REDIA, 104, 2021: 209-215 http://dx.doi.org/10.19263/REDIA-104.21.24
- Received 30 August 2021 Accepted 26 October 2021
EMILIANO MORIa,- ANDREA VIVIANObc - LEONARDO BRUSTENGAd - FRANCESCO OLIVETTIe - LUCA
PEPPUCCIf - CHIARA PUCCIg - DAVIDE SENSERINIh - UMBERTO SERGIACOMIi - CRISTIANO SPILINGAj -
PIO FEDERICO ROVERSIc - GIUSEPPE MAZZAc
DISTRIBUTION AND GENETIC ANALYSIS OF WILD-LIVING EURASIAN
BEAVERS IN CENTRAL ITALY
a) Consiglio Nazionale delle Ricerche, Istituto di Ricerca sugli Ecosistemi Terrestri, Via Madonna del Piano 10,
50019 Sesto Fiorentino (FI), Italy. E-mail: firstname.lastname@example.org. ORCID ID: 0000-0001-8108-7950.
b) Dipartimento di Scienze Agrarie, Alimentari e Agro-ambientali, Produzioni Agroalimentari e Gestione degli
Agroecosistemi, Università degli Studi di Pisa, Via del Borghetto 80, 56124 Pisa, Italy. E-mail:
email@example.com. ORCID: 0000-0002-2970-3389.
c) CREA Research Centre for Plant Protection and Certification, Via di Lanciola 12⁄a, Cascine del Riccio, 50125,
Firenze, Italy. E-mail: firstname.lastname@example.org; email@example.com. ORCID ID Giuseppe
d) Dipartimento di Chimica, Biologia e Biotecnologie, Università degli studi di Perugia, Via del Giochetto, 06126
Perugia, Italy. E-mail: firstname.lastname@example.org.
e) Via di Villa Ricotti 26, 00161, Roma. E-mail: email@example.com.
f) Vocabolo Scanzano 501, 06056, Massa Martana – Perugia, Italy. E-mail: firstname.lastname@example.org.
g) Free-lance wildlife technician, Str. di Pilli, 1 - 53035 Monteriggioni - Siena, Italy. E-mail:
h) Free-lance wildlife technician, Loc. Defizio, 58036, Roccastrada, Grosseto, Italy. E-mail:
i) Regione Umbria, Servizio Programmazione Faunistica Venatoria, Osservatorio Faunistico Regionale, Corso
Vannucci 96, 06121 Perugia, Italy. E-mail: email@example.com
j) Studio Naturalistico Hyla S.r.l., Via Baroncino, 11, 06069 Tuoro sul Trasimeno (Perugia), Italy. E-mail:
*Corresponding Author, Emiliano Mori: firstname.lastname@example.org
Mori E., Viviano A., Brustenga L., Olivetti F., Peppucci L., Pucci C., Senserini D., Sergiacomi U., Spilinga C.,
Roversi P.F., Mazza G.- Distribution and genetic analysis of wild-living Eurasian beavers in Central Italy.
The presence of the Eurasian beaver Castor fiber L. has been recently confirmed with two separated populations
in Tuscany (Central Italy) and probably represents the result of an unofficial release. In late spring and summer
2021, seven reliable records of Eurasian beaver have been collected in Umbria and other neighbouring regions,
implying that the distribution of this large rodent is even wider than previously reported. In this short work, we
updated the distribution of this protected species in Central Italy, by collecting and mapping all the confirmed
occurrences. Beavers were proved to be present throughout the Tiber (Tevere) river basin in both provinces of
Umbria, and another individual has been road-killed in the Marche region, near the border with Tuscany. Other
single signs of presence occurred in Emilia Romagna and Latium. The only hair sample we were able to collect
confirmed it as the Eurasian beaver species. No reliable evidence is available on the number of free-ranging beavers
in Central Italy, and systematic monitoring is needed. Before any management and conservation action, further data
are required concerning distribution range, potential origin, social perception, and the effects on the ecosystems.
KEY WORDS: Castor fiber; distribution assessment; riverine ecosystems; Rodentia; species release.
Within the dynamics of ecological systems, assessing
species distribution, the modification of habitat types,
and the interactions with human activities are pivotal for
conservation and management purposes.
The variety of life at every hierarchical level and
spatial scale of biological organisation from genes to
ecosystems is widely known as “biodiversity”. The
Anthropocene is the historical period we are currently
living in, characterised by a phase of severe decline of
biodiversity worldwide, mostly due to habitat loss and
fragmentation, and biological invasions (DIRZO et al.,
2014; BELLARD et al., 2016). As a response, the
European Union Biodiversity Strategy, adopted by the
European Commission, has set out several targets and
actions to limit the loss of biodiversity by 2030: all
European countries should maintain and restore
biodiversity (MAMMOLA et al., 2020). In this context,
reintroductions, i.e. translocations of individuals of a
species in an area where the species is extinct but it was
present in historical times, could be a useful tool to face
the biodiversity crisis. The International Union for the
Conservation of Nature (IUCN) has designed detailed
guidelines applying to the whole spectrum of
conservation translocations, including legal permissions,
selection of stocks, and stock health status (IUCN/SCC,
2013). Furthermore, all extinction causes should have
been removed from the release-area, to increase the
success of the translocation (IUCN/SSC 2013;
DROUILLY & O’RIAIN, 2021). Official and unofficial
releases have helped the range expansion of several
threatened or locally extinct species in many areas of the
world (CROMSIGT et al., 2018; ZIELKE et al., 2019; BODE,
210 MORI ET AL. REDIA, Vol. 104, 2021
2021). Amongst the most reintroduced species in Europe,
the Eurasian beaver Castor fiber Linnaeus, 1758
underwent a severe population decline during Medieval
times (e.g. SALARI et al., 2000) because of hunting for fur
and meat, as well as because of habitat loss, with only
1200 individuals remained between France and Mongolia
at the beginning of 1900 (HALLEY & ROSELL 2002;
HALLEY et al., 2021). Since then, reintroduction events
followed by natural spread triggered the recovery of the
species in most of its original range, up to a current global
population estimate of 1.5 million individuals (WRÓBEL,
2020; BOUROŞ et al., 2021; HALLEY et al., 2021).
Together with legal reintroductions conducted
following IUCN guidelines, several unauthorised
releases of Eurasian beavers of unknown origins have
occurred in Europe, e.g. in Belgium and Spain (DEWAS
et al., 2012; GIRLING et al., 2019). Most of these releases
have been successful and, after reintroductions, Eurasian
beavers have rapidly expanded their ranges to new areas
where they were historically extirpated (DEWAS et al.,
2012). The Eurasian beaver survived in northern Italy to
the early 1800s (HAINARD, 1949; PSENNER, 1971). After
reintroduction programs conducted in Austria and
Switzerland from the 1970s - 1990s, at least two
individuals of this rodent crossed the Italian borders in
2018 while spreading (PONTARINI et al., 2019; PUCCI et
al., 2021). In both cases, the involved streams flowed out
of Italy into the Danube watershed, on the northern side
of the Alps.
Later on, in early 2021, some wildlife technicians and
members of the provincial police noted some
unequivocal signs of beaver presence in two areas of
Tuscany (Central Italy), one in the municipalities of
Civitella-Paganico, Murlo, Montalcino, Buonconvento,
and Monticiano (provinces of Grosseto and Siena), and
the other near Sansepolcro (province of Arezzo). In the
first area, over 10 km of rivers (Ombrone river basin with
tributaries, i.e. Merse, and possibly Farma) were
characterised by the beaver presence; in the second one
(Tevere, i.e. Tiber river basin with its tributary, Cerfone),
signs of presence covered at least 6-7 km of the river
(PUCCI et al., 2021). These areas are separated by one
another by over 110 km in a straight line. After the
publication of PUCCI et al. (2021) and the media echo,
further data on beaver presence have been collected in
Umbria and other regions. In this note, we aimed to
determine beaver distribution in Central Italy through on-
site investigations in areas characterised by signs of the
presence of this species (i.e. gnawed trunks).
Monitoring of beaver occurrences has been carried out
in April-July 2021, and the authors verified all the
received reports through direct on-site visits to assess the
presence of signs of the presence of the Eurasian beaver.
We also checked on social networks and websites for
wildlife occurrence data collection (i.e. iNaturalist:
www.inaturalist.org, Ornitho: www.ornitho.it), but no
further record was found on these platforms.
Beaver guard hairs were found on a bramble bush
between Sansepolcro and Città di Castello and stored at -
20°C in labeled plastic bags (HERR & SCHLEY, 2009). A
further beaver hair sample was collected at the Poppi Zoo
Park (Arezzo) from the only individual still present (i.e.,
an adult male born in 2011). The Poppi Zoo Park is the
only one officially recognized structure in Italy hosting
Eurasian beavers together with the “Oasi di
Sant’Alessio” in Pavia (PUCCI et al., 2021). Once in the
lab, hair roots were plucked from the fur, and DNA was
extracted following a phenol-chloroform purification
protocol: hair roots have been grinded using liquid
nitrogen, a mortar, and a pestle (NERVA et al., 2021).
DNA pellets were re-suspended in 150 μL of elution
solution and cleaned with the DNA Clean and
Concentration kit (Zymoresearch, CA, USA). A 450 bp
fragment of the mtDNA cytochrome-b gene was
amplified by PCR using beaver-specific primers (KUEHN
et al., 2000), through a 2720 Thermal Cycler (Applied
Biosystems). Sequencing was conducted using the
dideoxy-chain termination method (SANGER et al., 1977)
with the forward primer. Genetic sequences were cleaned
at the 5’ and 3’ ends by looking at the chromatogram. To
conclude, the obtained sequence was compared to the
ones deposited in the NCBI database using BLASTn to
confirm the correct origin of the obtained sequences
(ALTSCHUL et al., 1990). Alignments of cytochrome-b
sequences were performed through ClustalX and Mega7
softwares (HIGGINS & SHARP, 1988; KUMAR et al.,
We found a total of 12 occurrence points (i.e.
coordinates confirmed by photos of gnawed trunks
and/or photos of individual beavers) from Umbria.
Amongst those, seven belonged to the Eurasian beaver,
two to the coypu Myocastor coypus (Molina, 1782), and
3 were unidentifiable signs of presence. Direct visits to
all of these sites confirmed the presence of the Eurasian
beaver in the Val Tiberina (i.e. the Tevere river valley) in
Umbria (Fig. I), both in Perugia (municipalities of Città
di Castello and Deruta) and Terni (municipalities of
Guardea and Alviano: see Figure 1a). A further road-
killed individual was observed near Mercatello sul
Metauro (province of Pesaro Urbino, Marche region) in
late June 2021, and a sign of presence (i.e. a gnawed
trunk) was recorded in Porretta (Bologna, Emilia
Romagna region) in August 2021, with no other evidence
of presence (Fig. I, 2). Occurrences of Eurasian beaver
from Latium were only confirmed at the immediate
borders with Umbria, thus requiring further research
DISTRIBUTION AND GENETIC ANALYSIS OF WILD-LIVING EURASIAN BEAVERS IN CENTRAL ITALY 211
Fig. I - 1) Map of the confirmed occurrences of the Eurasian beaver in Umbria. 2) Summary of beaver distribution in
Individuals and signs of presence were observed in the
abovementioned sites (Fig. II). Areas of beaver presence
are covered with deciduous riparian woodlands
composed by Salix alba Linnaeus, 1753, Populus nigra
Linnaeus, 1753 and Populus alba Linnaeus, 1753.
Beavers have been present in Umbria for at least one
year, given the age of the vegetation regrowth over
gnawed trunks. We detected 55 gnawed trunks (about
60% on Salix alba, 40% on Populus spp.), with an
average diameter of 15±5 cm.
The genetic sequence of cytochrome-b from
Sansepolcro (Val Tiberina, province of Arezzo) clustered
within the variability of the Western clade of C. fiber,
whereas the sample of the only individual currently
present in the Poppi Zoo Park clustered within the
Eastern clade of the same species (Fig. III).
The Eurasian beaver may attract human and media
attention more than other smaller, cryptic mammal
species which have recently been (re)discovered in Italy
(DONDINI et al., 2014; MORI et al., 2020). The appeal of
this large rodent to humans may be due to its size,
morphology, and famous cartoons that may have
stimulated empathy towards this species (e.g. “Don
Chuck Monogatari” and “Papa Beaver’s Story”).
212 MORI ET AL. REDIA, Vol. 104, 2021
Fig. II - Signs of presence and individual of Eurasian beaver in Val Tiberina, Umbria (photos A. Viviano, S. Galletti, F.
Olivetti, and L. Peppucci).
Fig. III - Neighbour-Joining tree on mtDNA sequence of the cytochrome-b of beavers including all available sequences
from GenBank and our samples. Italian sequences are shown in bold; a sequence of Hystrix cristata is also included as
an outgroup (grey).
In this brief report, we reported the presence of the
Eurasian beaver throughout the Umbria region, with
evidence of occurrences in four municipalities out of both
Perugia and Terni provinces. Further records from North-
Eastern Tuscany (Porrena, Arno river), North-Eastern
Latium (Tevere river), and Emilia-Romagna were only
based on single signs of presence (i.e. one gnawed trunk
per area) and require further researches, possibly during
the cold months to confirm the beaver presence
(ELMEROS et al., 2003). All new records confirmed in
this work come from the Tevere river basin, being one
record near the confluence with the Paglia river and one
other in Cerfone river. The origin of these populations of
Eurasian beaver in Central Italy, where the original
population became extinct in Medieval times (SALARI et
al., 2019), is still unknown. The simultaneous appearance
of beavers in different areas of Central Italy after
centuries being unnoticed, and molecular data bring us to
discard the hypothesis of a local “native” population. In
fact, a genuinely native Italian population would not be expected to show a so close similarity to the Western
clade, which derived from an Ice Age refuge in Iberia
DISTRIBUTION AND GENETIC ANALYSIS OF WILD-LIVING EURASIAN BEAVERS IN CENTRAL ITALY 213
(MARR et al., 2018). Most likely, given the known Ice
Age vegetation of Italy, beavers survived the last Ice Age
in an Italian refuge, possibly with a genotype similar to
the extinct Danube clade (MARR et al., 2018). On the
other hand, it is not possible to establish whether they
represent an unofficial release, an escape from confined
areas, or a natural population. However, the individual
still present in the only zoo park hosting beavers in
Central Italy belonged to a different clade with respect to
those sampled in the wild in Central Italy (see MARR et
al., 2018). Moreover, it is impossible to report the actual
number of individuals occurring in these areas. However,
we can confidently believe that populations of beavers in
Central Italy will increase (HALLEY & ROSELL, 2002);
the rivers they are on are relatively small compared to
those in Central Europe, so there may be little or no ‘lag’
time before the period of exponential population growth
begins (HALLEY & ROSELL, 2002).
The IUCN strongly discourages introductions or
reintroductions conducted without feasibility studies and
no molecular analysis of released stocks, because of the
damage they may cause to native ecosystems (KLEIMAN,
1989; HALLEY et al., 2009). The activity of beavers could
locally alter the riparian vegetation structure, in turn
influencing other components of the ecosystems,
including the diversity and abundance of invertebrates,
amphibians, and wading birds (NOLET et al., 1994;
ROSELL et al., 2005; BASHINSKIY, 2020). For this reason,
beaver alteration of heterogeneity and connectivity of
habitats needs further research in our study area
(BASHINSKIY, 2020). Where released, if impacts are
evident and affect the native biodiversity directly or
through the interaction with other native/alien species
(CAMPBELL-PALMER et al., 2016), management
strategies should also consider drastic actions including
individual removal (cf. GARGIONI et al., 2021).
Conversely, where reintroduced, Eurasian beavers are
reported to improve the hydrogeological status of
European rivers, to increase local species richness, and to
mitigate environmental pollution (ROSELL et al., 2005;
KEMP et al., 2012; PUTTOCK et al., 2017; CAMPBELL-
PALMER et al., 2021). If on one side the Eurasian beaver
is protected by the Habitats Directive (1992/43/EC,
Annex IV) and several national laws, human-wildlife
conflict may arise when releases are not authorised and,
sometimes, even where these operations are authorised,
at least in public perception (HALLEY & ROSELL, 2002).
Although crop damages by Eurasian beavers are
negligible in Europe (MIKULKA et al., 2020), particularly
when compared to those exerted by the wild boar Sus
scrofa Linnaeus, 1758 and the red deer Cervus elaphus
Linnaeus, 1758, illegal removal of animals perceived as
causing a nuisance may occur. However, from the legal
point of view, species protected according to the Habitats
Directive (92/43/EEC) should be treated as alien species
if released by humans where they are not naturally
present (for example the common chameleon Chamaeleo
chamaeleon Linnaeus, 1758 in Southern Italy: BASSO et
al., 2019). Natural range expansion to new areas, and any
official reintroductions, require completion of the
standard forms required for each six-year reporting
period (GENOVESI et al., 2014; STOCH & GENOVESI,
2016). In the case of Eurasian beaver in Central Italy,
conservation and restoration of riparian vegetation and
physical protection systems for trees may be used to
prevent damage (HALLEY & ROSELL, 2002).
Unauthorised (re)introductions are not to be condoned
(IUCN/SCC, 2013) and should be reprobated. In most
cases, however, introduced populations of protected
mammal species in Italy, including those listed in the
Annexes of the Habitats Directive as well as the Eurasian
beaver, have been left unmanaged (e.g. mouflons Ovis
aries Linnaeus, 1758 in continental Italy: LOY et al.,
2019; Alpine marmots Marmota marmota (Linnaeus,
1758) in the Apennines, both of them originally not
native of the study area: LOY et al., 2019; Eurasian otters
Lutra lutra Linnaeus, 1758 where originally native, i.e.
in Valsavarenche and, introgressed with the Asian
subspecies L. l. barang F.G. Cuvier, 1823, in the Ticino
Valley: PRIGIONI et al., 2009; FERRARI et al., 2017; alien
red deer in Monte Penna - Castell’Azzara, Grosseto and
in several other areas in Italy; alien crested porcupines
Hystrix cristata Linnaeus, 1758 in the Varese province,
Western Liguria and Sardinia regions: MORI et al., 2013).
Removal strategies for unofficially released Eurasian
beavers have, where attempted, been both ineffective and
expensive (e.g. experience from Spain: HALLEY et al.,
2020), and should be considered only in case of
undeniable proofs of human-mediated releases or, by
way of derogation to the Habitats Directive, in case of
severe and proven crop damage. As to Spain, 15 years
after the introduction, the Eurasian beaver is considered
as naturalised and, following the decision of the
European Commission, it cannot be removed anymore
and it requires monitoring following the Habitats
Directive. However, no complaint linked to this
increasing, unofficially-released population (occurring
throughout Ebro River up to Zaragoza) is recorded yet,
with only minor damages to trees in river groves
(Eugenio Fernandez, personal communication 2021). For
Central Italy, it would be important to currently
concentrate on how to manage the species, considering
also human attitudes. Different countries in Europe with
similar cultural landscapes have very different public
attitudes towards beavers, i.e. from very negative to very
positive, with little relationship to local beaver ecology
(CURRY-LINDAHL, 1967; SIEMER et al., 2013; AUSTER et
al., 2021). Imposing strong management actions even if
beavers have been unofficially released could thus trigger
a chronic, expensive, and emotionally tiring problem.
We would like to thank Andrea Boscherini, who first
discovered some possible signs of presence along the
Tiber river, Stefano Galletti for the photo of the beaver.
Thanks are also due to national and international
colleagues Duncan Halley, Davide Sogliani, Elena
Tricarico, James Wallace, Roisin Campbell-Palmer and
Eugenio Fernandez, who kindly provided us with useful
recommendations for field work, and on this MS. We
thank the Mattoni family (Poppi Zoo Park) and Giovanni
Mazza for sampling the individual at the Poppi Zoo Park,
214 MORI ET AL. REDIA, Vol. 104, 2021
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