Diurnal defecation rate of moose in southwest Finland

DIURNAL DEFECATION RATE OF MOOSE IN SOUTHWEST FINLAND Juho Ma ala 1 and An i Uo ila 2 1 Finnish Fo es Resea ch Ins i u e Me la, P.O. Box 68, FI-80101 Joensuu, Finland; 2 Uni e si y o Helsinki Hyy iälä Fo es S a ion, Hyy iälän ie 124, FI-35500 Ko keakoski, Finland. ABSTRACT: An accu a e measu e o de eca ion a e is essen ial o applica ion o pelle g oup coun s in moose (Alces alces) popula ion es ima es. We measu ed he win e ime, diu nal de eca ion a e o moose by acking 7 GPS-colla ed and 22 uncolla ed moose in sou hwes Finland. The mean de eca- ion a e was 23.5 ± 4.2 pelle g oups/d, one o he highes alues epo ed. The mean de eca ion a e did no di e be ween he acking me hods (GPS s. uncolla ed moose); limi ed sample size p e- cluded conclusions abou sex and age di e ences. The de eca ion a e was no co ela ed wi h calenda week, leng h o accumula ion pe iod, o numbe o diu nal beds. Ou esul s a e app op ia e o use in sou hwes Finland when using he pelle g oup me hod o assess moose popula ion densi y. ALCES VOL. 49: 155–161 (2013) Key wo ds:Alces alces, Finland, de eca ion a e, moose, pelle g oup, acking. Coun ing ecal pelle g oups o moose (Alces alces) has been widely used o es i- ma e habi a u iliza ion, eeding beha io , and popula ion ends and densi y (see F anzmann e al. 1976b, Fo bes and The- be ge 1993, Hä könen and Heikkilä 1999, Rönnegå d e al. 2008, Månsson 2009, Mån- sson e al. 2011a, b). Reliable moose popula- ion es ima es a e no always ealized om pelle g oup coun s (Rönnegå d e al. 2008), bu he me hod's use ulness has been no ed (Ne 1968, Lau enschlage and Jo dan 1993, Månsson e al. 2011b), despi e some unce ain y (Ne 1968). Howe e , o suc- cess ully es ima e moose popula ion densi y using a pelle g oup coun , i is c i ical o use an accu a e de eca ion a e in he su ey a ea and ime pe iod (Timme mann 1974). Ea lie s udies ha e used 2 main me h- ods o es ima e he de eca ion a es o moose: 1) ack moose in snow-co e ed e - ain (Joyal and Rica d 1986, Ande sen e al. 1992), and 2) es ima e he numbe o pelle g oups in a closed a ea o island whe e he numbe o moose is known (Jo dan e al. 1993). De eca ion a es ha e occasionally been es ima ed by compa ing he ae ial cen- suses and pelle g oup coun s in speci ic a eas (Rönnegå d e al. 2008). Moose enclo- su es could also be u ilized, bu esul s om domes ic moose can be a ec ed by ood quali y and beha iou ha a e dissimila o na u al condi ions. GPS adio-colla s enable in ensi e and accu a e acking by iden i y- ing speci ic indi iduals and he beginning and end poin s o hei speci ic acks, p o- iding ideal condi ions o measu e de eca- ion a es o ee- anging moose. Moose de eca ion a es a y by age, sex, habi a , ood quali y, season, and yea (Des- Meules 1968, F anzmann e al. 1976a, Olde- meye and F anzmann 1981, Joyal and Rica d 1986, Ande sen e al. 1992, Månsson e al. 2011b). La ge a ia ions in de eca ion a e ha e been epo ed in di e en a eas; o example, in No h Ame ica a ia ion was 9.6–32.2 pelle g oups/d (Timme mann 1974), and in no he n Eu ope a es a ied Co esponding au ho : Juho Ma ala, Finnish Fo es Resea ch Ins i u e Me la, P.O.Box 68, FI-80101 Joensuu, Finland. +358 40 801 5275. juho.ma a[email p o ec ed] 155 om 14–26.9 pelle g oups/d (Ande sen e al. 1992, Remm and Luud 2003, Rönnegå d e al. 2008). These da a emphasize he impo ance o using a ea-speci ic a es when using he pelle g oup me hod o es i- ma e moose popula ion densi y. Ou main objec i e was o o mula e a gene al es ima e o he win e ime, diu nal de eca ion a e o moose in sou hwes Fin- land by acking bo h GPS-colla ed and uncolla ed moose in snowy e ain. We also compa ed hese 2 acking me hods and looked o di e ences in de eca ion a es be ween sex and age. STUDY AREA Moose we e acked in 2 sepa a e a eas app oxima ely 100 km apa in sou hwes Finland (Fig. 1). Uncolla ed moose we e acked in he O i esi-Kangasala a ea (∼WGS84 61°36′N, 24°22′E) and GPS- colla ed moose in he Loppi-Hy inkää a ea (∼WGS84 60°38′N, 24°35′E). Bo h a eas a e loca ed in he sou he n bo eal ege a ion zone wi h Sco s pine (Pinus syl es is) and No way sp uce (Picea abies) as he domi- nan ee species. Fo es co e was 78% o he o al land a ea in O i esi-Kangasala and 71% in Loppi-Hy inkää (Me la 2012). METHODS T acking o Uncolla ed Moose We ac i ely sea ched o uncolla ed moose in hei known habi a s du ing esh snow condi ions be ween Decembe and Ap il, 1999–2003. The accumula ion pe iod o pelle coun s began when moose we e seen o lushed, enabling da a collec o s o accu a ely ime hei coun by loca ing esh es ing places, pelle g oups, o acks in Fig. 1. S udy a ea loca ions and s a ing poin s o moose acking pe iods in sou he n Finland. In he O i esi-Kangasala a ea some o he s a ing poin coo dina es o uncolla ed moose a e ough es ima es and o e lap because hey could no be sepa a ed a he map scale. 156 DIURNAL DEFECATION RATE OF MOOSE –MATALA AND UOTILA ALCES VOL. 49, 2013 he snow. The pelle g oups we e coun ed he ollowing day, un il moose led om he coun e . The end o he accumula ion pe iod was de e mined in a simila manne as he s a ing poin . The accumula ion pe iods a ied om 8–31 h. Moose we e classi ied as ei he adul s o cal es by isually obse ing hem and hei pelle s, bedding places, and beha io . Thei sex could be de e mined wi hou isual con- ac by analyzing hei u ina ion me hods, as bulls u ina e in on o he hind hoo es. We measu ed he diu nal de eca ion a e o 5 bulls, 3 cows, and 3 cal es. Addi ionally we measu ed 3 cows wi h win cal es and 1 cow wi h a single cal , wi hou sepa a ing he pelle g oups o he cows and cal es. In o al, we we e able o eco d he diu nal de e- ca ion a es o 22 uncolla ed moose and coun he numbe o beds o 7 indi iduals. T acking o GPS-Colla ed Moose The Finnish Game and Fishe ies Resea ch Ins i u e (FGFRI) p o ided loca- ion da a o he GPS-colla ed moose. The FGFRI implemen ed GPS-colla ing p oce- du es in acco dance wi h Finnish legisla ion, wi h pe mission om he Na ional Animal Expe imen Boa d o Finland. An indi idual GPS-colla ed cow was acked once and ano he on 5 sepa a e occasions, while 1 cow wi h win cal es was acked once, and one cow wi h a single cal wice; al oge he , 7 indi iduals we e acked. The GPS-colla ed moose we e acked a ew days a e esh snow all in Decembe – Ma ch 2010. The coun e wen o he mos ecen loca ion o moose acks which we e usually ∼12 h old. The pelle g oups we e coun ed along he moose ack by ollowing i agains he o iginal cou se o he moose; he coo dina es o he pelle g oups, beds, and u ina ion si es we e loca ed wi h hand- held GPS de ices. The accumula ion pe iod inished when an indi idual ack became mixed wi h o he s o sunse p ecluded acking. The du a ion o he accumula ion pe iod was de e mined by iden i ying he ime associa ed wi h he closes loca ion o he acks wi h GPS colla da a. The accumu- la ion pe iods anged om 6–47 h. Da a Analysis A leas 20 pelle s we e equi ed o make a pelle g oup. We p ocessed he pelle g oup da a o 29 moose (7 GPS-colla ed, 22 uncol- la ed) o calcula e he diu nal de eca ion a e (numbe o pelle g oups p oduced pe indi- idual in 24 h) om he accumula ion pe i- ods o indi idual bulls, cows, and cal es. Mean alues we e calcula ed o he g oup o cows and cal es when i was impossible o iden i y cal om cow pelle g oups; ana- lyses o 25 sepa a e cases we e used o calcu- la e he diu nal de eca ion a e. Fo compa ison, we sampled he mean alues o diu nal de eca ion a es o 3 sex and age classes: 1) bulls, 2) cows, and 3) cal es and cow-cal es. The las class was equi ed because we we e only able o ack 3 indi idual cal es, which was insu icien o any easonable analysis. We also com- pa ed he mean de eca ion a es o he uncol- la ed and GPS-colla ed moose. Due o limi a ions in he linea i y and homogenei y o a iances in he da a, we used he K uskal-Wallis es o mul iple samples and he Mann-Whi ney es o pai ed sam- ple compa isons. Fu he mo e, we sea ched o possible co ela ions be ween he diu nal de eca ion a e and 1) he calenda week o he accumula ion pe iod, 2) he du a ion o he accumula ion pe iod, and 3) he numbe o diu nal beds o pa o he samples. All s a is ical calcula ions we e pe o med using he S a is ical Package o he Social Sciences (SPSS) 17.0 so wa e. Da a ela ing o home ange o moose du ing he accumula ion pe iod (calcula ed using he Home Range Tools o A cGIS® e sion 1.1 wi h he Minimum Con ex Polygon me hod), leng h o he moose ack ALCES VOL. 49, 2013 MATALA AND UOTILA –DIURNAL DEFECATION RATE OF MOOSE 157 du ing he accumula ion pe iod, and he numbe o diu nal u ina ions we e also col- lec ed om he GPS-colla ed moose, bu insu icien sample sizes p ecluded hei u i- liza ion in he analysis. RESULTS The diu nal de eca ion a e anged om 12.2–32 pelle g oups wi h an o e all mean o 23.5 ± 4.2 (SD; Table 1, Fig. 2). The mean alues o bull, cow, and cal /cow-cal g oups we e di e en (K uskal-Wallis es : χ 2 = 9.9, d = 2, P= 0.007; Table 1). The cal and cow-cal g oup had he highes mean a e, bu was s a is ically di e en only om he cow g oup (Table 1). The bull g oup had he lowes mean a e, bu also he lowes and highes absolu e alues (i.e., he wides ange; Table 1). The cow and bull g oup a es and he a es o he uncolla ed and GPS-colla ed moose we e no di e en (Table 1). The diu nal de eca ion a e was no ela ed o he calenda week o he accumula- ion pe iod (Fig. 2, Table 2). The GPS- colla ed indi idual which was acked 5 imes be ween 17 Janua y and 23 Feb ua y 2010 showed no ends du ing his pe iod. The de eca ion a e did no co ela e o he du a ion o he accumula ion pe iod o o he numbe o diu nal beds (Table 2). The mean alues o he o he a iables we e: diu nal numbe o beds = 8.0 ± 4.2 (SD, n = 15), he diu nal u ina ion a e = 1.0 ± 1.0 (SD, n = 9), he a ea o home ange du - ing he accumula ion pe iod = 95,267 m 2 ± 122,399 (SD, n = 9), and leng h o ack du - ing he accumula ion pe iod = 955 m ± 832 (SD, n = 6). DISCUSSION The mean de eca ion a e was conside ed high (23.5 ± 4.2 pelle g oups/d) bu simila o he a e age a e measu ed in ela i ely good moose habi a in sou he n No way (22.9 pel- le g oups/d; Ande sen e al. 1992). Lowe de eca ion a es we e measu ed in sou he n Sweden (14 pelle g oups/d; Rönnebe g e al. 2008) and on Isle Royale, No h Ame ica (20.9; Jo dan e al. 1993). Many s udies ha e epo ed lowe de eca ion a es in Alaska and Canada (see DesMeules 1968, F anzmann e al. 1976a, Oldemeye and F anzmann 1981, Joyal and Rica d 1986). The high alues epo ed in ou s udy a ea a e no un easonable when compa ing he s a us o he moose popula ion in Finland o o he No dic coun ies. The Finnish moose popula ion is lowe and o a highe p oduc i e s a e compa ed o hose in Table 1. Mean alues o he win e ime, diu nal de eca ion a e and compa isons be ween moose g oups in sou he n Finland. Pelle g oups (#/ind/24 h) Mann-Whi ney Tes Mean Min Max n SD Tes agains U- alue P G ouping by moose ype: Bulls 20.1 12.2 32.0 5 7.3 Cow 12 0.152 Cows 22.3 18.9 24.0 9 2.2 Cal + g oups 10 0.002 Cal and Cow-cal g oups 25.9 23.0 31.3 11 2.2 Bull 11 0.061 G ouping by acking me hod: GPS-colla ed 23.5 20 26.7 9 1.8 Uncolla ed 64 0.647 Uncolla ed 23.4 12.2 32.0 16 5.2 All moose 23.5 12.2 32.0 25 4.2 158 DIURNAL DEFECATION RATE OF MOOSE –MATALA AND UOTILA ALCES VOL. 49, 2013 Sweden and No way (La sund e al. 2003, Tiilikainen e al. 2012). These di e ences p esumably imply be e o aging habi a , highe nu i ional condi ion, and a esul an highe de eca ion a e in Finland. Because de eca ion a es a e in luenced by ood qual- i y and a ailabili y and show la ge a ia ion (Ande sen e al. 1992), i ollows ha a ea- speci ic de eca ion a es should be measu ed when using pelle g oup coun s o es ima e popula ion densi y. The ex emes in de eca ion a e a ied la gely in ou s udy, bu mos obse a ions, especially o he GPS-colla ed indi iduals, we e simila o he mean indica ing he gen- e al eliabili y o hese da a and hei applica- ion o es ima e popula ion densi y. The ex eme alues could esul om local o a- ging condi ions o om longe mo emen s associa ed wi h unin ended dis u bances while acking he uncolla ed moose. Because he de eca ion a e can co ela e wi h age and sex, hese ela ionships should be aken in o accoun i a change in popula- ion s uc u e occu s (F anzmann e al. 1976a). Ou da a indica e ha he cal and cow-cal g oup had sligh ly highe de eca- ion a es han indi idual cows as also epo ed by DesMeules (1968), bu opposi e ha o Joyal and Rica d (1986). Bulls and cows we e no di e en in ou analyses; howe e , gene al conclusions abou sex and age di e ences canno be made due o he limi ed sample size. Using an accu a e de eca ion a e is c i i- cal when applying pelle g oup coun s in moose popula ion densi y es ima es. De eca- ion a es ha e no been published p e iously in Finland, and gi en he conside able a ia- ion in simila da a om no he n Eu ope, we conside ou esul s he bes a ailable o local use. Fu he mo e, simila de eca ion a es we e ob ained o bo h egions in ou s udy, making he mean alues applicable o all o sou he n Finland. The lack o co e- la ion be ween calenda week and de eca ion a e indica es a s able accumula ion a e h oughou win e , sugges ing ha hese a es migh be applicable o sp ing ime pel- le g oup coun s. Va ia ions in he absolu e da a and means should be aken in o accoun when calcula ing con idence limi s o he inal moose densi y es ima e. ACKNOWLEDGEMENTS We hank M . J. Sillanpää o acking he GPS-colla ed moose and M . H. Ma ila 50 Fig. 2. The diu nal de eca ion a e o moose in sou he n Finland ela i e o he ime o he acking pe iod (calenda weeks). Moose ypes a e classified as: 1) indi idual bulls, cal es, and cows, and 2) cow-cal g oups (when indi iduals could no be sepa a ed du ing acking); cow1 = cow wi h single cal , cow2 = cow wi h 2 cal es. Table 2. Pea son co ela ions be ween he diu nal de eca ion a e (pelle g oups/moose/24 h) and empo al a iables in sou he n Finland; no di e ences we e ound. Calenda week Accumula ion pe iod (h) Beds/ moose/ 24 h Coe icien −0.144 −0.086 −0.403 P- alue 0.492 0.682 0.136 n25 25 15 ALCES VOL. 49, 2013 MATALA AND UOTILA –DIURNAL DEFECATION RATE OF MOOSE 159 o acking in he Kangasala a ea. D . J. Pusenius om he Finnish Game and Fish- e ies Resea ch Ins i u e is g ea ly acknowl- edged o o ganizing he GPS-colla ing and hus enabling da a a ailabili y on exac moose loca ions o ou s udy. We also hank he Finnish Fo es Resea ch Ins i u e, he Uni e si y o Helsinki, and he Minis y o Ag icul u e and Fo es y o p o iding inan- cing o ou wo k. M.Sc. M. 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