Sushkin's studies (1924, 1925) of features of the h.o.r.n.y 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 pa.s.serine 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 a.s.sumes 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 a.s.sumes that such patterns may be used as indicators of relationship throughout the entire oscinine group. These two a.s.sumptions, then, serve as the basis for his hypothesis concerning relationship and phylogeny within this a.s.semblage. 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 h.o.r.n.y 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 a.s.sociated with food-getting may be subject to rapid evolutionary change. Finally, in attempting to cla.s.sify 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 a.s.sumes that since palato-maxillaries seem to be absent in the majority of pa.s.serine 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 a.s.sumes, therefore, that palato-maxillaries were evolved to provide for a more effective action of the _m. pterygoideus_. The need for such action could be a.s.sociated 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 a.s.sociated 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 Pa.s.seriformes, 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 quant.i.tative 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 a.s.semblage formed by the richmondenines, emberizines, and tanagers. Actually, the carduelines show less serological resemblance to this a.s.semblage 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 a.s.semblage, serologically, than do the carduelines.

Although the Carduelinae const.i.tute 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 cla.s.sification include the subfamilies Pa.s.serinae and Estrildinae in the Family Ploceidae. _Pa.s.ser_, 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 h.o.m.ogeneity of the Family Ploceidae as presently recognized by most ornithologists. If the Pa.s.serinae 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 const.i.tute 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 a.s.semblage, 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 cla.s.sifications place the Estrildinae and Pa.s.serinae 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 cla.s.sifications; 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 d.i.c.kcissel, _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 a.s.signed (Subfamily Richmondeninae). _Spiza_, serologically, is closely related to all species of the richmondenine-emberizine-thraupid a.s.semblage.

Within this a.s.semblage 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 a.s.semblage. 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 const.i.tute 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 cla.s.sification 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 Pa.s.serinae: |Subfamily Pa.s.serinae | Subfamily Pa.s.serinae 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 pa.s.serine 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 a.n.a.lysis 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 const.i.tute an a.s.semblage distinct from the other species studied. The Carduelinae are excluded from the a.s.semblage and serologically are most closely related to the Estrildinae. The estrildines, serologically, do not closely resemble _Pa.s.ser_, Subfamily Pa.s.serinae, although recent cla.s.sifications 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 Pa.s.serinae. 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.