11>< 1101< 0( Fi" .. ~o<,!><'"(:;momlloiu Eooo',,<m.EdIt<d bl' IlO>O W \\<,n..,.j l)mJ A ~to"l.<.n

() 19SJ SCOPE, Pl.l>h>h«l bj' loll" Wd<l if< Sort> lIJ

CHAPTER 9

Post-fire Succession ofSmall·mammal andBird Communities

J.F. Fox

AIISTIIACT

Chang.. In the <eolog~ of ""an·m.mm.1 anJ "fd rommuni,i.. JurinS p<»'.!ir","""""ion at. r.".,,'ed for the :"oMh Amen,,"n boT..1fores, 'nJ 'undr., Data fromth" h..r"m" a...ummarized '~'''<ma'i",1I~'. "mph"';"i"! rompori""" of the rom­muni'~' prop<"i<"> "f ....II-m.mmal' .nd bi,J ....mbl""., lrom .arll p.N-f".Ih,ou~h 10 malur. >u""o<"onal ,tag<>, 10 ~ene,.I .• brwd tr.nd 01 ,ncrea", ,n ,~.,d;\'e"'t), >","".> r",~""". numbe".•nd con'um,"g hI"ma» i, l"u"J I", bnJoommun,"es Ihrough "-'=»'00. "h,l. mom.., de<:,e ..." For "".1I·m.mm.1 oom·mu",t,e>, numbel'>. biom.... ood COn'Um,"! bIom... are h'gher '" lbe berh ".,e, ..dIouer ,n lbe >!>ruh and ..pl,nc"age>, ,hoo ,n malure f,,'e,". >p<C1es d"'ef'>l'l of >maUmammal, "' h,gh and in,".'';"! op '0 ,be >!>rub ".ge. and ""'., in the ...pli"! 'la~e

tha" 'n mato," I"'e,'- "!lie... trend, un<loub!edl~ refl." <han~.. ,n 'he grounJ,,,~.t.'ion .nd the hel.ro!"n"i'~' of plant communnie, (botli '<",cally throu~h the're<: ".nop~. and bo'iwntallj', "'fle<tiOi p,,,,~,",,») "'ith '0"".";00.

9.1 I"TRODUCTIO",

Thi' chapter ron,iders chang.. In ,he rommun"l ecolog~ of ~all·m.mmaland bird a"embl.gos during pos'·fire '·egel'li'e ,u",e"ion ;n NorlhAmerican boreal fore'l. M,d.I.'itude e~rcriene<: wi'h plant suece"ion andecolog;e.1 theoro' suggcsls thal during boreal foresl posl·fjre success,on. lhc"e"d""e rommunity should respond "';lh (I) increased biomass. prQduc­tion. and rommun;ly ,esp;ralion. and or (2) decreased species di>·e";l)". Inthe general lheories of ecos)'S'em rcoo>'cT)' (adum. 1969: VilOusek andReiners. 1975: Bormann and Likens. 1979). p"ma0' prodUClion is thought 10peak somc time in m,d-.uccc"i"n. Earll' sueee"ional 'tage' may exhibitmOrC rapid energy flux .nd nutrient cycling. smallcr organisms, and grcatermetaboli'm per unil h'''mass. Specie, di"ersit)' mal be 10W"1 carly )nsucce.~ion and greatesl dUrIng mid',uccession Or SOme time before lhe'"hm",,' ,n mOre rcc<:nt "ie"s (Bormann and L,kcn•. 197<;: Botkin and Sobel.1975).

'" Th~ Rol~ of Fi" in ,\'orrhrm Circumpol,l' £(O$J'"''''

Firc u~uall\ rele.se, a put", of .".il.bl. nutrients. ,.n.ned in highnIlrogen and phosphorus eontenlS of plant. during the earl) post-tire years(Bendel!. 197J; !>la,Lean rI al.. Chapter 7. thiS ,-olume). An carl) increase infenihll" and pWrluCli'"j or rdat"'. 8ro"th "1<"$ could red""," s~cies

di'ers;tj of con,umers under some conditions (Riebe",l!. 197J: Abram,k~·.

197R). especially when the ;ncr.a",d produc'ion is cuncent,.'ed ,n on1) onepan of 'he resource 'pc,trum In .ludic, u,ing fertilizer or food supplement'.speCIe, dn'.";'} decre.",d m'•• the short term for mammal' (Abram,kj'.1978: Grant rI al.• 1977). ;n""m (Kirchner. 1971: Hurd and Wolf. 197J). andplants (Mellinger and McNaughton, 1975).

Bendell", (197~j analysis of th~ ~arli~r 'ludies on I""t-fir~ bird andmammal communiti~s led him 10 th~ following condu,ion" Communil;e,displayed conSiderable "ah,lil), ,n a, much a, th~ num~r of indi'iduabchanged linl~_ d~'pil~ marke<l chang~s in ,pecies coml""ition. There weren~t gains in num~r of species, For mammals. tne mean gain waS 'mall and ofdoublful significance. and for birdS. the no" 'pecies were mosrl) gmund· andshrub.forag~...; 80% of the specie< exhib;t~d no change in populal;on 1~\"c1s

Bendell (1974) emphasi2~d the slabi];t)' of populations and sugg"ted lhat'intrinsic" popular ion regulation ,,'as rh~ cau",", _Biomas, and encrg) no" "crcuam,n~d;n onl) on~ study (Bock and l)neh. 1970). in "hieh numbe... ofbirds remained "inuaTl) un<:hanged afreT fire. while biomass and energ) fl""increased_ With the exccplion of Ih,s last reporl, BcndeWs infcr~nccs runcounter to the general bod) of ccologicaltheOf) and experimental re,ulls on,uccession, This led u, 10 Teexamine the a'-ailable dala more sySlema"eall)'.emphas,ztng romparlson of lhe SIXcies divers, I)'. biomass. and cn~rg) flo" ofposl-f;r~ and unhurned eommunil;". To increase lhe dala ba'" I""t-Iogg;ngand olher successional 'tud;es "-ere considered in addilion to the I""l-firedala,

9.2 Mf.THOOS

C~nsuses of small mammal, and birds before and afler fire. logging. and olh~r

dtsturbances are often nor d"C<:lly comparable because of 'he '"fIe') "fCtn,US methods u<cd, Onl) breeding-bird surW)'S made h) lhe territor)mapping melhod hO"e been used in the folio" ing labl~s. "ilh datastandardized to a uniform basis of 100 ha. Small·mammal lrapping studiesusually gn'e only r~lali"e eSlimare,. "hich are n<)l referable to numbers perunit area. Tho<. ,,'ilhin an) gi'-en stud) of small mammals. ral;<>< ofcommuni'y propcnies such as number of indi"idual. or biomass bel"eenpost.fire and ma'ure Slages were calculated, These ratios "'ere lhen used forromparison among sludies, To aid compar;son, successional "egetat,on stages"-ere classified into \Cry carl)" (1-3 )'ea"j. herb (usually 2--8 years). shrub(usuall)' 2~15 )'~a,,). sapling (""mellm.. as earl)' as 5-7 )ears). and

Small-mammal and Bird Cum",,,,,,,,~, lSi

",rondaf\-,ucu"ional forC't as "ell as mature or 'climax' fure't lne-ecategories. based on struclU,e of tbe sueee"Iunat ''''ge•. arc likely 10 be moredlrectl' rolated to prooucti,'Ily and SUucture uf "oimal eommuni,ic' tb,1n I>

'bc numbor of year> aftcr di\lurbanee.IlK.ma\S and consumIng hiomalS (Salt. 1957) arc estimaled b~ mult'pl,'ng

tbe index of abundance b}' body "eigbt (IV) or by all"". rc'pec""cly. wheroaly" " an empmeal relation bet"een bod}' "'eight and ba",1 metaholtsmCon,uming bioma.. e~uals 70\\·'"' for 'mall mammals (Klciber. 1%1) and1191\"-" and 7H,5\\" ,J for p",scrtne and nanpa"'rme bllds rC'Spectl>cl~

(LaStew,k, and D"wson. 1%7; Zar. 1%8). Follo"ing Salt (1957). consumingbioma,s of passcrine> "as actuall~' calculaled as 11"' ". (amlnlng lbccoo'lam).•nd far nonpa'serine> the body "cight used ""' no, 'heir O"n. b,,'a I<elgb, (11'.) "bleb. "ben used in the p..",rlne equalion 1291\" '''. ga'e ab.,al metabolic rate equal 10 that e,peeled for the nonpa""ioe: 11'...(0,602811-"-")1","', Also. 'be COm'am (0.602B) in thi' laner equa""n 1<",adJuSled for use ,,"b welghl In gramme' mstead of kilogrammes. Thusb",m"" and con,umlng blomas< "ere eakulaled In grammes 100 ba and lhenonpa»"ine cons"mlng bioma« "as adja"ed 10 be metabulically equi' 'lent10 lhe pas",,,ne data, Basal metabolism " onl~' one componenl 01 ani'"metaboli,m or of the animal's actual energy budget In the wild The I""'erlaw of b"al mctabulism was used '0 weight different anlmar. cont"bu"nn, 10

rommunit~ metaholism. "'thout Impl"ing that It doe> th" accuralely or lhat IIestimates ael"al commUnll) metabolism.

Body "eights "cre "",mbled fram Banfield (1974) far 'mall mammal' .ndfrom man~ different """fCeS for bud,. SpecIes d"e,,'"'' were e,,!cul;lted ;1>

(::'p;)_l ("here p' is the proouC1"n~ of the Ith specie, (Hurlberl. 1971:M.),. 1975). Speci", richness equal, the number of 'pee;'" rcpre",nle<l byrelall"e den'Hy e,Umales, The rotio of >pccle, d"et'l'Y- 10 'peei", "chnes>"as med a, an ,"de~ of e,enn.ss, Wi,b "II Sp'"I''' e4""II) rcp"<entcd.dl"c"it} e4u.ls rlehnes•. its maximum possible '''lue The sIX'cies <li'e"",Inde~ can IherefOr< be conSidered as 'he numller of e4uall) rcpr"<cntedspecie, "hleh would g;\C lhe same ::'p;. Speel" d"e"," in each c"", "",calculated lrom eon,umin~ biom,,, rather th,," oumber'. bee""'" 'hi' shouldbetter approximale lhe relati'e utilization of niche 'pace Or r"<l"tcC'

9.3 S.\I,\1.1. MA,\L\I,\I};

9.3.1 GCOlraphicall)istribution ofStudic<

Almost .lIbure.1 studies of 'mall mammals ,wsu, fire or logging com,' fromlhe '.r) >outhern boreal faresl. "ith no published da'a from the apenspruce-lichen woodland (I'csotundra. Hudson..n life zone). Data are a"ail·

""abl~ from jac~ pin~ forest in Minne"'I" (JS'N Latitude) (Ahlgren. 1966:KreIling and Ahlgren. 197~) and ",uthu,lern Mani10ba (5lfN) (Simm, andBuckner. 1973); ,pmcc_lir fore,I' ,n norlh"c'(Nn Maine (~8"N) (Richen'_lY74). <outhern Ontario (41)"N) (Simms. 1979). Minne"'t. (#N) (Krcftingand Ahlgren. 1974). and 1'0'.,," Brun,,,,id and the Gaspe Pcnin,,,I.(~N) (I>\o"i" 1955): and black <pruce fore,ts \0 Ontario (49"1') (Martelland Rad,-an) •. 19TIl. For inte'ior Ala'h (6Y1\ 1there is unpubli<hed work inblad' spruce fore<1 burn' (We'!. 197~. 19791. In lundra. on the Macken/icRiwf Delta near Inu,-j\;. Wern (1975) Ir~ppcd 'mall mammal< in. burn fornne \car. and;n the SOUlhern Yuk"n (fltJ-{,]<N) K",hs and \\;og.[< {1\176)compared rodenl communi1ies in man) "egetatio~ type'.

9.3.2 AI:l(J"~t. Communit~ Pro!"'.t;..

The"<C ,tudie' off.r nIne earl,.,uccc"ional (1-7 'car<. herb or hcrlH.hrub".gc) ",catment plot' lor compari",n .. ith mature boreal 10re<1 Table \1,1,ho.. , that ,pecic, richne.. and ,pecic, di\C"il, rna, either i~erc,,,e ordeerea'" o"cr that In mature lore't. but that number<. hloma". andcon'umln! bioma» In e"rl~' '''=»ional Slages a,e inereased in m",tin'tances, Later ,,,cce,,i"n.1 st.g.. domInated by 'hrul>< Or tree saplings rna,..hibil I"wee numher<. b,oma". and con'umlng bioma" lhan malu,e forest'lage,. but there "ere no ,ignifieant difference' The ratios 01 rommunit,pro!"'nies from succe><ional 10 mature 'tage' are 'h"wn in Table 11.2.Pmpertic", of thc 1M' herh "ages combIned "ere ,ignl(;"ntl,' different {P <t1.0251 Imm those of tbe 1"0 wood, stage< tal.en together. for numher<.biom,,,,. and <lmsumLng bi"ma'>S. bUl n<l1 fo' speeics di,efSll". SpecIe,"ehne" usuall' did not change ,'cr; much. "ith the same "-'t of speCIe, beIngpfCscnt in most su""""ional stage>. but rcla1l'O abundanee' and idemit, of

Tabl< 9, [ ""mtoer of ,lud,c> '" treotment plot> In "h"'h propertlC' of'mall·mammal comm""'''e, eitbef Increase<! or dtcrea""d from e.,ll,"',,:essiona[ (h<:rb or berb.dHub stag<1 10 m>luI< fore"

•""""

,,,,,,,,

Ch.ngc in eommnoi') plope")'SismfLO""""[""f.''''' De<:rea>e \;0 change [e,.I·

"S'

"11,0.'

"00.'

("ommun,,'

SpeCIe' nchne"Specic> di,'.",';""rnl>o"Biorn.»("on,nrnmS biorna«

"S<s.,f",ancc " '",,<d ~ "h ,I>< bonum'" "i.",~,,,,,, on ,I>< "r«'0l... <>I <q.ol,h.""" "f ;oc" .....oJ ~<""'~S· "'" "Inif"".,

S",all-",a",,,,al mId Bird CO"'''''''''li~, 15'1

Table 9,2 ~lean f.ti", of ,mall_mammal commumt,. pfopertie' from".,iou, ,u""»looal slage, to m.tuTe "and>

Communil,. pfO!","" Velyearlyhelb Herb Shrllb Sapling

I'Umbef of "udie. • .' " ."Specie< di"el<lly ,~ I. ~7 1.71 O.MNumber> US ,~

"~ WMean body ,ize 1.21 ,~ 0.87 0.95Con,uming biom." ,~ ,. 0.8! O.W

-Ahl*"" 11%1>. reor 1. ._11. Kreft,", ,nd AhI~,," 1I111~. K'dr e,.... re." I,nd 2.•_11. ~"".II 'nd Rod"n!; (t9"J, ._l). Sim, 'nd Il"c~ne, 1197.1. .-1)'Roc""", (I91~.•_1), Kref"", "d AhI~,," (11I1~. Koel! Creek. ,e." 1-1,.-1). Simm. (1'11'9.•_1), A"I~e" 11%6. yeo> J.• -1),'Krefti.i ",4 Ahll!". {191<, ._1, H"rt L"e l"''' 3-9. K«ly ere,. y...,l>--ll),'Mom' (1951. "_~, plol> 1.1.),11 """, 1>.1.~, Il. ,,'pc",,'e"'1

the dominant ,peeies changed with lime, Larger herbi"orou'- mammal' 'uehas tree Stluierel,. hare,. and rabbil' were nOl censu,ed ,n the.. report,. Theseom",ioo, "ill have lil110 effecl on lhe specie, di,-ersi" bUI could greml)' allercon,uming bioma,s and hioma..

Mean My weighl ("hich follo'" mean MY' ,ize) lend, to he Ie" onhumod or loggod ,ile. (P < 0.05). reflecting a shill 10 dominance b)Perom}'.,,,, manin'/aw". Ho"e"er. mean wcight i' somelimes incTeased h~

gaIn, in chipmunk (Tom,os or E,(/o",io,) abundance (Ahlgren, 1966: Kreftingand Ahlgren. 1974: Mancil and Radvanyi. 1977; Black and Hoo'-en. 1974;Ga,h,,'iler. 1970). Thu'. in earh 'uoce"ion thero is oflon a bimodal sIzedi,1Tihulion for .mall mammal•. wilh peak. at the 1"'0 size extrome. (deermice and chipmunks)_ Curiousl). lhe same phenomenon occurred inPiedmont old field ,ucce"ion of bird COmmUnilie' (JohnSlOn and Odum.1956: 1>kNaughton and Wolf. 1973. p, 377). and it can he postulated Ihat lhi.refleels lho seed size dimihulion. a bimodalit\ of seod don.ilie,. or 1"'0ahernati'-e S1Ialogie. of exploiling seed' rebled to pmchines, of lhe resource.ChIpmunk, mal depend upon palche. of l",e seed pcoduced in quanlit~ hytree'S ,u,,'i"ing fire Or logging. and may al'" rO'pond positi'oly 10 palche. ofshmb co\'er. logs. or olher a.peel' of habilat Slrueluro

Through ,ucce"ion. lhere is a trend for smaller deer mICe 10 be replaced b~

larger \'ole' (C1c,hriollomy.) and forost-dwolling chipmunks (Kreiling andAhlgron. 1974: G"hwiler: 1970), Thi. lead' to an ineroa,e in My .ize and adecrease in energ,. consumed pcr gramme of bIomass as judged from basalmetahali,m Thus lhere i' an Increase in Iho efficieney of energy ulili.ation.ahhough mOle dalu on Ihe aClual energ) bUdgets of lhese small mammalscould aher thiS conclusion

9.3.3 Spoor;'" Mque""" and Natural Hi~lorl

Small mammals in ~arl~' ,ucressional 'tag~' or po't-f"o reeo"en arcdominat~d by se~d-eate" (P~romyscus, Tam,a,. £Ulamias), Grani",re, ma~'

eontinue to dominate until fore't de,'elops. Or mal be replaced b) fol,o",re,(Microrw.) if a eo,'er rich In grasses and sedges de,'elops. Later. a, the fore,tde"elops. ClClh.ionomys. ha"ng a more omnivorou, di~t of foliage. seed"rOOl<. and b.ark. replace, the earl) sucressional mice or 'ole' (Gashwiler.1970, Kreftir.g and Ahlgren. 1~74. Ahlgren. 1966: Martell and Rad\Jnyi.19n, Sim, and Buckn~r. 1973; Riehen,. 1974, Morris. 1955)_ Fore't·dwelhngchipmunk, mal' also return. posslbl) because 'here arc seed· rich patches toe.ploit. ther. are larger seed~ in fore,t,. or because of habitat ,Iructure (e, g,.the mi>!ure of open and "egetated patche, d,,-eloping in mid-succession),

Puom}wu. manic"lalus is a uni"ornlll'onh American in,ader of loned.burned. and olher di'lurbed area, ueept in Ala,ka and the taIga of Canada.This mouse can be "er) abundant one monlh after a fire (Te,i,. 19563,Te<lcr. 1965). and in such ca""" pre,umabl) forage, for the seed' once storedin the fore't floor ",hieh are exposed b)' fire_ The carll' dominance of deermice and ['Crhaps SOme ,hrc"'~ prob.abl) depend' upon produClion b) c.d~'

~u="ional pl.nt,. "'hlCh propagate more b) s«d than '-egetati'e reproduc­tion (Ahlgren. 1966: Krefting and ,\hlgren. 1974), Allhough man) studi,..~how greater numbers of deer mice in burned and loned area, than In

forem. Sulli"an (1979) ~uue'ted Ihal 'hi' m.; be • seasonall) transiente«eo. and thaI cut, and burns ma, be beha,-ioral 'sink~' for ammal,di,pcrsing from forest,: ho"'e,"". the ~,idence i~ amhiguou,.

Deer mice seem unique in tolerating or ""Iecting habitat' "ith little rowr.and also ['Crsisting in herb and shrub stage' thaI arc nol grass dominated(Krefling and Ahlg..n. 1974; Gash",'i1". 1970; Ahlgren. 1966: T~"s. 1956b)In burned areas or old field' ",'here grasse, or ""dges dominate. microtine,.especially Microtus pmlU}'il'anicus. are relati"ely more abundant thanP~rom}'scus (Ahlgren. 1966; Slmm~. 1979, Coo•. 1959, Pearson. 1'159,Beck"'ith. 1954). Vegetation co"er and den,it)" may be more important tomicrotines than the presence of gr~sses (LaBue and Darnell. 195'1: Birnc)' ~I

al.. 1976).There was no dea' trend in shrew ,peeie, eompo<;ition or abundance

relati,," 10 other small mammal,. in response '0 fire and logging. Shrew,requ,,~ high humidit) near tbe ground surface (Getz. 1961c). and thelo"-humiditj microclimate of poorl)" '-eget.ted earl~ ,ucre,sional habitat,(Bendell. 1974) uplains the low post·fire densit) of shrew,.

9.3.4 Small Mammals in tM Far North

Ver)" little ha~ been publ;~hffl on po<;,.dl,turhanec response of sm.llmammals in taiga or tundra. Wein (1975) found f~wer small mammals on

Sm~II'",ammal ~nd BiTd CQm""lIli"~s '"burned lhan on unburned ~hmb lundra of Ihe Machnzie Ri,w Delt",dominance b>' CMthrioMmys TulillS "'as reducc<l and abundane<: of ,\/penlUyl,'anicus "'a, increased ,ligbtly In "urn~. I>lid·I .."ude 10 boreal 'tudie'(Table 9.2) showed thai "'creases \n numbel"< generall> ocrur early in\uCCess,on. !l" nol dca' ho" these tundra dala can be accounted fOT. bUll""producti"il}' or low w"er may be implicated. Reco"e,) of vegelation mal berelati'e1}' slow compared with boreal Silt>.

The fo,e" 'ole (Cleth,ionom,'s gappen). common lhroughoUlthe borealforest. i' replae<:d in the Hudsonian hfez<:>ne of "e"ern Canada and ,n AI.,kab; C. T",il"•. C "'til"" " found In boln forested and unfore'led habital>(Banfield. 19(4). West ( 1979) ,oo"ed lhal the habitat separation between ('.gapperi. limited to foresls. and .IliaOl'" spp, of more open hab,l'" (Granl.1972: Cameron. 19(,4: Morri~. 1%9) docs nm hold fOT C. rUlil"•. "hichcolonIZed a burned bjaek 'prue<: foro" in inleTior Ala'ka a, the dominant"ole. accompanied hy three ,lliclOtus spp

The dur mOme i, rare or a""'nt ,n Inc Far North and therofore b notpresent as a dominant colonizer on burn' (Banfield. 1974: Manell andPearson. 1978: Wein. 197$) in tundra. in for"Hundra. or ,n Ala<bn fo,o",It doc•. ho"c\Cr. appear as far as 61<1" a, a dominant ,pecies: Krebs andWingate (1976) reported deer mie<: and C. Tlllil,,, dominant in ,'inuall)' allhabiMs m (he Kluane tegion of lhe southern Yukon (6G-(iloN lalilude)Mirro"" pe"IU}'lm"irus ,,", foremO>I in ma"h and mcado". "hile IIoecOIlOmus "as dominant in tundra, Vegetation in thi' region i' a fire mosaicbUI no Sland ages Or suceess,onal sc4uence ,,'ere a'i\lgned to Ihe", "~ud} plol'b; K,e1" and Wingate (1976). Ne'erthcless a ,uc,e','onal inlCtprClation ispos,ible,

On t,..o sile'. designated 'open ,pruce,"'ill"" . and '" illo,..· . p,obabl, more"'e<:nl burn•. deer m'e<:. and C. rotil~s ~re co·dominant. thu•. C. rolilu.ptobabl)' in"ade' bum' her. as il doc' in interior AI",ka. There" no ob,iou,relation,hip bet""ccn Krcl>< and Wingale', d~"ifical'onof hab'la" as 'open'or 'closed' and rel~ti"e abundance of deer mice "ersu, C r~lIlus, I lo,..e>or. itcan be inferred thal deer mice dom,nance occurs in ,or; open habitat> thatare earl}' in primaf)' .u="ion (e.g. a grass'fi,e"eod area. a beach ,idge. anda Drya, d"''''nwndi commUnil}') or in habital' "here lhere is a greaterproduct,on of seed, and ""rrie. (e.g, a'pen. babam poplar. and ShephtTdlaca"adrn"i,·dominaled forest»,

9.3.5 Elf"''' ul Logging hrsus Fi/"t'S

Some studi.. of post·fire .mall-mammal populations in"oh'cd posl.loggingburning of sl"h and duff (Gash,..ile,. 1970). "hile Olhe" ,n"ol"ed for"l fire'in unloggcd foresl (Kreiling and Ahlgren. l'lU), Resulting differences inhabital 'lruclure such ~s fallen limbs. standing dead limber. etc .. rould ha>e

162 Th. R"" al Fir< i" SaTlh,,,, Circampolar Em"""''''

"nporlant effec" upon UllliZ<llion h\ mammals (Da\i>. 1977) and hird.(Hagar 1960), For example. cnipmunk- use logs Or downcd limb, as runwaysand lookout', There appears to h.\'e becn lillie COrrdall\e stud\ of eff~l'ofdiffering tn"" of di,'uroonce,

Ahlgren (1966) compared .mall mammal> On Minnesola jack pIne loggedare3< On "hi,h 'lash" as cilher hurned or lell unburne,!. Srecies compositionOn the logged area, resembled that On the control (unloUed) area. thoughpopulaHon! "cre higher. and ,n the third lear M. ~nnsJ/"Q"ic". Increased.probably bec.use of more ground I.yer CO\'cr in the loUed .rea Or .nups"ing in Ibe \'ole', populalion cycle. On lhc hurned arca. decr mICedominated In thc fif>tt"o \'ca,-;;. and C. /(ap~ri In thc th"d ycar, ChIpmunk,"ere m",t ahundant On thc 10Ued are. in ....rs I and 2. but On the burn in\e.r ." thi, change is n01 re.dily expl.ined. but i' rerhaps a result of thedC\elopment of .ufficienl CO\'cr and~ resourtts in the hurn,

Martell and Rad\3n\; (1977) comp.red ,m.1I mammal';n logged .rea, inOnt.r;o black ,pruce for"ts and found lhal s<:arificd areaS lended to

resemble burn. more than un>c.rificd are... Voles (M, ~"tl$.AQ",rw)

,nereased Qn bulh cUls. ,,'hile deer mice increa>ed On the scarified are. andchipmunks (E. "''''imus) incre.",d On the un",ar;fied area. Thi' r"ult seem,analQgou' 10 lh., of Ahlgren (1966). suggeStmg 'hat dcer mle<: ,ncrcase ,nmme oren habitats or "here seeds Qf colonizing plants are .bundanl. "hilechipmunks mal resl'clnd to h.bitat fcatures Ii.e sla'h and logs. Or a m"turc Qfopen and brushy arcas,

9.3.6 ~'ulu ... Sludi""

Imp'''' cmenlS could t>.. m.de in future studies Qf 'mall mammal' b\ relatingpopul"tiQn differences lQ fU<>d rc<;OurttS and habnat ,"uCture, PaS! studic~

ha\e failed to rephe." e'rerimcnt.1 hums .nd other habilat modifi,atiQns,Also expe,imental habitat .1,cMion. b}' herbicide apphealiQn (Black andHoo'en. 1974). slash remo"al ami addiuon. Or sowing of herbs and g....sses."ould be panieularl\ helpful in formul.tlng and testing hl'pothc..' <.>n therel.tion of ,m.lI-mamm.1 .bundance to habilat ,truClute. Because the'ariabilit}'. amQng studies. in <mall·mammal sequence and liming is perhapsrel.ted to ,'arl.t;"ns in ,eget.ti\,c ,ucre"iQn. our predie",'e abilit) dependsupon kno"'ing the qu.mitati'-. relalions bet"een the ,'egetatiQn and the'nimal' in\'Qh'.d, Soed production. ,ize,. and patchiness are fai'I}' cas}' tQ'tudy but ha\C becn neglcelCd ,n lhc "olks re",e,,'cd, Habilat and \ cgetation,,,ucture ha\'e ree<:ntl\ t>..en ;nlen,i'-el)' studied in relation lQ ,mall·mamm.1habit.t dislribuliQn (Grant. 1972, Richen,. 1974, M'Closkcy. 1975:M'CI,,,.e) and Field"ick. 1975: Birne\ n al.. 1976: Miller and Getz, 1977:Morri,. 1979). Applicalion of similar techniques will horefull} become morecOmmon in ,ucre!Slonal and post.disturooncc Studies, For small forest

'""",mmal•. nocM >q>aratlOll b~' habl.a, 0. mw:rohabitat. rather than food~kc"o" P" S<'. appea~ to ~ the rule (Eadle. 1953. B.anf"'ld. 1lJ'i4,J:otrlC"OII. 1949: Gru. 196Ia.b,cl_ MIlkr and G<:u (1977) r""nd 1....1 1MnumM. of ..oodl plant opccies "'"2'1 ~IaIN ""th unaIl rrwnmal~t,b,crs"~~ ~~ed th.at food <b"e""l~ "''». mtial factor: bcr.,c-'t<. lboocorrelalion found "'2< no! i""""""'ent ...,th mteroIutbotat "'pan'''''' M lhepnn,,,,atc QU,,", of both >peaeS d'H"'l) and ~.,.ed diet. (M"CIo$kel.1975 . .\otorns. 1m).

'.-t BIRD CO.\.I.\.IL~ITlr..s

9.4.1 GOOllraphkall);'lribut;on of Studies

No published sludics of post·fir. lmd «Immunities in the spruce-fir or theoorcal for.'1 "ere found. and th. follo",ng rcm~f~s ~rta;n 10 dala frombreeding-bird «nSu",. onl), A '-cr) lhOTOUgh stud) Irom earl) post-fir. tomalur. forCSI .tagts ""as r~ntll carried (>Ill (In Alaska's Kena; PeniQ.ula at6lJ....61<N (Ou,nlan. 1979).

Sescral >ludlC$ <k>cribc bird rommuntlOCS 1-10 )urs after locg,ng. but onl~on~ fall> ,n lh~ Canadian lif~ zone Al'tknoo ~ •. (1917) ..udocd blrdromunllle'. In P"""'~,·I",., corrilI<ln In the oat-lllc"~ region (36"1 InT~nl'lC'SS«. ]';'dlO'~ (1971) ~b«Ithc dfccu of 1Ulde1"Sl0t~~ '~""""-al InCalifornia Sequoia f<.>r'$ (36"3.):"1. Haga' (1960) eumiDffllogcd are~ InDoupa;..fir fOtC1olS of the northern Caltlonna CO&SI Range (41 .... ). FBlW'CbatJd Ohman (1978) ..1Khed 1oggI", effectS In IlIUcG rorufer (Doup·fi,.ponikrosa pmc) lOfnl In AfIlOlU (33'1'). Webb rr •. (1977) ..udoednorthern ...."""''00<1 fornl'l at four Ie'els 01 cull,ng '"'~""t~ in lIOe,," Yorlt"sAd'rondack \Ioun'a"" (-U '). atJd Knsk' (1979) ..udiW bird popuJallomin recent clca, cuts and through a suc:<:eS$Kmal SCM in th~ .....rillme COtllfe,fmcsu of SOUtMll5lem Alaska ($6'1'1)

Man} stud,C$ ha,'~ dc",,;!>ed bird rommUnll'es du"ng plant SUC«"lOfI Ot;n ..egelallon types "'hich can be placed ,n a succcs,,;onaJ scquence: t""o he Inthe Hud'(l",an hfe ZOne: Thebnge (1976). in Kluane I"at;onal Park. Yukon(6O--<>1'N). and Splndler and K~,,~I (1980). In the upper Tanana Valle}.interior Alaska (63'N). Although th;! rev;e ... is hm,t~d 10 Nonh America. anoutstanding and romptehen,ne stud}' f,om Finland ",-ill al§() be u",d,Haapanen (1%5). 6lJ-6.l°N. AtIOlher benchmnk study of i),td-rommunl1}su~'on .Ilould be recall¢d. Lack's stlKi) of the Breckland .,·ifauna In

tdation to affotCOlallon (Lack. 19.13. 1939: Lxk and Uck. 1951). There ,.al'" an nallent re" ...... of bt~nl..b..d «"nsu'!C'l in mature ronifer fOt'C'St.througlooolt "onh Amenca (W>em. 1975), Thru sturhcs from the Canad'an

'"life zone arc included her., Stcwart and Aldrich (1~9, 1952). from theroount~ill! of cenlr~l Wl!'Sl Virginia (2&<N) ~nd extreme northern ~hme

(~rN). re,pc:C1i'.I~. and Sail (1957) ftom Grand Telon t'allonal Park inWyoming (43'Nl· Thre. studies from lh. ITaosition life zone. norlhernhardwood for.st. arc Martin (1%0) from Algonquin Park. Ontario(4S-46'N). Kendeigh (19./6) from nonheastem New York (4Z·30'N). and"endeign (1948) from northern 10".' MIChigan (46'1"). 01her sluJieseOnj;idered range from 34" to 41·1' latilude. ,n Ih. oak-hickor~ "oll,beech-maple regions of the eastern United Slales (Aldrich. 1943. Shug.nand James. 1973: Brewer. 1951\; Kor. \%1;), The last tWO concernedsucce,sion On mip m,neS.

9.4.2 AggrtgaleCommunilj Properl~

Table 9.3 ,how, succession,lt,.od, in bird-community properties. a\'eragedOver ,,-,'·cral s,ud,es, S,gnifieant trends exi!t for !ped., richne's and di' .rsi')'(P < 0.05 and P < 0.005). al alrcad)' sho",n for s,milar ,·ege'atlon eategoricsby Tramer (1969)... ho analysed hundred' of cen,usc, 'aken from Am",c""Birds and Auduoo"'. Field ,"mrs. Mean ..eigh' and consumin~ biomass p<"runi' "'eig)" also exh'bit si~nifieant 'rend, (P < O.(J]). The la,e-w,cessionalinereasc in species richncss rclati'-e to species di\'ersity has !><-en noted anddi><:usud in man) pre\'ious studies.

Numl>ers and consummg biomass of birdS are 'cr,lo.. carll in sucrcs,ion(I;~nifi,ant for numl>ers onl). at P < 0.05). bUl,nerea"" rap'dl) and reach ama,imum b)' the sapling stage, This parallels trend' in plan, production andfohage hi<)mass. whICh rea,h and m"intain ,heir peak wcll beforc sucre"ion i,completed and bcfore biomass stora~e peak, (O"ngton. 1%2. Bra) andGorham. 1%-1). One m,ght then uplain trends in numbers and biomass",respan""s '0 prima~ producti,·,')' ... hilt 'he contlnucJ ,nCreas<: in sptti",riehnc" and di'c"it) could be cxplained b),hc cominuing 'ertleal red",,,_bution of fohage bioma... and perhaps b) an ,nerease in hori~onlal spa,ialheterogenei,y in 'he ma,urc forcst (Shafi and Yarranlon. 1973, Bormann andLiken,. 1979). The rela,ion,hip bet ..'..n bird ,pecies diversi,) and foliagehe,ght dtSlrtbu,ion has been well e!1ablished (MacAnhur. 1%-1: Willson.1974: Kar. 1\168). Spindler and Ke"eI (19&:1) corrcla'ed bird numbers "i'hprimar} product;'it) di'ided by canol') ,olume. "h'Ch mal cominue toincrease ahcr primary' production has peaked,

In contra,t to 'he summa!)' in Table 9.3. dat. for indi'idual studies ~i"e 'heimpression tha' there mal bc significant trends ,n numbe" and b,omass "ilhsU=5\ion. Spindlcr and Kessel (1980) stud,ed replica,ed samples of blackand .. hilt spruce stands (7 eaCh). deCiduous aspen Or birch woods (5). m,xeddeciduou.....oniferous stands (~). and shrub "'p<"' (7), The)' found a decreasefrom ,hrub 10 deCiduous '0 conifer sl.nds in numbers (486. 298. 237 pairs 100

Small·mammal a"d HJTd C()m""",i"~. '"T.ble 9.3 Il"d communtty I"operl;c. for differen' ,"""e,,,on.1 ,1>,..

S"""",,,,,n.1 "'j!"

Commum'l properl~' V.l) SerondiI'} M.'ure•.,1, Iterb ~.. Sal'hn~ t",.", lore", '" 1l!17 17.116 17 1~ 17,90 ~ l-ki

Spe.,e; ".hne" W 200 ~~I 11.03 no 7.48 10.17, ," " " " ", ,~ .,77 ,m ,m 926 9.~:

Species d,,'e""j " OM U ,~ 276 "" 29.., , • " " " ":>"mbcr of!,,'" per

, m ~ "" $10 ,~ m

" 137 "' "' .. '",~

lOOha , , • • '" ",Con'uminj! biorn."

, 2117 ..~SO 6132 6756 ~118 >-l:l6

" :.53~" "" 1321 ~, ~,

(.m'IOO ....l , , , • , • '"K.I.,,,. m...bo". , 28.2 330 "0 36,~ 37.1 36~

demand' • " '" 1,2 -,2 $,6 "'", , • , • '", 1. 50 12,1. 1035 1021 9.0S o,5~

,\1o.n bod~ ,if<' • ,,-"~ 2 78 22.. UO H2

", , , , • •

"Stud;<, tnN '0 rom",1< ..bk ;""ook<! K.~_h 11......). SI<~.n and ...klt.._ 11Q.>9, l';<2). Soli(laI71· lb."".." (1'llI.'l1. 1(.. (1111«), ";eo, (I'I7S' Thebe,~< 119"'6\. Spondlet ,nd K",,'I(1'lllV1

'C.i<ul",d., """urn,n, l>oom'''I'''' OM o<,~~"

'(aleul'IN" """,m,n, """""', ~ "m",,'

ha) and ,n consum,ng bl(lmar.s (liS(,. 699. 555~m HI) hal. Nuh of "h,ch areSIgnificant (P < 0,01): "end' of indi'idual 'ludic' are diseu,...d bdo" and,ho"n ,n Table \/.5. In lhe m"ed s'and,. ,herc "ere 331 pairs 100 h. and 1Mgm 100 h•. ,ligh,ly higher Ihan for deciduous "and, Tramer (11'6\1) aboreponed specie, richness and di'e"i') '0 be ,Iigh,h greater ,n mix«J forc',,-

Species di...rsit),. numbers. and ronsummg biom." can be dra"icall;rcduced in mid-->tIC't:~"ion b; th. d.'·elOflme~t of a unrfQfm. dcn'" Stand of'Il'eS ha"ng hule undetS1or~). Haapanen 096,S) re'.aled this for Ihe p,neth'cket §lage of a p,ne fOIl"1 sere,n Finland. and 'he '.me effect appe." inposl·fire succe"ion on AI.ska', Kenai Penin,ula (willo,,·btrch-whi,c spruce'u<Xession) (Quinlan. 19"79), In the Rock\ Moonta,~,. <pecics di,cl"u;'.numbers. and consuming biomar.s of birds "cre eXlfemcl, 10" in a homog­cn~ou,. thickly stocked lodgepol. p'~c stand. a t;'p,cal post·fire t)'pe. andthe.. quan'i'ics mcrcased in lodgepole---spru<x and ma'ure spruce fore<lS(Salt. 1957). Howe,er. loung post·fi,e aspen "and, wi'h a prolific undc,·

'''' The Role of f"irr In SOr/hull Circumpolar Eros,.."",.

'lore)' held ~ much larger and mOre Sj'Ccies·nch bird communl1)' {Salt. 1957).Spl'C1e~ dil'l'f<;lt~. num~rs. and blOmas~ ha\e $Ornctime§ dec~aj.Cd in thesapling 01 secondar; for.'t stag'" in other seres (e.g .. Johnslon and Odum.1956: Shugan and hrne'S. 1973: Theberge. 1976). sOl"'rimposed upon ageneral Incrcas,ng trend. These oornmun"l "bollienech" are probably" mOrelike" 10 de,-dop in ronifer slands .fler fire and coning: oon,[c,.., >Cern p,oneto establi'h a thick 'land and to exclude 'hrub> and herb> b)' root and lightcompetition. especiall; ;n the boreal forcsl where the common deciduo'"lree, hJW high light rC'luiremcnK Since tit"." no pronounced decrea,e inproduction al the ero" n-cl""ure Slag. of fore.! plamation growth (0\ ington.1%2: flra) and Gorham. 1%-1). ,hc hi'd_poPula'ioo ool1koeeL i, hkcl~ to bea re,ult of" reduction in foliage height di"e"i". undemore) co'er (Hooper~I al.. 1973. Flack. 1976, Curtis and Rlplc). 1975). and related changc, in a,a"e'Y of re""uree,. Indudmg n"lIng ,"es.

Con,rar) '0 the early. proml,ing tc,ults of Salt (1957). thcre appears 10 beno uniform trcnd ,o"ard increasing bioenergetic cfficicnc~. "hich ,hould"p!"'ar a, a dccre'''e in relati,·c mc'aoolic demand. wi'h ,uccc"ion. Rela,i"cmetaoolic demand imtead appears to increa,. "ith SU"-'CS>lon (I' < O.Ul.Table 9.3). This i, probabl) relatc<l'o the largcr bird ,izC'i in ea.l) s,age, of'U"-'C",on. and an increase in ,,·..blers in ma,ure forcS!>: mean bod) ,iledeerca"" through succe"ion (Table 9.3, P <: ,WI). Larger bird, ",II havelower ba,al metaoolic rate> pcr u"i, biomas>. '" lo"er rela,i,-c metabolicdcmand "'ould be expected for them Thcre is a dangcr. howner. ,hal trends"i'hin indi' idual >tudic-; "ill differ from thc a'eraged data of Table 9.3: mcanIlod) ,i,e wa' lo"er in lhc ma'ure 'tage a, compared with ,he ncx! carlieronein ,ix ou, of nine studi«. but in most Cascs the difference wa, small and nOl,ignificant.

",onp,.,scrinc' orc bener represcnted in more open babit.". and arc al>o,he larger bltd' of a WmmUnl1) (faaborg. 1977). for fif'een studie. (Table9,~(a)). 'h'" proportion of ,pecic, riehne..., accoun'cd for h) nonpasserine,,,", con.iSlentl) grea,er in early .uccc<>ional (berb and ,hruh) ,'ago> than inrnaturc 'lag« (fourtcen Out of .i"cen Ca'"""S. P - 4),002), Openne". orhabi'a' structure. i, clearl) more important than ,u"-'o>'ional age. "on­pas,.';"e, a1s.o con>titule a greatcr frac'ion of sp.:cie, in ,undra (e.g,Theberge. 1976) and ,n 'Cf' old ,hruh and ma"h nani,"" in thc <ubare,ic(Spindler and Ko".J. 1900), The trend is not mono'one decreasing. ho"c,'cr:lhc nonra".r;ne fra,·'ion appea" to reach a miOlmum ..,mc time in 'he ,hrubor >ocondar)-fore>! Slage' (Theberge 1976, SpIndle, and Ke,""'1. 19l1O, Sah.1957, Kat. l%ll: Johnston and O<lum, 1956: Shugart and Jame,. 1973:Bcck""h. 195-1. and Table 9A(a)),

For ,mall mammal~. grani"".. domlna'c the earl}' .u"-'e'\Sional andpos,-fire >lage,. Sced-ea,ing bi,,!> reach 'hei' mUlrnurn specie~ richness.numbers. and consuming biomas> in 'he herb and shrub >lage._ M'",

Small·mammal and Hird Comm"""". '"Tobie 9,~ (.) Pe,cen'age of the specie' ncb..... conl,ibuted by nonpass.erin... and(b) pe,,:.n'a~e of 'he ro",urnin~biorn,,, ",,"'rib"'ed bJ' ·~r.ni,w..· (spano"'•. qu.;!.60",)

Su"".,,,,,,,.1 ".geP"'Hhm..

\e~' seronda~ ~1a,ure open t>lacle'TI~' lte,b Shrub s..1>hn~ fo,." Ime" ~-

(., , 33% '" 16',,' "y~.

< " "~. " 3~.

• " " " "'" • "" ~, 3~'" "" n'l" l~'" '"• '" " " " ~~. , '"• , • " '" '" '"

,'$opl,"~ • .o "'''<''''''r~ r"",,1 ".~, rom.,..d

s<l'<I-eaters in Ihe'" cases a,e f tingillids. although I/IlwO case< in the sou,hemUnited States quail dominale, All ftingilhds ....p' the chipping Spart(WIWete ,ounled a, hreeding_season g,anivo,e,. "hi,h "a~ a ne.e.,a,y expedientbut almoSl ~u'c1y incorreCl, Thu~. the operational definilion for 'grani\ore"~h"uld ~ ta'en a, ·s!"'rrow<. quai!. .nd do\·e', In ten 'lUdle~ (T.bk 9 A{b}).grani\ore~ a\erage<:! 35% 01 the ,on,uming bioma", In the <hrub ~tage \cr~u\

1J% in mature habital, (Theberge. 1976; Spindler and Kessel. 1980: SIC,,'arl.nd Ald'ich. 1949. 1952: Salt. 1957: Johnst"n .nd Odum. 1956: Shugon andJam.,.. 1973: Kar. 1968: 8<><:. and Lynch. 1970: Hagar. 1960). The trend 01decrease in consuming biomas~ in Table 9.J(b} i' significam a, P < 0,01(..eluding blac' ~pruce lorell,) Even though the tOlal ,on~uml/lg biomas,in"eased (Table 9.3). "hich could 10"'et the percentage 01 granimres hadIheir biom• ., ~tal'ed constant. granivore consuming biomass decreasedIhreelold (955 gO' versus 31~ gO' per 100 h•. with <tand.rd errOrS 12J and 611./1.17 and 11).Su~sional and habilat relalions 01 indi"idual spcr;., are '00 numerouS

and generally too wen 'n(WI'n 10 meri, dis<:ussion here, Successional andposl·di"urbance chang.. in feedIng and nes,ing guilds ha" been adequatel~

discussed by Sal' (1957). Haapanen (1965). Willson (197~). Kendeigh (194:».Kat (1968). Boc' and Lynch (1970). Franzreb and Ohman (19711). andQuinlan (1979). and. for the most part. exhibit changes predictable from lhedC\dopmen! of ,'egela"on,

•

'1',,1>10".5 ('"mp"""'," of b"d mmmunlty P"'1",,'i<:, ,Iun,,! <'<'''y;lom ,1<,'cl"I~',,"'" in ,.,,",,1 <t."fe",,,, I"""" w"u, n\l(I-I,[" ,,,Iedec,d,,,",, ''''c,''

S,,,w,,,,'n;,1 ,,,,~,. ",Comm,,""y ll<,',<1",,", M",u,,, 0,,,,,, hl"d >~"sio" 1l""''''''Y Shlllh "," ~h,,,d """k, '1"""" Rd"",,,,,' ~Com/_m", fo",,, ...AI,,-,~. SD' 6,4Y ~,1I7 h.l~ ~27 '" Sp.,"lIcr .".1 >, ,,. 272 .HI 2X.\ :!II! K",,,,I ,

M II 'O\l) 1l..'\.l1J 7'""1 oJ'llI -, t 1')l\I," ,Yu~,," SI) ') ..~) 7.2X , 71J 7 I,j 11...·l><.~,· ,, ,~ ~, ." '" ( 1~7h) 0

M ,~, 27'12 ,-"!2 WM "Comic, ~"""'"'Y M.,'u,,, l"'I~ing """Ier lore" 0

Wyonllng SD Y,48 S,B • .00 10.87 s",l (IYS71,0, 121 I W W M iM 95.11 ~, ,,~ .124.

W"", Virgin'" SD n,ll/! H),II') 1.\ ..15 St"".,,, ,,,,,, ,, ,., ,~ !!It1 ,\I,ln.:h

.[" ';(01, I ,,~ 510.1 ( II'/<))

.\I""", SD 'M ~.7X 1U.~!.8.XJ IHI< 5.'12 SI""." a,,,1, ,~ ..... ,.1 1067.S76 m n~ ,\Idm-h ," "~ 635.1 76tll).44~7 "" 2.171' ( I~<;l) ,

•

Taole Y.S em",1 ","Su"",,,,,,,,,,,I,,,,F" rrom""",,')' Se",,,,1;" ~ M"",,,'

11.0£"''' 1""'1""'Y lie,!> ~h,ub S;'l""'~ fo','" I""·,, Rd,'"'''''' ,Iffl.",'<I/""". [ore," CMich'g"" '" DO 9.-ll' 9tH Y.~.l Ke"ocLIl!' l, ,~ ,~ 272 .W> ( ''!-Ill)

," ,,% ,m N~ .10.14 ~

III,n,~, SlJ YY! l~hJ 17..J7 1. ..17 K.r (1'IN<1 ,,'" Y27 117~ "" 0

" II 17~ 111']1 I I '/.1] 1l1,.!'1 ,,.".,"'~" ~I) ~,Ol '00 ~ I" "'" IVIll ~h"~;o,, "".I

I, ,,' 4B 11'1 W) "')2 h,,,,-"- I 1'17.11

" 421" 011011 10.17,1 \X7~ 7~.1

Geo'g'" ~Il I.~I ~, 1'1 ~A~ '1.111, III.]~ j"h",lon "",I, W 21.1 212 ~, ~ 0<1"", (IY>61M II<JI 248'J 1<J16 4.1'13 ~j..l1

• SD _ '1'<'6<, ""·<My. N "U",OCf'. M_ """"'"""I~.

170 7'h~ Rol~ of fiT~ III .\or/han Ci",/"poIaT EroI"'I~mJ

9A.J r",,·di,'urban« Respon.u

There are ronsiderable differen"'" bet"ccn unburncd and burncd arcas 1-10)ears ahcr Iog,ging in al~I.1I of the studic< rC"ic"'cd hcre, Ho"e'e,. nol all'tudie' of bird communilics in the firM 10 years after disturbance agr•• "ilhthe "end' of Table 9.3. U,ing »'mbnl, to ,how wh.rher e<>mmun,,).pro~rties are grealer ( .,.) or Ie" (-) in the disturbed ,'egetalion than in lhemalure or und,,'urbed 'land<;. and li,,,ng lhese in thc order of s~cie,

d" ..",), and numbers. ,,, >Iud,es sho" changes of (+.-). (-.+). (-.+).(-.-). (+_+). (+. no ehange) for the respect,,'e studies b) Webb n 01.(1977). Anderson rI al. (1977). Hagar (1960). Fr.nzreb and Ohm.n (197iS).La" renee (19M). and Bod and Ly'neh (1970). Perhaps the ,a.,ety ofresuh,IS to be e'~eted. considering lhe tremendous range of ,'egetation "'pesrepre"'nled. and hence thc possible "arictl of difference' in habital "ruc'urcbet"een malure and "CO, cry stage'. 1'-0 earl) post-dIsturbance studlC< fromlhe oore.1 forest "ere «presented here

9.~A Rird Cummuniti.. in lh.II<,,,,,"1 ror..1and Ihe rar :<:urth

When 'u,...·.',ion " a<lumed to be a sequencc from dcciduou, 'hrub tod"',duous tree to conifer tree. there appears to be a fundamcnlal differcncebet"een bnrea! and mid-latitude rcgiom. characterized b) conifer ""US

dc"<'idu"us dimas forells (Tahle 9.S), Species d,,'ersi'y' . numbers. and biomassof b"d communities all decrease from ,hrub to deciduous 10reS! 10 coniferou,forc'l "age" differ.nces in numbers and spec'ic'·ric'hness had alread} been,hown b}' Udva,d) (1957) and Tramer (1969). The csplana"o" hc' in the10" or produeti';" t}'pieal of mature spruce <lands e<>mp.ared "ith deelduou,stand' in the bnrcal fore't (Van Cle'e er al.. 1980). and perha!", in "fUelur,,1di' .r"l) ,,,,d 'recie, di' e,,"y of lhc vegetation. "hich maY' be greater in tallshrub ""nds than in even_aged I"J'OSt.fire bitch and Pop"l", 't"nd"

Ilow",e," ',uccc"ional de\Olopment" (i,e,. <land gro"th) of one ,egeta.uon ly'pc su,h as pu... red spruce (Slewart and Aldrich. 1949). as~n (flack.1976). or pine (H"apanen. 196.5) .tands. or "ithln either shrub. deCIduous. orconifer lype' (Spindler and Kessel. 19S0)). is generally accompanied byincre""ng 'recios d,,'ers,ty'. number>. and e"nsuming biomass It alsoappear>. from the r",ull, of Spindler and K"""I (1\180). that hoth agc "lth,nOne vegetation t)·~ and differences among tH'" account for signifieanlpo"ions of the 'arianee in bird communlt} pro~rties, WhIle man) ~peClCS

are ,hared among ..getation lYpeS. (from lO% to 30% of the .peele> "erefound In onl) one ,'egetat;on t)'~ (Spindler and Kessel. 1980» since onlysmall areas "'ere cen,used. this dissimilamy rellee!5 relat;,'e abundance morethan species absences.

Small-mammal mlit Bird Commu"",~_, '"K~S\le", (1979) re«nl slud~ of bird communil~ succe,sion in "'eslern

hemlock--.;ilka spruce fore'I' in ""'Ihea'iern Alaska is nol comp..able "'ilhIhe siudies in Table 9.5 since the dala "'ere based upon frequency ofencounler "lth bird ,pecie'_ The abundance ,nde' and specie, di\'C"il~

peaked al years 17 and .J(J (sapling to pole si,e). Species di' e"ily ",a. almosl.. high ,n 'he old gro"'lh (150+ .year) "age. bul Ihe populal,on index "'a.relati"el~ 10"'. perhaps rellecting lo",er deteCiabilily of the bird., Shrub cowrpeaked in years 11 and I'. ",hile 'hrub frequency ",a. k'''csi in the 17. and.J(J.year slag... H,gh co"er '" 'Ih the 10" fre'luenq suggeSts .hrub dumping atIhe I,me of maJIimum bird numbers and .pecie. di'ers.it~. 11 i. nol dear'" helher 'ertieal eanopy de'-elopment, sllrub co"er and spatial helerogeneily.produeli"il~'. or a combmation of the.. is r••pon.ible for Ih ...,. Irends inoommunil~ propeni",.

There are some ob"K>u, management impheation, to the.. re.uh•. Specie.d"e""y and mher properue. do nol unifonnly inerea.. all the "a) 10

e,"')',tem maturily. and di"ers;t), numbers. and consuming biomass ean bemarkedl~ 10"'er in mature northNn ..osystem•. Mi.<cd conifer---<1eeiduou.type. may pru\'ide Ihe maximum in .uch propenic,. but a mixture of d,.tinct'-.getation typo. i. needed 10 pro'-ide habital for Ihe full complemenl of'peCles,n a reg,on Thai thiS Should be Ih. case IS in line "'"h ronsideralion ofIhe habital and ne"ing requ"ements of the bird ,pecies (e.g., Godfrey.1966).

9.~.5 latitude altd Bird Community' Similarity

Theberge (1976) remarked on the great o"erlap and ,imilarit} among IhesouthNn Yukon b"d communilie' he .Iudicd. in romp<orison ",ith mid·latilude regions. Similarilie. among hi. sampl... are ob' iou., but he presenledno qaanlllati\'C comparison "'ith other region,. so one i. pro'-id~d here (Table9,6). Similarity "alue, ""ere romputed for .hrub and deciduous Irce stands;conifer fore.t. were not available for comparison in alll",,·latitude studie•.and if lhey had been. il i. suspectcd Ihat .h"'b-oonifer ,imilaril)' wouldcxh,bil a less pronounced lalitudinallrend, There appears 10 be an increase ,n,imilanly w;lh Iatilude (, _ O.7~." _ 9), The author is nol sati.fied "'ith Iherather subjecti,e deci.ions necessary to choose romparabl~ stand. in Ihe.hrub I)'PO. nor ",-ilh the .mall sample. and urge. mOre examination of thi,apparent trend before il is a<:<:cpled, The Theberge Sludy yielded aninexplicably high ,'alne. much higher e"en Ihan Spindler and Ke...r. datafrom farther north (Table 9.6): I"ith that value ncluded. r _ O.S3, If thi'relalionship ,s tenlati"el~' accepled. it suggests Ihat some p<lrt of Ihe lowe,species den.ilies of bird. in the ,uNrClie is due to greate' similarity in .peeie<composlllon In different habilal Ij'pt'!. AnQther component of nonhcrnspecies den.ily i' Ihe "ariel)' in habitatty1'C' existing in Ihe "egelalion mosaic

The Role of fire In Northan Circumpolur EeOJp,em$

Toble 9.6 Perc.ntage >Imilon'y b...d on bird numbers bet..·..,n bi,d communi,i., of.~ru~ '·mu. deCIduous f01'fi1 Ilands

Pereen'.g. utilude..milo.')" (':-) SI."", oomp.,.d Rd.,e"".

" " T..,,, ..ill"",' ,'."u' ,..." Th<berge (1976)I"'plar

" " T.11 ",ub ,·e"u, ~duoo, $pondle' ,nd K..... I(1980)

" " Tall cto..ed bru'" ,'.,.,'" Hoapo""n (1965)hard..<>Od

" " S<'ub me.do...."u, $011(1957)nOllond ..pen

" " Iloj: .lllrub ,'."'''' >«ondary Aldn<h{I9-l3)fore"

" ~ Earl)' .h,ub "'''u, mJlore Kar(I96/j)for.", ~ Open ...phn! ,.,,'" me". SlIugan and lame> (1973)fore", " Xeric 'hrub "e,,", oak Odum (1950)fore",

" 2O-)Ur g......shrub .."u, John""n .nd O<lum (1956)<>OK hick"'}' l<>r<>'

"Th< inde, oI .. md.n,~'"",,~ ~'.. (2 ~ M,n In.(;). n,I'I)) (V. + ".) ~'~'" ,\ • ~ n.t.!-

of the ,ubarctic: .I'hough sor.1 ,cget,lIon·tyl'" "arie,y is not e"mlned here.i' ..emS plausible '~O1 it de"'usos wilh Inerea.ing I"ilude.

YCl another component of ,ped., den,itl i, .. ;lhin-habi'al di'ersi,)': Table9.S sugg'''ted 'hat within·habita, species d;, ersit) w.s lower on 'he sub.re,;csites, This d,fference has yel to ~ a>.igned causo< ,uch a. lower product" i,)'(num~ .... and con.uming biomass are al<o lower in T.ble 9.5) "e ....us .impler"ege,ation Structure. The au,hor·s imprCloSion is 'ha, subarc,ic habnuts "hib"much Ie« Slructural \ariet)' bo'h '·Nliealll and horizontally: interior Alaskablack sp'u,,",. white .pru,,",. a'pen. birch. and bal.am popl.r 'tand. are mostl~'

ewn·aged (pos,·fire Or post-flooding) .nd are mosll)' pu,e s,ands, Produc·t;,·;t)' IS al<o lo"er lhan for s'milar nee species farther WUlh (V.n Cleve t, "I..19801·

Tentati,'el). one may infe' tha, .uccess;onal ,urno"e, in bird .peciescomposi,ion is less important in thc NOrth, Ho"e,'er. Table 9.6 refcrs onl)' 10deciduou~ sores. and <peci.,turno,·cr can ~ much grealcr in a deciduou.·to­coniferous sore (e,g .. Haapanen. 1965: Spindler and Kessel. 1980): thefire·telated patchwo,k of coniferous and deciduous types con,ributes much to

O\'crall .pecie. di'·e"it) in thc North (SpIndler and Kessel. 1980),

Small·mammal a"d Bird Comm'lIlilieJ

9.5 D1SCUSSIO~

For small mammals. numbe~. biomass. and c(lnsummg biomass arc hIgher inthe herb stage. and lower in the shrub and sapling stage,. than in matureforests: species dlYc"ily is hIgh and increasing through Ihe shrub 'lage. and isdepre,..ed in the sapling slage, relati"e to malure forest These trendsundoubtedl~' reflect changcs in lhe ground la)'er ,·egeta.,on. "hich pro,idesfood and coyer for lhe ,mall mammals of Ihe fore'l floor. Had tree squirrelsbeen Included. perhaps numbc". biomass. and consuming biomass "<Juldha.. exhibiled a continual increase through su=s'ion.

A, e.peclcd. for bi'd "'mmuni'ies lhere IS a broad trend of increase inspecies richness. species di'·ersit~·. numbers. and consuming biomass. and.dec rea", in biomass through succession. bUl thc trend " nol monolonlC. Forbird assemblages. speci.s di,·.~'t) increa>ed significantl) through succession.as e,peCled from pre"ious studies. Bowenr. change! in ,pecies di"ersit,­species richness. numbcrs. and consuming biomass from the 'hrub to maturestages "ere not significant when a'eraged among sludies. There "uesometimes significant changes among these stages in Ind,,,dual .tudie•. bUIthere "as also tremendous ,a,iabilily among studi.. and usually noreplicalion of habilat t~'pcs. Bird numbers and con,uming biomass increased>ig..fica.. ly onl) to lhe shrub Or saphng Slage. after "h"h lhere "'as a slightdecrease in the sapling or secondary fore'l stage and a slight increase again inthe mature foresl. In lhe 'hrub to malure foresl slages. ,,'hen numbe~ andconsuming biomass ".re essenlially slable. ,pecies di"e~it) and SpeCICSrichness increased onl~' ,Iightl) Bod~' size decreased and. corresponding!).rclatiye melabolic demand increased. thr<Jugh succession. Salt (1957).howeyer. reported opposite trends in th,ee seres. Tables 9.2 and 9.3 appearto bc the firsl altempl at a unIfied SU'ye) of lhis kind.

The conclu'ion of Bendell (1974) lhal population densilie! do not dllf.rbetween mature and post·fire habilatS i' not !>orne out for small mammals(Table 9,2) in the shrub O1ag.: and m Ihe herb stage. numhers and consumingbiomass are less for bird' (Table 9.3) and greater fo' 'mall mammal,. rcla'i' eto mature fore'l Species di'e~ity is clea,ly greater for 'mall mammals andless for birds earl~' in succe"ion. wjth mammals declinIng and birds increa,ingin diH'S11) In mid·succession. as the shrub and tree lay'~rs develop. It ISespeciall) Int~reSllng that changes afler the shrub slage are relali"ely small inlermS of aggregat~ measures. although considerable species lUrno''Cr i. stilloccurring.

Inler·dass competiti,,~O\'erlap i' probabl~' greatest bc1W~~n bird and smallmammal granivores in early succession: bolh lhese groups decr~a>e throughsuccession I" spile of the e"idence for large mammals (Bendell. 197J, Fox.1978). lhere IS 'till no solid e"idence lhal supposed pul>es of nutrienl' andprima'} production aher disturbance dircctl}' cause increased small mammal

'"and bird populations early in iu=s,ion. On the qualilali,e c,",deneea\'ailab\c. the jnil;.! high numbers of small mammals CQuid result fromgrealer ,..,lali,-. allocation 10 wed produclIon b) carl) suc.:essional herb!;(Ahlgren. 1%6: Krcfting and Ahlgren. 197~). or could reflect [fansien,in,a,;ons (Sulli'an. (979), Habitat facto.. other than food could be the,mponant determinant' of small·mammal and bird [lOpula,ion" field ex­periments arc needed (0 rewl,·. thiS quesuon

There arc.' least tWO promising lines 01 in'lui~ nol exploited b~ Ihe c;tcd,ludics, bearing upon lhe conlrol of species dj'orsi!} and numbers in post-fireand ""ral habitat., Firsllhcrc is a con",nsu, (Cuni, and Ririe)'. 1975) aboutIhe imponance of lhe herb and ,hrut> laicr of forests and woodlands 10hruding_bird num~", and di,·.rsill, In on. forest 'Iudy (Preston and Norris.1~7) more tban 70% of all n~t' WefC less tban 2 m from the ground. HO\,pcr.11'1. (1973) found ,hfUb co'er to ~ tbe single most imponant correlate ofbird spee;"'; .ichness in campground are.. "'bich are both forested andpark·like. Presence or absence of "'·ell-de'eloped fore'l undemorel corre·late. with imcr-sl.nd and inter-,..,gional differences in bird ,pceie, richness inoth•• case. (Cuni! and Ripley. 1975: Odum. 1950: Ste"'·an and Aldrich.1952. S.lt. 1957: ~laek. 1976). Of the pa,serine, li'l.d in Godfrey (1%6) andlik.ly to occur in bor.al ser... discussed h•••. aboUl 50% art limited '0nesling on or near the ground. in shrubs. or ~low 5 01 in lrees. Aboul.3l)..4()% may ~ flexible ,n ne,ting_heighl preference and 2(}..[I)% n." abo,'e4 01 in u •••. A majoril\' of th. smalL foliag.-searching insect;'·or•• lik.ly to~nefil from loti.ge profile de,·elopment in lhe lree canopy also prefe••o neslup 105m abo"e lhe ground, It seem, lha' neslIng and foraging guilds 'hould~ con'idered simultaneou,ly as interacting nich. dimensions. ralher lhanseparately. as i. SO often done, It also appea'" lhal full use of lhe foliagep.ofile b) b•••d,ng b"ds could be IImit.d bl ,,'ailability of ne.ting SileS. i.e,.b~ ,hrub co"er and profile. The aUlhor i' nol a""ar. thaI succe"ionallrendsand posl-dislurbance diff.rences in forest bird numbe.. and di'ersill ha'·.been rel.led 10 shrub CO"er (e.cepl Flack. 1976 and Hooper fr al.. 1973) oranalysed ,n lh. joinl niche space of foraging '."us nesling siles, The effect ofd.creased di'e"itl and numbe.. somelimes appearing dUfing the .anopyelosure Slage in mid-sucees'ion mal' ",.n be e.plalO.d by lh. ehm,na"on ofunder<lorel' herbs and shrubs al lhal lime. Experim.nlal manipulalion ofshrub co"er and palchiness could help leM h'-'POlhcse. aboul limil.lion byncs.inll silcs 'ersu, foll.ge profile and p"mar~ p,oduclion.

A second line of inquiry concern, spalial hclerog.neity of \Cg.lalion a, ilaffecls bolh num~rs and ,pecie. di"e"ill (Ro.h. 1976), Two sludies ofdisturbed areas suggest .natogies 10 post_fi", ".gelation. Fif'lt. Hooper f' 1'1( 1973). sludied de"eloped campground ,iles in lhe soulhern AppalachianMounlains and found that bird ,pecie. richness increased ",'ilh admixture ofconiferous and deciduo-us lree .pecies (also appar.m in Spindler .nd Kessel.

Small-mammal and Bird Comnll",i,,~.

1980; Tramer, 1969) and wi'h 'hrub co'·... More interes'ingly. SpeC1~S

richne.. was greater in 'he presence of large dense shrub dumps than with a'imilar percentage co"er of smaller dumps. In park-lit. area•. the size of thedumps in addition to rotal cover was an imponan, factor; larger. denserdumps pro,-ide more ne'hng CO,'er lhan lhe same ,",,)Iume of more uniforml)dimibuled foliage, Thu,. shrub and sapling clumping and spalial hetero­geneity may be one imporlant factor in determining bird numben anddiversil)' during post-fire reco'-ery, Se<:ond. Forman rl ai, (1976) sludled blld'pecie< den,i,;e, in forest 'i,land,' or ,,"<IOdlo,s in lhe mid,t of agricultural or.uburban land' in Ne" Jersey. Forest i,land' has'e more ,peci.. than theequi'-alent area in continuous forest; lhe added specie< are mainl)' 'edge''pec'e, and mostly' are insecti,,"'e', Th" could be explained by lhe obscrs'a­lion that lerrirories "hrink' "hen bird' are out of sight of compe,itors alongone Or more lerritorial b-oundaries (Ud,'ardy. 1957)_ Further.lhere aTe more<pecie' of bird. on one large island lhan in so'eral ,mall i'lands ha,'ing lhesame lo,al area. forman el 01. (1976) dISCUS' lhese re,ult, in 'he lighl of lhelheo') of island biogeography and .peeies-area curs'es. Our specifIC intereSlhere is lhal degree of post-fire horizontal spatial helerogenell)". Or '-egelationpale hines>. may have importanl effeels upon posl-diSlurbanee bird com­muni'ie', There seem 10 ha'e been few allemp" since MacArlhm tI 01,(l962) 10 study the horizontal "ariation in foliage structure in relarion to birdspecie, di"ersity and numben. ailhough venieal dIstribUtion ha, been s'ud'edextcn,i'el)'. Ho"'es'er. ROlh (1976) has 'ho"'n lhat horizontal spatial hetero­genell)' Or palehineu is relared ro bird species di,crsit) in fOreSl and brush.and can aeeounl for speeie, di"ersity differences not well accounted for bycorrelalions wilh foliage heighl di'-ersity, funhec application of thesemethod, in posl-diS'urbance and successional studies is highly deSirable.especially in the Iighl of the high dcgree of patchiness (and the con.inuingchange, in patchiness) "'hich may characterize reco"ering boreal eeos)·"em.(Shafi and Yarranlon. 1973; Borkin and S<mcl. 1973),

Finall~.. Simultaneous. quantitati"e stud ie, of plan, and anImal eommunn)ehanges are rare. integrated studies of successional de"elopment of plantspecies. primary production. habitat 'truClure. ammol food resourceS andanimal populatio,," 'hould be undertaken, In any son of 'ludy. repllcarion isneeessar~_ E,penmemal method,. such as pro'ioon of or remO'lal of foodand co'.... and u~ of herbIcides and fenilizers. ,,'ill pro"ide more insight inlocontrols owr community de"elopment than ,,'111 repeti'ion of the general typeof studie, cited here,

9_6 AC"-,,OWLE:[)G[.\!E:-'TS

Preparalion of Ihis cbapler was supponcd by NSF gram DEB·78-11594. 'Thestructure and funerion of a black spruce forest in relation to other fire-

Thf Rolf of fi1f ill SOr/hun Circ~mpol~l Eco.ysln",

affected taiga ecosy".m,·. Particularly for discussions aboUI laiga foroS1 plam

sucrmion and post-fire de\'e\opment. the author is indebled 10 lhe follo""ngrollfague, for their insighlS. although the) should not be held accountablf (01hi. "ie"": J. P. Bryant. F, S. Chapin. C. T. D)rne... J. lsi Footf. B. Nieland.K Van Cle"•. L. A. Vi.fCC~. and J. C. Zasada The author i' especiallj'Indebted 10 Mike Spindler. Brina Ke"el. and Sue Ann Ouinlan for u>e oflhcir dala. and to an anonymous rc,'i.w'er. who pro,'idcd many u",fulrommenl•. some of whieh arc incorporaled ,nlo lhi, chaplcr.

9.7 REFERE~CI::S

AbTll!"'k)·. Z. (19·~) Small mamm.) commun,,> «0101)": <hanl'" In 'pea« d"''''ljin 'espon'" '0 manipul.ted produetil'll). Orrologi<> • .J.I. 11 :l-123,

Ahlgren, C.E, (1%6) Small m.mmal' .nd reforc,l.hon follow'ing pre",,,bed blJmtng.J. Fo! .. 64. 61-Hi18,

Aldri<h. l. W, (1~3) Biolo!"e.1 .u"..)" ollhe bot' and ."amp< in norrhea"ern OhIO.Am. MIdi. Nal 30. 3-16--102,

Andcrson. S.H ..\I.nn. K...nd Shul'''' H II.. Jr (1971) The effe" 01Ir.n,mi",ion-lme rorrioor> on b"d population•. Am, Midi, Sal .. 97. 211'>-221.

B.nfield. A.W.F, (1973) Th. Mammab cl Callodo. Un;"e",,) of TcronlO Pre".To",nto. 4311 1'(>.

Beek"i'h. S.l. (195-1) E.rologie.1 .occe»ion on .b,ndoned farm I'ml> .nd 11<rel,'ion.hip '0 w'ildlifo m'n'gement. Eeol, MOIlOg1.. 2-1, 3-19--376,

Bendell. l.F. (1973) Elle<t> of fire on birds and mammal'.;n T,T. Kozlow"ki at><!e.E, Ahlgren (eds). Fi" and E<"O,,·jl<llll. A<ademit Pre><. New York. pp.7l-1.l3.

Il"ne). E,e.. Gr.nl. \.\ .E .. and B...d. D. D (1976) Impon.oo: of "egeta",e ""cr 10eyel., of M"";Jlus popul."on,. £",,1(8)'. 57. l{).ll-lOSI.

Block. 1i.C.. and ll00'cn, E.I1. ()<l74) Re,pon>t of ,m.ll·mammal rommun"i<> 10hobn.t eh.nif' In " ...tf'n O'f&"". ,n 11.e. Blact (<<I). ""ildli/~ aru1 ForniM.~"&,,,,~m i~ Ik~ PaCific NCr/h·m. SclIoo) of Fore"')'. Orego~ SlateUni"fn;'). CO" .Iho. 236 1'(>.

Bock. C.E...t><! LYll<h. J.F. (1970) 8r<edlnl b"d popula""n. of burned andunbIJmed ronlfer lore." in ,he Sie". Se'-.da. CQJld",. n. 182-1119,

Borm.nn, F.H. and Liken,. G.E. (1979) POII"~ alld Pmens i~ • J""."dE<:c,,,,'~m. Springe" Now York, 153 1'(>,

Ilotkin. 0.11 .. and Sobol. ~u, (1975) Slabilir~' in time."ay)'ing e",.)·"em•. A", Nm.•109.6lS-&l6.

Bra)'. J.R .. at><! Gorham. E. (1961) L,ne, prod....ion;n fore." 01 ,b. w"rld. Adv.Eccl. R.... 1. 101-157.

Bre,..r, R. (19S8) Breedm8-I",d popul'''on. of 'trlp-m,ned land ;n Pc,,)' Coun'y.lilinai•. £cal"8) 39. S-ll-545

Comoron. A,W. (1961) Compe';'i"e ..eluSIon be'..... n ,he r<>denl genera IfirIQ!IUand Clnltricllcmp. E,'clal"'". IS. 6JO-6J4

Coot. S.F, (1959) 1'h< eff=. 01 fi,e On a popula'ion of small roden ... £ccJog)·. -10.102-108.

Curti•. R,t.. and Rlple)', T, fl. (1975) Wat<r managfmen' ;>faCio.., and 'heir effect onnona.me bird babi, .. ,'alu« in a de<iduOll' Io'e" comm"n"). In D.R. Smilh(ed). SJntp<>Si"", "" Ma""8'_"1 al F"">I alld RailS' Halula" 10' -"oIl8a_

Small-mammul u/Jd Ri,d CQmmomlli~J

B"d,. liS Fot, SeIV.. G.n Te<~ R.p. WO·I. pp. 12lI-l.1.Da"",. PI' (1917) C."od ,.spon.. '0 fo'e,' li,e and c1ean;u",ng ,n """he.st.m

W~..>n"n•. J. ""IJI. M~"",g",,,,.71;5-788.Eadoe. W R. (1953) R••po".. of .lIoeml'" '0 ,'e!ClOtl,·. <0"<1. J. Mam"",l. J.l.

26J-2t>.lFaabo'g. J. M. (1977) M<lObohc rale., r.",urce, .•nd 'he ",""u,renee of ""np.o."""n••

in 'err.."i.1 ",'ian rommuni,ie,. Am, ,\'al" Ill. '!O}-9J6Flack. J.A,D. (1976) B"d P"pUlallOfl> of A'pe" flJTe'" in W<f"'" \',mh Ammo".

Am. Omi'h. Union. Ornith ~'onogr, 16.97 pp.Form.n. R.TT.. G.lIi, A E..•nd Leek. C.F, (1976) FOf<>t >I,e and I"an d,,"""! ,n

Ne.. J"..y ",'00<110" .. i'h ",m. land ,,"" apphcahon•. Ot«>k>,il1. ~. 1-8.Fox, J ,F, (1978) For••' fir., Ind file 'no""l1oe h",~anada I~n. cyck. Ouologol1. JI.

)-19--37-1.Franzreb. ICE.. and (}bma". R.O. (l978) 1lI¢ dr.e" of umber un'e,un! on

br.e<l,n. b"<l> ,n a m,,,,<I-wn,I«o.. lor.st. Condo,. 110. 431-141.Oosh,,·,I«. J.S. (1970) Plan' and mammal change; in a dear"" in ..·.,..."'n".l

Of<gon. Eeolog)·. 51. 1018-1016.Gel<, L.L. (19611) No'" 00 tile local di'tribu'ion of Nrcm)'''''' I'aeopu' and bp",

"ud'OIIieu,. Am, Midi, N", .. (;$, .86-500.G<". L.L. (l96lb) FactO" influencing 'he local di"ribuuon 01 ,\fiero<'" and

5,"n~pIOmH in ""'them Mich,!.n, Eeolo,.,'. U. 110-119,G.". l.L, (196lc) F..,o>r> inllu....ng tile local dL>lribuuOll of ,1".,,-.. Am. .Ilodl.

o\'~,. (;$. 67-88.Godl,.). E. W, (I\lt\t» Th~ B"'/h of Canada. National ~lu..um of Can.d•. Otta" •

•28 pp,Granl, P. R. (1972) In'.r<pectflc "",mpetLllon among rodent>. Anna. R,". £coJ. 5",,, ..

3. 7'1-106.Granl. W.E .. Ff<och. N.R.•nd S,,·ift. 0 ..\1. (1917) Re"l""'''' of. small mammal

communi')' '0 ....,...nd nitro~n trea'ment> in. >bonpas> puin. "",,)-.tem. JM.mmal,. 58. 637---{61.

Hoap.on.n, A (1965) Bird fauna ol'he Finnl,h fore... ,n r.lotion '0 fOrt.,.u",,...,oo.Ann. Zoo!. Fmn .. 2. 153-196.

llagar. D. (1960) The in'<rrelation>hlp> of logging, bird>. and umber regel\<r."on 'n'M Doup...fir region of ""nh'e"em C.I,lorn,a. £<:0/011.". 41. 116-12S.

!lOOper. R.G __ Cra~-rord. H .S.. and HaTl"",. R.F. (1973) BlId de",,,) and di,'o,,," .,rel.,cd '0 vc~ota'ion in fore>! ,e<,..t;onal area>. J. fo' .. ?l. 71k-7tn.

lI"'d. L.E,. and Wolf. L. (197.) S'ability in rela'ion '0 nn'".n' e",ichment In..fhropod con'am." 01 old field ...""".."nol """l.tem•. Eeoc ,\f0ll08". oW.46S--lS2.

Hurlb<". S.H. (1971) Th. I\OI>COl\CCpt of speco.. <I,,·."'t) a <"tiquc and ",me.I'ema,ive p.oumete". £coWg". S~. 577-581>.

lame""'. E. W (1949) Som. facto" ,nfl"....n; lh. local dL>lribution and obund"neeof "'oodland 'mall mammal, in centr.1 N.... Yo'k. ), ,I'ommol" .!O. nl-BS,

John"on. D.W, and Odum. E.P. (19S6) Boe.dlng b"d popul.llon, in ,ol.uon '0plan' 'l>CC<:..ion on file Piedmon' of Georgia.. £<:0/01.". 37. 50-<>2.

KOf. l.R. (1968) Habitat and a"an di"enl1y on st"p-m",.d land ,n ..st«nttallIIinoi•. ConJOF. 7G. 348-3-57.

K.ndei~h. S.c. (1946) Boee<l"'ll bird. of lh. b<.<h-maple---bemlock comma""),E<ol08y·27,226-24S

K.nd.igh. S.C. (1948) BlId population••nd biotic "",mmuniu", in nortllem lo...erMIChigan. uoWg)'. n, 101-114.

K....er. W.B, (197'9) Bird population r"I"""'" to dear""ttLng In tOo To"~a..

'"" .. ,,,,,.1 rOT." of South."" AI".... US For. SUI'., AI","a RqIOM. Rep_ So. 7122pp.

"'1"",•. 8,,\1 (911) Re,pon>c of br.e<i.n~ b"d I'Ol"'lallOll' to hab,t., ella.g., ,n aGi.nl SeqOOl' 10'0>1. Am. Mrdl. ,'al .. 35. 13S-1S2.

K"chne,. T B (1917) The efle<t' of resource .""ehment on lh. dll'ersi')' of pl.n"and arlhropo<k in a sIIong.... prai"•. £(011)1:)', 58. 13.u-l.l.l.4

KI..I>o,. ~1. (1%1} fu fi" of Llf_. John 1'.,1.,-. S ... York. ~S4 w-Kre"'. CJ .. and "",ogat•. I. (1976) Small mommal communltt... of lh. Klu.ne

ReSion. Yukon Te""o'). Con. Fkld ,\'at.. 9(1. 379-389.Kref"ng, LW and Ahlgren CE. (1974) Small mammal, .od ,'ego,."on ,h.ng..

• fler fir< in a mIXed ronif<r-hardwood for.". Ecoiogj'. SS. 1J91_139ll.LoBu" _L and D.m<II. R.J>1 (]'WJ) EIf= of habita' di'turbooc< on • ...,.11 mammal

J'OPUlation. J. .If"",,,,,,/, . .lCI. ~l.l-436,

uel;. D, (1933) Hab"., selection in b;r<f<, .. "h speci.1 rer.f.""" 10 the .ffoc" ofallorestall"" on the Brockland a,·,f.una. J. Anim. Ewl,. l. 239-262,

Lack. D. (1939) furth", eha"g". on the Brockl.nd a"fauna ""used by affo'e'lta'ion. JAnim Ero/.. 8 . 277-285,

l.<>ck. D.. and lack. D. (1951) Fu"he' ehan~... ,n bi,d hfe ..used b~' affore"a,i"". JAnim. EooJ.. ZOo In-IW,

La,",,,,·,k,. R.C .. and Da""",,". W. R. (1%1) A ,,,..,x,mina""" of the ,ela,ion be'''''""n"."dard me'aboli< 'ate a"d bod) ""e'ght i" btrd•. Condo•. 69. 12-23,

La...-ren«. G.E. (1966) Erology of Yert"b,ate .nim.l. in ,,,I.tI,,,, to ehaparr.1 flre ,n,he Sierra """ada foolhill>. Ero/"8\ ~1. 218-291

~rl""'key. R T. (1915) Habi'a' ,I>Ctt>Yon and ,oden' di"ribu'ion. J. Mammal .. 56.950-9S5.

~rClo>key, R.T .. and Fkldw"k. 8. (1975) Ee<>logical .epa,.tion of ')'mpatrie,ode"". J. Ma"'mul.. 56. 119-129

.\taeAnhur. R. H, (lllbll En'ironme""1 lactors .ff...,"! b"d .pea", di\'.""~'. Am.~.-u,.. \18 • .l81_391

.\laeArthur. R. H, ..\\lKArthnr. J. w__ ond I''''<r. J (l962) On b'rd ,pe",.. d"'erM"II Pre<l,ellon of b"d con,u, f,o", habi'at mea,ure",e"". Am. Sal.. 9(" 167_174

.\IaTl"II. A.M . and Rad"an),. A. (1971) Cban~e' i" 'mall mammal populat",n, aftere~"""lIm8of Nonhe,n Ontario black 'p"",e fo,est. Call. Fi,ld .\·Ul.. 91. ~1-46.

\\artell. A M .. and Pea"",n. A.).1. (I91S) The 'mall mammal, 01 the \\aekenIle Del'aReglO1l. North""".. Teni,one,. Canada. An'"·c. 31. 475-l1l8.

M.ft,n. N.D. (1960) An anol;-,." of bird populal101ls in ,el.,ion to 10''''' ,ue.,.",,,," In

Algo"'lui" Prm'i""ol Pork. Ontario, Ero/h~'·. ~l. 126-1010.),1.). R.M, (191S) 1'.1Ie,,,, of >pea" .b<lnd.nce.nd di'·e.... ')'. in \\.L. Cod)'.nd

J M Dlamo"d (ed,). £eolol}' und F.\'OI~lion of Comm~nirj... H.", ..dUni,·eI>lt}·I"res>. C.mbridge. M..... pp 81_1W.

\\e".ughlOn. S.J,. and .....olf. L.L. (1~73) (j'n"al Ecology. lIoll. Rineh.rt andWin,lOn. Ne'" York. 710 pp.

\lellin~er. M V,. and MeN.ugl""". SJ, (1915) Stru"'u'e and IUnel''''' of ,,,,,co"lOnal.,.""",1.. p1.nt eommun'tI.. ,n "".tral Xe'" York. Ero/. .110111>1"" ~,. 161-182,

~hll",. D.I t.. .nd Get>. L.L. (1m) Fact"', ,"nue"",ng loeal d",n!>utlon .nd "1'<"''''d"......'y of I",e" .mall mammal. in Nc..· England. Cun. J. Zooi.. 5S. 1I06-S1•.

\\o"i•. D,W. (1m) \lrerohabi'at ntili",i"" and .peci... Jimibution 01 'ympa'n<""all mammals in ",u,h~'e"em Ontario. Am. MIdi. Nut.. [OJ. 373-J8.I

\\o,ri•. R.D. (l\l69) Compo'i'i'" exclu,;"" be'''''""n .ltic""..,..nd Clcth""""",ys \n'he »pen parkland of S..kat<he",·an. J. Mamm"'.. SO. 2\11-301.

Small-mammal and B"d Commu"",~. '"Mo.-ri>. R F, (19~~) Pvpulation "udio, on ",me ,mall fOfe>! mammal, m ,,""em

C.nada.l. Mammal,. 36. 21-35,Odum. E.P. (l950) B;rd I'O?"I'lIons of the If,ghlanJ:. (North CaJOJina) Plate,u m

rela"on to plan' ,uro:",on and ",',an in"aMan. Ecol<>gJ. JJ. 587-605.Odum. E. P. (969) The .trategj' of eros""em de~lopmen1. Sc~nu. 16-1. 262-27ll.(hm~ton. J.D. (1962) O""n,i"'''''e erolocy .nd ,he ,•.-oodl.nd erosj"'em CQ<l~p'.

Ad,· EooI. R",.. I. 103-192Pe."",n. P. G, (1959) Small m.m"",b IUId old fidd ,UC<'eMion on ,he Piedmont of N...'

Jet'><j. uol<>gJ. "0. 24\L-2S5Pre"on. r. W, ••nd No.-ri>, R, T. (1947) Nesting h.ights of oreedin~ birds. Ua/"8)',

28.141_173,OUlnl.n, S Eo (1979) Eff«" 01 ""nnolled Oumln8 and 'u«"Uon of "'hite '!"uu

fore'" on b-reedin~ b;rd communities. Ken.i Penin'ul., Alaska. US for, St"'..ChugMh Nou'"",,1 fort:it. U""ab/. Rep. 64 pp.

R,elten,. V.B (197~) Numbers .nd habItat affin,,,e> of >m.1I mammal. ,n nor'h­.....tefn ~to'ne, Can. Field SOl.. 88.191-1\16,

Ri.be",U. J.r. (1974) Poradox of enriellment in compe',,,"e 'j..'ems. EcoI"lY. 55.183-187.

Roth. R.R. (1976) SI"'''al heter",e""1j and bird speei.. d,,'ersily. EcoI"l)'. 57.T73-782

Salt. G. W, (1957) An analy.i, of a"itan"a, in ,h. Telon Moun,.in••nd l.ck",n Hole.Wyomm8. COfIdor, 511. 3H-393.

Sh.fi. M.l and Va"anton. G.A. (1973) V.gelation.1 hetefogeneity during."""",d.'} (postfire) 'UoCC<'»ion. Can. J. fIo<.. 51. 73--90.

Shupn. H.1l Jf. and 10...... D. (1'17]) Erologocal .ucrewon of breedin~ birdpopulot;o", in nonh..·..'em ATkan,",. Aak, 90. 62_T7.

S,mms. O,A (197'9) North Amc,;"an ..·....d., "SOUr<'< uliliza,ion and di"ribu'ion.CO" I. Z<>aI.. 57. ~5:!O

Sim,. Il, P, .•nd Buokner, C.Il, (1'173) The effec' of ckar rutlong and burning of Pm",banbl.ono fore". on the popul.tion' of .m.1I m.mm.b in "'u,he"lemManitoba. Am. M,dl. SOl.. 90. 2ZS-2.l1.

Spindler. ~t.A .. and K....1. B. (19&1) A'i.n popul.tion, and hab,t., u'" ,n inleriOTAI.,ka taiga. S)·e>is. IJ. 61-11).1

Sl...·.". R,E .. and Ald';"h. J.W (1949) Breedl"i blrd popul."on, in the 'pnlceregion of lhe ceolrol Appalo<h,a",. EcoIOK>__ .10. 7S-81

Sle"an. R E...nd Aldrich. l,W, (1952) Ecological .ludi.. of breeding blrdpopulaM'" In Nonhern Mame. £(01"8)', JJ, 2l6-B8

Sulli,'.n. T. P. (197'9) Demographj' of I'Ol'Illa1io'" of deer mice in """tal forest andcleaHu' (Ioucd) h.bita", Can. J. Zoo!.. $1. 1636-16-18,

Te'ter, J.R, (I96S) Effect. of a ronlrolled bum on "".11 m.mm.l, in a Mmne"'l'oak-"","n.... Am. Midi No', 74. 2-10-143

T..lS. L., Jl (I9S60) Eff= of a 'lash burn on fore" mice. J. Wild/. M.n'W~ .. 20,""-""

T,,'i•. L.. Jr (19S6b) R«pon", of """II m.mmal, 10 iouJng of Dougla,-fir. JMammal __ J1. 189-196,

1heberge. J. B. (1'176) B,rd population, in ,he Kluo... Mount..n,. South""'1 Yukon.Canada . ...-i'h 'pecial rererence to ,'ege,.tion and fire C"". J. Z<>aI __ $-I.1346-1356,

Tr.mer, E.J (196'J) Bird .peei.. dll·."I,y: componenl' of Shannon', formul •.EroIQfY. 5(1. 'fiJ..Jm.

,.,Ud\·"dy. M.D.F. (1957) An ~,·.l"atio" of qu.n'ila';" ,,,,,lie. in b",l<. Cold SP""S

Hu,/)ar Sj'mp. Quan'. BIoi.. 22. 301_311Van Cleve. K.. Dyme... C.T".nd Vierc,k. L.A, (l980) Notrien' cytling in interior

AI..ko flood p1ai",.1><1 its Telation'bip '" ,.~ne,"lion .nd 'oMequcnt fores.dc,-elopm<nt. in M ,\,,,".yand II. .~1 Van "'Idh"''''" (ells). For'" Rq"'t'Qtioll01 Hi&h LI"irudts.. US For. Sc"'" Gen. T«!I. Rep. PNW-l07. P!'. 11-18,

Vi,o....,k. P.~' and R.i ..... W.A (197S) Ero<y~,.m OII<CC...,., and "o'nenlro'cotion,. h)-po<IIesi', BioS<mKt. H. 316-3!!1

Webb. W,L.. Behrend. D F ••n<1 ~,""". B. (1977) Effea ofloWng on sonlb"dpopulalions ,n I ""rlhern h..d....ood fOf.," 1\ ,Idl. Morn>g', !!. 35 PI'·

W.in, R. W. (197$) VegetallOn '='-"j '0 >"toe M,d.. and l"ro.H"n<lra .fler fir•.Can. IXp. Indian Aff, Nonh. [k"tlop .. ALUII Rtp. 7s_75-&. liS pp,

We't, S,D. (1974) Post·bo,n popula1ion "'ron" of ,he """hern r.d·l»ck.d 1'01•.Ck,lIricltmn)" rulll.... In in'enor AI..k•. MS 11>csi•. Uni,'.,,;,), of AI..k.,F,"book<, 66 pp.

W.... S.O, (1979) H.h,u., respon;e> of ml<T01ine rodeo" '0 ceon.l AI.,k.n lorn'~. PhD 1lIe>i•. Un,,'.rsi')' of C.liforni•. Ikrkeky. ll~ pp

\\,."'. l.A, (l97S) A'''n com"'uni"... enerl'.'i....rId fur><tio'" in coniferou. for..'h.hit,.". in 0 R. Smi,h (.d). Pro<:, S,""'p. Ma""8'_01 of Foml a'l<1 Ran8'/labila" for Non8""" Birds. US For. Se" . G.n, T.ch, R.p. WO·\. pp,226--W

W,lhoo. M.F. (1974) A,'l.n commun")' or~,o,,"io" and hab"., >"'''''ure. Ecology.55, 1017~1029

Zar. HI, (1968) S,.nd.rd mer.bot"", rompa"""'" be''''eon orders of bud•. Condor.70. 278.