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from anoxia. Drops of haemolymph were collected from a puncture in the neck, or by cutting off one of the legs. Analysis of lactate was carried out according to Strom (1949). RESULTS AND DISCUSSION The survival of beetles stored in nitrogen at O0 C for various time intervals is shown in Table 1. Although the material is limited, it appears that al1 species have a high capacity for survival under anaerobic conditions. Most of the adults of P. borealis survided 156 days in nitrogen, which is a longer period than previoiisly recorded (Conradi-Larsen & Ssmme 1973b). Both Byrrhus fiilula L. (Byrrhidae) and Otiorrhyncus dubius Strsm (Curculionidae) survived 120 days of anoxia. Only one specimen of Amara alpina (Carabidae) and a few specimens of Lepyrus arcticus (Curculionidae) were available, but the results Table 1. Survival of nitrogen at O0 C. adult beetles stored in No. of beetles Days in Dead or Species N2/0° Alive moribund Pelophila borealis 117 3 1 156 14 4 Amara alpina 51 1 0 Byrrhi~s pilz~la 30 2 O 48 4 O 58 2 O 120 2 o Lepyru~ arcticu~ 48 1 O 58 2 O Otiorrhynchus dubitis 30 2 O 58 1 O 120 3 o Table 11. Content of lactate in the haemolymph of adult beetles stored in nitrogen at O0 C. Species Conc. of Days in lactate (mM) n Nn/OO x SE Pelophila borealis 4 117 37.1 0.65 Amara alpina 1 5 1 13.3 Byrrhus pilula 4 4 8 20.2 +_ 1.44 Lepyrus arcticus 2 4 8 18.3 Otiorrhynchus dubius 1 51 46.7 indicate a high capacity to survive under anoxia in these speGes as well. During anoxia lactate was accumulated in the haemol~mph of al1 species (Table 11). The concentration varied from species to species, but appeared to be exceptionally high in O. dubius. Further investigations are necessary to see if other end ~roducts of anaerobic metabolism are formed. It is concluded that al1 of the s~ecies studied can survive extended periods of anoxia. Anaerobic metabolism during anoxia is demonstrated by the presence of lactate in their haemolymph. Since oxygen deficiencies are likely to occur in the natural habitats of the beetles, anaerobic metabolism appears to be an important part of winter survival in al1 species. As demonstrated by Pasche & Zachariassen (1973), adults of the longicorn beetle Rhagium inauisitor. which overwinters under the bark of dead coniferous trees, can survive long periods of low oxygen tension. In general it may be expected that anaerobiosis in insects is more common than previously recognized. A large number of insect species oveiwinter in moist habitats of various kinds, and may experience periods of anoxia during the winter. Further studies should be carried out on overwintering and anaerobic metabolism in species from different habitats. ACKNOWLEDGEMENTS 1 am grateful to cand. real. A. Fjellberg for identifying the specimens of Byrrhus pilula, and to Mrs. 1. Tambs-Lyche for technical assistance. REFERENCES Conradi-Larsen, E.-M. & Semme, L. 1973a. Anaerobiosis in the overwintering beetle Pelophila borealis. Nature 245, 388-390. Conradi-Larsen, E.-M. & Semme. L. 1973b. The overwintering of Pelophila borealis Payk. 11. Aerobic and anaerobic metabolism. ,Vorsk cnt. Tidsskr. 20, 325-332. Lien, L., Ustbye, E., Hogstad, O.. Haande, K. M., Haande, P. S., Hagen A., Skar, H.-J., Skartveit, A. & Svalastog, D. 19i.I. Bird surveys in the high mountain habitats at Finse and Stigstuv, Hardangervidda. South Norway, 1967-72. Norw. J. Zooi. 22, 1-14.
Anawobiosis 157 Dstbye, E. 1969. Records of Coleoptera from the to hypoxia and hypercapnia in adult Rhagium Finse area. Norsk ent. Tidsskr. 16, 41-43. inquisitor L. (Col., Cerarnbycidae). Norsk ent. Dstbye, E. & Ssmme, L. 1972. The overwintering Tidsskr. 20, 323-324. of Pelophila borealis Payk. 1. Survival rates Strom, G. 1949. The influence of anoxia on lactate and cold-hardiness. Norsk ent. Tidsskr. 19, utilization in man after prolonged muscular 165-168. work. Acta physiol. scand. 17, 440-451. Pische, A. Ek Zachariassen, K. E. 1973. Tolerante Received 30 April 1974
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250 A. Lillehammer Muller, K. 1955.
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