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Home , Allophyes, Bruchus rufimanus, Carabus granulatus, Carabus monilis, Dromius quadrimaculatus, Gastrallus

OB ERVATIO 0 ' THE E TO:\tOFA A OF \VE T PALAEARCTIC WITH PARTICt:'LAR REFERENCE TO LO GOO~ ~1ARSH, \VORCE TERSHIRE,

Whitehead, P.F. Moor Ley!>. Lntle Combenon. Pcl"hore. Worce,Ler!>hire WR I 0 3EH. England

The tree of life is inextricably linf..ed to the life of the tree ... J>. E Whitehead

Separating the wood from the trees In recent ) ear' auenuon hru. focussed on the role played h~ mature. senescent and monbund tree~ a..~ support syste~ for invertebrate,, man) ol wh1ch. b) relerence to the fo,sil record exemplify10g the•r changed di\LrihUiion' in \pace and time ObJects of lan era/., 1999: Franc, 1995, 1999: Harding and Ro,e, 19K6: Key, 1996. Read eta/., 2001. Trave. ::?.003!. Amongst oak-as'>Oeiated imenetmue,, the Hemut Om1odemw eremito (Scopoli) I' the subject of a Eumpean Umon Habitat Dtrecuve <92/43 CEEl. and funding from the European Umon L1fe ProJect ha!> been employed Lo ~afeguard 11. Bnta10. and the are linered With anc1ent und \Cter.m trees. Bnush veteran tree~ have earned a great deal of reverent ial regard (e.g. Pakenham, 1996). but there are no grounds to uppose that Bntam\ anc1ent tree stocks are unique. ;md many such tree~ and their site~ awan des~:ription and 'tudy throughout the reg10n. One only has LO cons1der the number ul \Clemo Ohve (Olea europaeu L10naeus) and other treeJ. m C or English farm-orchards) or es,entiall) equidistant (e.g.

171 figure I Dunwmhc Park. F.a't Yorbhire. England. E'tabJi,hcd an \D 1713. \\ ath ,ub,tanual Victonnn infiuen"e, tha' parkland '' '~nt: ot the nchc,a 'lie~ lor arboreal imcrtcbrJtC' 10 northern England Septcmhcr :! 1994

1,\II photo' lly author)

Figure 2. Elmlcy Ca,tlc Det:r Park. Bredon Hill, Worceo;tcr-h•rc. England. EmparkcJ tn AD I :!34 Iolio\\ ing 'orman tradations of selecting an ulrcad) ha,tonc "te, the tree' centre around an Iron Age hilllort. Brellon Hill i' one uf the forcmo t ate lor 10\'crtebrute biodiversity in the Bntash ble . Ckwbcr 29 1995. Qum unx-defincd orchards, managed oak \\OOdland~ nf l.xtramadura, Spam: pre,umed 'acum-orcharch' uf ~.:entral Europe). Thetr gro\!oth, mas and \olume rna) be optimal, lmngmg benefib to thetr a'socmted entomofaunas. both m term of compo Ilion and taged 'u~.:ce"ton. In 'orne large-scale managed s> terns ~ompki and Jan,"e\!otl't 12001) lbt 11564 'peLll'' ol antmab, tmm Btalo\\ ieza Forest that ~traddlc~ and Bclaru'. Where 'uc.:h nch pnsune lore''' are draped aero" d!,scctcd tupograph) a.nd n r.mgc of altnude , combmauon' of proce,,e, may te... d to c\lcn'l\ e lragmcntauon of the fore't edge. In Turk~y. the high Tauru~ :\tountams arc dotted 10 places \!otth e'traordtnaJ') andent pollari.led and/or coppiccd junipers: such trees arc either ehmauc l'olate,, hut rn

Fi~ure 3. ll".tn10U licld\, Lycia.. Turke). A bioucall} ru:h 'culture-,a\annah' 11o1th oak Qunnn puhelcf/1.1. May 23 1997.

173 Fi~rc 4 Bialo" 1e1a hlfe~t. nonh-east Poland. The Unwldrcltkt lucanlll beetle Ceroch111 c IJn 10/llt'lmu> • •m and1cator ~pecies of mtact \lo ell·presened arboreal ')'tems Scptemb<:r '27 1998 the tlead\Hxxl 111\Crtcbr:lle fauna of such tree-. rna_> e:~.ceed that 111 tho'e \\lthm the cluseJ fore't can\>p). In Balkan montane ')'tcm~. de"~:cated hulks of cedar.. htter the hii!-Jtle Uf>!-lu~ of the fore.,t core: forest edges in the'e pla~.;es are d1flu,c and wbjeCt to rapid!) \acillatmg climates, rather than tl10'e created b:,; the tnreSl lt,clt. R.1gged forest edges ma) interdigitate "'uh \Ubalpme herb-domi­ nated 'Y'tems: the ')han ecology 1s therefore no more intact than that of the anthroposy-.Jems ulreml) cited. In south-central Europe oak-dominated forest '' broken b) xerothemlit: c1>nditions: in such place' the forest noor " penetrated by stepp1c herbs and '\Cmthenmc invertebrates. Jenlk ( 1979) illustmtes well the inllu.:n~c of clunatc on fore~t structure. What climalllally·fragmcntc:u forest has in ~:ommon '' llh arboreal anthropo~ystem~ (Figs 3, 5. 11, 12) is an abilit} to su,taiO popul.ttmn' ol man) rare tmenebrate' (03JOl 19fl5, Trmt', "!.IXH) One reason lor this is that arboreal fungi v. hich prepare li\ 10g tree for invertebrate coloni ation may he nhle to de\elop 'trong populati<'n'; 'pore ma) perhap' b

174 figure 5 \\md"->r, Berk,h1re, England. the pnme Bnu'h sue fnr oak-a' ocmted 1mertcbrate,. An anctent oak Qut'n'll.\ robur. the tanned he:trtvHxl(f Y.eakcned and fragmented h) ha.,ic.llom)t:Ollne fungi. inducing bole 'hear llus group Ill entomologl\l~. mdudmg Dr ba Sprecher. Dr Peter Zach, and IO'Ide the role. tr P.M . Hammond, recorded the -.carcc beetle<> Pmcrat nu t1bwi1S (larvae), Ampcd111 mn./uw/1\ (larvae). Dryoplulwnu cnrtwalls and P/at\7111r nluulnn c.•n June 27 2002.

\vhere the1r commensal\, the ant' La1iu1 }itlilllllmU\ tLatrclllcJ and /.i.llllll /Jrunncu.\ Latreille resp.:cuvely. proliferate. On the Gred.. 1'land ol Th!Nh, open­ grown \eteran \lak tree~ are uncommon, but large bolated e\.tlllplc' ol Qunw.1 mlllprmm Wehb ,u.,win key geogmphical population' ol the longhorn beetle: Ceramhn rl'lllllfllll Brulle among'' other' The rare. h1ghl} 'pccmh,l'd /:un/1111 mwata Ench,on !Coleoptera StaphylinidaeJ. regarded a' ~tcnot{lpil' to oak tot the 'uhgenus Quctt"lll J Y.here 11 occurs :.ubcorticall) .md cnmmen..,;tll) Y.llh thl' ant u111111 bnmne1n, 1' ahu knoY.n from a smgle hedcgroY. ha\\lhom and 11 mgle culll\ated apple tree m Bntam. 10 th1-. ln,tance \\llh the ant/.awH mgt r l..tnnaeu' IWhuc:head, per . ob .), a notc:d b} Rc:mer I 1909) A \tudy on 11 mgle c:xotic Turke) Oak (Qw rrur cem.r l.mnacu ) (\\ hlte­ head. 1996b) planted 10 p~ture m Worce,ter-hirc, England. re\ealed o nch entomoluunn \\ llh a consen ation \-alue quanulit.>d a< greater than that of the enure Fore t of Dc:an 10 adJacent Glouce,ter-hlre A CUf'Of) tud). spannmg !'(lme 40 nunutes, of an opcn-groy. n Qunnts pube{Cf'm \\~lldeno\\ 10 an anul'nt agn­ '}'tem m L)cta, Turke~ . dunng Ma) 1997. re\ealed '"'" 'pecaes of be~·tle and one 'pecie' of :-.:eumptera nc:v. to -.c1ence (\\'hllehead. unpuh 1. In 'uch c:":\treme ~a'e tt ma) he rea.m,auon 'lll'Ce'' nf UniCI/drdd;t 10\ ertehr.lle' 10 relation to tree den'll) fRamu,, ::!OO ~ l In the .:entre

175 of the htstonc Cit) of Bath. Somerset. :1 veteran Cau ..· a t.m \\'ingnut Ptemcarw frcLumfoiiCI (!..nmhl Spach 'urrxm' the \\ 1de~pread tk.tthHxlll \lrltuwtu' 11//tl\lll ( f ourem) 1 (Cnleopterd. Elatcmlae). The 'nuauon 1 no dtflerent m the 1\earcuc Rcg1un In the rnmhhng DisnC)\\Orld comple" of Orlando, Ronda. dead\\OI.ld cl.lleml .md other hectic 'un 11e m v,olated tree' (J.i\. and P.r. Whnehcad. per . ob,,), Some Fumpean to\\n' and c1ue,. whkh ha1e lx:en httle·nHllltlied ,m.::e thc1r met.he\al e'tabJi,hment. pre-.ent fe\\ barrie~ to ~tcnoh1p1l' ltrbore,tl tmenehmh: . In l999mer J()()(JO exn hole~ of the 'uhcortll'al bl'ctle Gtlllrolflt, 1/IWUirgllrattll (Colcnpterd. Anobudael were ob,encd <111 a 'mgk tree m the then I W·)C;tr-Qid arh11retum at Ban,ka Stlil\ nica. Slt11akia I Whttehc,td, 1999 J. uml m Brau,l.thl, l/nraldrelila m'ect~ occur v.ithtn the cunnurhatinn (\'htdumr r ranc. per,. comrn. l. The Kcmcrtnn btatc 111 Worce~ tershire. Englund. c'tend' m cr 250 hcct

The imertebr utcs of oal.: Oak 1 also noted lor 11, great biodil'ei'Stt) ; Europe. n t' 1k arc thought to t.e coloni ed b) c-6000 'pcc1e' of m'ect (Dajoz. 20()()) anJ 111 )Jm 1k1.1 .11 lca't 286 'flCCie of par:l\IIIC \\a~p~ occur on oak (Patocka t1 ol 1<)1}9) "nne 225 'pedes of aphu.h arc dependant on the Quann (Biacknnn 1nd t.~bWp . 199~) . Sune' of 1m enehrdte' ..:har.leteri'e the \anou' ubfam1he of nak throughout the1r ranges, nlthough data from the more outl)1ng pan' ol the Palaearct1c Regton 1 ollen unpuhlbhed or SC"d.Ot). lmenehrntc- are able w colonise all pan of oak trees, and some gall-forming cympid \\a p alternate betw ccn arboreal and terreo;triaJ generation' a' a mean of 01 ercornmg sea,onal chmauc \artauon,. M1te' (Acanna) figure \CI') pmmmcntl) 111 nncu:nt \\ocldland h1ocoenose (T'ru hmg. 1994: Tm e. 2003 ). although the sp.:ctah I knov.ledgc requ1red ha! tended to limn intere-.t in the group Lanne t>l the 1.1g hl:etle UIC't/1111\ cenu~ Ltnnacu' (Coh:optera· Lut:amdiic) Iced mthc rhtl\"phcrc 1Spred1er , 2003 J. their pre encc olten onl) re., calcd \\hen the hl:etle~ appe.tr. 0\\en (2000) added s1gntlicantly to knowledge ol the entomolauna ol the oaJ.: rh110 phcre. ln\ertcbrates :.t'~octated w1th other part' ol the tree ma) he equull)

176 unohLTU,I\C :->om~ oak mirid bug\ rna~ 'pend up to II munths a' mae." h1h1 ome ad\CNI)-selcctcd deachHxld beetle' rna) pend three or four )!!at'S a' larvae. but onI) three 10 1\ v. ed.' a' beetle,. Ad\ ef'JI) -selected 'pecrc ( H nsu Green lade. 19 31. \\htch rna) aJ,o mdude mile' ICro ,Je). 1977). are s:ud to he ·K-~Iected • Some K-selectcd lxetle' fonn clo..ed pupulauon v.llhm the tree, and do not need to lea\e 11 to matntain 'iable pupulauon . Numcrou' pcc1e ol lanae feed both mtemall) and extemall) on oak fohage. although man) of thee are pol:.phagou' Patock:l eta/. tl999J cue 249 'pccu:s of on SJO\ :Wan oak': th1s ligure n e lurlher :.outh '"llh o\ er 300 spc~;ic! hemg rccotded from Span1'h oak .... Mom 11974) rc' lt:\\c:d the cntnmofuuna ut Brill'h 11ak. In the j,land group' of thC' kdlt~rmnc.m Sea ba'm and m Turk~). \\CC'\ II' of the genu' Phyllobitn and PolwlnHII\ arc t.'pcc.all) sp..:ciosc: thL'..C urc nx11lccdcr.. U.\ lar.ae. but along \\llh man) other hecth:s. they swarm in 'pnng on the flower> ot Quercus cal/iprino.1. QuerC'II\ nwcijem Lmnacus and Quercu1 ih~\ l.innacus. Few 10\CCh. such a' the strikmg red and blad: click hectic \mped111 carclmalt.l (SchiOdte) cColeoptcru: Eluteridae 1. ha\e e\ohed an ohhgate ~omm1tmcnt to oak. Amongst moth,, Plnl/odelma wilt r ijolia IDuponchel) cLepidoptera: La<;iocamp1dac l nl l·mncc. Spam and North A Inca teed' on oak-, ol both the -,ubgencra Qut'n 111 and Sc/cmpln11tNin·•. hut ..tpp; rent!) on nn nther tree~. The Oak Hav. k \1oth Hanunha quat IH (Dem' ;.n\1 Sc.:hitfemtullcr} cl.ep•dopte a Sphmgtdae) occur' mrcl) on other ho,ttrce~. hut lA 't ampa querr IH cL•nn;.eu') Lcp1dopter..t: La •ocamp•t.lac) chooses I rum at lca't :!0 other generJ ot v.txld~ plant' Man~ lfl\crtebrate' arc not known to di~tingm'h bet\\ecn tht.> age and the cont.lmon ol the1r ho't tree,. Dn·ophihH fl(/1 oqt~adnmaculatul cDe Gecr 1 and HarptK't'rtt tlmracica Crallenl Cllcm•• ptera: Miridac 1 occur on oak tree' ol almo't uny age. a' t.lo man) ... pcc•c' ol gall-lonning qmr•d v.a'P'· Lun .1c of the Purpk fluiNrcak INm::eph) m' quernt 1 ( Lmnacus 11 ( Lcp1doptcm: L)c:tcmd~) IX'Cur> cquall} rcachlJ on both \lgomu' and 1etcr:m oak trees>

177 (DeGeer) l Orthoptera· Ieltlgonudae) tx-cur.- on , 1: pccmlly of the suhgenu, Qu• r. , • 111 a "Ilk 'anet} ot age' and com.huon~ Ca.;e-bcanng moth I an uc ot the genu l.uffia (l..ep•doptt!ra. p,,d idae) often prefer 1 olated \Ctcran oak trees. "1th their nch flora ot hchen,, the Zebra Spider .~a/lit 111 <.l'braneu~ lC 1.. K1x-h) (Araneae: Snluculac) nlso ha' a c tear preferem:-c ('o marked that the author ha' reterrcc.l to it as the \eteran Tree Sp•der) lor the d~pl~ corrugatt!d lu:hen-co\ered bar!.; nl expo'>t'd anc1ent .md \Cteran oak tree' \1.-lth the1r li11er'il) nf nuaocli· mate,, ·•' do the predatol) larvae of Jfalucluus btpultularm (lumaeu'' (Co· lcoptcro· Malachildac J Wce\lb of the genu' Curwlio !Coleoptera: Curcuhomc.l.Ic) r.mgc: '-"•del} UC:TO" llJ(.; ~UO£CI1CTCI, Ill this genus C. \'ifiOSII\ foabm:ius 'tanc.J, apart i.l\ an mquilmc 111 gall, ot the C) mpid Biorhr:.u pal/ida IOh\lcr) (H) mc•wptera: Cympidac). Lcat-numng \I. eel ib of the genu-. Ordtt•.\tC\ (Cnlcuptcr.t· Curc:uliomdacl haw ulsu colonised exotic planted deciduous in lud" tmportant predator' m Wc~tcm P<~lacan.:tic oal.:\1.-oods. Arhorc.II ground beetle' of the genu' Dmmru' (Colcoptcr.J· Carahidac) arc knoY.n to breed on ancient Cork Oak IQut'rt"ltS 111her Lmna~uq in Britam r\Vhuehcad, per . obs.). Antheraea pem\1 (Guenn·Ment:lillel rl..ep•· doptera: Satumudae) pnnide, an intere,ting example of an urigmall) F.a't Palaearctic pec1e' e tablhhed on Holm Oak Quaru~ tle.x 111 the Balearic Island ... . The 'tnkmg nuchenorrh) n.. han Ledra au rita (Lmn:~eu' I I Hcrmptem Lcdndae l occurs uncommon!) on apple IMahH 'PP·l and maple (Acer .;pp.l tree'

Figure 7. Gcrakn,, 7.ak) nthlh, Greece. Coa,tally-mtluenc~ Q~~t•n 111 con ijaa \l.tlOdland, ~upportrng a dhllncll\e 'u1te ol •menehrate\. The hou,e, con,truetcd m Phc~:ene hme) andstone..once accommodated a fanul) of 1:! p<:(lplc. M.t~ 19 1999.

178 (\\'hnchead, 1991), hut more u ually on dc~Jccmed \eteran Pedunculate Oak Quercu mbur I mnaeu,, and 10 the ~outhem Europe on I lolm Oak Quercu5 rler

\\ood, t~ d~clinl', and the entomofauna \\'hat change most sJgnJficantly v.1th the age of the oak tree 1 the conc.huon and tructure of barl.: and v.ood. and. ot .. oursc. the ~.onfonnauon of the tree 1to;clf Th1~ " reflcctcJ m change~ in the enotomotauna of y.ooJ, panicularly \\hen 111 degraded h) arboreal fungi. A' a tree age,, ~o JLS energ) reqUJremenh mcreasc Annual nng~ dccrca~ 10 \\ldth "'th age becau'c ol the energy requ1red b) a -;emle tree to lay them dm\ n 0\ cr an incrca"ngl) large hole.: cm:umlercn~.e. As oak tree' age the.:) e\pcnd con"derable amount\ of energ) in tilling their redundant ,e,,ch \\Jth compte" aromatic comJXlUnJ, and tanmn' tTiwm.t,, :!CXXII. Tanned \~c.lc.ld "eflc.:cli\ely in'>(.>cticidal. but apart lrorn tung1. 'nmc.: in\cc.:l !:mac. ut hectb in particular. ha\'C found ways in v.hich 10 htl'alo.. these compte>. organic molecules, oltcn by gut-contained celluloly uc COl) mcs. l..af\ ae of lhl' Goal '~,1oth Co1111.1 c!l\1111 (1 innaeusl (Lepidoptera Cn"idae), llcrmJI Beetle D1mm/amtr er,·mrra I Ramu' and !Iedin. 2001 l. chafer.. nf the gl'nu' ,\It unmrc 1111 !Coleoptera: S~.aro~hac!c..laeJ tTau11n. :!000. m pre": Whnl'head. 20031 .•tnd longhorn beetles ot the genus Camnhyx !Coleoptera: Ceramby .. lc..l:tc) are all able to ex1't m the lanncc.J hean .... ood of oak: the larvae and adult~ :.tllnctimes 'mell ol leather. Th y are the largl'r agenh of excavation and c.:ulll\ alum t•f.,.. nod and the produc:twn lrom 11 c.•l \\c.IOJ mould. degraded or rec)clcd tanned hcan\\ood I fig. 8). Wtxld mnulc.J h a cumparnthely inert material, frequcntl) the c.Jehgmlil'd pnlduct of fungallU:tJ\ II) • .,., h1ch "hen not de,u:cated. as for e:~.runple by fragmcmauon of the tree, may remrun m ,;ru for decade,,

Fi~ure 8 Llan.lrth, ~1nnmouth,lurc . Wales. Queen Europc.m hclml'l \hpu cmbm, ahnuttn con,trucl u Wtnter cell m Y.c.x>dmould llcc.J, tn !allen nat.,; pollard Quernl\ mh11r. Nmemhcr II :wo~.

179 \\1xxl mould he(], 'UP{l\"'11 large numbers of Ill\ crtcornlcs, man) ol '' h11.:h are scarce or r.1rc gencrnll). \le,.ander 1200::!) e'llmaled thai nf "JM)(l 10\Crtc­ bnue a SOCilllcd "1th \I.CXxl decay proce~'e" m Bntam. ~8 '1 ha\e lonna! conser\'auun talus olone f1'rm or another. \Vhcn an anc1cnt oak tree tall' Y.ood mould contained vv 1thm il rna~ per,isl for det."l!dc ... and pro\ id1ng 11 doe' no1 dc,u;~.;atc. '-~- 111 continue to support rare imertc:bmtes Some msects, uch as the beclle Orwoda m01ul11a Keaau !Coleop1era Staph~hmdac) h.l\e hccomc spcc1ah ed and some examples rna) become '-tllllCY. hal dcp•gmcnh.'O. ,,, a CC'-(Xlll'e to continual darl..ne''· populations in ''<~ble h.lnllat '-~- irlun a 'mglc tree can prooabl) 'uf\1\C mddini1ely Some beetle' li\111!! m h1gh-,olume oak \Hxxl mould lll.IY ubo exh1b11 vunauons 10 eye size and 'omatic rngmcntallon Ill rela11on to example' ot the same occurring in other hahllats nc:arh) (Whitehead. per,, obs.) A rare overlooked habitat is that of wood mould dcvclopmg cxlcmally. This i' a phenomenon thai occurs on oak trees hea\ il> cnloni,ed h) fossmial macru-111\ertchratc laJ'\ae. Lanae of Cerumbyx cerdo Linnaeus !Coleoptera: Ccrambyc•dael and other beetle~ 111 particular, may perforate the tree to such an

Figure 9. Malee, me, Lago di Garda, . Chaler beetle Potc11ia cuprea at floy,ers ol D ,g.,..oc.xl Comm fOn!1111nea. ~1a) 7 2000.

extent thai fr:c. -~ds de\ clop on the ground agmn<>t the bole. These bed, ma} uppon an enure fauna of other tn\ertebrate,, partkularl). in central Europe, larue ol ::arnbacid or cctonud chafen. lfi~- 91. Decorticated wood I f ig_ 101 perforated h) the cx11 hole' nl anoh1id bcctJe, b the pec1al hahiwt of the rare oak-d\\elling £uwem1{ capucma Ahren' 1>d to enter the gallene' ol anohild beetle' 111 thc1r quest for pre). Al:ls of 'amlali~m ma} also, m

ISO a pen.er:.e w a), enrkh the entomotauna ot oak tree • m the same WD) that dramau~ally 'udden rnechanl­ cal damage, or the instantancou-. txliling of \UP bi lightnmg .,triJ..e. can pn>duce di.,tinc:t launal fac1c., , Trees are nut 'de.,1gned' to be hollow or even cancellou' m the -.en"«! that txmes are. ~o that their hollow mg IS an entire!)' pa'>'I\C: prt><:C'>S (~1atthcck. 1998). The dramauc impact of basldiOm)COtlne fungi on the hi\tologu;al structure of wood has been well Figure 10. Longdon Marsh, Worcestershire. England. illustrated b) Open grown boundary oaJ.. tree u~cd a' MatthecJ.. and hving lence p(l-.t. Decortication following crown Kubler ( 1997. lig. fro~gmentation. noung abo to,., of .. ecliments and 96). Lo-., of wood .. ub.,equently damaged and deca)ed root base-,. ma:. 111 ill lea't one Odober II 1000. 'cn-.c pn>\c benelicial to the tree: 11 the annual nngc; are breached a.' a result, the tree', O\crall energ) requirement ma) be reduced to the e\tent that what rcm:un-. ol the tree may retnm '1gour. E1ther WD). the in,enebrate fauna of dead or deca)ed wood owe' C\CT)thmg to the 1mpact <111 1t of fung1. (.\.turra). 1974) badena, the \aganes of chmatc and ph)'lologiLal ~tre'' · Thoma' (.~CX)(J) 1mphed that the to,, of hcamH>od m \Cteran tree., rna) re~uh in energ) reqchng proce,-.c-. that confer llenetits What 1s clear 1s that the stored energ) made D\aJlahle 1n the wood ol trees 1' of unmen-.c benefit to the ')net:olog) of the tree and to 11' h1ococntN!s. Wood pa H ' could nut mamtain the di\er..lt} of hk that we as\OCiah: with \etcran tree' ttxla)'. and thl'> pro, ides another rc:bon ''h) the 'upport role of

181 \Cteran tree . oak m parucul.tr. m hiologtcal ')'tCill! 1 potenttall} fur greater than that unpl) of tree' in !!Cncral.

The utili'>ummer ot 2tXl2. u un cy .,.. as made (Whuehe.td, :!IXI:!J nt the tn\ertebrate~ of Lnngdnn .\1ar,h, \\on:e,ter,htrc, 'ltu.ttc:d at 52"(16'=--: 02 l6'W !Figs 10.11.121. The sHe Y.as u"qutred 111 the late 199(}, b) the Worccster,htre Wildlife Tru,t, .,.. ith the ohjecti\e nl rc:~tnnng It as a Y.etland. Longdun Mar;h is n flu\ial basin near the wnllucnc:c ol the major Engh~h nudland me1s. the Se,em and A\on. It ex.tench mer an area nl c:7km • and ts flnMcd, as low a\ 11m 0 D., by allu\ ial stlts. The hi\tory and dc\t!lnpmcnt of the l.ongdon Mar'h basm is complex. The Severn Estuary ha' one nl the largest tidal ranges in the world and in the past. partic:ul:trly prior to river management regtmes. uual mlluence .,.. a..s felt a..' far upstream us Lnngdon. It is hdie\cd thai Lnngdun Marsh Lie\ eloped as a natural resenotr at lllncs 111 both htgh udes .md high mer lloods. although thc cutting of thc high-sided basin m

Figure I I. l.ongdt!ll Marsh. Wurce,ter,hire. England. Agm-~·n,)Sttm Y.llh woodland block,, uutgnw.n hedgerow' of willo.,.. Salit alba and '::ialn tral(ilr.f and open groY.n oak trees Qutltrt~ robur. Mahcrn llifJ., Wtth ouluded fronlm the ouckground. Jul) 20 2CXI2 . liS:! pro\ idl! I!\ tdcn\:e of a thud 'tgnili~ant regtonal 'ite (lullm.. mg Wlutl!head. I996a; Lou er al.. 1999) lor 10\ ertebrate-nch open-gm\\11 tr<.-e set m an :mthrupngent<. or 'cultural' land·u~c matnx (Table 1). The~ oak-a"ocmted 10\Crtehrate, extend mer 17 taxonumk orders. 101 fanuhC' and atlea~t 211 pectc \\ hnehead (:!

fiRurt.· 12 Longdon Mar,h. \\nrt·e,h:r-hin:, l:ngland. M<~ture and J'll,t·maturc de,iccated oak tree' Quan11 mhur ,u,t.unmg fa,lldtOU\ im.:rt.:br;ttc' in an agricultural land-u'e matn\ (Tahlc, I and 2). Suboptimal condition~ 10htb1ting UCll\ C tree J:!fO'II. th October fi 2()(Xl

Figure 13 'l1ddc II!} Wtx"1ll. \\orcestcr htre. England If) grophtlou' grnund beetle Camhur grcmulafl/1, a '(lCc.:te' \\.htch ma} v.mtt:r grcgam•u'l} under damp lotN~ har!.. tlf nak Qucrn11 mhur. Nm ember I X 200 I 00 ~ Order l'omil) S peci~ RDB N AW Niche

• Col McJ,,ndryidac AlxlcrJ btflc.\uo'a (Curt•~. J!Q!JI • h l\l.lg' I ep Gcornc~ndac AbrJ.\;J.!; gru,-.ulariata 1L.. 17581 foliage, A ran lhcndud;t Achac.u-.mca lun,ua ICicn:k, 17571 hark I cp NoctuJtl.Jc Acmnicta P'' CL., 17581 bark. I anal WC<)(lO Lcp Nocnnd.Jc Acronicta runuw. (1 ~ . 1758) barJ... Col Cncdncllidac Adalia dcccmpunrtata ( L., 17 'it\ l hark **Col Adcndac Aderu~ oculatu~ (PJ.... 1798) Nb J foliage. t\\ tgs u col Adcrid;Jc Adcru' populncu~ (Cr m Pt.. 17lJ6J Nb J t\\igs ( 'nl BuptC\tidac Agrilus hrgutlatus F. 1777 Nb 2 e>.ll holes Col I hlll'rJUUC Agnotes lincatus (L.. 1767) wood mould (bits) Col l ·lattndac Agnotcs nhscun1s (l. , 17581 wood mould (bits) ( ol l'l;uuid.tc Agnotcs sputator (L.. 1758) wood mould (hits) l.cp NoctuJd.ll: Allophyes ox.ym:anthac tl . 17'\8) lohage llcm Ctl.adclhdae Allygus mnttus IF.. 179-tJ barJ... l krn ( ll.tdclhdac Allygu'> modestus Scou. I R76 barJ... Col Tcnchrinnidac Alphituhius d1apcrinus !Pl., 1796) v.ood mnuld (hils) (\11 Car.Jtudac Amam aulica 1P1, 1796) v.nud rnuuld (hils) A ran Amaurohudae Amaurobius t.:nestr.ths (St.rocm, 17681 barJ..., undcr Cui Staphyhntdac Am•-.cha anal•' !GrJ\ Hi02l foliage Col \tapn) h nid.Jt' Ami'>cha lnn:ipaw Muh. &. Rc)', IX7~ \l.ood mould U ('nl llatcndac Ampedult cardmah~ (SchtiXhe. IH65) 2 Wt)(xl mould llar\a). harJ... 1 cp Nnctutdac Amphip)'ra pyr.mudea ( 1 .17581 fohagc II an ac l Col Scraptudac Ana\pll> humcralJ, (f . 1775) hole Col \~.:r aptiu.lac Ana~Jl" rcgimhart1 Sch1k. I !195 foliage Col linaptmlae Ana..spts thoractca IL.. 1758) h hark Col <. .trahida~ i\nt.:hiccra dol'>alc !Pontopp.. 176 ~~ harJ.... under Col <\nuhtidae Anohtum punctatum (De Gcer. 17741 \\lxxJ Col Staph) hmd.te AnOtylu~ rugo'u~ IF. 1775) wood. fallen II em \nthocorulac Anthocnris confusus Reuter. I !!84 tollage llcm <\nthocund.te A nthocons ncmorah' 1F . 1794) foliage Cui Denncsuda~ Anthrcnu~ fuscus 01..17!N harJ.... under (lanacl. hole Col Curculwmd.tc Archanu~ P> rrhoccras M sh. 1802 fohage Col Ocnncsudal' Anthrenus vcrha-.ct (L.. 17671 baric Jan ac Cui Staphyhnidae Astcnus tmmaculatus Stc . I !!'B N w

';JC Vt oe Cui Ccr ylonuJac: Cerylon hrstcmides barl.., umlcr ;, Clll Chrysomd rdac Chactocncma concinna fungus. laf\ ae Col Cud.te Cts pygmaeus (:\-1-.h . 180::!) barl... under: brand1. fallen Cui Crmhtdae Cli\i11a fnssor folmgc C nl I at11dndac Conrnrcara gabbosa (llhst 1791) tollage Col Chry,nmdidac Crcpidodcra plutus (Latr.. 1804) Joliage Col Cnptophagu.lac Cryptophagus pseudodcntatus Bruce. 1934 barl... under CluJ C ryptopadac Cryplops hortcnsis Lcac.:h. 1814 wood mould: harl... under c•• , [),•rn•c't idac Cte,ia.' 'erra Stc . 18 .~0 harl.: Col lll'tl!ndac Dt!ndrophilus pum:tatu' (Jih,t. 179::!) wood mould Palp;Jt Phal.1ng11dac: D1cranopalpu' ramusu' Sunon. 1909 foliage D1p T1puhdae Dictemd.a h1ma~.:ulata cL. 1761 l wood mould clanac. pupal'! Col <.,taph) hnidac Dmaraea angu,tula cGyll.. 1!1101 hart.:. under ·11~111 Pornpil•dac D1pogon ... ubintenncdiUs !Magrctu. IH!I6! hole Col Annhudac Dorcawma Chi),Omchna Strn. I x.n :! foliage Cui Anooiid.ll' DorcalC Stcnops1>etdac Gmphopsocus cructatu.' (l... 1768) foliage Col Anobitd.tc Hadrobregmu~ dcnticolh' (Cr. tn Pz. 1796> Nb \\ood • llym Ch.llmhdac lfaluchella rufipes 101.. 1790! INa) hart.. •Col Staphyhmd.te Hapalarea pygmaca CPk. 1800) wood mould (btl') A ran D)sdcndac llarpactca homhcrgt CScop .. 1763 J bark wood muuld (hils) Col Catabidac ll arpalu~ aflims (Schrank 178 1) Dtp Syrphtdac llelophilu\ pcndulus (L.. 1758) bark hark, under Col JlydmphtJid~tC llclophorus aqualtcus (L.. 1758) Col Anohmlac llemtcoelus lulvicomi\ (Stm .. 1817) tw t g~ wood mould (hils) Col rlatcndac llemtcreptdtu\ lnrtus (llbst, 1784) wood mould (htts) Col IIi stendal' I hster unicoi or L.. 1758 wood mould Chits) Clll Cun.:ulillnu.lac 11 )-pcr.l zoilus (Scop .. 1763) fohage A ran l Ill) phml.tc llypomma romutum (Blackwall, 1833) bart..; fohagc ••Jiem Ci~:adcllidac lassus l;mio (L., 1761) wood mould (btl\) Col Rh} nchlttdac lnvohulus c~eruleu' (De G, 1775) Arthru l'otonudac lsutomuru~ palustns (Muller, 1776) wood mould ..Col Clcmlac Kill) netes caeruleus (DeGecr. 1775) Nb 3 bark Lcp !'lnctmdae Lacanohia ulcrat·ea ( L.. 1758) fohage. Cui Tcncbrtuntdae hirta (L .• 1758> fohage barl.:; wcxxl mould (ne,tl • Hym l·urmicid.tc La,tus brunneus (Latr.. 1798) :\a wood mould !nest) II) Ill rumlll.'tdac La.siu\ Oavuo; (F.. 178.2) all parts • llym l·unntctdac Lasius fuliginosus (l.. atr., 1798) Col Staphyltntdac Lathrobtum gemtnum Kr.. 1857 roots ..Col Ccrarnbyc tdac Lctopus nebulosus (L.. 17'i8) hark. twig' Col ( 'ar.Jh1da~ Lt:Jstu' <.,punharhJs IF. 1775 I hark ,..Z}gt:n I ~pi,nl.ltldac Lepisma \accharina L.. 175S bar!.. under Orth Tctugunud.ll' Lcptoph)C' punctatissJma (B harl. ( ol Malachllll.tc ~1.1lachius b•pustulatu' (L.. 1751'11 barl... lan.Jc Col Canthaml.1c Millthinus fnmtaJt.., I hh . 180:!) Nh harl.. pup;! Col Canthamlac Malth harl.: Col \JipJUd,JC NLLmphtlrus humator (Cikdll-.ch, 17o7J \H'Icxlmould lhll'l *Aran \ranl'Jdac Ul:lenea umhmtJca I( Jerel.:. 1757 l barl.:. under Col St.lph) ltmdac Ck)pus olen' ~luller. 1704 \\IXKI mould lhatq llcm C'l\lld.u: Oharu' pan1cri Loc\\, I SIO 'b branch Palp.ll Phal.111gudac Oltgolophus Uldcn' cC.l •. Koch. I !\~(1) harl.: Col (.'lcnd.ae Opalo mul11' IL.. 17'ilit Nh J harl.:: "ll(l() mould 1hth l c,,, Cur' ullllntdae Orche,te' ulna (I... 175Xl hark Col Cur..:uh<•n•llac Orchcstc' pllo'u' (F.. 17H I 1 foliage Col ( urcuhl·ulus (Reuter. I 879) bark, undet • Col Col) lophtdar Orthoperus nigresl·en., Ste 1829 ;'l;h twigs Cut Chi) some hdar Out..:ma oiN:ura tSte 1811) \H>nd mould thll'} Pulmon i'.omudae O)l;ychtlu' alhanus 1~1tller, 18221 hark. under A ran Thonw.uJ.te Otyptila pr..lltwla IC L..Koch. llf\71 hark H)nt ve,plllae Pam"e'pula \Ulgari-. (l •. , I 7S!il hark under, WOIUS mtm·atu\ (Rat/., 18 n) IWI!_!S A tan Scgc,lltidac Scgcstria \CilllCulata ( l.. I 7'iX) harJ.. Col Cun.:uhnntdlium castaneum (llh't. J7l)7) \\ 1l

labh.• I. 1 ongdon Mar,h. Worce,tcr... hm!, England. 20

It j, oh~ncd that spec1es recorded from fungi are lc:v. . Under no circum­ 't:m.:es 'hnuld this he con\lrued a' implymg that fungi are of llule 1mponancc in Jn\ertehrate ecnlng). 7)<"7, ol all imenebrote occurrences were a"oc1ated v.ith habitah prepared for them by lungi: only 7.0% of occurrence' can he ascribed to purel) v.ond·.Jssl~iated spcc1es. and many of the obligate 'pcc1es 1n that case arc anobi1d beetles or other 'pecics a~soctated wtth the1r acu' It). It 1s unrxmam al\o to bear in mimi that few b10-assessors are able to work unilom1ly aero" a variety of taxonomK groups. so that in the present ca..e. infom1atton on mites is lackmg (Table I 1 ~C\enhcless the entomological usefulness ol the lntcracll\c: systems suswincd b) oak trees 1s v. ithout que uon. as i'> the biological \'alue ol v.ood per \f , Recent 'tud1es !Ha:~-.e. Topp and Zlch. 1998) ha\e suggested that a mimmum ol ..Wm of dead and dieing v.ood mcludmg fallen v.ood, 'hould be retamed v.1th111 each he~,;tare nf lore't to maintain biodiversll). :md that as man) a.s 18.000 md1' idual 1menehr.ne' rna) he ... upponed b) each cuh1c metre ol dead v.ood At Longdnn \1ar-h, m common v. ith other site' dommated h) open-gmv. n tree,, much ol th1c; laun.1 I' ab,cnt, and the bulk of the dead v.ood j, cont~uncd "1thm the II\ ing tree.

All facultathe Obligate

Foliage 'i2 (~04~) 26

:255 138 1'\J 117 (46t;i)

Table 2. l..ongdon \1ar,h, \\'orce,te~hrre. England. 2000 and 2

193 The relauvel) large number of facultative occurrcn e for bark and ~ood mould re,ult lrnm thl" u'e of these re~ources ac; 'heher- h) a mnge of -.~de, , or b) terrestnal predatol) invertebrates. The bark predatof" mclude the carah1d beetle Pttrmllchul maar: thi~ is a rare finding of a 'pec1e> nearl) ul~a)' ~ubterranean 10 fi"ile .;edimenL<,. Wood-mould predator~ include the ~car~c Carab11'i mnmli'i, much dechned but as'ociated ~1th old ummproved gra"lands and open ~uodland\ , Thi' Iauer occurs as an inten~ely green form. noted alc;o 10 an example of Carabu.'i l(rtlnulaws (fig. lJ).

A ckno,~ledgeme n ts I thank the many friends and coUeagues who have heightened my perception of b10~ystems over the ye~. particularly Dr. Peter Zach of the Slovak Academy uf Sc1ences. The Worcestershire Wildlife Trust k10dly pcrmmed reference to a study. and Mr. K. McGee (Drakes Broughton. Worccstcrshirc) kindly provided figs 9 and I 3. I also mention English Nature. who R<; the government's statutory agency. ha~ done ~o much to he1ghten appreciauon of arboreal '}'stem' m England.

Reference Alexander. K.N.A. 1999. The invertebrates of Britain's wood pastures. Bntr1h ~1,7/d/ife I 1: 10!1-1 17. Aleunder. K.!'i.A. 2002. The Invertebrates of hvwg and deca:.1ng umber 10 Bntain and Ireland - a provisional armotated checklist. Brrti1h Natllrt' Research Report 467. Peterborough. Blackman. R.L. and Eastop. V.F. 1994. Aphids on the world\ trt'cl. CAB lntemauonal, Wallingford. Buckland. PC and Dinnin. M. H. 1993. Holocene ~oodlands. the to,~il evidence In K1rb). K.J. and Drake, C.M. (ed'>). Dead wood maners: the ecology and conservation of saproxylic invcnebratcs m Britam English Nawre Scrence 7 6-20. Crossley. D.A.. Jr 1977 Onbaud mites and nutnent cychng In Dmdal, D.L (ed J. Bwlog'> of oribaud mites. SUNY-CESF. Syracu'e 71 8S Dajoz. R. 1965. Catalogue des Col~opteres de Ia Foret de 13 Ma\sam.. . 1-te Mt/ieu 15 (supplement): 1-207. DaJOL R 2000. lmecH and forests. Intercept Ltd, Andover. Elias S A 199~ Quaterrwn msecrs and rhetr mnronmena. Sm1th,onian ln~tnut1on Pre,s, Wa.-.hington and London. Fashlng. N.J. 1994. L1fe·h1~tory pauerns of asugmaud mhabnam' of Y.atrer-filled treeholes. In : Hout:k.. ~1.A. (ed.). Afius Chapman and Hall, London Fowle~. A.P.• Alexander, K.N.A. and Ke}. R.S 1999. The Saprox}hc Quain~ Index: evaluaung ~ooded habitat for the conservauon ol dcad·Y.Olld Coleoptera. Coleoprerist. 8 {3): 121-1~1 Franc. V. 1992. M}rrnecoph1lous beetle:. of Slo\akia ~•th 'pec1al reference to their endangerment and pe!".pecthe~ tor protectiOn. Acta. Unn Carol. Bwlogtca 36. 299-324. Franc. V. 1994. On the occurrence and biomdJcame value of 'everal rare 'pec1e' of the family Tetratomidae and Melandryidae (Coleoptera) m Slov.lkia. Biolo!lill Brattslm 49 (5 ): 723-728. Franc, V. 1997. Old tree~ m urban environments- refug1a for rare and endangered

19~ beetle!> I Coleoptera I. Acta lmner.5tta11s Carolmue BtologH'a.. 41 {3): 273-283. Green,)ade, P.J.M. 19 3. Ad\e,..,lty :.elecuon and the habttattemplet. Amencan Nawralw 122: 352-365 GutoY.,kl, J.:\.1 and Jaro,LCY.IC/, B. leili.J 2001. Catalogue oJ the fauna vj 8wlm11ezu Pnmnaf Forest. Forest Re~earch ln\l!tute 10 V.ar;:aw. Haase. V., Topp. W, and L..lch. P 1998. Etchen-TothoiL im Wirt-chafL,Y.ald als Leben,raurn flir) loh10nte lno,ekten. Zeuschrift fur Ok11logte und Naturschur: 7· 137-153. Hammond P"\1. and Hardmg, P.T. 1991 Saprox)lic 10\ertebrate assemblage' 10 Bnush Y.oodland': thetr consenauon significance and 11~ evaluation In· Read. J. led.) Pollard and 1eteran tree manaf'(ement. Corporauon of London. 30-37. llard10g, P.T. and Alexander K.N A 1994. The use ol saproxylic invertebrates in the selecuon ;md evaluauon of areas of rehc fore~t m pa~ture-woodlands. Brauh Journal of Emomolog\ and Natumllfuum 7

195 10 hulhm oaks 10 $\\eden In: Bo\\en. C.P. led.), Prr/(t't'dlllgl oj tlu.' :wcond pan f.uropetm cml}erence on ~apro."f)IIC heetlt•s, Rowlllollmw\, Lm~t•rmy ofwmlon, June :!002. People·~ Trust for End:mgered Spec1es, London. England. ~6 . Ramus, 1: and Hedin. J. 2001. The dispersal rate ol a tx.>ctle. Osmudemweremua. h' 10g 10 hollo\\ trees Oecolo~w I :!6: 363-370. Read, H, Fortang AS. \1arciau. R., Paino, H • Ander;,on, 1.. and Tard). B (eds.). :!001. Tcn/, jor prc!>en'ITif!. \\"1/(>dlam/ Bwdna~lf\ . Na<.:one:~. Reuter. E. 1909 fauna Gt•nnamca. D1e Kafer des Delll~cht'n Rerc he\. Band II. Lull Verlag Stuugan. Shotwn. F.W. and Osborne, P.J . 1965. The fauna ot the llomian lntergl.telal deposits at Neche! b. Birm10gham. Philo.wphica/ Tra111uc twm of the Rmal SoC'im· of London 8248: 353-378. Spe1ght. M. C. D. 1989 Saproxyltc invenebrates and the1r consenauon. Ntuun and £mmmment .~ene\ 4:!. Council of Europe. Stra\txmrg Sprecher. E. :!0(!3. The ~tatus of Lucanus cen·us 10 Sw1t1erland In Bowen. C.P. (ed.). PmceetiiiiJ!I of the second pan-European conft!fi'IIU 011 lclflrrJ\Yirc heetlc1, Rom/ Hollmrm. Unil·ersity of London, Jwu• 2002. Penple ' ~ Trust for Endangered SpeLies. London. England. 6-8. Taghanu.A. V. 199'\. Coleotten Carab1d1 del Parco 'ai'lonale d'Abruno. L.11tt prelunmar1 clet:li on;tmuml l'i1·en11 del Parco Na:.umalt• d' ~hm;::o I : 1- 1:!. Tau1m, P. :!000. Le genre Aleum.wctu.s KJ.rb~. 1827. Contribution a a connru,ancc et prec1~1on ~ur Ia d.Jstnbuuon de' espece~ (Cnleopter.t, Cetoni1dae, Tnchi10ae, Tnchiini). L'£ntonwlogiltt' 56 (6)' 231-2R I. TauLin. P. In pres,, Quelque~ localites coonue pour le~ espO;e~ tran\·ai es d'Aieumsflctul and the1r rare 10\Crtebrate,. Entomologi:,t'l Ga;me 43:303-30-t Whitehead, ~F. 1995. Stonwphi' graffii CholodkO\'kY (Homnptera· Aph1d1dae· Lachninae) new to the Bnush fauna. Entomoloqm'f Gtl:('(lt• 46:161 -16:!.

196 Whnehcad, P.I· 1996a The tmertebr..ue fauna of the Kemerton E'tatc. Worccqershtre. Part H. Arboreal habita~ and Beggar Boy' ~ctlantl . Jounra/ of the Ro)cll A~?ricu/wral Soe~eo of Enf!land 157: 170-190. Whttehead. P.F 1996b. Eucnemis capucina Ahren~. 181~ (Col .. Eucnemttlac) at three ue' m \\orce,ter..htre \\tth a remarkable beetle launa on Quurur Ct'rris L. at one tte. Enumwlo(l,ist's month/\ \fa1.1a:me 132·1X7-195. Whitehead, P.F 1996c. The northemmo t Bntish record ol EuT)u'a Hmwta Crich on. 1837 CColeoptera, Staph)hnidae). Entnllu,[of?ill'l Ga:me 47: :!55-256 Whuehead, P.F 1997. Beetle faunas of the European angto~pcnn Uma/d; problems and comple>.llte!>. Brologra, Bratislam 5212: 147-152. Whnehead. P.E 1999a. The Worce~ten,lure population of Ga.ltmllu! immarRIIlUtr/S (MUller, 1821) (Col.. Anobtidae). Entcmrolo~:i.lf '.1 Month/\ Ma~ta:inc 135 193-196 Whitehead. P.F. 1999b. Observations on. and a redscription of. Rlr\nclwnuH lrirtellur tr•·liller, 1862) (Coleoptera. Curcuhonrdae). a little-known Palaearcuc weevtl Emnmolngw's Ga:elfe 50 205-208. Whuehead, P.E :!000. SIO\akian Querc:eta; thetr temporal de\elopment and Unmldrelikt Coleoptera m relauon to arboreal refugta. Emomola~.:w's Ga:elfe 51 : 117-124. Whttehead. P.r. :!00~ . Lonfl,don Manh. ~\vrcesfl'nh~re: 2002 imatebrate lllfll?\ with pamcu/ar rejtrt'nct to arboreal specrel. Unpubh,hed report commh­ 'ionetl by Worce~ter,htre \\'ildhfe Tru~t. Whttehead, P.E 2003. The "

197