© Entomologica Fennica. 31 May 2011

Lepidoptera recorded on snow in Central Poland

Agnieszka Soszyfiska—Maj & Jaroslaw Buszko

Soszynska-Maj, A. & Buszko, J. 201 1: recorded on snow in Central

Poland. — Entomol. Fennica 22: 21—28.

Sixteen species ofLepidoptera, belonging to 10 families, were recorded on snow in Central Poland. Two episodes of mass occurrence on snow were observed: adults of Operoplztera brumata and larvae ofEuthrixpotatoria. Twelve species were recorded from snow for the first time in general. Three ecological groups of snow active Lepidoptera were distinguished: 1) autumn and winter active , 2) overwintering species which could be periodically activated from diapause, and 3) early spring active species. The ecology of winter active Lepidoptera is discussed. All were classified as chionoxenes, while 0. brumata was considered as the most regularly recorded on snow and most likely to be met in winter.

A. Soszynska—Maj, Department ofInvertebrate Zoology & Hydrobiology, Uni— versity ofLodz', Banacha 12/16, 90—23 7 Lodz', Poland; E—mail.‘ agasosz@bio— l.ani.lodz.pl J. Buszko, Department of Ecology, Nicolaus Copernicus University, Gagarina 9, 87—100 Toran, Poland; E—mail.‘ [email protected]

Received 5 May 2010, Accepted 27 October 2010

1. Introduction Hagvar & Greve 2004, Soszynska 2004, Soszyn- ska-Maj, unpubl.), and stay inactive when the In areas with a snow cover during winter, certain weather is disadvantageous. cold tolerant poikilothermic organisms are more The invertebrate fauna active on the snow sur- or less winter active. This is an evolutionary and face (supranivean fauna) consists of regular, ecologically explicable behaviour. Even at very however weather depending, snow active species low air temperature in winter, the temperature of and includes the following ecological groups: soil and litter below snow (the so-called sub- chionobionts, specialized in snow activity and nivean space) can remain close to 0°C. Due to its winter reproduction, stenothermic; chionophiles, high air content, snow has a high insulation ca- regularly snow active but occur also in other sea- pacity and low thermal conductivity. Snow with sons, eurytherrnic; chionoxenes, occur on snow low density and a certain thickness provides the only accidentally but adapted to low temperature greatest insulation and enables subnivean activity (Pruitt 1978). ofinvertebrates throughout the winter (Aitchison The invertebrate fauna present on snow is 1974, 2001). Certain ectothermic organisms, strongly weather dependent. Very few papers such as Boreas sp., Chionea sp. or some spring- have been published as a result ofregular investi- tails and spiders, use the favourable weather con- gations, not only accidental trips. Well known for ditions on mild winter days with low air pressure, their snow surface activity are springtails (Col- when the temperature is stable, for migration, lembola), scorpionflies (Mecoptera), flies (Dipte— predation and even copulation (Aitchison 2001, ra) from the families Trichoceridae, Myceto- 22 Soszyiéska—Maj & Buszko ° ENTOMOL. FENNICA Vol. 22 philidae, Limoniidae, Chironomidae and Phori- from 30 to 65 days of the year (unpublished data dae; beetles (Coleoptera) especially from the from Lodz Meteorological Station). families Cantharidae, Carabidae and Staphylini- Lepidoptera were sampled in nature reserves dae; as well as stoneflies (Plecoptera), caddish- and within the largest forest areas. Investigations flies (Trichoptera) and bugs (Heteroptera) (Tah- were made particularly in the dominating forest vonen 1942, Szulczewski 1947, Ackefors 1964, associations: mixed forests containing old Aitchison 2001, Soszynska & Durska 2002, , spruces and firs, riparian forests (Cir— Hagvar & Greve 2004, Soszynska 2004, Hagvar ceo—Alnetum) in vicinity of springs, acidophilous 2010). The snow activity of spiders is also well woods (Luzulo pilosae—Fagetum) contain- known (Aitchison 2001, Hagvar & Aakra 2006). ing 100-years old beeches and dry-ground forests Activity of Lepidoptera on snow has not ear- (Tilio—Carpinetum). lier been studied consistently. However, the win- ter activity of Geometridae and the problem of winglessness ofwinter moths (Roff 1990, Sattler 3. Material and methods 1991, Peterson & Nilssen 1998, Tammaru et al. 2001), as well as the influence ofweather factors This research is a part of more general investiga- on the activity of is well studied tion of the snow-surface active fauna of Central (Buszko & Nowacki 1991). In Poland, the winter Poland during four autumn-spring periods from activity of Lepidoptera was studied by Adam- 1998 to 2002. Few data came from other winters. czewski (1936, 1964) and by Buszko and No- Semi-quantitative samplings were made, if at all wacki (1990, 1991). The data only about Lepi- possible, every day during winter with snow doptera on snow in Poland were presented by cover, from November to March, in the tempera- Szulczewski (1947) and Soszynska-Maj (2008). tures ranging from —10 to +10°C in comparable The aim ofthis paper is to present Lepidoptera daily periods of time (1.5 h). Colder and warmer recorded on the snow surface during regular in- days were skipped. On account of weather-de- vestigations in Central Poland, to describe the pending activity ofthis fauna, short day in winter weather conditions under which they were active, and lower temperature in the morning, the samp- and to discuss their winter strategy. ling was made between 9.30 am. and 3.00 pm. The association ofthe occurrence ofsnow-ac-

tive invertebrates with weather conditions was in- 2. Study area vestigated, noting air temperature, humidity and the depth of the snow cover each time. The air The study area is located in Central Poland, in the temperature and humidity were measured 1 m vicinity of Lodz, one of the biggest cities in P0- above the snow surface in the shade. land. According to the principles established by The seasons were defined according to Klysik Kondracki (2000), which correlate with physico- (2001) for Central Poland. Autumn was defined geographical division ofEurope, the area is called as October, early winter as November until mid- the Lodz Upland. It is the northernmost upland December, winter as the period between mid-De- area in Poland, with maximum of 284 m above cember and March, early spring as March, and sea level, surrounded with lowlands and charac- spring as April. terized by postglacial landforms, lowest tempera- air and snow-cov- ture, highest humidity longest 4. Results ered period in Central Poland. Partly it is pro- tected within Lodz Uplands Landscape Park (Ku- Sixteen species of Lepidoptera, belonging to 10 rowski 1998). families were identified on snow, most for the The snow cover appears regularly in Central first time. Altogether 139 adults and 2 larvae of Poland, but its onset and duration vary among Lepidoptera were collected and numerous larvae years. In winter seasons 1998—2002, the first ofEuthrixpotatoria were observed on snow. One snow falls was noticed between the mid-Novem- species, Caryocolum junctella, was found in ber and the end ofDecember, and snow cover laid Central Poland for the first time. The others are ENTOMOL. FENNICA Vol. 22 ° Lepidoptera on snow in Poland 23

Table 1. No. of collected specimens of Lepidoptera on snow in Central Poland with dates, air temperatures and sampling sites.

Family Species No. Date T (°C) Site

Gracillariidae Phyl/onorycter agile/la (Zeller, 1846) 132 28 Xll 1999 0 Wolbérka res.

Ypsolophidae ustel/a (Clerck, 1759) 1S? 7 II 1999 —2 Las Lagiewnicki 1S? 8 III 2004 +2 Molenda res.

Lyonetiidae Lyonetia clerkel/a (Linnaeus, 1758) 132 24 XI 1999 +2 Las Lagiewnicki 16 25 XI 1999 +1 Wiaczyn res.

Gelechiidae Caryoco/umjuncte/Ia (Douglas, 1851) 1S? 1 l 2001 —2.5 Parowy Janinowskie res.

Tortricidae Tortricodes alterne/Ia (Denis et Schiffermller, 1775) 132 28 Ill 1995 0 Las Lagiewnicki 132 30 Ill 2001 +1 Molenda res.

Pieridae Gonepteryx rhamni (Linnaeus, 1758) 132 19 IX 1997 —2 Las Lagiewnicki

Lasiocampidae 1 potatoria (Linnaeus, 1758) 19 l 2006 — Szczekociny 22 8 III 2006 —4 Spala

Geometridae Archiearis pan‘henias (Linnaeus, 1761) 132 28 Ill 2001 —1 Galkéw res. Agriopis leucophaearia (Denis et Schiffermiiller, 1775) 132 30 Ill 2001 +6 Struga Dobieszk. res. Colotois pennaria (Linnaeus, 1761) 132 28 X 1997 0 Las Lagiewnicki Alsophi/a aceraria (Denis et Schiffermller, 1775) 132 28 XI 1997 —2 Las Lagiewnicki 26 23 XI 1999 —1 Grady n. Linda res. brumata (Linnaeus, 1758) 7657323 19 XI 1999 0 Molenda res. 15S? 21 XI 1999 —1 Galkéwek res. 832 23 XI 1999 —2 Dabrowa Grotnicka res. 66 23 XI 1999 —1 Grady nad Linda res. 19S? 24 XI 1999 +1 Las Lagiewnicki 232 27 Xll 1999 +1 Park Zdrowie — Lédz 132 12 Xll 2001 0 Kruszewiec res. 132 21 Xll 2002 —3 Wiaczyn res. Operophtera fagata (Scharfenberg, 1805) 16 20 XI 1997 —1 Las Lagiewnicki 132 27 XI 1993 —6 Las Lagiewnicki

Noctuidae rubiginosa (Scopoli, 1763) 1S? 6 III 1995 +2 Las Lagiewnicki Conistra vaccinii (Linnaeus, 1761) 132 20 XI 1997 —1 Las Lagiewnicki

Nolidae revayana (Scopoli, 1772) 132 26 II 1999 +2 Las Lagiewnicki 16 25 XI 1999 +1 rez. Wiaczyn 16 3 l 2001 0 Rogéw — Arboretum 132 31 l 2004 —1 Las Lagiewnicki

1 Numerous larvae were observed, J. Tabor & L. Maksalon pers. comm. 2 Larvae. 3 Many more females were observed, but not collected. 24 SOSZJ/fiskmMaj & Buszka a ENTOMOL,

Only Oparophtera brumata

D Other Lepidoptera recorded on snow specimens of No.

E i O 2/X 1/XE 2/Xi ’l/XH 2/XH ”M 2/! 1/” 2/” mu 2””

Seasonal dynamics

M6 "-5 ...2 ml 0 ”i 2

Air tam psmtum (0C)

90" 80" 70" 60" 50'" 40" ofspscimsns 30“ No- 20" 10"

50 6O 70 80 90 100 Humidity (%)

Fig. ”L Lepidoptera on snow in Central Poland in four

winter seasons 1998-4002. specimens __.. a. Seasonal dynamics. of First and last halves of No. months in Arabic numerals preceding months in Roman

numerals. “- b ...__ d. of 15 20 25 30 Lepidoptera in relation to Snow cover {cm} environmental variables.

Widely distributed in Poland. (Buszko & Nowacki share in collected material (6325 specimens of 2000) all invertebrates collected from snow). They were Detailed data of sampling site? date, tempera one of the least numerous orders of snow active ture and number of collected specimens are pren- .) but with almost 20% constancy in the sented in Table l. Lepidoptera had only 2.2% Whole material. ENTOMOL. FENNICA Vol. 22 ° Lepidoptera on snow in Poland 25

Most ofthe collected lepidopteran species, as emerging species. Autumn, winter and early well as of the specimens belonged to the family spring active species differ in their ecological re- Geometridae. One species, Operophtera bruma— action to winter conditions and snow, which is ta, clearly predominated in the collected material expressed in different behavior as well as in mor- (83.6%). Wingless females of this species were phological and physiological adaptations. Lepi- found on snow more often than winged males. doptera collected on snow in Central Poland were Two episodes ofmass occurrence ofmoths on classified into three ecological groups consider- snow were observed. In November 1999, more ing the time of emergence, short or long life, and than hundred individuals of wingless females of hibernation: 1. autumn and winter active moths, 0. brumata were found walking on snow at 0°C, 2. overwintering species which could be periodi- while the males were sitting on trunks of nearby cally activated from diapause, and 3. early spring trees (the same was observed in other days). Hun- active species. The characteristics ofthese groups dreds of caterpillars of were is presented below. seen in January 2006 walking on snow (J. Tabor One species collected on snow in Central Po- and L. Maksalon pers. com.). land belongs to the group ofautumn active moths. The largest catches ofLepidoptera took place Colotois pennaria (Geometridae) has one gene- in early winter and early spring. However, the ration, flies in the autumn until the first frost, usu- early autumn peak was made by the great number ally from September to early December and ofthe dominant species, 0. brumata. Lepidoptera overwinters as an egg (Waring & Townsend were found throughout the whole winter period 2006). To winter moths belong: Alsophila acera— from November to March (Fig. 1a). Lepidoptera ria, Operophtera brumata and 0. fagata (Geo- occurred on snow from —6 to +6°C, with the peak metridae). These species overwinter as eggs and at —1 and +1°C (Fig. lb). All catches were made their larvae appear in April and are thus spring- on overcast weather. They preferred air humidity feeding. 0. brumata has one generation and flies above 70% (with peak on 90%) (Fig. 1c), and the from October to January, while 0.fagata mainly snow cover was usually at least 20 cm deep (Fig. in November (Waring & Townsend 2006). This 1 d). behavior is connected with larval polyphagy, fe- male winglessness, short-lived adults which do not survive winter, do not feed and their mouth 5. Discussion parts are usually reduced (Sattler 1991, Tammaru et al. 2001). Males of 0. brumata are capable of Before this study, five species of Lepidoptera flying even in —2.6°C (Buszko & Nowacki 1991) have been recorded on snow in Poland: three but their flight is slow and poorly controlled. Fe- geometrids, Operophterafagata (in December at males emerge from pupae in early winter with —1°C), Apocheima pilosaria (in March at +2°C) fully developed eggs and mate with males on tree (Szulczewski 1947) and 0. brumata, as well as trunks. Protandry is characteristic for 0. bruma— Ypsolopha ustella () and Tortri— ta, males emerging one week ahead offemales to codes alternella () (Soszynska-Maj maximize mate availability (Peterson & Nilssen 2008). Four others have been recorded in Finland 1998). Newly hatched larvae are ready to exploit in January: Aclerz's hastiana (Tortricidae), fresh foliage in the spring. Females of the winter Caloptill'a elongella (Gracillariidae) (Levander active moths make up the largest category among 1913), Epz'plema crenana (Geometridae) at — Lepidoptera with reduced wings. Fligthlessness 3,5°C and larvae of a noctuid moth Agrotis sp. is possible in stable habitats that provide steady (Tahvonen 1942). In our study, Lepidoptera were resources (e.g. forest), with dispersal in the larval active on snow when temperature ranged from —6 stage. It brings advantages such as increasing to +6°C (Fig. 1b). The highest number of speci- crypsis and reducing body size to minimize heat mens was observed at 0°C, 90% of air humidity loss (Sattler 1991). Sattler (1991) as well as Pe- and snow depth over 15cm (Fig 1b—d.). terson and Nilssen (1998) suggested that cold Continental winter is so severe that it divides season Geometridae emerge and mate in early the cold season invertebrates into late and early winter because of reduced predation pressure. 26 Soszyfiska—Maj & Buszko ° ENTOMOL. FENNICA Vol. 22

Predatory beetles from families Staphylinidae vaccinii and C. rubiginosa. These three moths and Carabidae, and red wood ants in particular have also been observed during winter on tree (Tammaru at al. 2001) are dormant, insectivo- trunks in Germany (Hannig et al. 2006). On Brit- rous mammals hibernate, migratory birds are ab- ish Isles, C. vaccinii is one ofthe commonest win- sent and resident populations of birds change ter species, it flies from September to May and their diet from insects to seeds (Peterson & can be found visiting flowers (Kimber 2010). Nilssen 1998). However, insectivorous mammals Noctuidae stay in diapause in various hidden from family Soricidae as well as beetles from places: in litter, under bark of trees or in hollows families Staphylinidae and Carabidae are winter of trees during periods of unfavorable weather. active and create subnivean winter food chains However, Noctuidae can break the diapause (Aitchison 1984). Peterson and Nilssen (1998) when the air temperature is increasing (Buszko & concluded that 0. brumata emerges in a narrow Nowacki 1991). They are not adapted to stay ac- time period just before winter, as a result of two tive in such low temperatures like Geometridae. opposing selective forces — migrating ofinsectiv- Both sexes feed, have fully developed mouth, and orous birds and first snowfalls. Birds can easily live up to nine months in adult stage. Thanks to prey on the wingless females and snow cover pre- well developed wings, Noctuidae are capable to vents emergence (Peterson & Nilssen 1998). fly in low temperatures at +0,5°C to search for However, the first snow in Central Poland used to food, hiding place and oviposition sites (Adam- fall at the end of November during the observed czewski 1950, Sattler 1991). Their longevity re- peak of activity of 0. brumata. Other studies are quires supplementary feeding in low tempera- needed to find the precise time of emerging of 0. tures, so there is no example of flightlessness in brumata in Central Poland. Waring & Townsend this group of species (Sattler 1991). Long period (2006) pointed that both sexes of 0. brumata can of frost induces an obligatory diapause, which be found resting or walking up tree trunks after must come to the end without a break. C. rubigi— dark. In November 1999 in “Molenda” reserve nosa is well adapted to low temperature and is not more than hundred wingless females were ob- used to overwinter in diapause. It is most active in served walking on snow at 0°C at noon while December and January (Adamczewski 1950, males were seen resting on tree trunks. Hannig et Buszko & Nowacki 1991). al. (2006) observed this species on tree trunks in L. clerkella has two or more generations and Germany in November and December. flies all year round in Europe, and N. revayana The second group of snow-active Lepidop- occurs from September to July. Both overwinter tera comprises species overwintering the cold as adults and reappear in the spring. Y. ustella season in the adult stage. In contrast to the pre- overwinters as an adult and can be found on wing vious group, they have long adult phase, both during this period (Kimber 2010). This moth sexes have normally developed wings and mouth drinks water from the wet snow, which was ob- parts: Ypsolopha ustella (Ypsolophidae), Caryo— served in laboratory after collection on snow colum junctella (Gelechiidae), Canistra rubigi— (Fig. 2.). Similarly to other moths recorded on nosa, C. vaccinil' (Noctuidae), Gonepteryx snow it did not fly but walked on snow surface. It rhamni (Pieridae), Lyonetia clerkella (Lyonetii- is known from literature that many species from dae) and Nycteola revayana (). Gracillaridae, Gelechidae and Tortricidae also In general, a minority of Lepidoptera passes belong to this group, as well as from several other the cold season as adults and only some of them families of Lepidoptera with species overwinte- may be active in winter. Buszko and Nowacki ring as adults (Sattler 1991). (1991) reported that in Poland about 100 species The last group consists of short-lived species overwinter as adults, among them 20 belong to which overwinter as pupae and adults appear in Noctuidae. The above mentioned authors re- early spring: Agriopl's leucophaearia, Archiearis corded 12 species of Noctuidae to be active in parthenias, Tortricodes alternella. Females ofA. winter in central Poland, with three species ob- leucophaearia are wingless and mate with males served during the whole cold period: Eupsilia on tree trunks, and adults are active from early transversa (never seen on the snow), Canistra February to mid March (Kimber 2010). This spe- ENTOMOL. FENNICA Vol. 22 ° Lepidoptera on snow in Poland 27

correcting the English version of the manuscript. We are grateful to Sigmund Hagvar and an anonymous referee for their comments on the manuscript.

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