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Abbreviations ANOVA Analysis of Variance AUC Area Under Curve Abbreviations ANOVA Analysis of variance AUC Area Under Curve Cmax Maximum concentration DRM Doramectin endectocide Macrocyclic lactone antiparasitic against both (some) endoparasites and ectoparasites epg Eggs per gram of faeces FEC Faecal egg count GABA Gamma-amino butyric acid IVM Ivermectin lpg Larvae per gram of faeces ML Macrocyclic lactone MXD Moxidectin po Per os sc Subcutaneous s.e.m. Standard error of the mean tmax Time when maximum concentration is reached after drug administration 14 Rangifer, Special Issue No. 11, 1999 England that he owned 600 reindeer, six of which Introduction were valuable decoy animals used to lure wild reindeer. Reindeer husbandry obviously developed Reindeer, the circumpolar cervid from such hunt with decoy animals (in Vorren & The reindeer genus Rangifer comprises only one Manker, 1976; Skjenneberg, 1984). species, R. tarandus living in the northern The contemporary estimate of the total Rangifer hemisphere, both in the Palaearctic (Eurasian) and population of the world is nearly 8 million animals, Nearctic (American) areas (Banfield, 1961). It half of them semi-domesticated (Staaland & inhabits most of the circumpolar land areas not Nieminen, 1993). About 20% of the semi- covered by permanent ice. The southernmost domesticated reindeer are in Fennoscandia reindeer (apart from those introduced to the (Finland, Sweden and Norway) and 75% in Russia, southern hemisphere) graze in China (50 °N) and and the rest in North America and Greenland, with the northernmost on Svalbard, Greenland and arctic some thousand wild descendants of released semi- islands of Canada, even north of 80 °N. According domesticated reindeer on Iceland, South Georgia to current systematics, there were 12 subspecies of (54°30'S, 37°0'W) and Kerguelen (48°15'S, R. tarandus, but two of them are now extinct (in 69°10'E). Small-scale reindeer herding is also Nieminen, 1994). There is not full agreement about practised in Scotland and Japan (in Nieminen, the number of subspecies, other authors (e.g. Tyler 1993b). & Røed, 1993) count 7 subspecies. The most The main reindeer product is meat, but also important reasons for the evolution of the hides possess a considerable income in Finland subspecies have apparently been the separation of (Nieminen, 1992). In Russia reindeer are still the Asian and American continents and the important draught animals, and they are also milked repeated glacial periods (in Nieminen, 1994). The and used in riding at some locations (Nieminen, Nearctic wild reindeer is called caribou. 1995). Reindeer racing is becoming popular in The semi-domesticated reindeer descends from Fennoscandia, so much so that doping tests have the wild Eurasian mountain or tundra reindeer R. t. been introduced (Nieminen et ai, 1997). Velvet tarandus. There are still some 33 000 wild moun• antlers constitute the major reindeer product in tain reindeer in southern Norway, where they are Alaska (Nieminen & Muhonen, 1996). important game animals (Skogland, 1994). As late Reindeer calves are born mostly in May and as in 1970 it was estimated that there were also 22 early June with a body weight of 4-7 kg, and grow 000 wild reindeer on the Kola peninsula, but the fast during the summer, commonly reaching body number decreased to 2700 in 1984 (in weight of 50 kg or more in September-October. Syroechkovskii, 1995). The semi-domesticated Reindeer seldom deliver twins. In many areas, reindeer has probably also genes from the wild calves are ear-marked in midsummer. To do it, the forest reindeer, R. t. fennicus, with which it readily flock must be rounded up. The next round-ups take cross-breeds. Some years ago, there were about 1 place in autumn or winter, when animals are 000 wild forest reindeer in Finland, partly in a close selected for breeding, and others are slaughtered. neighbourhood of the reindeer husbandry area Those selected to live are often treated with (Kojola, 1995), but the population is growing. A antiparasitics (Kemppainen et al., 1997). relatively recent estimate of the population of this subspecies in Russian Karelia is 6 000 to 10 000 Finnish co-operative reindeer husbandry animals (in Nieminen, 1993a). The wild reindeer The Finnish reindeer husbandry organisation was has always been a popular game animal wherever it founded in the 18th century. The basic unit, the shares habitat with humans. Reindeer hunt was reindeer herding co-operative, is called paliskunta, practised in central Europe before the last Ice Age, which name was first used in the beginning of the and in Norway there is evidence of hunt 10 000 to 19th century. Reindeer are owned by individual 12 000 years ago. herders within a co-operative, but the pastures are common. The co-operatives are mostly separated The semi-domesticated reindeer from each other my means of fences. Presently The reindeer has been crucial to the Saami and there are 56 co-operatives, 40 in Lapland and 16 in other northern peoples and cultures, enabling the the province of Oulu. The Finnish paliskunta has settlement of the barren arctic and subarctic counterparts in Sweden, sameby, and in Norway, regions. The first report on European reindeer reinbeitedistrikt, or siida. The organisation husbandry is obviously from AD 892, when the structures differ, as in the Scandinavian countries Norwegian chieftain Ottar informed King Alfred of the level of co-operation is lower, and reindeer Rangifer, Special Issue No. 11, 1999 15 husbandry is limited to the indigenous Saami ground, e.g. conservation, forestry, agriculture and people. In Finland anybody (European Economic tourism. Area citizens) living within the reindeer husbandry area and being a member of the local co-operative PARASITE FAUNA OF THE REINDEER AND has a legal right to own reindeer (Huttu-Hiltunen, ANTIPARASITIC MEASURES 1993). In Finland, there are 6800 reindeer owners, and about 800 families live primarily on reindeer A parasite is a symbiont living on the cost of the husbandry, but another 1000 get a considerable part other counterpart of the symbiosis, the host. of their living from reindeer (Anon., 1998). The Reindeer harbour a variety of different parasites reindeer husbandry area now comprises 115 000 (see Halvorsen, 1986). Although viruses, many square kilometres, or slightly over one third of the bacteria and fungi are definitively parasitic, they are country. not handled with in this thesis. So, the parasites The co-operative reindeer herding system has here are (invertebrate) animals living of the proved flexible and has continuously adapted new reindeer. Furthermore, those not practically methods to intensify production. The total number controllable with current macrocyclic lactone (ML) of reindeer in Finland was 37 000 in 1845 (in endectocides are not discussed at all. ML Nieminen, 1993a). During the first half of the endectocides have efficacy against nematodes and 1970s the number oscillated between 100 000 and arthropods, which phyla contain many important 150 000. Because of the launch of new techniques, reindeer parasites. It is worth specially noticing that such as calf slaughter, winter supplementary ML endectocides do not have efficacy against feeding, and antiparasitic treatment, as well as the cestodes, such as Echinococcus and Moniezia use of motorised herding, the amount increased tapeworms, and trematodes, such as the rapidly, reaching 200 000 to 250 000 during the Paramphistomum rumen fluke and liver flukes. The latter half of the 1980's. Between 1970 and 1976, knowledge on the efficacy of ivermectin against the annual number of animals slaughtered averaged reindeer parasite species is discussed in a later 38 000. It peaked in 1990-1991, when 160 000 chapter (page 27), and because other ML reindeer were slaughtered. After that, both the endectocides are newer products, there is no former overwintering population and the amount literature on the use of them in reindeer. slaughtered have diminished. During 1996-1997, 202 000 reindeer were counted alive, 28 000 of Losses estimated to be due to reindeer parasites which were calves. Eighty-eight thousand reindeer (gain of treatment) were slaughtered, whereof 61 000 calves (Anon., It is perhaps principally erroneous to discuss 1998). While calves contributed to less than one parasite-induced losses in reindeer, as the parasites third of the slaughter reindeer in the early 1970's, certainly belong to the ecosystem the reindeer live the current proportion is about three fourths in, almost like weather and vegetation, and are not (Kemppainen et ah, 1997). so easily separable additions. It is therefore better to Carrying capacity of the Finnish reindeer debate in terms of possible gains caused by pastures was clearly exceeded during several antiparasitic treatment. In older literature, however, decades (Kojola & Helle, 1993; Kumpula et ah, attempts to estimate the costs of parasitism were 1997). It has been extended with increased often made. Nordkvist (1967) estimated warbles supplementary feeding and pen-feeding where and throat bots to consume 15-20% of the reindeer winter pastures are scarce (Helle & Kojola, 1993; production income in Sweden, and Saval'ev (1968) Kemppainen et ah, 1997; Nieminen et ah, 1998). In approximated warbles alone to reduce the income addition, the increased slaughter of calves keeps the of the Soviet reindeer husbandry by 25-30%. overwintering population low. To minimise Thinking about such high losses of production, parasite-induced damage to the condition of the control measures might easily appear worthwhile. animals, antiparasitic treatment is used. This may There is, however, very little scientific evidence indirectly help in keeping the overwintering that organophosphate treatments would have breeding stock as small as possible. About 80% of improved the survival or weight gain of reindeer, overwintering reindeer are treated yearly with although several experiments showed a high degree ivermectin (Anon., 1993). Future challenges are in of efficacy against warbles and throat bots sustainability; balancing reindeer husbandry and its (Nordkvist 1967; Klement'eva, 1975; Nieminen et productivity needs with the carrying capacity of the ah, 1980; Persen et ah, 1982).
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