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1. A century ago, overhunting had reduced Eurasian beaver Castor fiber populations to c. 1200 animals in scattered refugia from France to Mongolia. Reintroductions and natural spread have since restored the species to large areas of its original range. Population has more than tripled since the first modern estimate in 1998; the minimum estimate is now c. 1.5 million. 2. Range expansion 2000–2020 has been rapid, with large extensions in western and south-central Europe, southern Russia, and west and central Siberia. Beavers are now re-established in all countries of their former European range except for Portugal, Italy, and the southern Balkans; they occur broadly across Siberia to Mongolia, with scattered populations father east. About half of the world population lives in Russia. Populations appear to be mature in much of European Russia, Belarus, the Baltic States, and Poland. 3. There is a significant population of North American beaver Castor canadensis in Finland and north-west Russia. Most other 20th-Century introductions of this species have become extinct or been removed. 4. Recent DNA studies have improved understanding of Castor fiber population prehistory and history. Two clades, east and west, are extant; a third ‘Danube’ clade is extinct. Refugial populations were strongly bottlenecked, with loss of genetic diversity through genetic drift. 5. Future range extension, and large increases in populations and in impacts on freshwater systems, can be expected. Beavers are now recolonising densely populated, intensely modified, low-relief regions, such as England, the Netherlands, Belgium, and north-west Germany. They will become much more common and widespread there in coming decades. As beavers are ecosystem engineers with profound effects on riparian habitats, attention to integrating beaver management into these landscapes using experience gained in other areas – before the rapid increase in population densities and impacts occurs – is recommended
Content may be subject to copyright.
This paper is the third review – including short updates
the fifth paper – intended to summarise Eurasian beaver
Castor fiber distribution change and population develop-
ment (Nolet & Rosell 1998, Halley & Rosell 2002, 2003,
Halley et al. 2012; a population estimate for Europe was
also recently presented in a short communication, Wrobel
2020). The status of the North American beaver Castor
canadensis in Eurasia is also reviewed. Information avail-
able is copious, but very widely scattered. Some is available
in peer-reviewed papers, but most is in national- or
Population and distribution of beavers Castor fiber and
Castor canadensis in Eurasia
Duncan J. HALLEY*Norwegian Institute for Nature Research, PO Box 5685 Sluppen, Trondheim,
NO-7485, Norway. Email:
Alexander P. SAVELJEVDepartment of Animal Ecology, Russian Research Institute of Game
Management and Fur Farming, 79 Preobrazhenskaya Str., Kirov, 610000, Russia. Email: saveljev.
Frank ROSELLDepartment of Natural Sciences and Environmental Health, Faculty of Technology,
Natural Sciences and Maritime Sciences, University of South-Eastern Norway, Gullbringveien 36, Bø i
Telemark, NO-3800, Norway. Email:
Mammal Review 51 (2021) 1–24 © 2020 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution
in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made.
beaver, Castor fiber, Castor canadensis,
distribution, Eurasia, population,
*Correspondence author
Received: 4 May 2020
Accepted: 17 June 2020
Editor: DR
doi: 10.1111/mam.12216
1. A century ago, overhunting had reduced Eurasian beaver Castor fiber popula-
tions to c. 1200 animals in scattered refugia from France to Mongolia.
Reintroductions and natural spread have since restored the species to large
areas of its original range. Population has more than tripled since the first
modern estimate in 1998; the minimum estimate is now c. 1.5 million.
2. Range expansion 2000–2020 has been rapid, with large extensions in western
and south-central Europe, southern Russia, and west and central Siberia.
Beavers are now re-established in all countries of their former European
range except for Portugal, Italy, and the southern Balkans; they occur broadly
across Siberia to Mongolia, with scattered populations father east. About half
of the world population lives in Russia. Populations appear to be mature in
much of European Russia, Belarus, the Baltic States, and Poland.
3. There is a significant population of North American beaver Castor canadensis
in Finland and north-west Russia. Most other 20th-Century introductions of
this species have become extinct or been removed.
4. Recent DNA studies have improved understanding of Castor fiber population
prehistory and history. Two clades, east and west, are extant; a third ‘Danube’
clade is extinct. Refugial populations were strongly bottlenecked, with loss of
genetic diversity through genetic drift.
5. Future range extension, and large increases in populations and in impacts on
freshwater systems, can be expected. Beavers are now recolonising densely popu-
lated, intensely modified, low-relief regions, such as England, the Netherlands,
Belgium, and north-west Germany. They will become much more common and
widespread there in coming decades. As beavers are ecosystem engineers with
profound effects on riparian habitats, attention to integrating beaver manage-
ment into these landscapes using experience gained in other areas – before the
rapid increase in population densities and impacts occurs – is recommended.
Mammal Review ISSN 0305-1838
D. J. Halley, A. P. Saveljev and F. Rosell
Population and distribution of beavers in Eurasia
2Mammal Review 51 (2021) 1–24 © 2021 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
regional-level reports, popular articles, and other sources
not searchable in scientific databases.
The Eurasian beaver is on the threshold of large exten-
sions in geographic range, and exponential growth in
population, in many parts of western and south-central
Europe, including some of the most heavily populated
and human-modified regions of the world. This, the amount
of information relevant to understanding beaver popula-
tion development accumulated since the last review, and
the large changes in population and distribution since the
last update, make a new review timely.
We collect sources on the Eurasian population and dis-
tribution of both species of beaver, and patterns of popu-
lation development, on an ongoing basis. Principal sources
are searches of Internet science literature databases, general
Internet searches, grey literature, conferences, and personal
correspondence. Maps of beaver distribution and the popu-
lation table are updated as new information is received.
Beaver populations develop on watersheds (Hartman
1994a, 1994b, 1995) and should ideally be considered on
a watershed scale. However, almost all information is
available at a national or sub-national scale. Here, we
present data by country, following the structure of Halley
and Rosell (2002) for ease of comparison. ‘Beaver’ means
the Eurasian beaver unless otherwise stated.
Recent advances in DNA studies have deepened and modi-
fied understanding of the species’ post-Ice Age and bot-
tleneck period population histories.
Previously, eight subspecies, one for each 19th to early
20th-Century refugium, were identified (see, e.g. Lavrov
1983, Nolet & Rosell 1998). The reality of this subdivision
has long been doubted (e.g. Halley & Rosell 2002, Halley
2011, Rosell et al. 2012, Horn et al. 2014), and it is no
longer tenable in the light of modern genetic studies. For
example, “Ancient British beavers … formed part of a
large, continuous, pan-Western European clade that har-
boured little internal substructure” (Marr et al. 2018).
Senn et al. (2014) considered that “A likely suggestion is
that divergence in mtDNA haplotypes (into ‘western’ and
‘eastern’ phylogroups) did indeed arise following popula-
tion retreat into glacial refugia during the last glacial
maximum (~25000 ya), but that introgression following
secondary contact of re-emergent populations caused sub-
sequent mixing of divergent haplotypes in contact regions”.
This agrees with genetic evidence from a wide range of
other species indicating recolonisation of Europe from two
main Ice Age refugia in the Balkans and southern France/
Iberia (e.g. Hewitt 2000).
In addition, Durka et al. (2005) identified an extinct
‘Danube’ clade, neither ‘western’ nor ‘eastern’, while both
Biedrzycka et al. (2014) and Munclinger et al. (in prepara-
tion) identify DNA markers apparently present in no known
refugial population. These are likely to be explained (though
other explanations are possible) by the survival of one or
more populations in north-central Europe in addition to
the eight known. Saveljev et al. (2011) also found genetic
differentiation in beavers from the Pripet refuge, corre-
sponding to different rivers in which beavers survived,
flowing separately from the Pripet Marshes area.
Further genetic research deepening or modifying un-
derstanding of Castor fiber population history can be ex-
pected. We name, as appropriate below, the refugia
reintroduction stock came from. This allows assignment
to the earlier ‘subspecies’ if so desired.
Halley and Rosell (2002) provide information on beaver
source populations, and details of location and size of
reintroductions, before 2002. European distribution in 2002
is shown in Fig. 1; current distribution in Fig. 2. Asian
distribution in 2002 is shown in Fig. 3, and current dis-
tribution in Fig. 4. Dates of extinction, legal protection,
and reintroductions, and current population estimates for
each country are given in Table 1. Fig. 5 shows the mini-
mum population estimates from 1998 to 2019.
In Austria, 40 Eurasian beavers were reintroduced (several
sites) in 1970–90 (Halley & Rosell 2002). Beavers have also
immigrated to Austria from Bavaria, Germany, in large num-
bers. Beavers from Austria have colonised adjacent regions
of Slovenia, Hungary, Slovakia, the Czech Republic,
Switzerland, and Italy. The Universität für Bodenkultur, Wien,
maintains a website on beavers in Austria (https://www.dib. hung/proje kte-aktue lle-infor matio nen/
der-biber -casto r-fiber -in-oeste rreic h/). The population was
estimated as 7600 in 2017. The latest available map (Appendix
S1 and see website) dates from 2012. Nine North American
beavers were also introduced in the 1980s, but no beaver
carcasses examined in later years have been of this species;
it appears to have become extinct (Sieber 2000).
Beavers survived in the Pripet, Berezina, and Neman Rivers,
Belarus, and were reintroduced widely in the Soviet period.
Population and distribution of beavers in Eurasia
D. J. Halley, A. P. Saveljev and F. Rosell
Mammal Review 51 (2021) 1–24 © 2020 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
Beavers from these three populations differ significantly
from one another in morphological and biological char-
acters (Samusenko & Fomenkov 1983). Recent genetic data
indicate that combining all beavers of Belarus stock into
one subspecies, Castor fiber belorussicus, is not valid (Saveljev
et al. 2011).
Beavers are now distributed throughout Belarus. The
National Statistical Committee (http://www.belst
by/en/) publishes annual population figures in English.
The figure for 2017 was 51100. Figures suggest a peak-
ing and decline in population in recent years (2011:
60500; 2012: 64400; 2013: 62000; 2014: 63400; 2015:
58300; 2016: 51300).
Litvinov et al. (2012) describe the modern populations
of beavers. Belarus is developing a national management
strategy, aimed at achieving more intensive hunting and
use of beaver derivatives – meat and castoreum (G. Yanuta
personal communication).
Although vagrants from German reintroductions occurred
previously, beavers in Belgium descend mainly from
unofficial releases of 101 individuals in 1998–2000 to
several sites, mainly in the Ardennes, and an unofficial
release in the Dijle and Laan Rivers of Flanders in 2003
(all openly publicised). Other beavers have spread into
Flanders from the Netherlands, also in 2003. All source
to the mixed stock population in Bavaria (Verbeylen
The Flanders population in 2018 was c. 155 territories,
minimum 400 beavers (J. van den Bogaert, and Belgium
Beavers Working Group, personal communications); in
Wallonia c. 650 territories or 1800–2000 beavers
(Bernaerts 2016; J.-P. Facon, Gouvernment Wallonie,
and Belgium Beavers Working Group, personal com-
munications). Distribution in 2016 is shown in
Appendix S2.
Bosnia and Herzegovina
Forty beavers were released on the Semešnica and
Sokočnica Rivers, south of Banja Luka, Bosnia and
Herzegovina, in 2005 and 2006 (Trbojević & Trbojević
2016). Beavers have also spread throughout the Sana
and parts of the Korana and Una rivers in north-west
Bosnia from Croatia, to the lower Drina River in the
east from Serbian reintroductions, and along the Bosnian
side of the Sava River, from Croatia and Serbia. Trbojević
and Trbojević (2016) estimated the 2016 population at
c. 140. Population growth rates, and the scattered dis-
tribution away from the reintroduction sites (Appendix
Fig. 1. Beaver distribution in Europe in 2002. Black = refugia where Eurasian beaver was never extinct. [Colour figure can be viewed at]
D. J. Halley, A. P. Saveljev and F. Rosell
Population and distribution of beavers in Eurasia
4Mammal Review 51 (2021) 1–24 © 2021 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
S3; Trbojević & Trbojević 2016; Figs 1 and 2), suggest
that beavers are still in the ‘establishment’ phase (Hartman
1994a, 1994b, 1995, Halley & Rosell 2002). Rapid in-
creases on all of these rivers can be expected.
A recent study (Boev & Spassov 2019) documents former
beaver presence throughout Bulgaria and extinction in
1750–1850. Spread from Romania is likely in the near
A native population is found in Xinjiang in the far north-
west, in the upper part of the Ulungur (Wulungu) river
(upper Ob watershed; Lavrov & Lu 1961). Beavers also
live on the partly Mongolian Chinge and Bulgan River
tributaries. These populations are threatened by unsustain-
able water and soil exploitation, forest destruction, and
agricultural expansion. There are more than 60 irrigation
dams on the Ulungur River. The dam closest to the
Mongolian border completely isolates Mongolian from
Chinese subpopulations.
50 km of the Bulgan was designated as the Bulgan
River Beaver Reserve in 1980. In 1992, beavers were
translocated from the Bulgan to the Irtysh River to re-
duce risks of genetic bottlenecking, but results were
unsatisfactory (Huang 1993). A 2007 survey on the Bulgan
showed 145 families with 508–645 beavers (Chu & Jiang
Beavers are Critically Endangered in China (Jiang et al.
2016). The population is estimated as <600.
Beavers were reintroduced to Croatia from Bavaria in
1996–98. Forty-eight were released on the Sava River near
Ivanic Grad and 29 on the Drava at Legrad. Nine were
released on the Česma, a Sava tributary near the main
release site (Grubesić et al. 2001, Cavric 2016). Beavers
are now widespread in the Croatian Drava and Sava River
basins and have spread into Slovenia, Bosnia and
Herzegovina, Hungary, Slovenia, and Austria (on the Mura,
via Slovenia). The population in Croatia is estimated at
10000 (Tomljanović et al. 2018).
Czech Republic
History and status of beavers in the Czech Republic are
reviewed by Vorel et al. (2012) and Vorel and Korbelova
(2016). Recolonisation from Austria on the Morava River
(Danube watershed) began in 1988 and from Germany
on the Elbe in 1992. There has also been recolonisation
in the south-west Czech Republic from Bavaria since 1993.
These movements were supplemented by reintroductions
Fig. 2. Beaver distribution in Europe in 2020. Black = refugia where Eurasian beaver was never extinct. [Colour figure can be viewed at]
Population and distribution of beavers in Eurasia
D. J. Halley, A. P. Saveljev and F. Rosell
Mammal Review 51 (2021) 1–24 © 2020 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
in the 1990s (Halley & Rosell 2002). Beavers are now
widespread in Moravia and appear near to capacity on
the Morava watershed, which drains most of the region
(Vorel & Korbelova 2016). They are also widespread in
the south-west, with scattered populations in central and
north-western regions (Appendix S4). Population size for
the Czech Republic is estimated at >6000. Rapid spread
and linkage of populations are predicted in the coming
decade (Vorel & Korbelova 2016). Development has con-
formed to the typical pattern: rapid spread within a wa-
tershed to high-quality sites, with infilling of steadily less
optimal habitat thereafter (John et al. 2010, Barták et al.
2013). The rate of spread into unoccupied stretches of
main rivers approximates 20 km/year during the rapid
expansion phase (John et al. 2010, Bártak et al. 2013).
A national management plan was adopted in 2003 (Vorel
& Korbelova 2016). Natura 2000 sites, 1.2% of the Czech
Republic, are completely protected; in most of the country
(86%), management measures are permitted as necessary;
and the ‘south Bohemian fishpond basin’ (13%) with large
numbers of earth retention dykes is zoned as incompatible
with beaver settlement.
Elbe beavers were reintroduced at Klosterheden in Jylland,
Denmark, in 1999, and at Sjælland near Copenhagen in
2009–11. In Jylland (Appendix S5; Elmeros 2017), the
population has spread to a number of stream systems in
the northern and central peninsula. The beaver population
in Jylland numbered 98 territories, or an estimated 166–202
animals, in 2017 (Elmeros 2017). Taidal (2018) reported
13 territories (c. 50 individuals) in Sjælland.
A number of releases inside enclosures in England have
been made since 2002. Beavers have been living wild on
the River Otter in Devon, south-west England, since at
least 2008, where breeding is confirmed from 2014. Their
origin was unknown, and the government initially planned
to remove them. This changed, partly as a result of elec-
toral politics (the government wished to avoid controversies
that might lose votes in an election), and the proposal
of the Devon Wildlife Trust for a five-year ‘trial’ of the
beavers’ effects on the landscape was accepted in 2015
(Crowley et al. 2017). This involved supplementing the
population with further releases. Thirteen family groups
are now established (Devon Wildlife Trust 2016, Brazier
et al. 2020). Brazier et al. (2020) provide a comprehensive
report on the population, its development, and impacts.
In 2018, it became clear that beavers were living wild
elsewhere in England: on the Tamar River, the Kent Stour,
and the Wye on the Welsh border, and in the Somerset
levels (Swaile et al. 2018; Appendix S6). Most are Bavarian
stock (Campbell-Palmer et al. 2020), though the Kent Stour
animals appear to be descended from enclosed Norwegian
beavers at Ham Fen. Ten populations living in enclosures,
and 13 projects in development, fenced and free-living, were
identified (Campbell-Palmer et al. 2020; Appendix S6).
Fig. 3. Beaver distribution in Asia in 2002. Black = refugia where Eurasian beaver was never extinct. [Colour figure can be viewed at]
D. J. Halley, A. P. Saveljev and F. Rosell
Population and distribution of beavers in Eurasia
6Mammal Review 51 (2021) 1–24 © 2021 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
Natural England is reviewing evidence from the River Otter
trial and will make a recommendation on the desirability
of further reintroduction (Campbell-Palmer et al. 2020).
Beavers were reintroduced to Estonia in 1957 and spread
naturally from releases in Russia and Latvia. They are
now present throughout the country, except for parts of
the Baltic islands. The harvest has increased steadily to
2017 (7038 animals), and the number of families increased
from 2256 in 2012 to 2494 in 2015 (Veeroja & Männil
2018). Hunters’ estimates of the population, however,
declined from a peak of 20000 in 2008 to 14000 in 2016
(Veeroja & Männil 2018). The most recent estimate is
12–13000 individuals in 2016 (Sjöberg & Belova 2020).
Beavers were the Hunting Association’s ‘animal of the
year’ for 2019 ( -loom-2019-kobra s/).
The government of Finland collects high-quality data on
beaver populations, available online in English (LUKE
2019); see also Fig. 6. Eurasian beavers from Norway, and
North American beavers from the USA, were released in
1935–37 (Lahti & Helminen 1969). The North American
beaver population is most dense in Pohjois-Karjala, Etelä-
Savo and Pohjois-Savo, but the species also occurs in other
parts of eastern and central Finland, and sporadically in
Lapland (Fig. 6). It is spreading westwards in southern
Finland, and there is a risk that it will spread further
into areas populated by the Eurasian beaver. At present,
in south-west Finland, the two species are in contact in
western parts of Pirkanmaa Province. North American
beavers have also spread eastward from Finnish Lapland
into Murmansk region, Russia (Fyodorov & Krasovsky
Eurasian beavers have recently invaded Finnish Lapland
(Finland north of roughly 66°N) from Sweden though
the border Tornë watershed (Tornionjoki). Sixty-two bea-
vers were reintroduced, mainly as widely scattered pairs,
from 1960 to 1993 on Swedish tributaries (Mangi 2008).
They remained uncommon until the 1990s, when the rapid
increase phase of population development commenced.
Beavers are now strongly established on the lower and
middle Tornë in Finland and on the adjacent lower Kemi
(Kemijoki) watershed. A few groups are established on
the upper Ounas tributary of the River Kemi (Fig. 4).
The Kemi is the longest river in Finland, c. 550 km
for the main course. The watershed drains central and
southern Finnish Lapland from 66 to 68°N and a small
area of Russia (Fig. 6), an area of about 55000 km2. The
climate is harsh, but the topography is largely flat or
gently undulating. Rivers are tortuous and lakes abundant;
the amount of suitable habitat for beavers is very large.
More or less simultaneously, the North American beaver
has colonised the upper watershed in the east and south-
east, though established groups are widely scattered. Also,
there are, or appear to be, a few groups of North American
beavers remaining in the Tornë and Ounas rivers from
Fig. 4. Beaver distribution in Asia in 2020. Black = refugia where Eurasian beaver was never extinct. [Colour figure can be viewed at]
Population and distribution of beavers in Eurasia
D. J. Halley, A. P. Saveljev and F. Rosell
Mammal Review 51 (2021) 1–24 © 2020 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
Table1. Dates of extinction (extirpation), legal protection, reintroductions or translocations, and most recent population estimates for Eurasian beavers
Castor fiber by country.
Country Extirpation Protection
Reintroduction and/
or translocations
Most recent
population estimate Source/comments
Austria 1869 – 1970–90 7600 Kollar and Seiter (1990), Univ. für Bodenkultur Wien
Belarus Remnant 1922 1948 51100 National Statistical Committee of the Republic of
Belarus (2018)
Belgium 1848 1998–99 2200–2400 Dewas et al. (2012), www.bever werkg 2019
Bosnia and
? ? 2006 140 Trbojević and Trbojević (2016)
Bulgaria 1750–1850 ? Boev and Spassov (2011)
China Remnant 1991 600 Chu and Jiang (2009), Jiang et al. (2016)
Croatia 1857? – 1996–98 10000 Grubesic et al. (2001), Kralj (2014), Cavric (2016),
Tomljanović et al. (2018)
Czech Republic Mid 18th
1800–10, 1991–92,
>6000 Jitka Uhlikova pers. comm., Vorel et al. (2012), Vorel
and Korbelova (2016)
Denmark c.500 BC 1999, 2009–11 216–252 Asbirk (1998), Elmeros (2017), Taidal (2018)
England 18th Century? 2016† c. 150† Coles (2006), D. Gow pers. comm. (2019)
Estonia 1841 1957 18000 Timm, Estonian Environmental Information Centre
pers. comm., Estonian Hunter’s Association (2019),
Veeroja and Männil (2018)
Finland 1868 1868 1935–37, 1995 3300–4500‡ Lahti and Helminen (1969), Natural Resources
Institute Finland (Luke) (2019)
France Remnant 1909 1959–95 >14000 Dewas et al. (2012), ONCFS (2018)
Germany Remnant 1910 1936–40, 1966–89,
35000 Schwab et al. (1994), G. Schwab pers. comm. (2018)
Hungary 1865 1980–2006 14600–18300 Bajomi (2011), Čanády et al. (2016), Bajomi et al.
Italy 1541 proposed 1 Nolet (1996), Messagero Veneto 30/11/2018
Kazakhstan 1915 1963–86 5500 Saveljev (2005)
Latvia 1870s 1927–52, 1975–84 100000–150000 Belova et al. (2016)
Liechtenstein ? ? 50 Fasel (2018)
Lithuania 1938 1947–59 121000 A. Ulevicius pers. comm. (2019), Belova et al. (2016)
Luxembourg ? ? c. 75 Herr et al. (2018)
Moldova ? ? ? Status uncertain
Mongolia Remnant ? 1959–2002; 2012,
800 Samiya (2013), Saveljev et al. (2015), Adiya et al.
Netherlands 1826 1988–2000 2300–3800 Kurstjens and Niewold (2011), Dijkstra et al. (2018),
Dijkstra (2019)
Norway Remnant 1845 1925–32, 1952–65 >80000 Rosell and Pedersen (1999), Parker and Rosell (2003),
F. Rosell pers. comm. (2020)
Poland 1844 1923 1943–49, 1975–2000 124622 Janiszewski and Misiukiewicz (2012), Rozkrut (2018)
Portugal c. 1450 0 Antunes (1989)
Romania 1824? 1998–99 2145–2250 Ionescu et al. (2010), Pașca et al. (2018)
Remnant 1922 1927–33, 1934–41,
700000 Ognev (1963), Federal State Statistics Service, Russian
Research Institute of Game Management and Fur
Farming (2019); see text
152800§ Russian Fur Union (2016)
Central FD 153750 Russian Fur Union (2016)
Volga FD 168070 Russian Fur Union (2016)
South FD 7660 Russian Fur Union (2016)
Ural FD 58100 Russian Fur Union (2016)
Siberian FD 80780 Russian Fur Union (2016)
Far Eastern FD 800§ Russian Fur Union (2016)
D. J. Halley, A. P. Saveljev and F. Rosell
Population and distribution of beavers in Eurasia
8Mammal Review 51 (2021) 1–24 © 2021 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
an earlier introduction. Whether these are genuinely North
American beavers or Eurasian beavers settled at old North
American beaver sites is unclear. DNA testing is underway
(K. Kauhala, personal communication).
The scene seems set for rapid increase of both species,
eventually meeting along a front within the Kemi basin
somewhere in central Lapland. Castor canadensis is clas-
sified as an invasive species in Finland’s National Strategy
on Invasive Alien Species, so a logical strategy would be
to attempt to eliminate, or at least reduce, North American
beaver on the Kemi, to favour recolonisation by Eurasian
beaver (Parker et al. 2012). Strategic reintroductions of
Fig. 5. Minimum total Eurasian beaver Castor fiber population estimates 1998–2020. The sources are indicated. [Colour figure can be viewed at]
Country Extirpation Protection
Reintroduction and/
or translocations
Most recent
population estimate Source/comments
Scotland 16th Century 2009 c. 319–547 Coles (2006), Campbell-Palmer et al. (2018)
Serbia 1903? 2003–4 240 Cirovíc pers. comm. (2012), Smeraldo et al. (2017)
Slovenia 1750? ? 300–400 Grubešić pers. comm. Jursic et al. (2017)
Slovakia 1858 1995 7700–9600 Čanády et al. (2016)
Spain 17th Century 1980s 2003 450–650|| Cena et al (2004), Sáenz de Buruaga (2017)
Sweden 1871 1873 1922–39 130000 Hartman (1994a, 1995), Belova et al. (2016)
Switzerland 1820 1956–77 2800 C. Angst pers. comm. 2012, BAFU (2016)
Ukraine Remnant 1922 46000 Safonov and Pavlov 1973, Matsiboruk (2013a),
Matsiboruk (2013b)
Wales 16th Century Feasibility study
c. 15 Coles 2006; Jones pers. comm. (2009), D. Gow pers.
comm. (2019)
*1876 extirpation of animals descended from 1800–10 reintroduction in S Bohemia (Vorel & Korbelova (2016).
Based on subfossil remains. Philological evidence from place names suggests a remnant may have survived as late as the 11th Century.
First licensed release, to River Otter; to supplement a population apparently deriving from escapes. Populations on other rivers are apparently the
result of escapes from fenced enclosures. Population figure is for wild-living beavers only.
Also c. 10300–19100 Castor canadensis. Source: al-resou rces/game-and-hunti ng/beave rs/.
§Population estimate includes some Castor canadensis; Russian North-west FD < 20000, Far East FD < 200.
||Navarra only, see text.
Table 1. (Continued)
Population and distribution of beavers in Eurasia
D. J. Halley, A. P. Saveljev and F. Rosell
Mammal Review 51 (2021) 1–24 © 2020 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
Eurasian beavers to sites in the Kemi basin would assist
this process.
This would be technically straightforward, and much
cheaper than managing a later situation resembling that
in south-west Finland. Finland is legally required to (at
least) contain the spread of North American beaver when
feasible. Article 19 of the European Union Regulation on
the Prevention and Management of the Introduction and
Fig. 6. Distribution of Eurasian beavers Castor fiber and North American beavers Castor canadensis in Finland in 2018. The catchment of the Kemi
watershed is indicated. Adapted with permission from LUKE (2019). [Colour figure can be viewed at]
D. J. Halley, A. P. Saveljev and F. Rosell
Population and distribution of beavers in Eurasia
10 Mammal Review 51 (2021) 1–24 © 2021 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
Spread of Invasive Alien Species states that measures must
be taken ‘aimed at the eradication, population control or
containment of a population of an invasive alien species’
(Genovesi & Shine 2003, European Union 2014). Human
social and regulatory factors are the main barrier to man-
agement (K. Kauhala, personal communication).
Beavers survived in very small numbers in the Rhone
delta, France. There have been 26 reintroductions (273
individuals) from this source within France (Dubrulle &
Catusse 2012). The most recent accounts of population
and distribution are Dewas et al. (2012), Dubrulle and
Catusse (2012) and ONCFS (2018). ONCFS (now Office
français de la biodiversité) publishes a regularly updated
distribution map (Appendix S7) online at http://carmen.
The current population estimate for France is > 14000,
in effect unchanged from the 10–15000 reported by Halley
et al. (2012). A new population estimate based on a
2019 survey will be published in 2020 (ONCFS 2018)
and will be significantly higher. Range has extended sig-
nificantly on most occupied watersheds. In 2009,
10500 km of watercourse was permanently occupied; in
2018, 15000 km (ONCFS 2018); in 2019, 16500 km
(Yoann Bressan, Office français de la biodiversité, personal
communication). On much of the Rhone the population
is mature; some marginal sites have been abandoned
(Dewas et al. 2012).
Small populations are now established on the Saône and
Doubs (major northern tributaries of the Rhone) and on
the Garonne (84811 km2), ‘certainly’ to a 25 km stretch of
the Tarn tributary downstream of Albi, 55 km north-east
of Toulouse, and ‘probably’ to a 20 km reach above Rabastens,
a further 40 km downstream. These are below the hydro-
barrages on the upper Tarn, which until recently restricted
spread (ONCFS 2018). A population was recently discovered
on the lower Vère, a small side tributary of the Aveyron
tributary of the Garonne west of Mountauban (ONCFS
2017). Large-scale colonisation of both the Saône and Doubs,
and the Garonne above the barrage at Golfech (65 km
north-east of Toulouse), should occur over the next 20 years
(Dewas et al. 2012, ONCFS 2017). Colonisation of the city
of Lille and the Avesnois in Nord Pas de Calais, NW France,
was confirmed in 2020 (
Eleven beavers from north-east France, and 101 from
throughout France, were genetically tested in 2011 and
2018, respectively. All were Castor fiber, and all except
one of Rhone stock. The exception was an animal tested
in 2018, descended from Elbe beavers, found on the Saar
in extreme north-eastern France (Bressan et al. 2018).
An adult beaver was filmed on the River Nive near
Ustaritz in south-west France in late 2018 (Onda news-
paper, 11 January 2019; e-un-
casto r-sur-les-bords/). It may have been from the Ebro,
which would involve a crossing of the (formidable) foothills
of the Pyrenees.
Beavers on the River Elbe, Germany, descend from the
refugium on that river system. The population in Bavaria,
now found throughout the state, is of highly mixed origin,
but not including Elbe beavers (see Halley & Rosell 2002).
Bavaria has been the source for most reintroductions
throughout Europe since the 1970s, including to the re-
mainder of Germany. The current population is estimated
at 35000.
Beavers are now found throughout most of eastern and
southern Germany, with strongly established disjunct popu-
lations in the west (Appendix S8; Stolz 2017). Rapid ex-
pansion in range, followed by rapid population increase,
will almost certainly take place in western Germany, and
into remaining unoccupied areas of the south and east,
over the next 2-3 decades.
A comprehensive website on beavers in Germany and central
Europe is available at http://www.biber manag
Government organisations have suggested a beaver rein-
troduction feasibility study in Greece (Alexandros
Karamanlidis, personal communication).
Beavers immigrated to Hungary from Austria from the
1990s, along the River Danube (Boszér 2000). In 1996–2008,
more than 200 beavers were reintroduced in the areas of
Gemenc and Hanság, on the River Tisza (the main wa-
tershed of eastern Hungary, entering from Transcarpathian
Ukraine and exiting to Serbia) and on the River Dráva
(which forms part of the border with Croatia).
Reintroduction history is described by Bajomi (2011) and
Bajomi et al. (2016). Most were sourced from Bavaria.
Beavers are now found along the entire course of the
Hungarian Danube, and most of the Tisza and some of
its tributaries. The population is estimated at 14600–18300
(Čanády et al. 2016).
In 2018, a beaver settled on the Gaililz, a Danube head-
stream (via the Austrian Drava), at Tarvisio in extreme
Population and distribution of beavers in Eurasia
D. J. Halley, A. P. Saveljev and F. Rosell
Mammal Review 51 (2021) 1–24 © 2020 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
north-east Italy, the first Italian record for 450 years
(Messagero Veneto newspaper, 30 November 2018). The
Alps are a formidable barrier to spread into the rest of
Italy, as for other populations established around the land
border of Italy.
Beavers were part of the Kazakh fauna until the early
20th Century. The modern population spread into the
lower Ural watershed from 1963, from reintroductions
upstream near Orenburg, Russia (Karagoyshin 2000). A
second population in eastern Kazakhstan descends from
migration and translocation from the Russian Altai in the
1980s (Berber 2008). In 1996, beavers were removed from
the Kazakh Red Data Book. The current range is an almost
continuous strip of northern Kazakhstan. Population and
distribution are increasing; hunting has been permitted
since 2009 (Baytanaev et al. 2012).
Beavers are present throughout Latvia, as a result of re-
introductions from Norway and Russia. The population
estimate peaked at 89474 in 2009–10 and declined to 58000
by 2017–18 (Zemkopibas Ministrija 2018). Estimated popu-
lation size in 2016–17 was 58000 (State Forest Service
2018) but, according to expert opinion, the actual popula-
tion size could be as high as 150000 (J. Ozolins, cited in
Sjöberg & Belova 2020). See Lithuania for a likely cause
of the discrepancy.
Beavers spread from Switzerland in 2008. In 2017–18, the
population was estimated at 50 (Fasel 2018).
Beavers descended from reintroductions from Russia are
present throughout Lithuania. The official population es-
timate is 40600 (Lietuvos Respublikos Aplinkos ministerija
2017, Raskauskaite & Simkevicius 2017), but this is a
hunters’ association estimate, reported to the game and
wildlife authorities. Hunting associations (which have the
hunting rights) must pay landowners on a population basis
if they do not ensure the population is ‘optimal’. This
provides a motive for underestimating population size.
The real population appears to be roughly stable, with
local declines and increases observed, and is estimated at
c. 100000 (A. Ulevicius, personal communication) and
101265 (Sjöberg & Belova 2020).
Herr et al. (2018) review population history and current
distribution of beavers in Luxembourg. Groups of North
American beavers, established in the early 21st Century,
apparently from zoo escapes in German Saarland, have
been removed. There are currently c. 20 established Eurasian
beaver territories, or about 75 individuals (Appendix S9),
mainly descended from immigrants from Belgium; but
also Germany and possibly France.
A small population of Sino-Mongolian beavers survived
on the Bulgan River in the extreme west of Mongolia
and neighbouring Xinjiang in China. Local translocations
of Bulgan beavers have taken place, to the Khovd (1959–85,
39 animals), the Tes (in 1985, 1988 and 2002, 37 animals),
and to Voronezh (four, in 1962; Stubbe & Dawaa 1983,
Samiya et al. 2012, Saveljev et al. 2015).
Native beavers are today found in three isolated popula-
tions: Bulgan, Khovd, and the Uvs Lake basin (includes
the Tes River and adjacent small rivers in Mongolia and
Russia). The total population is estimated at 600, of which
the Tes population is the largest (Samiya 2013, Saveljev
et al. 2015).
Fourteen beavers from Bavaria, Germany, and 27 from
Kirov, Russia, were brought to Ulaanbaatar, Mongolia, in
2012 (Samiya et al. 2012, Adiya et al. 2015), to improve
water retention in regional rivers, and 16 Kirov beavers
(9 males, 7 females) from seven families were released
on the Zaan (Tuul River tributary, Selenga basin). From
2012 to 2019, 38 beavers from a captive breeding facility
(containing Kirov and Bavarian animals) were released
into the wild on the Zaan and Tuul. The facility contained
44 individuals in 20 family groups in 2019 (S. Shar, per-
sonal communication). Further releases in north and east
Mongolia are planned.
Monitoring attempts have not produced useable estimates
of the wild-living population in Mongolia.
The current status of beavers in the Netherlands is reported
by Dijkstra (2019). The population is estimated at 3500.
Beavers continue to expand in both population and range,
by expansion from reintroductions, immigration from
Belgium (mainly along the River Maas), and from Germany
in the Drenthe Province. In 2012–14, the range increased
by 30% per year in terms of 5x5 km squares occupied.
In 2015–17, this slowed to 11% per year (Dijkstra et al.
2018). Distribution (Appendix S10) is mapped in the
D. J. Halley, A. P. Saveljev and F. Rosell
Population and distribution of beavers in Eurasia
12 Mammal Review 51 (2021) 1–24 © 2021 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
National Flora and Fauna Database at https://www.versp
reidi ngsat
Beavers survived in Telemark in south-east Norway and
were reintroduced at numerous sites from the 1920s to
the 1990s (Rosell & Pedersen 1999). The species now
occupies the entire country south of Saltfjellet (an ocean-
edge tundra/ice cap plateau) on the Arctic Circle, and
east of the mountain chain dividing Atlantic from
Skagerrak watersheds south of Trondheim (Appendix
Populations reintroduced north of Saltfjellet in the 1960s
and 1970s at Rago National Park in Nordland, and in
the Porsanger/Tana region of Finnmark, are now extinct.
Occasional vagrants, probably from Sweden, are reported
in the area.
Four small populations have become established by
natural spread on Atlantic-draining rivers in the south-
west fjord region of Norway. This must have involved
overland travel across the watershed divide. In the long
term, beavers will probably establish themselves in pockets
of suitable habitat in much of fjordland Norway. Beavers
are also advancing slowly into the region along the coast,
both from the south and north. Colonisation patterns
indicate dispersal through brackish and salt water, especially
where it is sheltered (Halley et al. 2013).
In Norway, beavers are hunted as small game. Hunting
rights are owned by the landowner; there is no system
of ‘compensation’ for beaver impacts. This and that beavers
are not highly prized as a hunting object, in combination,
renders the species of low salience, underlining the large
human social factors involved in ‘wildlife-human conflicts’
(Parker & Rosell 2003). In consequence, there are no full-
time beaver managers in Norway and no funding for
monitoring. The population estimate (80000) is, in these
circumstances, an educated guess.
Beavers were widely reintroduced in Poland from the Soviet
Union, mainly from 1975, with internal translocations from
1990 to 2000. They are now found throughout Poland.
The population shows signs that it may be leaving the
rapid increase phase of development, from 500 animals in
1975 to 125000 in 2017. Figures in Rozkrut (2018) show
a levelling off of population in 2015–17. In the Vistula
Delta, where beavers reappeared in the mid-1990s, studies
indicate suitable habitat is now saturated and the popula-
tion stable (Zwolicki et al. 2019). While a longer series is
needed to confirm the trend, it is likely that populations
may decline moderately over the next 20 years.
A Polish website offering beaver management services claims
annual damage to private landowners of 6.6 million
However, despite the 23% population increase since 2015,
there has been an 11% decrease in reported conflicts.
Ministry of Environment compensation payments totalled
5192000 in 2017 (Wróbel & Krysztofiak-Kaniewska
2020). Wróbel and Krysztofiak-Kaniewska (2020) suggest
that the decrease in reported conflicts may be due to
landowners becoming used to beavers and developing
better management skills. Use of flow devices is now
widespread (Wróbel & Krysztofiak-Kaniewska 2020).
The Polish Union of Hunters does not want to hunt
beavers, apparently because the obligation to compensate
for damage would pass to them (Saveljev et al. 2020).
Nevertheless, looking ahead, Polish specialists have already
published a guide to the integrated use of beaver resources
and hunting products (Misiukiewicz 2018).
The most recent source on beavers in Romania is Pașca
et al. (2018). Reintroductions from Bavaria were carried
out from 1998 to 2003. Ninety-one beavers were released
in the Olt River, 56 in the Mureș and 35 in the Ialomița
(Ionescu et al. 2010). In 2014–17, populations were esti-
mated at 1565 on the Olt, 330 on the Mureș, and 214
on the Ialomița. In 2010, beavers, presumably from the
Ialomița, colonised the upper Danube Delta near Tulcea,
and later the main delta. In the north, beavers colonised
Maramureș in 2008, from the Hungarian Tisza via Ukraine
(Chiș 2015). There were six active territories in 2017
(double the 2015 figure), or c. 25–30 animals. (V. Chiș,
personal communication). The Romanian population is
estimated at 2145–2250 (Pașca et al. 2018). Signs of va-
grants have also been recorded on the Someș and Criș
Rivers of western Transylvania, tributaries of the Tisza;
colonisation can be expected in the short to medium term.
Russian Federation
Although greatly reduced in numbers and very patchily
distributed, in the 19th Century beavers survived in the
Russian Federation in the forest zone at least as far east
as the easternmost tributaries of the River Lena, and along
many rivers in the steppe zone (Ognev 1963). It has
been suggested that beavers did not live in rivers draining
to the Sea of Okhotsk or the Pacific, except the upper
Amur (Ognev 1963). However, beaver remains at human
sites dated from 800 to 1500 years ago in the Ussuri
headwaters about 50 km north of modern Vladivostok
(Kuzmin 1995, 1997) indicate they were naturally present
in that region.
Population and distribution of beavers in Eurasia
D. J. Halley, A. P. Saveljev and F. Rosell
Mammal Review 51 (2021) 1–24 © 2020 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
Population and distribution are expanding. A map is
available at map. While previously,
range expansion was most active in the mountains of
southern Siberia (Saveljev 2003), recent major expansion
has shifted to southern European Russia, and the northern
periphery in Siberia.
In European Russia, beavers have expanded down the
Volga to Volgograd. Range has significantly expanded in
the Rostov region along the River Don, to its mouth.
Most inhabit the northern Rostov region, on the Seversky
Donets, Kalitva, and Chir, and the middle Don. Population
increases are expected on the lower Don, as well as colo-
nisation of rivers around the Sea of Azov. Previously,
beavers swimming in the sea were recorded only in the
Dnieper estuary (Igor Sheygas, personal communication),
but dispersing beavers have recently been recorded in the
Sea of Azov. Migrants emerge from the mouth of the
Don and have settled south along the coast (Valery Stakheev,
personal communication).
The Don River delta and Tsimlyansk area may also be
colonised by beavers from Ukraine (see below). The Rostov
region population is estimated at >2300 (Stakheev et al.
2018). Beavers have also spread to the northern part of
Kalmykia (a typical steppe region between the Rivers Don
and Volga).
Beavers have begun to settle in large cities (St. Petersburg,
Kazan, Kirov and others). At least 20 beaver territories
are recorded in Moscow (see
In the western Urals, beavers have settled almost to
the mountain headwaters of the Rivers Pechora, Ilych and
Vishera. Permanent settlement on the Vishera River can
be found up to 555 m above sea level, near the transition
to alpine tundra (Saveljev et al. 2011).
Large-scale expansion is occurring in Siberia. Beavers
from the Ob have penetrated northwards to the Taz basin
in recent years. Average annual movement of the expan-
sion front was about 40 km per year (Yan Kizhevatov,
personal communication).
On the Yenisei watershed, the northernmost stable
population is found in the Yeloguy. Beavers are regular
in the lower reaches of the Podkamennaya Tunguska. To
the south, the entire middle and upper Yenisei basin is
now inhabited, with increasing density upstream, both in
protected and in hunting areas (Trenkov 2015). Beavers
also live in the upper Angara in Irkutsk Region, and in
Buryatia. An isolated population of Belarusian origin has
existed in the Amur basin near Khabarovsk for 55 years;
population and range are increasing. In contrast, the nearby
population of Castor canadensis on the Obor is in decline
(Oleinikov 2013).
In northern European Russia, range dynamics are less
active. Interesting changes have occurred in the contact
zone of the two species. Eurasian beavers have displaced
North American beavers in areas of southern Karelia in-
habited by Castor canadensis since their release there in
the late 1960s. The closest distance between colonies of
different species is 10 km. Conversely, in north-eastern
Karelia (Kemsky District), North American beavers have
penetrated 70 km into Arkhangelsk region and are colo-
nising areas inhabited by Eurasian beavers (Danilov &
Fyodorov 2016). Castor canadensis from Finnish Lapland
are moving east into Murmansk region (Kataev 2018,
Fyodorov & Krasovsky 2019).
Between 2005 (Saveljev 2005) and 2015, the number
of beavers in Russia increased annually, although growth
rates gradually decreased. Borisov (2011) estimated popula-
tions as 528900 in 2008, 611800 in 2009, and 628300 in
2010. In 2015, the population was estimated at 622000
(Russian Fur Union 2016). See Table 1 for a breakdown
by Federal District.
According to the Russian Research Institute of Game
Management and Fur Farming, the population of both
beaver species together reached 696070 in 2018. The Federal
State Statistics Service ( r/11194)
reported 24342 beavers harvested in the hunting season
Given the general trend, and that the number of North
American beavers in the north-western regions does not
exceed 20000, and in the Russian Far East no more than
200, the total population of the Eurasian beaver in Russia
in 2020 can be estimated as at least 700000, about half
of the world population.
Detailed investigation of reintroduction began in Scotland
in 1993, resulting ultimately in a ‘trial’ reintroduction of
Norwegian beavers on the Knapdale Peninsula in western
Scotland from 2009, at a site with strong natural barriers
to spread. In the same year, however, it became clear
that a wild population was established on the Rivers Tay
and Earn, the largest watershed in Scotland. The source
appears to have been escapes from enclosures, and beavers
are of Bavarian origin (Campbell-Palmer et al. 2020). The
first beaver observed and photographed in the wild was
at the Tay/Earn confluence in April 2001 (Hugh Chalmers,
personal communication). However, the territory at
Rannoch/Dunalastair on the upper river (Territory 49 in
Appendix S12), separated from the main population by
three large hydroelectric dams along 25 km of river unoc-
cupied by beavers despite much high-quality habitat, is
unlikely to have resulted from natural spread from the
lower river, where all fenced enclosures were located
(Campbell et al. 2012).
Entrenched controversy surrounding land use in Scotland
has strongly influenced both the official beaver
D. J. Halley, A. P. Saveljev and F. Rosell
Population and distribution of beavers in Eurasia
14 Mammal Review 51 (2021) 1–24 © 2021 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
reintroduction and management issues. It has also led to
much detailed research and review on various aspects of
beaver biology, particularly on issues of local controversy
such as beaver damming and migratory fish (Kemp et al.
2012, Beaver-Salmonid Working Group 2015). There has
also been comprehensive monitoring (Campbell-Palmer et
al. 2018); see also Gaywood (2018) for review.
The Knapdale population remains restricted in range
and numbers. In 2019, it was augmented by seven animals
translocated from the Tay (Scottish Natural Heritage 2020).
The Tay and Earn population has continued to expand
in range, and especially in population (Appendix S12),
despite 82% of individuals being as closely related as first
cousins, presumably due to small numbers of founding
animals (Cambell-Palmer et al. 2020), and significant
numbers being legally killed by landowners, as they were
unprotected. Beavers have also colonised the Forth wa-
tershed to the south, where they are widely but thinly
spread in high-quality habitat (Appendix S12). The wa-
tershed is small, and considerable growth in population
can be expected in the near future. The 2017–18 popula-
tion on the Tay/Earn and Forth was estimated at 114
active territories, or 319–547 individuals (Campbell-Palmer
et al. 2018).
A small population in Beauly, near Inverness, descended
from escapes, remains present despite the removal of five
individuals in 2018. Tay beavers have recently spread west
to rivers north of Knapdale (R. Campbell-Palmer, G. Dowse,
Scottish Beavers, personal communications).
Beavers were given European Protected Species status
in Scotland from 1 May 2019, and “the species will be
allowed to expand its range naturally” (Scottish Natural
Heritage, press release, February 2019). From that date
to 31 December 2019, 87 beavers were killed under gov-
ernment licence (Scottish Natural Heritage 2020).
Seventy-five beavers from Bavaria were reintroduced to
Serbia, at Obedska Bara and Zasavica nature reserves on
the lower Sava River, in 2004–2005. Both have established
local populations. Smeraldo et al. (2017) report spread
upstream to the Bosnia–Croatia border and to the lower
Drina on the Bosnia–Serbia border. There appears to have
been little downstream spread into the Danube, though
habitat and hydrology appear suitable for transit and set-
tlement; possibly Belgrade city (at the Sava-Danube con-
fluence) has hindered spread in that direction. In recent
years, the species has spread into northern Vojvodina from
Hungary along the Tisza (Appendix S13). From 2004–2013,
expansion of range was very rapid, 70.9 ± 12.8 km/year
(mean ± standard deviation, n = 10; Smeraldo et al. 2017),
typical of the early expansion phase on large river systems
(Hartman 1995, Fustec et al. 2001, Halley & Rosell 2002).
The population is estimated at c. 240. Rapid increase in
population and range can be expected in the next
20–30 years.
Recent history and status of beavers in Slovakia are re-
viewed by Čanády et al. (2016). Slovakian rivers drain
southwards to the Danube in, or on the border with,
Hungary. Beavers immigrated to Slovakia along the Danube
from Austria from 1977 and from Poland to north-east
Slovakia from 1993 (Valachovič 2012). Populations are
concentrated in the west and north-east, with recent es-
tablishment from the Hungarian Tisza on its Hornád
tributary in south-east Slovenia (Appendix S14). Central
Slovakia remains uncolonised, but contains much suitable
habitat. The population is estimated at 7700–9600
(Valachovič 2012).
Beavers from the Croatian reintroduction reached the Krka
tributary of the River Sava, Slovenia, in 1998 (Vochl &
Halley 2017, Jursic et al. 2017). Later, beavers entered the
southern Dravinja (Drava) river from Croatia (downstream)
and the north of the same river from Austria (upstream;
Deberšek 2012). Hydro-barrages have so far prevented
colonisation of the middle course. Since 2003 beavers have
colonised the whole of the Slovene Mura, a Drava tribu-
tary, from Croatia (Deberšek 2012). The national popula-
tion was estimated in 2015 as “at least 60 families with
300 to 400 individuals” (Ministry of Environment, quoted
in Dnevnik newspaper, 29 March 2018) and “70 families”
(Slovenske Novice newspaper, 10 April 2018). However, a
detailed survey of the Krka estimated the population on
that river alone at 50 active territories, 168–392 individuals
(Jursic et al. 2017).
Eighteen individual beavers of Bavarian origin were released
in Spain, near the confluence of the Ebro and its Aragon
River tributary in 2003 (Cena et al. 2004). The release
was unofficial, and Spanish authorities obtained a written
derogation from the European Union Habitats Directive
which allowed them to attempt extermination (albeit on
the incorrect grounds that the species was ‘outside its
natural distribution’ on the Ebro).
The issues both of the release and the attempts to
remove the animals generated considerable heat, but
little light on population status. The political geography
of the Ebro is complex, and the environment is the
Population and distribution of beavers in Eurasia
D. J. Halley, A. P. Saveljev and F. Rosell
Mammal Review 51 (2021) 1–24 © 2020 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
responsibility of the regions. Information obtained by
conservation groups through Freedom of Information
laws revealed that in La Rioja, 26 males and eight fe-
males were trapped in 2010, in Navarra 20 males and
16 females in 2008–2009, and in Aragon 10 males and
five females (all older than three years old) in 2009–2010,
using Bailey live traps and humane snares. A male in
his second year was captured by the firemen of Zaragoza
during the same period. The reason for the bias in the
sex ratio is not clear. Later, a similar request to the
European Commission revealed these figures were in-
complete: from 2008–2017, 102 beavers were trapped
in La Rioja, 83 in Navarra and 31 in Aragon (total 216;
European Commission Environment Directorate 2018,
Annexe, Echegaray et al. 2018).
The Ebro River basin contains a great deal of high-
quality habitat, in which beavers can live unobtrusively.
Trapping never extended to Euskadi (the Basque country),
in which beavers were already present. As predicted (see
http://www.iberi anatu blog/tag/beave rs-in-
spain/), the trapping attempt failed, and little attempt was
made after 2014 (16 animals trapped 2015–17, Echegaray
et al. 2018). The environmental economics journal Ballena
Blanca (No 6, April 2016) reported 100 individuals trapped
(an underestimate based on then released figures) at a
total cost of 131000. The population in Navarra alone
was estimated in 2014 as 450–650 (Government of Navarra
2015, unpublished report). If this is correct, the popula-
tion on the entire Ebro system must now be in excess
of 1000 individuals.
The main text of the European Commission Environment
Directorate (2018) letter reversed the European Union’s
position and stated beavers were a “historically autochthonous
and renaturalised species in Spain”, and that Spain “must
adopt the necessary actions to comply with the provisions
of the (Habitats) Directive”, legally protecting the species.
This decision is not as yet integrated into regional law.
Distribution in Spain (Appendix S15) is better known
than population. Beavers are now found from Miranda de
Ebro to Caspe on the main river, a straight-line distance
of over 300 km through which the river meanders strongly.
Beavers are established throughout the main Ega, Arga,
Irati and Aragon tributaries and their side streams in Navarra
(G. Berasategui, personal communication) and Euskadi
(Sáenz de Buruaga 2017). Newspaper reports, trapping loca-
tions (Echegaray et al. 2018, Fig. 2), and Internet videos
indicate beavers are also present on the Jalon (as far up-
stream as Arcos de Jalon), Alama, Cidaco, Iregua, Najarilla,
and Cicados tributaries. Signs of dispersing individuals have
been noted in several places on the lower River Ebro in
Catalonia. Genetic studies indicate the population is of
mixed genetic origin (Põdra & Aguilar Gómez 2015).
Beaver reintroductions in Sweden commenced in the 1920s
and were widespread except in the south (Hartman 1994a,
1994b, 1995). Recent expansion includes colonisation of
the region around Uppsala, and the lower Tornë watershed
on the border with Finland, the latter the result of many
but mostly very small (one pair) reintroductions, 1960–
1993, on the Swedish side of the watershed (Mangi 2008).
Range expansion on the southern margins has been slow.
Beavers now occupy the whole of Sweden apart from the
far north (roughly, north of 67°N) and the south (roughly,
south of 58°N); the population is estimated at 130000
(Belova et al. 2016).
A searchable mapping database of the Swedish distribu-
tion is available (Shah & Coulson 2019); however, the
range mapped (largely through citizen science reports) is
an underestimate, particularly in the sparsely inhabited
north, where there is an observer bias near towns with
academic institutions (G. Hartman, personal
Online maps of beaver distribution in Switzerland are
available at index.php?nuesp
=70807 (Appendix S16), with a comprehensive website
at fachs telle.html
Minnig et al. (2016) provide a detailed account of re-
introduction history, 141 individuals at 30 sites in 1956–77.
Population was estimated at 2800 in 2015 (BAFU 2016).
Beavers are now distributed widely on the Rhine and
Rhone watersheds, with recent settlement on the Inn
(Danube basin) from populations in Austria.
In the late 1930s, fewer than 100 Eurasian beavers survived
in the Pripet Marshes. In the Rovno region, there was a
colony of North American beavers brought from Poznań,
Poland, in 1933 and 1934, but by the mid-1950s, this
population had disappeared (Marin 1954). All beavers in
Ukraine today are Castor fiber.
20th-Century restoration is well documented (Safonov
& Pavlov 1973). Recent expansion has been rapid. From
2001 to 2010, the number of beavers in Ukraine increased
by more than three times and in some areas up to 7.3
times (Matsiboruk 2013a). Average annual population
growth was 24% (Matsiboruk 2013b). Beavers have begun
to colonise both eastern Ukraine (Tokarsky 2009,
Brusentsova & Ukrainskiy 2015) and southern Ukraine
(Volokh 2011).
D. J. Halley, A. P. Saveljev and F. Rosell
Population and distribution of beavers in Eurasia
16 Mammal Review 51 (2021) 1–24 © 2021 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
The beaver’s geographic range now occupies half of
Ukraine, including areas of intensive human activity
(Tokarsky et al. 2012). By 2011, the population was close
to 50000 (official statistics have not been published since
The population in Transcarpathian Ukraine, derived
from the Hungarian Tisza reintroduction, is in rapid growth.
Beavers reached the region in 2003 and doubled in popu-
lation from 25 to 50 territories from 2009 to 2012 (Bashta
& Potish 2012), with further expansion later (Barkasi 2016).
Populations on the upper Dniester, on the eastern slope
of the Ukrainian Carpathians, were expanding in the same
period (Barkasi 2016).
Recent summers in Ukraine have been very arid, and
the winter of 2019/20 was snowless. This affected habitat
quality in beaver-inhabited rivers. Predation patterns have
also changed: in the Chernobyl Exclusion Zone, wolf Canis
lupus excrement was formerly dominated by the remains
of wild boar Sus scrofa, but now it is almost entirely
composed of beaver hair (V. Smagol, Institute of Zoology,
NAS Ukraine, personal communication).
The history of beavers in Wales is reviewed by Coles
(2019). Reintroduction was assessed as feasible in 2009
(Halley et al. 2009, Jones et al. 2011). There were four
known fenced populations in 2018 (Swaile et al. 2018),
with field signs in the wild on the River Dyfi in west
Wales; beaver signs are widespread on the River Wye on
the English border, though a 2018–19 survey found no
strong evidence of beavers living in established territories
(R. Campbell-Palmer, A. Leow-Dyke, personal communica-
tion). Population size is unknown.
Beaver populations have continued to spread and grow
in size, confirming the prediction (Halley & Rosell 2002)
that the species will, within a few decades, be a fairly
common animal throughout most of its former range.
The current ‘minimum population estimate’ is 1479863
(which can be rounded as 1.5 million), 3.5 times the 1998
estimate (Nolet & Rosell 1998), and 2.5 times the 2002
estimate (Halley & Rosell 2002), using the same methods
of adding together the estimates for each country and
using the lower limit where a range is given. This is not
meant as anything other than a general guide.
Population development during recolonisation was re-
viewed by Halley and Rosell (2002). Developments since
have followed the same general pattern (e.g. John et al.
2010, Bártak et al. 2013, Zwolicki et al. 2019): colonisa-
tion of new watersheds by natural spread or reintroduction;
a period of relatively slow population growth, but rapid
range extension, followed by a phase of rapid population
increase; then a period of population decline as habitat
not capable of supporting beavers permanently is inhabited
and temporarily exhausted; and finally, rough stability.
The size of the watershed influences the length of these
stages, which are shorter in smaller watersheds; as do hu-
man interventions, in particular hunting and the pattern
of hunting (Parker & Rosell 2003, 2012).
In much of European Russia, Belarus, Ukraine, Poland,
Latvia, and Lithuania, populations appear to be nearing
maturity, with increases peaking in Poland and declines
reported in Belarus according to official statistics.
However, these trends are general, and the accuracy of
the statistics and the potential influence of biasing fac-
tors are open to question. Nevertheless, there is a general
consistency across the region which indicates that the
period of rapid population growth has probably now
In the Nordic countries, range expansion has slowed
in recent years (though not stopped), and the popula-
tion is mature in many areas. The main exception is
Finnish Lapland, currently under invasion from the west
by Castor fiber and from the south and east by Castor
canadensis (Fig. 4). In Norway, reintroduced populations
of Castor fiber north of the Arctic Circle are now ap-
parently extinct. Habitat patches are in general small
and, except in Finnmark, the terrain is very rugged,
inhibiting dispersal. At present, the northern limit of
both Castor fiber and Castor canadensis coincides roughly
with 67°N all the way from the Norwegian Sea to the
White Sea, even the populations of both species colonis-
ing the Kemi watershed in Finnish Lapland are mostly
south of this limit or very close to it. The only estab-
lished population to the north is in central Murmansk
Oblast at about 68°30N, the status of which is little
Castor canadensis is present to the shore of the Arctic
Ocean on the Mackenzie Delta, Canada, and in the ad-
jacent Alaskan north slope at 69°N (Tape et al. 2018), in
a winter climate much more severe that similar latitudes
in the Gulf-Stream influenced Nordics. In climatically
modern times (subfossils dated to c. 1350 before present),
beavers occurred to at least 70°N at Varanger in extreme
north-east Norway (Horn et al. 2014). Therefore, it seems
unlikely that climate is limiting beaver populations in the
north. The current populations all derive from refugia
much further south and have limited genetic diversity,
which may influence their ability to adapt to the extreme
conditions of the Arctic. However, the apparent pattern
may be a temporary effect of reintroduction history. The
Kemi watershed extends through most of Finnish Lapland
from 65°30N to 68°30N in largely flat terrain, with no
Population and distribution of beavers in Eurasia
D. J. Halley, A. P. Saveljev and F. Rosell
Mammal Review 51 (2021) 1–24 © 2020 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
barriers to dispersal (Fig. 6), so whether there is a real
climatic effect limiting northern distribution at around
67°N will soon become evident.
In the south, the Mediterranean Sea forms a natural
southern limit to the range in the west; it has formed an
effective barrier to most mammal dispersal for at least
5.5 million years (Groves & DiCastri 1991). Beavers for-
merly occurred in steppe and desert regions south to the
Tigris/Euphrates (Legge & Rowley-Conwy 1986), where
rivers flowing in from the north provided a dispersal route
and riparian vegetation permitted. River systems populated
by crocodiles or alligators – north to the Yangtze, Indian
subcontinent, and Nile – would presumably be unsuitable
for beavers.
Differences in chromosome number (Castor fiber 48, Castor
canadensis 40), anal gland secretion differences, failed
crossbreeding attempts, and lack of observations of hybrids
in the wild all confirm that Castor fiber and Castor canadensis
are distinct species which cannot hybridise (Lavrov & Orlov
1973, Lavrov 1983, Rosell & Sun 1999).
Because of the early history of Castor fiber and Castor
canadensis in Finland and north-west Russia, it has some-
times been assumed that the relationship is one of straight-
forward dominance of canadensis over fiber. Recent data
suggest that this is an oversimplification, without identifying
a general pattern which would allow prediction of the
outcome. In some places, such as France and Luxembourg,
wild-living Castor canadensis populations have been re-
moved (Dewas et al. 2012, Herr et al. 2018). However,
Castor canadensis were also introduced in the Styr River
(Dnieper watershed) in north-west Ukraine in 1924 and
to East Prussia (now Poland) in 1926. Beavers of unknown
origin were also released here in 1942–43 (Panfil 1971)
and four Voronezh Castor fiber in 1961 (Klarowski 1983).
In 1979, a survey revealed that all animals in the region
were now Castor fiber (Zurowski 1980, cited in Saveljev
1989). In 1933–34, seven North American beavers were
released in the Rovno region, then in Poland (now in
Ukraine), and survived to 1956 but then died out (Dezhkin
1960). A similar decline in the Castor canadensis colony
in the Russian Far East (see above) is currently evident.
In Austria, 12 individuals of Castor canadensis were rein-
troduced, along with 40 Eurasian beavers, in the 1980s
(Sieber 2000). In each of these cases, canadensis has dis-
appeared, apparently through intraspecific competition with
fiber in the latter case at least (Sieber 2000).
The species meet at the eastern and western edges of
the Finnish – north-west Russian range of Castor canadensis.
At present, fiber is slowly losing ground to canadensis in
south-west Finland (LUKE 2019), and on the line of
contact between the White Sea and Lake Onega, but
canadensis has lost ground to fiber by c. 50 km on the
Karelian isthmus north of St. Petersburg (Saveljev 1989,
Danilov et al. 2011, Danilov & Fyodorov 2015, 2016).
We agree with Danilov and Fyodorov (2015, 2016) that
the factors which can ‘tip the balance’ are likely to be
complex and may include local biotic conditions and the
provenance and genetic diversity of founder populations.
The results of interspecific competition are also likely to
be influenced by the relative sizes of each population,
and so by the number of dispersers, once beavers have
spread throughout a river system and begin to compete
for space (Parker et al. 2012).
For management, a policy of active preference for Castor
fiber over Castor canadensis, especially where populations
are not yet ‘consolidated’ (such as on the Kemi, see Fig. 6
and section on Finland), is clearly preferable, rather than
hoping that fiber will prove to be the dominant species.
European Union member states are required to extermi-
nate, control or contain North American beaver and other
invasive species (European Union 2014).
Recolonisation of the beaver’s former range and, to a
lesser extent, re-establishment of the population in Europe
north and east of Poland–Ukraine inclusive is, with excep-
tions, substantially complete; and within a few decades it
will be complete. In Fenno-Scandia, spread into southern
Sweden, western Norway, and north of the Arctic Circle
can be expected.
In contrast, in most of continental Europe west of
Poland – Ukraine, and in Great Britain, populations
are in the early colonisation or rapid increase phases,
though they are at or near maturity in some regions
such as Bavaria and the lower Rhone. Both range and
population size will increase rapidly in this region in
coming decades, roughly one third of the land mass
of Europe with well over two thirds of its human
Beavers remain absent from most of their former range
in southern Europe – Portugal, Spain excepting the Ebro,
Italy, and the Balkans from Bulgaria – Montenegro
In Asia, there are extensive regions of Russia, especially
eastern Siberia, where beavers remain absent; scope for
further increases in range and population is correspond-
ingly large. Beavers of Voronezh/Belarus origin released
in Siberia are now in contact with native populations in
Western Siberia (Castor fiber pohlei) and the upper Yenisei
(Castor fiber tuvinicus). Management may be necessary if
maintenance of unmodified forms of the two autochtho-
nous gene pools is to be maintained.
D. J. Halley, A. P. Saveljev and F. Rosell
Population and distribution of beavers in Eurasia
18 Mammal Review 51 (2021) 1–24 © 2021 The Authors. Mammal Review published by Mammal Society and John Wiley & Sons Ltd.
The Danube basin is the world’s most international
watershed, draining part or all of 19 countries. Thirty
years ago, beavers were confined to the upper reaches
from Bavaria to Vienna. As a result of reintroductions
and natural spread, the species is now present and rapidly
increasing on the main river as far as the Hungarian–
Serbian border near Mohacs, and on the main tributaries
Morava, Tisza, Sava, Drava, and Mures. There are no
serious barriers to further spread on the Danube basin
above the Iron Gates barrages, on the Serbia–Romania
border about 150 km south-east of Belgrade. There are
many rapidly expanding beaver populations, increasingly
linked. Downstream of the Iron Gates, the Olt and Ialomița
populations are also in rapid expansion, and a population
is now established in the Danube Delta. Populations are
relatively small at present, about 1800 animals in 2017
(Pașca et al. 2018), and the area of unoccupied habitat
is very large. Absent further reintroductions, it will take
rather longer for populations in the lower basin to expand
throughout, but, as with the upper basin, there are few
barriers to spread.
The western extremity of the North European Plain
– northern France, Benelux, north-west Germany and
England – is one of the most densely populated, human-
modified regions in the world. The terrain is flat or low
relief. Beaver populations are now widely established in
the region, and the period of rapid population increase
is under way or about to commence. Management is likely
to become an increasingly salient issue in coming decades.
Other than in the Netherlands, there is little sign at pre-
sent that it will be other than reactive.
“How will the increasing beaver populations across
Europe, and their associated impacts on flow regimes,
water quality and biodiversity, affect river catchment res-
toration?” was recently identified as a ‘priority question’
for landscape restoration in Europe (Ockendon et al. 2018).
Understanding, adapting to, and utilising these impacts
should be a priority for river catchment management in
general. Attention to this and to framing public debate,
on the basis of the detailed information and experience
already available (Thompson et al. in press), would be
both wise and cost-effective. As most large watersheds,
and most beaver populations, in central and western Europe
are international, European Union mechanisms for research
and development of management methods would perhaps
be most appropriate.
We are indebted to the many individuals who have con-
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online version of this article at the publisher’s web-site.
Appendix S1. Beaver distribution in Austria in 2012.
Appendix S2. Beaver distribution in Belgium in 2016.
Appendix S3. Beaver distribution in Bosnia & Herzegovina
in 2015.
Appendix S4. Beaver distribution in the Czech Republic
in 2015.
Appendix S5. Beaver distribution in Jylland, Denmark in
Appendix S6. Beaver distribution in England and Wales
in 2018.
Appendix S7. Beaver distribution in France in 2019.
Appendix S8. Beaver distribution in Germany in 2017.
Appendix S9. Beaver distribution in Luxembourg and
adjacent regions.
Appendix S10. Beaver distribution in the Netherlands in
Appendix S11. Distribution of beavers in Norway in 2019.
Appendix S12. Distribution of beaver territories in Scotland,
Tay and Forth watersheds.
Appendix S13. Beaver distribution in Serbia in 2015.
Appendix S14. Beaver distribution in Slovenia in 2015.
Appendix S15. Distribution of beavers in Spain (Ebro
watershed) in 2019.
Appendix S16. Distribution of beavers in Switzerland in 2019.
... Reintroductions of beavers were partly to maintain the beaver population, but also to exploit the ecosystem engineering activities of beavers and thus to rewild landscapes and to bring back natural ecological processes (Law et al., 2017;Marr et al., 2018;Halley et al., 2021). The beaver has proven particularly successful in repopulating new territories following reintroductions (Halley et al., 2021, Bouros et al., 2022. ...
... Reintroductions of beavers were partly to maintain the beaver population, but also to exploit the ecosystem engineering activities of beavers and thus to rewild landscapes and to bring back natural ecological processes (Law et al., 2017;Marr et al., 2018;Halley et al., 2021). The beaver has proven particularly successful in repopulating new territories following reintroductions (Halley et al., 2021, Bouros et al., 2022. In Europe, reintroduction programs have restored the beaver population from only 1200 individuals in the early 20th century to more than 1.2 million individuals in 2020 (Wróbel, 2020;Halley et al., 2021). ...
... The beaver has proven particularly successful in repopulating new territories following reintroductions (Halley et al., 2021, Bouros et al., 2022. In Europe, reintroduction programs have restored the beaver population from only 1200 individuals in the early 20th century to more than 1.2 million individuals in 2020 (Wróbel, 2020;Halley et al., 2021). There are several good examples of the natural expansion of beaver populations after releases (Bouros et al., 2022; but see also Ceña et al., 2004). ...
Full-text available
Ecosystem engineering species, such as beavers, may help the restoration of biodiversity. Through the building of dams and lodges and altering the natural hydrology, beavers change the habitat structure and create multiple habitats that facilitate a wide variety of other organisms including terrestrial invertebrate communities. Here we study the effect of beaver reintroduction in Klosterheden in Denmark on biomass of flying invertebrates and diversity of moths. Further, aerial photos were used to assess riparian structure and productivity using the normalized difference vegetation index (NDVI). Our findings show that the presence of beavers affected flying invertebrate biomass, but that this was dependent on time of the year. Further, a strong effect of presence of beavers was found on diversity of moths. The results also show an increase in vegetation productivity and structural heterogeneity at sites with presence of beavers. Overall, our results demonstrate the importance of beavers as important ecosystem engineers that affect invertebrate species composition and abundance, as well as riparian structure and productivity.
... This species was widely distributed in Palearctic, but populations started declining during the Middle Ages, up to the 19th century, mainly due to hunting for fur, meat and castoreum (a secretion produced for scent marking and historically used by humans for medical purposes) (Halley and Rosell 2002;Halley et al. 2021). Some estimates account for only 1200 individuals surviving, divided into eight isolated populations in the Palearctic, of which five of them in its Westernmost part (Nolet and Rosell 1998;Halley et al. 2021). ...
... This species was widely distributed in Palearctic, but populations started declining during the Middle Ages, up to the 19th century, mainly due to hunting for fur, meat and castoreum (a secretion produced for scent marking and historically used by humans for medical purposes) (Halley and Rosell 2002;Halley et al. 2021). Some estimates account for only 1200 individuals surviving, divided into eight isolated populations in the Palearctic, of which five of them in its Westernmost part (Nolet and Rosell 1998;Halley et al. 2021). In recent times, legal protection and reintroductions (started since 1922), together with natural recolonization, allowing the species to recover (Halley and Rosell 2002;Halley et al. 2021). ...
... Some estimates account for only 1200 individuals surviving, divided into eight isolated populations in the Palearctic, of which five of them in its Westernmost part (Nolet and Rosell 1998;Halley et al. 2021). In recent times, legal protection and reintroductions (started since 1922), together with natural recolonization, allowing the species to recover (Halley and Rosell 2002;Halley et al. 2021). ...
Full-text available
The Eurasian beaver is a keystone species and landscape-capable ecosystem engineer, which went close to extinction until the 19th century. Recently, thanks to legal protection and reintroduction programs, the species has recolonized much of its past range. However, in some countries this process did not occur. Objectives. Our objective is to model the potential distribution of the Eurasian beaver for current and future conditions, on a continental scale, at river and sub-basin level. We focus on the protected areas of Italy and Portugal for possible reintroductions. Methods. The study area is Europe, with a subset focusing on Italy and Portugal. We produce species distribution models for current and future conditions using climate change scenarios and predicting changes in river flow, including topographic and human disturbance variables. We then deepen suitability-related issues within Italian and Portuguese protected areas. Results. We find that the Eurasian beaver current suitability is comparable to its known distribution, although some potentially-suitable spots occur in Italy (where there are two occurrences), while the Iberian Peninsula and the Balkan countries host scattered suitable spots. Future scenarios predict a general lowering of suitability in Central and Northern Europe. Portuguese protected areas generally host unsuitable territories, while the Italian ones have reported a tangled scenario, depending on the biogeographical sector. Conclusions. Our results may support the large-scale management of the beaver, both for countries already hosting this species and those planning a reintroduction. The framework used may be applied to other species, and for different topics, from biogeography to conservation.
... It became extinct in the Iberian Peninsula around the XVIIth century [1,22]. Recently, it was returned to Spain through an unofficial release in the Ebro River basin in 2003 [38], and it is now considered a protected species in the country [11]. ...
... No habitat suitability studies have been undertaken for the beaver in Portugal or Spain. Yet based on known past presence, habitat requirements [33], populations in similar climates [38] and foraged tree species [17], it is likely that suitable habitats exist [74], with the northern wetter areas potentially preferable to southern drier ones. ...
... Wildlife translocations are valuable conservation tools for recovering imperiled species, reducing human-wildlife conflict, and restoring degraded ecosystems (Germano et al., 2015;Mengak, 2018;Novak, Phelan, & Weber, 2021). American and Eurasian beavers (Castor canadensis and C. fiber) have been translocated for over 70 years after extensive extirpation from much of their historical ranges during the fur trade of the 1700s-1800s (Baker & Hill, 2003;Halley, Saveljev, & Rosell, 2021). Translocations of American beavers (hereafter, beavers) in the United States often focus on removing nuisance individuals from conflict situations where they would otherwise be lethally removed and using them instead as ecosystem engineers for riparian restoration; they may increase the large woody debris contribution and the number of dams in the system to initiate process-based restoration and improve degraded systems (Naiman, Johnston, & Kelley, 1988;Pollock et al., 2014;Pilliod et al., 2018). ...
... Previous beaver studies have tied success to the translocation of entire colonies or opposite-sex pairs, often requiring multiple release efforts Petro et al., 2015;Brick & Woodruff, 2019). Persistent release efforts have also contributed to the successful translocation establishment of Eurasian beaver (Dewas et al., 2012;Halley et al., 2021). ...
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Wildlife translocation facilitates conservation efforts, including recovering imperiled species, reducing human–wildlife conflict, and restoring degraded ecosystems. Beaver (American, Castor canadensis; Eurasian, C. fiber) translocation may mitigate human–wildlife conflict and facilitate ecosystem restoration. However, few projects measure outcomes of translocations by monitoring beaver postrelease, and translocation to desert streams is relatively rare. We captured, tagged, and monitored 47 American beavers (hereafter, beavers) which we then translocated to two desert rivers in Utah, USA, to assist in passive river restoration. We compared translocated beaver site fidelity, survival, and dam‐building behavior to 24 resident beavers. We observed high apparent survival (i.e., survived and stayed in the study site) for eight weeks postrelease of resident adult beavers (0.88 ± 0.08; standard error) and lower but similar apparent survival rates between resident subadult (0.15 ± 0.15), translocated adult (0.26 ± 0.12), and translocated subadult beavers (0.09 ± 0.08). Neither the pre‐ nor the post‐translocation count of river reaches with beaver dams were predicted well by the Beaver Restoration Assessment Tool, which estimates maximum beaver dam capacity by river reach, suggesting beaver‐related restoration is not maximized in these rivers. Translocated beavers exhibited similar characteristics as resident subadult beavers during dispersal; they were more vulnerable to predation and many emigrated from the study sites. High mortality and low site fidelity should be anticipated when translocating beavers, but even so, translocation may have contributed to additional beaver dams in the restoration sites, which is the common goal of beaver‐assisted river restoration. Multiple releases at targeted restoration sites may eventually result in establishment and meet conservation objectives for desert rivers. A translocated American beaver was released into the Price River, Utah, after it was fitted with a VHF tag on its tail.
... In recent years, the population of the Eurasian beaver Castor fiber has increased significantly in both numbers and distribution; at present there are one and a half million beavers recolonizing the species' historical range throughout the northern hemisphere (Halley et al., 2021). Beavers are ecological engineers, substantially modifying the existing environment by altering its biotic and abiotic components (Jones et al., 2010). ...
... The beaver population of Poland, at present increasing in number and range, is estimated at 120 000 individuals, and is common throughout the country (Halley et al., 2021). The current situation is the upshot of a reintroduction programme conducted since 1975, when beavers were still rare in the country (Żurowski and Kasperczyk, 1988). ...
The engineering activities of the Eurasian beaver Castor fiber have far-reaching effects on the components of an environment and therefore modify the functioning of the ecosystem. The wetlands thereby created are the most conspicuous effect of beaver activity and attract water-related species. However, there is some evidence suggesting that beavers influence not only aquatic ecosystems but also the terrestrial habitats adjacent to these wetlands and the organisms occurring there. Because the impact of beavers on terrestrial birds is still poorly understood, this study evaluates the assemblage of birds wintering on beaver sites (N = 65) and paired reference sites (N = 65) in temperate forests of central Europe. We investigated the correlations between beaver presence, parameters of wetland areas, terrestrial vegetation characteristics, distance from the water's edge and bird species richness and abundance. We found a greater species richness and abundance of wintering birds on beaver sites than on watercourses unmodified by this ecosystem engineer (by 38% and 61%, respectively). Species richness and abundance were higher in the terrestrial habitats near the edges of beaver ponds, but for some species this tendency also held in forests growing at some distance from beaver wetlands. Greater species richness was related to beaver presence, but also increased with a more open canopy and greater forest floor diversity, whereas bird abundance was correlated only with canopy openness. The beaver sites attracted primary cavity nesters, secondary cavity nesters and frugivorous species. This study provides evidence that the engineering activities of beavers during the growing season have a delayed cascading effect on the richness and abundance of the bird assemblage in terrestrial habitats in winter. This indicates that beaver ecosystem engineering should be seen as having a potential for carry-over effects, in which the consequences of beaver activity become apparent in subsequent seasons. Birds are considered to be ecological indicators, so our results highlight the importance of beavers for the distribution of terrestrial organisms at the local scale, and therefore the functioning of ecosystems beyond the immediate wetland area.
... Der Biber ist ein Beispiel für erfolgreiche Naturschutzbemühungen, bei denen es aber auch zu Konflikten kommt. Gezielte Umsiedlungen und andere Schutzmaßnahmen haben insbesondere in Süd-und Ostdeutschland zu stabilen BiberpopulaƟonen geführt (Halley et al. 2021). ...
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Zusammenfassung Die menschliche Perspektive in Mensch-Wildtier-Konflikten zu verstehen, ist wichtig für ein ganzheitliches Naturschutzmanagement, da solche Konflikte oft über die direkten Mensch-Tier-Begegnungen und deren negative Folgen hinausgehen. Stattdessen handelt es sich zumeist um Konflikte zwischen Menschen bzw. Interessengruppen („Stakeholder“), die verschiedene Ansichten zu Wildtieren und deren Management vertreten. Als „Human Dimensions of Wildlife“ (HDW) werden die menschlichen oder gesellschaftlichen Aspekte in Bezug auf Natur und Wildtiere bezeichnet. Als eine Teildisziplin der Naturschutzsozialwissenschaften („Conservation Social Sciences“) stellen sie einen inter- und transdisziplinären Ansatz dar, der vielfältige Sichtweisen zur Lösung komplexer Sachverhalte in das Naturschutzmanagement einbezieht. In diesem Kapitel stellen wir gängige HDW-Konzepte vor, illustrieren auf der Grundlage einschlägiger Literatur deren analytisches Potenzial und veranschaulichen praktische Anwendungen in Fallbeispielen zu Wölfen, Wildschweinen, Bibern und Fledermäusen. Abstract Understanding people in the context of human-wildlife conflict is important for integrated conservation management, as these conflicts often go beyond people and their direct encounters with or negative impacts from wildlife. They often include social conflicts among people or groups of people (i.e. stakeholders) that have different views about wildlife and how it should be managed. “Human Dimensions of Wildlife” (HDW) is a sub-discipline of “Conservation Social Sciences”, an inter- and transdisciplinary approach including multiple perspectives to solve complex conservation and management matters. Here, we provide an overview of frequently used HDW concepts and illustrate their predictive potential with evidence from the literature, while their practical application is demonstrated with examples and case studies on wolves, boars, beavers and bats.
... In the eighteenth century, it had been exterminated mainly not only due to the damage it caused but also due to the harvesting of castoreum, meat, and fur. The recent return of the beaver began in the 1980s when the first beavers recolonized the Czech Republic along the Morava River from Austria, where the beaver was reintroduced (Halley et al. 2021;Šafář 2002). Over time, beavers also moved into the Czech Republic along the Elbe from Germany and were reintroduced to other places in the Czech Republic during the 1990s (Kostkan and Lehký 1997). ...
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Over the last few decades, the beaver has settled most of the European continent including the Czech Republic. Until recently, it was a species typically found in lowland floodplain forests. However, as the population grew, it began to spread into the agricultural landscape, where it has less favorable living conditions. In the Czech Republic, beavers were known to concentrate especially in large forest complexes; little information is so far available about their settlement of the agricultural landscape. At the same time, beaver management in the Czech Republic is applied according to zones with different levels of protection for this species; therefore, the assessment of beaver populations in different environmental conditions is essential. The article focuses on beaver population density, home range length, distribution, and dispersion in the four main types of environments, as well as on differences in food selection. Beaver population density is the highest in forest localities with small streams; the beaver finds the most suitable living conditions there. On average, the agricultural landscape is less populated. While population density in large rivers in the agricultural landscape is comparable to that in large rivers in the forest due to environmental conditions that can be similar, small streams in the agricultural landscape are inhabited 3.3 times less than small streams in forests because they are generally unsuitable environments with limited resources. The lengths of home ranges depend on the environment, the density of beavers, and especially on the availability of food resources. Likewise, the distances that beavers search for food differ, thus affecting their spatial activity.
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Beavers and their dams, once common in small streams throughout Eurasia and North America, are returning to their original range. Beaver dams are special habitats in the aquatic-terrestrial interface, but their macroinvertebrate fauna is hardly known. This study aims to qualify and quantify the invertebrate fauna of beaver dams, taking into account the maintenance status of the dams. Nine different areas covering the top, middle and base zones of eight maintained and eight abandoned beaver dams were sampled using a specially designed suction device. The invertebrate community of beaver dams proved to be diverse and predominantly rheophilic. Community composition reflects higher flow velocities in streamside habitats; this gradient is more pronounced in maintained than in abandoned dams. Shredders are the most abundant feeding type, but they are less frequent in streamside habitats, in particular in those of maintained dams, where passive filter feeders prevail. Terrestrial/Semiaquatic taxa colonising hygropetric areas were found in streamside habitats especially of abandoned dams. Our results underline that beaver dams provide a wide variety of environmental conditions and habitat types, which enhance biodiversity in small streams and their floodplains. Beaver dams in riverine landscapes should therefore be much more valued in floodplain conservation and management.
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Wild boar ( Sus scrofa ) seriously threatens agricultural production and rural livelihoods worldwide. Human-wild boar conflict has become an important research topic. We investigated the impact of wild boars on local livelihoods and rural community’s responses to the impact, based on a case study in Fu’an City, Fujian Province, Southeast China. We conducted a questionnaire survey of 152 rural households in six villages of Fu’an City. Results demonstrated that the overall impact of wild boars on livelihoods was significant; however, there was impact heterogeneity depending on the mentioned factors. The impact lied on the scale of agricultural output value and the extent of livelihood dependency on susceptible crops. Different impacts resulted in discrepant countermeasures of rural households in dealing with wild boars. The magnitude of economic losses caused by wild boars to a great extent determined the selection of prevention measures. Rural households with more economic losses tend to adopt costly and effective prevention measures, such as fencing and human guarding. Recommendations were put forward to address the human-wild boar conflicts in China.
Widely available ‘fish‐finder’ echo‐sounding devices are beginning to be used in bathymetric studies to estimate geomorphic change. To date, however, there have been no applications in shallow and complex wetlands, where changes in sediment storage are notoriously dynamic in time and difficult to describe accurately in space. Therefore, in this study, we tested the performance of an ‘off‐the‐shelf’ fish‐finder for mapping bathymetry in a shallow beaver pond. We tested fish‐finder sonar depth readings against a traditional‐sounding lead‐line method across 21 paired Sampling Points with a minimum depth of 0.31 m and a mean of 0.65 m. Spatial accuracy of the unit was also tested against a differentially corrected Global Navigation Satellite System (GNSS) receiver. Measured depths to pond bottom from the fish‐finder were on average within 5%, although significantly 0.015 m (SD = 0.034) less than those obtained by the lead‐line method. Spatial accuracy, however, varied greatly compared to the corrected GNSS receiver readings, with a mean discrepancy of 2.7 m (SD = 1.5) but up to 6.2 m. Given the close match of depth readings between the two methods, we conclude that sonar is a suitable, cost‐effective, and less‐intrusive method than existing techniques, even in moderately vegetated shallow waterbodies. Methods do need to be adopted to account for poor spatial precision with ‘off‐the‐shelf’ fish‐finder models, but this can be rectified with survey design or using a secondary GNSS. Application of this technology will allow rapid one‐off surveys or repeated monitoring of depth, bedform and sediment accumulation in otherwise hard‐to‐access or disturbance‐sensitive wetlands, such as beaver ponds and water treatment or flow attenuation wetlands.
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Founder genetic composition can affect reintroduction success, especially as the number of animals released tends to be small and therefore less genetically diverse than their source populations. Numerous translocations and reinforcements of beavers, Castor fiber, have occurred with little regard to geographic and/or genetic origin. Beaver reintroduction to Britain has been haphazard and currently disjointed populations of varying status exist – from sanctioned wild releases, unlicensed populations and naturalistic enclosed projects. This study investigated the genetic composition of two originally unofficially released beaver populations in Britain - Tayside, east Scotland, and River Otter, Devon, to provide data to support decision on their future management. From both wild populations (n = 34Tayside, n = 9Devon) all were confirmed as Eurasian beaver. The vast majority, origin was likely assignable to Germany and the mixed founder population of Bavaria. Eighty-two percent of the Tayside individuals examined at 275 loci were at least as closely related as first cousins, with pairwise estimates of relatedness at 26 loci indicated that the Devon beavers were more closely related on average. So far there is no evidence to suggest that beavers are failing to adapt to the British environment despite their reduced genetic founder based, however attention to genetic augmentation and longer-term management of genetic diversity should be factored into comprehensive restoration plans for the species across Britain. Many recent reintroductions are relying on serial founder events from an already limited founder base and that is counter to best practice in reintroduction planning.
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Beaver was once abundant throughout the Baltic Sea Region . Intensive hunting and capture, together with changes in human land use, led to the total extinction of the species in the watersheds of the Baltic Sea basin. Beaver populations are now considered to have reached densities causing substantial damage levels, e.g. in the south eastern Baltic Sea countries. The content of this book includes: • General information on beaver populations and management needs around the Baltic Sea. • Country-specific legislation and policy concerning beaver management, damage control and hunting/trapping. • Examples of raising the value of beaver as a resource for various purposes. • Presentation of a “Beaver Tool” for facilitating management decisions. • Good practices from seven countries. • Results on the beavers’ role for water quality. • Suggestions for reduction of leakage of nutrients and hazardous substances to watersheds and the Baltic. • Help in implementing EU directives and other conservation, environmental and water protection documents. • Suggestions for national / regional species management plans.
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The management of biological resources, which have important utilitarian value, should be based on current knowledge about their distribution, abundance, and taxonomic status. The data on the dynamics of distribution ranges and species abundance are easily available in the literature. However, it would seem that the simplest information - the lists of game species and their regional differences - is difficult for mammalogists to access. We compared the modern lists of hunting mammals of Estonia, Latvia, Lithuania, Poland, Belarus, and the bordering region of the Russian Federation with respect to their quantitative composition, structure and taxonomic verification. General trends and national features of the lists are noted. The recent and historical changes in the lists of objects of legal hunting are analysed. Управление биологическими ресурсами, имеющими важное утилитарное значение, должно базироваться на современных представлениях об их распространении, численности и таксономическом статусе. Но если в литературе достаточно данных о динамике численности и ареалов охотничьих видов, то, казалось бы, самая простая информация - списки видов дичи и их региональные различия - практически недоступна для териологов. Мы провели сравнение современных списков охотничьих видов млекопитающих Белоруссии, Латвии, Литвы, Польши, Эстонии и граничащих с ними районов Российской Федерации относительно их количественного состава, структуры и таксономической верификации. Отмечены общие тенденции и национальные особенности составления списков. Проанализированы исторические и современные изменения перечней объектов легальной охоты.
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The Eurasian beaver (Castor fiber) is an intensely expansionary species. Species reintroductions, which were conducted in various parts of Europe, as well as the rate of natural increase have resulted in the growth of the number of individuals. The dynamic development of the beaver population in Europe means that the available data concerning the quantity of beavers becomes outdated very quickly. The purpose of this manuscript is to update the available information about the beaver populations in all countries of Europe. The information was collected in the second half of 2019. Some of the data collected include generally available studies and articles. For some countries, the author had difficulty obtaining any data; therefore, personal communication was employed with various governmental or scientific units in the given country. The outcomes of the conducted analyses were figures about the Eurasian beaver population throughout Europe. It was found that the Eurasian beaver population in Europe numbered nearly 1,222,000 individuals.
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Abstract. The paper summarizes numerous scattered data on the former distribution of the Eurasian bea­ ver in Bulgaria, many of them unpublished till now. Data on fossil, subfossil and subrecent records (Late Pleistocene to 19th century AD) from 43 localities situated in 19 of the total of 28 provinces, and 209 bone/teeth finds of at least 11 skeletal elements are presented. Geographical, altitudinal and chronological distribution as well as the representation of particular rivers and skeleton elements is presented and analyzed. About 56 percent of the sites were situated between 100 and 300 m. a. s. l. Five sites revealed Paleolithic occurrence, 20 Neolithic, 12 Chalcolithic, six Bronze Age, two Iron Age. The six subrecent sites are dated to the last 2000 years. The species distribution declined about and after the turn of BC/AD. The 43 localities were situated along 28 Bulgarian rivers. All finds are related to lowland areas and lower courses of the rivers. Evidence for the last record in the country is dated 1750-1850 AD.
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In the medieval period, beavers were widespread throughout Poland. In the 13th century, the number of animals began to decline. After the Second World War, it was assumed that Eurasian beavers were no longer present in Poland. In 1974, the programme “Active protection of European beavers in Poland” was launched. Reintroduction was a very important element of the active protection of beavers. According to an inventory carried out in 1977, there were estimated to be over 1000 individuals. Long-term reintroduction efforts by hunters and scientists have caused beavers to be present throughout Poland. In recent years, the beaver population has increased significantly. This situation has created conflicts between farmers, foresters and beavers. In Poland, the beaver is considered a partly protected species. This manuscript discusses the dynamics of the beaver population in Poland and needed actions for future sustainable management.
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Article is devoted to the dynamics of European beaver Castor fiber in the Ukrainian Carpathians. Beaver expansion has like explosive character in the Transcarpathian part of the region. Beaver was noted here at the first time in 2003-2005, and in 2007 the region was inhabited by at least 14 settlements. The beavers distributed up the river Tisa at about 100 km during 2007-2009. 25 beaver settlements have been found in 2009 and about 50 ones in 2012 at the territory of Transcarpathia. Expansion of beavers continues in the upper basin of the Dniester. Supposedly, beaver population included about 400 ins. in the region of Ukrainian Carpathians. Distribution has mostly canalized character, namely along the major rivers and their tributaries. Geographical barriers - namely mountain ridges may restrict the movement of animals into the territory of the neighboring river basin systems. Key words: European beaver, Castor fiber, distribution, expansion, Ukrainian Carpathians
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The article discusses the role of the Green Belt of Fennoscandia (GBF) in the dispersal of beavers. The study is focused primarily on old-growth forests inside GBF, which have been preserved in the Kostomukshsky Strict Nature Reserve and Kalevalsky National Park. The specific characteristics of these areas – the abundance of waterbodies and the ex-tremely poor food resources for beavers – accelerated the dispersal of Canadian beavers from Finland towards the east and south of Karelia. Aspen, birch and willow, on which beavers mostly feed in winter, rarely occur in the north of GBF and are consumed by beavers in the first 2–3 years of their stay at a waterbody. The animals are therefore im-pelled to change their places of residence frequently and move over long distances. Surveys carried out in the Kostomukshsky Nature Reserve in 2018 showed that over 80 % of known colonies have been abandoned by beavers. In 2001, this proportion was 60 %. Most of the inhabited colonies were re-occupied, as a rule by single animals, who settled in an abandoned lodge temporarily. On average, the share of settlements occupied by single beavers was 45 %. The distribution of beaver settlements in old-growth north-taiga forests is very uneven, since it is more dependent on forage resources than on favorable habitat. It our surveys, 73.3 % (n = 75) of the settlements were situated in river stretches with water-logged birch and birch-spruce forests growing along their banks. A half of all settlements situated at watercourses were concentrated near highways, railways, power lines, i. e. where aspen and birch stands have formed after logging. We revealed a high building activity of beavers living in the north.
• We aimed to recognise beaver‐produced ecosystem services and quantify their theoretical value for the entire Northern Hemisphere. Activity of the Eurasian beaver Castor fiber and the North American beaver Castor canadensis in the landscape provides ecosystem services and disservices. Services produced by beaver activity include water purification, moderation of extreme events, habitat and biodiversity provision, nutrient cycling, greenhouse gas sequestration, recreational hunting and fishing, water supply, and non‐consumptive recreation. Beaver‐produced services have not been compiled, analysed, or quantified previously. • Each service we evaluated is worth millions to hundreds of millions of US dollars (USD) annually. Habitat and biodiversity provision (133 million USD), along with greenhouse gas sequestration (75 million USD), are particularly valuable services in absolute terms, while non‐consumptive recreation (167 USD ha⁻¹) and habitat and biodiversity provision (133 USD ha⁻¹) have the largest annual per‐hectare values. • Our results can be used to broaden decision‐making and management perspectives, as we offer value estimates to wildlife managers and municipality planners for assessing local site‐specific beaver wetland values and the opportunities for their realisation. Implementing Payments for Ecosystem Services schemes offer a concrete way for societies to benefit from beaver‐produced services while concurrently compensating beaver‐produced losses accrued to landowners. Building such schemes offer long‐term realisation of ecosystem services and damage mitigation. This would lead to increased societal well‐being and increased conservation interest and efforts.
Technical Report
Background There is a long, documented history of the Eurasian beaver Castor fiber and its former abundance throughout Britain (Coles 2006), and it is generally believed to have become extinct in Scotland, through over-hunting, by the 16th century (Kitchener and Conroy 1997). In 2009, an official trial reintroduction of beavers, the Scottish Beaver Trial (SBT), took place in mid-Argyll, though it later became apparent that a larger number of beavers existed through unauthorised releases in the Tayside area of Perthshire, with confirmed reports of their presence from around 2006. For the purposes of this survey ‘Tayside’ refers to the total catchments of the Rivers Tay and Earn. The largest beaver population in Scotland currently occurs in Tayside. A full survey was first undertaken in 2012 (Campbell et al. 2012). Scottish Government announced in November 2016 that it was minded to retain the Eurasian beaver in Scotland. The decision was informed by 20 years of work on beavers and beaver reintroduction issues, summarised in the SNH ‘Beavers in Scotland’ report (Gaywood 2015). The 2012 survey estimated that there were 38-39 groups of beavers present in the Tay catchment, equating to approximately 146 individual beavers (range 106 - 187). Further records were collated and presented in the final report of the Tayside Beaver Study Group (TBSG) published in 2015 (TBSG 2015), and more recent records received from the public were collated by SNH. The TBSG report highlighted the need for a resurvey of the Tay catchment to help inform decision-making for beaver management and clarify the current conservation status of the species. The need for a resurvey has also been confirmed by the current Scottish Beaver Forum. This report describes the findings of this survey, using density mapping of field signs and reports of beaver activity; makes comparisons with the 2012 survey; and estimates the current population size and distribution. It included surveys in catchments adjacent to Tayside, including sections of the Forth/Teith, Tay coastal catchments and the South Esk. Recommendations for future research and potential management strategies are also made. Main findings  Beaver activity was recorded throughout large parts of Tayside.  Beavers are spreading in distribution and are present outside the catchments of the Tay and Earn. Small numbers of territories occur within the Forth catchment from Loch Achray in the Trossachs, parts of the Teith and Devon, and the main stem of the Forth near Stirling.  No evidence of beaver presence was found on the South Esk nor in several freshwater bodies associated with the lower Forth and Forth estuary, including Loch Leven.  Distribution in Tayside ranged from as far north as Dunalastair Water, extending out to the River Dochart and River Lyon in the west, over to Forfar Loch in the east and down to Loch Earn in the south.  114 active beaver territorial zones were identified in this study, giving a conservatively estimated number of approximately 433 beavers (range 319 – 547). This number is based on a previously reported European mean group sizes of 3.8±1.0 animals per territory, which was also used in the 2012 SNH survey. Some identified zones may constitute multiple families and additional active territories, along with dispersing singletons, are likely to exist both within Tayside, especially on minor watercourses, and outside of the Tayside catchment which it was not feasible to cover during this survey.  Out of the 114 beaver territories defined using the 2017/2018 survey data, 100% were contained within the areas identified as ‘Potential Beaver Woodland’ and 95% were contained within the ‘Potential Core Beaver Woodland’, as defined by previous SNH GIS mapping exercises (Stringer et al. 2015).  Potential management issues were recorded at a total of 159 points, across 21 territories, ranging from dam building, collapsed burrows, tree felling, crop feeding and damage to fence lines.  A total of 86 dams, or sites where dams had been removed, were recorded. Of these, 41 dams occurred within one private estate.  There was an increase in both beaver distribution and density compared to the 2012 survey although spatial variability was evident, with areas of expansion and infilling, along with smaller areas of habitat abandonment potentially through culling.