fig_18.png" width="225" height="300" title="" alt="" /></td> + <td><a name="Fig_19"></a><img src="images/fig_19.png" width="220" height="298" title="" alt="" /></td> + </tr> + <tr> + <td><a name="Fig_20"></a><img src="images/fig_20.png" width="220" height="298" title="" alt="" /></td> + <td><a name="Fig_21"></a><img src="images/fig_21.png" width="221" height="299" title="" alt="" /></td> + </tr> + <tr> + <td colspan=2 class="center"><img src="images/key_18_21.png" width="316" height="49" title="" alt="key" /></td> + </tr> + </table> +</div> + +<p><span class="smcap">Figs. 18-21.</span> Graphs of precipitin reactions illustrating serological relationships. For further explanation, see text, pp.<a href="#Page_190"> 190-193</a>.</p> + +<p><span class="smcap">Fig. 18.</span> Serological relationships of <i>Spinus</i> and <i>Poephila</i> with the richmondenines. <span class="smcap">Fig. 19.</span> Serological relationships of <i>Carpodacus</i> and <i>Spinus</i> with <i>Richmondena</i> and <i>Piranga</i>. <span class="smcap">Fig. 20.</span> Serological relationships of <i>Poephila</i> and Richmondena with the carduelines. <span class="smcap">Fig. 21.</span> Serological relationships of <i>Richmondena</i> and <i>Spinus</i> with the estrildines.</p> +<p> </p> +<p> </p> + +<a name="Fig_22"></a> +<p><span class="pagenum"><a name="Page_198" id="Page_198">[Pg 198]</a></span></p> + +<div class="center"> + <img src="images/fig_22a.png" width="441" height="285" title="Relationship Model" alt="Relationship Model" /><br /><br /> + <img src="images/fig_22b.png" width="450" height="300" title="Relationship Model" alt="Relationship Model" /><br /><br /> +</div> + +<p><span class="smcap">Fig. 22.</span> Two views of a model illustrating serological relationships among fringillid and related birds. For further explanation, see text, pp.<a href="#Page_193"> 193-194</a>.</p> + +<div class="center"> + <table width="60%" summary="List of Genera"> + <tr><td colspan=3 class="center">Genera</td><td> </td><td>Pi</td><td> . . . . </td><td><i>Piranga</i></td></tr> + <tr><td>C</td><td> . . . . </td><td><i>Carpodacus</i></td><td> </td><td>Po</td><td> . . . . </td><td><i>Poephila</i></td></tr> + <tr><td>E</td><td> . . . . </td><td><i>Estrilda</i></td><td> </td><td>R</td><td> . . . . </td><td><i>Richmondena</i></td></tr> + <tr><td>J</td><td> . . . . </td><td><i>Junco</i></td><td> </td><td>Sn</td><td> . . . . </td><td><i>Spinus</i></td></tr> + <tr><td>M</td><td> . . . . </td><td><i>Molothrus</i></td><td> </td><td>Sz</td><td> . . . . </td><td><i>Spiza</i></td></tr> + <tr><td>Pa</td><td> . . . . </td><td><i>Passer</i></td><td> </td><td>Z</td><td> . . . . </td><td><i>Zonotrichia</i></td></tr> + </table> +</div> +<p> </p> +<p> </p> + +<a name="Fig_23"></a> +<p><span class="pagenum"><a name="Page_199" id="Page_199">[Pg 199]</a></span></p> + +<div class="center"> + <img src="images/fig_23a.png" width="437" height="276" title="" alt="Relationship Model" /><br /><br /> + <img src="images/fig_23b.png" width="454" height="321" title="" alt="Relationship Model" /><br /><br /> +</div> + +<p><span class="smcap">Fig. 23.</span> Two additional views of the model shown in fig. 22 illustrating serological relationships among fringillid and related birds. For further explanation, see text, pp.<a href="#Page_193"> 193-194</a>.</p> + +<div class="center"> + <table width="60%" summary="List of Genera"> + <tr><td colspan=3 class="center">Genera</td><td> </td><td>Pi</td><td> . . . . </td><td><i>Piranga</i></td></tr> + <tr><td>C</td><td> . . . . </td><td><i>Carpodacus</i></td><td> </td><td>Po</td><td> . . . . </td><td><i>Poephila</i></td></tr> + <tr><td>E</td><td> . . . . </td><td><i>Estrilda</i></td><td> </td><td>R</td><td> . . . . </td><td><i>Richmondena</i></td></tr> + <tr><td>J</td><td> . . . . </td><td><i>Junco</i></td><td> </td><td>Sn</td><td> . . . . </td><td><i>Spinus</i></td></tr> + <tr><td>M</td><td> . . . . </td><td><i>Molothrus</i></td><td> </td><td>Sz</td><td> . . . . </td><td><i>Spiza</i></td></tr> + <tr><td>Pa</td><td> . . . . </td><td><i>Passer</i></td><td> </td><td>Z</td><td> . . . . </td><td><i>Zonotrichia</i></td></tr> + </table> +</div> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_200" id="Page_200">[Pg 200]</a></span></p> + +<p>Within the richmondenine-emberizine-thraupid assemblage, <i>Junco</i> +and <i>Zonotrichia</i> constitute a sub-group apart from the others. +<i>Piranga</i> and <i>Richmondena</i> show close serological correspondence. +The present taxonomic position of <i>Spiza</i> in the Richmondeninae, +which has been questioned by Beecher (1951a:431; 1953:309), is +corroborated at least insofar as the serological evidence is concerned. +Certainly, serological correspondence of <i>Spiza</i> with the richmondenine-emberizine-thraupid +assemblage is greater than with any +other group of birds tested.</p> + +<p>It is obvious that the serological affinities of the carduelines do +not lie with the richmondenines, emberizines, or thraupids. The +carduelines show greater serological correspondence with the +estrildines than they do with any of the other groups tested. Further +serological investigation involving other species, however, is +necessary before the nearest relatives of the carduelines can be determined +with certainty.</p> + +<p>The two estrildines tested (<i>Estrilda</i> and <i>Poephila</i>) show close +serological relationship. Their nearest relatives, serologically, seem +to be the carduelines. The classification (Wetmore, 1951) that +places <i>Passer</i> in the same family with the estrildines is not upheld by +the serological data available. <i>Passer</i> is not, serologically, closely +related to any of the birds tested. It is of interest that Beecher +(1953:303-305), on the basis of jaw musculature, places <i>Passer</i> and +the estrildines in separate families (Ploceidae and Estrildidae, respectively).</p> + +<p><i>Molothrus</i> shows greater serological correspondence to the richmondenine-emberizine-thraupid +assemblage than to any of the other +birds tested. It is definitely set apart from this group, however, and +its position, serologically, is compatible with that based on evidence +from other sources.</p> + +<p>There seems to be but little argument among ornithologists that +icterids, fringillids, and ploceids constitute families which are distinct +from one another. If, then, the serological differences between +<i>Molothrus</i> (Icteridae) and <i>Richmondena</i> (Fringillidae), between +<i>Molothrus</i> and <i>Zonotrichia</i> (Fringillidae), and between <i>Richmondena</i> +and <i>Poephila</i> (Ploceidae) are indicative of family differences, +there are four families represented by the birds involved. +<i>Molothrus</i> represents one family; <i>Piranga</i>, <i>Richmondena</i>, <i>Spiza</i>, +<i>Junco</i>, and <i>Zonotrichia</i>, a second; <i>Estrilda</i>, <i>Poephila</i>, <i>Carpodacus</i>, +and <i>Spinus</i>, a third; and <i>Passer</i>, a fourth.</p> +<p> </p> +<p> </p> + +<a name="Conclusions"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span><br> +<p><span class="pagenum"><a name="Page_201" id="Page_201">[Pg 201]</a></span></p> +<div class="caption2h smcap"><span class="smcap">Conclusions</span></div> + + +<p>The heterogeneity of the Family Fringillidae has been emphasized +by many authors. The relationships of the species now included +in this Family have been the subject of much discussion and constitute +an important problem in avian systematics.</p> + +<p>Sushkin's studies (1924, 1925) of features of the horny and bony +palates have served as a basis for the present division of the Family +into subfamilies. Recently, Beecher (1951a, 1951b, 1953) and +Tordoff (1954) have used these features and others which they +thought to be of value in an attempt to clarify the relationships of +the species involved.</p> + +<p>Beecher's work (1951a, 1951b, 1953) on jaw-musculature is a +valuable contribution to our knowledge of the anatomy of passerine +birds. His myological studies were so thorough and his presentation +so detailed that students who disagree with his interpretations +can draw their own conclusions. Beecher (1951b:276) points out +that there are two basic types of skeletal muscle—those with parallel +fibers and those with pinnately arranged fibers. The muscles with +pinnate fibers seem to be more efficient, each muscle having a +greater functional cross section for its bulk than does one with +parallel fibers. He assumes that muscles with parallel fibers are +more primitive, phylogenetically, than are those with fibers arranged +pinnately. Since his study of the jaw muscles of the Icteridae +(1951a) revealed that patterns of jaw-musculature within this +Family remain constant regardless of the methods used in procuring +food, he assumes that such patterns may be used as indicators +of relationship throughout the entire oscinine group. These two +assumptions, then, serve as the basis for his hypothesis concerning +relationship and phylogeny within this assemblage. Beecher +(1951b:278-280; 1953:310-312) maintains that within the Family +Thraupidae there are two main lines which lead with almost no disjunction +to the Carduelinae and Richmondeninae. The thraupid-richmondenine +line involves a shift in the nature of the <i>m. adductor +mandibulae externus superficialis</i>, which becomes more pinnate in +the richmondenines. This results in greater crushing power. The +thraupid-cardueline line involves a shift in emphasis from the the <i>m. +adductor mandibulae externus medialis</i> to the <i>m. pseudotemporalis +superficialis</i> and the forward advance of the insertion of the latter. +This, also, promotes greater crushing ability. He states that features +of the horny palate and of the plumage provide further evidence of +close relationship of these groups. He includes, therefore, the +<span class="pagenum"><a name="Page_202" id="Page_202">[Pg 202]</a></span> +Thraupinae, the Carduelinae, and the Pyrrhuloxiinae (=Richmondeninae) +in the Family Thraupidae. Beecher (1953:307) indicates +that the patterns of jaw-musculature of the Parulinae (wood warblers) +and Emberizinae (buntings) are similar and suggests that the +buntings had their origin from the wood warblers. He includes +these subfamilies, therefore, in the Family Parulidae.</p> + +<p>Beecher's reasoning may be criticized on several points. It may +be, as he suggests, that muscles with parallel fibers evolved earlier, +phylogenetically, than did muscles with pinnate fibers, but he does +not give adequate consideration, it seems to me, to the possibility +that parallel fibers may also have evolved secondarily from pinnate +fibers. Since Beecher (1951a) found that patterns of jaw-musculature +within the Family Icteridae were conservative, he is reluctant +to admit the possibility of convergence among any of the other +families. Differences in patterns of jaw-musculature are, however, +functional adaptations and like the bill, which is also associated with +food-getting may be subject to rapid evolutionary change. Finally, +in attempting to classify the oscines, he has relied almost entirely on +a single character—the pattern of jaw-musculature.</p> + +<p>Tordoff's attempts (1954) to clarify the relationships of the fringillids +and related species are based chiefly on features of the bony +palate. He assumes that since palato-maxillaries seem to be absent +in the majority of passerine birds, their occurrence in certain nine-primaried +oscine groups indicates relationship among these groups. +He points out that these bones, when present, are important areas +of origin of the <i>m. pterygoideus</i> which functions in depression of +the upper jaw and in elevation of the lower jaw. He assumes, therefore, +that palato-maxillaries were evolved to provide for a more +effective action of the <i>m. pterygoideus</i>. The need for such action +could be associated with a seed-eating habit. All richmondenines +and emberizines possess palato-maxillary bones either free or fused +to the prepalatine bar, but there is no trace of these bones in the +carduelines. Carduelines, furthermore, possess prepalatine bars +that are characteristically flared anteriorly. This condition does not +exist in the richmondenines or in the emberizines.</p> + +<p>Tordoff points out, also, that the irregular, erratic migrations of +the New World Carduelinae are unlike the more regular migrations +of the richmondenines and emberizines. The carduelines, furthermore, +are more arboreal in their habits than are these other groups +and exhibit a decided lack of nest sanitation during the later stages +of nesting, a situation which contrasts with that found in the Richmondeninae +<span class="pagenum"><a name="Page_203" id="Page_203">[Pg 203]</a></span> +and Emberizinae. He suggests, therefore, that the +carduelines are not so closely related to the richmondenines and the +emberizines as previously has been thought.</p> + +<p>Since there are only two cardueline genera, <i>Loximitris</i> and <i>Hesperiphona</i>, +endemic to the New World and at least 10 genera with +many species endemic to the Old World, Tordoff (1954:15) suggests +an Old World origin for the carduelines. He strengthens his +argument for this hypothesis by pointing out that in features of the +bony palate and in habits the carduelines resemble the estrildines of +the Family Ploceidae.</p> + +<p>Tordoff (1954:29-30) states that the tanagers not only merge with +the richmondenines but also grade imperceptibly into the emberizines. +He includes, therefore, the Richmondeninae, Emberizinae, +and Thraupinae in the Family Fringillidae. He suggests that the +carduelines are ploceids, closely related to the Subfamily Estrildinae, +on the basis of structure of the bony palate, geographic distribution, +social behavior, and habits such as nest-fouling and nest-building.</p> + +<p>Tordoff, like Beecher, has based his interpretations chiefly on one +feature—structure of the bony palate. Since this feature also is +associated with food-getting, the possibilities of convergence of distantly +related species with similar habits and divergence of closely +related species with different habits may not be excluded.</p> + +<p>The hazard of unrecognized adaptive convergence cannot, of +course, be excluded from most fields of taxonomic research, but +some features of morphology and biochemistry are notably more +conservative than others and undergo slower evolutionary change. +Such features are often of utmost importance in distinguishing the +higher taxonomic categories.</p> + +<p>Most ornithologists are aware that, within the Order Passeriformes, +patterns of musculature in the leg have evolved at a slow +rate and exhibit little variation within the Order. Differences which +do occur, therefore, probably are significant, especially those that +are consistent between groups of species. As I have pointed out +earlier (p.<a href="#Page_184"> 184</a>), there are no significant differences in leg-musculature +between the Richmondeninae, Emberizinae, and Thraupidae. Indeed, +it is difficult to define these groups on the basis of leg-musculature. +If these groups are of common origin, the lack of distinct +boundaries between them is not surprising. A muscular band which +extends from the <i>pars interna</i> of the <i>m. gastrocnemius</i> around the +front of the knee is present in every emberizine species that I studied +and in the Genus <i>Piranga</i>. With the exception of <i>Spiza</i> none of the +richmondenines possesses this band.</p> + +<p><span class="pagenum"><a name="Page_204" id="Page_204">[Pg 204]</a></span> +The significant differences in leg-musculature which have been +discussed above (pp.<a href="#Page_183"> 183-184</a>) distinguish the carduelines from the +New World finches and tanagers. Even the cardueline <i>Leucosticte</i> +and the emberizine <i>Calcarius</i>, which resemble one another in general +adaptations and in several myological features of the leg (p. +<a href="#Page_183"> 183</a>), agree in significant features of the musculature with the respective +groups to which they belong. The carduelines agree in +the major features of leg-musculature with the ploceids which I +studied.</p> + +<p>The use of serological techniques in taxonomic work has two +main advantages. The biochemical systems involved in such investigations +seem to be relatively slow to change in response to external +environmental influences, and the quantitative nature of the +results obtained makes possible objective measurement of resemblances +among species.</p> + +<p>I have pointed out (p. <a href="#Page_200">200</a>) that the carduelines are excluded, +serologically, from the distinct assemblage formed by the richmondenines, +emberizines, and tanagers. Actually, the carduelines show +less serological resemblance to this assemblage than do the estrildines, +and most ornithologists agree that the Estrildinae are not at +all closely related to the Richmondeninae, Emberizinae, and Thraupidae. +<i>Molothrus</i>, representing a family (Icteridae) recognized as +distinct from the Family Fringillidae, also more closely resembles +the fringillid assemblage, serologically, than do the carduelines. Although +the Carduelinae constitute a distinct group serologically, +they show greater serological resemblance to the estrildines of the +Family Ploceidae than to any of the other species tested. At least +the carduelines and the estrildines form a group as compact as the +subfamilies of the Fringillidae. Thus, the serological data correlate +well with those obtained from the study of the leg-musculature.</p> + +<p>Present systems of classification include the subfamilies Passerinae +and Estrildinae in the Family Ploceidae. <i>Passer</i>, however, is less +closely related to the estrildines serologically than are the carduelines, +and is less closely related to the estrildines than <i>Molothrus</i>, an +icterid, is to the fringillids. This raises a question as to the homogeneity +of the Family Ploceidae as presently recognized by most +ornithologists. If the Passerinae and the Estrildinae are placed in +a single family, the serological divergence among members of this +group is certainly greater than it is in the Family Fringillidae. Additionally, +Beecher (1953:303-304) found that the estrildines possess +a pattern of jaw-musculature different from those in other ploceids.</p> + +<p><span class="pagenum"><a name="Page_205" id="Page_205">[Pg 205]</a></span> +The combined evidence from jaw-musculature and serology has +caused me to conclude that the estrildines should be excluded from +the Family Ploceidae (<a href="#Page_207">see below</a>).</p> + +<p>In an attempt to clarify the relationships of the Fringillidae and +allied groups, I here review briefly the evidence which has been +presented. From his studies of jaw-musculature (1951a, 1951b, +1953) Beecher concludes that the Pyrrhuloxinae (=Richmondeninae), +the Carduelinae, and the Thraupinae are closely related. +He places these groups in the Family Thraupidae. He excludes the +Emberizinae from this group and places them with the wood warblers +in the Family Parulidae. He suggests that the estrildines constitute +a family (Estrildidae) separate from the Family Ploceidae.</p> + +<p>From his studies of certain features of the bony palate Tordoff +(1954:25-26, 32) concludes that the richmondenines, the emberizines, +and the tanagers have a common origin and places these +groups in the Family Fringillidae. He excludes the carduelines from +this assemblage, suggests that they are closely related to the estrildines, +and includes them as the Subfamily Carduelinae in the +Family Ploceidae.</p> + +<p>In this paper I have presented data obtained from the study of +certain features of morphology and biochemistry which I think are +less subject to the influence of environmental factors than those +features studied by recent workers. It is significant that the data +obtained by use of serological techniques and those obtained from +the study of leg-musculature point to the same conclusions. On the +basis of these data I have drawn several conclusions concerning the +relationships of the groups which I studied.</p> + +<p>The richmondenines, emberizines, and tanagers are closely related +and should be included in a single family, Fringillidae. The +Carduelinae and the Estrildinae are closely related subfamilies. Although +most recent classifications place the Estrildinae and Passerinae +in the Family Ploceidae, the serological evidence indicates +that these groups are not closely related. Beecher (1953:303-304) +drew the same conclusion from his study of jaw-musculature (see +above). I suggest, therefore, that the Carduelinae and the Estrildinae +be placed in a family separate from the Ploceidae and that +the name Carduelidae (rather than Estrildidae) be used for this +group. At present, neither is an accepted family name. Because +<i>Carduelis</i> Brisson 1760 is an older name than <i>Estrilda</i> Swainson +1827 and because <i>Carduelis</i> seems to be a centrally located genus +in the family, I have chosen the former (although the International +<span class="pagenum"><a name="Page_206" id="Page_206">[Pg 206]</a></span> +Rules of Zoological Nomenclature do not specify that priority must +apply in forming family names).</p> + +<p>I have been unable to study any of the species included in the +subfamilies Fringillinae (not Fringillinae of Tordoff, see 1954:23-24, +and below) and Geospizinae of recent classifications; thus these +groups have not been discussed above. Beecher (1953:307-308) +includes <i>Fringilla</i> in the Subfamily Carduelinae; he includes the +geospizines in a separate family, Geospizidae, and states that they +are derived from the emberizines. Tordoff (1954:23-24) found that +in features of the bony palate <i>Fringilla</i> and the geospizines resemble +the emberizines and, on this basis, includes them in the Subfamily +Fringillinae.</p> + +<p>The Dickcissel, <i>Spiza americana</i>, possesses certain features which +merit special discussion. Beecher (1951a:431; 1953:309), on the +basis of jaw-musculature, considers it an icterid. To be sure <i>Spiza</i> is +in many ways an aberrant member of the group to which it is now +assigned (Subfamily Richmondeninae). <i>Spiza</i>, serologically, is +closely related to all species of the richmondenine-emberizine-thraupid +assemblage. Within this assemblage its nearest relatives +are the richmondenines. <i>Spiza</i> differs from the other richmondenines +studied and resembles the emberizines and tanagers in the +possession of the muscular band which extends from the <i>pars interna</i> +of the <i>m. gastrocnemius</i> around the front of the knee. This band, in +<i>Spiza</i>, is smaller, however, than in any of the other species. No +icterid dissected possesses such a structure. Tordoff (1954:29) +states that <i>Spiza</i> is typically richmondenine in palatal structure and +makes the suggestion, with which I agree, that <i>Spiza</i> is a richmondenine +and may be closely related to the ancestral stock which gave +rise to the fringillid assemblage. The serological position of <i>Spiza</i>, +approximately equidistant from the other fringillids (Figs.<a href="#Fig_22"> 22</a>,<a href="#Fig_23"> 23</a>), +and the presence of the small muscular band around the front of +the knee constitute evidence supporting the central position of <i>Spiza</i>.</p> + +<p>After consideration of evidence from the studies of external morphology, +ethology, myology, osteology, and serology, I propose here +an arrangement of the groups which I have studied and submit for +comparison the arrangements (of these groups) proposed by +Beecher and Tordoff. The names of subfamilies that I have been +unable to study are included in my classification and are placed in +brackets.</p> +<p> </p> +<p> </p> + +<p><span class="pagenum"><a name="Page_207" id="Page_207">[Pg 207]</a></span></p> + +<table width="100%" cellpadding=4 class="center" summary="Family Associations"> +<tr><td class="bt bb" width="33%">Here proposed</td><td class="bt bl bb" width="33%">Proposed by Tordoff (1954) on the basis of the bony palate:</td><td class="bt bl bb" width="33%">Proposed by Beecher (1953) on the basis of jaw-musculature:</td></tr> +<tr><td class="bb"><img src="images/bar_single.png" width="1" height="3" title="bar" alt="bar" /></td><td class="bb bl"><img src="images/bar_single.png" width="1" height="3" title="bar" alt="bar" /></td><td class="bb bl"><img src="images/bar_single.png" width="1" height="3" title="bar" alt="bar" /></td></tr> +<tr><td class="smcap text_lf"> Family Ploceidae</td><td class="smcap bl"> Family Ploceidae</td><td class="smcap bl"> Family Ploceidae</td></tr> +<tr><td class="text_lf">[Subf. Bubalornithinae]</td><td class="bl">Subf. Bubalornithinae</td><td class="bl"> </td></tr> +<tr><td class="text_lf">Subfamily Passerinae: distinguished from the Estrildinae by patterns of jaw-musculature (Beecher, 1953:303-304) and on the basis of comparative serology of saline-soluble proteins.</td> +<td class="bl vtop">Subfamily Passerinae</td> +<td class="bl vtop">Subfamily Passerinae</td></tr> +<tr><td class="text_lf">[Subfamily Ploceinae]</td><td class="bl">Subfamily Ploceinae</td><td class="bl">Subfamily Ploceinae</td></tr> +<tr><td class="text_lf">[Subfamily Viduinae]</td><td class="bl">Subfamily Viduinae</td><td class="bl">Subfamily Viduinae</td></tr> +<tr><td class="bb"><img src="images/bar_single.png" width="1" height="3" title="bar" alt="bar" /></td><td class="bb bl"><img src="images/bar_single.png" width="1" height="3" title="bar" alt="bar" /></td><td class="bb bl"><img src="images/bar_single.png" width="1" height="3" title="bar" alt="bar" /></td></tr> +<tr><td class="smcap text_lf"> Family Carduelidae</td><td class="bl"> </td><td class="bl"> </td></tr> +<tr><td class="text_lf">Subfamily Estrildinae: similar to the Carduelinae in features of the bony palate and habits (Tordoff, 1954: 18-22) and in patterns of leg-musculature and comparative serology of saline-soluble proteins.</td><td class="bl vtop">Subfamily Estrildinae</td><td class="smcap bl vtop">Family Estrildidae</td></tr> +<tr><td class="text_lf">Subfamily Carduelinae: distinguished from the Fringillidae by features of the palate, geographic distribution, migration patterns, and habits (Tordoff, 1954: 14-18) and by patterns of leg-musculature and comparative serology of saline-soluble proteins.</td><td class="bl vtop">Subfamily Carduelinae</td><td class="bl vtop">[In Thraupidae below]</td></tr> +<tr><td class="bb white"><img src="images/bar_single.png" width="1" height="3" title="bar" alt="bar" /></td><td class="bb bl"><img src="images/bar_single.png" width="1" height="3" title="bar" alt="bar" /></td><td class="bb bl"><img src="images/bar_single.png" width="1" height="3" title="bar" alt="bar" /></td></tr> +<tr><td rowspan=2 class="text_lf"><span class="smcap"> Family Fringillidae:</span> all members of this family show similarities in features of the bony palate (Tordoff, 1954: 22-23), patterns of leg-musculature, and in comparative serology of saline-soluble proteins.</td><td rowspan=2 class="smcap bl vtop"> Family Fringillidae</td><td class="bl vtop"> <span class="smcap">Family Parulidae</span><br />Subfamily Parulinae<br />Subfamily Emberizinae</td></tr> +<tr><td class="smcap bl vbot"> Family Thraupidae</td></tr> +<tr><td class="text_lf bb">Subf. Richmondeninae<br />Subfamily Thraupinae<br />Subfamily Emberizinae<br />[Subfamily Fringillinae]<br />[Subfamily Geospizinae]</td><td class="bl bb vtop">Subf. Richmondeninae<br />Subfamily Thraupinae<br />Subfamily Fringillinae<br />(including Emberizinae<br />and Geospizinae)</td><td class="bl bb vtop">Subfamily Pyrrhuloxiinae<br />Subfamily Thraupinae<br />[In Parulidae above]<br />Subfamily Carduelinae</td></tr> +</table> +<p> </p> +<p> </p> + +<a name="Summary"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span><br> +<p><span class="pagenum"><a name="Page_208" id="Page_208">[Pg 208]</a></span></p> +<div class="caption2h smcap"><span class="smcap">Summary</span></div> + +<p>It has long been recognized that the Family Fringillidae includes +some dissimilar groups. Specifically, the relationships of the subfamilies +Richmondeninae, Emberizinae, and Carduelinae of the +Family Fringillidae are poorly understood. Data from two recent +studies, one on patterns of jaw-musculature and the other on features +of the bony palate, emphasize the dissimilarity of these subfamilies +but have given rise to conflicting concepts of the relationships +of subfamilies within the Family.</p> + +<p>This paper reports the results of studies involving morphological +and biochemical features that I consider less sensitive to external +environmental factors than are features which have been studied +previously. Patterns of leg-musculature were chosen for study because +earlier work showed that muscle patterns in the legs of passerine +birds are highly stable and vary but little. Variations, therefore, +which are consistent in separating groups of species should be +significant. Serological techniques were used because the biochemical +systems involved seem to be relatively slow to change in +response to environmental influences and because the data obtained +may be used in a highly objective manner to measure resemblance +among species.</p> + +<p>Individual differences in the patterns of leg-musculature were +found to be slight and involved mainly the sizes and shapes of +muscles. For this reason variations involving origin, insertion, or +relative position of a muscle, were judged significant. In leg-musculature +the Richmondeninae, the Emberizinae, and the Thraupidae resemble +one another closely. Several differences in muscle pattern +were found, however, which distinguish these groups from the +Carduelinae. The leg-musculature of the carduelines closely resembles +that of the Ploceidae.</p> + +<p>Serological techniques involved the extraction of saline-soluble +proteins from the tissues of the species to be studied. These extracts +were carefully processed and were used as antigens. Formolization +of the antigens was necessary as a means of preventing denaturation +of the proteins by enzymatic activity. Antisera were produced in +rabbits. The method of testing involved turbidimetric analysis of +the precipitin reaction. Utilizing the values for the precipitin tests +a model was constructed which showed the relationships of the +eleven species used in these tests. From a study of the model and +the data used in its construction, it was determined that the Richmondeninae, +Emberizinae, and Thraupidae constitute an assemblage +<span class="pagenum"><a name="Page_209" id="Page_209">[Pg 209]</a></span> +distinct from the other species studied. The Carduelinae are +excluded from the assemblage and serologically are most closely related +to the Estrildinae. The estrildines, serologically, do not closely +resemble <i>Passer</i>, Subfamily Passerinae, although recent classifications +place these two subfamilies in the Family Ploceidae.</p> + +<p>Upon consideration of all evidence now available—from external +morphology, ethology, myology, osteology, and serology—several +hypotheses regarding the relationships of the groups studied are set +forth. The richmondenines, emberizines, and tanagers are closely +related subfamilies and are here included in the Family Fringillidae. +The Estrildinae and Carduelinae are closely related subfamilies, but +neither group is closely related to the Passerinae. The estrildines +and carduelines, therefore, are placed in a separate family, the +Carduelidae. In some ways, <i>Spiza</i> is an aberrant member of the Subfamily +Richmondeninae but should be retained in that subfamily. +It is suggested that <i>Spiza</i> is a primitive richmondenine closely related +to the ancestral fringillid stock.</p> +<p> </p> +<p> </p> + +<a name="Literature_Cited"></a> +<span class="pagenum"><a href="#toc">[↑ TOC]</a></span><br> +<p><span class="pagenum"><a name="Page_210" id="Page_210">[Pg 210]</a></span></p> +<div class="caption2h smcap">Literature Cited</div> +<p> </p> + +<span class="smcap">American Ornithologists' Union</span><br /> +<div class="reference">1931. Check-list of North American birds. Fourth edition. Lancaster, Pa., xix + 526 pp.</div><br /> + +<span class="smcap">Beecher, W. J.</span><br /> +<div class="reference">1951a. Adaptations for food-getting in the American blackbirds. Auk, 68:411-440, 11 figs.</div> +<div class="reference">1951b. Convergence in the Coerebidae. Wilson Bull., 63:274-287, 5 figs.</div> +<div class="reference">1953. A phylogeny of the oscines. Auk, 70:270-333, 18 figs.</div><br /> + +<span class="smcap">Berger, A. J.</span><br /> +<div class="reference">1952. The comparative functional morphology of the pelvic appendage in three genera of Cuculidae. Amer. Mid. Nat., 47:513-605, 29 pls.</div> + +<span class="smcap">Boyden, A.</span><br /> +<div class="reference">1942. Systematic serology: a critical appreciation. Physiol. Zool., 15:109-145, 12 figs.</div><br /> + +<span class="smcap">Chapin, J. P.</span><br /> +<div class="reference">1917. The classification of the weaver-birds. Bull. Amer. Mus. Nat. Hist., 37:243-280, 10 pls., 9 figs.</div><br /> + +<span class="smcap">Cumley, R. W.</span>, and <span class="smcap">Irwin, M. R.</span><br /> +<div class="reference">1941. Pictorial representation of the antigenic differences between two dove species. Jour. Hered., 32:178-182, frontispiece, 2 figs.</div> +<div class="reference">1941. Interaction of antigens in dove hybrids. Ibid., 429-434, 3 figs.</div> +<div class="reference">1944. The correlation between antigenic composition and geographic range in the Old and New World of some species of <i>Columba</i>. Amer. Nat., 78:238-256, 1 fig.</div><br /> + +<span class="smcap">DeFalco, R. J.</span><br /> +<div class="reference">1942. A serological study of some avian relationships. Biol. Bull., 83:205-218.</div><br /> + +<span class="smcap">Fisher, H. I.</span><br /> +<div class="reference">1946. Adaptations and comparative anatomy of the locomotor apparatus of New World vultures. Amer. Mid. Nat., 35:545-727, 13 pls., 28 figs.</div><br /> + +<span class="smcap">Gadow, H.</span>, and <span class="smcap">Selenka, E.</span><br /> +<div class="reference">1891. Vögel, vol. I, Anatomischer Theil. In Bronn's Klassen und Ordnungen des Thier-Reichs, Sechster Band, Vierte Abtheilung. Leipzig, 1008 pp., 59 pls.</div><br /> + +<span class="smcap">Garrod, A. H.</span><br /> +<div class="reference">1873. On certain muscles in the thigh of birds and their value in classification. Proc. Zool. Soc. London, Part I:626-644, 6 figs.</div> +<div class="reference">1874. On certain muscles in the thigh of birds and their value in classification. Ibid., Part II:111-123.</div><br /> + +<span class="smcap">Greenberg, D. M.</span><br /> +<div class="reference">1929. The colorimetric determination of serum proteins. J. Biol. Chem., 82:545-550.</div><br /> + +<span class="smcap">Hellmayr, C. E.</span><br /> +<div class="reference">1935. Catalogue of birds of the Americas. Field Mus. Nat. Hist., Zool. ser. 13, pt. 8, vi + 541 pp.</div> +<div class="reference">1936. Catalogue of birds of the Americas. Ibid., 13, pt. 9, v + 458 pp.</div> +<div class="reference">1937. Catalogue of birds of the Americas. Ibid., 13, pt. 10, v + 228 pp.</div> +<div class="reference">1938. Catalogue of birds of the Americas. Ibid., 13, pt. 11, vi + 662 pp.</div><br /> + +<p><span class="pagenum"><a name="Page_211" id="Page_211">[Pg 211]</a></span></p> + +<span class="smcap">Howard, H.</span><br /> +<div class="reference">1929. The avifauna of the Emeryville shellmound. Univ. California Publ. Zool., 32:301-394, 3 pls., 54 figs.</div><br /> + +<span class="smcap">Hudson, G. E.</span><br /> +<div class="reference">1937. Studies on the muscles of the pelvic appendage in birds. Amer. Mid. Nat., 18:1-108, 26 pls.</div><br /> + +<span class="smcap">Irwin, M. R.</span><br /> +<div class="reference">1953. Evolutionary patterns of antigenic substances of the blood corpuscles in Columbidae. Evol., 7:31-50.</div><br /> + +<span class="smcap">Irwin, M. R.</span>, and <span class="smcap">Cole, L. J.</span><br /> +<div class="reference">1936. Immunogenetic studies of species and of species hybrids in doves, and the separation of species-specific substances in the backcross. Jour. Exp. Zool., 73:85-108, 1 fig.</div><br /> + +<span class="smcap">Leone, C. A.</span><br /> +<div class="reference">1949. Comparative serology of some brachyuran crustacea and studies in hemocyanin correspondence. Biol. Bull., 97:273-286, 3 figs.</div> +<div class="reference">1953. Some effects of formalin on the serological activity of crustacean and mammalian sera. Jour. Immun., 70:386-392, 2 figs.</div><br /> + +<span class="smcap">Libby, R. L.</span><br /> +<div class="reference">1938. The photronreflectometer—an instrument for the measurement of turbid systems. Jour. Immun., 34:71-73, 1 fig.</div><br /> + +<span class="smcap">Martin, E. P.</span>, and <span class="smcap">Leone, C. A.</span><br /> +<div class="reference">1952. Serological relationships among domestic fowl as shown by comparisons of protein preparations from corresponding organ systems. Trans. Kansas Acad. Sci., 55:439-444, 1 fig.</div><br /> + +<span class="smcap">McGibbon, W. H.</span><br /> +<div class="reference">1945. Further division of contrasting antigens in species hybrids in ducks. Genetics, 30:252-265.</div> + +<span class="smcap">Sasaki, K.</span><br /> +<div class="reference">1928. Serological examination of the blood-relationship between wild and domestic ducks. Jour. Dept. Agri., Kyushu Imp. Univ., 2:117-132.</div><br /> + +<span class="smcap">Simpson, G. G.</span><br /> +<div class="reference">1944. Tempo and mode in evolution. Columbia Univ. Press, New York, xviii + 237 pp., 36 figs.</div><br /> + +<span class="smcap">Sushkin, P. P.</span><br /> +<div class="reference">1924. [On the Fringillidae and allied groups.] Bull. British Ornith. Club, 45:36-39.</div> + +<div class="reference">1925. The evening grosbeak (Hesperiphona), the only American genus of a Palaearctic group. Auk, 42:256-261, 2 figs.</div><br /> + +<span class="smcap">Tordoff, H. B.</span><br /> +<div class="reference">1954. A systematic study of the avian family Fringillidae, based on the structure of the skull. Univ. Michigan Mus. Zool. Misc. Publ. No. 81:1-42, 77 figs.</div><br /> + +<span class="smcap">Wetmore, A.</span><br /> +<div class="reference">1951. A revised classification for the birds of the world. Smithsonian Misc. Coll., 117(4):1-22.</div><br /> + +<p> </p> + +<p> +<i>Transmitted June 8, 1954.</i><br /> +</p> +<p> </p> + +<div class="center"> +<img src="images/square.png" width="16" height="17" title="square" alt="square" /><br /> +25-4632 +</div> +<p> </p> +<p> </p> + + +<p><span class="pagenum"><a name="Page_i" id="Page_i">[Pg i]</a></span></p> + +<div class="caption2">UNIVERSITY OF KANSAS PUBLICATIONS<br> +MUSEUM OF NATURAL HISTORY</div> + +<p>Institutional libraries interested in publications exchange may obtain this +series by addressing the Exchange Librarian, University of Kansas Library, +Lawrence, Kansas. Copies for individuals, persons working in a particular +field of study, may be obtained by addressing instead the Museum of Natural +History, University of Kansas, Lawrence, Kansas. There is no provision for +sale of this series by the University Library, which meets institutional requests, +or by the Museum of Natural History, which meets the requests of individuals. +Nevertheless, when individuals request copies from the Museum, 25 cents should +be included, for each separate number that is 100 pages or more in length, for +the purpose of defraying the costs of wrapping and mailing.</p> + +<p>* An asterisk designates those numbers of which the Museum's supply (not the Library's +supply) is exhausted. Numbers published to date, in this series, are as follows:</p> + +<table summary="UKMNH_Pubs"> +<tr><td class="text_rt vtop">Vol. 1.</td><td class="text_rt vtop">1.</td><td class="justify">The pocket gophers (Genus Thomomys) of Utah. By Stephen D. Durrant. +Pp. 1-82, 1 figure in text; August 15, 1946.</td></tr> + +<tr><td> </td><td class="text_rt vtop">2.</td><td class="justify">The systematic status of Eumeces pluvialis Cope, and noteworthy records of other amphibians and reptiles from Kansas and Oklahoma. By Hobart M. Smith. Pp. 85-89. August 15, 1946.</td></tr> + +<tr><td> </td><td class="text_rt vtop">3.</td><td class="justify">The tadpoles of Bufo cognatus Say. By Hobart M. Smith. Pp. 93-96, 1 figure in text. August 15, 1946.</td></tr> + +<tr><td> </td><td class="text_rt vtop">4.</td><td class="justify">Hybridization between two species of garter snakes. By Hobart M. Smith. Pp. 97-100. August 15, 1946.</td></tr> + +<tr><td> </td><td class="text_rt vtop">5.</td><td class="justify">Selected records of reptiles and amphibians from Kansas. By John Breukelman and Hobart M. Smith. Pp. 101-112. August 15, 1946.</td></tr> + +<tr><td> </td><td class="text_rt vtop">6.</td><td class="justify">Kyphosis and other variations in soft-shelled turtles. By Hobart M. Smith. Pp. 117-124, 3 figures in text. July 7, 1947.</td></tr> + +<tr><td> </td><td class="text_rt vtop">*7.</td><td class="justify">Natural history of the prairie vole (Mammalian Genus Microtus). By E. W. Jameson, Jr. Pp. 125-151, 4 figures in text. October 6, 1947.</td></tr> + +<tr><td> </td><td class="text_rt vtop">8.</td><td class="justify">The postnatal development of two broods of great horned owls (Bubo virginianus). By Donald F. Hoffmeister and Henry W. Setzer. Pp. 157-173, 5 figures in text. October 6, 1947.</td></tr> + +<tr><td> </td><td class="text_rt vtop">9.</td><td class="justify">Additions to the list of the birds of Louisiana. By George H. Lowery, Jr. Pp. 177-192. November 7, 1947.</td></tr> + +<tr><td> </td><td class="text_rt vtop">10.</td><td class="justify">A check-list of the birds of Idaho. By M. Dale Arvey. Pp. 193-216. November 29, 1947.</td></tr> + +<tr><td> </td><td class="text_rt vtop">11.</td><td class="justify">Subspeciation in pocket gophers of Kansas. By Bernardo Villa R. and E. Raymond Hall. Pp. 217-236, 2 figures in text. November 29, 1947.</td></tr> + +<tr><td> </td><td class="text_rt vtop">12.</td><td class="justify">A new bat (Genus Myotis) from Mexico. By Walter W. Dalquest and E. Raymond Hall. Pp. 237-244, 6 figures in text. December 10, 1947.</td></tr> + +<tr><td> </td><td class="text_rt vtop">13.</td><td class="justify">Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. By Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, 1 figure in text. December 10, 1947.</td></tr> + +<tr><td> </td><td class="text_rt vtop">14.</td><td class="justify">A new pocket gopher (Thomomys) and a new spiny pocket mouse (Liomys) from Michoacán, México. By E. Raymond Hall and Bernardo Villa R. Pp. 249-256, 6 figures in text. July 26, 1948.</td></tr> + +<tr><td> </td><td class="text_rt vtop">15.</td><td class="justify">A new hylid frog from eastern Mexico. By Edward H. Taylor. Pp. 257-264, 1 figure in text. August 16, 1948.</td></tr> + +<tr><td> </td><td class="text_rt vtop">16.</td><td class="justify">A new extinct emydid turtle from the Lower Pliocene of Oklahoma. By Edwin C. Galbreath. Pp. 265-280, 1 plate. August 16, 1948.</td></tr> + +<tr><td> </td><td class="text_rt vtop">17.</td><td class="justify">Pliocene and Pleistocene records of fossil turtles from western Kansas and Oklahoma. By Edwin C. Galbreath. Pp. 281-284. August 16, 1948.</td></tr> + +<tr><td> </td><td class="text_rt vtop">18.</td><td class="justify">A new species of heteromyid rodent from the Middle Oligocene of northeastern Colorado with remarks on the skull. By Edwin C. Galbreath. Pp. 285-300, 2 plates. August 16, 1948.</td></tr> + +<tr><td> </td><td class="text_rt vtop">19.</td><td class="justify">Speciation in the Brazilian spiny rats (Genus Proechimys, Family Echimyidae). By João Moojen. Pp. 301-406, 140 figures in text. December 10, 1948.</td></tr> + +<tr><td> </td><td class="text_rt vtop">20.</td><td class="justify">Three new beavers from Utah. By Stephen D. Durrant and Harold S. Crane. Pp. 407-417, 7 figures in text. December 24, 1948.</td></tr> + +<tr><td> </td><td class="text_rt vtop">21.</td><td class="justify">Two new meadow mice from Michoacán, Mexico. By E. Raymond Hall. Pp. 423-427, 6 figures in text. December 24, 1948.</td></tr> + +<tr><td> </td><td class="text_rt vtop">22.</td><td class="justify">An annotated check list of the mammals of Michoacán, Mexico. By E. Raymond Hall and Bernardo Villa R. Pp. 431-472, 2 plates, 1 figure in text. December 27, 1949.</td></tr> + +<tr><td> </td><td class="text_rt vtop">23.</td><td class="justify">Subspeciation in the kangaroo rat, Dipodomys ordii. By Henry W. Setzer. Pp. 473-573, 27 figures in text, 7 tables. December 27, 1949.</td></tr> + +<tr><td> </td><td class="text_rt vtop">24.</td><td class="justify">Geographic range of the hooded skunk, Mephitis macroura, with description of a new subspecies from Mexico. By E. Raymond Hall and Walter W. Dalquest. Pp. 575-580, 1 figure in text. January 20, 1950.</td></tr> + +<tr><td> <span class="pagenum"><a name="Page_ii" id="Page_ii">[Pg ii]</a></span></td><td class="text_rt vtop">25.</td><td class="justify">Pipistrellus cinnamomeus Miller 1902 referred to the Genus Myotis. By E. Raymond Hall and Walter W. Dalquest. Pp. 581-590, 5 figures in text. January 20, 1950.</td></tr> + +<tr><td> </td><td class="text_rt vtop">26.</td><td class="justify">A synopsis of the American bats of the Genus Pipistrellus. By E. Raymond Hall and Walter W. Dalquest. Pp. 591-602, 1 figure in text. January 20, 1950.</td></tr> + +<tr><td> </td><td colspan=2 class="justify">Index. Pp. 605-638.</td></tr> + + +<tr><td class="text_rt vtop">*Vol. 2.</td><td colspan=2 class="justify" style="padding-left:2em; text-indent:-2em">(Complete) Mammals of Washington. By Walter W. Dalquest. Pp. 1-444, 140 figures in text. April 9, 1948.</td></tr> + + +<tr><td class="text_rt vtop">Vol. 3.</td><td class="text_rt vtop">*1.</td><td class="justify">The avifauna of Micronesia, its origin, evolution, and distribution. By Rollin H. Baker. Pp. 1-359, 16 figures in text. June 12, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">*2.</td><td class="justify">A quantitative study of the nocturnal migration of birds. By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. June 29, 1951.</td></tr> + + +<tr><td> </td><td class="text_rt vtop">3.</td><td class="justify">Phylogeny of the waxwings and allied birds. By M. Dale Arvey. Pp. 473-530, 49 figures in text, 13 tables. October 10, 1951.</td></tr> + + +<tr><td> </td><td class="text_rt vtop">4.</td><td class="justify">Birds from the state of Veracruz, Mexico. By George H. Lowery, Jr., and Walter W. Dalquest. Pp. 531-649, 7 figures in text, 2 tables. October 10, 1951.</td></tr> + +<tr><td> </td><td colspan=2 class="justify">Index. Pp. 651-681.</td></tr> + + +<tr><td class="text_rt vtop">*Vol. 4.</td><td colspan=2 class="justify" style="padding-left:2em; text-indent:-2em">(Complete) American weasels. By E. Raymond Hall. Pp. 1-466, 41 plates, 31 figures in text. December 27, 1951.</td></tr> + + +<tr><td class="text_rt vtop">Vol. 5.</td><td class="text_rt vtop">1.</td><td class="justify">Preliminary survey of a Paleocene faunule from the Angels Peak area, New Mexico. By Robert W. Wilson. Pp. 1-11, 1 figure in text. February 24, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">2.</td><td class="justify">Two new moles (Genus Scalopus) from Mexico and Texas. By Rollin H. Baker. Pp. 17-24. February 28, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">3.</td><td class="justify">Two new pocket gophers from Wyoming and Colorado. By E. Raymond Hall and H. Gordon Montague. Pp. 25-32. February 28, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">4.</td><td class="justify">Mammals obtained by Dr. Curt von Wedel from the barrier beach of Tamaulipas, Mexico. By E. Raymond Hall. Pp. 33-47, 1 figure in text. October 1, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">5.</td><td class="justify">Comments on the taxonomy and geographic distribution of some North American rabbits. By E. Raymond Hall and Keith R. Kelson. Pp. 49-58. October 1, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">6.</td><td class="justify">Two new subspecies of Thomomys bottae from New Mexico and Colorado. By Keith R. Kelson. Pp. 59-71, 1 figure in text. October 1, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">7.</td><td class="justify">A new subspecies of Microtus montanus from Montana and comments on Microtus canicaudus Miller. By E. Raymond Hall and Keith R. Kelson. Pp. 73-79. October 1, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">8.</td><td class="justify">A new pocket gopher (Genus Thomomys) from eastern Colorado. By E. Raymond Hall. Pp. 81-85. October 1, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">9.</td><td class="justify">Mammals taken along the Alaskan Highway. By Rollin H. Baker. Pp. 87-117, 1 figure in text. November 28, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">*10.</td><td class="justify">A synopsis of the North American Lagomorpha. By E. Raymond Hall. Pp. 119-202, 68 figures in text. December 15, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">11.</td><td class="justify">A new pocket mouse (Genus Perognathus) from Kansas. By E. Lendell Cockrum. Pp. 203-206. December 15, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">12.</td><td class="justify">Mammals from Tamaulipas, Mexico. By Rollin H. Baker. Pp. 207-218. December 15, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">13.</td><td class="justify">A new pocket gopher (Genus Thomomys) from Wyoming and Colorado. By E. Raymond Hall. Pp. 219-222. December 15, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">14.</td><td class="justify">A new name for the Mexican red bat. By E. Raymond Hall. Pp. 223-226. December 15, 1951.</td></tr> + +<tr><td> </td><td class="text_rt vtop">15.</td><td class="justify">Taxonomic notes on Mexican bats of the Genus Rhogeëssa. By E. Raymond Hall. Pp. 227-232. April 10, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">16.</td><td class="justify">Comments on the taxonomy and geographic distribution of some North American woodrats (Genus Neotoma). By Keith R. Kelson. Pp. 233-242. April 10, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">17.</td><td class="justify">The subspecies of the Mexican red-bellied squirrel, Sciurus aureogaster. By Keith R. Kelson. Pp. 243-250, 1 figure in text. April 10, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">18.</td><td class="justify">Geographic range of Peromyscus melanophrys, with description of new subspecies. By Rollin H. Baker. Pp. 251-258, 1 figure in text. May 10, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">19.</td><td class="justify">A new chipmunk (Genus Eutamias) from the Black Hills. By John A. White. Pp. 259-262. April 10, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">20.</td><td class="justify">A new piñon mouse (Peromyscus truei) from Durango, Mexico. By Robert B. Finley, Jr. Pp. 263-267. May 23, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">21.</td><td class="justify">An annotated checklist of Nebraskan bats. By Olin L. Webb and J. Knox Jones, Jr. Pp. 269-279. May 31, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">22.</td><td class="justify">Geographic variation in red-backed mice (Genus Clethrionomys) of the southern Rocky Mountain region. By E. Lendell Cockrum and Kenneth L. Fitch. Pp. 281-292, 1 figure in text. November 15, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">23.</td><td class="justify">Comments on the taxonomy and geographic distribution of North American microtines. By E. Raymond Hall and E. Lendell Cockrum. Pp. 293-312. November 17, 1952.</td></tr> + +<tr><td> <span class="pagenum"><a name="Page_iii" id="Page_iii">[Pg iii]</a></span></td><td class="text_rt vtop">24.</td><td class="justify">The subspecific status of two Central American sloths. By E. Raymond Hall and Keith R. Kelson. Pp. 313-337. November 21, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">25.</td><td class="justify">Comments on the taxonomy and geographic distribution of some North American marsupials, insectivores, and carnivores. By E. Raymond Hall and Keith R. Kelson. Pp. 319-341. December 5, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">26.</td><td class="justify">Comments on the taxonomy and geographic distribution of some North American rodents. By E. Raymond Hall and Keith R. Kelson. Pp. 343-371. December 15, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">27.</td><td class="justify">A synopsis of the North American microtine rodents. By E. Raymond Hall and E. Lendell Cockrum. Pp. 373-498, 149 figures in text. January 15, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">28.</td><td class="justify">The pocket gophers (Genus Thomomys) of Coahuila, Mexico. By Rollin H. Baker. Pp. 499-514, 1 figure in text. June 1, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">29.</td><td class="justify">Geographic distribution of the pocket mouse, Perognathus fasciatus. By J. Knox Jones, Jr. Pp. 515-526, 7 figures in text. August 1, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">30.</td><td class="justify">A new subspecies of wood rat (Neotoma mexicana) from Colorado. By Robert B. Finley, Jr. Pp. 527-534, 2 figures in text. August 15, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">31.</td><td class="justify">Four new pocket gophers of the genus Cratogeomys from Jalisco, Mexico. By Robert J. Russell. Pp. 535-542. October 15, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">32.</td><td class="justify">Genera and subgenera of chipmunks. By John A. White. Pp. 543-561, 12 figures in text. December 1, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">33.</td><td class="justify">Taxonomy of the chipmunks, Eutamias quadrivittatus and Eutamias umbrinus. By John A. White. Pp. 563-582, 6 figures in text. December 1, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">34.</td><td class="justify">Geographic distribution and taxonomy of the chipmunks of Wyoming. By John A. White. Pp. 584-610, 3 figures in text. December 1, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">35.</td><td class="justify">The baculum of the chipmunks of western North America. By John A. White. Pp. 611-631, 19 figures in text. December 1, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">36.</td><td class="justify">Pleistocene Soricidae from San Josecito Cave, Nuevo Leon, Mexico. By James S. Findley. Pp. 633-639. December 1, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">37.</td><td class="justify">Seventeen species of bats recorded from Barro Colorado Island, Panama Canal Zone. By E. Raymond Hall and William B. Jackson. Pp. 641-646. December 1, 1953.</td></tr> + +<tr><td> </td><td colspan=2 class="justify">Index. Pp. 647-676.</td></tr> + + +<tr><td class="text_rt vtop">*Vol. 6.</td><td colspan=2 class="justify" style="padding-left:2em; text-indent:-2em">(Complete) Mammals of Utah, <i>taxonomy and distribution</i>. By Stephen D. Durrant. Pp. 1-549, 91 figures in text, 30 tables. August 10, 1952.</td></tr> + + +<tr><td class="text_rt vtop">Vol. 7.</td><td class="text_rt vtop">*1.</td><td class="justify">Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, 73 figures in text, 37 tables. August 25, 1952.</td></tr> + +<tr><td> </td><td class="text_rt vtop">2.</td><td class="justify">Ecology of the opossum on a natural area in northeastern Kansas. By Henry S. Fitch and Lewis L. Sandidge. Pp. 305-338, 5 figures in text. August 24, 1953.</td></tr> + +<tr><td> </td><td class="text_rt vtop">3.</td><td class="justify">The silky pocket mice (Perognathus flavus) of Mexico. By Rollin H. Baker. Pp. 339-347, 1 figure in text. February 15, 1954.</td></tr> + +<tr><td> </td><td class="text_rt vtop">4.</td><td class="justify">North American jumping mice (Genus Zapus). By Philip H. Krutzsch. Pp. 349-472, 47 figures in text, 4 tables. April 21, 1954.</td></tr> + +<tr><td> </td><td class="text_rt vtop">5.</td><td class="justify">Mammals from Southeastern Alaska. By Rollin H. Baker and James S. Findley. Pp. 473-477. April 21, 1954.</td></tr> + +<tr><td> </td><td class="text_rt vtop">6.</td><td class="justify">Distribution of Some Nebraskan Mammals. By J. Knox Jones, Jr. Pp. 479-487. April 21, 1954.</td></tr> + +<tr><td> </td><td class="text_rt vtop">7.</td><td class="justify">Subspeciation in the montane meadow mouse, Microtus montanus, in Wyoming and Colorado. By Sydney Anderson. Pp. 489-506, 2 figures in text. July 23, 1954.</td></tr> + +<tr><td> </td><td class="text_rt vtop">8.</td><td class="justify">A new subspecies of bat (Myotis velifer) from southeastern California and Arizona. By Terry A. Vaughn. Pp. 507-512. July 23, 1954.</td></tr> + +<tr><td> </td><td class="text_rt vtop">9.</td><td class="justify">Mammals of the San Gabriel mountains of California. By Terry A. Vaughn. Pp. 513-582, 1 figure in text, 12 tables. November 15, 1954.</td></tr> + +<tr><td> </td><td colspan=2 class="justify">More numbers will appear in volume 7.</td></tr> + + +<tr><td class="text_rt vtop">Vol. 8.</td><td class="text_rt vtop">1.</td><td class="justify">Life history and ecology of the five-lined skink, Eumeces fasciatus. By Henry S. Fitch. Pp. 1-156, 26 figs. in text. September 1, 1954.</td></tr> + +<tr><td> </td><td class="text_rt vtop">2.</td><td class="justify">Myology and serology of the Avian Family Fringillidae, a taxonomic study. By William B. Stallcup. Pp. 157-211, 23 figures in text, 4 tables. November 15, 1954.</td></tr> + +<tr><td> </td><td colspan=2 class="justify">More numbers will appear in volume 8.</td></tr> +</table> + + + + + + + + + +<pre> + + + + + +End of the Project Gutenberg EBook of Myology and Serology of the Avian +Family Fringillidae, by William B. 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Stallcup + +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: Myology and Serology of the Avian Family Fringillidae + A Taxonomic Study + +Author: William B. Stallcup + +Release Date: October 19, 2010 [EBook #33914] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK MYOLOGY AND SEROLOGY OF THE *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +http://www.pgdp.net + + + + + + + + + + + + ================================================================== + UNIVERSITY OF KANSAS PUBLICATIONS + MUSEUM OF NATURAL HISTORY + + + Volume 8, No. 2, pp. 157-211, figures 1-23, 4 tables + + ---------------------- November 15, 1954 ---------------------- + + + Myology and Serology + of the Avian Family Fringillidae, + A Taxonomic Study + + BY + WILLIAM B. STALLCUP + + + UNIVERSITY OF KANSAS + LAWRENCE + 1954 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + Editors: E. Raymond Hall, Chairman, A. Byron Leonard, + Robert W. Wilson + + + Volume 8, No. 2, pp. 157-211, figures 1-23, 4 tables + Published November 15, 1954 + + + + + + UNIVERSITY OF KANSAS + Lawrence, Kansas + + + + + + PRINTED BY + FERD VOILAND, JR., STATE PRINTER + TOPEKA, KANSAS + 1954 + [Union Label] + 25-4632 + + + + + Myology and Serology + of the Avian Family Fringillidae, + A Taxonomic Study + + BY + WILLIAM B. STALLCUP + + + + +CONTENTS + + + PAGE + + INTRODUCTION 160 + + MYOLOGY OF THE PELVIC APPENDAGE 162 + General Statement 162 + Materials and Methods 163 + Description of Muscles 164 + Discussion of Myological Investigations 175 + + COMPARATIVE SEROLOGY 185 + General Statement 185 + Preparation of Antigens 186 + Preparation of Antisera 188 + Methods of Serological Testing 188 + Experimental Data 190 + Discussion of Serological Investigations 190 + + CONCLUSIONS 201 + + SUMMARY 208 + + LITERATURE CITED 210 + + + + +INTRODUCTION + + +The relationships of many groups of birds within the Order +Passeriformes are poorly understood. Most ornithologists agree that +some of the passerine families of current classifications are +artificial groups. These artificial groupings are the result of early +work which gave chief attention to readily adaptive external +structures. The size and shape of the bill, for example, have been +over-emphasized in the past as taxonomic characters. It is now +recognized that the bill is a highly adaptive structure and that it +frequently shows convergence and parallelism. + +Since studies of external morphology have failed in some cases to +provide a clear understanding of the relationships of passerine birds, +it seems appropriate that attention be given to other morphological +features, to physiological features, and to life history studies in an +attempt to find other clues to relationships at the family and +subfamily levels. + +This paper reports the results of a study of the relationships of some +birds of the Family Fringillidae and is based on the comparative +myology of the pelvic appendage and on the comparative serology of +saline-soluble proteins. Where necessary for comparative purposes, +birds from other families have been included in these investigations. + +It has long been recognized that the Fringillidae include dissimilar +groups. Recent work by Beecher (1951b, 1953) on the musculature of the +jaw and by Tordoff (1954) primarily on the structure of the bony +palate has emphasized the artificial nature of the assemblage although +these authors disagree regarding major divisions within it (see +below). + +The Fringillidae have been distinguished from other families of +nine-primaried oscines by only one character--a heavy and conical bill +(for crushing seeds). Bills of this form have been developed +independently in several other, unrelated, groups; as Tordoff (1954:7) +has pointed out, _Molothrus_ of the Family Icteridae, _Psittorostra_ +of the Family Drepaniidae, and most members of the Family Ploceidae +have bills as heavy and conical as those of the fringillids. The +ploceids are distinguished from the fringillids by a single external +character: a fairly well-developed tenth primary whereas in +fringillids the tenth primary is absent or vestigial. Tordoff +(1954:20) points out, however, that this distinction is of limited +value since in other passerine families the tenth primary may be +present in some species of a genus and absent in others. The Genus +_Vireo_ is an example. Furthermore, at least one ploceid +(_Philetairus_) has a small, vestigial tenth primary, whereas some +fringillids (_Emberizoides_, for example) possess a tenth primary +which is rather large and ventrally placed (Chapin, 1917:253-254). +Thus, it is obvious that studies based on other features are necessary +in order to attain a better understanding of the relationships of the +birds involved. + +Sushkin's studies (1924, 1925) of the structure of the bony and horny +palates have served as a basis for the division of the Fringillidae +into as many as five subfamilies (Hellmayr, 1938:v): Richmondeninae, +Geospizinae, Fringillinae, Carduelinae, and Emberizinae. + +Beecher (1951b:280) points out that "the richmondenine finches arise +so uninterruptedly out of the tanagers that ornithologists have had +to draw the dividing line between the two groups arbitrarily." His +study of pattern of jaw-musculature substantiates this. He states +further that the cardueline finches arise without disjunction +from the tanagers. He suggests, therefore, that the two groups of +"tanager-finches" be made subfamilies of the Thraupidae and that a +third subfamily be maintained for the more typical tanagers. He states +that the emberizine finches are of different origin, arising from the +wood warblers (1953:307). Beecher (1951a:431; 1953:309) includes the +Dickcissel, _Spiza americana_, in the Family Icteridae, chiefly on the +basis of jaw muscle-pattern and the horny palate. + +Tordoff (1954:10-11) presents evidence that the occurrence of +palato-maxillary bones in nine-primaried birds indicates relationship +among the forms possessing them. He points out that all fringillids +except the Carduelinae possess palato-maxillaries that are either free +or more or less fused to the prepalatine bar. He points out also that +in all carduelines, the prepalatine bar is flared at its juncture with +the premaxilla, and that the mediopalatine processes are fused across +the midline; noncardueline fringillids lack these characteristics. In +addition to the above he cites differences between the carduelines and +the "other" fringillids in the appendicular skeletons, in geographic +distribution, in patterns of migration, and in habits. Tordoff +concludes, therefore, that the carduelines are not fringillids but +ploceids, their closest affinities being with the ploceid Subfamily +Estrildinae. On the basis of palatal structure, the Fringillinae and +Geospizinae are combined with the Emberizinae, the name Fringillinae +being maintained for the subfamily. The tanagers merge with the +Richmondeninae on the one hand and with the Fringillinae on the other. +On this basis, Tordoff (1954:32) suggests that the Family Fringillidae +be divided into subfamilies as follows: Richmondeninae, Thraupinae, +and Fringillinae. The carduelines are placed as the Subfamily +Carduelinae in the Family Ploceidae. + +From the foregoing, it is apparent that the two most recent lines of +research have given rise to conflicting theories regarding +relationships within the Family Fringillidae. The purpose of my +investigation, therefore, has been to gather information, from other +fields, which might clarify the relationships of these birds. + +Since the muscle pattern of the leg in the Order Passeriformes is +thought to be one of long standing and slow change, any variation +which consistently distinguishes one group of species from another +could be significant. With the hope that such variation might be +found, a study of the comparative myology of the legs was undertaken. + +The usefulness of comparative serology as a means of determining +relationship has been demonstrated in many investigations. Its use in +this instance was undertaken for several reasons: comparative serology +has its basis in biochemical systems which seem to evolve slowly; its +methods are objective; and its use has, heretofore, resulted in the +accumulation of data which seem compatible, in most instances, with +data obtained from other sources. + +I acknowledge with pleasure the guidance received in this study from +Prof. Harrison B. Tordoff of the University of Kansas. I am indebted +also to Prof. Charles A. Leone without whose direction and assistance +the serological investigations would not have been possible; to +Professors E. Raymond Hall and A. Byron Leonard whose suggestions and +criticisms have been most helpful in the preparation of this paper; +and to T. D. Burleigh of the U. S. Fish and Wildlife Service for gifts +of several specimens used in this work. Assistance with certain parts +of the study were received from a contract (NR163014) between the +Office of Naval Research of the United States Navy and the University +of Kansas. + + + + +MYOLOGY OF THE PELVIC APPENDAGE + + +General Statement + +In an excellent paper in which the muscles of the pelvic appendage of +birds are carefully and accurately described, Hudson (1937) reviewed +briefly the more important literature pertaining to the musculature of +the leg which had been published to that date. A review of such +information here, therefore, seems unnecessary. + +Myological formulae suggested by Garrod (1873, 1874) have been +extensively used by taxonomists as aids in characterizing the orders +of birds. Relatively few investigations, however, involving the +comparative myology of the leg have been undertaken at family and +subfamily levels. The works of Fisher (1946), Hudson (1948), and +Berger (1952) are notable exceptions. + +The terminology for the muscles used in this paper follows that of +Hudson (1937), except that I have followed Berger (1952) in Latinizing +all names. Homologies are not given since these are reviewed by +Hudson. Osteological terms are from Howard (1929). + + +Materials and Methods + +Specimens were preserved in a solution of one part formalin to eight +parts of water. Thorough injection of all tissues was necessary for +satisfactory preservation. Most of the down and contour feathers were +removed to allow the preservative to reach the skin. + +In preparing specimens for study, the legs and pelvic girdle were +removed and washed in running water for several hours to remove much +of the formalin. They were then transferred to a mixture of 50 per +cent alcohol and a small amount of glycerine. + +All specimens were dissected with the aid of a low power binocular +microscope. Where possible, several specimens of each species were +examined for individual differences. Such differences were found to be +slight, involving mainly size and shape of the muscles. The size is +dependent partly on the age of the bird, muscles from older birds +being larger and better developed. The shape of a muscle (whether long +and slender or short and thick) is due in part to the position in +which the leg was preserved; that is to say, a muscle may be extended +in one bird and contracted in another. For these reasons, descriptions +and comparisons are based mainly on the origin and insertion of a +muscle and on its position in relation to adjoining muscles. + +Birds dissected in this study are listed below (in the order of the A. +O. U. Check-List): + + SPECIES + + _Vireo olivaceus_ (Linnaeus) _Leucosticte tephrocotis_ + _Seiurus motacilla_ (Vieillot) (Swainson) + _Passer domesticus_ (Linnaeus) _Spinus tristis_ (Linnaeus) + _Estrilda amandava_ (Linnaeus) _Loxia curvirostra_ Linnaeus + _Poephila guttata_ (Reichenbach) _Chlorura chlorura_ (Audubon) + _Icterus galbula_ (Linnaeus) _Pipilo erythrophthalmus_ + _Molothrus ater_ (Boddaert) (Linnaeus) + _Piranga rubra_ (Linnaeus) _Calamospiza melanocorys_ + _Richmondena cardinalis_ (Linnaeus) Stejneger + _Guiraca caerulea_ (Linnaeus) _Chondestes grammacus_ (Say) + _Passerina cyanea_ (Linnaeus) _Junco hyemalis_ (Linnaeus) + _Spiza americana_ (Gmelin) _Spizella arborea_ (Wilson) + _Hesperiphona vespertina_ (Cooper) _Zonotrichia querula_ (Nuttall) + _Carpodacus purpureus_ (Gmelin) _Passerella iliaca_ (Merrem) + _Pinicola enucleator_ (Linnaeus) _Calcarius lapponicus_ (Linnaeus) + + +Description of Muscles + +The descriptions which follow are those of the muscles in the leg of +the Red-eyed Towhee, _Pipilo erythrophthalmus_. Differences between +species, where present, are noted for each muscle. The term thigh is +used to refer to the proximal segment of the leg; the term crus is +used for that segment of the leg immediately distal to the thigh. + + +_+Musculus iliotrochantericus posticus+_ (Fig. 2).--The origin of this +muscle is fleshy from the entire concave lateral surface of the ilium +anterior to the acetabulum. The fibers converge posteriorly, and the +muscle inserts by a short, broad tendon on the lateral surface of the +femur immediately distal to the trochanter. It is the largest muscle +which passes from the ilium to the femur. + +Action.--Moves femur forward and rotates it anteriorly. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus iliotrochantericus anticus+_ (Fig. 3).--Covered laterally +by the _m. iliotrochantericus posticus_, this slender muscle +has a fleshy origin from the anteroventral edge of the ilium +between the origins of the _m. sartorius_ anteriorly and the _m. +iliotrochantericus medius_ posteriorly. The _m. iliotrochantericus +anticus_ is directed caudoventrally and inserts by a broad, flat +tendon on the anterolateral surface of the femur between the heads of +the _m. femorotibialis externus_ and _m. femorotibialis medius_ and +just distal to the insertion of the _m. iliotrochantericus medius_. + +Action.--Moves femur forward and rotates it anteriorly. + +Comparison.--No significant differences noted among the species studied. + + +_+Musculus iliotrochantericus medius+_ (Fig. 3).--Smallest of the +three _iliotrochantericus_ muscles, this bandlike muscle has a fleshy +origin from the ventral edge of the ilium just posterior to the origin +of the _m. iliotrochantericus anticus_. The fibers are directed +caudoventrally, and the insertion is tendinous on the anterolateral +surface of the femur between the insertion of the other two +_iliotrochantericus_ muscles. + +Action.--Moves femur forward and rotates it anteriorly. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus iliacus+_ (Figs. 4, 5).--Arising from a fleshy origin on +the ventral edge of the ilium just posterior to the origin of the _m. +iliotrochantericus medius_, this small slender muscle passes +posteroventrally to its fleshy insertion on the posteromedial surface +of the femur just proximal to the origin of the _m. femorotibialis +internus_. + +Action.--Moves femur forward and rotates it posteriorly. + +Comparison.--No significant differences among the species studied. + + +_+Musculus sartorius+_ (Figs. 1, 4).--A long, straplike muscle, the +_sartorius_ forms the anterior edge of the thigh. The origin is +fleshy, half from the anterior edge of the ilium and from the median +dorsal ridge of this bone and half from the posterior one or two free +dorsal vertebrae. The insertion is fleshy along a narrow line on the +anteromedial edge of the head of the tibia and on the medial region of +the patellar tendon. + +Action.--Moves thigh forward and upward and extends shank. + +Comparison.--In _Loxia_ and _Spinus_, only one-third of the origin is +from the last free dorsal vertebra. In _Hesperiphona_, _Carpodacus_, +_Pinicola_, and _Leucosticte_, only one-fifth of the origin is from +this vertebra. + + +_+Musculus iliotibialis+_ (Fig. 1).--Broad and triangular, this muscle +covers most of the deeper muscles of the lateral aspect of the thigh. +The middle region is fused with the underlying _femorotibialis_ +muscles. In the distal half of this muscle there are three distinct +parts; the anterior and posterior edges are fleshy and the central +part is aponeurotic. The origin is from a narrow line along the iliac +crests--from the origin of the _m. sartorius_, anteriorly, to the +origin of the _m. semitendinosus_ posteriorly. The origin is +aponeurotic in the preacetabular region but fleshy in the +postacetabular region. The distal part of the muscle is aponeurotic +and joins with the _femorotibialis_ muscles in the formation of the +patellar tendon. This tendon incloses the patella and inserts on a +line along the proximal edges of the cnemial crests of the +tibiotarsus. + +Action.--Extends crus. + +Comparison.--In _Vireo_ the central aponeurotic portion of this muscle +is absent. + + +_+Musculus femorotibialis externus+_ (Fig. 2).--Covering the lateral +and anterolateral surfaces of the femur, this large muscle has a +fleshy origin from the lateral edge of the proximal three-fourths of +the femur. The origin separates the insertion of the _m. +iliotrochantericus anticus_ from that of the _m. ischiofemoralis_ and, +in turn, is separated from the origin of the _m. femorotibialis +medius_ by the insertions of the _m. iliotrochantericus anticus_ and +_m. iliotrochantericus medius_. Approximately midway of the length of +the femur this muscle fuses anteromesially with the _m. femorotibialis +medius_. Distally, the _m. femorotibialis externus_ contributes to the +formation of the patellar tendon which inserts on a line along the +proximal edges of the cnemial crests of the tibiotarsus. + +Action.--Extends crus. + +Comparison.--No significant differences noted among the species studied. + + +_+Musculus femorotibialis medius+_ (Figs. 2, 4).--The origin of this +muscle, which lies along the anterior edge of the femur, is fleshy +from the entire length of the femur proximal to the level of +attachment of the proximal arm of the biceps loop. Laterally this +muscle is completely fused for most of its length with the _m. +femorotibialis externus_ and contributes to the formation of the +patellar tendon, which inserts on a line along the proximal edges of +the cnemial crests of the tibiotarsus. Many of the fibers, +nevertheless, insert on the proximal edge of the patella. + +Action.--Extends crus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus femorotibialis internus+_ (Fig. 4).--One of the most +superficial muscles lying on the medial surface of the thigh, this +muscle is divided, especially near the distal end, into two parts, +lateral and medial. The origin of the lateral part is fleshy from a +line on the medial surface of the femur; the origin begins proximally +at a point near the insertion of the _m. iliacus_. The medial, bulkier +part of the muscle has a fleshy origin on the medial surface of the +lower one-third of the femur. The two parts fuse to some extent above +the points of insertion and insert on the medial edge of the head of +the tibia. + +Action.--Rotates tibia anteriorly. + +Comparison.--Two parts of this muscle variously fused; otherwise, no +significant differences in the species studied. + + +_+Musculus piriformis+_ (Fig. 3).--This muscle is represented by the +_pars caudifemoralis_ only, the _pars iliofemoralis_ being absent in +passerine birds as far as is known. The _pars caudifemoralis_ is flat, +somewhat spindle-shaped, and passes anteroventrally from the pygostyle +to the femur. The origin is tendinous from the anteroventral edge of +the pygostyle, and the insertion is semitendinous on the +posterolateral surface of the shaft of the femur about one-fourth its +length from the proximal end. + +Action.--Moves femur posteriorly and rotates it in this direction; +moves tail laterally and depresses it. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus semitendinosus+_ (Figs. 2, 3, 5).--The origin from the +extreme posterior edge of the posterior iliac crest of the ilium is +fleshy and is aponeurotic from the last vertebra of the synsacrum and +the transverse processes of several caudal vertebrae. The straplike +belly passes along the posterolateral margin of the thigh. Immediately +posterior to the knee, the muscle is divided transversely by a +ligament. That portion passing anteriorly from the ligament is the _m. +accessorius semitendinosi_ (here considered a part of the _m. +semitendinosus_) and is discussed below. The ligament continues +distally in two parts; one part inserts on the medial surface of the +_pars media_ of the _m. gastrocnemius_ and the other part fuses with +the tendon of insertion of the _m. semimembranosus_. + +The _m. accessorius semitendinosi_ extends anteriorly from the above +mentioned ligament to a fleshy insertion on the posterolateral surface +of the femur immediately proximal to the condyles. + +Action.--Moves femur posteriorly, flexes the crus and aids in +extending the tarsometatarsus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus semimembranosus+_ (Figs. 3, 4, 5).--This straplike muscle +passes along the posteromedial surface of the thigh. The origin is +semitendinous along a line on the ischium, from a point dorsal to the +middle of the ischiopubic fenestra to the posterior end of the +ischium, and from a small area of the abdominal musculature posterior +to the ischium. The insertion is by means of a broad, thin tendon on a +ridge on the medial surface of the tibia immediately distal to the +head of this bone. The tendon of insertion passes between the head of +the _pars media_ and _pars interna_ of the _m. gastrocnemius_ and is +fused with the tendon of the _m. semitendinosus_. + +Action.--Flexes crus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus biceps femoris+_ (Fig. 2).--Long, thin, and somewhat +triangular, this muscle lies on the lateral side of the thigh just +underneath the _m. iliotibialis_. Its origin is from a line along the +anterior and posterior iliac crests underneath the origin of the _m. +iliotibialis_. Anterior to the acetabulum the origin is aponeurotic, +and the edge of this aponeurosis passes over the proximal end of the +femur. The origin posterior to the acetabulum is fleshy. The most +anterior point of origin is difficult to ascertain but it lies near +the center of the anterior iliac crest. The most posterior point of +origin is immediately dorsal to the posterior end of the ilioischiatic +fenestra. Behind the knee the fibers of this muscle converge to form +the strong tendon of insertion which passes through the biceps loop, +under the tendon of origin of the _m. flexor perforatus digiti II_, +and inserts on a small tubercle on the posterolateral edge of the +fibula at the point of the tibia-fibula fusion. + +The biceps loop is tendinous and the distal end attaches to a +protuberance on the posterolateral edge of the femur at the proximal +edge of the external condyle. The proximal end attaches to the +anterolateral edge of the femur immediately proximal to the distal end +of the loop, which extends posterior to the femur. The distal arm of +this loop is connected with the tendon of origin of the _m. flexor +perforatus digiti II_ by a strong tendon. + +Action.--Flexes crus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus ischiofemoralis+_ (Fig. 3).--Short and thick, this muscle +arises directly from the lateral surface of the ischium between the +posterior iliac crest and the ischiopubic fenestra. The area of origin +extends to the posterior edge of the ischium. The insertion is +tendinous on the lateral surface of the trochanter opposite the +insertion of the _m. iliotrochantericus medius_. + +Action.--Moves femur posteriorly and rotates it in this direction. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus obturator internus+_ (Figs. 4, 7).--Lying on the inside of +the pelvis and covering the medial surface of the ischiopubic +fenestra, is this flat, pinnate, leaf-shaped muscle. The origin is +fleshy and is from the ischium and pubis around the edges of this +fenestra; none of the fibers arises from the membrane stretched across +the fenestra. Anteriorly the fibers converge and form a strong tendon +that passes through the obturator foramen and inserts on the +posterolateral surface of the trochanter of the femur. + +Action.--Rotates femur posteriorly. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus obturator externus+_ (Fig. 7).--Short and fleshy, this +muscle consists of two parts which are not easily separable but which +may be traced throughout its length. The parts are more nearly +distinct at the origin. The dorsal part arises directly from the +ischium along the dorsal edge of the obturator foramen. The larger +ventral part arises directly from the anterior and ventral edges of +the obturator foramen. The fibers of the dorsal part pass anteriorly, +cover the tendon of the _m. obturator internus_ laterally, and insert +on the trochanter around the point of insertion of the latter muscle. +The fibers of the ventral part pass parallel with the tendon of the +_m. obturator internus_ and insert on the trochanter immediately +distal and posterior to the tendon of the latter muscle. + +Action.--Rotates femur posteriorly. + +Comparison.--In _Passer_, _Estrilda_, _Poephila_, _Hesperiphona_, +_Carpodacus_, _Pinicola_, _Leucosticte_, _Spinus_ and _Loxia_, this +muscle is undivided and, in its position, origin, and insertion, +resembles the ventral part of the bipartite muscle described above. +The origin is from the anterior and ventral edges of the obturator +foramen and the insertion is on the trochanter of the femur +immediately distal and posterior to the insertion of the _m. obturator +internus_. In all other genera examined, the muscle is bipartite. In +_Chlorura_ the dorsal part is larger and better developed than it is +in the other genera. + + +_+Musculus adductor longus et brevis+_ (Figs. 3, 4, 5).--Consisting of +two distinct, straplike parts, this large muscle lies on the medial +surface of the thigh, posterior to the femur. + +The _pars anticus_ has a semitendinous origin on a line that extends +posteriorly from the posteroventral edge of the obturator foramen to a +point half way across the membrane that covers the ischiopubic +fenestra. The insertion is fleshy along the posterior surface of the +femur from the level of the insertion of the _m. piriformis_ distally +to the medial surface of the internal condyle. + +The _pars posticus_ originates by a broad, flat tendon on a line +across the posterior half of the membrane that covers the ischiopubic +fenestra. The insertion is at the point of origin of the _pars media_ +of the _m. gastrocnemius_ on the posteromedial surface of the proximal +end of the internal condyle of the femur. There is a broad tendinous +connection with the proximal end of the _pars media_ of the _m. +gastrocnemius_. The anterior edge of the _pars posticus_ is overlapped +medially by the posterior edge of the _pars anticus_. + +Action.--Flexes thigh; may flex crus also and may extend +tarsometatarsus. + +Comparison.--In _Vireo olivaceous_, the origin of this muscle does not +extend the length of the ischiopubic fenestra. The origin, +furthermore, is along the dorsal edge of the ischiopubic fenestra and +not from the membrane covering the fenestra. Finally, in this species, +the origin of the _pars posticus_ is fleshy. + + +_+Musculus tibialis anticus+_ (Figs. 2, 5).--Lying along the anterior +edge of the crus, a part of this muscle is covered by the _m. peroneus +longus_. The origin is by two distinct heads, each of which is +pinnate. The anterior head arises directly from the edges of the outer +and inner cnemial crests. The posterior head arises by a short, strong +tendon from a small pit on the anterodistal edge of the external +condyle of the femur. This tendon and the proximal end of the muscle +pass between the head of the fibula and the outer cnemial crest. The +two heads of the muscle fuse at a place slightly more than one-half of +the distance down the crus. At the distal end of the crus this muscle +gives rise to a strong tendon which passes under a fibrous loop +immediately proximal to the external condyle in company with the _m. +extensor digitorum longus_ and which passes between the condyles of +the tibia and inserts on a tubercle on the anteromedial edge of the +proximal end of the tarsometatarsus. + +Action.--Flexes tarsometatarsus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus extensor digitorum longus+_ (Figs. 3, 5, 8).--Slender and +pinnate, this muscle lies along the anteromedial surface of the tibia. +The origin is fleshy from most of the region between the cnemial +crests and from a line along the anterior surface of the proximal +fourth of the tibia. Approximately two-thirds of the distance down the +crus the muscle gives rise to the tendon of insertion which passes +through the fibrous loop near the distal end of the tibia in company +with the _m. tibialis anticus_. The tendon then passes along beneath +the supratendinal bridge at the distal end of the tibia, traverses the +anterior intercondylar fossa, and passes beneath a bony bridge on the +anteromedial surface of the proximal end of the tarsometatarsus. The +tendon continues along the anterior surface of the tarsometatarsus to +a point immediately above the bases of the toes and there gives rise +to three branches, one to the anterior surface of each foretoe. The +insertions of each branch are on the anterior surfaces of the +phalanges as shown in Fig. 8. + +Action.--Extends foretoes. + +Comparison.--This muscle is weakly developed in _Leucosticte_ and +_Calvarius_; the belly is slender and extends only half way down the +crus before giving rise to the tendon of insertion. The functional +significance of this variation is difficult to understand. The +convergence in muscle pattern shown by these two genera, however, is +in all probability the result of similarities in behavior patterns. +These birds perch less frequently than do the other birds studied. +Thus, the toes are neither flexed nor extended as often; the smaller +size of the _m. extensor digitorum longus_ may have resulted in part +from this lessened activity. Except for the variations just noted, +there are no significant differences among the species studied; even +the rather complex patterns of insertion are identical. + + +_+Musculus peroneus longus+_ (Fig. 1).--Relatively thin and straplike, +this muscle lies on the anterolateral surface of the crus and is +intimately attached to the underlying muscles. The part of the origin +from the proximal edges of the inner and outer cnemial crests is +semitendinous but the part of the origin from the lateral edge of the +shaft of the fibula is tendinous. Approximately two-thirds the +distance down the crus the muscle gives rise to the tendon of +insertion. Immediately above the external condyle of the tibiotarsus +this tendon divides. The posterior branch inserts on the proximal end +of the lateral edge of the tibial cartilage. The anterior branch +passes over the lateral surface of the external condyle to the +posterior surface of the tarsometatarsus and there unites with the +tendon of the _m. flexor perforatus digiti III_. + +Action.--Extends tarsometatarsus and flexes third digit. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus peroneus brevis+_ (Figs. 2, 3).--Lying along the +anterolateral surface of the tibia, this slender, pinnate muscle +arises from a fleshy origin along this surface and along the anterior +surface of the fibula from a point immediately proximal to the +insertion of the _m. biceps femoris_ to a point approximately +two-thirds of the way down the crus. Near the distal end of the tibia +the muscle gives rise to the tendon of insertion that passes through a +groove on the anterolateral edge of the tibia just above the external +condyle. Here the tendon is held in place by a broad fibrous loop and +passes under the anterior branch of the tendon of insertion of the _m. +peroneus longus_ and inserts on a prominence on the lateral edge of +the proximal end of the tarsometatarsus. + +Action.--Extends tarsometatarsus and may abduct it slightly. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus gastrocnemius+_ (Figs. 1, 4).--The largest muscle of the +pelvic appendage, it covers superficially all of the posterior +surface, most of the medial surface, and half of the lateral surface +of the crus. The muscle originates by three distinct heads. + +The _pars externa_ covers the posterolateral surface of the crus, is +intermediate in size between the other two heads, and arises by a +short, strong tendon from a small bony protuberance on the +posterolateral side of the distal end of the femur immediately +proximal to the fibular condyle. The tendon is intimately connected +with the distal arm of the loop for the _m. biceps femoris_. + +The _pars media_ is the smallest of the three heads and lies on the +medial surface of the crus. The head of the _pars media_ is separated +from the _pars interna_ by the tendon of insertion of the _m. +semimembranosus_ and originates by a short, strong tendon from the +posteromedial surface of the proximal end of the internal condyle of +the femur. The proximal portion of the _pars media_ has tendinous +connections with the tendon of the _m. semitendinosus_ and with the +_pars posticus_ of the _m. adductor longus et brevis_. + +The _pars interna_ is the largest of the three heads and covers most +of the medial surface of the crus. This head in its proximal portion +is distinctly divided into anterior and posterior parts, the former +overlapping the latter medially. The origin of the posterior part is +fleshy from the anterior half of the tibial head. Some of the fibers +of the anterior part arise directly from the inner cnemial crest while +its remaining fibers arise from the patellar tendon (Fig. 1) and form +a band that extends around the anterior surface of the knee, covering +the insertion of the _m. sartorius_. + +Approximately half way down the crus, the three heads give rise to the +tendon of insertion, the _tendo achillis_, which passes over and is +tightly bound to the posterior surface of the tibial cartilage. The +insertion is tendinous on the posterior surface of the hypotarsus and +along the posterolateral ridge of the tarsometatarsus. This tendon +seems to be continuous with a fascia which forms a sheath around the +posterior surface of the tarsometatarsus holding the other tendons of +this region firmly in the posterior sulcus. + +Action.--Extends tarsometatarsus. + +Comparison.--Study of the _pars externa_ and _pars media_ reveals no +significant differences among the species dissected. The _pars +interna_, however, is subject to some variation which is described +below. + + _Pars interna_ bipartite + + _Vireo_ _Chlorura_ + _Seiurus_ _Pipilo_ + _Icterus_ _Calamospiza_ + _Molothrus_ _Chondestes_ + _Piranga_ _Junco_ + _Richmondena_ _Spizella_ + _Guiraca_ _Zonotrichia_ + _Passerina_ _Passerella_ + _Spiza_ _Calcarius_ + +The two parts of the _m. gastrocnemius_ are most distinct in _Vireo_. +_Icterus_, _Molothrus_, _Richmondena_, _Guiraca_, and _Passerina_ lack +the fibrous band that passes around the front of the knee. In _Spiza_ +this band of fibers is smaller than in the other species. + + _Pars interna_ undivided + + _Passer_ _Pinicola_ + _Estrilda_ _Leucosticte_ + _Poephila_ _Spinus_ + _Hesperiphona_ _Loxia_ + _Carpodacus_ + +In _Leucosticte_, although the _pars interna_ is undivided, there is a +band of fibers which extends around the front of the knee (see +discussion, p. 183). + + +_+Musculus plantaris+_ (Fig. 5).--Small and slender, this muscle lies +on the posteromedial surface of the crus, beneath the _pars interna_ +of the _m. gastrocnemius_ and originates by fleshy fibers from the +posteromedial surface of the proximal end of the tibia immediately +distal to the internal articular surface. The belly extends +approximately one-sixth of the way down the crus and gives rise to a +long, slender tendon that inserts on the proximomedial edge of the +tibial cartilage. + +Action.--Extends tarsometatarsus. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus flexor perforatus digiti II+_ (Figs. 3, 9).--This is a +slender muscle which lies on the lateral side of the crus beneath the +_pars externa_ of the _m. gastrocnemius_ and is intimately connected +anteromedially with the _m. flexor digitorum longus_ and +posteromedially with the _m. flexor hallucis longus_. The origin is by +a strong tendon from the lateral surface of the external condyle of +the femur at the point of origin of the _m. flexor perforans et +perforatus digiti II_. This tendon serves also as the origin of the +anterior head of the _m. flexor hallucis longus_. The tendon connects +also by a broad tendinous band with the distal arm of the loop for the +_m. biceps femoris_ and by a similar band with the lateral edge of the +fibula immediately distal to the head. The tendon of insertion passes +distally, perforates the tibial cartilage near its lateral edge, +traverses the middle medial canal of the hypotarsus (Fig. 6), and +passes distally to the foot. At the distal end of the tarsometatarsus +the tendon is held against the medial surface of the first metatarsal +by a straplike sheath. The tendon then passes over a sesamoid bone +between the first metatarsal and the base of the second digit and is +bound to this bone by a sheath. The tendon inserts mainly along the +posteromedial edge of the proximal end of the first phalanx of the +second digit, although the termination is sheathlike and covers the +entire posterior surface of this phalanx. This sheathlike termination +is perforated by the tendons of the _m. flexor perforans et perforatus +digiti II_ and the branch of the _m. flexor digitorum longus_ that +inserts on the second digit. + +Action.--Flexes second digit. + +Comparison.--In _Vireo_ this muscle is larger and more deeply situated +than it is in the other species examined and has no connection with +the _m. flexor hallucis longus_. + + +_+Musculus flexor perforatus digiti III+_ (Fig. 5).--Long and +flattened, this muscle lies on the posteromedial side of the crus +beneath the _m. gastrocnemius_. The belly is tightly fused laterally +with the belly of the _m. flexor hallucis longus_ and posteriorly with +the belly of the _m. flexor perforatus digiti IV_. The origin is by a +long, strong tendon from a small tubercle just medial to, and at the +proximal end of, the external condyle of the femur. Below the middle +of the crus this muscle terminates in a strong tendon which perforates +the tibial cartilage near its lateral edge. In this region the tendon +is sheathlike and wrapped around the tendon of the _m. flexor +perforatus digiti IV_. These two tendons together pass through the +posterolateral canal of the hypotarsus (Fig. 6). Immediately distal to +the hypotarsus the two tendons separate, and the tendon of the _m. +flexor perforatus digiti III_ receives a branch of the tendon of the +_m. peroneus longus_. The tendon passes distally over the surface of +the second trochlea, and its insertion is sheathlike on the posterior +surface of the first phalanx, and on the proximal end of the second. +In the area of insertion this tendon is perforated by that of the _m. +flexor perforans et perforatus digiti III_ and by that of the _m. +flexor digitorum longus_ to the third digit. + +Action.--Flexes digit III. + +Comparison.--In _Passer_, _Estrilda_, _Poephila_, _Hesperiphona_, +_Carpodacus_, _Pinicola_, _Leucosticte_, _Spinus_, and _Loxia_ the +edges of the sheathlike tendon are thickened at the points of +insertion, so that the tendon appears to have two branches which +insert along the posterolateral edges of the first phalanx and are +connected medially by a fascia. + + +_+Musculus flexor perforatus digiti IV+_ (Fig. 3).--Extending along +the posterior edge of the crus, this slender muscle lies beneath the +_m. gastrocnemius_. The belly is fused with those of the _m. flexor +hallucis longus_ and _m. flexor perforatus digiti III_. Its origin is +fleshy from the intercondyloid region of the distal end of the femur +and has a few fibers arising from the tendon of origin of the _m. +flexor perforatus digiti III_. Near the distal end of the crus the +muscle gives rise to the strong tendon of insertion which perforates +the tibial cartilage near its lateral edge and in this region is +ensheathed by the tendon of the _m. flexor perforatus digiti III_. The +two tendons pass together through the posterolateral canal of the +hypotarsus (Fig. 6). The tendon continues distally along the +tarsometatarsus and the posterior surface of digit IV. The tendon +bifurcates at approximately the middle of the first phalanx. A short +lateral branch inserts on the posterolateral edge of the proximal end +of the second phalanx. The long medial branch is perforated by a +branch of the _m. flexor digitorum longus_; the distal end is +flattened, has thickened edges, and inserts over the posterior +surfaces of the distal end of the second phalanx, and over the +proximal end of the third phalanx. + +Action.--Flexes digit IV. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus flexor perforans et perforatus digiti II+_ (Figs. 2, +9).--Small and spindle-shaped, this muscle lies on the posterolateral +side of the crus immediately beneath the _pars externa_ of the _m. +gastrocnemius_. The origin is fleshy and arises in company with the +_m. flexor perforans et perforatus digiti III_ from a point on the +posterolateral surface of the distal end of the femur between the +point of origin of the _pars externa_ of the _m. gastrocnemius_ and +the fibular condyle. The belly extends approximately one-fourth of the +way down the crus and gives rise to the tendon of insertion which +passes distally and superficially through the posterior edge of the +tibial cartilage. The tendon traverses the posteromedial canal of the +hypotarsus (Fig. 6) and continues along the posterior surface of the +tarsometatarsus. Between the first metatarsal and the base of the +second digit the tendon is enclosed by the medial surface of a +sesamoid bone. This tendon then perforates that of the _m. flexor +perforatus digiti II_ at the level of the first phalanx and in turn is +perforated by the tendon of the _m. flexor digitorum longus_ at the +proximal end of the second phalanx. The insertion is on the posterior +surface of the second phalanx. + +Action.--Flexes digit II. + +Comparison.--In _Passer_, _Estrilda_, _Poephila_, _Hesperiphona_, +_Carpodacus_, _Pinicola_, _Leucosticte_, _Spinus_, and _Loxia_ the +proximal portion of this muscle is more intimately connected with the +posterior edge of the _m. flexor perforans et perforatus digiti III_ +than it is in the other species examined. + + +_+Musculus flexor perforans et perforatus digiti III+_ (Fig. 2).--Long +and pinnate, this muscle lies on the lateral surface of the crus +beneath the _m. peroneus longus_ and _pars externa_ of the _m. +gastrocnemius_. There are two distinct heads. The origin of the +anterior head is fleshy from the proximal edge of the outer cnemial +crest and from the internal edge of the distal end of the patellar +tendon. The posterior head arises by a tendon from the femur in +company with the _m. flexor perforans et perforatus digiti II_, is +connected also with the tendon of origin of the _m. flexor perforatus +digiti II_, and is loosely attached to the head of the fibula. Fibers +from the belly of the muscle attach throughout its length to the +lateral edge of the fibula, and the muscle is tightly fused also with +adjacent muscles. The tendon of insertion is formed approximately +one-half the way down the crus. The tendon perforates the posterior +surface of the tibial cartilage and passes through the posteromedial +canal of the hypotarsus (Fig. 6). At the base of the third digit the +tendon ensheathes that of the _m. flexor digitorum longus_ and the two +together perforate the tendon of the _m. flexor perforatus digiti +III_. Immediately distal to this perforation the tendon of the _m. +flexor perforans et perforatus digiti III_ ceases to ensheath that of +the _m. flexor digitorum longus_. The latter passes beneath that of +the former. Near the distal end of the second phalanx the tendon of +the _m. flexor digitorum longus_ perforates that of the _m. flexor +perforans et perforatus digiti III_. The latter inserts on the +posterior surface of the distal end of the second phalanx and the +proximal end of the third. + +Action.--Flexes digit III. + +Comparison.--In _Passer_, _Estrilda_, and _Poephila_, and in all the +cardueline finches examined the proximal portion of this muscle is +more intimately connected with the anterior edge of the _m. flexor +perforans et perforatus digiti II_ than it is in the other species +examined. + + +_+Musculus flexor digitorum longus+_ (Figs. 3, 5).--This strong, +pinnate muscle is deeply situated along the posterior surfaces of the +tibia and fibula. There are two distinct heads of origin. The lateral +head arises by means of fleshy fibers from the posterior edge of the +head of the fibula. The medial head arises by means of fleshy fibers +from the region under the ledgelike external and internal articular +surfaces of the proximal end of the tibia. Neither head has any +connection with the femur in contrast to the condition, described by +Hudson (1937: 46-47) in the crow, _Corvus brachyrhynchos_, and in the +raven, _Corvus corax_. Near the point of insertion of the _m. biceps +femoris_ the two heads fuse. The common belly is attached by fleshy +fibers to the posterior surface of the tibia and fibula for two-thirds +of the distance down the crus. Near the distal end of the crus the +muscle terminates in a strong tendon which passes deeply through the +tibial cartilage and traverses the anteromedial canal of the +hypotarsus (Fig. 6). About midway down the tarsometatarsus this tendon +becomes ossified. Immediately above the bases of the toes it gives +rise to three branches, one to the posterior surface of each of the +foretoes. These branches perforate the other flexor muscles of the +toes as described in the accounts of those muscles and insert as +follows: The branch to digit II inserts on the base of the ungual +phalanx and by a stout, tendinous slip on the distal end of the second +phalanx (Fig. 9). The branch to digit III inserts on the base of the +distal end of the third phalanx and a stronger slip to the distal end +of the second or proximal end of the third. The branch to digit IV +inserts on the base of the ungual phalanx, with one tendinous slip to +the distal end of the third phalanx and another to the distal end of +the fourth. + +Action.--Flexes foretoes. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus flexor hallucis longus+_ (Fig. 3).--Situated immediately +posterior to the _m. flexor digitorum longus_, the belly of this +large, pinnate muscle is intimately connected anteriorly to that of +the _m. flexor perforatus digiti II_. The _m. flexor hallucis longus_ +arises by two heads which are separated by the tendon of insertion of +the _m. biceps femoris_. The smaller anterior head arises from the +same tendon as does the _m. flexor perforatus digiti II_. The larger +posterior head arises by means of fleshy fibers from the +intercondyloid region of the posterior surface of the femur along with +the _m. flexor perforatus digiti III_ and _IV_. The two heads join +just distal to the point of insertion of the _m. biceps femoris_. +There is no trace of a tendinous band connecting the two heads as +there is in the crow and in the raven (Hudson, 1937:49). Near the +distal end of the shank the muscle gives rise to a strong tendon which +perforates the tibial cartilage along its lateral edge and passes +through the anterolateral canal of the hypotarsus (Fig. 6). The tendon +crosses over to the medial surface of the tarsometatarsus, passes +distally, and perforates the sheathlike tendon of the _m. flexor +hallucis brevis_ between the first metatarsal and the trochlea for +digit II. The tendon continues along the posterior surface of the +hallux and has a double insertion; the main tendon attaches to the +base of the ungual phalanx and a smaller branch inserts on the distal +end of the proximal phalanx. + +Action.--Flexes hallux. + +Comparison.--In _Vireo_ this muscle has only the posterior head of +origin and is not connected with the _m. flexor perforatus digiti II_. +The muscle is proportionately smaller and weaker than in any of the +other species studied. + + +_+Musculus extensor hallucis longus+_ (Fig. 4).--One of the smallest +muscles of the leg, the origin is fleshy from the anteromedial edge of +the proximal end of the tarsometatarsus. The belly is long and slender +and terminates distally in a slender tendon which passes distally +along the posterior surfaces of the first metatarsal and the first +digit. The insertion is on the base of the ungual phalanx. Near the +distal end of the proximal phalanx, the tendon passes between two +thick bands of fibro-elastic tissue which insert also on the ungual +phalanx. These bands of tissue function as automatic extensors of the +claw. + +Action.--Extends hallux; action must be slight. + +Comparison.--In _Vireo_ this muscle is proportionately larger and +better developed than it is in any of the other species examined. + + +_+Musculus flexor hallucis brevis+_ (Fig. 4).--This minute muscle has +a fleshy origin from the medial surface of the hypotarsus. The short +belly terminates in a weak, slender tendon which passes down the +posteromedial surface of the tarsometatarsus and into the space +between the first metatarsal and the trochlea for digit II. In this +region the tendon envelops the tendon of the _m. flexor hallucis +longus_ and inserts on the distal end of the first metatarsal and on +the proximal end of the first phalanx of the first digit. + +Action.--Flexes hallux; action must be slight. + +Comparison.--The small size of this muscle makes it exceedingly +difficult to study. The muscle is larger in _Vireo_ than in any of the +other species examined. This may be correlated with the smaller size +of the _m. flexor hallucis longus_ in this species. The muscle does +not seem to be so well developed in the cardueline finches as it is in +the other species. + + +_+Musculus abductor digiti IV+_ (Fig. 2).--Extremely small, delicate +and difficult to demonstrate, this muscle arises in a fleshy origin +immediately from underneath the posterior edge of the external cotyla +of the tarsometatarsus. The tendon of insertion is long and slender +and inserts along the lateral edge of the first phalanx of digit IV. + +Action.--Abducts digit IV. + +Comparison.--No significant differences noted among the species +studied. + + +_+Musculus lumbricalis.+_--Semitendinous throughout its length, this +muscle arises from the ossified tendon of the _m. flexor digitorum +longus_ at a point immediately proximal to the branching of this +tendon. The insertion is on the joint pulleys and capsules at the base +of the third and fourth digits. + +Action.--Hudson (1937:57) states that: "Meckel (_vide_ Gadow--1891, p. +204) considered this muscle as serving to draw the joint pulley behind +in order to protect it from pinching during the bending of the toes. +It perhaps also tends to flex the third and fourth digits." + +Comparison.--No significant differences noted among the species +studied. + + +Discussion of the Myological Investigations + +Simpson (1944:12) and others have emphasized that different parts of +organisms evolve at different rates. Beecher (1951b:275) in stating +that "... the hind limb is very similar in muscle pattern throughout +the Order Passeriformes and seems to have become relatively static +after attaining a high level of general efficiency ..." implies that +the muscle pattern of the leg must be one of long standing and slow +change. This concept was emphasized by Hudson (1937) who found but +little variation in muscle pattern among members of the several +families of passerine birds. The concept is further confirmed by the +present investigation. The intricate patterns of origin and of +insertion seem to remain almost the same throughout the order in spite +of adaptive radiation which has occurred. + +Two major differences in patterns of leg-musculature, however, were +found among the species studied, and these differences are significant +since they are consistent between subfamilies. The muscles involved +are the _m. obturator externus_ and the _pars interna_ of the _m. +gastrocnemius_. + +The _m. obturator externus_ is bipartite, consisting of dorsal and +ventral parts, in the passerine species studied by Hudson (1937) and +in all of the species examined by me except the ploceids and the +cardueline finches. In the ploceids and cardueline finches this muscle +is undivided and resembles in its position, origin, and insertion only +the ventral portion of the muscle found in the other birds studied. It +is difficult to imagine what advantage or disadvantage might be +associated with the bipartite or with the undivided condition. The +action of this muscle is to rotate the femur (right femur clockwise, +left femur counterclockwise), and certainly the greater mass of the +bipartite muscle could lend greater strength to such action. The +possible significance of this is discussed below. + + List of Abbreviations Used in Figures + + Abd. dig. IV _M. abductor digiti IV_ + Acc. _M. accessorius semitendinosi_ + Add. long. _M. adductor longus et brevis_ + Anterolat. can. Anterolateral canal of hypotarsus + Anteromed. can. Anteromedial canal of hypotarsus + Bic. fem. _M. biceps femoris_ + Bic. loop Loop for _m. biceps femoris_ + Ext. cot. External cotyla + Ext. dig. l. _M. extensor digitorum longus_ + Ext. hal. l. _M. extensor hallucis longus_ + Fem. tib. ext. _M. femorotibialis externus_ + Fem. tib. int. _M. femorotibialis internus_ + Fem. tib. med. _M. femorotibialis medius_ + F. dig. l. _M. flexor digitorum longus_ + F. hal. brev. _M. flexor hallucis brevis_ + F. hal. l. _M. flexor hallucis longus_ + F. p. et p. d. II _M. flexor perforans et perforatus digiti II_ + F. p. et p. d. III _M. flexor perforans et perforatus digiti III_ + F. per. d. II _M. flexor perforatus digiti II_ + F. per. d. III _M. flexor perforatus digiti III_ + F. per. d. IV _M. flexor perforatus digiti IV_ + Gas. _M. gastrocnemius_ + Iliacus _M. iliacus_ + Il. tib. _M. iliotibialis_ + Il. troc. ant. _M. iliotrochantericus anticus_ + Il. troc. med. _M. iliotrochantericus medius_ + Il. troc. post. _M. iliotrochantericus posticus_ + Int. cot. Internal cotyla + Isch. fem. _M. ischiofemoralis_ + Midmed. can. Midmedial canal of hypotarsus + Obt. ext. _M. obturator externus_ + Obt. int. _M. obturator internus_ + P. ant. _Pars anticus_ + P. ext. _Pars externa_ + P. int. _Pars interna_ + P. med. _Pars media_ + P. post. _Pars posticus_ + Per. brev. _M. peroneus brevis_ + Per. long. _M. peroneus longus_ + Pirif. _M. piriformis_ + Plan. _M. plantaris_ + Posterolat. can. Posterolateral canal of hypotarsus + Posteromed. can. Posteromedial canal of hypotarsus + Sar. _M. sartorius_ + Semim. _M. semimembranosus_ + Semit. _M. semitendinosus_ + Tib. ant. _M. tibialis anticus_ + Tib. cart. Tibial cartilage + + [Illustration: FIG. 1. _Pipilo erythrophthalmus._ Lateral view of + the superficial muscles of the left leg, x 1.5.] + + [Illustration: FIG. 2. _Pipilo erythrophthalmus._ Lateral view of + the left leg showing a deeper set of muscles. The superficial + muscles _iliotibialis_, _sartorius_, _gastrocnemius_ and + _peroneus longus_ have been removed, x 1.5.] + + [Illustration: FIG. 3. _Pipilo erythrophthalmus._ Lateral view of + the left leg showing the still deeper muscles. In addition to + those listed for figure 2, the following muscles have been + wholly or partly removed: _iliotrochantericus posticus_, + _femorotibialis externus_, _femorotibialis medius_, + _biceps femoris_, _semitendinosus_, _tibialis anticus_, + _flexor perforans et perforatus digiti II_, and _flexor + perforans et perforatus digiti III_, x 1.5.] + + [Illustration: FIG. 4. _Pipilo erythrophthalmus._ Medial view of + the superficial muscles of the left leg, x 1.5.] + + [Illustration: FIG. 5. _Pipilo erythrophthalmus._ Medial view of + the left leg showing a deeper set of muscles than those seen + in figure 4. The following superficial muscles have been + removed: _iliotibialis_, _sartorius_, _femorotibialis internus_, + _obturator internus_, _adductor longus (pars posticus)_, + _gastrocnemius_, and _peroneus longus_, x 1.5.] + + [Illustration: FIG. 6. _Pipilo erythrophthalmus._ Proximal end of + left tarsometatarsus and the hypotarsus, x 4.] + + [Illustration: FIG. 7. _Pipilo erythrophthalmus._ Lateral view of + proximal end of left femur and a portion of the pelvis, x 3.5.] + + [Illustration: FIG. 8. _Pipilo erythrophthalmus._ Upper surfaces + of the phalanges of the foretoes of the left foot showing + insertions of the _M. extensor digitorum longus_, x 3.] + + [Illustration: FIG. 9. _Pipilo erythrophthalmus._ Medial view of + the second digit of the left foot, showing insertions of the + flexor muscles, x 3.] + +The division of the _pars interna_ of the _m. gastrocnemius_ into +anterior and posterior parts has not been reported by previous authors +yet the division is quite distinct in those birds in which it occurs. +Hudson (1937:36) points out that in some non-passerine birds the _pars +interna_ is double, but that in these species the _m. semimembranosus_ +inserts between the two parts. This is not the condition in those +species studied by me. Only the ploceids and the cardueline finches in +the present investigation fail to show such a division. The undivided +muscle in these birds resembles, in its origin and position, the +posterior portion of the muscle found in those species showing the +bipartite condition. The greater mass of the bipartite muscle probably +makes possible a stronger extension of the tarsometatarsus. + +Thus, the divided or undivided conditions of the _m. obturator +externus_ and the _pars interna_ of the _m. gastrocnemius_ seem to be +correlated with the degrees of strength of certain movements of the +leg. It is conceivable that these differences in structure are +correlated with the manner in which food is obtained, the birds having +the bipartite muscles being those which spend the most time on the +ground searching and scratching for seeds and other sorts of food. +Yet, in _Leucosticte_, a cardueline, and in _Calcarius_, an +emberizine, whose foraging habits are rather similar, the structure is +unlike. _Leucosticte_ does resemble the emberizines and also _Piranga_ +and _Spzia_ in the extension of a band of muscle fibers from the _pars +interna_ of the _m. gastrocnemius_ around the front of the knee. A +band of muscle fibers of this sort strengthens the knee joint and +gives still more strength to the _pars interna_. This condition has +been reported in a number of birds by Hudson (1937) and is, in all +probability, an adaptation for greater strength of certain leg +movements. The development of this band in _Leucosticte_ seems to +parallel that in the other birds studied and does not indicate +relationship, since in _Leucosticte_ this band arises from the +undivided muscle which (as stated above) resembles only the posterior +portion of the bipartite muscle described for the other birds. In the +latter, the muscular band arises from the anterior part of the muscle. + +Minor differences in muscle pattern, like those already mentioned, are +consistent also between subfamilies, but correlation of these minor +differences with function is difficult. There is the implication, +however, that in all the groups except the carduelines and ploceids, +the emphasis is on greater strength and mobility of the leg. In the +carduelines that were studied the origin of the _m. sartorius_ does +not extend so far craniad as in the other species. In the latter, at +least half of the origin is from the last one or two free dorsal +vertebrae; in the carduelines no more than one third of the origin is +anterior to the ilium. It is conceivable that the more craniad the +origin, the stronger the forward movement of the thigh would be. + +In _Passer_, _Estrilda_ and _Poephila_, and in all the cardueline +finches examined, the bellies of the _m. flexor perforans et +perforatus digiti II_ and the _m. flexor perforans et perforatus +digiti III_ are more intimately connected than they are in the other +species studied. Thus, the amount of independent action of these +muscles in _Passer_, in the estrildines, and in the carduelines +probably is reduced. + +In _Passer_, the estrildines, and the carduelines the edges of the +sheathlike tendon of insertion of the _m. perforatus digiti III_ are +thickened; as a result the insertion appears superficially to be +double but closer examination reveals that there is a fascia stretched +between the thickened edges. In the other species examined, the +insertion is sheathlike throughout and there are no thick areas. I +cannot explain this on the basis of function. The difference, however, +is obvious and constant. + +Aside from the differences noted above, there were variations of +muscle pattern that seem to be significant only in _Vireo olivaceus_. +In this species the central, aponeurotic portion of the _m. +iliotibialis_ is absent. The origin of the _m. adductor longus et +brevis_ is from the dorsal edge of the ischiopubic fenestra and not +from the membrane covering this fenestra. The origin of the _pars +posticus_ of this muscle, furthermore, is fleshy and not tendinous as +it is in the other species. The _m. flexor perforatus digiti II_ is +larger and more deeply situated in _Vireo_ and has, furthermore, no +connection with the _m. flexor hallucis longus_. The latter muscle is +smaller and weaker than in any of the other species and has only one +(the posterior) head of origin. The _m. flexor hallucis brevis_, on +the contrary, is larger than in the other birds, compensating, +probably, for the small _m. flexor hallucis longus_. In those +differences, however, which separate the carduelines and ploceids from +the other birds studied, _Vireo_ resembles, in every instance, the +richmondenines, emberizines, tanagers, warblers, and blackbirds. + +On the basis of differences in leg-musculature the species which are +now included in the Family Fringillidae may be separated into two +groups. One group includes the richmondenines and the emberizines; the +other, the carduelines. The muscle patterns of the legs of the birds +of the first group are indistinguishable from those of _Seiurus_, +_Icterus_, _Molothrus_, and _Piranga_, and except for the differences +noted are similar to those in _Vireo_. The carduelines, on the other +hand, are similar in every point of leg-musculature to the ploceids +which were studied. Thus, the heterogeneity of the Family +Fringillidae, as now recognized, is emphasized by differences in the +muscle patterns of the leg. + + + + +COMPARATIVE SEROLOGY + + +General Statement + +The application of serological techniques to the problems of animal +relationships has been attempted with varying degrees of success over +a period of approximately fifty years. Few of the earlier studies were +of a quantitative nature, but within the past decade, satisfactory +quantitative serological techniques have been developed whereby +taxonomic relationships may be estimated. The usefulness of +comparative serology in taxonomy has been demonstrated in +investigations of many groups wherein results obtained have, in most +instances, been compatible with the results obtained by more +conventional methods, such as comparative morphology. As Boyden +(1942:141) stated, "comparative serology ... is no simple guide to +animal relationship." However, the objectiveness of its methods, the +fact that it has its basis in the comparisons of biochemical systems +which seem to be relatively slow to change in response to external +environmental influences, and the fact that the results are of +quantitative nature favor, where possible, the inclusion of data from +comparative serology along with that from more conventional sources +when an attempt is made to determine the relationships of groups of +animals. + +The application of serological methods in ornithology has not been +extensive. Irwin and Cole (1936) and Cumley and Irwin (1941, 1944) +used two species of doves and their hybrids and demonstrated that a +distinction between the red cells of these birds could be made by use +of immunological methods involving the agglutinin reaction. McGibbon +(1945) was able to distinguish the red cells of interspecific hybrids +in ducks by similar methods. Irwin (1953) used similar techniques in +his study of the evolutionary patterns of some antigenic substances of +the blood cells of birds of the Family Columbidae. Sasaki (1928) +demonstrated the usefulness of the precipitin technique in +distinguishing species of ducks and their hybrids. This technique +was used successfully also by DeFalco (1942) and by Martin and +Leone (1952). Working with groups of known relationships, these +investigators showed that the "accepted" systematic positions of +certain birds were confirmed by serological procedures. The precipitin +reaction, however, has never been applied to actual problems in avian +taxonomy prior to the present study. + + +Preparation of Antigens + +Although most previous work in comparative serology in which +precipitin tests were used has involved the use of whole sera as +antigens, Martin and Leone (1952) indicated that tissue extracts are +satisfactory as antigens and that serological differentiation can be +obtained with these extracts and the antisera to them. I decided, +therefore, to use such extracts in these investigations, since the +small sizes of the birds to be tested made it impracticable to obtain +enough whole sera. + +Most of the birds used were obtained by shooting, but a few were +trapped and the exotic species were purchased alive from a pet dealer. +When a bird was killed, the entire digestive tract was carefully +removed to prevent the escape of digestive enzymes into the tissues +and to prevent putrefaction by action of intestinal bacteria. As soon +as possible (and within three hours in every instance) the bird was +skinned, the head, wings, and legs were removed, and the body was +frozen. Each specimen, consisting of trunk, heart, lungs, and kidneys, +was wrapped separately and carefully in aluminum foil to prevent +dehydration of the tissues. The specimens were kept frozen until the +time when the extracts were made. + +When an extract was to be prepared, the specimen was allowed to thaw +but not to become warm. In the cold room with the temperature of all +equipment and reagents at 2 deg.C., the specimen was placed in a Waring +blender with 0.9 per cent aqueous solution of NaCl buffered with M/150 +K_{2}HPO_{4} and M/150 Na_{2}HPO_{4} to a pH of 7.0. The amount of +reagent used was 75 ml. of saline for each gram of tissue to be +extracted. The tissues were minced in the blender, allowed to stand at +2 deg.C. for 72 hours, and the tissue residues removed by centrifugation +in a refrigerated centrifuge. Formalin was added to a portion of the +supernatant in the amount necessary to make the final dilution 0.4 per +cent. This formolization was found to be necessary to inhibit the +action of autolytic enzymes over the period of time required to +complete the investigations. The effects of formolization on the +antigenicity and reactivity of proteins are discussed later. It was +necessary to sterilize and clarify the "native" (unformolized) +extracts; this was done by filtration through a Seitz filter. These +"native" substances were used only in the early stages of the +investigation (see below). The filtrate was bottled and stored at 2 deg.C. +In the early stages of this investigation clarification of the +formolized extract was accomplished by the same sort of filtration. It +was determined, however, that centrifugation in a refrigerated +centrifuge at high speeds (17,000g) served the same purpose and was +quicker. The formolized extracts were bottled and also stored at 2 deg.C. +(although refrigerated storage of the formolized extracts does not +seem necessary). For each extract the amount of protein present was +determined colorimetrically by the method of Greenberg (1929) with a +Leitz Photrometer. + +Species for which extracts were prepared and the protein values of the +extracts are listed in Table 1. Extracts of some species were used +throughout most of the experiment; extracts of others were used only +when needed for purposes of comparison. + + TABLE 1.--Species from Which Extracts Were Prepared and Injection + Schedules for Extracts Against Which Antisera Were Produced + + ==========================+==========+================================= + | Protein, | + SPECIES | gms. per | Injection schedules for + | 100 ml. | production of antisera + --------------------------+----------+--------------------------------- + _Myiarchus crinitus_ | 0.65 | Series 1: Intravenous, 0.5, 1.0, + (Linnaeus) | | 2.0, and 4.0 ml. + --------------------------+----------+--------------------------------- + _Passer domesticus_ | 1.40 | Series 1: Subcutaneous, 0.5, + | | 1.0, 2.0, and 4.0 ml. + --------------------------+----------+--------------------------------- + _Estrilda amandava_ | 0.45 | [A]Series 1: Intravenous, 0.5, + | | 1.0, 2.0, and 4.0 ml. + | | + | | [A]Series 2: Subcutaneous, 0.5, + | | 1.0, and 2.0 ml. + | | + | | Intraperitoneal, 8.0 ml. + --------------------------+----------+--------------------------------- + _Poephila guttata_ | 0.56 | [A]Same as for _Estrilda_. + --------------------------+----------+--------------------------------- + _Molothrus ater_ | 0.65 | Series 1: Intravenous and + | | subcutaneous, respectively, 0.5 + | | and 0.5 ml., 1.0 and 1.0 ml., + | | 3.0 and 1.0 ml., 5.0 and 3.0 ml. + | | + | | Series 2: Subcutaneous, 0.5, + | | 1.0, 2.0 and 4.0 ml. + --------------------------+----------+--------------------------------- + _Piranga rubra_ | 0.50 | Same as for _Molothrus_. + --------------------------+----------+--------------------------------- + _Richmondena cardinalis_ | 0.70 | [A]Same as for _Estrilda_. + --------------------------+----------+--------------------------------- + _Richmondena cardinalis_ | 0.60 | Same as for _Spinus_. + --------------------------+----------+--------------------------------- + _Passerina cyanea_ | 0.45 | Antiserum not prepared. + --------------------------+----------+--------------------------------- + _Spiza americana_ | 0.70 | Same as for _Molothrus_. + --------------------------+----------+--------------------------------- + _Carpodacus purpureus_ | 0.50 | Antiserum not prepared. + --------------------------+----------+--------------------------------- + _Spinus tristis_ | 0.49 | Series 1: Intravenous, 0.5, 1.0, + | | 2.0, and 4.0 ml. + | | + | | Series 2: Intravenous, 0.5, 1.0, + | | 2.0, and 4.0 ml. + | | + | | Series 3: Subcutaneous, 0.5, + | | 1.0, 2.0, and 4.0 ml. + --------------------------+----------+--------------------------------- + _Pipilo erythrophthalmus_ | 0.92 | Antiserum not prepared. + --------------------------+----------+--------------------------------- + _Junco hyemalis_ | 0.56 | Same as for _Spinus_. + --------------------------+----------+--------------------------------- + _Spizella arborea_ | 0.48 | Same as for _Spinus_. + --------------------------+----------+--------------------------------- + _Zonotrichia querula_ | 0.48 | Same as for _Spinus_. + --------------------------+----------+--------------------------------- + _Zonotrichia albicollis_ | 0.92 | Antiserum not prepared. + (Gmelin) | | + --------------------------+----------+--------------------------------- + + [A] Antiserum prepared against formolized antigen. + + +Preparation of Antisera + +All antisera were produced in rabbits (laboratory stock of +_Oryctolagus cuniculus_). Three methods of injection of antigen were +used in various combinations: intravenous, subcutaneous, and +intraperitoneal. Injection schedules used in the production of each +antiserum are listed in Table 1. Both formolized and "native" antigens +were used. Each rabbit received one or more series of four injections, +each injection being administered on alternate days and doubling in +amount: 0.5 ml., 1.0 ml., 2.0 ml., and 4.0 ml. In all but two +instances more than one series of injections was necessary to produce +a useful antiserum. More than two series, however, resulted in little +or no improvement of the reactivity of the antiserum. + +The injection-series were separated by intervals of eight days. On the +eighth day after the last injection of each series, 10 ml. of blood +were withdrawn from the main artery of the ear of the rabbit, and the +antiserum was used in a homologous precipitin test to determine its +usefulness. If the antiserum contained sufficient amounts of +antibodies to conduct the projected tests, the rabbit was completely +exsanguinated by cardiac puncture, by using an 18-gauge needle and a +50 ml. syringe. The whole blood was placed in clean test tubes and +allowed to clot. It was allowed to stand at 2 deg.C. for 12 to 18 hours so +that most of the serum would be expressed from the clot. The serum was +then decanted, centrifuged to remove all blood cells, sterilized in a +Seitz filter, bottled in sterile vials, and stored at 2 deg.C. until used. + + +Methods of Serological Testing + +The precipitin reaction is the most successful of the serological +techniques thus far devised for systematic comparisons. The reaction +occurs because antigenic substances introduced into the body of an +animal cause the formation of antibodies which precipitate antigens +when the two are mixed. The antisera which are produced show +quantitative specificities in their actions; therefore, when an +antiserum containing precipitins is mixed with each of several +antigens, the reaction involving the homologous antigen (that used in +the production of the antiserum) is greater than those reactions +involving the heterologous antigens (antigens other than those used in +the production of the antiserum). Furthermore, the magnitudes of the +reactions between the antiserum and the heterologous antigens vary +according to the degrees of similarity of these antigens to the +homologous one. + +The method of precipitin testing follows that outlined by Leone +(1949). The Libby (1938) Photronreflectometer was used to measure the +turbidities developed by the interaction of antigen and antiserum. +With this instrument parallel rays of light are passed through the +turbid systems being measured. Light rays are reflected from the +suspended particles to the sensitive plate of a photoelectric cell; +this generates a current of electricity which causes a deflection on a +galvanometer. The deflection is proportional to the amount of +turbidity developed and readings may be taken directly from the scale +of the instrument. + +The reaction-cells of the photronreflectometer are designed to operate +with a volume of 2 ml.; therefore, this volume was used in all +testing. In every series of tests the amount of antiserum was held +constant and the amount of antigen was varied. The volume for each +antigen dilution was always 1.7 ml., and to this was added 0.3 ml. of +antiserum to make up a volume of 2 ml. + + TABLE 2.--Percentage values obtained from analyses of precipitin + reactions. Numerals represent relative amounts of reaction between + antigens and antisera. Homologous reactions are arbitrarily valued + as 100 per cent, and heterologous reactions are expressed + accordingly. _Comparisons are meaningful only if made within each + horizontal row of values._ + + Table headings: + Col A: _Estrilda amandava_ + Col B: _Poephila guttata_ + Col C: _Piranga rubra_ + Col D: _Richmondena cardinalis_ + Col E: _Spiza americana_ + Col F: _Spinus tristis_ + Col G: _Junco hyemalis_ + Col H: _Zonotrichia querula_ + + ========================+============================================== + | ANTISERA + ANTIGENS +-----+-----+-----+-----+-----+-----+-----+---- + | A | B | C | D | E | F | G | H + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Passer domesticus_ | 75 | 74 | 73 | 66 | 81 | 72 | ... | 81 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Estrilda amandava_ | 100 | 88 | 75 | ... | 79 | 72 | 53 | ... + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Poephila guttata_ | 95 | 100 | 77 | 67 | 87 | 81 | ... | ... + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Molothrus ater_ | 66 | 54 | 69 | 65 | 86 | 75 | 69 | 75 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Piranga rubra_ | ... | ... | 100 | ... | ... | ... | ... | 89 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Richmondena cardinalis_| 75 | 80 | 91 | 100 | 98 | 65 | 88 | 91 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Spiza americana_ | 65 | 68 | ... | 71 | 100 | 64 | 67 | 80 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Carpodacus purpureus_ | 70 | 71 | 71 | 61 | 89 | 93 | 53 | 70 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Spinus tristis_ | 72 | 74 | 73 | 60 | 89 | 100 | 60 | ... + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Junco hyemalis_ | 64 | 56 | 74 | 65 | 87 | 68 | 100 | ... + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + _Zonotrichia querula_ | 65 | 71 | ... | 67 | 89 | 75 | ... | 100 + ------------------------+-----+-----+-----+-----+-----+-----+-----+---- + +Antigens were diluted with 0.9 per cent phosphate-buffered saline +solution. Tests were run in standard Kolmer test-tube racks, each test +consisting of 12 tubes. Each dilution was made on the basis of the +known protein concentration of the antigen. The first tube contained +an initial dilution of 1 part protein in 250 parts saline and each +successive tube contained a protein dilution one-half the +concentration of the preceding tube, ranging up to 1:512,000. Saline +controls, antiserum controls, and antigen controls were maintained +with each test to determine the turbidities inherent in these +solutions. These control-turbidities were deducted from the total +turbidity developed in each reaction-tube, the resultant turbidity +then being considered as that which was caused by the interaction of +antigens and antibodies. The turbidities were allowed to develop over +a 24-hour period. In the early stages of this investigation the +reactions were allowed to take place at 2 deg.C. in order to inhibit +bacterial growth. + +Later tests were carried out at room temperatures, and bacterial +growth was prevented by the addition to each tube of 'Merthiolate' in +a final dilution of 1:10,000. + + +Experimental Data + +Corrected values for the turbidities obtained were plotted with the +turbidity values on the ordinate and the antigen dilutions on the +abscissa. The homologous reaction was the standard of reference for +all other test reactions with the same antiserum. By summing the +plotted turbidity readings, numerical values are obtained which are +indices serving to characterize the curves. Such values were converted +to percentage values, that of the homologous reaction being considered +100 per cent. These values, plus the curves, provide the data by means +of which the proteins of the birds may be compared. Plots +representative of the precipitin curves are presented in Figs. 10 to +21. For convenience each plot represents only several of the 10 curves +obtained with each antiserum. + +A summary of the serological relationships of the birds involved in +the precipitin tests is presented in Table 2, in which percentage +values are presented. Since the techniques involved in testing were +greatly improved as the investigation proceeded, the summary is based +solely on those tests run in the later stages of the investigation. +For reasons which will become apparent in later discussion, it should +be emphasized that in Table 2 comparisons may be made only within each +horizontal row of values. + + +Discussion of the Serological Investigations + +One of the problems met early in this investigation was instability of +the proteins in the extracts that were prepared. Extracts in which no +attempt was made to inactivate the enzymes present proved +unsatisfactory. It was necessary to maintain the temperature of the +"native" antigens at 2 deg.C, and all work with such antigens had to be +performed at this temperature. This arrangement was inconvenient; +furthermore, inactivation of the enzymes was not complete even at this +low temperature, and some denaturation of the proteins took place as +evidenced by the gradual appearance of insoluble precipitates in the +stored vials. + +The preservatives, 'Merthiolate' and formalin, were used in an attempt +to inhibit the autolytic action of the enzymes present. Formalin, when +added to make a final dilution of 0.4 per cent, proved to be the more +satisfactory of the two preservatives and was used throughout most of +the work. Formalin caused slight denaturation of some of the proteins, +but this effect was complete within a few hours, after which any +denatured material was removed by filtration or centrifugation. The +proteins remaining in solution were stable over the period necessary +to complete the investigations. + +The addition of formalin reduces the reactivity of the extracts when +they are tested with antisera prepared against "native" antigens and +causes changes in the nature of the precipitin curves. This effect has +been pointed out by Horsfall (1934) and by Leone (1953) in their work +on the effects of formaldehyde on serum proteins. Their data indicate, +however, that even though changes in the immunological characteristics +of proteins are brought about by formolization, the proteins retain +enough of their specific chemical characteristics to allow consistent +differentiation of species by immunological methods. In the tests +which I performed, the relative positions of the precipitin curves, +whether native or formolized extracts were involved, remained +unchanged (Figs. 10, 11). _All data used in interpretation of the +serological relationships were obtained from tests in which formolized +antigens of equivalent age were used._ + +Only three antisera were produced against formolized antigens, all +others being produced against "native" extracts. The formolized +antigens seemed to have a greater antigenicity, in most instances, +than did those which were unformolized, and precipitin reactions +involving antisera produced against formolized antigens developed +higher turbidities. The antisera produced against formolized antigens +were equal to but no better than those prepared against "native" +extracts in separating the birds tested (Figs. 12, 13). + +The rabbit is a variable to be considered in serological tests. Two +rabbits exposed to the same antigen, under the same conditions, may +produce antisera which differ greatly in their capacities to +distinguish different antigens. It is logical to assume, therefore, +that two rabbits exposed to different antigens may produce antisera +which also differ in this respect. This explains the unequal values of +reciprocal tests shown in Table 2. Thus, in the test involving the +antiserum to the extracts of _Richmondena_, a value of 71 per cent was +obtained for _Spiza_ antigen, whereas in the test involving +anti-_Spiza_ serum, a value of 98 per cent was obtained for +_Richmondena_ antigen. In Table 2, therefore, comparisons may be made +only among values for the proteins of birds tested with the same +antiserum. + +Since the amount of any one antiserum is limited, there is, of +necessity, a limit as to the number of birds used in a series of +serological tests. Therefore, although the results reveal the actual +serological relationships of the individual species, interpretation of +the relationships of the taxonomic groups must be undertaken with the +realization that such an interpretation is based on tests involving +relatively few species of each group. It is reasonable to assume, +however, that a species which has been placed in a group on the basis +of resemblances other than serological resemblance would show greater +serological correspondence to other members of that group than it +would to members of other groups. Specifically, in the Fringillidae +and their allies, there seems to be little reason to doubt that +genera, and even subfamilies, are natural groups. This is illustrated +in tests involving closely related genera: _Richmondena_ and _Spiza_ +(Figs. 14, 15, 18), _Estrilda_ and _Poephila_ (Fig. 21), _Spinus_ and +_Carpodacus_ (Figs. 12, 17, 19, 20). In each of these tests the pairs +of genera mentioned show greater serological correspondence to each +other than they do to other kinds involved. This point is illustrated +further by a test (not illustrated) involving _Zonotrichia querula_ +(the homologous antigen) and _Zonotrichia albicollis_. Although this +test was one of an earlier series in which difficulties were +encountered (the data, therefore, were not used), it is of interest +that the two species were almost indistinguishable serologically. + +The serological homogeneity of passeriform birds is emphasized by the +fact that the value of every heterologous reaction was more than 50 +per cent of the value of the homologous reaction, except in the test +involving the anti-_Richmondena_ serum and _Myiarchus_ (Fig. 13) in +which the value of the heterologous reaction was 45 per cent. Because +most ornithologists consider these genera to be only distantly related +(they are in different suborders within the Order Passeriformes), the +relatively high value of the heterologous reaction emphasizes the +close serological correspondence of passerine birds and indicates that +small consistent serological differences among these birds are +actually significant. The possibility that some of the serological +correspondence is due to the "homologizing" effect of formalin on +proteins should not be excluded. I think, however, that this effect is +not entirely responsible for the close correspondence observed here. + +An additional point to consider in interpretation of the serological +tests is that the techniques used tend to separate sharply species +that are closely related whereas species that are distantly related +are not so easily separated. In other words, comparative serological +studies with the photronreflectometer tend to minimize the differences +between distant relatives and to exaggerate the differences between +close relatives. + +In analyzing the serological relationships of the species used in this +study, it becomes obvious that two or more series of tests must be +considered before the birds can be placed in relation to each other. +For example, the data presented in Fig. 14 indicate that _Spiza_ and +_Molothrus_ show approximately the same degree of serological +correspondence to _Richmondena_. This does not imply necessarily that +_Spiza_ and _Molothrus_ are closely related. If Fig. 15 is examined, +it can be determined that _Richmondena_ shows much greater serological +correspondence to _Spiza_ than does _Molothrus_. Thus, an analysis of +both figures serves to clarify the true serological relationships of +the three genera. By reference to other series of tests involving +these three birds a more exact determination of their relationships +may be obtained. + +To illustrate this point by a hypothetical example, two species might +seem equidistant, serologically, from a third species. Additional +testing should indicate if the first two species are equidistant in +the same direction (therefore, by implication, close relatives) or in +opposite directions (therefore, distant relatives). A single test +supplies only two dimensions of a three dimensional arrangement. + +It is impossible to interpret and to picture the serological data +satisfactorily in two dimensions; therefore, a three-dimensional model +(Figs. 22, 23) was constructed to summarize the serological +relationships of the birds involved. Each of the eleven kinds used +consistently throughout the investigation is represented in the model. +By use of the percentage values (Table 2), each bird was located in +relation to the other birds. Where possible, averages of reciprocal +tests (Table 3) were used in determining distances between the +elements of the model. In this way seven of the birds were accurately +located in relation to each other. Lacking reciprocal tests, the +positions of the other birds were determined by the values of single +tests (Table 4). Although these birds were placed with less certainty, +at least four points of reference were used in locating each species. +At least one serological test is represented by each connecting bar in +the model. The lengths of the bars connecting any two elements were +determined as follows: a percentage value (Table 3 and Table 4) +representing the degree of serological correspondence between two +birds was subtracted from 100 per cent; the remainder was multiplied +by a factor of five to increase the size of the model and the product +was expressed in millimeters; a bar of proper length connects the two +elements involved. + +From the model it is observed that, _Molothrus_ and _Passer_ excluded, +the birds fall into two distinct groups: one includes _Piranga_, +_Richmondena_, _Spiza_, _Junco_, and _Zonotrichia_; the other includes +_Estrilda_, _Poephila_, _Carpodacus_, and _Spinus_. + + TABLE 3.--Reciprocal Values Used to Determine Distances Between + Elements of the Model; Each Value Represents the Average of + Serological Tests Between the Species Involved + + Table Headings: + Col A: _Estrilda amandava_ + Col B: _Poephila guttata_ + Col C: _Richmondena cardinalis_ + Col D: _Spiza americana_ + Col E: _Spinus tristis_ + Col F: _Junco hyemalis_ + Col G: _Zonotrichia querula_ + + ==========================+====+====+====+====+====+====+====+ + | A | B | C | D | E | F | G | + --------------------------+----+----+----+----+----+----+----+ + _Estrilda amandava_ | .. | 92 | .. | 72 | 72 | 59 | .. | + --------------------------+----+----+----+----+----+----+----+ + _Poephila guttata_ | 92 | .. | 74 | 78 | 78 | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + _Richmondena cardinalis_ | .. | 74 | .. | 85 | 63 | 77 | 79 | + --------------------------+----+----+----+----+----+----+----+ + _Spiza americana_ | 72 | 78 | 85 | .. | 77 | 77 | 85 | + --------------------------+----+----+----+----+----+----+----+ + _Spinus tristis_ | 72 | 78 | 63 | 77 | .. | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + _Junco hyemalis_ | .. | .. | 77 | 77 | .. | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + _Zonotrichia querula_ | .. | .. | 79 | 85 | .. | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + + TABLE 4.--Single Values Used to Determine Distances Between Elements + of the Model; Each Value Represents a Single Test Between the + Species Involved + + Table headings: + Col A: _Estrilda amandava_ + Col B: _Poephila guttata_ + Col C: _Piranga rubra_ + Col D: _Richmondena cardinalis_ + Col E: _Spinus tristis_ + Col F: _Junco hyemalis_ + Col G: _Zonotrichia querula_ + + ==========================+====+====+====+====+====+====+====+ + | A | B | C | D | E | F | G | + --------------------------+----+----+----+----+----+----+----+ + _Passer domesticus_ | .. | 74 | 73 | .. | 72 | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + _Molothrus ater_ | .. | 54 | .. | 65 | .. | 69 | 75 | + --------------------------+----+----+----+----+----+----+----+ + _Piranga rubra_ | .. | 77 | .. | 91 | 73 | 74 | .. | + --------------------------+----+----+----+----+----+----+----+ + _Carpodacus purpureus_ | 70 | 71 | .. | 61 | 93 | .. | .. | + --------------------------+----+----+----+----+----+----+----+ + + [Illustration: FIGS. 10-13. Graphs of precipitin reactions + illustrating effects of formalin on antigenicity and reactivity + of the extracts. For further information, see text, pp. 190-193. + + FIG. 10. Reactions of unformolized antigens of _Richmondena_, + _Zonotrichia_, and _Molothrus_ with anti-_Richmondena_ serum. + FIG. 11. Reactions of formolized antigens of _Richmondena_, + _Zonotrichia_, and _Molothrus_ with anti-_Richmondena_ serum. + FIG. 12. Reactions of anti-_Richmondena_ serum prepared against + native antigen with antigens of _Richmondena_, _Zonotrichia_, + _Carpodacus_, and _Spinus_. + FIG. 13. Reactions of anti-_Richmondena_ serum prepared against + formolized antigen with antigens of _Richmondena_, _Zonotrichia_, + _Poephila_, _Spinus_, and _Myiarchus_.] + + [Illustration: FIGS. 14-17. Graphs of precipitin reactions + illustrating serological relationships. For further explanation, + see text, pp. 190-193. + + FIG. 14. Serological relationships of _Richmondena_, _Spiza_, and + _Molothrus_. + FIG. 15. Serological relationships of _Richmondena_, _Spiza_, and + _Molothrus_. + FIG. 16. Serological relationships of _Carpodacus_ with the + richmondenine-emberizine-thraupid assemblage. + FIG. 17. Serological relationships of _Carpodacus_ and _Spinus_ with + _Richmondena_ and _Junco_.] + + [Illustration: FIGS. 18-21. Graphs of precipitin reactions + illustrating serological relationships. For further explanation, + see text, pp. 190-193. + + FIG. 18. Serological relationships of _Spinus_ and _Poephila_ with + the richmondenines. + FIG. 19. Serological relationships of _Carpodacus_ and _Spinus_ + with _Richmondena_ and _Piranga_. + FIG. 20. Serological relationships of _Poephila_ and Richmondena + with the carduelines. + FIG. 21. Serological relationships of _Richmondena_ and _Spinus_ + with the estrildines.] + + [Illustration: FIG. 22. Two views of a model illustrating + serological relationships among fringillid and related birds. + For further explanation, see text, pp. 193-194. + + Genera Pi . . . . _Piranga_ + C . . . . _Carpodacus_ Po . . . . _Poephila_ + E . . . . _Estrilda_ R . . . . _Richmondena_ + J . . . . _Junco_ Sn . . . . _Spinus_ + M . . . . _Molothrus_ Sz . . . . _Spiza_ + Pa . . . . _Passer_ Z . . . . _Zonotrichia_] + + [Illustration: FIG. 23. Two additional views of the model shown in + fig. 22 illustrating serological relationships among fringillid + and related birds. For further explanation, see text, + pp. 193-194. + + Genera Pi . . . . _Piranga_ + C . . . . _Carpodacus_ Po . . . . _Poephila_ + E . . . . _Estrilda_ R . . . . _Richmondena_ + J . . . . _Junco_ Sn . . . . _Spinus_ + M . . . . _Molothrus_ Sz . . . . _Spiza_ + Pa . . . . _Passer_ Z . . . . _Zonotrichia_] + +Within the richmondenine-emberizine-thraupid assemblage, _Junco_ +and _Zonotrichia_ constitute a sub-group apart from the others. +_Piranga_ and _Richmondena_ show close serological correspondence. +The present taxonomic position of _Spiza_ in the Richmondeninae, +which has been questioned by Beecher (1951a:431; 1953:309), is +corroborated at least insofar as the serological evidence is +concerned. Certainly, serological correspondence of _Spiza_ with the +richmondenine-emberizine-thraupid assemblage is greater than with any +other group of birds tested. + +It is obvious that the serological affinities of the carduelines do +not lie with the richmondenines, emberizines, or thraupids. The +carduelines show greater serological correspondence with the +estrildines than they do with any of the other groups tested. Further +serological investigation involving other species, however, is +necessary before the nearest relatives of the carduelines can be +determined with certainty. + +The two estrildines tested (_Estrilda_ and _Poephila_) show close +serological relationship. Their nearest relatives, serologically, seem +to be the carduelines. The classification (Wetmore, 1951) that places +_Passer_ in the same family with the estrildines is not upheld by the +serological data available. _Passer_ is not, serologically, closely +related to any of the birds tested. It is of interest that Beecher +(1953:303-305), on the basis of jaw musculature, places _Passer_ and +the estrildines in separate families (Ploceidae and Estrildidae, +respectively). + +_Molothrus_ shows greater serological correspondence to the +richmondenine-emberizine-thraupid assemblage than to any of the other +birds tested. It is definitely set apart from this group, however, and +its position, serologically, is compatible with that based on evidence +from other sources. + +There seems to be but little argument among ornithologists that +icterids, fringillids, and ploceids constitute families which are +distinct from one another. If, then, the serological differences +between _Molothrus_ (Icteridae) and _Richmondena_ (Fringillidae), +between _Molothrus_ and _Zonotrichia_ (Fringillidae), and between +_Richmondena_ and _Poephila_ (Ploceidae) are indicative of family +differences, there are four families represented by the birds +involved. _Molothrus_ represents one family; _Piranga_, _Richmondena_, +_Spiza_, _Junco_, and _Zonotrichia_, a second; _Estrilda_, _Poephila_, +_Carpodacus_, and _Spinus_, a third; and _Passer_, a fourth. + + + + +CONCLUSIONS + + +The heterogeneity of the Family Fringillidae has been emphasized by +many authors. The relationships of the species now included in this +Family have been the subject of much discussion and constitute an +important problem in avian systematics. + +Sushkin's studies (1924, 1925) of features of the horny and bony +palates have served as a basis for the present division of the Family +into subfamilies. Recently, Beecher (1951a, 1951b, 1953) and Tordoff +(1954) have used these features and others which they thought to be of +value in an attempt to clarify the relationships of the species +involved. + +Beecher's work (1951a, 1951b, 1953) on jaw-musculature is a valuable +contribution to our knowledge of the anatomy of passerine birds. His +myological studies were so thorough and his presentation so detailed +that students who disagree with his interpretations can draw their own +conclusions. Beecher (1951b:276) points out that there are two basic +types of skeletal muscle--those with parallel fibers and those with +pinnately arranged fibers. The muscles with pinnate fibers seem to be +more efficient, each muscle having a greater functional cross section +for its bulk than does one with parallel fibers. He assumes that +muscles with parallel fibers are more primitive, phylogenetically, +than are those with fibers arranged pinnately. Since his study of the +jaw muscles of the Icteridae (1951a) revealed that patterns of +jaw-musculature within this Family remain constant regardless of the +methods used in procuring food, he assumes that such patterns may be +used as indicators of relationship throughout the entire oscinine +group. These two assumptions, then, serve as the basis for his +hypothesis concerning relationship and phylogeny within this +assemblage. Beecher (1951b:278-280; 1953:310-312) maintains that +within the Family Thraupidae there are two main lines which lead with +almost no disjunction to the Carduelinae and Richmondeninae. The +thraupid-richmondenine line involves a shift in the nature of the _m. +adductor mandibulae externus superficialis_, which becomes more +pinnate in the richmondenines. This results in greater crushing power. +The thraupid-cardueline line involves a shift in emphasis from the the +_m. adductor mandibulae externus medialis_ to the _m. pseudotemporalis +superficialis_ and the forward advance of the insertion of the latter. +This, also, promotes greater crushing ability. He states that features +of the horny palate and of the plumage provide further evidence of +close relationship of these groups. He includes, therefore, the +Thraupinae, the Carduelinae, and the Pyrrhuloxiinae (=Richmondeninae) +in the Family Thraupidae. Beecher (1953:307) indicates that the +patterns of jaw-musculature of the Parulinae (wood warblers) and +Emberizinae (buntings) are similar and suggests that the buntings had +their origin from the wood warblers. He includes these subfamilies, +therefore, in the Family Parulidae. + +Beecher's reasoning may be criticized on several points. It may be, as +he suggests, that muscles with parallel fibers evolved earlier, +phylogenetically, than did muscles with pinnate fibers, but he does +not give adequate consideration, it seems to me, to the possibility +that parallel fibers may also have evolved secondarily from pinnate +fibers. Since Beecher (1951a) found that patterns of jaw-musculature +within the Family Icteridae were conservative, he is reluctant to +admit the possibility of convergence among any of the other families. +Differences in patterns of jaw-musculature are, however, functional +adaptations and like the bill, which is also associated with +food-getting may be subject to rapid evolutionary change. Finally, in +attempting to classify the oscines, he has relied almost entirely on a +single character--the pattern of jaw-musculature. + +Tordoff's attempts (1954) to clarify the relationships of the +fringillids and related species are based chiefly on features of the +bony palate. He assumes that since palato-maxillaries seem to be +absent in the majority of passerine birds, their occurrence in certain +nine-primaried oscine groups indicates relationship among these +groups. He points out that these bones, when present, are important +areas of origin of the _m. pterygoideus_ which functions in depression +of the upper jaw and in elevation of the lower jaw. He assumes, +therefore, that palato-maxillaries were evolved to provide for a more +effective action of the _m. pterygoideus_. The need for such action +could be associated with a seed-eating habit. All richmondenines and +emberizines possess palato-maxillary bones either free or fused to the +prepalatine bar, but there is no trace of these bones in the +carduelines. Carduelines, furthermore, possess prepalatine bars that +are characteristically flared anteriorly. This condition does not +exist in the richmondenines or in the emberizines. + +Tordoff points out, also, that the irregular, erratic migrations of +the New World Carduelinae are unlike the more regular migrations of +the richmondenines and emberizines. The carduelines, furthermore, are +more arboreal in their habits than are these other groups and exhibit +a decided lack of nest sanitation during the later stages of nesting, +a situation which contrasts with that found in the Richmondeninae and +Emberizinae. He suggests, therefore, that the carduelines are not so +closely related to the richmondenines and the emberizines as +previously has been thought. + +Since there are only two cardueline genera, _Loximitris_ and +_Hesperiphona_, endemic to the New World and at least 10 genera with +many species endemic to the Old World, Tordoff (1954:15) suggests an +Old World origin for the carduelines. He strengthens his argument for +this hypothesis by pointing out that in features of the bony palate +and in habits the carduelines resemble the estrildines of the Family +Ploceidae. + +Tordoff (1954:29-30) states that the tanagers not only merge with the +richmondenines but also grade imperceptibly into the emberizines. He +includes, therefore, the Richmondeninae, Emberizinae, and Thraupinae +in the Family Fringillidae. He suggests that the carduelines are +ploceids, closely related to the Subfamily Estrildinae, on the basis +of structure of the bony palate, geographic distribution, social +behavior, and habits such as nest-fouling and nest-building. + +Tordoff, like Beecher, has based his interpretations chiefly on one +feature--structure of the bony palate. Since this feature also is +associated with food-getting, the possibilities of convergence of +distantly related species with similar habits and divergence of +closely related species with different habits may not be excluded. + +The hazard of unrecognized adaptive convergence cannot, of course, be +excluded from most fields of taxonomic research, but some features of +morphology and biochemistry are notably more conservative than others +and undergo slower evolutionary change. Such features are often of +utmost importance in distinguishing the higher taxonomic categories. + +Most ornithologists are aware that, within the Order Passeriformes, +patterns of musculature in the leg have evolved at a slow rate and +exhibit little variation within the Order. Differences which do occur, +therefore, probably are significant, especially those that are +consistent between groups of species. As I have pointed out earlier +(p. 184), there are no significant differences in leg-musculature +between the Richmondeninae, Emberizinae, and Thraupidae. Indeed, it is +difficult to define these groups on the basis of leg-musculature. If +these groups are of common origin, the lack of distinct boundaries +between them is not surprising. A muscular band which extends from the +_pars interna_ of the _m. gastrocnemius_ around the front of the knee +is present in every emberizine species that I studied and in the Genus +_Piranga_. With the exception of _Spiza_ none of the richmondenines +possesses this band. + +The significant differences in leg-musculature which have been +discussed above (pp. 183-184) distinguish the carduelines from the New +World finches and tanagers. Even the cardueline _Leucosticte_ and the +emberizine _Calcarius_, which resemble one another in general +adaptations and in several myological features of the leg (p. 183), +agree in significant features of the musculature with the respective +groups to which they belong. The carduelines agree in the major +features of leg-musculature with the ploceids which I studied. + +The use of serological techniques in taxonomic work has two main +advantages. The biochemical systems involved in such investigations +seem to be relatively slow to change in response to external +environmental influences, and the quantitative nature of the results +obtained makes possible objective measurement of resemblances among +species. + +I have pointed out (p. 200) that the carduelines are excluded, +serologically, from the distinct assemblage formed by the +richmondenines, emberizines, and tanagers. Actually, the carduelines +show less serological resemblance to this assemblage than do the +estrildines, and most ornithologists agree that the Estrildinae are +not at all closely related to the Richmondeninae, Emberizinae, and +Thraupidae. _Molothrus_, representing a family (Icteridae) recognized +as distinct from the Family Fringillidae, also more closely resembles +the fringillid assemblage, serologically, than do the carduelines. +Although the Carduelinae constitute a distinct group serologically, +they show greater serological resemblance to the estrildines of the +Family Ploceidae than to any of the other species tested. At least the +carduelines and the estrildines form a group as compact as the +subfamilies of the Fringillidae. Thus, the serological data correlate +well with those obtained from the study of the leg-musculature. + +Present systems of classification include the subfamilies Passerinae +and Estrildinae in the Family Ploceidae. _Passer_, however, is less +closely related to the estrildines serologically than are the +carduelines, and is less closely related to the estrildines than +_Molothrus_, an icterid, is to the fringillids. This raises a question +as to the homogeneity of the Family Ploceidae as presently recognized +by most ornithologists. If the Passerinae and the Estrildinae are +placed in a single family, the serological divergence among members of +this group is certainly greater than it is in the Family Fringillidae. +Additionally, Beecher (1953:303-304) found that the estrildines +possess a pattern of jaw-musculature different from those in other +ploceids. + +The combined evidence from jaw-musculature and serology has caused me +to conclude that the estrildines should be excluded from the Family +Ploceidae (see below). + +In an attempt to clarify the relationships of the Fringillidae and +allied groups, I here review briefly the evidence which has been +presented. From his studies of jaw-musculature (1951a, 1951b, +1953) Beecher concludes that the Pyrrhuloxinae (=Richmondeninae), +the Carduelinae, and the Thraupinae are closely related. +He places these groups in the Family Thraupidae. He excludes the +Emberizinae from this group and places them with the wood warblers +in the Family Parulidae. He suggests that the estrildines constitute +a family (Estrildidae) separate from the Family Ploceidae. + +From his studies of certain features of the bony palate Tordoff +(1954:25-26, 32) concludes that the richmondenines, the emberizines, +and the tanagers have a common origin and places these groups in the +Family Fringillidae. He excludes the carduelines from this assemblage, +suggests that they are closely related to the estrildines, and +includes them as the Subfamily Carduelinae in the Family Ploceidae. + +In this paper I have presented data obtained from the study of certain +features of morphology and biochemistry which I think are less subject +to the influence of environmental factors than those features studied +by recent workers. It is significant that the data obtained by use of +serological techniques and those obtained from the study of +leg-musculature point to the same conclusions. On the basis of these +data I have drawn several conclusions concerning the relationships of +the groups which I studied. + +The richmondenines, emberizines, and tanagers are closely related and +should be included in a single family, Fringillidae. The Carduelinae +and the Estrildinae are closely related subfamilies. Although most +recent classifications place the Estrildinae and Passerinae in the +Family Ploceidae, the serological evidence indicates that these groups +are not closely related. Beecher (1953:303-304) drew the same +conclusion from his study of jaw-musculature (see above). I suggest, +therefore, that the Carduelinae and the Estrildinae be placed in a +family separate from the Ploceidae and that the name Carduelidae +(rather than Estrildidae) be used for this group. At present, neither +is an accepted family name. Because _Carduelis_ Brisson 1760 is an +older name than _Estrilda_ Swainson 1827 and because _Carduelis_ seems +to be a centrally located genus in the family, I have chosen the +former (although the International Rules of Zoological Nomenclature do +not specify that priority must apply in forming family names). + +I have been unable to study any of the species included in the +subfamilies Fringillinae (not Fringillinae of Tordoff, see 1954:23-24, +and below) and Geospizinae of recent classifications; thus these +groups have not been discussed above. Beecher (1953:307-308) includes +_Fringilla_ in the Subfamily Carduelinae; he includes the geospizines +in a separate family, Geospizidae, and states that they are derived +from the emberizines. Tordoff (1954:23-24) found that in features of +the bony palate _Fringilla_ and the geospizines resemble the +emberizines and, on this basis, includes them in the Subfamily +Fringillinae. + +The Dickcissel, _Spiza americana_, possesses certain features which +merit special discussion. Beecher (1951a:431; 1953:309), on the basis +of jaw-musculature, considers it an icterid. To be sure _Spiza_ is in +many ways an aberrant member of the group to which it is now assigned +(Subfamily Richmondeninae). _Spiza_, serologically, is closely related +to all species of the richmondenine-emberizine-thraupid assemblage. +Within this assemblage its nearest relatives are the richmondenines. +_Spiza_ differs from the other richmondenines studied and resembles +the emberizines and tanagers in the possession of the muscular band +which extends from the _pars interna_ of the _m. gastrocnemius_ around +the front of the knee. This band, in _Spiza_, is smaller, however, +than in any of the other species. No icterid dissected possesses such +a structure. Tordoff (1954:29) states that _Spiza_ is typically +richmondenine in palatal structure and makes the suggestion, with +which I agree, that _Spiza_ is a richmondenine and may be closely +related to the ancestral stock which gave rise to the fringillid +assemblage. The serological position of _Spiza_, approximately +equidistant from the other fringillids (Figs. 22, 23), and the +presence of the small muscular band around the front of the knee +constitute evidence supporting the central position of _Spiza_. + +After consideration of evidence from the studies of external +morphology, ethology, myology, osteology, and serology, I propose here +an arrangement of the groups which I have studied and submit for +comparison the arrangements (of these groups) proposed by Beecher and +Tordoff. The names of subfamilies that I have been unable to study are +included in my classification and are placed in brackets. + + ------------------------+----------------------+----------------------- + | Proposed by Tordoff | Proposed by Beecher + Here proposed: | (1954) on the basis | (1953) on the basis + | of the bony palate: | of jaw-musculature: + ========================+======================+======================= + FAMILY PLOCEIDAE | FAMILY PLOCEIDAE | FAMILY PLOCEIDAE + | | + [Subf. Bubalornithinae] |Subf. Bubalornithinae | + | | + Subfamily Passerinae: |Subfamily Passerinae | Subfamily Passerinae + distinguished from the | | + Estrildinae by patterns | | + of jaw-musculature | | + (Beecher, 1953:303-304) | | + and on the basis of | | + comparative serology of | | + saline-soluble proteins.| | + | | + [Subfamily Ploceinae] |Subfamily Ploceinae | Subfamily Ploceinae + | | + [Subfamily Viduinae] |Subfamily Viduinae | Subfamily Viduinae + | | + FAMILY CARDUELIDAE | | + | | + Subfamily Estrildinae: |Subfamily Estrildinae | FAMILY ESTRILDIDAE + similar to the | | + Carduelinae in features | | + of the bony palate and | | + habits (Tordoff, 1954: | | + 18-22) and in patterns | | + of leg-musculature and | | + comparative serology | | + of saline-soluble | | + proteins. | | + | | + Subfamily Carduelinae: |Subfamily Carduelinae | [In Thraupidae below] + distinguished from the | | + Fringillidae by features| | + of the palate, | | + geographic distribution,| | + migration patterns, and | | + habits (Tordoff, 1954: | | + 14-18) and by patterns | | + of leg-musculature and | | + comparative serology | | + of saline-soluble | | + proteins. | | + | | + FAMILY FRINGILLIDAE: all| FAMILY FRINGILLIDAE | FAMILY PARULIDAE + members of this family | | Subfamily Parulinae + show similarities in | | Subfamily Emberizinae + features of the bony | | + palate (Tordoff, 1954: | | + 22-23), patterns of | | + leg-musculature, and | | + in comparative serology | | + of saline-soluble | | + proteins. | | FAMILY THRAUPIDAE + | | + Subf. Richmondeninae |Subf. Richmondeninae | Subfamily + | | Pyrrhuloxiinae + | | + Subfamily Thraupinae |Subfamily Thraupinae | Subfamily Thraupinae + | | + Subfamily Emberizinae |Subfamily Fringillinae| [In Parulidae above] + |(including Emberizinae| + [Subfamily Fringillinae]| and Geospizinae) | Subfamily Carduelinae + | | + [Subfamily Geospizinae] | | + ------------------------+----------------------+----------------------- + + + + +SUMMARY + + +It has long been recognized that the Family Fringillidae includes some +dissimilar groups. Specifically, the relationships of the subfamilies +Richmondeninae, Emberizinae, and Carduelinae of the Family +Fringillidae are poorly understood. Data from two recent studies, one +on patterns of jaw-musculature and the other on features of the bony +palate, emphasize the dissimilarity of these subfamilies but have +given rise to conflicting concepts of the relationships of subfamilies +within the Family. + +This paper reports the results of studies involving morphological and +biochemical features that I consider less sensitive to external +environmental factors than are features which have been studied +previously. Patterns of leg-musculature were chosen for study because +earlier work showed that muscle patterns in the legs of passerine +birds are highly stable and vary but little. Variations, therefore, +which are consistent in separating groups of species should be +significant. Serological techniques were used because the biochemical +systems involved seem to be relatively slow to change in response to +environmental influences and because the data obtained may be used in +a highly objective manner to measure resemblance among species. + +Individual differences in the patterns of leg-musculature were found +to be slight and involved mainly the sizes and shapes of muscles. For +this reason variations involving origin, insertion, or relative +position of a muscle, were judged significant. In leg-musculature the +Richmondeninae, the Emberizinae, and the Thraupidae resemble one +another closely. Several differences in muscle pattern were found, +however, which distinguish these groups from the Carduelinae. The +leg-musculature of the carduelines closely resembles that of the +Ploceidae. + +Serological techniques involved the extraction of saline-soluble +proteins from the tissues of the species to be studied. These extracts +were carefully processed and were used as antigens. Formolization of +the antigens was necessary as a means of preventing denaturation of +the proteins by enzymatic activity. Antisera were produced in rabbits. +The method of testing involved turbidimetric analysis of the +precipitin reaction. Utilizing the values for the precipitin tests a +model was constructed which showed the relationships of the eleven +species used in these tests. From a study of the model and the data +used in its construction, it was determined that the Richmondeninae, +Emberizinae, and Thraupidae constitute an assemblage distinct from the +other species studied. The Carduelinae are excluded from the +assemblage and serologically are most closely related to the +Estrildinae. The estrildines, serologically, do not closely resemble +_Passer_, Subfamily Passerinae, although recent classifications place +these two subfamilies in the Family Ploceidae. + +Upon consideration of all evidence now available--from external +morphology, ethology, myology, osteology, and serology--several +hypotheses regarding the relationships of the groups studied are set +forth. The richmondenines, emberizines, and tanagers are closely +related subfamilies and are here included in the Family Fringillidae. +The Estrildinae and Carduelinae are closely related subfamilies, but +neither group is closely related to the Passerinae. The estrildines +and carduelines, therefore, are placed in a separate family, the +Carduelidae. In some ways, _Spiza_ is an aberrant member of the +Subfamily Richmondeninae but should be retained in that subfamily. It +is suggested that _Spiza_ is a primitive richmondenine closely related +to the ancestral fringillid stock. + + + + +LITERATURE CITED + + +AMERICAN ORNITHOLOGISTS' UNION + + 1931. Check-list of North American birds. Fourth edition. + Lancaster, Pa., xix + 526 pp. + + +BEECHER, W. J. + + 1951a. Adaptations for food-getting in the American blackbirds. + Auk, 68:411-440, 11 figs. + + 1951b. Convergence in the Coerebidae. Wilson Bull., 63:274-287, + 5 figs. + + 1953. A phylogeny of the oscines. Auk, 70:270-333, 18 figs. + + +BERGER, A. J. + + 1952. The comparative functional morphology of the pelvic + appendage in three genera of Cuculidae. + Amer. Mid. Nat., 47:513-605, 29 pls. + + +BOYDEN, A. + + 1942. Systematic serology: a critical appreciation. + Physiol. Zool., 15:109-145, 12 figs. + + +CHAPIN, J. P. + + 1917. The classification of the weaver-birds. Bull. Amer. Mus. + Nat. Hist., 37:243-280, 10 pls., 9 figs. + + +CUMLEY, R. W., and IRWIN, M. R. + + 1941. Pictorial representation of the antigenic differences + between two dove species. Jour. Hered., 32:178-182, + frontispiece, 2 figs. + + 1941. Interaction of antigens in dove hybrids. Ibid., 429-434, + 3 figs. + + 1944. The correlation between antigenic composition and geographic + range in the Old and New World of some species of _Columba_. + Amer. Nat., 78:238-256, 1 fig. + + +DEFALCO, R. J. + + 1942. A serological study of some avian relationships. + Biol. Bull., 83:205-218. + + +FISHER, H. I. + + 1946. Adaptations and comparative anatomy of the locomotor + apparatus of New World vultures. Amer. Mid. Nat., + 35:545-727, 13 pls., 28 figs. + + +GADOW, H., and SELENKA, E. + + 1891. Voegel, vol. I, Anatomischer Theil. In Bronn's Klassen und + Ordnungen des Thier-Reichs, Sechster Band, Vierte Abtheilung. + Leipzig, 1008 pp., 59 pls. + + +GARROD, A. H. + + 1873. On certain muscles in the thigh of birds and their value in + classification. Proc. Zool. Soc. London, Part I:626-644, + 6 figs. + + 1874. On certain muscles in the thigh of birds and their value in + classification. Ibid., Part II:111-123. + + +GREENBERG, D. M. + + 1929. The colorimetric determination of serum proteins. + J. Biol. Chem., 82:545-550. + + +HELLMAYR, C. E. + + 1935. Catalogue of birds of the Americas. Field Mus. Nat. Hist., + Zool. ser. 13, pt. 8, vi + 541 pp. + + 1936. Catalogue of birds of the Americas. Ibid., 13, pt. 9, + v + 458 pp. + + 1937. Catalogue of birds of the Americas. Ibid., 13, pt. 10, + v + 228 pp. + + 1938. Catalogue of birds of the Americas. Ibid., 13, pt. 11, + vi + 662 pp. + + +HOWARD, H. + + 1929. The avifauna of the Emeryville shellmound. Univ. California + Publ. Zool., 32:301-394, 3 pls., 54 figs. + + +HUDSON, G. E. + + 1937. Studies on the muscles of the pelvic appendage in birds. + Amer. Mid. Nat., 18:1-108, 26 pls. + + +IRWIN, M. R. + + 1953. Evolutionary patterns of antigenic substances of the blood + corpuscles in Columbidae. Evol., 7:31-50. + + +IRWIN, M. R., and COLE, L. J. + + 1936. Immunogenetic studies of species and of species hybrids in + doves, and the separation of species-specific substances in + the backcross. Jour. Exp. Zool., 73:85-108, 1 fig. + + +LEONE, C. A. + + 1949. Comparative serology of some brachyuran crustacea and + studies in hemocyanin correspondence. Biol. Bull., + 97:273-286, 3 figs. + + 1953. Some effects of formalin on the serological activity of + crustacean and mammalian sera. Jour. Immun., 70:386-392, + 2 figs. + + +LIBBY, R. L. + + 1938. The photronreflectometer--an instrument for the measurement + of turbid systems. Jour. Immun., 34:71-73, 1 fig. + + +MARTIN, E. P., and LEONE, C. A. + + 1952. Serological relationships among domestic fowl as shown by + comparisons of protein preparations from corresponding organ + systems. Trans. Kansas Acad. Sci., 55:439-444, 1 fig. + + +MCGIBBON, W. H. + + 1945. Further division of contrasting antigens in species hybrids + in ducks. Genetics, 30:252-265. + + +SASAKI, K. + + 1928. Serological examination of the blood-relationship between + wild and domestic ducks. Jour. Dept. Agri., Kyushu Imp. + Univ., 2:117-132. + + +SIMPSON, G. G. + + 1944. Tempo and mode in evolution. Columbia Univ. Press, New York, + xviii + 237 pp., 36 figs. + + +SUSHKIN, P. P. + + 1924. [On the Fringillidae and allied groups.] Bull. British + Ornith. Club, 45:36-39. + + 1925. The evening grosbeak (Hesperiphona), the only American genus + of a Palaearctic group. Auk, 42:256-261, 2 figs. + + +TORDOFF, H. B. + + 1954. A systematic study of the avian family Fringillidae, based + on the structure of the skull. Univ. Michigan Mus. Zool. + Misc. Publ. No. 81:1-42, 77 figs. + + +WETMORE, A. + + 1951. A revised classification for the birds of the world. + Smithsonian Misc. Coll., 117(4):1-22. + + +_Transmitted June 8, 1954._ + + +25-4632 + + + + +UNIVERSITY OF KANSAS PUBLICATIONS + +MUSEUM OF NATURAL HISTORY + + +Institutional libraries interested in publications exchange may obtain +this series by addressing the Exchange Librarian, University of Kansas +Library, Lawrence, Kansas. Copies for individuals, persons working in +a particular field of study, may be obtained by addressing instead the +Museum of Natural History, University of Kansas, Lawrence, Kansas. +There is no provision for sale of this series by the University +Library which meets institutional requests, or by the Museum of +Natural History which meets the requests of individuals. However, +when individuals request copies from the Museum, 25 cents should +be included, for each separate number that is 100 pages or more +in length, for the purpose of defraying the costs of wrapping and +mailing. + + * An asterisk designates those numbers of which the Museum's supply + (not the Library's supply) is exhausted. Numbers published to date, + in this series, are as follows: + + Vol. 1. 1. The pocket gophers (Genus Thomomys) of Utah. By Stephen D. + Durrant. Pp. 1-82, 1 figure in text; August 15, 1946. + + 2. The systematic status of Eumeces pluvialis Cope, and + noteworthy records of other amphibians and reptiles from + Kansas and Oklahoma. By Hobart M. Smith. Pp. 85-89. + August 15, 1946. + + 3. The tadpoles of Bufo cognatus Say. By Hobart M. Smith. + Pp. 93-96, 1 figure in text. August 15, 1946. + + 4. Hybridization between two species of garter snakes. + By Hobart M. Smith. Pp. 97-100. August 15, 1946. + + 5. Selected records of reptiles and amphibians from Kansas. + By John Breukelman and Hobart M. Smith. Pp. 101-112. + August 15, 1946. + + 6. Kyphosis and other variations in soft-shelled turtles. + By Hobart M. Smith. Pp. 117-124, 3 figures in text. + July 7, 1947. + + *7. Natural history of the prairie vole (Mammalian Genus + Microtus). By E. W. Jameson, Jr. Pp. 125-151, 4 figures in + text. October 6, 1947. + + 8. The postnatal development of two broods of great horned + owls (Bubo virginianus). By Donald F. Hoffmeister and + Henry W. Setzer. Pp. 157-173, 5 figures in text. + October 6, 1947. + + 9. Additions to the list of the birds of Louisiana. By George + H. Lowery, Jr. Pp. 177-192. November 7, 1947. + + 10. A check-list of the birds of Idaho. By M. Dale Arvey. + Pp. 193-216. November 29, 1947. + + 11. Subspeciation in pocket gophers of Kansas. By Bernardo + Villa R. and E. Raymond Hall. Pp. 217-236, 2 figures in + text. November 29, 1947. + + 12. A new bat (Genus Myotis) from Mexico. By Walter W. + Dalquest and E. Raymond Hall. Pp. 237-244, 6 figures in + text. December 10, 1947. + + 13. Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. + By Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, + 1 figure in text. December 10, 1947. + + 14. A new pocket gopher (Thomomys) and a new spiny pocket + mouse (Liomys) from Michoacan, Mexico. By E. Raymond Hall + and Bernardo Villa R. Pp. 249-256, 6 figures in text. + July 26, 1948. + + 15. A new hylid frog from eastern Mexico. By Edward H. Taylor. + Pp. 257-264, 1 figure in text. August 16, 1948. + + 16. A new extinct emydid turtle from the Lower Pliocene of + Oklahoma. By Edwin C. Galbreath. Pp. 265-280, 1 plate. + August 16, 1948. + + 17. Pliocene and Pleistocene records of fossil turtles from + western Kansas and Oklahoma. By Edwin C. Galbreath. + Pp. 281-284. August 16, 1948. + + 18. A new species of heteromyid rodent from the Middle + Oligocene of northeastern Colorado with remarks on the + skull. By Edwin C. Galbreath. Pp. 285-300, 2 plates. + August 16, 1948. + + 19. Speciation in the Brazilian spiny rats (Genus Proechimys, + Family Echimyidae). By Joao Moojen. Pp. 301-406, + 140 figures in text. December 10, 1948. + + 20. Three new beavers from Utah. By Stephen D. Durrant and + Harold S. Crane. Pp. 407-417, 7 figures in text. + December 24, 1948. + + 21. Two new meadow mice from Michoacan, Mexico. By E. Raymond + Hall. Pp. 423-427, 6 figures in text. December 24, 1948. + + 22. An annotated check list of the mammals of Michoacan, + Mexico. By E. Raymond Hall and Bernardo Villa R. + Pp. 431-472, 2 plates, 1 figure in text. December 27, 1949. + + 23. Subspeciation in the kangaroo rat, Dipodomys ordii. + By Henry W. Setzer. Pp. 473-573, 27 figures in text, + 7 tables. December 27, 1949. + + 24. Geographic range of the hooded skunk, Mephitis macroura, + with description of a new subspecies from Mexico. + By E. Raymond Hall and Walter W. Dalquest. Pp. 575-580, + 1 figure in text. January 20, 1950. + + 25. Pipistrellus cinnamomeus Miller 1902 referred to the Genus + Myotis. By E. Raymond Hall and Walter W. Dalquest. + Pp. 581-590, 5 figures in text. January 20, 1950. + + 26. A synopsis of the American bats of the Genus Pipistrellus. + By E. Raymond Hall and Walter W. Dalquest. Pp. 591-602, + 1 figure in text. January 20, 1950. + + Index. Pp. 605-638. + + *Vol. 2. (Complete) Mammals of Washington. By Walter W. Dalquest. + Pp. 1-444, 140 figures in text. April 9, 1948. + + Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and + distribution. By Rollin H. Baker. Pp. 1-359, 16 figures + in text. June 12, 1951. + + *2. A quantitative study of the nocturnal migration of birds. + By George H. Lowery, Jr. Pp. 361-472, 47 figures in text. + June 29, 1951. + + 3. Phylogeny of the waxwings and allied birds. By M. Dale + Arvey. Pp. 473-530, 49 figures in text, 13 tables. + October 10, 1951. + + 4. Birds from the state of Veracruz, Mexico. By George H. + Lowery, Jr., and Walter W. Dalquest. Pp. 531-649, + 7 figures in text, 2 tables. October 10, 1951. + + Index. Pp. 651-681. + + *Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466, + 41 plates, 31 figures in text. December 27, 1951. + + Vol. 5. 1. Preliminary survey of a Paleocene faunule from the Angels + Peak area, New Mexico. By Robert W. Wilson. Pp. 1-11, + 1 figure in text. February 24, 1951. + + 2. Two new moles (Genus Scalopus) from Mexico and Texas. + By Rollin H. Baker. Pp. 17-24. February 28, 1951. + + 3. Two new pocket gophers from Wyoming and Colorado. + By E. Raymond Hall and H. Gordon Montague. Pp. 25-32. + February 28, 1951. + + 4. Mammals obtained by Dr. Curt von Wedel from the barrier + beach of Tamaulipas, Mexico. By E. Raymond Hall. + Pp. 33-47, 1 figure in text. October 1, 1951. + + 5. Comments on the taxonomy and geographic distribution of + some North American rabbits. By E. Raymond Hall and Keith + R. Kelson. Pp. 49-58. October 1, 1951. + + 6. Two new subspecies of Thomomys bottae from New Mexico and + Colorado. By Keith R. Kelson. Pp. 59-71, 1 figure in text. + October 1, 1951. + + 7. A new subspecies of Microtus montanus from Montana and + comments on Microtus canicaudus Miller. By E. Raymond Hall + and Keith R. Kelson. Pp. 73-79. October 1, 1951. + + 8. A new pocket gopher (Genus Thomomys) from eastern Colorado. + By E. Raymond Hall. Pp. 81-85. October 1, 1951. + + 9. Mammals taken along the Alaskan Highway. By Rollin H. + Baker. Pp. 87-117, 1 figure in text. November 28, 1951. + + *10. A synopsis of the North American Lagomorpha. By E. Raymond + Hall. Pp. 119-202, 68 figures in text. December 15, 1951. + + 11. A new pocket mouse (Genus Perognathus) from Kansas. + By E. Lendell Cockrum. Pp. 203-206. December 15, 1951. + + 12. Mammals from Tamaulipas, Mexico. By Rollin H. Baker. + Pp. 207-218. December 15, 1951. + + 13. A new pocket gopher (Genus Thomomys) from Wyoming and + Colorado. By E. Raymond Hall. Pp. 219-222. + December 15, 1951. + + 14. A new name for the Mexican red bat. By E. Raymond Hall. + Pp. 223-226. December 15, 1951. + + 15. Taxonomic notes on Mexican bats of the Genus Rhogeessa. + By E. Raymond Hall. Pp. 227-232. April 10, 1952. + + 16. Comments on the taxonomy and geographic distribution of + some North American woodrats (Genus Neotoma). By Keith R. + Kelson. Pp. 233-242. April 10, 1952. + + 17. The subspecies of the Mexican red-bellied squirrel, + Sciurus aureogaster. By Keith R. Kelson. Pp. 243-250, + 1 figure in text. April 10, 1952. + + 18. Geographic range of Peromyscus melanophrys, with + description of new subspecies. By Rollin H. Baker. + Pp. 251-258, 1 figure in text. May 10, 1952. + + 19. A new chipmunk (Genus Eutamias) from the Black Hills. + By John A. White. Pp. 259-262. April 10, 1952. + + 20. A new pinon mouse (Peromyscus truei) from Durango, Mexico. + By Robert B. Finley, Jr. Pp. 263-267. May 23, 1952. + + 21. An annotated checklist of Nebraskan bats. By Olin L. Webb + and J. Knox Jones, Jr. Pp. 269-279. May 31, 1952. + + 22. Geographic variation in red-backed mice (Genus + Clethrionomys) of the southern Rocky Mountain region. + By E. Lendell Cockrum and Kenneth L. Fitch. Pp. 281-292, + 1 figure in text. November 15, 1952. + + 23. Comments on the taxonomy and geographic distribution of + North American microtines. By E. Raymond Hall and + E. Lendell Cockrum. Pp. 293-312. November 17, 1952. + + 24. The subspecific status of two Central American sloths. + By E. Raymond Hall and Keith R. Kelson. Pp. 313-317. + November 21, 1952. + + 25. Comments on the taxonomy and geographic distribution of + some North American marsupials, insectivores, and + carnivores. By E. Raymond Hall and Keith R. Kelson. + Pp. 319-341. December 5, 1952. + + 26. Comments on the taxonomy and geographic distribution of + some North American rodents. By E. Raymond Hall and Keith + R. Kelson. Pp. 343-371. December 15, 1952. + + 27. A synopsis or the North American microtine rodents. + By E. Raymond Hall and E. Lendell Cockrum. Pp. 373-498, + 149 figures in text. January 13, 1953. + + 28. The pocket gophers (Genus Thomomys) of Coahuila, Mexico. + By Rollin H. Baker. Pp. 499-514, 1 figure in text. + June 1, 1953. + + 29. Geographic distribution of the pocket mouse, Perognathus + fasciatus. By J. Knox Jones, Jr. Pp. 515-526, 7 figures in + text. August 1, 1953. + + 30. A new subspecies of wood rat (Neotoma mexicana) from + Colorado. By Robert B. Finley, Jr. Pp. 527-534, 2 figures + in text. August 15, 1953. + + 31. Four new pocket gophers of the genus Cratogeomys from + Jalisco, Mexico. By Robert J. Russell. Pp. 535-542. + October 15, 1953. + + 32. Genera and subgenera of chipmunks. By John A. White. + Pp. 543-561, 12 figures in text. December 1, 1953. + + 33. Taxonomy of the chipmunks, Eutamias quadrivittatus and + Eutamias umbrinus. By John A. White. Pp. 563-582, + 6 figures in text. December 1, 1953. + + 34. Geographic distribution and taxonomy of the chipmunks of + Wyoming. By John A. White. Pp. 584-610, 3 figures in text. + December 1, 1953. + + 35. The baculum of the chipmunks of western North America. + By John A. White. Pp. 611-631, 19 figures in text. + December 1, 1953. + + 36. Pleistocene Soricidae from San Josecito Cave, Nuevo Leon, + Mexico. By James S. Findley. Pp. 633-639. December 1, 1953. + + 37. Seventeen species of bats recorded from Barro Colorado + Island, Panama Canal Zone. By E. Raymond Hall and William + B. Jackson. Pp. 641-646. December 1, 1953. + + Index. Pp. 647-676. + + *Vol. 6. (Complete) Mammals of Utah, _taxonomy and distribution_. + By Stephen D. Durrant. Pp. 1-549, 91 figures in text, + 30 tables. August 10, 1952. + + Vol. 7. *1. Mammals of Kansas. By E. Lendell Cockrum. Pp. 1-303, + 73 figures in text, 37 tables. August 25, 1952. + + 2. Ecology of the opossum on a natural area in northeastern + Kansas. By Henry S. Fitch and Lewis L. Sandidge. + Pp. 305-338, 5 figures in text. August 24, 1953. + + 3. The silky pocket mice (Perognathus flavus) of Mexico. + By Rollin H. Baker. Pp. 339-347, 1 figure in text. + February 15, 1954. + + 4. North American jumping mice (Genus Zapus). By Philip H. + Krutzsch. Pp. 349-472, 47 figures in text, 4 tables. + April 21, 1954. + + 5. Mammals from Southeastern Alaska. By Rollin H. Baker and + James S. Findley. Pp. 473-477. April 21, 1954. + + 6. Distribution of Some Nebraskan Mammals. By J. Knox Jones. + Pp. 479-487. April 21, 1954. + + 7. Subspeciation in the montane meadow mouse, Microtus + montanus, in Wyoming and Colorado. By Sydney Anderson. + Pp. 489-506, 2 figures in text. July 23, 1954. + + 8. A new subspecies of bat (Myotis velifer) from Southeastern + California and Arizona. By Terry A. Vaughn. Pp. 507-512. + July 23, 1954. + + 9. Mammals of the San Gabriel Mountains of California. + By Terry A. Vaughn. Pp. 513-582, 1 figure in text, + 12 tables. November 15, 1954. + + More numbers will appear in volume 7. + + Vol. 8. 1. Life History and Ecology of the Five-Lined Skink, Eumeces + fasciatus. By Henry S. Fitch. Pp. 1-156, 26 figures in + text. September 1, 1954. + + 2. Myology and Serology of the Avian Family Fringillidae, + a Taxonomic Study. By William B. Stallcup. Pp. 157-211, + 23 figures in text, 4 tables. November 15, 1954. + + More numbers will appear in volume 8. + + + + + * * * * * + + + Transcriber's Notes + + The text presented is essentially that in the original printed + document with the exception of some minor punctuation changes and + the typographical correction detailed below. Some of the tables + split between paragraphs in the original and they were moved and + the paragraphs restored into one. The captions for Figures 10-13 + and 14-17 were reformatted to enhance readability. + + + Empasis Notation + + _Text_ - Italics + + +Text+ - Bold + + + Typographical Corrections + + Page 187, Table 1 Item 5: Intavenous => Intravenous + + + * * * * * + + + + + +End of the Project Gutenberg EBook of Myology and Serology of the Avian +Family Fringillidae, by William B. 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