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This article isprotected rights by All copyright. reserved. 10.1111/1365 doi: lead todifferences version between this andthe ofPleasecite article Version Record. this as been and throughtypesetting, proofreadingwhich may thecopyediting, process, pagination This article has for accepted been publication restricts the best growth to open areas. In shade, saplings a tolerant but adults are only moderately so. This, however, it grows best where competition for space and sunlight is limited associated with woodland than any other British species. Like other British species, most shade 2 disease, history, and conservation. environment, structure and physiology, phenology, of the British Isles behaviour. The main topics are presented within the standard framework of the Crantz 1 Summary Europaea * † Running head: Sciences,School Life Keele of University, Peter A.Thomas Biological List Vasc. Pl. Br. ISLES* BIOLOGICAL FLORAOF BRITISH THE Article : . . Correspondence Accepted Nomenclature Article Sorbus tor This account

(Wild

and .

- tolerant member of the in the British Isles and as a result it is Wales, and F minalis S BiologicalFlora ervice lora lora theof British Sorbus present of vascular Isles (1992)

: distribution, habitat, communities, responses to biotic factors, responses to † author - 2745.1285

- is ) s information on all aspects of the biology of a n uncommon temperate and Mediterranean .

E that are relevant to understanding its ecological characteristics and mail n 7

o.

: 75, 28,24

follows Stace ( p.a.thomas@ , I mostly small tree (but can reach 33 m) native to lowland sles: Staffordshire ST5 5BG, ST5 UK Staffordshire

Sorbus torminalis and undergoneand buthas not review fullpeer combi keele.ac.uk 2010 floral and characters, her ned regions ) and, for non

nd with its

young

of mainland . It is low adults form a

- British species,British competitive

. Seedlings are shade

Biol more closely bivores and sapling

ability ogical Flora No.

Flora (L.) the

, bank, 286

This article isprotected rights by All copyright. reserved. identification: of long shoots less lobed and even more variable alternate, viscous with olive yearsbecoming c. 1981). 30 fromold(Harper scaly lateral, angled branches in reaching canopy tree clusii ( ServiceWild Tree biology, soils altitudinal distribu Key extent to which these losses will be offset by increases due to climate change is unknown. Britain through the size of most populations makes it susceptible southerly and 4. mammals. range, pollinated. Seed production is reliable only in warm years, especially at the edge of its number of other 3 limestone. wide range of soil showing torminalis Crataegus .

Accepted ArticleThis tree is very tolerant of short droughts Sorbus torminalis - - scarious margins words

green to brown

Roem., although even then . and Stipules narrow reproduction and 30 very variable

: - Europe despite its wide range of hi loss or 33 climatic limitation, communities, conservation, ecophysiology, geographical and .

lowland distribution. It faces no particular Torminalis Torminalis clusii broadly ovate m

of open coppiced areas Sorbus display .

tion, germination, herbivory, mycorrhiza, parasites and diseases, reproductive reaction

Suckers profusely. Crown narrow or Che

is a sexual, diplo . of

Buds ovoid and

L., in shape

temperate species to form , embryo dormancy but most wil linear, glandular ciliate

quers/Chequer seed viability is low.

extensive growth only when released. full sparse white hairs Pyrus torminalis (pH 3.5 , 5. sun 5 :

(Roem.) Roberts - leaves of short shoots are simple, 10.0(11) x (4.5)5.0

forests to light - 8.0) but obtuse i d, non . . As a consequence it appears to be declining throughout Barkgre microspecies. The hermaphrodite are p tree ,

grows

3 - , rarely slightly tomentose (L.) Ehrh. but only moderately tolerant of frost, hence its . apomictic species to habitat loss and fragmentation

- . storical uses and the high value of its timber 6 mm long The are , reaching 1 y

- & Phipps best on green to dark brown on young browngreen on todark stems, , - and more rarely 10.5(12) cm when Shoots , . Aria torminalis individual l germinate the first spring after falling. Leaves of short shoots are used for Sorbus torminalis ,

calcareous

bright green to yellowish

shade ) is 2

initially tomentose, later glabrous, p mm long rimarily dispersed by carnivorous

that has hybridised with a usually Sorbus torminalis d , 0.8

broad but r threats

a clay -

1.3

co , falling early. otherwise glabrous and ound, ovate a

ly

(L.)Beck. subordinate - (1.5) s dominant

although the small

ovoid, deeply and thin soils over (L.) Crantz , particularly

times as

, tolerates a , with canopy tree Torminaria Torminaria bud scales , sub Leaves

rimarily long as many lobed;

- .

The

This article isprotected rights by All copyright. reserved. torminalis apomictic 2005). Sorbus torminalis large number of small sclerids (70 μ the ski the base white rapidly brownfruit. uponremovalfrom turningdark but (23%) with dense hairs an deciduous during ripening lenticels brightestripe: are before they in size within the same corymb (Bednorz 2007a); mm by a fleshy genus up a few reddish S across, calyx and stalks very woolly. Vivid teeth; petiole broadly cuneate; finely serrate with small strongly asymmetric above, greyish or almost so divergent (15)20 wide epals

Accepted Article. can have u

Two carpels each with 1 long

with autumn colour of reds and ye n (c. 400 per fruit), a Sorbus

- narrowly triangular to deltate, acuminate, sparsely to densely tomentose, 60(100)% to the midrib at the centre of the leaf) InIsles,British the Sorbus torminalis Flowers h

, a few as long as the petals , , obovoid, 7

with species shows great inleaf variation morphology range its across - , long petioles 9 on the underside - long and slender

brown glands on (Pietzarka p to triangular acute - green below large few veins ermaphrodite;

, also has six out of a total ofaroundout of 40native longe ( . the

Seeds resembleapple pips, ;

5 , r than wide

density is the highest in any (4 d 7% hairless (Bednorz 2007a). -

Lehmann & Roloff

is m 10 flavonoids notfoundinother European and the distinctive brown fruits with a large number of lenticels in ; multi Sorbus torminalis Sorbus -

, - 5 lateral pairs) leaf apex acuminate, , any remaining hairs restricted to the veins, glossy dark green

2 ovules; i 2 (20) (15)20 to acuminate initially reddish - ost readily identified the 3 styles i nflorescence - layered

fruits in a margins llow. m in diameter) around the seed (Aldasoro

(except smallest fruits which are - . 50(58)

In , most fruits (c. 70%) without hairs, some nferior free to base or

cu , .

lobes divided ≤half way to midrib ticle and the low water content (49%) caused by the

Fl sparsely tomentose when young Petals 4

corymb ( mm long (Price & Rich 2007;

a

owers 2009). - is oneof yellow later lax leaf base

6 corymb - from - creamy white , 9 x 4

7 x 7 x F Sorbus Sorbus Oršanić in contrast to all other members all otherof to in contrast the joined half to three quarters ruit , only the lowest two lobes are distinctivel 3 other - The f - 8 mm

a pome with the 2 carpels enclosed rounded, truncate or cordate, rarely 4 mm, obov 4 mm, of turning

hollow hollow

sp four sexual species (Rich , forward pointing, et al 30 Sorbus ruits are n ecies; , curved, blunt, round - 50 flowers, , . 2009)

10 brown Sorbus at unusualbeing apex, in sub

Aldasoro -

but by 13 mm at , ormally globose) e, flattened, initially . F

the wing diploid only with numerous

Rich species (McAllister et al

ruits becoming up to 20 cm

et al (but variably cut in two forms et al sometimes with . 2010)

et al diameter (8 appressed , single ,

. 1998) - of the way non very variable ) like lobes at . 1998 12 . 2010). , - -

c. 12 glabrous 18 - .

Sorbus seed . . ( small ) 20 ; y

) ed

This article isprotected rights by All copyright. reserved. in and M Lloyd(1981),and (1977). (1974) Wilmott (1977), Pigott ‘running accountsroots’. Regional a number species, including ofuseful fruit andits wood(Roper 1993) except al Westmorland, the WealdValley andthe Wye Sorbus torminalis altitudinal I. Geographicaland distribution Kollmann 2004a). open outcrops, tends tooccurinancient wood commonly 2012) leaves of fruit observedatripening. the time stresses thatthe primarily alsofruit shape shape but and onleaf (1960) lobes completely separated lobes caucasica et al 1997), and forma tomentose below atas correctly maturity(possibly forma known appear widespread tobe . 2010). editerranean zone Acceptednorthwest Article northern Irannorthern . 2010). , deciduous forests ,

and forma nativerange inawide of as anuncommon intr used the variability thespecies toidentify within Sorbus torminalis The Sorbus torminalis and often most

It is Diapulis native Two other have forms been largely absent from absent largely although natural its probablyBay northernis (Rich around limit Morecambe variability of formsis such thatidentification of requiresalarge number

perincisa .

pinnatifida

is In extends it south the distribution of distribution

(= s in the scattered

of Europe of

S. orientalis S.

and at theand e : forma : as ascattered tree

is strongly with associated is widely lower elevation throughout distributed west

( (Kov Borb oduction.

(Borb throughout

, (Fig

. E (Fig semitorminalis and easternSyriaLebanon into and inthe woods, scrub andwoods, scrub anda 1997; Aldaroso 1997;Aldaroso anda ás S

arly (e.g. planters 1679) Cook

Scotland, Ireland, Scotland, . dges of rides,dges clearings ofor

Schönbeck ás) ás) that take these plantings accountthat take into these

. & Fekete& torminalis

. 1)

2)

S. torminalis

K ,

, extending eastwards into continuing á separated basedseparated onextremesof shape: leaf forma into into rpáti with lowest leaf pinnae cut to withlowest leafrpáti pinnaecut to the lowlands of the lowlands l and on calcareousand calcareous on rocky soils, in

) B ) the Iberianthe peninsula areas whereareas light -

has been Temesy (Borbás) Jávorka withleavesfinely leaf indumentum,mostly leaf eck

hedgerows onclay mostly or limestone , as away ‘ of north ,

whose woodland woodland the Islethe ofMan et al ) ,

obscured

35 forms 35 w

which

Wales andWales England ards to . 2004;Bednorz 2006). leaves forest margins (Rasmussen forest margins

in Britain (Pellicer has leaves fluxes fluxes thickening woods’ fromthickening certainly mollis Northumberland and by extensive plantingforby extensive of the Cauca have ,

and MoroccoAlgeria and S. torminalis andIsles, theChannel areas highest such

Beck the lowest the

are given by Harper from from

east east advocated planting with hardlyany temperate and temperate sus Mountains sus Mountains the midrib(Rich , especially in – ( .

Kovanda Meusel

based

Ká et al pair of rp

, . á

Jäger Jäger ti

He & .

It et This article isprotected rights by All copyright. reserved. Acceptedis below1000mm largely where restrictedtoareas temperatures July mean are16 above Cisneros >800 with the optimum 6.5 to Sorbus torminalis European and ItConord 2010). suboceanic Rasmussen inEurope & distribution (Rodwell Kollman 1991; Sorbus torminalis (A) CLIMATIC TOPOGRAPHICALLIMITATIONS AND II. Habitat foundonly is mountains, it between 1300 Ferrazzini Espahbodi TurkeyIranMountains, min 2200 and (Kausch 2300min ascends Sicil , 1250min 720min to Article380 (Mad & Dimitrov and (Dinca 2000; is it personal communication) of338min limit Shropshire Rich Ardenneand from Scandinavia Pietzarka & Weinert - restricted to 1000 m in (W Switzerland 1000 min , withparticularly et al ě

17 ra Sorbus torminalis ,

, 2003; ° Tichá &Tichá Řepka . 2010)

C, withanoptimumC, of8.5 Lehmann& Roloff

2008 et al 1965;

Lorrai sits at sits . 2007b; Rasmusen & de Kollmann 2004a; Oria Rueda 2008). ;

s requirement for sharesthat it requirement warmth to such asimilar is distribution s below 60 m. below 60 , Saravi Demesure

with around 80% ofoccurrenceswith around belowelevational 150manda maximum e is athermophile, withabroad continental southern or sub performs bestwithin

(Kevin Walker personal communication). xcept forxcept Sorbus ne,

the current distribution distribution the current high and south toBurgundy south and

et al mm Paganová 2007

. 2013), is primarilya lowlandspecies At thenorthern range ofits mainland Europe limits inDenmark in -

densities

. 2008;Mad Musch ( ( Wohlgemuth González 1994; 1993;Pleines

Botanical Further between isfound it 300 south, 2009 ohlgemuth 1994). 1993;Pleines

100 ) & . - 1000 m in Ital 1000 min

In thenorth ° to the

( . Oddou C D

; those areas a with Genova 2008), Society of Society of ,

inc ě

Tabandeh Tabandeh - and annual600 precipitation between ra 1800 m (Pietzarka 1800 m

, ă 2000 e

centre Tichá &Tichá Řepka ast - y, Muratorio 2004 and and

1400 m in Lebanon, min m 1400 1500 of of it ; Britain & Ireland&Britain Kevin data; Walker

Franche Paris y (Rasmussen et al of extends tothe Balticextends and Pietzarka S. torminalis 175

.

in in the regions Champagne of 2007;

mean - the - - Blecken vonSchmeling1994b; Comté 760 m

, 2013

,

It p

LehmannRoloff & ; Lehmann & Brit

Bednorz Belletti Elsewhere

annual range of temperature robably indry, evolved open ). In). Moroccan the &

ish Islesish in the

(Drapier 1993a).(Drapier with about25 million

- Kollmann 2007) and 650 m in theBalkans650 min In is Greatit Britain, ° C and mean and annualC ppt ,

Monteleone & Urbaniak& 2005 , n 2008;Fady& n

Rey&Vinãs de

S. torminalis Roloff (Roper 1993; (Roper mediterranean is largelyabsent in the - 1500 mm 1500 mm

2009). 2009

& - ).

; -

This article isprotected rights by All copyright. reserved. flood Leuschner increasinglydry restrictedto atthe soils sharingmoderately to tovery soils, a requirement similar dry 1994b Řepka that are moderately to nitrogen deficient fertile & Buschmann although does it Camacho d Sorbus torminalis ( windthrow (Kausch slopes ( only important torminalis were notimportant. high that thepresence of aftera fire in ( towards the edgeitsrange south pr occurs of it facingand slopes,particularly easternedgeat itsrange the northern of ( undoubtedlysouthern is distribution by limited drought years,cooler can even insouthern Britain,be seed (Davis veryThe production 1976). poor InRoper 1993).Britain primarily summers bythatre cool limited Sorbus torminalis woodland and reproductive its performance isstrongly(Termena affectedby climate 1972). B) SUBSTRATUM B) De Dominicis Drapier 1993b ’ Accepteda Article nne 1990; Laniernne 1990; rock cover (as were otherdry (asspecies cover rock conditions), tolerantof were ed butavoidspermanently peaty waterlogged or (Maděra soils )

2013) . It. The requirement anddry for conditions tend warm Espahbodi 2006

, can is restricted tothe drier,(Wohlgemuth steeper slopes 1993)

Köckemann &Köckemann Buschmann

although it performs best on deep fertile soils soils performsalthough it fertile bestondeep Pinus nigra

also 2009). ; ). in hot, dry areas such as Iran dry occursin hot, asit areas where such onnorthand northeastfacing & García increasing

Barluzzi is tolerant of is et al cope - S Slope Slope Blecken vonSchmeling 1994b). only moderately et al

. It similarly is of tolerant t - . 2008) orminalis , Abad Delgado Alonso,Gómez

with awide range of abundant fruit isonly produced . 1990; ,

angle Q l 1983 y well from uercus ilex . Sorbus torminalis different ) only seems to become important on the wettest soils when seemsonly important become soils onthe wettest to Savill 1991; Savill . was Siles Siles

frost and cold resistantand and cold thenortherlyfrost is distribution associated with

2009 et al slight and eastern end end eastern soil typesoil duce successfulduce seed ( production

. Q. faginea )

. ( l Drapier 1993b; soil soil 2010 yacidic tobasicsoils It imarilycooler, on dampernorth , a wide rangea of

humus will survivewill on

s moisture although best on does it i , growing3.5 ofpH onsoils s verywind firm ) modelled the of range its

a

-

& Rodríguez forest

southerly ex or loess following

s torestrict acerbated temperatures by higher (Kausch García in southeast Taxus baccata soil fertility,soil coping withsoils

- soils soils (Grundmann &(Grundmann Roloff 2009 enriched aspect, recovery ofvegetation although altitude and slope warm summers warm

, - andfrom rarely suffers (Leuschner

Blecken vonSchmelingBlecken López Espinosa Tichá & Řepka Tichá that are S. torminalis . ( Similarly, aspect is D shallow

inc (Maděra Drapier 1993b & , and is ă 2000

te

Allué - 2009) facing slopes mporar , 8.0

and found Köckemann

soil , ,

( to south to south ) Tichá & Tichá and in Favre

although 2013). s

il and a y - ;

S.

; This article isprotected rights by All copyright. reserved. ovina also been recordedin derived oncalcareousash woodland from overand soils clay limestone(Pollard 1973). It particularly is McLaughlin (Williams, & clearings, S III. Communities 2006 typesfound in hasbeen Lowersea Wyeand Valley, disturbed habitat equally of soils dry west Shropshire and Nottinghamshire Yorkshire toSouth contrasts abundance withits on draining theFruitgravel naught, bear andthe soils], Tree compared very sadly” toclay. sandy soils. MeasuresOld Redglacial Sandstone andgravels Shales, 1993), avoiding (Roperthedriest strata produce bands moist much more locally the gravelsand or Oxford species (Pietzarka likely due tothe 2007 rich, from Acceptedorbus torminalis Article Gault Clay south of ; ; Maděr claysriver vall and clays and -

Maděra Agrostis capillaris Agrostis InBritain, Despite wide its tolerance, and Barton woodland edges woodland

Cook (1679) notedthat (1679) Cook a , , . It. much is

Tichá & Řepka& Tichá Řepka& Tichá from limestone.

associated withassociated woodland relic hedgerows , greater tolerance

, such as along the Severn Estuary at Chepstow and Lydney, alongsuch andas Estuaryat theSevern Chepstow and i S. torminalis

is mostly a treeof

Lehmann & Roloff on more acidic damp soils ofhilly acidicon more soils undulating damp gravel or Clay coastal andinlandcliffs, land hillsides, rocky the Gowerthe Peninsula

Shropshire growing and onrocks scree inbase ern s insoutheast Englands e mainland Europe ( - ybottoms Rumex acetosella less common on

. These include t (Rasmussen England

cliffs of South Wales cliffs Wales ofSouth

2013 2013 is The best growthThe best of shallow calca locally abundant of S. torminalis

wherea there is

). (Roper 1993 ). S. torminalis

Harrison 1986; S. tormina open deciduous deciduous open and the C and the

The contrasting The

2009).

&

Kollmann 2004a) Kollmann 1993). (Roper Oria calacerous he Culm measures England,he ofsouthwest Culm Coal

grassl

erous soils over soils theMagnesianerous

arboniferous where the lis

de Rueda de shows a marked derived preference for soils ; Brewis, Bowman& 1996 ; Rose

does fruit not

and ( to droughtcanopy dominant than most (Roper 1993). on the on the S. torminalis

continuous Kausch woodlands

Boulder Clay and p U1; Rodwell 1992) byLocktonU1; Rodwell 1992)& referencefor Italso foundtoalesser on is degree yarefree deposits ofsuperficial ,

nutrient Mart (species - limestone and Blecken v

í well Gravel on“sharp [free suppl nez , theedgesof rides and

hedgerows inBritainhedgerows and A similar preference A similar for soil is - - slips and similar open similar and and slips rich found on

- de Azagra rich ancient hedges)ancient rich - drier drier y of poor lowland poor lowland s on Schmeling Weald of the Peakof District the

virtually absent fromvirtually absent water ( calcareous is soils terraces where

deep, Limestone of ,

London & Paganová

). Á n the nutrient Festuca It lvarez

1993). occurs occurs , It has This

of -

, -

This article isprotected rights by All copyright. reserved. also a Quercion pubescenti sylvestris sylvatici notably western Europe petraea robur where frequentare most associates its Cardiganshire andBirmingham 2010) the (Chater region (Trueman Quercus robur (Pigott & Huntley 1978) is quickly but Like othertrees, these occasional cordata woodland hederacea common, albeit swamp primarily Anglia inEast along with Phragmites australis 1977) 1974) Kirby including 2006) frequently used been as woodland. Hermy Halliday( (2015) Whild

Acceptedmore Article , and

. , minor component of mixed deciduousof forest( component mixed minor

EppingForest (Lloyd 1977)

Incontinental Europe, InBritain, Despite needis its it for openness, et al , , frequent along the Pulmonario – excelsior Carpinus betulusCarpinus

Carpinetum

and 1997

(W8) sub Due woodland, closeassociation toits withancient Populus tremula can reachcan a (Rodwell large size 1991). as. (1999)it one of132species lists indicator European of olddeciduous

and heathland with Cephalanthero - - ). communit Pteridium aquilinum Pteridium still still S. torminalis in the southeast

S . –

as ascarce – torminalis

Fagetum melittetosum Fagetum

sub petraeae and and such Shropshire2003) (Whild such an an . -

y On more acidprobably frequently with associated it more soils, community of

Lathyro

and north A of , . Fagus sylvatica – ncient

As such can it from inthe zonation be woodlandto found occurs primarily ina (Trueman Fraxinus excelsior Fraxinus Pinetum sylvestrisPinetum

S. torminalis S. ( sylvestris

W is arare and western side of the community western side ofthe along with other occasionalalongwith otheras treessuch and around themeresand ofCheshire Shropshire. Juniperus S. torminalis ohlgemuth 1993 – , Lincolnshireand (Peterken, 1983)

Quercetum W scattered treescattered - Rubus fruticosus oodland Corylus avellana Corylus et al component on drier (but still damp) areas of damp)component (but the areas ondrier still Alnus glutinosa also

, hornbeam woodlands,particularly

is typically with associated scrub onrockscrub Lake outcrops the in District by . 2013).

and , pine

I a common component of oldwoodlandsand

does not usually not does invade ne ndicator in ndicator (Rodwell 1991; Lockton (2015) 1991; Whild &(Rodwell , and mixed oakand woodlands mixed , – , Carpinus betulus Carpinus Acer campestre ; in Moreover, l

variety of Pleines 1994

Aceri platanoidis woodlands the ,

L woodland reported as (W10), for , ancashire (Pigott and Westmorland – Primula vulgaris Hyacinthoides non

Carex paniculata many calcareous ike

S around Mor around orbus such as

). areas –

Tilia cordata Tilia In Germany, Central is it

Mercurialis perennis Mercurialis (Rodwell 1991) & , Poulton –

Derbyshir torminalis Tilietum platyphyllosTilietum of Britain Britain of Quercus robur Molinio beech woodlands, beech w – Betula

Glechoma ecambe Bay woodland

- stands very scripta of

the Reade Gallo Gallo , – e (Willmot (Willmot e

S

spp., (Rose 1999; has Pinetum Pinetum .

alliance . It is t , orminalis In Quercus

(W5) , 2013) Tilia Tilia Q. more

.

, )

This article isprotected rights by All copyright. reserved. only Sorbus torminalis IV. Response to bioticfactors on calcareous2004)and (Loidi soils merging into component of theassociation that are coppiced frainetto northwest Greece, along with in acidic below the Lebanon Alno range including forests ofbeech Iran, isfoundonwest it woodlands and torminalis d Szymura2012) genistetosum tinctoriae present the in the rare of thermophilous association woodlands of communities shrub communities limestone (Gebauer dominated by Accepted Articlery - mesic associations theorder in – reaches its maximum height maximum wherereaches its open in conditions light Fagetum Aroun FurthereastTurkey northern in In o EasternEurope

Q and S. torminalis

Cedrus libanii woodland is a uercus ( Velichkov sub Q. pubescens

in the Festuco d the Mediterranean, d theMediterranean, open, the class Querco Fagus sylvatica defining

. In . Slovakia, ( Espahbodi Espahbodi -

( Mediterranean dominated by forests

petraea Rackham 2003; 2003; Rackham is apoor competitor S. torminalis Fraxinus ornus Fraxinus , in the class species rich, species

Horna Corno

, is restricted to

- of Poland of species ofspecies Zlatanov & Popov

and southwest forest

woodland

– ne of thelargest of ne concentrations of woodlands in - et al & Brometea Quercetum Querco cerridis S. torminalis Carpino orientalis , Leuschner –

Tilia Rhamno

( Fagetea . 2007a; is Rich Rich Carpino slightly acidic oak forestslightly acidicoak ( the mesophilous alliancethe mesophilous also a

Medak 2011). Kwiatkowski ( S. torminalis Vukeli

s spp. and various Q. ilex

mixed deciduous forestmixed limestone at on Sorbo torminalis et al in the

, (Rasmussen &(Rasmussen Kollman 2004a).

- in upland mixed woodlands in upland mixed . In. Poland

readily out - facing slopesbetween1500 (Prudič Rad & Shafiei 2010).

– Prunetea rare of oftheshrub component layer 2012; Seidel 2012; Seidel the orderQuercetalia Fagetum

. 2010).

is also versiona soil the of component of thebasic ć 2007;

, order in –

Baričević &Baričević Šapić Carpinetalia orientalis northeast Spain

1998; 1998; is afairly constant 2003:

Quercetum cerridis Korkmaz

Simila

andgrassland as acolonistdry in Quercetalia robori - , , competed trees by other Slovakia Rusco Acer – et al Roleček Bednorz 2007b; Quercetum r l

y –

Luzu Quercus pubescens , . 2013). It. 2013). alsocan occur in tall spp. on loess underlain by spp. onloessby underlain et al

Fagetum Carpino betuli in central , isaminor it in central and theCzech Republic S. torminalis

At its south At its

( pubescentis 2005).

lo luzuloidis . 2011) Sardans 2010)

fluxes

, - on though infrequent

1800 m altitude inam altitude 1800 ,

- and in upland mixed in upland mixed ( petraeae Fagetum OrientalFagetum Rosati Rosati

. acidic soils acidic . In.

From here downinto

are high

Maděra & in – - the mountainsof

, particula Peñuelas eastern , Acerion hirkani occu

– a et al Quercetum c

nd sousually Quercus in northern . 1500m, and

rs in et al . 2010)

andalso is limit in in limit Q. faginea ,

2007). . rly those S

it formsit 2012; species .

and , -

This article isprotected rights by All copyright. reserved. a reasons Kaczmarek prevent theof naturalregeneration exclosures root suckersgrowinga of3.8 mean cm Kollmann (2007) fencing found roedeer that toexclude abundance of northeast France However, 6), withfew chemical defences than some other digestive( inhibitors torminalis (Harper 1981; 2007; 1993b saplings by arebrowsed readily invertebrates ( infertile that (VIII(C)) and are are those readilypredated by viable thinning under low f correlated (2007)suckers.& Rasmussen found Kollmann that component of oldwoodlands. et al d’a Pietrzak Central Europe forest trees (W or

number light ofother

Accepted Article nne ohlgemuth w .

ater 2006 ; Collet Collet

The management oftraditional loss woodland beenas themain has suggestedone of R 1990; Drapier Ewald responsible forresponsible

2014). leading to lower egenerationalso is vu ,

,

nutrients Boulanger . Certainly,a ofauthorssevere thatbrowsing number enough suggest to isusually w ) although, as noted in III,noted in ) although,as isalsoa it

against largeon a herbivores 1 asvery low(2 . diffuse light

2012; Forexample, ith et al

, a Termena 1972 1993;

Nicolescu

Zander s preferred species preferred where seedlings of other species where ofother seedlings As such, some regard , tem

. 2008) and suckersareeven prone more t

S. torminalis Maděra

and particularly

Collet Collet l

et al ength 1993 & -

S. torminalis S. demanding species, fluxe

et al Jander . (2009) sucker

seedlings et al b; Demesure , b et al ; s . 2009). asal diameter Lloyd 1977;Veli S. torminalis

lnerable influences. tobiotic since here . individuals individuals

.

2012). small mammals, rabbits and ungulates mammals,rabbits small (

1994;

dens 2008 of state S.

tend tobe Cornus light

becoming

ity. torminalis it as an early asit Borchard ) d Schüte 2000

in height or

et al ,

that in their studies in mixed oak that intheir studies inmixed woodland in

root suckers root suckers leading to where

‘were alwaysavoided’

and sprout is susceptible toself mainly as a resultmainlyas of and .

2000 more abundantmore and onshallow č rarer in Europe over the last century, inEuroperarer the over last as shade al et Rosa ka 1993; Rich 1993;Rich ka (Bednorz 2009 are conditions weakenconditions thedominance of in one seasonin - b ;

root to mid ; Hoebee . self less abundant Biedenkopf

( - canina

were taller but thinner 2011 tolerant a ge - - sucker ( thinnin 6 scalewith

Capreolus capreolus , suggesting that suckers , suggesting that - o browsing anddeer by rodents ) succe The majority seedsare of usually

. In. &contrast, Rasmussen categorised the defencecategorised the of

et al ,

et al compared to 0.6cm outside compared to if ; d - g

, competition betweencompetition Bednorz

ensity a c change from s . non , pe Ammer

sional species ( sional species

. Competition was higherCompetition (

mammals 2006; Oddou Szymura, Szymura, . 2010). Seedlings and Ellenberg rhaps duerhaps tothe - Taxus baccata reproducing

was was Harper 1981;

, & Kaźmierczak &

negatively , M , birds

Szymura & Szymura with acidic soils

ü L.) et al ller - oppicing to F Muratorio , resulted in resulted avre self

- compete and . Starck

being 1991). Drapier root root - other

- with S.

in

This article isprotected rights by All copyright. reserved. seedlingsclustered tobe similarly population of597 Kaźmierczak & Kaczmarek trees. trees;individual ata largerscalemore thepatternwas distance trees, of2.5 mfrom the considerably theBalticislands in Sea, (K across 157ha Huntingdonshire (some trees tall)have upto20m 70individual around recorded been where typical Nationalwell studiedpopulationatMonksWood Nature thea is Reserve in S. torminalis 130 eastern Weald between are mostly possible, where &Dimitrov Ginova 2008; Muratorio range ( are usuallyinthe range of found thatonly sites 11% of ( 100 individuals of individuals S GREGARIOUSNESS (A) V. Response more closed highforests ( orbus evin Accepted Articleroot - ha areacoppice of detail XI in On steep slopes,On steep suckers togr tended

Pleines 1994 suckers O Walker, personal communication). Walker, personal t orminalis n a small 2004 across of range. thewhole its M

in size in size tree

to environment of ancient ash/oakBoulder woodlandof and onOxford the1970s Clay since with population size often size with population being than fewer 10trees ; Pleines Oddou . s per hectare

er S. torminalis is ;

M from 6to from scale, suckers growing from suckers rootsscale, inevitably tendtobe e Demesure Demesure xtremely non ü - with ller Ashford inKent inEast andRobertsbridge Sussex -

Muratorio 1994 Rasmussen &(2007) found Kollmann 0.1 to Aas & 2011; Kohles , S.

- Ammer &Ammer Nüßlein ( ( standardsAustria, insouthern 2012 8

torminalis ;

( 207 Hoebee

DBH, diameter at breast atDBH, diameter height, out of73) trees et reflecting the concentrationreflecting of the suckers 40 individuals ha 40 individuals . ) al

suckers -

et al investigated natural regeneratiinvestigated natural gregarious, as found .

in a 1.72 ha fein a1.72ha 2000 et al .

has been and as cultivated planted such 2004 had

a 100 m ow perpendicular totheow perpendicular slope. ; any woodlands have onewoodlands just any . 2006) Hochbichler 2003 Hochbichler ; >100 >100

Belletti 2000; 2000; Maděra - 2 - . 1 ; nced in plot

S S. torminalis for exa with most beingwith most at the lower this end of uckers were clumped highly

Szymura 2012 random pattern reflecting between the ,

et al Monteleone spatially isolated,

mple, Hochbichler .

2012). ;

> 7 cm). > 7 north that Demesure

in Poland, Bednorzin Poland, (2007c) trees. densities Population o n

and rarely onlyvery over clump & ) Locally densities high from seedfrom ina

; this is discussed in is ; this Poland Ferrazzini around thebase of around

(2003) (2003) Possibly more individual orclump individual Bednorz

- s var s (Roper 1993) scattered Musch &Oddou clumped.

and found ied recorded in

2008; up toa ,

the On . Ina . 36 - This article isprotected rights by All copyright. reserved. population was as classified shade dominant and39%‘intermediate’growing were the or canopy; towards 22% only ofthe (2012) found that34% of component of thecanopy oaks. DBH Germany, & Kunz found Pyttel, Bauhus th (2013) (Gonzalez 37 Europe Kahle 2004; von Schmeling showinggrowth annual or little reproduction conditions &Schmidt Meyer suckers forests (Belletti disturbancegaps creating ( hedgerows, relic theformer wood including hedges mark woodlands that boundariesof woodlands suchas coppice, cracks seedlinga can survival with exemplified suckers no more 1 than regeneration of such as tolerance high temperature (V(C)) of and drought but, given partial its tolerance shade Sorbus torminalis (B) PERFORMANCE VARIOUS HABITATS IN Pollard 1973;Pollard

Accepted- Article 54% Quercus , Further more east in continental conditions

and most were <10 m tall withonly mtall and were most <10 treesand soshorter occasional than >20 m the ( ; for example, in mixed oak woodlands in example, inmixed ; for On deeper Roper 1993). on

can be outcompeted becan outcompeted of ,

rocky they 400 m DeconchatBalent &

on Rich Rich castaneifolia Rasmussen

, 1994 south facingsouth other tree speciesother will persist will

Monteleone outcrops 2 ,

et al

has been classifieda forest specialist as( 2004; more nutrient plots plots b;

. 2010). lso behigher - Prudič Prudič 2 mhigh 2

as Hochbichler 2003;

. . Onshallow or rocky soils, in sometypes of

S. torminalis

This is particularlyThis is so &

an as asapling bank (Zare , sandy soils near the Caspian Sea the, sandy near soils the edges of rides, and woodland marginsasand the edges ofrides,in aswell

byspecies competitive more & Kollman 2004 understory 1997 As such israrelya it canopy tree Ferrazzini

- 2009)

, , rich but

d but alive. In study,d butalive. of treesfruit, this bore ofthese 35% Tabari &Tabari Espahbodi S. torminalis ;

due toprotection (

Stăn with VI(E) soils, soils, .

individuals wereco In a escu shrub

some 2008) woodland ) S. torminalis b; Maděra Quercus petraea Quercus , Schrötter 2001) lesshighlySchrötter butin shaded until a until 10 from poor ,

Şofletea &Şofletea Popescu , and , and

growth may restricted topatches be of of the densest growth thedensestof .

within oak within

S. torminalis southwest France , it , it In moreconditions shaded at 65%of competi cano only

is . In. Czech Republic the

- from predators

where 20 years20 age of et al. et also oftenfound 2002).

py performs op bestin

7% su tive abilities tive abilities - de Albuquerque &de 2010 Albuquerque Rueda

- dominant with theoaks,dominant with 3%were ch and as beech oak 2012) gap opens up S. torminalis

conditions are and beech woodland in southwest woodland insouthwest of plots as a of as plots

On rocky where becomes a more frequent

in cool western areas in coolwestern

and where thereis and where

, 1997

S. torminalis S. torminalis within

or as mature as trees, or of anyofhabitat -

( dominated in ; Hochbichler outcrops IV

treescm were <7

( canopy open Kausch ) too harsh for

, and high its crevic en seedlings Maděra areas within situations situations (Geb , s

occurred in

tree - es

is eed andeed Blecken mixed , as ,

mixed and

et al ,

of

2003 and .

)

; This article isprotected rights by All copyright. reserved. ( Conord 2010). shows a strong north while Bednorz climates; conditions S. torminalis (Schirone 35 planted 1 alsoseenconditions is towardsmortality the ofitsrange south the of innorthSardinia: Germany to0.006% corresponding declinein centre itsrange Germany. This of incentral and alsosoils, increasingly range alkaline ofits intheeast soils in the easternedge increasinglyrestricted tomoderately ofitsrange, being and dryvery dry (Balandier pattern:similar 23% for (18 cm) and smaller growing atfirst the end ofseas meadow vegetation gapscan by belimited still poorcompetitive its ability.F Light also needed is for production seed (VI(E)). growth but (Rasmussen & Kollmann positively Sea regenerative werealways 81% were dominant orco Rasmussen &2008) Kollmann

Accepted- Article 40 cm high, but this was40 cm ona high, p butthis , root ern Genetic diversity in K

Poland ö - sucker densitysucker RasmussenKollmann &(2004b) ,

year

ckemann (2008) correlated with lightcorrelated with , Salis (Dimitrov &(Dimitrov G

they tall were also Frochot Quercus petraea

success along woodland edalong woodland under beech were and damaged individuals) (<65 upto50%were small by dry -

, old seedlings of At thenorthern edgethe range, diversity of between populations decrease with latitude decrease

Kaźmierczak & Kaczmarek . &

Vessella

in central France, &

with of Sourisseau S. torminalis S. (rangingfrom 100m 207suckers 6to S. torminalis in Slovakia. regional - 2007). Suckers were inova 2008) dominant andthe 2% provides

S. torminalis 2011).

(14 cm).and followeda Survival between September May under the lowest light flux densityflux S. torminalis

ges dominant and since

abundance of abundance of 2009). ar with theMediterraneanar specialist

Nevertheless, in thedryBalkans,Nevertheless, of populations in southern

. evidence that

, asdoes it for manywoody species

A Itscope ability wi to compared to6%for . Reproduction appears. Reproduction to gene S. torminalis on (4 cm and 3cmcm respectively) and on (4 t the northern end of rangeinthe onislands Baltic its

is generally between (VIII(A)) high populations but

noted flow (

below the canopy deciduous ofthebelow fore mixed restriction in soil types insoil restriction (

was >78% over 12 years,>78% reachingwas survivors over 12 just 2012 also S. torminalis S. of treesof that

poor repr outbreeding However, . Light the in appears to be important ) S. torminalis

found an population expanding and tallest fluxes Carpinus betulus Carpinus ollowing ofseeds into direct sowing

under high light th C. betulus C. oduction in a Danish population inaDanish oduction , presumably toetiolation. due establishment woodland into , and even

had

) from 0.25%cover

- between populations is between populations 2

) occupies a be and basal diameterand were basal the Slovakia better in more continental in better is associated witha is associated

large and 37% forand Quercus ilex require than than

in Europe were significantly st fluxes fluxes narrower narrower ,

number of fruit than inthe Prunus avium ,

hot, dry, open

is higher with due torapid P. avium

in

(Fady &(Fady niche at in

st

This article isprotected rights by All copyright. reserved. Acceptedcommunication) detrimental due shaded theincreased in conditions shading (Rodney without plastic Wilhelm herbaceous species edges ( decline conversioncoppice high of into forests is forests, century (Lloyd 1977 in Europe with deeper habitatsextensively hasbeen modified by human corridors (Hoebee would benefit (Biedenkorf treesfrom inastandoronly standswith few are genetic a trees used,the diversity lowering Article geographicalthis patte also because clonal perennationgreater inthe north,and thecentre in between outbreeding populations but Oddou locus ina population small diversity ( inthe north lower Muratorio France due to comparison, trees) (Finkeldey populations, also drift comparatively their lo lowdue to . Populations inSwitzerland. Populations andGermany parts of showsimilar - Maděra tree shelters

There doubt isno By contrast,genetic populations diversity within

of Muratorio shelters &

are found S. torminalis et al

Ducos 1996 soil et al genetic very is centre between populations ofitsrange diversity lowinthe in the larger populations and increasedlarger populations from therestoration connectivity increased through or re

. 2001c e

. t al (Dupraz 1994) (Dupraz in mainland Europe central populations arecentral largerand populations

. 2007). attributed toreproductive of (generally isolation thesmallpopulations <100 Seedlings are et al et al et al : after 4 yearsafter 4 : is consideredbeneficial to . ) wher 2012) and . . 2000;Rotach 2000; ). The rn tends tobeby diluted rn planting .

.

) – 2007)

2001b

.

The management for implication forest that is

e coppicepracticwas management that thefrequencyand abundance of see XI. In France, southern

) compared to creation ofespecially forests, high by transformation 1.8 . ; , although

very sensitivevery to the removalof - Rasmussen & 4.6 alleles locus per ,

n

Forest management invo trees inside shelters were shelters trees inside g -

term isolation from isolation each andterm (Szymura 2012). (Szymura 2012). , as many with , may shelters species, prove 10.7 alleles per locus in Biedenkopf S S. torminalis

Kollmann 2008 Kollmann competing(especially canopytrees beech) and gene flow .

t herbicide

orminalis

often cited planting, certainly o hold ) and ( inSwitzerland Indeed ,

highergenetic if seeds fromif seeds number alimited of Ammer shows a rever

(Demesure

l treatments (Oria de Ruedatreatments 2008 (Oria

ving of thecreation

has been seen to c ( ed until the end of ed the end until Crave 1985;Wilhelm . 190cm as one oftheas reasons one for the , m ) S. torminalis . This is partly. Thisis greater due to

ost of &

Franceet (Hoebee al. higher levels ofgenetichigher levels

M

ly high ly Helliwell, personal et al ü n more productive sites co

se pattern with a with se pattern the larg ller variation - creation of isolated m . - 6.4 alleles pared to Starck 2000

in different diversity between benefit from e

woodland the

. However, populations

a populations and ;

2007). By2007). Oddou

nineteenth 1993; 130 cm lan

per dscape llelic

2006; - ) .

This article isprotected rights by All copyright. reserved. centre (oceanic)and tothe western edge of itsrange ( conditions. al height southwest Germany correlated with crown ( diameter BednorzPoland, growth mm averaged 1.9 southwest Germany, & Kunz foundincrement Pyttel, Bauhus thatannual (2013) radial Germany although recorded Ramussen(2007) tree mean (Sevrin & 1993 Keller as years recorded years (Demesure 20 Rö to 15 growth & Oprea Schüte 2000) to torminalis Blecken vonSchmeling1994 debilitating effect oncompeting speciessuchas forest and species, reaches Haralamb 1967 . (2004) - 1

Accepted Articlehrig 1972; 30 years30 - yr 2 mm inwoodlandwhile 2 mm -

(Sevrin 25 cm - in metres , 1 Maximum ageMaximum of d Stem Heightgrowth generally is as slow described

become

with annual growth rateswith annual of between 20 at around 25 yearsat andth around remained constant 25

. ageHowever, appears tobe ofthe2012). tree as

found Age is also a large factor in diameter growth isalsoalarge Age factor indiameter from 35

At the 13.4 mat20 (Sevrin &(Sevrin 1993) Keller butin yr -

2.2 mm 2.2 mm & 1993) Keller Schü iameter growth - 1 s

; Stă

a mean of , (y = 0.799 at 50 yearsat old 50 - DBH was relaDBH was very small oreven tocease small appears ( very Musch &Musch Oddou Kaźmierczak &Kaźmierczak Kaczmarek northern and eastern te 2001 - 60 cm y 60 cm found a p found nescu yr ;

Hochbichler 2003;2004 Kahle - 1 - S. torminalis

49 years,49 15.9mat 50 during the first 8 yearsduringfirst 8afterestablishment, the then decreased to<0.7 ; , 21 cm

a

Kahle 2004 x + 1.125, x r Şofletea & Pop - maximum ondrymaximum where sites low 1 . b; Schrö

ositive relationshipositive between varies with competition varies with individuals in theopen in individuals can have in Germany (Kausch ted to age:ted to and 5cm r 2 yr -

= 0.830) whereas= 0.830) Muratorio (2004). I (2004). Muratorio - 1

in tter 2001).

is r ). edge 2 northeast Romania andnortheast Molda

highly

In height trees, reach is maximum shaded = 0.71) yr DBH = 16 ×DBH = ln(age) more escu - s 1

of its at 70 yearsat 70 old( (2012) found(2012) that ring wid

-

- variable 1997

79 years older than79 and trees 18.7min open open Quercus 100 cm , Growth

and innorthFrance - main Blecken vonSchmeling , ereafter. live for 200 live

Pyttel, Kunz & Bauhus (2013)Pyttel, in Bauhus & Kunz ; Ramussen 2007), but is oftenbut is quickerassociated than , conditions Crave

; n and annualbe as ringwidthmay low . In.

a ths Sjöman

and dependentand on France meanhave been heights yr range more important also

spp. and DBH - Quercus petraea of 1

1985 in the first 10

After 50 After varies across therangevaries across of up Cra

, water availabilitygreaterwater a has − 39(

r , trees live for 100 trees livefor ing widths of widths ing

- stronglymost DBH was in centimetres in Nielsen &Oprea growth for continues to 16.1 300 years 300 ; ve

Fagus sylvatica Fagus Drapier

1985 r , 2 -

v

70 years 70 Oddou = 0.73

i

whereas a mm determinant determinant ; Bamberger; 1990; ( -

growing 18 years,18 falling Sjöman 1993 (

Crave 1985; woodland in

1993, in central - ) 2.5 Muratorio . ,

and c diameter - - trees inthe ). 200 years200

4.0 mm 4.0 mm

2012) 2012) , ( ed within 1994b; In north

tree Kausch Nielsen of 80 140 S.

- et 100 - This article isprotected rights by All copyright. reserved. Acceptedshoot (Barnola, Durand & Parmentier 1993 lateral, In open (A) MORPHOLOGY VI. Structure physiology and 36 plantsinvestigate likely prolifically tosucker canopyground tothe written about diseases ( droughtreduced lead can to torminalis drought laurifolius Cistus embolism intense droughts H 0.05 mol Articlefollowing period a through dry was isohydric Ellenberg 1988; or more co particularly frosts terminal can leading early unhardened autumn buds kill toforking Rasmussen 2007) downto tolerate lowwinter temperatures Seedlings by be can but damaged frost (C) DROUGHT, EFFECT OFFROST, ETC. 2012 Drapier measured ) and even up to 400 yearsand upto400 even (

angled branches

Sorbus torminalis S

conditions 1993 in orbus torm , Hemery , similar , similar

due due - and evergreen forest oak in 2 do O m a; Roper1993 the minant shoots in northeast Spain Spain in northeast to a certain degree control (VI(E)).to acertain stomatal through A , - González .

2 , fire tolerance of Martínez monspessulanum

, the crown is round et al loosing > s in degree - inalis 1 d

in would suggestwould above intolerance bythe had had . 2009). ,

July to<0.01inAugust ( while under shade it tends towards being narrow with a main vertical

is is , resprouted afterprescribed -

- Rey& Vinãs Cisneros de ; Vilalta after fire 75% of maximum hydraulic ofmaximum conductivity during a75% summer 31 to

tolerant ofshort very growth to (Pietzarka Demesure

Quercus ilex till

Favre ( tolerant ofdroughts at - Haralamb 1967; Haralamb 1967; S. torminalis 34 ° northeast Spain August. Stomatal conductanceAugust.dropped Stomatal however, from et al

than intheother >0.5 m mmol and .

ed Indeed, - C , - once cold hardened, d’a . (2002) found that it was. (2002)thatit found

, ovate with Musch & Oddou &Musch Lehmann& Roloff

potentially a greater susceptibilitypotentiallyato pestsand ( Schüte ; nne

Drapier , - term 1993b) (Drapier flooding Acer monspessulanum Acer Quevedo

1990;

but the thin barkbut the thin andcloseness typical ofthe

2000) Martínez - 2 . Foliar senescence higher in was González

MPa 1993 strong

seven

Hurt & 2004 Kantor 2003) which achieves it by being that last that management , .

Rodrigo& Espelta - However, - c 1 Muratorio

;

- s

orthotropic development orthotropic development species tested.species Vilalta Sevrin -

1 sapling 2009). ,

in July high and remained - Rey& de Vinãs Cisneros a few ( months ground portion vulnerable &

et al spring frosts s and s

fire Spain. innortheast , Hydraulic conductivityHydraulic 2004

Arbutus unedoArbutus Keller1993 . 2002 older ; ). Thus prolongedThus

Maděra

(2007) found all .

to xylem Littlehas been Haralamb 1967; 1967; Haralamb ). ,

but it is highly is but it trees

But i and ).

The et al

S of and n more can

. into two into

many

. 2003;

c

. This article isprotected rights by All copyright. reserved. Boulet Lachard & 1993 Mansouri the wood becomesredder sometimes with There isusually as 325kg m C content at m C 355kg groundandnot durable is the incontact with However, (Wagenführ1996). m of clear stem and a canopy diamet Wormley Wood, Broxbourne, Herts was reported in 2011 to be 28.5 m tall, 78 cm DBH with 3 2006; 1993 Schmeling 1993 France and Germany (Crave F high 1993 (Drapier qualitylogs twisting Cisneros pruned, epicormic few br produce (Crave stems merchantable spacedbranchesreadily inthe open, and needed dead brashingis fall donot ofplantations to Vinãs & Cisneros led toinferior remaining specimens (Kausch singleare less trunked fre trees now (2009) as being of formsalso are Europe seenform inmainland withthe shrubby described by found at Monks edgesclay onboulder old. canopy is composed of a mix of Accepted necessaryormative pruning issometimes Article In ; Orša Demesure -

Britain, there are two main The woodi Sorbus torminalis In shade, self 1985; D G

and leaning

ercourt 2003 ni ć ). - et al 3 rapier 1993c; Wilhelm 1993). Wilhelm rapier 1993c;

)

- in

The HuntingdonshireWood, little differentiationlittle between

2000). Musch & Oddou and up to 1.0

2003; s moderatelydense Quercus robur ‘ . 2009). chandelier -

, towards the lighttowards readiness toforknoted above, atreadiness <3m,as pruning of dead branches is rapid leaving a clean trunk ( ( , and , and - Lanier 3

The bark is high woodland spec compared tothat of O an can readily reach 30 m in mainland Europe and even 32 ria de

; Trees in Britain do not reachTreesBritain abuts donot size insuch

sprawling multi ches(Lanier are produced Hochbichler 2003 et al - ’ 1.4 m DBH (

c; Sevr habit long can shootspurs andyears short shortshoots; be upto66 R - .

is comparatively thin, decreasing is comparativelythin, canopy

Muratorio and 320kg m C ueda 1990; er of 14 m (Henry Girling, personal

quent duetorepeatedquent 1985; with

is a constraint inthe productionconstraint of veneeris a and other at m 640kg in ,

Mart Sevrin & Keller&1993; Wilhelm (HenryGi forms: multiple forkmultiple a core brown Drapier 1993 Drapier - to preventdevelopment of Drapier 1993 stemmed shrub stemmed - 2004 í

Blecken 2006). vonSchmeling& Kellner the light ) nez

and ofless and commercial value & Keller 1993; medium tolarge tree

; de Azagra - - 3 Kausch 3 r

in ( ling, personal communication Safdari Safdari - et al Taxus baccata coloured s and lateral shoots. The upright, s and lateral shoots. c; in older, slow a

;

cutting -

Sevrin . B Kausch s

1990;

lecken with no dominant leaderwith nodominant & et al

Kausch and the frequent slight frequent and the Á sapwood and heartwood, and sapwood but l . 2008),tobirch, similar of

& Keller varez González 1993 ies with dense with wood,ies such -

with increasing Blecken v Blecken v the on Schmeling s

(Puhe & Ulrich 2001).

- the ch the

- favouring woodland communication grown trees B ;

best treesbest whichhas

2006) Ramussen 2007 lecken pecimen at 1993). 1993). the woodcarbon the González , andelierform

Nicolescu Rey& Vinãs de (Raguin on

but when well

- v

33 Schmeling on )

When When

& . Both (Sevrin, (Sevrin, d m iameter

, , such as , such Kellner

& in ). Rey de

et al

).

. This article isprotected rights by All copyright. reserved. thatpossible t concluded that of leaveshairs (37 numerous with (54%) or truncate measured) shoots. and 66.71 angle and between themidrib tree observed vary dependingtypeand uponthe p ofshoot in Poland. decreasing distance the between (Bean 1980;Rich underside Lehmann & Roloff system 50 23.1 cm) discussed by (2009). Wilhelm La Mansouri (1993), Safdari Wagenführ (1996). Physical, 7 the earlywood( longμm and 20 the beginning ofthe many there but porous diffuse isso woods, from 8% 12to of orminalis μm) - Accepted Articlechau 70 cm , butha

bordered (Lachau pits d & Mauro d The morphologically most variable y One ( The base from often were ofleaves shortmost shoots truncate Herman 1971

increasing to40 that leaves were smaller onshortthat leavescompared were shoots mechanical and chemicaland mechanical properties

° in , and generallymoreand variable , cordate , However, (1960) that even Kárpáti individual has out within pointed trees, leaves

d a longer petiole (mean petiole d a longer in long respectively) lobes project shoots, more thebottom and onlong short so and are depth some ear leaves on short shoots bettershort shoots characteriseleaves oninterpopulational variability in Sevrin, Lachaud& 1993) Sevrin, Mansouri - - 27 old seedlings been have log (39%)and rarely cuneate (8%). et al , u

forms of

therefore increasingly strong (33%) (33%) μm wide inlate and earlywood wide μm sset Wood structuregiven aregrowth details Lachaud and by&Wood Sevrin, 2009;

earlywood

volume ). . 2010).

(1996 On porous soi , -

Fig. 60 cm in or cuneate Sigarody& Ahmed S. torminalis d & Mauro best used for identification best used for

(Kahle )

the vein in the lowest pairthe vein oflobes thelowest in (mean was larger of71.80

3). L 3). , Bednorz (2006)thatle found

and silvicultural management toachieve value high trunks is (Safdari second second - 39%), a(54%)and hairs 39%), nohairs few eaves dissected are more 2 of 3.72

- 2004). 2004). (27 3 year3 ls, roots mayls, roots reach 1

in outline in outline and farand u and %)on while sset 1996

et al based onleaf morp seen metimes character and - Growth ring Growth osition onthe tree.osition third old seedlings (Schüte 2000) . 2008)

- of with a ( reaching

2011 3.50 cm 3.50 S. torminalis S. towards the south lateral increasing veinswith distance south ; No differencein thewas proportion found ,

; Pietzarka respectively a narrow band of vessels . ),

of of

maximum long leaveswere shoot mostly cordate Vessels Lachaud (1993 & Mansouri S. torminalis S. . , respectively lateral roots

On the strength of this variationOn thethis is strength it of boundaries

- 2 m depth (Schüte 2000).2 mdepth(Schüte aves and a lobes had more , more pubescent onthe , more pubescent are to hology (e.g. ,

are LehmannRoloff & woo long shoots long

Indeed, jo , root

with higher vessel areawith higher in mostly so i ned large( by numerous d and east east and

are summaris create create as in are indistinct, depth is leaf shape ( ). ). slightlylarger pores

(40% ofleaves(40% On long the On shoots Bednorz (2006) . (6

forma Once

- o at the topofthe a heart litary 9%). Bednor of range its f 32.

roots reach perincisa , 0 cm

ed in 2009 939 ) and root Pietzarka

-

992 (mean - )

. S

z .

at c ° . ,

This article isprotected rights by All copyright. reserved. fromcomparatively seed be can rare produced Sorbus torminalis PERENNAT(C) is suspected ofAMwiththetree forming ( pratense Glomeromycota) Acaulospora viridis symbiont on Burgundy truffle( not correlated chemistry. tosoil Simila chemistry was colonisa correlation foundbetween root butno autumn (32.4 fungi ofunk Moradi Sorbus Ha (B) MYCORRHIZA (BednorzPoland 2007a, Bednorz produced that fruit was constant width more than length characteristic by which to that fruit the lengththan widthin constant more is within (SE, with individual represent seen normal populations variation inmost ( Borb

Accepted &rley record Harley (1987) Article n ás

r = 60)

populations andgreater variationpopulations between l Idžojtić, densityStomatal species native to

et al yfungal densityfromsurrounding soil theroots AMspore (56 & Fekete&

in pot cultures, has cultures,therhizosphere beenin pot within of found . Other than there. Other this, profusely nown species in northern Iran. innorthern species nown . (2014) observed 51.0 in length S. torminalis - 55.3% of roots) wasroots) initiallybe55.3% to thought due of seasonal tochanges soil in

stoma Zebec & , which has formed AM associations with with hasassociations formed AM , which ) B ) I

Tuber uncinatum Tuber responds well to ON: REPRODUCTION ON:

from roots from Palenz., Oehl,Palenz., Azcón eck) eck)

and and on matureafter leaves (24daysbudopening) measuring

range Britain Drvodelić and

distinguish in southern Chinaalthough are given. results specific not 19.13 ± 0.33 µm wide 19.13 ±0.33µm forma s in mainland Europe

S. torminalis

from from are few that are also a Wehrelen et al -

co

68.5% of 1 cm long be 1cm to colonised root68.5% sections of

Chatin (2006) noted , particularlyat the edges range ofits mollis 171 ppic between . 2006b).

geographic trendscertainly fruit in orseed traits, in - - 220 mm ing Aguilar &Aguilar G.A. Palenzuela et al

: Ascomycota, Lower colonisation Beck

as being only ectomycorrhizal, unlike otheras being ectomycorrhizal, only

and pollard and rbuscular mycorrhiza in Poland in Poland Sorbus . (2009) report. (2009) ontrialsestablish to

)

(

some encompassed within Bednorz’s st encompassed Bednorz’s within Čaňová - British IslesBritish 2

but less so in Britainbut less so

. (

ofletea

et al sp variation in fruit and leaf morphology infruit and variation leaf ecies despite ofseeds variable numbers ing

Pezizales

et al . 2014). Silva Sorg

(Harper 1981). 1981). (Harper . McAllister (2005) suggests

tion and tion soil et al but Bednorz (2007a)but Bednorz foun . 2012) and of rootsfungi by

hum vulgare ( Diversisporales

S. torminalis . 2005; ) l (AM). l (AM). is

as an ectomycorrhizalas an . thus a useful thus a

- 81 spores g 81 spores .

S

Čaňová and indeepand shade ince establishment chemistry. However, Suckers and

29.20 ±0.50 in Spain and in Spain in the , Trifolium Trifolium

et al

- the 1

udy may

soil) wassoil) are byAM . 2012) d

, This article isprotected rights by All copyright. reserved. 1977) Drapier Kollmann 2004 5 within & Roloff typically 1993; Wilhelm 1981), especially roots the are surface the when ( near ordamaged expected for treeseedlings. found thattheannualBal ofsuckers mortality on increasing their thetime, suckers fare than better tree seedlings environmentsa inshaded undercanopy, 10 Kollmann 2007) suckering allows its torminalis wereindividuals Similarly, innort 1.3 m,only 43% Dume the centre itsEuropean r of ( difficult (HenryGirling,personal buteven communication) early as such planters SussexWest issuggestedit trees thatmanyfrom including originate suckers, those 94 Rasmussen &(2008) observed that Kollmann from the majority of 1679) advocated planting elm,and cherry, poplar - Accepted– Article 100% 15 years15 Savill (Crave 1985; root .

1989 Unlike coppice shoots, most suckers grow suckersUnlike coppice intact trees with canopies most from shoots, S

1993 - uckers are more thangrow seedlings tolerant shade quickly and inthe inheight

1

arise from roots 10 fromarise roots 2009). The distanceparent2009).from The ofsuckersthe isvery tree variable, of tr suckers

5 m(Germain1993; K.K.Rasmussenunpubl.data cited inRasmussen & colonises disturbed areasvegetative primarily bysuckering. ) . On

c ( ees were of sucker wereees of origin Evelyn ;

established treesestablished b Germain 1993; Germain 1993; & relative abundance . )

h France, h from Demesure suckers. showed three

, more occasionally , more (Roper 1993; (Roper Ducos 199

long

1664 sites i

- signs oforiginating from term persistence inotherwise ; Oddou ange,

Ll - n southwest Germany, 6 15 cmolder deep than10 ontrees ; oyd 1977) Hoebee Rich Rich Pleines 1994; Pleines 1994; in these 1991) there islessreliance onsuckering - Muratorio Muratorio (Nicolescu et al

25

. and althoughgrowth slows the et al - Even in . marginal marginal 30 m( . 2010). Investigation. 2010). suckers has shownthat - Propagation from canBritain Propagation suckers be in very Mush & Oddou Mush . 2006 et al Rasmussen & Kollmann 2004 at the northern end northern ofrangeat its the inDenmark, et al Favre the Britthe tic tic . ( ) areas tend S. torminalis suckersBauhus & (Pyttel, Kunz . 2009). and exceptionally110m( upto 2004 with anaverage of241 treestaller than Sea islandswas than 17%,lower - d’a s i uboptimal ) found that sh Islesweaklysh suckering where is it

nne 1990; Laniernne 1990; - Muratorio (2004) suggest (2004) Muratorio Rasmussen (2007) & Kollmann to be -

20 years20 (Pietzarka from “running roots”. Nearer Drapier 1993 Drapier produced vegetativelyproduced

sites (Rasmussen ( Rameau o

in Epping Forestin Epping nly nly considerably afterthis

Certainly, repeated 8

et al

stems b, 2007 ,

Mansion but most but most . c;

1990;

(Harper (Harper Wilhelm Cook Cook of , L

2013). ). Lehmann ed that loy & 185 & often and

d first

S. This article isprotected rights by All copyright. reserved. ratio of & (2 Gillon accumulate than otherof species levelsheavy higher the genus. metalsthefruits of in resistant tothe influenceof heavy metal ions. fertility.Itsoil alsolargely is indiffe responsefound. were that water availability growth. relationship between Monthly p at may somesites.This attributable to be (JulyAugust) and northern ofdistribution limit Ra droughts (V(C)) Sorbus torminalis (E) PHYSIOLOGICAL DATA species, diploid sexual ( et al reportedas stable Siljak f 2 (D) CHROMOSOMES 2009 nodal segments and Siljak rom Spain (Aldasoro n

Accepteds Article = mussen . 2012) and 2C = 1.49 ± 0.02 (SD) pg (1C = 728.61 Mbp)inB . 2012)±0.02(SD) andpg(1C = = 2C 728.61 1.49

, 34 - Máchová level of - Yakovlev Nutrients Sorbus Yakovlev

40.2 (Bailey The highest recipitation recipitation 2C = 009) (

2007 to ploidy ( ploidy

torminalis 42.7

. However, low availabil water . However, et al

et al

found that the leaffound that litter had et al

1.612 arebe limiting since unlikelyto ) found thatannualgrowth) found alongringa tree et al

performs bestondeep, in the previous yearprevious in the

, embryos . 2009 . 2008). . 2008). growth rates withintermediate were precipitation, associated which isf

spring and summer precipitation ofthe spring and summer . (2010), possibly. (2010),

. 2010). The genome The . 2010). is is thekeygrowth incontrolling et al Pellicer Pellicer was lesscriticalwas growth although for atapositive was somesitesthere

S. aria ±

0.002 can be micro . 2004) ) .

Sorbus torminalis was negatively tom correlated (

Chalupa 1992; Chalupa and et al airly

(SD) pg

and a t S. aucuparia . 2012). typical of rent pH soil to , and to , - o propagated ( f hybrid originf hybridBritish since riploid riploid 1C =792.180 water stress dueto stress water warm, warm,

The genome size of size genome The

1.28 B slightly larger than slightly Kol'Tsova other other spring and summer months in the monthsin spring and summer

ity andity high temperature attut is d

(Pellicer (2 moist fertile but moist soils S. torminalis - n 1.31% Nand 54.7

iploid but tetraploids but haveiploid been reported in vitro (Drapierandprobably 1993b) is , forest trees in France.forest trees in = 51) Grenier .

No latitudinal gradients climatic in Mbp

(1980) et al

has been

from tips shoot ean temperature monthly summer

) increased evapotran previous growingprevious season

& de M

. 2012). from material English transect from transect from

can toleraterangea wide of S. torminalis records that records that that of osnia and Herzegovinaosnia and recorded inSerbia by - arch individuals

52.5% C giving C a C:N 52.5%

is also tolerant s

two two

reduce

1993; 1993; , axillarybuds, the

S. torminalis other has been centre to

Malá growth. present yearpresent s show a very piration. suggesting main

fairly (Pellicer

of short et al David and the

can .

This article isprotected rights by All copyright. reserved. fluorescenceratio (F was investigated by Čaňová appears tobe m 1.57 than for F. sylvatica platyphyllos 7.2 μ compensation point reached 5.96 lower than leaves of many species tolerant shade respond from rapidly shade torelease Yet, t upon whethersurvival measured.growth is or (Oršani young,graduallywhen becoming light more the demanding ofitslife over first decade variation can be Řepka 2007 Á 1988 1999; being shade 1988). with an Ellenberg value orminalis lvarez Accepteday theslowgrowth inshade explain ( help Article 0.797 ±0.007 it canit survive inaseed b, ; mol CO mol mol m mol

Pyttel, L

Nevertheless, t 2013 Pyttel, & Kunz found Bauhus thatmean (2013) m Sorbus torminalis ć c anier anier

shade intolerantshade 2006;

; et al

understorey trees insouthwest Germany understorey trees Kotar 2001

rapidlyyears, growthits 8 slowsataround values reported forreported values -

- ) 2

suggesting a partial shadesuggestinga was light point The f tolerance. compensation demanding (Gonzalez (Gonzalez demanding

or

2 ± & 2013 Bauhus Kunz and et al s . 2009) associated with associated

m -

P 1 0.65 Acer pseudoplatanus

aganová - for shade leaves partially 2

very shade very

.

s

(SE, n= 40) 1990; -

1 was atwas μmol CO μmol v factor. Another isthat , resp ; /F he

Oddou o for light of 4 whereas all other British et al et ) was) 3.968±0.080

light requir

S. domestica 2007 is the most shade F attributed to ectively ehr a light ling/sapling bank with limited limited ling/sapling bank with . - . (2012)

intolerant ( intolerant - 2 a lowphotochemical efficiency:c

Muratorio , 2008

1993 other m – . These values are notaslow values . These in

-

2 low compared to other low compared to ) was higher than ) was than higher flux densityflux

s ) et al ; e , -

) woodland trees woodland

1 S and maximumphotochemical efficiencyPSII (F of ment but partially shade , , years150 wheneven old(Ra , . Darkrespiration

the fact that chütte shade similar tothe shade . 2010) Belletti et al similar -

of tolerant member of the genus in the British Isles the

Pyttel, Kunz & Kunz BauhusPyttel, 2013).

intolerant (E intolerant – 2001; 2001; . 2006; Rich Rich . 2006; S. torminalis

Pyttel, Kunz & Kunz Pyttel, Bauhus(2013) of 25.4 μ of low ,

shade classification , shade(Mü tolerant

to Monteleone

( est of S. torminalis Fraxinus excelsior 8.53 O . in tolerant ( Moreover ria de as domany F. sylvatica

mol PAR m PAR mol of shade and sunleaves ± 0.65 (SD) μmol CO ± 0.65(SD) μmol the Sorbus aximum aximum

vans

- et al annual has been variouslyreported tolerant tolerant R Sorbus

Sorbus ueda & Haralamb

1988; . 2010 , values for, values

hlorophyll ahlorophyll fluorescence was

Ferrazzini is more tolerant ofshade species tested; initial species initial tested; growth the ability with , shade photosynthesis

and ller

Mart

Fagus Carpinus

of - species testedspecies s 2 species are 5 mussen 2007) mussen ; Maděra Savill 1991; 1991; Savill

s , - S.

- Kroehling T. platyphyllos T. Quercus petraea 1

- í 1967;

intolerant species. species. intolerant nez for sunleaves torminalis

sylvatica leaves shade 2008 Slow growthSlow

de betulus noted , 2 Ellenberg

Tichá & Tichá m Azagra ) ( - . -

7 (Ellenberg . - of 0.73 and 0.73 B & This 2

, as do

.

ednorz s that depends sun The light Franz Franz ar lower , - 1 Tilia Tilia )

and

as and

was &

v

S and /F

to .

m - ) This article isprotected rights by All copyright. reserved. Accepted discerni late ofgrowth withasecond May, early June butsmaller burst variation least Poland inphenology,at in 2005). Aswithmany temperature precipitationrather than themaindriver trees, is ofannual October & earlyofNovember fall(Bednorz Urbaniak tothe October withleaf middle from Urbaniak 2005; Nov ripens late, rather 2005) typically east quite short;in tight those withpetals budto falling same are found tree onthe sequentiallyfrom late older indi Leaf ArticleVII. Phenology species testedZiegler& (Weiler 1981). acid triterpenoids have from (HuneckVerylevels identified been the bark 1962). ofabscisic low also been reported uniqueto as being 2015 Tzard palmitoleic, stearic, linoleic and oleic, linolenic acids campesterol, and andacids sitosterol, fattypalmitic, stigmasterol including myristic, vitexin, fruits A number ofcompounds phenolic (F) BIOCHEMICAL DATA

ember - (30 nmoll ). b i

of but synchronous between trees(Klein , Loukis, & Philianus uds I In trees n thes ble growth between2.5 months ble the in apigenin, Sorbus torminalis viduals (Pietzarka

begin t begin and and ubgenus Torminaria - fruits under the shade of a the under shade 1

Rich Rich )

o occasionally from lateoccasionally July from butusually have measured been quercetinluteolin, and torminaloside swell inMarchfirstdevelop leaves and may still bemay onthetree inJanuary still et al April April .

including 2010) , ern 1992; Olszewska & 2011 Roj

into May into Lehmann& Roloff

Poland it is only itis Poland 5 . Lea , f ha canopy, shoot elongationcanopy, shoot lavone O lavone caffeic acid and p and caffeic acid ve

S. t ves show autumn c autumn ves show ( Oddou in phloem exudate in phloem

been isolated been isolated orminalis (Bednorz & Urbaniak 2005). ,

Desassis - - (Barnola Muratorio Muratorio glycosides and theC

2009) -

14 days (Challice & Anumber of Kovanda 1978).

(Tsitsa from

. & , sterols such as cholesterol, - ,

coumaricacid ( Flowers Durand& Parmentier Snow & Snow 1988; Bednorz& 1988; & Snow Snow

ol Oddou et al from though to September

, thelowest ; Hasbal per tree ouration inmid the leaves in - Tzardi . 2006) has been reported to early through April toMay in in Sept . The flowering. Theis period - within awithin tree Muratorio ,

Yilmaz (Bednorz & Urbaniak(Bednorz - glycoside vitexin haveglycoside vitexin & Loukis&

, so that flowers from of an , e inflorescences flavanoids such as flavanoids

mber/October - - y treeof the14

Ozden & Can September to 2008) open

1993) 1991; . F occur in ruit .

Tsitsa and with

no -

This article isprotected rights by All copyright. reserved. 2004b, 2007 et al Krzakowa genetically isolatedstands in small, st Muratorio (Demesure The rate Bednorz 2000; high abortion of rates and (Termena (Bednorz diameter, 3(4) aided by sequential grains awithin 22 corymb. floweringelliptic, are Pollen 2004 and bees bumble including fliessmall the but 2005). andin dominant woodland edge treesunde (particularly ≥50cm DBH in trees high 50 Roloff ( starts Musch & Oddou Flowering comparativelyearly begins FLORAL(A) BIOLOGY VIII. Floral seed characters and Haralamb ands

Accepted Article- 70% of trees flowered in any one year.70% inanyof treesflowered production Pollen related is one size totree 100%, withamean . ; 2006, 2007 until treesuntil reach , reaching Rasmussen & Kollmann 2004b;

2009). Not all trees flower every year;2009). Notall treesflower every inSwit As expected from a largely O Flowersare (Hymenoptera, pollinated by

of self of utcrossing isthepr et al ,

1972

2002 - 1967; Klein div Musch ;

Oddou - . 2005). erse colporate, - ) Rasmussen Rasmussen ; fertilis

2 and - ;

Bednorz, MyczkoBednorz, & Kosiński &

Favre Muratorio

- Oddou large open flowersattract and twoSwiss 29% in & within andwithin between populations

Austerlitz - was found& bywas to (2007) Rich Price

Muratorio, Oddou

any 15 ation is usually ation is Pollen viabilityPollen -

of 75%. of d’a (Hoe - - and of mediumsize Muratorio yearsyearsgood30 seed oldwith inbred bees (Prat & Daniel 1993; bees& 1993; (Prat Daniel

& nne 1990;Savill edominant 2004 - Muratorio 2004 bee Kollmann 2004b

2005

Klein seeds ) although andfrequentabundant se et al

out ) although selfing become et al

stands

. - low

in life is affected byandis affected both temperature humidity

Oddou & breeding tree, 2007). mating system, produced

which physically hold more flowers physically holdmorewhich . Austerlitz , from 2001 r reduced competition (Odduoreduced r

with 1991 and occurs in trees <10 yearsand ( occurs<10 intrees old ;

- compared toother species

Rasmussen &Rasmussen Kollmann 2004 Muratorio ; a 2004, 2004, Klein

, 2004, 2006 , ; is rareapomixis ; other generalist other pollinators 0 ; a

-

Oršanić 2% maximum individual selfingmaximum of rate 54%

with stronggene( flow 2005; z S Demesure , 2006;

erland, Hoebee

. with variation between mother treeswith variation between Desassis due

torminalis Diptera and Coleoptera) et al

Angelone vary England around between 50 et al to

occurring Demesure ;

s . 2006). high Rasmussen & K

more common inisolated more . 2009;Pietzarka

- & Musch & Oddou &Musch

( Oddou has be been found to self Wojciechowski &Wojciechowski et al ed production ed production

Pollen Pollen et al every2 - et al incompatibility andincompatibility within the genuswithin - . 2007; . 2007; - Muratorio Muratorio . (2007) notedthat Bednorz . 2000 ) andalso is high - such as b 40 µm 40 µm dispersal ; - ollmann 3 years3 Oddou Demesure , Espahbodi , mostly

- Lehmann &Lehmann a Muratorio ;

does not in beetles Hoebee &

et al 2008) - is

. - .

et This article isprotected rights by All copyright. reserved. wider range whohad pollen and more long to 53trees). donors’) and var affected an of122 by to betweeneffective ‘pollen distance size trees(reducing thepopulation Oddou been (Hoebee demonstratesreduced butstill amountofpollen to4% had thatsmall this travelled 1.25 Muratorio for a largeSwiss 2008). adult of10 (range 1 a km dispersal likely. is slowly Austerlitz pollen movement some (especially with flying long bumblebees) distances. distance pollen movement with amean providing of6cloudalso pollen donors aroundgiven long thepollen and a tree, main patterns V(B) subject to been found tobe lower Oddou French compared and Spanish populations) to Chester Groenhof 1989; common sexuallyreproducing species diploid al

median distance Accepted Article . 2008 is possible . transported

- - Long dispersal thepopulationtobe37 was fromfound pollen distance outside than exponential with distancethan exponential with from thetree From microsatelliteoak stand studies innorthFrance, inamixed Demesure

Muratorio Muratorio ) et al . et al , greater Sorbus torminalis et al

Klein

Conversely, forConversely, female success, isolatedtreesgreater a diversity received of . (2007) seven DNA identified types differentBritain plastid in . 2007). :

preferential matingpollen dispersal between local neighbouring treesfrom . (2004) concluded that of the dispersal ( Oddou

Chester Chester i

> ability in pollen productionability between trees(reducing in theeffective population stand (Hoebee - &

genetic drift Musch &Musch Oddou et al (2008) (2008) of of 6 km Paternity analysisthat confirms

- Austerlitz 23)

c - visited visited Wirth Wirth distance pollen distance . 500 m . 500 - . (2001 Muratorio between (Bednorz populations . ( et al

Oddou found that of upto2.5km.

et al in Britain more . 2007) and lower than a

and and 2005). However, smaller and stand in more isolated ). - et al . (2005) suggests that . (2005) suggests Muratorio Inwith scattered areas populations,

et al pollen an averageeffective numberper ofpollen maternal donors loss loss - Muratorio male . 2007) and 40% . 2007)forFrance and a large standin (Oddou , seemingly because . 2003 has genetic alower diversity of - producing trees reproductive successreproductive geneticdiversity ( , This is This is consistent social with bees providing

Klein ) although

, the ( S. aria S. 2004

25 & long ,

S. torminalis

suggesting thatl ) found thatpollen exchangefound showstwo )

Austerlitz haplotypes throug identified

, and S. torminalis up to5%

- Myczko & Kosiński the distance pollendistance dispersal ofupto2.2 .

they attracted they Very isolatedtrees, however, S. aucuparia average is distance Kučer

of 172 of 2005

of

in mainland Europe: ová S. torminalis

pollen decreased than the otherthan the two , ong pollen distance gene flow

Oddou et al S. torminalis

pollina (Proctor,& Proctor Klein . 2010) .

Modelling by (sharedwith 2006) and - Muratorio , c tors from a

has inevitably

Desassis . 750m, pollen had h Europe byh Europe

s –

trees was this wasthis see more more - 39% -

6 km et al - local local had

with &

. -

This article isprotected rights by All copyright. reserved. backcrossingminor with not appear withthe tobackcross et al directions, and 2015). species England inclose proximityonlimestonesinnorthern Sorbus rupicola porrigentiformis al 1997; S. aria confusa S S (B) HYBRIDS ( Quercu higher inlight, than wind (G geneto track flow equallyge determining important in found inwoody plants Muratorio mediated in different that treesgreatera received ( diversity of pollen pollinators pollinators was strong tended tobepollinatedmore by Moreover, forfewer, trees. when local competition poorer seed by production due presumably tolower visitation pollinators. Prat & 1993; Daniel . x . x orbus . 2010;

Accepted ArticleST 30% aof seeds within . 2001 decipiens ) was 0.33, similar to theunderstorey) was to 0.33,similar species Fay

I into into s t

n cent orminalis Gremli

sp. (0.84) andsp. (0.84)

Pellicer geneflow are equally propagating important in genes the of b et al , determined bythe et al fruits S. torminalis ) .

The hybrids haveTheand hybrids lowerfertility andthey seed thanthe pollen parents ralLeinemann Germany, (=

. 2002; . 2001,2004) give the in France 75% of hybridsin France have

,

(Syme) Hedl. x S S

E.F. Warb. on thesame tree et al

. . x . x

is the femalediploid parent of

through seeds, andthrough for x tomentella semiincisa Oddou

. 2012 Chester in a Frenchin population suggesting thatrare Fraxinus excelsior Fraxinus

( - S Sell 1989; Sell dispersed of seeds . singlefruit torminalis ; Stace, Preston & Pearman 2015 - sensu lato Muratorio size of the canopy ofthenumbersize and offlowers et al

Gand.

Borbas S. torminalis parents

( . 2007; Oddou netic diversity. Prat & pollen/seed migrationpollen/seed as 2.21,one ratio ofthe lowest

, were found to were found in Franc

introgressed into introgressed into S et al and et al

(Price & Rich 2007) &Rich isevidence(Price though there of .

S. torminalis x

- (0.86). Price & Rich 2007; & Rich Price Muratorio Muratorio long Oddou vagensis . (2010) found. (2010) hybridization Daniel 1993; Daniel S . 2001 Salix caprea Salix , . x could alsoexist the S. torminalis e (Oddou - distance distance eximia hybrids in

attractiveness an of Crataegus monogynaCrataegus - Others have G measured b

Muratorio Petit Petit ;

shar Wilmott BednorzKrzakowa &2002). et al

genetic between distance populations

Kovanda - et al S.

Muratorio e 11%father and ofseeds thesame dispersal

Aas (0.09) than butmuch lower in . 200

torminalis

the as thefather (Oddou . (2003) used chloroplast DNA ) ) (Stace, & Preston Pearman et al et al due tothe occurrence ofboth , , Rich Rich

6). and subgenus S. latifolia ,

caused ofby introgression

. 200 Both pollen . 1994; of pollen and seedof pollen are another hybridanother of et al et al

individual S. torminalis ( Rich 6).

, . 2001 and and . 2010 occurred resulted in resulted Tormaria

Less isolated trees Rudow &Rudow Aas Theyalso found (Lam.) Pers ST et al Rubus

as low as 0.13 as low - b ;

and seed ). Robertson

- . 2014) to

Muratorio in both in both , including , including

; ( spp.,

larger Oddou S. . ,

. d S - o . x . x et - This article isprotected rights by All copyright. reserved. yearevery other seed. Evenoptimal warm under and dry C (C) PRODUCTION SEED AND DISPERSAL ( a species ownright inits Greece Malosorbus florentina and apomictic triplod hybridsnorthwestas Hungary triploid involving (2012, 2015) also reports KovandaLepší 1987,1988, danubial danubialis Lepší Vít et P. portae are reported aria Sorbus cordigastensis between not been diploid) In1989). cases other decipiens that theycontain f microspecies between both &Briggs Smith Wilmott

Accepted conditionslimatic affect the ability Article

S. sanubialis and - bohemicae A Sorbus intermedia Sorbus torminalis (Christensen 1995) (Christensen ,

S. torminalis is S. aucuparia

S. torminalis recorded

breeding member of witha the rare hybrid between (Bechst.) I (Bechst.)

and Prodan(Jáv.)

that involve

in Britain in parents and thehybrid( parents and

and S. torminalis

. U 2002; Chester

(Jáv.) Prodan(Jáv.) agg. ineastern Slovakia. S lavone O lavone

in

M. Lepší, P. M. et Vít . nder less condit nder optimal S. albensis dolomiticola rmisch in Petzold & Kirchner inPetzold rmisch Sorbus

(for example, (for ,

gametes and unbalanced subgenus

in Bavaria. N. Meyreported been byAas has S. aria (Zucc.)Browicz (= have been suggested been have tohave

S. torminalis and

S. barabitsii , native to the Baltic has also been female identified parent as of the Malus florentina , -

glycosides althoughQian ) and et al.2009,Velebil 2012,Vít (Robertson S. torminalis

and S M. Lepší,LepšíM. Boublík, P. etVít et al .

torminalis Mikoláš sp.nov.arisingMikoláš as ahybrid between S. barrandienica In various theCzech Republic, S. torminalis . S.

2007) of

Proctor & Proctor ,

x , otherwise unique to , otherwise Cs. NémethCs. and including

, houstoniae Sorbus torminalis

conditions in S. aria Newton &Newton Ennos ), Sorbus , et al

i (Zuccagni) which is which and ons, particularlyedge range,its at the of

S. eximia Boublík

region . (2008) gives. (2008) convincing evidence that

(Nelson Malus sylvestris aggregate Groenhof 1992 S. torminalis S.

S

x

.

an

Vit, M. Lepsi Vit, M. Lepsi etP. Rich omissa florentina likely , S. torminalis , isconsidereda hybrid tetraploid triple d

Kovanda (between S. milensis Britain

S. devoniensis C.K. S. polgariana -

)

Jones & Ko & Aria

, these would require u these would require

to (

Lepší,&Lepši 2004 given the geographic overlapgiven the Velebil Wilmott 1934; Wilmott

S. torminalis Schneid. produce

most treesmost produce only fr (1996). Mikoláš reports the male gametes gametes male , (Zucc.) Hedlund) hles

Briggs& Smith ). ).

has been de ( M. Lepší,K. Boublík, P.

hybrid A number of other A number

a hybrid a hybrid

as aparent

, (2011) as ahybridas of(2011) S. bohemica

). Cs. NémethCs. flower E. F. Warb. (SellWarb. F. E.

apomictic apomictic ; these include

S. bristoliensis S. aria S. 2012)

Nelson between

s, fruit and viable and these have s ( Jankun &Jankun cribed on theevidence S. torminalis nreduced

. , 2002).

Kovanda, Németh or native to - in Jones triploid S. , S.

it fruitsit named x named x

this is is this uit uit S.

,

S.

(i.e. s S.

This article isprotected rights by All copyright. reserved. Espahbodi but amean seed (Bednorz to experimentally self Rasmussen &(2004 Kollmann Espahbodi mg mg, to 990 edge0.07 and ofitsrange(2.2 ± number ofseeds perat fruit the centre of ofthe range spring ( found todepend inSwitzerland uponspring weather; fruit Switzerland Italyin 1987; Similar fruits 2seeds with and 2 norm is with a 0.13 (SE, 1988).Snow fruits in common butfew maturefruits & are(Rasmussen Kollmann 2007). produced years25 Poland ( innorthern 2 evenInyears 1993).andgood lessfrequently (Roper Croatia Switzerland seed have occur SE - 3 years3 apar

Accepted Articlethe Rosaceae, Rasmussen & Kollmann (2004 , givinga, fruit moisture content of 49 ,

n and morefruit per(Davi seeds n

duringfirst pollination. after 8weeks the n overall

= 4000) in Iran in (Espahbodi = 4000) - Espahbodi that year.that unstated 1170 mg& (Rasmussen inDenmark Kollmann 2004 Sorbus torminalis Fresh from fruit 765mg varied ( massinSpain of Fruits become (Belletti norms havefound been acrossrange of the 1 seedfruit per , Myczko& Kosiński n et al et al

= 27) in the Wye= 27)inthe( Valley 0.17 to2.63± (Bednorz In the n England, t (Barengo

) . (2007a)noted a correlation positive seed between massand sound mean Iran,. (2007a)noted that in mass wascorrelated fruit negatively withDBH. mean of 1.9 ±0.076( mean of ,

dry dry seeds per fruit seeds per This may also explain whyThis may Rasmussenalso & explain Kollmann (2004 Monteleone et al mass of20 mass

- . 2007a darker as they ripen and soften after asripen after anddarker they soften 2007a pollinated) yielded heavierfruits.pollinated) slightly , butsamples offruits3different from trees - , 3% with 3seeds were a (butwhich

s Rudow &Rudow Schwab eeds ; Conwentz 1895 Conwentz

) buthigher have numbers reported; been u

& Hoebee umber of

b - in 2001 2006 are ) 21 mg Ferrazzini

recorded 1.9 ±0.07 et al s 1976). As an example,t an s 1976).As some of the biggest in thegenus biggestin some of the ) n . et al . 2007a). Dry mass fruit

Published d Published = 174) (Price & 2007). Rich appears but only

fertile seed per seed fertile fruit - 59% 1987; (Herrera

. 2007 that fruits resulting fruits from outbreeding(as that opposed 2008) –

). 2001;

SE, SE,

At thenorthern itsrange,flowering end of is

typical b

0.6 ± 0.6 ).

) observed) and occasionally n ry are extremely massvalues seed variable

Warm summersWarm

Oršani S. torminalis unstated n

Hoebee S. torminalis = 30) in Leigh= 30)in nearBristol, Woods 0.17 in2002 (Herrera 1987; Herrera 1987;

ć bletting, usually

et al b ll shrivellednon ll and 75 – was heseeds numberfruit of perwas ) and

et al Aldasoro respectively). - in Spain in Spain 85 . 2006, 2009) . 2006,2009) , from Spain

found from tovary 1.37± %

1240 mg (range 1240 mg 890 .

5

compared tothe north l following a in 2014 in 2014 ( , similar , similar In Staffordshire, the

ead tomore abundant - 2007 abortion of immatureabortion 6 in Poland 6 inPoland Aldasoro Aldasoro Aldasoro Aldasoro varied from 345 et al p to4 ) found found ) b by showed 8 . 1998). ) found a higher in size

to

As iscommon

fallingevery to seeds per fruit frost

cold, wet Iran (Herrera et al - and 1.28 viable). et al et

to (Snow & . 1998). -

. 1998) 15% of S. aria ± 0.16 - 1710 - 390 r ed

This article isprotected rights by All copyright. reserved. temperatures andgro longer & comparativelyand most late monogyna especially tobrighter incomparison fruitsnearerground carried theas in suggested,the including that Rasmussen &2004 Kollmann datatreebirds (K.K.presumed citedwere Rasmussen tobeby unpubl. in and taken 20% only the northernrange of ofthe edge S. torminalis Juniperus canina oxycedrus, Rosa Her endozoochorous ofseeds dispersal likely is tobeineffective feedfruits of onthe philomelos through1989).gut the (Herrera The abundant high sclereidshelpprotect and fruit content fibre during the seeds their passage such as readily eatendespitebeing by carnivores ( carnivores 10.9 ±12.4(SD) seeds of ( Demesure (L. the range tobedue suggested here tocomplete and were maturation so than those from the (2004 seed massremained constant Ingermination. were Croatia, fruits inlength toincrease seen altitude with Iran,mean fruit mass in 1987, Conwentz

AcceptedKollm Article ), badg rera (1984) <20% foundrera that b

1989) ) observed Although of a bird number Fruit Juniperus oxycedrus, Ficus carica Juniperus oxycedrus, ann, 2004a). In this context, In context, ann, 2004a). this er er -

Musch &Musch Oddou

byHerrera (1989) ( Brehm 1895) Snow & 1988). Snow Snow Moreover, at

Meles meles

are dispersed

fruits were takenand by only blackbirds these1.5% just formed observed

and that while fruits atthecentrerangethat while of of ) , blackbird , blackbird rang S. torminalis domest

that l

e edge,e theseedswere larger. but foundnocorrelation between and size percent seed

(L. S. torminalis primarily by primarily - wing arge,

) and stone Muratorio frugivorous have passerines

. brown of colouration thefruitsattractive not tobirds is ic pigscattle 1993). (Roper and (Orša b Of recovered 556seed only5.2% were damaged. ( ). Various causes T

. of large

S. torminalis

season ( scented, fibrousscented, merula

Snow &Snow 1988; Snow such as redwing and ni ć seed dispersal

et al

Sorbus torminalis 2004 per scat were found in carnivorous mammals marten marten

-

due to climate change due to

low in protein andmineralslow inprotein sized fruit fruit sized L.

and . ) ) aided byaided ) 2009)

and robin and

in Denmark

Sorbus domest Sorbus Martes foina Martes the northof the

for poorhavefor dispersal been birds by fruits .

( By contrast, by birdsmay from plants such as Turdus iliacus Turdus The smaller fruits theThe centre from smaller of other wild other such as ( Herrera 1984,1989) that fall to the ground whenthat fallare tothe ground ripe Erithacus rubecula

migrated bytime this wereIn taken by birds. study, this ,

75% of fruitsfell from75% simply the of its .

( In Spain, southeast I

Erxleben ica

the such as n southeast Spain, n southeast S. torminalis . range A further factor limiting bird Rasmussen & Kollmann

in addition to in addition improve with improve scats of scats

L. . This included species. This S. torminalis ) , songthrush red ) lower lower Ruscus aculeatus, Ruscus ( Herrera 1989; Crataegus Crataegus fox fox the abovethe (226

were L. ,

s of Vulpes vulpes ) tar

( age S. torminalis - are to known the Herrera Rasmussen 499 m)but their for example, a mean of ch smaller

boar fruit ripen

higher ( , content. T.

diet. At

.

s

This article isprotected rights by All copyright. reserved. yeargermination,showing first higher the germination. the thatthelower the delayed year.By contrast Espahbodi ( yearsBritain twoor more of continental winters, followed germinationmore was uniform obtained by alternating two resulting inmost stratificationmoist at 2 embryo dormancy 1993 viabilityseed sorted can sound of (1977)Wilmott found thato Few GERMINATION(D) OF VIABILITY SEEDS: Wojciechowicz (2007a), andontogeny and theepidermis discussed by development is fruit & of Bednorz (2004), Musch & Oddou seedlings inthe torminalis found at 2004 torminalis underneath f where presumably is this largeoaks roostafter because thrushes birds does, however,appear France, tohappen;in seedlings of shreddedattached and skins pulp ( dispersalthe seeds thatmany is are taken of by AcceptedRoper 1993 Article Pyrrhula pyrrhula fully ). ;

Orša

The m seedMost Maciejewska a by

formed formed

mean distance fruits ( 150 ni et al ) 18 ć . orphology of fruits andorphology seeds fruits of

Delayed germination hasDelayedfound dryIn alsobeen inhot Iran,climates. - et al - 300 . (2007a) found 42% germinationgermination. (2007a)found 2000and 42% thefollowing 14% in 20 weeks at 3 20 weeks (2009). centre - , seeds

Muratorio S. OddouS. seed is , aided byaided , for seeds sownin 2001,germinationyear wasfor 20%andthe seeds 26% after, . 2006; . 2006;

(L.) m in m in radius.But disperse someseeds muchfurther since dispers - RutkowskaBednorz (2004), &

s of of germination takes placeproduction springfollowing but inthe seed - germinating

) 3 are produced and

and to a lesser extent marshand toalesserextent tits

of the ° may beforeis elapse dormancy Var C forC ( 3months - ed Muratorio, 174 m nly 5% of seed tested inDerbyshirenly seed 5%of we tested 2004 chemical inhibition fromchemical inhibition

470 ha forest originated from470 ha outside thestand forest originated

° , over shortover distances.

C Bekci &Bekci Dinçer

; ( to the tree(

easily be >60% and>60%easily be Muller Oddou from

within four months (Var fourwithin months

personal observation

the mother treethe mother - & the Muratorio Muratorio Devillez 1979 of

Laroppe Snow & 1988). Snow Snow viability of viability of S. torminalis

seed predators,particularly bullfinch seed 2010

In France

et al ) 1993 Debnorz . the seed coat, reach 94

Sorbus torminalis , with ( seed collectedseed

; Devillez . Poecile palustris broken

). 2004). -

are de

week periods atweek 3 periods

Under the long, cold conditions theUnder long, , cited in

,

Bekci& Dinçer genetically established seedlings were et al S. torminalis - 97 , allowing germination , allowing s % ( %

cribed by Aldaroso cribed by Aldaroso . (2006b) and. (2006b) Bednorz , Some limited dispersal by limited Some re

that can becan overcome by that Fraipont & Tissot Oddou

Muller fertile

is usually low is usually

related clumps of seeds have a deep

(L -

. Muratorio are common ( 17% of eeding on & However, Demesure

.) ° 2010) C andC 20 ) Laroppe , leaving the , leaving the

es although ; At

et al

1980 et al - S. ° C . in . S. ) , This article isprotected rights by All copyright. reserved. 35 cm germination 4.9 upper). upper),cm lower,and height 41 (52 cm upper) yearslargerafteras two from the lower ainto common garden1500 m at emergence (38 altitude early performance October. reportedgoodgermination when intact whole (1993)germination found decreased >50% if fruit by (Cook 1679; Laroppe Parmentier storage, sowingdepth exposure ofhydrated 20 to seeds - germination of mineral leaf orthin soil litter. Nevertheless,can seeds drying withsome cope since that germination whenwas wasthebettersupply seed with incontact highest water of mid (2003)Efthimiou found thatletting‘dry seeds considerably’ youngseed or pre wassoil mulch which covered suggeststhatburial witha cm ( stratification highe 5 1999 Winkler

Accepted Article° deep emergence increased C - Octoberin Decembergermination<1% and resulted sowing in r altitudes r and

diameter. G c

(1993) and(1993) ermination is . 16kmapart Espahbodi Espahbodi

7, 8 or 10% moisture ( 7, 8or 10% moisture

Espahbodi ( . The highest . The highest 1993 Roper 1993 ( Orša ) and somayInIran, northern burial. from benefit burial of seeds %) than the higher site (45%). Survival of 1 %) higher thanSurvival site(45%). the S. torminalis site (89.7%) thansite (89.7%) (80.5%). thehigher in

), - emergent emergent Croatia, period seeds andrequired of dormant were a more longer ni between seedbetween from Drapier

Kausch et al ć

and otherand al et et al improved in Iran in

measured . ;

germination rate ( Yagihashi . 2007 . ( - 2009)

( 1993 seedling Blecken von Schmeling &Blecken Kellner vonSchmeling was foundto 2007 .

(Espahbodi (Espahbodi Trees from the lower siteshowed thelower Trees significantly from lower seedling b in vivo

Muller - if the fruitremoved is ) b was 25 . a

found geneticfound ), by root collarby diameter root Seeds inthedark germinate will following stratification ) also foundthat

, °

Kausch

under hot and dry hotand conditunder Hayashida

C ( C the trees growingtrees

recommendat & Muller opposite, beingopposite, inseedlings significantly higher be et al Laroppe (43%) (43%) - Blecken vonSchmelingBlecken similar after two y aftersimilar two fruits were afterfruits sownimmediately harvest in . 2002). This effect when. 2002). was found the not theyalso

allyfixed & & over two yearsover two was inseed

Laroppe Miyamoto 1993) DBH of at Seedlings from the lower ions canBarnola be found in were

1600 from around theseedsfrom had more branches (8.4lowerhad more branches

. better

(10.98 lower mm differences in germination anddifferences ingermination - Dormancy can be reDormancy be can left intact left

year 1993 - the in Greece 1800 mand

ions 1998 ears whetherears stored at 4 - imp ( pa old 2006) ). . rent tree affectedrent tree Further d Further ). , and Certainly, roves waterroves tothe supply seedlings , a Muller & LaroppeMuller & (1993), . lthough Röhrig lthough between in collection

Röhrig (1972) noted (1972) Röhrig at at

2100 etails of site

from 25 trees Muller &

Takos &Takos transplanted transplanted tosowing prior

- site , 9.54 induced by induced

up to - 2300 m ,

were also Durand &

seed mm 1.5

( site 1972 ° C or C - 3.5 - , )

This article isprotected rights by All copyright. reserved. & (, the Malvern Hills on on theleaves Dysaphis aucupariae linked in mainland Europe common in torminalis (Coello shoots (Fabricius) Ceph (Hymenoptera, ( Torymus druparium variety ofchalcid wasps 2011) and ofmitesAn extensivelist feeding on Eriophyes inthe familymites marsh are predators tits knownseed (VIII(C)). mammals, rodents anddeer.Thrushesfeeda certain to IV,As notedin and seedlings saplings (A) diseaseIX. Herbivory and cotyledonsare 2004) are similar Seedlings of (E) SEEDLING MORPHOLOGY Henry Girlin

AcceptedJerzak Article S. torminalis FEEDERS ORPARASITES ANIMAL .

to extensive prematureto extensive fall leaf The root starts witharapidly system developing the taproot time andby the Insectsfeedingon recorded Sorbus torminalis

2010)

Ripka &Ripka Szabó torminalis torminalis is to Hungary (Ripka 2001 S res

(Lanier 1993b). g the later true leaves,the later but true ) is orbus , personal communication). , personal expanded, thetaprootexpanded, can 8 be et al .

tricted tothe (Table 1) , the type location (Emmet 1976). Eriophyidae . 2013).

at the beginning theat the su of torminalis that

and ( B uckton) that forms ( can result trees inexcellent seed

Hymenoptera , notably is fairly toinsect pests(Chaluparesistant 1992) buta Phytoptus pyri

( 2010) A numberaphids (Hemiptera, of

Rosaceae A number of miners leaf

have also

are found on idae) can causeidae)canof inconsequential withering . ) and a ) and

Stigmella torminalis torminalis S.

an occasional leafan occasional two small, ovaltwo small, cotyledons

S. torminalis

torminalis initially (

of of , Lanier 1993b or phids ofthephids genus Tenthredinidae

var. S. torminalis Sorbus The shootsawfly reddish to yellowish to reddish pseudogalls rolledor twisted S. torminalis torminalis - mmer, sometimes inlarge number,

14 cm long14 cm (Schüte 2000). without the characteristicwithout

in the British I in the British

(Table 1).(Table

in Hungary (2010, isgivenby Ripka Phyllonorycter leucographella ).

miner in Poland (Walczakminer inPoland These include the wild serviceThese aphid include the areby readily browsed small (Lepidoptera, micro extent on extent )

that that are restricted to affect

only rarelyonly set

in Britain (Table 1)notabl Dy

Aphis pomi is restricted to Aphididae s aphis S. torminalis sles are given 1. sles are inTable

(Fig. 4) fruits J anus compressus anus

are frequently observed , and bu ) . The primaryThe leaves .

De GeerDe ting l found on found

obing obing

- S. mot

. any S. torminalis

Most notableMost is S. torminalis torminalis ll number of number (Amann (Amann finches and hs)

, viable is also and are

Baraniak S.

(Zeller) are found

y

seed

in A .

This article isprotected rights by All copyright. reserved. valleysavoiding wetlands start ofthe across Europe( haplotype suggests distribution that refugia yearsBPalthough, (Godwin woody 1975)other retreated have like most will it to species, records of that s early Fossil species asTorminariagroup of theTertiary the period. found been have in Sorbus X. History torminalis Enterobacteriales are importantoffireblight hosts ( sedis Kausch galligena Armillaria Sorbus torminalis (C) DISEASES al Andrianova Phycomycotaand Ascomycota) found on torminalis torminalis S. torminalis Isles national surveys a number despite of Yıldırım Sorbus torminalis (B) PLANT PARASITES outh Accepted Article. (2009) reportsaquatic ofonseeds on26 species fungi of found S. torminalis ) w

causewilting

est - was arepresentative ofamesophytewidespread temperatefloraas warm thatwas further further Blecken vonSch

& ) and ( apple scab

and centralfrom Pliocene China theMiocene and S. torminalis by Grove(1935) (Ascomycota, (

sp Holocene, Anon

Karakas (2001) a(2001)

p. in the BritishI south , Demesure

which

) evolved indry, woodlands open

2013

is prone toinfection by fungal ( diseases w in

ill supportill and diebackand ofbranches

2012) in the warmerclimatethe ofthe number offungalon number parasites S. torminalis

).

can be fatal 1993c). (Draper

Europe during the last glaciation. duringEurope Evidence thelast chloroplast DNA from in the British IslesInterglacialin the British warm from the Hoxnian

meling 2000). Bo -

Musch &Musch Oddou (Roper 1993) particularly inthe warmer between period 8000 but Venturia . sles is given in Table 2 given inTable sles is

tryosphaeriales Badurowa S. torminalis the Erwinia amylovora amylovora Erwinia epiphytic hemiparasite

is thought have to re slowly

S. torminalis inaequalis

& Species of of Species

. A S. torminalis Badura (1968) - ) was recorded) also fading leaves of on

Muratorio Muratorio has been not inthe British reported as a host list of list ) Mediterranean S. torminalis S. use

survived inanum

(Gabri . Verticillium Verticillium

S Europeancanker of ( apple fungi Phyllosticta sorbi S. torminalis . torminalis (Burrill)

2004 leaves insoutheast Poland,

ė

list anumberlist of fungi (primarily and moulds slime lian mistletoe mistletoe Drapier 199 ).

(Gabriė (Butin 1989 : As

1961) and certainly will include Proteobacteria - Nees colonised along

in the the as aminorhost S. tormi berdi of Viscum . There are. There possible

lian 1978) climate warmedatclimate the (Ascomycota,

West 3b ). ) including n Sorbus en fferentrefug alis associated ,

d. v

Nectria and islikely it . c

in Poland. warmer . Czeczuga

( ar. 130 000

Table 2; ( and species S. Turker Incertae with - ia river river S.

,

et This article isprotected rights by All copyright. reserved. AcceptedGirling, (1994). torminalis planted onshooting est importantthe most forestry2000) andregardedas is genus the treewithin as astandardcoppice systems forvaluable its tree in torminalis However charcoal 1821; Drapier (Phillips c; 1993a, rather than Sorbus torminalis USES humans ( although was it planted, extensively consistent extensive with local occasional movement pollen and long introductionforits fruits ( Articleorigin 1975). (Godwin Dorset Sorbus torminalis eastedge(Kevin aroundcommunication) thenorthern ofMorecambe Bay Walker, personal last century Birks 1982) (Baker 1885; Cumb Woods, (GodwinNeolithicBP at around1975)and pollen hasRoudsea indeed found 5900 been pollen 1997 woodland (Roper 1993) an torminalis years5000 BP ;

Rasmussen during (Salisbury & 1940) (Salisbury Jane

Thereare suggest Due The timber of

personal communication) andpersonal although communication) , other native are species better Demesure

edges is w in forestrygiven by is Nicolescu being might rarer it than otherwise be. for to the tree’s the Holocene. It the Holocene. been knowntohave is

ria the value of wood. its the value idely be to one held ofthe later when d like and

in

& was forme charcoal hasbeen -

Musch &Musch Oddou mid

Kollmann 2004 on it reachedasit faras north S. torminalis other other ates for its fruits (Conwentz 1895). A reviewuse of for fruitsates (Conwentz its ofthe

steep - comparative scarcity,arecomparative more ofits thansporadicrecords there no ions that its distri ions thatits Flandr thermophile r Roper 1993) ly widely coppiced possibly toreadily widelyly coppicedability toits due sucker south ia

but n

has a fine yellowhas afine with an colour attractive (Henry sheen deposits deposits - there are doubts facing b shows little evidenceshows little oflong recovered - The been woodhas w ). Muratorio but other populations remainbut other populations

s in both respectsin both

. However, the genetic. However, the diversityEurope across is (II(A)) will havewill survived(II(A)) bution some in p Demesure slopes slopes (6680 et al southern (Ku southern tree species to tree from the Iron the from Age fort hill it is it . (2009) and by. (2009) inagroforestry Guitton 2004 I - wherecompetition was reduced n central Europe 5150 BP) now - as to larger pieces of Musch &Musch Oddou ). present

and this may account forand may this account

little used little whether idely used foridely and usedfirewood arts of was Europe by enhanced . It. in the British Isles in the British the since arriv became extinct hereextinct became inthe - distance movement by distance e the wood the wood

it in Britain, on thelimestones tothe in the British Isles in the British tzelnigg cooler periods cooler timber timber - was frequently planted distance distance - Muratorio at at Sorbus ( was oflocal Maiden Castle, 1995) M ller in c dispersal

.

S. 2004 ( entral S. . It also was Kollmann along Sorbus et al ). , and , .

.

This article isprotected rights by All copyright. reserved. decoction ( expectorant as an torminalis widespread they antioxidant lower have propertiesfruits ofmany than treesincluding Erol Rivera 1999; Loukis 1991; and vasorelaxant and 1955), name. hence Thebeen have specific fruits also the make board spirits, of cider. make the also been raw(Mustafa eaten and has been germanica (Sowerby& Sowerby 1878) (Kausch drying acrossBritain, Europeincluding Kellner 2006; Blecken von w instruments, measuring instruments beautiful wood 2004 Franke Europe

Accepted Articleas

15 as anti sometimes dyedsometimes ). I ).

drink wine th w

, ( The formerly Innin Chequers drunk The fruits

it is one of the most highly isone oftheit most n Evelyn 1664;Roper1993).Evelyn as hung outside to advertise presence(Mabeyas its 1996). hungoutsideto

Dagenbach & Hauff Dagenbach & - century (Hellwig 1997). T Blecken von Schmeling &Blecken Kellner vonSchmeling 2006). deed, in

he namefruits derivedfrom possibly he the - ) , brandy and, brandy (Facciola vinegar 1998). leaves s

diarrhoeal, . astringent tannins Sorbus aucuparia . There isalongeconomic InGermanythe fruit was Schmeling1994 also Tsitsa Rich Rich in

medicines

1900 at theExhibition Paris World 1900 are have eaten been boiledtotreat been processedand intojellies the world the world et al et al - Tzardi, Loukis Tzardi, & Philianus

and used asand anebony substitute hi

diuretic, anti . 2005; . 2005; gh in vitamin C gh invitamin . 2010) . , and also as antioxidan , and also as , often treated (Düll 1959)

1990; Uthoff 2002 Tuzlacı b

et al in (Serteser K ltr ; .

Klumpp & 1998 Kiristis

It usedwas crossbow for alsowidely and making rifle the fruits and for cogsand for However, the . 2012)

priced timbers and ingreatpriced timbers ( demand - , arrows It used alsobeen forof has wine themain presses screw inflammatory, antiinflammatory,

& 1999) Erol 2007 macerated in a sweet porridge in macerated andsour

s, particularly inthe Weald history and et al .

. It hasbeenused for have been Seeds were eaten in Lower were Saxony in eaten Seeds certainly since ; in a similar way to in asimilar

, and for veneersfor , and have as food,particu been used Pietzarka . 2009;Olszewska 2011) since it shrinks little and on shrinkslittle does split it not since of use of the of ‘chequers’ fruit main main ;

1992;

diabetes anddiabetes Demesure cons . t ’

Despite the abundant uses, thename,Despite theabundant uses, agents ( use tobrewdrinkan alcoholic , used culinary hasbeen fruits useof the , ( S. torminalis

erves Germain 1993; Lanier Germain 1993a; 1993;

Lehmann& Roloff Klumpp & 1998; Kiristis - diabetic, broncho vasoprotective, ;

Kausch used to - (Facciola 1998) Musch & Oddou Musch Grigson 1955; , wood settle

fine furniture, musicalfine furniture, the medlar( colic ( against choleraagainst

They have also have They been used to - Blecken von Schmeling&Blecken

tormina and Kent, and - was carving K lt . In Turkey, voted the most Drapier 1993 Drapier

r 2007 the muchmorethe 2009

Tsitsa l or colic (Grigson

; the fruits have ar and turnery to make a form

- wherechequer a Muratorio

l

(Düll 1959) y bletted once and )

) and a Tuzlacı although - S. Tzardi &

dysentery or flavour

Kausch b ; stocks , t

o & and -

- ,

This article isprotected rights by All copyright. reserved. al Blecken reproduction large that thedecline incoppicing importantly,most changing management practices and increasingpopulation size and the rarest trees al the Czech Republic considered IUCNConcern using threat criteria ( personal communication), tall treesgoodare ofbeing form ( such as France eastern species (Keith woodlands and since consequently the1940s light theabundanceofmore reduce coppiceride andmanagement Indeed, ismore it improved of knowledge thespecies distribution ofthe due surveys totargeted 1993). (Roper periods has apparently the rarest trees forest inGermanyofthe making up<1% area Sorbus torminalis XI. Conservation ‘serves’, the‘serve’ O plural from of ‘service’, is Drapier 1993 Lef

Accepted. 2004; . 2010 Article ex situ è - vre scale p scale

The mainthreats to Fewconsidercountries European 1930

)

von Schmeling1994

et al Oddou and in Poland and inPoland

conservation ( as not opulation decline

- ( . 2012) 1960 and 1987 b E Rö

et increased i increased in the country (Szymura 2012) ; Ewald ndangered an indication ofusefulan its indication -

hrig Muratorio, al

likely to have declined in many Britishlikely tohavecessation woodsdue declinedinmany totheof

is by nature its a . 2009). T (

Stoltze Stoltze

for example inGreatfor where Britain Least has as classified it been 1972 , , starting in1988(Allegrini it Zander Zwierzyński n frequency,Englandand such as Scotland between in has been

or leading becoming tothe species increasingly shrubby. ; S. torminalis

Franke Klein & - with here is N a 88 (Rich& 1996) Woodruff ,

; of frag Pihl 1998 Pihl which has toincrease been shading inBritish shown ear ear Kleinschmit 1998 & preferentially S. torminalis

a

Jander & n uncommon T mentation), increased browsing , protected by law protected by law

concomitant concomitant Dagenbach & HauffDagenbach& also hreatened (Cheffings & Farrell&(Cheffings 2005). Nevertheless

ld Austerlitz

& E

are be woodland( considered clearance to ; evidence that it is that it evidence nglish

Bednorz S. torminalis ness but rather is derived from the Latinness rather but from isderived the 1994) Pietzarka , ,

in harvested and( notreplacedharvested

and state bodiesand areactively engaged in

increased . ‘syrfe’(Grigson 1955). 2005; Angelone

tree places inmost rise in It has alsobeen thatparticula suggested the northof range, its the

; , 2012).

. Giard & Mariel M ller , As notedinVI(E), m

since 1946 Lehmann& Roloff

to beinneedconservation efforts of denser

1990;

fragmentation and reducedfragmentation and although this mayalthough this to be due et al becoming rarer

(Schü

high forest has led foresthigh has . 2000 Drapier 1991; Drapier 1991; (Bednorz 2007c et al , especially by deer,

1993) ; f te 2001). te . 2007; ; Oddou or example, it isone of example, it or

central Germany

2009 anyconsiderauthors ,

and centraland Europe in some areas in Henry Girling

Rasmussen

In some areas it - ; , Muratorio Kausch

Úradníč it is it )

- , the two

demanding directly as one of as one of Sorbus reducing -

ek and, in situ rly and et , via , et to

This article isprotected rights by All copyright. reserved. (2009 specific habitat plans management (Belletti done indiscrim et al provenance countered collect byseed planting may limit ( species such ineffect as beech, simulating conditions thinning (Müller (Úradníce Schmeling 1994 Fehrbe inforestryextensively used 1993d; more (Drapier 1993; the high cost ofse the species reduce reduction inopencoppiced woodland Europe across reproduction deep shade. Bauhus reason.this and stimulating suckering 1993), undoubtedly and (Wilhelm inthepast has for planted been readily spreadthrough incoppice by suckering,aided roo oflogs theskidding damaging coppicedso much because theopen needs woodlands it conditions not serious threat to VI(E)) &Kunz Bauhus (2013) charge counterthis bythat stating Schü (F Lehmann & Roloff 2007 Kleinschmit

Accepted Articleavre . 2007) although this will dilute genetic dilute . 2007) will although especially these in if this populations uniqueness te ; ) sugges Paganová -

and d’a Increasing of isolation 2001;

(2011) found frequent recruitment, (primarily by suckering) over the past 80 years found recruitment, (primarily(2011) frequent past in the 80 by suckering)over ,

Monteleone

k 2010 gene flow nne

as

Perennation is byPerennation is intheshade: suckers as possible to ensuregenetic diversityas maximum to possible (Wirth However, tolerating despite conditions shaded

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a , Ammer &, Ammer Nüßlein S. torminalis 2007 ). atelywi ; Demersure ; Demersure s eds (Jacobée 1993), has limited its planting, its haseds despitecalls limited 1993),for (Jacobée frequent to it

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This article isprotected rights by All copyright. reserved. Amann, G.(20 Allegrini,&(1993)D. M. Mariel, Perpétuer Giard, C., ressource et la en améliorer locale Alford Aldasoro, Aedo, F.P.Aldaroso, Garmendia,Hoz, & C., (2004) J.J., F.M.,La Navarro, Revision Del C. von Verjüngung und Häufigkeit Verbreitung, (2011) M. Kohles, & G. Aas, A References kindlya draft. commented complete on Ripka Itranslating am documents. various and writeBullard ofFellowship as Forest, a part atHarvard and Harvard Acknowledgements and poor seed ( dispersal eastwards m maincompetitors suchits as seed orchardsgenotypes. to protectthe ‘best’ protect isolatedpopulations and oftheestablishment the ‘best’ the84 archives ofclonal and

Acceptedas Article , G., Maier, J., Baltisberger, M. & Metzger, S. (1994) Morphology, (1994) S. Metzger, & M. Baltisberger, J., Maier, G., ,

for their unstinting ,

Climate changeClimate may benefit Melsungen,Germany. alisiers:projet un Haut Wolfe systematics (, Europe Africa: and in and Rosaceae) morphological inNorth analysis Monographs of 59 cordigastensis torminalis and cytology D University is gratefullyUniversity is providing for acknowledged toread, and time space think .V. - Sorbus J.J., J.J., 71

a

. (1984) y due amongstbe individuals limited topoor seed widely spaced production

Publishing, Aedo, 04)

subgenera and Aria Torminaria(Rosaceae (L.) Crantz. Bäume und Sträucher des Waldes und Sträucher des Bäume .

Systematic Botan Systematic , A Colour Atlas of Fruit Fruit of Atlas A Colour

C., C., 69

erdcin f yrd between hybrids of reproduction (Kordigas , 1 Hemery Navarro

sharing information of London, UK. - 14 Fagus sylvatica

8. - Botanica Saônois. Saônois.

t , C.& Muñ et - eler) n e nrlce Frankenalb. nördlichen der in Mehlbeere) al S. torminalis very gratvery y . 2009 ,

23 Revue Helvetica , 189 ). Pests: Their Recognition, andControl Biology

oz Garmendia oz

eful eful (Paganová 2007)

Forestière F Forestière - 212 , and , and

due toit being drought more tolerant to , 104 . David Colman

. Neumann especially ,

195

, F. (1998), F. obs aria Sorbus rançaise - – 214.

Maloideae). but migration northwards andbut migration - for Neudamm Verlag,Neudamm to

, HenryGirling and Rodney Helliwell who Rodney the help provided in help the , 45 The GenusThe , 383 L) rnz and Crantz (L.)

isozyme variation, variation, isozyme Systematic Botany Systematic - 387. Tuexenia Sorbus

Sorbus than Géza

,

31 . S. , This article isprotected rights by All copyright. reserved. AcceptedBednorz,L. Morphological (2007a) fruits and variability ofseeds of Bednorz,L. Morphological (2006) of variability ofleaves Bean, (1980) W.J. Battut,de& A.,Grenier,March,G. (1993) E. Micropropagation de Barnola,& Durand, P., Pa P. Barengo,& N.,Rudow,A. Bamberger Balandier Baker ArticleBailey, J.P. Badurowa Austerlitz Anon (2013) (2) 7/20 PM Angelone, S., Andrianova , J.G. (1885) , J.G. Poland. Poland. Poland. Revue la morphogenèse l'alisier torminal. de and Schweizer Alpennordseite Forst vegetation composition cover inaff crops Sorbus Reserve fungi Molecular Ecology Sork, V.L.(2004) Mol population (Ukraine, Crimea). , P., , P., , .

, F , M ecular

Eidgen ,

U

. Kay, McAllister, H.T.C.G. (2008) & numbers Q.O.N., Chromosome Rich, in T , Dick on Zeitschrift

. & . . Forestière F Forestière Frochot Hilfiker,

.

L.Isles. British (Rosaceae) inthe V (1990) Elsbeer Ergebnisse des Dendrobiology Acta Societatis BotanicorumPoloniae “ leaves and needles fromleaves and needles

. Kamień Śląski Badura

(2001) Phyllotrophic(2001) fungi"Cape mitosporicnature Mart'ian" of reserve Ecol össische Hochschule Technische dynamics Trees and ShrubsHardy Isles inthe British Trees A Flora of theEnglishLake of District A Flora ,

C , H. &, H. .

W ogy K., , , orestation: 45 .

, Dutech Using geneticUsing markerstoestimate L Erwinia amylovoraErwinia

, , rança

Mikologiya iFitopatologiya Mikologiya Holderegger,

S . , 13 16

(1968) (1968) 817 rmentier, C. (1993) Recherches surla C.(1993)rmentier, croissance préliminaires et chwab, P. Sourisseau define thelocal define , , 1291 , . 937 ” 57 Forest Ecology andManagement – ise . . 818. , Ekologia Polska , 3 Swiss AgencySwiss LandscapeEnvironment, forand Forests

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A Microclimate by and availability resource induced 45 9 - - . 1301. , Klein 14. 54.

comparative studycomparative ontheoccurren

, 2 ( , A. (2009) (2009) , A. 2001

R.,

different species ofgrow treesdifferent species 84

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, 28 genetic )

. Revue - E

Förderung seltenerFörderung Baumarten Kaiserstuhl. Herkunftsversuchs im EPPO Bulletin . 8 K . Watsonia

. , Oddou Improvement with of seeding directtree - ,

Forestière F Forestière Seria A Seria Z rich structure , 75 , . George Bell & Sons, London. GeorgeBell & Sons,

A. 35

, 233 - Sorbus torminalis ,

, 1 & Muratorio , B 27 , the pollen dispersal curve.the pollendispersal

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Sorbus torminalis Sorbus - – , 260. n

, 45 S S ). Murray,London. ing within the within ing 1716 . .E. , ce ofce microscopic . ,

Smouse, P.E. & P.E. Smouse,

45 ( 2007

- (L.) Cratzin , 261 1724

auf der .

) Allgemeine - . . Regional 278.

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This article isprotected rights by All copyright. reserved. AcceptedBiedenkopf Belletti, P., Bednorz,L.& M.K.(2009) Wojciechowicz, Development ofthemultilayered epidermis Bednorz,L., Maciejewska R., Walkowiak, Bednorz,L.& A. Urbaniak, Bednorz,L., Bednorz,L.,L.Isozyme& (2004) P. Myczko, Kosiński, polymorphism genetic and structure ArticleBednorz,L., Maciejewska Bednorz and structure spatial of Analyses (2012) L. Kaczmarek, & K. Kaźmierczak, L., Bednorz, Bednorz,L. (2009 Bednorz,L.genetic of Conservation (2007c) resources of Bednorz,L.wild service The (2007b) tree Forests the artificial ofwild service regeneration tree( markers. forest covering fruit of 3 variability species Polish ofthe (L.) National and Crantz)inPoznań Wielkopolski Park. service tree( Journal Genetics ofApplied of thepopulation of Poloniae morphology ofthe Sorbus torminalis Poland). of growth of measures selected 360. to protect service wild ( tree Dendrobiologia of communities Poland. , -

9. L .

, Monteleone,

& S Myczko, L.& (2006 P. Myczko, Kosiński,

species, .

, Ammer , Krzakowa

33 , Dendrobiology Eur 74 ) Jak chronić) Jak jarząb( brekinię , 1 , 315 Sorbus torminalis opean -

12. wild , ,

Sorbus torminalis 58 C I.

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(L.) Crantz.

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Polish speciesgenus of the Polish

. & Rutkowska, I.Rutkowska, & , 3 J service Sorbus torminalis ournal of of ournal (2005)the wild service Phenology of tree( .

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(2002) in Phosphoglucose isomerase(PGI) polymorphism 7. M

, Dendrobiology ü 67

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tree [ , 59 Sorbus torminalis , 45 - Starck For Acta Societatis BotanicorumPoloniae

(L.) inPoland. Crantz) - , 321 of the genusof the 65. Sorbus torminalis

Sorbus torminalis Sorbus , Wrońska P. ( P. est

- Sorbus torminalis (Rosaceae). Rutkowska,I. & K.(2006b)Rutkowska,I. Moliński, Seed

– , 2008

Res 324

G (L.) from Crantz theBytyńForest (Po ) Genetic variabilityand structure ofthe wild , . Sorbus torminalis Sorbus

57 (2007) Geneticaspects(2007) seed for harvests of earch . )

, 49 A - Pilarek, D. & Fujiki,&Pilarek, T.(2005) D.Pollen Sorbus

) inP population genetic study Sorbus torminalis Sorbus Sorbus - , Dendrobiology 54. 54. 127

Sorbus torminalis (L.) Crantz],

olish forests?]olish

,103 in Forest District Jamy (northern (northern Jamy District Forest in (Rosaceae).

(L.) in plant Crantz

L. Silvae Genetica Dendrobiology - Acta Societatis Botanicorum Acta Societatis 114. ) w polskich [How lasach?

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Sorbus torminalis Sorbus 62 using Dendrobiology [L.] C [L.] Sylwan , 11

, in Poland. in Poland.

71 , RAPD -

in rantz). 16. , 55 , 121 53

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This article isprotected rights by All copyright. reserved. L.Cheffings, & Farrell, (2005) C.M. Chater, A.O. Chalupa, V. &Challice, Kovanda, M.(1978) J.S. I.,Čaňová, Ďurkovič C Butin British Mycological Society (2016) Brewis, A., Bowman,& P. F.(1996) Rose, Boulanger Borchard Birks Biological Records Centre

Acceptedampbell Article , , H.

H Species Germany, Agriculture andFore wild service tree[ Europe. Photosynthetica and photochemicalefficiency 135. andconcept species Maloideae (R subfam. in 2. übe http://www.fieldmycology.net/FRDBI/FRDBI.asp Colchester, UK. Ecology andManagement Ranking temperate woodys Indicators foranimal interactions: a new Possibilities plant value? defenseindicator Reserve http://www.brc.ac.uk/dbif/ . J , , C.S. &, C.S. D ( . , V., F 1989 B

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J 50 Joint NatureJoint Peterborough, Committee, Conservation UK. -

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pecies along a gradient ofbrowsing alongpecies a by deer forest history offorest RoudseaNational Wood Nature history The FungalThe Database Records of , .

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Y . P Dupouey Sorbus aucuparia . , S . Harley Books, 12 Biologie . Britain and Ireland andBritain

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This article isprotected rights by All copyright. reserved. De (1983)Barluzzi, Dominicis,C. V,& Coenological Rueda, andde&Forestloss fragmentation Albuquerque, M.(2010) F.S. woody effectson D David, J. Czeczuga, E.,Godlewska,A. &Mazalska, B. B.,Aquatic fungi and (2009) Muszyńska, C M.(1679)Cook, Piboule,Leroy, C., Collet, A., & O. Frochot,Advance H.(2008) Becquey, Gonin,P. J., Coello, J.P., Ortisset, J., CIE (1969) Christensen, K.I.× (1995) Chester, Fay, M.,Cowan, (2007) R.S., & M.F. T.C.G. Parentageendemic Rich, of rave,(1985) M.F. Acceptedavis, T.A. Article oak woodsinthe hill 479 plant species Great richnessBritain. in Welsh Bulletin Change Biology qualitylitter onthelife 467. fungus Printed for P coppice pseudoplatanus Catalonia, Spain. NaturalCatalan& Environment, GovernmentofCatalonia ForestCentre, Ownership Neighbouring Regions Silviculture ofthe MainValuableBroadleaved inthe Species Pyrenean & Fund Nature,pp.358 Greuter Wide XX, for World W.) Book A.,Snogerup, of D.,Strid, Rareand ofGreece ThreatenedPlants Phitos, (ed. S. Societyof the Linnean Isles:(Rosaceae) British DNA. species inthe evidenceplastid from - F. (2009) & ofelevated effects Combined and D. reduced Gillon, temperatures leaf .

W. (1976) wi W. The Distribution Maps of Pests, No. 87 MapsofPests,No. Distribution

- like organisms growing like organisms taxa. of131plant onseeds - with The Manner of Raising, Ordering; Raising, andFruit of andImprovingThe Manner Forest

Un frutier mecoUn frutier . Parker,London. - standards forest. standards ,

25 , seedlings broadleaved dominate treea in mixed regeneration former

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MinistryLivestock, ofAgriculture, andFisheries, Food history parameters ofhistory parameters a saprophagous macroarthropod.

- 165 , 291 nnu: l’alisier torminal. nnu: l’alisier torminal. .

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Forest Ecology andManagementForest Ecology . , ,

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CAB International,CAB 81 Baiges, T.& M.(2013) Piquè, Vegetatio

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Trees Acer , 451 Global

- L. - . This article isprotected rights by All copyright. reserved. (195 Düll, R. DrapierN. (1993 D DrapierN. Drapier,N. Drapier,N. (1991) D I. Dimitrov, &Investigations andstatus of (2008) thedistribution health Genova, F. on Devi Devillez De Demesure Demesure rapier,N. (1993c) inca

Accepted ArticleB.,mesure, llez, F., Fraipont, J. F.,llez, Fraipont, J. & M. Tissot, ,

L Germany Forestière Française Forestière Française Forestière Française botaniques etgénéralités. forestières en Sorbus torminalis 1 de desBulletin laClasse Sciences Acad surleleurs milieu positions de aria Sciences Acad (L desForetsNational L’alisier torminal: InternationalInstitute, GeneticResource Rome for GeneticandU Conservation Science of , F. (1979) . 82

.) ( , B., , - a scatteredforest tree: temperate

2000 Musch, B.Oddou Musch, & . (1993

(1993 Crantz (L.) 9) Oddou, Le Guerroué, , ) Unsere Bastarde Ebereschen undihre

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a ) ) , , 63 ) Connaissancegenre du

de laprotection À propos Influence de lastratificationsurleset graines de les fruits

Écologie de l'alisier de torminal, Écologie Écologie et intérêt divers de sylvicole Écologie S Recherche desylviculture pour d’éléments l’alisier torminal. . general.

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45 45 45 Revue Revue , 51 - rantz. inPreslavska Mountain. rantz. Muratorio, S. , , 321 , Lucchi

L Revue 229 345 . et – 63. Forestière F - - –

354 243. (1980) S 334. culture et appliqué culture desprétraitements

, G., . se Forestière F Forestière B.

Corminaria torminalis L.et de : Wild: Service T . ,

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EUROFORGEN Technical GuidelinesEUROFORGEN Technical L. 14 . A. Ziemsen. A. Wittenberg Verlag, .)

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Bulletin Bulletin Technique de l'Office (L . 501 Forest Science -

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- b ) This article isprotected rights by All copyright. reserved. Evelyn, (1664) J. E Espahbodi Espahbodi, Espahbodi Espahbodi, K I.A.Emmet, M.A.,Watkinson, & M.R. Wilson, Emmet, Emmet, V., Ellenberg, Düll,R.,Wirth, Werner,H.E., H., Weber, W. Ellenberg LeDupraz, C.(1994)chêne peut et le blé:l'agroforesterie

Accepted (1988) vans, J. Article HMSO, London HMSO, interrelationship populations ofwildservice tree( J and Biology on wildservice (2007 S.M. 33 torminalis Dehghan 55 ( T & Emmet, A.M.), Butterflie & Emmet, A.M.), ofGreatButterflies Britain andIreland, V Zeigerwerte von Press, Cambridge Européennesgrandescultures? de Sorbus torminalis A. A.M. (1976 alali he effect of ol. 1 , , 107 , 47

, K., , K., M. H K., . (1988) . (1988) , S.G. (2008), S.G. . . - I.A., Watkinson, & M.R. Wilson,

, Mirza 55 ( - Mirzaie Hosseini Mirzaie ed. 119 - Natural Natural Regeneration ofB s of Greats of Britain andIreland, . Shooraki

. Sylva b Bulgarian AcademyBulgarian AppliedPhysiology and Plant ofSciences, General Heath, J.) Heath, J.) . , ) )

9 sowing depth Vegetation E Vegetation ei Seed . , 426 . Jo. Martin & Allestry,London.. Jo. Ja. - – - ( , UK .

Na Nodoushan Sorbus torminalis , S.M., , S.M., Nodoushan Pflanzen Pflanzen Silvae GeneticaSilvae , UK. pp. 244 pp. 212

, Y.( Geneticgrowth variationcharacteristics inearly of in source effects emergence,and seed seedling on survival doushan, - 430. .

a Harley,UK,Books, pp.171 Colchester,

mountainous 2007a) T 2007a) Mirzaie

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Harley,Books, UK. Colchester, Harley,Books, UK. Colchester, soil protectionsoil for – Emadian, Emadian, Tabari Sorbus torminalis ree age Nodoushan Tabari ,

Revue Française Forestière ) s 57 roadleaves eedlings , nursery

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Forestry Commission BulletinForestry 78, Commission Lyonetiidae. , Gracillariidae. Tabari

- seed Sabbagh, S

elle intéresser les exploitations elleexploitations intéresser les (L.) Crantz)

(4th edn). &D. ( Paulißen, , M., , M., germination tree of service , M., - - Heliodinidae Heliodinidae Dehghan Jalali Akbarinia

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This article isprotected rights by All copyright. reserved. Geb García G Gabri Gabrielian Franke Finkeldey, Sperisen,Bonfils,& R.,Mátyás,(2000) C. G., P. Strategien zurAuswahl LaFehr, de l'alisier torminal R.(1993) place F F Fady,& geneticB. ofMacroecologicaland species Conord, C.(2010) diversity patterns in Facc Ewald, Zander, C., M. avre ay, M. Acceptedarcía Article , iola, S.

M - ė d

- lian, , López, Abad Alonso, J.J., Gómez Delgado, Gómez Alonso, J.J., Abad M.& der Bestandesstruktur. 161 Guadalajara.la detección ennaturales Utilidad deenclaves de interés. Cartografíaplantas vasculares,sector Alta Alcarria, en de detalladaun de la Península Ibérica Botanicheski Garden, Edinburgh 45 einh 137 forstlicher der Genreservate Schweiz. in d'un en forestier Suisse. public domaine Watsonia (2002) etdes desForêts Eaux (ENGREF) B vascular Mediterranean basin. plantsofthe California, Brandenburg.

. ’ F., Gernand A , Schmidt anne ibliographique . , , E.T. (1961) , E.T. , Dagenbach 166 - -

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Flora I , France , Western Asiaandthe Him Rodríguez (2009) Espinosa, V.M. 567 Diversity and Diversity waldprojekt Liebenburgwaldprojekt , Spain. – ois 573 R., Kitchen,R., & C. Rich Sorbus latifolia latifolia Sorbus Notes lustrada del CentroyNortedelustrada la

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30 75

, , This article isprotected rights by All copyright. reserved. AcceptedHalliday, J. Guitton, Grundmann Grove Grigson,(1955) G. Ladet, M., Gonzalez, Deconchat, G.(2010) Cabanettes, A.,Alard,Balent, M., & S., D. Article M.,Deconchat,Gonzalez, M.& G.M.,ReyGonzález, I.C.Vinãs, de&O.(2003) Cisneros, Godwin, H.(1975) Germain,B. (1993) Unréseau d'alisierà Regeneration 30kmdeNotre torminal. placettes de Gebauer ,

W Furness (v.c.69),Furness (v.c.70) of North Cumberland andparts Française Germany D Resistance Plant Crop Factors andAbiotic toBiotic Cambridge,UK. have orNearlySpores Colourless Colourless Melanconiales. Sphaeropsidales, tothe Volumeend ofthewhich Sphaerioideae 1. Knowledge ofthe FungiImperfecti Belonging tothe Sphaeropsidales and 274 richness: woody Anexample for plants. Relative 201 the importance andenvironmental conditions spatial configuration. of Madrid, Spain. Plantaciones de Especies Productorasdede Madera Calidad. Cambridge,UK. Française Dameen de forêt régionale Paris Ferrières de (Seine 4 forest withvariable abundance stands , T., 43 . G. (1997) . V B - . , 305 , 2 L. Essais de (1994) plantations agroforestières duSud. enCorse . & , .

Horna, B (1935) (1935) - 14 . M Furk - contribution ofedge zones topatcheffect and size interior onspecies , pp.53 . 318. , , . & .

46 4 The Englishman’s Flora The Englishman’s Flora ofthe British History 5 A Flora of Cumbria: Comprising Cumbria: of theVice A Flora , V. &V. British British , 96 , 335 Roloff C

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Balent, G. (2009) Woody of forestBalent, composition G.(2009) plant layers: -

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U se of forest treeseclimate ofunder species change - Fungi (Coelomycetes). our to Fungi AContribution .

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Manual de Selvicultura para Manual deSelvicultura

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This article isprotected rights by All copyright. reserved. Dü Hoebee, U., Arnold, S.E., Hochbichler Herrera, (198 C.M. Herrera, Vertebrate (1987) C.M. Herrera,bird A frugivores, ofavian study (1984) C.M. Hermy,Firbank, M.,Honnay,L., O., Grashof H Hemery, J.R.,Aldinger, M.E.,O’Connor, E.,H.,Malvolti, G.E., Claessens, E., Clark, Hellwig Hasbal, G., Yilmaz Harper, G. service (1981) Middlesex. Wild and inHertfordshire Harley,& J.L. Harley, A E.L. check (1987) H

Acceptederman, Articlearalamb,(1967) A. R. (2007)R. (Niederösterreich). fruit characteristics, inundisturbed Mediterraneanhabitats. characteristics. in Mediterranean scrublands. the implicationsfor forestconservation. ecological between andforest ancient plantspeciesEurope, comparison other of 83 speciesaclimate:of inEuropechanging risksand areview opportunities. in &Raftoyannis, P.S. Brus,GrowingY., Savill, tree R.(2009) scattered broadleaved Archaeobotany century) G from and DrugAnalysis activities of Hertfor Phytologist Romania Lancaster, (v.c. Lancashire North 60). ,

J M , .

65 . (

, 1971 (1997) Plant remains(1997) from(15thand16thcentury) cesspits two a pond(13th

E. - 81. dshire Natural Historydshire Natural Society (2003) .

) Mating patterns andgeneMating contemporary flow patterns by pollen

Š , 9 - Sorbus torminalis umarska D umarska T.&and(2015) antiacetylcholinesterase Antioxidant Ozden, Can, A. 105 ) F

Cultura Speciilor F Cultura , Monographs Ecological rugivory and seed dispersal byrugivory seed and associated and carnivorous dispersal mammals,

Die Elsbeere( Die ö

6 (suppl.), 1 ttingen, southern Lower southern Germany.ttingen, Saxony, , 105 , Forst und Forst 23 ggelin, C., Gugerlggelin, C., UK. , 57 - 116. endrologija - dispersed plants of the Iberiandispersed ofthe a Peninsula: plants studyfruit of

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orestiere , . Stanbiro,Zagreb, 58 - list of mycorrhiza Flora. inthe British of list

- , and FieldClub Bokdam, C. & Bokdam, C.

647 , i, F., & Brodbeck,P. Rotach, S., Biological Conservation Biological 57

(3 – , 305 - 653. rd West Regional Studies West

edn.) - dispersed pla

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331.

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, , Lawesson, (1999) An J.E. Croatia. 28 54 Transactions (4) , 1 Oikos - , 17 23. nts, and their interactionnts, and Vegetation History and History Vegetation

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This article isprotected rights by All copyright. reserved. Kausch Kárpáti,Z. Die (1960) Kahle J J Jacobée Idžojtić,& M.,Zebec, Drvodelić, D. M. Hurt, Kantor, V. &Jeřáb břek (2004) P. ( Huneck,III.gewachsenem (1962) S. Die Triterpene. als Anwendungvon Aluminiumoxyd Hoebee Acceptedankun, A.& Kovanda,M. (1988) ankun, A.& Kovanda,M. (1987) Article ,

M - cormier)en Allemagne. Repert Ö Crantz) amBeispiel ei (Embryological in studies studies in propose. Solution to morphologyand fruits. ofleaves variabilitywild servicecontinental tree part of the of populationsin Hrvatskekontinentalnom dijelu obilježjimališća prema iplodova morfološkim Černými lesy, Prague,CzechR Woody soil andforest plants in Křivoklátsko area (ed.(Central Bohemia).] Anon) L. torminalis Crantz.) listnatá dřevina a naKrivoklátsku. její význam Journal ofChromatography derselbsttragendes in Diinnschichtchromatographie Adsorbens vonTriterpenen. 1 rare witha treespecies scatt structure of torminalis continuous and a isolatedpopulation of small forest thescattered , , F. (1993) Base génétique, F.Base (1993) des dans récolteslequart d'alises nord

Blecken v - S kologie, Bodenordnung undForsten,kologie,Bodenordnung Recklinghausen . 8.

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. , Menn orium Sorbus . Untersuchungen zum Wachstum derWachstum Untersuchungen zum on Schmeling Heredity Sorbus torminalis , ,

C 62 Sorbus . , Rotach , , 71

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331. 331. Preslia 99 - niger Mischbestäniger

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226 The ce , This article isprotected rights by All copyright. reserved. AcceptedKorkmaz, H.,Engin,Kutbay,& H.G. A., Kol'Tsova Kollmann, J. Köckemann &Klumpp, R. T.( Kiristis, Kleinschmit, J. Klein,& E.K.,Desassis, N. ArticleKirby,K. Keith, N S.A., Kausch Kausch Kausch Kausch - - - - Botany scrub, forest,vegetation andsteppe valley. ofthe Kızılırmak B 27 gapsforestc edges and in S of the 50 of Conservation Status Holz estimated jointlywith the dispersal kernel. Sorbus torminalis Key Proceedings Society ofthe Royal homogenisationTaxonomic years. over of 70 woodland plantcommunities Stuttgart aen em r erg von Schmeling, Germany. Bovenden, Verlag Bovenden, Kausch, Diplomarbeiten. Blecken & vonSchmeling,W. Kelln Blecken (2000) vonSchmeling,W. Blecken (1994b) vonSchmeling,W. Blecken vonSchmeling pecies iologicheskii Zhurnal Armeniiiologicheskii Zhurnal , 85 (2006) , RoyalBotanic Kew, Gardens, Surrey,UK.

. (ed. M , B. (2008) , ,

53 (1997) . – D ewton, A.C., Morecroft,ewton,&Bealey, A.C., Bullock, M.D.,C.E. J.M. A , 91.

. istribution istribution ( 35 , , Germany . PhD t PhD 1998

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, 121

Hypotheses onthe regeneration niche offleshy - ) 519. hesis Erhaltungwertvoller Edellaubbaumarten. undNutzung - Abundance, 165. Allgemeine ForstZeitschrift Allgemeine C

(L.) Crantz. IV.(L.) variance Crantz. interindividual ofmale Whole

ontent of trace infruitofdifferentelements ash mountain species Ancient Woodland IndicatorAncient Woodland R . , pp.558

, genus1998) The

: Natur Kleinen undUmwelt (Hrsg.):im Naturschutz

Oddou ange Georg

Sorbus ,

. entral Europe. W A - - Germany. August - Muratorio, S. Muratorio, . N 561

M (1994 iche iche in the UK acroecological ,

33 , Series Penguin

Yalçin, E. (2011)Yalçin, Asyntaxonomicalstudy onthe - B a , 95 Universität ) Fö) Der Speierling. 2.Auflage Der Speierling. readth and readth Sorbus

er, T. (2006)er, T. Die Elsbeere( Die - 99.

Verhandlungen rderung von Elsbeere und Speier rderung vonElsbeere und . (eds B , London,

( , Molecular Ecology 2008

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in Austrian Forestry. In:in Austrian Forestry. , . 49

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1449 , UK , Die Elsbeere Sorbus torminalis Sorbus torminalis - 3544. (AWI – .

- A 1450 Gesellschaft Gesellschaft bundant and bundant , Germany.

- s C Turkish Journal of Turkish fruited speciesfruited innatural . )

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order order tree, , - B. - . This article isprotected rights by All copyright. reserved. Lefèvre, Kraigher, Hubert,Koskela,Longauer, F.,J., J., H., L Lanier,LesL.maladies de (1993b). l'alisier torminal. Lanier, leL. sur Le marché torminal. boom (1993a) l'alisier de Lachaud Lachaud,& A.(1993) ducambium Mansouri, etxylogenèse Reprise S. d'activite chez PodgórskaKwiatkowski, (2003)ciepłolubna P. dąbrowa brekiniowa Kutzelnigg,H., 1995. Kučerová Kovanda, Observations on M.(1997) Kovanda, M.(19 Kotar, M.(2001) Hö

Acceptedanier Article , L.,, R.T., R.T., Postolache, Maaten Eysteinsson, Bednarova, Bozzano, torminal 344. 4 xylem maturation. vessels andcells other in xylem l'alisier torminal. Geobotanica Polonica forest Quercetum pp. 343 Biologia of serrei h II.adjacent , 5

, V., S , 319 Sorbus torminalis Rameau, J. Rameau, . & .

Westergren, Sorbo torminalis

Honec , T., – – Maurousset , 320. 349. Sorbus torminalis

84) Ahybridogeneous new 84) 65 M., ,

na Pogórzu Złotoryjskim. Pogórzu [Submontanena thermophilous oak 134 D., D., Paino, ,

T., F T., henwachstum der des Elsbeere Speierlings.henwachstum und 817 , M., , M., Parey Alizoti, - Sorbus torminalis Sorbus C., , Ryan, Bord 305 - Revue ForestièreRevue Française

r 821 M.,

ý Protoplasma Paule E.N., Keller, R.,Joly,H. , , Berlin, , Berlin, erhan lungeneroologis h dl, L

- ác in Eastern Europe 316

. . C., C., P., P., Wolter, ,

s, - J., J., of(1996) plasmodesmatadifferentiating Occurrence between 10 Quercetum

, L.,, Ozt . S. Bakys,

Steffenrem, Haverkamp, , 175

, (L.) Crantz. Germany Collin, ü Zhelev rk,

Sorbus F., F., - , 193. Sorbus torminalis

. R., R., K., 191 Yrj Illustrierte Flora vonMittel

in Złotoryja foothills.] Złotoryjafoothills.] in E., , P. &, P. Baldwin, Pandeva, , 220 ä

Sorbus

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- in Southwest Moraviain Southwest Republic) and (Czech n

A.,

I.,& E.(1990) Drapier N. Sevrin M.,

ä De Cuyper, as by determined markers microsatellite Revue Française Forestière ,

L.& Varel - G 226. Ivankovic, . ö , Preslia m

C., C., I.D., Revue Française Forestière 45 Zariŋa

a, ö ry

, 279 , M.C., M.C., Ballian,

- L.fateduring and Crantz their

, D. (2010) , o anis oanis hen in esells ha B., Parnuta,

R., R., , , I. Dynamic, (2012)

56 - M., De Vries,De 283. R Vessella, Olrik, , 169 evue Française Forestière

D., Konrad, Fragmenta et Floristica

G., Corminaria - europa Black

Sorbo torminalis Sorbo 172. Genetic differentiationGenetic D.C., D.C., Pilipovi

S.M.G., F.,

H., , - Samuelsson, Sch . 42 Volosyanchuk,

(Ed. G.Hegi), - Koziol, wild servicewild , 13

č üler, S., L’alisier , ,

, A., 16 – 4 34. 5 , , 343 19

C., C., - .

S., S., 22 – ,

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This article isprotected rights by All copyright. reserved. Accepted Máchová, Novák,Malá,Hauserová, Cvrčková, J., Karady, J., O.,Mikulík, P., E., M., H., Maděra, and of &Distribution ecological S. (2013) Tichá, requirements P. Řepka Maděra,& Šenfeldr, Kohoutek, M. P., M., Maciejewska Máchová (2004)Loidi, J. Lockton, A. & S. Whild, ArticleLloyd Leuschner,& B. breadth,(2009) Abundance, C.,Köckemann, Buschmann, H. and niche Lepší, Boublík,Lepší,P., K.& (2008) Vít,P., Suda, J. M., Lepší, Boublík,Lepší,P., F. K.& (2009) Vít,P., Kolář, M., Leinemann , E.G. (1977) E.G. Greplová, Strnad, M.& Micropropagation J., Doležal, K.(2009) of wildservice tree torminalis (Czech Republic). regeneration of Acta Societatis BotanicorumPoloniae observations onthe Lesnického Výzkumu jeřábu břeku 126. Botanical Society,Shrewsbury, UK. Na range.their distribution niche occupationof Europeaninthe treespecies centre Central andmargin at the of hybridogenousBohemia. from northwestern of Sorbus bohemica. Sorbus Jagdzeitung genetic inthe genus patterns variation Biology conservationgeneticforest of resources European in33 countries. , turalist P ,

. L., Kahlert , - Malá Rutkowska, I.L. Bednorz, &Rutkowska, SEMandmicroscope stereoscope (2004) albensis , Südwesteuropäisc 27 ,

(L.) inthe Czech Crantz Republic. , 56 ,

, The Wild ServiceWild TreeThe . [Optimalization of wildservicemicropropagation.] tree J 373 9/10 . , , 22 Cvrčková , twonew endemic recognizedrevision apomictic ona based species Sorbus torminalis ,

- K., Arenhövel 384 - , 169 (2015) 28. Dendrobiology

seeds of the Polish species oftheseeds Polish of thegenus , . Preslia

54

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Forest Ecology andManagement 174

, 218 , The F , H h e sommergrüne Laubmischwälder.e sommergrüne . . ,

& - 81 222. lora ,

Dostál , 63 W., Voth W.,

, in Hádecká National planinka NatureReserve

68

Řepka, R. (2012)structureR. Řepka, The and population Sorbus torminalis torminalis Sorbus and V -

89. , 63 , Sorbus , 73 J

. ( - egetation of Shropshire of egetation , , 294 68.

2009 W. W. Preslia

Dendrobiology in Thuringia. & - 300. 300. )

Hosius Optimalizace mikropropagace

Sorbus milensis ,

Sorbus portae 80 in Epping Forest. , 22 ,

B.Insights (2010) into ,

258 9 Allgemeine Forst und und Forst Allgemeine Sorbus – , 244. 6 , 1

Tuexenia 9 , 59 248 .

Shropshire

Conservation Conservation

- L. (Rosaceae). , a new bohemicae - – 68. Zprávy 1259 The ,

24 Sorbus .

, Lond 113

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This article isprotected rights by All copyright. reserved. Németh Nelson Q.,Syla,Pajazita, Mustafa, A.,Hajdari, B., B., An Quave,& C.L. (2012) Pieroni, A. M Müller, Ammer, N S., C.& Muller, C Moradi, Matinizadeh, M.,Etemad, Shirvany, M., V.,Naji,A., H.R.,Abdul Mik M (20Medak, J. Martí McAllister, H. (2005) Mabey, R.(1996)

Acceptedü Articleeusel ller oláš, nez - - , H.,J Act the genus Evolution ethnobotanicalthe Gollak region, survey Kosovo. of um ihren Erhalt, undHolzverwertung.Waldbau Anbau, Forstwissenschaftliches Centralblatt silvicultural decisions Forestière Française Biogeosciences andForestry does vary across not nutrientsand enzyme withsoil activities different Sayah, Arbuscular (2014) mycorrhizal S. with fungal symbiosis Sorbus Zentraleuropäischen Flora ( sativae Mediter roots and stems ofnine Gardens, UK. Richmond, Kew, during organogenesis. ( Kroehling , Cs. (2012)new, Cs. Two E.B., TheJones, Briggs,& A.G.(2002) origin D.Smith, ofintermediate species of Aposeri foetidae Sorbus tor

- V . Vilalta a Botanica Hungarica & Laroppe,& E. .

(1996) äg 11) Šume pitomog kestena zeljem sprasećim

s.l. from easternSlovakia. ass. nova) u Hrvatskoj [Forestsass. ofsweet nova) withodorouspig uHrvatskoj chestnut er , ,

, Sorbus , J Flora Britannica Flora

59 E. & E. minlais . S , Prat Sorbus dolomiticola Sorbus . & , 739 The Genus The

W Franz . - ,

castanetum sativae castanetum Theoretical and Applied Genetics andApplied Theoretical ( - E

einert, 1993 754. [L.] the Crantz): regulative ofdifferent role cytokinins aromatic üß . , Oliveras ,

, 45 – (2000)lein, S. stand Analyses of Journal of Plant Growth Regulation Growth ofPlant Journal Sorbus

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Conservation et germination Conservation des et semences. ,

E. 253 Sorbus (1999) undSpeierling Elsbeere Bayern. Bem in 54 . Veb . Sinclair

(1965) , , 131

- 8 , (Rosaceae) species from the Bakony Mts, Hungary.Mts, (Rosaceae)Bakony species from the 260.

, I ranean woody species.

. & .

308 Mikoláš Gustav : Thaiszia - Mountain AshandOtherRowans Mountain 144

Vergleichende der Chorologie Pi - , -

313 Stevenson, London,Stevenson, UK. ass. nova) inCroatia]. ass. nova) 119 ñ . ol

F . , agenus newof species hybridogenous the Quercus petraea Quercus ,

, 32

ischer , J

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(2002) Xylem properties hydraulic of , 1 - 42. -

12. V

erlag

, ( Aposeri foetidae Genetic Crop Resources and 105 Oecologia

, Jen structure as a tool structure as Corminaria , 953

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28 a Šumarski Lis Šumarski Lis Sorbus torminalis , Germany -

, 341 963.

Sorbus torminalis , 133 - -

, Hamid, H.& . RoyalBotanic 348. - 12 Revue castanetum , 19

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This article isprotected rights by All copyright. reserved. Oddou Oddou Oddou O Oddou Oddou Oddou Oddou Nicolescu,Hochbichler, J., E.,CoelloGomez, V.N., Németh, (2015) Cs.

Acceptedddou Article ------55 Pollen Botany Mating and patterns theecological maternal neighborhood. patterns thewild of pollen flow in successfrom inferred parent Sorbus torminalis 2754 flow a within populationofa scattered treespecies. analysis populations. procedureincontinuous flow inthe wildservice tree, Environmental C Star scale investigation of of seed a in dispersal scattered forest tree species, genegenetic logging flow and stru historyonthelocal Mol B literature review. Ecologyofand wildservice silviculture tree( Lakesurroundings Balaton, Hungary of Muratorio, Demesure S., Muratorio Muratorio, S., Petit, R. LeMuratorio, R. Petit, S., J., B., Guerroue, E.K.,Demesure Muratorio, Klein, S., E.K. Muratorio, Klein, S., Muratorio, S. Muratorio, Houot, S., Muratorio .

, 112 (2001b) Microsatellite primers for(2001b) Microsatellite ck, G. & (eds)ck, G. Schubert,of Genetic Forest R. Response S ecular .

- ,

3

versus seed 93 Sorbus torminalis - 1135. , ,

, 1650

Ecol S S S . . , Guesnet , Aligon orbus &

ogy

Klein, E.K. - onditions. Die Bodenkultur Die

1659. (L.) Crantz. I (L.) Crantz.

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Notes mediated gene flow forestmediated inascattered tree species. pelsoensis

, population genetic structureforpopulation chlo M.

C ,

D . - - , Decroocq , Musch, B., R.&Musch, Pélissier, Gouyon, P. L., Demesure .

, Ozdemir & 1 L. Crantz. Springer ,

– Sorbus torminalis ofgene(2008) Comparingestimates directvs.indirect 297 Austerlitz, F.Austerlitz, offspring analysis.

( I S . –

orbus Pollen dispersal and dispersal heterogeneityPollen inmating - 299 , service torminalis (Rosaceae).III. tree,Sorbus

60 , , ,

S - 271 Dordrecht, pp Molecular Ecology Sorbus torminalis E . , 35 Musch, B.Musch,

.

. Petit, R.J. & R.J. Demesure,B.. Petit, , Plomion - subgenus . Musch, B. &(2003) B. F.Pollen Austerlitz, Musch, Studia Botanica Hungarica Botanica Studia

species ( species -

44. (2005) Guesnet,& B. D. Demesure, (2001 Ravagni, & S. V. (2009) Giulietti, Sorbus torminalis Sorbus

Molecular Ecology

(L.) Crantz. I.(L.) Crantz. paternity Evaluating the Molecular Ecology , & tormaria

Pollen flowPollen inthe wil Sorbus tormin C

Austerlitz, F. Austerlitz, Molecular Ecology . , Lamant cturea scattered of tree

– and related species. , 280 13 roplast DNA. In: Mü roplast DNA. ), aspecies new), from the American JournalAmerican of , 3689

ystems to ,

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- (L.) Crantz): . & . H. (2004) Impacts (2004) H. of alis

, ( 2006 12 -

3702. (2001c) Patterns(2001c) , Mush ) based onmulti ,

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17 Real Evolution - Demesure , 2743 , - – 3439. 60. A - - time 4452. a ller

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This article isprotected rights by All copyright. reserved. servicePeterken, (1983) G.F. Wild Pellicer Palenzuela Paganová Paganová Orša Orša O (2008)Oria de J.A. Rueda, ONF (1999) Olszewska Olszewska

Accepted Articleria

ni ni de ć, M., Drvodelić, D., Jemrić, T., Anić, I. T.,Anić,D., Jemrić, ć, & M.,Drvodelić, I.D., Anić,& ć, M.,Drvodelić,

Lin olnshire Na uralis olnshire Na Lin s’ Union 110 the in Andalucía (Spain) Acaulospora viridis forestry and CRANTZ ScienceHorticultural 495 (L.) Crantz)fruits andseedsfrom morphologicaland biological characteristics 706 of fruitgenus and ofthe seed (2006) género Cálamo desForêtsNational torminalis Pharmaceutica inflorescences, frui leavesand , J., , J., R , , ueda

, M.A.& , M.A.(2011) , J., , J., V. V. Sorbus , – . Clermont

1185 504. Leconservatoire génétiquea des ( ( , , J.A., , J.A.,

2008 2007 Sorbus Azcón , Palencia, Spain. Fuera de Serie

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Mart , S., Eco Ecology and -

Aguilar en España.

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scape. Drug Research logical requirements of wild logical of requirements I Houston í n nez

, a new species inthe Glomeromycetes from two

, Guía de vitro antioxidant activityantioxidant vitro content and total phenolic

. 38 , . Nova Hedwigia Nova

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34 de Phytochemistry Letters , 166 , , 9 Journal ofForestScienceJournal

, 138

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Investigaciones Sistemas y Forestales Recursos Agrarias: , L.,, d Barea 11. - istribution of istribution tree in central Lincolnshire.tree incentral - Á Sorbus domestica 168 Sorbus ts rboles y Phenolic constituents oftheinflorescences of -

151 Mikac Rich

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, 158 - , T.C.G. & , T.C.G.

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- This article isprotected rights by All copyright. reserved. AcceptedRasmussen (2004 Rasmussen, K.K.Kollmann, &J. Rasmussen, K.K.Dendroecological (2007) analy Rameau Raguin,F.Boulet & &Rad, Shafiei, J.E. (2010) A.B. Rackham, O.(2003) Quevedo,L., & (2007) Rodrigo,A. Espelta,Post J.M. ArticleQian, G. & Bauhus,(2013Pyttel, J. Kunz, P., J. & Bauhus,(2011)Pyttel, J. Kunz, Age P., J. growth of and wildservice tree( & Ulrich,Puhe, J. B. (2001) of therare tree 662. from phytosociological gradient. relevés along a latitudinal climate, latitude, habitat and conditions forest history. et Collines E Flora ofcommunities Caspian Forest Science dominant shruband treespeciesareas NE inMediterranean Spain. of 227 section 27 ServiceWild Tree ( Potsdam, 11/07, Scientificp. 64 TechnicalSTR Report Tree in Archaeology, Rings ClimatologyEcology, andVol.9.GFZ Potsdam, Theunissen M,SpieckerH,GärtnerI H, G, Heinrich(2011): Helle (Eds.) TRACE torminalis Europe Central Ecosystems:Developm Postglacial , -

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Acta De ent, Present Situation andFutureTrends in ent, Present Situation

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Journal ofPhysiology and Farkas , 373 - 1988: The BSBI Scheme. Monitoring Brevulacus Brevulacus species - 38. .

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L – . This article isprotected rights by All copyright. reserved. Salisbury Safdari, V., Sigarody, M.R.N Safdari, V., Ahmed, A.Rudow, & G.(1997) Aas, Rotach R Rosati Service of theWild Roper, P.(1993) The distribution Tree, (2005)Roleček, Vegetation types J. ofdry Röhrig, E. (1992Rodwell, J.S. (1991)Rodwell, J.S. Robertson L., Harris, Robertson, Bridle, J.R., S.A. A.,Rich,C., T.C.G.,Allen,A.M., Houston, Roberts,

Acceptedose Article , F. , L.,, , P. (2000), P. Ecology relation the to 2133 non wood with hand lens. and site Distribution, populationbiology. Research Überlegungen, Situationsan ancient nature woodland conservation. for 38 vascular a Mediterranean plantdiversity in the British 261. 27 Communities University Cambridge,UK. Press, theendemicevolution in onArran. Sorbustaxa in evolution andspeciation (2010) & S.J. Hiscock, ( , E. 1999 - , ,

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woody plant species in pulp andwoodyin pulp plantspeciespapers. . J. & Jane, J. . .

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, 267 . Cambridge University Cambridge, UK. Press, M., Palmer, J. &IdentificationPalmer,(2008) J. Baig,Iranian ofM., M.B. commercial

British Plant Communities Plant British Communities Plant British

veget , F.

A Watsonia - 325. W. (1940) MaidenW. significance Charcoals Castle and their from in Marignani - . 284. C ation and climatic prehistoric conditionsation in times . & . Pakistan Journal ofBotany Pakistan Sorbus latifolia

& Hybridization and polyploidycontinuing of as drivers

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Botanica HelveticaBotanica British WildlifeBritish

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40 Effects of fragmentation on Effects offragmentation Sorbus torminalis Sorbus torminalis , 1851 , . 19 , Forest andLandscape Snow 93

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, , 43 77 , , 241 2127 ridge wirt , 144 , – - , This article isprotected rights by All copyright. reserved. Siljak G.,Alcántara,Siles, Rey J.M., Lachaud,Sevrin, E., S. S Serteser, Gök,V.,Kargioğ, Bağci, A., M.& Antioxidant M., Y.,Özcan, D.(2009) Arslan, (1989) P.D. Sell, The Seidel Schü Schüte, G. (2000) Schrö Schirone (1991): P.S. Savill, &Sardans, (2007) Peñuelas,phosphorusand Droughtaccumulation J. J. potassium changes Saravi Acceptedevrin Article te tter - , D., , , E. Yakovlev ,

O. &Brown, (2010) S.C. species restoration. assemblages for Forestière Française relations avec lasylviculture et le sol. properties of some plants 17 and Management andexploration productivity inatemperate broad Fehrmann Elsbeere( Göttingen, Germany. ( Mecklenburg 7 Mediterranean forest Wallingford patterns inanMediterranean evergreen forest. Tree ( correlation between characters Ser of selected parent and trees inWild progenies A.T., Tabari G Sorbus torminalis , 81 , B., ,

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K 92 (2001) Vogelkirsche, Spitzahorn Vogelkirsche,und Elsbeere:(2001) Spitzahorn Baumarten wertvolle Drei in Sorbus torminalis Salis eller, R.(1993) - 399. . ,

Sorbus torminalis S L., (2013) auli he in al , A. &, A. . , Pustahija , UK The Silviculture F ofTrees usedinBritish ,

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growing Turkey. wild in Towardsgenome and a size chromosome database number , F. (2011), F. - -

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Beyer Forschungszentrum Waldökosysteme, Univer E

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Asian Journal of Plant Sciences Plant of Asian Journal Restoration EcologyRestoration attentoleranz vegetativerattentoleranz Verjü . Forst Forst Effectiveness oftheMiyawaki methodin Effectiveness , Revue Française Forestière , F., , - Bogunić, F.,Bogunić, Bašić, Muratović, N., Catrice, E., si u si er ngungskon e e Forst Forst . Wommelsdorf Wommelsdorf und Landscape andEcolLandscape und Functional Ecology Functional Holz -

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, , This article isprotected rights by All copyright. reserved. AcceptedTakos TabandehTabari M.,Nadoushan K. A., H.M.,Espahbodi Szymura,Influence T.H., M.& Szymura, M.(2014) relief Pietrzak, and ofland soil Szymura, does recent T.H.How reflect (2012) vegetation Stoltze &Stănescu,(1997) V.,ofletea,N. Popescu, O. Stace, & C.A.,Preston,C.D. Pearman, D.A. (2014) Stace, C.(2010) Article (1878) &Sowerby, J.E. Sowerby, J. ofletea Snow, H.,Nielsen,Sjöman, A.B.& innorthern(2012) for Oprea,Trees environments urban A. , I.A.&, G.S Efthimiou, B. , M.& species northern autumn sownina Sciences characteristics of tormin properties onstand structure ofovergrowncoppice forests oak origin with management? Copenhagen Energiministeriet, Miljøundersøgelser Danmarks og Skov Editura Ceres, BotanicalIreland, Britainand UK. Bristol, Societyof Cambr UK. (ed. stand characteristics. avium statcondiiile sălbatic Republic ofMoldavia. parts of Central Europe Letters of Balkan flora: C , N.,Târziu,L. D.,Spârchez,& G. Curtu,

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Snow, D. (1988) (1988) D.Snow,

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Dendrobiology S , , Denmark. . ionale idearboret 2760

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Sorbus tor Sorbus - – values in 343 taxa withnovelvalues in343taxa forvalues 242. 2763.

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. . This article isprotected rights by All copyright. reserved. Wagenführ, R.(1996) Baričević,Vukelić, &I. J., (2010) D. Šapić, Nomenclatural Ví Velička, výzkum Dnešní M.(1993) Moravě. naseverovýchodní pvodních dřevin I.,Velichkov,Zlatanov, T. & G.(2007)oftree Popov, deciduous thenative Response Velebil Bekci,Var,& B. M., Uthoff Úr Tuzlacı,II: E.&(1999) Turkish plants.Part(Isparta). Erol, M.K. folkmedicinal Eğirdir Turker Tsitsa Tsitsa Trueman,I., M.& (2013) Poulton, Reade, P. Termena t, P., Lep t, P., Accepted Articleadnícek - - , , A.U., , A.U., Tzardi, E., Loukis,Tzardi, E., A.& S. Philianos, Tzardi, E.

, Hacquetia the association Preslia species: revision of a biosystematics leskéhoea v emskéhoy ní muOavě. érieříro ě A: reforestation vegetation State Forest 'Byala in reka' Balkan Range)cutting (eastern toclear and Vltava river valley. Journal ofBiotechnology Sorbus torminalis Umweltvorsorge Republic Fitoterapia Viscum album Fitoterapia 62 Pisces Newbury, Publications, UK. of P

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, 28 Sorbus , (2002) Achtung, Sorbus!

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) rz, L. rz, programme (2012) Regional )

in the territory regional inPiła oftheforests directorate thestate

living on living Wasp larvae on fruits and seeds, on Rosaceae on seeds, fruitson and larvae Wasp Rosaceae primarily leaves, skeletonizing Sawflylarvae Flo to restricted Leafrolling, to restricted Leaves, of pest a Primarily to Restricted Commonon on primarily Galling, torminali Forms on primarily Galling, on primarily Galling, Galling notes Ecological Larvae, primarily Rosaceae Larvae,primarily hosts of widerange Larvae, rangeof hosts Wide Wide Rosaceae on seeds, fruitson and larvae Wasp heavy canbe infestation

(a case northernIran). forests of inthe study wers,leaves, rangeofhosts brown wart brown

s

Sorbus torminalis torminalis Sorbus

Rosaceae, larvae and adults and larvae Rosaceae, S. torminalis S. , S. torminalis S.

6 - Malus , paper 42. , paper like pustules on the leaves of of the on leaves likepustules

Sorbus S. aucuparia S. Pyrus S. aucuparia S.

Sorbus

, larvae and adults and larvae , of conservation andof restitution conservation

and and

a major host major a S. aucuparia S. Islesin the British

and and

Malus – 18

.

;

S. S.

4, 9 4, 11 11 Source 2 2 2 2 2 8 2, 2 2 2 2 3 2 2 11 11

, 10

This article isprotected rights by All copyright. reserved. (2007). Wilson ( Sources: 1 sorbi Contarinia Cecidomyiidae Diptera dispar Xyleborus rugulosus S. Scolytus ruficornis Magdalis conspersus A. chevrolati Anthonomus Curculionidae cupreus R. caeruleus Rhynchites Attelabidae Coleoptera padella Yponomeuta conjugella Argyresthia Yponomeutidae bradfordi Ectodemia torminalis S. mespilicola Stigmella Nepticulidae clerkella Lyonetia crataegi Bucculatrix Lyonetiidae P Schiffermuller) cydoniella Phyllonorycter scoticella P. anglicella Gracillariidae Lepidoptera trapezina Cosmia Noctuidae brumata Operophtera 1993c .

Accepted Articlemespilella

); 5, Emmet (1976); 6, ); 5,Emmet(1976);

mali (

1985b

(L.)

,

(Stainton) (M

(Wood) (Hubner)

(micro

(Bechstein) Alford 2, (1984);

Desbrochers

ü

ller, P.W.J.) ller, Kieffer

(Fabricius ); 8,

(L.) (L.)

(Stainton) (L.)

-

(L.) Zeller

Emmet

moths) (Degeer)

(Frey)

(L.)

Harper Zeller Desbrochers

(Denis & (Denis

)

( 1981 Biological Centre Records (2016 Emmet, Watkinson Emmet, Watkinson & Wilson

); 9, Lanier of pest a Primarily rangeof hosts Wide rangeof hosts Wide Rosaceae leaves on and buds Vegetative Rosaceae to Restricted Rosaceae to Restricted on Adults,primarily Rosaceae other on found shoots; Larvaeon Rosaceae on Widespread mineron Leaf suggestsit A to minerrestricted Leaf mineron Leaf Rosaceae miner,primarily on Leaf minerLeaf minerRosaceae on Leaf minerRosaceae on Leaf mineron Leaf minerRosaceae on Leaf Larvaeleaves on hosts of widerange Larvae, Galling, restricted to to restricted Galling, dult found on found dult ( 1993b

is

a food plant food a Sorbus torminalis Sorbus Sorbus S ); 10,

. t orminalis Pyrus

Prunus Sorbus

and and

Rich Rich S. torminalis S.

Malus ( 1985a

and and and mined fallen leaf minedleaf fallen and ) et al ;

3, CIE (1969); 4, 3, CIE (1969);

Sorbus

); 7, .

(

2010

Emmet, Watkinson &Emmet, Watkinson spp.

); 11

,

Ripka Ripka Drapier 5 2 2 2 6 2 2 6 2 2 2 1 1 2 2 2 2 2 2 2 2 2 , , , ,

5 7 7 7

This article isprotected rights by All copyright. reserved. Accepted squarrosa Pholiota Inocybe flavella cesatiiCrepidotus Clitocybe nebularis gallica Armillaria MoncalvoVilgalys & Ampulloclitocybe clavipes Amanita pantherina Agrocybe praecox Agaricales Basidiomycota D. fissa concentricaDaldinia Xylariales Enerthenema papillatum Stemonitida Bombardia bombarda Sordariales Venturia inaequalis Spilocaea Pleosporales Peziza micropus Pezizales Punctelia subrudecta ArticleLecanorales Nectria galligena Gibberella baccata Hypocreales Lachnum niveum H. fructigenus Hymenoscyphus caudatus Helotiales Septoria sorbi Capnodiales Ascomycota Species/classification and n 2 Table omenclature

Lloyd Fungi

sp.

Fr.

(Ces.) Fuckel (Bull.)Fr. and Slime moulds and Slime

P. Karst.P.

Pers.

(R. Hedw.) P. Karst. P. (R. Hedw.)

Bres.

from Marxm. & Romagn. (Pers.) Fayod(Pers.) (Rabenh.) Sacc.

(Wallr.) Sacc.(Wallr.) (Weigel) P.Kumm.

(Batsch) Kumm. P. (Cooke) G. Winter (DC.) Kumm. P.

(Bolton) Ces. &Not. (Bolton) De (Nyl.) Krog

(Batsch) J. Schröt.(Batsch) J.

the the

(Pers.) Rostaf.

(P. Karst.) Dennis (Pers.) Redhead, Lutzoni,(Pers.) Redhead, B

ritish Mycological(2016 ritish Society

associated with

Sorbus torminalis

Fallen wood Bark Bark Fallen branch Old seeds Dead leaves Fallen leaf Ecological notes Fallen wood,stump under treeWood Fallen twig dead Soil and soil Wood Soil Soil Soil Wood Wood Bark Bark Dying leaves Bark )

in the Isles.British

Data This article isprotected rights by All copyright. reserved. Accepted A. pomiformis Arcyria cinerea Trichiida Bremek. Paradiacheopsis solitaria Stemonitida E. minutum Echinostelium fragile Echinostelida Amoebozoa Tremella mesenterica Tremellales Brevicellicium olivascens Trechisporales ArticleThelephora terrestris Thelephorales Stereum rugosum nigricans Russula subdulcis Lactarius Russulales Polyporus leptocephalus Phlebia radiata fissilis Aurantiporus Polyporales Ochropsora ariae Pucciniales Phellinus igniarius Inonotus hispidus Hymenochaetales mesentericaAuricularia Auriculariales Pleurotus cornucopiae

de Bary

(Leers) Rostaf.

(Bull.) Pers. Fr.

(Pers.) Fr. (Pers.) (Bull.) Karst. P. (Fuckel) Ramsb. (Fuckel) (Bull.) Fr.

(L.)Quél.

(Pers.) Gray

(Berk. & H. M.A.Curtis) Jahn

Ehrh. Nann. Retz.

(Paulet) Rolland (Paulet)

(Dicks.) Pers. (Jacq.) Fr.

(Bres.)Hjortstam &Larss. K.H. (Nann.

-

Bremek.

- Bremek.) Nann.

-

Living trunk Living branch and leafSoil litter Wood Fallen wood Fallen wood Wood Living leaves Living tree Living wood Stump Standing tree Bark Bark Living bark Bark Living bark Fallen wood Dead attached branch

This article isprotected rights by All copyright. reserved. Accepted DMAP software. mainly Hydrology, and B of Society Botanical the of Ecology members by collected for Centre Centre, Records Biological ( 1970; 10 a in record one 1 Fig. Article Figures

The distribution of distribution The +

) non ) - native 1970 native

- km square of the National Grid. ( Grid. National the of square km Sorbus torminalis Sorbus

onwards

( ; x ) non )

in the British Isles. British the in - native pre native ritain and Ireland and ritain ● ) native 1970 native ) -

1970 . Mapped by Mapped .

Each dot represents dot Each

onwards , using Dr A Dr using , Colin Harrower Colin ; ( ; rm records from 

. native pre native

Morton’s at least least at ,

This article isprotected rights by All copyright. reserved. Accepted EUFORGENForest(European P Resources Genetic Fig. Article 2

The natural distribution area of natural distribution

Sorbus torminalis rogramme

in Europe. Map courtesy of

; www.euforgen.org ).

This article isprotected rights by All copyright. reserved. Accepted torminalis Fig. 3 Article

Examples oftheExamples v . Scale bar ,

1 cm. From 1 cm. ariation

in

Rich Rich lea

f shape found betweenf shape found of trees different et al .

( 2010 ).

Sorbus This article isprotected rights by All copyright. reserved. Accepted above ground. excavated appr Fig. 4 Article Seedlings of

oximately (a) 1, (b) 2 and (c) Sorbus torminalis

growing under parent trees at Keele University, UK, 6

weeks after the first appearance of the seedlings