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Papers in Ornithology Papers in the Biological Sciences

1971

Experimental Hybridization of the New World Quail (Odontophorinae)

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

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Johnsgard, Paul A., "Experimental Hybridization of the New World Quail (Odontophorinae)" (1971). Papers in Ornithology. 59. https://digitalcommons.unl.edu/biosciornithology/59

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 the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

EXPERIMENTAL HYBRIDIZATION OF THE NEW WORLD QUAIL (ODONTOPHORINAE) •

PAULA. J0tINSGARD

ALTHOUGHthe surprisingly high capacity for hybridizationamong species of the odontophorinegenera Colinus, Callipepla, and Lophortyxhas been recognizedfor sometime (Gray, 1958), no systematicattempt has been made to produceand study such hybrid combinations.Those that have occurredhave resultedunder natural conditionsor have been fortuitously producedby game breederswho were interestedneither in the hybrids themselvesnor in determiningtheir relative fertility. As a result, the only positiveinformation so far availableon the fertility of hybridsamong this groupis that of Shore-Baily(1913), who obtaineda broodof Fs hybrids from an originalcross between a male ScaledQuail (Callipepla squamata) and a female California Quail (Lophortyx californica). These second- generationbirds died before their fertility could be determined. Hubbard (1966) reporteda possibleback-cross hybrid involvingScaled Quail back- crossedto Gambel'sQuail (Lophortyxgambelii) taken in New Mexico. To my knowledge,no other case of successfulhybridization beyond the first generation has been reported, although hybrid females of various inter- genericcombinations have been known to produceeggs (Johnsgard, 1970). To gain more satisfactoryinformation on the possiblefertility of such hybrids,and to study behavioraland morphologicalfeatures of thesebirds, an effort has been made to breed interspecifichybrid combinationsinvolv- ing all of the North Americanspecies of thesethree genera. These include the ScaledQuail, Gambel'sQuail, California Quail, Elegant or Douglas Quail (Lophortyxdouglasii), and Bobwhite(Colinus virginianus). Also as the hybrid combinationhas twice been reported to occur in nature, the Mountain Quail (Oreortyx picta) was crossedwith the California Quail. All birdswere kept indoorsin roomsprovided with 17-hourphotoperiods and temperaturesranging from about 70 to 80.øF. Two typesof cageswere usedwith equal success,including standard wood and wire-bottomquail breedingpens measuring24 X 24 X 72 inches,and entirely open welded wire cagesapproximately 18 X 14 X 48 inches. Nest boxeswere also pro- vided, but the birds rarely usedthem. Eggswere collecteddaily, placed temporarily under cool storagein plastic bags, and set in an incubator at weekly intervals. The incubatorwas a forced-airmodel of commercialsize with automatic turning, set for dry bulb and wet bulb readingsof 99.3-

x Contribution No. 421 from the Department of Zoology, University of Nebraska, Lincoln, Nebraska 68508.

264 The Auk, 88: 264-275. April 1971 Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

April 1971] New World Quail Hybrids 265

100øFand 83-88øF respectively.This setting,which is that recommended for domestic fowl eggs, gave good hatching successwith an incubation periodof slightlyover 23 daysfor all quail speciesand hybrid combinations studied. Eggswere initially candledafter 7 days,when infertile eggsand dead embryos ("early embryonic death" in Table 1) were removed. Subsequentprehatching mortality was recordedas late embryonicdeath, includingfailure to completepipping. Establishmentof hybrid pairingswas usually achievedwith little diffi- culty. Wheneverpossible birds representingthe desiredcrosses were placed togetherbefore they had reachedsexual maturity, and in someinstances as soonas they could be accuratelysexed. Fully adult birds were also used, and with few exceptionsthey soontolerated the new partner. Femalesthat were already producingeggs sometimes did not interrupt their egg-laying schedulewhen placedwith a new male, althoughnone of the eggsdropped shortlyafter new pairingswere made proved fertile. Seriousinjury or the death of a newly introducedfemale occurredin only a few instances,and couldusually be avoidedby early separationof noncompatiblebirds. Pair formationin all the speciesappears to consistof ritualized aggressivepos- tures and inhibited attacks on the female, while the latter assumessubmis- sive posturessuch as crouching.Differences in the vocalizations,postures, and plumage characteristicsof the various speciesthat may serve as iso- lating mechanismsunder natural conditionsare subjectsof future studies and are not considered here.

RESULTS Egg production,fertility, and viability.--A summaryof incubationand rearingresults appears in Table 1. With few exceptionsthe numberof eggs incubatedis lessthan the total laid, as someeggs were cracked,used for albumenanalysis, or not incubatedfor other reasons.As the table shows, successwas achieved in rearing F• hybrids between the Bobwhite and Gambel'sQuail, the Bobwhite and ScaledQuail, the Scaledand Gambel's Quail, the California and Scaled Quail, and between the California and Mountain Quail. Except for the first-namedcross, all thesecombinations havepreviously been reported from the wild or captivity (Johnsgard,1970). The fairly substantiallosses of youngof both hybridsand parental types stemmedprimarily from two sources,cannibalism and accidentalchilling. Even weeklybill-trimming failed to controlcompletely losses from pecking, which causedthe most severelosses. In no casewas the death of any F• hybrid clearly attributed to a weaknessat hatching. The weakestchicks at hatchingwere thoseof the DouglasQuail, of which only 4 out of 32 suc- cessfullycompleted pipping and nonelived beyonda week. Likewiseboth the Scaledand Gambel'sQuail young showeda surprisinglyhigh incidence Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

266 PAUL A. JO•SGARD [Auk, Vol. 88

TABLE 1 RESULTSOF INCUBATION Ot• HYBRIDQUAIL EGGS AS COMPARED WlTlt PARENTALSPECIES

Total Early Late eggs embry- embry- incu- Infer- onic onic bated tile death death Hatched Reared

Parental Species Bobwhite (B) 28 14 3 0 11 7 Gambel's (G) 62 4 10 20 28 6 California (C) 76 35 39 2 0 0 Douglas (D) 35 1 2 28 4 0 Scaled (S) 83 16 16 26 1 6 Total Parentals 290 70(24.1%) 70(24.1%) 76(26.2%) 74(25.5%) 19

Hybrid Pairings (male X female) BXS 110 43 16 11 40 12 SXB 86 52 34 0 0 0 CXS 47 21 4 3 19 14 BXG 17 0 0 4 13 2 SXG 16 5 0 4 7 1 GXC 13 11 2 0 0 0 GXS 9 6 2 0 1 0 SXD 5 3 0 1 1 0 Mountain X C 34 27 0 4 3 1 Total F• Hybrids 337 168(49.8%) 58(17.2%) 27(8.0%) 84(24.9%) 30

Other Hybrid Pairings F•BG X F•BG 16 6 10 0 0 F•BG X G 28 0 28 0 0 F•GS XF, GS 33 26 7 0 0 S X F• GS 32 11 11 2 8 F• CG X C 32 19 13 0' 0 Fa BC X F_oBC 74 52 22 0 0 F.oBC X B 25 15 10 0 0 F•CSXF•CS 213 211 2 0 0 F• BS X F, BS 228 228 0 0 0 BW X F• BS 44 28 6 10 0 S X F• BS 15 15 0 0 0 Total Other Hybrids 740 611(82.6%) 109(14.7%) 12(1.6%) 8(1.1%) of late embryonicmortality. The low fertility and highincidence of early embryonicdeath in California Quail apparentlywas the result of a subfer- tile male in the singleintraspecies pair that was establishedfor California Quail. The "otherhybrid pairings" included in Table 1 consistmainly of attemptedF2 productionor of backcrosspairings. Records are alsoincluded Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

April 1971] New World Quail Hybrids 267 from pairingsof four F2 Bobwhite x California Quail hybrids that were obtainedfrom the San JoaquinGame Farm, Reedley,California. Accord- ing to the manager,R. Davis (in litt.), thesefour hybrids (two of eachsex) were the only onesthat hatchedfrom a very largenumber of eggslaid by a pair of F• birds reared at that farm. I was unable to obtain the F• birds, but did purchasethe F,,s. Both femaleslaid considerablenumbers of eggs (one produced21 eggsin 21 days, the other 103 eggsin 137 days), all of whichwere abnormally small (averageof 10 is 27.1 x 20.0 ram). Of the 74 eggsset, none exhibitedembryonic development beyond the very earliest stage. The only successfulsecond-generation hybrid production thus far achievedin our laboratoryhas been the hatchingof eight backcrosshybrids (none survivingbeyond 6 days) that resulted from pairing a male Scaled Quail to a female F• Gambel'sx ScaledQuail. Earlier attempts to obtain F: offspringfrom the F• generationwere unsuccessful.These F• birds had beenproduced by William S. Huey, who informedme (in litt.) that he too had beenunable to obtain F: birds,but did hatch a numberof chickswhen a ScaledQuail male was in the pen alongwith two F• femalesand a single F• male. Theseresults suggest that at least a limited degreeof hybrid fer- tility existsin Scaledx Gambel'sQuail crosses,for which there was only suggestiveevidence earlier (Hubbard, 1966). The F• Gambel'sx Scaled Quailfemale in our laboratoryhas laid typical-sizedeggs at normalintervals (31 eggsin 63 days,compared with three Gambel'sQuail that averaged41 eggsin 97 daysand three ScaledQuail that averaged32 eggsin 67 days). The possiblefertility of the Scaledx California Quail cross,as reported by Shore-Baily(1913), has not yet been confirmed.Three of the seven femalesreared have laid, but the eggshave shownno embryonicdevelop- ment beyond3 or 4 days or have beeninfertile. The eggsof the first breed- ing cyclewere slightly smallerthan normal (averageof 10 is 28.6 X 22.5 mm), while later eggslaid by the samefemales were of virtually normal size, averaging 32.6 x 24.7 min. The Bobwhitex Gambel'sQuail hybrid appearsdefinitely to be sterile. All the eggslaid by the only femalereared of this crosswere distinctly smallerthan normal(average of 10 is 24.7 x 19.5 mm). None of the eggs representingpotential F: offspringdeveloped beyond the earliestembryonic stages,and whenthe F• malewas later pairedwith a femaleGambel's Quail all of the resultingfertile eggsalso suffered early embryonicdeath. The Bobwhitex ScaledQuail hybridsthat have beenreared are appar- ently sterile,at leastinter se. All six femaleshave laid eggsbut mostof theseare distinctlysmaller than normal (average of 10 laid by threefemales is 21.5 x 16.6 mm), and only onefemale has produced eggs of nearlynormal size. Further, all the eggsthese females produced have provedeither infer- tile or have in any caseshown no detectableembryonic development. It is Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

268 P^uL A. Jo•t•sc^RD [Auk, Vol. 88 of interest that in this cross•as well as the Scaled X California cross,no obvious excessof males has been noted (collectively 14 males and 13 femaleshave been reared to maturity). Excessivemales are frequently encounteredin many intergenericgalliform hybrids and apparently are a reflectionof a sexchromosome imbalance, causing differential sexmortality or sexreversal (Sandnes and Landauer,1938). It has not beenpossible to determinethe fertility of the singlemale F• Mountain • California Quail that has beenreared, as the female California Quail it waspaired to has producedno eggs. The results of these studies to date are similar to those of Morejohn (1968), who foundthat in crossesbetween two congenericspecies of jungle- fowl (Gallus) the F• offspringhad high hatchabilityand werevigorous, but laid smaller-than-normaleggs and exhibitedhigh embryonicmortality, with no F2 birds survivingto maturity. He did, however,succeed in hatching and rearinga few backcrossoffspring. Inheritanceof plumage'characteristics.--Considering the diversearray of crestshapes, head patterning, and body feathercoloration in the New World quail, the plumagesof the hybridsare. of specialinterest. Earlier published illustrationsof adult hybridsamong this groupinclude photographs of both sexesof BobwhiteX ScaledQuail skins(McCabe, 1954), the skin of a male (probably a backcross)Scaled X Gambel'sQuail (Hubbard, 1966), and a mountedScaled X DouglasQuail (Banks and Walker, 1964). A photo- graph of the Bobwhite X Gambel'scross produced in our laboratory has appeared(Johnsgard, 1970), and monochromeillustrations of Mountain X California Quail hybrid maleshave been published(Peck, 1911; Peterle, 1951). A colorpainting of this samecross, in additionto male California X Gambel'sQuail and Gambel's• ScaledQuail, has also appeared(Hachi- suka, 1928). Finally the ScaledX California hybrids Shore-Bally (1916) rearedhave beenillustrated and a photographof a captive male Scaledx Gambel's Quail appeared on the cover of the Arizona Game and Fish Department's "Wildlife views" for November-December 1966. Before consideringadult plumages,the natal plumagesare worthy of attention. O'f the parentalspecies, the Bobwhitenatal pattern is easilythe most distinctive,with its broad, dark chestnutdorsal stripe that lacks a definite medial line and is indistinctly separatedfrom the mottled wood- brown and buff upperpartsby narrow cinnamonlines. The ScaledQuail and the two U.S. speciesof Lophortyx (as well as L. douglasii) are all extremelysimilar, and it is sometimesdifficult to identify downy youngof thesespecies. All have smallcrests and a conspicuouslight mid-dorsalline borderedon both sideswith fuscous,the latter in turn borderedwith wider buffy or cinnamon-bufflines that extend from the sidesof the tail to the neck. Mountain Quail downyyoung somewhat resemble Bobwhites in that Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

April 1971] New World Quail Hybrids 269

Figure 1. Diagram of adult male plumage patterns of six speciesof quail and eight intergenerichybrids, with lines connectinghybrid combinationsand parental species. Some variation occurred among the Scaled X California males which is not indicated here (see text). they usuallylack crestsat hatching(but acquirethem within a week), have a dark postocularstripe extendingdown the side of the neck, and have a distinctlychestnut rather than fuscousdorsal area that darkensto blackish laterally. The blackis in turn borderedby narrowbuffy-white lines, below whichare a secondpair of blackishlines, reminiscent of Lophortyx. In all casesthe hybrid downyyoung have been exactly intermediate between the parentals. Differencesin the adult male plumagepatterns of the hybridsare more difficult to presentwithout colorplates. They are largely limited to the head, lower breast,and flanks, all of which are utilized in sexualdisplay. Differencesin the dorsalsurface, wings, tail, and tail covertsare either relatively negligibleor are understandablein relation to differencesin the other areas mentioned. The head plumage conditionsare summarizedin Figure 1, which illustratesthe adult male headsof the six speciesstudied, all the F• hybrid combinationsobtained in this study, and two additional crosses(California x Douglas Quail and Scaled x Douglas Quail) that Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

270 PAUL A. JOHNSGARD [Auk, Vol. 88 have been bred previously. The drawing illustrating the first of these crosseswas basedon a specimen(No. 458926) in the AmericanMuseum of Natural History, while the ScaledX DouglasQuail drawingis basedon the photo publishedby Banks and Walker (1964). Not shownis the Cali- fornia x Gambel'sQuail cross,which not only has already been illustrated (Hachisuka, 1928) but also is of little interestbecause of the great simi- larity of the parental species. Although the drawingsin Figure 1 representthe breedingplumage, it is of interestthat, in someof the hybrid crossesreared, a distinctivesubadult plumageimmediately followedthe juvenal plumage. Thus the male Bob- white x Gambel's Quail temporarily exhibited an entirely white throat after molting its buffy juvenal plumage,but within a few weeksbegan replacingthese feathers with black in the chin region,a conditionthat has persistedin succeedingmolts. Similarly the male Mountain X California Quail initially had an entirely black throat immediately after its postjuve- nal molt, but this was later permanently altered to a chestnutthroat with a narrowblack border. These two instances,and one mentionedbelow, sup- port the view originally proposedby Dwight (1900) but questionedby Raitt (1961), namely that a limited prenuptialmolt occursin the chin and head regionin Colinus,Lophortyx, and probablyalso Oreortyx. In general the ultimate color and head pattern of the hybrids is intermediate between the parentalspecies, with a few interestingexceptions. In both the Scaledx Gambel'sQuail and the California x ScaledQuail malesthe chin and upper throat are predominantlychestnut, being somewhatbrighter in the former crossand virtually the samecolor as the crown of male Gambel'sQuail. In two of the six California X ScaledQuail malesthe throat becamealmost entirely black when the birds were between 6 and 7 months old. The crest and anterior portion of the crown of the Scaledx Gambel'sQuail male are chestnut,whereas in the California x ScaledQuail hybrid these areas are more fuscous. In both hybrid combinationsthe abdomenof the adult male is chestnuttoo, brighterthan in maleCalifornia Quail and similar to that of the castanogastrisrace of ScaledQuail. These color conditionshave pre- viouslybeen illustratedin Hachisuka's(1928) plate, althoughthe crest shapein that plate is not typical of our specimensor of the specimen Hubbard (1966) illustrated. Crest shapesand lengthsof the hybridsare also of interest,particularly amongthose involving distinctly crested and relativelycrestless species. For examplein the BobwhiteX ScaledQuail malesa crestcondition occurs that is strongly reminiscentof that found in the Crested Bobwhite (Colinus cristatus),which is reinforcedby a facial pattern muchlike thoseof such Central American races as C. c. leucopogon.Ohmart (1967) has already Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

April 1971] New World Quail Hybrids 271

BOBWHITE

RustBrown toYeUowish /!:..//•Oark Gray ß •ß OhvetoGrayish Bro.,,,n SCALED QUAIL

Figure2. Upper abdomenand flank feathersfrom malesof five speciesof quail and six intergenerichybrid combinations.Lines connect hybrid combinationswith parental types. pointedout that evenin the relativelybushy-crested Scaled Quail a pterylo- graphiccrest pattern of 10 feathersmay be detected,which assumes a con- spicuousform in the BobwhiteX ScaledQuail cross. In spiteof the strikinglydifferent body and flank patternsin malesof the parental species,examination of individualfeathers from the upper abdomenand flanksand thoseof the availablehybrid combinations(Figure 2) revealsthe relatively simplevariations in pigmentationconfigurations that are the basesfor these differences. Thus the distinctive "scaling" of the ScaledQuail breast and abdomenis achievedby the shiftingof the sub- terminal black band to the feather tip, whereasin Gambel'sQuail males this melanisticdeposition spreads over mostof the feather,as is also the casein the Mountain Quail. The distinctiveflank streakingof the Scaled, Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

272 Paul A. Jore•sG^aD [Auk, Vol. 88

Figure 3. Acrylamide gel separations of albumen from (top to bottom) Douglas, California, Gambel's, Scaled, Bobwhite, and Crested Quails. Anodal movement is from left to right, with major visible fractions present representingovoconalbumin, ovomu- coid, and a triple ovalbumin. An unknown fraction between cm 3 and 4 is also present.

Gambel's, and California Quails are producedby lighter shaft-streaks, whereasthe spottedflanks of the DouglasQuail (not illustrated) are the result of elongatedvane spots (Banks and Walker, 1964), and the barred flanks of the Mountain Quail are achieved by feather widening and an exaggerationof the transversebanding that may be detectedin the Bob- white. Interestingly,the male hybrids involvingthe Bobwhitewith the Gambel'sand California Quail have flanks that achievethe spottedcondi- tion of the DouglasQuail and alsoare nearly identicalwith the character- istic spottedflanks of the CrestedQuail. Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

April 1971] New World Quail Hybrids 273

Egg-whitepro•eins.--The pioneering survey by Sibley (1960) on the egg-whiteproteins of birds includedelectrophoretic patterns representing 42 speciesof Galliformes,including all of the parentalspecies considered in the presentpaper, as well as a Gambel'sx ScaledQuail F• hybrid sample. His resultsof electrophoresison a paper substrateindicated that the New World quail speciesstudied exhibit a dense,usually triple, ovalbuminpeak that undergoesconsiderable anodal movementat pH 8.6, a secondmajor ovoconalbuminfraction that travels only a short distance,and a third anodalcomponent between these that presumablycorresponds to the ovo- mucoidcomponent of domesticfowl (Gallusgallus) a.s illustrated by Sibley and Johnsgard (1959). The cathodally-migratinglysozyme fraction is readily visible in Sibley's paper separations,but the globulinsare not distinguishableon the profiles he published. With the possibleexception of the Mountain Quail, all the publishedprofiles were extremelysimilar and Sibleyattempted no detailedcomparisons among them. Throughthe cooperationof C. Michael Cowan,disc electrophoretic sep- arationsusing a polyacrylamidegel mediumhave so far been made for all the six speciesconsidered in this paper, as well as the Harlequin Quail (Cyrtonyx mearnsi), a Colombianrace of CrestedBobwhite (C. cristatus decoratus)and five intergenerichybrid combinations.The procedureused was that describedby Clarke (1964), exceptthat minor buffer modifica- tionsproduced an electrodebuffer of pH 8.3 and a gel buffer of 8.9 (Cowan, 1968). Thus the resultingseparations should be closelycomparable to those obtainedby Sibley at a pH of 8.6 and a paper substrate,and this appears to be the case. All samplesdone so far additionallyshow a distinctivebut minor fraction that migrates beyond the ovalbumin complex. This latter componentis triple in all speciesand their hybrids. The CrestedBobwhite appearsto be separablefrom the Bobwhiteon the basisof a reducedanodal ovomucoidmovement, and is closerto the other non-Colinusspecies in this respectthan is the Bobwhite. Hybrids betweenthe Bobwhite and the Gambel's,Scaled, and California Quails have ovomucoidcomponents with intermediatemigration properties, but their differencesare slight. All of the separationsare obviouslyextremely similar and suggestvery close evolutionaryrelationships (Figure 3).

ACKNOWLEDGMENTS

This study was financed by a National ScienceFoundation researchgrant (GB- 7666X) and the University of Nebraska. Assistancein data-gathering and care of birds was provided by Daniel Hatch, Alice Prososki, and Raymond Goldstein. Electro- phoretic separationswere obligingly performed by Michael Cowan of Nebraska Wes- leyan University. My sincere appreciation is extended to all these individuals and institutions. Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

274 PAV• A. JOttNSGARD [Auk, Vol. 88

SUMMARY Hybridizationexperiments among six speciesof New World quail cur- rentlyplaced in fourgenera (Colinus, Callipepla, Lophortyx, and Oreortyx) involvedthe establishmentof nine mixed-paircombinations. Seven of these combinationsresulted in the productionof F• offspring,and individuals representingfive intergenericcrosses have been reared to maturity. These F• individuals exhibited a hatchability at least as high as occurredin parental speciesmatings. In four of five intergenericcrosses, the hybrid femaleshave laid smaller-than-normaleggs in spite of averageor higher- than-averageegg production,and a high proportionof eggsfemales laid have been infertile or sufferedearly embryonicdeath. Limited hybrid fertility has been establishedfor one ColinusX Lophortyx and two Calli- pepla x Lophortyx combinations.Downy and adult plumagepatterns of hybrids are generallyintermediate and provide clues as to the probable evolutionof plumagediversity in the group. Egg-whiteproteins of eight odontophorinespecies and five hybrid combinationswere analyzedelectro- phoreticallyand exhibitedgreat similarity, further suggestingclose evolu- tionary relationships.

LITERATURE CITED

BANKS,R. C., ANDL. W. W•KER. 1964. A hybrid ScaledX DouglasQuail. Wilson Bull., 76: 378-380. CrARK, J.T. 1964. Simplified disc (polyacrylamide gel) electrophoresis.Ann. Ne•v York Acad. Sci., 121: 428. COWAN,C.M. 1968. The blood proteins of the bull shark Carcharhinus leucas. Un- publishedPh.D. dissertation,Lincoln, Univ. Nebraska. DwmtaT, J., JR. 1900. The moult of the North America Tetraonidae (quails, par- tridges and grouse). Auk, 42: 34-51; 143-166. GRAY,A.P. 1958. Bird hybrids. Farnham Royal, Commonwealth Bureau. HACtaISUKA,M. 1928. Variations among birds (chiefly game birds). Suppl., Publ. Ornithol. Soc. Japan, No. 12. HUBRARV,J.P. 1966. A possibleback-cross hybrid involving Scaled and Gambel's Quail. Auk, 83: 136-137. JOaNSCARD,P.A. 1970. A summary of intergenericNew World quail hybrids, and a new intergenerichybrid combination. Condor, 72: 85-88. McCABE, R.A. 1954. Hybridization between the Bob-white and Scaled Quail. Auk, 71: 293-297. MOREJOtaN,G.V. 1968. Breakdown of isolation mechanismsin two speciesof cap- tive junglefowl (Gallus gallus and Gallus sonneratii). Evolution, 22: 576-582. OtaMART, R. D. 1967. Comparative molt and pterylography in the quail genera Callipepla and Lophortyx. Condor, 13: 149-151. PECK, M.E. 1911. A hybrid quail. Condor, 13: 149-151. PETERiE, T.J. 1951. Intergeneric galliform hybrids: A review. Wilson Bull., 63: 219-224. RAITT,R. J., JR. 1961. Plumagedevelopment and molts of California Quail. Condor, 63: 294-303. Johnsgard in the Auk (1971) 88. Copyright 1971, University of California. Used by permission. http://elibrary.unm.edu/sora

April 1971] New World Quail Hybrids 275

SANDNES,G. C., AND W. LANDAIYER.1938. The sex ratio in the cross of Phaslanus torquatus • X Gallus domesticus •. Amer. Naturalist, 72: 180-183. S•OR•-BA•¾, W. 1913. Fertility of hybrid quail. Bird Notes, 4: 326. S•OR•-BA•¾, W. 1916. Hybrid Squamata X Californian Quail. Bird Notes, 7: 149. SmLE¾,C.G. 1960. The electrophoreticpatterns of arian egg-whiteproteins as taxo- nomic characters. Ibis, 102: 215-284. SmLE¾,C. G., Am) P. A. JO•NSOA•D. 1959. An electrophoreticstudy of egg-white proteins in twenty-three breeds of the domestic fowl. Amer. Naturalist, 93: 107- 115.

Department of Zoology, University of Nebraska, Lincoln, Nebraska 68508. Accepted 25 February 1970.