1
Does o es agmen a ion a ec he same way all g ow h- o ms?
Glo ia RODRÍGUEZ-LOINAZ1*, Ibone AMEZAGA1,2, Mi en ONAINDIA1,3.
1Depa men o Plan Biology and Ecology. Uni e si y o he Basque Coun y, P.O. Box
644. 48080 Bilbao, Spain;
2 E-mail: [email protected]; 3 E-mail: [email p o ec ed];
* Co esponding au ho ; Fax: 34-94-6013500; Phone: 34 94 601 2559;
E-mail: [email p o ec ed]s
This documen is he Accep ed Manusc ip e sion o a Published Wo k ha appea ed in inal o m in Jou nal o En i onmen al Managemen
94 :125-131 (2012). h ps://doi.o g/10.1016/j.jen man.2011.06.024
© 2012. This manusc ip e sion is made a ailable unde he CC-BY-NC-ND 4.0 license h ps://c ea i ecommons.o g/licenses/by-nc-nd/4.0/
2
Does o es agmen a ion a ec he same way all g ow h- o ms? 1
2
F agmen a ion o na u al habi a s is one o he main causes o he loss o 3
biodi e si y. Howe e , no all plan species show a uni o m esponse o habi a 4
agmen a ion due o di e ences in species ai s. We s udied he e ec o pa ch size 5
and isola ion on he biodi e si y o ege a ion in he mixed-oak o es s in he no h o 6
he Ibe ian Peninsula. The aim was o e alua e whe he all he g ow h- o ms o 7
ege a ion a e equally a ec ed by o es agmen a ion in o de o imp o e he 8
managemen s a egies o es o e his ype o ege a ion. 9
This s udy has shown ha he e ec o he a ea and spa ial isola ion o he 10
pa ches was no he same o he di e en g ow h- o ms. F agmen a ion had a mainly 11
nega i e e ec on he ichness and di e si y o o es specialis species, especially e ns 12
and he baceous g ow h- o ms. Mo eo e , he p esence and/o co e o woodland 13
he baceous species (such as Lamias um galeobdolon and Hellebo us i idis) and o 14
woodland e ns (namely Asplenium adian um-nig um, Asplenium ichomanes, 15
Polys ichum se i e um, D yop e is a inis) we e nega i ely a ec ed by pa ch size, 16
possibly due o he educ ion o habi a quali y. This species ha e been eplaced by 17
mo e gene alis species (such as Ca damine p a ensis, Ci sium sp., Pulmona ia 18
longi olia o Rumex ace osella) in small pa ches. Pa ch isola ion had a nega i e e ec 19
on he p esence o o es specialis species (namely, L. galeobdolon, F angula alnus, 20
Hype icum and osaemum, A. adian um-nig um and A hy ium ilix- emina) and a o ed 21
he coloniza ion o mo e gene alis species such as Ci sium sp., Calluna ulga is, E ica 22
a bo ea o Ulex sp. Hence, in his egion a special a en ion should be gi en o he 23
conse a ion o o es specialis species, especially e ns and he bs. In a conse a ion 24
3
policy ocused on his o es specialis species, which a e he mos aluable species in 25
o es ed ecosys ems, la ge o es s should be p omo ed. 26
27
Keywo ds: Pa ch size / deg ee o isola ion / species ails / o es specialis species 28
4
1. In oduc ion 29
The excessi e des uc ion and agmen a ion o na u al and semi-na u al habi a s 30
on he Ea h’s su ace is ecognized as one o he p incipal causes o he loss o wild 31
biodi e si y (D’eon and Glenn, 2005; Fische and Lindenmaye , 2007; Haines-Young, 32
2009; Ha ison and B una, 1999; Hobbs, 2000; Me e and Ca oll, 1997; Wilcox and 33
Mu phy, 1985; Wood e al., 2000). The e ec s o habi a agmen a ion on biodi e si y 34
ha e been s udied o se e al decades, esul ing in a as li e a u e on his opic, and, 35
despi e con inued deba e abou he ela i e impo ance o habi a agmen a ion and 36
habi a loss (Fah ig, 2003; Hanski & Gaggio i, 2004), i is mos ly clea ha he size and 37
spa ial dis ibu ion o habi a emnan s al e s he pa e ns o species dis ibu ion and 38
abundance wi hin a landscape (Ewe s and Didham, 2006). 39
The p ocesses o educ ion, spa ial di ision and inc eased isola ion o habi a s 40
caused by agmen a ion a e associa ed wi h a educ ion in he abundance, dis ibu ion 41
and iabili y o species closely linked o hese habi a s (Bende e al., 2005; Fah ig and 42
Me iam, 1994; Kleye e al., 1996; Kup e e al., 2006). Howe e , no all plan species 43
show a uni o m esponse o habi a agmen a ion. Fo ins ance, a numbe o s udies 44
ha e shown ha he na u e o he species-a ea ela ionship desc ibing species loss om 45
habi a agmen s is con ounded by di e ences in species ai s (Cagnolo e al., 2006; 46
Ewe s and Didham, 2006; Gode oid and Koedan, 2003; Kolb and Diekmann, 2005). 47
Some s udies show ha habi a agmen a ion a ec ed plan s wi h speci ic dispe sal 48
modes (Kolb and Diekmann, 2005; Taba elli e al., 1999), low equency o occu ence 49
and high habi a speci ici y (Hill and Cu an, 2001; Iida and Nakashizuka, 1995). Plan 50
species wi h di e en g ow h- o ms (woody s. he baceous; sho -li ed s. long-li ed) 51
can p esen di e en esponses o agmen a ion. Woody plan s g ow mo e slowly and 52
de o e he la ge pa o hei pho osyn hesis o he p oduc ion o s uc u al ma e ials o 53
5
long- e m su i al (Chapin, 1991). Meanwhile, he he baceous plan s g ow and die 54
mo e apidly and de o e he la ge pa o hei pho osyn hesis o ep oduc ion and apid 55
u n o e . These cha ac e is ics can make he species espond di e en ly o 56
agmen a ion and, i hey a e a ec ed, ha e di e en esponse imes (Ewe s and 57
Didham, 2006). In ac , i has been pos ula ed ha sho -li ed species like he bs should 58
be mo e sensi i e o edge e ec s which would a ou colonisa ion by ude al species 59
(Cagnolo e al., 2006). In luence om su ounding ege a ion may ac ually inc ease he 60
o al species ichness o agmen ed woodlo s, bu educe he ac ion o habi a 61
specialis s (Ha ison, 1999).Thus, an assessmen o he e ec o agmen a ion on plan 62
communi ies should be based no only on species ichness bu also on species ype, 63
which can be de ined in e ms o conse a ion alue o ecological ai s (Honnay e al., 64
1999a; Hill and Cu an, 2001). 65
In he no h o he Ibe ian Peninsula he po en ial ege a ion is mixed-oak 66
o es s, domina ed by Que cus obu L. wi h F axinus excelsio L. and Cas anea sa i a 67
Mille (Onaindia e al., 2004). Howe e , since he beginning o he 20 h cen u y mos o 68
he po en ial a ea has been e o es ed by as g owing exo ic species, namely Pinus 69
adia a and Eucalyp us globulus, ha ha e mainly a ec ed o es specialis species 70
(Amezaga and Onaindia, 1997). The aim o his esea ch was o es whe he he spa ial 71
con igu a ion o hose o es s, namely size, o m and deg ee o isola ion o he pa ches, 72
a ec s in he same way he ege a ion as a whole o a ies o di e en g ow h- o ms 73
(he baceous, e ns, climbe s, sh ubs and ees) and o es specialis species (Aseginolaza 74
e al., 1988). 75
76
6
2. Me hods 77
2.1. S udy A ea 78
This s udy was ca ied ou in he U daibai Biosphe e Rese e (UBR) (a ea 220 79
km2) loca ed in he no h o he Ibe ian Peninsula (43º19´N, 02º40´W) (Figu e 1). The 80
UBR is one o he mos impo an na u al a eas o he Basque Coun y (No he n Spain) 81
due o, among o he ea u es, i s unique and di e se landscape which includes a c aggy 82
coun yside occupied by meadow land, oak g o es, deciduous woods and, especially, 83
pine plan a ions. 84
The po en ial ege a ion o he 80% o he UBR is mixed-oak o es s, domina ed 85
by Que cus obu L. wi h F axinus excelsio L. and Cas anea sa i a Mille (Onaindia 86
e al., 2004). Th oughou he 20 h cen u y, hese na i e mixed-oak o es s we e hea ily 87
agmen ed and, as a esul , oday hey co e only abou 6% o he o al a ea o he 88
U daibai Rese e (Rod íguez-Loinaz e al., 2011) as has happened wi h o he na u al 89
o es s in o he pa s o he wo d (Schessl e al., 2008). A e wa ds, he adi ional use o 90
imbe and coal p oduc ion was abandoned and he emaining o es pa ches s a ed a 91
p ocess o egene a ion (Michel, 2006). 92
93
2.2. Pa ch selec ion and ege a ion sampling 94
A o al o 33 pa ches o mixed-oak o es si ua ed in he UBR we e selec ed by 95
means o he land use map a a 1:10 000 scale (Figu e 2). The selec ion was made as a 96
unc ion o size, since a p incipal objec i e was o es ablish i he di e si y o he 97
ascula plan species was a ec ed by he size o he pa ch. The e o e, 18 pa ches o a 98
size be ween wo and h ee hec a es and 15 pa ches o a size be ween en and hi y 99
hec a es we e selec ed. The e was no di e ence on al i ude, slope, soil ype o 100
geog aphical loca ion be ween small and la ge pa ches (small pa ches: mean al i ude: 101
7
133±18.89 m, slope: 25±2.10 %, pH: 4.65±0.10, UTM_X: 525762±687, UTM_Y: 102
4.7984 106±1534 and la ge: mean al i ude: 174±17.36 m, slope: 30±2.61 %, pH: 103
4.76±0.11, UTM_X: 526363±962, UTM_Y: 4.7965 106±1693). This selec ion was 104
de e mined a e analysis o he dis ibu ion o pa ch sizes gi en ha hese we e he only 105
sizes ha occu ed in signi ican numbe s. The ollowing indices we e de e mined o 106
each pa ch: a ea, dis ance o he nea es pa ch o mixed-oak o es s (edge o edge) 107
(NND, measu e o he deg ee o isola ion) and he ac al dimension (FD, measu e o 108
he o m) (Mc Ga igal e al., 2002), o which he -LATE so wa e was used (Lang and 109
Tiede, 2003). 110
Since sampling e o and numbe o species eco ded a e usually ela ed 111
(Magu an, 1988; Hill e al., 1994; Lomolino, 2001), he a ea sampled was kep 112
cons an in all si es in o de o a oid sampling a e ac s on he e ec s o habi a 113
agmen a ion (Hill e al., 1994). In each o he pa ches (la ge and small) one plo o 114
25m x 25m was de e mined app oxima ely in he cen e o each pa ch in o de o 115
minimise possible edge e ec s. Wi hin each plo , i e sub-plo s o 2x1m we e 116
delinea ed. One was in he cen e and he o he ou sepa a ed by 12m, making a c oss 117
wi h an a m unning wi h he slope and he o he pe pendicula o i . The numbe o 118
sub-plo s was de e mined acco ding o he me hod o he species/a ea cu e (Ken and 119
Coke , 1992). In hese sub-plo s he pa e n o ege a ion du ing June and July 2005 120
was s udied. In each sub-plo , plan species we e iden i ied and he pe cen age co e o 121
each plan species, calcula ed h ough isual es ima ion, was de e mined. In o de o 122
de e mine pe cen age co e , i e di e en s a a (le els) we e conside ed, i.e. 0-0.20, 123
0.20-1, 1-3, 3-7, >7 m, ollowing B owe and Za (1977) and Onaindia e al. (2004). 124
Thus, he i s s a um co esponded o he baceous plan s, he second o lowe sh ub-125
like plan s, he hi d o highe sh ub-like plan s, he o h o he lowe ee canopy and, 126
8
inally, he i h o he highe ee canopy. The o al pe cen age co e o each plan 127
species was ob ained by adding up i s pe cen age co e in each o he i e di e en 128
s a a. In addi ion, he co e o ees as an indi ec measu e o quan i y o ligh was 129
measu ed, as ligh condi ion is one o he main ac o s in o es habi a s (Sa lö -He lin 130
and F y, 2000) and i is known o a ec ege a ion (Amezaga e al., 2006; Bo chsenius 131
e al., 2004). 132
Summing he co e in he i e sub-plo s, he o al co e o each species in he 133
sampled a ea was ob ained. Using hese da a he indices o ichness (S) and Shannon 134
(H´) and Simpson (1-D) di e si y we e calcula ed. These indices we e ob ained o he 135
o e all ege a ion, he di e en g ow h- o ms p esen (he baceous, e ns, climbe s, 136
sh ubs and ees), he o e all o es specialis species and inally o he di e en 137
g ow h- o ms wi hin he o es specialis species. To classi y a species as o es 138
specialis he “Illus a ed keys o he lo a o he Basque Coun y and bo de ing 139
e i o ies” (Aseginolaza e al., 1988) was used. In his book he na u al habi a o e e y 140
species is desc ibed. All hose species whose na u al habi a was desc ibed as nemo al 141
o es , beech o es , oak o es o humid and shaded si es in o es , we e classi ied as 142
o es specialis species. 143
Besides, he o e all ege a ion simila i y in ela ion o pa ch size and dis ance o 144
he nea es missed-oak o es pa ch was calcula ed using he So ensen´s communi y 145
simila i y index. As he dis ance o he nea es pa ch was a con inuous a iable, he 146
compa ison was pe o med among he i e pa ches wi h he smalles (<50 m) NND and 147
he i e pa ches wi h he la ges (> 200 m) NND. 148
149
9
2.3. S a is ical analysis 150
As pa ch indices (pa ch size, pa ch isola ion, ac al dimension) we e no 151
co ela ed (Spea man ank co ela ion, P>0.05), a Gene al Linea Model (GLM) was 152
pe o med o analyze he e ec s o agmen a ion on he ichness and di e si y o he 153
ege a ion. In his model he size (la ge o small) was in oduced as a ac o and he 154
ac al dimension (FD), deg ee o isola ion (NND) and co e o ees we e in oduced as 155
co- a ian s. 156
Ha ing analyzed he e ec s o agmen a ion on ichness and di e si y, he 157
e ec o size and isola ion (dis ance o he nea es pa ch o mixed-oak o es ) on o e all 158
species composi ion was es ed by means o he semi-pa ame ic pe mu a ional 159
mul i a ia e analyses o a iance (he ea e PERMANOVA) de eloped by Ande son 160
(2001). Indica o Species Analysis (ISA; Du ene and Legend e, 1997) was used o 161
de e mine he cha ac e is ic species wi hin pa ch size. Only species wi h P<0.05 we e 162
conside ed (assessed using Mon e Ca lo andomiza ions wi h 999 pe mu a ions and 163
INDVAL> 25). 164
165
3. Resul s 166
3.1. Vege a ion s uc u e 167
A o al o 110 plan species o which 53 (27 o es specialis ) we e he baceous, 5 168
(4 o es specialis ) climbe s, 18 (6 o es specialis ) ees, 23 (7 o es specialis ) sh ubs 169
and 11 (7 o es specialis ) e ns we e ound in his s udy (Table 1). O hese 110 170
species, 84 we e ound in he la ge pa ches and 90 in he small ones. 171
The ege a ion simila i y esul s showed ha 78% o he species we e he same 172
o he la ge and small pa ches and 50% o he pa ches wi h he smalles and la ges 173
NND. Those species only p esen in he la ge pa ches we e usually (80 %) o es 174
16
6. Acknowledgmen s 316
We g a e ully acknowledge inancial suppo om he Spanish Minis y o 317
Educa ion and Science (CGL2005-08046-C03 01 and CGL2008-05579-C02-01), he 318
Depa men o Uni e si ies, Resea ch and Educa ion o he Basque Go e nmen 319
(G oups bu sa ), UNESCO Chai o he Uni e si y o he Basque Coun y and Basque 320
Go e nmen (Indus y Depa men -E o ek P ojec ). We would also like o hank Si 321
Ma ack Goulding o his con ibu ion o he edi ing o his documen .322
17
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21
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Figu e 1: Loca ion o he s udy a ea. 483
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Figu e 2: Map o land uses and localiza ion o s udied mixed-oak o es s in he s udy a ea. 486
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