Home range patterns of male fallow deer - Springer Link

Acta Theriologica 49 (3): 393–404, 2004. PL ISSN 0001–7051

Home range patterns of male fallow deer Dama dama in a sub-Mediterranean habitat Sara DAVINI, Simone CIUTI, Siriano LUCCARINI and Marco APOLLONIO*

Davini S., Ciuti S., Luccarini S. and Apollonio M. 2004. Home range patterns of male fallow deer Dama dama in a sub-Mediterranean habitat. Acta Theriologica 49: 393–404. From 1996 to 2000 the home ranges of 14 male fallow deer Dama dama (Linnaeus, 1758) were studied in the San Rossore Preserve (Italy) using radio-telemetry. Mean size of annual home ranges was 588.9 ± 278.9 ha, calculated by MCP, and 337.5 ± 178.9 ha, using Kernel method, and was larger than that reported in published literature to date. The size of the seasonal home range estimated with the MCP method was 90.6 ± 129.1 ha during spring, 73.7 ± 67.9 ha in summer, 465.0 ± 230.6 ha in fall, and 65.6 ± 60.6 ha in winter. The Kernel method gave 84.7 ± 140.2 ha in spring, 61.3 ± 64.6 ha in summer, 306.0 ± 170.5 ha in fall, and 46.5 ± 44.0 ha during winter. The seasonal analysis suggested that bucks tended to occupy the same particular area from winter to summer, which was related to rich trophic resources, even despite of anthropic disturbance. During autumn, males reached the rutting site (a lek) that was 4 km distant from the areas occupied during the other three seasons. The lekking behaviour was the main factor influencing home range size. Dipartimento di Zoologia ed Antropologia Biologica, UniversitB degli Studi di Sassari, Via Muroni 25, I-07100 Sassari, Italy, e-mail: [email protected] Key words: Dama dama, home range, radio-tracking

Introduction Evaluation of home range sizes has an important role in ecological research and allows for the understanding of spatial behaviour of a species. The spatial behaviour of mammals is influenced by several factors: metabolic needs, body mass, feeding habits, and mating system (McNab 1963, Harestad and Bunnell 1979, Cameron and Spencer 1985, Lindstedt et al. 1986, Clutton-Brock 1989, Sandell and Liberg 1992). Despite its extensive geographical distribution, comparable to those of red deer Cervus elaphus and roe deer Capreolus capreolus, little work has been devoted to the ecology of fallow deer Dama dama (Linnaeus, 1758) and only two publications have concentrated its spatial behaviour (Rand in Putman 1986, Nugent 1994). On the contrary, many papers have dealt with the use of space by the other two western European cervid species (red deer: Georgii 1980, Georgii and Schröder 1983, Catt and Staines 1987, Carranza et al. 1991, Koubek and Hrabe

* Corresponding author [393]

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1996, Cuartas et al. 2000; roe deer: Bideau et al. 1983, Cederlund 1983, Maublanc 1986, Danilkin 1996, Van Laere et al. 1996, Andersen et al. 1998, San José and Lovari 1998). Hence, it could be useful to study space use of fallow deer in Mediterranean and sub-Mediterranean habitats, its native environment, to provide a basic knowledge about this species. Moreover, although some studies have dealt with mating system of fallow deer (Espmark and Brunner 1973, Braza et al. 1986, Langbein and Thirgood 1989, Moore et al. 1995), also with reference to the lekking behaviour (Clutton-Brock et al. 1988, Apollonio et al. 1989, Apollonio et al. 1992), no studies have shown the use of space by males adopting this latter mating system. This work reports data on home ranges of male fallow deer in a sub-Mediterranean habitat over a four-years period. In our study area, the fallow deer adopts a mixed mating system: males in this population could defend territories located in the western side of the Estate that were either single, clumped in leks, or satellite to leks (Apollonio et al. 1992, 1998a). Males remain at the rutting site from the first days of September untill the last days of October (Apollonio et al. 1992). The aims of the present paper are: (1) to evaluate male fallow deer annual and seasonal home range sizes; (2) to evaluate the influence of lekking behaviour on male home range size, considering that males had to extend their ranges in order to reach the rutting site during the rutting period; (3) to investigate other possible ecological influences of the male home range size.

Study area This research took place within the San Rossore Preserve (43°43’N, 10°19’E). The 4654-ha study area lies along the Thyrrhenian Sea coast, west of Pisa, Italy. It is bounded by the Serchio river to the north, the Arno river to the south, the Thyrrhenian Sea to the west, and is fenced on the east. The soil is sandy and the ground is flat. The area available to the deer consists of open or bushy, grassland, deciduous and mixed woods (Quercus spp., Fraxinus spp., Populus alba, Alnus glutinosa), pine woods (Pinus pinea and Pinus pinaster), and marshes; the remainder is sandy littoral vegetation. Ten different types of vegetational cover are present at San Rossore, indicating a heterogeneous environment (Fig. 1). This diversity is confirmed by a Shannon index value of 2.69 (maximum possible value = 3.32). Some typologies are more represented, especially mixed deciduous forest and domestic pine, though threy are fragmented throughout the study area, as are others. The eastern side of the Estate (a 549.5-ha area, Fig. 1) is characterized by intense anthropic disturbance caused by the presence of farmers, workmen and horse riding. Moreover the two entrances to the Estate, the two inhabited areas, and the main road are localized in this part of the study area. In our study site, analyses of the diet of adult male fallow deer were made through the identification of the rumen contents (Bruno and Apollonio 1991), and habitat selection of adult males were evaluated by direct observation (Apollonio et al. 1998b). Following these results, it is to be noted that the disturbed area contains almost only cover types selected by deer (deciduous mixed woods, meadows, and oak plantations) subdivided on few fragments (Fig. 1). The disturbed area proved therefore to be a more homogeneous and less fragmented environment (Shannon index value = 1.99) than the other areas of the Estate (Shannon index value excluding disturbed area = 2.68). The climate is sub-Mediterranean, with mild winters and warm, summers. Large predators are absent and the wild boar Sus scrofa is the only other ungulate. The estimated populations were 973, 1256, 870, and 799 in the springs of 1997, 1998, 1999, and 2000, respectively.

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Material and methods This study was performed from April 1996 to March 2000. A total of 14 male deer were radiolocated during this period by discontinuous radio-tracking. During the yearly deer control operations in the Preserve, deer were caught, aged by tooth eruption, wear (Lowe 1967, Chapman and Chapman 1997), weighed and measured. Animals were then eartagged, radiocollared and released. Ten male deer were tagged in this way in 1996–1997, but one of them died in 1997. Another four deer were radiocollared in 1997–1998, so the most comprehensive data were collected in 1998–1999, when 12 males were radiolocated. Because 1 deer disappeared and 4 males died (natural death), in 1999–2000 tracking was focused on 7 animals. Televilt transmitters worked on the 151 MHz waveband. Televilt RX-8910HE receivers, and four element hand-held Yagi antenna were used. The location of radiocollared animals was determined by triangulating bearings obtained from three different reference points (Springer 1979, Kenward 1987, White and Garrott 1990) with the “loudest signal” technique (Springer 1979). Each month, at least 12 locations per animal were obtained, uniformly distributed over the 24 hours. Subsequent locations were at least 12 hours apart. Locations were plotted onto a 1:10 000 digitized scale map of the study area (Springer 1979, Kenward 1987). Accuracy of fixes was determined in the field by placing test transmitters in various habitats and taking fixes on this (Harris et al. 1990). Accuracy of bearings within the central “telemetry area” (Cederlund 1983) was less than ± 25 m for fair signals. B

Dis turbed area PP

1 km

MP W PW

P DW

F enced areas

ME DCZ

DMW

OP MA

Fig. 1. Study area map showing fragmentation of different types of vegetational cover: PW – pine woods, DMW – deciduous and mixed woods, ME – meadows, PDW – mixed pine and deciduous woods, MPW – maritime pine woods, MA – marshes, DCZ – degraded coastal zone, B – beach, OP – oak plantations, PP – poplar plantations, and localization of fenced areas and of disturbed area.

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Home range size was determined for whole years (from April to March each study year = annual range) and for seasons within each year (seasonal range). The different seasons were made up of the following months: winter = December–February, spring = March–May, summer = June–August, autumn = September–November. Using Ranges V software (Kenward and Hodder 1996), both the Minimum Convex Polygon (MCP; Mohr 1947, Southwood 1966) and the Kernel method (Worton 1989) were used to calculate annual and seasonal home range size, using 95% of available locations. We then calculated home range overlaps between seasons (MCP 95% polygons); overlaps were also calculated between seasonal and annual home range of subsequent years. The linear distance between successive seasonal and annual range centre of activity, as calculated by Kernel method (Kenward and Hodder 1996), was determined. Using MapInfo Professional 5.0 (MapInfo 1998), percentage of fixes of male fallow deer made in the disturbed area (both during and outside the rutting period) were evaluated. To evaluate the level of significance of the results, non parametric statistics were used (Sokal and Rohlf 1995). The SPSS 11.0 program (SPSS 2001) was used for statistical analysis. Using data for each year separately, Friedman tests were used to test variation of home ranges sizes and of distances between centres of activity, and Wilcoxon tests to compare a specific season with each other when significant variation was found. In all tests significance was set at p < 0.05 and was accordingly corrected, using the sequential Bonferroni method (Rice 1989, Sokal and Rohlf 1995), for Wilcoxon tests.

Results Annual home ranges

Thirty-three annual home range sizes were estimated (Table 1). Annual home ranges were not significantly different over the 4 years (Friedman test: n = 5, df = 2 2 3, c = 6.360, p = 0.095 using MCP; c = 5.400, p = 0.145 using Kernel). The two methods gave considerably different results for annual home range size, with the MCP estimates being larger than the Kernel ones (Fig. 2). These differences were statistically significant for 3 of 4 years: in the 1st year (Wilcoxon test: z = –2.547, n = 9, p = 0.011), in the 2nd year (z = –2.197, n = 7, p = 0.028), and in the 3rd year of study (z = –2.803, n = 10, p = 0.005). Home range size varied considerably among individuals in 1999–2000 when two of seven males had ranges less than one fifth larger than the other five. In 1999–2000, males no. 530 and no. 0 (their annual home range being, respectively, 60.8 ha and 33.6 ha, using MCP 95%, 59.1 ha and 25.8 ha, using Kernel 95%), unlike all the other monitored males, did not reach the rutting area in fall.

Table 1. Male fallow deer annual home ranges (mean ± SD) recorded in the San Rossore Estate from 1996 to 2000. n – number of bucks. Annual home range (ha) Method

MCP 95% Kernel 95%

1996–1997 n=9

1997–1998 n=7

1998–1999 n = 10

1999–2000 n=7

703.0 ± 229.0 445.0 ± 196.2

541.2 ± 134.1 274.4 ± 87.1

691.9 ± 331.2 366.3 ± 167.2

342.8 ± 228.2 221.7 ± 176.1


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Fixes MCP 95% Kernel 95% Lek

1 km

Fig. 2. Example of annual home range of a male fallow deer (male no. 2 monitored in 1999–2000) evaluated using both MCP 95% and Kernel 95% methods.

Annual home range overlap (%)

100

75

50

25

1s t 1s t/4 th 4t h/ 1s t 2n d/ 3r d 3r d/ 2n d 2n d/ 4t h 4t h/ 2n d 3r d/ 4t h 4t h/ 3r d

rd

d/ 3r

1s

t/3 1s

d/ 2n

1s

t/2

nd

t

0

Pairs of years Fig. 3. Annual home range overlaps (MCP 95%) between years (1st – 1996–1997, 2nd – 1997–1998, 3rd – 1998–1999, 4th – 1999–2000). Box-plots represent the interquartile range, which contains the 50% values. The whiskers represent the highest and lowest values containing the 80% values, excluding outliers (o) and extreme values (+). The line across the box indicates the median.

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Moreover, both males showed a considerable reduction in their home range size during autumn (for male no. 530 it was 4.3 ha and 2.5 ha; for no. 0, autumn home range size was 8.5 ha and 7.9 ha, MCP and Kernel methods, respectively). Excluding these two males, in 1999–2000 home ranges averaged ± SD 461.1 ± 129.8 ha (MCP 95%) and 293.4 ± 154.5 ha (Kernel 95%) For all males, each annual home range was overlaid on each home range of the other 3 years to determine the extent of variation of the areas occupied from 1996 to 2000 among individuals: mean overlap was 73.8 ± 24.1%, showing a high degree of site-fidelity over the 3-year period (Fig. 3). Annual centres of activity were close to each other: mean distances between the ranges were 322.9 ± 463.5 m, and there 2 was not a significant trend over the years (Friedman test: c = 5.158, n = 5, df = 2, p > 0.050). Seasonal home ranges

Size of seasonal home ranges showed a characteristic pattern every year (autumn > spring > summer = winter). One hundred and forty-seven home ranges were calculated using both methods (MCP and Kernel, Fig. 4). Friedman test showed that home range size vary between seasons during the first 3 years 2 2 (1996–1997: n = 8, df = 3, c = 17.550, p = 0.001 using MCP, c = 16.950, p = 2 2 0.001 using Kernel; 1997–1998: n = 7, df = 3, c = 14.829, p = 0.002 using MCP, c 2 = 16.200, p = 0.001 using Kernel; 1998–1999: n = 10, df = 3, c = 19.224, p < 0.001 2 using MCP, c = 17.400, p = 0.001 using Kernel). On the contrary, in 1999–2000 2 the size of the seasonal ranges did not undergo significant change (n = 7, df = 3, c 2 = 2.486, p = 0.478 using MCP, c = 4.543, p = 0.208 using Kernel), even though it

Home range size (ha)

600

M CP 95% K er nel 95%

300

0 Spring

Summer

Autumn

Winter

Seasons Fig. 4. Male fallow deer seasonal home ranges evaluated using both MCP 95% and Kernel 95% methods. Data from 1996 to 2000: in 1999–2000 males no. 0 and 530 were excluded.


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followed the same trend. This result was again affected by the presence of the two individuals (no. 0 and 530) that did not reach the lek during autumn 1999. The differences among the seasonal home range were significant when they were 2 2 excluded (n = 5, df = 3, c = 9.240, p = 0.026 using MCP, c = 10.680, p = 0.014 using Kernel). Seasonal home ranges were compared within each year: using both MCP and Kernel methods, from 1996 to 1999 autumn ranges proved significantly larger than winter (Wilcoxon test: 7 < n < 11, –2.934 < z < –2.521, 0.003 < p < 0.018), spring (7 < n < 11, –2.934 < z < –2.521, 0.003 < p < 0.025), and summer ones (7 < n < 12, –3.059 < z < –2.366, 0.002 < p < 0.018). No difference was found comparing spring, summer and winter range sizes (7 < n < 11, –1.820 < z < –0.140, p > 0.050 in all instances, using both MCP and Kernel methods). The two methods gave considerably different results for autumnal home range size, with the MCP estimates being larger than the Kernel ones. These differences were statistically significant for 3 of 4 years: in the 1st year (z = –2.240, n = 8, p = 0.025), in the 3rd year (z = –2.903, n = 12, p = 0.004), and in the last year of study (z = –1.960, n = 8, p = 0.050). For each deer the seasonal home range was overlaid on the others home ranges within the same year. From 1996 to 2000 the largest overlaps obtained were those of spring, summer, and winter on autumn home range (60.0 ± 26.1%, 61.6 ± 30.3%, 70.4 ± 23.4%, respectively). This was expected, as autumn range sizes were by far the largest of all four seasons. During the first 3 years, the lowest values of overlap were obtained for autumn ranges on the other seasonal ranges (on spring 9.1 ± 6.6%, on summer 11.6 ± 12.5%, on winter 8.8 ± 6.3%), while for the last year greater values were recorded (but always < 36%). In spite of their small home range size, spring, summer, and winter ranges overlapped about 50% (spring with summer 51.2 ± 27.9%, spring with winter 60.4 ± 25.4%, summer with winter 44.7 ± 30.1%). Distance between autumn centre of activity and all the other seasonal centres was greatest (2273.0 ± 1540.7 m), while spring, summer, and winter centres of activity were similar to each other (spring–summer 602.3 ± 628.7 m, spring–winter 320.9 ± 314.6 m, summer–winter 586.1 ± 593.3 m). For each deer, in different years were overlapped home ranges of the same season (Fig. 5): areas used by male deer throughout consecutive springs, autumns and winters were consistently overlapped averaging 61.0 ± 25.9%, 68.1 ± 27.2%, 56.3 ± 30.0%, respectively. Overlap among the four summer ranges had a mean of 47.9 ± 28.8%. Differences in mean distances between centres of activity of the same season 2 from 1996 to 2000 were tested with Friedman test (6 < n < 7, df = 2, 0.667 < c < 3.739, 0.154 < p < 0.717), confirming that these distances did not vary during the study. The low values obtained for spring, summer and winter ranges suggested that their centres of activity were close (402.1 ± 388.5, 515.5 ± 394.2, 364.8 ± 386.2 m, respectively). On the contrary, distances among autumn centres of activities proved high throughout the four years (1918.6 ± 1662.5 m).

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Spring

Summer

1 km

Autumn

Winter

Study area

43°43'N, 10°19'E

Lek 1996-1997 1997-1998 1998-1999 1999-2000

Fig. 5. Seasonal home range overlaps (male no. 3, monitored from 1996 to 2000) and study area.

We evaluated the percentage of fixes made in the disturbed area. We found high percentages of fixes of males made in disturbed area in spring, summer and winter (from 1996 to 2000: 85.5% ± 17.1%), while low values were recorded during autumn (from 1996 to 2000: 35.4% ± 14.3%). All comparisons between percentages of fixes recorded in the disturbed area in autumn and the other three seasons proved to be significant (7 < n < 12, z < –2.366, p < 0.018).


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Discussion The annual home range sizes are larger than those reported in published literature to date. Nugent (1994) reported a mean annual home range, calculated by the 100% MCP method, of 309.5 ha for the male fallow deer of the Blue Mountains in New Zealand. Home range for male fallow deer in the New Forest, southern England, calculated over May–September, varied in size from 55 to 260 ha (MCP 100%), with a mean value of 107 ha (Rand in Putman 1986). Annual range sizes recorded at San Rossore are strongly influenced by the large autumn home ranges. During this season males leave the areas occupied throughout the other three seasons and reach the lek located on the western side of the Preserve (Apollonio et al. 1998a). Male fallow deer monitored during the period of study were either territorial lek males or non-territorial lek males. Irrespective to the strategy adopted, males reached the lek area during the rutting season and remained there from the first days of September to the last days of October. The display sites of males contain no significant resources, thus the autumn movement to the rutting area was solely influenced by the reproductive urge. The examples of males no. 530 and no. 0 in 1999–2000 (when they did not reach the lek) showed that to low values of autumn ranges corresponded low annual home range values. Therefore, in our study site the lekking behaviour is a basic factor influencing home range size. Consequently, the reproductive system adopted by a deer population may account for space use interpretation. The use of two distinct areas by males (during and outside the rutting period, respectively) leads to a distribution of fixes which influences home range size evaluation, according to the method used. In these cases the MCP method estimated larger home ranges than Kernel ones because it considered intermediate areas used by deer in their movements between breeding and non-breeding grounds. On these occasions the Kernel method seems to be better to evaluate actual home range sizes. Excluding rut, male deer were sedentary in all seasons. Seasonal analysis revealed small home range outside the rutting period comparable to those reported by Nugent (1994) in the Blue Mountains (New Zealand). Interestingly thirteen female fallow deer monitored at the same time of the present study showed, in spite of a smaller body size, larger annual and seasonal home ranges than males outside the rutting period (Ciuti et al. 2003). The smallest female home ranges were found during summer, averaging 126.0 ± 102.4 ha (Kernel 95%), when their movements patterns were restricted by the presence of fawn, born in late May or early June (Ciuti et al. 2003). However, during summer female home ranges resulted twice as large than male ones. These data contrast with the concept that given constant home range productivity, larger animals (in this case males with respect to females) with higher absolute metabolic requirements will use a larger home range to meet their metabolic needs.

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Why seasonal male fallow deer home ranges resulted so small outside the rutting period? Why male home range sizes proved to be smaller than female ones even if males are characterized by a higher body mass? We think this is because adult males outside the rutting period exploited areas located on the eastern side of the Preserve. These areas are constituted of deciduous woods, shrubs, and meadows providing adequate availability of food. According to Main and Coblentz (1996), polygynous male ungulates tend to engage in foraging and behavioural patterns that maximize body condition before rut. Males exploit areas where nutritious resources are abundant in all seasons, excluding rut, because they need to compete successfully for mates. On the other hand, these areas are characterized by intense human disturbance as they area inhabited by people and most frequently visited by tourists. Female deer avoided to use these highly disturbed areas (Ciuti et al. 2003). This way, males taking the risk of being disturbed, escaped the competition with females and used the best foraging grounds. In turn, the abundant food resources allowed them to use smaller ranges than females. Acknowledgements: We are grateful to the Segretariato della Presidenza della Repubblica, to the Regional Gonverment of Tuscany and to game keepers of San Rossore Estate. We thank dr J. Borkowski and two anonymous referees who made valuable comments on this paper.

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