SSttaattuuss aanndd ddiissttrriibbuuttiioonn ooff TTyyppee 11 RReedd CCrroossssbbiillll ((LLooxxiiaa ccuurrvviirroossttrraa )):: aann AAppppaallaacchhiiaann CCaallll TTyyppee??

MATTHEW A. YOUNG • CORNELL LAB OF • 159 SAPSUCKER WOODS ROAD, ITHACA, NEW YORK 14850 • OMAY6NCORNELL.EDUP KEN BLANKENSHIP • 2400 BARRETT CREEK BOULEVARD #827, MARIETTA, GEORGIA 30066 • OKENHBLANKENSHIPNCOMCAST.NETP MARILYN WESTPHAL • P. O. BOX 1427, HENDERSONVILLE, NORTH CAROLINA 28793 • OMJWESTPHNRET.UNCA.EDUP STEVE HOLZMAN • 349 JEFFERSON ROAD, BISHOP, GEORGIA 30621 • OSTEVE_HOLZMANNYAHOO.COMP

records of our own, and also adduce the broad - er natural history of this Type, using primarily recent records of calling that have been identified by audiospectrographic analysis. We review evidence that Type 1 is not a specialist on a single of (as some Types appear to be), and from that we infer that wan - derings away from the core breeding range in Appalachia are driven by cone crop failures of several species. We further suggest that wan - dering Type 1 birds, which have medium-sized bills among the Red Types, settle most readily in areas where cone sizes and types are most similar to those found in core range. We urge observers to audio-record Red wherever they are detected and to note the species of conifer in which they are observed, to increase our understanding of their complex natural histories.

Background: Types and their natural histories Red Crossbill ( Loxia curvirostra ) has been di - vided into ten “call types” (hereafter Types 1- 10) in ; in addition to differ - ences in flight calls, small but significant dif - ferences in bill morphology, ecology, and ge - netics distinguish these Types (Groth 1993c, Benkman 1999, Parchman et al. 2006, Irwin 2010). Groth (1993c) was the first to assert that the Types meet criteria for full-species sta - tus under both phylogenetic and biological species concepts and that they should there - fore be recognized as taxonomic species. More recent studies continue to inform discussion of Red Crossbill’s taxonomic status. Sewall (2009, 2011) and Sewall and Hahn (2009) present ev - idence to support the ideas that Red Crossbill This Red Crossbill was audio-recorded and confirmed as Type 1, a Type that appears to be predominantly an Appalachian . flight calls are an example of reliable signaling This individual was photographed along Persimmon Road, Clayton, Georgia on 5 July 2008. Photograph by Darlene Moore. and that adults show limited learning of vocal - izations. Benkman (2007) and Benkman et al. Abstract Northwest have at times called this association (2009) argue that Types are effectively repro - Red Crossbill ( Loxia curvirostra ) of Type 1 has into question. In this paper, we review evi - ductively isolated. Interbreeding between been considered chiefly an Appalachian bird, dence in support of this association (n=186 Types seems to be relatively rare, despite sym - though detections of this Type as far away from Appalachian recordings of Type 1, n=14 non- patry of several Types and widespread wander - Appalachia as New England and the Pacific Appalachian recordings of Type 1), including ing by many Types. In several published stud -

554 NORTH AMERICAN BIRDS STATUS AND DISTRIBUTION OF TYPE 1 RED CROSSBILL: AN APPALACHIAN CALL TYPE?

ies, interbreeding between various Types has been estimated at up to 5% (Groth 1993b, Summers et al. 2007, Benkman et al. 2009). In the South Hills of Idaho, where the resident population (Type 9) occurs sympatrically with Types 2 and 5, interbreeding was found to be less than 1% among 1704 paired crossbills (Benkman et al. 2009). Similarly, in a Euro - pean study of sympatrically occurring Red (or Common) Crossbills ( L. curvirostra ), Crossbills ( L. pytyopsittacus ), and Scottish Crossbills ( L. scotica ), Summers et al. (2007) Figure 1. A male Type 1 Red Crossbill with a fledgling visible in the left part of the frame (at left); the same male is shown found evidence of hybridization in approxi - feeding a different fledgling (right). These birds were audio-recorded and photographed in Rabun County, Georgia on 25 mately 5% of pairs, mostly between the larger- April 2009. Images from video by Ken Blankenship. billed and . Benkman (1993a) further predicted the exis - conclusively documented for a few; definitive In a study of sympatrically occurring Types 1 tence of a Sitka ( P. sitchensis )-associated evidence of specialization for a primary and 2 in the mountains of Virginia and North Type—and indeed Irwin recently (2010) de - conifer is lacking for Types 1, 6, and 7. Carolina in the 1980s, Groth (1993b) found scribed Type 10 in the coastal Sitka Spruce Benkman (2007) suggested that Type 1 might that all 60 monitored birds paired with mem - forests of northern California. specialize on Red Spruce ( P. rubens ), a species bers of their own Types. The evolution of dietary specialization in largely confined to the Northeast (defined Types differ in bill depth and palate groove crossbill Types is readily understandable in here as the area from the Adirondack Moun - structures, and Benkman (1993a) proposed insular populations such as those of New - tains northward and northeastward to the that each Type is most efficient at feeding on foundland Red Crossbill (Benkman 1989, Canadian Maritimes). However, Red Spruce single key species of conifer, although not Benkman 1993b, Benkman and Parchman is also found in small isolated pockets along obligatorily. Other authors have proposed that 2009) or ( L. megapla - the spine of the central and southern Ap - certain Types have core zones of occurrence ga ) (Benkman 1994); it is also clear how is - palachians from Pennsylvania to North Car - where they regularly breed and are most com - land-like geography, such as Idaho’s South olina, and Groth originally thought that mon (Dickerman 1987, Knox 1992, Kelsey Hills, could give rise to differentiated cross - medium-billed Type 1 might be primarily an 2008, Young 2010a). While most Types do ap - bill populations, particularly where relatively Appalachian bird, but he later recorded two pear to forage most efficiently on a single stable cone crops are found (Benkman et al. Type 1 birds in British Columbia and set species of conifer when in their core zone of oc - 2009). Such specialization in other Types is aside the concept of an Appalachian Red currence (Benkman 1993a), they also feed on less clearly motivated and indeed is not yet Crossbill (Groth 1993c). seeds of multiple coniferous species and can be seen switching to those that provide Table 1 . Estimated conifer seed usage through the cone cycle year (June through May) for Type 1 Red Crossbill, as well the highest energy yields throughout the cone as usage of insects and bird feeding stations. These estimates are based on our field observations, as well as on our un - derstanding of cone-ripening phenologies and Type 1’s bill morphology. Southeast (SE) and Northeast (NE) are abbrevi - cycle (Benkman 1987, 1988). In North Ameri - ated herein. ca, Benkman (1993a) indicates that Type 2 ap - pears to be most efficient at feeding on Pon - Conifer/Area Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr May derosa ( scopularum ), Type White Spruce (NE) X X X X X-x X-x X-x x-o x-o x-o 3 on Western Hemlock ( heterophylla ), Red Spruce (NE and SE) X X X X X-x X-x x-x x-o x-o x-o x-o Type 4 on Douglas-fir ( Pseudotsuga menziesii European (Southern New York) x x x x x x x x x x x x menziesii ), and Type 5 on Lodgepole Pine ( Pi - White Pine (East) x X X X X-x X-x x-o x-o nus contorta ). Benkman (1993b) suggests that Type 8 feeds most efficiently on Black Spruce Eastern Hemlock (East) x x x x x x x x x x x ( ). And Benk - Red Pine (NE) x-X X X X man et al. (2009) note that Virginia Pine (mostly SE) x-X X X X Type 9—which has already Pitch Pine (mostly SE) x-X X X X been proposed as a full Shortleaf Pine (SE) x-X X X X species, South Hills Crossbill (L. sinesciurus )—feeds most Loblolly Pine (Georgia) x-X X X X efficiently on an isolated pop - Norway Spruce (Southern New York) x x x x-X X X X X ulation of Lodgepole Pine. Insects x x Feeders o-x o-x o-x o-x x x Figure 2. Audiospectrograph of Type 1 Red Crossbill, considered diagnostic, KEY: recorded 19 June 1983 in Montgomery X = used often County, Virginia by Jeffrey Groth. The x = used less frequently dark horizontal marks on the right sides X-x = birds are using resource less and starting to transition to other conifers of image are for kHz lines 3, 5, and 7. x-X = birds are transitioning from other conifers to a new conifer Image used and adapted from Groth o-x = birds could start utilizing alternative resource (e.g., feeders), depending on availability of others (1993a) with permission of the author. x-o = birds have exhausted resource

VOLUME 65 (2011) • NUMBER 3 555 STATUS AND DISTRIBUTION OF TYPE 1 RED CROSSBILL: AN APPALACHIAN CALL TYPE?

Table 2 . Totals of Red Crossbills from Red Spruce areas of the Appalachians in North Carolina during mid- through late May surveys conducted by Mar - ilyn Westphal. Note the absence of Red Crossbills on most May surveys. Date / Location Black Mountains Balsam Mountains Buncombe May 2006 18 0 0 May 2007 0 0 0 May 2008 0 (no survey) 0 May 2009 0 0 0 May 2010 1 0 0 May 2011 32 2 0 History of Type 1 in the Appalachian Mountains McNair (1988) provides a thorough review of historical Red Crossbill breeding records from the southern Appalachians, from the Virginias to Georgia. Griscom (1937) was among the first to suggest explicitly that a quasi-resident crossbill inhabit - ed the Appalachians. Subsequently, Stupka (1963) and Simpson (1974) found Red Crossbills breeding regularly in the moun - tains of North Carolina and Tennessee, and Scott’s (1981) ob - servations led him to suggest that a mostly resident population existed on Shenandoah Mountain in the Virginia Appalachians. Later in the 1980s, as he began to describe the various dif - ferences in Red Crossbill Types, Groth (1988) noted that Type 1 was consistently findable in the western mountains of Vir - ginia and North Carolina. In 1993, Douglas Gross document - ed Type 1 breeding in Pennsylvania (Hess et al. 1998), and in 1995, Type 1 was recorded in mature, second-growth forests of White Pine and Eastern Hemlock ( Tsuga canadensis ) in Rabun County, Georgia (Oberle and Forsythe 1995). Type 1 has been reported in 1998-2000 at Caesar’s Head, South Carolina, very close to the North Carolina border (Pitts 1988; Andrew Farnsworth, pers. comm.), although these have not yet been verified by recordings. Type 1 has also been confirmed in Georgia in 2008-2010 (Blankenship et al. 2010), with nesting confirmed in April 2009 (Figure 1). Type 1 was recorded at Bald Knob Ridge, North Carolina in July and August 2009 (by M. Westphal) and confirmed by audiospectrographic analysis (M. Young). In addition, Type 1 has been documented in southern New York annually 2004-2011, sometimes in large Figure 3. a) Type 1 flight call, recorded by Gregory F. Budney and Matthew A. Young at Georgetown, New York, 5 August 2006 (Macaulay Library of Sounds #137497). b) Type 1 flight call, recorded by Grant McCreary in Dawson County, Georgia, 29 May 2008 (see Blankenship et al. 2010). c) Type 1 flight call, recorded by Matthew A. Young at Pharsalia, New York, 26 July 2008. d) Type 1 flight call, recorded by Ken Blankenship in Rabun County, Georgia, 25 April 2009 (see Blankenship et al. 2010). e) Type 1 “variant” flight call (see Young 2010a), recorded by Matthew A. Young at Fabius, New York, 28 March 2009 (Macaulay Library of Sounds #139452). f) Type 1 flight call, recorded by Marilyn Westphal at Bald Knob Ridge, North Carolina, 22 August 2009. g) Type 1 flight call, recorded by Matthew A. Young at Georgetown, New York, 5 September 2009 (Macaulay Library of Sounds #139448). h) Type 1 flight call, recorded by Matthew A. Young at West Fulton, New York, 17 January 2010 (Macaulay Library of Sounds #139455). i) Type 1 flight call, recorded by Matthew A. Young at Ger - man, New York, 30 July 2010 (Macaulay Library of Sounds #139454). j) Type 1 flight call, recorded by Mike Nelson along Clingman’s Dome Road, Great Smoky Mountains National Park, North Carolina, 1 March 2011. k) Type 1 flight call, recorded by Diane Lepkowski and William Leigh in western Rockingham County, Virginia (within several miles of the West Virginia border), 1 May 2011. l) Type 1 flight call, recorded by David Caton and David Kirschke at Erwin, Tennessee, 19 May 2011. m) Type 1 flight call, recorded by Ken Blankenship at Hale Ridge, Georgia, 27 June 2011. n) Type 1 flight call, recorded by Matthew A. Young at Otselic, New York, 2 July 2011. Audiospectrographs by Matthew A. Young.

556 NORTH AMERICAN BIRDS STATUS AND DISTRIBUTION OF TYPE 1 RED CROSSBILL: AN APPALACHIAN CALL TYPE?

numbers and often breeding (e.g., in 2004, 2007, and 2009) (Young 2011). Thus, al - though the evidence is patchy, there are now confirmed records of Type 1 Red Crossbills nesting along the length of the Appalachians, from northern Georgia to southern New York. Since Groth began to audio-record Red Cross - bills in the early 1980s, we have located 186 recordings of Type 1 from southern New York to northern Georgia, whereas in exhaustive analyses of thousands of recordings from across the continent, we know of only 14 non- Appalachian recordings of Type 1 (Table 3). Red Crossbill Types are defined, and con - clusively documented, primarily by distinctive flight calls. All of the records noted above, with the exception of the South Carolina records, have been verified to refer to Type 1 Red Crossbill by audiospectrographic analysis, currently the only way to confirm identifica - tion of this Type. Type 1’s flight call shows a downward modulated component (Figures 2, 3). Type 2, which has often been documented Figure 4. Range of Type 1 Red Crossbill, based on published literature, documented records, and cone-ripening phenologies. in the vicinity of Type 1, has a similar flight Data are deficient from areas of the Northeast, Great Lakes, and West but also from areas here designated part of or near the “core zone of occurrence,” including Pennsylvania and western Maryland. Map created by Steve Holzman. call. The audiospectrograph of Type 1’s flight call almost invariably commences with an ini - conelets that mature the following year in and early spring are also the times of the year tial upward component, and the downward June and July. These species are also relative - when Red Crossbills start to show up at bird part descends more quickly than in Type 2’s ly inefficient at holding seed except in the feeding stations (Benkman 2010), where they flight call (Groth 1988, 1993c). Both Types best cropping years. Hard-coned such take primarily sunflower seed. Eastern Hem - can have secondary elements toward the end as Red ( P. resinosa ), Shortleaf ( P. echinata ), lock cones are used during the year as well, of the flight call, but these elements tend to be Pitch ( P. rigida ), Virginia ( P. virginiana ), particularly when other conifers have failed. much more common and prominent in Type Lodgepole, and Ponderosa develop immature In North Carolina, Type 1 has rarely been 1. To differentiate between these two Types green cones during May and June. These found during May surveys of Red Spruce with certainty, audiospectrographic analysis is cones do not start to mature until the follow - stands (Table 2), a conifer proposed as its key essential. In comparison to the flight call of ing year in September and October. Hard- food source during spring (Benkman 2007). Type 2, that of Type 1 is described as a quick - coned pines hold their seed longer than do This is not surprising, as Red Spruce cones er, drier, and sharper and is sometimes tran - other conifers, providing crossbill food dur - are mostly spent by spring. scribed as chewt-chewt-chewt (Evans and ing critical times of limited food supply in late In a typical year with average cone crops, O’Brien 2002, Young 2008a). Type 2 flight winter and early spring (Benkman 1993a). Type 1 in New York will often use Eastern calls sound a bit more robust, huskier, lower Red Crossbills may nest almost throughout White Pine cone crops for early nesting during in pitch, and more modulated than those of the year (Adkisson 1996). In New York, and January–February (Benkman 1987; M. Young, Type 1 (Young 2008a). probably in other areas, Type 1 Red Crossbills pers. obs.). In March and April, Type 1 in New appear to nest primarily in two peaks (Young York switches to primarily Red Pine and Nor - Ecological associations and movements 2008b, 2011): during late June –early Septem - way Spruce ( P. abies ) (Young 2011), the only away from the southern Appalachians ber, in which newly developing cone crops two conifers in the Northeast that regularly To analyze the movements and ecological as - are utilized, and again in January–April, in hold seed during these lean times of the year. sociations of Type 1 Red Crossbills, an under - which the remaining seed from the previous In the southern New York counties of standing of the phenologies of the conifers year’s crop is utilized. Table 1 summarizes our Cayuga, Madison, Cortland, Onondaga and used by this Type is vital. ( Picea observations and inferences of seed use by Chenango, Type 1 Red Crossbills have been spp.), ( Larix spp.), and hemlocks Type 1. In the central and southern Ap - present since at least 2004, with widespread (Tsuga spp.), all soft-coned conifers, develop palachians, during an average cone crop year, nesting observed in 2004 and 2007 (Young cone crops in May and June, and their cones extending from June through the following 2011). They have been observed feeding and mature during the following July through May, Type 1 feeds on Red Spruce in June–Au - nesting in state forests where conifer planta - September (Benkman 1987). These soft- gust (McNair 1988, Groth 1993c, M. West - tions are found, including planted stands of coned conifers typically drop much of their phal, pers. obs.), then on Eastern White Pine Norway Spruce, Red Pine, European Larch ( L. seed by November and December except in in October–February, and finally on Loblolly decidua ), Eastern White Pine, and White excellent or “bumper” cone crop years. The (P. taeda ), Shortleaf, Virginia, and Pitch Pine Spruce. Also present in these areas are native semi-soft-coned white pines—Eastern White in March–May, when all soft-coned conifers Eastern White Pine, Eastern Hemlock, and a Pine ( P. strobus ), Western White Pine ( P. mon - have typically dropped their seed (Groth few pockets of native Red Spruce. Norway ticola ), and Limber Pine ( P. flexilis )—form 1993c; Blankenship et al. 2010). Late winter Spruce is perhaps the most widely planted

VOLUME 65 (2011) • NUMBER 3 557 STATUS AND DISTRIBUTION OF TYPE 1 RED CROSSBILL: AN APPALACHIAN CALL TYPE?

Table 3. Recordings of Type 1 Red Crossbills verified by audiospectrographic analysis from locations outside the Appalachians. Discussion Recordings are of one individual unless otherwise noted. Forest compositions differ markedly between eastern and Audiospectro- Location Date Recordist Catalog # graphic Analysis Notes western North America, and crossbill density and diversity Algonquin Provincial (richness) are also quite differ - Park, Ontario 3/20/1959 William W. H. Gunn Macaulay # 58166 Matthew A. Young Type 4 also present ent between the two regions, Manning Provincial 6/2/1962 William W. H. Gunn Macaulay # 58169 Matthew A. Young being lower in the East, higher Park, British Columbia in the more conifer-rich West. , Several Type 10 In the Pacific Northwest, par - 6/12/1962 Robert C. Stein Macaulay # 12984 Matthew A. Young Maine also present ticularly in Washington and Thompson Plateau, Types 2, 3, 4, and 5 British Columbia, conifer diver - British Columbia 8/8/1987 Jeffrey G. Groth Groth #jM635 (MVZ 171360) Jeffrey G. Groth also present sity and density are both high, Thompson Plateau, Types 2, 3, 4, and 5 and this area also has the high - British Columbia 8/8/1987 Jeffrey G. Groth Groth #aF636 (MVZ 171365) Jeffrey G. Groth also present est density and diversity of crossbill Types (Groth 1993c). near Neilton, Several Type 3 also Washington 8/29/1989 Tom Hahn Tom Hahn in area In the East, the central and southern Appalachians have Several Type 10 the highest density and highest Patricks Point, California 12/11/2003 Ken Irwin Ken Irwin also in area diversity of conifers; soft-, Several Type 10 Patricks Point, California 3/16/2004 Ken Irwin Ken Irwin semi-soft-, and hard-coned also in area conifers can be found in close Just north of Patricks Three Type 1 with proximity in this region (Parch - 1/7/2005 Ken Irwin Ken Irwin Point, California several Type 10 man et al. 2006). The Northeast Madawaska, New York Four Type 10 also lacks a diversity of hard-coned 8/10/2008 Matthew A. Young Matthew A. Young in area conifers, with only Red Pine be - About eight Type 1 ing widespread. This lower di - versity may be one reason that Near North Sandwich, Matthew A. Young birds with about New Hampshire 7/31/2009 Robert Ridgely (see Young 2010b) 24 Type 10, about the Northeast appears to sup - eight Type 2, and port no large numbers of a resi - one Type 3 dent Red Crossbill Type; Type About 12 Type 10 10, and to a lesser extent, Type Sevey Corners, New York 3/4/2011 Matthew A. Young Matthew A. Young also present 2, are found in small quasi-resi - About 24 Type 10 dent numbers, with Types 3, 1, East Hancock, Maine 8/4/2011 Doug Hitchcox Matthew A. Young also in area and 4 only occurring occasion - ally. To form quasi-residency, Several Type 3 also Mary’s Peak, Oregon 8/15/2011 Doug Robinson Matthew A. Young in area crossbills need either a single conifer that produces a reliable tree in the area, and unlike native spruces, Northwest, to coastal Sitka Spruce (Benkman cone crop over a large enough area from year to this species holds its seed for as long as 14 1993a) and Western Hemlock forests (Groth year, or an area that has high diversity of months. Seed of Norway Spruce is most ac - 1993c), where they are thought to be rare conifers that provide seed from year to year cessible for the crossbills beginning in Janu - and, thus far, have been confirmed by very (Benkman 1987). In the East, no widespread ary, which coincides with a period when oth - few audio-recordings (Table 3; Irwin 2010, T. species of conifer appears to produce a reliable er seed becomes scarce in the region. Hahn, pers. comm.; C. Benkman, pers. cone crop suitable for a medium-billed crossbill If central and southern Appalachian Red comm.). Cross-continental wanderings are such as Type 1, but the diversity of conifers in Spruce crops are poor at the start the cone documented in other Types, e.g., Types 3 and the central and southern Appalachians appears crop year in June and July, then Type 1 Red 10 have appeared in the Northeast and Penn - to provides sufficient cone crops, in most years, Crossbills are apparently forced to move, and sylvania during irruptions (Groth 1993c, for Type 1 to be resident in this region, though the primary zone of irruption is likely to be Hess et al. 1998, Young 2011). It would be flocks surely wander widely within the Ap - the Northeast, specifically from central Mas - logical for Type 1 to emigrate to the Pacific palachians (Groth 1993c, Blankenship et al. sachusetts and the Adirondacks through On - Northwest, where Sitka Spruce and Western 2010, Young 2011). tario and the southern Maritime provinces, Hemlock provide food resources similar to Dickerman (1987) suggested that a medi - where soft-coned conifers are common. Once Red Spruce and Eastern Hemlock in the Ap - um-billed Red Crossbill subspecies had disap - in the Northeast, they use Red Spruce and palachians, and although there are no returns peared from the Northeast because of the log - White Spruce ( P. glauca ) during their June– of audio-recorded Type 1 Red Crossbills that ging of virgin Eastern White Pine forests in the September nesting period (Benkman 1987; were banded in Appalachia and recovered in late 1800s early 1900s. His review of museum M. Young, pers. obs.). If cone crops of many the Pacific Northwest, for instance, studies in specimens led him to conclude that a “North - species fail widely, which occurs very infre - (Newton 2006) verify that Red Cross - eastern subspecies” had become nearly extinct. quently (Koenig and Knops 2000, Kelsey bills tend to return to natal areas after periods However, there is no acceptable evidence that 2008), Type 1 may wander as far as the Pacific of irruption due to food shortage. such a bird existed, and because Types of Red

558 NORTH AMERICAN BIRDS STATUS AND DISTRIBUTION OF TYPE 1 RED CROSSBILL: AN APPALACHIAN CALL TYPE?

Crossbills are currently distinguished primarily to northern Georgia, in particular those areas Sounds at the Cornell Lab of Ornithology pro - by flight calls, it is not feasible to evaluate Dick - dominated by Red Spruce and in close prox - vided recordings for analysis, and we thank erman’s hypothesis in terms of known Types. In imity to Eastern White Pine, Eastern Hem - Curator Greg Budney at the Macaulay Library the Northeast, three medium-billed Types (1, 4, lock, and hard-coned pines, appear to form for extensive help in the field and in the li - and 10) are known to occur, with Type 10 be - the core zone of occurrence for Type 1 (Figure brary. Finally, we thank Julie Siler for assis - ing the most frequently recorded in this region 4). As Benkman (2007) first noted, Red tance in editing earlier versions of this paper. (Young 2010b). In addition to the Northeast’s Spruce is clearly one of the most important Finally, we thank Julie Siler and P. A. Buckley lack of a diversity of hard-coned conifers, it conifers for Type 1, and so surveys for Type 1 for extensive editorial assistance. may be the case that the occasional presence of should be focused in July and August, when White-winged Crossbill ( L. leucoptera ) and Red Spruce is commonly used for nesting. For Literature cited four other competing crossbill Types in this re - this Appalachian crossbill to survive, it is crit - Adkisson, C. S. 1996. Red Crossbill ( Loxia gion (Young 2011), along with abundant squir - ical that extensive areas of Red Spruce and curvirostra ), in: The Birds of North America rels (cf. Parchman and Benkman 2008), could other key conifers such as Eastern White Pine Online , A. Poole, ed. Cornell Lab of Or - limit Type 1’s ability to make regular use of and Eastern Hemlock be maintained through - nithology, Ithaca, New York. . medium-billed crossbills are only resident in supplemental plantings of these conifers Benkman, C. W. 1987. Food profitability and the wettest areas of the Pacific Northwest, would be useful as well. In the northern edge the foraging ecology of crossbills. Ecologi - where conifers hold their seeds through the of this Type’s core range, central-southern cal Monographs 57: 251-267. winter. The small- to medium-billed Types 3, 4, New York, existing plantings of Norway ——-. 1988. Seed-handling ability, bill struc - and 10 are regularly found in the Pacific North - Spruce, Red Pine, European Larch, and other ture, and the cost of specialization for west; thus, although this region does appear to conifers could be managed to benefit the crossbills. Auk 105: 715-719. provide occasional refuge to Type 1 during ir - crossbills that nest here. Acid rain, logging, ——-. 1989. On the evolution and ecology of ruptions, competition with other crossbill outbreaks of Hemlock Wooly Adelgid ( Adelges island populations of crossbills ( Loxia ). Types would likely limit the ability of Type 1 to tsugae ) (Tingley et al. 2002), and global warm - Evolution 43: 1324-1330. establish a stable population in this region. ing, which alter forest composition, could ——-. 1993a. Adaptation to single resources Based on all verified records of Type 1, as well threaten the existence of Type 1 Red Crossbill, and the evolution of crossbill ( Loxia ) di - as the distribution of conifers known to be used which may prove, with further study, to be a versity. Ecological Monographs 63: 305-325. by Type 1 in areas with records, we propose an distinct species. Observers interested in docu - ——-. 1993b. The evolution, ecology, and de - approximate range map that illustrates the core menting Red Crossbill Types anywhere are en - cline of the Red Crossbill of Newfound - zone of this species’ distribution along with couraged to study Groth’s (1993a) descrip - land. American Birds 47: 225-229. what appear to be primary and secondary tions of calls and Young’s (2008a) primer on ——-. 1994. Comments on the ecology and zones of irruption (Figure 4). vocalizations, and to correspond with status of the Hispaniolan Crossbill ( Loxia Matthew Young. It is worthwhile to publish leucoptera megaplaga ), with recommenda - Conclusions any analyzed crossbill recordings in state and tions for its conservation. Caribbean Jour - Groth’s (1993c) original contention that Type provincial ornithological journals. nal of Science 30: 250-254. 1 is an Appalachian crossbill appears to be Red Crossbills represent an ecological puz - ——-. 1999. The selection mosaic and diver - supportable. The older literature points to the zle for biologists and birders alike, an oppor - sifying co-evolution between crossbills and existence of a distinct quasi-resident Red tunity for pioneering field work for those in - lodgepole pine. American Naturalist 153: Crossbill in the central and southern Ap - clined to explore some of North America’s lit - 75-91. palachians, and in the past 25 years, the vast tle-birded montane habitats. Every recording ——-. 2007. Red crossbill types in Colorado: majority of Type 1 sightings and audio record - adds an important piece to the puzzle, espe - their ecology, evolution, and distribution. ings have come from this area, but there are cially when accompanied by notes on behav - Colorado Birds 41: 153-163. very few recordings from other parts of the ior and ecology, including tree species used Benkman, C. W., and T. L. Parchman. 2009. continent, and those were apparently made for foraging and nesting. The conservation of between crossbills and black only during irruption years, probably when these attractive finches will depend in large pine: the importance of competitors, forest cone crops of multiple species failed in the measure on our understanding their complex area, and resource stability. Journal of Evo - Appalachians. In the conifer-rich central and distributions and ecological associations, and lutionary Biology 22: 942-953 southern Appalachians, Type 1 would experi - birders can make critical contributions to Benkman C. W., J. W. Smith, P. C. Keenan, T. ence relatively little competition with other their conservation by recording crossbill calls L. Parchman, and L. Santisteban. 2009. A Types, except at the northern edge of their and by reporting their findings. new species of Red Crossbill (Fringillidae: core zone of occurrence, where other Types Loxia ) from Idaho. Condor 111: 169-176. are more frequent during irruptions. Finally, Acknowledgments Benkman C. W., T. L. Parchman, and E. T. genetic studies (Groth 1993c, Parchman et al. We thank Craig Benkman, Jeffrey Groth, Tom Mezquida. 2010. Patterns of co-evolution 2006) support the distinctiveness of Type 1: Hahn, Ken Irwin, and Nathan Pieplow for dis - in the adaptive radiation of crossbills. An - of the three distinct clusters of crossbill Types, cussions of the Red Crossbill complex. Jeffrey nals of the New York Academy of Sciences two clusters have core breeding areas in the Groth has provided extensive assistance with 1206: 1-16. West, and the other cluster includes only analysis of flight calls, and Craig Benkman has Blankenship, K., M. A. Young, and S. Holtz - Type 1. Parchman et al. (2006) found Type 1 supplied the authors with extensive details man. 2010. Red Crossbills: breeding to be most genetically distinct. about the natural history of the various Red records and call types in Georgia and the The Appalachians from southern New York Crossbill Types. The Macaulay Library of southeastern United States. Oriole 74: 1-14.

VOLUME 65 (2011) • NUMBER 3 559 STATUS AND DISTRIBUTION OF TYPE 1 RED CROSSBILL: AN APPALACHIAN CALL TYPE?

Dickerman, R. W. 1987. The “old northeast - the Red Crossbill. Western Birds 41: 10-25. Parchman, T. L., and C. W. Benkman. 2008. ern” subspecies of Red Crossbill. American Kelsey, T. R. 2008. Biogeography, foraging ecol - The geographic selection mosaic for pon - Birds 41: 188-194. ogy, and population dynamics of Red Cross - derosa pine and crossbills: a tale of two Evans, W. R., and M. O’Brien. 2002. Flight bills in North America . Doctoral disserta - squirrels. Evolution 62: 348-360. Calls of Migratory Birds: Eastern North tion. University of California, Davis. Pitts, I., Jr. 1988. Red Crossbill nesting at Cae - American Landbirds . Old Bird, Inc., Tru - Koenig, W. D., and J. M. H. Knops. 2000. Pat - sar’s Head, South Carolina. Chat 52(3): 55- mansburg, New York. terns of annual seed production by north - 56. Griscom, L. 1937. A monographic study of ern hemisphere trees: a global perspective. Sewall, K. B. 2009. Limited adult vocal learn - the Red Crossbill. Proceedings of the Boston American Naturalist 155: 59-69. ing maintains call dialects but permits pair- Society of Natural History 41: 77-210. Knox, A. G. 1992. Species and pseudospecies: distinctive calls in red crossbills. Groth, J. G. 1988. Resolution of cryptic the structure of crossbill populations. Bio - Behavior 77: 1303-1311. species in Appalachian Red Crossbills. logical Journal of the Linnaean Society 47: ——-. 2011. Early learning of discrete call Condor 90: 745-760. 325-335. variants in red crossbills: implications for ——-. 1993a. Crossbill diagnosis page. McNair, D. B. 1988. Review of breeding reliable signaling. Behavioral Ecology and . the southern Appalachian Mountains and Sewall, K. B., and T. P. Hahn. 2009. Social ex - ——-. 1993b. Call matching and positive as - adjacent regions. The Migrant 59: 105-112. perience modifies behavioural responsive - sortative mating in Red Crossbills. Auk Newton, I. 2006. Movement patterns of Com - ness to a preferred vocal signal in red cross - 110: 398-401. mon Crossbill Loxia curvirostra in Europe. bills, Loxia curvirostra . Animal Behaviour ——-. 1993c. Evolutionary differentiation in Ibis 148: 782-788. 77: 123-128. morphology, vocalizations, and allozymes Oberle, M. W., and D. M. Forsythe. 1995. Pos - Simpson, M. B. 1974. Red Crossbill observa - among nomadic sibling species in the sible breeding by Red-breasted Nuthatch tions in western North Carolina. Chat 51: North American Red Crossbill ( Loxia curvi - and Golden-crowned Kinglet in Georgia 60-61. rostra ) complex. University of California and South Carolina. Oriole 60: 52-55. Scott, F. R. 1981. Evidence for a resident cross - Publications in Zoology 127: 1-143. Parchman, T. L., C. W. Benkman, and S. C. bill population on Shenandoah Mountain. Hess, P., M. R. Leahy, and R. M. Ross. 1998. Britch. 2006. Patterns of genetic variation Raven 52: 61-64. Pennsylvania’s crossbill winter of 1997-98. in the adaptive radiation of New World Stupka, A. 1963. Notes on the Birds of the Great Pennsylvania Birds 12: 2-6. crossbills (Aves: Loxia ). Molecular Ecology Smoky Mountains National Park. University Irwin, K. 2010. A new and cryptic call type of 15: 1873-1887. of Tennessee Press, Knoxville, Tennessee. Summers, R. W., R. J. G. Dawson, and R. E. Phillips. 2007. Assortative mating and pat - terns of inheritance indicate that the three 7th ANNUAL crossbill taxa in are species. Jour - nal of Avian Biology 38: 153-162. Tingley, M. W., D. A. Orwig, R. Field, and G. Eagle Expo Motzkin. 2002. Avian response to removal of a forest dominant: consequences of hem - lock woolly adelgid infestations. Journal of Feb. 99–11,–11, 22012012 Biogeography 29: 1505-1516. Morgan City, La. Young, M. A. 2008a. Introduction to differ - ences in crossbill vocalizations. . ——-. 2008b. Red Crossbill ( Loxia curviros - tra ), pp. 612-613 in: McGowan, K. J., and Boat tours to view eagle nests K. J. Corwin, eds. The Second Atlas of Breed - ing Birds in New York State . Cornell Univer - Presentations by wildlife sity Press, Ithaca, New York. professionals and photographers ——-. 2010a. Type 5 Red Crossbill in New York: first confirmation east of the Rocky Reception, photography Mountains. North American Birds 64: 343- workshop and much more 346. ——-. 2010b. First documented occurrences of Red Crossbill Types 1, 2, 3, and 10 in The America’s Wetland Birding New Hampshire. New Hampshire Bird Trail on the Cajun Coast offers Records 4: 43-50. an opportunity to add over ——-. 2011. Red Crossbill ( Loxia curvirostra ) 300 species to your life list. Call Types of New York: a closer look at For more birding information, their , flight call vocalizations, (800) 256-2931 call us or visit our website. www.cajuncoast.com and ecology. Kingbird 61: 106-123. n

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