Red Crossbill

Loxia curvirostra



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1. Griscom, L. (1937a). A monographic study of the Red Crossbill. Proc. Boston Soc. Nat. Hist. 41:77-210.

2. Groth, J. G. (1988). Resolution of cryptic species of Appalachian Red Crossbills. Condor 90:745-760.

3. Groth, J. G. (1993b). Evolutionary differentiation in morphology, vocalizations, and allozymes among nomadic sibling species in the North American Red Crossbill (Loxia curvirostra) complex. Univ. Calif. Publ. Zool. 127:1-143.

4. Benkman, C. W. (1993a). Adaptation to single resources and the evolution of crossbill (Loxia) diversity. Ecological Monographs 63:305–325.

5. Benkman, C. W. (2003). Divergent selection drives the adaptive radiation of crossbills. Evolution 57 (5):1176-1181.

6. Irwin, K. (2010). A new and cryptic call type of the Red Crossbill. Western Birds 41 (1):10-25.

7. Benkman, C. W., and R. E. Miller (1996). Morphological evolution in response to fluctuating selection. Evolution 50:2499–2504.

8. Benkman, C. W., W. C. Holimon, and J. W. Smith. (2001). The influence of a competitor on the geographic mosaic of coevolution between crossbills and lodgepole pine. Evolution 55 (2):282-294.

9. Young, M., and T. Spahr. (2017). Crossbills of North America: Species and Red Crossbill call types. eBird [online].

10. Parchman, T. L., C. A. Buerkle, V. Soria-Carrasco, and C. W. Benkman (2016). Genome divergence and diversification within a geographic mosaic of coevolution. Molecular Ecology 25:5705–5718.

11. Smith, J. W., and C. W. Benkman (2007). A coevolutionary arms race causes ecological speciation in crossbills. American Naturalist 169 (4):455-465. doi: 10.1086/511961.

12. Benkman, C. W., J. W. Smith, P. C. Keenan, T. L. Parchman, and L. Santisteban. (2009). A new species of the Red Crossbill (Fringillidae: Loxia) from Idaho. The Condor 111:169-176.

13. Young, M., and T. Spahr (2017). Crossbills of North America: species and Red Crossbill call types. eBird. [online] URL: http://

14. Benkman, C. W. (1987a). Crossbill foraging behavior, bill structure, and patterns of food profitability. Wilson Bulletin 99:351-368.

15. Young, M. A., D. A. Fifield, and W. A. Montevecchi (2012). New evidence in support of a distinctive Red Crossbill (Loxia curvirostra) type in Newfoundland. North American Birds 66:29-33.

16. Koenig, W. D. and J. M. H. Knops. (2001). Seed-crop size and eruptions of North American boreal seed-eating birds. Journal of Animal Ecology 70 (4):609-620.

17. Benkman, C. W. (1990). Foraging rates and the timing of crossbill reproduction. Auk 107:376-386.

18. Hahn, T. P. (1995). Integration of photoperiodic and food cues to time changes in reproductive physiology by an opportunistic breeder, the Red Crossbill, Loxia curvirostra (Aves: Carduelinae). J. Exp. Zool. 272:213-226.

19. Hahn, T. P. (1998). Reproductive seasonality in an opportunistic breeder, the Red Crossbill, Loxia curvirostra. Ecology 79 (7):2365-2375. doi: 10.1890/0012-9658(1998)079[2365:rsiaob];2.[2365:rsiaob];2

20. Santisteban, L., C. W. Benkman, T. Fetz, and J. W. Smith (2012). Survival and population size of a resident bird species are declining as temperature increases. Journal of Animal Ecology 81 (2):352-363. doi: 10.1111/j.1365-2656.2011.01918.x.

21. Smith, J. W., C. W. Benkman and K. Coffey. (1999c). The use and misuse of public information by foraging Red Crossbills. Behavioral Ecology 10 (1):54-62.

22. Smith, J. W., S. M. Sjoberg, M. C. Mueller, and C. W. Benkman (2012). Assortative flocking in crossbills and implications for ecological speciation. Proceedings of the Royal Society B-Biological Sciences 279 (1745):4223-4229. doi: 10.1098/rspb.2012.1500.

23. Snowberg, L. K., and C. W. Benkman (2007). The role of marker traits in the assortative mating within Red Crossbills, Loxia curvirostra complex. Journal of Evolutionary Biology 20 (5):1924-1932. doi: 10.1111/j.1420-9101.2007.01372.x.

24. Sewall, K. B. (2009). Limited adult vocal learning maintains call dialects but permits pair-distinctive calls in Red Crossbills. Animal Behavior 77:1303-1311.

25. Parchman, T. L., P. Edelaar, K. Uckele, E. T. Mezquida, D. Alonso, J. P. Jahner, R. W. Summers, and C. W. Benkman. (2018). Resource stability and geographic isolation are associated with genome divergence in western Palearctic crossbills. Journal of Evolutionary Biology 31:1715-1731.

26. Phillips, A. R. (1977). Sex and age determination of Red Crossbills (Loxia curvirostra). Bird-Banding 48 (2):110-117.

27. Clement, P., A. Harris and J. Davis. (1993a). Finches and sparrows. Princeton, NJ: Princeton Univ. Press.

28. Dwight, J., Jr. (1900). The sequence of plumages and moults of the passerine birds of New York. Annals of the New York Academy of Sciences 13:73-360.

29. Ridgway, R. (1901). The birds of North and Middle America, Part 1. Bulletin of the United States National Museum 50.

30. Tordoff, H. B. (1952b). Notes on plumages, molts, and age variation in the Red Crossbill. Condor 55:200-203.

31. Oberholser, H. C. (1974). The Bird Life of Texas. University of Texas Press, Austin, TX, USA.

32. Cramp, S. and C. M. Perrins. (1994a). The birds of the Western Palearctic, Vol. 8: Crows to finches. Oxford: Oxford Univ. Press.

33. Svensson, L. (1992). Identification Guide to European Passerines. British Trust for Ornithology, Beech Grove, United Kingdom.

34. Jenni, L., and R. Winkler. (1994). Moult and Ageing of European Passerines. Academic Press, New York, NY, USA.

35. Pyle, P. (1997). Identification Guide to North American Birds, Part I: Columbidae to Ploceidae. Slate Creek Press, Bolinas, CA, USA.

36. Humphrey, P. S., and K. C. Parkes (1959). An approach to the study of molts and plumages. Auk 76:1-31.

37. Howell, S. N. G., C. Corben, P. Pyle, and D. I. Rogers (2003). The first basic problem: A review of molt and plumage homologies. Condor 105:635-653.

38. Howell, S. N. G. (2010). Peterson Reference Guide to Molt in North American Birds. Houghton Mifflin Harcourt Company, Boston, MA, USA.

39. Howell, S. N. G., C. Corben, P. Pyle, and D. I. Rogers (2004). The first basic problem revisited: Reply to commentaries on Howell et al. (2003). Condor 106:206-210.

40. Stone, W. (1896). The molting of birds with special reference to the plumages of smaller land birds of eastern North America. Proceedings of the Academy of Natural Sciences of Philadelphia 43:108-164.

41. Pyle, P. (1997). Molt limits in North American passerines. North American Bird Bander 22:49-89.

42. Herremans, M. (1988). Measurements and moults of irruptive Common Crossbills (Loxia curvirostra curvirostra) in central Belgium. Gerfaut 78:243-260.

43. Tordoff, H. B. (1954a). Further notes on plumages and molts of Red Crossbills. Condor 56:108-109.

44. Cornelius, J. M., N. Perfito, R. Zann, C. W. Breuner, and T. P. Hahn. (2011). Physiological trade-offs in self-maintenance: plumage molt and stress physiology in birds. Journal of Experimental Biology 214:2768-2777.

45. Benkman, C. W. (1988a). A 3:1 ratio of mandible crossing direction in White-winged Crossbills. Auk 105:578-579.

46. Benkman, C. W. (1996). Are the ratios of bill crossing morphs in crossbills the result of frequency-dependent selection? Evolutionary Ecology 10 (1):119-126. doi: 10.1007/bf01239352.

47. Newton, I. (1972). Finches. Engelwood-Cliffs, NJ: Taplinger.

48. Parchman, T. L., and C. W. Benkman (2002). Diversifying coevolution between crossbills and black spruce on Newfoundland. Evolution 56 (8):1663-1672.

49. Summers, R. W., R. J. G. Dawson, and R. E. Phillips (2007). Assortative mating and patterns of inheritance indicate that the three crossbill taxa in Scotland are species. Journal of Avian Biology 38 (2):153-162. doi: 10.1111/j.2007.0908-8857.03798.x.

50. Marquiss, M., and R. Rae. (2002) Ecological differentiation in relation to bill size amongst sympatric, genetically undifferentiated crossbills Loxia spp. Ibis 144:494–508. 

51. Cornelius, J. M., and T. P. Hahn (2012). Seasonal pre-migratory fattening and increased activity in a nomadic and irruptive migrant, the Red Crossbill Loxia curvirostra. Ibis 154 (4):693-702. doi: 10.1111/j.1474-919X.2012.01266.x.

52. Benkman, C. W. (2012). White-winged Crossbill (Loxia leucoptera), version 2.0. In The Birds of North America (A. Poole, Editor), Cornell Lab of Ornithology, Ithaca, New York, USA.

53. Monson, G. and A. R. Phillips. (1981b). "The races of the Red Crossbill, Loxia curvirostra, in Arizona." In Annotated checklist of the birds of Arizona, 223-230. Tucson: Univ. of Arizona Press.

54. Payne, R. B. (1987). Populations and type specimens of a nomadic bird: comments on the North American crossbills Loxia pusilla Gloger 1834 and Crucirostra minor Brehm 1845 Occas. Pap. Mus. Zool. Univ. Mich. 714.

55. Dickerman, R. W. (1986b). A review of the Red Crossbill in New York State, Part 1. Historical and nomenclatural background. Kingbird 36:73-78.

56. Dickerman, R. W. (1986a). A review of the Red Crossbill in New York State Part 2. Identification of specimens from New York. Kingbird 36:127-134.

57. Benkman, C. W. (1999). The selection mosaic and diversifying coevolution between crossbills and lodgepole pine. American Naturalist 153:S75-S91. doi: 10.1086/303213.

58. Groth, J. G. (1993a). Call matching and positive assortative mating in Red Crossbills. Auk 110:398-401.

59. Robb, M. S. (2000). Introduction to vocalizations of crossbills in north-western Europe. Dutch Birding 22 (2):61-107.

60. Summers, R. W., D. C. Jardine, M. Marquiss, and R. Rae (2002). The distribution and habitats of crossbills Loxia spp. in Britain, with special reference to the Scottish Crossbill Loxia scotica. Ibis 144 (3):393-410. doi: 10.1046/j.1474-919X.2002.00064.x.

61. Förschler, M. I., and E. K. V. Kalko (2009). Vocal types in crossbill populations (Loxia spp.) of southwest Europe. Journal of Ornithology 150 (1):17-27. doi: 10.1007/s10336-008-0312-z.

62. Edelaar, P., M. Robb, K. van Eerde, K. Terpstra, R. Bijlsma, and E. Maassen (2004). Are there several species of 'Common' Crosbill Loxia curvirostra in the Netherlands? Limosa 77 (1):31-38.

63. Edelaar, P., K. van Eerde, and K. Terpstra (2008). Is the nominate subspecies of the common crossbill Loxia c. curvirostra polytypic? II. Differentiation among vocal types in functional traits. Journal of Avian Biology 39 (1):108-115. doi: 10.1111/j.0908-8857.2008.04230.x.

64. Piertney, S. B., R. Summers, and M. Marquiss (2001). Microsatellite and mitochondrial DNA homogeneity among phenotypically diverse crossbill taxa in the UK. Proceedings of the Royal Society B-Biological Sciences 268 (1475):1511-1517. doi: 10.1098/rspb.2001.1015.

65. Benkman, C. W. (1989c). On the evolution and ecology of island populations of crossbills. Evolution 43:1324-1330.

66. Tremblay, J., M. Robert, D. Hynes, M. Young, and B. Drolet (2018). Range extension of the threatened Red Crossbill (Loxia curvirostra percna) in Canada: New insights from Anticosti Island, Québec. Avian Conservation and Ecology 13(1):10.

67. Howell, T. R. (1972). Birds of the lowland pine savanna of northeastern Nicaragua. Condor 74:316-340.

68. Browning, M. R. (1990). Taxa of North American birds described from 1957 to 1987. Proc. Biol. Soc. Wash. 103:432-451.

69. Edelaar, P., D. Alonso, S. Lagerveld, J. C. Senar, and M. Björklund (2012). Population differentiation and restricted gene flow in Spanish crossbills: not isolation-by-distance but isolation-by-ecology. Journal of Evolutionary Biology 25: 417–430.

70. Hall, K. S. S. (2005). Do nine-primaried passerines have nine or ten primary feathers? The evolution of a concept. Journal of Ornithology 146 (2):121-126.

71. American Ornithologists' Union (1998). Check-list of North American Birds, 7th edition. American Ornithologists' Union, Washington, DC, USA.

72. Ponge, J.-F., D. Zuccon, M. Elias, S. Pavoine, P.-Y. Henry, M. Théry, and É. Guilbert (2017). Ancestrality and evolution of trait syndromes in finches (Fringillidae). Ecology and Evolution 7 (23):9935-9953. doi: 10.1002/ece3.3420.

73. Knox, A. G. (1990b). The sympatric breeding of Common and Scottish Crossbills Loxia curvirostra and L. scotica and the evolution of crossbills. Ibis 132:454-466.

74. Edelaar, P. (2008). Assortative mating also indicates that Common Crossbill Loxia curvirostra vocal types are species. Journal of Avian Biology 39 (1):9-12. doi: 10.1111/j.0908-8857.2008.04231.x.

75. Tallman, D. A. and R. L. Zusi. (1984). A hybrid Red Crossbill-Pine Siskin (Loxia curvirostrax Carduelis pinus) and speculations on the evolution of Loxia. Auk 101:155-157.

76. Davidson, P. J. A., R. J. Cannings, A. R. Couturier, D. Lepage and C. M. Di Corrado (Editors). (2015). The atlas of the breeding birds of British Columbia, 2008-2012. Bird Studies Canada, Delta, British Columbia.

77. Sinclair, P. H., W. A. Nixon, C. D. Eckert, and N. L. Hughes (2003). Birds of the Yukon Territory. UBC Press, Vancouver, BC, Canada.

78. Godfrey, W. E. (1986). The Birds of Canada. Revised Edition. National Museums of Canada, Ottawa, ON, Canada.

79. Federation of Alberta Naturalists (2007). Atlas of breeding birds of Alberta: A second look. Federation of Alberta Naturalists, Edmonton, AB, Canada.

80. Smith, A. R. (1996). Atlas of Saskatchewan Birds. Saskatchewan Natural History Society Special Publications no. 22, Regina, Saskatchewan, Canada.

81. Cadman, M. D., D. A. Sutherland, G. G. Beck, D. Lepage, and A. R. Couturier. (2007). Atlas of the Breeding Birds of Ontario 2001-2005. Bird Studies Canada, Environment Canada, Ontario Field Ornithologists, Ontario Ministry of Natural Resources, and Ontario Nature, Toronto, Ontario, Canada.

82. Cyr, A. and J. Larivee. (1995a). Atlas saisonnier des oiseaux du Quebec. Sherbrooke, Canada: Les Presses de l'Univ. de Sherbrook et la Societe de Loisir Ornithologique de l'Estrie, Inc.

83. Benkman, C. W. (1993c). The evolution, ecology, and decline of the Red Crossbill of Newfoundland. Am. Birds 47:225-229.

84. Benkman, C. W., A. M. Siepielski, and T. L. Parchman (2008). The local introduction of strongly interacting species and the loss of geographic variation in species and species interactions. Molecular Ecology 17 (1):395-404. doi: 10.1111/j.1365-294X.2007.03368.x.

85. Stewart, R. L. M., K. A. Bredin, A. R. Couturier, A. G. Horn, D. Lepage, S. Makepeace, P. D. Taylor, M. Villard and R. M. Whittam. (2015). Second atlas of breeding birds of the Maritime Provinces: Environment Canada, Natural History Society of Prince Edward Island, Nature New Brunswick, New Brunswick Department of Natural Resources, Nova Scotia Bird Society, Nova Scotia Department of Natural Resources, Prince Edward Island Department of Agriculture and Forestry.

86. Gilligan, J., D. Rogers, M. Smith, and A. Contreras. (1994). Birds of Oregon: Status and Distribution. Cinclus Publications, McMinnville, OR, USA

87. Wahl, T. R., B. Tweit and S. G. Mlodinow. (2005). Birds of Washington: Status and Distribution. Oregon State University Press, Corvallis, OR, USA.

88. Szeliga, W., L. Benner, J. Garrett, and K. Ellsworth. (2014). Call types of the Red Crossbill in the San Gabriel, San Bernardino, and San Jacinto Mountains, southern California. Western Birds 45:213-223.

89. Payne, R. B. (1972). Nuts, bones, and a nesting of Red Crossbills in the Panamint Mountains, California. Condor 74:485-486.

90. Small, A. (1994). California Birds: Their Status and Distribution. Ibis Publishing Company, Vista, CA, USA.

91. Alcorn, J. R. (1988). The birds of Nevada. Fallon, NV: Fairview West Publ.

92. Floyd, T., C. S. Elphick, G. Chisholm, K. Mack, R. G. Elston, E. M. Ammon, and J. D. Boone (2007). Atlas of the Breeding Birds of Nevada. University of Nevada Press, Reno and Las Vegas, NV, USA.

93. Marks, J. S., P. Hendricks, and D. Casey (2016). Birds of Montana. Buteo Books, Arrington, VA, USA.

94. Faulkner, D. W. (2010). Birds of Wyoming. Roberts and Co, Greenwood Village, CO, USA.

95. Wickersham, L. E. (2016). The Second Colorado Breeding Bird Atlas. Colorado Bird Atlas Partnership and Colorado Parks and Wildlife, Denver, CO.

96. Corman, T. E., and C. Wise-Gervais. (2005). Arizona Breeding Bird Atlas. University of New Mexico Press, Albuquerque, NM, USA.

97. Hubbard, J. P. (1978). Revised check-list of the birds of New Mexico. New Mexico Ornithological Society Publication 6.

98. Lockwood, M. W. and B. Freeman. (2014). The TOS handbook of Texas birds. College Station, TX: Texas A&M University Press.

99. South Dakota Ornithologists' Union (1991). The birds of South Dakota, 2nd edition. South Dakota Ornithologists' Union, Aberdeen, SD, USA.

100. Mollhoff, W. J. (2016). The Second Nebraska Breeding Bird Atlas. Miscellaneous Publication No. 122 of the University of Nebraska-Lincoln and Bulletin of the University of Nebraska State Museum, Volume 29, Lincoln, NE, USA.

101. Janssen, R. B. (1987). Birds in Minnesota. University of Minnesota Press, Minneapolis, MN, USA.

102. Cutright, N. J., B. R. Harriman and R. W. Howe. (2006). Atlas of the breeding birds of Wisconsin. Waukesha, WI: Wisconsin Society of Ornithology.

103. Chartier, A. T., J. J. Baldy, and J. M. Brenneman (Editors) (2011). The Second Michigan Breeding Bird Atlas. Kalamazoo Nature Center, Kalamazoo, Michigan, USA.

104. McGowan, K. J. and K. Corwin. (2008). The second atlas of breeding birds in New York State. Ithaca: Comstock/Cornell University Press.

105. Renfrew, R. B. (2013). The Second Atlas of Breeding Birds of Vermont. University Press of New England, Lebanon, NH, USA.

106. Foss, C. R. (1994a). Atlas of breeding birds in New Hampshire. Dover, NH: Audubon Soc. of New Hampshire/Arcadia.

107. Young, M. A. 2010b. First documented occurrences of Red Crossbill Types 1, 2, 3, and 10 in New Hampshire. New Hampshire Bird Records 4:43-50.

108. Adamus, P. R. (1987). Atlas of Breeding Birds in Maine, 1978-1983. Augusta: Maine Dept. Inland Fisheries and Wildlife.

109. Young, M. A., K. Blankenship, M. Westphal, and S. Holzman. 2011. Status and distribution of Type 1 Red Crossbill (Loxia curvirostra): an Appalachian Call Type? North American Birds 65: 554-561.

110. Boyle, W. J., Jr. (2011). The birds of New Jersey: Status and distribution. Princeton, NJ: Princeton University.

111. Warren, R. C., J. A. Jackson and T. L. Darden. (1977). Nesting of the Red Crossbill in Mississippi. Am . Birds 31:1100.

112. Nicholson, C. P. (1997). Atlas of the Breeding Birds of Tennessee. University Press of Tennessee, Knoxville, TN, USA.

113. James, D. A., and J. C. Neal. (1986). Arkansas Birds: Their Distribution and Abundance. University of Arkansas Press, Fayetteville, AR, USA.

114. Howell, S. N. G., and S. Webb (1995). A Guide to the Birds of Mexico and Northern Central America. Oxford University Press, NY, USA.

115. Jones, H. L., and D. Gardner. (2004). Birds of Belize. University of Texas Press, Austin, TX, USA.

116. Monroe, B. L., Jr. (1968). A distributional survey of the birds of Honduras. Ornithological Monographs 7.

117. Gallardo, R. J. (2014). Guide to the Birds of Honduras. Mountain Gem Tours, Tegucigalpa, Honduras.

118. Chavarría-Duriaux, L., D. C. Hill, and R. Dean (2018). Birds of Nicaragua: A Field Guide. Cornell University Press, Ithaca, NY, USA.

119. Mezquida, E. T., J.-C. Svenning, R. W. Summers, and C. W. Benkman (2018). Higher spring temperatures increase food scarcity and limit the current and future distributions of crossbills. Diversity and Distributions 24 (4):473-484. doi: 10.1111/ddi.12694.

120. Young, M. A. (2011). Red Crossbill (Loxia curvirostra) Call Types of New York: a closer look at their taxonomy, flight call vocalizations, and ecology. Kingbird 61:106-123.

121. Young M. A. 2010. Type 5 Red Crossbill (Loxia curvisrotra) in New York: first confirmation east of the Rocky Mountains. North American Birds 64:343-346.

122. Stevenson, H. M., and B. H. Anderson (1994). The Birdlife of Florida. University Press of Florida, Gainesville, FL, USA.

123. Formozov, A. N. (1960). La production de graines dans les forêts de conifères de la taiga de l'URSS et l'envahissement de l'Europe occidentale par certaines espèces d'oiseaux. Proceedings of the International Ornithological Congress 12:216-229.

124. Herremans, M. (1982). Notes on measurements and moult of irruptive Red Crossbills (Loxia curvirostra curvirostra) in central Belgium. Le Gerfaut 72:243-254.

125. Voous, K. H. (1978). The Scottish crossbill Loxia scotica. Br. Birds 71:3-10.

126. Newton, I. (1970). "Irruptions of crossbills in Europe." In Animal populations in relation to their food resources., edited by A. Watson, 337-357. Oxford, England: Blackwell Scientific.

127. Newton, I. (2006). Movement patterns of Common Crossbills Loxia curvirostra in Europe. Ibis 148 (4):782-788. doi: 10.1111/j.1474-919X.2006.00585.x.

128. Benkman, C. W. (2017). Matching habitat choice in nomadic crossbills appears most pronounced when food is most limiting. Evolution 71 (3):778-785. doi: 10.1111/evo.13146.

129. Senar, J. C., A. Borras, T. Cabrera, and J. Cabrera (1993). Testing for the relationship between coniferous crop stability and Common Crossbill residence. Journal of Field Ornithology 64 (4):464-469.

130. Benkman, C. W., T. Fetz, and M. V. Talluto (2012). Variable resource availability when resource replenishment is constant: The coupling of predators and prey. Auk 129 (1):115-123. doi: 10.1525/auk.2011.11069.

131. Benkman, C. W. (1987b). Food profitability and the foraging ecology of crossbills. Ecol. Monogr. 57:251-267.

132. Austin, Jr., O. L. (1968c). Life histories of North American cardinals, grosbeaks, buntings, towhees, finches, sparrows, and allies. U.S. Natl. Mus. Bull. 237, Pt. 2.

133. Peterjohn, B. G. (2001). The Birds of Ohio with Breeding Bird Atlas Maps. Wooster Book Company, Wooster, OH, USA.

134. Robbins, M. B. (2018). The Status and Distribution of Birds in Missouri. University of Kansas Libraries, Lawrence, KS, USA.

135. Benkman, C. W. (1992a). Newfoundland crossbill's twist of fate. Nat. Hist. 101:39-42.

136. Cornelius, J. M., C. W. Breuner, and T. P. Hahn (2010). Under a neighbour's influence: Public information affects stress hormones and behaviour of a songbird. Proceedings of the Royal Society B-Biological Sciences 277 (1692):2399-2404. doi: 10.1098/rspb.2010.0164.

137. Cornelius, J. M., G. Perreau, V. R. Bishop, J. S. Krause, R. Smith, T. P. Hahn, and S. L. Meddle (2018). Social information changes stress hormone receptor expression in the songbird brain. Hormones and Behavior 97:31-38. doi: 10.1016/j.yhbeh.2017.10.002.

138. Summers, R. W., D. C. Jardine, M. Marquiss, and R. Proctor (1996). The biometrics of invading Common Crossbills Loxia curvirostra in Britain during 1990-1991. Ringing and Migration 17 (1):1-10.

139. Alonso, D., and J. Arizaga (2011). Seasonal patterns of breeding, moulting, and body mass variation in Pyrenean Common Crossbills Loxia curvirostra curvirostra. Ringing & Migration 26 (1):64-70.

140. Benkman, C. W. (2007). Red Crossbill types in Colorado: Their ecology, evolution, and distribution. Colorado Birds 41 (3):153-163.

141. Young, M. A. (2011). Status and distribution of Type 1 Red Crossbill (Loxia curvirostra): an Appalachian Call Type? North American Birds 65:554-561.

142. Adkisson, C. S. (1996). Red Crossbill (Loxia curvirostra), version 2.0. In The Birds of North America (A. Poole, Editor), Cornell Lab of Ornithology, Ithaca, New York, USA.

143. Benkman, C. W. (2016). The natural history of the South Hills Crossbill in relation to its impending extinction. The American Naturalist 188:589-601.

144. Parchman, T. L., and C. W. Benkman. (2008). The geographic selection mosaic for ponderosa pine and crossbills: a tale of two squirrels. Evolution 62:348-360.

145. Perry, Jr., J. P. (1991). The pines of Mexico and Central America. Portland, OR: Timber Press.

146. Lundelius, E. L., R. W. Graham, E. Anderson, J. Guilday, J. A. Holman, D. W. Steadman and S. D. Webb. (1983). "Terrestrial vertebrate faunas." In Late-Quaternary environments of the United States. Vol. 1: The late Pleistocene., edited by S. C. Porter, 311-353. Minneapolis: Univ. of Minnesota Press.

147. Steadman, D. W., and J. Franklin (2017). Origin, paleoecology, and extirpation of bluebirds and crossbills in the Bahamas across the last glacial-interglacial transition. Proceedings of the National Academy of Sciences of the United States of America 114 (37):9924-9929. doi: 10.1073/pnas.1707660114.

148. Benkman, C. W. (2017). Crossbills were unlikely resident in the Bahamas; thus, there was no population to be extirpated. Proceedings of the National Academy of Sciences of the United States of America 114 (47):E10031-E10032. doi: 10.1073/pnas.1716928114.

149. Nethersole-Thompson, D. (1975). Pine crossbills: a Scottish contribution. Berkhamsted: T. & A. D. Poyer.

150. Johnson, N. K. (1958). Notes on the Red Crossbill in Nevada. Condor 60:136-138.

151. Knox, A. G. (1983). Handedness in crossbills Loxia and the Akepa Loxops coccinea. Bull. Brit. Ornithol. Cl. 103:114-118.

152. Tordoff, H. B. (1954b). Social organization and behavior in a flock of captive, nonbreeding Red Crossbills. Condor 56:346-358.

153. Benkman, C. W. and A. K. Lindholm. (1991). The advantages and evolution of a morphological novelty. Nature 349:519-520.

154. Fowells, H. A. (1965). Silvics of forest trees in the United States. Wash., D.C: Agricultural Handbook Number 271, U.S.D.A.

155. Kelsey, T. R. (2008). Biogeography, foraging ecology and population dynamics of Red Crossbills in North America. Ph.D. dissertation, University of California, Davis, CA.

156. Benkman, C. W., and T. L. Parchman. (2013). When directional selection reduces geographic variation in traits mediating species interactions. Ecology and Evolution 3 (4):961-970. doi: 10.1002/ece3.518.

157. Benkman, C. W., T. L. Parchman and E. T. Mezquida. (2010). Patterns of coevolution in the adaptive radiation of crossbills. Annals of the New York Academy of Sciences 1206:1-16.

158. Benkman, C. W., and E. T. Mezquida (2015). Phenotypic selection exerted by a seed predator is replicated in space and time and among prey species. American Naturalist 186 (5):682-691. doi: 10.1086/683131.

159. Mezquida, E. T., and C. W. Benkman. (2005). The geographic selection mosaic for squirrels, crossbills and Aleppo pine. Journal of Evolutionary Biology 18:348-357.

160. Benkman, C. W. (1988b). Flock size, food dispersion, and the feeding behavior of crossbills. Behav. Ecol. Sociobiol. 23:167-175.

161. Hofslund, P. B., W. H. Marshall and G. Robinson. (1955). Wasteful feeding by White-winged Crossbills. Wilson Bulletin 67:215.

162. Benkman, C. W. (1989b). Intake rate maximization and the foraging behaviour of crossbills. Ornis Scand. 20:65-68.

163. Groth, J. G. (1990). Cryptic species of nomadic birds in the Red Crossbill (Loxia curvirostra) complex of North America. Phd Thesis, Univ. of California, Berkeley.

164. Génard, M. and F. Lescourret. (1987). The Common Crossbill Loxia curvirostra in the Pyrenees: some observations on its habitats and on its relations with conifer seeds. Bird Study 34:52-63.

165. Richardson, F. (1938b). Red Crossbills feeding on juniper galls. Condor 40:257.

166. Bartlett, E. (1976). Crossbills feeding at chimney-stacks. British Birds 69:312.

167. Aldrich, E. C. (1939). Notes on the salt feeding habits of the Red Crossbill. Condor 41:172-173.

168. Marshall, W. H. (1940). More notes on salt feeding of Red Cross-bills. Condor 42:218-219.

169. Bennetts, R. E. and R. L. Hutto. (1985). Attraction of social fringillids to mineral salts: an experimental study. J. Field Ornithol. 56:187-189.

170. Dawson, W. R., V. H. Shoemaker, H. B. Tordoff and A. Borut. (1965). Observations on metabolism of sodium chloride in the Red Crossbill. Auk 82:606-623.

171. Bailey, A. M., R. J. Niedrach and A. L. Baily. (1953). The Red Crossbills of Colorado. Publs. Denver Mus. Nat. Hist. 9:1-64.

172. Dawson, W. R. and H. B. Tordoff. (1964). Relation of oxygen consumption to temperature in the Red and White-winged crossbills. Auk 81:26-35.

173. Dawson, W. R. and C. Carey. (1976). Seasonal acclimatization to temperature in cardueline finches I. Insulative and metabolic adjustments. J. Comp. Physiol. 112:317-333.

174. Carey, C., W. R. Dawson, L. C. Maxwell and J. A. Faulkner. (1978). Seasonal acclimatization to temperature in cardueline finches II. Changes in body composition and mass in relation to season and acute cold stress. J. Comp. Physiol. 125:101-113.

175. Sewall, K. B. (2011). Early learning of discrete call variants in Red Crossbills: Implications for reliable signaling. Behavioral Ecology and Sociobiology 65 (2):157-166. doi: 10.1007/s00265-010-1022-0.

176. Keenan, P. C., and C. W. Benkman (2008). Call imitation and call modification in Red Crossbills. Condor 110 (1):93-101. doi: 10.1525/cond.2008.110.1.93.

177. Sewall, K. B., and T. P. Hahn. (2009). Social experience modifies behavioural responsiveness to a preferred vocal signal in Red Crossbills, Loxia curvirostra. Animal Behaviour 77:123-128.

178. Mundinger, P. C. (1979). Call learning in the Carduelinae: ethological and systematic considerations. Syst. Zool. 28:270-283.

179. Adkisson, C. S. (1981). Geographic variation in vocalizations and evolution of North American Pine Grosbeaks. Condor 83:277-288.

180. Pieplow, N. (2017). Peterson Field Guide to Bird Sounds of Eastern North America. Houghton Mifflin Harcourt Publishing Company, NY, USA.

181. Summers, R. W., and S. T. Buckland (2011). A first survey of the global population size and distribution of the Scottish Crossbill Loxia scotica. Bird Conservation International 21 (2):186-198. doi: 10.1017/s0959270909990323.

182. Ross, W. M. 1948. Notes on the Crossbill. Scot. Nat. 60:147-156.

183. Benkman, C. W. (1997). Feeding behavior, flock-size dynamics, and variation in sexual selection in crossbills. Auk 114 (2):163-178.

184. Benkman, C. W. (1988e). Why White-winged Crossbills do not defend feeding territories. Auk 105:370-371.

185. Benkman, C. W. (1992b). "White-winged Crossbills." In The birds of North America, no. 27, edited by A. Poole, P. Stettenheim and F. Gill. Washington, D.C: Acad. Nat. Sci., Philadelphia, PA, and Am. Ornithol. Union.

186. McCabe, T. T. and E. B. McCabe. (1933). Notes on the anatomy and breeding habits of crossbills. Condor 35:136-147.

187. Jardine, D. C. (1994). Brood patch on a Common Crossbill Loxia curvirostra still in juvenile plumage. Bird Study 41:155-156. doi: 10.1080/00063659409477212.

188. Snowberg, L. K., and C. W. Benkman (2009). Mate choice based on a key ecological performance trait. Journal of Evolutionary Biology 22 (4):762-769. doi: 10.1111/j.1420-9101.2009.01699.x.

189. Kleven, O., B.-A. Bjerke and J. T. Lifjeld. (2008). Genetic monogamy in the Common Crossbill (Loxia curvirostra). Journal of Ornithology 149 (4):651-654.

190. Møller, A. P., and J. V. Briskie (1995). Extra-pair paternity, sperm competition and the evolution of testis size in birds. Behavioral Ecology and Sociobiology 36 (5):357-365. doi: 10.1007/s002650050158.

191. Ciach, M., and M. Bylicka (2009). Relative abundance and flock size of wintering Common Crossbills Loxia curvirostra L. in Western Carpathians: A habitat influence. Polish Journal of Ecology 57 (1):181-189.

192. Benkman, C. W. (1988c). On the advantages of crossed mandibles: an experimental approach. Ibis 129:288-293.

193. Petty, S. J., I. J. Patterson, D. I. K. Anderson, B. Little and M. Davison. (1995). Numbers, breeding performance, and diet of the sparrowhawk Accipiter nisus and merlin Falco columbarius in relation to cone crops and seed-eating finches. Forest Ecology and Management 79:133-146.

194. Clouet, M. (2000). The breeding biology of the Common Crossbill Loxia Curvirostra in the Central Pyrenees. Bird Study 47:186-194. doi: 10.1080/00063650009461174.

195. Smith, C. C. and R. P. Balda. (1979). Competition among insects, birds and mammals for conifer seeds. Am. Zool. 19:1065-1083.

196. Coombs-Hahn, T. P. (1993). Integration of environmental cues to time reproduction in an opportunistic breeder, the Red Crossbill Loxia curvirostra.. Phd Thesis, Univ. of Washington, Seattle.

197. Lawrence, L. de K. (1949b). The Red Crossbill at Pimisi Bay, Ontario. Can. Field-Nat. 63:147-160.

198. Summers, R., E. Humphreys, M. Newell, and C. Donald (2002). Nest-site selection by crossbillsLoxiaspp. in ancient native pinewoods at Abernethy Forest, Strathspey, Highland. Bird Study 49:258–262.

199. Snyder, D. P. and J. F. Cassel. (1951). A late summer nest of the Red Crossbill in Colorado. Wilson Bulletin 63:177-180.

200. Snyder, D. E. (1954b). Nesting study of Red Crossbills. Wilson Bulletin 66:32-37.

201. Watson, A., M. Marquiss, and R. Summers (2009). Abundance of crossbills, siskins and cone-crops. Ornis Fennica 86 (1):38-40.

202. Brightsmith, D. J., E. A. Hobson, and G. Martinez (2018). Food availability and breeding season as predictors of geophagy in Amazonian parrots. Ibis 160 (1):112-129. doi: 10.1111/ibi.12515.

203. Ternovskij, D. V. (1954). The winter breeding of crossbills. Bjullet. M. O-va Isp. Prirody, Otd. Biol. 59:37-40.

204. Lutmerding, J. A. and A. S. Love (2017). Longevity records of North American birds. Version 2017.1. Patuxent Wildlife Research Center, Bird Banding Laboratory, Laurel, MD, USA.

205. Alonso, D., and J. Arizaga (2013). The impact of vagrants on apparent survival estimation in a population of Common Crossbills (Loxia curvirostra). Journal of Ornithology 154 (1):209-217. doi: 10.1007/s10336-012-0887-2.

206. Benkman, C. W., J. S. Colquitt, W. R. Gould, T. Fetz, P. C. Keenan, and L. Santisteban (2005). Can selection by an ectoparasite drive a population of Red Crossbills from its adaptive peak? Evolution 59 (9):2025-2032.

207. Cornelius, J. M., M. Zylberberg, C. W. Breuner, A. C. Gleiss, and T. P. Hahn. (2014). Assessing the role of reproduction and stress in the spring emergence of haematozoan parasites in birds. Journal of Experimental Biology 217:841–849.

208. Edelaar, P., and K. van Eerde (2011). Non-random infection across individuals and populations supports that parasites can change morphology within an adaptive radiation. Journal of Zoology 283 (2):135-142. doi: 10.1111/j.1469-7998.2010.00755.x.

209. Pomeroy, D. E. (1962). Birds with abnormal bills. British Birds 55 (2):49-72.

210. Summers, R. W. (2011). Patterns of exploitation of annually varying Pinus sylvestris cone crops by seed-eaters of differing dispersal ability. Ecography 34 (5):723-728. doi: 10.1111/j.1600-0587.2010.06498.x.

211. Siepielski, A. M., and C. W. Benkman (2005). A role for habitat area in the geographic mosaic of coevolution between Red Crossbills and lodgepole pine. Journal of Evolutionary Biology 18 (4):1042-1049. doi: 10.1111/j.1420-9101.2005.00902.x.

212. Rosenberg, K. V., J. A. Kennedy, R. Dettmers, R. P. Ford, D. Reynolds, J. D. Alexander, C. J. Beardmore, P. J. Blancher, R. E. Bogart, G. S. Butcher, A. F. Camfield, A. Couturier, D. W. Demarest, W. E. Easton, J. J. Giocomo, R. H. Keller, A. E. Mini, A. O. Panjabi, D. N. Pashley, T. D. Rich, J. M. Ruth, H. Stabins, J. Stanton, and T. Will (2016). Partners in Flight Landbird Conservation Plan: 2016 Revision for Canada and Continental United States. Partners in Flight Science Committee.

213. Sauer, J. R., D. K. Niven, J. E. Hines, D. J. Ziolkowski Jr., K. L. Pardieck, J. E. Fallon, and W. A. Link (2017). The North American Breeding Bird Survey, Results and Analysis 1966–2015. Version 2.07.2017. USGS Patuxent Wildlife Research Center, Laurel, MD, USA.

214. Dickerman, R. W. (1987). The "old northeastern" subspecies of Red Crossbill. Am. Birds 41:188-194.

215. Erskine, A. J. (1992a). Atlas of Breeding Birds of the Maritime Provinces. Nova Scotia Museum, Halifax, Nova Scotia, Canada.

216. Reinikainen, A. (1937). The irregular migrations of the crossbill (L. c. curvirostra) and their relation to the cone crop of the conifers. Ornis Fenn. 14:55-64.

217. Castro, J., J. M. Gómez, D. García, R. Zamora, and J. A. Hódar (1999). Seed predation and dispersal in relict Scots pine forests in southern Spain. Plant Ecology 145 (1):115-123. doi: 10.1023/a:1009865703906.

218. Svärdson, G. (1957). The "invasion" type of bird migration. Br. Birds 50:314-343.

219. Mineau, P., and L. J. Brownlee. (2005). Road salts and birds: an assessment of the risk with particular emphasis on winter finch mortality. Wildlife Society Bulletin 33:835-841.[835:RSABAA]2.0.CO;2

220. Benkman, C. W. (1993b). Logging, conifers, and the conservation of crossbills. Conservation Biology 7 (3):473-479.

221. Holimon, W. C., C. W. Benkman, and M. F. Willson (1998). The importance of mature conifers to Red Crossbills in southeast Alaska. Forest Ecology and Management 102 (2-3):167-172. doi: 10.1016/s0378-1127(97)00157-6.

222. Helle, P. (1985). Effects of forest fragmentation on bird densities in northern boreal forests. Ornis Fennica 62:35-41.

223. Helle, P. and O. Järvinen. (1986). Population changes of north Finnish land birds in relation to their habitat selection and changes in forest structure. Oikos 46:107-115.

224. Tingley, M. W., D. A. Orwig, R. Field and G. Motzkin. (2002). Avian response to removal of a forest dominant: Consequences of hemlock woolly adelgid infestations. Journal of Biogeography 29 (10-11):1505-1516.

225. Westerling, A. L., H. G. Hidalgo, D. R. Cayan and T. W. Swetnam. (2006). Warming and earlier spring increase western U.S. forest wildfire activity. Science 313 (5789):940-943.

226. Enright, N. J., J. B. Fontaine, D. M. J. S. Bowman, R. A. Bradstock, and R. J. Williams (2015). Interval squeeze: Altered fire regimes and demographic responses interact to threaten woody species persistence as climate changes. Frontiers in Ecology and the Environment 13 (5):265-272. doi: 10.1890/140231.

227. Westerling, A. L., M. G. Turner, E. A. H. Smithwick, W. H. Romme, and M. G. Ryan (2011). Continued warming could transform Greater Yellowstone fire regimes by mid-21st century. Proceedings of the National Academy of Sciences of the United States of America 108 (32):13165-13170. doi: 10.1073/pnas.1110199108.

228. Logan, J. A., J. Regniere, and J. A. Powell (2003). Assessing the impacts of global warming on forest pest dynamics. Frontiers in Ecology and the Environment 1 (3):130-137. doi: 10.1890/1540-9295(2003)001[0130:atiogw];2.[0130:atiogw];2

229. Coops, N. C., and R. H. Waring (2011). A process-based approach to estimate lodgepole pine (Pinus contorta Dougl.) distribution in the Pacific Northwest under climate change. Climatic Change 105:313–328.

Alonso, D., J. Arizaga, R. Miranda, and M. A. Hernández (2006). Morphological diversification of Common Crossbill Loxia curvirostra populations within Iberia and the Balearics. Ardea 94 (1):99-107.

Benkman, C. W. (1988d). Seed handling ability, bill structure, and the cost of specialization for crossbills. Auk 105:715-719.

Björklund, M., D. Alonso, and P. Edelaar (2013). The genetic structure of crossbills suggests rapid diversification with little niche conservatism. Biological Journal of the Linnean Society 109 (4):908-922. doi: 10.1111/bij.12097.

Borrás, A., J. Cabrera, and J. C. Senar (2008). Local divergence between Mediterranean crossbills occurring in two different species of pine. Ardeola 55 (2):169-177.

Edelaar, P., and K. Terpstra (2004). Is the nominate subspecies of the Common Crossbill Loxia c. curvirostra polytypic? I. Morphological differences among years at a single site. Ardea 92 (1):93-102.

Hahn, T. P., J. Wingfield, R. Mullen and P. Deviche. (1995). Endocrine bases of spatial and temporal opportunities in Arctic-breeding birds. Am. Zool. 35:259-273.

Hynes, D. P., and E. H. Miller (2014). Vocal distinctiveness of the Red Crossbill (Loxia curvirostra) on the island of Newfoundland, Canada. Auk 131 (3):421-433. doi: 10.1642/auk-13-224.1.

James, P. C., T. W. Barry, A. R. Smith and S. J. Barry. (1987b). Bill crossover ratios in Canadian crossbills Loxia spp. Ornis Scand. 18:310-312.

Massa, B. (1987). Variations in Mediterranean crossbills Loxia curvirostra. Bulletin of the British Ornithologists' Club 107 (3):118-128.

McNair, D. B. (1988e). Review of breeding records of Red Crossbill and Pine Siskin in the southern Appalachian Mountains and adjacent regions. Migrant 59:105-113.

Mezquida, E. T., and C. W. Benkman (2014). Causes of variation in biotic interaction strength and phenotypic selection along an altitudinal gradient. Evolution 68 (6):1710-1721. doi: 10.1111/evo.12394.

Miller, A. H. (1932d). Fossil passerine birds from the Pleistocene of Carpinteria, California. Univ. Calif. Publ. Bull., Dept. Geol. Sci. 21:169-194.

Parchman, T. L., C. W. Benkman and S. C. Britch. (2006). Patterns of genetic variation in the adaptive radiation of New World crossbills (Aves: Loxia). Molecular Ecology 15 (7):1873-1887.

Pulliainen, E. (1971). Winter nutrition of crossbills (Loxia curvirostra and L. leucoptera) in northeastern Lapland in 1969. Ann. Zool. Fennici 8:326-329.

Pulliainen, E. (1972). Summer nutrition of crossbills (Loxia pytyopsittacus, L. curvirostra and L. leucoptera) in northeastern Lapland in 1971. Ann. Zool. Fenn. 9:28-31.

Questiau, S., L. Gielly, M. Clouet, and P. Taberlet (1999). Phylogeographical evidence of gene flow among Common Crossbill (Loxia curvirostra, Aves, Fringillidae) populations at the continental level. Heredity 83:196-205. doi: 10.1038/sj.hdy.6885510.

Tordoff, H. B. and W. R. Dawson. (1965). The influence of day length on reproductive timing in the Red Crossbill. Condor 67:416-422.

Young, M. A. 2010. First documented occurrences of Red Crossbill Types 1, 2, 3, and 10 in New Hampshire. New Hampshire Bird Records 4: 43-50.

Yuri, T. and D. P. Mindell. (2002). Molecular phylogenetic analysis of Fringillidae, "New World nine-primaried oscines" (Aves: Passeriformes). Molecular Phylogenetics and Evolution 23 (2):229-243.

Recommended Citation

Benkman, C. W. and M. A. Young (2019). Red Crossbill (Loxia curvirostra), version 2.0. In The Birds of North America (P. G. Rodewald, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA.