11° The Importance of Veteran Trees for Saproxylic Insects Juha Siitonen"* and Thomas Ranius* "Natural Resources Institute Finland, Vantaa, Finland: ?Department of Ecology, ‘Swedish University of Agricultural Sciences, Uppsala, Sweden 11.4. Introduction (Old toes ~ often referred to as ancient oF veteran — have aways atacted atention, but recently dhere as been aval of interest in them from an ecological an conservation pee spective. Ancient tees ae old individuals that have leat passed beyon maturity and often show features such ascavites or hallow trunks, Dark loss aver sections ofthe trunk and large {quantity of dead wood in the eanopy. The tem, ‘Veteran tree’ includes younger individuals that have developed simular characteristics a= a result of adverse growing cantons or in- jury (Woodland Trust, 2055; Lonsdale, 2013). Neteran tres are defined as being of interest biologically culturally or aesthetically because oftheir age, size or condition (Read, 200) ‘Allarge old tree has been described as an arboreal megalopolis for saproxylic species (Speight, 1989). We therfore address the following questions: +) What i it about those trooe that ie {important ~ in terms of their structure land where they occur for invertebrates? + Which invertebrate taxa inhabit these + How are invertebrate species occur rence pattems and longterm persistence “Ema: ha sitonenmetiat affected by the abundance, quality and Spatial distribution of old trees? + What practices should be encouraged to further the conservation of saproxylic species associated with ole tees? 11.2. What Are Saproxylic Species? Speight (1889) defined saproxylic species as those that depend, during some part of their lifecycle, upon the dead or dying wood of moribund or dead trees (standing or fallen), upon wood-inhabiting fungi, or upon the presence of ether saproxylic species However, ‘dead wood occurs fegulary in mature living ‘wees, and heartood decay and hollowing is normal part of the life cjele of almast all Tongived deciduous tres (Alexander, 2008), Hollow trees are not necessarily moribund, So the ecological definition of saproxylic species should also include species that depend, luring come part of tie Ife eyes, upon wounded or decaying woody material From, living tres (StoKand eal, 2012), ‘Studies of sub-fossl insect remains (@vckland_ and Dinnin, 1983; Whitehouse, 2006; Olsson and Lemdabl, 2008) suggest that ‘many saproxylic species that are nov rare or ‘©CAE Itratonal 2015, Euape's Changing Woods and Fares: From Wood 0 to Managed Landscapes (eds Ks. Katy and, Wane)Europe’s Changing Woods and Forests ~From Wildwood to Managed Landscapes Edited by Keith J. Kirby Department of Plant Sciences, Oxford, UK and Charles Watkins School of Geography, Nottingham, UK Ekologibiblioteket @_.x Ek (CABIis a ading name of CAB Intemational capt cant Noswworthy Way 35 Chauncy Street Wallingford Site 1002 Oxfordshire OX108DE Boston, MAQ2IIL UK USA, ‘Tel +44 (1491 32401 ‘Tel: +1 800552 3085 (tol free) Fae +44 ()1491 823508 Exmail cabi-naoaesbiorg Exmail:infod@sabiorg Website: worwcab.org ‘© CAB Internation 2015, All rghts reserved. No part ofthis publication smay be reproduced in any form or by any means, electronically, mechanically, by photocopying, cording or otherwise, without the prior permission of {he copyright owners. catalogue recond for this bookis available fom the British Library London, UK. Libraty of Congress Cataloging-in-Publication Data Europe's changing woods and forests: from wildwood tomanaged landscapes / edits: Keith J. Kirby, Department of Plant Sciences, Oxford, ‘Charles Watkins, choo! of Geography, Notingham pages. Includes bibliographical references and inde, ISBN 978-17806487-3 (hbk: alk paper) 1. Forest ecology~Europe- History. 2. Forest management-Earope-Listry. 3, Forests and forestry Europe-History. I Kithy KJ, editor SD177.858 2015 634.72004-de23 2014081667 ISBN-15:978 1780643973 Commissioning editors: Vicki Bonham and Nicki Dennis Eitoral assistant Emma McCann Production editors Lauren Povey and Emma Ross “Typeset by SP, Pondichery, India Printed and bound in the UK by CPI Group (UK) Ltd, Croydon, CRO4YY.11° The Importance of Veteran Trees for Saproxylic Insects ‘Juha Siltonen’* and Thomas Ranius* ‘Natural Resources Insitute Finland, Vantaa, Finland: “Department of Ecology, ‘Swedish University of Agricultural Sciences, Uppsala, Sweden 11.4 Introduction (1d trees ~ often refered to as ancient of vyeleran ~ have always atracted attention, but recently there has been a nevival of interest in them from an ecologies] and conservation per- spective. Ancient trees are eld individuals that have larly passed beyond maturity and often shove fentures such as cvitisorholiow trunks, bark loss aver sections ofthe trunk anda lage ‘quantity of dend wood in th canopy The term “Yeteran tee” includes younger individuals that have develope similar characteristics as a result of adverse growing coalitions or in jury (Woodland Test, 2008; Lonsdale, 2013) eteran trees are defined as being of interest iological culturally oe aesthetically because ‘oftheie age, size or condition (Read, 200), “lange old tree has been described as an arboreal megalopolis for saproxylic species (Speight, 1989). We therefore address the falloving questions: What is i about Use trees that is Important = in terms of their structure and where they occur for invertebrates? 1+ Which invertebrate txa inhabit these + How are invertebrave species occur ‘rence palterns and long-term persistence Emak joha torent affected by the abundance, quality and Spatial distribution of old tees? + What practices should be encouraged to further the conservation of saproxylic species associated with old tees? 14.2. What Are Saproxylic Species? ‘Speight (1980) defined saproxylic species as those that depend, during some part of their lifecycle, upon the dead or dying wood of ‘moribund oe dead trees (standing, o fallen), lupon wooxtinhabiting fungi, oF upon the presence of other saproxylic species. However, flead wood occurs regulaly in mature living tees and heartwood decay and hollowing is normal part of the life cjcle of almast all Tonglived deciduous trees (Alexander, 2008) hollow tres are not necessarily moribund. So the ecological definition of saproxylic species should also include species that depend, ‘ning come part of their Ile cycles, upon wounded or decaying woody material from living tees (Stokland ea, 2012) ‘Studies of subfossil insect remains (Buckland and Dinnin, 1993; Whitehouse, 2006; Olsson and Lemahl, 200) suggest that ‘many saproxylic species that are now rare or ‘©.CAEiarnatona! 2015, Europe's Changing Woods and Fass: Fram Wdwood 0 {eManeged Landscapes (ods Ks. Kiby and. Wars)Imponanc of Veteran Tees for Saproxye Insects wa regionally extinct were common across the Tandecape inthe pre-Neclithic period. Some of the fist species te disappeat, such as Rhyses sulcatus and Prstomis mandiuleris, are inhabitants of old-growth forests with ‘minimal human influence. Others have su vived inthe cultural habitats created by ha- san in particular in wood:-pastures (Hartel tal, Chapter 5), but awe now endangered or extinct regionally or acnss western Europe as ‘whole, Inthe Europen Rad List of Saree Bales (Nieto and Alexander, 2010), 1% of bout 450 species aseised were considered ‘heatened inal of Europe, while atthe Euro- pean Union (EU) level, 14% were threatened, primarily through habitat los in relation to Jogging and wood harvesting and the decline of veteran trees throughout the landscape 113. Veteran Trees in Past and Present Landscapes Veteran tees ocurin diferent types ofnatural habitats including closed old-growth forests. Under the original natural conditions, the density of wide-cowned veteran toes would have been higher in store open woodland where there was less competition from young {growth for example, nwa forests where only patches of higher ground are suitable for tee growth, rocky catzrops and mountain slopes in lod pain forests where regular dis- farances keep the stands open, and in areas that are heavily grazed by lange herbivores. ‘Agriculture and ax increasing impact of humans spread into Euzope starting from the Neolithic period some 8000 years ago. BY about 3000 years ago, natural forests had Deen cleared from most cultivable lowland areas all over Enope (Kaplan ef al, 2008; Kirby and Watkins, chapter 4). However, traditional land-use forms, sach 35 g732ing, and the coppicing anc pollarding of tees, ‘reated new types of cutiral woodland habi- tats that were to some degree analogous to natural habitats and alowed the survival of many saproxylic species iltonen, 2012). ‘OF particular impertance were the vari- ou forms of wood-pastures (Bergmeiere a, 2010; Hartel tal, Chapter 5; Plate 5). These range from forests grated only occasionally by cattle to permanently grazed grasslands ‘with scattered toes (Rackham, 1988), such as are often found in parks. From the medieval period onvvard, parks were commonly estab lished around castles and manor houses Many ofthe oldest parks were originally used. for hunting, but Inter landscaped to create more altractive setting, sometimes incorpo ating veteran tees already on thesite Fetches, Chapter 9). Postmedieval landscape parks that include older features host much richer saproxylic beetle faunas than parks lacking such continsity (Harding and Alexander, 1994, Alexander, 1998), but this richness also de pends on whether the parks contain some Semi-natural vegetation or are lose to poten fal source areas. Hellow lime trees (Tita spp) in old manor park in southern Sweden have been found to host as many specialist and rect-listed saproxylic beetle species a similar "tees in open wond- pasture or overgrown for- ser wood-pastures lonsel, 2012) In Estonia, ‘old manor parks have about thre timesas high ' basal aren of lage (>40 cm) trees as mature forest patches in the surrounding landscape; the proportion of brosdleaved tres (Ta spp., (Quercus rabur and Fraxinusexclsin) is also bout tree times that found in the forests (Cahmas and Lire, 2013). ‘Other cultural habitat types important for saproxylic species include orchards and venus, the latter able toot saproxylic spe- ‘lists such asthe hermit betle (Osmaderma ‘remita) (Pate 5) and the rose chafer species Prataetia marmorala(Oleksa el, 2006, 2013). Hrdgerows form extensive networks in old agricultural landscapes (Baudry etal, 2000) fd the large pollard tres in them are i portant habitats for saproxylic species such a5, the hermit etl (Dubois a, 2008). Veteran frees may also be found along fell margins, riverbanks and old vad, 11.4 Important Structures ‘and Associated Species in Old Trees 11441 Microhabitat diversity, “The importance of old trees for saproxylic species lies in the large number and variety of special structures they have in comparison42 4 Sito and. Rane ‘with younger tres. Microhabitats (distinct parts of a tree that host different species Assemblages) include cavities with wood mold, waterilled rot hoes, dead bark, ex posed wood, sap flows fruiting bodies and iycelia of fungi, dead branches and dead 0s Sltonen, 2012) (Fig. 11), ‘On completely dead tees, these micro- habitats gradually disappear a6 a result of de- composition. Ip lemperte forests the Hine required fora dead medium-sized tee to de~ ‘compose tnt ite no longer a visible stu ture is only a few decades, Most saproxylic species ae able to use dead trees only during 4 particular part of the decay. succession (StoKland and sitonen, 2012) and so the wine ‘dow of ime during which a dead ree cons tutes suitable habitat may be only fev years to a few decades, Speces feeding om fresh poe tive tres ony ring the fist summer following te death, after which they need to colonize new hosts, In contrast, deadwood microhabitats in old living tres can last for much longer periods; cavities filled with ‘wood mould, which start develop in tees that have reached their maturity, may then persist for centuries within the lve tre. Tree species, grovth rate, age and diam- ctor ofthe tee, a well as the environment in ‘which the te is growing all aft the number ff microhabitats per tee (Winter and Moller, 2008; Halland Bunce 2011; Vidot otal. 2011: Regnery tal, 203), Wide-rowned toes grove jing. in open conditions swally support a higher number of microhabitats than other ‘wise similar trees growing in closed forest. 11.42 Tree cavities and thelr Invertebrates (Cavities are formed primarily by the action ‘of fungi that decay the dead heartwood in mature living tees (heat rots). Heartwood Fig, 11. Mirohabtats na votoran roo. A dea sun-exposed limb, woodpecker hols, dead ‘tached banchos nthe eaopy,a branch cay: E polpore uty bodes , a turkey, G, ‘fan ranch on te rours Ha baeal cay | an open wound sounded by elu sue =p ‘natn: K dad oot nt sl (Drawing by Juha Sitonen}Importance of Vtoran Troos for Saprone Insects v3 forms the inner core ofthe trunk and larger branches. Hear-rot fungi do not grow in the functional saprsood forming the outer wood layer, so only the inner tunk will become soft as a result of deca Damage t, oF breakage Of lange branches provides saproxylic iver tebraes with acest the exposed heartwood, ‘Cavities then devaop through the combined section of hear-o! lung invertebrates and the breadown of tha dacaying wood Tnastudy in Sweden, less than 1% of oak (Q.robur tees up 100 years old had cavities. By the age of 200-30 years, about 50% of the trees bore cavities, while tees older than 400 years were al hollow. Cavities form earlier in ast growing res ‘han in slow-growing tres, probably because fst growing tees have big ier branches that she ealier (Ranius eta, 2000), Some shore ved broadleaved species, such a8 lls (Sli spp) poplars (Populns spp), maples (Ace spp) and horse chestnut (tsculs hippocitane) start to become hole Tow at much younger ages. Asa cavity develops and becomes large, it becomes structurally ‘more complex anithe diversity oftheinverte- brates if supports increases, Wood mould stars toaccumilatin the bottom ofthe cavity nd isthe principal substrate of invertebrates living in cavities. ood mould isa mixture of decaying wood, dead leaves and debris falling Ino the cavity, the residues of bird or mammal nests and dzoppings, an fungi al fasspro- luce by the laveebrates themselves. The cavity mcald fauna is taxonomic ‘very diverse, and across Europe includes sev feral thousand species. Many’ are generalists thatarealsofoundin other dead-sood micro- habitats, bur othe are specialists (Sitonen, 20128), The most important groups are beetles (Coleoptera), gna and fies (Diptera) and theiedepenclent pans wasps (Hymenoptera) Ants (Hymenoptera: Formicidae), springtails (Cllemola}, mites (Acar), spiders (Araneae), harvestmen (Opiliones), pseudoscorpions (Peeudescorpicnida) woodhice (sopoda),centi- pds (Chilopoda)and milipedes (Diplopeda) re also frequently found. This species assem- Dlage forms a conplex food veeb of wood mould consumers and fangivores their spe~ alized. predator and parasitoid species, and scavengers feeding on dead insects. Most beetle species associated with tee cavities seem to prefer th seddish brown, crumbly mold typical of, for example, oaks decayed bythe angus aciporaesuphanes. The moist, dark wood mould typical of several other broadleaved toe species, eg. elms (Uimus spp) ashes (Frainus spp.) and horse chest- ‘ul, seems to be favoured by dipterans (Ane dlerson, 1999; Alexander 2002) ‘Large chafers Scarabaeidae, Cetoninae) suchas the hermit beetle, Protea and Gnor- rue pp. are among the most emblematic dnd functionally important species inhabiting ‘cavities, Therlarvae tunnel into the decaying ‘vals ofthe cavity, thereby expanding it arc {eating lange amounts of nittogen-eniched frass (Jonsson ta, 2008) Ober typical beetle families found in cavities include comb-cawed betes (lleeidae, click betes (later), and darkling betes (Tenebrionidae) (Kelner~ Pillault, 1974; Martin, 1989; Ranius and Jansson, 2000) "Among the specialist species, the hermit beetle is the most intensively studied and hence i commonly used here as an example. However there ae other cavity specialist spe ces thatare much rarer, which may therefore bbe more demanding in thelr habitat require= ‘mens, Even the hermit beetle may vary in its esuirements; molaclar geneti studies in cate that it isa complex of atleast four distinct species (Audis eal, 2009). veia curs in western Europe wp to southern Sweden in the north O, ambia is distributed through fensiem Furope to central Russia and up to Southern Finland; O. cistine is confined to Sicily; and 0. tsa is found in Greece and the European parts of Turkey. The taxonomic rank of the southem Taian hermit beetles (CO. talc) remnins unresolved. This speci- ‘ation may have occured in temperate forest Tefugia in the Kalian and Ballan peninsulas land Sicily before and during the Pleistocene (Kirby and Watkins, Chapter 3), Similar pat- tems of genetic diversity and eryptie species ‘might be foun in other saproxsic specs thus ‘emphasizing the desirabty of preserving Species throughout their distabution, 41.43. Other mlorohabitats Sap flows can also be long lasting features of veteran tees, They may be cused ether by awe 4. Sitonen and, Ranios conic infection of anaerobic bacteria, or by repeated damage to hark resulting from in- femal fractures, Many saproxslicspecies feed ‘on them but there aresome that specialize on sch floes, particulary among the Diptera. Dead branches host specialized fang) and the insets feeding on them, Individal branches dlecay andl fall opi, but within the coven fof an old tee new dead branches appear regularly. SaayUasonmual uiting bodice fof important heart-ot fungi of broadleaved, trees such aL sulpharews in oaks and Palypo- ‘rus spuansns in elms and ashes, appear and decay within a coupl: of months, but these tung produce new faiting bodies on thet Ist tee each yeat for decades. Many saproxylic species depend cn the conditions created by ther species. Galleries ‘made by wood-and bark-feeding insets pro- vide microhhitas fora wide array of assoc fated species to the pint tha some, such as the great Capricorn teetle (Cerambyr certo), can be considered to ke ecosystem enginces ‘The leva tunnels ané adult exit hoes ofthis beetle are sed by ater beetle species and, by weakening the te, they open the way’ to ‘other saproxylic species (Buse eta, 2008). Emergence holes of cod boring beetles, such as cerambycidsand arobids, are used by soi- tary bees and wasps that are unable to dig out their on nesting holes Diferent species select holes of diferent sizes, orn diferent positions and under different conditions, in a manner thatis analogous to tat of hole-nestng binds, Dut ata miniature scale. Social insects that Did their nests in cavities provide habitats for a range of commensal such as the large rove beetle species, Veins dattus, which i found in hornet (Visya cabo) nest, 41.9. Effects of Environmental Factors on the Invertebrate Fauna 11.54 Effects of tree characteristics on spocies assemblages, ‘The habitat requirements of saproxylic in sects have been described from field observ tions (Palm, 1958) Sone are generalists ancl, can occur as larvae in any deciduous tree species, but most favour one or a few tee specs anda limited number are very stricted in their host choice (lonsell ef aly 1998; Stokland, 2012). All tre species host at least some specialist saproxylic species, but in nor- thern Europe oaks (Q. bi, Q. petra) are ‘he most important both in tems of total spe cies richness, umber of specialist species and number of red-listed species. Oaks tend to frow langer and older than moet othor tee species and therefore have more of the micro- Iabitats associated vith old trees for longer periods “Thete ate only afew quantitative studies ‘on host-tree associations of saproxylic spe- cies. Milberg al. 2014) compared saproxylic bel faunas betcen four tre pecs (ab, ‘Acer pltanoides,F.excelsor and T.cordata) and found thatthe great majority of species were not specialized to tee species. However, some beetle species clearly preferred a particular host-ree species, and more species wer aso- dated with Q. ror than with any of the other taee species studied, [Eage trees generally hacbour more sap- roxyle species than smaller ones. In a study of nine hollow-specaist invertebrates, six hha ahigher occurrence probability ina given volume of wood mould in large hollow tees (Ganius ta, 2071). lagge tree contains, on average, a langer number of more varied ticrohabitats, which may be larger and more complex because they have had a longer time to develop. Also, a lage trunk has a more ‘stable intemal microclimate than thin tuk, land a larger tree has existed for longer and provides more opportunity for species clon leation, The relative importance ofthese fac tors in driving the positive tee size-species richness relationship isnot known. [Exposure to sun is important fr saprox- sic fauna (Palm, 1959), and more saproxylic Insect species are report in tres that re ‘exposed to sun (Ranius and Jansson, 2000, ‘oda et a, 2009; Koch Widerberg etal, 2012; Horsk and Reb), 2013). Greater exposure to ‘sun gives rise to higher temperatures, so less time is needed for larval development: con- versely, exposure to sun makes the wood drier, which may be a limiting factor forthe ‘occurrence of speces,espealy in epions with warm and dy climate. Asa result, speciesImportance af Veteran Trees or Saproxye Insects 45 may be more abundant in sun-exposed te towards their norther distribution limit, but lack such a pattem father south (Chiari ff aly 2012), There could be an interaction ‘vith how and where tres grow as well For {stance beech (Fagus eatin often occurs in ‘more denseandshadec stands han oak, which right lead to the invertebrates associated ‘with beech being beter adapted to shaded Stuslidune (Gitenfe and Daranoveski, 1992). Even im beesh-dominated forests though, more species appear to be associ- ated with warm than with cool sites (Lachat etal, 2012), 11.52 Effects of surrounding landscape on species assemblages ‘Trees die and so the rcritment of new! co horts of veteran trees and their colonization by saproxylic invertebrates, must take place ‘The probability ofa species finding the right habitat ina tre andi long-term persistence at landscape seale depend not only on the ‘Characteristics of individual tees, but also on the density of suitable tees inthe surrourel ing landscape (Gvesleup-Thygeson etal, 2010; Gatmark ea, 2011; Rane al, 2011; Bergman ot al, 2012). At the landscape level, local extinctions from trees must be balanced by the colonization of new trees. ‘Ametapopulationstudy was arid outin southem Swesen 08 herait Beetles living in hallow oaks in areas with oak wood-pastures that ae a few klometes apart (Rani, 2007). Thelarval development of his species only cur in the cavities of dt oaks, each re as fffectvely a habitat pztch that potently hare bu a local populition. The probability of fccurrence ofa population increased with the ‘umber of aaks with avites containing wood ‘mould. (potential dispersal sures) inthe surroundings, withthe strongest relationship ‘blaine when including only tres within ius of 200 m (Rani ca, 2011), Dispersal was fonsiderably more common within this dis- tance than farther away. The dispersal ate anc range were assessed by capture-recapure and telemetry (Plate 5), wich confirmed that most dispersal movements oy the beetle tok place to the nesrest tee and that mast individuals remained in their natal tree throughout their time (Hedin a, 208) Th the Swedish study, the population size of hermit beetle per tree was relatively stablebetween years in comparison with tat ‘of many other insects. However, population Size varied a lot between trees because the ‘quality of the hollow trees differed widely Inainly as a result of differences in the vol lume of wood mould (Ranive, 2007) The tinction risk was higher in toes with stall amounts of wood mould than in tees with high-quality cavities. Cavities usually de~ ‘velop in oaks when they are about 200-300 ‘years old (Ranius eta, 2008), and after that fhe oaks may remain atleast for another 100, years of mote (Fig 112) in the study area, the oldest oaks were up to 500 years ol “Therefore, at many sites species persistence relies heavily om a fe hollow tres of very high quality, and the overall population is vulnerable to extinction if these tees aze lost, ordeterorate Tn ely, similar study of hermit beetles revealed dispersal distances about ten times longer than in Sweden and many more dis persal events (Char ea, 23a). Thus, mot population dynamics may vary in diferent parts of the distribution ofa species. This may Sepend on temperature differences although ‘complicating factor is that the allan hermit beetles may be a different species or subspe les:’0 talcum. Dispersal biology isa key facto for spe- cies persistence and occurrence pattens in Tanelcapes with various levels of abit fag ‘meniaon, Species adapted to naturally shor ‘duration habitats are expected to have evolved | more opportunistic and wider ranging dis- persal strategy than species adapted to more Stable and long-duration habitats. Foe these, a better strategy may be to remain at the Fame habitat patch and spend fewer re sources (or fake risks) on dispersal (Travis and Dytham, 199) ‘Many common deadwood microhabitats (eg. igs and small dead branches) are «ephemeral and species adapted to such condi tions are strong colonizers. Spies developing in recently dead tees, such as bark beetes, tendalso to have good dispersal ability. Incon- Irast, species associated with longer lating6 4: Sitonen an. Ranios Fig. 112. Storoogen the stork oak, Quercus ‘ebur In Segerspis Horio, Denmark: a). & [ogra by Gur rm "880 poraying the giant holo oak stag nan 2560 cary 1800; () pores ne sam 180 years later n a photopaph taken by Ole Marino 1974, hen shows beech regrowth follning te coseation of yazng. At ho tino the ota was taken the oe was abou 0-200, oor and wa el ae, ard specalizea proxyie boot spaces, such as bestles Amped cardinals ard A. ort andthe ‘aring beste Tenebrio coecus, wer sting {hanallw (Ole Marin, posanal communica}. ‘The tee Is now dea siceohahitats such as avis in ol ving trees, appenr to he poorer colonizers (NUSson and Baranowski, 197; Hedin eal, 2008). They fare, consequent kly tobe particularly vul> erable tothe habitat Fagmentation that has been caused by the severe declines in the abundance of veteran tes across Europe over recent centuries, There may even have been recent selection against strong dispersal ten- lence in species occupying rare and widely sdsperse niches 11.53 Catering for the needs of the ‘adults as wel as the larvae Many species that are saproxylic in the Iar- val stage depend on nectar a8 adults. Sitch Species include, for example, many long> horn beetles (Cerambycidae), jewel beetles Guprestidae) and hoverdes (Syxphidac). “Therefore, the landscape needs to contain an adequate supply of flowers rough shrubs such as hawthom (Crataegus spp.) or herbs ‘uch as umbels, thistles or composites 1154 Survey methods ‘A problem with assessing the conservation ‘needs of saproxylicspecies has been develop ing survey methods with known efficiencies, (A traditional method of sampling inverte- brates living in caviles is to spread wood mould on a shite canvas, and to search for adult individual, fragments and larvae. This ‘works vrell for some larger species, sch a8 the hermit beetle and the click beetle species Crepidophorus muliltus, in which fragments remain identifiable for many years. Other ‘conventional methods inelade Might intecep- tion traps placed inside hollow trunks o out- side cavities, pitfall traps buried inthe wood mould, and traps bated with mould grass and animal bones. ll ofthese methods are Selectively efficient for a particular set of spe- cies and thus complement one other (Ranius and Jansson, 2002). ‘More novel sampling techniques have been developed recently, partly prompted by research on the saproxylic species included in the Anmexes ofthe 1992 European Union Habitat and Species Directive. A new type of emergence trap has made it easier to survey for the volt cick beetle (Linonises ilies) (Gouixand Brustl, 2012). Dispersal of the her~ mit beetle has been studied using marking- release-recaptre stadies(Ranius and Hedin, 2001) and radiotelemetry (Dubois and Vignon, 2008; Heine a, 2008; Chae etal, 20133), “Another new branch of research wiizes ‘pheromones. The pheromanecf the hermitboctie {-decalactone) ean be synthetically produced (Larsson ef a, 2005; Svensson eal, 2009) andEe Importance of Veteran Tres for Seponyic Insets 1 SO ated ith thispheromona can bemeore flit etting te presence of thee Siebert than conventoralmethods- Pero bce tapping mcs have been used In otis of tne fccurancepaters an sbute Siesol the ert ee at heat patch es well an in sce of Ste diapers thd Steno, 2011 Svensson a, Soitchun eta 200. The pheromone an Ein actas on nraraet Gr hr re are ese atone wih hall tee oro Zipline lage ik beetle spose Bt fo tigen vhichisa predator of heer ie {i tlrcon and Sexson, 21) Th pecs Sethe cfc survey. Some specs ae eat 10 detect fom their gleresThe eat Caprese proiuers unmisakebl, lage ext hos on EB cals tr i inbubite, Thene indict the umber of invidals that have emerged fram pra ec whih atop {oases popson ss fhe specie thd incl of octets an srvronmenial eins on brewing es (Deseo 207 ete, 201) ‘he mt eet pec eles Aoownigecf he mae fe op ses tl sina ye opses Te Senate of much pst oi sk io undeestnating he mio co Pied sis. Modeling Oe habitat egeieenis Uiapeds based on nenive sping ina Ste are, then collating dts on habit avalbity eequeny aggregation and us iyo pont Hot wo) over ete egos hy dow more consent prediction be osinhated by apes Gata, 201) 11.8 Current Situation in Europe Large od tres area globally declining habi- tat feature (Lindenmnayer etal, 2012). At the ‘current rate of os, nos ofthe wood-pasture systems that were analysed by Gibbons etl (@ins), including southern Spanish holm oak (Ge) wood-pastires, would lose all of hele veteran es within the next 90-180 years, The mortality tate of existing tees far exceeds the recruitment of new veteran tees, Existing management recommendations fo- fusing.on recruitment vill not reverse this teend. The more critical factor i the curent mortality rate of veteran tres, which should bekept t around 05% per year (Gibbons a, 2008). Veteran oak ros often have a mortality of about 1% a year and if they are situated in high-density forests the morality becomes ‘even higher (Drobyshev ea, 200). “The threats to cata habitats and their veteran toes fall into two broad categories: intensification of land use and the cessation of traditional management. The intensifict- tion of land use results in the fling of trees valuable forthe saproxylic insect fauna or in increased tree mortality: the cessation of trad- itional management such a grazing leads to regrowth and closed-canopy thickets. Old trees that used to grow in open conditions ‘cannot adapt toshading by younger trees and tend to die beensse of competition (Hartel tal, Chapter 3), These sues can be ilus- trated bythe case study from southem Sweden, reported in Box ILL 11.7 Howto Preserve the Specialized ‘Saproxylic Species? 41.7.4 Management for increasing ‘habitat amount and quality “The priority inthe short tem isthe survival of existing Veleran tres: Where traditional agri- Culture maintained the old trees and this has ‘now been abandoned, the maintenance of tee ‘Quality is likely to require the reinstatement of {ctive management. Haymaking and grazing by calle should be revived to keep the sur- ‘oundings fee from competing young growth, Orbushes and young trees should be removed ‘mechanically. This removal must be done regularly iftis et fr toolong, the coaltions for tees as wel as for thelr asocated species, Dbecome unstable, which may increase tee ‘mortality. Ties traditionally managed by pol- Tarding should be e-pollarded if possible, as ppollartng prolongs thelifespan of trees (Read, 2ond) and sa increases the rate of eavity fore mation (Sebek ea, 2013) (Fg. 12). ‘Where there are mature tes o tres ap- prosching maturity, these can be encouragec, to develop into the next cohort of veteran8 4 Sitonen and. Rani Box 11.1, History of etn ake (Quercus spp) a southern Sweden, Lagos ss of veteran oaks in sue Sweden started more than 200 years ago (lasson and Nisoon, 2002), High-qually oak oer Became an impor strtoi resource or balding warshipsin | bt cntryanosk wes dear toe te propery of he satya voyaldecre in 1558, Dung the 17h century, lage bale Heet were consti, ahd the cloctve fling ot high-uaty oaks ld to.an almng option df ak stocks reponaly. An assessment of he inberresources was needed {orlonpemm planing, so somprehensve vents covering ene provinces were main the eaty ‘7302 in those vero, mature oaks wore caslied as beng of ood quay, pal decayed, 00 eespod ete wna, ovmushapan non mary ranches thas ervdl a elerene panto essessin th atbeequen changes. ost of ong ok oss the quay ange grow on meadows clos villages. As the al popu Ist gre these toes became an abstalefo the nonscabon fag. By he end of he 1th entry, del esermentagalst ak was widespread In 1785, a royal doce gave the peasants the Wht the land the nobly had had bot, but wh ene excopton - he dere excluded ‘night oe cak and bos In tesponso, the pensar cated fr anand survey to determine ‘thc tres wero self the navy and which could be rleases om sate contol. Dung he 1780s, ‘rosin wore surveyed and good tees marked wih a crown slp. ‘Desi the ban ona aks, the cling of ranches, ceberae damaging lees, leg! posal ‘tees snd alo laa fling based on exemptions wore widespread. When the naomi survey of ‘aks aerated nthe art 1820, te numberof aks it for raval use had been reduced by over ‘0% betwecn 1790 and 125, During the eat 1h century, an average of 7.00 aes a yar wore ‘ole logy, between 180 and 1895 at east ne aca Ralf mon mature oaks were cut legally, and ‘as many ff not mor) ex legal Te fling ban en oak was ay abolished in 1880, “The dramatic deaine mati oaks noble caused 29 egal and immediate deine inthe populabns of oakassocated sapronic species, and many reponal extincons. However. as he Colonization extincon prcassee of species esseiled wih veteran toes ae iow, the slow ds- “pparance of species hes continued and may sil be happening in landscapes where the rent ‘erat of potent hot tee tooo t alow lorem perstanoe. A sty onthe occurence pat {emf iehens and woodinnabling fang asrocod wih ol oaks n suthom Sweden showed Tt, tn 2 species sued, the ocourenos of 18 species was postive ltd oo curet dens of teks bun aden, the Sosurence of 11 species was expsied by ho storia dons of oaks in “890 (Rani et af, 2008) ‘noes, The formation of important microhabi- tats on these tees might be speeded up by slamaging them (Cavalli ana Mason, 2002), for ‘ample hy creating entrances fr decay fungi by dling holes, ehe praring of branches or the removal of patches of bark. Fungi have Also been inoculated int these injures, but it will take several decadesbefore its possible to fee the outcome, Also, rot all allows are of equal habitat quality Ranius, 2007), As yet, we now litle about how to manipulate tees 0 that they become ‘good! hollow trees. For some species ts possible to create atifcial substrates, In Sagland, tre trunks have been filled ith sss eaves and dead animals, and used asa breeding substrate by the rare violet click beet (Whitehead, 2003). (Other species, suchas the hermit bocleand the rose chafer Proactia martorata, have been brea in the laboratory in plistc containers with -savedust and leaves as substrate, Broeding 9pe- ‘Ges in the laboratory and the inoculation of Sites with tific substates could bea means ff producing individuals tobe used for trans- location totablesites where they donot car- renily occur This sort of study has never been Conducted in a systematic way, but as many species havea very fragmented distribution it ‘could bean efficient conservation measure for the preservation of individual species. Therele- vance of sich a method could increase if we ned to consider changes in the potential range of species sulting fram climate change 11.7.2. Management for securing ‘spatiotemporal continuity Conservation efforts are expensive, so which sites anal landscapes should be given prion?Importance of Veteran Tres for Saponyic Insects150 4 Sitonen and Ran 5 clase is relate to she dispersal biology of 11.8 Future Prospects the species concert; for beetles inhabiting hollow trees this can vary between 200 and Increasing public awareness of the import- 5000 m (Ranius ef aly 200 Bergman etal, ance of veteran trees to biodiversity is a key 2012), These distances do not diteety reflect issue for sympathetic management, There i the dispersal capacity of individual beetles, an increasing interest in ol trees, because of Dut they indicate the sale at which hosttee thei aesthetic, cultural and historical values. fonnectvity is important at the population Utilizing real examples ofthe often colourful level and intriguing creatures that are associated ‘ites that previously lauboused a lage with them adda another dimension to this umber of ol wees, but from which many and may, inthe Tong run lead to greater pro have now gone, may sill maintain more tection ofthese species and habitats, species than might be expected or that can swith other groups of onganisms there be maintained (without action) in the longs the uestion of how the saproxylic species fn. This surplus of species is often referred inhabiting old tres will respond to climate to as an ‘extinction debt. Positive correl- change. In principle, a warming. climate tions have, for insance, been found be- might benefit species atthe northern pars of tween the historical abundance of old tees their range. However, the effects of climate fand the current occurrences of lichens and change are likely to be mainly deleterious fungi (Ranius ef a, 2008), Sites and land- ‘This is because the morality of old tres can scapes with a high historical density of old be expected to increase as. result of more fr trees, where there issill a particulasly rich quent drought and storm events. A warmer fauna, should be a high prionty for restor- climate could aso enhance invasive pest and ation efforts to secire long-term species pathogen species that increase the mortality persistence ff veteran tees References [Albert J Plate, M. an Cie, (2012) Vert statisti an mleohabitat selection by the re ‘Capricorn beetle Conmy ed) (Coleopers:Cerambycise) ix open-grown, veteran oaks. Epa Journal of Enomoy108, 955-9. ‘Aes ENA. (1998 The inks been forest history a biodiversity: he invertebrate fun of ‘lent pasturewoordandsin Brita aod ie consereton ne Kisby KJ and Watkins C (a) The Ea Td History of Emap ores, CAB eternatoal, Walrond UK pp. 73-9. [loa RNA. (2M0 Te orate sing ond Decaying Tbe Brin and lad a Proviso “ttt Chess English Nature Research Reports 47, Engh Nature, eteborough UK. ‘lexan NA. Ci) Tee blogy and prone Coleoptera issues of deations andl conservation Tangtage Reed Loa Teta Vi 8 1-5 [Anderson FH. 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Ore Rate ere No.7 Peterborough UR: PE. an oman Tn?) Modeling Mab 2 TS iebuton of an nda eae cn ofr prongs comet Biogial Coneraton 137 Eat pn al at sean, 8.2) An endangee ngs ese Sn) vith ld oaks and its cm mn ne cone aaa eas) au Resse fo saponin Con a NATIT sts Boe dela on Me wt Reports 2 SE a rg Connroe dle Beet FO pe Garo Nasr Ba arvana Eg eee ee or ye ot Projection hae ow itep=Be-DO%CO. ca eacon paaceed 27 Novernet 204) can pane CN, Za A, Main, Aut Fand {g0utabitt ofamendangered Fela ena NER 99-07 Cee Cat tn pal Ga es, nd Rake © Oa Dispersal patterns nS Car mara vei Meieanean won I ‘Cortona Dior 6, 309-315 en A Ln tA and Care Ce ba ee 0 a itn, Cora Meenas rapa beter dang a eth, lf set Conseretine 17, 171-18 ont Besar Se nena Heese Kain Mr an Lan (2008) admit of leaks combining emp = ae ing approaches 0 suppor tel ‘espa none san. as of Foret ce 9 a om Ft ale of cain ot OT (Coleoptent oi, an Me tora Rr ce Le Tet anne ee, Yk ann Yoo Pa ea asc the ss GE, en Does pan bei One eta Sepa 9) (CSP Coe czar nce amcor Pin 9. rasan and Nason SG. 202) te aoa ent and young nobimen.Theerroneta se Pan Nepean ent Sree ESN etry 769-87 Cee ee 0) eels EE tl tha a gl Tf 1 11 Dash Cen rr J Maing, AD. ton Pe and ne Be Le eo tered os asta nae, eS ily 2138-1318 a at rane N00) How weimproe aan ices of nck Ae pata fw el cup Be poe alg end gee 262,257-2905- Emergence wa, new mei tae Ls ST (Coleopte: bo a eos res, Bier ond Contr 221 08 a om a Nr ten eee a 1 antares ean ine Eg uk Fat ln a 0-2. 1 eae NA C9) The seo TN lection a eval ing a Ae rnd, uh oral of re Natal 077 Poulan T2226, 1 eT an, Gand Sith 1. 2009 Resse sere aflying este assed by ay Bier and Comers 17, 63-684, ns teat (2013) Te chines feck ees scent pasture woodland benefits fom Ne Sure oun of et Coseetion 17, 30318 on park ono att fo spent ees eT and Csr, “9-60 en an ti, 8 (99) Stl aint oe proxi nvete aM. 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Plan Bowe, (od) rctedng ofthe Send Pour Confrence ‘Sip Bete English Nature Peterborough, UK pp. 1. iichones NI (206) The Holocene Bris and eh ncn forest fossil bet foun zplcatons for treat ory, Motiversty and faunal colazatin. Quateray Sine Reis 25, 17-178. Wines and Malle © 08) Micra n lomand beech forests as monitoring to formate terrain foes Ely ond Managment 25, 1251-1261, Woon! st i) Anco Te Cue Wi Are Ait, Veo and Othe rs of Spent? Ancient Pease olan hues Guntur, Ce svabae st hip: /weecodandtustorK/ mesa Thua4ss13 ppt 2014 ancien tee guide t-deutionspatbs35e9 7619001970334 5 18 (sce 27 November 213).