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<I>Colias Palaeno</I>

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<I>Colias Palaeno</I> CryoLetters 35 (3), 247-254 (2014) © CryoLetters, [email protected] OVERWINTERING OF THE BOREAL BUTTERFLY COLIAS PALAENO IN CENTRAL EUROPE Pavel Vrba1, Matthias Dolek2, Oldřich Nedvěd1,3*, Helena Zahradníčková3, Cristiana Cerrato4 and Martin Konvička1,3 1 Faculty of Science, University of South Bohemia, Ceske Budejovice, Czech Republic; 2 Büro Geyer und Dolek, Wörthsee, Germany; 3 Institute of Entomology, Biology Centre, Academy of Sciences of the Czech Republic, Ceske Budejovice, Czech Republic; 4 Department of Life Science and System Biology, University of Turin, Torino, Italy. *Corresponding author email: [email protected]. Abstract BACKGROUND: Colias palaeno (Linnaeus, 1761) (Lepidoptera: Pieridae) is a butterfly with boreal distribution with declining populations in peat bogs and subalpine habitats in Central Europe. OBJECTIVE: We investigated the cold tolerance of overwintering caterpillars from one mountain population from Czech Republic (960m a.s.l.) and one alpine population from Italy (2000m a.s.l.). METHODS: We measured supercooling point (SCP), lower lethal temperature (LLT) and content of cryoprotectants. RESULTS: The caterpillars were freeze-avoiding, with lower LLT close to their very low SCP (–25 to –27°C). The mountain population accumulated high concentrations of glycerol (5% fresh mass) and sugars (trehalose 0.8%, glucose 0.2%), while the Italian alpine population only moderate amounts of glycerol (0.3%) and sugars (trehalose 0.5%, glucose 0.3%) without effect on their cold hardiness. Larvae that overwintered at +5°C had a lower body mass than those overwintering in natural conditions, indicating a metabolic weight loss, but both groups survived equally well. CONCLUSION: We hypothesize that the high concentration of glycerol contributes to the high desiccation tolerance. Keywords: cold hardiness, cryoprotective polyols, insect conservation, mountain ecology INTRODUCTION mountains (33, 11). Supercooling capacity and protection against chill injury are enhanced by Low temperatures affect insect survival, the synthesis and accumulation of protective particularly in extremely cold environments, substances, including glycerol, glucose, sorbitol such as mountains. Insects have to complete and trehalose (37, 31). Glycerol functions as an their development there in a limited part of the antifreeze, decreasing the freezing temperature, season (32) and have to deal with temperature as a cryoprotectant binding free water (26) and extremes during overwintering (15, 25). On the against desiccation generally. other hand, predictable snow cover in such Model species distribution and activity regions insulates ground-dwelling organisms Colias palaeno (Linnaeus, 1761) against low ambient temperatures, improving (Pieridae), is a Boreal zone butterfly, inhabiting their survival (8, 27, 28, 43). Similarly, humid a vast circumpolar range in northern Eurasia, Sphagnum tussocks in boggy habitats buffer from approximately the 50° parallel northwards. against microclimatic fluctuations (41). In addition, it forms multiple relic populations Selection of an appropriate microhabitat is situated more southerly, in higher-elevated and important for survival (22, 45). From a cooler areas of the temperate zone (42). conservation point of view, recent changes of C. palaeno occurs in a wide range of habitats, temperature and (or) precipitation regimes provided that its host plant, Vaccinium should be risky for species inhabiting limited uliginosum L., is present. In temperate Central extrazonal habitats, e. g. cold peat bogs or high 247 Europe, C. palaeno occurs as a glacial relic at We used hibernating caterpillars of two main habitat types: cold peat-bogs, i.e. C. palaeno, originating from two distinct azonal oligotrophic wetlands in submountain localities in the Czech Republic and Italy. The and mountain elevations, from approximately Czech mountain population (Šumava Mts., 400m altitude onwards, and open, sometimes 49°02'N, 13°40'E, 960 m a.s.l.) inhabits a drier, heathlands at and above the timberline in mosaic of Sphagnum-bog, wet grasslands and high-elevations of the Alps (5, 6, 39). Larvae waterlogged Picea abies – Betula pubsecens are commonly reported to overwinter on dry woodlands. Here, fertilized females (n = 45) leaves attached to the stems by silk fibre or were captured in July 2010, and transported to falling with the leaf to the ground, mainly in the an outdoor rearing facility. The Italian alpine lower parts of the host plant (15, 23) or leave population (Alpi Lepontine, 46°10'N, 8°10'E, the host plant and overwinter at Sphagnum 2000 m a.s.l.) inhabits dwarf shrubland habitats covered ground (Dolek et al., unpublished). at the timberline containing the host plant Many of the southern relic populations of Vaccinium uliginosum.We directly collected 30 C. palaeno are declining at present (7, 30). This third-instar larvae from V. uliginosum leaves in particularly applies for populations inhabiting September 2011, and transported them to the Sphagnum-bogs within submontane and same rearing facility. mountain elevational zones. The species is thus The rearing facility, located in lowland extinct in Belgium as well as in northern and semi-natural conditions (at 400 m a.s.l.) some central states of Germany.Since the 1990s, consisted of wired cages (50 x 50 x 100cm) a loss of about 50% of localities occurred in this covered by nylon mesh. Each of cages contained southern Bavaria (6). Concurrently in the Czech a flower pot with a V. uliginosum plant. Republic, the species lost about 40% of its mid- From the mountain population, we released 20th century distribution. The species does not up to five females into each cage and fed them seem to be threatened in the Alps (9). Central with a 5% sucrose solution. After the larvae European populations and high altitude alpine hatched, they were let to develop without populations are different subspecies, C. p. disturbance, except for watering host plants. In europome (Esper, 1778) and C. p. europomene the first decade of November, after the larvae (Ochsenheimer, 1808), respectively. Direct stopped feeding and entered diapause, being exchanges of individuals between the prealpine rolled in dry leaves of V. uliginosum attached to bog and hill area and the Alps are unlikely. stems, we transferred pots to an air-conditioned The reasons implicated to explain the loss room with constant 5°C temperature and about of lower-elevation populations include habitat 40% r.h. The plants were regularly sprayed with degradation (drainage, successional change) (8), water to avoid desiccation. Larvae from the and climatic warming (cf. 34). No information alpine population were already showing signs of regarding the ecophysiological limits of the entering diapause when released to the cages. species’ persistence is known. Obtaining such After a few days in the rearing facility, they information may shed some light on persistence were treated identically as the mountain ones. patterns of northern species in more southerly Cold hardiness measurements latitudes, and thus on their future prospects. In January, we removed individual larvae In this paper, we investigated the lower from the host plant and measured their cold thermal survival limits of C. palaeno diapausing hardiness. The supercooling point (SCP) was larvae. We compared larvae from two contrast measured individually in sixteen individuals per southern populations, from mountain peat bogs population using a line recorder with hand-made in Czech Republic and from timberline in the thermocouples (14), which were attached to the Alps, Italy. We measured how these two body of the experimental caterpillars (10) and populations differed in their cold hardiness, cooled at rate 1 degree per minute. After the expressed as supercooling point, lower lethal exotherm appeared on the line recorder, the temperature, postdiapause survival and body larva was kept in the cooling device until its content of cryoprotectants. body temperature decreased again to the value of crystallization temperature. It was then MATERIALS AND METHODS warmed up, removed from the chamber and kept in a Petri dish at +5°C for 24 hours. Survival Rearing experimental individuals was subsequently checked at 20°C as presence 248 of spontaneous movement or reaction to twigs with young leaves. Their body mass was mechanical stimuli. measured after the transfer and then after one To measure lower lethal temperature and two weeks. Pupation and adult eclosion (LLT), groups of ten larvae were put on dry were recorded, sex of the adults was filter paper in a Petri dish, and placed in determined. Differences between the two incubator providing slow cooling to the set up treatments were analyzed by t-test with temperature measured inside the dish by resistor independent samples (Statistica 10). thermometer. Series of temperatures close to and above SCP were used. Larvae were RESULTS removed after 24 hours and treated as above to check their survival. LLT50 was calculated using Cold hardiness the logistic regression method in Statistica 10 Overwintering larvae of Colias palaeno did (38) package. We also measured lethal time (Lt) not survive SCP measurements with freezing of at selected temperature above SCP using the their body fluids, and thus exhibited the freeze- same method as for LLT, with exposure times avoiding strategy. Mean SCP (±SD) were – from one to seven days. Again, Lt50 was 24.8±3.9°C (mountain population) and – calculated using logistic regression. 26.8±2.7°C (alpine population);
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