Metapopulation perspective to institutional fit: maintenance of dynamic habitat networks

Copy igh © 2017 by he au ho (s). Published he e unde license by he Resilience Alliance. Fab i ius, H., A. Jokinen, and M. Cabeza. 2017. Me apopula ion pe spec i e o ins i u ional i : main enance o dynamic habi a ne wo ks. Ecology and Socie y 22(2):30. h ps://doi.o g/10.5751/ES-09203-220230 Resea ch Me apopula ion pe spec i e o ins i u ional i : main enance o dynamic habi a ne wo ks Henna Fab i ius 1,2, A i Jokinen 3 and Ma Cabeza 1 ABSTRACT. Species li ing in me apopula ions depend on connec ed habi a ne wo ks o hei su i al. I habi a ne wo ks expe ience as empo al dynamics, species conse a ion equi es p e en ing habi a discon inui ies ha could lead o me apopula ion ex inc ions. Howe e , ew ins i u ional solu ions exis o he main enance o spa io empo ally dynamic habi a ne wo ks ou side o p o ec ed a eas. To explo e his o en neglec ed p oblem, we s udied he ins i u ional i o alse hea h i illa y (Meli aea diamina) conse a ion in Finland om he pe spec i e o conse a ion ins i u ions’ abili y o manage ea ly successional habi a a ailabili y o his endange ed species. We iden i ied ou ins i u ional a angemen s ha enable e ec i e conse a ion managemen o dynamic habi a ne wo ks: (1) acknowledgmen o habi a dynamics, (2) moni o ing o and esponding o changes in he habi a ne wo k, (3) managemen o esou ces o luc ua ing esou ce needs, and (4) scaling o ac i i ies h ough lexible collabo a ions. These a angemen s p o ide he ins i u ional lexibili y needed o esponding o empo al changes in habi a a ailabili y. Key Wo ds: conse a ion; dynamic habi a ne wo ks; alse hea h i illa y; unc ional i ; habi a o e g ow h; ins i u ional i ; Meli aea diamina; me apopula ion dynamics; spa ial i ; empo al i INTRODUCTION Conse a ion o dynamic habi a ne wo ks Species li ing in me apopula ions depend on he con inuous a ailabili y o connec ed habi a pa ch ne wo ks o hei su i al (Hanski 1994, Hanski and O askainen 2000, Cabeza 2003). Habi a pa ch ne wo ks may unde go g adual deg ada ion (Mo elli i e al. 2010) o loss (Bulman e al. 2007) o habi a pa ches because o human land use ac i i ies. Addi ionally, clima e change may cause habi a pa ch ne wo ks o shi g adually owa d he poles (Radchuck e al. 2014). In pa icula , habi a pa ch ne wo ks can be dynamic o species ha occupy ea ly successional habi a s, in which case indi idual habi a s may eme ge and disappea depending on local dis u bance pa e ns and he p ocess o succession (Johs e al. 2011). As he in ensi ica ion o human land use has caused changes o dis u bance egimes ac oss ecosys ems (Ly le and LeRoy Po 2004, Enck and Oda o 2008, Lou i al e al. 2011), he a ailabili y o ea ly successional habi a s has dec eased in many ecosys ems (Thompson and DeG aa 2001, Halada e al. 2011), and his has caused he endange men o ea ly successional species (Askins 2001, De me s 2003, Long 2009). These de elopmen s ha e esul ed in spa ially explici conse a ion planning needs o species ha li e in spa io empo ally changing habi a ne wo ks (Van Tee elen e al. 2012). When conse a ion plans ha e o be designed o dynamic habi a ne wo ks, speci ic complexi ies eme ge because o hei ansi ional na u e. I habi a pa ches a e sho li ed, empo al a ia ions in habi a egene a ion a es may cause empo al luc ua ions in habi a a ailabili y (Moilanen e al. 2014). Such luc ua ions ha e been shown o inc ease ex inc ion isks o species ha li e in dynamic habi a ne wo ks because hey can cause synch onized collapses o local popula ions (Bough on and Mal adka 2002, Johs e al. 2011, Van Tee elen e al. 2012). Conse a ion o dynamic habi a ne wo ks may he e o e equi e managemen o cons an spa io empo al habi a a ailabili y o he species o in e es , which may equi e acili a ing main enance ac i i ies a imes and loca ions ha bes minimize habi a discon inui ies, i.e., empo al gaps in habi a a ailabili y (Hanski 1999). These main enance ac i i ies can include me hods ha delay o es a succession o inc ease he a ailabili y o he successional s age o in e es , e.g., mowing o g azing. Howe e , ew ins i u ional solu ions ha e been de eloped o he pu pose o such spa io empo al planning. Ag i-en i onmen al schemes (Eu opean Commission 2005, Kleijn e al. 2006, A ponen e al. 2013) a e used o main ain habi a ne wo ks o demanding ea ly successional species, bu hese ins i u ions a e es ic ed in hei abili y o p io i ize spa ial connec i i y o habi a s (Schou en e al. 2013). Fi e managemen ins i u ions (Richa ds e al. 1999, Keeley and Fo he ingam 2001, S ewa e al. 2005) p o ide examples o ins i u ions ha aim a p o iding con inuums o ea ly successional habi a s. Thei design and pe o mance ha e been s udied om a spa ially implici pe spec i e (Hansen 2014). Ins i u ional i o habi a ne wo k main enance o ganiza ions Ins i u ional i has been used as a concep ual amewo k o s udy how well ins i u ions a e aligned wi h he eali ies hey manage o go e n (Young 2003, Folke e al. 2007, Cox 2012). Fi depends on he go e nance o managemen a ge s o ins i u ions and on he poli ical and cul u al se ing (Halle e al. 2013) and may en ail mul iple di e en measu es o i simul aneously (Lebel e al. 2013). In he case o na u al esou ce managemen , an ins i u ion’s pe o mance may be measu ed in e ms o i s abili y o inc ease he spa ial esilience o he sys em agains pe u ba ions (Beng sson e al. 2003, Allen e al. 2016), which educes empo al a iabili y in he low o he esou ce in o he economy (Walke e al. 2004, Folke e al. 2007, Os om 2007). Fi 1Depa men o Biosciences, Uni e si y o Helsinki, 2Swedish Species In o ma ion Cen e, Swedish Uni e si y o Ag icul u al Sciences, 3Facul y o Managemen (En i onmen al Policy), Uni e si y o Tampe e Ecology and Socie y 22(2): 30 h ps://www.ecologyandsocie y.o g/ ol22/iss2/a 30/ has been es ima ed quali a i ely based on in e iews and policy documen s (Munck a Rosenchöld e al. 2014), by calcula ing simila i y me ics be ween ecological sys ems and policy desc ip ions (Eks om and Young 2009), o by coun ing i measu es om expe ques ionnai e eplies (Lebel e al. 2013). Ins i u ional i has been measu ed om spa ial (Folke e al. 2007, Moss 2012, Lebel e al. 2013), empo al (Pé ez-No d ed e al. 2008, Munck a Rosenschöld e al. 2014), and unc ional (Eks om and Young 2009) pe spec i es. Func ional i measu es whe he he key unc ions o he managed sys em a e add essed by he managing ins i u ion (Eks om and Young 2009), whe eas spa ial i e e s o misma ches in he spa ial ex en o adminis a ion e sus ha o managed ecological sys ems (Folke e al. 2007). Tempo al i e e s o he iming o ins i u ional ac i i ies in ela ion o he empo al pa e ns in he biophysical en i onmen hey manage o espond o (Munck a Rosenschöld e al. 2014). Pé ez-No d ed e al. (2008) sugges he use o concep s o empo and phase o cha ac e ize he leng h and iming o o ganiza ional and en i onmen al cycles, such as cycles o echnological imp o emen o go e nmen al egula o y ac i i ies, o assess hei empo al i . Munck a Rosenschöld e al. (2014) use a wide concep o imescapes om Adam (2000), which includes he s udy o spa io empo al pa e n, sequence, and ime ame o o ganiza ional ac i i ies. Spa ial and empo al i ha e also been analyzed simul aneously (Cash e al. 2006). Pas wo ks ha e p oposed ha in social-ecological sys ems p one o pe u ba ions and dynamic changes, high ins i u ional i equi es adap i e go e nance (Folke 2006). This equi es an unde s anding o ecosys em dynamics, o ins ance o he eedback mechanisms ha a e c i ical in he main enance o he spa ial esilience o he sys em (Allen e al. 2016), managemen p ac ices ha espond o eedback, adap i e capaci y o espond o su p ise, and ins i u ional lexibili y (Folke 2006, Folke e al. 2007). In mul iscale ins i u ions, adap i e capaci y may equi e in e play o ac o s o di e en scales, amewo ks, and egimes (Paa ola e al. 2009) and he exis ence o nes ed ins i u ions (Folke 2006, Folke e al. 2007) ha can b idge he unc ions o di e en ac o s a he igh ime o c ea e social-ecological esilience (Lebel e al. 2006, Olsson e al. 2007, G een e al. 2015). High i also equi es accep abili y o he ins i u ions in he eyes o esou ce use s and landowne s (Hukkinen 2012, Fa ell and Thiel 2013, Hiedanpää 2013), he a ainmen o which may equi e c ea ing mechanisms ha u ilize hei knowledge in he unc ioning o he ins i u ions (Olsson e al. 2007). Powe s uc u es wi hin mul ile el go e nance ins i u ions can main ain ins i u ional mis i (Folke e al. 2007). Pé ez-No d ed e al. (2008) sugges ha empo al i o ins i u ions depends on whe he in e nal empo al cycles o he ins i u ions can be easonably synch onized wi h, o en ained o (Jokinen 2006), hei ex e nal en i onmen o whe he dominan cycles wi hin go e nance o ganiza ions ac as pace s, also called zei gebe s (Pé ez- No d ed e al. 2008), ha main ain mis i . We se ou o iden i y ins i u ional a angemen s ha enable e ec i e conse a ion managemen o dynamic habi a ne wo ks. We assessed he pe o mance o conse a ion ins i u ions om he pe spec i e o hei abili y o p o ide habi a s ha suppo s able, esilien popula ion sizes o he species being conse ed. This, based on he me apopula ion heo y, equi es habi a ne wo ks o adequa e connec i i y (Cabeza 2003) and capaci y (Hanski and O askainen 2000) o e ime. We ca ied ou a case s udy o he conse a ion o a Finnish endange ed bu e ly, he alse hea h i illa y Meli aea diamina (Lang, 1789), o which he p ima y documen ed cause o endange men in Finland is he successional dynamics o i s habi a s, i.e., “o e g owing o meadows and o he open habi a s” (Rassi e al. 2010:40). We used hema ic in e iews and conse a ion epo s o iden i y ypes and imings o ac i i ies in he Finnish na ional-le el and egional- le el go e nmen al conse a ion o ganiza ions wi h ega d o alse hea h i illa y conse a ion. Finally, we collec ed expe es ima es on he po en ial need o habi a main enance o endange ed Finnish species o es ima e he gene alizabili y o ou indings. Based on he case s udy, we iden i ied key ins i u ional a angemen s ha enable e ec i e conse a ion managemen o dynamic habi a ne wo ks. METHODS Case s udy: alse hea h i illa y conse a ion in Finland The alse hea h i illa y is a mois meadow specialis ha has, like many bu e ly species, su e ed om ag icul u al mode niza ion and d ainage o mois soils wi hin he ecen decades (Liinalaakso 2000, an Swaay e al. 2006). The alse hea h i illa y is lis ed as an endange ed species (Rassi e al. 2010), a p o ec ed species, and a species unde s ic p o ec ion (Minis y o he En i onmen 1997) in Finland, he majo cause o i s endange men being he as successional dynamics o i s habi a s, i.e., “o e g owing o meadows and o he open habi a s” (ca ego y N), in he absence o adi ional land uses. Conse a ion planning o he alse hea h i illa y is complica ed by he species’ dependence on dis u bed habi a s, ei he human-main ained mois meadows o a dynamically changing ne wo k o sui able ea ly successional allows. Mos alse hea h i illa y habi a s appea on p i a e ag icul u al land. Because alse hea h i illa y habi a s a e seldom occupied by o he endange ed species o egional in e es , hey a e no he ocus o habi a - ocused conse a ion p og ams. The expec ed li e ime o an unp o ec ed alse hea h i illa y habi a has been es ima ed o be 13.1 yea s, he quali y o he habi a si e s a ing o dec ease 2-3 yea s a e si e main enance because o he successional dynamics o he habi a s (Fab i ius and McB ide 2017) and habi a s equi ing main enance, e.g., mowing o g azing, a leas e e y 5 yea s (In ke 2003). False hea h i illa y conse a ion in Finland ocuses on he egional-le el Cen es o he Economic De elopmen , T anspo and he En i onmen (ELY cen e s), which hold he go e nmen al au ho i y o s ee and moni o local and egional land use planning, o es ablish conse a ion a eas on p i a e land, and o en o ce habi a dema ca ions and pe mi s o excep ion ela ed o he habi a s o endange ed species (Minis y o he En i onmen 2015, Cen e o Economic De elopmen , T anspo and he En i onmen 2016). Based on he alse hea h i illa y dis ibu ion in Finland (Wahlbe g 1998), ma e s ela ed o he species a e deal wi h in he ELY cen e s o Pi kanmaa (Pi kanmaa egion), Sou h Os obo hnia (Os obo hnia and Sou h Os obo hnia egions), and sou hwes Finland (Sa akun a egion). A he cen al le el o adminis a ion, he Minis y o he En i onmen o mula es he en i onmen al policies o he Finnish go e nmen , s ee s and esou ces he ELY cen e s in na u e conse a ion, and Ecology and Socie y 22(2): 30 h ps://www.ecologyandsocie y.o g/ ol22/iss2/a 30/ guides he wo k ca ied ou in he Finnish En i onmen Ins i u e (SYKE), which conduc s esea ch and compiles en i onmen al da a. The Na u al He i age Se ices uni o Me sähalli us (Finnish Fo es y Se ice) is esponsible o na u e conse a ion on s a e-owned land (Minis y o he En i onmen 2015). Thema ic in e iews and conse a ion epo s We ca ied ou hema ic in e iews be ween No embe 2014 and Feb ua y 2015 o de ail he goals, p io i ies, decision-making p ocesses, collabo a ions, and p ac ices o alse hea h i illa y conse a ion in Finland. The esponden s included h ee senio ad ise s o he egional ELY cen e s ha we e in cha ge o he alse hea h i illa y conse a ion in hei espec i e egions (Pi kanmaa, Sou h Os obo hnia, and sou hwes Finland), wo ci il se an s in he Minis y o he En i onmen esponsible o he unding decisions ega ding go e nmen al conse a ion unds, and a biologis a he Na u al En i onmen Cen e o SYKE who was esponsible o egional p io i iza ion mee ings o endange ed species. The in e iewees co e ed key ac o s, and hey we e well awa e o he basic ecology and conse a ion challenges o he alse hea h i illa y because o collabo a ion be ween he species-speci ic esea che s and adminis a ion since Wahlbe g (1997, 1998). We conduc ed he in e iews by ollowing a eady-made hema ic in e iew ame (Appendix 1) ha was di e en o he h ee g oups o in e iewees, i.e., he ELY cen e s, he Minis y o he En i onmen , and SYKE, bu allowed he discussion o ollow opics b ough up by he esponden s. Besides hema ic in e iews, we s udied 14 epo s o he egional en i onmen al adminis a ion ega ding he moni o ing and es o a ion o alse hea h i illa y habi a s. We used con en analysis o examine he in e iew da a and conse a ion epo s wi h ega d o he decision-making p ocesses, goals, p ac ices, collabo a ion s uc u es, and dependencies ela ed o alse hea h i illa y conse a ion. Fi s , we iden i ied wha in o ma ion was a ailable o conse a ion manage s wi h ega d o changes in he numbe s, sizes, quali ies, and loca ions o habi a pa ches in he alse hea h i illa y habi a ne wo k o hei espec i e adminis a i e a eas. Nex , we iden i ied cyclical ac i i ies aking place in he conse a ion o ganiza ions ega ding he iden i ied habi a ne wo k dynamics and asked wha se he empo and phase o hese ac i i ies. We also examined how hese o ganiza ions applied o mal legal amewo k e sus adap i e go e nance in conse a ion (Ga mes ani e al. 2013). Finally, we asked he ollowing ques ions o e alua e he ins i u ional i o conse a ion: A e conse a ion ins i u ions able o iden i y and espond o dec easing habi a quali y and a ailabili y wi hin such ime ames ha can p e en habi a discon inui ies o alse hea h i illa y me apopula ions? A e conse a ion ins i u ions able o p io i ize, execu e, o acili a e habi a main enance, es o a ion, o gene a ion a loca ions ha a e impo an o he spa io empo al connec i i y o he habi a ne wo k? Based on ou indings, we iden i ied key ins i u ional a angemen s ha enable e ec i e conse a ion managemen o dynamic habi a ne wo ks. Expe es ima es on he gene alizabili y o he s udy case To assess how common he conse a ion challenges iden i ied in ou case s udy a e in he ield o endange ed species conse a ion in Finland, we es ima ed how many o he endange ed species in Finland, also h ea ened by he successional dynamics o hei habi a s, would be conse ed using he same p ocesses as he alse hea h i illa y because o a simila legal and conse a ion s a us. Fo his pu pose, we ex ac ed he numbe s o endange ed species, belonging o he ca ego ies ulne able (VU), endange ed (EN), and c i ically endange ed (CR), om he Finnish Red Lis o Endange ed Species 2010 (Rassi e al. 2010) ha a e also h ea ened by successional habi a dynamics (p ima y cause o endange men N) and ha a e ca ego ized as species unde s ic p o ec ion. We hen asked wo axon-speci ic expe s o iden i y species in he esul ing lis ha end o occupy habi a s ha all ou o common habi a -speci ic conse a ion p og ams and would he e o e need species-speci ic conse a ion measu es o e icien conse a ion. RESULTS We ound unc ional, spa ial, and empo al mis i s in he ins i u ions ha manage habi a a ailabili y o he alse hea h i illa y. Based on hese indings, we iden i ied ou key ins i u ional a angemen s ha enable e ec i e main enance o dynamic habi a ne wo ks: (1) acknowledgmen o habi a dynamics, (2) moni o ing o and esponding o changes in he habi a ne wo k, (3) managemen o esou ces o luc ua ing esou ce needs, and (4) scaling o ac i i ies h ough lexible collabo a ions. Fo each a angemen , we sugges key ques ions ha can be used by conse a ion manage s, public o icials, and esea che s o assess he ex en o i and causes o po en ial mis i s in ins i u ions ha main ain dynamic habi a ne wo ks. The ques ions ha e been o mula ed so ha , in each ca ego y, an inc easing numbe o posi i e eplies sugges s inc easing i . Following he ques ions, we sugges guidelines o imp o ing he ins i u ional i wi h ega d o he ques ions p esen ed. Acknowledgemen o habi a dynamics Fo alse hea h i illa y conse a ion, a key sou ce o ins i u ional mis i a ose om he limi ed acknowledgmen o empo al habi a dynamics by o mal conse a ion ins i u ions. The legisla i e basis o alse hea h i illa y conse a ion, i.e., he species’ s a us as a species o s ic p o ec ion, was linked o a policy ool ha enabled egional ELY cen e s o use a p ocedu e o o icial habi a dema ca ions o es ic des uc i e ac i i ies a alse hea h i illa y habi a s (Minis y o he En i onmen 1996). Howe e , he esponden s conside ed habi a dema ca ions a con o e sial ool o conse ing si es p one o successional habi a dynamics because imposing limi a ions o p i a e landowne s’ land use igh s was expec ed o de e io a e p i a e landowne s’ a i udes owa d habi a main enance. The s a us as a species o s ic p o ec ion also enabled he Minis y o he En i onmen (1996) o p epa e a p og am o e i ing he species’ popula ions, bu such a p og am had no been p epa ed. None o he s a uses o he alse hea h i illa y in oked such o mal conse a ion policies ha would ha e p o ided access o esou ces o ools o ackling habi a dynamics o o gene a ing new habi a s o compensa e habi a loss. I was no un il he shi om legal amewo k o mo e in o mal adap i e go e nance a he na ional-le el and egional-le el policies ha successional dynamics a alse hea h i illa y habi a s was acknowledged as he main conse a ion challenge o he species, and i was add essed by he egional-le el au ho i ies in he o m o habi a quali y assessmen s and habi a es o a ion plans o deg aded si es. These ac i i ies we e suppo ed by Ecology and Socie y 22(2): 30 h ps://www.ecologyandsocie y.o g/ ol22/iss2/a 30/ egional p io i iza ion mee ings o endange ed species, an ou come o a Na ional Species Conse a ion Ac ion Plan p epa ed in 2010-2011 (Minis y o he En i onmen 2016, SYKE 2017). The mee ings p o ided guidance and commi men om na ional-le el ac o s, such as SYKE, Minis y o he En i onmen , Finnish Fo es y Se ice, and axon-speci ic expe s o he na ional endange ed species wo king g oups, in o egional-le el conse a ion planning. Fo alse hea h i illa y conse a ion, he mee ings p o ided suppo o he p io i iza ion o ELY cen e s’ conse a ion asks and commi ed key na ional- le el ac o s o he egional execu ion o alse hea h i illa y conse a ion ac i i ies on s a e-owned land. Howe e , he ag eemen s as such did no p o ide esou ces o he ELY cen e s o ca ying ou he p io i ized asks. The case o he alse hea h i illa y demons a es ha conse a ion p og ams may lack ools, p ocesses, and o mal policies o ackling habi a dynamics e en i hey ha e been designed o conse e species ha li e in dynamic landscapes. This can lead o a unc ional mis i , i conse a ion ins i u ions a e op imized o ackling di e en h ea s han wha a e he p ima y causes o endange men o he species in ques ion. We sugges he ollowing ques ions o assessing he le el o acknowledgmen o habi a dynamics in conse a ion ins i u ions: 1. A e habi a dynamics acknowledged as a key conse a ion challenge o he managed sys em bo h in he o mal and in o mal conse a ion policies? 2. Does such acknowledgmen ex end o all ele an spa ial scales o policy making, e.g., go e nmen al le el, egional le el, and local le el? 3. Has such acknowledgmen led o de ined a ge s and ask p io i iza ion ha help wi h esponding o habi a dynamics? 4. Has such acknowledgmen led o he p o ision o ope a ional ools and esou ce alloca ion ha help wi h esponding o habi a dynamics? I o mal conse a ion policies o in o mal policies a he go e nmen al le el do no acknowledge habi a dynamics as a conse a ion challenge, egional- o local-le el ac o s may ha e es ic ed oppo uni ies o ackle habi a dynamics because o he ask p io i iza ions and esou ce alloca ions de ined in he go e nmen al-le el decision making. The e o e, a aining high ins i u ional i may equi e ha acknowledgmen o habi a dynamics is ex ended o hose le els and o ms o go e nance ha con ol local-le el ask p io i iza ion and esou ces (Folke e al. 2005, Va n and Vedeld 2012). Ope a ional ools and esou ces o ackling habi a dynamics could be in oked i he challenge o habi a dynamics was acknowledged in he documen ed causes o h ea , h ea ac o s (Rassi e al. 2010), h ea ening p ocesses (Ho mann e al. 2008), o ecommended conse a ion ac ions (Rod igues e al. 2006) in he espec i e Red Lis documen a ion o in he objec i es o eco e y o a species-speci ic eco e y plan (McDonald e al. 2015). Al e na i ely, in o mal go e nmen al- le el policies, such as in o ma ion sha ing and decision-making p ac ices ha a e no de ined in he legisla ion, could be e ised so ha such a ge s, p io i ies, ope a ional ools, and esou ces a e ensu ed ha enable he managemen o habi a dynamics. A mo e ho ough solu ion could equi e a e o m o conse a ion policies o allow adap i e managemen o complex sys ems (G een e al. 2015). Moni o ing o and esponding o changes in he habi a ne wo k The esponden s did no ha e clea and uni ied iews on how p oac i ely ELY cen e s should collec in o ma ion on he alse hea h i illa y and on changes in he numbe s, sizes, quali ies, and loca ions o i s habi a pa ches o assess he adequacy o i s habi a ne wo k, because he legisla ion did no explici ly pose esponsibili ies on his ma e o any o ganiza ion. Regional au ho i ies esponsible o hea h i illa y conse a ion, howe e , collec ed in o ma ion o alse hea h i illa y sigh ings and he loca ions o alse hea h i illa y habi a s wi h he help o con ac o s and olun ee s, such as ci izen expe s om he local En omological Socie y. In Pi kanmaa, whe e he conse a ion his o y o he species was decades long, egional au ho i ies su eyed he meadow ne wo k wi h he help o con ac o s once a decade. The su eys included a quali a i e classi ica ion o habi a si es acco ding o he u gency o hei es o a ion. Su eys we e ollowed by habi a es o a ion p og ams ha included con ac s o habi a landowne s and making habi a es o a ion plans wi h coope a i e landowne s. Res o a ion p og ams mo ed annually om one sub egion o ano he o gain syne gies om coope a ing wi h mul iple landowne s a he same ime pe sub egion; his enabled in o ma ion-sha ing mee ings and he change o expe ise be ween in ol ed landowne s. The decadal hy hm o habi a ne wo k su eys in Pi kanmaa was d i en by esou ce a ailabili y in ela ion o he ex en o he habi a ne wo k o be main ained. Based on exis ing knowledge o habi a succession a alse hea h i illa y meadows (In ke 2003, Fab i ius and McB ide 2017), he empo o habi a dynamics a alse hea h i illa y habi a s was, howe e , o en as e han his. Consequen ly, epo s o habi a ne wo k su eys indica ed ha la ge p opo ions o habi a s had al eady been des oyed in he p ocess o habi a succession by he imes o he su eys (Kekkonen and Rönkä 2009). This gene a ed a empo al mis i o alse hea h i illa y conse a ion because in o ma ion lags p e en ed he iming o habi a main enance ac i i ies wi hin such empo al windows o oppo uni y (Folke e al. 2007) ha would ha e p e en ed local ex inc ions o alse hea h i illa y popula ions a deg aded habi a s. The spa io empo al agg ega ion o he esul ing habi a es o a ion p og ams (Fig. 1) was ano he po en ial sou ce o spa io empo al mis i . Synch onized es o a ion o nea by habi a s was likely o gene a e a spa io empo al pa e n in which habi a s loca ed close o each o he had synch onized successional dynamics. Acco ding o pas s udies o dynamic habi a s, such spa ial agg ega ion o habi a dynamics can ei he inc ease o dec ease species’ me apopula ion sizes in dynamic landscapes, depending on hei li e his o y ai s (Johs and D echsle 2003, Hinsch and Poe hke 2007, Elkin and Possingham 2008). I was no known which one was he case o he alse hea h i illa y. E ec i e managemen o habi a ne wo k dynamics bene i s om he collec ion o spa ial and empo al esilience assessmen da a (Allen e al. 2016) ha can be used o build knowledge on he sys em’s dynamics and c i ical h esholds. This knowledge can hen be used o de e mine he igh igge s and scales o ac ion o conse a ion managemen in he case o pe u ba ions and Ecology and Socie y 22(2): 30 h ps://www.ecologyandsocie y.o g/ ol22/iss2/a 30/ g adual sys em changes. We sugges he ollowing ques ions o assessing he i o moni o ing and esponding o changes in dynamic habi a ne wo ks: Fig. 1. Annual unding by he Minis y o he En i onmen o he Pi kanmaa ELY cen e o alse hea h i illa y conse a ion in 2005-2014. Dashed lines depic he lowe and uppe bounds o es ima ed annual unds o he yea s 2005-2010, o which exac eco ds pe species we e no a ailable. The inc eased unding in 2011-2014 e lec s a la ge-scale habi a es o a ion p og am ha ollowed he 2009 habi a ne wo k su ey and consis ed o si e main enance, landowne con ac s, and he de elopmen o new, expe imen al habi a main enance me hods. ELY cen e , Cen e o he Economic De elopmen , T anspo and he En i onmen . 1. Do conse a ion manage s collec in o ma ion on habi a ne wo k dynamics? 2. Is he a e age ime in e al o ecei ing o collec ing such da a sho e han he expec ed li e ime o mos empo a y habi a s? 3. Is in o ma ion o empo al habi a dynamics collec ed using such s anda dized measu es ha ensu e he compa abili y o da a ac oss yea s and su eyo s? 4. Do conse a ion manage s ecei e in o ma ion om landowne s o olun ee s on changes in landowne ship, land use ype, o des uc ion o habi a s? 5. Is he e a sys ema ic me hod o sea ching o eme ging habi a s? 6. A e conse a ion manage s able o an icipa e changes in he habi a ne wo k based on si e main enance his o ies and/o modeled empo al dynamics o habi a s? E ec i e managemen o habi a ne wo k dynamics equi es imely in o ma ion on changes in he habi a pa ch ne wo k so ha conse a ion managemen plans can be i e a ed based on up- o-da e da a on he sys em s a e (G een e al. 2015). Such in o ma ion may be collec ed ia egula pa ch ne wo k su eys ha a e o ganized wi hin ime in e als ha lea e a window o oppo uni y o ca y ou main enance ac i i ies be o e local popula ion ex inc ions ake place a deg ading habi a pa ches. The imeliness o a ailable in o ma ion could be u he inc eased by se ing up mechanisms ha enable landowne s o epo changes conce ning habi a pa ches loca ed a hei p ope ies, such as habi a main enance ac i i ies, habi a des uc ion, changes in landowne ship, and changes in land use plans a he p ope y (Pocewicz e al. 2008, Daye e al. 2016). Addi ionally, depending on he habi a ype in ques ion, he sea ch o new, eme ged habi a pa ches could be ca ied ou ei he by conduc ing egula ield su eys o by s udying land use eco ds ha may e eal p obable loca ions o habi a eme gence. The use o s anda dized measu es enables he accumula ion o such da a on pa ch and pa ch ne wo k dynamics ha is compa able ac oss yea s and su eyo s and ha can hus be used o p edic ing sys em dynamics, h esholds, and u u e de elopmen . Fo ins ance, i epea ed s anda dized measu emen s o habi a sizes and quali ies can be complemen ed wi h sys ema ic eco ds o habi a main enance and land use his o ies, p edic i e models can be de eloped o es ima ing how managemen in e en ions impac habi a quali ies and succession (Pöy y e al. 2006, Taylo and Mo ec o 2009). Such p edic i e models could be used o es ima e he annual need o main enance esou ces o he sys em and o de ine sui able ime in e als o u u e habi a ne wo k su eys. Managemen o esou ces o luc ua ing esou ce needs In alse hea h i illa y conse a ion, bo h esou ce needs and esou ce a ailabili y luc ua ed in ime. Annual esou ce needs a he h ee ELY cen e s depended on which ac i i ies ela ed o he alse hea h i illa y, i.e., species moni o ing, habi a ne wo k su eys, o es o a ion p og ams, we e planned o he gi en yea and how many habi a si es we e planned o be main ained o es o ed using he esou ces o he ELY cen e s. Al hough no ea ma ked unding exis ed o hese ac i i ies, he budge o he Minis y o he En i onmen con ained wo clauses ha could be used o co e cos s o ac i i ies ela ed o he alse hea h i illa y. Resou ces a ailable in he Minis y o he En i onmen o he wo clauses luc ua ed annually depending on he s a e budge and on empo ally luc ua ing p io i ies, such as he execu ion o la ge-scale conse a ion p og ams o upda ing he Finnish Red Lis o Endange ed Species, which usually ook place o 3 yea s e e y 10 yea s. The annual unding ecei ed by he ELY cen e s was he e o e o en su p isingly small, some imes e en less han hal o he sum applied o alse hea h i illa y conse a ion. Consequen ly, ELY cen e s had o p io i ize di e en species in di e en yea s o ge adequa e unding o execu ing species- speci ic p ojec s. Access o main enance unding o alse hea h i illa y habi a s also a ied acco ding o he ype o habi a owne ship, a me s a us, landowne ’s abili y o ca y ou habi a main enance independen ly, and dema ca ion s a us o he habi a (Fig. 2). The 5-yea o 10-yea ag i-en i onmen al suppo con ac s p o ided an impo an sou ce o unding o he main enance o alse hea h i illa y habi a s by landowne s, gi en he p opo ionally la ge amoun s o unding a ailable in he ag i-en i onmen al suppo and he long e m o he suppo con ac s. Howe e , his Eu opean Union–based sou ce o unding was only a ailable o Ecology and Socie y 22(2): 30 h ps://www.ecologyandsocie y.o g/ ol22/iss2/a 30/ a me s en i led o a ming suppo . The u iliza ion o ag i- en i onmen al schemes o he main enance o alse hea h i illa y habi a s also depended on whe he habi a s o be main ained we e loca ed on a p ope y ha me he condi ions o he suppo and whe he habi a s we e la ge enough so ha he a ea-based ag i-en i onmen al suppo co e ed habi a main enance cos s. The applica ion p ocess also equi ed high landowne mo i a ion and ime in es men om he pe sonnel o he ELY cen e , which limi ed he usage o his sou ce o unding. The unp edic abili y and in lexibili y o a ailable main enance unding he e o e caused a spa io empo al mis i in he managemen o habi a ne wo k dynamics o alse hea h i illa y conse a ion because i limi ed conse a ion manage s’ possibili ies o plan and s ee esou ce alloca ion. Fig. 2. Sou ces o habi a main enance unding o a alse hea h i illa y habi a depending on he ype o landowne , dema ca ion s a us and a me s a us o he habi a , and he landowne ’s abili y o ca y ou habi a main enance independen ly. ELY cen e , Cen e o he Economic De elopmen , T anspo and he En i onmen . E ec i e managemen o habi a ne wo k dynamics would equi e ha he esou ces a ailable o habi a main enance can be alloca ed ac oss ime and space acco ding o wha minimizes habi a discon inui ies o he species o in e es . We sugges he ollowing ques ions o assessing he i o managing esou ces o luc ua ing esou ce needs: 1. Do conse a ion manage s ha e da a on how spa ial o empo al luc ua ions o he habi a ne wo k a ec he species o in e es ? 2. Do conse a ion manage s alloca e main enance esou ces acco ding o long- e m spa ial conse a ion plans? 3. Can conse a ion manage s bu e and dis ibu e managemen esou ces ac oss yea s o maximize he e ec i eness o esou ce use? 4. Can conse a ion manage s alloca e main enance esou ces lexibly in e ms o loca ion o maximize he e ec i eness o esou ce use? Being able o alloca e main enance esou ces ac oss ime and space o he species o in e es equi es an unde s anding o how he species’ me apopula ion dynamics a e a ec ed by di e en spa io empo al pa e ns o habi a a ailabili y. Wi hou de ailed s udies o he managed sys em, such pa e ns a e no sel -e iden . Fo ins ance, spa ial agg ega ion o dis u bances may o may no alle ia e he de imen al e ec s o local habi a des uc ion o dis u bance-dependen species depending on species-speci ic ai s (Johs and D echsle 2003, Hinsch and Poe hke 2007, Elkin and Possingham 2008), and in ensi e main enance o a smalle habi a ne wo k may o may no be bene icial in compa ison o c ea ing new habi a s (Ross e al. 2008). Ideally, mul iple ealis ic scena ios o habi a ne wo k dynamics could be de eloped o he landscapes being managed, and he me apopula ion dynamics o he species o in e es could be modeled using hese al e na i e scena ios. As in many cases, esou ce limi a ion may p e en he de elopmen o applica ion o such p edic i e ools, bu exis ing models ha ha e been de eloped o species o simila li e his o y ai s could be used as guidance o de e mining e ec i e spa io empo al managemen s a egies. In he absence o any es ima es o a species’ me apopula ion esponses o landscape dynamics, managemen could aim a maximizing pa ch li e imes while minimizing changes in habi a numbe s and loca ions whene e possible because hese s a egies ha e gene ally been shown o main ain highe popula ion sizes han apidly changing habi a ne wo ks (Johs e al. 2011, Van Tee elen e al. 2012). A he same ime, conse a ion managemen on p i a e land akes place in he eali y o mul iple cons ain s, such as esou ce sho ages and he eliance on landowne coope a ion (Opdam e al. 2008), which may p e en ollowing he mos e ec i e managemen s a egies. Fo such cases, he e is a need o p ac ical decision-making ules o ools ha enable planning o he iming and placemen o habi a main enance ac i i ies gi en such cons ain s o unde unce ain y. Conse a ion manage s’ abili y o alloca e esou ces ac oss ime and space could be imp o ed i main enance budge s we e lexible in ime, and manage s could decide whe he o sa e o ollowing yea s o spend he ull budge (see, e.g., D echsle e al. 2007). This would enable manage s o u ilize mo e esou ces du ing yea s o need o la ge amoun s o main enance unding o o la en he luc ua ions o a ailable conse a ion unds ac oss yea s. Imp o ing conse a ion manage s’ abili y o alloca e main enance esou ces lexibly in e ms o loca ion would equi e ei he a gene al-le el alloca ion o unds o he main enance o he habi a ne wo k, which would educe manage s’ dependency on applying o al e na i e sou ces o unding, o , in he case o ea ly successional habi a s, mo e lexible ules o he use o ag i- en i onmen al suppo . Annual unding alloca ions should also be handled a such scale o go e nance ha allows lexible de e mina ion o he igh scale o ac ion (G een e al. 2015), o ins ance ealloca ion o esou ces be ween managemen egions in he case o la ge pe u ba ions a pa icula egions. Scaling o ac i i ies h ough lexible collabo a ions In he h ee ELY cen e s, esou ce sho ages es ic ed pe sonnel esou ces ha could be spen on alse hea h i illa y conse a ion, and mos esou ces wen in o handling bu eauc a ic ma e s o being in con ac wi h landowne s. The e o e, alse hea h i illa y dis ibu ion and habi a ne wo k su eys, as well as habi a Ecology and Socie y 22(2): 30 h ps://www.ecologyandsocie y.o g/ ol22/iss2/a 30/ Table 1. Numbe s o endange ed species in he Finnish Red Lis o Endange ed Species 2010 (Rassi e al. 2010) in all axa (le ), in Lepidop e a (middle), and in ascula plan s ( igh ) o which he majo cause o endange men is habi a su oca ion ( h ea code N), g ouped acco ding o hei endange men s a us: c i ically endange ed (CR), endange ed (EN), and ulne able (VU). The able lis s he numbe s o species h ea ened by habi a o e g ow h in he espec i e axa ha a e classi ied as species unde s ic p o ec ion and would, based on he opinions o wo axon-speci ic expe s, also equi e species-speci ic conse a ion plans like he alse hea h i illa y. All axa Lepidop e a Vascula plan s CR EN VU To al CR EN VU To al CR EN VU To al To al numbe o endange ed species h ea ened by habi a o e g ow h 107 388 389 884 30 151 92 273 9 47 37 93 O which, species unde s ic p o ec ion 54 192 25 271 17 74 1 92 5 29 2 36 O which, species in need o species- speci ic conse a ion plans (es ima ed by expe 1) 6 21 1 28 O which, species in need o species- speci ic conse a ion plans (es ima ed by expe 2) 2 16 1 19 main enance planning and in o ma ion sha ing o landowne s, we e handed o ex e nal con ac o s o olun ee s. The amoun o con ac ed wo k a ied annually depending on he amoun o unding a ailable, bu also acco ding o he landowne ship o habi a s o be main ained: Landowne s a ied om ac i e a me s o e i ed a me s and o people li ing o wo king in u ban a eas, and habi a s we e ei he main ained by landowne s hemsel es o needed o be main ained by con ac o s. Collabo a ion, when i was needed, had o be scaled up as : In o ma ion on unding alloca ions by he Minis y o he En i onmen was usually a ailable in Feb ua y, and sp ing ac i i ies a main ained si es had o be ca ied ou be o e he s a o he g ow h season in Ap il o May. Addi ionally, ELY cen e s needed expe ienced con ac o s ha could ca y ou he needed asks independen ly. This did no cause majo p oblems in Pi kanmaa, whe e he e was no sho age o expe ienced con ac o s and whe e plans and con ac s could be scaled as acco ding o he amoun o a ailable unding based on known bes p ac ices om pas collabo a ions. In he o he wo adminis a i e egions, which had ewe ac o s in he ci il socie y, expe ienced con ac o s did no exis , and he sho - e m na u e o annually con ac ed wo k complica ed he de elopmen o expe ise among he ew po en ial collabo a o s. The sho - e m alloca ion o main enance esou ces caused a isk o empo al mis i in he execu ion o main enance ac i i ies because he sho ime ames in which p ojec s had o be es ablished could ha e ou paced he empo o inding con ac o s. This, in u n, could ha e led o de iciencies in he execu ion o planned main enance ac i i ies. Such mis i could be alle ia ed ia he buildup o collabo a ion ne wo ks ha enable as scaling o ac i i ies o by widening he ime ames o esou ce a ailabili y o conse a ion manage s. We sugges he ollowing ques ions o assessing he i o scaling ac i i ies h ough lexible collabo a ions: 1. Do conse a ion manage s ha e access o an adequa e supply o expe ienced in-house pe sonnel o ex e nal con ac o s ha can be in ol ed in habi a main enance ac i i ies when needed? 2. Is he ime ame a ailable om unding alloca ions o he p e e ed ime o execu ion o main enance ac i i ies adequa e o ac i a ing a scalable wo k o ce? Ha ing he possibili y o dis ibu e main enance esou ces lexibly o e mul iple yea s would enable longe e m planning o esou ce use and longe planning ho izons o scaling up ope a ions. Thus, i would also enable he c ea ion o long- e m managemen con ac s, commi men , and capaci y building wi h local con ac o s. The case o he alse hea h i illa y in ela ion o o he species Acco ding o he es ima es o wo axon-speci ic expe s, Finland has a leas 19 and 28 endange ed species in he s udied axa o ascula plan s and Lepidop e a, espec i ely, ha a e simila o he alse hea h i illa y wi h espec o hei s a us as endange ed species (CR, EN, o VU), s a us as species o s ic p o ec ion, main cause o endange men , and es ima ed need o species- speci ic conse a ion plans (Table 1). These es ima es co espond o 10.2% and 20.4% o all endange ed species in he espec i e axa ha a e h ea ened by successional habi a dynamics. As hese numbe s es ima e he numbe s and ac ions o species o which conse a ion is likely o aise simila challenges and po en ial ins i u ional mis i s as hose iden i ied o he alse hea h i illa y, he implica ions and ecommenda ions we iden i ied a e o gene al applica ion and impo ance. CONCLUSION The ou ins i u ional a angemen s iden i ied in ou case s udy, i.e., acknowledgmen o habi a dynamics, moni o ing o and esponding o changes in he habi a ne wo k, managing esou ces o luc ua ing esou ce needs, and scaling o ac i i ies h ough lexible collabo a ions, s ess he impo ance o ins i u ional lexibili y and adap i e go e nance in he conse a ion o dynamic habi a ne wo ks. High pe o mance in he conse a ion o dynamic sys ems equi es he abili y o espond as o po en ially eme ging habi a discon inui ies ha could be de imen al o species being conse ed. Ou indings hus ag ee wi h many p e ious indings on he e ec i e managemen o social-ecological sys ems, which highligh he impo ance o en i onmen al moni o ing (Cox 2012), adap i e capaci y (Folke e al. 2007, Ga mes ani e al. 2013, Allen e al. 2016), c oss-scale Ecology and Socie y 22(2): 30 h ps://www.ecologyandsocie y.o g/ ol22/iss2/a 30/ in e ac ions (Hansen 2014), and he u iliza ion o landowne s’ knowledge (Olsson e al. 2007, G een e al. 2015) in he a ainmen o high ins i u ional i in hese con ex s. As demons a ed by he expe es ima es on he gene alizabili y o ou case s udy, habi a dynamics may be an o e looked conse a ion challenge ha calls o new kinds o ins i u ional a angemen s ha can main ain habi a a ailabili y o species li ing in dynamic landscapes. In he e a o clima e change, such lexibili y may p o e bene icial o a la ge ex en o species han hose li ing in ea ly successional habi a s. In he ace o po en ially la ge-scale pe u ba ions, such as possible ex eme clima e e en s, conse a ion manage s could aim a inc easing, no only main aining, sys em esilience by inc easing he he e ogenei y and connec i i y o habi a pa ch ne wo ks and acili a ing hei spa ial ex ension owa d he di ec ion o p edic ed sui able clima es o he u u e. T ans o ma ion om adi ional conse a ion managemen , which is based on igid ules and managemen o sys em pieces as independen uni s (Allen e al. 2016), is, howe e , likely o equi e legal e o m be o e he adap i e managemen o complex sys ems would be ully enabled in conse a ion ins i u ions (G een e al. 2015). 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Should he alloca ion o habi a main enance unding o endange ed species be based on some hing else han he ou comes o he p io i iza ion mee ings? I yes, wha ? Species o s ic conse a ion 1. Wha does he conse a ion esponsibili y o he species o s ic conse a ion conce n mean o you, wha is he conc e e goal o he conse a ion ac i i ies and on who’s esponsibili y is i o plan (a) he mee ing o he a o able conse a ion s a us, (b) main enance ac i i ies needed, (c) o e all conse a ion planning? 2. Which unding ins umen should be used o co e he habi a main enance cos s o endange ed species ha a e h ea ened by habi a o e g ow h? Fo (a) p i a e ag icul u al land, (b) p i a e conse a ion a eas, (c) publicly owned land? 3. Wha ools o esou ces should be enables o he ELY cen e s, so ha he a o able conse a ion s a us o he alse hea h i illa y could be me in a socially meaning ul way? 4. Fo how many o he species is habi a o e g ow h he p ima y h ea ?