Distribution and Color Variation of Gyrfalcons in Russia
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j. RaptorRes. 26(2):81-88 ¸ 1992 The Raptor ResearchFoundation, Inc. DISTRIBUTION AND COLOR VARIATION OF GYRFALCONS IN RUSSIA DAVID H. ELLIS, CATHERINE H. ELLIS, GREY W. PENDLETON U.S. Fish and Wildlife Service,Patuxent Wildlife ResearchCenter, Laurel, MD 20708 ANDREI V. PANTELEYEV ZoologicalInstitute, Academy of Sciences,19903d St. Petersburg,Russia IRENA V. REBROVA ZoologicalMuseum, Moscow Lomonosov State University,103009 Moscow K-9, Russia YURI M. MARKIN CentralOrnithological Station, Oka StateReserve, 391072 Po Lakash,Spassk Region, Ryazan District, Russia ABSTRACT.--Gyrfalcon(Falco rusticolus) museum specimens in Moscow (73) and St. Petersburg(132) were dividedinto four colorclasses (gray, light gray, white gray, and white) and four longitudinalbelts representingmajor physiographicregions of northern Russia. Gray variants predominatedin the west and central regions.White birds were most commonin extremeeastern Siberia, but were occasionally found even west of the Ural Mountains. Frequencieswere as follows: European Russia 4% white, 50% gray (the remainderwere intermediates);western Siberia 0% white, 58% gray; central Siberia 15% white, 42% gray; and easternSiberia 47% white, 33% gray. Remarkably, in the easternmostsubregion, white birds predominatedeven near the southernmostextension. Becausethe northernmostportions of the species'range in continentalRussia are in central Siberia where white variants were rare, we proposethat a better predictorof the white variant is longitude,not latitude. White birds were most frequent at the easternreaches of both the Palearcticand Nearctic. The bestenvironmental correlates of this distribution pattern may be the southwardbending thermal isoclines proceedingeastward toward Greenlandor Kamchatka,where bothland massesare bathedby coldoceanic currentsof Arctic origin. By contrast,the western reachesof both land massesare bathed by warm currents. In these western reaches,Gyrfalcon summer distribution is displacednorthward and dark variants predominate. The breedingrange of the Gyrfalcon, determinedby mapping the locationsof the specimenswe examined,differs little from the range proposedin 1951. Distribuci6ny variaci6ndel coloren el Gerifalte en Rusia EXTRACTO.--Esp•cimensde Gerifalte (Falcorusticolus), existentes en museosde Moscfi (73) y San Petesburgo(132), fueron separadosen cuatroclases de colores(gris, gris claro, blancogrisiceo y blanco) dentrode cadauna de las cuatroregiones longitudinales que representanlas mayoresregiones fisiogrificas del norte de Rusia. Las variantesgrises predominan en las regionesdel oestey del centro.Las aves blancasson mils comunesen el extremoeste de Siberia, y hastason ocasionalmente encontradas al oeste de los Montes Urales. Las frecuenciasson las siguientes:Rusia europea4% blanco,50% gris (el resto estabaen el intermedio);Siberia occidental 0% blanco,58% gris; Siberia central 15% blanco,42% gris; y Siberia oriental 47% blanco, 33% gris. Es de notar que aves blancaspredominan aun cerca de la extensi6nsur de la subregi6nextremo-oriental. Las porcionesdel extremonorte de la regi6n de Rusia continental,donde se halla esta especie,estiin en Siberia central dondelas variantesblancas son raras. Por estoproponemos que un mejor pronosticador de la variante blancaes la longitudy no la latitud. Las avesblancas son mils frecuentesen los extremos orientalestanto del Paleirtico comodel Neirtico. E1 mils importantefactor de esteambiente, que corre- lacionacon estatendencia de distribuci6n,es el isoclinotermal que se curva hacia el sur y que continfia hacia el este,en direcci6nde las regionesde Groelandiao Kamchatkaque sonbafiadas por corrientes ocelnicasfrlas de origen irtico. En contraste,el oestede estasdos regioneses bariadopor corrientes cilidas. En estaszonas occidentales, la distribuci6nde verano del F. rusticolesse desplazahacia el norte; y predominanlas variantesoscuras. 81 82 DAVID H. ELLIS ET AL. VOL. 26, NO. 2 E1 fireade reproducci6nde estehalc6n, basada en lossitios donde fueron colectados los esp6cimens de museo,no es muy diferentede la propuestaen 1951. [Traducci6nde Eudoxio Paredes-Ruiz] In the four decadessince publication of Demen- for dates (from the Julian to Gregorian calendars)and tiev's (1951) monograph on the Gyrfalcon (Falco locations(from pre- to post-revolutionaryregions). Spec- imens that lacked sufficient data were excluded from our rusticolus),little new informationhas been published analysisas were the few birds with spuriousdata. For in English on this speciesfor the former SovietUnion example,we excludedZIAS No. 75403 (which by color (comprisingnearly 50% of the species'breeding and sizeis a large second-yearfemale Gyrfalcon)because range). Dementiev's(1951) southernlimit of breed- its original label identified it as a male PeregrineFalcon ing is well substantiatedby museumspecimens, but (F. peregrinus);its collectiondate and localeare alsobetter matchesfor a breedingperegrine. did not loop southto includethe middleUral Moun- This study required that we deal with the difficulties tains where he (Dementiev 1960) reportedthat Gyr- presentedby the possibletaxonomic affinities of the Gyr- falconsbred a century earlier as far south as the falcon,Saker (F. chefrug),and Altay (or Altai) Falcon (F. Chusovaya,Ufa and Byelaya Rivers (seeFig. 1). rusticolus/cherrugaltaicus). Our responsewas to include Recent decadeshave seena trend in the published only birds that were clearly Gyrfalconsof the non-Altay Falcon type,judging either by collectionlocation and date, literature toward simplifying(i.e., eliminating un- or by plumage.This separationwas difficult for only a dulations) and compressingDementiev's (1951) few immatures,those which had the spottedtail pattern breedingrange without offeringsupporting data (e.g., thoughttypical of Sakers.As Cade (1982) indicated,some Cade 1982, Flint et al. 1984). These changes,if Sakersalso have Gyrfalcon-likebarred tails. We includedonly one specimen(ZIAS No. 75497) that correct, would signal population declinesand/or had previouslybeen treated as an Altay Falcon (cotypeof abandonment of formerly occupied range. The Gennaialorenzi: considered synonymous with F. r./c. al- boundarywe report is little changedfrom Demen- taicus,Sushkin 1938, Vaurie 1961). This specimenwas a tiev's 1951 map, thereforewe affirm the accuracyof large,gray femaleGyrfalcon and lackedthe brown dorsal backgroundcolor that characterizeseven the most Gyr- his map and recommendits continueduse until more falcon-likeof the supposedAltay types.Its location,near information is obtained. the southern end of the Ural Mountains and over 1000 Dementiev (1951, 1960) noted the general trend km from the mountainsof Central Asia where the Altay for Gyrfalconsto be lighter in colorand larger pro- Falcon reportedlyoccurs (Dementiev and Gladkov 1951), ceedingeastward acrossEurasia. Portenko (1972) and collection date (October or November 1900) are a goodmatch for a migrating Gyrfalcon. also noted that the white variant predominateson Recognizingthat colorvaries along a continuum(Vau- the Chukchi Peninsula.We provide frequenciesof tie 1961), we nonetheless,for comparisonpurposes, grouped color variants for the breadth of Russia based on all the specimensinto eight age/color classesas follows: examination of 205 specimens(91 adult and 114 white variants (adult and juvenile) were birds with white as a dorsalbackground color; white gray birdswere adults juveniles)in the two primary bird collectionsin Rus- with dorsaldark barring approachingthat of gray adults sia, the University of Moscow ZoologicalMuseum but with very little gray in an otherwisewhite dorsal (UMZM) and the ZoologicalInstitute of the Aca- background;white brown birds,the correspondingjuvenile demic Sciences(ZIAS) in St. Petersburg. In dis- variant,were palebrown dorsally with light tips and edges cussingcolor variation, we amplify Vaurie's (1961), and were lightly streakedwith brown ventrallyon a pale creambackground color; light gray birds(and light brown Cade's(1982), and Palmer's(1988) observationsthat juveniles)had the heavilybarred dorsal pattern of normal Gyrfalconplumage is infinitelyvariable along a con- gray adults(or brown mantle if juveniles)but their dorsal tinuum from the whitest individualsto dark gray or backgroundcolor, and oftenthe hue of their light and dark "black" birds. To recognizecolor phasesor, more bars, was much lighter than the gray variant (or brown variant if juvenile); gray (or brown if juvenile) variants precisely,color morphs (e.g., white, gray, brown, differed from light gray (or light brown) birds in having black) is misleading. It is instructive, however, to darkergray (or brown) pigmentationdorsally. Gray adults report regional trends in the occurrenceof light, often had a light gray wash ventrally, especiallyon tibias dark, and intermediate birds. and lowerbelly, while juveniles had darkerventral streak- ing and a darker buff wash than other variants.Birds in mixed plumage(molting) providedevidence that the four METHODS juvenileclasses molt into correspondingadult classes(e.g., This study is based primarily on examination of the white brown juveniles becomewhite gray adults). Ex- Gyrfalcon skins, specimen tags and cataloguesat the amplesof adults (and two juveniles) representingmost of UMZM (73 specimens)and the ZIAS (132 specimens). our light and dark variantsare illustratedin Weick's(1980) Great effort was expendedto translateold Russianscript Plate 39. Juveniles,although often showing a grayishbloom JUNE 1992 COLORVARIATION IN RUSSIANGYRFALCONS 83