Landscape and climate determine patterns of spread for all colour morphs of the alien ladybird Harmonia axyridis

Aim: Understanding why some areas are more vulnerable to invasion by invasive alien species than others represents a key challenge in invasion biology. We investigated the roles of landscape, climate, proximity to initial invasion sites, recording intensity and colour pattern polymorphism in explaining the spread of the invasive ladybird Harmonia axyridis across Great Britain. Location: The British mainland and inshore islands (Anglesey, Isle of Wight and the Inner Hebrides). Methods: Patterns in time to invasion of H. axyridis were quantified at a hectad level (10 km grid squares) from over 23,000 records. Bayesian spatial survival models were developed relating spread patterns to concurrent patterns in climate, land cover and recording intensity. Separate models were developed for melanic (f. conspicua and f. spectabilis) and non‐melanic (f. succinea) colour forms that were hypothesized to differ in their response to climate factors, particularly temperature, and subsequent spread. Results: Harmonia axyridis spread more rapidly into hectads containing a high proportion of urban land cover and marginally more slowly into hectads containing high conifer cover. Invasions were recorded more rapidly in hectads with high recording intensity, or that were close to initial invasion sites. Harmonia axyridis colour morphs spread to similar extents, although increased sunshine significantly enhanced the spread of the non‐melanic form (f. succinea). Correspondence between observed and predicted times of invasion from survival models was substantial (Pearson's rp for all H. axyridis colour forms = 0.834). Main conclusions: The ability of H. axyridis to use a wide range of habitats and prey species and overwinter inside buildings may have allowed this species to establish rapidly in urban areas. Native ladybird species in urban areas may suffer combined negative impacts of habitat deterioration and competition from H. axyridis. Spatial survival models help us to identify areas vulnerable to invasion and to understand the impacts of multiple environmental drivers on the spread of invasive alien species, given detailed data on time to invasion.