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Home , Cervinae, Dama (genus), European fallow deer, Mule deer, Odocoileus, Oryx

SEASONAL AND CIRCADIAN CHANGES IN THE HOME RANGES OF REINTRODUCED PERSIAN FALLOW

AMIR PERELBERG,1,2Department of Zoology, Tel Aviv University, Tel Aviv 69978, DAVID SALTZ, Mitrani Department of Ecology, Jacob Blaustein Institute for Desert Research, Ben Gurion University, Sde Boqer Campus, 84990, Israel, and Science Division, Israel Nature and Parks Authority, Jerusalem, Israel SHIRLI BAR-DAVID, Department of Zoology, Tel Aviv University, Tel Aviv 69978, Israel AMIT DOLEV, Department of Zoology, Tel Aviv University, Tel Aviv 69978, Israel YORAM YOM-TOV, Department of Zoology, Tel Aviv University, Tel Aviv 69978, Israel

Abstract:The ( mesopotamica)—one of the rarest deer in the world—has been grad- ually reintroduced, using individuals from a captive-bred population, in northern Israel since September 1996. As of October 2000, >80animals were in the wild population. We studied seasonal and circadian attributes of deer home ranges to assess the success of the reintroduction in terms of behavioral adjustments to the wild. We used radiotelemetry to determine locations and analyzed home ranges with the adaptive kernel method. We defined 3 seasons: fawning (Mar–Jun), rut (Jul–Oct), and winter (Nov–Feb). For females (n= 16), rut home ranges were signif- icantly larger than winter home ranges (449±45ha [mean ±SE] vs. 384±36ha, P1,15= 0.013). During fawning, female home ranges were intermediate (424±51ha). Males (n= 5) increased their home ranges in rut season (820±162ha [mean ±SE], P<0.012) and shifted their locations toward the release point. In winter, males significantly decreased their home ranges (584±158ha, P<0.012), shifted their locations away from the release point, and almost no overlap of core areas was noticeable (1.8%overlap). In fawning (the time of casting and regrowth), males con- tinued to shift away from the release point and decreased home ranges (358±66ha, P= 0.049) with almost no over- lapping of core areas (0.06%overlap). No statistically significant differences were found between day home ranges (males [n= 5]: 621±220ha [mean ±SD], females [n= 16]: 402±164) and night home ranges (males: 482±145, females: 389±183), although day core areas tended to be larger (in all males and 12of 16females). All docu- mented aspects of seasonality in female and male home ranges are in accordance with the annual reproduction cycle, and are related to seasonal food availability. These results, combined with previous works, suggest that so far, the reintroduced Persian fallow deer have adjusted well to living in the wild and that the chances of achieving a self- sustaining wild population are good. However, further research for extended period should verify these conclusions. JOURNAL OF WILDLIFE MANAGEMENT 67(3):485–492

Key words:adaptive kernel, Cervidae, circadian, Dama mesopotamica, home range, Israel, Persian fallow deer, radio- telemetry, reintroduction, seasonality.

Reintroductions have become common proce- based on how the reintroduced adapt to dure in wildlife conservation, but often have lim- their new environment. ited success (Sarrazin and Barbault 1996). Sever- Reintroduction (as defined by the Internation- al criteria have been offered for evaluating the al Union for the Conservation of Nature and Nat- success of a reintroduction: achieving a mini- ural Resources [IUCN] 1995), in contrast to mum viable population (Beck et al. 1994), suc- translocation, is the release of captive-bred ani- cessful breeding of the first wild-born mals into parts of their former range. Because (Kleiman et al. 1991), and a recruitment rate that these animals are captive-bred and reared and are is higher than adult death rate over 3years (J. unfamiliar with the release area, their behavior in Craig and C. Reed [unpublished] cited in Cade the wild (i.e., home-range establishment, space-use and Temple 1995). All these criteria require long- patterns, and responses to seasonal and term monitoring or direct observation. Over the changes) is an important indicator of their adap- short term and for species that inhabit densely tation to the new environment. Because most vegetated areas and are not easily visible, post- reintroduced species are rare or extinct in the release monitoring must rely on remote tech- wild, often no data exist regarding their “normal” niques (e.g., radiotelemetry). Thus, other criteria behavior. However, the expected patterns of behav- must be used to evaluate reintroduction success ior can be derived from related species’ behaviors that appear to be consistent within the taxon. One important criterion in understanding the 1Present address: Department of Psychology, Univer- acclimation level of the reintroduced animals to sity of Haifa, Haifa 31905, Israel. life in the wild and the viability of the reintro- 2E-mail: [email protected] duced population is their spatial adjustments as 485 486 PERSIAN FALLOW DEER HOME RANGES• Perelberg et al. J. Wildl. Manage. 67(3):2003 expressed by different characteristics of their home-range attributes of other deer (Cervidae) home ranges (Saltz et al. 2000). Home range is species (i.e., [Dama dama], defined as the area where an animal performs its deer [ hemionus], normal activity, foraging for food, mating, and [Ozotoceros bezoarcticus], [ elaphus], nursing of offspring (Otis and White 1999). The [ capreolus], [Cervus home range is an important component of an nippon], and white-tailed deer [Odocoileus virgini- animal’s fitness because it facilitates the animal’s anus]) with similar ecological characteristics to knowledge of the area, allowing efficient assess how the reintroduced population has accli- exploitation of resources and shelters, and find- mated to the wild. We considered the existence of ing mates (Sjöåsen 1997). seasonal changes that agree with patterns found Seasonal home ranges are especially important in other deer species as an indication of a suc- (Nugent 1994) because they indicate the animals’ cessful adaptation to the wild. ability to adjust to a changing environment. In many deer species, home ranges are stable on an STUDY AREA annual basis but change in response to changes We reintroduced Persian fallow deer in the 2,400 in environmental conditions (e.g., seasons; Thir- ha Nahal Kziv Nature Reserve, along the Kziv good 1995, Leeuwenberg et al. 1997, Borokowski stream, in the upper west Galilee region, north- and Furubayashi 1998), the animals’ energetic ern Israel. The reserve is well connected by nat- status (e.g., lactation; Tufto et al. 1996) or repro- ural areas to other protected regions. Our study duction cycle (e.g., rutting; Koubek and Hrabe area encompassed the range occupied by the 1996). Even when home ranges are not stable deer, covering an area of about 220km2, between among years, clear seasonal components can be 32°95¢–33°05¢N latitude and 35°08¢–35°19¢E longi- found in the size and location of deer home tude. The area is a gradual slope from the ranges (Nugent 1994). Mediterranean shore region eastward, toward the We described the seasonal changes in the home hilly upper Galilee. A few creeks cut through the ranges of captive-bred Persian fallow deer (see hills from east to west. Except for the Kziv stream, Randi et al. 2001for species designation) rein- most creeks are ephemeral, flowing only in the troduced into northern Israel. The species, previ- winter. Artificial water resources (e.g., pipe leaks, ously abundant throughout western Asia, is cur- troughs, agricultural watering) are scat- rently endangered. In 1950, the Persian fallow tered throughout the region. Several nature deer was considered extinct. However, in 1956, a reserves, characterized mostly with Mediterranean remnant population of approximately 24individ- vegetation, are scattered in the area. Dominant uals was discovered in , from which a captive vegetation consists of open scrubland of Kermes breeding-core was established in (Chap- (Quercus calliprinos)and spiny broom (Calyco- man and Chapman 1997). In 1976, the Israel tome villosa) shrubs; dense forest of Kermes oak, Nature Reserves Authority (currently Israel Bay laurel (Laurus nobilis), buckthorn(Rhamnus Nature and Parks Authority [INPA]) established punctata), mock privet (Phillyrea latifolia), and sar- a breeding core of Persian fallow deer from 7 saparilla (Smilax aspera); and riparian zones along (2M, 5F) deer imported from Iran and Germany streams dominated by Oriental planetrees (Pla- for future reintroduction (Saltz 1996). In 1996, tanus orientalis). The settlements in the area are the reintroduction project, managed by INPA, agricultural villages (except for 1city), and about began in the upper west Galilee region, northern 20%of the land is used for agricultural orchards Israel. Bar-David (2002) estimated adult yearly (mostly apple and peach trees), fields, hen hous- survival for the reintroduced population to be es, and cattle sheds (Bar-David 2002). around 95%, and conservatively estimated repro- duction success to be 0.3fawns/female/year. METHODS Other than the reintroduced population, only 1 remnant wild population estimated at about 24 Reintroduction Process animals exists in the wild, in Iran (Karami and The reintroduction is an ongoing project that Heidemann 1994). began in September 1996. It consists of biannual Due to the rarity of Persian fallow deer, no data transfers and releases into the wild of approxi- exist on their behavior, social structure, or space- mately 12deer (6M, 6F) each time (Saltz 1998). use patterns (Chapman and Chapman 1997). We transferred Persian fallow deer from the Consequently, we compared our findings with the breeding core to an 11-ha habituation enclosure J. Wildl. Manage. 67(3):2003 PERSIAN FALLOW DEER HOME RANGES• Perelberg et al. 487 in Nahal Kziv Nature Reserve. Before transfer- The rut of the Persian fallow deer begins in ring the deer to the enclosure, we deployed August and peaks by the end of this month or the radiocollars on nearly all females (30released, 27 beginning of September. Pregnancy duration is collared) and adult males (37released, 7col- 229days on average, and parturition occurs lared). After approximately 3months in the between the end of March and mid-May. Females enclosure, we released the deer to the wild. produce 1fawn and can give birth every year (Saltz 1996). We used the annual reproduction cycle as Radiotelemetry our guideline for defining seasons. We recognized We used radiocollars with mortality sensors 3main seasons: fawning (Mar–Jun), rut (Jul–Oct), (Telonics, Mesa, , USA). Females were and winter (Nov–Feb). This definition takes into equipped with a 0.5-kg-unit (2–3yr life expectan- account the behavioral changes that appear prior cy) and males with a 1-kg-unit (5yr life expectan- to the beginning of the true season itself (such as cy). We studied the space-use patterns by tracking the behavioral changes of the females before radiomarked deer 2–3times per week, with “bio- fawning, or of the males before entering the rut). logically independent” locations (i.e., ³6hr sepa- The adaptive kernel method is sensitive to sam- ration between consecutive locations of the same ple size, and a small number of locations produce animal, as defined by Dolev et al. [2002]), from an inflated home range (Seaman and Powell September 1996until April 2000. We conducted 1996). Seaman et al. (1999) recommended using tracking with an automatic frequency scanning ³30locations, and preferably ³50. Because we receiver and a directional hand-held 3-element had <50locations per season for several animals, Yagi antenna. One person located animals by tri- we checked whether these limitations applied to angulating from 2known sites, measuring the our data. Applying lower limit without changing bearing with a prismatic compass with 1°resolu- home-range assessment reliability allowed the tion. To minimize the measuring error caused by inclusion of more animals in our analysis. We animal movement, we defined an ad-hoc limit of chose 6females with 50seasonal locations. For minimum angle difference of 20°, and maximum each animal, we generated a random sample of time of 30min between 2bearings of the same 20, 25, 30, 35, 40, and 45locations from the orig- animal (Kenward 1987, Harris et al. 1990, inal 50locations. Every random sample size was Schmutz and White 1990, White and Garrot 1990, repeated 10times to generate a distribution, and Saltz 1994). We assessed telemetry error in the then compared to the other samples. manner described by White and Garrot (1990). Because most deer are active at night as well, We calculated 2parameters: error arc and trian- home ranges based on daytime locations alone gulation location error (i.e., the distance from the might cause a misleading description of home- true location of the transmitter to the measured range size and habitat use (Beyer and Haufler location) by placing transmitters at random loca- 1994, Thirgood 1995, Kernohan et al. 1996). To test tions in the study area not known to the observer. for differences between the day and night home- range size and location, we located 21deer (5M, Home-range Estimates and Comparisons 16F) during the day and night, from September We calculated the home ranges using the adap- 1999until April 2000. We obtained 24–29daytime tive kernel method (Worton 1989). We defined and nighttime locations per individual during this the 95%isopleth of the utilization distribution as period. Although samplesize was smaller than rec- the animal’s home range, the 50%isopleth as the ommended, because the comparison itself was “core area” (CA), and the 10% isopleth as the important (and not the home-range size) we con- “center of gravity” (CG) in cases where we need- sidered our sample size sufficient(Harris et al. ed a point measure of the home range (Worton 1990, Seaman and Powell 1996). 1989, Harris et al. 1990, Seaman and Powell We compared seasonal home-range sizes using 1996). We used the HOME RANGERsoftware HOME RANGERsoftware (Hovey 1999). Season- (Hovey 1999), with 0.8coefficient for the smooth- al home ranges of 16females and 5males were ing parameter (Worton 1995). To assess differ- analyzed. Each animal had 1–3years of data col- ences among seasonal home ranges, we evaluated lection. We omitted the first 6months after home-range size and location (as measured from release, when home ranges were not stabilized the release point, which was used as a reference (Dolev et al. 2002). point) for each season, and spatial overlap To test whether the presence or absence of a between successive seasons. fawn had influence on the females’ home-range 488 PERSIAN FALLOW DEER HOME RANGES• Perelberg et al. J. Wildl. Manage. 67(3):2003 size, we used data from direct observations or berof non-orthogonal tests (e.g., for 3dependent video filming of 7females with known maternal tests: a” = 0.05/3= 0.0167; Sokal and Rohlf 1995). status in 1999(3mothers, 4barren). Every season was separately tested. RESULTS To assess changes in home-range location, we used the CG of the seasonal home ranges. We cal- General culated the distance of each seasonal CG from the After 8releases, 67(37M, 30F) captive-born release point, which was used as a reference point. deer existed in the wild, 34of which (7M, 27F) Data were collected from 17females (4–9seasons were radiomarked. In addition, an unknown of data collection each) and 4males (4–7seasons number of fawns were born in the wild. We ana- of data collection each). We tested seasonal home- lyzed 5,996telemetry locations obtained from range shifts by calculating the distance between March 1997until February 2000. We calculated CG of individuals in successive seasons and com- 339test triangulations. The error arc measured paring these values between the different season was 0°±11.3°(mean ±SD), and the triangulation pairs (e.g., winter–fawning vs. fawning–rut). To location error was 284±237m (mean ±SD). assess seasonal dispersal/clustering trends with reference to the release point, we compared the Sample Size for Home-range Calculation distance of home-range CG from the release point In 4of the 6animals, we found no difference in between successive seasons for each animal. home-range size among different numbers of loca- To measure home-range overlap between suc- tions. Two animals showed a difference (P <0.05) cessive seasons we used the IDRISI(Clark Labs between 20and 25locations to 40and 45. There- 1999) Geographic Information System software. fore, we applied the limit of ³30seasonal loca- This software produces a raster map of the sur- tions per animal as our sample size lower limit. face, projects the home ranges on the map, and then analyzes the amount of overlap between the Circadian Differences in Home-range areas. For each deer, the 2successive seasonal CA Characteristics were overlapped, and the percentage of the over- We found no differences (t-test for dependent lapping area of the second season on the first sea- samples; P4= 0.22for males, P15= 0.69for son area was measured (i.e., if the first season was females) between day home ranges (males: 621± fully encompassed inside the second season area, 220ha [mean ±SD], n= 5; females: 402±164ha, a 100%overlap was recorded). We collected data n= 16) and night home ranges (males: 482±145 from 17females (4–9seasons of data collection ha, females: 389±183ha). The day CA of males each) and 4males (4–7seasons of data collection was larger than the night CA (Friedman’s ANOVA each). To evaluate CA overlap among males, we for randomized blocks; P1<0.025). In females, we collected data for 4males during 1year only found no difference between day and night CA (Mar 1999–Feb 2000). (P15= 0.3). In both males and females, percent We used Bartlett’s c2test for homoscedasticity overlap between day and night home ranges was as a variance equality test, and c2for goodness-of- high (89±14%[mean ±SD]). In most cases (4of fit to test normal distribution fit (Sokal and Rohlf 5males and 10of 16females), the night home 1995). Where both tests were nonsignificant, we ranges were roughly encompassed by the day used parametric tests (1-way analysis of variance home ranges, within boundaries of radioteleme- [ANOVA], 1-way ANOVA with repeated mea- try error. Further results relied on this finding sures, 2-way ANOVA, t-test for independent sam- and are based only on daytime locations. ples, t-test for dependent samples, and planned comparisons when needed). Where 1of these Seasonal Differences in Home-range Size tests was significant, we used a nonparametric test Home-range variation was different among (Friedman’s ANOVA for randomized blocks, females (n= 16), but not among males (n= 5). Kruskal-Wallis ANOVA by ranks, Wilcoxon We found a difference in home ranges among matched pairs test, Mann-Whitney U-test, and seasons for both females and males (Table 1). multiple comparison by STP test). When non- For females, home-range size was larger in the parametric post-hoc comparisons were necessary, rut compared to the winter (Table 1). In all sea- we used a Bonferroni adjustment for redefining sons, home ranges of females accompanied by the significance level: a” = a/k, where a” is the fawns (n= 3) were larger than home ranges of new significance level, a= 0.05, and kis the num- barren females (n= 4; Table 2). J. Wildl. Manage. 67(3):2003 PERSIAN FALLOW DEER HOME RANGES• Perelberg et al. 489

Table 1. Seasonal home-range size (ha) of radiomarked Persian fallow deer, Israel, Mar 1997–Feb 2000 (Fawning: Mar–Jun; Rut: Jul–Oct; Winter: Nov–Feb).a

Fawning Rut Winter

n Area SE PFawning-Rut Area SE PRut-Winter Area SE PWinter-Fawning Malesb 5 358 66 0.012 820 162 0.012 584 158 0.049 Femalesc 16 424 51 0.22 449 45 0.013 384 36 0.23

PFawning PRut PWinter Among malesd 0.15 0.21 0.66 Among femalesc 0.001 0.001 0.001 Between genderse 0.67 0.006 0.55

a Each animal was radiotracked 1–3 yr. b Wilcoxon matched pairs test, Bonferroni limit: a= 0.0167. c 2-way analysis of variance. d Kruskal-Wallis analysis of variance by ranks. e Mann-Whitney U-test.

For males, home-range size in the rut was sig- between fawning to rut CG, and rut to winter CG nificantly larger than home range in both fawn- (Table 3). Males showed no significant difference ing and winter (Table 1). Comparing seasonal in the seasonal distance of CG (Table 3). home-range size between genders, we found that Female home-range CG shifts were greater males’ home ranges were larger than females’ from fawning to rut (369±94m [mean ±SE]), only during the rut (Table 1). compared with winter to fawning shift (40±34m; Wilcoxon matched pairs test;P <0.001), but not Seasonal Differences in Home-range significant compared with rut to winter shift (–31 Location ± 110m; P = 0.068). Males showed the opposite Among females with at least 3years in the wild trend. In CG shifts from fawning to rut, males (n= 10), we found a difference in the distance of showed a negative trend of dispersal (i.e., their the CG from the release point (t-test for inde- CG was closer to the release point than the CG in pendent samples;P8= 0.002). Four females never the former season; –233±144m [mean ±SE]), a changed their home-range location after it was trend significantly different from that shown in stabilized (0.9±0.1km [mean ±SE]), and 6 the other seasons (1-way ANOVA and planned females showed a gradual and directional shift in comparisons;P1,9= 0.015). their home range (2.3±0.3km), dispersing away from the release point. The second group of Spatial Overlap between Seasonal Core females was significantly younger at the time of Areas release (t-test for independent samples;P8= 0.02) For females, CA overlap between winter and than the first group (3.67±2.25yr [mean ±SD] fawning was significantly higher than in other vs. 7.75±2.06yr, respectively). We found no evi- seasons (Table 4). For males, CA overlap between dent directional shifts in male home ranges. fawning and rut was lower (and the variance was In females, distance between winter to fawning smaller) than the overlap between the other sea- CG was significantly smaller than the distance sons, but not significant (Table 4). However, only 4 males were tested, so the nonsignificant result may be due to lack of statistical power. Among males, Table 2. Seasonal home-range size (ha) of radiomarked Per- overlap of CA in each season was noticeable only sian fallow deer according to maternal status, Israel, Mar during rut (18.9±9.5%[mean ±SE]; Friedman’s 1999–Feb 2000 (Fawning: Mar–Jun; Rut: Jul–Oct; Winter: ANOVA for randomized blocks; P<0.005). Over- Nov–Feb). 2 lapof CA during other seasons was negligible n Fawning Rut Winter (0.06%overlap in fawning and 1.8%in winter). Area SE Area SE Area SE Mothers 3 559 99 724 165 572 149 DISCUSSION Barren 4 256 23 323 50 367 78 Pa 0.017 0.044 0.24 The interaction between water and energy demands, reproductive status, and resource avail- a 1-way analysis of variance. ability varies over time and space, and dictates 490 PERSIAN FALLOW DEER HOME RANGES• Perelberg et al. J. Wildl. Manage. 67(3):2003

Table 3. Average shift (m) of center of gravity (CG) of radiomarked Persian fallow deer home ranges between successive sea- sons in Israel, Mar 1997–Feb 2000 (Fawning: distance between winter to fawning CG; Rut: distance between fawning to rut CG; Winter: distance between rut to winter CG).a

Fawning Rut Winter

n Area SE PFawning-Rut Area SE PRut-Winter Area SE PWinter-Fawning Malesb 4 342 114 0.570 495 120 0.57 614 97 0.570 Femalesc 17 239 42 0.001 404 55 0.50 322 53 0.001

PFawning PRut PWinter Among genderse 0.5 0.5 0.5

a Each animal was radiotracked 4–9 seasons. b 1-way analysis of variance with repeated measures and planned comparisons. d 2-way analysis of variance.

much of the seasonal and circadian home-range the mother and the growing fawn, and possibly dynamics of deer (Putman 1988). In hot, dry also reflecting the tendency of females to search summers and cool, wet winters of the Mediter- for the mating partners. We also found that the ranean region, the home ranges of large herbi- home ranges of lactating females were larger vores are expected to be the greatest in autumn, than the home ranges of barren females, in all when resources are most limited (before winter seasons, reflecting the higher nutritional needs green-up) and when the rut takes place (for food of lactating females. availability study see Dolev 1999). Our current Male home ranges were the smallest in fawning work suggests that distinctive seasonal differences season, the time of antler casting and regrowth. exist in home-range size and location of Persian This phenomenon is familiar from other deer fallow deer that are in agreement with the differ- species (Putman 1988). ences expected in behavior of a cervid of this size Our findings that younger females tended to under the given climatic regime. As in other deer have nonstable home ranges are complementary species (Hayes and Krausman 1993), circadian to previous studies on European fallow deer characteristics of home-range location and size (Nugent 1994). Adult females of European fallow revealed that diurnal and nocturnal home ranges deer tended to be stationary and did not change are similar, and that relying on daytime locations their home-range locations between seasons or alone reliably represents Persian fallow deer years. Conversely, some adolescents had a grad- home ranges. The females’ home ranges were ual shift of home ranges and a gradual expansion smallest in the winter when food is plentiful. In in seasons and years (Nugent 1994). A previous fawning, the females’ home ranges mildly in- study of Persian fallow deer (Dolev et al. 2002) creased, possibly to account for the growing found that young females disperse faster from needs of the lactating mother but tempered by the release point than adult females. the abundance of food typical of that time of year Many generations of and low (spring). In rut, the females’ home-range sizes genetic variance of the captive-bred Persian fal- were the largest, reflecting the decrease in the low deer population (Saltz 1996) have raised con- availability of food (mid-summer to autumn), cerns regarding the ability of the reintroduced together with the increasing energetic needs of population to adjust to living in the wild. Our

Table 4. Radiomarked Persian fallow deer core areas (CA) overlap (%) between successive seasons in Israel, Mar 1997–Feb 2000 (Fawning–Rut [FR]: overlap between fawning and rut CA, Rut–Winter [RW]: overlap between rut and winter CA, Winter–Fawning [WF]: overlap between winter and fawning CA).a

FR RW WF

n Overlap SE PFR-RW Overlap SE PRW-WF Overlap SE PWF-FR Malesb 4 26 4 0.187 52 19 0.187 52 16 0.187 Femalesb 17 39 6 0.316 45 3 0.018 53 3 0.018

a Each animal was radiotracked 1–3 yr. b 1-way analysis of variance with repeated measures and planned comparisons. J. Wildl. Manage. 67(3):2003 PERSIAN FALLOW DEER HOME RANGES• Perelberg et al. 491 study was based on a radiomarked sample that troduction process is successful and that the represented 90%of released female (27/30) and chances of achieving a self-sustaining wild popu- 19%of released male (7/37) Persian fallow deer. lation are good. In polygamous species, females are considered more important for the growth of the herd, and ACKNOWLEDGMENTS thus were given more attention. The extensive We thank our colleagues D. Meir (Tap), A. Ali, data collected on almost all females, the fact that S. Maklade, Y. Maklade, and A. Laurie from the population growth fits Saltz’s (1996) expected Israel Nature and Parks Authority for their invalu- curve (according to Bar-David 2002), combined able help and support in this project. We thank with direct observational data (Dolev 1999, Bar- the 2anonymous reviewers of this manuscript for David 2002) and videotaping (Perelberg 2000), their illuminating remarks. The Persian fallow all support our findings regarding the adaptation deer reintroduction program in Israel is carried of these deer to the wild. out and funded by the Israel Nature and Parks We conclude that although little is known Authority, with additional funding from The San- about the ability of reintroduced species to Diego Zoological Society, The Schussheim Foun- acquire behavioral characteristics typical of wild dation, and the Keren Kayemet Le’Israel. populations, our results indicate that the Persian fallow deer population shows home-range ecolo- LITERATURE CITED gy patterns that are in accordance with those BAR-DAVID, S. 2002. The reintroduced population of the expected from the annual breeding cycle and Persian fallow deer—modeling population growth in time and space. Dissertation, Tel-Aviv University, Tel- seasonal conditions, and correspond to the Aviv, Israel. behavior of other deer as described in the litera- BECK, B. B., L. G. RAPAPORT, M. R. STANLEYPRICE, AND ture. Furthermore, because home-range dynam- A. C. WILSON. 1994. Reintroduction of captive born ics (e.g., size, location, stability, seasonal shift, sea- animals. Pages 265–286inP. J. S. Olney, G. M. Mace, and A. T. C. Feistener, editors. Creative conservation: sonal overlap) reflect an animal’s ability to adapt interactive management of wild and captive animals. to a novel environment and changes within it, we Chapman & Hall, New York, New York, USA. suggest that it can be used as a short-term indica- BEYER, D. E., JR., ANDJ. B. HAUFLER. 1994. Diurnal versus tor of reintroduction success. Such findings may 24-hour sampling of habitat use. Journal of Wildlife help improve future reintroduction protocol. Management 58:178–180. BOROKOWSKI, J., ANDK. FURUBAYASHI. 1998. Home range MANAGEMENT IMPLICATIONS size and habitat use in radio-collared female sika deer at high altitudes in the Tanzawa Mountains, Japan. We found that a tendency exists for older Annales Zoologici Fennici 35:181–186. females to remain relatively stationary in com- CADE, T. J., ANDS. A. TEMPLE. 1995. Management of threatened species: an evaluation to the hands-on parison to younger ones. Consequently, we rec- approach. Ibis 137:161–172. ommend that in future releases in other sites, the CHAPMAN, D., ANDN. CHAPMAN. 1997. Fallow deer: their his- first groups should consist of older females that tory, distribution and biology. Second edition. Coch-y- will establish a core population in proximity to Bonddu Books, Machynlleth, Powys, United Kingdom. the release site. After the first group is estab- CLARKLABS.1999. IDRISI: geographic analysis and image processing software. Clark University, Worces- lished, younger females can be released to ex- ter, Massachusetts, USA. pand the population boundary ranges. DOLEV, A. 1999. Population expansion in space, habitat We found that adult males tended to maintain use and vegetation influence in reintroduced Persian territoriality throughout the year, as indicated fallow deer (Dama dama mesopotamica). Thesis, Tel- Aviv University, Tel-Aviv, Israel. from the very low overlap between core areas. ———, D. SALTZ, S. BAR-DAVID, ANDY. YOM-TOV. 2002. Younger males do not present an immediate Impact of repeated releases on space use patterns of threat to the adults males’ status, and thus their Persian fallow deer. Journal of Wildlife Management gradual integration in the herd hierarchy should 66:737–746 be more natural than when an adult male is HARRIS, S., W. J. CRESSWELL, P. G. FORDE, W. J. TREWHELLA, T. WOOLLARD, ANDS. WRAY. 1990. Home range analy- released. Therefore, we recommend preferring sis using radio-tracking data—a review of problems the release of young males over adult males. and techniques particularly as applied to the study of Concerning all aspects, the reintroduced Per- . Review 20:97–123. sian fallow deer apparently have adjusted well to HAYES, C. L., ANDP. R. KRAUSMAN. 1993. Nocturnal activ- ity of female . Journal of Wildlife Manage- living in the wild. Our results, combined with pre- ment 57:897–904. vious works (Dolev 1999, Perelberg 2000, Dolev et HOVEY, F. W. 1999. The Home Ranger V1.5. Ursus soft- al. 2002, Bar-David 2002), suggest that the rein- ware, Revelstoke, , . 492 PERSIAN FALLOW DEER HOME RANGES• Perelberg et al. J. Wildl. Manage. 67(3):2003

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