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Home , California quail, Callipepla, Colinus, New World quail, Quail, Scaled quail

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1970

A Summary of Intergeneric New World Hybrids, and a New Intergeneric Hybrid Combination

Paul A. Johnsgard University of Nebraska-Lincoln, [email protected]

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Johnsgard, Paul A., "A Summary of Intergeneric Hybrids, and a New Intergeneric Hybrid Combination" (1970). Papers in Ornithology. 77. https://digitalcommons.unl.edu/biosciornithology/77

This Article is brought to you for free and open access by the Papers in the Biological Sciences at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in Ornithology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Johnsgard in CONDOR (January 1970) 72(1). Copyright 1970, University of and Cooper Ornithological Society. Used by permission.

A SUMMARY OF INTERGENERIC NEW WORLD QUAIL HYBRIDS, AND A NEW INTERGENERIC HYBRID COMBINATION

PAUL A. JOHNSGARD Department of Zoology University of Nebraska Lincoln, Nebraska 68505

The exceedingly close affinities of the quail x LOPHORTYX and have genera , Callipepla, Lophortyx The range of the overlaps fairly been for some time and have re- recognized extensively with that of the Gambel Quail (L. been additional cently emphasized by morpho- gambelii), primarily in New (Campbell Hudson et al. bio- logical (Holman 1961; 1966), and Lee 1953; Ligon 1961), but also in western chemical and (Sibley 1960), pterylographic along the Rio Grande (Texas Game, evidence. Holman has (Ohmart 1967) Indeed, Fish, and Oyster Commission 1949), south- that these three taxa best be suggested might eastern (Phillips et al. 1964), and ad- as et al. regarded subgenera. Phillips (1964) jacent Mexico (Leopold 1959). Introductions with Cal- pragmatically merged Lophortyx of the Scaled Quail into central Washington lipepla; and Delacour (1961-1962) synony- have also resulted in a small amount of con- mized not but also only Lophortyx, Oreortyx tact with the (L. californicus), and with retained Philortyx, Callipepla, yet and two hybrid specimens have been reported Colinus as a Ohmart separate . Although from that area (Jewett et al. 1953). Shore-baily the of (1967) questioned validity generically (1912, 1913, 1914, 1917) bred and reared a and he separating Lophortyx Callipepla, sug- number of such hybrids, and reported them to that new taxonomic gested any reorganization be fertile. There is apparently no natural con- should also include consideration of such re- tact between the Scaled Quail and the Douglas lated as Colinus. genera Quail (L. douglasii) in northern Mexico, but One useful source of evidence for judging several examples (all apparently sterile) of such distinction is the occurrence possible generic a hybrid combination have been obtained in of both and ob- naturally occurring artificially captivity (Banks and Walker 1964). tained and their relative hybrids, fertility. Wild hybrids between the Scaled Quail and before taxonomic Thus, advocating any Gambel Quail have been known to occur for the New World changes among quail genera, some time; apparently the earliest published a review of recorded intergeneric hybrids record is that of Bailey (1928), who described would seem to be in . a male hybrid shot from a covey of Gambel 1916 in Grant New CALLIPEPLA x COLINUS Quail during County, Mexico. Hachisuka (1928) also illustrated this Natural sympatry between the Bobwhite (Co- specimen. More recently Hubbard (1966) de- linus virginianus) and the Scaled Quail (Cal- scribed an apparent back-cross to Gambel lipepla squamata) exists across a fairly broad Quail, taken in Grant County in 1930. Phillips zone extending from northern Mexico through et al. (1964) reported three hybrid specimens west-central Texas (McCabe 1954), the Okla- from different localities in southeastern Arizona homa panhandle (Schemnitz 1964; Sutton and Lowe (1964) mentioned wild hybrids from 1967), possibly extreme southwestern one of the same areas. Captive hybrids have (Johnston 1964), and southeastern been reported by Plath (1953) and have also (Hoffman 1965; Bailey and Niedrach 1965). been obtained by the author (male Scaled x Wild hybrids have been reported in Texas female Gambel). According to J. S. Ligon, such from Concho County (McCabe 1954), Stone- female hybrids lay only sterile (Sibley wall County (Sutton 1963), and Motley County 1961), but eggs that developed nearly to hatch- (Schemnitz 1964). Probable hybrids have also ing (when the incubator failed) have been pro- been seen in (Sutton 1967). Captive duced by a hybrid pair at the Arizona- hybrids have also been produced on a variety Desert Museum (L. W. Walker, pers. comm.). of occasions (Gray 1958). Although the hybrid x females are known to produce eggs, attempted LOPHORTYX OREORTYX inter se matings and back-cross matings with The area of geographic overlap between the Bobwhites have not yet proven successful (0. picta) and the California (Sutton 1963). Quail is considerable and includes much of [85] The Condor, 72:85-88, 1970 Johnsgard in CONDOR (January 1970) 72(1). Copyright 1970, University of California and Cooper Ornithological Society. Used by permission.

86 PAUL A. JOHNSGARD

California, Oregon, and western Washington (Grinnell and Miller 1944; Jewett et al. 1953; :x:i~aii S-Z The 2U, Masson and Mace 1962). earliest record ..Xii. dii$ of a hybrid between these is that of Wh'H-', .41: . .... Xii-ii;;iiJ' h?:;:r:ri:.:NXi~J Peck (1911), who described a specimen taken zmtrx in 1911 in Harney County, Oregon. Hachisuka ...... (1928) provided an illustration of this hybrid 9~i~si~'r~sb~~i~ii~i~ii~i...... ;~~~iilJ~r~ specimen. Peterle (1951) published a repro- duction of a painting by L. A. Fuertes of a presumably wild hybrid in the collection of II L. M. Loomis, the skin of which is no longer .... extant...... There do not to be be-

hybrids . appear any . .:------tween Oreortyx and Lophortyx produced in ...... :..a captivity, nor are there any naturally occurring or captive-bred hybrids known that involve the Mountain Quail and Callipepla or Colinus. With the possible exception of the introduced Bobwhite in northern Oregon (Masson and FIGURE 1. Pair of F1 Bobwhite X Gambel Quail Mace 1962), the current range of the Mountain hybrids, at 140 days of age. Quail would preclude any natural hybridiza- tion with these species. Interestingly, Holman (1961) suggests that Oreortyx may actually be Gambel Quail was established. The were more closely related to Callipepla than to not allowed visual contact with others of their Lophortyx, which points out the fallacy of dis- species, but auditory communication was pos- tinguishing genera primarily on the basis of sible. They were placed on a 17-hr photoperiod crest condition. and were minimally disturbed, although this was a few feet from office LOPHORTYX x COLINUS pair kept only my desk. The female began to produce eggs in Natural sympatry between these two genera is early February, and while several were used essentially non-existent, although there may be for other purposes, a total of 17 eggs were some overlapping of marginal ranges of the artificially incubated. All of them proved to Bobwhite and the Gambel Quail in extreme be fertile, but of the first group of five eggs western Texas near El Paso (Peterson 1960). only a single individual hatched, the other Additionally, there is some potential contact four having died just prior to pipping. This between the nearly extinct Masked Bobwhite individual also died two days later, perhaps (C. v. ridgwayi) and the Gambel Quail or as a result of faulty incubation procedures. Douglas Quail in western Mexico (Leopold The 12 remaining eggs all hatched between 30 1959). Furthermore, introduction of the Bob- March and 16 April and resulted in apparently white into Washington, Oregon, and Idaho has healthy and normal chicks. Various accidents resulted in some probable contact with the and cannibalism gradually reduced this num- California Quail. Finally, introduction of the ber, but two individuals were ultimately reared Bobwhite and the California Quail into to their first winter (figure 1). also produced a relatively short-lived sympatry between them and resulted in the only known TAXONOMIC IMPLICATIONS naturally occurring intergeneric hybridization The primary criteria for the genera Lophortyx, between Colinus and Lophortyx. Aiken (1930) Callipepla, and Colinus have been variations described three probable Bobwhite x Califor- in the number of rectrices (14 in Callipepla, nia Quail hybrids taken in 1895 near Salt Lake 12 in the others), the degree of sexual dimor- City. Compton (1932) likewise described a phism, and the condition of the crest (Ridgway male hybrid that apparently resulted from and Friedmann 1946). The first of these criteria keeping a female Bobwhite in a pen with is weakened by the fact that some individuals California Quail. of at least two species of Lophortyx may have As part of a general study on the behavior as many as 14 rectrices (Clark 1898; Ohmart and relationships of odontophorine quail, I 1967). The degree of sexual dimorphism is of have attempted to produce various intergeneric little or no significance at the generic level, hybrids involving Colinus, Lophortyx, and Cal- since it is the result of various factors such as lipepla. In October 1967 a mixed pair involv- the intensity of sexual selection and the rela- ing an immature male Bobwhite and female tive requirements for species and sexual recog- Johnsgard in CONDOR (January 1970) 72(1). Copyright 1970, University of California and Cooper Ornithological Society. Used by permission.

INTERGENERIC NEW WORLD QUAIL HYBRIDS 87

crosses of the Douglas Quail with the Bob- white and Gambel Quail, although Ridgway and Friedmann (1946) believe that "Lophortyx leucoprosopon" may have been described on the basis of two captive-bred birds represent- Bobwhite ing the latter hybrid combination. At least one intergeneric cross (Scaled x California Quail) has been reported to be fully fertile, and one ScaledQuail other Callipepla x Lophortyx combination is apparently occasionally fertile. The fertility of _. the Colinus x Lophortyx cross is as yet un- Gambel'sQuaill known. Continued comparative studies on all these and their are and ulti- CaliforniaQuail species hybrids planned, mately specific taxonomic recommendations are contemplated. At present, however, it is Z PresumedWild Hybrids Reported clearly apparent that serious consideration should be given to merging the genera Lophor- tyx and Callipepla with Colinus, and that the HybridsReported From Captivity [ question of the validity of Oreortyx and Philor- tyx should be kept open for further assessment. and CaptiveHybrids; Infertile SWildor Fertility Not Established SUMMARY A review of hybridization records among the SHybrids Reportedto be at least New World quail indicates that the five North Fertile Occasionally American species of Colinus, Callipepla, and have been in at least [:7 ApparentlySympatric at Present Lophortyx hybridized Sor in the RecentPast eight of the ten possible combinations. On the basis of inadequately distinctive morphological FIGURE 2. Summary of interspecific hybrid records the several known cases of among species of Colinus, Callipepla, and Lophortyx. criteria, naturally Sympatric contact has also been indicated, even in occurring intergeneric hybridization, and the situations of slight geographic overlap or possible eco- intergeneric fertility of at least one such hy- logical barriers to extensive contact. brid combination, it is suggested that Lophor- tyx and Callipepla be merged with Colinus. ACKNOWLEDGMENTS nition signals (Sibley 1957). Finally, although crest shape and length among the species of This study was financed in part by a National Science and varies Foundation research grant (GB-7666X), and in part by Lophortyx Callipepla considerably, of Nebraska it the of these University Zoology Department funds; pterylographic arrangements feath- is Study No. 394 from the Department of Zoology. I ers are almost identical (Ohmart 1967). Differ- express my appreciationto both of these organizations. ences in the relative proportions of tail-to-wing LITERATURE CITED lengths and the variations in scapular and ter- tial spotting are likewise unconvincing generic AIKEN,C. E. H. 1930. A Bobwhite X California criteria. Quail hybrid. Auk 47:80-81. the BAILEY, V. 1928. A hybrid Scaled X Gambel's Quail Collectively considering three species of from . Auk 45:210. the one of and Lophortyx, species Callipepla, BAILEY,A. M.,AND R. J. NIEDRACH. 1965. Birdsof the single North American species of Colinus, Colorado. Vol. 1. Denver Museum of Natural there are 10 intra- and intergeneric hybrid History, Denver. R. ANDL. W. WALKER.1964. combinations possible. At least eight of these BANKS, C., A hybrid Scaled X DouglasQuail. Wilson Bull. 76:378- have been actually achieved, including five 380. from presumably wild birds and seven from CAMPBELL,H., ANDL. LEE. 1953. Studieson quail captivity, while four combinations have oc- malaria in New Mexico, and notes on other curred both in and under natural aspects of quail populations. New Mexico Dept. captivity of Fish and Game, Project W-41-R. conditions (figure 2). Remarkably, four of the CLARK, H. L. 1898. The -tracts of North five recorded combinations that have occurred American and quail. Proc. U.S. Natl. Mus. naturally are intergeneric on the basic of cur- 21:641-653. rent nomenclature. So the COMPTON, L. V. 1932. A probable hybrid between far, only possible the California Quail and the Texas Bob-white. combinations that remain unrecorded are Condor 34:48. Johnsgard in CONDOR (January 1970) 72(1). Copyright 1970, University of California and Cooper Ornithological Society. Used by permission.

88 PAUL A. JOHNSGARD

DELACOUR, J. 1961-1962. The American PECK, M. E. 1911. A hybrid quail. Condor 13: (Tribe Odontophorini). Avicult. Mag. 67:12-18; 149-151. 68:15-19. PETERLE, T. J. 1951. Intergeneric galliform hybrids: GRAY, A. P. 1958. hybrids. Commonwealth a review. Wilson Bull. 63:219L-224. Agri. Bur., Royal Banks, England. PETERSON, R. T. 1960. A field guide to the birds of GRINNELL, J., AND A. H. MILLER. 1944. The distri- Texas. Houghton Mifflin, Boston. bution of the birds of California. Pacific Coast PHILLIPS, A. R., J. T. MARSHALL, AND G. MONSON. Avifauna, no. 27. 1964. The birds of Arizona. Univ. Arizona Press, HACHISUKA, M. 1928. Varieties among birds (chiefly Tucson. game birds). Suppl. Publ. Ornithol. Soc. Japan, PLATH, K. 1953. A diary for 1952 of new arrivals in no. 12. the Bird Department of the Brookfield Zoo at HOFFMAN, D. M. 1965. The Scaled Quail in Colo- Brookfield, Ill. Avicult. Mag. 59:104-107. rado. Tech. Publ. no. 18, Colorado Dept. of RIDGWAY, R., AND H. FRIEDMANN. 1946. The birds Game, Fish and Parks. of . U.S. Natl. Mus., Bull. 50. HOLMAN, J. A. 1961. Osteology of living and fossil Pt. 10. New World quails (Aves, ). Bull. SCHEMNITZ, S. D. 1964. Comparative ecology of Florida State Mus., Biol. Sci. 6(2):131-233. Bobwhite and Scaled Quail in the Oklahomapan- HUBBARD, J. P. 1966. A possible back-cross hybrid handle. Amer. Midland Naturalist 71:429-433. involving Scaled and Gambel's Quail. Auk 83: SHORE-BAILY, W. 1912. Quail hybrids. Bird Notes 136-137. 3:322. HUDSON, G. E., R. A. PARKER, J. V. BERGE, AND P. J. SHORE-BAILY,W. 1913. Fertility of hybrid quail. LANZILOTTI. 1966. A numerical analysis of the Bird Notes 4:3268. modificationsof the appendicularmuscles in vari- SHORE-BAILY,W. 1914. Hybrid quail. Bird Notes ous genera of gallinaceous birds. Amer. Midland 5:31-32. Naturalist 76:1-73. SHORE-BAILY,W. 1917. Hybrid breeding. Bird Notes JEWETT, S. C., W. P. TAYLOR, W. T. SHAW, AND J. W. 8:13-16. ALDRICH. 1953. Birds of Washington State. SIBLEY, C. G. 1957. The evolutionaryand taxonomic Univ. Washington Press, Seattle. significance of sexual dimorphism and hybridiza- JOHNSTON, R. F. 1964. The breeding birds of Kansas. tion in birds. Condor 59:166-191. Univ. Kansas Publ., Mus. Nat. Hist. 12(14): SIBLEY,C. G. 1960. The electrophoretic patterns of 575-655. avian -white proteins as taxonomic characters. LEOPOLD, A. S. 1959. Wildlife of Mexico. The game Ibis 102:215-284. birds and mammals. Univ. California Press, SIBLEY, C. G. 1961. Hybridization and isolating Berkeley. mechanisms. p. 69L-88. In W. Frank Blair [ed.] LIGON, J. S. 1961. New Mexico birds and where to Vertebrate speciation. Univ. Texas Press, Austin. find them. Univ. New MexicoPress, Albuquerque. SUTTON, G. M. 1963. Interbreeding in the wild of LOWE, C. H. 1964. The vertebrates of Arizona. the Bobwhite (Colinus virginianus) and Scaled Univ. Arizona Press, Tucson. Quail (Callipepla squamata) in Stonewall County, MCCABE, R. A. 1954. Hybridization between the northwestern Texas. Southwestern Naturalist 8: Bob-white and Scaled Quail. Auk 71:293-297. 108-111. MASSON,W. V., AND R. U. MACE. 1962. Upland SUTTON,G. M. 1967. Oklahoma birds. University of game birds. Bull. no. 5, Oregon State Game Oklahoma Press, Norman Comm., Portland. TEXAS GAME, FISH, AND OYSTER COMMISSION. 1949. OHMART, R. D. 1967. Comparative molt and ptery- Principal game birds and mammals of Texas. lography in the quail genera Callipepla and Lo- phortyx. Condor 69:535-548. Accepted for publication 19 September 1968.