Effect of the forest-mine boundary form on woody colonization and forest expansion in degraded ecosystems

A icle E ec o he Fo es -Mine Bounda y Fo m on Woody Coloniza ion and Fo es Expansion in Deg aded Ecosys ems Ca olina Ma ínez-Ruiz 1,2,* , Ana I. Milde 1,3, Daphne López-Ma cos 1,4 , Pila Zaldí a 4and Belén Fe nández-San os 3   Ci a ion: Ma ínez-Ruiz, C.; Milde , A.I.; López-Ma cos, D.; Zaldí a , P.; Fe nández-San os, B. E ec o he Fo es -Mine Bounda y Fo m on Woody Coloniza ion and Fo es Expansion in Deg aded Ecosys ems. Fo es s 2021,12, 773. h ps:// doi.o g/10.3390/ 12060773 Academic Edi o : Richa d Dodd Recei ed: 1 May 2021 Accep ed: 8 June 2021 Published: 11 June 2021 Publishe ’s No e: MDPI s ays neu al wi h ega d o ju isdic ional claims in published maps and ins i u ional a il- ia ions. Copy igh : © 2021 by he au ho s. Licensee MDPI, Basel, Swi ze land. This a icle is an open access a icle dis ibu ed unde he e ms and condi ions o he C ea i e Commons A ibu ion (CC BY) license (h ps:// c ea i ecommons.o g/licenses/by/ 4.0/). 1Á ea de Ecología, Dp o. de Ciencias Ag o o es ales, E.T.S. de Ingenie ías Ag a ias de Palencia, Uni e sidad de Valladolid, Campus La Yu e a, A da. de Mad id 50, 34071 Palencia, Spain; [email p o ec ed] (A.I.M.); [email p o ec ed] (D.L.-M.) 2Sus ainable Fo es Managemen Resea ch Ins i u e UVa-INIA, E.T.S.II.AA., Campus La Yu e a, A da. de Mad id 50, 34071 Palencia, Spain 3Á ea de Ecología, Facul ad de Biología, Uni e sidad de Salamanca, Campus Miguel de Unamuno s/n, 37071 Salamanca, Spain; [email p o ec ed] 4Á ea de Bo ánica, Dp o. de Ciencias Ag o o es ales, E.T.S. de Ingenie ías Ag a ias de Palencia, Uni e sidad de Valladolid, Campus La Yu e a, A da. de Mad id 50, 34071 Palencia, Spain; pila [email p o ec ed] *Co espondence: ca [email p o ec ed] Abs ac : We e alua ed he ecological signi icance o he bounda y o m be ween wo pa ches wi h con as ing ege a ion (mine g assland and adjacen o es ) on woody coloniza ion and o es expansion in open-cas coal mines in No he n Spain. Woody coloniza ion and b owsing aces we e measu ed on h ee mine si es, along 24 ansec s ha we e laid ou pe pendicula o he o es -mine bounda y and classi ied acco ding o hei shape (conca e, con ex, s aigh ). Mine si es we e colonized om he close o es by woody species, whose coloniza ion in ensi y depends on he bounda y o m. The o e all coloniza ion in ensi y dec eased wi h inc easing dis ance o he o es and di e ed depending on he bounda y o m. The mo e in ense coloniza ion was ound in conca e bounda ies and he s onges dec ease in con ex bounda ies close o he o es , whe eas s aigh bounda ies showed an in e media e coloniza ion pa e n. Conca e bounda ies eached highe woody co e in he basal s a a o he mines han con ex (up o 2 m) o s aigh bounda ies (up o 1 m) om 11 m o he o es edge, mainly by he p esence o dense pa ches o Cy isus scopa ius (L.) Link, wi h a sca e ed o e s o y o Genis a lo ida L. These sh ubs migh educe he b owsing in ensi y and ac as nu se plan s acili a ing he es ablishmen o Que cus pe aea (Ma .) Liebl. in mine a eas a g ea e dis ances om he o es edge. The o es -mine bounda y o m does no a ec he o es e ical s uc u e ha is homogenous and does no help explain he woody coloniza ion pa e n in he mines. We conclude ha edge cha ac e is ics ha e a s ong po en ial o be used in he es o a ion o na i e o es s based on na u al p ocesses. The implica ions o ou esul s o sessile oak ( Que cus pe aea (Ma .) Liebl.) o es expansion along edges in agmen ed Medi e anean o es landscapes we e discussed. Keywo ds: bounda y o m; b owsing pa e n; agmen a ion; Que cus pe aea o es s; sub-Medi e - anean clima e; su ace coal-mining; e ical s uc u e; woody coloniza ion 1. In oduc ion The bounda y o m be ween di e en ecosys ems o landscape elemen s can be widely a ied [ 1 ], including conca e, con ex, and s aigh shapes. Con exi ies a e cha ac e ized by a pa icula species and mic oen i onmen composi ion because o he exposu e on h ee sides o he su ounding ma ix. On he con a y, conca i ies ha a e less exposed and ha e an amelio a ed mic oclima e became a eas wi h highe exchange a es due o coloniza ion p ocesses [ 2 ]. Such bounda y o ms doub less a ec impo an ecological Fo es s 2021,12, 773. h ps://doi.o g/10.3390/ 12060773 h ps://www.mdpi.com/jou nal/ o es s Fo es s 2021,12, 773 2 o 18 p ocesses and dynamics in he icini y [ 1 , 2 ], since he bounda y o m de e mines hei oles as habi a , il e , o condui [ 3 ]. As a esul , he bounda y o m can de e mine pa ch expansion o con ac ion [ 4 ], because adjacen pa ch shapes in e digi a e each o he , and in e ac signi ican ly wi h he o ien a ion o di ec ional o ces in he landscape, such as wind and species dispe sion om sou ce a eas [ 3 ]. Many o hese e ec s emana e om he ela ion exis ing be ween he shape o pa ches, in e ac ions wi h he su ounding ma ix, and anspo wi hin a pa ch [3–5]. The e o e, edges c ea ed by habi a agmen a ion media e he a es o low o ene gy, ma e [ 6 ], and o ganisms [ 7 ]. How edges media e hese luxes is de e mined by he edge pe meabili y, which measu es he endency o an edge o inhibi o enhance he mo emen o o ganisms, ma e ial, o ene gy ac oss i [ 6 , 8 ]. Fo example, he deg ee o pe meabili y o a o es -g assland edge (wi h a high deg ee o ege a ion con as ) could de e mine he mo emen o o ganisms, including seed mobili y, and hus in luence he in asion o woody species and o es expansion om he o es owa ds he inside o he g assland [9,10]. Al hough implica ions o edge pe meabili y in e ms o o es conse a ion, egene - a ion, and wildli e managemen a e clea [ 6 , 9 , 11 ], ew s udies assess he in luence o he bounda y o m on o es expansion, and pa icula ly, in deg aded a eas such as hose a - ec ed by opencas coal mining (bu see [ 3 ]). In he Eas e n USA, he o es -mine bounda y o m exe ed a powe ul con ol on woody coloniza ion, esul ing in opposi e beha io s o conca e s. con ex bounda ies, wi h highe coloniza ion in ensi y nea conca e bounda ies and coloniza ion eaching a highe dis ance [ 3 ]. Such a woody coloniza ion pa e n was ela ed o he ac i i y o animals h ough he quan i ica ion o he bi o y in ensi y, al hough o he hypo heses could be conside ed, such as di e en amoun s o o es in he p oximi y ha in oduce di e ences in mic o-en i onmen al condi ions on he mine [ 12 , 13 ]. Besides, he o es e ical s uc u e on edges can in luence he unc ion o edges as ba ie s o seed lux [ 13 ], and i can become a ec ed by he bounda y o m o e ime [ 14 ]. Thus, he bounda y o m as a spa ial cha ac e is ic signi ican ly a ec s se e al o he a iables; conca i ies o co es ha e an amelio a ed mic oclima e, mo e adjacen co e o he bi o es, and mo e po en ial p opagule sou ce in he o es pe uni a ea o mine su ace [ 15 , 16 ], all o which doub less in e ac wi h he basic immig a ion p ocess [3]. In No he n Palencia (Spain), as in o he a eas in he wo ld, opencas coal mining plays an impo an ole in he dynamics o landscape agmen a ion [ 7 ]. In he eclaimed mining su aces, he baceous species a e usually employed o a o soil s abiliza ion since o en he e is a lack o soil seed bank in he opsoil [ 17 ]. The ini ial esul is a pa ch o g assland in a o es ma ix. The g assland is hen colonized by woody species om he adjacen o es o dispe sed by wind o animals om su ounding a eas [ 7 ]. This p ocess is s ongly a ec ed by in e ac ions be ween he su ounding o es ma ix and he g assland pa ch and is p obably in luenced by he geome y o he pa ch, which can be desc ibed as a combina ion o se e al bounda y o ms [ 3 ]. The o es ma ix, as a seed sou ce, pa ly de e mines he species composi ion o woody colonize s [ 7 ]. B owsing animals om he o es ma ix selec i ely eed in he pa ch, e a ding woody coloniza ion, and addi ionally a ec ing species composi ion [ 18 ]. G azing can di ec ly limi he ec ui men o woody species [ 11 , 19 – 21 ], and indi ec ly by speci ically a ec ing he ew seeds dispe sed o he high-quali y mic osi es [22,23]. In his con ex , ou gene al aim was o e alua e he ecological signi icance o he bounda y o m be ween wo pa ches wi h a high deg ee o ege a ion con as ( o es and g assland) on woody coloniza ion o eclaimed coal mines in No he n Spain. Speci ically, we examined he ollowing ques ions: (1) A e he in ensi y and colonizing dis ance o woody species highe in conca e han in con ex bounda ies? (2) Does he o es e ical s uc u e a ec such a woody coloniza ion pa e n in he mines? (3) Does he bi o e b ows- ing play a key ole in de e mining he coloniza ion pa e ns? The esul ing in o ma ion may o e s ong possibili ies o landscape conse a ion [ 24 ] and es o a ion o dis u bed si es, such as lands mined o coal all a ound he wo ld, based on spon aneous p ocesses such as coloniza ion and succession [25,26]. Fo es s 2021,12, 773 3 o 18 2. Ma e ials and Me hods 2.1. S udy A ea This s udy examines h ee close si es, all su ace-mined o coal, in he p o ince o Palencia (No he n Spain), nea he illage o Gua do (1110 m a.s.l.; see Figu e A1). They lie a 42 ◦ 47 0 N, 4 ◦ 50 0 W in he “Mon aña Palen ina” a ea on p edominan ly limes one o Paleozoic age, wi h he p esence o clay and o a lesse deg ee o sand [ 7 ]. The clima e is sub-humid Medi e anean [ 27 ] wi h a mean annual ain all o 973 mm, ainy seasons in au umn and sp ing, and acu e summe d ough s h ough July and Augus wi h only 8% o he annual ain all [ 28 ]. The mean annual empe a u e is 9.2 ◦ C, he mean minimum o he coldes mon h (Janua y) is − 2.7 ◦ C, and he mean maximum o he wa mes mon h (Augus ) is 25.9 ◦ C. Mos o he a ea su ounding he si es is o es ed wi h ela i ely di e se ege a ion associa ed wi h he sessile oak (Que cus pe aea (Ma .) Liebl.) o es s. The na u al soils a e mainly Incep isols wi h a udic soil mois u e egime and mesic soil empe a u e egime [29,30]. S udy si es we e eclaimed a ound 16 yea s be o e ou au umn sampling. They we e eg aded o app oxima ely he o iginal con ou , and he su ace was co e ed wi h 30 cm o ine- ex u ed ma e ials amended wi h ca le manu e (30 /ha) and e ilize (8 N:15 P:15 K; 150 kg/ha). The ine- ex u ed ma e ials we e a mix u e o opsoil and sedimen s om deepe pa s o he nea es opencas pi s; his mix u e had a clay loam ex u e, wi h a pH o 6.5, an elec ical conduc i i y o 114.3 µ S/cm, easily oxidizable ca bon o 19.8 g/kg, a ailable phospho ous o 9.7 mg/kg and an e ec i e dep h o 10–15 cm [ 30 ]. These soils can be classi ied as Li hic Udo hen s (sensu [ 29 ]) and hey ha e a e y low wa e -holding capaci y compa ed o he na u al soil in he o es (1.0–3.4 s. 19.8 7 ± 1.52) [ 30 ]. A e opsoiling, mines we e e ege a ed wi h a comme cial seed mix u e (210 kg/ha) o g asses and legume he bs (80:20 by weigh ). When sampled, he h ee si es we e colonized by wel e na i e sh ub species om he su ounding o es s, mainly Cy isus scopa ius (L.) Link and Genis a lo ida L., wo legume sh ub species no especially abundan in he o es edge. The a ea was being g azed by wild animals om he su ounding landscape (dee , oe dee , and wild boa ), and li es ock (ca le and ho ses) [7]. The h ee eclaimed mine si es had simila geome y and slope: Si e I: 325 × 185 m, eigh ansec s sampled om no heas o eas edges, slope a ying om 20 o 28 ◦ ; Si e II: 100 × 100 m, six ansec s sampled om no heas o eas edges, slopes 20–25 ◦ ; Si e III: 100 ×370 m, en ansec s sampled om no hwes o sou hwes edges, slope ca. 22◦. 2.2. Vege a ion Sampling The sampling design ollowed ha p oposed by [ 3 ]. T ansec s 74 m long, ex ending 63 m on o he mine si e and 11 m in o he o es , we e laid ou pe pendicula o he o es - mine bounda y. Nine een 2 × 2 m plo s, cen e ed a 4 m in e als along each ansec , we e sampled, i.e., h ee plo s in o he o es and 16 in o he mine. An ini ial ansec poin was es ablished a he sou heas e nmos co ne o each mine si e, and subsequen ansec s we e es ablished a ound he o es ed bounda y in a clockwise ashion a 30 m in e als a la ge si es, and a 15 m in e als a he smalle si e. The ini ial ansec poin was omi ed om sampling a each si e o minimize a possible bias in he s a ing poin selec ion. None o he selec ed ansec s c ossed a eas dis u bed by e osion ha could a ec land ecoloniza ion (see Figu e A2 o mo e sampling de ails). A each ansec , he o es -mine bounda y o m was eco ded as conca e, s aigh , o con ex (i.e., a conca e bounda y is a p ojec ion o g ass- ege a ed mine in o he o es ). To de e mine he bounda y o m, a angen was d awn a he midpoin o a smoo hed a c connec ing in lec ion poin s on each co e o lobe. Bounda ies we e de ined as s aigh i he angle be ween he angen and a line connec ing he midpoin wi h he in lec ion poin was <11 ◦ 19 0 [ 3 ]. The 24 sampled ansec s included nine conca e, eigh con ex, and se en s aigh bounda ies. Wi hin each 2 × 2 m plo , he o al co e o woody ege a ion a se en s a a (0–25 cm, 25–50 cm, 0.5–1 m, 1–2 m, 2–4 m, 4–8 m, and >8 m) was eco ded o cha ac e ize he e ical Fo es s 2021,12, 773 4 o 18 s uc u e. The numbe o indi iduals o each woody species was also no ed, eco ding he p esence o b owsing e idence; he size class was also eco ded o indi iduals in o he o es edge (11 m). Sepa a e s ems a g ound le el we e egis e ed as indi iduals. Fi e size classes we e used: (1) ≤ 0.3 m heigh ; (2) >0.3 and ≤ 0.6 m heigh ; (3) >0.6 m and ≤ 2 m heigh ; (4) >2 m heigh , wi h a diame e a b eas heigh (dbh) <10 cm; (5) >2 m heigh , wi h dbh ≥ 10 cm. The p esence o a cu s em o wig was eco ded as e idence o b owsing [ 3 ]. B owsing in ensi y (pe cen o b owsed s ems o he o al o colonizing s ems) was hen calcula ed. 2.3. Da a Analysis Linea Mixed Models (LMM) wi h he Res ic ed Maximum Likelihood me hod (REML; [ 31 ]) we e used o assess he e ec o bounda y o m on he e ical s uc u e o woody ege a ion in he o es edge ( he 11 m s ip o he o es close o mine), and in he mines conside ing he dis ance o he o es edge. In bo h cases, a null model conside ing he andom e ec o plo s wi hin ansec s was es ed wi h he al e na i e model ha , in he i s case, included he ixed e ec o he ype o bounda y, and, in he second case, he ixed e ec s o he ype o bounda y plus he dis ance o he o es edge. The Akaike in o ma ion c i e ion (AIC; [ 32 ]) was used o e i y whe he he al e na i e model was mo e pa simonious, i.e., smalle alues o AIC. The ANOVA was applied o es he signi i- can di e ences be ween he null and he al e na i e models. A e wa d, wo king o e he model ma ix, con as s we e ca ied ou o es di e ences be ween ixed ac o le els [ 33 ]. Consequen ly, he Bon e oni co ec ion was used o adjus he signi icance le el o each - es [34,35]. Linea Mixed Models we e also used o assess he in luence o he bounda y o m and dis ance o he o es edge on global woody coloniza ion in ensi y (s ems/m 2 ) and o he six mo e abundan species in he mines, as well as on b owsing in ensi y in he mines. Then, we compa ed h ee di e en models: (i) a null model (wi hou he a iable “dis ance o he o es edge”); (ii) a model wi h a linea dis ance e ec (densi y = a × Dis ance + b); and (iii) a model wi h an exponen ial decay dis ance e ec ( densi y = a ×exp[−Dis ance] + b ). The bes model was chosen by he lowes AIC, and he ANOVA was applied o es he signi ican di e ences be ween he null and he al e na i e models. The s eng h o he “edge e ec ” (how and by how much he abundance dec eases owa ds he inside o mines) is desc ibed by he ype o model suppo ed ( om no e ec —null model, o mode a e e ec —linea model, o s ong e ec —exponen ial decay model) and, o a gi en ype o model, by he alues o he slope pa ame e es ima es “a”. S a is ical analyses we e implemen ed in he R so wa e en i onmen ( e sion 4.0.3; [36]) using he nlme package o LMM ( e sion 3.1–137; [37]). 3. Resul s 3.1. Ve ical S uc u e and Composi ion o Woody Vege a ion in he Fo es Edge Fo es edge woody co e nea o he mine was simila in he h ee bounda y o ms (F [2,20] = 1.33, p= 0.29), bu he e was a signi ican e ical s uc u e (F [6,442] = 28.91, p< 0.001 ) sligh ly in luenced by he bounda y o m (bounda y o m by s a um in e ac ion; F [12,442] = 2.38, p< 0.01; Figu e 1). In conca e bounda ies, he lowes s a um (0–25 cm) showed he highes co e ha was signi ican ly highe han ha in he nex wo s a a (25–50 cm and 0.5–1 m). In s aigh bounda ies, he highes co e was ound in he s a um o 2–4 m and signi ican ly di e ed om he emainde s a a, excep om he immedia ely highe (4–8 m) and he lowes (0–25 cm). In con ex bounda ies, he highes co e was ound in he s a a o 2–4 m and 4–8 m and signi ican ly di e ed om he emainde s a a, excep om he lowes (0–25 cm). Abo e 2 m high, Que cus pe aea domina ed (100% o s ems wi h dbh ≥ 10 cm and 63% o s ems wi h dbh < 10 cm) and C a aegus monogyna,Rosa canina, and Genis a lo ida we e he main accompanying species wi h dhb < 10 cm (see Table A1 in Appendix A ). Be ween 60 cm and 2 m high, Q. pe aea also domina ed (72% o s ems) wi h Rosa can- Fo es s 2021,12, 773 5 o 18 ina, Ligus um ulga e L. ,E ica a bo ea L., and Rubus ulmi olius as he main accompanying species. Below 2 m high, he pe cen age o s ems o Que cus pe aea was also impo an (ca. 26%), bu close o hose o o he sh ub species, such as Rosa canina o Cy isus scopa ius, and wi h se e al accompanying species. Fo es s 2021, 12, x FOR PEER REVIEW 5 o 19 s a a o 2–4 m and 4–8 m and signi ican ly di e ed om he emainde s a a, excep om he lowes (0–25 cm). Figu e 1. Ve ical s uc u e o woody ege a ion in he o es edge nex o he mine (mean co e and s anda d e o in each s a um) acco ding o he bounda y o m (conca e, s aigh , o con ex). Di - e en le e s indica e signi ican di e ences be ween all pai wise compa isons wi h Bon e oni’s es (p < 0.05); i.e., among s a a wi hin each bounda y o m, and among bounda y o ms o each s a um. Abo e 2 m high, Que cus pe aea domina ed (100% o s ems wi h dbh ≥ 10 cm and 63% o s ems wi h dbh < 10 cm) and C a aegus monogyna, Rosa canina, and Genis a lo ida we e he main accompanying species wi h dhb < 10 cm (see Table A1 in Appendix A). Be ween 60 cm and 2 m high, Q. pe aea also domina ed (72% o s ems) wi h Rosa canina, Ligus um ulga e L., E ica a bo ea L., and Rubus ulmi olius as he main accompanying spe- cies. Below 2 m high, he pe cen age o s ems o Que cus pe aea was also impo an (ca. 26%), bu close o hose o o he sh ub species, such as Rosa canina o Cy isus scopa ius, and wi h se e al accompanying species. 3.2. Ve ical S uc u e o Woody Vege a ion in he Mines The LMM analysis o woody co e mines showed h ee signi ican wo-way in e ac- ions (Table 1 and Figu e 2). Fi s , he e ical s uc u e o woody ege a ion in he mines depended on he o es -mine bounda y o m (Figu e 2A). In conca e bounda ies, he co e was signi ican ly highe in he lowe ou s a a (0–2 m) han in he uppe h ee. In s aigh bounda ies, he co e o he s a a be ween 25 cm and 2 m was highe only han ha in he las s a um p esen (4–8 m) since no ege a ion exceeding 8 m was ound. In con ex bounda ies, he highes co e ound in he lowes s a um only di e ed signi i- can ly om ha in he las wo. The co e in he i s ou s a a (0–2 m) was signi ican ly highe in conca e han in s aigh o con ex bounda ies (in e media e alues o he s aigh bounda y o m), whe eas he co e was simila in he h ee bounda y o ms abo e 2 m. Table 1. The summa y esul s o linea mixed models es ing he e ec o bounda y o m (conca e, s aigh , and con ex), s a um (7 le els), and dis ance (16 le els) on he co e o woody ege a ion colonizing he mines. The F- alues o he ixed ac o s and hei signi icance a e p esen ed. d F p In e cep 1 2220 284.03 <0.0001 Bounda y o m 2 20 21.51 <0.0001 S a um 6 2220 61.56 <0.0001 Dis ance 15 2220 30.47 <0.0001 Bounda y o m × s a um 12 2220 11.90 <0.0001 Bounda y o m × dis ance 30 2220 5.00 <0.0001 S a um × dis ance 90 2220 3.14 <0.0001 Bounda y o m × s a um × dis ance 180 2220 0.87 0.8833 0 1020304050607080 0-25 cm 25-50 cm 0,5-1 m 1-2 m 2-4 m 4-8 m >8 m Con ex a b a b b b ab 0 1020304050607080 0-25 cm 25-50 cm 0,5-1 m 1-2 m 2-4 m 4-8 m >8 m Woody co e (%) S aigh b ac a bc b b ac 0 1020304050607080 0−25 cm 25−50 cm 0.5−1 m 1−2 m 2−4 m 4−8 m >8 m S a a Conca e ab ab ab ab b b a ab ab Figu e 1. Ve ical s uc u e o woody ege a ion in he o es edge nex o he mine (mean co e and s anda d e o in each s a um) acco ding o he bounda y o m (conca e, s aigh , o con ex). Di e en le e s indica e signi ican di e ences be ween all pai wise compa isons wi h Bon e oni’s es (p< 0.05); i.e., among s a a wi hin each bounda y o m, and among bounda y o ms o each s a um. 3.2. Ve ical S uc u e o Woody Vege a ion in he Mines The LMM analysis o woody co e mines showed h ee signi ican wo-way in e ac- ions (Table 1and Figu e 2). Fi s , he e ical s uc u e o woody ege a ion in he mines depended on he o es -mine bounda y o m (Figu e 2A). In conca e bounda ies, he co e was signi ican ly highe in he lowe ou s a a (0–2 m) han in he uppe h ee. In s aigh bounda ies, he co e o he s a a be ween 25 cm and 2 m was highe only han ha in he las s a um p esen (4–8 m) since no ege a ion exceeding 8 m was ound. In con ex bounda ies, he highes co e ound in he lowes s a um only di e ed signi ican ly om ha in he las wo. The co e in he i s ou s a a (0–2 m) was signi ican ly highe in conca e han in s aigh o con ex bounda ies (in e media e alues o he s aigh bounda y o m), whe eas he co e was simila in he h ee bounda y o ms abo e 2 m. Table 1. The summa y esul s o linea mixed models es ing he e ec o bounda y o m (conca e, s aigh , and con ex), s a um (7 le els), and dis ance (16 le els) on he co e o woody ege a ion colonizing he mines. The F- alues o he ixed ac o s and hei signi icance a e p esen ed. d F p In e cep 1 2220 284.03 <0.0001 Bounda y o m 2 20 21.51 <0.0001 S a um 6 2220 61.56 <0.0001 Dis ance 15 2220 30.47 <0.0001 Bounda y o m ×s a um 12 2220 11.90 <0.0001 Bounda y o m ×dis ance 30 2220 5.00 <0.0001 S a um ×dis ance 90 2220 3.14 <0.0001 Bounda y o m ×s a um ×dis ance 180 2220 0.87 0.8833 Second, he o es -mine bounda y o m also in luenced he coloniza ion pa e n o woody ege a ion by inc easing he dis ance o he o es edge (Figu e 2B). The co e was signi ican ly highe in con ex han in conca e o s aigh bounda ies a 3 m. Be ween 3 and 7 m, he e was a signi ican co e dec ease in con ex and conca e bounda ies (much s onge in con ex bounda ies) whe eas i was g adual in s aigh bounda ies. F om 7 m, he co e did no di e wi h he dis ance o any bounda y o m. Ne e heless, he co e Fo es s 2021,12, 773 6 o 18 was signi ican ly highe in conca e han in s aigh o con ex bounda ies om 23 o 39 m, al hough he di e ences we e no always signi ican . Fo es s 2021, 12, x FOR PEER REVIEW 6 o 19 (A) (B) (C) Figu e 2. (A) Ve ical s uc u e o woody ege a ion in he mines acco ding o he bounda y o m. (B) Woody co e in he mines along wi h he dis ance om he o es -mine limi acco ding o he bounda y o m. (C) Ve ical s uc u e o woody ege a ion in he mines acco ding o he dis ance om he o es edge. Mean co e and s anda d e o a e shown. Di e en le e s indica e signi ican di e ences be ween all pai wise compa isons wi h Bon e oni’s es (p < 0.05). 0 10203040 0−25 cm 25−50 cm 0.5−1 m 1−2 m 2−4 m 4−8 m >8 m Con ex c cc bd bd bc bc bc bc 0 10203040 0−25 cm 25−50 cm 0.5−1 m 1−2 m 2−4 m 4−8 m >8 m S aigh bd bc cd c c ce Woody co e (%) 0 10203040 0−25 cm 25−50 cm 0.5−1 m 1−2 m 2−4 m 4−8 m >8 m S a a Conca e bd a ae bc bd ae a 0 10 20 30 40 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 Co e (%) Dis ance (m) Conca e S aigh Con ex 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 conca e e cij ade i cdhi ad i cei cdgh ce cd cdg ac ad i ac ad j aghj adj s aigh ce cde ad i acd ad i ai a a ai ai adj ahj ahj ahj ahj a con ex bacadjad jaaaaaaadjaaaaa 0 10 20 30 40 50 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 Co e (%) Dis ance (m) 0-25 cm 25-50 cm 0.5-1 m 1-2 m 2-4 m 4-8 m >8 m 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 0-25 cm b ac ad ad ad ad ad ad ad ad ad ad ad ad ad ad 25-50 m ac c g ac ce g adg c g ac ac adg adg acg adg adg adg adg adg 0.5-1 m ac cdh ac ch ac cdi ac ac ac ac ac ac ac ac ac ac 1-2 m bc cd ad cde ade ade ade ade ade cde ade ade ade ade ae ae 2-4 m b adgh aeghi adgh aeg ae aegh aeghi aeghi aeghi ae aeghi ae ae ae ae 4-8 m b >8 m ac Figu e 2. ( A ) Ve ical s uc u e o woody ege a ion in he mines acco ding o he bounda y o m. ( B ) Woody co e in he mines along wi h he dis ance om he o es -mine limi acco ding o he bounda y o m. ( C ) Ve ical s uc u e o woody ege a ion in he mines acco ding o he dis ance om he o es edge. Mean co e and s anda d e o a e shown. Di e en le e s indica e signi ican di e ences be ween all pai wise compa isons wi h Bon e oni’s es (p< 0.05). Fo es s 2021,12, 773 7 o 18 Thi d, he e ical s uc u e o woody ege a ion in he mines also depended on he dis ance om he o es edge (Figu e 2C). A 3 m o dis ance om he o es edge, he co e was simila in he i s (0–25 cm), i h (2–4 m), and six h (4–8 m) s a a bu highe han ha in he second (25–50 cm), hi d (0.5–1 m) and se en h (>8 m) s a a; he co e was in e media e in he ou h s a um (1–2 m). A 23 m, he co e in he i h s a um ( 2–4 m ) was signi ican ly lowe han ha in he second (25–50 cm) and hi d (0.5–1 m) s a a; he co e was in e media e in he i s (0–25 cm) and ou h (1–2 m) s a a. The co e was simila in all he s a a o he emaining dis ances. 3.3. Bounda y Fo m E ec on To al Woody Coloniza ion (Densi y) in he Mines We ound ha woody densi y declined exponen ially wi h an inc easing dis ance om he o es edge owa ds he inside o mines (F [1,342] = 32.45, p< 0.0001; Table 2); o y-one pe cen o all colonizing s ems we e wi hin 13 m om he o es edge. Such an edge e ec was e iden in he h ee bounda y ypes (Table 2; Figu e 3) bu i s magni ude a ied depending on he bounda y o m. In all cases, he null models (no edge e ec ) o he linea models (linea dec ease in abundance owa ds he in e io o mines) we e he leas suppo ed (Table 2). The e was also a signi ican bounda y o m x exp[-dis ance] in e - ac ion (F[2,342] = 6.09, p= 0.0025). The slope pa ame e ha es ima es compa isons among bounda y ypes indica ed ha he exponen ial decay in woody coloniza ion in ensi y in he mines wi h inc easing dis ance o he o es edge was s onge in con ex (slope ± SE: 2.16 ± 0.35) han in s aigh bounda ies (1.23 ± 0.39) bu i was no signi ican in conca e bounda ies. In gene al, woody densi y in conca e bounda ies (4.18 ± 0.32 s ems/m 2 ) was 1.4 imes g ea e han in s aigh bounda ies (2.89 ± 0.35 s ems/m 2 ; = 2.59, p= 0.0175), and 2.1 imes g ea e han in he con ex bounda ies (2.03 ± 0.34 s ems/m 2 ; = 4.33, p= 0.0003 ). Table 2. Resul s o Linea Mixed Models (LMM) desc ibing how woody densi y in mines a ied wi h inc easing dis ance o he o es edge. The bes models (lowes AICs) a e highligh ed in bold. p indica es he signi ican di e ences be ween he null and he al e na i e models a e he ANOVA. Model d. . AIC ∆AIC p Global Null 3 2030.24 35.54 Lineal 8 2016.43 21.73 <0.0001 Exp. decay 8 1994.70 0.00 <0.0001 Bounda y ype Conca e Null 3 802.36 8.49 Lineal 4 807.52 13.65 0.079 Exp. decay 4 793.87 0.00 0.049 Con ex Null 3 608.36 46.85 Lineal 4 598.55 37.04 <0.001 Exp. decay 4 561.51 0.00 <0.001 S aigh Null 3 612.17 13.10 Lineal 4 612.97 13.90 0.007 Exp. decay 4 599.07 0.00 0.010 3.4. Bounda y Fo m E ec on Main Woody Colonize s o Mines The Que cus pe aea (Figu e 4A) and C a aegus monogyna (Figu e 4D) densi y declined exponen ially wi h inc easing dis ance o he o es edge (Table 3) bu wi h di e en in ensi y depending on he bounda y o m (Table A2 in Appendix A). The Que cus pe aea decay was s onge in con ex (slope ± SE: 1.49 ± 0.21) han in conca e (0.56 ± 0.12) and s aigh bounda ies (0.71 ± 0.29). The C a aegus monogyna decay was he s onges in conca e bounda ies (slope ± SE: 0.40 ± 0.12), in e media e in he con ex (0.16 ± 0.04), and no signi ican in he s aigh . Fo es s 2021,12, 773 8 o 18 Fo es s 2021, 12, x FOR PEER REVIEW 8 o 19 Figu e 3. A e age woody densi y (s ems/m2) and s anda d e o acco ding o he dis ance om he o es edge (m) in he h ee bounda y ypes (conca e, con ex, and s aigh ). 3.4. Bounda y Fo m E ec on Main Woody Colonize s o Mines The Que cus pe aea (Figu e 4A) and C a aegus monogyna (Figu e 4D) densi y declined exponen ially wi h inc easing dis ance o he o es edge (Table 3) bu wi h di e en in- ensi y depending on he bounda y o m (Table A2 in Appendix A). The Que cus pe aea decay was s onge in con ex (slope ± SE: 1.49 ± 0.21) han in conca e (0.56 ± 0.12) and s aigh bounda ies (0.71 ± 0.29). The C a aegus monogyna decay was he s onges in con- ca e bounda ies (slope ± SE: 0.40 ± 0.12), in e media e in he con ex (0.16 ± 0.04), and no signi ican in he s aigh . Table 3. Resul s o Linea Mixed Models (LMM) desc ibing how he densi y o main woody colo- nize s o he mines a ied acco ding o bounda y o m, dis ance o he o es edge, and hei in e - ac ion. F- alues o ixed e ec s and hei signi icance (* p < 0.05; ** p < 0.01; *** p < 0.001) o he bes model (lowes AIC; see Table A2 in Appendix A) a e shown. Bes Model Bounda y Fo m Dis ance Bounda y Fo m × Dis ance Que cus pe aea Exp. decay 1.17 119.19 *** 13.36 *** Cy isus scopa ius Exp. g ow h 13.47 *** 7.23 ** 2.57 * Genis a lo ida Exp. decay 0.68 1.58 0.52 C a aegus monogyna Exp. decay 0.63 29.94 *** 8.70 *** Rosa canina Exp. decay 0.88 7.31 ** 2.84 Rubus ulmi olius Exp. decay 1.54 9.67 ** 2.12 Unlike p e ious species, he Cy isus scopa ius densi y inc eased exponen ially wi h inc easing dis ance o he o es edge (Table 3; Figu e 4B; Table A2 in Appendix A) bu wi h di e en in ensi y depending on he bounda y o m (Table A2 in Appendix A). The inc ease was only signi ican in conca e bounda ies, whe eas i was no signi ican in con- ex and s aigh bounda ies. So, om a ce ain dis ance (23 m), di e ences in densi y we e ound be ween conca e and con ex o s aigh bounda ies (Figu e 4B). The Genis a lo ida densi y was simila ega dless o bounda y o m and dis ance (Table 3; Figu e 4C). The Rosa canina (Figu e 4E) and Rubus ulmi olius (Figu e 4F) densi y declined exponen ially wi h inc easing dis ance o he o es edge ega dless o he bounda y o m (Table 3). As a whole, Que cus pe aea showed he mos ex eme pa e n wi h 94% o he s ems wi hin 3 m om he o es edge. In addi ion, C. monogyna was pa icula ly es ic ed o he nea he o es edge (60% o he s ems wi hin 7 m om he o es edge). R. canina, and R. ulmi olius we e pa icula ly abundan nea he o es edge, and a a he dis ances (Rosa canina a 39 m o 63 m, and Rubus ulmi olius a 19 m). Ne e heless, o e all coloniza ion beyond 11– 15 m om he o es edge was p edominan ly by C. scopa ius and G. lo ida. 0 2 4 6 8 10 12 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 Densi y (s ems/m 2 ) Dis ance (m) conca e con ex s aigh Figu e 3. A e age woody densi y (s ems/m 2 ) and s anda d e o acco ding o he dis ance om he o es edge (m) in he h ee bounda y ypes (conca e, con ex, and s aigh ). Fo es s 2021, 12, x FOR PEER REVIEW 9 o 19 Figu e 4. A e age woody densi y (s ems/m2) and s anda d e o (SE) o six main woody species (A−F) colonizing he mines ( hose ha accoun ed o mo e han 2% o all he colonizing s ems), acco ding o he dis ance o he o es edge (m) in he h ee bounda y ypes (conca e, con ex, and s aigh ). 3.5. In luence o B owsing Only 2.2 ± 0.5% o he s ems colonizing he mine showed e idence o b owsing com- pa ed o 25.6 ± 2.6% in he o es , he di e ence being s a is ically signi ican (F[1.414] = 214.96, p < 0.0001). Que cus pe aea, Cy isus scopa ius, and C a aegus monogyna we e he mos b owsed species (67%, 28%, and 5% o all b owsed s ems, espec i ely) wi h simila b owsing in ensi y in o es and mines and ega dless o he bounda y o m (s a is ical analysis esul s a e omi ed o simplici y). O e all b owsing in ensi y (pe cen o b owsed s ems o he o al o colonizing s ems) in he mines declined exponen ially wi h he dis ance o he o es edge (F[1,229] = 22,55, p < 0.0001). Pa icula ly, 52% o he b owsed s ems we e wi hin he i s 7 m om he o es edge, and e idence o b owsing was conside ably educed beyond 15 m. The e was also a signi ican bounda y o m x exp[-dis ance] in e ac ion (F[2,229] = 4.48, p = 0.0123; Figu e 5). In all cases, he null models (no edge e ec ) o he linea models (linea dec ease in abun- dance owa ds he in e io o mines) we e he leas suppo ed (Table A3 in Appendix A). The exponen ial decay was s onge in con ex (slope ± SE: 5.90 ± 1.58) han in conca e bounda ies (2.88 ± 1.36), and i was no signi ican in s aigh bounda ies. 0 2 4 6 8 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 Densi y (s ems/m 2 ) Dis ance (m) (B) Cy isus scopa ius 0 2 4 6 8 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 Densi y (s ems/m 2 ) Dis ance (m) conca e con ex s aigh (A) Que cus pe aea 0 1 2 3 4 5 6 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 Densidad (ind/m 2 ) Dis ancia al bo de del bosque (m) (D) C a aegus monogyna 0 1 2 3 4 5 6 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 Densi y (s ems/m 2 ) Dis ance (m) (C) Genis a lo ida 0 1 2 3 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 Densi y (s ems/m 2 ) Dis ance (m) ( F ) Rubus ulmi olius 0 1 2 3 3 7 11 15 19 23 27 31 35 39 43 47 51 55 59 63 Densi y (s ems/m 2 ) Dis ance (m) ( E ) Rosa canina Figu e 4. A e age woody densi y (s ems/m 2 ) and s anda d e o (SE) o six main woody species ( A−F ) colonizing he mines ( hose ha accoun ed o mo e han 2% o all he colonizing s ems), acco ding o he dis ance o he o es edge (m) in he h ee bounda y ypes (conca e, con ex, and s aigh ). Fo es s 2021,12, 773 9 o 18 Table 3. Resul s o Linea Mixed Models (LMM) desc ibing how he densi y o main woody colonize s o he mines a ied acco ding o bounda y o m, dis ance o he o es edge, and hei in e ac ion. F- alues o ixed e ec s and hei signi icance (* p< 0.05; ** p< 0.01; *** p< 0.001) o he bes model (lowes AIC; see Table A2 in Appendix A) a e shown. Bes Model Bounda y Fo m Dis ance Bounda y Fo m ×Dis ance Que cus pe aea Exp. decay 1.17 119.19 *** 13.36 *** Cy isus scopa ius Exp. g ow h 13.47 *** 7.23 ** 2.57 * Genis a lo ida Exp. decay 0.68 1.58 0.52 C a aegus monogyna Exp. decay 0.63 29.94 *** 8.70 *** Rosa canina Exp. decay 0.88 7.31 ** 2.84 Rubus ulmi olius Exp. decay 1.54 9.67 ** 2.12 Unlike p e ious species, he Cy isus scopa ius densi y inc eased exponen ially wi h inc easing dis ance o he o es edge (Table 3; Figu e 4B; Table A2 in Appendix A) bu wi h di e en in ensi y depending on he bounda y o m (Table A2 in Appendix A). The inc ease was only signi ican in conca e bounda ies, whe eas i was no signi ican in con ex and s aigh bounda ies. So, om a ce ain dis ance (23 m), di e ences in densi y we e ound be ween conca e and con ex o s aigh bounda ies (Figu e 4B). The Genis a lo ida densi y was simila ega dless o bounda y o m and dis ance (Table 3; Figu e 4C). The Rosa canina (Figu e 4E) and Rubus ulmi olius (Figu e 4F) densi y declined exponen ially wi h inc easing dis ance o he o es edge ega dless o he bounda y o m (Table 3). As a whole, Que cus pe aea showed he mos ex eme pa e n wi h 94% o he s ems wi hin 3 m om he o es edge. In addi ion, C. monogyna was pa icula ly es ic ed o he nea he o es edge (60% o he s ems wi hin 7 m om he o es edge). R. canina, and R. ulmi olius we e pa icula ly abundan nea he o es edge, and a a he dis ances (Rosa canina a 39 m o 63 m, and Rubus ulmi olius a 19 m). Ne e heless, o e all coloniza ion beyond 11–15 m om he o es edge was p edominan ly by C. scopa ius and G. lo ida. 3.5. In luence o B owsing Only 2.2 ± 0.5% o he s ems colonizing he mine showed e idence o b owsing com- pa ed o 25.6 ±2.6% in he o es , he di e ence being s a is ically signi ican (F[1.414] = 214.96 , p< 0.0001). Que cus pe aea,Cy isus scopa ius, and C a aegus monogyna we e he mos b owsed species (67%, 28%, and 5% o all b owsed s ems, espec i ely) wi h simila b ows- ing in ensi y in o es and mines and ega dless o he bounda y o m (s a is ical analysis esul s a e omi ed o simplici y). O e all b owsing in ensi y (pe cen o b owsed s ems o he o al o colonizing s ems) in he mines declined exponen ially wi h he dis ance o he o es edge (F [1,229] = 22,55, p< 0.0001 ). Pa icula ly, 52% o he b owsed s ems we e wi hin he i s 7 m om he o es edge, and e idence o b owsing was conside ably educed beyond 15 m. The e was also a signi ican bounda y o m x exp[-dis ance] in e ac ion (F [2,229] = 4.48, p= 0.0123; Figu e 5 ). In all cases, he null models (no edge e ec ) o he linea models (linea dec ease in abun- dance owa ds he in e io o mines) we e he leas suppo ed (Table A3 in Appendix A ). The exponen ial decay was s onge in con ex (slope ± SE: 5.90 ± 1.58) han in conca e bounda ies (2.88 ±1.36), and i was no signi ican in s aigh bounda ies. Fo es s 2021,12, 773 16 o 18 Table A3. Resul s o Linea Mixed Models (LMM) desc ibing how b owsing in ensi y in mines a ied wi h inc easing dis ance o he o es edge. The bes models (lowes AICs) a e highligh ed in bold. p indica es he signi ican di e ences be ween he null and he al e na i e models a e he ANOVA. Model d. . AIC ∆AIC p Global Null 3 1999.93 57.41 Lineal 8 2002.52 60.00 0.192 Exp. decay 8 1942.52 0.00 <0.0001 Bounda y ype Conca e Null 3 792.76 24.75 Lineal 4 797.77 29.76 0.008 Exp. decay 4 768.01 0.00 <0.0001 Con ex Null 3 644.06 16.8 Lineal 4 641.37 14.11 0.0304 Exp. decay 4 627.26 0.00 <0.0001 S aigh Null 16 545.25 8.05 Lineal 551.61 14.41 0.0368 Exp. decay 537.20 0.00 0.0015 Re e ences 1. Fo man, R.T.T.; Moo e, P.N. Theo e ical ounda ions o unde s anding bounda ies. In Landscape Bounda ies: Consequences o Bio ic Di e si y and Ecological Flows; Hansen, A.J., di Cas i, F., Eds.; Sp inge : New Yo k, NY, USA, 1992. 2. Fo man, R.T.T. Land Mosaics: The Ecology o Landscape and Regions; Camb idge Uni e si y P ess: Camb idge, UK, 1995. 3. Ha d , R.A.; Fo man, R.T.T. Bounda y o m e ec s on woody coloniza ion o eclaimed su ace mines. Ecology 1989 ,70, 1252–1260. [C ossRe ] 4. Fo man, R.T.T.; God on, M. Landscape Ecology; John Wiley & Sons: New Yo k, NY, USA, 1986. 5. Fo man, R.T.T. Eme ging di ec ions in landscape ecology and applica ions in na u al esou ce managemen . In P oceedings o he Con e ence on Science in he Na ional Pa ks; He mann, R., Bos ed -C aig, T., Eds.; US Na ional Pa k Se ice and he Geo ge W igh Socie y: Fo Collins, CO, USA, 1986; pp. 59–88. 6. López-Ba e a, F.; Manson, R.H.; González-Espinosa, M.; New on, A. E ec s o a ying o es edge pe meabili y on seed dispe sal in a neo opical mon ane o es . Landsc. Ecol. 2007,22, 189–203. [C ossRe ] 7. Milde , A.I.; Fe nández-San os, B.; Ma ínez-Ruiz, C. Coloniza ion pa e ns o woody species on lands mined o coal in Spain: P elimina y insigh o o es expansion. Land Deg ad. De elop. 2013,24, 39–46. [C ossRe ] 8. S amps, J.A.; Buechne , M.; K ishnan, V.V. The e ec s o edge pe meabili y and habi a geome y on emig a ion om pa ches o habi a . Am. Na . 1987,129, 533–552. [C ossRe ] 9. López-Ba e a, F. Es uc u a y unción en bo des de bosques. Ecosis emas 2004 ,1, 55–68. A ailable online: h ps://www. e is aecosis emas.ne /index.php/ecosis emas/a icle/ iew/581 (accessed on 13 Decembe 2020). 10. Ries, L.; Fle che , R.J.; Ba in, J.; Sisk, T.D. Ecological esponses o habi a edges: Mechanisms, models and a iabili y explained. Annu. Re . Ecol. E ol. Sys . 2004,35, 491–522. [C ossRe ] 11. López-Ba e a, F.; New on, A.C.; Manson, R.H. Edge e ec s in a opical moun ains o es mosaic: Expe imen al es s o pos -dispe sal aco n emo al. Ecol. Res o . 2005,20, 31–40. [C ossRe ] 12. Cadenasso, M.L.; Picke , S.T.A. Linking o es edge s uc u e o edge unc ion: Media ion o he bi o e damage. J. Ecol. 2000 ,88, 31–44. [C ossRe ] 13. Cadenasso, M.L.; Picke , S.T.A. E ec o edge s uc u e on he lux o species in o o es in e io s. Conse . Biol. 2001 ,15, 91–97. [C ossRe ] 14. Ha pe , K.A.; Macdonald, S.E.; Bu on, P.J.; Chen, J.; B oso ske, K.D.; Saunde s, S.C.; Esseen, P.A. Edge in luence on o es s uc u e and composi ion in agmen ed landscapes. Conse . Biol. 2005,19, 768–782. [C ossRe ] 15. McQuilkin, W.R. The na u al es ablishmen o pine in abandoned ields in he Piedmon Pla eau Region. Ecology 1940 ,21, 135–147. [C ossRe ] 16. Ha on, T.J.; Ca pen e , A.T. An empi ical es o he mass e ec de e minan o species ichness. Vege a io 1986 ,68, 33–36. [C ossRe ] 17. González-Alday, J.; Ma s, R.; Ma ínez-Ruiz, C. Soil seed bank o ma ion du ing ea ly e ege a ion a e hyd oseeding in eclaimed coal was es. Ecol. Eng. 2009,35, 1062–1069. [C ossRe ] 18. Pa men e , R.R.; MacMahon, J.A.; Waaland, M.E.; S eube, M.M.; C isa ulli, C.M. Reclama ion o su ace coal mines in wes e n Wyoming o wildli e habi a . Reclam. Re eg. Res. 1985,4, 93–115. 19. Gómez, J.M.; Ga cía, D.; Zamo a, R. Impac o e eb a e aco n- and seedling-p eda o s on a Medi e anean Que cus py enaica o es . Fo . Ecol. Manag. 2003,180, 125–134. [C ossRe ] Fo es s 2021,12, 773 17 o 18 20. To oba-Balmo i, P.; Zaldí a , P.; Alday, J.G.; Fe nández-San os, B.; Ma ínez-Ruiz, C. Reco e ing Que cus species on eclaimed coal was es using na i e sh ubs as es o a ion nu se plan s. Ecol. Eng. 2015,77, 146–153. [C ossRe ] 21. Cos a, A.; Villa, S.; Alonso, P.; Ga cía-Rod íguez, J.A.; Ma ín, F.J.; Ma ínez-Ruiz, C.; Fe nández-San os, B. Can na i e sh ubs acili a e he ea ly es ablishmen o con as ed co-occu ing oaks in Medi e anean g azed a eas? J. Veg. Sci. 2017 ,28, 1047–1056. [C ossRe ] 22. Callaway, R.M. E ec o sh ub on ec ui men o Que cus douglasii and Que cus loba a in Cali o nia. Ecology 1992 ,73, 2118–2128. [C ossRe ] 23. Rousse , O.; Lepa , J. Posi i e and nega i e in e ac ions a di e en li e s ages o a colonizing species (Que cus humilis). J. Ecol. 2000,88, 401–412. [C ossRe ] 24. Mu cia, C. Edge e ec s in agmen ed o es s: Implica ions o conse a ion. T ends Ecol. E ol. 1995,10, 58–62. [C ossRe ] 25. P ach, K.; Hobbs, R.J. Spon aneous succession e sus echnical eclama ion in he es o a ion o dis u bed si es. Res o . Ecol. 2008 , 16, 363–366. [C ossRe ] 26. P ach, K.; Walke , L.R. Fou oppo uni ies o s udies o ecological succession. T ends Ecol. E ol. 2011 ,26, 119–123. [C ossRe ] [PubMed] 27. MAPA. Ca ac e ización Ag oclimá ica de la P o incia de Palencia; Minis e io de Ag icul u a Pesca y Alimen ación: Mad id, Spain, 1991. 28. Sigcha, F.; Palla icini, Y.; Camino, M.J.; Ma ínez-Ruiz, C. E ec s o sho - e m g azing exclusion on ege a ion and soil in ea ly succession o a Subhumid Medi e anean eclaimed coal mine. Plan Soil 2018,426, 197–209. [C ossRe ] 29. Soil-Su ey-S a . Keys o Soil axonomy, 12 h ed.; USDA-Na u al Resou ces Conse a ion Se ice: Washing on, WA, USA, 2014. [C ossRe ] 30. López-Ma cos, D.; Tu ión, M.B.; Ma ínez-Ruiz, C. Linking soil a iabili y wi h plan communi y composi ion along a mine-slope opog aphic g adien : Implica ions o es o a ion. Ambio 2020,49, 337–349. [C ossRe ] [PubMed] 31. Richa ds, A. Tes ing ecological heo y using he in o ma ion heo e ic app oach: Examples and cau iona y esul s. Ecology 2005 , 86, 2805–2814. [C ossRe ] 32. Akaike, H. In o ma ion heo y as an ex ension o he maximum likelihood p inciple. In Second In e na ional Symposium on In o ma ion Theo y; Pe o , B.N., Csaki, F., Eds.; Akademiai Kiado: Budapes , Hunga y, 1973; pp. 267–281. 33. Pinhei o, J.; Ba es, D. Mixed-E ec s Models in S and S-Plus; Sp inge : New Yo k, NY, USA, 2000. 34. Sokal, R.R.; Rohl , F.J. Biome y: The P inciples and P ac ice o S a is ics in Biological Resea ch, 3 d ed.; W.H. F eeman and Company: New Yo k, NY, USA, 1995. 35. González-Alday, J.; Ma s, R.; Ma ínez-Ruiz, C. The in luence o aspec on he ea ly g ow h dynamics o hyd oseeded species in coal eclama ion a eas. Appl. Veg. Sci. 2008,11, 405–412. [C ossRe ] 36. R De elopmen Co e Team. R: A Language and En i onmen o S a is ical Compu ing; R Founda ion o S a is ical Compu ing: Vienna, Aus ia, 2011; A ailable online: h p://www. -p ojec .o g (accessed on 1 Feb ua y 2020). 37. Pinhei o, J.; Ba es, D.; DebRoy, S.; Sa ka , D.; R-Co e-Team. Package “nlme.” nlme: Linea and Nonlinea Mixed E ec s Models; R Package Ve sion 3.1-137; R Founda ion o S a is ical Compu ing: Vienna, Aus ia, 2018; A ailable online: h ps://CRAN.R- p ojec .o g/package=nlme (accessed on 1 Feb ua y 2020). 38. Vallada es, F.; Balague , L.; Ma ínez Fe i, E.; Pé ez-Co ona, E.; Man ique, E. Plas ici y, ins abili y and canaliza ion: Is he pheno ypic a ia ion in seedlings o scle ophyll oaks consis en wi h he en i onmen al unp edic abili y o Medi e anean ecosys ems? New Phy ol. 2002,156, 457–467. [C ossRe ] [PubMed] 39. Chambel, M.R.; Climen , J.; Alia, R.; Vallada es, F. Pheno ypic plas ici y: A use ul amewo k o unde s anding adap a ion in o es species. In es ig. Ag a . Sis . Recu . Fo . 2005,14, 334–344. [C ossRe ] 40. Tá ega, R.; Cal o, L.; T abaud, L. E ec o high empe a u eas on seed ge mina ion o wo woody Leguminosae. Vege a io 1992 , 102, 139–147. [C ossRe ] 41. Alday, J.G.; Zaldí a , P.; To oba-Balmo i, P.; Fe nández-San os, B.; Ma ínez-Ruiz, C. Na u al o es expansion on eclaimed coal mines in No he n Spain: The ole o na i e sh ubs as sui able mic osi es. En i on. Sci. Pollu . Res. 2016 ,23, 13606–13616. [C ossRe ] 42. Hosho sky, M. Elemen s ewa dship abs ac o Cy isus scopa ius and Genis a monspessulana. In Sco ch B oom, F ench B oom; The Na u e Conse ancy: A ling on, VA, USA, 1986. 43. Malo, J.E. Po en ial ballis ic dispe sal o Cy isus scopa ius (Fabaceae) seeds. Aus al. J. Bo . 2004,52, 653–658. [C ossRe ] 44. Mo eno-Ma cos, G.; Gómez-Gu ié ez, J.M.; Fe nández-San os, B. P ima y dispe sal o Cy isus mul i lo us seeds. Pinineos 1992 , 140, 75–88. [C ossRe ] 45. Malo, J.E.; Suá ez, F. He bi o ous mammals as seed dispe se in Medi e anean dehesa. Am. Na . 1995 ,123, 338–353. [C ossRe ] [PubMed] 46. Oso o, K.; Celaya, R.; Ma ínez, A.; Zo i a, E. Pas o eo de las comunidades ege ales de mon aña po umian es domés icos: P oducción animal y dinámica ege al. Pas os 2000,30, 3–50. 47. Milde , A.I. S udy o Mechanisms ha P omo e Fo es Res o a ion in Deg aded A eas: Facili a ion, Fo es Edge Shape and P opaga ion S a egies. Ph.D. Thesis, Uni e si y o Salamanca, Salamanca, Spain, 2016. 48. Baue , H.J. Ten yea s’ s udies o biocenological succession in he exca a ed mines in he Cologne dis ic . In Ecology and Reclama ion o De as a ed Land; Hu nik, R.J., Da is, G., Eds.; Go don and B each: New Yo k, NY, USA, 1973; pp. 271–284. Fo es s 2021,12, 773 18 o 18 49. Pe e son, D.J.; P asa d, R. The biology o Canadian weeds. 109. Cy isus scopa ius (L.) Link. Can. J. Plan Sci. 1998 ,78, 497–504. [C ossRe ] 50. Foga y, G.; Facelli, J.M. G ow h and compe i ion o Cy isus scopa ius, an in asi e sh ub, and Aus alian na i e sh ubs. Plan Ecol. 1999,144, 27–35. [C ossRe ] 51. C uz, O.; Ga cía-Du o, J.; Ri ei o, S.F.; Ga cía-Ga cía, C.; Casal, M.; Reyes, O. Fi e se e i y d i es he na u al egene a ion o Cy isus scopa ius L.(Link) and Salix a ocine ea B o . Communi ies and he ge mina i e beha iou o hese Species. Fo es s 2020 , 11, 124. [C ossRe ] 52. Rod íguez-Calce ada, J.; Pa dos, J.A.; Gil, L.; Reich, P.B.; A anda, I. Ligh esponse in seedlings o a empe a e (Que cus pe aea) and a sub-Medi e anean species (Que cus py enaica): Con as ing ecological s a egies as po en ial keys o egene a ion pe o mance in mixed ma ginal popula ions. Plan Ecol. 2008,195, 273–285. [C ossRe ] 53. Do Ama al F anco, J.; Que cus, L. Flo a Ibe ica; Cas o, S., Laínz, M., López González, G., Mon se a , P., Muñoz Ga mendia, F., Pai a, J., Villa , L., Eds.; Real Ja dín Bo ánico, CSIC: Mad id, Spain, 1990; Volume 2, pp. 15–36. 54. Ol ho , A.; Ma ínez-Ruiz, C.; Alday, J.G. Dis ibu ion pa e ns o o es species along an A lan ic-Medi e anean en i onmen al g adien : An app oach om o es in en o y da a. Fo es y 2016,86, 46–54. [C ossRe ] 55. Gómez, J.M.; Pue a-Piñe o, C.; Schupp, E.W. E ec i eness o oden s as local seed dispe se s o Holm oaks. Oecologia 2008 ,155, 529–537. [C ossRe ] 56. Mülle , S.C.; O e beck, G.E.; P adenhaue , J.; Pilla , V.D. Plan unc ional ypes o woody species ela ed o i e dis u bance in o es -g assland eco ones. Plan Ecol. 2007,189, 1–14. [C ossRe ] 57. Díaz-He nández, R.; Vicen e Villa dón, J.L.; Ma ínez-Ruiz, C.; Fe nández-San os, B. The e ec s o na i e sh ub, encing, and aco n size on he eme gence o con as ing co-occu ing oaks in Medi e anean g azed a eas. Fo es s 2021,12, 307. [C ossRe ] 58. Alday, J.G.; San ana, V.M.; Ma s, R.H.; Ma ínez-Ruiz, C. Sh ub-induced unde s o y ege a ion changes in eclaimed mine si es. Ecol. Eng. 2014,73, 691–698. [C ossRe ] 59. Bu schel, P.; Huss, J. G und iß des Waldbaus; Pa ey: Be lin, Ge many, 2003. 60. Ca lo, T.A.; Ga cía, D.; Ma ínez, D.; Mo ales, J.M. Whe e do seeds go when hey go a ? In eg a ing dis ance and di ec ionali y o a ian seed dispe sal in he e ogeneous landscapes. Ecology 2013,94, 301–307. [C ossRe ] [PubMed] 61. C uz-Padilla, M. Ecogene ic S udy o Plan Popula ions in Al e ed Media: Coal Mining. Ph.D. Thesis, Uni e si y o León, León, Spain, 1993. 62. Williams-Line a, G. Vege a ion s uc u e and en i onmen al condi ions o o es edges in Panama. J. Ecol. 1990 ,78, 356–373. [C ossRe ] 63. Ma ínez-Ruiz, C.; Ma s, R.H. Some ac o s a ec ing successional change on u anium mine was es: Insigh s o ecological es o a ion. Appl. Veg. Sci. 2007,10, 333–342. [C ossRe ] 64. Ma ín-Sanz, R.C.; Fe nández-San os, B.; Ma ínez-Ruiz, C. Ea ly dynamics o na u al e ege a ion on oadcu s o he Salamanca p o ince (CW Spain). Ecol. Eng. 2015,75, 223–231. [C ossRe ] 65. Schmi z, O.J. Scaling om plo expe imen s o landscapes: S udying g asshoppe s o in o m o es ecosys em managemen . Oecologia 2005,145, 225–234. [C ossRe ] 66. Ro hley, K.D. Use o mul iobjec i e op imiza ion models o examine beha iou al ade-o s o whi e- ailed dee habi a use o es ha es ing expe imen s. Can. J. Fo . Res. 2002,32, 1275–1284. [C ossRe ] 67. Amma , H.; López, S.; González, J.S.; Ranilla, M.J. Seasonal a ia ions in he chemical composi ion and in i o diges ibili y o some Spanish leguminous sh ub species. Anim. Feed Sci. Technol. 2004,115, 327–340. [C ossRe ] 68. K ame , K.; G oo B uinde ink, G.W.T.A.; P ins, H.H.T. Spa ial in e ac ions be ween ungula e he bi o y and o es managemen . Fo . Ecol. Manag. 2006,226, 238–247. [C ossRe ] 69. Bobiec, A.; Kuijpe , D.P.J.; Niklasson, M.; Romankiewicz, A.; Solecka, K. Oak (Que cus obu L.) egene a ion in ea ly successional woodlands g azed by wild ungula es in he absence o li es ock. Fo . Ecol. Manag. 2011,262, 780–790. [C ossRe ]