Entomol Gener 33 (1/2): 079–089 Stuttgart: 2011-05 ISSN: 0171-8177

EGT-Nr 1.367 VII

Monitoring of Lucanus cervus by means of Remains

of Predation (Coleoptera: Lucanidae)
Alessandro Campanaro, Ilaria Toni, Sönke Hardersen & Donato A. Grasso

Received: 2010-11-08 Accepted: 2010-12-02

Campanaro A [Dep Biol Biotecnol “Charles Darwin”, Sapienza Univ Roma, IT-00185 Roma], Toni I,
Hardersen S [Cent Naz Biodivers For ‘Bosco Fontana’, Corp For Stato, IT-46045, Marmirolo (MN)]
& Grasso D A [Dip Biol Evol Funz, Univ Parma, IT-43100 Parma]: Monitoring of Lucanus cervus
by means of Remains of Predation (Coleoptera: Lucanidae). – Entomol Gener 33 (1/2): 079–089;
Stuttgart 2011-05 ––– [Note]
In Italy, Lucanus cervus (Linnaeus 1758) generally lives in oak forests characterized by the
presence of old trees. Such habitats are critically rare and fragmented. L cervus is listed in Annex II of
the Habitats Directive and according to article 17, needs to be monitored by member states. However,
methods for monitoring populations of L cervus have so far only been proposed for single countries
and these are mostly not quantitative. In Italy survey data for L cervus are scarce and no national
monitoring protocol has been proposed. At Bosco Fontana, a Nature Reserve in Northern Italy, remains
of predation of L cervus can easily be found along forest roads and it seems that the main predator
is the hooded crow (Corvus cornix cornix). The preliminary study presented here, which is based on
the collection of these remains, had the following aims: quantify the number of remains which can be
sampled at Bosco Fontana, investigate the temporal pattern of the remains and estimate the phenol-
ogy of L cervus, analyze the spatial distribution of the remains found and analyze the results with a
view to monitoring L cervus by means of collecting remains of predation. The survey was carried
out by searching all remains of predation of L cervus, from May to August 2008. In this period the
road network of the reserve (23 km in total) was walked on a weekly basis. A total of 306 remains
were collected (95 % males), which confirmed that these searches of remains were an efficient way of
monitoring male L cervus. The remains were distributed non-homogeneously and most were found in
a restricted number of roads. The data collected were used to estimate that L cervus was on the wing
in the reserve from the first week of May to the first week of August. The implications of the results
for monitoring L cervus are discussed.
Key words: Lucanus cervus (Linnaeus 1758) – Habitats Directive – hooded crow – monitoring –
phenology –predation

Campanaro A [Dip. Biol Biotecnol “Charles Darwin”, Sapienza Univ Roma, IT-00185 Roma], Toni I,
Hardersen S [Cent Naz Biodivers For ‘Bosco Fontana’, Corp For Stato, IT-46045, Marmirolo (MN)]
& Grasso D A [Dip Biol Evol Funz, Univ Parma, IT-43100 Parma]: Monitoraggio di Lucanus cer-
vus attraverso resti di predazione (Coleoptera: Lucanidae). – Entomol Gener 33 (1/2): 079–089;
Stuttgart 2011-05 ––– [Nota]
Il cervo volante, Lucanus cervus, è una specie associata, in Italia, a foreste con esemplari vetusti
di querce, un habitat raro e frammentato. L cervus è una specie inserita nell’allegato II della Direttiva
Habitat, pertanto, ai sensi dell’art. 17 della direttiva stessa, necessita di monitoraggio da parte degli
Stati membri.

0171–8177/11/0033-0079 $ 2.75
© 2011 E. Schweizerbart’sche Verlagsbuchhandlung, D-70176 Stuttgart

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 080 – Alessandro Campanaro et al

Nonostante ciò, metodi di monitoraggio delle popolazioni sono stati proposti soltanto per sin-
goli paesi e questi raramente permettono di ottenere informazioni quantitative sulle popolazioni di L
cervus. In Italia i dati di popolazione su cervo volante sono scarsi e nessun protocollo di monitoraggio
a livello nazionale è stato sinora proposto. Nella Riserva Naturale di Bosco Fontana, situata in Italia
settentrionale, vengono abitualmente rinvenuti, lungo le strade forestali, resti di predazione di L cervus,
presumibilmente da parte di cornacchia grigia (Corvus cornix cornix). Da tale osservazione scaturisce
lo studio qui presentato che ha i seguenti obiettivi principali: quantificare il numero di resti che pos-
sono essere raccolti a Bosco Fontana, analizzare la distribuzione temporale dei resti per stimare la
fenologia di L cervus e analizzare i risultati del campionamento dei resti in un’ottica di monitoraggio
della specie. Il campionamento dei resti di predazione è stato effettuato nel 2008, da maggio ad agosto.
In questo periodo il sistema di strade forestali della Riserva (23 km in totale) è stato completamente
percorso ogni settimana. Un totale di 306 resti è stato rinvenuto, per lo più maschi (95 %). Un numero
così elevato permette la candidatura di tale metodo quale possibile strumento di monitoraggio della
popolazione degli individui di sesso maschile di L cervus a Bosco Fontana. La distribuzione spaziale
dei resti è risultata essere non omogenea, poiché la maggior parte di essi è stata rinvenuta in un numero
molto ristretto di strade. I dati ottenuti inoltre hanno permesso di stimare il periodo di attività degli
adulti di cervo volante nella riserva, dalla prima settimana di maggio alla prima settimana di agosto.
Le implicazioni dei risultati saranno discusse alla luce dell’utilizzo di tale metodo come strumento di
monitoraggio della popolazione di L cervus.
Parole chiave: Lucanus cervus (Linnaeus 1758) – cornacchia – Direttiva Habitat – fenologia – moni-
toraggio – predazione

1 Introduction
The European stag beetle Lucanus cervus (Linnaeus 1758) is considered a flag-ship spe-
cies for saproxylic insect conservation, a role which has recently been criticized [Thomaes,
Kervyn, Beck et al 2008]. The species typically lives in oak forests characterized by the
presence of old trees and deadwood, but significant populations have been recorded in urban
environments where remnants of historic larger forests or old trees are present [Hawes 2005,
Thomaes et al 2008]. The larvae of L cervus feed on decaying wood and development usu-
ally last 3–5 years [Franciscolo 1997]. Stag beetle larvae live in a variety of wood types:
Pinus, indigenous oaks Betula, Alnus, Carpinus, (Quercus sp), Fagus, Juglans, Salix, Populus,
Ulmus, Prunus, Malus, Pyrus, Tilia, Fraxinus, non-native Mahonia sp. [Franciscolo 1997,
Hawes 2005, Kervyn 2006, Rink & Sinsch 2008].
The distribution of this species in Europe is relatively well known: eg Belgium [Thomaes et al
2008], Germany [Müller 2001], Netherlands [Huijbregts 2002], Slovenia [Jurc, Ogris, Pavlin et
al 2008] and Spain [Méndez 2003]. In the UK, the “National Stag Beetle Survey” has been carried
out twice in 1998 and 2002 [Joint Nature Conservation Committee 2007]. In Italy, L cervus is manly
distributed in the northern and central part of the country [Bartolozzi & Maggini 2006].
Lucanus cervus is protected at various national and international levels; amongst others, the
species is listed in the Bern Convention [Council of Europe 1979] and in the Annex II of the Habitats
Directive [Council of The European Community 1992]. The main threats for the European stag beetle
are the loss and fragmentation of habitat and the availability of dead wood [Dudley & Vallauri
2004]. Monitoring the conservation status of the species is, according to Art 11 of the Habitats Di-
rective, an obligation and the results of the monitoring shall be included, every 6 years, in national
reports according to Art 17. These reports represent the main tool to verify the status progress of the
measures taken under the Directive. The 2nd Art 17-Report for Italy, concerning the period 2001–2007,
has recently been published and the conservation status of L cervus has been evaluated as ‘unfavorable
bad’ for all the biogeographical regions [La Posta, Dupré & Bianchi 2005].

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To accomplish the obligations contained in Art 17 of the Habitats Directive, standardized monitor-
ing programs are required. We now briefly analyze and summarize the proposed methods for studying
populations of L cervus in European countries.
Regarding Germany, Müller [2001] proposed a first scheme. Currently the following methods
are suggested for Germany [see Malchau 2006]:
– In order to gather first qualitative data, a survey of local naturalists, forest owners, hunters and
forest personal is to be conducted;
– Identification of ‘sap-trees’;
– Identification of brood-trees for ‘emergence-holes’ and egg-laying females;
– Searching for the presence of beetle remains and dead specimens;
– Using of traps with attractants.
In the Netherlands L cervus is too rare to allow for specific monitoring in the field. The method
proposed [Smit & Krekels 2006] consists in placing public announcements in local media and in
specialized magazines in order to collect all observations by private people. A photograph for recogni-
tion and a distribution map are also provided to the public. The data are collected in a database which
lists all sites where L cervus has been observed.
In the UK, Hawes [2005] presented data from belt transect surveys of stag beetle road casualties
in the County of Suffolk, without however suggesting a standard method.
In Spain, the main interest in conservation of stag beetle comes from amateur entomologists,
namely the Grupo de Trabajo sobre Lucanidae Ibéricos/Working Group on Iberian Lucanids (GTLI).
They indicate that: survey should start in July and should be done every 4 days for a total of seven
repetitions [GTLI 1996, 2005].
In Slovenia [Vrezec & Kapla 2007] the method adopted is based on the Spanish protocol [GTLI
2005]. A transect at the forest edge is selected and here beetles found on the ground or observed flying
are registered. The transect surveys are carried out from the second half of June to late July between
20.30 and 21.30, for a total of three surveys per transect per season [Vrezec pers. com.].
In France, region of Wallonne, the survey methods proposed consist of transect count of flying
beetles, interception traps and the search for dead individuals or remains on the ground or in owls
regurgitated pellets [Kervyn 2006].
In Italy a protocol for monitoring L cervus was proposed within the LIFE Project LIFE03NAT/
IT/000119 in the Regional Park of Colli Euganei, Province of Padova, Veneto. According to the action
F3 – Monitoring, surveys are repeated every 15 days in the period: second half of June – July [Costan-
tini, Lasen & Minelli et al 2004; Lasen, Minelli & Rallo et al 2005, 2006, 2007]. The protocol
consisted in searching for adults in late afternoon and captures by baited traps (sugar/alcoholic baits).
Data obtained with the variety of approaches presented are unlikely to provide a reliable and
consists monitoring tool for L cervus in Europe. Only a reliable and repeatable standard, applied in
all European countries, with a known level of detectability, would generate data on population size,
threat level and conservation status of this protected beetle species. The results of such a standard-
ized monitoring could be compared in space and time. Sampling design, field and analytical methods
should make it possible to discover real changes with adequate sensitivity, ie to be able to distinguish
real changes from natural variation and measurement errors [Framstad 2008]. However, systematic
quantification of L cervus is problematic [Malchau 2006] and therefore population sizes are difficult
to assess. One of the main reasons is that the species has a relatively short flight period and has its
activity peak just before dusk. For example, in Germany the main flight activity was recorded between
20.00–21.00 [Rink & Sinsch 2007] and in Italy between 20.00 and 21.30 (pers obs). Additionally,
individuals often rest in inaccessible places. These traits make it particularly difficult to obtain good
estimates of population size of L cervus [Malchau 2006; pers obs].
At the Nature Reserve Bosco Fontana the presence of the European stag beetle L cervus has been
known since Piatella [2002], but conservation status and population size have not been assessed.
However, every year a number of remains of predation are found incidentally by visitors and local
staff and this fact lead to the idea of investigating the systematic collection of remains as possible tool
for monitoring the species in the reserve.

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The present work is the first report on spatial-temporal aspects of L cervus distribution and
monitoring in Italy based on a standardized survey using remains of predation, a method that has
already been applied [Hawes 2005, Lagarde, Corbin & Goujon et al 2005, Harvey & Gange
2006] and that has been suggested as one of a number of methods for the monitoring of L cervus in
other countries, eg Germany [Malchau 2006] and France [Kervyn 2006].
In particular, the aims of the investigation were:
Quantify the number of remains which can be sampled at Bosco Fontana and quantify the search
effort;
Investigate the temporal pattern of the remains and estimate the phenology of L cervus at Bosco
Fontana;
Analyze the spatial distribution of the remains found;
Analyse the results in relation to possible monitoring schemes using remains of predation.

2 Material and methods

The study was conducted in the Nature Reserve and Site of Community Importance (IT20B0011)
‘Bosco Fontana’, municipality of Marmirolo, Province of Mantua, Lombardy, Italy. The Reserve covers
an area of 233 ha, 198 of which are occupied by a “primary ancient wood” [Mason 2004], classified
as Quercum-Carpinetum boreoitalicum Pignatti 1953.
The survey was carried out by slowly walking all forest roads of the reserve at a constant speed
and searching visually for remains of L cervus. The total length of forest roads was 23 km. The remains
of L cervus were searched once a week for 16 weeks (from 12.05.2008 to 28.08.2008). The survey
was carried out during morning hours (7:30–12:30) and always by the same person. The route walked
was changed haphazardly every week, with the aim of varying the order and the time of the day in
which specific roads were covered. For every L cervus found, the following data were recorded: date,
time, geographical coordinates (using a detailed map). All remains of L cervus found were collected
by hand and preserved, dry, in paper bags.
The field work was terminated after the 16th week as no freshly dead specimens were collected
from week 13 onwards. The authors estimate that all these specimens had been dead for more than 1
week. This approximation is based on the observation that no fatty tissues or muscles were present,
which had probably been consumed by scavengers. Additionally, head and pronotum were always
separated.
The chi-square test was used to test if the spatial distribution of remains deviated from a ho-
mogenous distribution. For every road the observed number of remains was compared with the one
expected if the beetles were homogeneously distributed. The expected number of remains per meter
was calculated by dividing the total number of remains found by the total length of forest roads.

3 Results
The total number of remains of L cervus collected was 306 and 94.8 % (290 remains)
of these were males. Only 5.2 % (16 remains) of the beetles were females and for this reason
the latter were excluded from further analysis.
The number of remains collected during the study period is shown in Fig 1. During
the first week of survey (12.–18.05.2009) 4 specimens were found, during the last week
(25.–31.08.2009) 5 specimens were collected. The maximum number of specimens collected
was 56 in the 5th (09.–15.06.2009) and 8th (30.06–06.07.2010) week.

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 Monitoring of Lucanus cervus – 083

Fig 1: Number of remains of Lucanus cervus per week (from 12.05.2008 to 28.08.2008) [Coleoptera:
Lucanidae].

The distribution of remains, which is shown in Fig 2, was not homogenous as the
number of remains observed in the forest roads is significantly different from the expected
number (χ2 = 1.979,761, df = 74, P < 0.001). For example, in 20 % (n = 18) of the forest
roads a total of 85 % (n = 247) of all remains were found. In the S-E part of the Reserve a
higher number of remains was collected than was expected (Fig 2).
The effort devoted to carry out the present survey can be expressed as the total distance
covered (370 km) and the total time of the survey (192 hours). Thus for every km covered
an average of 0.43 male remains were found and every hour an average of 1.59 remains
were collected.

4 Discussion
The total number of remains collected (306) is high, especially if the relatively small
size of the study site (233 ha) is considered. It can be concluded that the collection of remains
of predation allowed to ascertain the presence of L cervus at Bosco Fontana with a high
level of reliability. It should be mentioned here that during the survey only a very limited
number of live adults (0 males, 4 females) were observed. This can be imputed to the time
of the surveys, which was always carried out during daylight hours, a time of the day well
separated from the main period activity [eg Rink et al 2007].

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Remain
Forest track
River
Grassland

Fig 2: Map of the Nature Reserve Bosco Fontana with the locations of remains of Lucanus cervus
males collected during the study.

Even though investigations carried out by other authors employed different methods,
some quantitative figures are given here, in order to facilitate categorizing the number of
remains collected; Clark [1977] collected 573 specimens in the town of Colchester (UK)
over a period of 3 years. Hawes [2008] collected 138 specimens in a village garden in UK,
while Rink et al [2007] collected 56 specimens over 3 years in a rural area in Germany.
Lagarde et al [2005] collected 93 remains of predation in the forest of Chizé (France).
Predation of L cervus, observed during the present study, was attributed mainly to
hooded crows (Corvus corone cornix); this is based on direct observations carried out by the
staff of the Reserve (2008). The exploitation of insects and in particular of beetles as food
source by the hooded crow is well known [see Cramp & Perrins 1994, Roselaar 1994,
Horgan & Berrow 2004]. The predation by birds is also supported by the morphology of
bite-marks on the elytra, which were typical of beak bites and not of predatory mammals.
The last census of the hooded crow conducted at ‘Bosco Fontana’ resulted in 15–20 couples
[Longo & Nadali 2001]. We assume the hooded crows catch the stag beetles on trees and
subsequently fly to open ground (forest roads) where they consume the prey, a behavior also
reported by Glutz von Blotzheim & Bauer [1993] for insect preying birds in general.

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Birds have an advantage to consume stag beetles on the forest roads and not in the
forest for purely mechanical reasons and because here the visibility of a potential predator
is higher (a vast literature exists about visibility, vigilance and foraging efficiency for birds:
eg Lazarus & Symonds [1992], Whittingham & Evans 2004, Devereux, Mckeever &
Benton et al [2004]. This behavior of hooded crows, which leads to the remains of preda-
tion being left on forest roads, facilitates their detection.
The sex-ratio of the specimens collected by means of remains of predation was highly
biased towards males: the number of males collected was 290 from a total of 306 specimens
(sex-ratio 94.8 % males, 5.2 % females). This value is not in line with data in literature.
Hawes [2005] reported sightings of 11,523 specimens of L cervus during surveys conducted
in 1998 and 2002 and the sex ratio was: 54.3 % males, 45.7 % females. However, when
only road casualties are considered, the sex ratio of female to male beetles was 3:1 [Hawes
2005]. The ratio observed at Bosco Fontana is probably a result of the collection method and
does not reflect the sex-ratio of the population. Nevertheless, the highly skewed sex ratio of
the remains collected contains important biological information on behavioral differences
between the sexes. Males should be at much higher risk of predation by birds because they
are much more active and more commonly seen in flight [Rink et al 2007, Hawes 2008]
and so are highly visible to predators [Harvey et al 2006].
The phenology of L cervus in Italy has to our knowledge not been systematically inves-
tigated before. According to the data collected during the present study, in 2008 the flight
period of male L cervus at Bosco Fontana started in early May. Given that the first survey
was successful, it seems likely that emergence already began at least one week earlier. The
end of adult activity was covered by the investigation, but it cannot be determined reliably
for methodological problems, as old remains were still found after the end of the flight period
and remains collected in a given week can be the result of a predation event which occurred
weeks earlier. However, the condition of the remains gives some clues to their age. The here
reported observations indicate that first all fatty tissues and muscles are eaten (mainly by
ants) and later head and pronotum separate. Considering these observations, it is possible
to estimate the end of the flight period for L cervus at Bosco Fontana. The last remain with
head and pronotum still connected was found between 18 and 24 August 2008. This time
period is likely to coincide with the end of the activity of adults. Thus, one may estimate
that the flight period of L cervus males at Bosco Fontana in 2008 ranged from the first week
of May to the first or second week of August.
The phenology inferred for Bosco Fontana is very similar to the data reported by
Sprecher-Uebersax & Durrer [1998] for Switzerland. Here specimens were observed
from May to July. In Asturias (northwestern Spain), imagoes are reported to appear form
the middle of June and are present to the end of August or early September [Álvarez Laó
& Álvarez Laó 1995]. In Slovenia adults have been collected from late March, to late
September, with a continuous presence ascertained from early May to late August [Vrezec
2008]. Vrezec [2008] reported the period for the 2nd and 3rd quartile of records to be to
emerge 30.06.–23.07. and this phenological period has shifted towards spring by 16 days since
1900. Calculating the 2nd and 3rd quartile for the phenological data obtained at Bosco Fontana
gives the following dates: 13.05.–07.06.2008. The difference might partly be explained by
the known protandry of L cervus [Percy, Bassford & Keeble 2000, HAWES 2008]. The
data on the phenology reported here are in contrast with those reported by Franciscolo
[1997] for Italy, who gives the activity period as ranging from mid-July to the end of August.
During the weeks 6 and 7, in the peak of the flight period, relatively low numbers of beetles
were collected (Fig 1). These weeks were characterized by heavy rain and it is known that
under bad weather conditions L cervus flying activity is reduced [Rink et al 2007].

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Thus, the low number of remains collected during this time span is very likely a con-
sequence of the poor weather rather than a decline in population size.
The spatial distribution of remains of L cervus was not homogenous and no temporal
variation in the spatial distribution pattern was observed, by visual inspection of the distri-
bution of the remains collected on a weekly basis. Reasons for the clumped distribution of
the remains could be caused by:
a) the method employed;
b) the behavior and the distribution of the predator;
c) the biology of L cervus.
It seems unlikely that the method employed is responsible for the observed pattern as the
routed walked was changed haphazardly every week and thus any possible temporal differences
in the presence of remains and any decline in attention by the operator during the searches
would not translate in clear spatial patterns. The vegetation cover of the forest roads is likely
to be negatively correlated to the visibility of L cervus remains. However, due to management
actions (mowing) and vegetation changes during the season, the visibility varied in a number
of forest roads during the survey. In some roads which offered a very good visibility during
the study period, very few remains were found. It thus seems unlikely that vegetation cover
of forest roads was responsible for the distribution of the remains collected (Fig 2).
Larvae of L cervus, feed on dead wood of roots or of trunks in contact with the soil,
where they move before entering in the pupal stage [Franciscolo 1997, Rink et al 2008].
The distribution of this micro-habitat at Bosco Fontana is not known, but given the current
knowledge about the forest structure [Giannelle, Travaglini & Mason et al 2007] it
seems likely that suitable sites for larvae are not homogenously distributed. Forest composi-
tion and forest structure [Mason 2004] are also likely to influence the distribution of adult
male stag beetles. Visual comparisons of the distribution of remains with detailed maps of
tree species in the surrounding forest [Dalponte, Giannelle & Bruzzone 2007] did not
reveal any correlations.
In 2009 and 2010, transect counts of flying males of L cervus showed that the high-
est density was found in the forest roads where the largest number of remains of predation
had been found (pers obs). We therefore assume that the highly clumped distribution of the
remains reported in Fig 2 is closely related to the real distribution of the males of L cervus
at Bosco Fontana.
On the basis of the results presented, it is possible to suggest a new monitoring proto-
col for Bosco Fontana, which focuses on the period of the year when the highest number of
remains was found and on the area where most were collected in order to optimize the ratio
of costs (in terms of time spent and distance covered) and results (in terms of remains col-
lected). This new protocol is based on the assumptions, that distribution of stag beetle and
abundance of hooded crows remain stable at Bosco Fontana.
The new protocol:
– Duration: from the last week of May to the last week of July (total 10 weeks);
– Frequency: one weekly survey;
– Course: reduced to 13 roads (4.2 km).
From the data collected in 2008 it is possible to extrapolate that the protocol presented
above would have resulted in the collection of 80 % of the remains while employing only
16 % of the time.
We suggest that similar protocol can be applied in other forests in Italy (and in Europe),
considering the local phenology of L cervus, and might provide a valid tool for monitoring,
as collection of remains has already been carried out in other regions of Europe for other
purposes (Lagarde et al 2005, Harvey et al 2006).

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The influence of factors like forest composition, forest structure, road characteristics,
morphology of the terrain and presence of members of the family Corvidae should be ad-
equately taken into account.

4.1 Conclusions

The methods published for monitoring of L cervus in Europe are highly heterogeneous,
do often not consist of a single standard method and thus repeatability of the results is likely
to be low. A standardized count of remains, as tested here, might provide a valid method
which is easily repeatable in time. The survey carried out at Bosco Fontana was considered to
have generated sufficient data to ascertain the presence of L cervus at Bosco Fontana reliably
and to analyze the phenology of male stag beetles. With variants appropriately designed, the
method might be applied in other forests and should be evaluated in correlation with other
quantitative monitoring methods as a standard monitoring tool for populations of L cervus.
The proposed method has some advantages in comparison to others:
– The impact on the population is minimal or non-existent, a very important aspect when
working on rare and protected species;
– The method is easily applicable even by not-specialized staff/personnel;
– The method is relatively cheap and does not require specialized equipment;
– The method allowed collecting a large number of remains of males;
– The method can be standardized.
In this context it is important to point out that a highly aggregated spatial distribution
of remains is possible and that this property makes standardized monitoring, using remains
of predation, more complicated. If one re-considers the example given in the results section
from another point of view, it shows that leaving out only 20 % of the forest tracks (eg to
reduce the work load) could result in missing 85 % of all remains.

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Acknowledgements
We thank Franco Mason for giving us the opportunity to carry out this study. We are indebted to Luca
Bartolozzi for providing important information regarding ecology of stag beetles in Italy. Our
thanks go also to our colleagues of CNBFVR – Bosco Fontana for their help during the research
and especially to Emma Minari for technical support.
Authors’ addresses: Alessandro Campanaro1,2, (corresponding author), Ilaria Toni2, Sönke Hardersen2
& Donato A. Grasso3
1 Dipartimento di Biologia e Biotecnologie “Chares Darwin”, Sapienza Università di Roma, P.le

Aldo Moro 5, Roma, IT-00185.

2 CNBFVR – Centro Nazionale per lo Studio e la Conservazione della Biodiversità Forestale “Bosco

Fontana” di Verona, Corpo Forestale dello Stato, Strada Mantova 29, Marmirolo (MN), IT-46045.
3 Dipartimento di Biologia Evolutiva e Funzionale, Università degli Studi di Parma, Via Usberti

11/A, Parco Area delle Scienze, Parma, IT-43100.

E-mail: ale.naro@gmail.com

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