Coevolution between crossbills and black pine: the importance of competitors, forest area and resource stability


Craig W. Benkman, Department of Zoology and Physiology, Program in Ecology, Box 3166, University of Wyoming, Laramie, WY 82071, USA.
Tel.: (307) 766 2978; fax: (307) 766 5625; e-mail:


Studies of predator-prey interactions have found that geographically structured coevolution has played an important role in the adaptive diversification of crossbills (Loxia spp.). We extend those studies by considering common crossbills (L. curvirostra) in the Mediterranean where they rely on seeds in the cones of black pine (Pinus nigra). On the continent, where tree squirrels (Sciurus vulgaris) are present, enhanced defenses against crossbills were most evident in larger areas of pine forest. On islands in the absence of tree squirrels, crossbills and black pine have coevolved in a predator-prey arms race on Cyprus but not Corsica. In contrast to other conifers that island endemic crossbills rely upon, black pine does not hold seeds in its cones year round. Consequently, key to the strong crossbill–pine interaction on Cyprus is likely the presence of an alternative conifer that provides seeds during early summer when black pine seeds are scarce.