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Systematics of the White-Throated Towhee (Pipilo Albicollis)

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Systematics of the White-Throated Towhee (Pipilo Albicollis) SHORT COMMUNICATIONS 457 BAGNARA, J. T., T. D. TAYLOR, AND M. E. HADLEY. LERNER, A. B. 1953. Metabolism of phenylalanine 1968. The dermal chromatophore unit. J. Cell and tyrosinase. Adv. Enzymol. 14:73-128. Biol. 38:67-79. LEWIN, V. 1963. Reproduction and development of ENGELS, W. L. 1959. The influence of different day young in a population of California Quail. Con- lengths on the testes of a transequatorial migrant, dor 65:249-278. the Bobolink (D&&onyx oryzioorus), p. 759- PFEIFFER, C. A., C. W. HOOKER, AND A. KIRKBAUM. 766. In Photoperiodism and related phenomena 1944. Deposition of pigment in the sparrows’ in plants and animals. Publ. 55, Amer. Assoc. bill in response to direct applications as a Adv. Sci., Washington, D.C. specific and quantitative test for androgen. FISHER, R. A., AND F. YATES. 1949. Statistical tables. Endocrinology 34 : 389-399. Hafner Publ. Co., Inc., New York. 112 p. POTTER, W. P., AND K. S. NORRIS. 1969. Lizard FLETCHER, R. A. 1971. Effects of vitamin A defi- reflectivity change and its effect on light trans- ciency on the pituitary-gonadal axis of California mission through body wall. Science 163:482- Quail, Lopho~rtyx californicus. J. Exp. Zool. 176: 484. 25-34. RAITT, R. J., AND R. D. OHMART. 1966. Annual cycle HADLEY, M. E. 1972. Functional significance of of reproduction and molt in Gambel Quail of vertebrate epidermal pigmentation. Amer. Zool. the Rio Grande Valley, southern New Mexico. 12: 63-76. Condor 68:541-561. HADLEY, M. E., AND W. C. QUEVEDO. 1966. Verte- RALPH, C. L. 1969. The control of color in birds. brate epidermal melanin unit. Nature 209:1334- Amer. Zool. 9:521-530. 1335. RALPH, C. L., D. L. GRINI~ICH, AND P. F. HALL. HALL, P. F. 1966. Tyrosinase activity in relation to 1967. Studies of the melanogenic response of plumage color in Weaver Birds. (Streganuru regenerating feathers in the Weaver Bird: Com- padisues ) , Comp. Biochem. Physiol. 18:91- parison of two species in response to two gonado- 100. tropins. J. Exp. Zool. 166:283-288. HAMILTON, H. L. 1941. Influence of adrenal and sex hormones on the differentiation of melano- SERVENTY, D. L., AND A. J. MARSHALL. 1956. Factors phores in the chick. J. Exp. Zool. 88:275-305. influencing testis coloration in birds. Emu 56: 219-222. JONES, R. E. 1970. Effect of season and gonadotropin on testicular interstitial cells of California Quail. STEEL, R. G. D., AND J. H. TORRIE. 1960. Principles Auk 87:729-737. and procedures of statistics. McGraw-Hill Book JUHN, M., AND R. G. GUSTAVSON. 1930. The produc- Co., New York, 481 p. tion of female genital subsidiary characters and STURKIE, P. D. 1965. Avian physiology. Comstock plumage sex characteristics by injection of human Publ. Assoc., Cornell Univ. Press, Ithaca. 766 p. placental hormones in fowls. J. Exp. Zool. 56: WILLIA~~S, C. R. 1967. The breeding biology of 31-66. California Quail in New Zealand. New Zealand KRUMREY, W. A., AXD I. 0. Buss. 1969. Observations Ecol. Sot. Proc. 14:88-99. on the adrenal gland of the African elephant. J. Mammal. 50:90-101. Accepted for publication 30 January 1974. SYSTEMATIC3 OF THE in intensity, in general, to the pigmentation of these WHITE-THROATED TOWHEE areas in adults), whereas all juveniles I saw of P. (PZPZLO ALBZCOLLZS) fuscus and Aberts’ Towhee, P. uberti, (which, with P. uZbico,llis, form the “brown towhee” group) com- KENNETH C. PARKES pletely lacked dorsal streaking. In M. kieneri the Carnegie Museum dorsal markings appeared as bars (actually transverse Pittsburgh, Pennsylvania 15213 expansions at the tips of somewhat suppressed streaks). The one juvenile of P. ulbicollis then avail- The White-throated Towhee, Pipilo ulbicollis, is a able (Moore Collection 32696) showed a “faint Mexican endemic species of limited distribution. barring” on the dorsum. Two additional juveniles Most of the published information about this species now before me (Carnegie Museum 141364; A. R. can be found in two papers, those of Davis ( 1951) Phillips 5195), as well as several in first prebasic and Marshall ( 1964). Davis used the specific name molt that retain some dorsal feathers of the juvenal Pipilo v.dus, but see Stresemann ( 1954). plumage, confirm this. The terminal barbs of the Marshalls’ careful studies of vocalizations indicate juvenal dorsal feathers have a few blackish barbules that PipiZo albicollis is less distinct from the Brown at the very tip, giving the effect of a faintly darker, Towhee, P. fuscus, than I had believed when I wrote narrow transverse barring or scalloping. This is pre- of this genus and its relatives in 1957 ( Parkes 1957). sumably what Marshall (1964:354) meant in stating I called attention to certain striking similarities be- of P. albicollis that its “marks are broader than long.” tween P. ulbicollis and Melozone kieneri and proposed As stated in my 1957 paper and confirmed by addi- that, judging from skins, P. ulbicollis formed a link tional specimens, the ventral markings of juvenile between P. fuscus and M. kieneri. Marshall found ulbicollis resemble those of the juvenile Melozone that some of the Mexican races of P. fuscus exhibited kieneri I examined in being irregularly distributed and some of the characters that I had thought were con- vaguely shaped dense spots, quite unlike those of any fined, in this group, to P. ulbicollis and M. kieneri, but other Pipilo. Marshall deprecated my statement that the latter two species still share characters not found P. fuscus and P. uberti differed from P. erythrophthal- in the other forms. One of the characters I invoked mus and P. chlorurus in completely lacking dorsal was the pattern of the juvenal plumage. I found that streaks. He found that “many juvenal fuscus [he did in true Pipilo (i.e., the erythrophthulmus group plus not mention abetii] are liberally streaked above and chlorurus) the juvenal plumage is distinctly streaked below; and that juvenal kieneri are usually streaked, above and below (the ventral streaking corresponding not spotted below.” My 1957 findings were based 458 SHORT COMMUNICATIONS in large part on borrowed material; at this point I can Davis emphasized the need for additional collecting say o& that I do not doubt Marshall-word,’ but to provide seasonally comparable samples. Paynter that the 53 iuveniles of P. eruthrovhthalmus and 8 ( 197O:lSO) listed parvirostris in the “Peters” Check- of P. chlorurk in the Carnegie”Muskum are all char- list with a query, citing Davis ( 1954) on the un- acterized by distinct longitudinal dorsal streaking. certainty of its validity. There is no trace of such streaking in the Carnegie On hand at Carnegie Museum are specimens of sample (part of a substantially larger series examined Pipilo albicollis recently collected by Allan R. Phillips, in 1957) of six P. fuscus and four I.’ aberti juveniles. Robert W. Dickerman, Otto Epping, and the writer. Marshall speaks several times of the “Canyon For comparison, 28 specimens, including 23 of the Towhee” (Pipilo fuscus part) as the “ancestor” or type series of paruirostris, were borrowed from the “ancestral stock” of the White-throated Towhee. I Moore Laboratory of Zoology. A list of specimens have long objected to the assignment of a living examined will be found at the end of this paper. species as the “ancestor” of another living species; In studying variation in this species, Davis ( 1951) but this objection aside, Marshalls’ interpretation did not wholly segregate his specimens by age class. leaves no room whatsoever for a possible relationship In discussing the molts and nlumaees of the brown between Pipilo albicollis and Melozone kieneri. He towhee group in general, he- wrote (p. 3) : “Birds suggests that Melozone may be related to Atlupetes undergoing the postjuvenal (first fall) [= first pre- and the chlorurus-eruthrovhthalmus eroun of Pivilo basic] molt retain the primaries and secondaries, and (rather than to P. afbicolks), and, f&her, that ;the‘ usually the rectrices.” A few lines later, however, he brown towhees may be connected with some members wrote “. rectrices are often replaced in the post- of the heterogeneous Aimophila.” I do not find these iuvenal molt. .” Insofar as P. albicollis is con- suggestions persuasive, based as they are on sub- cerned, the statement about the retention of juvenal jective impressions of a few vocalizations plus, in the remiges is too dogmatic. At least some individuals first case, “a black shiny bill plus addiction to densest do replace remiges. A. R. Phillips 7891, “windows understory brush” and, in the second, “brown colora- cover sk[ull] roof,” 20 November, has several short, tion and occurrence at edges of brush.” Marshalls’ sheathed remiges. Moore 31112, 17 September, is paper, nevertheless, is a major contribution to our molting from the narrow rectrices typical of juveniles knowledge of relationships among the brown towhee to the broad rectrices of the first basic plumage (see segment of Pipilo (if, indeed, they are truly members Davis 1951:3, fig. 1). It is also in heavy wing molt. of that genus. as Marshall and I and several other The old remiges are scarcely worn at all and are authors have wondered). obviously of the juvenal generation, whereas the old Mayr and Short (1970:82) state that “albicollis remiges of adults undergoing wing molt are generally and fuscus meet barely, if at all [italics mine], in worn and frayed. Virtually all of the parvirostris southern Mexico.” They cite Davis ( 1951) to sup- specimens from Moctum, Oaxaca, taken in .September port this statement, although Daviss’ map clearly and October (including Moore 31112) are moltina shows svmuatrv of the two snecies in almost half of remiges; the chances of all of these having been , & the range of albicollis and his text cites three localities adults, at that time of year, are slim, but unfortunately of sympatry.
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