Dew-worms in white nights: High-latitude light constrains earthworm (Lumbricus terrestris) behaviour at the soil surface
Soil is an effective barrier to light penetration that limits the direct influence of light on belowground organisms. Variation in aboveground light conditions, however, is important to soil-dwelling animals that are periodically active on the soil surface. A prime example is the earthworm Lumbricus terrestris L. (the dew-worm), an ecosystem engineer that emerges nocturnally on the soil surface. In the summer, the northernmost populations of L. terrestris are exposed to a time interval with no daily dark period. During a two-week period preceding the summer solstice, we studied the constraints that boreal night illumination imposes on L. terrestris surface activity by comparing their behaviour under ambient light with artificially-induced darkness. Looking for evidence of geographical divergence in light response, we compared the behaviour of native L. terrestris (Jokioinen, S–W Finland; 60°48′N) with two markedly more southern populations, from Preston (Lancashire, UK; 53°47′N) and Coshocton (Ohio, USA; 40°22′N) where the nights have a period of darkness throughout the year (total latitudinal range ca. 2300 km). Under ambient light conditions, L. terrestris emergence on the soil surface was diminished by half compared with the darkened treatment and it peaked at the darkest period of the night. Also mating rate decreased considerably under ambient light. The native dew-worms were generally the most active under ambient light. They emerged earlier in the evening and ceased their activity later in the morning than dew-worms from the two more southerly populations. The differences in behaviour were, however, significant mainly between native and UK dew-worms. In the darkened treatment, the behaviour of the three earthworm origins did not differ. Under the experimental conditions light condition was the dominant environmental factor controlling surface activity, but elevated night-time air temperature and humidity also encouraged dew-worm emergence without discernible differences among geographical origins. Our results show, that in boreal summer, the high level of night illumination strongly limits soil-surface activity of dew-worms. Considering the important regulatory role of L. terrestris in many ecosystem processes, this can have significant corollaries in dew-worm impacts on the environment. Although evidence for geographical differentiation in behaviour was obtained, the results point to phenotypic flexibility in L. terrestris light response.