Cryptic Species The White-breasted Nuthatches 4UFWFO(.MPEJOPX -POHNPOU$PMPSBEP In Brief THNMPE!BPMDPN D uring 2013, the American Ornithologists’ Union (AOU) evaluated a proposal to “split” the Whitebreasted Nuthatch (Sitta carolinensis) into any- where from two to four species. The proposal did not pass (AOU 2013a). Members of the AOU’s Committee on Classification and Nomenclature of North and Middle American Birds were, however, nearly unanimous in their desire to revisit the question soon, after more data have been gathered (AOU 2013b). What does this mean for North American birders? Certainly more identification challenges, but also an opportunity to contribute to our understanding of the current and changing ranges of these birds. Background A key geological event was the formation, 14–2½ million years ago, of the “Basin and Range” district of western North America. This event promoted the splitting of forests of the U.S. and Canada into three distinct regions: Eastern (east of the Mountain West, and extending north to include much of the continent’s northern reaches); Interior West (from the Rocky Mountains west to the eastern slopes of the Sierra Nevada and the Cascades, south to the mountains of northern Mexico, and including the Davis and Chisos mountains of Texas); and Pacific (from the crests of the Sierra Nevada and the Cascades west to the Pacific Coast). A more complete summary is given by Graham (1999). These distinctions were reinforced by “recent” Quaternary Period glacial cycles (Van Devender 1990, Graham 1999) and sub- This article presents an overview, based on recent scientific literature and the author’s field and museum work, of our present knowledge of the identification and geographic distribution of the White-breasted Nuthatches of North America. The bird currently classified as the White-breasted Nuthatch exhibits considerable geographic variation, and may well comprise multiple species. Three distinctive populations—the “Carolina,” “Rocky Mountain,” and “Slender-billed” nuthatches—occur in the ABA Area. Field identification is problematic where the ranges of these populations overlap (Carolina and Rocky Mountain, mid-continent; Rocky Mountain and Slender-billed, farther west). Additional challenges involve vagrancy, hybrids, and natural variation within each group. Fortunately, careful study in the field often permits a credible identification, especially if a nuthatch is both seen and heard; photos and audio can be very helpful too. 26 # * 3 % * / (t.":+ 6 / & sequently led to many of the tripartite groupings of species and subspecies encountered in North America north of Mexico (Yellow-bellied, Rednaped, and Red-breasted sapsuckers; Blue-headed, Plumbeous, and Cassin’s vireos; etc.). White-breasted Nuthatches divide nicely along these lines, with Eastern (S. c. carolinensis), Interior West (S. c. nelsoni), and Pacific (S. c. aculeata) subspecies-groups (Grubb and Pravosudov 2008) represented. Spellman and Klicka (2007) analyzed a single mitochondrial DNA (mtDNA) gene and concluded that there are four major groupings (or “clades”) of White-breasted Nuthatches; they split the Interior West group further into Rocky Mountain and Sierra Nevada/Cascade groupings, S. c. nelsoni and S. c. tenuissima, respectively. Gene flow between these two Interior West groups is not trivial. Another study, looking at one mtDNA gene and 19 nuclear DNA genes, found the same four groups (Walstrom et al. 2012), although sampling near contact zones was limited. Issues with gene flow and sample size were major reasons why the AOU decided not to split the White-breasted Nuthatch into multiple species. Furthermore, checklist committee members seemed inclined to split the White-breasted Nuthatch into three species (keeping the Interior West group as one) rather than four (AOU 2013b). No common names were proposed for these possible new species. Therefore, I follow the AOU’s (1910) nomenclature, using Carolina Nuthatch for the carolinensis group, Rocky Mountain Nuthatch for the nelsoni group, and Slender-billed Nuthatch for the aculeata group. Given current knowledge, the Rocky Mountain (nelsoni, in the strict sense) and Sierra Nevada/Cascade (tenuissima) taxa are not reliably separable in the field, so I will discuss them as a single entity, the Rocky Mountain Nuthatch (see Grubb and Pravosudov 2008). Distribution of Proposed Species Carolina Nuthatch This nuthatch is largely resident from the Peace River region of northeastern British Columbia east through central Alberta, Saskatchewan, and Manitoba to Prince Edward Island and south to eastern Texas, northern Louisiana, and Georgia, plus a small area of the Florida Panhandle (Dunn and Al- "#"03(#*3%*/( UFSUJBMT HSFBUFS DPWFSUT nBOLT Fig. 1. *OUIJTQPTFBGFNBMF8IJUFCSFBTUFE /VUIBUDITVCTQFDJFTcarolinensisi$BSPMJOB /VUIBUDIw TIPXTBMMPGUIFWJTVBMDIBSBD UFSTEFTDSJCFEJOUIJTBSUJDMF'JFME TFQBSBUJPOPGUIFUISFFNBJO8IJUF CSFBTUFE/VUIBUDIQPQVMBUJPOT SFRVJSFTBOBQQSFDJBUJPOBOE VOEFSTUBOEJOHPGUIF FõFDUTPGGFBUIFSXFBS QPTUVSFMJHIUJOH BOEOBUVSBM WBSJBUJPO Photo by © Garth McElroy. 27 W H I T E - B R E A S T E D N U T H ATC H E S 2a 2b 2c Fig. 2.5IFTFQIPUPTEFQJDUUIFOBSSPXFSEBSLDSPXOPGUIF3PDLZ.PVOUBJO/VUIBUDI'JHB DPNQBSFEUPUIFCSPBEFSDSPXOTPG$BSPMJOB 'JHC BOE4MFOEFSCJMMFE'JHD OVUIBUDIFT5IFDPOUSBTUJOHUFSUJBMBOEHSFBUFSDPWFSUTPGUIF$BSPMJOB/VUIBUDIBSFBMTPFWJEFOUFWFO XJUIBEPSTBMiUPQEPXOw WJFX5IF3PDLZ.PVOUBJO/VUIBUDI'JHB TIPXTEBSLFSUFSUJBMTUIBOBWFSBHFBOEOBSSPXMZEBSLCVUOPUCMBDL HSFBUFSDPWFSUDFOUFSTJODPOUSBTUUPUIFCSPBECMBDLHSFBUFSDPWFSUDFOUFSTPGUIF$BSPMJOB/VUIBUDI'JHC 5IF4MFOEFSCJMMFE/VUIBUDI 'JHD TIPXTBQMBJOXJOHOPUSBSFGPSUIBUUBYPOCVUQFSIBQTEVFJOQBSUUPXFBSPOUIJTJOEJWJEVBMFig. 2a. Rocky Mountain Nuthatch. Chico Basin Ranch, El Paso County, Colorado; 2 October 2013. Photo by © Steven G. Mlodinow. Fig. 2b. Carolina Nuthatch. 5IJTJTUIFTBNFCJSE BTJO'JHD Crow Valley Campground, Weld County, Colorado; 21 September 2012. Photo by © Steven G. Mlodinow. Fig. 2c. Slender-billed Nuthatch. Ridgefield, Clark County, Washington; 12 January 2009. Photo by © Scott Carpenter. derfer 2011, eBird data accessed 26 March 2014). The Carolina Nuthatch is at least partially migratory, with wintering birds westward to the Front Range of Colorado (Mlodinow 2014), southwest to central Texas (Phillips 1986, Lockwood and Freeman 2014), and south to the Gulf Coast from Louisiana to Florida (Stevenson and Anderson 1992, Turcotte and Watts 1999, eBird data accessed 26 March 2014). Additionally, large numbers have been noted migrating past Cape May Point (T. Leukering, personal communication). A range shift appears to have occurred over the past century, with Carolina Nuthatches establishing themselves as breeders along the South Platte River in northeastern Colorado, starting approximately in the 1980s (Mlodinow 2014), and retreating as breeders from central and most of northern Florida since at least the 1950s (Stevenson and Anderson 1992). Vagrants have been recorded on Sable Island, Nova Scotia (Phillips 1986), in easternmost New Mexico (Baumann et al. 2013), and in southern Florida (Stevenson and Anderson 1992). Rocky Mountain Nuthatch This nuthatch is largely resident from south-central British Columbia and the Washington Cascades south into Nevada and easternmost southern California and southeastern Arizona. The range extends east to southernmost Alberta, the southwestern half of Montana, Wyoming, and westernmost Texas, 28 then into the cordilleras of Mexico south to Oaxaca. The Rocky Mountain Nuthatch also occurs east from the Rocky Mountains along the southernmost edge of Colorado east to westernmost Oklahoma, plus there are isolated populations in the Sierra de la Laguna of Baja California Sur, the Black Hills of South Dakota (Spellman and Klicka 2007, Grubb and Pravosudov 2008, Dunn and Alderfer 2011), and the Pine Ridge of Nebraska (R. Wright, personal communication). The Rocky Mountain Nuthatch is at least a partial migrant, with a few occurring regularly to the eastern border of Colorado (Mlodinow 2014), into the Nebraska panhandle (eBird data accessed 26 April 2013), and in the lowland areas of New Mexico (Dunn and Alderfer 2011). Vagrants have occurred in southwestern Kansas (Phillips 1986) and San Diego County, California (Unitt 2004). Slender-billed Nuthatch This nuthatch is largely resident from southwestern Washington through western Oregon, most of California, and into northwestern Baja California; there also is an isolated population in Sierra de San Pedro Mártir of Baja California (Spellman and Klicka 2007, Grubb and Pravosudov 2008, Dunn and Alderfer 2011). An isolated population in the southern Puget Sound area of Washington has been extirpated since at least the 1990s (Wahl et al. 2005). Vagrants have occurred in central Arizona # * 3 % * / (t.":+ 6 / & and on Vancouver Island, British Columbia (Phillips 1986). Identification Call notes Identification criteria for White-breasted Nuthatch taxa typically emphasize the differences in the call notes, which is reasonable, as these birds’ calls are generally distinct from one another. The most distinctive is the Rocky Mountain Nuthatch, which usually utters a rapid, stuttering, polysyllabic di-di-di-di-di burst that carries with it a sense of urgency. Carolina and Slender-billed nuthatches both give calls that recall the Red-breasted Nuthatch, being somewhat nasal and usually emitted at a more leisurely pace than the calls of the Rocky Mountain Nuthatch: a throaty, emphatic yarnk...yarnk... for the Carolina, and a shriller, more wavering yurrr...yurrr... for the Slender-billed. With both Carolina and Slender-billed nuthatches, these notes are typically given one at a time, monosyllabically. Excited Carolina and Slender-billed nuthatches are prone to speeding up the rate of their calling, thus approaching the Rocky Mountain Nuthatch, and occasional Rocky Mountain Nuthatches will call in a more leisurely fashion, thus sounding more like the other two taxa. Indeed, recorded calls attributed to one taxon in the field have proven to be of another via subsequent analysis of sound spectrograms of the calls (T. Floyd, personal communication). The songs of White-breasted Nuthatches—soft, whistled sequences, sounding like a distant Northern Flicker—are less variable among the three main groups, and are not discussed here. Fig. 3.5IFTFQIPUPTTIPXEJõFSFODFTJOUIFTJEFBOEnBOLQBUUFSOT PGUIFUISFFHSPVQTPG8IJUFCSFBTUFE/VUIBUDIFT/PUFUIFNFEJ VNEBSLHSBZPGUIF3PDLZ.PVOUBJO/VUIBUDI'JHB WTUIFXIJUF TJEFTPGUIF$BSPMJOB/VUIBUDI'JHC "OPUIFS$BSPMJOB/VUIBUDI 'JHD IBTEBSLFSnBOLTCVUSBUIFSQBMFTJEFTXJUIPSBOHFoCVõ GFBUIFSTNJYFEJO5IF4MFOEFSCJMMFE/VUIBUDI'JHE TIPXTQBMF HSBZnBOLTXJUIBUPVDIPGPSBOHFoCVõ"MTPXPSUIOPUJOHJTIPX DMPTFMZUIFCMBDLDSPXOPGCPUI$BSPMJOB/VUIBUDIFT'JHTCoD BQQSPBDIUIFFZFBTXFMMBTUIFUIJOCMBDLMJOFFYUFOEJOHCBDL GSPNUIFFZFPOUIJTUBYPO5IFIFBENBSLJOHTPGUIF4MFOEFSCJMMFE /VUIBUDI'JHE BSFTJNJMBSUPUIPTFPGUIF$BSPMJOB'JHTCoD JODMVEJOHBEBTIPGCMBDLCFIJOEUIFFZF/PUFUIBUUIFCJMMPGPOFPG UIF$BSPMJOB/VUIBUDIFT'JHC TFFNTUPNBUDIUIFMFOHUIPGUIF CJMMPGUIF3PDLZ.PVOUBJO/VUIBUDI'JHB Fig. 3a. Rocky Mountain Nuthatch.5IJTJTUIFTBNFCJSEBTJO'JHB Conifer, Jefferson County, Colorado; 28 December 2012. Photo by © Steven G. Mlodinow. Fig. 3b. Carolina Nuthatch. Wray, Yuma County, Colorado; 1 September 2013. Photo by © Steven G. Mlodinow. Fig. 3c. Carolina Nuthatch. Ovid, Sedgwick County, Colorado; 3 November 2013. Photo by © Steven G. Mlodinow. Fig. 3d. Slender-billed Nuthatch.5IJTJTUIFTBNF CJSEBTJO'JHE Portland, Multnomah County, Oregon; 10 May 2013. Photo by © Scott Carpenter. ABA.ORG/BIRDING 3a 3b 3c 3d 29 W H I T E - B R E A S T E D N U T H ATC H E S Table 1%JTUJOHVJTIJOH$IBSBDUFSTPG8IJUFCSFBTUFE/VUIBUDI(SPVQT Carolina Rocky Mountain Slender-billed Greater Coverts Pattern Prominent black or blackish centers with contrasting grayish edging Dusky to gray centers with broad grayish edging; little or no contrast Typically gray centers, sometimes dusky, with little to no contrast; plainest of all three Outermost Tertial Pattern Black on inner and outer webs Black to dusky on inner webs, dusky on outer webs Black to dusky on inner webs, dusky on outer webs; averages paler than Rocky Mountain Upperparts Coloration Paler gray Darker gray Medium gray Dark Crown Width Broad Narrow Broad Facial Pattern Black of crown approaches eye more closely than Rocky Mountain; many birds have small black dash or line extending back from eye Black of crown farther from eye, typically forming arc over face; no black mark behind eye Black of crown approaches eye fairly closely; occasionally small black dash back from eye Side/Flanks Coloration Pale gray to white, sometimes suffused with buff Medium blue-gray, rarely with buff wash Pale gray to white, sometimes suffused with buff Bill Length Averages shorter and stouter Averages longer and more slender Averages shorter and more slender Also, all White-breasted Nuthatches give weak, mumbling, nasal notes when foraging: plink, pleek, etc. Audio recordings and sound spectrograms of typical calling sequences are available on the Birding magazine website: aba.org/birding Visual characters The standard identification literature has given short shrift to visual identification cues, but I have found that these are surprisingly useful. The discussion in Wood (2006) provides a good starting point. To help clarify the usefulness of various visual characters, I studied approximately 150 specimens each of Carolina and Rocky Mountain nuthatches and 50 Slender-billed Nuthatch at The Field Museum of Natural History in Chicago and at the University of Puget Sound’s Slater Museum of Natural History. Also reviewed were more than 100 photos of each taxon online from locations at which only one is likely—for example, Illinois for Carolina Nuthatch. I found the three most useful characters to be the following: (1) the pattern of the greater secondary coverts (hereafter, greater coverts), (2) the outermost tertial pattern, and (3) the width of the dark crown. Two other characters were useful: (4) the side/flank coloration and (5) the facial pattern. I was able to find no consistent difference in vent (between the legs) pattern, and I believe that the differences in back color are probably of little value in the field. Fig. 1 provides an overview of terminology (greater coverts, tertials, flanks), and Table 1 summarizes useful and proposed differences. Greater Coverts Carolina Nuthatches have blackish centers to their greater coverts that contrast markedly with the feathers’ edges. Rocky Mountain Nuthatches are variable in this character, ranging from rather uniformly marked greater coverts to coverts with dusky (but not black) centers. In my museum and photographic studies, I found that birds from the northwestern portion of 30 this taxon’s range seem to show the plainest greater coverts, whereas birds from Arizona and New Mexico average darkercentered greater coverts. Some Rocky Mountain Nuthatches have almost blackish centers to their greater coverts, but these are never as dark as those of Carolina Nuthatches, and when present, the dark center is narrower. Slender-billed Nuthatches usually have plain greater coverts, with little or no contrast. These differences would seem to make the identification of Carolina Nuthatches straightforward. However, feather positioning can confound interpretation, as the scapulars or side feathers sometimes partly veil the greater coverts, making the dark centers less apparent. Also, the centers of the greater coverts are less evident if these feathers are crowded together and more evident if they are spread apart. Tertials On all three taxa, only the inner web of the smallest (innermost) two tertials is black or dusky. On the largest (outermost) tertial, Carolina Nuthatches are black on the inner and outer webs, Rocky Mountain Nutatches are dusky (occasionally black, especially on southern birds) on the inner web and dusky on the outer web, and Slender-billed Nuthatches are gray to dusky (rarely blackish) on both the inner and outer webs. Again, feather positioning can make this character difficult to assess. Crown and Face Carolina Nuthatches have a rather broad dark crown when viewed from above. In Rocky Mountain Nuthatches, the crown stripe is relatively narrow. Slender-billed Nuthatches have a crown width similar to that of Carolina Nuthatches, but may average a bit narrower. When viewed from the side, the broader crown of Carolina and Slender-billed nuthatches tends to bring the crown edge closer to the eye than in Rocky Mountain Nuthatches. Be advised that the posture of a bird can affect the ap- # * 3 % * / (t.":+ 6 / & pearance of crown width, particularly with side views. Many, perhaps most, Carolina Nuthatches have a narrow black line extending backward from the eye, sometimes short, sometimes rather long. This mark is seemingly never present on Rocky Mountain Nuthatches. Some Slender-billed Nuthatches appear to show a very brief dark mark behind the eye. The H-word Sides/Flanks I owe a debt of gratitude to The Field Museum of Natural History and the University of Puget Sound’s Slater Museum of Natural History for their generosity in allowing me to ponder their specimens ad nauseam. Additionally, Scott Carpenter and David Irons are to be thanked for making so many images of Slender-billed Nuthatch available to me. Finally, but not least, many thanks to Tony Leukering, Mark Lockwood, and Michael O’Brien for helpful comments on an earlier version of this manuscript. The sides of Carolina and Slender-billed nuthatches are whitish or pale blue–gray, sometimes washed in orange–buff. The sides of Rocky Mountain Nuthatches are pale to medium blue–gray, sometimes nearly as dark as the back, rarely washed in orange–buff. Bill There is extensive overlap in bill size, with bill length ranging from 15.4 to 19.5 mm in Carolina Nuthatch, from 16.9 to 21.7 mm in Rocky Mountain Nuthatch, and from 16.3 to 19.2 mm in Slender-billed Nuthatch (Pyle 1997). In Colorado, where Carolina and Rocky Mountain nuthatches occur, I have found bill length to be of little use in the field. Slender-billed Nuthatches do, however, appear to have a shorter and, in particular, slimmer bill; bill shape of this taxon may well be useful for field identification. Where the ranges of different Whitebreasted Nuthatch populations intersect, it is quite possible, if not likely, that some hybridization occurs. One should always be open to the possibility of hybridization when confronted with potential vagrants or in regions where the ranges of two taxa approach or overlap one another. Acknowledgments 4a 4b 4c Literature Cited "NFSJDBO0SOJUIPMPHJTUT6OJPO<"06> The AOU Check-list of North American Birds SE FE "NFSJDBO 0SOJUIPMPHJTUT 6OJPO /FX:PSL "NFSJDBO 0SOJUIPMPHJTUT 6OJPO <"06> B 'JGUZGPVSUI TVQQMFNFOU UP UIF AOU Check-list of North American Birds. Auk o 4d Fig. 4.5IFTFQIPUPTTIPXEJõ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õVTFEJOUPUIFEBSL CMVFoHSBZTJEFTPGPOFPGUIF3PDLZ.PVOUBJO/VUIBUDIFT'JHB BOJOGSFRVFOUCVUOPU PWFSMZSBSFUSBJUFig. 4a. Rocky Mountain Nuthatch.5IJTJTUIFTBNFCJSEBTJO'JHB Conifer, Jefferson County, Colorado; 28 December 2012. Photo by © Steven G. Mlodinow. Fig. 4b. Rocky Mountain Nuthatch. Patagonia, Santa Cruz County, Arizona; 23 February 2013. Photo by © Steven G. Mlodinow. Fig. 4c. Carolina Nuthatch. 5IJTJTUIFTBNFCJSEBTJO'JHC Crow Valley Campground, Weld County, Colorado; 21 September 2012. Photo by © Steven G. Mlodinow. Fig. 4d. Slender-billed Nuthatch.5IJTJTUIFTBNFCJSEBTJO'JHE Portland, Multnomah County, Oregon; 10 May 2013. Photo by © Scott Carpenter. ABA.ORG/BIRDING 31 W H I T E - B R E A S T E D N U T H ATC H E S Fig. 5.5IFBVUIPSGPVOEUIBUCPUI$BSPMJOBBOE3PDLZ .PVOUBJOOVUIBUDIFTBQQBSFOUMZTQFOUUIFTVNNFSPG BU#BSS-BLF"EBNT$PVOUZ$PMPSBEP5IJTJOEJ WJEVBMQIPUPHSBQIFEBUUIBUTJUFUIFGPMMPXJOHXJOUFS IBTUFSUJBMNBSLJOHTMJLFUIPTFPGB$BSPMJOB/VUIBUDICVU UIFEBSLJTIHSBZTJEFTMBDLJOHCVõBSFUZQJDBMPGB3PDLZ .PVOUBJO/VUIBUDI5IFDSPXOQBUDITFFNTJOUFSNFEJBUF JOXJEUIBOEUIFSFBQQFBSTUPCFBUJOZCMBDLNBSLCFIJOE UIFFZF5IFHSFBUFSDPWFSUTBSFIBSEUPKVEHFBTUIFZ BSFQBSUMZWFJMFECZUIFTDBQVMBSTCVUUIFZBQQFBSCMBDL DFOUFSFEBTPOB$BSPMJOB/VUIBUDI5IJTCJSETDBMMXBT JOEFDJQIFSBCMFUPUIFBVUIPS5IJTDPNCJOBUJPOPGGFBUVSFT TVHHFTUTUIBUUIJTCJSEJTB$BSPMJOB/VUIBUDIY3PDLZ .PVOUBJO/VUIBUDIIZCSJEBarr Lake, Adams County, Colorado; 30 December 2012. Photo by © Steven G. Mlodinow. Final Thoughts "NFSJDBO0SOJUIPMPHJTUT6OJPO<"06>CAOU Committee on Classification and Nomenclature–North and Middle America: Votes on Proposals 2013-CUJOZVSMDPN"06QSPQPTBMT #BVNBOO.+4..D/FXBOE$88JUU.PSQIPMPHJDBMBOE NPMFDVMBSFWJEFODFDPOmSNUIFmSTUEFmOJUJWF&BTUFSO8IJUFCSFBTUFE /VUIBUDISitta c. carolinensis GPS/FX.FYJDPWestern Birdso %VOO+-BOE+"MEFSGFSFETThe National Geographic Field Guide to the Birds of North America/BUJPOBM(FPHSBQIJD4PDJFUZ8BTIJOHUPO (SBIBN " Late Cretaceous and Early Cenozoic History of North American Vegetation0YGPSE6OJWFSTJUZ1SFTT/FX:PSL (SVCC5$BOE771SBWPTVEPW8IJUFCSFBTUFE/VUIBUDISitta carolinensis JO"1PPMFFEThe Birds of North America OnlineUJOZVSM DPN#/"8#/V $PSOFMM-BCPG0SOJUIPMPHZ*UIBDB -PDLXPPE . 8 BOE # 'SFFNBO The TOS Handbook of Texas BirdsOEFE5FYBT".6OJWFSTJUZ1SFTT$PMMFHF4UBUJPO 1IJMMJQT"3The Known Birds of North and Middle AmericaQBSU "31IJMMJQT%FOWFS 32 1ZMF1Identification Guide to North American BirdsQBSU* 4MBUF$SFFL1SFTT#PMJOBT 4QFMMNBO ( . BOE + ,MJDLB 1IZMPHFPHSBQIZPGUIF 8IJUFCSFBTUFE /VUIBUDI Sitta carolinensis %JWFSTJmDBUJPO JO /PSUI "NFSJDBO QJOF BOE PBL XPPEMBOET Molecular Ecologyo 4UFWFOTPO).BOE#)"O EFSTPO The Birdlife of Florida 6OJWFSTJUZ PG 'MPSJEB 1SFTT(BJOFTWJMMF 5VSDPUUF8)BOE%-8BUUTBirds of Mississippi.JTTJTTJQQJ%F QBSUNFOUPG8JMEMJGF'JTIFSJFTBOE1BSLT+BDLTPO 6OJUU1San Diego County Bird Atlas. 1SPDFFEJOHTPGUIF4BO%JFHP 4PDJFUZPG/BUVSBM)JTUPSZOP 7BO%FWFOEFS53-BUF2VBUFSOBSZWFHFUBUJPOBOEDMJNBUFPG UIF4POPSBO%FTFSU6OJUFE4UBUFTBOE.FYJDPQQoJO+#F UBODPVSU537BO%FWFOEFSBOE14.BSUJOFETPackrat Middens: The Last 40,000 Years of Biotic Change 6OJWFSTJUZ PG "SJ[POB 1SFTT 5VDTPO 8BIM53#5XFJUBOE4(.MPEJOPXFETBirds of Washington. 0SFHPO4UBUF6OJWFSTJUZ1SFTT$PSWBMMJT 8BMTUSPN+8+,MJDLBBOE(.4QFMMNBO4QFDJBUJPOJOUIF 8IJUFCSFBTUFE/VUIBUDISitta carolinensis "NVMUJMPDVTQFSTQFD UJWFMolecular Ecologyo 8PPE$-/VUIBUDIFTGBNJMZ4JUUJEBFQQoJO+"MEFS GFSFEComplete Birds of North America/BUJPOBM(FPHSBQIJD4PDJFUZ 8BTIJOHUPO Recent evidence suggests that three or four species are represented by what we currently label the White-breasted Nuthatch. Most individuals of these “new” species are identifiable with careful study of calls and plumage. Correct identification requires that observers understand the effects of posture, plumage wear, and lighting—as well as the somewhat subjective nature of human hearing and seeing. As always, an appreciation for variation in field characters—aural and visual—is essential. Photos and audio recordings may be necessary for establishing the identity of birds well out of range. Many such “cryptic” species have been recognized over the past decade or two, mainly in the tropics, but also right here in the ABA Area. Perhaps the most famous is the Gunnison Sage-Grouse, described to science only in 2000. A very recent example is that of the erstwhile Sage Sparrow, now considered to comprise two species, the Bell’s and Sagebrush sparrows; conceivably, a third species is involved. Genetic data are certainly part of the elucidation of cryptic species, but phenotypic characters play a role, too. For example, the differences between what we today know as the Bell’s and Sagebrush sparrows were well understood by the ornithologists of the 19th century. And the recent discovery of the Gunnison Sage-Grouse was based on a broad suite of evidence: behavioral, vocal, physiological, and of course genetic. By being more aware of birds at the subspecies—or apparent subspecies—level, rather than simply at the species level, savvy birders can contribute to our understanding of biogeography as well as identification criteria. # * 3 % * / (t.":+ 6 / &
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