Lepidoptera: Diurnal and Nocturnal Life-Styles
Thus, whilst farmland abandonment is generally reported to be relevant with regard to butterfly declines, moth declines are rarely linked to farmland abandonment. This observation makes sense if we consider their contrasting life-histories. Butterflies are day-flying ectotherms that need direct sunlight in order to raise their flight muscle temperature to optimal levels, most often well above ambient temperatures. By contrast, most moths are nocturnal and endothermic, raising their body temperatures above ambient levels by generating internal heat via muscle activity. As such, most butterflies and (ectothermic) day-flying moths occur typically in open, sun-lit habitats, whereas a majority of moths are to a greater or smaller degree linked to wooded habitats. For example, even in a largely deforested region (woodland cover < 5 %) of Flanders, 58 % of in total 499 macro-moth species observed in 1980–2012 use shrubs or trees as foodplants (Sierens and Van de Kerckhove 2014). However, the situation is far from black and white as some butterflies are woodland specialists and many nocturnal moths require more open habitat conditions. Still, based on their contrasting thermoregulatory requirements it is clear why in general but terflies are perceived to be more susceptible than moths to farmland abandonment, which is typically accompanied by scrub and forest encroachment, shading out formerly sun-lit biotopes (van Swaay et al. 2010).
Although day-flying Lepidoptera are numerically the exception to the nocturnal norm, sound conservation strategies need to be inclusive of both. For example, in a context of temperate landscapes under intense human land-use, Merckx et al. (2012a) recommend zoned woodland management for the effective conservation of both life-history strategies. Their research showed that the late-successional deciduous woodland biotope is characterised by high numbers of both individuals and species of moths, being especially important for some scarce and specialist species of conservation concern (see also Baur et al. 2006), while coppicing and ride widening, which open up dense forest structures, are valuable woodland conservation tools for Lepidoptera species with an affinity for more open biotopes (see also Fartmann et al. 2013). The mechanism behind the pattern of increased lepidopteran species richness at the woodland-scale, due to such zoned management, involves an increased structural and micro-climatic diversity, and, more generally, increased habitat resource diversity (Merckx et al. 2012a). The importance of habitat heterogeneity at such larger scales is further highlighted by the results obtained from a recent study on butterfly richness of semi-natural meadows in Estonia; using a sample of 22 meadows with a total of 56 butterfly species, the research showed a positive correlation between butterfly diversity on local meadows and forest cover in the landscape directly surrounding (i.e. 250 m radius) these meadows, whilst meadow cover in the surrounding landscape at various spatial scales actually impacted the butterfly diversity of these meadows in a negative way (Ave Liivamägi pers. comm.). For the Mediterranean region too, Verdasca et al. (2012) showed that whilst regular management (i.e. removing understory vegetation) in oak stands has a positive effect on butterfly assemblages, undisturbed stands are nevertheless needed by some butterfly species. A study by Baur et al. (2006) on the abandonment of subalpine semi-natural grasslands in Transylvania, Romania, found that whilst vascular plants reached highest species richness in yearly-mown hay meadows, diurnal Lepidoptera were actually most species-rich in meadows abandoned for three years or more (see also Dover et al. 2011 and references therein), and nocturnal Lepidoptera and Gastropoda were most species-rich in young (20–50 years) and mature (50–100 years) forests, respectively. Because the complementarity of species composition increased with successional age in all four taxonomic groups, and because the proportion of red-listed nocturnal Lepidoptera increased with successional age too, their results indicate the high conservation value of all stages of grassland succession, and especially so the later seral stages up to mature woodland. Hence, all these studies highlight that although semi-natural open biotopes may locally reach high diversity levels (for some taxa), their partial abandonment at the landscape-scale has a role to play for biodiversity in general.