BALTIC FORESTRY THERELATIONSHIP BETWEEN THE RED DEER AND EUROPEAN BISON I.../ 0. BIELOVA= The relationship between the Red deer and European bison over the winter period

OLGA BIELOVA Lrtlzzrarzian Forest Researdch Ir~stitzrte Girionys I, LT-4312 Katri~asdistr: Lit/1tiunia

Bielova 0. 1997. The relationship between the Red deer and European bison ovcr the winter per~od.Rnltic F'oresrt:~;2: 4 146.

Thc relations between the wide-spread game herbivores, such as the red deer, and rare herbivores, such as the European bison, which had entered the Lithuanian Red Data Book, were studied in their common habitats. All signs of animal location and movements were recorded. The methods of the selective observations, sample plots and belt transect route one were used. Red deer preferred foraging in the remote places from the location of European bisons. The supplemental feeding in winter docs not restrain the damage caused by deer on the pfeferablc spccics of trees and shrubs. Deer avoid the direct contact with bisons. Indifferent relations usually predominate in the common habitats. The bison is the nearest competitor of the deer. Becausc of thc philopatria and common wintering territory of both spccies under unfavourable foraging conditions the relations between the species arc taking the character of certain con~petitiontill aniensalisni.

Key words: Red deer, European bison, common habitat, avoidance, competition.

Introduction surrounding lands. This period was unfavourable for animals. The optimal density is 1 bison per 150 hectares Red deer are wide-spread and well-known game of forest area and 1 bison per 5-7 hectares in enclosure. animals in Lithuania. They belong to the Middle Europe The experience has shown that the breeding of free subspecies Cervrrs elaphus hippelaplirrs. There are many bisons in comparatively small stands is not real in works concerning the ecology of red deer and their Lithuania (BaleiSis et al., 1987). Because of enclosure importance connected with the impact of deer on the reconstruction in 198 1, all animals were released frorn the forest vegetation (Padaiga, 197 1,1990,1994; etc.). Deer enclosures to the forest. Then the bisons rambled about prefer the stands covering not less than I000 hectares, the forest by the radius of 4-1 1 km. Some animals were and usually step aside from the stands covering less lost. The remaining animals were enclosed again. In 1985 than 500 hectares (BaleiSis, skerys, 1984). The foraging there were 19 bisons (3 males and 16 females). It is depends on the stand characteristics, seasonal known that more than 400 species of used plants are in conditions, animal density, population indices, the the bison diet, including 50 species of trees and shrubs. habitat carrying capacity. In snowy winter shoots and The most preferable species are Populus trernzrla L., bark of preferable deciduous species comprised up to Sor.bus aircuparia L., Ft-asinus sp., Acer sp., as well 92.3% of deer diet while these of coniferous species Sali.~sp., Eonymus europaeus, Padus racenlosa Lam., constituted from 26 to 46.5% and more (Liet.Fauna ..., Q~rercusrobus L., Tilia sp. Betula sp., Picea abies L., 1988; Padaiga, 1990). Woody species constituted from Piize sylvestris L. are rare in the diet. The bisons prefer 2.5 to 25.4%, small shrubs from 13.5 to 57.6%, and herbs the shoots with diameter of 2 cm and more. The bark is - frorn 11.2 to 79.5% (Liet.Fauna ..., 1988). stripped off in the spring and winter then the air The European bison (Bison honns~rsL.) was a usual temperat~lreis not less than -1 5°C (Liet.Fauna ..., 1988). game animal in ancient Lithuania. In the XVIth century The bisons from the PaSiliai forest prefer cultural plants. the abundance of bisons diminished, and finally they In the enclosures the bisons preferred Alnus gl~itinosa vanished. The first bison breeding ground was L., Betzila sp., Fraxinzrs sp., Padzts 1-acenzosa Lam., established in 1969 in the PaSiliai forest (The Paneveiys Populus tremula L., Quercus robzrs L., Sorbus district). In 1979 there were 14 free bisons. The animals azrcuparia L.,, and other woody plants (BalejSis et al., migrated through the PaSiliai forest and pastured on the 1987). In the two enclosures usually there are 14-15

1997, VOL. 3, NO. 2 ISSN 1392-1355 = BALTIC FORESTRY THERELATIONSHIP BETWEEN THE RED DEER AND EUROPEAN BISON I.../0. BIELOVA- bisons. A total of 15 animals are in the surrounding deer density in the forest was 30 animals per 1000 forest too. The free animals adapted well to different hectares, and the total of 84 deer foraged there (Padaiga, seasonal changes and successfully survived in incle- Bielova, 1994). ment winters of recent years. However, the bison ecology has not received due consideration in Lithuania (Baleisis et a1,1987). Of importance to the works Results connected with the management of red deer population, to the improvement of the European bison herd and also The tracks of red deer concentrated near the to fi~rtherextension of studies on the bison are the sample plots with attractive foods in the northern part of questions on the relations of rare bisons with other the territory tested. There were the main moving animals such as abundant red deer. directions of deer to the yearly supple~nentalforaging places. The deer began to attend these places at the end of November when the air temperature was lower than Material and methods O°C, and it snowed a little. They avoided tlie places of free European bison location in the southern part of the Trials were established in the PaSiliai forest (the territory investigated. Here bisons moved from the Panevkiys forest enterprise) to test attraction of red deer yearly supplemental feeding places where last year the into certain places near the breeding-ground of enclosed animals had no assistance in the winter period. Every bisons. The area of forest is 2,800 hectares. The reaction day they gathered in the mentioned part near their late to the supplementa1 attractive foods was studied near the native place, namely, the enclosures. The most con- mentioned enclosures and within the track of 2 km around centrations of deer tracks were in the remote places. this place by the method of selective observation on the Within the whole tested period from November to the wintering territory of animals. The daily route observa- second part of April the deer held themselves apart the tions were carried out in 9-10 hours within the day. The bison concentration place. The behaviour of deer was total time of observations was 195 hours over the later stereotype: animals moved to the usual feeding places, autumn-middle winter period. A total of 40 observations and the solitary approaches to the attractive food were carried out every month from the second part of patches took place on the way. The supplemental January till second part of April. All signs of animal feeding places were attended by animals one after location and movements on the mentioned territory were another. While the free herd of bison always had rallied recorded including the direct contacts with animals. The in the same part of the territory, the deer began to attend attractive foods were presented in the places of yearly only at some distance, and did not come to place of the supplemental feeding (3 locations) within the territory bison locality nearer than up to 48-70 In. The use of the tested, and along the enclosure perimeter on the strip of attractive food in the northern part of the territory 100x 1 m (n=80 sample plots) every 5 meters. The route comprised up to 90% (Tables 1, 2). unit was 100x4 m. Particular transect belts were set along In the second period of the studies (January-April) the roads - forest block lines. This territory was covered sugar beet rootlets were given as attractive food (the with transect belt net every 100 m. The total observed sugar beet rootlets are usual supplement food for game territory covered 196 hectares. animals in the study forest). The use of this food made In accordance with the 1997 census there are 30 up from 98 to 100%. In the southern part of the territory bison in the Paiiliai forest (10 males, 14 females and 6 the deer tracks turned aside from the bison locality place young animals) at present. In the tested period there on average up to 45 m (the bisons usually came from were about 19 animals in the forest and also 15 bisons south-eastern and southern part of the territory). Later (10 females and 5 males) were enclosed. The census on the behaviour of deer has changed slightly. However, (Padaiga, Bielova, 1994) by the method of pellet-group the northern part of the territory was always attended by counting has shown that the deer were gathered in the deer (Figure 1). The use of the attractive food consti- northern (namely on the territory where the bison tuted up to 100%. A total of 10 tracks were permanent. enclosures were located), and western parts of the stand. In the situation of disturbance (such as human Pine and birch stands growing on Vaccinio-rny~tilloscr approaching) the deer allowed humans to approach up sites predominate there. Deciduous stands with spruce to 250 m, and ran to the spruce thicket. Bisons allowed grow on Aegopodiosa, Oxalidosa sites. The average them to approach up to 20-25 m, and retreated clamor-

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Table 1. The use of tested territory by red deer and European bison within the first study period

Frequency of the tracks, n per day, in diffcrent part of the territory No of southern part northern part surrounded forest blocks* test deer bison deer bison deer bison

* - surrounded forest blocks with places of yearly supplemental feeding of game animals ** - the strip of 48-70 rn not used by deer up to the places of bison locality

Table 2. The use of the attractive food by red deer and European bison on the tested territory

The usc of the attractive food, of total the quantity, in different parts of the territory No of southern part northern part surrounded forest blocks* test deer bison deer bison deer bison

0 0 85 0 single single 4 3 0 75 0 90 0 90 0 5 0 0 11 0 so 0 ously and hastily. In April the average frequency of the of certain migration is not their characteristic feature. deer tracks in the southern part of the territory was 41100 However, the seasonal movements or micromigrations m and in the northern part 1611 00 m. The direction of the within their habitats are so peculiar to deer. While there moving remained the same as in March. In the places of is no shortage of fodder the deer stay late at the places attractive fodder (sugar beet rootlets) the deer stripped of a sufficient forage in the forest in the vicinity of off the bark of 75% and more of ashes on average at the farmlands. The favourable living conditions i~sually distance of up to 12 m from the feeding place. It shows arouse a tolerance to the neighbours. This tolerance is that the supplemental feeding within the winter period the beginning of the social instinct. Eventually the does not restrain the damage caused by deer on the tolerance becomes the need to group (Bielova, 1995a). preferable tree species as ash, especially at the end of The tradition of the local red deer population is moving winter and in early spring. along the well-known tracks to the places of better foraging and supplemental feeding. This behavioural dominance began to display at the second part of Discussion November on the territory tested. Eventually, the moving of deer becomes more intensive, especially in The Lithuanian deer are not highly social (that is January-February. The moving activity depends on the the social ani~nalwhich is prone to grouping and to weather conditions, disturbances of anthropogeneous making common cause with other relatives). The tradition and other origin (Bielova 1995a,b). Over the tested

1997, VOL. 3, NO. 2 ISSN 1392-1355 = BALTIC FORESTRY THERELATIONSHIP BETWEEN THE RED DEER AND EUROPEAN BISON I.../0. BIELOVA-

Spruce+ P~nc

B~rch+Pme+ Ash

Pme+ Spruce+ B~rch

Fig. 1. Thc schcmc of thc rcd dccr and Europcan bison location on thc tcstcd tcrritory

LEGEND

Ms - mislctou (Viscrtrn alba); Sh - location of thc sugar bcct routlcts; RD - concentration placcs of rcd dccr:

BISON - location placc of Europcan bison;

XXXXX - usc of thc attractive food; . M - mcadow; B - barlcy; Pinc. Sprucc. Birch.. .. - prcdominntcd !rcc spccics; BL-R -block linc - forcst road; C - land - improvcmcrit canal; Snrucc - sprucc thickct > - thc main dircction of rcd dccr movcmcnt

1997, VOL. 3, NO. 2 ISSN 1392-1355 = 44 - BALTIC FORESTRY THERELATIONSHIP BETWEEN THE RED DEER AND EUROPEAN BISON /...I0. BlELOVA- period the need of animals for forage prevails. The 1997). However, the nearest competitor of the red deer characteristic feature of the deer is certain conservati- could be the European bison in their common habitats. vism to the habitat objects and fodder. The usual fodder In bison diet there is up to 100% of tree and shrub (natural and supplemental ones) in the habitat determin- species that are well used by red deer. However, the ed the stability of the deer feeding - the foraging in the differences between the feeding needs, intensity, usual places of supplemental feeding predominates. feeding conditions, and especially between the abun- However, the supplemental feeding by all means does dance of animals, decrease the possible competition. In not restrain the damage caused by deer on the accordance with the common number of used plant preferable species of forest plants (in this case, it was ash species the co~npetitivenessof deer (then the plants bark stripping) directly near the place of the attractive used by deer comprise 100%) with roe deer is found to food. Such intensive damage has not been caused only be 16.3%, with elk 13.1% and with bison 47% In their due to foraging favourability within the tested winter common habitat (~ostaket al., 1983; Bielova, 1997). The period but also due to disturbances of anthropogeneous European bison is the potential competitor - amensal of origin (hunting, machinery, transport, etc.). The repeated deer within the winter period. disturbances can result in frustration and eventually in stress (Lochman, 1976; Missbach, 198 1; Ozoga, Verme, 1982; Bielova, 1993). One of the disturbance could be the Conclusions attendance of bisons in the usual places of deer moving especially under the conditions of forage shortage. 1. The tradition of the local red deer population is Usually the relations between the deer and bison are moving along the well-known tracks to the places of not distinctive, and they avoid contacts because of better foraging and supple~nentalfeeding. indifference. Say, animals tried to meet their needs, but 2. Behavioural dominance of moving began to they failed to do it, then next time their memory display at the second part of November and becomes suspends the need and incentive to avoid an unpleasant especially intensive in Janua~y-February. situation (Bielova, 1995a) (the place is occupied, and 3. The prevailing need of animals is hunger in the deer slightly goes round). The bisons occupied the same winter period. However, the certain conservatism to the habitat on the territory investigated. Free European fodder and other habitat objects is still characteristic bisons usually occupied 10,000 hectares, and more feature of red deer in any season. frequently attended the area of 4,700 hectares (BaleiSis 4. The usual fodder determined the stability of the et al., 1987). The herd of 10-16 animals within the deer feeding and predominance of the foraging in the pasturing occupied about 1 hectare. Their daily territory usual places of supplemental feeding. However, the is 3-5 hectares, and the length of daily track is about 1.3 supplemental feeding by all means does not restrain the km. In winter the bisons usually attend the territory near damage caused by deer on the preferable species of the special supplemental feeding and near the bison forest plants especially directly near the place of the breeding-ground as well (BaleiSis et al, 1987). In the food. period of our study the free bisons started attending 5. The relations between the deer and bison are not the territory of their supplemental feeding and of the distinctive, and they avoid contacts because of bison enclosures in the second part of November and indifference. stepped aside at the end of March. This behaviour 6. The free bisons started attending the territory of corroborates relatively settled life and a philopatria of their supplemental feeding and their native place in the the bison. Bisons yearly return on the well-known second part of November and stepped aside at the end territory. The loss of supplemental fodder in the usual of March. This behaviour corroborates relatively settled places caused intensive feeding of bisons on the life and a philopatria of the bison. attractive new food. It shows a more flexible feeding 7. The feeding behaviour of the bison is more behaviour of the bison in comparison to the deer. Our flexible in comparison to the deer. study of niches of herbivores has shown that the niches 8. The differences between the feeding needs, mostly overlapped during the winter-spring period, the intensity, feeding conditions, and especially between the niches of the European hare and roe deer, roe deer and abundance of animals, decrease the possible competi- red deer, red deer and elk are inostly overlapped (Bielova, tion. However, between the European bison and red deer

1997, VOL. 3, NO. 2 ISSN 1 392- 1355 = BALTIC FORESTRY THERELATIONSHIP BETWEEN THE RED DEER AND EUROPEAN BISON I.../0. BIELOVA- the potential colnpetition is possible up to amensalisln in Biclova 0. 1997. Augaled%i~lgyviinq ekologiniq niSq ir miSko ckosistcmq biojvairoves rySys [Thc relationship bctwccn their colnmon habitats under the unfavourable living tlic forcst ecosystem biodivcrsity and ecological nichcs of conditions. hcrbivores]. MiSkininkyste [Silviculture]. 39,l: 28-38 (in Lithuanian). I~oclunanJ. 1976. Prezimovaci obory pro jclcni zvcr. Lcstictvi, 22,6: 463-484 (in Czech). References Lictuvos Fauna. iinduoliai. 1988 [Lithuanian Fauna. Vertcbratcs]. Vilnius, pp. 26 1-267 (in Lithuanian). BaleiSis R., Ski.rys J. 1984. Rcd dccr introduction, Misshach K. 198 1. Wildscliaden, Nchrungsbcdarf und Fiitterung transferring and marking in Lithuania. Proc. Sci. Acad, dcs Damwildcs. Unscrc Jagd, 3 1, 10: 298-299 (in Gcr~nan). L~tli..B, 3(87)- 89-96 (in Russian). Ozoga J., Verme L. 1982. Physical and rcproductivc charac- BalciSis R., Bluznia P., BalT-iauskas L. 1987. Lictuvos tcristics of a supplcmcntally-fcd wli~tc-tailcd dccr Iicrd. J. kanop~n~aitverys [Lithuanian ungulates]. V~lnius,200 pp. Wildlifc Managerncnt, 46, 2: 28 1-301. (~nL~thuanian). Padaiga V. 197 1. Elniniq pasiskirsty~nas miikc ticmq [Dccr Bielova 0. 1993. Aptvarai clriiq 2icmojimui [Thc cnclosurcs for distribution in the forcst in wintcr]. Girios, 12: 17-1 8 (in dccr overwintcring]. Musq girios, 10: 17 (in Lithuanian) Lithuanian). Bielova 0. 1995a. Ar iicmotq taurusis elnias aptvaruosc [Would Padaiga V. 1990. McdZiokles Ckio pagrindai. Paskait~i kon- rcd dcer overwinter in thc enclos~~rcs].McdZ. ir ivcj., 3: 7-8. spcktai [Thc fi~ndamentalsof thc gamc managcmc~it.Notcs Bielova 0. 1995b. On tlic ethological considcration of solnc of Iccturcs]. Kaunas-Akadcmija, pp. 45-46 (in Lithuania). forcst protcction mcasurcs against daniagc causcd by rcd Padaiga V., Bielova 0. 1994. Tauri~lj~lclniq leikymo aptva- dccr. Baltic-Scand. Joint Symp. Environment and Ungula- ruosc iicmq tcihnologija ir cfcktyvumas [Thc tccli~iology tcs, 1995, pp. 18. and cfficicncy of rcd dccr ovcrwiritcring in cnclosurcs]. F. Bielova 0. 1995c. Pagrindinit1 augaledziq gyviinq vaidmuo Rcp. Kaunas-Girionys, 60 pp. (in Lithuanian). igsaugojant iikiniq miikq biologi~iq jvairovq [The i~nportanccof tlic main hcrbivorcs duc to protcction of thc ~nanagcdforcst biodivcrsity]. F.Rcp. Kaunas-Girionys. 108 pp. (in Lithuanian)

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