Is the alpine divide becoming more permeable to biological invasions? - Insights on the invasion and establishment of the Walnut Husk Fly, Rhagoletis completa (Diptera: Tephritidae) in Switzerland

The Walnut Husk Fly, Rhagoletis completa Cresson (Diptera: Tephritidae), is native to North America (Midwestern US and north-eastern Mexico) and has invaded several European countries in the past decades by likely crossing the alpine divide separating most parts of Switzerland from Italy. Here, we determined its current distribution in Switzerland by sampling walnuts (Juglans regia L.) in ecologically and climatically distinct regions along potential invasion corridors. R. completa was found to be firmly established in most low altitude areas of Switzerland where walnuts thrive, but notably not a single parasitoid was recovered from any of the samples. Infested fruit was recovered in 42 of the 71 localities that were surveyed, with mean fruit infestation rate varying greatly among sites. The incidence of R. completa in Switzerland is closely related to meteorological mean spring temperature patterns influencing growing season length, but not to winter temperatures, reflecting survival potential during hibernation. Importantly, areas in which the fly is absent correspond with localities where the mean spring temperatures fall below 7 degrees C. Historical data records show that the natural cold barrier around the Alpine divide in the central Swiss Alps corresponding to such minimal temperatures has shrunk significantly from a width of more than 40 km before 1990 to around 20 km after 2000. We hypothesize on possible invasion/expansion routes along alpine valleys, dwell on distribution patterns in relation to climate, and outline future research needs as the incursion of R. completa into Switzerland; and, more recently, other European countries, such as Germany, Austria, France and Slovenia, represent an example of alien species that settle first in the Mediterranean Basin and from there become invasive by crossing the Alps.