ALCES VOL. 45, 2009 SipkO – REiNTRODUCTiON OF LARGE HERBiVORES iN RUSSiA 35 STATUS OF REINTRODUCTIONS OF THREE LARGE HERBIVORES IN RUSSIA Taras P. Sipko Institute of Ecology and Evolution RAS, Leninskii pr. 33, 119071 Moscow, Russia. ABSTRACT: Reintroductions of muskoxen (Ovibus moschatus), European bison (Bison bonasus), and moose (Alces alces) have occurred recently in Russia. Although the process of capturing and moving muskoxen was problematic in remote areas, the reintroduction of animals from Canada and the USA successfully restored this extirpated species, and the current population in northern Russia serves as a source for further transplants. European bison populations were stagnant and suffered from inbreeding in Russia prior to reintroduction of captive animals from throughout Europe. The popula- tion in Orlovskoye polesie National park has experienced population growth with improved genetic potential. Of concern is that reintroductions in other areas of Russia were unsuccessful and the global population of European bison is not improving. Moose from the penzhina River area in Russia were successfully reintroduced to the kamchatka peninsula where they were absent for >400 years. The population is growing and dispersing across the peninsula from the transplant sites, and is among the largest physically in Eurasia. ALCES VOL. 45: 35-42 (2009) Key words: Alces alces, Bison bonasus, population, reintroduction, restoration, Ovibus moschatus, Russia. The primary goal of reintroducing large herbivores in Russia is to restore biological diversity in northern ecosystems. A secondary goal is to provide a dependable and renewable food supply for residents of northern Russia. in this paper i summarize reintroduction ef- forts with muskoxen (Ovibus moschatus), European bison (Bison bonasus), and moose (Alces alces) that were undertaken for different ecological reasons and circumstances. prior to the reintroduction efforts muskoxen were extirpated, the resident population of European bison was stagnant and suffered from inbreed- ing associated with a small founder population, and moose, although increasing in certain areas of Russia, were absent for centuries from the proposed reintroduction area. Shorter and ear- lier descriptions of these efforts can be found in "Re-introduction News" a newsletter of the iUCN (Sipko et al. 2006, Sipko and Gruzdev 2006, Sipko and Mizin 2006). REINTRODUCTION OF MUSKOXEN IN NORTHERN RUSSIA Background and Approach A considerable part of Russia’s landmass borders the Arctic Ocean and has severe cli- matic conditions including long periods of cold temperatures. Remains of muskoxen discov- ered on the Taimyr peninsula were 2000-4000 years old indicating that they inhabited the region within relatively recent geological time (Vereshagin and Barishnikov 1985). previous research indicated that this region was capable of supporting >2 million muskoxen without damage to the fragile northern ecosystem. Re- introduction of muskoxen into suitable habitat was expected to have a positive impact on the ecological community because of increased utilization of vegetation resulting in faster turnover of energy at all trophic levels. Be- cause Yakushkin (1998) found that immature male muskoxen dispersed up to 800 km from REiNTRODUCTiON OF LARGE HERBiVORES iN RUSSiA – SipkO ALCES VOL. 45, 2009 36 their natal area, the plan called for herds to be reintroduced within 600-700 km of each other to encourage genetic interchange. Sites along the shoreline of the Arctic Ocean were selected for the first reintroduc- tion that occurred in 1974 when 10 animals were delivered from Canada (Banks island) to the eastern part of the Taimyr peninsula. it was successful and eventually muskoxen spread north, east, and south (putorana plato) of the Taimyr peninsula. The population was estimated at 2,500 in 2002 (Sipko et al. 2003), and was nearly 4,000 by 2005. A second rein- troduction on Vrangel island in 1975 used 20 muskoxen obtained from the USA (Nunivak island, Alaska). population growth was slow because mortality was high in the initial ac- climatization period. By 2003, the population was 750 animals (Gruzdev and Sipko 2003), and recent estimates indicate that the popula- tion has stabilized at 800-850. Additional reintroductions were achieved by relocating muskoxen from the Vrangel island and Taimyr peinsula populations (Table 1). The objective was to capture and relocate muskoxen that were 0.3-3.5 years old, however, most captured animals were 0.5 years old. Vrangel island is a nature preserve and only vehicles with low-pressure tires and snowmobiles are allowed. Once located, muskoxen were surrounded by people with dogs and selected animals were chemically immobilized with a dart gun. Sedated ani- mals were isolated from the herd and placed into containers for transportation to a hold- ing enclosure. After the capture quota was met, a helicopter transported them to another enclosure on the mainland where they were placed in individual containers, loaded onto an airplane, and transported to the reintroduc- tion site or temporary holding enclosure with local transportation equipment. The methods of capture, handling, and containment on Taimyr island were the same as at Vrangel island. A helicopter was used to locate and deliver muskoxen either to a temporary hold- ing enclosure or directly to the reintroduction site. They were usually kept in the temporary enclosure for a period before release. DISCUSSION The initial reintroduction of muskoxen to northern Russia in the 1970s proved to be successful, and set the stage for further rein- troductions in other parts of Russia (Sipko et al. 2007). in addition to those captured for subsequent reintroductions, 81 muskoxen were also captured for zoological parks and domes- Location Region Year Number 1st Breeding 2008 population East Taimyr peninsula krasnoyarsk 1974, 1975 30 1975 ~6500 Wrangel island Chukotka 1975 20 1977 ~ 800 Bulun Yakutia 1996 24 1999 >300 Anabar Yakutia 1997, 2000 41 2000 >150 Begichev island Yakutia 2001, 2002 25 2003 >50 Allaikhov Yakutai 2000 11 2004 64 Taas-Yrach Yakutai 2001-2003 18 2004 12 Tamma Yakutai 2002, 2003 22 2004 0 polar Ural Yamal 1997, 1998, 2001, 2003 63 1999 108 kolima Magadan 2004 22 None 20 Total 284 >8000 Table 1. A summary of the location, history, and status of reintroduced herds of muskoxen in northern Russia. ALCES VOL. 45, 2009 SipkO – REiNTRODUCTiON OF LARGE HERBiVORES iN RUSSiA 37 tication experiments. Regular surveillance of muskoxen is hampered by the remoteness of the reintroduction areas. Surveys conducted in Yakutia in 2005 indicated that 347 muskoxen were in 4 reintroduction areas, a >3x popula- tion increase in 10 years. The fastest growth rate occurred in the Allaikhov herd where 18 calves were produced in 3 years. The muskoxen population in the Bulun region split into 2 nearly equal sized herds; one herd dispersed 120 km west to the delta of the Lena River where it resides currently. Reintroductions of muskoxen in northern Russia are problematic. Difficult working conditions, remote locations, and numer- ous animal transfers with different modes of transportation meant that animals needed skilled animal care specialists to accompany them. The overall mortality rate was 10-15% during the process of capture and containment prior to release. Muskoxen from the Taimyr peninsula adapted well to relocation sites in central Sibe- ria, and those from Vrangel island established viable herds in eastern Russia. presumably, the introduction of muskoxen from 2 separate populations will have a positive impact on genetic variability that influences survival, productivity, and stability of new herds. A program for additional introductions is underway, and plans are in development to introduce muskoxen to the mountain ranges of northern Asia. REINTRODUCTION OF EUROPEAN BISON IN CENTRAL RUSSIA Background and Approach The global population of European bison has not expanded in the past 15 years, remain- ing at approximately 3,000. Small herds scattered in free-roaming populations and captive-rearing facilities typically consisted of 5-7 ancestors from the 12 original animals that were founders of all contemporary bison (Belousova 1993). These circumstances caused high inbreeding coefficients in the Lowland populations (44%) and moderate coefficients (26%) in the Lowland-Caucasian line (Olech 1998). Recent studies indicated that inbreeding occurred over a much longer period of time, thus, the actual inbreeding coefficient is probably much higher. As a result, phenotypic expressions of inbreeding depression are evident in certain populations (Sipko 2002). it is estimated that an effective population size of 500 individuals is required to preserve genetic polymorphism that enables a popula- tion to adapt and evolve in a constantly chang- ing environment to prevent extinction (Soule and Wilcox 1980). An effective population must have an adequate sex ratio and sufficient mature and sexually active animals that com- prise 25-35% of the population. Thus, estab- lishing a population of 1,500-2,000 should ensure long-term viability and survival of a species. However, at least 2 geographically isolated populations are deemed necessary to reduce the risk of disease or unforeseen events that might decimate a population. Russia has sufficient geographical area of suitable habitat to accommodate multiple distinct populations of European bison. Two areas of appropriate size and ecological conditions were selected for the reintroductions; importantly, they also offered protection as designated wildlife reserves. One area was in the European broadleaf forest of the Bryansk-Oryol-kaluga region in the European (i.e., central) part of Russia. it was a large, contiguous forest tract extend- ing from the boundary of Ukraine along the Desna River (Black Sea basin) northeast to the Oka River (Caspian Sea basin), and had few natural or artificial barriers to impede bison movement; railways and highways bisected the region in only one location. The forest was 30-50 km wide and stretched >400 km. The eastern section was known as the Oka Defense Line of the State of Moscow during the 14th-17th centuries. These types of frontier forests acted as a line of defense REiNTRODUCTiON OF LARGE HERBiVORES iN RUSSiA – SipkO ALCES VOL. 45, 2009 38 until the middle of the 18th century and were strictly protected from harvesting and use by people. As a result, this tract of contiguous broadleaf forest was one of only a few areas that remained largely intact in its natural state, and is a protected natural area. Designated sections that contributed to this forest tract included 1) the Desnaynsko- Starogutsky National park (Ukraine) with 162 km2 area situated on the boundary of Ukraine and Russia, 2) the Bryansky Les Biosphere Reserve with 1230 km2 in the Bryansk region of Russia, and 3) the adjacent protected areas of the Oryol and kaluga regions in the north including Orlovskoye polesie National park (777 km2), kaluzhskie Zaseki State Nature Reserve (185 km2), and Ugra National park and Biosphere Reserve (986 km2). The second area used to reintroduce bison was the Ust-kubenskoye Hunting Facility and surrounding region located about 400 km north of Moscow in the Vologda region (Vologodskai oblast) on the 590 N latitude parallel. The landscape is a series of raised terrace plains at 110-200 m elevation within the Severnai Dvina River drainage; 64% was forest dominated by conifers (55%, mostly Abies spp.). The Ust- kubenskoye Hunting Facility is 260 km2 with the Russkii Sever National park (1664 km2) at the western boundary. The southern border begins at the shore of Lake kubenskoe, and the northern and western boundaries adjoin 8,000 km2 of federal forest lands. This area was considered optimal because European bison have survived there long-term without human support. Further, resident animals have shown evidence of twinning that is uncom- mon in bison, suggesting high habitat quality. Logging has produced large areas of second- growth forest with a shrub-layer suitable for foraging, and most agricultural areas were abandoned and these regenerating lands also provide rich food resources for bison. Bison used in the reintroduction were from captive breeding centers in Russia and West Europe (Table 2) to potentially enhance their future genetic viability. Their gene pools were quite distinct because bison from Russia and West Europe have been isolated for almost 100 years. Bison were transported in group or individual containers to temporary enclosures and released 1-2 months later. Location Region Year Number 2008 population Comments Cherga Mountains Altay 1982-1984 12 34 Orlovskoye pollesye National park (OpNp) Oryol 1996-2001, 2006 75 143 Successful kaluzhskie Zaseki State Nature Reserve kaluga 2001 0 NA Dispersal from OpNp petrovskoe Hunting Facility kaluga 2,007 9 9 Ust-kubenskoye Hunting Facility Vologda 1991, 1994 5 24 Vilikoozerskoe Hunting Facility Vladimir 1989, 1994, 2002, 2004, 2007 25 15 4 bison present at 2nd transplant Muromskij sanctuary Vladimir 2001-2004 13 16 Sknjatinskoe Hunting Facility Tver 1986, 1991 33 3 Unsuccessful Branskij Les State Nature Reserve Bryansk 1999-2000 11 0 Unsuccessful Total 183 >225 Table 2. A summary of the location, history, and status of reintroduced herds of European bison in northern Russia. ALCES VOL. 45, 2009 SipkO – REiNTRODUCTiON OF LARGE HERBiVORES iN RUSSiA 39 DISCUSSION Our method of transporting bison over long distances in individual containers proved successful. Based on our experience, using large containers containing several animals, as done with cattle, was problematic and should be avoided if possible. Bison often injured each other during group transport, resulting in high injury and mortality rates during the reintroduction effort. The bison population in the Orlovskoye polesie National park has the greatest genetic potential compared to other bison groups in the world (Table 3). The population is growing rapidly with 20 calves born in 2005. Animals have dispersed to areas adjacent to the park and regularly appear in the kaluzhskie Zaseki State Nature Reserve. it appears that 3 separate herds have formed from the original popula- tion. Additional releases into these areas are needed in order to quickly establish optimally sized populations. The region situated between the Volga and Oka Rivers contains a large bison population. However, the region has much industry and transport and communication lines and this lowland area has little coniferous forest. The Vilikoozerskoe Hunting Facility, Muromskijj Sanctuary, and the Sknjatinskoe Hunting Fa- cility located here have insufficient area for further expansion of the population. Slow population growth and numerous mortalities are evident, and further reintroductions are not considered worthwhile. The release of bison into the Bryansky Les State Nature Reserve proved to be unsuccess- ful. Long migration patterns and poaching in the Russia-Ukraine border areas resulted in their demise. There is need to supplement the bison population in the Ust-kubenskoye Hunting Facility. Also, the introduction of captive bison from the Netherlands into the Bukovina population in East Carpathian had limited success. The introduced males were unable to compete during the rut with local bulls native to the rugged mountain conditions. it is concerning that European bison numbers are not increasing worldwide. The European bison pedigree book (2002) noted that overall growth was weak with 172 births and 112 deaths overall. Arguably, there are insufficient animals to successfully establish new viable populations (Sipko and kazmin 2004). A new reintroduction effort in Yakutia will focus on a captive breeding and release program. in 2006, 30 wood bison were do- nated and transported from Alberta, Canada and relocated in a fenced enclosure 120 km from the city of Yakutsk. Successful births occurred in 2008 (6) and 2009 (7); 4 of the original animals have died. The long term plan is to establish a free-ranging population in Yakutia through gradual release of young bison. REINTRODUCTION OF MOOSE TO THE KAMCHATKA PENINSULA Background and Approach The kamchatka peninsula has been occupied and developed by Russians since the 17th century with no evidence of moose Country Source Number Russia prioksko-Terrasnyjj zapovednik 21 Okskijj zapovednik 24 Zoo Rostov na Donu 1 Zoo St. peterburg 1 Belarus Belovezhskaja pushha 2 Netherlands Natuurpark Lelystad 14 Germany Springe 6Zoo Dortmund Tierpark Chemnitz Switzerland Tierpark Dahlholzly 4 Wildpark Langenberg Finland Zoo Helsinki 1 Belgium Han-Sur Lesse 1 Total 75 Table 3. Origin of European bison transplanted in the Orlovskoye pollesye National park, Russia. REiNTRODUCTiON OF LARGE HERBiVORES iN RUSSiA – SipkO ALCES VOL. 45, 2009 40 inhabiting this region during that time. The wildlife of kamchatka is low in diversity when compared to the mainland and both lynx (Felis lynx) and squirrel (Sciurus vulgaris) appeared only in the 20th century (Valentsev and Mosolov 2004). However, archeological evidence indicates that moose were present during the 11th-16th centuries in southern and eastern areas of the peninsula (Vereschsgin and Nikolaev 1979); this information lead to the interest of reintroducing moose on the kamchatka peninsula. Moose have continuously inhabited north- eastern mainland Russia, but populations have been low in recent centuries. Growth of these populations has been documented only since the mid-20th century. The human popula- tion has been localized in small settlements leaving vast tracts of land without hunting or poaching pressure. Further, the whole region was involved in a wolf extermination (poisoning) program. As a consequence, the moose population in the mountain taiga sec- tor of the penzhina River Basin expanded to 2000 animals by 1974 (Fil 1975), thus was considered a suitable donor population for the reintroduction. After explorating the peninsula, the kam- chatka River valley was considered most suit- able for moose. To the east, west, and south were mountain ranges that protected this area from deep snow, and food resources in the val- ley appeared adequate for moose. Although the southern kamchatka region had considerable food resources for moose, deep snow and high risk of poaching were considered problematic. Snow depth frequently reaches 120 cm but is not uniform throughout the region because of the local effect of wind. Because many rivers do not freeze due to frequent thaw cycles and volcanic heat sources, the southern area of the kamchatka peninsula was considered to have highest potential as moose habitat. The reintroduction occurred in 2 stages, each with unique characteristics. The first stage was in 1977-1988 when 63 moose were captured along the tributaries of the penzhina and Belaya Rivers (pavlov 1999) and subse- quently moved to the kamchatka River valley in the central part of the peninsula. Captures were selective for 9-month old calves that were caught with the aid of a heli- copter; they were pushed toward the treeless part of the bottomland and immobilized by darting from the helicopter. They were trans- ported by helicopter to an enclosure where they remained 5-7 days, after which they were im- mobilized and placed in individual containers and transported by helicopter to the release site; transport took approximately 9 h. Moose were kept in an enclosure at the release site for 15 days to allow them to recover and acclimatize to the local environment; they were released immediately if snow depth exceeded 60 cm. The first birth was documented in 1979. The second stage of the reintroduction oc- curred in March and April, 2004-2005 (Table 4) when 26 moose were captured, transported, and released in the southern part of the kam- chatka peninsula. Most were 11 months old; 4 animals were 2 years of age. Captures oc- curred in the kamchatka River basin where tall forest cover reduced the effectiveness of the helicopter. When moose were detected, the helicopter drove them towards openings in the forest where snowmobiles were used to overtake the animals. They were immo- bilized, hobbled, and transported by sled to an individual holding container where they remained until fully recovered (1-3 h), then loaded into the helicopter and transported to the reintroduction site; transportation time was 5-6 h. They were immediately released in the Golygina and Udochka River valleys. Most moose were released on thermal terrain that was snow-free during winter because of volcanic heat. A cow moose with accompany- ing calf was observed in autumn 2005. DISCUSSION A stable moose population appears to be established and a few animals have even ALCES VOL. 45, 2009 SipkO – REiNTRODUCTiON OF LARGE HERBiVORES iN RUSSiA 41 dispersed to the western coast of kamchatka. A census conducted in the central kamchatka region in 2004 estimated the population at 1698-1775 animals (Sipko et al. 2004). Deep snow exceeding 150 cm in winter 2004-2005 caused considerable concern about winter mortality of released moose. However, they overwintered successfully with only a single mortality to a bear (Ursus arctos) the follow- ing spring. This successful reintroduction of moose is important for developing tourism and hunting on the kamchatka peninsula. Steady population growth has been real- ized in the southern area of the peninsula where twinning predominates. physical measure- ments of 32 moose indicate that individuals are realizing larger growth than their donor population from central kamchatka. it is pre- sumed that there is optimal forage production and availability on the volcanic substrate, and that moose are using accessible marine grass (seaweed). However, deep snow is of concern relative to selective and heavy browsing pres- sure during winter. Moose inhabiting the kamchatka pen- insula are among the largest of all Eurasian specimens, and this seems characteristic of the local population. Body weight of large bull moose ranges from 600-750 kg, outside spread of antlers are 161.5-181 cm (n = 6; the largest bull was killed in 2002 in Ust-kamchatskyi district), and 11 month-old calves weigh 220-325 kg (n = 10). The large size of these animals posed problems during immobiliza- tion, handling, and transport. increased drug volumes created considerable difficulty dur- ing recovery from drug-induced shock, and their large size caused handling problems during translocations by helicopter. Moose appeared to be more sensitive to capture and transportation problems than muskoxen and European bison; much time and effort was spent in recovering them to a normal physical state. One important factor lending success to the second stage of the reintroduction was that the moose had improved resistance to capture and handling stress. in previous ef- forts capture mortality often exceeded 50%, whereas mortality was insignificant in the second stage of the reintroduction. ACKNOWLEDGEMENTS My most sincere thanks are extended to V. G. Tikhonov and S. S. Egorov for the Source Year Release Area Number of Moose 2008 population Total Female penjinskii: palmatkina, Essoveem, Chichill, and Belaya (White) Rivers 1977 Milkovskii 4 3 1978 Milkovskii 9 5 1979 Milkovskii 12 5 1980 Milkovskii 12 4 1981 Milkovskii 26 14 1982 Milkovskii and Elizovskii ~2000 Chukotka: Anadyr River valley 1988 Smirnihovskii: Sakhalin island region 10 7 10-15 kamchatka peninsula: Milkovskii area 2004 Ust’-Bolsheretskii 11 6 2005 Ust’-Bolsheretskii 15 7 >45 Table 4. A summary of the location, history, and status of moose reintroductions on kamchatka pen- insula, Russia. 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