No! Vol. No. December 1994 itI. odonalol., 4, 4. pp. 57-76. I, 63

Survival of adult Somatochlora arctica (Zetterstedt), a

summer species, on snowfields (Anisoptera: Corduliidae)

K. Reinhardt

Hauptstrasse 38, D-09244 Oberlichtenau, Germany

Abstract – - On Mutnovskaya Sopka, Kam-

E. alt. 1300 7 individ- chatka. Russia, m, living

uals were found on the snow surface, July 21 -23,

1991. Two specimens were taken for a closer

examination. One of these survived the at tempe-

rature of about -l°C to -5°C. Arguments for and

against the need for cold hardness are discussed,

and it is concluded that lack speciali-

sation. but can avoid freezing. More males sur-

vived than females.

Introduction

Arthropod fallout and records on snow

surfaces have been noted from various geo-

graphic regions, viz. North and South America

(summarizedin EDWARDS, 1987), northern Eu- Fig. 1. Location of the study site

rope (KA1SILA, 1952; ELTON, 1925; ASH-

MOLE et al.. 1983), Central European mountains (Alnus kamtschatica). Alpine meadows extend to

LISTON & Australia for this there (e.g. LESLIE, 1982), (ED- about 1600 m. Unusually area, were

June and the first WARDS, 1974), Central Asia (SWAN, I96l)and snow falls throughout two

northern even Africa (ASHMOLE & ASH- weeks of July 1991. Only patches of vegetation

field above the sites for MOLE. 1988). During a trip on the Kam- appeared snow and no poten-

tial of chatka peninsula in northeastern Asia, development dragonflies were found.

Ponds of melt were observed on snowfields, amongst them li- water were frozen at night during

adults S. arctico. This is the first record the when ving of, study period, temperatures were ap-

of adult corduliid dragonfliessurviving frost, al- proximately -1”C to -5"C.

though it has been reported in larvae of Sorna-

tochlora semicircularis (WILLEY & EILER. Observations

1972), S. árctica (STERNBERG, 1989) and S. On the 21 st-23rd and 26th-27th July 1991, a large

JOHANSSON & number of found alpeslris(STERNBERG. 1989; insects were near a plateau on

The observations shown These NILSSON, 1991). are the snow, at about 1300 m a.s.l. insects

in detail because very little is known about the were found in obvious pathways. In particular,

effect of snow on dragonfly survival strategies, moths (Geometridae), chrysomelid beetles (Me-

particularly their adult lasoma ae)). carabid beetles (;Leistus during stages. tremul spp.

and Nebria — R. Predel well spp. leg), as as

and Locality hymenopterans (Cimbicidae Symphyta) oc-

The Kamchatka peninsula (eastern Russia) is a curred on the snow surface. Furthermore, two

subarctic area roughly situated between 155" E dead specimens ofAeshna subarctica elisabethae

and E. the 165° The precise observation area, Djakonov and about 40 both dead and alive indi-

volcano active Mutnovskaya Sopka (2323 m). is viduals of S. arctica were found between 21st

Below and at about 58° N and 158° E (Fig. I). 800 23rd July. The two specimens of A. subarc-

birch m, forest (Betula ermani) predominate. tica had folded legs, those of S. arctica showed

alder Above this (up to 1200 m) there is an zone a different pattern. The legs of all dead specimens 64 Notul. odonalol.. Vol. 4, No. 4, pp. 57-76, December 1, 1994

of fallout were extended. Some specimens tippedon their important components are

sides whilst others were tippedon their abdomens members ofthe Homoptera, Dlptera, Coleoptera,

their heads. those and or Of specimens recovered, Hymenoptera (EDWARDS, 1987), occa-

their small melted also and CRAWFORD which were tipped on heads, sionally spiders bugs (e.g.

hollows were noted directly under the head re- & EDWARDS, 1986). According to EDWARDS

gion. No individuals appeared to be injured. (1972, 1987), EDWARDS & BANKO (1976),

and PAPP Of the 40 individuals observed, only 7 were MANN, EDWARDS & GARA (1980)

the alloch- still alive, sitting free on the snow. Surprisingly, (1978) two guilds are always present,

A all ofthe surviving specimens were males. total thonous fallout and the attracted predator guild.

of 23 6 13 collected the The of the and has (10 , 5) were during activity predators scavengers

study period. been noted within different adult arthropodcom-

On the evening of 26th July I searched specifi- munities, notably in collembolans (e.g. AITCHI-

BLOCK & LEI- cally for living individuals located on the top of SON, 1989; ZETTEL, 1980;

AITCHISON, the snow. Two individuals were located and ex- NAAS, 1981), spiders (e.g. 1989;

amined. Althoughone specimen showed signs of HUHTA & VIRAMO, 1979; KOPONEN, 1983,

and in the Boreidae the physical weakness, the other individual appeared 1989), beetles, among

be Both to in arelatively healthy condition. speci- mecopterans (e.g. KOPONEN, 1983). Although

returned the substrate and carnivorous, S. arctica can be mens were to snowy hardly assigned to

examined after the nocturnal frost. On examina- the latter group because of the rare occurrence

Also the tion early the next morning, it became evident of ground feeding (CORBET, 1962).

about 2-8 above the that the weak indvidual had died. The surviving flightactivity takes place m

in dragonfly was placed an area which received ground rather than on the surface (M1DTTUN,

the The maximum exposure to morning sunlight. 1977).

observed 15 It also be incorrect dragonfly was at min intervals. One may to use the term fallout

hour after exposure to the direct sunlight, the in this situation for two reasons:firstly, although

dragonfly had flown away. From intitial observa- not observed, the fallout must have come from

tions it canbe calculated that this individual was lower altitudes due to the occurrence of birch

the substrate least and the associated with frosty for at breeding cimbicid wasps Symphyta.

10 hours. Some ofthe dragonfliesprobably accompanying

An increase in the number offlying specimens the hymenopterans did not die by freezing. Sec-

showed normal and individuals basking on alder shrubs during ondly, they activity. EDWARDS

with rise in in earlier that be the day was clearly associated a tem- (1987), an study, suggested "to

in the environment perature levels. This is in general agreement with fit alpine an organism must

the observations ofMIDTTUN (1977)who recor- either accommodate or evade the extrema”. It is

ded that flight activity in western Norway never interesting to note that S. arctica might follow

occurred below air of 14-15°C. both it does temperature Only strategies, although not belong to

the fallout attracted but males were identified with certainty. either or predators to a

ED- third group, the temporary residents (sensu

Discussion WARDS, 1987). The extended legs recognized

Relatively few published records exist of dragon- in Somatochlora might implicate a landing on

in fly mortalities on snowfields. Ofthose available, the snow, thus a trial and error behaviour this only Sympetrum flaveolum (FUDAKOWSKI, cold withstanding. In , however, the fold-

could cold 1930), Sympetrum sanguineum (LISTON & ed legs mean an overcome by during

LESLIE. 1982), and Aeshna juncea mongolica the flight. The evidence suggest that S. arctica

resistance. This infer the (WOJTUSIAK, 1974) have been mentioned in possesses cold may also

detail. Herewith A. subarctica and Somatochlora production of a cryoprotectant by individuals

found within the whole arctica can be added to this list. geographical area

Results from previous studies suggest that covered by this species. Evidence for this may

role the dragonflies do not play an important in come along three lines. Firstly, 7 ofthe 40 speci- arthropod fallout (SPALDING, 1979). The most mens (17%) of iS. árctica survived the low frost Vol. No. December Notul. odonatol., 4, 4, pp. 57-76, 1, 1994 65

at night. Secondly, sub-zero temperatures are not dead individuals.

unusual at the end ofAugust in some higher parts Although females of S. arctica are generally

of central Europe (e.g. the Harz and Krkonose more difficult to observe (Dr. K. Sternberg, pers.

the in northern males found mountains, or Alps), Norway or comm.), only were to have survived

on the Kamchatka peninsula. Thirdly, since there the frost conditions. Using pitfall traps on snow,

are morphological differences between geogra- KOPONEN (1983, 1989) found in spiders and

there and in that 85-100% phic populations (SCHMIDT, 1957), may flies some mecopterans of

also be physiological differences. Mechanisms those active on the snow were males. Like KO-

for hardiness insects cold in usually depend on PONEN (1983, 1989), the present paper cannot

low mole- of this Differ- high concentrations of a substance of give an explanation phenomenon.

the male cular weight (BAUST & ROJAS, 1985). ences in physiological adaptation of

While other observations of cold hardiness in insects to frost conditions and its implication for

dragonflies have been concerned with larvae the ecology of dragonflies remain unstudied, al-

(BEUTLER, 1989; DABORN, 1971; JOHANS- though differences in the allocation ofenergy for

SON & & GIL- be NILSSON, 1991; SAWCHYN reproductive processes may implicated.

LOT, 1975; WILLEY & EILER, 1972), it is rare

I for adult dragonfliesto survive under frost condi- Acknowledgements – am grateful to the fol-

in Other who tions, except Sympecma. exceptions are lowing gave helpful comments on earlier restricted to the subpolar and the alpine regions. drafts or helped in the English version: Mr S.

CORBET (1962) defines the terms spring and ROTH, Drs C. BEAN, R. JODICKE, A. MAR-

The be D. K. H. summer species. summer species may TENS, MIDDLETON, STERNBERG,

into and late For and referee. Mr R. PREDEL divided early summer forms. WILDERMUTH, a

the latter, frost contacts in the temperate latitudes led the field trip. Dr C. BEAN helpedpreparing

are not unusual in species flying until the late the figure.

autumn (JODICKE, 1991), including e.g. ovipo-

the of - sition on ice cover ponds (BISCHOF, References AITCHISON, C.W., 1989, Aquilo

while the surrounded 24: - 1983 Libellula 2: 1992), or emergence waters 83-89; ANSELIN, A., ,

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- PETERS. G., mi .Die Edellibellen Europas Received April 14. 1994

Aeshnidae Ziemsen, Wittenberg Luther-