scieee Open visual document viewer

Forest Loss in Protected Areas and Intact Forest Landscapes: A Global Analysis

Heino, Matias,Kummu, Matti,Makkonen, Marika,Mulligan, Mark,Verburg, Peter H.,Jalava, Mika,Räsänen, Timo A.

Abstract

In spite of the high importance of forests, global forest loss has remained alarmingly high during the last decades. Forest loss at a global scale has been unveiled with increasingly finer spatial resolution, but the forest extent and loss in protected areas (PAs) and in large intact forest landscapes (IFLs) have not so far been systematically assessed. Moreover, the impact of protection on preserving the IFLs is not well understood. In this study we conducted a consistent assessment of the global forest loss in PAs and IFLs over the period 2000-2012. We used recently published global remote sensing based spatial forest cover change data, being a uniform and consistent dataset over space and time, together with global datasets on PAs' and IFLs' locations. Our analyses revealed that on a global scale 3% of the protected forest, 2.5% of the intact forest, and 1.5% of the protected intact forest were lost during the study period. These forest loss rates are relatively high compared to global total forest loss of 5% for the same time period. The variation in forest losses and in protection effect was large among geographical regions and countries. In some regions the loss in protected forests exceeded 5% (e.g. in Australia and Oceania, and North America) and the relative forest loss was higher inside protected areas than outside those areas (e.g. in Mongolia and parts of Africa, Central Asia, and Europe). At the same time, protection was found to prevent forest loss in several countries (e.g. in South America and Southeast Asia). Globally, high area-weighted forest loss rates of protected and intact forests were associated with high gross domestic product and in the case of protected forests also with high proportions of agricultural land. Our findings reinforce the need for improved understanding of the reasons for the high forest losses in PAs and IFLs and strategies to prevent further losses.

Full text

RESEARCH ARTICLE Fo es Loss in P o ec ed A eas and In ac Fo es Landscapes: A Global Analysis Ma ias Heino 1 , Ma i Kummu 1 *, Ma ika Makkonen 2 , Ma k Mulligan 3 , Pe e H. Ve bu g 4 , Mika Jala a 1 , Timo A. Räsänen 1 1Wa e and De elopmen Resea ch G oup, Aal o Uni e si y, Tie o ie 1E, 02150, Espoo, Finland, 2Na u al Resou ces Ins i u e Finland (Luke), Jokiniemenkuja 1, 01301, Van aa, Finland, 3Depa men o Geog aphy, King's College London, S and, London, WC2R 2LS, Uni ed Kingdom, 4En i onmen al Geog aphy G oup, VU Uni e si y Ams e dam, de Boelelaan 1087, 1081 HV, Ams e dam, he Ne he lands *[email p o ec ed] Abs ac In spi e o he high impo ance o o es s, global o es loss has emained ala mingly high du - ing he las decades. Fo es loss a a global scale has been un eiled wi h inc easingly ine spa ial esolu ion, bu he o es ex en and loss in p o ec ed a eas (PAs) and in la ge in ac o es landscapes (IFLs) ha e no so a been sys ema ically assessed. Mo eo e , he impac o p o ec ion on p ese ing he IFLs is no well unde s ood. In his s udy we conduc ed a con- sis en assessmen o he global o es loss in PAs and IFLs o e he pe iod 2000–2012. We used ecen ly published global emo e sensing based spa ial o es co e change da a, being a uni o m and consis en da ase o e space and ime, oge he wi h global da ase s on PAs’ and IFLs’loca ions. Ou analyses e ealed ha on a global scale 3% o he p o ec ed o es , 2.5% o he in ac o es , and 1.5% o he p o ec ed in ac o es we e los du ing he s udy pe iod. These o es loss a es a e ela i ely high compa ed o global o al o es loss o 5% o he same ime pe iod. The a ia ion in o es losses and in p o ec ion e ec was la ge among geog aphical egions and coun ies. In some egions he loss in p o ec ed o es s exceeded 5% (e.g. in Aus alia and Oceania, and No h Ame ica) and he ela i e o es loss was highe inside p o ec ed a eas han ou side hose a eas (e.g. in Mongolia and pa s o A ica, Cen al Asia, and Eu ope). A he same ime, p o ec ion was ound o p e en o es loss in se e al coun ies (e.g. in Sou h Ame ica and Sou heas Asia). Globally, high a ea- weigh ed o es loss a es o p o ec ed and in ac o es s we e associa ed wi h high g oss domes ic p oduc and in he case o p o ec ed o es s also wi h high p opo ions o ag icul u al land. Ou indings ein o ce he need o imp o ed unde s anding o he easons o he high o es losses in PAs and IFLs and s a egies o p e en u he losses. In oduc ion Fo es s play a c ucial ole in sus aining li e on ea h. They main ain ecological di e si y, egu- la e clima e, s o e ca bon, p o ec soil and wa e and p o ide esou ces and li elihoods o he wo ld’s popula ion [1–4]. Despi e he inc easing awa eness o he impo ance o hese ecosys- ems, global de o es a ion a es ha e emained ala mingly high o e he pas decades [2]. PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 1/21 OPEN ACCESS Ci a ion: Heino M, Kummu M, Makkonen M, Mulligan M, Ve bu g PH, Jala a M, e al. (2015) Fo es Loss in P o ec ed A eas and In ac Fo es Landscapes: A Global Analysis. PLoS ONE 10(10): e0138918. doi:10.1371/jou nal.pone.0138918 Edi o : Madhu Anand, Uni e si y o Guelph, CANADA Recei ed: Ma ch 16, 2015 Accep ed: Sep embe 4, 2015 Published: Oc obe 14, 2015 Copy igh : © 2015 Heino e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License, which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed. Da a A ailabili y S a emen : All ele an da a a e wi hin he pape and i s Suppo ing In o ma ion iles. Funding: The wo k was unded by Maa- ja esi ekniikan uki y. In addi ion, MK was unded by he Academy o Finland p ojec SCART (g an no. 267463) and MMa by he Academy o Finland p ojec G een Economy and Policies (g an no. 257919). PHV hanks he ERC g an ag eemen n . 311819 (GLOLAND) and he FP7 p ojec LUC4C o suppo . The unde s had no ole in s udy design, da a collec ion and analysis, decision o publish, o p epa a ion o he manusc ip . In yea 2010 o es co e ed a ound 40 million km 2 o 31% o he global land a ea acco ding o coun y epo s [2]. Es ima es o global o es loss a es wi hin he pas decade a y be ween 130,000 km 2 /y [2] and 177,000 km 2 /y [5]. A he same ime, some e o es a ion and na u al eg ow h ha e occu ed, leading o ne o es loss epo s anging be ween 52,000 km 2 /y and 115,000 km 2 /y , espec i ely. Al hough o es loss is s ill ema kably high, i has shown some signs o decline: FAO [2] epo s ha du ing he 1990s, he o es loss a e was 160,000 km 2 /y ; whe eas be ween 2000–2010, he a e was 130,000 km 2 /y . Acco ding o FAO [2] he opics we e he only domain whe e he a e o o es loss inc eased in he i s decade o he 21 s cen- u y compa ed o 1990s: de o es a ion inc eased in he opics by 2,101 km 2 /y on a e age ac oss he decade. De o es a ion in he opics accoun ed o 32% o global o es loss wi hin he pe iod o 2000–2012 [5]. Howe e , he e a e also posi i e signs in he opics: he a e o o es loss in he B azilian Amazon has declined in ecen yea s [6,7]. The main d i e s o global de o es a ion a e linked o expansion o ag icul u e, wood ex ac- ion, in as uc u e ex ension, popula ion g ow h, and expansion o ag icul u e [8–13]. The dom- inan d i e s, howe e , a y among he egions [8–13]. In addi ion o ag icul u e and popula ion g ow h, a me a-analysis o 117 de o es a ion s udies by Fe e i-Gallon and Busch [9]sugges s ha de o es a ion is gene ally lowe in high, s eep and we a eas while i is highe in a eas whe e o es s a e close o oads and u ban a eas. De o es a ion has also shi ed om a dominan ly s a e ini ia ed o an en e p ise d i en p ocess in he opics be ween 1970 and 2000 [14]. Ala ming de o es a ion a es combined wi h he inc easing awa eness o he impo ance o o - es s ha e esul ed in exponen ial g ow h o he wo ld’s p o ec ed a eas (PA) o e he pas decades [15–17]. Schmi e al. [18] es ima e he global o es co e using ea h obse a ion sa elli e da a om MODIS2005 and he ex en o p o ec ed o es using he Wo ld Da abase o P o ec ed A eas (WDPA) o he yea 2008. They ind ha 7.7% o global o es co e ell wi hin IUCN’s ou s ic es p o ec ion ca ego ies (I-IV) and 13.5% wi hin IUCN’s all six p o ec ion ca ego ies (I-VI) (IUCN p o ec ion ca ego ies a y om s ic ly p o ec ed a eas (I) o p o ec ed a eas wi h sus ain- able esou ce use (VI); o mo e de ailed ca ego y de ini ion see [19]).Schmi e al.[18] u he conclude ha o es p o ec ion a ied g ea ly be ween di e en egions and o es ypes, and ha o es p o ec ion in p io i y a eas, such as biodi e si y ho spo s, was insu icien . Conside able e o s o p o ec ion a e a ge ed o he p ima y o es s and la ge in ac o es landscapes (IFLs, i.e. unb oken expanse o na u al ecosys ems). These o es a eas play c ucial oles in sus aining ecological di e si y [2,20]. FAO [2] es ima es ha p ima y o es s accoun ed 14 million km 2 (36% o he global o al o es s) in yea 2010, ha ing dec eased ala ming 400,000 km 2 o e he pe iod o 2000–2010 (annual a e 0.4%). Po apo e al. [20] epo ha he ex en o IFL is 13.1 million km 2 . Vas majo i y o he IFLs a e ound ei he in dense T opi- cal and Sub-T opical o es s (45% o wo ld o al) o in Bo eal Fo es s (44%). The lowes p o- po ion o IFL was ound in Tempe a e B oadlea and Mixed Fo es s. Po apo e al. [20] u he ind ha 18.9% o IFLs a e unde p o ec ion o IUCN p o ec ion ca ego ies I-VI and only 9.7% o IFLs is s ic ly p o ec ed unde IUCN p o ec ion ca ego ies I-III. The p o ec ion e ec o PAs is, howe e , ques ioned. The PAs a e conside ed in many cases o be biased in hei loca ion, meaning ha PAs a e loca ed in a eas ha a e unlikely o ace land con e sion p essu es [21]. Joppa and P a [22] e eal ha a majo i y o he PA ne wo ks a e loca ed in high ele a ions, s eep slopes and a om oads and ci ies. Joppa and P a [21] a gue ha his bias has esul ed in o e es ima ions in he p o ec ion e ec o PAs. Local case s udies (e.g. [23]) suppo hese global indings. Joppa and P a [24] use ‘ma ching’app oach ha a emp s o a oid he o e es ima ion o p o ec ion e ec o PAs in 147 coun ies by com- pa ing p o ec ed and non-p o ec ed a eas wi h simila land cha ac e is ics. They ind ha ma ching educed he p o ec ion e ec in 80% o he coun ies compa ed o an assessmen wi hou ma ching. Al oge he hey ind ha he p o ec ion educed con e sion o na u al land P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 2/21 Compe ing In e es s: The au ho s ha e decla ed ha no compe ing in e es s exis . co e in 75% o he assessed coun ies. P o ec ed a eas a e also epo ed o become inc easingly isola ed, especially in opics [25]. This is ala ming because smalle p o ec ed a eas a e o en unde g ea h ea [26] and isola ion o hese a eas es ic s he habi a size, i.e. limi s he su - i al o a g ea numbe o auna and lo a species (e.g. [27,28]).Al hough he unde s anding o o es loss a a global scale wi h an inc easing spa ial esolu ion is g owing apidly [5], he global o es loss in PAs and in IFLs a e no assessed wi h de ailed and uni o m da ase s ha allow consis en o es ex en compa isons o e space and ime. So a only egional analyses exis a his le el, o example, o Indonesia [29]. Fu he mo e, he success o p o ec ion in p ese ing he IFLs is no ye well unde s ood. The e o e, in his s udy, we aim o conduc a consis en and spa ially explici assessmen o he global o es loss, pa icula ly wi hin p o- ec ed and in ac o es s by ocusing on he a ia ion among he coun ies be ween 2000 and 2012. Addi ionally, we aim o s udy whe he socio-economic indica o s can o e po en ial explana ion o hese obse ed global o es losses in PAs and IFLs. We hypo hesised ha i) he ex en o o es loss wi hin PAs and IFLs a ies s ongly among he coun ies bu is always less mani es ed wi hin PAs han ou side o hem, and ii) coun y le el indica o s o popula ion size, land use change and s a e economy can o e po en ial explana ion o he obse ed global o es losses in PAs and IFLs. The key e minology used in his s udy is explained in Table 1. Ma e ials and Me hods To conduc he assessmen , we combined ou global da ase s (Fig 1;Table 2): o es ex en based on he Global Fo es Change (GFC) da a [5], o es loss based on he GFC da a [5], he Wo ld Da abase o P o ec ed A eas (WDPA) [30] and he global La ge In ac Fo es Land- scapes (IFL) da a [20]. F om GFC we assessed he o es ex en and o es loss while WDPA and IFL da ase s we e used o de e mine how much o ha loss ook place in p o ec ed a eas and in ac o es landscapes espec i ely. The esul ing spa ial da a we e agg ega ed o coun y- scale and analysed wi h Weigh ed Leas -Squa es (WLS) eg ession analyses o assess whe he one o a se ies o socio-economic indica o s a e associa ed o o es loss pa e ns, possibly explaining hei occu ence. The o es loss esul s a e gi en a na ional and global scale and by geog aphical egions, while eg ession analyses we e conduc ed a na ional scale o co espond wi h he socio-economic indica o da a and he pu sued o es go e nance le el. Below he da a and me hods a e desc ibed in mo e de ail. Da a We used he GFC da ase [5] o map he o es ex en and o es loss (Fig 1A and 1B;Table 2). GFC da a a e based on Ea h obse a ion sa elli e da a and ha e a esolu ion o ~30 me e s a Table 1. Key e minology used in he s udy. Te m Defini ion Fo es Vege a ion alle han 5 m wi h 20% ee co e canopy h eshold using Global Fo es Change (GFC) da a by Hansen e al. [5]. Fo es loss ”S and eplacemen dis u bance o comple e emo al o o es canopy”as defined by Hansen e al. [5] in GFC da a. Fo es gain was no aken in o accoun . P o ec ed a ea (PA) P o ec ed a ea as in Wo ld Da abase on P o ec ed A eas (WDPA) [30]. P o ec ed o es Fo es wi hin PA (i.e. p o ec ed a ea). In ac o es landscape (IFL) ”An unb oken expanse o na u al ecosys ems wi hin a eas o cu en o es ex en ,wi hou signs o significan human ac i i y,and ha ing and a ea o a leas 500 km 2 ”as in IFL da ase by Po apo e al. [20]. In ac o es Fo es wi hin IFL (i.e. In ac o es landscape). P o ec ed in ac o es In ac o es wi hin PA (i.e. p o ec ed a ea). doi:10.1371/jou nal.pone.0138918. 001 P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 3/21 he equa o (i.e. 1 a c-second). The o es ex en in he GFC da a a e gi en as canopy co e pe - cen age pe ou pu g id cell o all ege a ion alle han 5 me e s o he yea 2000 [5]. The o - es loss in he GFC da ase is de ined as s and- eplacing dis u bance, o comple e emo al o ee canopy, which has occu ed be ween yea s 2000 and 2012 [5]. In his s udy, we used he same de ini ion o o es loss (Table 1). The GFC da ase p o ides g oss and ne o es loss da a. We chose o ocus on g oss o es loss ins ead o ne o es loss, as we belie e he o me e lec s be e he occu ence o o es dis u bances in p o ec ed and in ac o es s han he la - e . The use o ne o es loss could ha e masked some o he dis u bances h ough o es gain. I should be no ed ha he GFC da ase does no speci y he easons behind o es loss and hus, he epo ed o es loss inside p o ec ed a eas migh be due o di ec human ac ions (e.g. illegal logging, managemen s a egies) as well as na u al causes (e.g. o es i es, diseases, pes s Fig 1. Agg ega ed (A-B) and con e ed (C-D) da ase s used o he o es loss analyses. A) Fo es ex en in pe cen age o g id a ea a 1 km esolu ion, based on Global Fo es Change (GFC) da a [5]; B) Fo es loss in pe cen age o g id a ea a 1 km esolu ion, based on GCF da a [5]; C) P o ec ed a eas based on Wo ld Da abase o P o ec ed A eas (WDPA) [30], con e ed o 1 km esolu ion; and D) La ge In ac Fo es Landscapes (IFLs) based on da ase by Po apo e al [20], con e ed o 1 km esolu ion. doi:10.1371/jou nal.pone.0138918.g001 Table 2. A desc ip ion o he da ase s used in his s udy. GFC s ands o Global Fo es Change da ase . Da ase Spa ial ex en Time pe iod Spa ial esolu ion Re e ence GFC: Fo es loss La : 80N - 60S Lon: 90E - 90W 2000–2012 30 m (a equa o ; 1 a c-second x 1 a c-second) Hansen e al. [5] GFC: Fo es ex en La : 80N - 60S Lon: 90E - 90W 2000 30 m (a equa o ; 1 a c-second x 1 a c-second) Hansen e al. [5] Wo ld Da abase o P o ec ed A eas (WDPA) Global 2010 ( e e ence yea ) Vec o da a wi h a ying esolu ion IUCN and UNEP-WCMC [30] La ge In ac Fo es Landscapes (IFL) Global 2000 Vec o da a wi h app oxima e esolu ion o 1 km (scale: 1:1,000,000) Po apo e al. [20] doi:10.1371/jou nal.pone.0138918. 002 P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 4/21 and s o ms) [2,5,31,32]. Fo example, na u al o es i es a e epo ed o be he mos signi ican cause o o es loss in he bo eal o es s [33]. We used he WDPA da ase o map he p o ec ed a eas o he globe (Fig 1C;Table 2). The da ase is a join p ojec o IUCN and UNEP and i is s a ed o be he mos ex ensi e da ase on p o ec ed a eas con aining bo h na ionally and in e na ionally designa ed p o ec ed a eas [30]. The da ase is based on bo h au ho i y and communi y sou ces. We included only he p o- ec ed a eas epo ed by au ho i y sou ces in o ou assessmen . Fo he in ac o es a eas we used he global ex en o IFLs (Fig 1D;Table 2)[20]. The da a- se is c ea ed using a ious emo ely sensed da ase s and maps, including da a om MODIS and Landsa . They de ine IFL as an unb oken a ea o na u al ecosys ems wi hin o es s, wi hou signs o signi ican human ac i i y, and ha ing an a ea o a leas 500 km 2 . The e e ence yea o his da ase is 2000, i.e. he s a ing yea o ou analysis. Me hods: o es ex en and o es loss We conduc ed he analyses on 1 km esolu ion (a he equa o ; i.e. on 30 a c-second esolu- ion). All da ase s we e ei he agg ega ed ( o es ex en and o es loss) o con e ed (PA and IFL) o his esolu ion. The con e sion p ese ed he o iginal esolu ion o he inpu da a (Table 2) and he agg ega ion o coa se esolu ion was ca ied ou in a way ha o iginal in o - ma ion was no dis o ed (see below o mo e de ails). Be o e any da a agg ega ion, we ans o med he o iginal GFC o es co e da a in o boolean ype a 30 m esolu ion. Fo he ans o ma ion we used 20% ee canopy co e h eshold, simi- la ly o Po apo e al. [33], o de ine whe he he 30 m esolu ion g id cells a e classi ied as o es o no- o es . These boolean o es co e da a we e mul iplied by a su ace a ea o 30 m cell in ques ion. The o es ex en a ea da a (in km 2 ) we e hen agg ega ed in o 1 km esolu ion by sum- ming he su ace a ea o o es om 30 m g id cells wi hin each 1 km g id cell (no e: he a ea o 30 m g id cells a y o e la i udes in he o iginal WGS84 p ojec ion and his was aken in o accoun in a ea calcula ions). The GFC o es loss da a a e boolean da a in na u e (loss o no loss). These o es loss boolean da a we e mul iplied by a su ace a ea o 30 m cell in ques ion and agg ega ed o 1 km esolu ion simila ly as o es ex en (see abo e). Possible o es loss was only accoun ed in hose 30 m g id cells ha had ee canopy co e o e he 20% h eshold in he o es ex en da ase . As a esul we go he o es ex en and o es loss in km 2 o each 1 km g id cell wi hou losing any in o ma ion in he agg ega ion p ocess. To illus a e he agg ega ed da ase s, he o es ex en and o es loss in pe cen ages o g id cell a ea a e p esen ed in Fig 1A and 1B. IFL and WDPA da ase s (Fig 1C and 1D;Table 2) we e used o calcula e he ex en and loss o p o ec ed o es , in ac o es and p o ec ed in ac o es . We i s con e ed he ec o da a o bo h da ase s o as e da a wi h 1 km esolu ion, being he o iginal esolu ion o IFL da a while esolu ion o WDPA da a a ies be ween he en ies. A e his we used he agg ega ed GFC da a o calcula e he a ea o o es ex en and o es loss inside each ype o spa ial c i e ia wi hin an analysis uni (coun y, egion o global): i) o al o es ex en and loss, ii) o es ex en and loss wi hin p o ec ed a eas, iii) o es ex en and loss wi hin in ac o es landscapes, and i ) o es ex en and loss wi hin p o ec ed in ac o es landscapes. We epo bo h absolu e and ela i e alues o o es ex en and loss. By using hese esul s, we analysed he o es p o ec ion e ec o p o ec ed a eas. The e ec was es ima ed by calcula ing anomaly a ios o ela i e loss o p o ec ed and non-p o ec ed o es , in ac and non-in ac o es , and o p o ec ed in ac and non-p o ec ed in ac o es . The anomaly a ios we e calcula ed a na ional scale by di iding he ela i e o es loss in he p o ec ed a eas in ques ion by he ela i e o es loss in he co esponding non-p o ec ed a eas, e.g. [( ela i e loss o p o ec ed in ac o es / ela i e loss o non-p o ec ed in ac o es )–1]. P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 5/21 Thus he anomaly a ios p o ide in o ma ion on he e ec o p o ec ed a eas on o es loss as hey indica e whe he he ela i e o es loss is lowe (<0) o highe (>0) in he p o ec ed o es han in he non-p o ec ed o es . We u he iden i ied simila i ies in o es loss pa e ns be ween coun ies by clus e ing he coun y scale esul s wi h k-mean clus e ing me hod. In his me hod he clus e ing o he obse a ions (i.e. coun y- alues) is based on minimizing he sum o he squa ed Euclidean dis- ances be ween each obse a ion and he clus e -cen oid- alue. We assessed goodness o i in e ms o a ia ion wi hin clus e s. To de ine he op imal numbe o clus e s we used he SSE cu e. We pe o med wo di e en clus e analysis, each conside ing ou di e en pa ame e s. In he i s clus e analysis, we iden i ied clus e s based on absolu e and ela i e losses in o al o es and p o ec ed o es a eas while in he second clus e analysis, we conside ed absolu e and ela i e o es losses in in ac o es landscapes and p o ec ed in ac o es landscapes. I should be no ed ha while o o es ex en and in ac o es da ase s he e e ence yea is 2000, o p o ec ed a eas he co e age in yea 2010 was aken. The e e ence yea 2010 o p o- ec ed a eas was chosen because ha moniza ion o he e e ence yea was no easible. The WDPA da ase does no epo he es ablishmen yea o all p o ec ed a eas. This howe e did no comp omise he analyses as we ound ha he WDPA da a om he yea 2010 adequa ely e lec he p o ec ion s a us o he o es o e he s udy pe iod: only 8% o he wo lds p o ec ed a eas (whe e an es ablishmen yea is p o ided) we e es ablished a e he yea 2000. Me hods: he ela ionship be ween o es loss and socio-economic indica o s To s udy he ela ionship be ween obse ed o es loss and socio-economic a ibu es, we employed WLS (i.e. Weigh ed Leas Squa es) eg ession analysis, which allowed he o al ex en o o es ype in ques ion o be used as weigh s. The deg ee o which he socio-economic a i- bu es can accoun o he ela i e o es loss was analysed o each o he ou o es loss mea- su es conside ed in his s udy: o al o es loss, loss in p o ec ed o es , loss in in ac o es , and loss in p o ec ed in ac o es . Al oge he 11 commonly used socio-economic indica o s we e selec ed and hey a e lis ed in Table 3. The socio-economic indica o s we e, mo eo e , selec ed Table 3. The socio-economic indica o s used in he eg ession analysis. The indica o s we e used as independen a iables in he assessmen o he o es loss d i e s wi h Weigh ed Leas -Squa es (WLS) eg ession analysis. Va iable Da a ype Time pe iod Re e ence Popula ion densi y pe squa e km 2010 UN [34] Popula ion densi y g ow h % 2000–2010 UN [34] GDP g ow h % 2000–2010 Wo ld Bank [35] GDP pe capi a (PPP) In e na ional USD 2010 Wo ld Bank [35] Ru al popula ion o o al popula ion % 2010 Wo ld Bank [35] Popula ion g ow h % 2000–2010 Wo ld Bank [35] Co up ion pe cep ion index a 0.. 10 2010 T anspa ency In e na ional [36] Poli y index b -10.. 10 2010 Ma shall and Gu [37] Human De elopmen Index (HDI) c 0.. 1 2012 UNDP [38] Ag icul u e o o al land a ea % 2010 Wo ld Bank [35] Ag icul u al land a ea g ow h % 2000–2010 Wo ld Bank [35] a Low co up ion pe cep ion index co esponds wi h high co up ion and ice e sa b Low poli y index co esponds wi h low democ acy and ice e sa c High HDI co esponds wi h high human de elopmen and ice e sa doi:10.1371/jou nal.pone.0138918. 003 P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 6/21 o ep esen he widely acknowledged causes o global de o es a ion: popula ion size, land-use change, s a e go e nance and s a e economy. The selec ion was ca ied ou wi h a equi emen o a ailable homogeneous da a a he global scale. We b ie ly desc ibe he WLS eg ession analysis below while s ep-by-s ep desc ip ion is gi en in S3 Appendix. The dependen a iables we e log 10 – ans o med p io o analyses o ensu e he no mali y o hei dis ibu ions. Simila ly, he independen a iables we e ans o med wi h a link unc- ion in cases i imp o ed he linea i y o he ela ion wi h he analysed o es loss measu es. The link unc ion o each independen a iable was de ined sepa a ely o each ou dependen a iables. Despi e he ans o ma ions, he da a included some s a is ical ou lie s. Exclusion o hese ou lie s did no a ec he signi icance o he esul s and hus he epo ed esul s we e gained wi hou omi ing he s a is ical ou lie s. The mul iple eg ession models we e buil by s a ing wi h he ull model comp ising o all 11 independen a iables and ending wi h a model whe e no mul icollinea i y was de ec ed. Mul icollinea i y was de e mined by he Va ia- ion In la ion Fac o (VIF, a se limi o VIF <4) and he collinea independen a iable wi h he lowes explana o y powe was always excluded om he model. Resul s Ou analyses based on GFC da a [5] e ealed ha o es co e ed one hi d o he o al land a ea (app oxima ely 43 million km 2 ) in he yea 2000, and 19% o his o es (o e 8 million km 2 ) was unde some o m o p o ec ion (Table 4). Acco ding o ou analyses 25% (11 million km 2 ) o global o es s we e in ac , o which almos 35% was p o ec ed. A eas ha a e bo h in ac and p o ec ed co e ed hus a o al o almos 3.7 million km 2 (o 8.7% o he o al o es ex en ) (Table 4). The global o es loss based on GFC da a was 2.14 million km 2 be ween he yea s 2000 and 2012, being o e 5% o he o al o es ex en in yea 2000 (Table 4). This global o es loss a e co esponds o calcula ions by Hansen e al. [5]. Fu he , acco ding o ou analyses, o e 10% o he o al o es loss occu ed in p o ec ed a eas: 219,000 km 2 o p o ec ed o es (3% o he o al p o ec ed o es ) was los wi hin ha ime pe iod. We u he ound ha 269,000 km 2 (2.5%) o in ac and 55,000 km 2 (1.5%) o p o ec ed in ac o es we e los du ing he yea s 2000–2012 (Table 4). In he ollowing sec ions he esul s a e p esen ed in mo e de ail. The nume ic esul s a e also gi en o each coun y in S2 Table o he supplemen a y in o ma ion. I is wo h o highligh ha ou es ima es on o al o es ex en and o al o es loss on a global scale p esen ed in his sec ion a e gi en only o alida e ou analyses (i.e. compa ison o Han- sen e al. [5]) and o pu ou esul s on PAs and IFLs in o a con ex . Fo es ex en s and hei p o ec ion s a us In yea 2000 he absolu e o es ex en was la ges in Russia, China, Indonesia, Democ a ic Republic o he Congo, B azil, Uni ed S a es, and Canada, exceeding 1 million km 2 (Fig 2A). Toge he hese coun ies con ained o e 60% o he global o es ex en . When examining ela- i e o es ex en (Fig 2B), pa icula ly Scandina ia and Finland (No dic coun ies om he e on), and coun ies in cen al A ica and Sou heas Asia had a ema kable po ion o hei e i- o y unde o es . The coun ies wi h he la ges p o ec ed o es a eas we e mainly he same coun ies wi h he g ea es ex en o o es (Fig 2C). Howe e , when assessing he a io o p o- ec ed o es ex en and he o al o es ex en , he la ges p o ec ed o es a eas we e loca ed in Cen al Eu ope, Aus alia, and in some coun ies o Sou h Ame ica, Sub-Saha an A ica and he Middle Eas (Fig 2D), whe e o e 30% o he o es s we e p o ec ed ( hough hese o en ep- esen small a eas o o es ). P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 7/21 The coun ies wi h he la ges o al o es ex en had also he la ges a eas o in ac o es (o e 400,000 km 2 ), excep Indonesia, Uni ed S a es and China (Fig 2E). On he o he hand, a la ge pa o Eu opean, Middle Eas e n, No h A ican and Sou he n A ican coun ies did no ha e any in ac o es landscapes (Fig 1D). When compa ing he a ea o in ac o es o he a ea o o al o es , we ound ha in many coun ies o No h and Sou h Ame ica, Sou heas Asia, Middle A ica and Aus alia sha es o in ac o es o o al o es we e e y high (Fig 2F). In con as o hese coun ies, No dic coun ies had a la ge p opo ion o hei land unde o - es (Fig 2B) bu only less han 2% o ha o es was classi ied as in ac o es (Fig 2F). Only B azil had o e 1 million km 2 o p o ec ed in ac o es (Fig 2G). Russia, Canada, Uni ed S a es and no he n coun ies o Sou h Ame ica had o e 100,000 km 2 o hei in ac o es s unde p o ec ion. When examining he ela i e ex en o p o ec ed in ac o es (ex en o p o ec ed in ac o es e sus ex en o o al in ac o es ), we ound ha majo i y o in ac o es , o e 70%, we e p o ec ed in coun ies such as Thailand, New Zealand, Japan, Madagas- ca , and E hiopia as well as No dic coun ies (Fig 2H). On he con a y, in some coun ies less han 20% o he in ac o es was unde p o ec ion, including Russia, Canada and Democ a ic Republic o he Congo (Fig 2H). Loss o o al and p o ec ed o es The la ges absolu e o es loss alues we e encoun e ed in B azil, Canada, Uni ed S a es, Rus- sia and Indonesia. Each o hem expe ienced o es losses o o e 100,000 km 2 du ing he pe iod o 2000–2012 (Fig 3A). The la ges ela i e losses, o e 10% o hei o es ex en , we e Table 4. Fo es ex en in 2000 and i s loss o e he pe iod o 2000–2012. The o es ex en and loss a e calcula ed o geog aphical egions in absolu e [10 3 km 2 ] and ela i e alues [%]. Fo es ex en [10 3 km 2 ] Fo es loss [10 3 km 2 ] Region To al ex en P o ec ed ex en (o o al ex en ) In ac ex en (o o al ex en ) P o ec ed in ac ex en (o o al in ac ex en ) To al loss (o o al ex en ) P o ec ed loss (o o al p o ec ed ex en ) In ac loss (o o al in ac ex en ) P o ec ed in ac loss (o o al p o ec ed in ac ex en ) Aus alia and Oceania 1,101 251 (23%) 256 (23%) 71 (28%) 48 (4%) 12 (5%) 4 (2%) 2 (3%) Cen al Ame ica 1,038 221 (21%) 60 (6%) 47 (78%) 54 (5%) 11 (5%) 1 (1%) 0.4 (1%) Eas e n Asia 2,203 193 (9%) 32 (1%) 10 (33%) 70 (3%) 5 (3%) 1 (5%) 0.4 (4%) Eas e n Eu ope and Cen al Asia 8,759 1,067 (12%) 2,039 (23%) 300 (15%) 357 (4%) 43 (4%) 72 (4%) 10 (3%) La in Ame ica 9,665 3,322 (34%) 4,400 (46%) 2,415 (55%) 531 (5%) 41 (1%) 31 (1%) 9 (0.4%) Middle and Sou h A ica 6,768 1,054 (16%) 998 (15%) 212 (21%) 201 (3%) 22 (2%) 3 (0.3%) 0.4 (0.2%) Middle Eas 188 12 (6%) 4 (2%) 0.4 (9%) 3 (2%) 0.1 (0.4%) 0.004 (0.1%) 0.0003 (0.1%) No h A ica 219 15 (7%) - (-) - (-) 2 (1%) 0.2 (1%) - (-) - (-) No h Ame ica 7,298 837 (11%) 2,408 (33%) 418 (17%) 513 (7%) 47 (6%) 152 (6%) 32 (8%) Sou h Asia 592 84 (14%) 31 (5%) 8 (26%) 10 (2%) 1 (1%) 0.1 (0%) 0.03 (0.3%) Sou heas e n Asia 3,234 657 (20%) 478 (15%) 203 (42%) 271 (8%) 23 (4%) 3 (0.7%) 1 (0.3%) Wes e n Eu ope 1,481 353 (24%) 11 (0.8%) 9 (81%) 83 (6%) 12 (3%) 0.03 (0.2%) 0.01 (0.1%) Global 42,562 8,068 (19%) 10,717 (25%) 3,693 (34%) 2,144 (5%) 219 (3%) 269 (2.5%) 55 (1.5%) doi:10.1371/jou nal.pone.0138918. 004 P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 8/21 ound in some coun ies in no h-wes e n and sou he n A ica, and Sou heas Asia (Fig 3B). In hese coun ies he ela i e losses we e high in hei p o ec ed o es s oo (Fig 3D). Rela i ely high (>5%) p o ec ed o es losses occu ed in Aus alia, Uni ed S a ed and some Eu opean coun ies. Sou h Asia, Middle Eas and Cen al Asia showed, on he o he hand, small numbe s o bo h absolu e and ela i e p o ec ed o es losses (Fig 3C and 3D). Fig 2. Absolu e and ela i e o es ex en by coun y in yea 2000. Absolu e o es ex en s (le column) a e p esen ed in km 2 and ela i e o es ex en s ( igh column) a e in %. A and B) To al o es ex en ; C and D) P o ec ed o es ex en ; E and F) In ac o es ex en ; G and H) P o ec ed in ac o es ex en . doi:10.1371/jou nal.pone.0138918.g002 P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 9/21 assessed coun ies (76%), ela i e o es loss was smalle in p o ec ed a eas han ou side hem (anomaly a io <–0.125: Fig 5A). The p o ec ion, mo eo e , seemed o be pa icula ly e ec i e o educing o es loss in Sou h Ame ica, Sou heas Asia and Sub-Saha an A ica (Fig 5A). Ne e heless, he e ec o p o ec ion on o es loss was he opposi e in some coun ies in Mid- dle A ica, Eu ope, Middle Eas and Mongolia in Asia and hus ejec s ou i s hypo hesis o a uni ied posi i e p o ec ion e ec . On coun y scale, ou indings sugges posi i e news o B a- zilian Amazon whe e p o ec ion was ound o be e y e ec i e in p e en ing o es loss. This is in ag eemen wi h he ea lie esea ch indings on he B azilian Amazon [6,39]. In he case o in ac o es , he global esul s we e simila , i.e. p o ec ion was e ec i e in 43 o 60 o assessed coun ies (72%) (Fig 5B). La ges de ia ions o he global majo i y we e shown by Mongolia, Nepal and Aus alia. I is wo h o acknowledge ha he PAs a e no necessa ily always aimed o p o ec he o es s ands pe se, ins ead p o ec ion can ha e o he goals. P io o ou s udy, he e ec i eness o p o ec ion is globally analysed by Joppa and P a [24]. Ou s udy u he s eng hens hose indings, ye ou indings a e e ie ed da a-wise in mo e consis en manne by using empo ally compa able o es ex en da ase s (Table 2), which only became a ailable a e he s udy by Joppa and P a [24]. D i e s o o es loss Impo an ly, we also explo ed he d i e s o o es loss by looking a he s a is ical linkage be ween socio-economic indica o s and weigh ed o es loss (Table 5). Weighing he coun y speci ic o es losses by hei o al o es ex en in ques ion allowed ou analysis o indica e he d i e s ha a e exp essi e o he global o es ex en ins ead o ocusing on d i e s explaining he in e -coun y a ia ion. This globally uni o m assessmen showed o es loss ha ing a s ong connec ion wi h ag icul u al land ex en . Mo eo e ag icul u al land expansion is ecog- nised as one o he mos impo an p ocesses causing o es loss (see e.g. [8–13]) and ou analy- sis ag ees well wi h his, pa icula ly conce ning he p o ec ed in ac o es s (Table 5). Howe e , i is wo h o ecognise ha ag icul u e ela ed o es loss in ol es much mo e com- plex dynamics han ou eg ession analyses a e able o cap u e (see e.g. [40]). We u he ound s ong connec ion be ween losses in p o ec ed and/o in ac o es s and popula ion densi y and GDP o ully suppo ou second hypo hesis. While se e al global scale s udies ha e ound ha highe popula ion densi y inc eases o es loss (e.g. [41,42]), hese all epo o al o es loss. We did no ind signi ican ela ionship be ween o al o es loss and popula ion densi y, whe eas ou indings o losses in in ac and p o ec ed in ac o es s (Table 5) show opposi e di ec ion compa ed o hese exis ing s udies on o al o es loss. Ou indings also seem o disag ee wi h a me a-analysis by Po e -Bolland e al. [43]. Howe e , a di ec compa ison be ween ou esul s and hose o Po e -Bolland e al. [43] is discou aged by he di e en scopes and app oaches in he s udies. Unlike ou s udy, he me a-analysis assessed he e ec i eness o only 40 p o ec ed a eas in he opics. Addi ionally, popula ion densi ies a e measu ed by di e en manne s: in ou s udy popula ion densi ies we e coun y a e ages while he me a-analysis based i s da a on popula ion densi y in he immedia e neighbou hood o p o ec ed a eas. Ou desi e o single s a is ical analysis including mul iple explana o y a iables de e mined he selec ion o coun y a e ages on he popula ion me ic o ma ch wi h he a ailable le el o he o he a iables. All in all, ou indings oge he wi h hose by Po e -Bolland e al. [43] poin o a need o u he esea ch on he ole o popu- la ion dynamics in o es loss. In e ms o GDP, we did no ind a s a is ically signi ican ela ionship wi h o al o es loss (Table 5). This inding is in line wi h some p e ious s udies [44–46], whe eas some o he s ud- ies sugges ha lowe GDP could indica e highe o es loss (e.g. [47,48]). While he e ec o P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 16 / 21 GDP on o es loss in p o ec ed a eas has no been assessed p e iously on a global scale, i is e alua ed by Nagend a [49] on 56 p o ec ed a eas. He indings indica e ha he a es o land co e clea ing do no di e signi ican ly be ween coun ies wi h di e en le els o GDP pe capi a. We ound, in u n, high GDP pe capi a o indica e high weigh ed o es loss a e in p o- ec ed a eas (Table 5). This ela ion be ween GDP and weigh ed o es loss on p o ec ed a eas can possibly indica e di e en managemen s a egies in coun ies wi h a ying GDP, i.e. coun- ies wi h highe GDP may ha e mo e e sa ile managemen s a egies such as, hose ha aim o s eng hen biodi e si y o p o ide habi a s o speci ic species, compa ed o coun ies wi h lowe GDP. Ou assessmen , i.e. looking a he o es loss d i e s globally, na u ally used a he limi ed se o socio-economic indica o s on a coa se scale. This cons ic ion on indica o s was needed o he sake o homogeneous da a. Ano he s o y would be e ealed by mo e local and egional s udies ha could o e ui ul insigh s o his global s udy, as is o example shown by he global me a-analysis looking a he d i e s behind he e ec i eness o p o ec ion managemen [50]. Ou s udy p o ides a global empi ical assessmen , weigh ed by he o al o es ex en s in ques ion, ha can be used as a e e ence poin o mo e de ailed s udies a o he spa ial scales. Taking u he –subna ional esul s and compa ison among da ase s We acknowledge ha he coun y le el esul s p esen ed in his s udy do no e eal ine le el he e ogenei y. The e o e, as an example, we explo ed how he o es loss in PAs a ies wi hin a coun y. Taking Colombia as an example, in ou esul s we showed ha he mean a e o o es loss wi hin all p o ec ed a eas was 0.08% pe yea . In a ine scale assessmen using he same da ase (i.e. GFC) we ound ha he o es loss a ied among he indi idual Colombian PAs om ze o o 1.48%, he obse ed a es being low o e la ge PAs in he Amazon and gene ally highe o p o ec ed a eas in he Andes. The mean a e o annual o es loss pe PA was 0.12% wi h s anda d de ia ion o 0.19%. This highligh s he impo ance o s udy u he he impac o p o ec ion on o es loss in lowe spa ial scales. Fu he mo e, we ca ied ou he same analysis by using an al e na i e da ase , Te a-I [51], o explo e possible di e ences in a ailable da ase s. Te a-i da ase de ia es om GFC in i s use o MODIS (250 m esolu ion) ins ead o Landsa (30 m esolu ion) and 2004–2014 ins ead o 2000–2012. The ange in o es loss in he indi idual p o ec ed a eas (0% o 1.72%) was simi- la o he inding made by GFC bu he mean annual o es loss a e wi hin he PAs was one hi d (mean a e o 0.043% wi h s anda d de ia ion 0.14%) o he ound losses compa ed o GFC da ase indings. This compa ison e lec s di e ences in da a ype, me hodology and pe iod bu also indica es he unce ain y associa ed wi h hese da ase s and, hus calls o u - he compa isons be ween he widely used da ase s. Me hodological limi a ions and ways o wa d I is impo an o expand he discussion owa ds he limi a ions c ea ed by he used da ase s ha also ou s udy is na u ally subjec ed o. Fo example, o he o es co e and loss we used he GFC da ase [5], which can gi e de ia ing esul s compa ed o o he da ase s mainly due o he di e ences in me hods such as: de ini ion o o es , sepa a ion o an h opogenic om na u- ally occu ing o es loss, and handling o a mosphe ic dis o ions (e.g. cloud co e ) in map- ping he o es co e . Al hough, di ing deepe in o hese di e ences and he e ec s hey ha e on he indings is beyond he scope o his analysis, i would o e a ui ul opic o a compa a- i e s udy. In addi ion, Lee [52] aises a conce n on he IFL da ase [20] ha i s de ini ion o in ac ness o o es may no cap u e he socio-ecological a ia ion be ween di e en geog aphi- cal egions and may hus in oduce inaccu acies in coun y speci ic analyses. In e -compa ison P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 17 / 21 o global scale da ase s as well as mo e de ailed analyses a he coun y/ egional le el would be needed o un a el he s a e o global classi ica ion o IFLs. I should be no ed ha ou s udy ocused on o es loss, o mo e p ecisely on s and- eplac- ing dis u bance, igno ing he possible o es gain. This ocus ga e us mo e space o explo e he loss o o iginal habi a s wi hin p o ec ed a eas and in ac o es landscapes and he d i e s behind hese losses. A compa ison o ou indings o indings ha would be based on he ne o es loss, i.e. a loss assessmen including he plan ing o o es s and na u al o es egene a- ion (see [2,5] and e e ences he ein o u he in o ma ion), would allow acking o he o - es canopy co e pe se. Ou es ima ions on he e ec i eness o p o ec ion o educe o es loss wi hin PAs com- pa ed o he o es loss ou side hose a eas p o ide use ul in o ma ion he p o ec ion e ec a he coun y le el. A he coun y le el, we we e ne e heless no able o assess he leakage e ec o indi idual p o ec ed a eas, as done by e.g. Oli ei a e al [53]. As highligh ed by Ewe s and Rod igues [54] a compa ison o o es loss in he icini y o and inside p o ec ed a eas is essen- ial in unde s anding he e ec s o o es p o ec ion on a local scale. I is also wo hwhile o ec- ognise ha o es ed a eas migh ha e been p o ec ed o o he easons han p o ec he o es s ands pe se. The global da ase s used in his s udy could be u he employed o his kind o assessmen , compa ing he leakage e ec wi hin and among egions and u he mo e ex ending i all he way o global le el. Mos impo an ly, global da a on o es managemen goals pe each p o ec ion a ea would be highly needed o suppo mo e de ailed assessmen on he impac o p o ec ion on o es loss. Conclusions Ou global scale analysis indica es ha subs an ial a eas o p o ec ed (3%) and in ac o es s (2.5%) we e los o e he pas decade, and la ge pa o ha loss occu ed in coun ies wi h well-es ablished p o ec ion (e.g. in Aus alia and Oceania, and No h Ame ica). In e ms o compa ison be ween o es loss inside and ou side o PAs and IFLs we could no con i m ou i s hypo hesis, which s a ed ha o es loss would always be less mani es ed wi hin PAs han ou side o hem. Al hough he ela i e o es loss was smalle inside he PAs and p o ec ed IFLs in global a e ages and in majo i y o coun ies, he e we e nume ous coun ies ha showed an opposi e e ec . Also ou second hypo hesis was no en i ely con i med: while popula ion size and o he indica o s we e able o explain di e ences in weigh ed global o es loss wi hin PAs and IFLs, a la ge pa o he a ia ion emained unexplained. Mo eo e , he s a is ical associa- ions did no always co espond wi h ou causal unde s anding o o es loss p ocesses. Ou indings hus highligh ha p o ec ion o o es ed a eas does no always gua an ee a lowe a e o o es loss. The e is, indeed, a high geog aphical a ia ion in he e ec i eness o p o ec ion agains o es loss, a a ia ion ha is p obably a leas o some ex en explained by coun ies’ di e en means and in ensi ies o comba agains o es loss. Suppo ing In o ma ion S1 Appendix. C oss-co ela ion ables. C oss-co ela ion ables o socio-economic indices in cases o di e en o es loss ca ego ies. (PDF) S2 Appendix. Compa ison o ou indings o p e ious s udies: o es ex en and loss. (PDF) S3 Appendix. Weigh ed Leas Squa es (WLS) eg ession analysis s eps. (PDF) P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 18 / 21 S1 Table. Socio-economic da a a coun y le el. Socio-economic da a used in he WLS eg es- sion analysis. (XLSX) S2 Table. Coun y le el esul s. Fo es ex en and o es loss esul s p esen ed a coun y le el. (XLSX) Acknowledgmen s We a e g a e ul o CSC—IT Cen e o Science o p o iding he compu a ional acili ies o he analyses. We also hank he e iewe s o hei cons uc i e commen s and P o . Olli Va is, D . Joseph Guillaume, and o he membe s o ou esea ch eam a Aal o Uni e si y o hei encou agemen s and ui ul discussions. Au ho Con ibu ions Concei ed and designed he expe imen s: MH MK M. Mulligan TAR. Pe o med he expe i- men s: MH MK M. Mulligan MJ TAR. Analyzed he da a: MH MK M. Makkonen M. Mulligan MJ TAR. Con ibu ed eagen s/ma e ials/analysis ools: MH MK M. Makkonen M. Mulligan MJ TAR. W o e he pape : MH MK M. Makkonen M. Mulligan PHV TAR. Re e ences 1. Bonan GB (2008) Fo es s and Clima e Change: Fo cings, Feedbacks, and he Clima e Bene i s o Fo - es s. Science 320: 1444–1449 doi: 10.1126/science.1155121 PMID: 18556546 2. FAO (2010) Global o es esou ce assessmen 2010. Rome: Food and Ag cul u e O ganisa ion o he Uni ed Na ions. 3. Gibson L, Lee TM, Koh LP (2011) P ima y o es s a e i eplaceable o sus aining opical biodi e si y. Na u e 478: 378–381. doi: 10.1038/na u e10425 PMID: 21918513 4. Pan Y, Bi dsey RA, Fang J, Hough on R, Kauppi PE, Ku z WA, e al. (2011) A La ge and Pe sis en Ca - bon Sink in he Wo ld’s Fo es s. Science 333: 988–993 doi: 10.1126/science.1201609 PMID: 21764754 5. Hansen MC, Po apo PV, Moo e R, Hanche M, Tu ubano a SA, Tyuka ina A, e al. (2013) High- eso- lu ion global maps o he 21s -cen u y o es co e change. Science 342: 850–853. doi: 10.1126/ science.1244693 PMID: 24233722 6. Neps ad D, Soa es-Filho BS, Me y F, Lima A, Mou inho P, Ca e J, e al. (2009) The End o De o es a- ion in he B azilian Amazon. Science 326: 1350–1351. doi: 10.1126/science.1182108 PMID: 19965742 7. Goda J, Ga dne TA, Tizado EJ, Pacheco P (2014) Ac o -speci ic con ibu ions o he de o es a ion slowdown in he B azilian Amazon. P oceedings o he Na ional Academy o Sciences 111: 15591– 15596. 8. DeF ies RS, Rudel T, U ia e M, Hansen M (2010) De o es a ion d i en by u ban popula ion g ow h and ag icul u al ade in he wen y- i s cen u y. Na u e Geoscience 3: 178–181. 9. Fe e i-Gallon K, Busch J (2014) Wha d i es de o es a ion and wha s ops i ? A me a-analysis o spa- ially explici econome ic s udies. Washing on, DC: Cen e o Global De elopmen . 10. Geis HJ, Lambin EF (2002) P oxima e causes and unde lying d i ing o ces o opical de o es a ion. BioScience 52: 143–150. 11. Hosonuma N, He old M, De Sy V, De F ies RS, B ockhaus M, Ve cho L, e al. (2012) An assessmen o de o es a ion and o es deg ada ion d i e s in de eloping coun ies. En i onmen al Resea ch Le - e s 7. 12. Rudel TK (2005) T opical Fo es s. Regional Pa hs o Des uc ion and Regene a ion in he La e Twen i- e h Cen u y. New Yo k: Columbia Uni e si y P ess. 13. Rudel TK, DeF ies R, Asne GP, Lau ance WF (2009) Changing d i e s o de o es a ion and new oppo uni ies o conse a ion. Conse a ion Biology 23: 1396–1405. doi: 10.1111/j.1523-1739.2009. 01332.x PMID: 20078640 P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 19 / 21 14. Rudel TK (2007) Changing agen s o de o es a ion: F om s a e-ini ia ed o en e p ise d i en p ocesses, 1970–2000. Land Use Policy 24: 35–41. 15. Ju e-Bignoli D, Bu gess ND, Bingham H, Belle EMS, de Lima MG, Deguigne M, e al. (2014) P o ec ed Plane Repo 2014. Camb idge, UK: UNEP-WCMC. 16. Naugh on-T e es L, Holland MB, B andon K (2005) The ole o p o ec ed a eas in conse ing biodi e - si y and sus aining local li elihoods. Annu Re En i on Resou 30: 219–252. 17. Radelo VC, Beaud y F, B ooks TM, Bu sic V, Dubinin M, Kuemme le T, e al. (2013) Ho momen s o biodi e si y conse a ion. Conse a ion Le e s 6: 58–65. 18. Schmi CB, Bu gess ND, Coad L, Beloku o A, Besançon C, Bois obe L, e al. (2009) Global analysis o he p o ec ion s a us o he wo ld’s o es s. Biological Conse a ion 142: 2122–2130. 19. Dudley N (2008) Guidelines o Applying P o ec ed A ea Managemen Ca ego ies. Gland, Swi ze land: IUCN. 20. Po apo P, Ya oshenko A, Tu ubano a S, Dubinin M, Laes adius L, Thies C, e al. (2008) Mapping he wo ld's in ac o es landscapes by emo e sensing. Ecology and Socie y 13. 21. Joppa LN, P a A (2010) Reassessing he o es impac s o p o ec ion: The challenge o non andom loca ion and a co ec i e me hod. Annals o he New Yo k Academy o Sciences 1185: 135–149. doi: 10.1111/j.1749-6632.2009.05162.x PMID: 20146766 22. Joppa LN, P a A (2009) High and a : Biases in he loca ion o p o ec ed a eas. PloS ONE 4: e8253– e8273. 23. Ve bu g PH, O e ma s KP, Huigen MGA, de G oo WT, Veldkamp A (2006) Analysis o he e ec s o land use change on p o ec ed a eas in he Philippines. Applied Geog aphy 26: 153–173. 24. Joppa LN, P a A (2011) Global p o ec ed a ea impac s. P oceedings o he Royal Socie y B 278: 1633–1638. doi: 10.1098/ spb.2010.1713 PMID: 21084351 25. DeF ies R, Hansen A, New on AC, Hansen MC (2005) Inc easing isola ion o p o ec ed a eas in opical o es s o e he pas wen y yea s. Ecological Applica ions 15: 19–26. 26. Maio ano L, Falcucci A, Boi ani L (2008) Size-dependen esis ance o p o ec ed a eas o land-use change. P oc R Soc B June 7, 2008 275 1640 1297–1304 275: 1297–1304. 27. B ook BW, Sodhi NS, B adshaw CJA (2008) Syne gies among ex inc ion d i e s unde global change. T ends in Ecology & E olu ion 23: 453–460. 28. Tu ne IM (1996) Species loss in agmen s o opical ain o es : A e iew o he e idence. Jou nal o Applied Ecology 33: 200–209. 29. Ma gono BA, Po apo PV, Tu ubano a S, S ollle F, Hansen MC (2014) P ima y o es co e loss in Indonesia o e 2000–2012. 30. IUCN, UNEP-WCMC (2014) The Wo ld Da abase on P o ec ed A eas (WDPA). In: UNEP-WCMC, edi- o . Camb idge, UK. A ailable: www.p o ec edplane .ne . Accessed 7 July 2014. 31. Bonda WJ, Keeleyb JE (2005) Fi e as a global ‘he bi o e’: he ecology and e olu ion o lammable eco- sys ems. T ends in Ecology and E olu ion 20: 387–394. PMID: 16701401 32. Logan JA, Régniè e J, Powell JA (2003) Assessing he impac s o global wa ming on o es pes dynamics. F on ie s in Ecology and he En i onmen 1: 130–137. 33. Po apo P, Hansen MC, S ehman SV, Lo eland TR, Pi man K (2008) Combining MODIS and Landsa image y o es ima e and map bo eal o es co e loss. Remo e Sensing o En i onmen 112: 3708– 3719. 34. UN (2014) Wo ld Popula ion P ospec s: The 2012 Re ision. UN Popula ion Di ision. A ailable: h p:// esa.un.o g/unpd/wpp/Excel-Da a/popula ion.h m. Accessed 7 Augus 2014. 35. Wo ld Bank (2014) Wo ld Bank De elopmen Indica o s. Wo ld Bank. A ailable: h p://da a.wo ldbank. o g/indica o . Accessed 7 Augus 2014. 36. T anspa ency In e na ional (2010) Co up ion Pe cep ion Index. T anspa ency In e na ional. A ailable: h p://www. anspa ency.o g/cpi2010/. Accessed 14 June 2014. 37. Ma shall MG, Gu T (2014) Poli y IV P ojec , Poli ical Regime Cha ac e is ics and T ansi ions, 1800– 2013. Cen e o Sys ema ic Peace. A ailable: h p://www.sys emicpeace.o g/insc da a.h ml. Accessed 14 June 2014. 38. UNDP (2014) Human De elopmen Index 2012. Uni ed Na ions De elopmen P og amme. A ailable: h ps://da a.undp.o g/da ase /Table-1-Human-De elopmen -Index-and-i s-componen s/wxub-qc5k. Accessed 7 Augus 2014. 39. Nol e C, Ag awal A, Sil ius KM, Soa es-Filho BS (2013) Go e nance egime and loca ion in luence a oided de o es a ion success o p o ec ed a eas in he B azilian Amazon. PNAS 110: 4956–4961. doi: 10.1073/pnas.1214786110 PMID: 23479648 P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 20 / 21 40. Pagnu i C, Bauch CT, Anand M (2013) Ou look on a Wo ldwide Fo es T ansi ion. PLoS ONE 8: e75890. doi: 10.1371/jou nal.pone.0075890 PMID: 24130750 41. Ma he AS, Needle CL (2000) The ela ionships o popula ion and o es ends. Geog aphical Jou nal 166: 2–13. 42. Ma he AS, Needle CL, Fai bai n J (1998) The human d i e s o global land co e change: he case o o es s. Hyd ological P ocesses 12: 1983–1994. 43. Po e -Bolland L, Ellis EA, Gua igua a MR, Ruiz-Mallen I, Neg e e-Yankele ich S, Reyes-Ga cia (2012) Communi y managed o es s and o es p o ec ed a eas: An assessmen o hei conse a ion e ec i eness ac oss opics. Fo es Ecology and Managemen 268: 6–17. 44. Dame e O, Delaco e P (2012) On he economic ac o s o de o es a ion: Wha can we lea n om quan- ile analysis? Economic Modelling 29: 2427–2434. 45. Koop G, Tole L (1999) Is he e an en i onmen al Kuzne s cu e o de o es a ion? Jou nal o De elop- men Economics 58: 231–244. 46. Tole L (1998) Sou ces o De o es a ion in T opical De eloping Coun ies. En i onmen al Managemen 22: 19–33. PMID: 9419282 47. Ewe s RM (2006) In e ac ion e ec s be ween economic de elopmen and o es co e de e mine de o - es a ion a es. Global En i onmen al Change 16: 161–169. 48. Smi h RJ, Mui RDJ, Walpole MJ, Balm o d A, Leade -Williams N (2003) Go e nance and he loss o biodi e si y. Na u e 426: 67–70. PMID: 14603318 49. Nagend a H (2008) Do pa ks wo k? Impac o p o ec ed a eas on land co e clea ing. Ambio 37: 330– 337. PMID: 18828277 50. Le e ing on F, Lemos Cos a KL, Pa ese P, Lisle A, Hockings M (2010) A Global Analysis o P o ec ed A ea Managemen E ec i eness. En i onmen al Managemen 46: 685–698. doi: 10.1007/s00267- 010-9564-5 PMID: 20859627 51. Reymondin L, Ja is A, Pe ez-U ibe A, Tou al J, A go e K, Rebe ez J, e al. (2012) A me hodology o nea eal- ime moni o ing o habi a change a con inen al scales using MODIS-NDVI and TRMM. A ail- able: www. e a-i.o g. Accessed 16 Decembe 2014 52. Lee PG (2009) Cau ion agains using in ac o es -landscapes da a a egional scales. Ecology and Socie y 14: 1. A ailable: h p://www.ecologyandsocie y.o g/ ol14/iss11/ esp11/. 53. Oli ei a PJC, Asne GP, Knapp DE, Almeyda A, Gal án-Gildemeis e R, Keene S, e al. (2007) Land- Use Alloca ion P o ec s he Pe u ian Amazon. Science 317: 1233–1236. PMID: 17690260 54. Ewe s RM, Rod igues ASL (2008) Es ima es o ese e e ec i eness a e con ounded by leakage. T ends in Ecology and E olu ion 23: 113–116. doi: 10.1016/j. ee.2007.11.008 PMID: 18280005 P o ec ed and In ac Fo es Loss PLOS ONE | DOI:10.1371/jou nal.pone.0138918 Oc obe 14, 2015 21 / 21