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?T·E R(2)ROS, NORWAY Gl.L 737 ~V\SS PROCEEQINGS OF THE J-57 SECOND INTERNATIONAL REINDEER/CARIBOU SYMPOSIUM jq(q 17.-21. SEPTEMBER 1979 ?t·E R(2)ROS, NORWAY EDITED BY: EIGIL REIMERS ELDAR GAARE SVEN SKJENNEBERG . 0 i 0 0 ·10 ! l!) 1'- DIREKTORATET FOR VILT OG FERSKVANNSFISK :i M ,I M TRONDHEIM 1979 \ -664- Reimers, E., Gaare, E. and Skjenneberg, S. (eds.) 1980. Proc. 2nd Int. Reindeer/Caribou Symp., R(l)ros, Norway, 1979. Direktoratet for vilt og ferskvannsfisk, Trondheim. POPULATION ECOLOGY OF REINDEER ON SOUTH GEORGIA N. Leader-Williams British Antarctic Survey, Madingley Road, Cambridge CB3 OET, U.K. ABSTRACT Data from an ecological study are used to evaluate differences in population trends shown by the introduced reindeer of South Georgia, and those on other islands. On St Paul and St Matthew Islands, the populations increased exponentially in number, but rapidly declined within 20 to 40 years of their introduction. The two discrete herds on South Georgia increased up to maximum densities (23 animals km2) ~imilar to those on other islands, but the increase and subsequent decline (to present densities of 3 to 13 animals km2 50 to 65 years after their introduction) has been gradual, and reindeer from one herd emigrated to colonise a new area. There is selective grazing on preferred food plants in summer (with elimination of some species), but range quality, and thus body growth, are insufficient to allow conceptions in calves, and condition fluctuates widely between seasons. In winter reindeer depend exclusively on the resilient coastal tussock grass, Poa f/abe/lata, but grazing pressure has reduced winter range carrying capacity, probably ensuring a continuing slow decline in numbers. 1. INTRODUCTION Reindeer (Rangifer tarandus L.) have been introduced to several small arctic islands which lacked predators and on which the opportunity for migration was resticted, sometimes with dramatic results. On St Paul Island (Pribilof group) and ~t Matthew Island (NE Central Bering Sea) the original small populations reached high densities, but then cr~shed to very low numbers within 40 and 20 years, respectively, of their ,<i,ntroductions (Scheffer 1951; Klein 1968). The results of these introductions are of great interest as they did riot follow the general pattern of most ungulate eruptions occurring after liberation or introduction (Caughley 1970): Norwegian reindeer were introduced to two separate areas- of the subantarctic island of South Georgia (53°30' to 55°S and 35°30' to 38°30'W) by whalers in the early part of this century. Permanent ice and.inow cover c. 60% of its surface and large sea-entering glaciers act as barriers to movement. Snow lies seasonally down to sea-level, usually from June to September. The vegetation is typical of subantarctic tundra and comprises twenty-four native vascular species, c. 200 bryophytes and c. fifty lichens together with c. fifty largely non­ persistent aliens (Lewis Smith and Walton 1975). It is dominated by the coastal tussock grass Poa f/abe/lata, and is substantially different to that encountered by reindeer in the northern hemisphere. Ten reindeer were introduced to the Barff Peninsula (Fig. J) in 1911. This herd was originally confined to the Barff area, but in 1961-65, an unknown number of reindeer first spread into the area to the north of Royal Bay (Lindsay 1973) and formed a second herd. A third herd came from a separate import of seven rein­ deer to the Busen area in 1925. Reindeer calves are born in November and the rut occurs in late March (Olstad 1930; Bonner 1958). There are no native land mammals and no further interspecific competitors (except the brown rat, Rattus norvegicus) have been introduced. The reindeer have never been husbanded or actively mana­ ged. Results from the natural experiment which their introduction has created are evaluated in the present paper. Attention is focused upon two main points of interest; firstly, the ways in which reindeer have responded to this unusual habitat, and secondly, the differences between subantarctic and arctic island populations of introduced reindeer. 2. METHODS Historical data on reindeer numbers, size and feeding habits are derived variously from Olstad (1930), Bonner (1958) and Leader-Williams (1978). The method ot.collection and numbers of reindeer shot between 1973 and 1976 for the present study are detailed in Leader~Williams (1980a); selected results from part of this sample are used in the present paper. The ages of reindeer {n,ere determined by a combination of tooth eruption patterns \ and annulations in incisor cementum (Leader-Williams 1979). The total weight of unbled reindeer (exc;:epting ' -665- Fig. 1. Map of reindeer areas of South Georgia. Stippled areas represent permanent ice and snow. loss from the shot) was taken on a 200 kg spring balance; the total length was measured from the nose, along the spine to the ba~e on the tail with the animal lying on its right side with neck extended; hind foot length was measured from the 'point of the hock to the horn line of the hoof in a straight line. Rump fat qepth and kidney fat weight of each reindeer were measured as described by Dauphine (1976) and the! percentage 0f femur marrow fat was determined from a c. 2 g sample using the method of Neiland (1970). The diet of reindeer was assessed by quantitative analy~es of rumen contents (Leader-Williams et al. 1980). Vegetated areas within the reindeer ranges were traced from monochrome aerial photographs taken in November 1973 at a height of 1600 m. These tracings were transferred to the largest available maps (scale of 1 :50000 (Barff Peninsula and Busen area) or 1:25000 (Royal Bay area)) with cross-reference to landmarks visible on both aerial photographs and maps. Both total and vegetated areas available to reindeer were measured with a planimeter, the former probably to within.± 5% and the latter to within.± 10%. Corrections were not made for sloping terrain. 3. RESULTS AND DISCUSSION 3.1. Numerical change Ungulates introduced into previously unoccupied areas normally adjust to their new environment with a single eruptive oscillation (Riney 1964). Historical estimates of total reindeer numbers on South Georgia (Leader­ Williams 1978) and accurate counts of adult and yearling reindeer numbers made from 1972 to 1976 (Leader­ Williams 1980a) are shown in detail in Leader-Williams and Payne (1980). These data are summarised in Fig. 2, together with a summary of the available population estimates for St Paul Island (Scheffer 1951) and St Matthew Island (Klein 1959, 1968). Data are available forSt George Island (Scheffer 1951) but are not shown for reasons of scale; this herd reached a maximum of c. 220 reindeer in 1921 and fluctuated slowly thereafter between seventy and ten animals. Several estimates lack accuracy, at least for South Georgia (Leader-Williams 1978). However, Fig. 2 shows that the eruptive oscillations on each island differed (a) in the maximum number attained (the Barff and Busen herds also differeQ in this respect), (b) in the number of years after introduction that these peaks occurred, and most especially (c~ in their declines. Initially introduced ungulates increase in number in response to the discreRancy between carrying capacity of the environment and the numbers actually present (Riney 1964). The total areas available to introduced -666- Tab. 1. Areas of reindeer range on different islands and for each herd on South Georgia Island/Herd Total area Vegetated area occupied by (km2) reindeer (km2 ) Barff Peninsula 131 30 Royal Bay area 58 9 Busen area 124 11 St Matthew Island 332* St Paul Island 107* St George Island 91* 2 *Converted ~rom areas (miles ) in Klein (1968) and Scheffer (1951), with 1 mile = 2.59 km2 6000 6000 BUS EN BARFF --- BARFF AND ROYAL BAY e e e e e ST PAUL ..l ST MATTHEW 5000 I~ . 4000 . 3000 :Q 3000 w lD ::; :::> z ., '· , ___ _ 2000 : 2000 • .. ' •• : ••• • .. •• •• .. 1000 • . • .. 800 . ~­ .. ,.... ..~-· ., . .. .... ., .. ...... ,. .. ... ··-··-··-··-··. ..!· 1910 1920 1930 1940 1950 1960 1970 1980 Fig. 2. Summary of reindeer numbers on South Georgia, St Paul Island (from Scheffer 1951) and St Matthew Island (from Klein 1968) reindeer, which for present purposes can be substituted for carrying capacity, are shown in Tab. 1; the areas of actual vegetated range used by each herd are known only for South Georgia. If the eruptive oscillation of each herd is described in terms of density (Fig. 3), the maxima attained were similar (18 to 23 animals km2 of total area), except forSt George Island (2.4 animals km2, not shown) and the Busen herd. Scheffer (1951) could advance no reason for the differences between St Paul and St George Islands. In the Busen area reindeer have not negotiated four mountain passes leading into potentially available, but as yet ungrazed, range within their overall area (Leader-Williams 1978). If density is calculated using only occupied and grazed areas (Tab. 2), the peak densities in the Barff and Busen herds :fe similar. These were also similar to 1976 densities in the Royal Bay herd, which at that time had probably declined slightly from maximum numbers attained after emi­ gration .to this new area (Leader-Williams 1980a). Thus miximum numbers attaihed by introduced reindeer I generally have been proportional to the area available. 1 -667- 24 20 • "I • E •• . "'.. •• . ~ 16 ••• • ·;: • • ~ • • ! • • .. •• • .. 12 .. •• • .. • • . •• • . ?- •• .. 8 . •• .. •• .. •• .. ./\ • .· ./· ·., 4 ·.. ...· ,.,. ~· ·. .. .. .·. ,·· "'.· :. o L_~~---,,_~~~··T-~·~·~-~·~·-~··r-~·~·~:~.~~··_-__··---.--~·L---,------.-- 1910 1920 1930 1940 1950 1960 1970 1980 Fig. 3. Summary of reindeer densities on South Georgia, St Paul Island and St Matthew Island. Legend as shown in Fig. 2. Barff densities omitted during formation of Royal Bay herd. Tab. 2. Densities of reindeer on South Georgia, with compara7ive data from Svalbard (animals-km2) ;~;f >· Herd Total area Occupied and grazed area Peak 1976 Peak 1976 .
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