Red deer calf tagging programmes in Scotland an analysis Deer 12: 420 423. Mike Daniels and Colin McClean Deer Commission for Scotland, Knowsley, 82 Fairfield Road, Inverness IV3 5LH Introduction Since 1963 the Red Deer Commission and (after 1996) the Deer Commission for Scotland has conducted calf tagging programmes around Scotland, most recently on estates in the West Grampians (1981-1985) - W; Invermark (1986 1991) - I; Braeroy / Glenshero (1992 1999) - B and Strathconnon / Stathfarrar (2000 2001) - S. The aim of this paper is to review the information collected by calf tagging and its usefulness for deer management in Scotland. The original objectives of the calf tagging programme was to obtain information about the range of movement and the life-expectancy of deer... (Red Deer Commission, 1964). Subsequent programmes have been used to attempt to analyse deer dispersal and movement. Below we compare the results of recent programmes with those previously published in Red Deer Commission annual reports for the studies in North Ross (Red Deer Commission, 1982; 1983; 1989) and West Grampians 1967 1980 (Red Deer Commission, 1982), and evaluate the usefulness of calf tagging in contemporary deer management. Methods Calf tagging study sites were chosen to answer specific deer management questions related to movement and longevity on individual estates or within larger areas covered by Deer Management Groups. Calves were caught in June, in their first few weeks after birth. Hinds 1
were observed suckling their calves, the area noted, and when hinds moved off to feed, calves were captured in hand-held nets. Alternatively, large groups of hinds were moved from a hillside and the area systematically searched for resting calves by a team of stalkers. Small metal Ketchum cattle ear tags were inserted in each ear with a unique number, and the location and date of capture recorded (Red Deer Commission, 1964; 1989). These tags are not visible even using binoculars and are only detected once the animal has been culled. Subsequently, details of recoveries (date and six figure grid reference), from culled deer or natural mortalities, were reported to the Red Deer Commission / Deer Commission for Scotland. DATA ANALYSIS The distance between where deer were tagged and recovered (km) and the age deer were recovered (months) were natural log (x + 1) transformed prior to analysis, to normalise data (Fowler and Cohen, 1990). Residual maximum likelihood (REML) models, which allow for unbalanced data sets (Patterson & Thompson, 1971), were then used to estimate mean values. The Genstat REML option was used which approximates standard errors of the differences (SED) for parameters (Lawes Agricultural Trust, 1993). Deer tag number was used as the random effect, with area (B, I, S, W), year tagged and sex as fixed effects. Wald statistics from the REML routine were used to determine significant fixed effects and their interactions. The Wald statistic (W) was quoted along with the relevant degrees of freedom and the probability value (compared to a X 2 distribution) for the fixed effects (Lawes Agricultural Trust, 1993). Back transformed means are given with 95% confidence intervals (Zar, 1996). Correlations between transformed variables were compared using a t-test (Snedecor & Cochran, 1980) Results Between 1981-2001, 2380 red deer calves were tagged (52 % stags: 48% hinds) and 590 (25%) records of tagged animals were returned to DCS (Table 1). Thirty nine percent of 2
recoveries were stags and 61% were hinds. More deer were tagged (45%) and recovered (35%) from Invermark, than from any of the other study sites. Of the tags returned 544 had complete information from which the following results were obtained. Although this represents a large number of tags, the information is a relatively small sample of the number of animals originally tagged. The oldest stag was recovered after 151 months (12.5 years) and the oldest hind after 246 months (20.5 years) both were tagged in the West Grampians. The furthest distance recorded for a stag was 57.6 km and the furthest distance for a hind was 31.0 km both were tagged in Invermark. There was a significant effect of sex on the mean age tagged deer were recovered (w = 28.50, d.f. = 1, P < 0.001), with hinds recovered at an older age than stags (Table 2). There was also a significant effect of site (w = 20.32, d.f. = 3, P < 0.001) and year (w = 35.43, d.f. = 1, P < 0.001) on age of tag recovery, since the mean age deer were recovered was likely to be greater the earlier the year they were tagged (i.e. 1980 as opposed to 1990) and therefore deer recovered from Strathconnon-Strathfarrar were likely to be younger than those from other sites, as were deer tagged in 2000 as opposed to 1981. There were no significant interactions between site, year or sex (all P > 0.05) on the age of recoveries, suggesting that the effects of sex (i.e. hinds recovered at an older age than stags) and the link between recovery age and the length of time since tagging had taken place, were consistent across sites. There was a significant effect of sex (REML; w = 37.79, d.f. = 1, P < 0.001) on the mean distance deer were recovered from, with stags moving greater distances between tagging and culling than hinds (Figures 1 & 2; Table 2). Study site was also highly significant (w = 7.61, d.f. = 3, P < 0.001) with tags recovered furthest from calves tagged at Invermark and least from Strathconnon-Strathfarrar again related to the number of years since the tagging programme began. 3
There were no significant interactions between site, year or sex (all P > 0.05) on the distance of recoveries, suggesting that the effects of sex (i.e. stags recovered further than hinds) and the link between recovery distance and the length of time since tagging had taken place, were consistent across sites. There was a significant correlation between the natural log transformed distance stags moved and their natural log transformed age (r 200 = 0.35, t = 5.25, P < 0.01), suggesting that dispersal was age related (Figure 3). By contrast there was no significant correlation between the natural log transformed distance hinds moved and their natural log transformed age (r 344 = 0.02, t = 0.34, ns, Figure 4). Discussion The main aims of the Red Deer Commission / Deer Commission for Scotland calf tagging programmes were to investigate the longevity and dispersal distances of red deer calves from their natal territories based on tag recoveries, and to obtain known age jaws for ageing estate culls (Red Deer Commission, 1964). From the studies analysed here, an overall tag recovery of 25% was recorded, comparable to the recovery rate reported for North Ross - 30% - eleven years after the tagging programme was completed, and for West Grampians (1967 1980) 21% - one year after the tagging programme was completed (Red Deer Commission, 1982). Stags in this analysis were recovered at a younger age than hinds. This was in contrast to previous calf tagging programmes in North Ross and West Grampians (1967 1980) and a study of marked red deer in Norway, where the mean longevity of both sexes appeared to be similar (Red Deer Commission, 1982; 1989; Loison & Langvatn, 1998). The reasons for this are most likely to be related to different age selection culling policies, since all of the data collected here comes from managed populations. 4
With respect to distance moved, in all studies stags were recovered further from their tag site than hinds. In this analysis the overall mean distance moved from tag site was 5 km for stags and 3 km for hinds. This was comparable for the mean distances reported for previous calf tagging programmes in North Ross - 7 km versus 3 km - and West Grampians (1967 1980) 9 km versus 4 km, and consistent with previous information suggesting that hinds move over smaller distances than stags during their lifetime (Red Deer Commission, 1989). The analysis also provided evidence that ranging distance was age-related in stags but not for hinds, in accordance with an analysis of the data presented for North Ross and West Grampians (1967 1980). In conclusion, red deer calf tagging programmes in Scotland have consistently resulted in approximately 25% returns, due to the help and co-operation of estates and deer managers in reporting recoveries. While a large number of tags were recovered, the sample size of a quarter is relatively small and the results described should be treated with some caution, since the fate and distance moved of a large number of tagged animals remains unknown. Although longevity and movement data is constrained by both the date since marking and differences in culling policies on estates, recoveries have yielded interesting data on the extremes of life expectancy and movements, with red deer stags in Scotland recovered on average twice as far from their natal range as hinds. Calf tagging recovery programmes do not, however, provide data on daily, seasonal, annual or lifetime movements and as such their usefulness for deer management in Scotland is limited. Acknowledgements The authors wish to thank all the estates and RDC, DCS and estate staff involved in calf tagging over the years, especially: Dochie Cameron, Angus Cameron, Rhuaridh Campbell, Alan Corrigan, Iain Crichton, Ackie Dempster, Raymond Fraser, John Fraser, Jimmy Gordon, Craig Harbison, Colin Hendry, Bruce Hendry, Ronnie Hepburn, Graeme Kerr, Rory Lean, Alister 5
MacDonald, Iain Mackay, Murdo MacLean, George Macleod, Harry MacNeil, Ewen Macoll, Iain Macoll, Graeme Macrae, Lewis Macrae, Jimmy Phimster, Steven Potter,Colin Reid, Roddy Sinclair, Louis Stewart, Fred Taylor, Ab Taylor, Mike Watt, References FOWLER, J. & COHEN, L. 1990. Practical statistics for field biology. Milton Keynes: Open University Press: 83-90. LAWES AGRICULTURAL TRUST. 1993. Genstat 5 reference manual. Oxford: Claredon Press. LOISON, A. & LANGVATN, R. 1998. Short and long term effects of winter and spring weather on growth and survival of red deer in Norway. Oecolgia, 116, 489-500. PATTERSON, H. D. & THOMPSON, R. 1971. Recovery of inter-block information when block sizes are unequal. Biometrica, 58, 545 554. RED DEER COMMISSION. 1964. Annual Report for 1963. Edinburgh, HMSO. RED DEER COMMISSION. 1982. Annual Report for 1981. Edinburgh, HMSO. RED DEER COMMISSION. 1983. Annual Report for 1982. Edinburgh, HMSO. RED DEER COMMISSION. 1989. Annual Report for 1988. Edinburgh, HMSO. SNEDECOR, G.W. & COCHRAN, W.G. 1980. Statistical methods. Ames: Iowa State University. ZAR, J.H.1996 Biostatistical analysis. 3rd edition. London: Prentice Hall. 6
Table 1. Total numbers of deer tagged and recovered on the four study sites by year (B = Braeroy; I = Invermark; S = Strathconnon; W = West Grampians) total tagged by study site total recovered by study site total % recovered by study site study stags hinds total % stags % hinds stags hinds total % stags % hinds site year tagged tagged tagged tagged tagged recovered recovered recovered recovered recovered B 1992 27 23 50 54 46 3 3 6 11 11 B 1993 12 12 24 50 50 0 3 3 0 25 B 1994 14 15 29 48 52 1 1 2 7 7 B 1995 32 21 53 60 40 4 3 7 13 9 B 1996 30 41 71 42 58 1 12 13 3 40 B 1997 37 37 74 50 50 4 8 12 11 22 B 1998 42 38 80 53 48 6 5 11 14 12 B 1999 43 38 81 53 47 462 2 7 9 5 16 63 14 I 1986 99 88 187 53 47 20 37 57 20 37 I 1987 80 96 176 45 55 21 46 67 26 58 I 1988 81 72 153 53 47 24 35 59 30 43 I 1989 76 57 133 57 43 25 29 54 33 38 I 1990 109 82 191 57 43 28 31 59 26 28 I 1991 117 125 242 48 52 1082 24 59 83 21 50 379 35 S 2000 80 63 143 56 44 8 1 9 10 1 S 2001 69 63 132 52 48 275 1 0 1 1 0 10 4 W 1981 90 69 159 57 43 22 20 42 24 22 W 1982 70 76 146 48 52 14 24 38 20 34 W 1983 14 21 35 40 60 0 7 7 0 50 W 1984 64 70 134 48 52 15 21 36 23 33 W 1985 48 39 87 55 45 561 6 9 15 13 19 138 25 total 1234 1146 2380 52 48 229 361 590 19 29 7
Table 2. Predicted mean distance recovered from tag site and age recovered of red deer calves by study site and sex from REML analyses. Backtransformed means are shown with 95% confidence intervals in parentheses. Study site Braeroy Invermark Strathconnon-Strathfarrar West Grampians sex stags hinds stags hinds stags hinds stags hinds Mean age 19.1 72.6 35.9 49.8 0.0 0.0 14.3 63.9 (months) (0.0 44.8) (46.9 98.3) (10.2 61.6) (24.1 75.5) (0.0 25.7) (0.0 25.7) (0.0 40.0) (38.2 89.6) Mean distance 7.4 2.3 6.3 3.5 3.3 1.9 3.4 3.4 (km) (0.0 17.8) (0.0 12.7) (0.0 16.7) (0.0 13.9) (0.0 13.7) (0.0 12.3) (0.0 13.8) (0.0 13.8) 8
Figure 1. Location of 1234 stag calves tagged and 200 recovered between 1981 and 2001 from four study sites in Scotland. 9
Figure 2. Location of 1146 hind calves tagged and 344 recovered between 1981 and 2001 from four study sites in Scotland. 10
Figure 3. Distance recovered versus age for 200 tags from stags recovered between 1981 and 2001 from four study sites. Data are natural log transformed but axes are backtransformed labelled for ease of interpretation. 54 19 k m 6 2 0 2 6 19 54 147 months 11
Figure 4. Distance recovered versus age for 344 tags from hinds recovered between 1981 and 2001 from four study sites. Data are natural log transformed but axes are backtransformed labelled for ease of interpretation. 54 19 k m 6 2 0 2 6 19 54 147 months 12