Climatic dipoles drive two principal modes of North American boreal bird irruption

Pine Siskins exemplify normally boreal seed-eating birds that can be sparse or absent across entire regions of North America in one year and then appear in large numbers the next. These dramatic avian "irruptions" are thought to stem from intermittent but broadly synchronous seed production (masting) in one year and meager seed crops in the next. A prevalent hypothesis is that widespread masting in the boreal forest at high latitudes is driven primarily by favorable climate during the two to three consecutive years required to initiate and mature seed crops in most conifers. Seed production is expensive for trees and is much reduced in the years following masting, driving boreal birds to search elsewhere for food and overwintering habitat. Despite this plausible logic, prior efforts to discover climate-irruption relationships have been inconclusive. Here, analysis of more than 2 million Pine Siskin observations from Project FeederWatch, a citizen science program, reveals two principal irruption modes (North-South and West-East), both of which are correlated with climate variability. The North-South irruption mode is, in part, influenced by winter harshness, but the predominant climate drivers of both modes manifest in the warm season as continental-scale pairs of oppositely signed precipitation and temperature anomalies (i.e., dipoles). The climate dipoles juxtapose favorable and unfavorable conditions for seed production and wintering habitat, motivating a push-pull paradigm to explain irruptions of Pine Siskins and possibly other boreal bird populations in North America.