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BALTIC FORESTRY

POPULATION AND DISTRIBUTION OF EURASIAN ( CASTOR FIBER ) H. HALLEY ET AL. Population and Distribution of ( Castor fiber )

1 2 3 DUNCAN HALLEY , FRANK ROSELL AND ALEXANDER SAVELJEV

1 Norwegian Institute for Research, Tungasletta 2, N-7485 , , e-mail: [email protected]

2 Telemark University College, N-3800 B¸ i Telemark, Norway

3 Russian Research Institute of Management and Farming (VNIIOZ) of RAAS, 610000 Kirov,

Corresponding author: Duncan Halley

Halley, D., Rosell, F. and Saveljev, A. 2012. Population and Distribution of Eurasian Beaver ( Castor fiber ).

Baltic Forestry 18(1): 168–175.

Abstract

After being reduced to about 1,200 in eight isolated populations by the beginning of the 20th century,

Eurasian ( Castor fiber ) have powerfully recovered in range and population, through widespread reintroductions,

relaxation of persecution, and natural spread. Populations are now (2010) established in all countries within their former

natural range in Europe except for , , and the south Balkans (, Albania, , Macedonia, Kosovo,

Montenegro). In Asia, there are significant populations in West and Central Siberia; and small relict populations elsewhere

in west and south Siberia, and in Xinjiang ()/western . The current minimum world population estimate

for Eurasian beavers is 1.04 million, though this figure includes some populations of introduced North American beavers

( C. canadensis ) in the NW and Far Eastern Russian Federation, where the ranges of the two occur in close

proximity and to some extent overlap. Both populations and range are in rapid expansion, and in 2008 the species was

reclassified by IUCN as Least Concern , though some remain threatened. We present maps summarizing current

knowledge of the world distribution of Eurasian beaver and the Eurasian distribution of the introduced North American

beaver, and tables of the most recent known population estimates for each country, broken down by region for the

Russian Federation.

Keywords : beaver, Castor fiber , population, distribution, range, reintroduction

Introduction overtaken by events. Periodic summaries and updates of the most recent population and distribution data are

Eurasian beavers ( Castor fiber ) have, since the late therefore of use, both to current researchers and man-

19th century, staged a remarkable recovery both in agers seeking an overview of the many, widely scat- population and distribution. From being a critically tered, papers on the subject, and for future biologists endangered species reduced to about 1,200 animals in interested in following in detail a remarkable case study scattered enclaves, the species is now conservational- in conservation biology. In this paper, we present a ly secure and, aided by widespread reintroductions, summary of the information available on population and rapidly recolonizing much of its range, including areas distribution up to the end of 2010, in a similar format where it has not occurred for centuries or even millen- to information presented in Halley and Rosell 2002, 2003 nia. The pattern of re-expansion is not only interesting for ease of comparison. in itself, but also offers valuable insights in the fields of population biology and conservation . Until Distribution and populations recently, this expansion has not been well document- ed, but since the latter part of the 20th century increasing Distribution in Europe is shown in Figure 1 and amounts of information have become available, allow- in Asia in Figure 2. The continuous population rang- ing the preparation of reviews summarizing the then es from through , the , current status of the species (Macdonald et al. 1995, , , to central Siberia in the Russian

Nolet and Rosell 1998, Halley and Rosell 2002, 2003). Federation. There is a large disjunct population in

However, both population and distribution continue to Norway and , and smaller scattered disjunct expand rapidly, both through natural spread and new populations through the rest of mainland Europe. Dis- reintroductions, so that these reviews rapidly become junct reintroduced populations are also found on the

2012, Vol. 18, No. 1 (34) ISSN 2029-9230 168 BALTIC FORESTRY

POPULATION AND DISTRIBUTION OF EURASIAN BEAVER ( CASTOR FIBER ) H. HALLEY ET AL.

Soviet Union, almost 17000 beavers were translocat-

ed between 1927 and 2004, of which 12000 were to

sites in Russia and the remainder to Ukraine, Belarus,

Latvia, , Lithuania and in decreas-

ing order of numbers (Saveljev 2003).

In 2009, beavers were reintroduced to in 1 western and near Copenhagen in . In

2010 the existence of a wild- living population on the Tay

2 4 5 watershed in Scotland, probably resulting from escapes

of captive animals, became public. Sightings of free liv-

ing animals on the Tay date from 2001 (H. Chalmers pers.

comm.). Breeding has been confirmed in at least two

3 locations and suspected in at least six others; but both

the size and status of the population is unclear. This

population was not established by the licensing proc-

ess required in Scotland, and the authorities are currently

Figure 1. Distribution of both beaver species in Europe. (December 2010) attempting removal by live at

Traditional subspecies designations: 1 – Castor fiber fiber ; some of the known sites on the river. 2 – Castor fiber albicus ; 3 – Castor fiber galliae ; 4 – Castor Introduced populations of North American bea- fiber belarusicus; 5 – Castor fiber osteuropaeus . Dark grey ver are established in Finland and northwest Russia, shading represents the present range of Castor fiber (loca- and the Russian Far East (Savelyev and Safanov 1999; tions of relict populations are marked in black); light grey Safonov and Saveljev 1999). In 2009 a small popula- shading represents the range of Castor canadensis . Squares tion was discovered in Luxembourg, apparently as a are reintroduction sites where range has not yet spread sig- nificantly result of escapes from a wildlife park just across the

border in Germany (Table 1).

Table 1. Location and estimated minimum population sizes

of relict populations of Eurasian beaver ( Castor fiber )

Minimum Population population References size

Lower Rhöne, 30 Richard 1985 Telemark, Norway 60-120 Collet 1897 , Germany 200 Heideke and Hürig 1986 Pripet marshes, Belarus/Ukraine/Russia <300 Zharkov and Solokov 1967 Voronezh, Russia 70 Lavrov and Lavrov 1986 Konda-Sosva, Russia 300 Lavrov and Lavrov 1986 Upper Yenesei, Russia 30-40 Lavrov and Lavrov 1986 Urungu, Mongolia/China <100-150 Lavrov and Hao-Tsuan 1961

Figure 2. Distribution of both beaver species in Russia, Ka- Of recent reintroductions, that to the / zakhstan, Mongolia, and China. Traditional subspecies des- watershed of , which is considered illegal by the ignations: 1 – Castor fiber fiber; 6 – Castor fiber pohlei ; regional authorities affected, is the most poorly docu-

7 – Castor fiber tuvinicus ; 8 – Castor fiber birulai . Dark mented. Current distribution and populations are un- grey shading represents the present range of Castor fiber known, though probably rather wider and larger than (relict populations are marked in black); light grey shading mapped and given in the table, given known patterns represents the range of Castor canadensis of dispersal and population growth (Halley and Rosell

2002; Discussion) and the large amount of high quality periphery of the main Russian range, in central Asia, habitat available; breeding and the occupation of new and on the Amur of the Russian Far East. Although territories is known to have occurred. Trapping and natural spread has contributed significantly to range removal is apparently underway in one (possibly two) and populations, most of the expansion is due to re- of the four Spanish regions affected, but neither sys- introductions, of which at least 205 have been recorded tematically or widespread enough to have any realistic to distinct locations outside the former Soviet Union chance of substantially affecting the population. There

(FSU) (Halley and Rosell 2002). Within the former are reports of beaver signs found recently (autumn 2010)

2012, Vol. 18, No. 1 (34) ISSN 2029-9230 169 BALTIC FORESTRY

POPULATION AND DISTRIBUTION OF EURASIAN BEAVER ( CASTOR FIBER ) H. HALLEY ET AL.

downstream on the Ebro at Caspe, c.200 km SE of the descended solely from some of the smaller surviving known range (I. Jiminez pers. comm.), but this may rep- populations may be more susceptible as a population resent a wandering individual rather than establishment. to epidemic disease, due to lack of diversity in immune

The status in remains uncertain. Reports systems (Ellegren et al. 1993). Recently other fuller and of an observation on the Yumurtalik marshes (Adana interesting research of the major histocompatibility province) in the 1970s (Boessneck 1974, Kumerloeve complex, MHC , of the aboriginal beavers of

1975, Veron 1995, discussed in Savelyev 2000) were (albicus, galliae, fiber, tuvinicus, birulai, pohlei ) has discounted by Turkish wildlife experts (Özgün Emre been carried out (Babik et al., 2005), using the same

Can, Turkish Society for the Protection of Nature, pers. samples as used in Ducroz et al., 2005 and Durka et al., comm.; Halley and Rosell 2002). However, further sight- 2005. In all the relict populations except pohlei , the ings have been alleged (Dr.Yüksel Coskun, , Univ of MHC on the DRB gene is specific and monomorphic,

Dulche, pers. comm.), but their reliability is unclear and one form only in each relict population and a different requires confirmation. If the species is still extant in one for each. Only West Siberian beavers C.f.pohlei

Turkey, clearly it would represent an additional sur- show any variation of the immune system (4 variants viving subspecies (and ESU), a significant addition to of the DRB gene). The current minimum world popula- the genetic diversity of the species, and is probably tion estimate of 1 036 226 given here is conservative, critically endangered. Establishing the status, and, if calculated by adding together the lowest estimates for extant, population and distribution, in the area should each country. Note, however, that the population fig- clearly be priority. ures for the North West Russian and Far East Federal

Available information on relict population sizes is Districts of the Russian Federation are for Eurasian and given in Table 1, and data on date of extinction, legal North American beavers combined (those for Finland protection, reintroduction, and current population size do not include C. canadensis ). is presented in Table 2. All surviving European populations have grown

in numbers beyond the point where further loss of

Discussion genetic diversity through drift might be a conserva-

tion problem. The status of some of the Asian relict

Populations of the eight refugia in which Eurasian populations is unclear and would merit investigation. beavers survived the 19th century are currently each In most countries of the western and central Euro- described as separate subspecies. Recent mDNA stud- pean mainland, beavers occur at relatively low numbers ies (Ducroz et al. 2005) indicate that extant populations for the present, and there is much unused suitable hab- of the species divides into at least two Evolutionary itat. The range maps presented here are therefore prob-

Significant Units (ESUs). The western ESU comprises ably conservative, as newly colonising beavers tend to the traditional subspecies C.f. fiber, albicus, and gal- select prime habitat requiring little alteration, where their liae ; the eastern C.f. belarusicus, osteuropaeus, poh- presence may not immediately be noted by non-special- lei, tuvinicus, and birulai. While western ESU popu- ists. Very considerable expansion in both populations lations show very little differentiation and are clearly and range can be expected in the coming decades, es- a single ESU, eastern populations are more diverse and pecially in and the watershed. may be considered to be several ESUs. These findings A typical pattern of population development on a wa- should be taken into account by managers seeking tershed following natural or artificial recolonisation has reintroduction stock in line with IUCN guidelines. become evident in recent years, the evidence for which

Eurasian beaver families consist of, on average, is reviewed in Halley and Rosell (2002); the most de- about four individuals, of which only the adult pair tailed case study of range and population expansion is breeds. The minimum populations quoted should be in- found in Hartman (1995). At first, range expansion terpreted in this light; the Rhône remnant population, through the watershed is rapid, but population expan- for example, would seem to have been reduced to about sion is relatively slow. This seems to be because bea- six breeding pairs at minimum, well below the 25 pairs vers select the best habitat available, rather than set- often quoted as a minimum viable population (MVP). tling close to their natal . It is also difficult to

(That evidence on the ground indicates that this rule- find a mate in the vast, unoccupied stretches of a large of-thumb is generally too pessimistic, is fortunate for watershed, and it appears that beaver will wander widely conservation; see Caughley and Sinclair (1994) for re- while searching. The furthest known dispersal known, view of this issue). The small size of remnant popula- 500km, was registered in 1958 in the Irtysh watershed, tions, and resultant inbreeding and loss of genetic di- in the Omsk region of the USSR (Lavrov 1980 quoted versity, does not seem to have led to serious breeding in Saveljev et al. 2002); and a dispersal of 200 km in- problems in this species, but suggests that populations volving crossing a watershed divide has also been re-

2012, Vol. 18, No. 1 (34) ISSN 2029-9230 170 BALTIC FORESTRY

POPULATION AND DISTRIBUTION OF EURASIAN BEAVER ( CASTOR FIBER ) H. HALLEY ET AL.

Table 2. The history and present status of Eurasian beavers ( Castor fiber )

Cou ntry -xtirp at ion Prote ction Re-in trodu ction and/ Pres ent Referenc es/ comm ents

or trans lo ca t ions populat io n

si ze

! " 5 6

Austri a & 69 - 970 -90 ! 000 Sie >er  9&9, Ko lla r and

Se ite r  990 , Parz-Goll ne r

and Vogl  006 ; Parz-Goll ner

and Hü lzle r co nf . p r es.

Du>in giai

Bela rus remnant 9 9"& 65 "0 0 Dj oshkin and Sa fono v  97 ,

Bela rus Min is try of Stat is tics

 00 &

Bel giu m &"& - 99 & -99 &00 - 000 R u>> ers and Van d e n B oe gert

 006 ; DeMas pers comm .

Bosni a and ? ? 006 60 Kunov ac pers . c omm . 009 

Herze govin aa

Ch in a remnant 99 - 700 She ng et al.  990 , Smith

 999 

Croat ia & 57? - 996 -9& 600 G ru>e si c et al  006 ;

Gru>e si c pers . comm . 009 

Czech 7th c entu ry - 99 -9 , 9 96 000 - 500 Kostk an and L eh k ý  99 7,

Repu>lic Kostkan  0 0 , Vorel et al.

 00 & Denmark c.500 B C - 999 Skov - og Natu rs tyrel sen

 999 ; Kle in  99 9 ; -lmeros

et al.  00 7, 00 9 

-ngla nd & th c entu ry? - F ea si >ility stud y 0 Cole s 006 ; G oM pers .

complete d comm .

-sto ni a &" - 95 7 6 !00 - 7 500 L aa ne tu  9 95 , U le vicius

pers . comm ., T imm -s to ni an

-nvir o nmental Inf ormat io n

Ce ntre pers . comm .

Fin la n d & 6& & 6& 9!5 -!7, 9 95 500 - 500 -rmala AetJ = al.l.  999 , L aht i 6  995 , Hä rkü nen  999 ,-U

France remnant 909 959 -95 0 000 - 5 000 R ic ha rd  9&5, 9 &6, Offi ce

Nat iona le de la Cha ss e

 99 7, De Mas in pr ess 

German y remnant 9 0 9!6 -"0, & 000 - 0 000 SchMa> et al.  99 ",

966 -&9, 9 99 - 000 Ma cdo nald et al.  995 ,

SchMa> pe rs . c omm . 009 .

Hungary & 65 - 9&0 - 006 "9 Kolla r and Se ite r  990 ,

Bozsé r pers . comm .; Gru>e r 6 pers .comm .; -U

Ital y 5" - pr oposed 0 Nole t  9 96 

Kazak hsta n 9 5 - 96 ! - 9& 6 5 500 Karag ois h in  0 00 , Sa vel jev

 005 , B er>er  00 &

Lat via &! 0s - 9 7 -5 , 9 75-&" 00 000 – 50 Balod is  99 , 99 ", 9 95,

000 99 7, 99 &; Ozol ins and

Bau manis  000 

Lithua ni a 9!& - 9"7 -59 &6 000 - 000 Pal ion ene  965 , Mic kus

 995 , B al cia u sk as et al.

 999 ; U le v ic ius  000 ;

Ule vic ius pers .c omm . 009  ! Luxem> ou rg ? ? - 0 Schle y et al. conf . pr es.

Du>in giai

Moldo va ? ? - ? Statu s unce rta in

Mo ngol ia remnant ? 959 - 00 &00 L avrov  9& ! , Stu >> e and

1 DaMaa  9& ! , 9&6, Stu >> e Based on subfossil remains. Philological evidence from placenames suggests a remnant may have survived as late

t h as the 11 century.

2 Reintroductions to fenced enclosures at Ham , Kent, 2001 (6); Cotswolds Water Park in 2005 (6) (2009 at

population least 10, Rodell pers. obs.); and Martin Mere in 2007 (4). Enclosures are large enough that animals

live an essentially wild existence, but dispersal is prevented.

3 C. 15 C. canadensis established, probably escaped from zoo in Saarland (Schley et al pers. comm)

4 Current population of the official trial reintroduction in Knapdale. A small feral population of beavers, of un-

known size, is established in the wild on the Tay river system (see text).

2012, Vol. 18, No. 1 (34) ISSN 2029-9230 171 BALTIC FORESTRY

POPULATION AND DISTRIBUTION OF EURASIAN BEAVER ( CASTOR FIBER ) H. HALLEY ET AL.

Table 2. ! " 5 6

(continued) Mo ngol ia remnant ? 959 - 00 &00 L avrov  9&! , Stu >> e and

DaMaa  9&! , 9&6, Stu >> e

et al.  99 , Sha r  005  6 Nethe rla nds & 6 - 9&& - 000 !00 -!50 Nolet  99 ", 996 , -U

NorMay remnant &" 5 9 5-! , >7 0 000 R osell and Pede rs en  999 ;

95 -65 Parker and R osell  00 !

Pola nd &"" 9 ! 9"! -"9, ! 000 – " ZuroMski and Kasp erczy k

975- 000 000  9&6, 9&& , Z uroMski  99 ,

Ma c do n ald et al.  995 ,

Dz ieciolo Msk i and

GozdzieMski 999, Czech 5 pers . comm .; -U

Portu gal c. " 50 - - 0 Antu nes 9&9

Romani a & " ? - 99 &-99 6 0-690 T roidl and Io nescu  99 7,

SchMa> pers . comm .;

Io nesc u et al.  006  ; G . H erlo

pers . comm . 009 ; G .

Io nesc u et al. pers .comm .

009  6 Russi an remnant 9 9 7-!! , "95 700 >y Dj oshk in and Sa fonov  97 ,

Fede rat ion 9!" -" , Fede ral Dis tri ct Lavrov  9&! , Sa vel jev and

9"6- 005 see >elo M Sa fonov  9 99 , Sa fonov and

Sa vel iev  999 , Dezhk in

 999 ; Sa vel jev 00 !;

Lomanova 009 7 NoHJh-W As J 120 500 Lomanova 009

FAdAH= l

DEsJHE cJ

CAnJH= l F D 126 000 Lomanova 009

PHE volz hs k y 13 8 600 Lomanova 009

FD

SoKJh F D 7 500 Lomanova 009

UH= l F D 31 500 Lomanova 009

SEbAHE= n 70 8 00 Lomanova 009

FD 7 F=H 8 00 Sa vel jev 00 !; L omanova

E=sJAH n F D 009

" Scotla nd 6th centu ry - 009 Scott is h W ildl if e T rust pers . " comm . 009 

Se r>ia 90 !? - 00 !-" 50 Ciroví c 006 ; Cir ovíc

pers .c omm . 00 & 5 5 Slo veni a ? ? 999 0 Gru>esi c pers . comm .; -U

Slo vak ia & 5 - 995 5 0-700 Pachin ger and H ul ik  999 ;

Vala cho vic  99 7, Du>ha and 5 Ma jla n  99 7; -U th Spain 7 c entu ry 9&0s 00 ! >"0 Ce na et al  00 "; Ce na pers .

comm .

SMede n & 7 & 7! 9 -!9 > 00,000 F reye  97&, H artman  99 ",

995 

SMitzerla n d & 0 - 956 -77 600 Sto ck er  9&5, Ma cdo nald et

al.  995 , Win te r  99 7, S.

Ca pt, Ce ntre Su iss e de

Ca rto graphie de la F au ne

pers . comm . 00 7; Angst

c onf . pr es . Du>in giai

Uk rain e remnant 9 !5 " 0 Sa fonov and Sa ve l iev  999 

Uk rgoss tat is tik a  00 &

W ale s 6th centu ry - F ea si > ility s tud y 0 Cole s 006 ; Jones pers

completed comm . 009 

Minim Km ""  usin g lo Mest es timate fr om

PoFKlati on ea ch cou ntry; fin al fi gure

-I timate rou nded to ne a rest thou sand

5 Natural spread from Croatia.

6 EU: European topic centre on http://biodiversity.eionet.europa.eu/article17/

7 Both species ( C. fiber and C. canadensis ).

2012, Vol. 18, No. 1 (34) ISSN 2029-9230 172 BALTIC FORESTRY

POPULATION AND DISTRIBUTION OF EURASIAN BEAVER ( CASTOR FIBER ) H. HALLEY ET AL.

ported (Saveljev et al. 2002).Both were animals moving ticular, we would like to thank S. Asbirk; O. Boszér; into uncolonised waters. However, the longest distance K. Bevanger; S. Capt; D. Cirovic; A. Czech; R. Den- colonisation (i.e., settlement and breeding) has been nis; J.-F. Ducroz; M. Grubesic; G. Hartman; V. Kost- reported is apparently Fustec et al.’s (2001) record of kan; P. Lahti; H. Lea; R. Liobis; P. Rouland; G. colonisation of an area about 80 km away from the Schwab; J. Sieber; F. Tattersall; U. Timm; A. Ulevi- nearest other occupied area, on the in France. cius, A.Volokh,C. and G. Yanuta.

Some time later, depending on the size and topog- raphy of the watershed but often after about 10-25 Afterword years, populations reach a critical density for encoun- tering a mate, and the population then increases very The authors would be grateful, on a continuing rapidly. This is followed (on average 25-34 years after basis, for any corrections and/or updates to the infor- watershed colonisation in Hartman’s (1995) study) by mation we have regarding the progress of reintroduc- a phase of population decline as marginal habitats tion, range expansion, and population development of become exhausted; and then by rough stability. Eurasian beavers. All contributions will, of course, be

While in established populations average disper- acknowledged in any resulting publications. sal distance is usually much less than the extremes mentioned above, 3.9 km on the Azas river in south References Siberia, individuals disperse much further, one subadult male being found 85 km upstream (Saveljev et al. 2002). Babik, W., Durka, W. and Radwan, J. 2005. Sequence di-

This implies the potential for considerable gene flow versity of the MHC DRB gene in the Eurasian beaver (Castor fiber ). Molecular Ecology 14: 4249-4257. within continuous populations. Balciauskas, L., Trakimas, G., Juskaitis, R., Ulevicius, A. Incidences of beaver conflicts with humans tend and Balciauskiene, L. 1999. Atlas of Lithuanian mam- to intensify during the later stages of the rapid increase mals, amphibians and reptiles. Akstis Publications, Vilnius, phase, in part because then beaver more often take into Lithuania. (In Lithuanian with English summary). Balodis, M . 1992. Die Biber in Lettland (The beaver in Lithua- use more marginal sites requiring more beaver engi- nia). In: R. Schröpfer, M. Stubbe and D. Heidecke (eds.). neering, , , etc, which may conflict with Materialen des 2. Internationalen Symposiums Semiaqua- human landuses. In many countries, this phase of tische Säugetiere: 121-129. Martin-Luther-Universität, population development has been accompanied by the Halle/Saale, Germany. Balodis, M. 1994. Beaver population of : history, de- introduction of , aimed at least in part at ad- velopment and management. Proceedings of the Latvian dressing conflict issues (Halley and Rosell 2002). Academy of Sciences, section B . 7-8: 122-127.

Conversely, while beavers can and do cross land, Balodis, M . 1995. Beavers in Latvia. In: A. Ermala and S. and have been found up to 11.7 km away from the Lahti (eds.). Proceedings of the Third Nordic Beaver Symposium, 1992, Finland: 6-9. Finnish Game and Fish- nearest water body (Saveljev et al. 2002) watershed eries Research Institute, Helsinki, Finland. divisions do show a clear barrier effect for beaver Balodis, M . 1997. Recovery of the beaver in the inconstant expansion, which can be strongly isolating where nat- Latvian landscape. In: K. Pachinger (ed.). Proceedings ural or artificial habitat barriers, such as high moun- European Beaver Symposium, 15-19 September 1997: 3. Comenius University, Bratislava, Slovakia. tains or intensive farmland, intrude between water- Balodis, M. 1998. Beavers in the Gauja national park, Latvia. sheds. Depending on the management strategy, there- Proceedings of the Latvian Academy of Sciences 52: 49- fore, this suggests a policy of many reintroductions 5 3 . to many watersheds, or, conversely, the early removal Berber, A.P . 2008. Game resources of Kazakhstan. TaiS Print. House, Karagandy. 1-456. (in Russ.). of colonising individuals on watersheds where their Boessneck, J. 1974. Erganzungen zur einstigen Verbreitung presence is considered undesirable. Given the pattern des Bibers, Castor fiber (Linné, 1758). Saugetierkundli- of range expansion within watersheds, confining bea- che Mitteilungen (München). 22. 83-88. ver populations to a particular stretch within a water- Caughley, G. and Sinclair, A.R.E . 1994. Wildlife ecology and management. Blackwell Science, Cambridge, UK. shed will be impractical unless there are strong artifi- Dewas,M. in press. The situation of native and introduced cial barriers to expansion, such as man-made river beavers (Castor spp.) in France and neighbouring coun- barrages, or a heavy and directed hunting or trapping tries. Review. effort (Halley and Rosell 2002). Beaver populations Dezhkin, V. 1999. Beaver in the modern world: necessity of the population management on the national and the inter- should therefore be managed on a watershed scale. national level. In: Proceedings of the First European-Amer-

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i 1000 år [The first beaver young for 1000 years]. Press

release, 6/8/00.

Received 15 December 2010

Accepted 18 May 2012

2012, Vol. 18, No. 1 (34) ISSN 2029-9230 175