Scots pine forest in Central Europe as a habitat for Harmonia axyridis: temporal and spatial patterns in the population of an alien ladybird

Understanding of habitat favourability has wide relevance to the invasion biology of alien species. We studied the seasonal dynamics of the alien ladybird Harmonia axyridis (Coleoptera: Coccinellidae) in monoculture Scots pine forest stands in south-west Slovakia, Central Europe, from April 2013 to March 2015. Adult H. axyridis were collected monthly across seven randomly selected pine stands of different ages and canopy closure, from the lower branches of pine trees, and larvae were recorded qualitatively. Adults were recorded all year round, most abundantly in November and least abundantly in February. The relationship between the abundance of H. axyridis and selected forest stand characteristics was modelled using the negative binomial Generalized Additive Model with penalized spline component in month (seasonality) effect, year, canopy closure and age effects and the random effect of forest stand (sample area effect). The abundance of H. axyridis was significantly influenced by the age of stand and seasonality (with month granularity) for both closed and open canopy stands, whereas the effects of canopy closure and sample area were not significant. The bimodal pattern of seasonal dynamics of H. axyridis on Scots pine was common for closed and open canopy stands, with two peaks reflecting the cyclic movement of the species from and to overwintering sites. Harmonia axyridis utilized certain pine stands preferably for foraging during the growing season and certain stands for refuge during winter. The ladybirds were found in highest numbers in the 15 year old closed canopy stand (overwintering site). The occurrence of both adults and larvae in most stands indicated a suitability of Scots pine forest for ladybird breeding. The model of year-round dynamics of H. axyridis has been presented for the first time within the invaded range of the ladybird in Europe.