Front matter (continued 2)

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.

Here proposedProposed by Tordoff (1954) on the basis of the bony palate:Proposed by Beecher (1953) on the basis of jaw-musculature:

Family Ploceidae Family Ploceidae Family Ploceidae

[Subf. Bubalornithinae]Subf. Bubalornithinae

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 Passerinae Subfamily Passerinae

[Subfamily Ploceinae]Subfamily PloceinaeSubfamily Ploceinae

[Subfamily Viduinae]Subfamily ViduinaeSubfamily Viduinae

Family Carduelidae

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.Subfamily EstrildinaeFamily Estrildidae

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.Subfamily Carduelinae[In Thraupidae below]

Family Fringillidae: 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. Family Fringillidae Family Parulidae Subfamily Parulinae Subfamily Emberizinae

Family Thraupidae

Subf. Richmondeninae Subfamily Thraupinae Subfamily Emberizinae [Subfamily Fringillinae] [Subfamily Geospizinae]Subf. Richmondeninae Subfamily Thraupinae Subfamily Fringillinae (including Emberizinae and Geospizinae)Subfamily Pyrrhuloxiinae Subfamily Thraupinae [In Parulidae above] Subfamily Carduelinae

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.

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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.

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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.

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1873. On certain muscles in the thigh of birds and their value in classification. Proc. Zool. Soc. London, Part I:626-644, 6 figs.

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Greenberg, D. M.

1929. The colorimetric determination of serum proteins. J. Biol. Chem., 82:545-550.

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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.

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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.

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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.

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1944. Tempo and mode in evolution. Columbia Univ. Press, New York, xviii + 237 pp., 36 figs.

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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.

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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.

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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. 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.

* 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. 313-337. 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 of the North American microtine rodents. By E. Raymond Hall and E. Lendell Cockrum. Pp. 373-498, 149 figures in text. January 15, 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, Jr. 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 figs. 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.