ArticlePDF Available

Abstract and Figures

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.
Content may be subject to copyright.
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: emiliano.mori@cnr.it. 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:
a.viviano@studenti.unipi.it. 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: piofederico.roversi@crea.gov.it; giuseppe.mazza@crea.gov.it. ORCID ID Giuseppe
Mazza: 0000-0002-6644-1429.
d) Dipartimento di Chimica, Biologia e Biotecnologie, Università degli studi di Perugia, Via del Giochetto, 06126
Perugia, Italy. E-mail: leonardobrustenga@libero.it.
e) Via di Villa Ricotti 26, 00161, Roma. E-mail: cpitgst@gmail.com.
f) Vocabolo Scanzano 501, 06056, Massa Martana Perugia, Italy. E-mail: lucapep04@gmail.com.
g) Free-lance wildlife technician, Str. di Pilli, 1 - 53035 Monteriggioni - Siena, Italy. E-mail:
chiara.pucci80@gmail.com; chiara.pucci@epap.conafpec.it
h) Free-lance wildlife technician, Loc. Defizio, 58036, Roccastrada, Grosseto, Italy. E-mail:
davidesenserini@gmail.com
i) Regione Umbria, Servizio Programmazione Faunistica Venatoria, Osservatorio Faunistico Regionale, Corso
Vannucci 96, 06121 Perugia, Italy. E-mail: usergiacomi@regione.umbria.it
j) Studio Naturalistico Hyla S.r.l., Via Baroncino, 11, 06069 Tuoro sul Trasimeno (Perugia), Italy. E-mail:
info@studionaturalisticohyla.it
*Corresponding Author, Emiliano Mori: emiliano.mori@cnr.it
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.
INTRODUCTION
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).
METHODS
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.,
2016).
RESULTS
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
downstream.
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
Central Italy.
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).
DISCUSSION
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.
ACKNOWLEDGEMENTS
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
and Giulio Salamon for samples from the “Oasi di
Sant’Alessio”.
REFERENCES
ALTSCHUL S.F., GISH W., MILLER W., MYERS E.W.,
LIPMAN D.J., 1990 - Basic local alignment search
tool. - Journal of Molecular Biology, 215: 403-410.
AUSTER R.E., BARR S.W., BRAZIER R.E., 2021 -
Improving engagement in managing reintroduction
conflicts: Learning from beaver reintroduction. -
Journal of Environmental Planning and
Management, 64: 1713-1734.
BASHINSKIY I.V., 2020 - Beavers in lakes: a review of
their ecosystem impact. - Aquatic Ecology, 54: 1097-
1120.
BASSO R., VANNUCCINI M.L., NERVA L., MAZZA G.,
SENO M., MORI E., 2019 - Multiple origins of the
common chameleon in southern Italy. - Herpetozoa,
32: 11-19.
BELLARD C., CASSEY P., BLACKBURN T.M., 2016 - Alien
species as a driver of recent extinctions. - Biology
Letters, 12: 20150623.
BODE M., 2021 - Covert rewilding: Modelling the
detection of an unofficial translocation of Tasmanian
devils to the Australian mainland. Conservation
Letters, 14: e12787.
BOUROŞ G., PALADI V., CASSIR P., 2021 - First report of
the Eurasian beaver (Castor fiber Linnaeus 1758) in
Republic of Moldova. - North-Western Journal of
Zoology, 2021: e211701.
CAMPBELL-PALMER R., JONES S., PARKER H., ROSELL F.,
GOW D., SCHWAB G., DICKINSON H., 2016.- The
Eurasian beaver handbook: ecology and
management of Castor fiber. - Pelagic Publishing Ltd,
London, UK.
CAMPBELL‐PALMER R., PUTTOCK A., WILSON K.A.,
LEOW‐DYKE A., GRAHAM H.A., GAYWOOD M.J.,
BRAZIER R.E., 2021 - Using field sign surveys to
estimate spatial distribution and territory dynamics
following reintroduction of the Eurasian beaver to
British river catchments. - River Research
Application, 37: 343-357.
CROMSIGT J.P., KEMP Y.J., RODRIGUEZ E., KIVIT H.,
2018 - Rewilding Europe's large grazer community:
how functionally diverse are the diets of European
bison, cattle, and horses?. - Restoration Ecology, 26:
891-899.
CURRY-LINDAHL K., 1967 - The beaver, Castor fiber
Linnaeus, 1758 in Swedenextermination and
reappearance. - Acta Theriologica, 12: 1-15.
DEWAS M., HERR J., SCHLEY L., ANGST C., MANET B.,
LANDRY P., CATUSSE M., 2012 - Recovery and status
of native and introduced beavers Castor fiber and
Castor canadensis in France and neighbouring
countries. - Mammal Review, 42: 144-165.
DIRZO R., YOUNG H.S., GALETTI M., CEBALLOS G.,
ISAAC N. J., COLLEN B., 2014 - Defaunation in the
Anthropocene. - Science, 345: 401-406.
DONDINI G., TOMASSINI A., INGUSCIO S., ROSSI E., 2014
- Rediscovery of Mehely's horseshoe bat
(Rhinolophus mehelyi) in peninsular Italy. - Hystrix,
the Italian Journal of Mammalogy, 25: 59-60.
DROUILLY M., O’RIAIN M.J., 2021 - Rewilding the
world’s large carnivores without neglecting the
human dimension. - Biodiversity and
Conservation, 30: 917-923.
ELMEROS M., MADSEN A.B., BERTHELSEN J.P., 2003 -
Monitoring of reintroduced beavers (Castor fiber) in
Denmark. - Lutra, 46: 153-162.
FERRARI C., D'ALFONSO M., MORIS V., BATTISTI A.,
GIULIANO D., 2017 - Diet in a free ranging individual
of Lutra lutra in Valsavarenche. - Journal of
Mountain Ecology, 10: 7-12.
GARGIONI C., MONACO A., FICETOLA G.F., LAZZERI L.,
MORI E., 2021 - From the Andes to the Apennines:
rise and fall of a free-ranging population of feral
llamas. - Animals, 11: 857.
GENOVESI P., ANGELINI P., DUPRÈ E., ERCOLE S.,
GIACANELLI V., RONCHI F., STOCH F., 2014 - Specie
ed habitat di interesse comunitario in Italia:
distribuzione, stato di conservazione e trend. -
ISPRA, Serie Rapporti, 194/2014.
GIRLING S.J., NAYLOR A., FRASER M., CAMPBELL
PALMER R., 2019 - Reintroducing beavers Castor
fiber to Britain: a disease risk analysis. - Mammal
Review, 49: 300-323.
HAINARD R., 1949 - Le Mammiferes sauvages d'Europe.
II. Pinnipedes - Rongeurs - Ongules - Cetaces. -
Delachaux and Niestlè Editions, Paris, France.
HALLEY D.J., ROSELL F., 2002 - The beavers reconquest
of Eurasia: status, population development, and
management of a conservation success. - Mammal
Review, 32: 153-178.
HALLEY D.J., JONES A.L., CHESWORTH S., HALL C., GOW
D., JONES-PARRY R., WALSH J., 2009 - The
reintroduction of the Eurasian beaver Castor fiber to
Wales: an ecological feasibility study. - NINA
Report, 457: 1-66.
HALLEY D.J., SAVELJEV A.P., ROSELL F., 2021.
Population and distribution of beavers Castor fiber
and Castor canadensis in Eurasia. - Mammal Review,
51: 1-24.
HERR J., SCHLEY L., 2009 - Barbed wire hair traps as a
tool for remotely collecting hair samples from
beavers (Castor sp.). - Lutra, 52: 123-127.
HIGGINS D.G., SHARP P.M., 1988 - CLUSTAL: a package
for performing multiple sequence alignment on a
microcomputer. - Gene, 73: 237244.
IUCN/SSC, 2013 - Guidelines for Reintroductions and
Other Conservation Translocations. Version 1.0.
Gland, Switzerland: IUCN Species Survival
Commission, viiii + 57 pp
KELLER A., 1983 - Etude comparative des différentes
structures pileuses du Castor canadensis (Kuhl) et du
Castor fiber Linné (Mammalia, Castoridae). - Revue
Suisse Zoologie, 90: 183-189.
KEMP P.S., WORTHINGTON T.A., LANGFORD T.E., TREE
A.R., GAYWOOD M.J., 2012 - Qualitative and
quantitative effects of reintroduced beavers on stream
fish. - Fish and Fisheries, 13 : 158-181.
KLEIMAN D.G., 1989 - Reintroduction of captive
mammals for conservation. - BioScience, 39: 152-
DISTRIBUTION AND GENETIC ANALYSIS OF WILD-LIVING EURASIAN BEAVERS IN CENTRAL ITALY 215
161.
KUEHN R., SCHWAB G., SCHROEDER W., ROTTMANN O.,
2000 - Differentiation of Castor fiber and Castor
canadensis by noninvasive molecular methods. - Zoo
Biology, 19: 511-515.
KUMAR S., STECHER G., TAMURA K., 2016 - MEGA7:
Molecular Evolutionary Genetics Analysis version
7.0 for bigger datasets. - Molecular Biology
Evolution, 33: 18701874.
LOY A., ALOISE G., ANCILLOTTO L., ANGELICI F.M.,
BERTOLINO S., CAPIZZI D., CASTIGLIA R.,
COLANGELO P., CONTOLI L., COZZI B., FONTANETO
D., LAPINI L., MAIO N., MONACO A., MORI E., NAPPI
A., PODESTÀ M., RUSSO D., SARÀ M., SCANDURA M.,
AMORI G., 2019 - Mammals of Italy: an annotated
checklist. - Hystrix, the Italian Journal of
Mammalogy, 30: 87-106.
MAMMOLA S., RICCARDI N., PRIÉ V., CORREIA R.,
CARDOSO P., LOPES-LIMA M., SOUSA R., 2020 -
Towards a taxonomically unbiased European Union
biodiversity strategy for 2030. - Proceedings of the
Royal Society B, 287: 20202166.
MARR M.M., BRACE S., SCHREVE D.C., BARNES I., 2018
- Identifying source populations for the
reintroduction of the Eurasian beaver, Castor fiber L.
1758, into Britain: evidence from ancient DNA. -
Scientific Reports, 8: 1-8.
MIKULKA O., HOMOLKA M., DRIMAJ J., KAMLER J., 2020
- European beaver (Castor fiber) in open agricultural
landscapes: crop grazing and the potential for
economic damage. - European Journal of Wildlife
Research, 66: 1-10.
MORI E., SFORZI A., DI FEBBRARO M., 2013 - From the
Apennines to the Alps: recent range expansion of the
crested porcupine Hystrix cristata L., 1758
(Mammalia: Rodentia: Hystricidae) in Italy. - Italian
Journal of Zoology, 80: 469-480.
MORI E., BRAMBILLA M., RAMAZZOTTI F., ANCILLOTTO
L., MAZZA G., RUSSO D., AMORI G., GALIMBERTI A.,
2020 - In or out of the checklist? DNA barcoding and
distribution modelling unveil a new species of
Crocidura shrew for Italy. - Diversity, 12: 380.
NERVA L., IANNUCCI A., MENCHETTI M., ANDREONI A.,
CHITARRA W., MARTINI M., MUELLER N., PETERS
T.M.J., PESENTI E., VERBEYLEN G., ZOZZOLI R.,
MORI E., 2021 - Where do Chip and Dale come from?
Origins of invasive populations of the Siberian
chipmunk in Europe. - Mammal Research,
https://doi.org/10.1007/s13364-021-00569-4
NOLET B.A., HOEKSTRA A., OTTENHEIM M.M., 1994 -
Selective foraging on woody species by the beaver
Castor fiber, and its impact on a riparian willow
forest. - Biological Conservation, 70:117-128.
PONTARINI R., LAPINI L., MOLINARI P., 2019 - A beaver
from North-Eastern Italy (Castor fiber: Castoridae,
Rodentia). - Gortania, 40: 115-118.
PRIGIONI C., SMIROLDO G., REMONTI L., BALESTRIERI
A., 2009 Distribution and diet of reintroduced
otters (Lutra lutra) on the River Ticino (NW Italy). -
Hystrix, the Italian Journal of Mammalogy, 20: 45-
53.
PUCCI C., SENSERINI D., MAZZA G., MORI E., 2021 -
Reappearance of the Eurasian beaver Castor fiber L.
in Tuscany (Central Italy): the success of
unauthorised releases? - Hystrix, the Italian Journal
of Mammalogy, https://doi.org/10.4404/hystrix-
00445-2021
PUTTOCK A., GRAHAM H.A., CUNLIFFE A.M., ELLIOTT
M., BRAZIER R.E., 2017 - Eurasian beaver activity
water storage, attenuates flow and mitigates diffuse
pollution from intensively-managed grasslands. -
Science of the Total Environment, 576: 430-433.
PSENNER H., 1971 - Tiere der Alpen. Tyrolia-Verlag,
Innsbruck, Austria.
ROSELL F., BOZSER O., COLLEN P., PARKER H., 2005 -
Ecological impact of beavers Castor fiber and Castor
canadensis and their ability to modify ecosystems. -
Mammal Review, 35: 248-276.
SALARI L., MASSETI M., SILVESTRI L., 2020 - Late
Pleistocene and Holocene distribution history of the
Eurasian beaver in Italy. - Mammalia, 84: 259-277.
SANGER F., NICKLEN S., COULSON A.R., 1977 - DNA
sequencing with chain-terminating inhibitors. -
Proceedings of the National Academy of Sciences,
74: 54635467.
SIEMER W.F., JONKER S.A., DECKER D.J., ORGAN J.F.,
2013 - Toward an understanding of beaver
management as human and beaver densities increase.
- Human-Wildlife Interactions, 7: 114-131.
STOCH F., GENOVESI P., 2016 - Manuali per il
monitoraggio di specie e habitat di interesse
comunitario (Direttiva 92/43/CEE) in Italia: specie
animali. - ISPRA, Serie Manuali e Linee Guida,
141/2016, Roma, Italy.
WRÓBEL M., 2020 - Population of Eurasian beaver
(Castor fiber) in Europe. - Global Ecology and
Conservation, 23: e01046.
ZIELKE L., WRAGE-MÖNNIG N., MÜLLER J., NEUMANN
C., 2019 - Implications of spatial habitat diversity on
diet selection of European Bison and Przewalski’s
Horses in a rewilding area. - Diversity, 11: 63.
... In November 2020, another individual was camera-trapped in Val Pusteria (province of Bolzano), where two individuals might be currently present. Since 2019, the Eurasian beaver has also been present in Central Italy, with at least two populations in two river basins (Merse-Ombrone and Tevere rivers), where reproduction events have been confirmed for the first time since 2020 [22][23][24], proving their local establishment. ...
... Our study was conducted in river basins, namely Ombrone-Merse (municipalities of Sovicille, Murlo, Monticiano, Buonconvento, Montalcino, and Civitella-Paganico) and Tevere (municipalities of Anghiari and Sansepolcro) in the Tuscany region (Central Italy), where the occurrence of at least 6-8 Eurasian beavers has been confirmed since at least 2019-2020 [22,23]. Deciduous riparian woodland is the main habitat type (87%), mostly represented by Salix eleagnos Scop., Populus nigra L., Populus alba L., Alnus glutinosa L. (Gaertn.), ...
... In this work, we determined the patterns of activity rhythms of the Eurasian beaver in a Mediterranean ecosystem of recent reconquest [22][23][24]. The beaver was confirmed to be mostly a crepuscular-nocturnal species, as well as being present in the core of its European extent of occurrence [30][31][32][33]. ...
Article
Full-text available
Analyses of temporal partitioning and overlaps in activity rhythms are pivotal to shed light on interspecific coexistence between similar species or prey and predators. In this work, we assessednthe overlap of activity rhythms between the Eurasian beaver Castor fiber and its potential competitors and predators through camera trapping in an area in Central Italy. Interspecific overlaps of temporal activity patterns were estimated for the beavers, potential predators (the red fox Vulpes vulpes and the grey wolf Canis lupus), and a potential competitor, the coypu Myocastor coypus. The beavers showed a mostly crepuscular behaviour. Although high temporal overlap was observed between the Eurasian beavers and the red foxes and grey wolves, the activity of the beavers did not overlap with that of the predators. Accordingly, the beavers were more active on the darkest nights, i.e., avoiding bright moonlight.
Article
Full-text available
Through the Habitats Directive (92/43/EEC) and the financial investments of the LIFE projects, Europe has become an experimental arena for biological conservation. With an estimated annual budget of €20 billion, the EU Biodiversity Strategy for 2030 has set an ambitious goal of classifying 30% of its land and sea territory as Protected Areas and ensuring no deterioration in conservation trends and the status of protected species. We analysed LIFE projects focused on animals from 1992 to 2018 and found that investment in vertebrates was six times higher than that for invertebrates (€970 versus €150 million), with birds and mammals alone accounting for 72% of species and 75% of the total budget. In relative terms, investment per species towards vertebrates has been 468 times higher than that for invertebrates. Using a trait-based approach, we show that conservation effort is primarily explained by species' popularity rather than extinction risk or body size. Therefore, we propose a roadmap to achieve unbiased conservation targets for 2030 and beyond.
Article
Full-text available
The European beaver (Castor fiber) has extended its range into most Central European countries over the past 30 years, resulting in increased forestry damage and water management issues. As the number of beavers increases populations become established in new types of habitat. In the Czech Republic, for example, established beaver populations are now found on rivers flowing through agricultural landscapes, where living conditions differ significantly to those found in forest landscapes. To date, there have been no studies on the impact of beavers on agricultural production. The aim of this study was (1) to describe how beavers graze field crops, and (2) to estimate potential damage to agricultural production. This 2-year study examined five beaver territories in agricultural landscapes where crops were separated from watercourses by a narrow strip of bank vegetation. Beavers fed on all crop types grown in their territories throughout the growing period (May–October), peaking from mid-June to mid-July. The beavers clearly preferred oilseed rape, which (along with wheat and barley) represented the largest part of the grazed area. Rape was usually grazed at during vegetation growth and flowering, while cereals were usually grazed from the milk ripe kernel stage to harvest. Damage to agricultural production was up to €20–30 per ind./year. While beaver population density in the agricultural landscape remains low, damage to agricultural production is relatively insignificant; however, field crops clearly represent an important part of the beavers’ diet in such areas, helping them survive in such open landscapes.
Article
Full-text available
The genus Crocidura (Eulipotyphla, Soricidae) is the most speciose genus amongst mammals, i.e., it includes the highest number of species. Different species are distinguished by skull morphology, which often prevents the identification of individuals in the field and limits research on these species' ecology and biology. We combined species distribution models and molecular analyses to assess the distribution of cryptic Crocidura shrews in Italy, confirming the occurrence of the greater white-toothed shrew Crocidura russula in the northwest of the country. The molecular identification ascertained the species' presence in two distinct Italian regions. Accordingly, species distribution modelling highlighted the occurrence of areas suitable for C. russula in the westernmost part of northern Italy. Our results confirm the role of Italy as a mammal hotspot in the Mediterranean; additionally, they also show the need to include C. russula in Italian faunal checklists. To conclude, we highlight the usefulness of combining different approaches to explore the presence of cryptic species outside their known ranges. Since the similar, smaller C. suaveolens may be displaced by the larger C. russula through competitive exclusion, the latter might be the species actually present where C. suaveolens had been reported previously. A comprehensive and detailed survey is therefore required to assess the current distribution of these species.
Article
Full-text available
We present the latest version of the Molecular Evolutionary Genetics Analysis (MEGA) software, which contains many sophisticated methods and tools for phylogenomics and phylomedicine. In this major upgrade, MEGA has been optimized for use on 64-bit computing systems for analyzing bigger datasets. Researchers can now explore and analyze tens of thousands of sequences in MEGA. The new version also provides an advanced wizard for building timetrees and includes a new functionality to automatically predict gene duplication events in gene family trees. The 64-bit MEGA is made available in two interfaces: graphical and command line. The graphical user interface (GUI) is a native Microsoft Windows application that can also be used on Mac OSX. The command line MEGA is available as native applications for Windows, Linux, and Mac OSX. They are intended for use in high-throughput and scripted analysis. Both versions are available from www.megasoftware.net free of charge.
Article
Full-text available
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.
Article
Full-text available
The genus Castor first appeared in the Palaearctic region during the Late Miocene, while the current species, Castor fiber, is widely accepted to have emerged in the Early Pleistocene. In the Last Glacial Maximum (Late Pleistocene), the beaver disappeared from most of the Western Palaearctic, only surviving in a few relic areas including the south-eastern Alpine Chain as shown by new data. After the subsequent extended repopulation in the warmer phases of the Lateglacial and in the early Holocene, the species once again disappeared locally from several countries, including Italy, between the 17th and the 20th centuries. Direct or indirect persecution by humans seems to be the main cause of beaver extinction in Europe. In Low Medieval Italy, it is more likely that the disappearance of the beaver between the 16th and 17th centuries was due to habitat alteration and human population pressure. Numerous reclamations have been carried out since the late Middle Ages, mostly in the easternmost area of the Po Valley, the last beaver refuge in Italy. Eurasian beaver was common in the northern and widespread in the central part of Italy, but always absent in southern Italy, probably due to unfavourable hydrological conditions of watercourses in the latter.
Article
Full-text available
In Europe, the interest in introducing megaherbivores to achieve ambitious habitat restoration goals is increasing. In this study, we present the results of a one-year monitoring program in a rewilding project in Germany (Doeberitzer Heide), where European bison (Bison bonasus) and Przewalski´s horses (Equus ferus przewalskii) were introduced for ecological restoration purposes. Our objectives were to investigate diet and habitat preferences of Przewalski´s horses and European bison under free-choice conditions without fodder supplementation. In a random forest classification approach, we used multitemporal RapidEye time series imagery to map the diversity of available habitats within the study area. This spatially explicit habitat distribution from satellite imagery was combined with direct field observations of seasonal diet preferences of both species. In line with the availability of preferred forage plants, European bison and Przewalski´s horses both showed seasonal habitat preferences. Because of their different preferences for forage plants, they did not overlap in habitat use except for a short time in the colder season. European bison used open habitats and especially wet open habitats more than expected based on available habitats in the study area. Comparative foraging and feeding niches should be considered in the establishment of multispecies projects to maximize the outcome of restoration processes.
Article
Full-text available
Establishing true phylogenetic relationships between populations is a critical consideration when sourcing individuals for translocation. This presents huge difficulties with threatened and endangered species that have become extirpated from large areas of their former range. We utilise ancient DNA (aDNA) to reconstruct the phylogenetic relationships of a keystone species which has become extinct in Britain, the Eurasian beaver Castor fiber. We sequenced seventeen 492 bp partial tRNAPro and control region sequences from Late Pleistocene and Holocene age beavers and included these in network, demographic and genealogy analyses. The mode of postglacial population expansion from refugia was investigated by employing tests of neutrality and a pairwise mismatch distribution analysis. We found evidence of a pre-Late Glacial Maximum ancestor for the Western C. fiber clade which experienced a rapid demographic expansion during the terminal Pleistocene to early Holocene period. Ancient British beavers were found to originate from the Western phylogroup but showed no phylogenetic affinity to any one modern relict population over another. Instead, we find that they formed part of a large, continuous, pan-Western European clade that harbored little internal substructure. Our study highlights the utility of aDNA in reconstructing population histories of extirpated species which has real-world implications for conservation planning.
Article
Full-text available
Beavers are the archetypal keystone species, which can profoundly alter ecosystem structure and function through their ecosystem engineering activity, most notably the building of dams. This can have a major impact upon water resource management, flow regimes and water quality. Previous research has predominantly focused on the activities of North American beaver (Castor canadensis) located in very different environments, to the intensive lowland agricultural landscapes of the United Kingdom and elsewhere in Europe.
Article
Among invasive squirrels in Europe, the Siberian chipmunk Eutamias sibiricus, native to NorthEast Asia, shows the highest number of free-ranging populations in European countries, due to the intense pet trade it underwent between the 1960s and the 1980s. We describe 628-bp cytochrome b sequences from the Netherlands (N = 3), Belgium (N = 4) and Switzerland (N = 1), and through phylogenetic analysis show that they belong to the Korean subspecies. This confirms previous findings that the Korean subspecies (Eutamias sibiricus barberi) has been the most commonly introduced to Europe. Another subspecies (the Northern subspecies, Eutamias sibiricus sibiricus) has only been reported previously in an Italian free-living population. If the splitting of the Korean taxon as a proper species (Eutamias barberi) will be confirmed by analysis of nuclear markers, a revision of the European Regulation 1143/2014 should be conducted to explicitly ban the trade of newly identified taxa.