Myology and Serology of the Avian Family Fringillidae: a Taxonomic Study

Part 1

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================================================================== 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.

Part 2

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.

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