Life cycle and phenology of problematicus Herbst, 17 86 in (Coleoptera: Carabidae)

PALLEJ0RUM

J0rum, P.: Life cycle and phenology of Carabus problematicus Herbst, 1786 in Den­ mark (Coleoptera: Carabidae). Ent. Meddr 53: 27-30. Copenhagen, Denmark 1985. ISSN0013-8851.

The life cycle and the phenology of Carabus problematicus Herbst were studied in a Danish oak wood, Hald Ege, in central Jutland, by means of pitfall trapping and exa­ mination of ovaries. Adult were active from May to October, predominantly in May-June and August-September. The activity in spring was caused by beetles which had hibernated, and which had probably reproduced during the previous year. Newly emerged adults occurred from late July to early August; they reproduced in the autumn, mainly in late August and the first half of September. Consequently, in Denmark C. problemati­ cus appears to be an autumn breeder with larval hibernation.

Palle J0rum, N0rrem01levej 84, 8800 Viborg, Denmark.

Introduction climate in northern England (Houston Life cycles of carabid beetles are often 1981 ). described on the basis of seasonal activity The discrepancies between these reports variations of the adults. According to Lars­ might be due to variation in life cycle and son (1939) carabids are either »spring bree­ phenology with climate. It is possible, howe­ ders« or »autumn breeders« with adults or ver, that the conclusions reached by Larsson larvae hibernating, respectively. The annual (1939) and Lindroth (1945) were incorrect, activity pattern and the time of breeding, the more so as they were based chiefly on however, are influenced by climate and may inspection of museum collections of imagi­ accordingly vary in time and space. In nes and larvae, and not on examination of Patrobus atrorufus (Strom), Refseth (1980) the breeding condition of the beetles. Thus, found that the breeding period shifted from further studies of the life history pattern of autumn to spring with increasing altitude, C. problematicus in Scandinavia seem to be and in certain species which normally have required. The present account describes the an annual life cycle, development may last phenology and life cycle of the species in a two years in northern and alpine climates Danish oak wood habitat. (Forsskahl 1972, De Zordo 1979, Refseth 1984), as well as in temperate lowland habi­ Distribution and choice of habitat tats in years with unfavourable weather C. problematicus primarily occurs in (J0rum 1980). western . In Denmark it is widely Carabus problematicus Herbst is an distributed in Jutland, less so in the autumn breeder with overwintering larvae in easternmost parts. Except for Lres0 it is western and central Europe (e.g. Drift 1959, absent from the isles (Bangsholt 1983). Greenslade 1965, Hurka 1973). For Den­ According to Lindroth (1945) the species mark and Fennoscandia. Larsson (1939) and is decidedly xerophilous in Fennoscandia, Lindroth (1945) consider the species to be a occurring on heaths with Calluna, Empe­ spring breeder having summer larvae and trum, etc., sometimes also with sparse pine hibernating as adults. A biennial life cycle, growth. on the contrary, is described for a subarctic In Britain (Greenslade 1965) and in cen-

27 tral Europe (Gries et al. 1973) it is a euryto­ whether they were immature or old females pic woodland species living on different which already had laid eggs. kinds of soil and under highly varying humi­ dity conditions (Drift 19 59, Lauterbach Results 1964, Loser 1972); rarely does it occur on The activity pattern of C. problematicus in open heathland, where populations are pro­ Hald Ege is shown in Fig. I. The results are bably maintained only by migration from given as number of specimens caught per adjacent forest habitats (Boer 1970). 100 traps per day (activity density; cf. In Denmark C. problematicus is confined Thiele 1977). Catches of newly emerged to more or less dry, sandy soil; it occurs in beetles and of females with mature eggs are open country as well as in woodland (Han­ indicated. sen 1968). Beetles were active from May to October. Two peaks of activity were recorded, one in Study area and method May-June and one in August-September. The spring peak was due to old beetles The investigation took place in 1979 and which had hibernated and which had most 1980 in Hald Ege, central Jutland (UTM re­ probably reproduced the previous year. A ference: 32VNH25), an oak wood on poor, number of females caught during this period rather dry, sandy soil. Most parts of the contained conspicuous corpora lutea; but, wood have a layer of raw humus; only as stated above others were badly preserved minor areas have mull soil. Oaks (Quercus and their state of gonad development could robur L., Q. petraea Liebl. and Q.robur x not be determined. In 1979 spring activity Q. petraea) are the dominating trees, in so­ was high, starting about mid-May and me places growing in rather open, tall lasting until the beginning of July. In spring stands, but also forming low, dense, scrubli~ 1980, however, only a few specimens were ke stands. The underwood is especially rich trapped. in juniper (Juniperus communis L.), but al­ After a summer period with low catches so honeysuckle (Loricera periclymenum L.) an increase in activity was recorded in late and alder buckthorn (Frangula alnus Mill.) July and early August when newly emerged are common. The most predominant plants beetles occurred. The autumn peak of activi­ in the field layer are bilberry (Vaccinum ty coincided with reproduction. Females myrtillus L.), wavy hair-grass (Deschampsia containing eggs were most predominant in flexuosa (L.) Trin.), May lily (Maianthe­ the last half of August and in the first half of mum bifolium (L.) Schm.), common cow­ September, which, accordingly, was the wheat (Melampyrum pratense L.) and hairy main breeding period. Some mature females wood-rush (Luzula pilosa (L.) Willd.). could still be found in October. Beetles were sampled in pitfall traps, jars Larvae showed little above-ground activi­ 7.5 cm in diameter and 12 cm deep, contai­ ty. Only two third instar larvae were trap­ ning a 4% formaldehyde solution. The ped, both during 20 May-3 June 1979. samples were taken on 4 study plots with different vegetation; at each site 3-4 traps were used, placed about 20 m apart. The Discussion traps were emptied at irregular intervals, The present investigation shows that C. usually tvo or three times a month during problematicus has autumn propagation the summer. The trapping periods were and larval hibernation in Hald Ege. This April-December 1979 and April-October result does not correspond with information 1980. hitherto given on the life cycle in southern Females were dissected in order to deter­ Scandinavia (Larsson 1939, Lindroth 1945), mine their breeding condition (Schj0tz­ but it agrees well with findings from other Christensen 1961). Individuals with mature parts of Europe. eggs were easily recognized; many speci­ A detailed account of the life cycle in mens were in a poor state of preservation, Holland was given by Rijnsdorp (1980): probably because of too long sampling inter­ Newly emerged beetles occur in July and in vals, which made it impossible to decide the beginning of August. Larvae occur from

28 cf 1979 9 1979 30

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0 A M J J A s 0 A M J j A s 0 Fig. I. Activity density (beetles caught per 100 Fig. 1. Aktivitetstcetheden (biller fanget pr. 100 traps per day) of Carabus problematicus fcelder pr. dag af Carabus problematicus Herbst in 1979 and 198{). The total Herbst i 1979 og 1980. Det totale antal number of beetles caught was 246 in fangne biller var 246 i 1979 og 72 i 1980. 1979 and 72 in 1980. Signaturer Signatures Sorte area/er: For hanner, alle undtagen Black area: For males, all except newly nyligt klcekkede biller. hatched beetles. For females, immature For hunner, umodne samt post-repro­ as well as post-reproductive . duktive dyr. White area: Newly hatched beetles. Hvide area/er: Nyligt klcekkede biller. Hatched area: Females with ripe eggs. Skraverede area/er: Hunner med modne ceg. the end of October to the beginning of June. Besides larvae, a number of old adults hiber­ nate and resume ·activity in the following Sammendrag spring. Some of these beetles reproduce for a Livscyklus og frenologi hos Carabus proble­ second time in the autumn. The results con­ maticus Herbst, 1786 i Danmark (Coleopte­ cerning the life cycle of C. problematicus in ra: Carabidae) Hald Ege are in good accordance with those reported from Holland by Rijnsdorp. They L0bebillen Carabus problematicus Herbst er i also agree with data obtained from southern Mellemeuropa en efteriirsforplanter med overvin­ England by Greenslade (1965) and from tring pii larvestadiet. Derimod er den hidtil blevet central Europe by e.g. Kolbe (1968), Hurka anset for at have foriirsforplantning i Danmark og (1973) and Krause (1974). Fennoskandien, en formodning som har vreret ba­ On the basis of the present study, and seret pa registrering af imagines og larver i considering the fact that previously obtained museumssamlinger, men derimod ikke pii under­ results from Scandinavia were not based on s0gelser af gonadetilstanden hos imagines. inspection of gonads, it appears that in Den­ I iirene 1979 og 1980 har jeg unders0gt artens livscyklus i Hald Ege (EJ; NH25). Indsamlingeme mark C. problematicus is an autumn breeder foregik ved hjrelp af faldgrubefrelder med forma­ with larval hibernation. lin; freldeme blev t0mt ea. 2-3 gange pr. maned. Hunneme blev dissekeret, hvorved tidspunktet for Acknowledgment reglregning kunne fastlregges. I wish to thank Viborg Statsskovdistrikt for Imago var aktiv fra maj til oktober, isrer foriir permission to work in Hald Ege. (maj-juni) og efteriir (august-september), Fig. 1.

29 Fonhsaktiviteten, som var langt sterre i 1979 end Houston, W.W.K., 1981: The life cycles and age i 1980, skyldtes gamle, overvintrede biller, der of Carabus glabratus Paykull and C. problema­ formentlig havde forplantet sig det foregaende ar. ticus Herbst (Col. Carabidae) on moorland in Hejsommeren var pneget af srerdeles ringe loko­ northern England. - Ecol. Entomol. 6 : motorisk aktivitet. Den nye generation fremkom 263-271. hovedsagelig i slutningen af juli og i begyndelsen Hurka, K., 1973: Fortpflanzung und Entwicklung af august. Den heje aktivitet i efterarsmanederne der mitteleuropiiischen Carabus- und Proce­ faldt sammen med tidspunktet for forplantningen; rus-Arten.- Studie Csl. Akad. Ved. 9: 1-78. andelen af hunner med modne reg i ovarierne var Jerum, P., 1980: Life cycles and annual activity sterst i sidste halvdel af august og i ferste halvdel patterns of Pterostichus melanarius (Illig.) and af september. P. niger (Schall.) (Coleoptera: Carabidae) in a Larvernes aktivitet pa skovbunden var yderst Danish beech wood. - Ent. Meddr 48: 19-25. beskeden. Kun to 3.stadie-larver forekom i frelde­ Kolbe, W., 1968: Ober das Vorkommen boden­ materialet, begge i perioden 20.5.-3.6.1979. bewohnender Kiifer in einem Siegerliinder Hau­ Undersegelsen var vist, at C. problematicus i berg und dem angrenzenden Fichtenforst. - Hald Ege er efterarsforplanter med samme livs­ Decheniana 120: 225-232. cyklus som i Mellemeuropa. Denne livscyklus for­ Krause, R., 1974: Die Laufklifer der Siichsischen modes at vrere den for arten normale i Danmark. Schweiz, ihre Phiinologie, Okologie und Ver­ gesellschaftung (1). (Col. Cicindelidae et Carabi­ dae).- Faun. Abh. Mus. Tierk. Dresden 5, (2): 73-179. Reforences Larsson, S.G., 1939: Entwicklungstypen und Entwicklungszeiten der diinischen Carabiden. - Bangsholt, F., 1983: Sandspringernes og lebebil­ Ent. Meddr 20: 277-560. lernes udbredelse og forekomst i Danmark ea. Lauterb~ch, A.W., 1964: Verbreitungs- und akti­ 1830-1981. Dansk faun. bib!. 4. Kebenhavn. vitlitsbestimmende Faktoren bei Carabiden in Boer, P.J.den, 1970: On the significance of disper­ sauerlandischen Wiildern. - Abh. Landesmus. sal power for populations of carabid-beetles Naturk. Miinster26, (4): 1-103. (Coleoptera, Carabidae). - Oecologia (Berl.) 4: Lindroth, C.H., 1945: Die Fennoskandischen l-28. Carabidae. I. - Gi:iteborgs Vetensk. Samh. De Zordo, 1., 1979: Okologische Untersuchungen Hand!. (B) 4: 1-709. an Wirbellosen des zentralalpinen Hochgebirges LOser, S., 1972: Art und Ursachen der Verbrei­ (Obergurgl, Tiro!). III. Lebenszyklen und ZOno­ tung einiger Carabidenarten (Coleoptera) im tik von Coleopteren. - Veri:iff. Univ. Innsbruck Grenzraum Ebene-Mittelgebirge. - Zoo!. Jb. 118: 1-131. Syst. 99: 213-262. Drift, J. van der, 1959: Field studies on the surfa­ Refseth, D., 1980: Differences in seasonal activity ce fauna of forests. - Bijdr. Dierkde. 29: pattern and breeding time of Patrobus atrorufus 79-103. (Carabidae) in central . - Ho1arct. Ecol. Forsskahl, B., 1972: The invertebrate fauna ofthe 3:87-90. Kilpisjiirvi area, Finnish Lapland. 9. Carabidae, - 1984: The life cycles and growth of Carabus with special notes on ecology and breeding bio­ glabratus and C. violaceus in Budalen, central logy. - Acta Soc. Fauna Flora Fenn. 80: Norway.- Ecol. Entomol. 9: 449-455. 99-119. Rijnsdorp, A.D., 1980: Pattern of movement in Greenslade, P.J.M., 1965: On the ecology of so­ and dispersal from a Dutch forest of Carabus me British carabid beetles with special reference problematicus Hbst. (Coleoptera, Carabidae). - to life histories. - Trans. Soc. Brit. Ent. 16: Oecologia (Berl.) 45: 274-281. 149-179. Schjetz-Christensen, B., 1961: Forplantningsbio­ Gries, B. et al., 1973: Coleoptera Westfalica: logien hos Amara infima Dft. og Harpalus neg­ Familia Carabidae, Genera Cychrus, Carabus, lectus Serv.- Flora Fauna Arhus 67: 8-18. und Calosoma. - Abh. Landesmus. Naturk. Thiele, H.U., 1977: Carabid beetles in their envi­ Munster 35, (4): 1-79. ronments. A study on habitat selection by Hansen, V., 1968: Sandspringere og lebebiller.­ adaptation in physiology and behaviour. - Danmarks Fauna 76. Kebenhavn. Springer, Berlin.