.--The Green-tailed
Towhee, currently placed in the genus Chlorura Sclater, 1862, was originally described
by Audubon (1839). The twenty-second supplement (1947) to the American
Ornithologists' Union Check-List of North American birds designates the species as
Chlorura chlorura (Audubon). The species breeds from central Oregon and south-
central Montana to southern California, southeastern New Mexico, and central
western Texas. In general it is a bird of the mountains or, in the Great Basin, of
high plateau country. The purpose of the present paper is to present evidence to
show that this species is a true Pipilo allied to the type species, Pipilo erythroph-
thalmus, by way of the closely related Pipilo ocai, the Collared Towhee, of Mexico.
Few species of North American birds have been moved about generically more
than the Green-tailed Towhee. Ridgway (1901: 401-402) lists eight genera in the
synonymy of the species and there have been two others in use since 1901. The
species appeared most frequently in the literature under Pipilo prior to 1896 when
Ridgway placed it in a new monotypic genus, Oreospiza. He proposed this separation
in spite of his remarks favoring it as a Pipilo, which appeared in 1890 in a review
of a work by Salvin and Godman. Salvin and Godman (1879-1887) had placed the
species in Embernagra, and Ridgway correctly pointed out the line of relationship
to the true towhees via the rufous-capped Mexican forms currently known as P. ocai.
The original use of the genus Oreospiza for the Green-tailed Towhee appears on
page 439 of Ridgway's "Manual" (1896), and in the appendix (1896: 605) he writes
as follows:
"Page 439, Pipilo ohiorufus (=Oreospiza chlorura): This bird, which has been
referred by different authors to the genera "Embernagra" (i.e. Arremonops) and
Atlapetes, but which is really far more out of place in either than in Pipilo, I propose
to make the type of a new genus, Oreospiza, whose characters are intermediate
between, or rather a combination of, those of Pipilo and Zonotrichia."
This constitutes the original description of the genus.
Five years later Ridgway (1901: 399) wrote the following: "Oreospiga is inter-
mediate between Pipilo and Zonotrichia, though much nearer the former, with
which it agrees in its stout feet with long claws, rounded tail, and form of bill. Its
coloration, too, is not so abnormal for Pipilo as has been supposed, every feature of
color--rufous cap, white throat, yellow carpal edge, and olive-green upper parts--
being shared by some species of that genus, though by none in the stone conbination.
The wing, however, is very different from that of Pipilo, being quite the same in the
relative length of the primaries as that of Zonotrichia, that of Z. albicollis being
even more rounded."
In the statement quoted above with reference to color features, Ridgway's remark
that the rufous cap and other color characters are, "shared . .., though by none
in the same combination," is simply in error. Pipilo ocai combines these and other
color characters also found in the Green-tailed Towbee. It thus seems that Ridg-
way's only remaining basis for separation is that of wing shape which, in the Green-
tailed Towbee, is longer and more pointed than in Pipilo ocai and P. erythrophthalmus.
Long, pointed wings are characteristic of species which are strong flyers or which
make long migratory flights. Mayr (1942: 92, 95) discusses the correlation between
wing length and geographic distribution and points out that forms of cooler climates
tend (1) to have longer wings and (2) to be more migratory than forms of warmer
climates.
The Green-tailed Towbee is migratory, and it seems apparent that the relatively
longer wing is correlated with this habit.
Further changes in the generic position of the Green-tailed Towbee, following
Ridgway's description of Oreospiza, have been dictated solely by nomenclatural
problems. In 1915 Richmond proposed the genus Oberholseria to replace Oreospiza
Ridgway, 1896, because the latter was preoccupied by Oreospiza Keitel, 1857. The
change to Chlorura Sclater, 1862, was effected in 1947, as noted above, since Chlorurus
Swainson, 1839, was deemed not to preoccupy Sclater's name.
In view of the foregoing it can scarcely be argued that the basis for Oreospiza
Ridgway is entirely beyond question. It now remains to present the evidence in
favor of its inclusion with the true towhees in the genus Pipilo.
In two recent papers (1950; 1954) I have described the extensive hybridization
which occurs in Mexico between Pipilo erythrophthalmus and Pipilo ocai. Nearly
1,500 specimens showing evidence of hybridization between these two species have
been examined up to the present. The two species are strikingly different in color
pattern (1950: plate 11), and P. ocai is approximately 30 per cent heavier in weight
than P. erythrophthalmus. Moreover, these species show well-marked, but over-
lapping, ecological differences (1950: 152-153; 1954: 270-273). Their songs, although
similar, are easily distinguished, but the common call notes of the two are easily
confused. Their eggs are alike in color and pattern, and both build similar nests.
Both are species of dense underbrush and both scratch in the leaf litter in the same
manner. P. ocai is primarily a bird of the understory vegetation at the higher
elevations where the dominant trees are conifers. P. erythrophthalmus occurs most
abundantly in the brushy thickets at lower elevations where oaks are more frequently
dominant. Hybridization occurs at some of the points where the two species come
into contact.
The situation concerning the degree of relationship between ocai and erythroph-
thalmus is relevant here in order to establish beyond any doubt that these two are
properly considered congeneric. If this be granted then it remains only to demon-
strate the close relationship between P. ocai and the Green-tailed Towbee in order
to show the latter also to be a Pipilo.
In color pattern the Green-tailed Towbee is far more similar to P. ocai than the
latter is to/. erythrophthalmus. The Green-tailed Towbee looks like a diminutive
edition of ocai with the breast and facial areas gray rather than black. If size alone
is considered of importance it may be pointed out that one race of/. erythrophthalmus,
the Socorro Island population,/. e. carmani, is smaller than the Green-tailed Towbee.
The ecological preferences of the Collared and Green-tailed towhees are extremely
similar. Both occur in dense brushy vegetation, and in many parts of its range the
Green-tailed occupies the shrubby undergrowth associated with conifers.
The songs of the two species are recognizably different but show a remarkably
similar pattern. After becoming familiar with the song of/. ocai at many localities
in Mexico in 1946 and 1948, I heard the song of the Green-tailed Towbee near Irwin,
Bonneville County, Idaho, on July 10, 1949. The similarity in the two songs was
striking. The call notes are even more alike. Both utter a roewing note which
may be written, "zree" or "zew." This cat-like note is also characteristic of many
races of /. erythrophthalmus, especially the "Spotted Towbees" of the western
United States and Mexico.
In coloration and pattern of markings, the eggs of /. erythrophthalmus, J>. ocai,
and the Green-tailed Towbee are nearly identical. All of these species lay eggs
which have a whitish ground color with reddish markings.
The evidence presented above contains many points in favor of including the
Greea-tailed Towbee in J>ipilo, but there is another line of reasoning which, although
negative, bears on the question. The so-called brown towbees, lipilo fuscus,
rutilis, and aberti, are probably not so closely allied to/. erythrophthalmus as is the
Green-tailed Towbee. Davis (1951: 100-102) suggests that the brown towbees may
be more closely related to the genus Melozone and not congeneric with the red-eyed
towbees, ocai and erythrophthalmus. I am in agreement with Davis on this point
and propose, as a first step toward the correct generic alignment of these various
forms, that the Green-tailed Towbee be recognized as a true J>ipilo. If one is to
disagree with this proposal he must justify the inclusion of the brown towbees in
J>ipilo because they are without doubt less closely related to the type species, Pipilo
erythrophthalmus, than is the Green-tailed Towbee.
The name of the Green-tailed Towbee becomes Pipilo chlorurus (Audubon) if
this recommendation is accepted.
LITERATURE CITED
AMERICAN ORNITHOLOGISTS' UNION. 1947. Twenty-second supplement to the
American Ornithologists' Union check-list of North American birds. Auk, 64:
445-452.
AUDUBON, J.j. 1839. Ornithological Biography, vol. 5. (Edinburgh.)
DAvis, J. 1951. Distribution and variation of the brown towbees. Univ. Calif.
Publ. Zool., 52: 1-120.
KEITE, G.T. 1857. Verzeichniss der EuropSischen V6gel .... (etc.) (Berlin.)
MAYR, E. 1942. Systematics and the origin of species. (New York, Columbia
Univ. Press), 334 pp.
RICHMOND, C. W. 1915. Notes on several preoccupied generic names (Aves).
Proc. Biol. Soc. Wash., 28: 180.
RIDGWA3/4, R. 1890. Review of Salvin and Godman's Biologia Centrali-Americana:
Aves. Auk, 7: 189-195.
RIDGWA3/4, R. 1896. Manual of North American birds. Second edition. (Phila-
delphia, J. B. Lippincott), 653 pp.
RmGw,3/4, R. 1901. The birds of North and Middle America. Bull. U.S. Nat.
Mus., 50, part 1: xxx q- 715 pp.
S,w, O., and F. D. GoDM,. 1879-1887. Biologia Centrali-Americana. Aves.
Vol. 1 (London) xliv q- 512 pp.
Sc,xR, P. L. 1862. Catalogue of a collection of American birds. (London),
xvi q- 338, 20 pls.
SB3/4, C.G. 1950. Species formation in the red-eyed towhees of Mexico. Univ.
Calif. Publ. Zool., 50: 109-194.
SmokY, C.G. 1954. Hybridization in the red-eyed towhees of Mexico. Evolution,
8: 252-290.
8WAINSON, W. 1839. On the natural history and classification of fishes, amphibians
and reptiles. (London).
CAm,r:s G. Smr:3/4, Department of Conservation, Cornell University,
Ithaca, New York.