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 of Small·mammal and
Bird 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 from
th" 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", bnJ
oommun,"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, ..d
Iouer ,n lbe >!>ruh and ..pl,nc"age>, ,hoo ,n malure f,,'e,". >p<C1es d"'ef'>l'l of >maU
mammal, "' 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.mmal
and bird a"embl.gos during pos'·fire '·egel'li'e ,u",e"ion ;n Norlh
American boreal fore'l. M,d.I.'itude e~rcriene<: wi'h plant suece"ion and
ecolog;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)". In
the general lheories of ecos)'S'em rcoo>'cT)' (adum. 1969: VilOusek and
Reiners. 1975: Bormann and Likens. 1979). p"ma0' prodUClion is thought 10
peak somc time in m,d-.uccc"i"n. Earll' sueee"ional 'tage' may exhibit
mOrC rapid energy flux .nd nutrient cycling. smallcr organisms, and grcater
metaboli'm per unil h'''mass. Specie, di"ersit)' mal be 10W"1 carly )n
succe.~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~
'"
Firc
u~uall\
of Fi" in ,\'orrhrm Circumpol,l' £(O$J'"''''
rele.se, a put", of .".il.bl. nutrients. ,.n.ned in high
nIlrogen and phosphorus eontenlS of plant. during the earl) post-tire years
(Bendel!. 197J; !>la,Lean rI al.. Chapter 7. thiS ,-olume). An carl) increase in
fenihll" and pWrluCli'"j or rdat"'. 8ro"th "1<"$ 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) one
pan 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). and
plants (Mellinger and McNaughton, 1975).
Bendell", (197~j analysis of th~ ~arli~r 'ludies on I""t-fir~ bird and
mammal communiti~s led him 10 th~ following condu,ion" Communil;e,
displayed conSiderable "ah,lil), ,n a, much a, th~ num~r of indi'iduab
changed linl~_ d~'pil~ marke<l chang~s in ,pecies coml""ition. There were
n~t gains in num~r of species, For mammals. tne mean gain waS 'mall and of
doublful significance. and for birdS. the no" 'pecies were mosrl) gmund· and
shrub.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" "crc
uam,n~d;n onl) on~ study (Bock and l)neh. 1970). in "hieh numbe... of
birds remained "inuaTl) un<:hanged afreT fire. while biomass and energ) fl""
increased_ With the exccplion of Ih,s last reporl, BcndeWs infcr~nccs run
counter 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" of
posl-f;r~ and unhurned eommunil;". To increase lhe dala ba'" I""t-Iogg;ng
and olher successional 'tud;es "-ere considered in addilion to the I""l-fire
dala,
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') "f
Ctn,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 data
standardized to a uniform basis of 100 ha. Small·mammal lrapping studies
usually gn'e only r~lali"e eSlimare,. "hich are n<)l referable to numbers per
unit area. Tho<. ,,'ilhin an) gi'-en stud) of small mammals. ral;<>< of
communi'y propcnies such as number of indi"idual. or biomass bel"een
post.fire and ma'ure Slages were calculated, These ratios "'ere lhen used for
romparison 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-e
categories. based on struclU,e of tbe sueee"Iunat ''''ge•. arc likely 10 be more
dlrectl' 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. whero
aly" " an empmeal relation bet"een bod}' "'eight and ba",1 metaholtsm
Con,uming bioma.. e~uals 70\\·'"' for 'mall mammals (Klciber. 1%1) and
1191\"-" 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). consuming
bioma,s of passcrine> "as actuall~' calculaled as 11"' ". (amlnlng lbc
coo'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 a
b.,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. Thus
b",m"" and con,umlng blomas< "ere eakulaled In grammes 100 ba and lhe
nonpa»"ine cons"mlng bioma« "as adja"ed 10 be metabulically equi' 'lent
10 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""'er
law 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 II
estimates ael"al commUnll) metabolism.
Body "eights "cre "",mbled fram Banfield (1974) far 'mall mammal' .nd
from 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 by
relall"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"<cnted
specie, "hleh would g;\C lhe same ::'p;. Speel" d"e"," in each c"", "",
calculated lrom eon,umin~ biom,,, rather th,," oumber'. bee""'" 'hi' should
better 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,' from
lhe '.r) >outhern boreal faresl. "ith no published da'a from the apen
spruce-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, and
Buckner. 1973); ,pmcc_lir fore,I' ,n norlh"c'(Nn Maine (~8"N) (Richen'_
lY74). <outhern Ontario (41)"N) (Simms. 1979). Minne"'t. (#N) (Krcfting
and 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') (Martell
and Rad,-an) •. 19TIl. For inte'ior Ala'h (6Y1\ 1 there is unpubli<hed work in
blad' spruce fore<1 burn' (We'!. 197~. 19791. In lundra. on the Macken/ic
Riwf Delta near Inu,-j\;. Wern (1975) Ir~ppcd 'mall mammal< in. burn for
nne \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 or
deerea'" o"cr that In mature lore't. but that number<. hloma". and
con'umln! bioma» In e"rl~' '''=»ional Slages a,e inereased in m",t
in'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, beIng
pfCscnt 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"
("ommun,,'
Ch.ngc in eommnoi') plope")'
SismfLO""""
De<:rea>e \;0 change
[e,.I·
[""f.'''''
,
,,,
SpeCIe' nchne"
Specic> di,'.",';
""rnl>o"
Biorn.»
("on,nrnmS biorna«
,
"S<s.,f",ancc " '",,<d ~ "h ,I>< bonum'"
,h.""" "f ;oc" .....oJ
'~S· "'" "Inif"".,
~<""
,
,,
• "S'
"
"
" "
""
"r«'0l... <q.ol
11,0.'
00.'
"i.",~,,,,,,
on ,I><
<>I
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!",""
I'Umbef of "udie.
Specie< di"el<lly
Number>
Mean body ,ize
Con,uming biom."
Velyearly
helb
Herb
Shrllb
,~
•
I. ~7
1.71
US
,~
"~
0.87
0.8!
1.21
,~
.'
,.
,~
"
Sapling
."
O.M
W
0.95
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' 'ueh
as tree Stluierel,. hare,. and rabbil' were nOl censu,ed ,n the.. report,. These
om",ioo, "ill have lil110 effecl on lhe specie, di,-ersi" bUI could greml)' aller
con,uming bioma,s and hioma..
Mean My weighl ("hich follo'" mean MY' ,ize) lend, to he Ie" on
humod 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: Krefting
and 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 sIze
di,1Tihulion for .mall mammal•. wilh peak. at the 1"'0 size extrome. (deer
mice and chipmunks)_ Curiousl). lhe same phenomenon occurred in
Piedmont 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"'0
ahernati'-e S1Ialogie. of exploiling seed' rebled to pmchines, of lhe resource.
ChIpmunk, mal depend upon palche. of l",e seed pcoduced in quanlit~ hy
tree'S ,u,,'i"ing fire Or logging. and may al'" rO'pond positi'oly 10 palche. of
shmb 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 and
Ahlgron. 1974: G"hwiler: 1970), Thi. lead' to an ineroa,e in My .ize and a
decrease in energ,. consumed pcr gramme of bIomass as judged from basal
metahali,m Thus lhere i' an Increase in Iho efficieney of energy ulili.ation.
ahhough mOle dalu on Ihe aClual energ) bUdgets of lhese small mammals
could 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 arc
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,t
de"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·dwelhng
chipmunk, mal' also return. posslbl) because 'here arc seed· rich patches to
e.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 stored
in the fore't floor ",hieh are exposed b)' fire_ The carll' dominance of deer
mice 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) transient
e«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 than
P~rom}'scus (Ahlgren. 1966; Slmm~. 1979, Coo•. 1959, Pearson. 1'159,
Beck"'ith. 1954). Vegetation co"er and den,it)" may be more important to
microtines 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 the
lo"-humiditj microclimate of poorl)" '-eget.ted earl~ ,ucre,sional habitat,
(Bendell. 1974) uplains the low post·fire densit) of shrew,.
dominat~d
9.3.4 Small Mammals in tM Far North
Ver)" little ha~ been publ;~hffl on po<;,.dl,turhanec response of sm.ll
mammals 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 be
relati'e1}' slow compared with boreal Silt>.
The fo,e" 'ole (Cleth,ionom,'s gappen). common lhroughoUlthe boreal
forest. i' replae<:d in the Hudsonian hfez<:>ne of "e"ern Canada and ,n AI.,ka
b; 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"•. "hich
colonIZed 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 not
present as a dominant colonizer on burn' (Banfield. 1974: Manell and
Pearson. 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 and
Wingate (1976) reported deer mie<: and C. Tlllil,,, dominant in ,'inuall)' all
habiMs m (he Kluane tegion of lhe southern Yukon (6G-(iloN lalilude)
Mirro"" pe"IU}'lm"irus ,,", foremO>I in ma"h and mcado". "hile II
oecOIlOmus "as dominant in tundra, Vegetation in thi' region i' a fire mosaic
bUI 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 is
pos,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'on of hab'la" as 'open'
or 'closed' and rel~ti"e abundance of deer mice "ersu, C r~lIlus, I lo,..e>or. it
can be inferred thal deer mice dom,nance occurs in ,or; open habitat> that
are earl}' in primaf)' .u="ion (e.g. a grass'fi,e"eod area. a beach ,idge. and
a Drya, d"''''nwndi commUnil}') or in habital' "here lhere is a greater
product,on of seed, and ""rrie. (e.g, a'pen. babam poplar. and ShephtTdla
ca"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.logging
burning 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 in
habital 'lruclure such ~s fallen limbs. standing dead limber. etc .. rould ha>e
Th. R"" al Fir< i" SaTlh,,,, Circampolar Em"""''''
162
"nporlant effec" upon UllliZ<llion h\ mammals (Da\i>. 1977) and hird.
(Hagar 1960), For example. cnipmunk- use logs Or downcd limb, as runways
and lookout', There appears to h.\'e becn lillie COrrdall\e stud\ of eff~l' of
differing tn"" of di,'uroonce,
Ahlgren (1966) compared .mall mammal> On Minnesola jack pIne logged
are3< On "hi,h 'lash" as cilher hurned or lell unburne,!. Srecies composition
On the logged area, resembled that On the control (unloUed) area. though
populaHon! "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 .n
ups"ing in Ibe \'ole', populalion cycle. On lhc hurned arca. decr mICe
dominated 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 the
dC\elopment of .ufficienl CO\'cr and ~ resourtts in the hurn,
Martell and Rad\3n\; (1977) comp.red ,m.1I mammal';n logged .rea, in
Ont.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. and
chipmunks (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 ,n
mme oren habitats or "here seeds Qf colonizing plants are .bundanl. "hile
chipmunks mal resl'clnd to h.bitat fcatures Ii.e sla'h and logs. Or a m"turc Qf
open and brushy arcas,
9.3.6
~'ulu ...
Sludi""
Imp'''' cmenlS could t>.. m.de in future studies Qf 'mall mammal' b\ relating
popul"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 and
Hoo'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 the
rel.tion of ,m.lI-mamm.1 .bundance to habilat ,truClute. Because the
'ariabilit}'. amQng studies. in <mall·mammal sequence and liming is perhaps
rel.ted to ,'arl.t;"ns in ,eget.ti\,c ,ucre"iQn. our predie",'e abilit) depends
upon 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.1
habit.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 more
cOmmon 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 1M
numM. 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<. lboo
correlalion found "'2< no! i""""""'ent ...,th mteroIutbotat
M lhe
pnn,,,,atc QU,,", of both >peaeS d'H"'l) and ~.,.ed diet. (M"CIo$kel.
1975 . .\otorns. 1m).
"'pan''''''
'.-t
9.4.1
BIRD CO.\.I.\.IL~ITlr..s
GOOllraphkall);'lribut;on of Studies
No published sludics of post·fir. lmd «Immunities in the spruce-fir or the
oorcal for.'1 "ere found. and th. follo",ng rcm~f~s ~rta;n 10 dala from
breeding-bird «nSu",. onl), A '-cr) lhOTOUgh stud) Irom earl) post-fir. to
malur. forCSI .tagts ""as r~ntll carried (>Ill (In Alaska's Kena; PeniQ.ula at
6lJ....61<N (Ou,nlan. 1979).
Sescral >ludlC$ <k>cribc bird rommuntlOCS 1-10 )urs after locg,ng. but onl~
on~ fall> ,n lh~ Canadian lif~ zone Al'tknoo ~ •. (1917) .. udocd blrd
romunllle'. In P"""'~,·I",., corrilI<ln In the oat-lllc"~ region (36"1 In
T~nl'lC'SS«. ]';'dlO'~ (1971) ~b«Ithc dfccu of 1Ulde1"Sl0t~~ '~""""-al In
California Sequoia f<.>r'$ (36"3.):"1. Haga' (1960) eumiDffllogcd are~ In
Doupa;..fir fOtC1olS of the northern Caltlonna CO&SI Range (41 .... ). FBlW'Cb
atJd Ohman (1978) ..1Khed 1oggI", effectS In IlIUcG rorufer (Doup·fi,.
ponikrosa pmc) lOfnl In AfIlOlU (33'1'). Webb rr •. (1977) .. udoed
northern .... """''00<1 fornl'l at four Ie'els 01 cull,ng '"'~""t~ in lIOe,," Yorlt"s
Ad'rondack \Ioun'a"" (-U '). atJd Knsk' (1979) .. udiW bird popuJallom
in 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 In
the 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. an
outstanding 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
roount~ill!
of
cenlr~l
Wl!'Sl Virginia (2&<N)
~nd
(1~9,
1952). from the
extreme northern
~hme
and Sail (1957) ftom Grand Telon t'allonal Park in
Wyoming (43'Nl· Thre. studies from lh. ITaosition life zone. norlhern
(~rN). re,pc:C1i'.I~.
hardwood 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 sluJies
eOnj;idered range from 34" to 41·1' latilude. ,n Ih. oak-hickor~ "oll,
beech-maple regions of the eastern United Slales (Aldrich. 1943. Shug.n
and James. 1973: Brewer. 1951\; Kor. \%1;), The last tWO concerned
succe,sion On mip m,neS.
9.4.2
AggrtgaleCommunilj
Properl~
Table 9.3 ,how, succession,lt,.od, in bird-community properties. a\'eraged
Over ,,-,'·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 eategorics
by Tramer (1969) ... ho analysed hundred' of cen,usc, 'aken from Am",c""
Birds and Auduoo"'. Field ,"mrs. Mean .. eigh' and consumin~ biomass p<"r
uni' "'eig)" also exh'bit si~nifieant 'rend, (P < O.(J]). The la,e-w,cessional
inereasc in species richncss rclati'-e to species di\'ersity has !><-en noted and
di><: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 a
ma,imum b)' the sapling stage, This parallels trend' in plan, production and
fohage 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) and
Gorham. 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,ial
heterogenei,y in 'he ma,urc forcst (Shafi and Yarranlon. 1973, Bormann and
Liken,. 1979). The rela,ion,hip bet ..'.. n bird ,pecies diversi,) and foliage
he,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'h
primar} product;'it) di'ided by canol') ,olume. "h'Ch mal cominue to
increase ahcr primary' production has peaked,
In contra,t to 'he summa!)' in Table 9.3. dat. for indi'idual studies ~i"e 'he
impression tha' there mal bc significant trends ,n numbe" and b,omass "ilh
sU=5\ion. Spindlcr and Kessel (1980) stud,ed replica,ed samples of black
and .. hilt spruce stands (7 eaCh). deCiduous aspen Or birch woods (5). m,xed
deciduou.....oniferous stands (~). and shrub "'p<"' (7), The)' found a decrease
from ,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~'
,
Spe.,e; ".hne"
,
W
,
Species d,,'e""j
:>"mbcr of!,,'" per
lOOha
Con'uminj! biorn."
(.m'IOO ....l
K.I.,,,. m...
demand'
bo".
V.l)
•.,1,
'",
200
SerondiI'}
no
t",.",
17,90
7.48
" ,m" ,m"
926
Iterb
~
1l!17
17.116
11.03
~~I
Sal'hn~
17
1~
M.'ure
lore"
~ l-ki
10.17
" "
29..
," ,
""
, m • " " " m"
"" • '"
", 137, "', "'
'" "
, 2117 .. 6132•
, ~", ""• 1321, •
",, :.53
'"
28.2
37.1
330
"0
1,2
•" ", '", • , • "'
, 1.,,-50 12,1. 1035 1021 9.0S '"
•" , "~, 2 78, 22.., UO• H2•
,~
OM
.,77
U
~
~SO
,~
..
9.~:
276
$10
6756
,~
,~
~118
>-l:l6
~,
~,
36~
36,~
-,2
$,6
o,5~
,\1o.n bod~ ,if<'
'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",,'
"Stud;<, tnN
ha) and ,n consum,ng bl(lmar.s (liS(,. 699. 555~m HI) hal. Nuh of "h,ch are
SIgnificant (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 1M
gm 100 h•. ,ligh,ly higher Ihan for deciduous "and, Tramer (11'6\1) abo
reponed 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,ne
th'cket §lage of a p,ne fOIl"1 sere,n Finland. and 'he '.me effect appe." in
posl·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. and
the.. 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 the
sapling 01 secondar; for.'t stag'" in other seres (e.g .. Johnslon and Odum.
1956: Shugan and hrne'S. 1973: Theberge. 1976). sOl"'rimposed upon a
general Incrcas,ng trend. These oornmun"l "bollienech" are probably" mOre
like" 10 de,-dop in ronifer slands .fler fire and coning: oon,[c,.., >Cern p,one
to establi'h a thick 'land and to exclude 'hrub> and herb> b)' root and light
competition. especiall; ;n the boreal forcsl where the common deciduo'"
lree, hJW high light rC'luiremcnK Since tit"." no pronounced decrea,e in
production 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 be
a 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 be
no 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"c
metaoolic 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) ,ile
deerca"" through succe"ion (Table 9.3, P <: ,WI). Larger bird, ",II have
lower ba,al metaoolic rate> pcr u"i, biomas>. '" lo"er rela,i,-c metabolic
dcmand "'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: mcan
Ilod) ,i,e wa' lo"er in lhc ma'ure 'tage a, compared with ,he ncx! carlierone
in ,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. (Table
9,~(a)). 'h'" proportion of ,pecic, riehne..., accoun'cd for h) nonpasserine,
,,", con.iSlentl) grea,er in early .uccc<>ional (berb and ,hruh) ,'ago> than in
rnaturc 'lag« (fourtcen Out of .i"cen Ca'"""S. P - 4),002), Openne". or
habi'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 ,hrub
or >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 and
pos,-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
".ge
P"'Hhm..
\e~'
e'TI~'
(.,
,
<
•
•
•
'$opl,"~
• .o
lte,b
Shrub
33%
'"
"~.
16',,'
3~'"
n'l"
s..1>hn~
"" "
•
'"
seronda~
~,
""
, •" " """
'"
" '"
"'''<''''''r~
r"",,1
".~,
rom.,..d
fo,."
""
~~.
'"
~1a,ure
open t>lacl
Ime"
~-
"y~.
3~.
"
,
l~'"
'"
'"
,
'"
s<l'<I-eaters in Ihe'" cases a,e f tingillids. although I/IlwO case< in the sou,hem
United States quail dominale, All ftingilhds ....p' the chipping Spart(WI
Wete ,ounled a, hreeding_season g,anivo,e,. "hi,h "a~ a ne.e.,a,y expedient
but 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 and
Jam.,.. 1973: Kar. 1968: 8<><:. and Lynch. 1970: Hagar. 1960). The trend 01
decrease 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 had
Iheir biom • ., ~tal'ed constant. granivore consuming biomass decreased
Ihreelold (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 and
posl·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). and
Quinlan (1979). and. for the most part. exhibit changes predictable from lhe
dC\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,["
dec,d,,,",, ''''c,''
S,,,w,,,,'n;,1
~"sio"
Com/_m", fo",,,
AI,,-,~.
1l""''''''Y
","
,,.
~,1I7
h.l~
~27
272
.HI
2X.\
II 'O\l)
') .. ~)
1l..'\.l1J
7.2X
7'""1
, 71J
oJ'llI
SI)
,~
~,
M
,~,
27'12
,-"!2
WM
6,4Y
Comic,
Wyonllng
W"", Virgin'"
.\I""",
~h,,,d
M",u,,,
"""k,
,
,
SD'
M
Yu~,,"
Shlllh
,
SD
,
SD
",
"
"'I~ing
SD
Y,48
M
121 I
95.11
S,B
W
~,
,.,
n,ll/!
."
~"""'"'Y
'"
W
M
,,~
.124.
1.\ ..15
!!It1
,,~
'M
1U.~!.8.XJ
"~
635.1
1067.S76
76tll).44~7
'1""""
-,
'"
:!II!
Rd"",,,,,'
Sp.,"lIcr
K",,,,I
.".1
t 1')l\I,"
11...·l><.~,·
( 1~7h)
510.1
IHI<
m
""
"
>
...>~
,,
,
0
",l
0
s",l (IYS71
10.87
H),II')
,~
0,,,,,, hl"d
M.,'u,,,
lore"
~.7X
.....,.1
7 I,j
"""Ier
• .00
';(01, I
,~
,
,,,,~,.
ll<,',<1",,",
Comm,,""y
,,,Ie
St"".,,, ,,,,,,
,\I,ln.:h
( II'/<))
5.'12
n~
2.171'
SI""." a,,,1
,\Idm-h
( I~<;l)
0
i
,
.[,
,
•
Taole Y.S
,"
em",1
rom""",,')'
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EroI"'I~mJ
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•• "ilh
the "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 lhe
malure 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 "'pes
repre"'nled. and hence thc possible "arictl of difference' in habital "ruc'urc
bet"een malure and "CO, cry stage'. 1'-0 earl) post-dIsturbance studlC< from
lhe 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 to
d"',duous tree to conifer tree. there appears to be a fundamcnlal differcnce
bet"een bnrea! and mid-latitude rcgiom. characterized b) conifer ""US
dc"<'idu"us dimas forells (Tahle 9.S), Species d,,'ersi'y' . numbers. and biomass
of 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 the
10" 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,,1
di' .r"l) ,,,,d 'recie, di' e,,"y of lhc vegetation. "hich maY' be greater in tall
shrub ""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. or
conifer lype' (Spindler and Kessel. 19S0)). is generally accompanied by
incre""ng 'recios d,,'ers,ty'. number>. and e"nsuming biomass It also
appear>. from the r",ull, of Spindler and K"""I (1\180). that hoth agc "lth,n
One vegetation t)·~ and differences among tH'" account for signifieanl
po"ions of the 'arianee in bird communlt} pro~rties, WhIle man) ~peClCS
are ,hared among ..getation lYpeS. (from lO% to 30% of the .peele> "ere
found In onl) one ,'egetat;on t)'~ (Spindler and Kessel. 1980» since only
small areas "'ere cen,used. this dissimilamy rellee!5 relat;,'e abundance more
than 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 "'ilh
Ihe siudies in Table 9.5 since the dala "'ere based upon frequency of
encounler "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 cowr
peaked 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 at
Ihe 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 in
oommunil~ 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 be
markedl~ 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 of
Ihe 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 Ihe
southNn Yukon b"d communilie' he .Iudicd. in romp<orison ",ith mid·
latilude regions. Similarilie. among hi. sampl... are ob' iou., but he presenled
no qaanlllati\'C comparison "'ith other region,. so one i. pro'-id~d here (Table
9,6). 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)' would
cxh,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 Ihe
rather 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 an
inexplicably high ,'alne. much higher e"en Ihan Spindler and Ke... r. data
from 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 nonhcrn
species 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.
.. milo.')"
"
utilude
(':-)
"
""
"
""
"
"
"
",
,
~
,
~
"
"
SI."", oomp.,.d
T..,,, ..ill"",' ,'."u' ,..."
I"'plar
T.11 ",ub ,·e"u, ~duoo,
Tall cto..ed bru'" ,'.,.,'"
hard..<>Od
S<'ub me.do...."u,
nOllond ..pen
Iloj: .lllrub ,'."'''' >«ondary
fore"
Earl)' .h,ub "'''u, mJlore
for."
Open ...phn! ,.,,'" me".
fore"
Xeric 'hrub "e,,", oak
fore"
2O-)Ur g......shrub .."u,
<>OK hick"'}' l<>r<>'
"Th< inde, oI .. md.n,~'"",,~
~' ..
(2
~
Rd.,e"".
Th<berge (1976)
$pondle' ,nd K..... I(1980)
Hoapo""n (1965)
$011(1957)
Aldn<h{I9-l3)
Kar(I96/j)
SlIugan and lame> (1973)
Odum (1950)
John""n .nd O<lum (1956)
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,)': Table
9.S sugg'''ted 'hat within·habita, species d;, ersit) w.s lower on 'he sub.re,;c
sites, 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 Alaska
black 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 .pecies
composi,ion is less important in thc NOrth, Ho"e,'er. Table 9.6 refcrs onl)' 10
deciduou~ sores. and <peci.,turno,·cr can ~ much grealcr in a deciduou.·toconiferous sore (e,g .. Haapanen. 1965: Spindler and Kessel. 1980): the
fire·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 in
the herb stage. and lower in the shrub and sapling stage,. than in mature
forests: species dlYc"ily is hIgh and increasing through Ihe shrub 'lage. and is
depre,..ed in the sapling slage, relati"e to malure forest These trends
undoubtedl~' reflect changcs in lhe ground la)'er ,·egeta.,on. "hich pro,ides
food and coyer for lhe ,mall mammals of Ihe fore'l floor. Had tree squirrels
been Included. perhaps numbc". biomass. and consuming biomass "<Juld
ha.. exhibiled a continual increase through su=s'ion.
A, e.peclcd. for bi'd "'mmuni'ies lhere IS a broad trend of increase in
species richness. species di'·ersit~·. numbers. and consuming biomass. and.
dec rea", in biomass through succession. bUl thc trend " nol monolonlC. For
bird 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 mature
stages "ere not significant when a'eraged among sludies. There "ue
sometimes significant changes among these stages in Ind,,,dual .tudie•. bUI
there "as also tremendous ,a,iabilily among studi.. and usually no
replicalion 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 slight
decrease in the sapling or secondary fore'l stage and a slight increase again in
the mature foresl. In lhe 'hrub to malure foresl slages. ,,'hen numbe~ and
consuming biomass ".re essenlially slable. ,pecies di"e~it) and SpeCICS
richness 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 appear
to bc the firsl altempl at a unIfied SU'ye) of lhis kind.
The conclu'ion of Bendell (1974) lhal population densilie! do not dllf.r
between 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 consuming
biomass are less for bird' (Table 9.3) and greater fo' 'mall mammal,. rcla'i' e
to mature fore'l Species di'e~ity is clea,ly greater for 'mall mammals and
less for birds earl~' in succe"ion. wjth mammals declinIng and birds increa,ing
in diH'S11) In mid·succession. as the shrub and tree lay'~rs develop. It IS
especiall) Int~reSllng that changes afler the shrub slage are relali"ely small in
lermS of aggregat~ measures. although considerable species lUrno''Cr i. still
occurring.
Inler·dass competiti,,~ O\'erlap i' probabl~' greatest bc1W~~n bird and small
mammal granivores in early succession: bolh lhese groups decr~a>e through
succession 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' and
prima'} production aher disturbance dircctl}' cause increased small mammal
'"
and bird populations early in iu=s,ion. On the qualilali,e c,",denee
a\'ailab\c. the jnil;.! high numbers of small mammals CQuid result from
grealer ,..,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-fire
and ""ral habitat., Firsllhcrc is a con",nsu, (Cuni, and Ririe)'. 1975) about
Ihe imponance of lhe herb and ,hrut> laicr of forests and woodlands 10
hruding_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 of
bird spee;"'; .ichness in campground are .. "'bich are both forested and
park·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 in
oth•• 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) and
lik.ly to occur in bor.al ser... discussed h•••. aboUl 50% art limited '0
nesling 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,'e
4 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 nesl
up 105m abo"e lhe ground, It seem, lha' neslIng and foraging guilds 'hould
~ con'idered simultaneou,ly as interacting nich. dimensions. ralher lhan
separately. as i. SO often done, It also appea'" lhal full use of lhe foliage
p.ofile b) b•••d,ng b"ds could be IImit.d bl ,,'ail ability of ne.ting SileS. i.e,.
b~ ,hrub co"er and profile. The aUlhor i' nol a""ar. thaI succe"ionallrends
and 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) or
analysed ,n lh. joinl niche space of foraging '."us nesling siles, The effect of
d.creased di'e"itl and numbe .. somelimes appearing dUfing the .anopy
elosure Slage in mid-sucees'ion mal' ",.n be e.plalO.d by lh. ehm,na"on of
under<lorel' herbs and shrubs al lhal lime. Experim.nlal manipulalion of
shrub co"er and palchiness could help leM h'-'POlhcse. aboul limil.lion by
ncs.inll silcs 'ersu, foll.ge profile and p"mar~ p,oduclion.
A second line of inquiry concern, spalial hclerog.neity of \Cg.lalion a, il
affecls bolh num~rs and ,pecie. di"e"ill (Ro.h. 1976), Two sludies of
disturbed areas suggest .natogies 10 post_fi", ".gelation. Fif'lt. Hooper f' 1'1
( 1973). sludied de"eloped campground ,iles in lhe soulhern Appalachian
Mounlains and found that bird ,pecie. richness increased ",'ilh admixture of
coniferous 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 the
dumps in addition to rotal cover was an imponan, factor; larger. denser
dumps 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 and
diversil)' 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 the
equi'-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 along
one 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 lhe
same lo,al area. forman el 01. (1976) dISCUS' lhese re,ult, in 'he lighl of lhe
lheo') of island biogeography and .peeies-area curs'es. Our specifIC intereSl
here is lhal degree of post-fire horizontal spatial helerogenell)". Or '-egelation
pale 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 bird
specie, di"ersity and numben. ailhough venieal dIstribUtion ha, been s'ud'ed
extcn,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 by
correlalions wilh foliage heighl di'-ersity, funhec application of these
method, in posl-diS'urbance and successional studies is highly deSirable.
especially in the Iighl of the high dcgree of patchiness (and the con.inuing
change, 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 plant
species. primary production. habitat 'truClure. ammol food resourceS and
animal populatio,," 'hould be undertaken, In any son of 'ludy. repllcarion is
neeessar~_ E,penmemal method,. such as pro'ioon of or remO'lal of food
and co'.... and u~ of herbIcides and fenilizers. ,,'ill pro"ide more insight inlo
controls owr community de"elopment than ,,'111 repeti'ion of the general type
of studie, cited here,
9_6 AC"-,,OWLE:[)G[.\!E:-'TS
Preparalion of Ihis cbapler was supponcd by NSF gram DEB·78-11594. 'The
structure 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""ng
rollfague, for their insighlS. although the) should not be held accountablf (01
hi. "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 of
lhcir dala. and to an anonymous rc,'i.w'er. who pro,'idcd many u",ful
rommenl•. 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"''''lj
in '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" 01
Ir.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 10
eyel., 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<> 10
hobn.t eh.nif' In " ...tf'n O'f&"". ,n 11.e. Blact (<<I). ""ildli/~ aru1 Forni
M.~"&,,,,~m i~ Ik~ PaCific NCr/h·m. SclIoo) of Fore"')'. Orego~ Slate
Uni"fn;'). CO" .Iho. 236 1'(>.
Bock. C.E...t><! LYll<h. J.F. (1970) 8r<edlnl b"d popula""n. of burned and
unbIJmed ronlfer lore." in ,he Sie". Se'-.da. CQJld",. n. 182-1119,
Borm.nn, F.H. and Liken,. G.E. (1979) POII"~ alld Pmens i~ •
E<: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!IU
and 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 on
nona.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_
J""."d
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>.l
Faabo'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}-9J6
Flack. 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
Am. .Ilodl.
o\'~,. (;$. 67-88.
Godl,.). E. W, (I\lt\t» Th~ B"'/h of Canada. National ~lu.. um of Can.d•. Otta" •
,1".,,-..
• 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
nd nitro~n trea'ment> in. >bonpas> puin. "",,)-.tem. J
communi')'
M.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.
'0 ....,...
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"nee
of "'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 '0
plan' '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«nttal
lIIinoi•. 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...er
MIChigan. 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. 71
22pp.
"'1"",•. 8,,\1 (911) Re,pon>c of br.e<i.n~ b"d I'Ol"'lallOll' to hab,t., ella.g., ,n a
Gi.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 wKre"'. 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" of
allorestall"" 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. J
Anim Ero/.. 8 . 277-285,
l.<>ck. D.. and lack. D. (1951) Fu"he' ehan~... ,n bi,d hfe .. used b~' affore"a,i"". J
Anim. 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
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
di\'.""~'. Am.
,,,I.tI,,,,
.. \18 • .l81_391
~.-u,
.pea",
.\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, after
e~"""lIm8 of 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'a
ReglO1l. 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.", ..d
Uni,·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 and
Win,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. 103-192
Pe."",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-2S5
Pre"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~ bird
populot;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"lem
Manitoba. 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 inleriOT
AI.,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 'pnlce
region of lhe ceolrol Appalo<h,a",. EcoIOK>__ .10. 7S-81
Sle"an. R E . . .nd Aldrich. l,W, (1952) Ecological .ludi.. of breeding blrd
populaM'" 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 and
cleaHu' (Ioucd) h.bita", Can. J. Zoo!.. $1. 1636-16-18,
Te'ter, J.R, (I96S) Effect. of a ronlrolled bum on
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,
"".11
""-""
T,,'i•. L.. Jr (19S6b) R«pon", of """II m.mmal, 10 iouJng of Dougla,-fir. J
Mammal __ 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__ $-I.
1346-1356,
Tr.mer, E.J (196'J) Bird .peei.. dll·."I,y: componenl' of Shannon', formul •.
EroIQfY.
5(1.
'fiJ..Jm.
,.,
Ud\·"dy. M.D.F. (1957) An ~,·.l"atio" of qu.n'ila';" ,,,,,lie. in b",l<. Cold SP""S
Hu,/)ar Sj'mp. Quan'. BIoi.. 22. 301_311
Van Cleve. K.. Dyme... C.T".nd Vierc,k. L.A, (l980) Notrien' cytling in interior
AI..ko flood p1ai",.1><1 its Telation'bip '" ,.~ne,"lion .nd 'oMequcnt fores.
dc,-elopm<nt. in M ,\,,,".yand II. .~1 Van "'Idh"''''" (ells). For'" Rq"'t'Qtioll
01 Hi&h LI"irudts.. US For. Sc"'" Gen. T«!I. Rep. PNW-l07. P!'. 11-18,
Vi,o....,k.
P.~'
and R.i ..... W.A (197S)
Ero<y~,.m
OII<CC...,., and "o'nenl
ro'cotion,. h)-po<IIesi', BioS<mKt. H. 316-3!!1
Webb. W,L.. Behrend. D F •• n<1 ~,""". B. (1977) Effea ofloWng on sonlb"d
populalions ,n I ""rlhern h..d....ood fOf.," 1\ ,Idl. Morn>g', !!. 35 PI'·
W.in, R. W. (197$) 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.