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Rangi e , 35, (1) 2015
Composi ion o la e summe die by semi-domes ica ed eindee (Rangi e
a andus a andus) in di e en g azing a eas in no he nmos Finland
Pauline Beza d1, Sophie B illand1 & Jouko Kumpula2
1ONIRIS, Nan es-A lan ic Na ional College o Ve e ina y Medicine, Food Science and Enginee ing, FR- 44307
Nan es cedex 3, F ance.
2 Na u al Resou ces Ins i u e Finland (Luke), Ecosys ems and ecology, FIN-99910 Kaamanen, Finland (Co espond-
ing au ho : Jouko.K[email p o ec ed]).
Rangi e , 35, (1), 2015: 39-52
Abs ac : This s udy aims o in es iga e he die composi ion o semi-domes ica ed eindee (Rangi e a andus a andus)
in la e summe in di e en kinds o g azing condi ions in no he nmos Finland. The composi ion o die by eindee
was de e mined on he g ounds o mic ohis ological analysis o eces samples collec ed in ea ly Augus in di e en sea-
sonal g azing a eas (win e o summe /yea - ound g azing a eas) in h ee eindee managemen dis ic s. Al hough he
p opo ion o di e en plan g oups a ied be ween he s udied dis ic s, he quan i ied g oup o g ound lichens (which
also con ained small amoun s o mush ooms) was he mos abundan , a ying om 33.0 o 46.4% in he analyzed sam-
ples. In gene al, he e we e signi ican di e ences in he p opo ions o lichen be ween dis ic s, bu no be ween g azing
a eas. The p opo ion o lichen in samples inc eased signi ican ly when he amoun o lichen pas u e a ound a sample
si e inc eased. The p opo ion o dwa sh ubs and lea es in samples a ied om 24.9 o 37.9% and di e ed signi ican -
ly be ween dis ic s, bu no be ween g azing a eas. In he same way, he p opo ion o g aminoids a ied be ween 20.9
and 36.2% and di e ed signi ican ly be ween dis ic s and also be ween g azing a eas. Highe amoun s o g aminoids
in eces we e obse ed in summe /yea - ound g azing a eas han in win e g azing a eas. Finally, he p opo ion o b yo-
phy es a ied be ween 2.9 and 6.5% and was signi ican ly di e en be ween dis ic s, bu no be ween g azing a eas.
An inc ease in old and ma u e coni e ous o es a ound a sample si e signi ican ly inc eased he amoun s o b yophy es
in samples. The esul s indica e ha eindee adap hei summe die composi ion acco ding o he a ailabili y o ood
plan s. The esul s also show ha when eindee a e allowed o selec hei summe anges eely, eindee end o use
lichen pas u es in ensi ely also du ing summe , which causes a conside able educ ion in lichens due o g azing and
ampling. The e o e, a p ope seasonal pas u e o a ion sys em o p o ec lichen pas u es om g azing and ampling,
om ea ly sp ing o la e au umn, is an essen ial pa o sus ainable pas u e use in eindee he ding.
Key wo ds: die ; no he nmos Finland; eindee ; Rangi e a andus a andus; mic ohis ological analysis.
DOI 10.7557/2.35.1.2942
Rangi e , 35, (1) 2015
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40
In oduc ion
Semi-domes ica ed eindee usually o age in
la ge pas u e a eas, whe e hey change hei pas-
u es acco ding o hei seasonal g azing pa e n
(be ween summe , sp ing/au umn, and win e )
(S een, 1968). This g azing pa e n enables
semi-domes ica ed eindee o ob ain o age
mos sui able o each season, wi h high con-
en s o ca bohyd a es in win e and p o eins
in summe (Albon e al., 1992; Bjo k oll e al.,
2009). In summe , semi-domes ica ed eindee
can use he pas u e landscape o g azing ee-
ly; while in win e , g azing o eindee is mo e
con olled by eindee he de s. The summe
habi a p e e ence o eindee depends mainly
on ac o s connec ed o insec ha assmen , eg-
e a ion ypes, and plan g ow h (Ska in e al.,
2008), al hough many kinds o human ac i i-
ies can also dis u b o aging (An onen e al.,
2011).
The die composi ion o semi-domes ica ed
eindee a ies seasonally and is dependen on
he quali y and a ailabili y o di e en ood
plan s on pas u es (Bjo k oll e al., 2009). In
summe , eindee ood con ains a la ge a ie y
o he bs, g asses, sedges, ho se ail, dwa sh ubs
(e.g. bluebe y), lea es o deciduous ee, mush-
ooms, and lichens (Wes e ling, 1970; Sulka a
e al., 1983). The a ailabili y o g asses and
he bs is ela i ely high in mesic and submesic
o es and mi es, and he e o e he use o hese
pas u e ypes inc eases in summe (Kumpula e
al., 2007). In ensi e use o mos sui able sum-
me g azing a eas is connec ed o he eindee ’s
need o ul ill hei equi emen s o p o eins
and mine als om g een plan s du ing a sho
summe season. Du ing summe , eindee se-
lec a g ea a ie y o plan s and usually con-
sume only g owing pa s o plan s o hei new
g ow h. In no mal g azing condi ions, eindee
ha e no di icul ies du ing summe in ul illing
hei nu i ional equi emen s: p o eins, mine -
als, and o he necessa y ace elemen s (S een,
1968).
In con as , du ing win e , eindee need di-
e a y ene gy o locomo ion, digging, and he -
mo egula ion (S een, 1968; Wes e ling, 1970).
Reindee lichens (Cladonia spp.) a e he e o e
an essen ial pa o he die o eindee in
win e , as hey con ain a lo o easily diges ed
ca bohyd a es and also p omo e he diges ion
o mo e ib ous ood (Nieminen e al.,1989;
Aagnes e al., 1995). Reindee lichens can also
o m an impo an pa o he die in sp ing,
when he snow has al eady mel ed bu g een
plan s a e s ill sca cely a ailable (Opho e al.,
2013). Reindee can also use lichens in sum-
me (Wes e ling, 1970), bu he e a e e y ew
s udies a ailable on he consump ion a e o li-
chens by eindee du ing summe . I eindee
a e allowed o g aze on lichen pas u e a eas
du ing he summe season, his also conside -
ably a ec s he condi ion o lichen pas u es
due o g azing and ampling (Kumpula e al.,
2011;2014).
One o he mos impo an objec i es in
semi-domes ica ed eindee he ding is, how-
e e , o ensu e su icien a ailabili y o sui able
pas u es and ood plan s o eindee in each
season, since seasonal nu i ional condi ions,
dependen on he a ailabili y o na u al ood,
a ec he ep oduc ion, milk quali y, cal bi h
weigh , g ow h, and mo ali y o eindee (Rog-
nmo e al., 1983). In o e g azed o by o he
means de e io a ed o insu icien pas u es, he
nu i ional needs o eindee a e no ul illed.
Especially in Finland, supplemen a y win e
eeding o eindee has inc eased due o he lack
o na u al win e ood, in o de o keep he d
p oduc i i y high enough (Helle & Kojola,
1993). Especially he condi ion o lichen pas-
u es has de e io a ed ma kedly du ing ecen
decades. Since summe g azing may conside -
ably a ec he condi ion o lichen pas u es, i
is impo an o know which ac o s inc ease he
use o lichens by eely g azing eindee du ing
summe , and how much eindee use g ound li-
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Rangi e , 35, (1) 2015
chens in he summe in di e en kinds o g az-
ing condi ions. This kind o in o ma ion will
p obably help o de elop pas u e use sys ems
in eindee he ding in such a way ha he mos
impo an win e ood esou ces, lichen pas-
u es, a e g azed as op imally as possible.
The aims o his s udy we e o cla i y how
much lichen and o he main ood plan s semi-
domes ic eindee (Rangi e a andus a andus)
use in di e en kinds o dis ic s and g azing
a eas in he la e summe season, and whe he
he ype o pas u e a ea and he a ailabili y o
di e en pas u es a ec his selec ion and in-
c ease he use o lichens in summe . Fo his,
we analyzed he die composi ion o eindee
in ea ly Augus by using eindee ecal samples
collec ed in di e en g azing a eas in h ee ein-
dee managemen dis ic s loca ed in no he n-
mos Finland.
Ma e ials and me hods
The s udy dis ic s and hei eindee he ding sys-
ems and g azing a eas
The ecal samples o analyzing he composi-
ion o he die o semi-domes ica ed eindee
in he la e summe season we e collec ed in
ea ly Augus in di e en seasonal g azing a eas
in he Pais un u i, Muddusjä i, and Salli aa a
eindee managemen dis ic s in no he nmos
Finland (Fig. 1). One o hese dis ic s (Pais un-
u i) is loca ed in he moun ainous a ea whe e
he e a e no sepa a e seasonal g azing a eas.
The lack o a p ope seasonal pas u e o a ion
sys em allows eindee o g aze lichen pas u es
also in summe . The o he wo dis ic s (Mud-
dusjä i and Salli aa a) s udied a e loca ed in
moun ainous and pine o es a eas, and in bo h
o hese dis ic s a la ge win e pas u e a ea has
been sepa a ed by pas u e o a ion ences. The
win e s ock densi y (acco ding o he maxi-
mum allowed numbe o eindee ) in Muddus-
jä i is 2.6, in Pais un u i 2.2, and in Salli aa a
2.6 eindee /km2 land a ea, espec i ely. In he
Muddusjä i and Salli aa a dis ic s, which a e
loca ed in bo h moun ainous and pine o es
a eas, a dis inc win e g azing a ea has been
sepa a ed om o he seasonal pas u e a eas by
a pas u e o a ion ence. The main ege a ion
ypes in hese dis ic s a e o med by di e en
aged pine o es s, as well as moun ain bi ch o -
es s and open hea hs abo e he ee line. In
he Pais un u i dis ic loca ed in he moun-
ainous a ea, open hea h land abo e he ee
line and moun ain bi ch o es s domina e he
ege a ion, and eindee can g aze i ually in
he whole dis ic a ea all yea ound. In Mud-
dusjä i and Salli aa a, a ew eindee always
s ay in he win e g azing a ea du ing summe ,
which enabled he collec ion o ecal samples
om eindee in hese win e ange a eas, as
well.
Fecal sample collec ion
The ecal samples we e collec ed a e obse -
ing de eca ing eindee o by ecogni ion o
esh eces on pas u es. The obse ed eindee
we e om bo h male and emale/cal g oups
and p obably he unobse ed eindee ha le
esh ecal samples we e also o bo h sexes, al-
hough i was no possible o iden i y he gen-
de o hese eindee . In o de o ob ain esh
ecal samples, he age o eces was de e mined
om odo , mois u e, and he p esence o mu-
cus. The loca ion o each sample was de ined by
a GPS de ice. The numbe o samples collec ed
was 20 in he Muddusjä i dis ic (7 samples
in he win e g azing a ea and 13 samples in
he summe /au umn g azing a ea) and 25 in
he Salli aa a dis ic (10 samples in he win-
e g azing a ea and 15 samples in he summe /
au umn g azing a ea). In he Pais un u i dis-
ic , 25 samples we e collec ed in he summe /
yea - ound g azing a ea. A e collec ion, he
samples we e s o ed in plas ic bags in a eeze
(-20°C).
The p opo ions o he main pas u e ypes
and g ound lichen biomass ( o med mainly by
Cladina spp.) on lichen pas u es a ound each
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Figu e 1. The s udy dis ic s in he eindee managemen a ea o Finland and hei seasonal g azing a eas used in
summe /yea - ound o only in win e .
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Rangi e , 35, (1) 2015
sample loca ion wi hin a adius o 3.0 km
we e de e mined by A cMap 10.0 using he
eindee pas u e in en o y da a om he yea s
2005-2008 (Kumpula e al., 2009; Colpae e
al., 2012). In cases whe e pas u e o a ion o
ano he ence p e en ed eindee om g azing
ce ain a eas wi hin a ci cle a ound a sample lo-
ca ion, hese a eas we e emo ed om he pas-
u e analyses. We used his size o adius a ound
a sample si e o iden i y all po en ial main pas-
u e ypes a ailable o eindee du ing 2-3 days
be o e sample collec ion.
Handling o ecal samples
The samples we e p epa ed o analysis using
a me hod adap ed om Hansson (1970) and
Vi o and Sulka a (1985). Each sample was
hawed and hen c ushed wi h a mixe , and
insed wi h unning wa e h ough s aine s
wi h ape u es o 0.125 mm and 0.250 mm.
A small quan i y o sample sepa a ed be ween
hese s aine s was pu o one minu e in 70%
alcohol. Again a small pa o his sample was
colo ed by 1% me hylene blue o abou 15
seconds o aise he con as o cell issues and
hen o help o iden i y each ype o cell is-
sue. The colo ed sample ma e ial was insed
wi h unning wa e o one minu e and hen
le o d y o one minu e. Then he sample was
pu in 70% alcohol o one minu e and quickly
insed in 96% alcohol. The inal plan ma e ial
p epa a ion was hen sp ead on o a mic o slide.
Some d ops o Eupa al moun ing medium
we e added, and he mic o slide was co e ed
wi h a co e slip. F om each ecal sample, h ee
subsamples we e p epa ed in his way.
In he ea lie s udy on win e and sp ing die
composi ion o eindee , only one 0.125 mm
il e was used o inse c ushed samples (Opho
e al., 2013). In ha s udy, no plan pa icles
we e obse ed o be emo ed om win e and
sp ing samples using he 0.250 mm il e . This
is p obably because he s uc u e o win e and
sp ing eces is less ib ous and con ains smalle
plan pa icles han summe eces. This di e -
ence is due o he eindee die composi ion in
summe and win e (S een, 1968) and he di-
ges ibili y o plan s (Thomas e al., 1980).
Howe e , in o de o compa e he di e -
ences in he il e ing me hods o summe ecal
samples, 15 samples ( i e samples in each dis-
ic ) we e also p epa ed using only he 0.125
mm ape u e il e . In his way, we we e able
o compa e he di e ences caused by hese wo
il e ing me hods on he ela i e p opo ion o
each plan g oup in he samples.
Mic ohis ological analysis
The sample slide analyses we e pe o med wi h
a mic oscope (size o enla gemen x200). In
each subsample, i e di e en iews we e an-
domly chosen and s udied. In each iew, he
ela i e p opo ion o main ood plan g oups
was calcula ed in he g id, which had 25 c oss-
ing poin s. The plan pa icles we e iden i ied
in hese c ossing poin s and de ined in ou
classes: g ound lichen, g aminoids (g ass and
sedges), dwa sh ubs (Vaccinium spp., Em-
pe um nig um) and lea es (mainly willows and
bi ch), and b yophy es (mosses). The p opo -
ion o mush ooms could no be es ima ed
sepa a ely in he samples, as hey we e di icul
o iden i y, and due o a lack o a ailable mush-
ooms in pas u es a ha ime, he amoun s
we e also p obably e y small in he samples.
I is he e o e e y likely ha a ew mush ooms
in he samples a e included in he lichen class.
Due o a e y d y summe (Finnish Me eo o-
logical Ins i u e, 2013), he e we e no o e y
ew mush ooms obse ed wi hin he sampling
si es. Howe e , only wo weeks a e he ac ual
sample collec ion, he e we e al eady many
mo e mush ooms a ailable o eindee .
S a is ical analysis
S a is ical analyses we e based on he calcula ed
mean alues o plan s in each sample (a e age
o 15 windows) de i ed om mic ohis ological

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44
analysis. An independen sample - es was used
o analyze whe he he e is a signi ican di e -
ence be ween he p opo ions o plan g oups
in wo di e en il e ing me hods. Mul i a i-
a e ANOVA was i ed o de e mine whe he
a signi ican di e ence exis s in he p opo ion
o plan g oups be ween he eindee he ding
dis ic s. Co ela ion be ween di e en plan
g oups was analyzed using he Pea son co ela-
ion coe icien .
A mixed linea model (GLMM) was used o
analyze he e ec s o di e en backg ound ac-
o s on he p opo ion o di e en plan g oups
in ecal samples. These independen ac o s
we e s udy dis ic , g azing a ea, and he ol-
lowing pas u e a iables: p opo ions o lichen
pas u es, a bo eal lichen pas u es (ma u e and
old coni e ous o es ), dwa sh ub/g ass domi-
na ed pas u es, mi es, and ba e moun ain a eas
a ound he sample places. In GLMM, summe
and yea - ound g azing a eas we e handled as
one g azing a ea, and he win e a ea as ano he
g azing a ea. The es ima e o g ound lichen bio-
mass (kg/ha) on lichen pas u es a ound a sam-
ple loca ion was also included as one independ-
en a iable in he analyses.
All linea mixed models we e pe o med
using backwa ds s eps, in o de o emo e
non-signi ican ac o s om he inal models
acco ding o AIC- alues and P- alues. The in-
dependen explana o y a iables added a he
beginning o all analyses we e he ding dis ic
(Muddusjä i, Pais un u i, Salli aa a), g azing
a ea (win e and summe /yea ound), ela-
i e p opo ions o di e en pas u e ypes ( i e
classes), and lichen biomass on lichen pas u es
a ound a sample loca ion. The dependen a i-
able in each model was a p opo ion o ce ain
plan g oups in he samples. Since he lichens,
g aminoids, and dwa sh ubs/lea es in he
samples we e no mally dis ibu ed (Shapi o
Wilk es ; P- alue > 0.2), bu no he b yo-
phy es (Shapi o Wilk es ; P- alue < 0.05), we
made a log- ans o ma ion change o he b yo-
phy e g oup be o e analysis. A e his change,
he Shapi o Wilk es showed ha he b yo-
phy e g oup was also wi hin no mal dis ibu-
ion (P- alue = 0.09).
Resul s
P opo ions o di e en plan g oups in eces
The e we e signi ican di e ences in he ela i e
p opo ions o lichens, g ass and sedges (g ami-
noids), and dwa sh ubs/lea es in eindee
eces be ween he s udy dis ic s (ANOVA e-
sul s o lichens: F = 20.530, P = 0.001; g ami-
noids: F = 23.715, P = 0.001; dwa sh ubs:
F = 20.530, P = 0.001). Howe e , in he p o-
po ion o b yophy es (mosses), he e we e no
di e ences be ween he dis ic s (F = 0.206, P
= 0.814).
In gene al, lichens (which included a small
amoun o mush ooms) we e he mos abun-
dan ood i em g oup iden i ied in he ecal
samples (Fig. 2). Thei p opo ion was highes
in Muddusjä i (summe g azing a ea: 42.58
± 6.56%, win e g azing a ea: 46.35 ± 3.82%)
and lowes in Salli aa a (summe g azing a ea:
32.95±5.12%, win e g azing a ea: 33.62 ±
6.99%). G aminoids and dwa sh ubs/lea es
we e he nex mos abundan plan g oups in
he ecal samples (Figs. 3 and 4). The p opo -
ion o g aminoids was highes in Salli aa a
(summe g azing a ea: 36.19 ± 6.99%, win-
Figu e 2. The ela i e p opo ions (%) o g ound lichen
g oup (mean±SD) in la e summe eces o eindee in
each s udy dis ic and hei g azing a eas based on he
mic ohis ological analysis.
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Rangi e , 35, (1) 2015
e g azing a ea: 32.76 ± 7.97%) and lowes
bo h in Pais un u i (20.94 ± 7.11%) and in
he win e g azing a ea in Muddusjä i (20.93
± 5.36%) (Fig. 3). The ela i e p opo ion o
dwa sh ubs was highes in Pais un u i (37.85
± 6.41%) and lowes in Muddusjä i (summe
g azing a ea: 24.92 ± 5.59%, win e g azing
a ea: 26.25 ± 6.27%) (Fig. 4). The p opo ion
o b yophy es was clea ly lowes in he samples,
a ying be ween 2.9 and 6.5% in he di e en
g azing a eas in he dis ic s (Fig. 5).
The co ela ion analysis shows a signi ican
nega i e co ela ion be ween he p opo ions o
lichen and g aminoids (P = 0.002). In he same
way, he p opo ion o lichen was co ela ed
signi ican ly nega i ely wi h he p opo ion o
dwa sh ubs/lea es (P = 0.019). A signi ican
nega i e co ela ion be ween he p opo ions
o dwa sh ubs/lea es and g aminoids was also
obse ed (P = 0.001). The e was no signi ican
co ela ion be ween he p opo ions o b yo-
phy es and o he plan g oups (Table 5).
Figu e 3. The ela i e p opo ions (%) o g aminoids
(g ass and sedges) (mean±SD) in la e summe eces o
eindee in each s udy dis ic and hei g azing a eas
based on he mic ohis ological analysis.
Figu e 4. The ela i e p opo ions (%) o dwa sh ubs
and lea es (mean±SD) in la e summe eces o eindee
in each s udy dis ic and hei g azing a eas based on
he mic ohis ological analysis.
Figu e 5. The ela i e p opo ions (%) o b yophy es
(mosses) (mean±SD) in la e summe eces o eindee
in each s udy dis ic and hei g azing a eas based on
he mic ohis ological analysis.
Table 1. Dependence o lichen p opo ion (%) in ecal samples on he a iables le in he inal model.
GLMM wi h backwa ds s eps was used o he analysis.
Es ima e S anda d
E o
Min Max alue P alue
In e cep 28.076 1.861 24.361 31.791 15.090 0.001p
Dis Muddusjä i 12.449 1.581 9.293 7.875 15.605 0.001p
Pais un u i 1.750 1.634 -1.512 1.071 5.011 0.228
Salli aa a 0.000 0.000 - - - -
Lichen pas u e 0.300 0.092 0.117 0.484 3.265 0.002
Rangi e , 35, (1) 2015
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46
Fac o s explaining he p opo ions o plan g oups
in samples
The GLMM analysis showed ha he p opo -
ion o lichen g oup in ecal samples was sig-
ni ican ly dependen on he dis ic and on he
p opo ion o lichen pas u e a ound a sample
si e (Table 1). The p opo ion o lichen g oup
in ecal samples was highe in Muddusjä i
han in he Salli aa a dis ic (P = 0.001), bu
he e was no signi ican di e ence be ween he
p opo ions o lichens in he Salli aa a and
Pais un u i dis ic s. In gene al, he mo e li-
chen pas u e he e was a ound a sample si e,
he highe he p opo ion o lichens he e was
obse ed in he ecal samples (P = 0.002). The
p opo ion o g aminoids in ecal samples was
signi ican ly dependen (P = 0.001) on he dis-
ic and on he g azing a ea, being lowe bo h
in Muddusjä i and in Pais un u i han in he
Salli aa a dis ic (Table 2). In gene al, he
ela i e p opo ion o g aminoids (g ass and
sedges) was signi ican ly highe in summe /
yea - ound g azing a eas han in win e g azing
a eas (P = 0.006).
The p opo ion o dwa sh ubs/lea es in
ecal samples was signi ican ly dependen on
he dis ic (P = 0.001), being highe in Pais-
un u i han in Salli aa a (Table 3). The e was
no signi ican di e ence in he ela i e p opo -
ion o dwa sh ubs/lea es be ween Muddus-
jä i and Salli aa a. The ela i e p opo ion o
b yophy es in ecal samples was signi ican ly
dependen on he dis ic and on he p opo -
ion o a bo eal lichen pas u e (ma u e and old
coni e ous o es ) a ound he sample si e (Table
4). The p opo ion o b yophy es was signi i-
can ly di e en in Pais un u i and in Salli aa a
(P = 0.030). The mo e a bo eal lichen pas u e
he e was a ound a sample si e, he highe he
p opo ion o b yophy es was obse ed in he
ecal samples (P = 0.002).
Table 2. Dependence o g aminoids (g ass and sedges) p opo ion (%) in ecal samples on he
a iables le in he inal model. GLMM wi h backwa ds s eps was used o he analysis.
Es ima e S anda d
E o
Min Max alue P alue
In e cep 31.250 1.849 27.558 34.943 16.898 0.001p
Dis ic Muddusjä i -8.170 2.041 -12.244 -4.096 -4.004 0.001p
Pais un u i -16.258 2.096 -20.442 -12.074 -7.758 0.001p
Salli aa a 0.000 0.000 - - - -
G azing
a ea
Summe /
yea ound
5.948 2.091 1.773 10.124 2.844 0.006
Win e 0.000 0.000 - - - -
Table 3. Dependence o dwa sh ubs and lea es p opo ion (%) in ecal samples on he a iables le
in he inal model. GLMM wi h backwa ds s eps was used o he analysis.
Es ima e S anda d
E o
Min Max alue P alue
In e cep 28.794 1.370 26.060 31.528 21.022 0.001p
Dis ic Muddusjä i -3.411 2.055 -7.511 0.690 -1.660 0.102
Pais un u i 9.057 1.937 5.191 12.924 4.676 0.001p
Salli aa a 0.000 0.000 - - - -
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Rangi e , 35, (1) 2015
Di e ences be ween wo sample p ocessing me h-
ods
The independen sample - es esul s showed
ha he e was a signi ican di e ence be ween
he p opo ions o lichen g oup in he wo il-
e ing me hods (mean o one il e me hod
33.04 ± 1.629% and o wo il e s me hod
39.12 ± 2.029% ; = 4.712, P = 0.001). Us-
ing one addi ional il e wi h an ape u e o
0.250 mm, he p opo ion o lichen inc eased
by 15.5% ( om 33.04 o 39.12%) in he sub-
samples, compa ed o he me hod o using only
one il e wi h an ape u e o 0.125mm. Tha
means ha he di e ence be ween he ela i e
p opo ions o lichen in he il e ing me hods
amoun ed o 6.077 ± 4.995%. In o he plan
g oups, he e we e no signi ican di e ences be-
ween hese wo me hods (P > 0.110).
Discussion
The obse ed high amoun o lichen in he sum-
me die o eindee is somehow an unexpec ed
esul , since due o hei high ene gy con en ,
lichens a e conside ed o o m he o age g oup
impo an o eindee nu i ion mainly in win-
e (Wes e ling, 1970). Ou esul s, howe e ,
show ha lichen can also o m a conside able
pa o he eindee die in he summe season,
i eindee ha e ee access o lichen pas u es
du ing summe . Lichens p obably also com-
pensa e o some o he mo e common summe
o age g oups, i hese o age g oups a e no
su icien ly a ailable. We obse ed ha ein-
dee compensa ed o g aminoids wi h lichen
and dwa sh ub/lea es when he e was a lack o
g aminoids on pas u es bu when lichens and
dwa sh ubs, in con as , we e easily a ailable,
as in Pais un u i (see Table 5 and Figs. 2 and
3). I he e a e no a ailable mush ooms (which
o m an impo an pa o he eindee die in
la e summe and ea ly au umn), eindee a e
p obably also mo e eage o ea lichens.
Depending on plan a ailabili y and qual-
i y, eindee u ilize a wide a ie y o plan s in
summe (Bjo k oll e al., 2009). Usually, ein-
dee eed selec i ely on se e al dozens o ascu-
la plan s in summe (Nieminen e al., 1989).
Howe e , ou s udy shows ha in ce ain pas-
u e condi ions in summe , g ound lichens can
domina e he eindee die . Simila ly o ou
s udy, S aaland e al. (1995) also obse ed ha
du ing he summe season, when eindee had
high access o lichens, he selec ion o g asses,
o bs, and lea es d opped o 45-70%, and li-
chens o med he mos u ilized ood i em
g oup (lichen in ake a ied be ween 15-42%)
o eindee g azing in d y g aminoid hea h and
low alpine sh ub hea h.
The ela i e p opo ions o lichen, g ami-
noids, and dwa sh ubs/lea es in eindee eces
also a ied signi ican ly be ween he s udy dis-
ic s. In gene al, he p opo ion o lichen was
dependen on he p opo ion o lichen pas u e
a ound he sample si e, which indica es ha
when eindee g aze on an a ea domina ed by
lichen ype ege a ion, hey s ill ac i ely selec
Table 4. Dependence o b yophy es (mosses) p opo ion (log-%) in ecal samples on he a iables le
in he inal model. GLMM wi h backwa ds s eps was used o he analysis.
Es ima e S anda d
E o
Min Max alue P alue
In e cep 0.247 0.113 0.022 0.473 2.193 0.032
Dis ic Muddusjä i 0.14 0.099 -0.058 0.338 1.409 0.164
Pais un u i 0.273 0.123 0.028 0.517 2.225 0.030
Salli aa a 0.000 0.000 - - - -
A bo eal lichen pas u e 0.01 0.003 0.004 0.017 3.243 0.002