Composition of late summer diet by semi-domesticated reindeer (Rangifer tarandus tarandus) in different grazing areas in northernmost Finland

This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Bi gi a Åhman, Technical Edi o E a Wiklund and G aphic Design: H-G Olo sson, www. angi e jou nal.com 39 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 This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Bi gi a Åhman, Technical Edi o E a Wiklund and G aphic Design: H-G Olo sson, www. angi e jou nal.com 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- This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Bi gi a Åhman, Technical Edi o E a Wiklund and G aphic Design: H-G Olo sson, www. angi e jou nal.com 41 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 Rangi e , 35, (1) 2015 This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Bi gi a Åhman, Technical Edi o E a Wiklund and G aphic Design: H-G Olo sson, www. angi e jou nal.com 42 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 . This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Bi gi a Åhman, Technical Edi o E a Wiklund and G aphic Design: H-G Olo sson, www. angi e jou nal.com 43 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 Rangi e , 35, (1) 2015 This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Bi gi a Åhman, Technical Edi o E a Wiklund and G aphic Design: H-G Olo sson, www. angi e jou nal.com 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. This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Bi gi a Åhman, Technical Edi o E a Wiklund and G aphic Design: H-G Olo sson, www. angi e jou nal.com 45 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 This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Bi gi a Åhman, Technical Edi o E a Wiklund and G aphic Design: H-G Olo sson, www. angi e jou nal.com 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 - - - - This jou nal is published unde he e ms o he C ea i e Commons A ibu ion 3.0 Unpo ed License Edi o in Chie : Bi gi a Åhman, Technical Edi o E a Wiklund and G aphic Design: H-G Olo sson, www. angi e jou nal.com 47 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