ACKNOWLEDGMENTS Gagn(•,W.C. andL. W. Cuddihy.1990.Vegeta- tion. Pp. 45-114 in W. L. Wagner, D. R. The studywas funded by USGS/PIERC. For loan Herbst, and S. H. Sohmer. Manual of the of large box traps, advice on catchingbirds, or floweringplants of Hawaii.University of both, we thank T. Lum of Hawaii Division of Hawaii Pressand BishopMuseum Press, Forestryand Wildlife,J. Klavitterof the U.S. Fish Honolulu. and Wildlife Service, H. Hoshide and D. Hu of the Johnsgard, P. 1999. of the world. National Park Service, and G. Norman of the SmithsonianInstitution Press, Washing- VirginiaDepartment of Game and InlandFishedes. ton, D.C. C. Collins,L. Pratt,W. Sakai,and E. Spurrprovided Lewin,V. and G. Lewin.1984. The KalijPheas- helpfulreviews of thispaper. ant, a newlyestablished game birdon the Island of Hawaii. Wilson Bull. 96:634-646. LITERATURE CITED McClure,E. 1984. Birdbanding. Boxwood Press, Pacific Grove, CA. Backs,S.E., P.D. Major,C.H. Eisfelder,and J. S. Pratt,T. K. 2001. Socialmonogamy in the Kalij Thompson.1985. Techniques for trapping :skewed sex ratios,parental and restockingRuffed in Indiana. care, and cooperativebreeding. Poster at IndianaDepartment of NaturalResources, AmericanOrnithologists' Union meeting, Indianapolis. Seattle, WA. Bub,H. 1991. trapping and birdbanding. Schemnitz,S. D. 1994.Capturing and handling CornellUniversity Press, Ithaca, NY. wildanimals. Pp. 106-124 in T. A. Bookout,ed. Researchand management techniquesfor wildlifeand habitats. Fifth ed. The WildlifeSociety, Bethesda, MD. UsingWing Plumageto DetermineAge of Mountain

David J. Delehanty*and Nichole C. Turek Departmentof BiologicalSciences 8007 IdahoState University Pocatello, ID 83209 *Correspondingauthor Phone:(208) 282-4532 E-mail: deledavi @ isu.edu

ABSTRACT

We analyzed the occurrence of mottled versus quailshed their juvenal primary coverts during the solid-coloredupper greater primary coverts on the second(definitive) prebasic molt and replace them wingsof MountainQuail (Oreortyxpictus) to withsolid-colored adult primary coverts. All pdmary evaluatethe use of wing as an indicatorof covertsof quail >15 moage consistentlyare solid- MountainQuail age. MountainQuail retaintheir colored.Thus, colorpattern of primarycoverts juvenalpdmary coverts through the first prebasic providesa reliablemeans for aging Mountain moltand subsequently enter their first breeding Quail. Individualsthat possess one or more season with juvenal coverts still intact. Juvenal mottledcoverts are < approximately15 mo old, primarycoverts #1-7 are mottled,and juvenal whileindividuals with only solid-colored coverts are pdmarycoverts #8 (usually)and #9 are solid- > approximately15 mo old. Solid-coloredcoverts colored.At the end of their first breeding season, #8 and#9 are notinformative for aging quail. Page 116 NorthAmerican Bird Bander Vol 28 No 3 INTRODUCTION "The spotted juvenal primary-coverts,which accompanythe innereight juvenal primaries,are The last decade has seen substantialprogress in retaineda fullyear, and are lostat the secondfall usingsubtle plumage characteristics to determine moult." Sumner (1935), using Quail age in NorthAmerican (Pyle 1997). The (Callipeplacalifornica), was the first to report in ability to determinethe age of birds through detailthat the presenceof mottledprimary coverts plumagecharacteristics isespecially helpful to the can be used to age quail. This was followedby a studyof smallor declining populations (e.g., White prominentpaper by Leopold (1939) in which et al. 2002) because it adds a demographic mottled coverts among presumed yearling quail dimensionto data on reproductionand survivor- were observed for nine New Wodd ship. Mountain Quail (Oreortyx pictus) have includingMountain Quail. declinedsignificantly in distribution and abundance acrosstheir historicrange (Brennan1994). Field Leopold(1939) remainsthe principalreference for studies of MountainQuail breeding biologyand the use of mottled juvenal primary coverts as populationdynamics could be enhancedby the indicators of age among New Wodd quail. ability to determine age of individualswith However, for several reasons it presents confidence(Gutierrez and Delehanty1999). Very ambiguitieswhen applied to determiningage of young MountainQuail (0-2 mo) are recognized Mountain Quail. The age of the Mountain Quail easilyas juvenilesbased on theirsmall size and used to correlate age and covert color was not substantial mottling throughout their plumage. known independently,but rather was inferredby However, followingthe first prebasicmolt, aging comparingtheir covert color to an unstatednumber adult-sized Mountain Quail can be difficult. of museum specimens. Mountain Quail bearing juvenal-like primary coverts were assumed to be For more than a century, ornithologistshave less than approximatelyone year old, while those known that juvenile with solid-coloredpdmary covertswere assumed (Odontophoddae)retain certain wing to be more than ayearold (Leopold1939). Hence, throughthe firstprebasic molt, which occurs during the studywas circularand does notappear to have thefirst summerof life (age 2-5 mo),and holdthese been open to alternativeexplanations. Secondly, feathersuntil the second(definitive) prebasic molt the details of Mountain Quail covert color and the followingsummer (age 12-14 mo). Normally, retention were not provided and the general the two outermostprimary remiges (p9 and p10) patternfor juvenal primarycoverts, illustrated with are retained(Dwight 1900) and the presenceof a NorthemBobwhite ( virginianus) wing in relativelypointed, worn, or faded outer primaries which all nine coverts are buffy-tipped(Leopold has been used to judge the age of quail (Dwight 1939), does not reflect the covert pattem seen in 1900, Leopold1939, Larsonand Tabor 1980). New Mountain Quail. Among young Mountain Quail, World quail also retain juvenal upper primary some coverts are mottled with pale or buff spots covertsthrough the first prebasicmolt and carry and flecksat the tips,while other covertsare solid- these feathers throughthe first breeding season colored (Gutierrez and Delehanty 1999). Lastly, (Leopold1939). Using primarycovert retentionto among New World quail,the secondprebasic molt determinethe age of New Worldquail may be less follows a quail's first breeding season and the subjective than using outer primaries because pdmarycoverts molt sequentiallyfrom innermost adult primarycoverts are uniformlysolid-colored to outermost slightly ahead of the sequentially whereas juvenal primary coverts can be mottled moltingprimaries (Leopold 1939). Because some with pale flecks and spots. Mountain Quail exhibitsolid-colored innermost and outermostprimary coverts with mottledcoverts in The observation that New Wodd quail retain between, it would be helpfulto understandif this juvenal coverts often has been attributed pattem reflectsthe general replacementsequence incorrectlyto a study by Dwight(1900), although describedby Leopold(1939). this publicationdoes not address juvenal covert retention. However, van Rossem (1925: 420) We measured the frequency and location of reported a communicationfrom Dwight stating, mottled and solid-colored primary coverts of Jul - Sep 2003 NorthAmerican Bird Bander Page 117 Mountain Quail of known age. Our goal was to We comparedthe presenceor absenceof mottled cladfythe pattem of retentionof mottledpdmary primarycoverts to the age of quail and lookedfor covertsamong MountainQuail and to describethe consistentpatterns between plumage and age of use of pdmary covertsto determine the age of MountainQuail. In our analysis,we focusedon the Mountain Quail. abilityto detect age at life-stagesimportant to describingMountain Quail population dynamics. In METHODS particular,we lookedfor the abilityto detecthatch- year quail among autumn populationsof adult- We analyzedthe leftwing of 69 MountainQuail that sized .This would providea means for had beenfrozen for 1-9 yearsfollowing their deaths measuring autumn age ratios and annual andforwhich age was known independently of any reproductive success. We also looked for the plumage characteristics. Quail came from two abilityto discernbetween yeading breeders versus sources:a wild populationin the Mojave Desert of older individuals to better understand Mountain southem Califomia, and a captive research Quailbreeding biology. population.The captive populationconsisted of individualstaken from the Mojave population, RESULTS commerciallypurchased quail (R. Tybie, Doyle, CA), andthe progenyof wild and commercial quail. Mountain Quail exhibited a consistent and Bothgroups of quailwere the subjects of long-term interpretable pattern of solid-colored versus study(Delehanty 1997, Gutierrezand Delehanty mottledpdmary coverts (Table 1 ). Allcoverts on all 1999) unrelatedto plumagemorphology, and we quail>15 mo age (n = 22) were solid-coloredand usedquail that were collectedor salvageddudng matchedthe olive-brownplumage of the restof the the course of that research. upperwing. Similarly, all quail <10 mo age (n = 38) had 1-8 mottledcoverts. Typically, coverts #1 Likeother New Woddquail, Mountain Quail have through#7 weremottled on these young quail. In 10 pdmaryflight feathers.One upper pdmary all buttwo cases, covert #8 was solid-colored,and covertovedays the gap formed between each pair #9 was alwayssolid-colored. Quail 11-14 monold of adjacentprimaries. We numberedpdmary (n= 9), i.e.,yeading breeders in the latter stages of covertsstarting with the small,innermost pdmary their first breedingseason, were in transitional covert(#1) covedngthe gap betweenpl andp2 plumagethat accuratelyreflected the onsetof the through the outermost pdmary covert (#9) covering prebasicmolt that follows each breeding season. the gap betweenp9 and p10. For analysis,we One of the nine yeadingbreeders had not yet removedthe leftwing and recordedthe colorof all moltedits juvenal coverts.Six individualshad from ninecoverts. Covert color was recorded as being two to six new, solid-coloredcoverts appearing either solidolive-brown (color 28, Smithe 1975- sequentially in the #1 - #6 positions. Two 1981)or mottledwith buffy or cream(colors 124, individualsfrom June and July,respectively, and 54, respectively,Smithe 1975-1981) tips and fine approximately14-15 moage, had replacedall nine buffy or cream specklingon an olive-brown pdmary coverts. background. Table1. Percent frequency ofmottled primary coverts from the left wing of Mountain Quail. Quail are divided into ages thatreflect important lifestages: adult-sized quail that have not reached their first breeding season (ages 2-10 months), yearlingbreeders (ages 11-14 months), and individuals that have completed at least one prebasic molt following a breedingseason (ages > 15months). Every quail ages 2-10 months had at least one mottled covert (mean =6.3 + 1.45 SD, min.= 1,max, = 8).Yearling breeders had from 0-6 mottled coverts (mean = 2.4+ 2.13 SD). Coverts that were not mottled were solid colored.

Primary Coverts Age N #1 #2 #3 #4 #5 #6 #7 #8 #9 2-10 mos. 38 84.2 89.5 94.7 94.7 92.1 89.5 81.6 5.3 O 11-14mos. 9 11.1 11.1 33.3 33.3 55.5 44.4 55.5 0 O >_15rnos. 22 0 0 0 0 0 0 0 0 0

Page118 NorthAmerican Bird Bander Vol28 No 3 DISCUSSION appearanceeven thoughthey representthe first generationof coverts.Coverts #8 and #9 are adult- In our study, the presenceof even one mottled likejuvenal coverts that are retainedalong with the pdmarycovert indicated that the MountainQuail in mottledjuvenal coverts #1 -#7. A similarpattem of questionwas less than approximately15 mo of retained, mottledjuvenal coverts#1 -#7 and solid- age. Thissuggests that researcherscan identify colored coverts #8-#9 in yeading birds has been hatch-yearindividuals reliably among adult-sized described for Scaled Quail ( squamata; quailcaptured dudng autumn or winter.Similarly, Wallmo 1956, Smith and Cain 1984). any breedingbird with one ormore mottled pdmary covertis a yeadingbreeder. Pdmarycoverts #5 - Our findingsconfirm the astute general observa- #7 mightbe usefulespecially in detectingyeading tion made by Dwightmore than a centuryago (van breeders, as these are the last mottled coverts to Rossem1925) that all pdmarycoverts are retained be replaced. The exclusive presence of solid- duringthe firstprebasic molt. We modifythe details colored coverts indicated that the Mountain Quail of using pdmary coverts to age New World quail was 15 mo old or older. These quail had (Leopold 1939) specificallyfor use on Mountain experiencedat leastone breedingseason and had Quail. We emphasize that solid-coloredprimary undergoneat leastone prebasicmolt at the end of coverts #8 and #9 are not informativein revealing the breedingseason. the age of Mountain Quail. Also, because the primary covert molt sequence progressesfrom The patternsof mottledand solid-coloredpdmary innermost to outermost, it is possible to observe covertsthat we observedare explainedreadily by yearling breeders late in the breeding season with Mountain Quail molt sequence. Mountain Quail confusing transitional plumage. As yeading retaintheir juvenal primary coverts through the first breeders initiatetheir post-breedingprebasic molt, prebasic molt, which occurs during their first they will exhibit with solid-colored innermost summer of life, and enter their first breeding primarycoverts (due to the onset of the prebasic season with juvenal coverts still intact. Although molt), mottled middle pdmary coverts (due to several species of New Wodd quail undergoavery retention of juvenal coverts), and solid-colored limited prealtemate molt during eady spring outermost primary coverts (due to retention of involvingthe replacementof some head and neck adult-like juvenal coverts). Presence of mottled feathers(Dwight 1900), a prealtematemolt has not pdmarycoverts on a nestingadult reliablyindicates been documentedfor MountainQuail. If present,a a yeading breeder. If all the prirdarycoverts of a prealtemate molt does not appear to include breedingquail are solid-coloredand are also newly primary coverts. Mottled juvenal coverts are not molted, then the quail is >approximately 15 mo replaced with solid-coloredadult coverts until the age. When all pdmary coverts on a nesting adult second prebasic molt. are solid-colored, faded, and wom, this should reliably indicate that the quail is in its second or Because juvenal pdmary coverts #8 and #9 are more breeding season. solid-colored like adult pdmary coverts, the presence of solid-coloredcoverts #8 and #9 is not ACKNOWLEDGMENTS informativefor indicatingMountain Quail age. The presence of adult-like,solid-colored coverts #8 and Specimens were obtained from quail trappingand #9 in juvenilequail probablyresults from the highly bandingactivities carried out underthe auspicesof compressedmolt sequence typical of New Wodd the CaliforniaDepartment of Fish and Game, the quail. In New World quail,juvenal plumageis held Division of Wildlife (NDOW), and the very briefly. The molt from juvenal plumage into UnitedStates Navy (USN) at the China Lake Naval adult plumagecan beginbefore the moltfrom natal Air Weapons Station, and also from research down into juvenal plumage is completed (Dwight flocks maintained by the University of Nevada, 1900). Pdmary coverts #8 and #9 are among the Reno (UNR) and Idaho State University.The UNR last juvenal feathers to erupt (DJD, pers. obs.). Biological Resources Research Center, NDOW, They erupt in concertwith start of the firstprebasic USN, Quail Unlimited(QU), the Northem Nevada molt into adult plumage and are adult-like in Chapter of QU, Chukars Unlimited, and the

Jul. - Sep. 2003 NorthAmerican Bird Bander Page 119 National Fish and Wildlife Foundation provided Larson,J. S. and R. D. Tabor. 1980. Pp. 143-202 in financial and logistical support. We thank T. Wildlifemanagement techniques manual, Campbell,S. Eaton,S. Stiver,and C. R. Tracyfor 4th edition(S. D. Schemnitz,editor). The long-term logisticalsupport and S. Baab, C. WildlifeSociety, , D.C. Elphick, P. Pyle, J. M Reed, and an anonymous Leopold,A. S. 1939. Age determinationin quail.J. reviewerfor helpfulmanuscript reviews. We thank Wildl.Manage. 3:261-265. the manyvolunteers and students who participated Pyle, P. 1997. Identificationguide to North in quail trapping and flock maintenance across American birds. Slate Creek Press, many years. Bolinas, CA. Smith, D. S. and J. R. Cain. 1984. Criteria for LITERATURE CITED age classificationof juvenile Scaled Quail J. Wildl. Manage. 48:187-191. Brennan, L. A. 1994. Broad-scalepopulation Smithe,F. B. 1975-1981. Naturalist'scolor guide. declinesin fourspeciesof NorthAmerican Three Parts. Am. Mus. Nat. Hist., NY. quail:an examinationof possiblecauses. Sumner, E. L., Jr. 1935. A life historystudy of the Pp. 160-169 inSustainableecological California Quail, with recommendations systems:implementing an ecological for conservationand management.Cal. approachto landmanagement. Gen. Tech. Fish Game 21:167-256; 275-342. Rep. RM-247. USDA ForestService, van Rossem,A. J. 1925. Flightfeathers as age Rocky Mountain Forest and Range indicatorsin Dendrapagus. Ibis 67:417- ExperimentStation, Fort Collins,CO. 422. Delehanty, D. J. 1997. MountainQuail reproduc- Wallmo. O. C. 1956. Determination of sex and tion and restoration in the Groat Basin. age of ScaledQuail. J. Wildl.Manage. Ph.D. Dissertation, Univ. Nevada, Reno. 20:154-158. Dwight, J., Jr. 1900. The molt of the North White, G. C., A. B. Franklin, and T. M. Shenk. AmericanTetraonidae (, , 2002. Estimatingparameter of PVA and grouse).Auk 17:34-51; 143-166. modelsfrom data on markedanimals. Pp. Gutierrez,R. J. and D. J. Delehanty.1999. 169-190 in PopulationViability Analysis MountainQuail (Oreortyxpictus). In The (S. R. Beissingerand D. R. McCullough, birdsof NorthAmedca. No 457 (A. Poole eds). Universityof ChicagoPress, Chi- and F. Gill, eds). The Birdsof North cago, IL. America,Inc., Philadelphia,PA. News, Notes, Comments Erratum: Vol.28 No. 2, Inside Front Cover Erratum: Vol.27 No. 4, pages 127-129 Misidentification of Species Recommended Band Size for As many NABBreaders pointed out (a listtoo long Spotted Towhees: A Suggested to printhero), the insidecover of NABB28(2) in- correctlylabeled the cover illustrationas a Black- Revision by Rita R. Colwell and-white Warbler. The caption should have indicated that the bird illustrated was a Black- Seventeen females aro shown in the Tables on bumianWarbler. The ProductionManager apolo- p. 128, but, in error, only 16 were used for gizes to the artist and readers, and thanks Dr. calculations.The missingindividual was the bird GeorgeWest for providinga graphicof a Black- whose wing length was 81 mm and weight was and-white Warbler for the cover of this issue. 33.7 g. Correctedcalculations, including all 17 females,aro: wing lengthrange = 75-81 mm, mean Readersmay be interestedto knowthat Dr. West's = 77.5 mm; weight range = 33.7-41.4 g, mean = drawingsand sketches are availablefor purchase 37.75 g. The author thanks Walter Sakai for at: birch @ birchsidestudios.com pointingout this oversight. Robert Pantie Prod. Mgr. Page 120 North American Bird Bander Vol 28 No 3