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Journal for Nature Conservation 20 (2012) 101– 108

Contents lists available at SciVerse ScienceDirect

Journal for Nature Conservation

jou rn al h omepage: www.elsev ier .de / jnc

eview

anging on by the tips of the tarsi: A review of the plight of the criticallyndangered saproxylic beetle in European forests

akub Horáka,b,∗, Karel Chobotc, Jana Horákovád

Department of Biodiversity Indicators, Silva Tarouca Research Institute for Landscape and Ornamental Gardening, Kvetnové námestí 391, CZ-252 43 Pruhonice, Czech RepublicDepartment of Ecology, Faculty of Environmental Sciences, Czech University of Life Sciences, Kamycká 1176, CZ-165 21 Prague 6, Czech RepublicAgency for Nature Conservation and Landscape Protection of the Czech Republic, Nuselská 39, CZ-140 00 Prague 4, Czech RepublicDepartment of Forest Protection and Game Management, Faculty of Forestry and Wood Sciences, Czech University of Life Sciences, Kamycká 1176, CZ-165 21 Prague 6, Czech Republic

r t i c l e i n f o

rticle history:eceived 10 April 2011eceived in revised form4 September 2011ccepted 19 September 2011

eywords:ommercial forestry

a b s t r a c t

Since the beginning of the new millennium, many conservation biologists and forest managers havebeen discussing the future of European forests. Historical evidence shows that the diversity of saproxylicbeetles, a key measure of forest biodiversity, has declined at a frightening pace. Most of the data regardingspecies-rich forests were collected during a period when most European forests were managed usingtraditional management practices. We present extinction and genesis of relictual distribution of Cucujushaematodes, one of the three most endangered saproxylic beetles in the EU. We also analyse and comparethreats to its presence and extinction according to forest history, management and current conditions

ucujus haematodeson-intervention managementelictual occurrenceraditional management

in European forests. Our review showed that one of the main aims of conservation efforts relating tosaproxylic beetles should focus on the refinement of the profound effects of commercial forestry andon respect for forest history and traditional forest management. Traditional management practices andtheir principles present one solution to the problem of decreasing forest biodiversity. We believe thatour review can help stop the decrease of forest biodiversity in an era when people and large institutions

d abo

are increasingly concerne

im of this review

The main objectives of this review were the following: (i) toresent a survey of distributional data of the critically endangeredaproxylic beetle Cucujus haematodes Erichson, 1845 (Coleoptera:ucujidae) in Europe; and, (ii) to analyse and compare threats to

ts presence and extinction according to forest history, manage-ent and current conditions in European forests. We focus on the

tory of the decline of an interesting and rare saproxylic beetle thatmphasises the importance of continuous forest ecosystems andighlights the effects of declines in primary forest habitats. Thisaper presents a summary of the status of this beetle and bringsttention to the plight of insects that are frequently ignored inainstream conservation.

istory of forest management in Europe

The history of forest management in Europe is one of the best-ocumented human activities (Rackham 2006). It is well known

∗ Corresponding author at: Silva Tarouca Research Institute for Landscapend Ornamental Gardening, Kvetnové námestí 391, CZ-252 43 Pruhonice,zech Republic.

E-mail addresses: jakub.sruby@seznam.cz, jakub.sruby@gmail.com (J. Horák).

617-1381/$ – see front matter © 2011 Elsevier GmbH. All rights reserved.oi:10.1016/j.jnc.2011.09.002

ut nature and the environment.© 2011 Elsevier GmbH. All rights reserved.

that humans had a variable influence on different parts of the OldContinent. The human influence was largely attributable to thedegree of settlement and the cultural history of nations. Thus, thefirst large forest areas were deforested in Southern Europe in theAncient World, a pattern which has been documented in severalhistorical accounts (e.g., Burnet 2004).

The history of forest use and abuse in Western Europe has beenwell documented (Grove 2002). Forests had scarcely reached theirmaximum post-glacial extent when humans began clearing them.In the following millennia, forest cover was drastically reduced andthe structure and composition of the remaining forest fragmentswere greatly altered over time (Thirgood 1989).

The forest management systems in Central Europe were notmuch different from those in Western Europe. However, saprox-ylic organisms in Central Europe seemed to be less jeopardised dueto their distance from the maintained areas of Eastern Europe andslower movement of influence of agricultural revolution toward theeast of Europe. The oldest and best-known central European forestreserves are found there (e.g., Białowieza in Poland and Belarus,Zofín in the Czech Republic or Badín in Slovakia; e.g., Wesołowski2005).

Northern Europe has been negatively affected by forestpractices over the last centuries (e.g., due to factors such asclimate or settlement). However, the influence of intensiveforestry practices and fatal structural changes of woodlands in a

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02 J. Horák et al. / Journal for Natu

elatively short period of time is more negative (Lindbladh et al.007).

Populations of saproxylic organisms seem to be most stable inastern Europe. Recent studies have indicated the importance ofonserving this area (Bussler et al. 2005; Siitonen & Martikainen994). Furthermore, this area’s fauna has not been well studied,hich is likely the reason that there is little data from this area.

se, abuse and conservation of European forests

Recently, many authors have written about the ecosystemhanges and impacts caused by forestry (Grove 2002; Lindbladht al. 2007; Ranius & Jansson 2000; Simberloff 1999). Humanctivities have been the main factors influencing the condition ofuropean forests after most of the large herbivores died out. Humanctivities such as the following have created a spatial mosaic of dis-arate forest habitats – traditional forest management systems inhe form of wood pasturing, coppicing with or without standards,reaming, pollarding, litter gathering, charcoal burning, resin tap-ing, alternating forest and farm crops or floating. Recent studiesave suggested that many species benefit from traditional low-

mpact forest use (Hofmeister et al. 2004; Ranius & Nilsson 1997;utherland 2002; Sverdrup-Thygeson & Ims 2002; Vodka et al.009). Human effects on forests have intensified over time. In the

ast two centuries, and particularly at the beginning of the Industrialevolution, forest management became more uniform and highlyffective. Thus, modern forestry has had many negative effects,uch as the decline of many forest species.

Traditional (or historical) forest management vanished and waseplaced with new management concepts. Even though traditionalorest management systems persisted in large areas for millennia,t seems that today, due to many factors, people are not able to

able 1he main events in European forests that possibly affected saproxylic beetles.

Period Events

After glacial epoch (Whittle 1996; McNeely 2002) Impact of large herbivHunting as the main s

Neolit (6000–4000 BC) (Küster 2000; Vera 2000) First settlementsAgriculture and wood(fuel-wood), creamingMany microhabitats d

Eneolit (4000–3000 BC) (Mládek et al., 2006) Intensification of agriProbably first instance

Bronze age (3000–1000 BC) (Agnoletti & Anderson 2000;Billamboz 2003; Fanta 2007a; Harding 2000)

Permanent sedentaryFirst clear-cuttings fo

Iron age (1000 BC–0) (Fanta 2007a; Küster 2000; Mládeket al. 2006)

Organised pastures (eFirst haymakingLarger exploitation ofSettlements on non-a

Antiquity (3000 BC–5000) (Mládek et al. 2006) Housing of livestock iMiddle ages (5000–15 000) (Fanta 2007a; Haneca et al. 2005) Feudalism (the pastur10th–12th Century (Agnoletti 2000; Küster 2000) Establishment of perm13th Century (Fanta 2007b; Mládek et al. 2006) Fee for common right14th Century (Fanta 2007b) First efforts in forest p

First seedlings (humaIntroduction of saw to

15th Century (e.g., Szabó 2010) Lowlands almost defo16th Century (Putman 1986; Fanta 2007b) Energy crisis (insuffic

First tendencies to enFirst silvicultural met

17th Century (Fanta 2007b) 30-Years War (forestaFirst half of 18th Century (Fanta 2007b) First real economical

Cattle plague (strong

Second half of 18th Century (Gimmi & Bürgi 2007; Mládeket al. 2006)

Wood pasture prohibYear-long housing of

19th Century (e.g., Vrska et al., 2001) First forest protected

20th Century (e.g., Konvicka et al. 2004) Intensification of foreProlongation of rotati

nservation 20 (2012) 101– 108

restore these systems (Konvicka et al. 2004), even if this manage-ment supports the continuity of forests.

For example, in Central Europe, forest management hasoccurred on two extremes. The first was the profound effects ofcommercial forestry (i.e., high forest, plantations, clearcutting, sal-vage logging, etc.), which resulted in a general lack of dead woodin European forests (Kaila et al. 1997).

On the other extreme, non-intervention (hands-off) forest prac-tices were also implemented. Non-intervention practices weremostly implemented in response to the negative changes causedby commercial forestry (e.g., Cronon 1996). These practicesbecame more popular in the 19th century. The main objectivesof non-intervention practices concerned aesthetics and ethics,namely to conserve the last remains of virgin forests in Europe(Vandekerkhove et al. 2009). Conservationists aspire to restore orestablish new primeval forests (Hruska 2008).

There is evidence suggesting that virtually all forests have beensubstantially influenced by humans, and most forests have beenaffected for at least several thousands of years (McNeely 2002).Furthermore, if there is really no virgin state of forests, then thepreservation of natural processes in forests (i.e., non-interventionpractices) for reference purposes would be a waste of resources(Lawton 1997; Grove 2002).

Many threatened species are jeopardised by commercialforestry (Konvicka et al. 2008a; Ranius & Nilsson 1997). Speciesdeclines are usually attributed to the low availability of dead woodand the high tree densities in managed forests (see high forest inPeterken 1993), which affected the shading of species’ habitats. Itis therefore important to increase the supply of dead wood through

actions such as the establishment of non-intervention reserves(Vrska et al. 2006). However, this practice is restricted by severalfactors (Table 2) and is only possible in small areas. A high numberof specialised woodland species from multiple groups require open

oresource of food (one of the main factors why most large herbivores declined)

pasture, wood fires (for fields and boundaries), pollarding (browse), coppicing (building timber)ue to pressure of wood pasture out of vegetation period

culture and impact of wood pasture of livestock stabling, thus, litter gathering

settlements, long-time maintenance of farmland (permanent deforestation)r bronze melting.g., use of watchdogs), stables for livestock (still depending on pasture)

forest land for minesgricultural land (due to vicinage of mineral or natural sources)n the winter (decrease in wood pasture pressure)e was the main domicile of subjects)anent arable land with fallows (permanent deforestation of large areas)

s (pasture, litter gathering, etc.)rotection (unregulated felling was prohibited)

n-caused target tree species composition) the forestryrested, large exploitations in higher altitudes and wood floatingiency of fuel-wood)join wood pasturehods (e.g., plantings)tion of landscape)

tendencies in forest usereduction of wood pasture)ited (e.g., in Austria-Hungaria)livestockareas (established by nobility) and first non-intervention practicesstry and homogenisation of landscapeon period (end of coppicing and coppicing with standards)

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Conservation 20 (2012) 101– 108

103

Table 2SWOT analysis (based on studies such as Alexander & Butler 2004; Bogucki 1988; Buckley 1992; Cronon 1996; Peterken 1993, 1996; Szabó 2010; Thirgood 1989; Vera 2000; Vrska et al. 2006, 2009).

Strengths Weaknesses Opportunities Threats

Non-interventionLarge quantities of dead wood Generally without influence of large herbivores Observe natural processes Loss of sun-exposureSupply of different diameter, position and decay

stage dead woodLocking up for people Shelter without human stress factors Tendency to create artificial climax

Support forest continuity Small area High game stocksOften on localities with long-time human influence Changes in tree composition to shade-tolerant tree speciesNo economical production Unsafe for peopleLow support in recent legislative Tended to create only one type of habitatArtefact of recent time

CommercialMoney Artificial regeneration Affect “negative” nature disturbances Changes in tree compositionSafe for people Expensive tendings Potential to equalise Mono-culturingReal forest in human mind Disdain for dead wood Orderly forest Clear-cuts causing loss of forest continuityUniform trees for the timber industry Homogeneity of forests Loss of high diameter trees

Source of funds for corporations Large areas of salvage cuttingsDiseases and outbreaksShadingChanges in the spatial composition of dead wood

(horizontal and vertical)TraditionalMany non-wood producing roles of the forest Difficult natural regeneration Create sun-exposed dead wood Loss of know-howCreate mosaic of habitats Difficult use of modern logging and hauling machinery Useful for large areas Loss of lying dead woodCreate microhabitats Low support in recent legislative Potential of stabilityOpen and safe for people Often lower quality timber Simulate influence of large herbivoresSupport forest continuity Lower economic profitLong traditionSource of local revenuesMyriads of organisms have been accustomed to

this for millennia

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propetrovsk (Mamaev et al. 1977). The beetle was also knownto be present in many parts of European Russia, such as inMoskovskaya, Kirovskaya, Yaroslavskaya, Ul’yanovskaya oblast’

04 J. Horák et al. / Journal for Natu

abitats (Ranius et al. 2005; Vodka et al. 2009). Nevertheless, a for-st biodiversity strategy also should not ignore the critical habitatsreated by disturbances (Sverdrup-Thygeson & Ims 2002).

vents in forest history in Europe through the eyes of aaproxylic beetle

We documented the main events in forest management historyrom the viewpoint of the saproxylic beetle (Table 1). Our analysesllowed us to identify the main problems of recent forest manage-ent. Many of these are well known, such as the loss of dead wood

nd changes in the tree species composition (Jonsson et al. 2005),lthough some of these problems are still not widely accepted.

Table 1 illustrates that forest ecosystems and managementhange over time. Recent studies (Sutherland 2002; Vera 2000;

hitehouse & Smith 2004) have indicated that the European land-cape was historically much more open than previously realised.umans began to have effects on forests soon after the last Ice Age

McNeely 2002). Human effects were likely important to the declinend extinction of large herbivores and carnivores (e.g., Vera 2000).arge herbivores were often important factors in creating opennessn the forest landscape (Putman 1986). The first settlements werereated by deforestation and gave rise secondary forest habitatsfter their abandonment (Küster 2000). Wood pastures and tradi-ional activities such as creaming or litter-gathering helped to formuropean forests as early as 5000 BC (Harding 2000). The growinguman population intensified and extended its influence on forestabitats. Humans likely influenced larger areas than we previouslyealised. For instance, lowlands typically have more open forestsKonvicka et al. 2004). There are also many indications of humanctivities in highlands and mountains. For example, many moun-ainous areas were deforested to create meadows or pastures (e.g.,olany or Poloniny in the Carpathians; Janicki 2005). Large partsf highland forests were pastured by cattle (e.g., in terrain thatas difficult to access) or deforested for mines. This pattern beganuring the Iron Age and intensified during the Middle Ages. Dur-

ng the Industrial Revolution traditional management systems hadanished from most of the European landscape.

omparison of forest practices that affected saproxyliceetles

In support of our conclusions based on existing data, we used SWOT analysis (Table 2) to compare the different forest prac-ices. A SWOT analysis is a method of identifying the strengths,eaknesses, opportunities and threats of several entities such asrojects, businesses, actions, policies and politics, and uses a mul-idisciplinary approach. Furthermore, SWOT analysis is part oftrategic and long-term planning and can be used to detect prob-ems or drawbacks in certain projects (Armstrong 2006).

Commercial forestry has occurred for approximately two orhree centuries (Peterken 1993; Table 1). Surprisingly, non-ntervention practices were not used frequently in most Europeanorests (Peterken 1996; Table 2). Non-intervention practices couldave similar implications for saproxylics like too much care for non-

orest species (Konvicka et al. 2008b), because they often causehading of habitats. Furthermore, the conservationists who wereesponsible for the creation of non-intervention sites confess thatather than conserving genuineness or biodiversity, the emphasishould be on creating wilderness (Zajoncová & Vrska 2008). Euro-

ean forests have been affected by humans for millennia. Thus, theorests that we protect are the heritage of our ancestors. Practicallyverything is a consequence of various impacts of human beings oncosystems (Lawton 1997; McNeely 2002).

nservation 20 (2012) 101– 108

Cucujus haematodes – a vanishing saproxylic relict

The saproxylic genus Cucujus Fabricius, 1775 is distributedthroughout North America and Eurasia. Cucujus cinnaberinusScopoli, 1763 and Cucujus haematodes are both found in Europe.1

Whereas Cucujus cinnaberinus is distributed only in Europe, Cucu-jus haematodes is distributed throughout most of Eurasia (Horák& Chobot 2009). Although, Cucujus cinnaberinus is one of a fewanimals that have been focused on by mainstream nature conser-vation (e.g., Baillie & Groombridge 1996), Cucujus haematodes wasneglected. Furthermore, and contrary to the suggestion of manyauthors, this species is a relict of primeval forests (Burakowski et al.1986; Franc 2001; Gutowski et al., 2006; Horion 1960). Nowadays,the species is one of the three most endangered saproxylic beetlesin the European Union (Nieto & Alexander 2010).

Summary of historical and recent distribution of Cucujushaematodes in Europe

The distributional range of Cucujus haematodes extends west toCentral Europe and specifically to Germany. Erichson (1845) indi-cated that this beetle was distributed in many parts of Germany.However, the only known specimen is from Ruhpolding in theBavarian Alps, and the collection site of this specimen was indi-cated by Horion (1960) as being misidentified. Borchert (1938) alsodiscussed species collected in München, but this likely referred toa sample collected by Reitter (1911). Moreover, the records fromBavaria that do not include a voucher could be confused withCucujus cinnaberinus (H. Bussler, pers. comm.). Recent evidencesuggests that the distribution of Cucujus haematodes extends tothe Czech Republic and that one of the historical locations wherethe species was collected is close to Bavarian border. The specieswas historically present in more locations than it is currently. How-ever, only four sites from the Czech Republic were documented instudies (Karlova Studánka, Praded, Vsetec and Stramberk; Horáket al. 2009). The species was distributed in Poland throughoutmost of the country. Authors indicated many localities – Kłodzko,Puck, Ostróda, Gdansk and Rosztocze (Horion 1960), Białowieza(Burakowski et al. 1986), Duszniki-Zdrój and Szczeliniec (Zebe1852) and Ustron (Roger 1856). The species appeared to be rel-atively widespread in many largely undisturbed forest areas ofSlovakia (Borchert 1938; Horion 1960; Roubal 1936).

The southernmost part of the historical range in Europe wasPollino in southern Italy (Borchert 1938; Palm 1941; Ratti 1999),Taygetos in Greece (Horion 1960), Velebit in Croatia (Borchert1938) and Mok. Poljane in Bosnia & Herzegovina, which are bothlocated in the Dinaric Alps (Horion 1960).

In Northern Europe, the species was known to be distributedin Scandinavia and the Baltic. Borchert (1938) indicated that thespecies was known to occur in southern Finland, in the westernpart of Latvia (Kurland) and in Kronstadt in Russia. The species waslikely also distributed in Estonia (cf., Lilleleht 2001–2002).

There is little data from Eastern Europe due to the limitedknowledge of the fauna in this region. Horion (1960) mentioneda record from Baile-Tus nad in Romania, whereas Borchert (1938)mentioned Mahılëu in Belarus. In the Ukraine, the species occurredin Bolekhov (Horion 1960) and in Srednaiya Polost’ in Dne-

1 Third species, C. siculus Pic, 1894, was found only in Italy. This species was con-sidered by Ratti (1986) to be introduced and identical to the nearctic species C.clavipes (Fabricius, 1781).

J. Horák et al. / Journal for Nature Conservation 20 (2012) 101– 108 105

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Fig. 1. Distribution, extinction and genesis of relictual distribution

nd in the Respublika Tatarstan and Udmurtia (e.g., Borchert 1938;orion 1960; Mamaev et al. 1977).

Today, Cucujus haematodes is probably extinct in Southern andorthern Europe. The westernmost occurrence of Cucujus haema-

odes in recent years occurred in the easternmost part of the Czechepublic (Horák et al. 2009). In this location, the species occurs inne of the last remnants of traditional pasture woodland in Mionsí.his is the largest complex (170 ha) of a mixed fir and beech for-st in the Czech Republic (Vrska et al. 2000; Weissmannová 2004).he species also occurs in Białowieza. The Białowieza Forest is sit-ated on the border between Poland and Belarus (1600 km2) and

s the last large remaining fragment of the former deciduous andixed forest of the Northern Temperate Zone in Europe (Bobiec

002; Wesołowski 2005). Contrary to the status of this species inost of central Europe, there is evidence of recent records of the

pecies from central Slovakia (Franc 2001).Due to an intensification in forestry practices throughout the

ast two or three centuries, only remnants of former large scaleraditional forestry operations exist in Europe, most of which areound in Eastern Europe. Thus, the species likely occurred more fre-uently in Eastern Europe in recent times. For instance, the beetleeems to be relatively common in European Russia (A. A. Zaitsevers. comm.).

Cucujus haematodes seems to be relatively common in manyarts of its range in Asia. This is likely due to the maintenance of theosaic of managed and natural biotopes. For instance, the species

s widely distributed and is relatively common in the Far East onhe eastern border of its former distribution (Horák et al. 2011).

Our review illustrates the importance of documenting the distri-ution and decline of perhaps somewhat less charismatic species,ecause its occurrence in Europe shows a relictual distributionFig. 1).

Recent records (occurring since 1950) are represented by blackelds; records from before 1950 are dark grey. The distribution inuropean states is represented in light grey. Thick black lines indi-ate state borders, whereas thin grey lines represent the provinces

cujus haematodes in Europe (adapted from Horák & Chobot 2009).

of the states. We also show the distribution of the species in Cau-casus here, which is a disputed part of the Old Continent. However,the continuous occurrence (e.g., Nikitskiy et al. 2008) of the speciesillustrates its decline in the western parts of the distributional area.We used ArcView GIS (version 3.3; background layers provided byESRi, Inc.).

Synthesis

Our review shows the status of Cucujus haematodes populationsin Europe. This example of a vanishing saproxylic species is alarm-ing because the species is a typical forest organism. Moreover, thediversity of the organisms found in the forest is thought to be enor-mous. Forests are not only the home of rare saproxylic beetles, butalso of many species that are rare or extinct elsewhere. Thus, manyforest species have begun to decline steeply or have become extinctin their distributional areas (Wesołowski 2005).

Non-intervention reserves increase the supply of dead wood(Vrska et al. 2006), and it is indicated that sites with a high con-nectivity of dead wood pieces contained more species. However,Schiegg (2000) demonstrated that there was a higher diversity ofsaproxylic beetles at sites with spatially distributed dead woodthan at sites with clumped dead wood distributions. Moreover,the authors dealt with temporal continuity, the openness of forestsand traditional human intervention. For example, Ellenberg (1988)indicated that deciduous trees are almost completely intolerantof shade and produce relatively little shade. In addition, manysaproxylic fungi were indicated as being shade dependent. Contraryto general expectations, several saproxylic fungi species prefersun-exposed habitats (Lindhe et al. 2004). Furthermore, many ver-tebrates, including snakes and reptiles, are shade intolerant (Toddand Andrews 2008). Shading processes threaten organisms such

as day butterflies or higher plants (Anthes et al. 2008; Konvickaet al. 2008b; Rolecek 2005). Spitzer et al. (2008) demonstrated thepositive effects of stand openness created by traditional forestryon epigeic invertebrates and on many relict species that tend

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06 J. Horák et al. / Journal for Natu

o be associated with sparse stands. In addition, several Dipteraepend on sun-exposed dead wood (Bouget & Duelli 2004). Thus,hitehouse and Smith (2004) point out that if the primeval forestas uniformly dense, then it is unclear where all of the numerous

ight-demanding species of plants and invertebrates evolved.The distribution of Cucujus haematodes could provide one of the

nswers to many questions about the condition and history of Euro-ean forest ecosystems. The species is widely distributed and iselatively common on the eastern edge of its range in Eastern AsiaHorák et al. 2011). However, a strong decline in the populationsn the western edge of its range (along with favourable conditionsn the eastern parts of its range) seems to indicate that the histor-cal western border might be much further west than was initiallyhought. Furthermore, there are other indications of the plight ofhe forest organisms, such as the extinction of many saproxylic bee-les in the Great Britain (Whitehouse & Smith 2004). Buckland andinnin (1993) listed the species as being known from the Greatritain only from sub-fossils. None of these species are globallyxtinct, but most now survive only within tiny refuges elsewheren Europe (Speight 1989). Other rare saproxylic species are knowno be distributed in Eurasia, and these species have also declinedn the western border of their range in Europe. Boros schneiderianzer, 1795 and Pytho kolwensis Sahlberg, 1833 are stable on theastern edge of their distribution area, but in many parts of Europehey have declined or have become regionally extinct (Karalius &lazyte-Cereskiene 2009; Siitonen & Saaristo 2000). Furthermore,he congenial species Cucujus cinnaberinus have also declined in

any parts of its range (Horák et al. 2010). Thus, Cucujus haema-odes could have become extinct in many parts of Europe before itas recorded.

Recent commercial forest management practices are most likelyriving the massive decline of saproxylic beetles in the Old Conti-ent, particularly in the western regions. In Europe, only remnantsf temporally continuous, traditionally managed forest habitatsxist, and many of these remnants fail to satisfy the demands ofhe threatened saproxylic beetles, including the presence of deadood, higher degrees of sun exposure and respect for natural dis-

urbances.The habitat loss and range restrictions of many saproxylic

pecies throughout Europe are extremely alarming (e.g., Nieto &lexander 2010). At the beginning of the 19th century, the dis-

ribution of Cucujus haematodes extended as far as Bohemia (androbably even Bavaria) and the beetle was also distributed in North-rn and Southern Europe. This beetle is now extinct or has declinedrecipitously in most parts of Europe, and we should see that iteclined at smaller geographic scales, which may suggest a largerattern of future decline throughout the entire distribution areaor this species and for other rare forest species. This conclusions identical to that of Wilson et al. (2004) that rare species livingn large, patchy areas are more endangered than species living inmall, integrated areas.

Cucujus haematodes is critically endangered (not only rare) oregionally extinct in almost all of Europe. As traditional and con-inuous forest habitats are lost, so are their biological relics. Weope that this paper contributes to the conservation of the disparatepecies of the world’s natural heritage in an era when people andarge institutions are increasingly concerned about nature and thenvironment.

onclusion

Our review suggests that recent forest management in Europeas lead to the decline of rare saproxylic beetles. Conservationtrategies concerning saproxylic species could focus on main-enance and a return to or a simulation of traditional forest

nservation 20 (2012) 101– 108

management practices, which were used by our ancestors andcreated a mosaic of habitat types. Commercial forestry will likelycontinue in its current form due to many economical and social fac-tors. Thus, the main aim of conservation efforts of saproxylic beetlesshould focus on the refinement of the profound effects of commer-cial forestry and on respect for forest history and traditional forestmanagement.

Acknowledgements

We would like to thank V. Kubán for providing literature andnotes on an earlier draft of this manuscript and P. Svácha for pro-viding literature. We thank J. Mertlik, A. Sedlácek, C.-F. Lee, H.Bußler, V. Franc, A. A. Zaitsev and J. Ch. Vávra for contributinginformation, M. Boukal and L. Cízek for the fruitful discussions.The comments of J. Settele and an anonymous referee helped toimprove our manuscript. AJE corrected the English. This study wassupported by a grant from the IGA CZU, the project Monitoring ofPinus pumila (Pall.) Regel in the range of its natural distribution(www.pumila.cz) and by the CZ Ministry of the Environment (grantno. MSM 6293359101).

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