Study sites
The study was conducted during the months of May and June 2021 on Lipsi
Island, Greece (approximate area: 17 km2), which is
located within the eastern Aegean Sea (37° 17’ 44.7” N, 26° 46’ 45.5” E)
and used as a model small island ecosystem. While the island experiences
low-intensity urbanisation on a global scale (building density = 15-20%
in urban areas), the impacts of urban development on the natural
landscape of small island ecosystems can be comparatively greater than
in built-up areas and warrant investigation (Fernandes and Pinho, 2017).
Sampling over an entire year was not logistically feasible, so we chose
this timescale because previous studies in Mediterranean regions
indicated that peak butterfly activity and abundance should occur in May
and June (Fileccia et al., 2015; Hantson and Baz, 2013). The lower
abundance and richness observed in early spring and late summer is
mainly due to a reduction in flower diversity and, thus nectar sources
for pollinators (Hantson and Baz, 2013).
Nine urban and nine rural sites with clear separation were selected
(Figure 1). The minimum distance between study sites in urban areas was
75.3 m, which limited the double counting of individuals. Due to the
lack of trees, shade is restricted on Lipsi Island, thus, locations with
high light intensity and low shade were utilised to give an accurate
representation of the urban and rural habitats used by butterfly
species. Sites were chosen to represent the predominant land-use types
utilised by butterflies during one or several stages of their life cycle
(Grill and Cleary, 2003). The chosen rural habitats were shrubland,
olive groves, and meadows, while the urban habitats included
agricultural meadows, abandoned land, parks, roadsides, and olive
groves. The sites were similar in size to keep the sampling effort
consistent.