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Pantropical modelling of canopy functional traits using Sentinel-2 remote sensing data Remote Sensing of Environment

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Tropical forest ecosystems are undergoing rapid transformation as a result of changing environmental conditions and direct human impacts. However, we cannot adequately understand, monitor or simulate tropical ecosystem responses to environ ⁎ mental changes without capturing the high diversity of plant functional characteristics in the species-rich tropics. Failure to do so can oversimplify our understanding of ecosystems responses to environmental disturbances. Innovative methods and data products are needed to track changes in functional trait composition in tropical forest ecosystems through time and space. This study aimed to track key functional traits by coupling Sentinel-2 derived variables with a unique data set of precisely located in-situ measurements of canopy functional traits collected from 2434 individual trees across the tropics using a standardised methodology. The functional traits and vegetation censuses were collected from 47 field plots in the countries of Australia, Brazil, Peru, Gabon, Ghana, and Malaysia, which span the four tropical continents. The spatial positions of individual trees above 10 cm diameter at breast height (DBH) were mapped and their canopy size and shape recorded. Using geo-located tree canopy size and shape data, community-level trait values were estimated at the same spatial resolution as Sentinel-2 imagery (i.e. 10 m pixels). We then used the Geographic Random Forest (GRF) to model and predict functional traits across our plots. We demonstrate that key plant functional traits can be measured at a pantrop-ical scale using the high spatial and spectral resolution of Sentinel-2 imagery in conjunction with climatic and soil information. Image textural parameters were found to be key components of remote sensing information for predicting functional traits across tropical forests and woody savannas. Leaf thickness (R 2 = 0.52) obtained the highest prediction accuracy among the morphological and structural traits and leaf carbon content (R 2 = 0.70) and maximum rates of photosynthesis (R 2 = 0.67) obtained the highest prediction accuracy for leaf chemistry and photosynthesis related traits, respectively. Overall, the highest prediction accuracy was obtained for leaf chemistry and photosynthetic traits in comparison to morphological and structural traits. Our approach offers new opportunities for mapping, monitoring and understanding biodiversity and ecosystem change in the most species-rich ecosystems on Earth.
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;DL?HEDC;DJI FBKI J>; KI; E< J>; .;DJ?D;B :7J7 M?BB 7BBEM KI JE 79
9KH7J;BO FH;:?9J FB7DJ <KD9J?ED7B JH7?J :?IJH?8KJ?EDI J>7J 7H; FEJ;DJ?7BBO
=;D;H7B?I78B; 79HEII J>; JHEF?97B <EH;IJ 8?EC;
2. Methods
 *353A/A7<; =9<A@
2; 9EBB;9J;: L;=;J7J?ED 9;DIKI :7J7 <HEC  F;HC7D;DJ L;=;J7J?ED
FBEJI J>7J 7H; F7HJ E< J>; "BE87B 9EIOIJ;CI (ED?JEH?D= D;JMEHA " (
MMM=;CJHEF?97B<EH;IJIEN79KA />;I; FBEJI ;D9ECF7II M;J JHEF?97B
<EH;IJI I;7IED7BBO :HO JHEF?97B <EH;IJI 7D: JHEF?97B <EH;IJI7L7DD7 JH7D
I?J?ED7B L;=;J7J?ED />; I7CFB;: L;=;J7J?ED FBEJI >7L; 7D 7H;7 H7D=?D=
<HEC  JE  >7 M?J> CEIJ  8;?D=  >7 />; FBEJI KI;: 7H; BE97J;:
79HEII <EKH JHEF?97B 9EDJ?D;DJI 7D: IF;9?<?97BBO ?D J>; 9EKDJH?;I E< KI
JH7B?7 H7P?B "78ED ">7D7 (7B7OI?7D EHD;E <HEC >;H; EDM7H:I H;
<;HH;: JE 7I (7B7OI?7 7D: +;HK /78B;  $D ;79> FBEJ 7BB MEE:O FB7DJ
?D:?L?:K7BI M?J> 7 :?7C;J;H  9C 7J 8H;7IJ >;?=>J # EH 78EL;
8KJJH;II HEEJI M;H; C;7IKH;: 7D: J>;?H ;N79J =;E=H7F>?9 BE97J?ED M7I
H;9EH:;: I;; J>; $D:?L?:K7B JH;; 9HEMDII;9J?ED 8;BEM <EH CEH; :;
J7?BI $D JME FBEJI )31 7D: )31 ?D )EL7 37L7DJ?D7 >;H; ED
M7H:I H;<;HH;: JE 7I H7P?B)3 J>; # M7I C;7IKH;: D;7H =HEKD: B;L;B
7I ?I IJ7D:7H: ?D I7L7DD7 CED?JEH?D= FHEJE9EBI
 B;1A7<;/9 A?/7A@
2; 9EBB;9J;: FB7DJ <KD9J?ED7B JH7?J C;7IKH;C;DJI <HEC 7BB MEE:O
FB7DJI BE97J;: ?D ;79> E< J>;  L;=;J7J?ED FBEJI C;DJ?ED;: 78EL;
/78B;  BB JH7?JI
M;H; =7J>;H;: <HEC J>; " ( D;JMEHA 7D: M;H; 9EBB;9J;: <EBBEM?D= 7
IJ7D:7H:?I;: C;J>E:EBE=O 79HEII FBEJI !EH;IJ ?DL;DJEHO :7J7 M;H; KI;:
JE IJH7J?<O JH;; IF;9?;I 8O 87I7B 7H;7 :EC?D7D9; 7 FHENO <EH 97DEFO 7H;7
:EC?D7D9; />; JH;; IF;9?;I J>7J 9EDJH?8KJ;: CEIJ JE 87I7B 7H;7 78KD
:7D9; M;H; I7CFB;: M?J>  H;FB?97J; ?D:?L?:K7BI F;H IF;9?;I M?J>
7 =E7B E< I7CFB?D=  E< 87I7B 7H;7 79HEII J>; I7CFB?D= H;=?ED
?=>JO F;H9;DJ E< 87I7B 7H;7 M7I E<J;D 79>?;L;: ?D BEM :?L;HI?JO I?J;I
;= CEDJ7D; EH :HO <EH;IJI 8KJ EDBO 7HEKD:  M7I 79>?;L;: ?D IEC;
>?=> :?L;HI?JO I?J;I BEMB7D: >KC?: H7?D<EH;IJI !EH ;79> I;B;9J;: JH;;
7 IKD 7D: 7 I>7:; 8H7D9> M;H; I7CFB;: 7D: ?D ;79> 8H7D9>  B;7L;I
M;H; KI;: <EH JH7?J C;7IKH;C;DJI 2; EDBO ?D9BK:;: J>; IKD ;NFEI;:
8H7D9>;I ?D EKH 7D7BOI?I 8;97KI; M; M;H; ?DJ;H;IJ;: ?D J>; 8H7D9>;I
J>7J 9EKB: FEJ;DJ?7BBO 8; H;9;?L?D= :?H;9J IKD H7:?7J?ED 7D: J>KI I>EM
:?H;9J IF;9JH7B H;a;9J7D9; />?I H;FH;I;DJ;: 7 JEJ7B I7CFB; E<  ?D
:?L?:K7B JH;;I 79HEII J>; JHEF?9I /78B;  />; FB7DJ <KD9J?ED7B JH7?JI
9EBB;9J;: M;H; J>EI; H;B7J;: JE F>EJEIODJ>;J?9 97F79?JO 7J 8EJ> I7JKH7J
?D= *9ED9;DJH7J?ED  FFC * C7N 7D: 7C8?;DJ *9ED9;D
JH7J?ED  FFC * I7J B;7< 9>;C?IJHO D?JHE=;D F>EIF>EHKI 97H
8ED 97B9?KC FEJ7II?KC 7D: C7=D;I?KC 9EDJ;DJ 7D: B;7< CEHF>EBE=
?97B 7D: IJHK9JKH7B JH7?JI 7H;7 IF;9?`9 B;7< 7H;7 J>?9AD;II :HO C7II
<H;I> C7II 7D: M7J;H 9EDJ;DJ D EL;HL?;M E< J>; C;J>E:I <EH ?D:?L?:
K7B B;7< <KD9J?ED7B JH7?J C;7IKH;C;DJI ?I FHEL?:;: ?D J>; .KFFB;C;DJ7HO
$D<EHC7J?ED I;; <KBB JH7?JI 9EBB;9J?ED FHEJE9EB I;9J?ED !KHJ>;H :;J7?BI
E< C;7IKH;C;DJI <EH J>; +;HKL?7D D:;I 97CF7?=D 7H; =?L;D ?D (7HJ?D
;J 7B  7D: DGK?IJ ;J 7B  <EH J>; (7B7OI?7D 97CF7?=D ?D
EJ> ;J 7B  7D: <EH J>; ">7D7 7D: H7P?B 97CF7?=DI ?D *B?L;H7I
;J 7B  7D: "LEP:;L7?J; ;J 7B 
.EC; ?D:?L?:K7BI ?D J>; FBEJI B79A;: <KD9J?ED7B JH7?J L7BK;I /E 7II?=D
H;FH;I;DJ7J?L; JH7?J L7BK;I JE KDI7CFB;: ?D:?L?:K7BI M; :?: J>; <EBBEM
?D=  ?D:?L?:K7BI <HEC M>?9> JH7?JI M;H; C;7IKH;: A;FJ J>;?H EH?=?D7B
JH7?J ?D<EHC7J?ED  <EH ?D:?L?:K7BI M?J> DE JH7?J ?D<EHC7J?ED M; H7D
:ECBO I7CFB;: JH7?J L7BK;I <HEC EJ>;H ?D:?L?:K7BI <HEC J>; I7C; IF;9?;I
FH;I;DJ ?D J>; I7C; FBEJ  ?< J>; IF;9?;I M7I DEJ I7CFB;: ?D J>; =?L;D
FBEJ J>;D M; H7D:ECBO I7CFB;: 7D ?D:?L?:K7B <HEC J>; I7C; IF;9?;I J>7J
>7: JH7?J ?D<EHC7J?ED ?D EJ>;H FBEJI <HEC J>; I7C; H;=?ED /78B;  />?I
FHEJE9EB <EH JH7?J L7BK; 7BBE97J?ED 7BBEM;: KI JE MEHA M?J> J>; ;N?IJ?D=
H7D=; E< JH7?J L7BK;I 7J J>; IF;9?;I B;L;B 7D: 7LE?:;: JE 9H;7J; 7L;H7=;
L7BK;I F;H IF;9?;I 7:EJJ; ;J 7B  .9>D;?:;H ;J 7B  2; :?:
DEJ 7II?=D JH7?J L7BK;I JE J>; H;C7?D?D= ?D:?L?:K7BI 8;BED=?D= JE IF;9?;I
<HEC M>?9> DE JH7?J 9EBB;9J?ED M7I E8J7?D;: 7J J>; H;=?ED7B B;L;B
 ;27C72B/9 A?33 1?<D;@
/H;; 9HEMD BE97J?EDI 7D: IJHK9JKH7B 7JJH?8KJ;I M;H; H;9EH:;: <EH
;79> JH;; M>;H; 9HEMD 7H;7 7D: I>7F; M;H; C;7IKH;: 8O :?H;9J 9HEMD
`;B: C;7IKH;C;DJI ?D J>; 97I; E< FBEJI ?D (7B7OI?7 7D: +;HK I;; FHE
JE9EB 8;BEM EH 8O C;7DI E< H;=?ED7B B;L;B 7BBEC;JH?9 ;GK7J?EDI :;L;B
EF;: 8O .>;DA?D ;J 7B  7BB EJ>;H FBEJI $D J>; B7JJ;H 97I; J>;
9HEMDI I>7F; M7I 7IIKC;: JE 8; 9?H9KB7H />; :?H;9J `;B: 9HEMD C;7
IKH;C;DJI M;H; 7I <EBBEMI 7BB JH;;I  9C # ?;  C <HEC J>;
=HEKD: M;H; C7FF;: KI?D= 7 =HEKD:87I;: !?;B:(7F B7I;H J;9>DEBE=O
$! - 'J: %VBELT K +H7>O P;9> -;FK8B?9 #T:B ;J 7B  />;
!?;B:(7F J;9>DEBE=O M7I 87I;: ED 7 9EC8?D7J?ED E< $CFKBI;  .J7D
:7H: B7I;H H7D=;`D:;H M?J> ?D8K?BJ J?BJ I;DIEH <EH C;7IKH?D= L;HJ?97B
7D=B;I (7F.J7H CE:KB; $$ ;B;9JHED?9 9ECF7II 8EJ> '7I;H /;9>DEBE=O
$D9 EBEH7:E 0. 7D: J>; IF;9?7B?P;: C7FF?D= IE<JM7H; !?;B:(7F
L  $! - P;9> -;FK8B?9 />; J;9>DEBE=O M7I KI;: JE H;9EH: IF7
J?7B FEI?J?EDI E< JH;; IJ;CI ?D J>H;;:?C;DI?ED7B IF79; N O P9EEH:?
D7J;I 7I M;BB 7I JE C7F ?D:?L?:K7B >EH?PEDJ7B FHE@;9J?EDI E< JH;; 9HEMDI
?D J>; FBEJI />; >EH?PEDJ7B 9HEMD FHE@;9J?ED E< ;L;HO JH;; M7I E8
J7?D;: 8O C;7IKH?D= IF7J?7B FEI?J?EDI N 7D: O9EEH:?D7J;I E< I;H?;I
E< FE?DJI H7D=?D= <HEC  JE  FE?DJI :;F;D:?D= ED J>; I?P; E< J>;
9HEMD 7J J>; 8EKD:7HO E< 7 9HEMD FHE@;9J;: JE J>; >EH?PEDJ7B FB7D;
/>; I>7F; E< 9HEMD FHE@;9J?ED M7I IK8I;GK;DJBO ICEEJ>;: KI?D= J>;
ICEEJ> 9EDJEKH B?D;<;7JKH; E< !?;B:(7F IE<JM7H; L  #;?=>JI E<
7BB JH;;I M?J> #  9C M;H; C;7IKH;: 8O J>; $CFKBI; 7D: /HK+KBI;
 - B7I;H H7D=;`D:;HI 8EJ> '7I;H /;9>DEBE=O $D9 EBEH7:E 0.
/>KI ;79> ?D:?L?:K7B 9HEMD M7I 799KH7J;BO =;EBE97J;: H;D:;H?D= ?D<EH
C7J?ED 78EKJ ?JI I>7F; 7D: L;HJ?97B 7D: >EH?PEDJ7B FEI?J?ED
UNCORRECTED PROOF
5B7??3BA7L??3G3A/9 &3:<A3'3;@7;5<4;C7?<;:3;A EEE EEEE EEEEEE
Table 1
EBB;9J?ED :;J7?BI <EH L;=;J7J?ED FBEJI 7D: FB7DJ <KD9J?ED7B JH7?JI  JEJ7B E<  ?D:?L?:K7B JH;;I M;H; I7CFB;: <EH <KD9J?ED7B JH7?JI
'E97J?ED .F;9?;II7CFB;:<EHJH7?JI +BEJ9E:; .?P;>7 ;DJHE?:9EEH:?D7J;I 7J;E<9EBB;9J?ED
3 4 1;=;J7J?ED9;DIKI /H7?JI
KIJH7B?7   +     %KD;.;FJ;C8;H
 +   
-*   
-*  
">7D7  )&     *9JE8;H(7H9>=H7CC
)&   
*    
*   
"78ED  '+"    !;8HK7HO(7H9>
'+"  
()"   
H7P?B)3  )31    (7H9>(7O
)31   
1-   
)31    
)31   
)31   
)31   
)31   
)31   
)31   
)31   
)31   
)31   
H7P?B./       K=KIJ.;FJ;C8;H
   
   
   
   
+;HK  .+    FH?B)EL;C8;H
+)   
.+   
.+   
/-0   
24   
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.!  
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2; 97B9KB7J;: J>; 9ECCKD?JO M;?=>J;: C;7D E< ;79> JH7?J <EH ;79>
 e  C IK8FBEJ C7J9>?D= J>; >?=>;IJ F?N;B IF7J?7B H;IEBKJ?ED E< J>;
.;DJ?D;B ?C7=;HO 87I;: ED J>; C7II H7J?E >OFEJ>;I?I M>?9> IJ7J;I
J>7J J>; CEIJ :EC?D7DJ IF;9?;I :H?L; J>; ;9EIOIJ;C FHE9;II;I 8O C;7DI
E< J>;?H <KD9J?ED7B JH7?JI "H?C;  2; `HIJ =;EBE97J;: J>; L;=;
J7J?ED FBEJ M?J> ?JI 7BH;7:O C7FF;: JH;; 9HEMDI I;; FHEJE9EB 78EL;
JE J>; .;DJ?D;B ?C7=;HO 87I;: ED J>; 9EHD;H 9EEH:?D7J;I E< J>; FBEJI
/>?I ?I 7D ?CFEHJ7DJ IJ;F 7I =;EBE97J?ED ;HHEHI 8;JM;;D J>; L;=;J7J?ED
FBEJ 7D: J>; 9EHH;9J BE97J?ED ?D J>; I7J;BB?J; ?C7=; 9EKB: H;FH;I;DJ 7
B7H=; FHEFEHJ?ED E< 7 =?L;D FBEJ :;F;D:?D= ED J>; FBEJI 7H;7 />;D <EH
;79> E< J>; JH7?JI A 7D: F?N;BI = M; 97B9KB7J;: J>;?H 9ECCKD?JO B;L;B
M;?=>J;: C;7D 2( KI?D= J>; ?D:?L?:K7B JH;; 9HEMD >EH?PEDJ7B 7H;7
7I J>; M;?=>J?D= <79JEH !?=  7I <EBBEMI
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L7BK; E< ?D:?L?:K7B 7?D F?N;B ="?I J>; JEJ7B DKC8;H E< ?D:?L?:K7BI F;H
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2( M7I M;?=>J;: 8O ?JI FHEFEHJ?ED7B 9EL;H E< J>; 9EHH;IFED:?D= F?N;B
/>; JEJ7B DKC8;H E< F?N;BI KI;: ?D EKH 97B9KB7J?EDI 7H;  <EH KIJH7B?7
 <EH H7P?B )3 )EL7 37L7DJ?D7  <EH H7P?B ./ .7DJ7H;C 
<EH "78ED  <EH ">7D7  <EH (7B7OI?7 7D:  <EH +;HK
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2; =7J>;H;: .;DJ?D;B ?C7=;HO J>7J M7I 9BEI;IJ ?D J?C; 7D: I;7IED
JE J>; I7CFB?D= :7J;I E< <KD9J?ED7B JH7?JI 7D: L;=;J7J?ED 9;DIKI 79HEII
J>; JHEF?9I <EH ;79> E< J>; IJK:O BE97J?EDI /78B; . />; .;DJ?D;B
?C7=;HO M7I `HIJ I;B;9J;: KI?D= J>; KHEF;7D .F79; =;D9O  . .9?
;D9;#K8 9>EEI?D= ?C7=;I M?J> >?=> F?N;B GK7B?JO 7D: BEM 9BEK: 9EL;H
 JCEIF>;H?9 H7:?EC;JH?9 7D: JEFE=H7F>?9 9EHH;9J?EDI M;H;
7FFB?;: JE J>; I;B;9J;: ?C7=;HO ';L;B  KI?D= J>; .;DEH 7B=E
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7=;HO M?J> J>; L;=;J7J?ED FBEJI 7FF;7H;: <H;; E< 9BEK:I 7D: 9?HHKI ;<
<;9JI />; 78EL;C;DJ?ED;: IJ;FI 7BBEM;: KI JE E8J7?D B;L;B  ?C7=;HO
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Table 2
;I9H?FJ?ED E< FB7DJ <KD9J?ED7B JH7?JI 9EBB;9J;: 79HEII J>; JHEF?9I 7D: J>;?H H;B;L7D9; KD:;H 7 9>7D=?D= ;DL?HEDC;DJ
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B;7<
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97FJKH;7D:F>EJEIODJ>;J?979J?L?JO
27BA;H;J7B 2H?=>J;J7B
 %KD;7K7D:/7H7IEc V7P
;J7B #7MJ>EHD; >7L;;J
7B #K7D=;J7B
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M7J;HBEIID;J7II?C?B7J?EDH7J;B;7<B?<;IF7D
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97FJKH;H;B7J;:JEF>EJEIODJ>;J?997F79?JO7D:=HEMJ>
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C7OL7HO:;F;D:?D=ED;DL?HEDC;DJ7B9ED:?J?EDI7D:
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H;IEBKJ?ED 87D:I    M;H; DEJ KI;: 7I J>;I; 7H; :;I?=D;:
<EH 9?HHKI M7J;H L7FEKH 7D: 9BEK: :;J;9J?ED /78B;  7D:  M7I DEJ
KI;: 7I ?J 9EL;HI 7D EL;HB7FF?D= IF;9JH7B M?D:EM M?J> 87D:  7D: >7I
7 BEM;H IF7J?7B H;IEBKJ?ED .?D9; L;=;J7J?ED ?D:?9;I C7O ?D9H;7I; FH;:?9
J?ED 799KH79O M>;D CE:;BB?D= 9ECCKD?JO M;?=>J;: JH7?JI 27BB?I ;J 7B
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DKJH?;DJ B;L;BI ?D J>; B;7L;I 7D: J>;?H KI; E< J>; L?I?8B;JEH;: ;:=; IF;9
JH7B 87D:I
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<;H;DJ L;=;J7J?ED JOF;I 7D: :?<<;H;D9;I ?D 97DEFO IF;9JH7B 9ECFEI?
J?ED /E 9>7H79J;H?I; 97DEFO IJHK9JKH; M; 97B9KB7J;: J>; "H;O ';L;B
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C7JH?N J>7J ?I IF7J?7BBO :;F;D:;DJ />; 9EE99KHH;D9; C7JH?N :;F;D:I
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F?N;BI 7D: :;F?9JI J>; DKC8;H E< E99KHH;D9;I E< J>; H;B7J?EDI>?F 8;
JM;;D 7 F?N;B 7D: ?JI D;?=>8EKH <J;H JH?7BI M?J> IC7BB;H M?D:EMI
I?P;  e  M; EFJ;: JE KI; 7  e  F?N;B A;HD;B M?D:EM M>?9> M7I
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CE:;BB?D= IJ;F I;; I;9J?ED  8;BEM M?J>EKJ J7A?D= B7H=; F;H?E:I
E< J?C; <EH ?JI 97B9KB7J?ED />; J;NJKH; H;IKBJI E8J7?D;: M?J> J>; KI;:
A;HD;B M?D:EM M7I >?=>BO 9EHH;B7J;: JE J>; IC7BB;H A;HD;B M?D:EM
<?   $ 7I;: ED J>; "'( M; 97B
9KB7J;: JME L7H?78B;I J>7J 7H; B;7IJ 9EHH;B7J;: M?J> ;79> EJ>;H J>; D
JHEFO 7D: EHH;B7J?ED <EH ;79> E< J>; L;=;J7J?ED ?D:?9;I 2>?B; DJHEFO
C;7IKH;I J>; >ECE=;D;?JO B;L;B <EH 7 =?L;D 7H;7 J>; EHH;B7J?ED C;7
IKH;I FHE878?B?JO E< E99KHH;D9; E< J>; IF;9?`;: F?N;B F7?HI 79HEII J>; ?C
7=; #7H7B?9A ;J 7B  27BB?I ;J 7B  BB H;CEJ; I;DI?D= 7D7BO
I;I H;B7J;: JE J>; =;D;H7J?ED E< L;=;J7J?ED ?D:?9;I 7D: J;NJKH; C;JH?9I
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=?EDI 7D: 9EKB: 7J B;7IJ F7HJBO :;J;HC?D; J>; H;=?EDI L;=;J7J?ED 7D:
?DJH?DI?9 JH7?J 9ECFEI?J?ED 2; E8J7?D;: ?D<EHC7J?ED ED J>;I; J>H;;
9ECFED;DJI <EH ;79> I7CFB?D= BE97J?ED />; J>H;; 9ECFED;DJI M;H;
=HEKF;: 7I 8;BED=?D= JE ;DL?HEDC;DJ7B 9B?C7J; EH IE?BJ;HH7?D J;NJKH;
F# 97J?ED ;N9>7D=; 97F79?JO 7D: JEFE=H7F>O :H?L;HI /78B; 
!EH 9B?C7J; 7D: <EH ;79> I7CFB?D= BE97J?ED M; =7J>;H;: =H?::;:
:7J7 ED J>; C;7D 7DDK7B 9B?C7J?9 M7J;H :;`9?J (2 M>?9> ?I 7
C;JH?9 E< :HEK=>J ?DJ;DI?JO 7D: I;L;H?JO C;7D 7DDK7B C7N?CKC J;C
F;H7JKH; (/C7N IEB7H H7:?7J?ED .- 7D: IE?B CE?IJKH; .(
/78B;  BB 9B?C7J?9 :7J7 M?J> 7 IF7J?7B H;IEBKJ?ED E< Q AC M;H;
E8J7?D;: <HEC J>; /;HH7B?C7J; =H?::;: 9B?C7J; FHE:K9J 87JPE=BEK
;J 7B  /E 9>7H79J;H?I; J>; 9B?C7J?9 9ED:?J?EDI <EH
UNCORRECTED PROOF
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Fig. 1. ?7=H7C IKCC7H?I?D= J>; IJ;FI <EBBEM;: JE 7II?=D JH7?J L7BK;I F;H .;DJ?D;B F?N;B  !?HIJ J>; L;=;J7J?ED FBEJI 7H; :;`D;: 87I;: ED J>; " ( "BE87B DL?HEDC;DJ7B (ED?JEH?D=
:7J7I;J 7D:  <HEC ;79> L;=;J7J?ED FBEJ J>; 9EHD;H 9EEH:?D7J;I 7H; ;NJH79J;:  !HEC ;79> L;=;J7J?ED FBEJ J>; 34 FEI?J?ED E< ;79> IJ;C  9C # ?I ;NJH79J;: 7D:  J>; 9HEMD
>EH?PEDJ7B 7H;7 ?I 97B9KB7J;: 87I;: ED J>; FHEJE9EB :;I9H?8;: ?D J>; C;J>E:I I;9J?ED  />;D J>; .;DJ?D;B ?C7=;HO <EH J>; IJK:O 7H;7 ?I FHE9;II;: JE B;L;B  KI?D= J>; . .)+ JEEB8EN
7D:  J>; L;=;J7J?ED FBEJ ?I EL;HB7?: ?D J>; .;DJ?D;B ?C7=; 87I;: ED ?JI 9EHD;H 9EEH:?D7J;I $D J>?I B7IJ IJ;F  ;79> F?N;B :;`D;I 7 IK8FBEJM>?9> ?I J>; KD?J KI;: JE 97B9KB7J; J>; JH7?J
9ECCKD?JO M;?=>J;: C;7D 87I;: ED J>; 9HEMD 7H;7 E< J>; JH;;I J>7J 7H; 9EDJ7?D;: 8O J>7J F?N;B $D  ;H;<;HI JE 7 =?L;D JH;; ?D 7 =?L;D F?N;B JH7?J 7H;FH;I;DJI 7 =?L;D JH7?J 7D: E7D: F
7H; L7BK;I <EH J>7J JH7?J />; ?C7=; KI;: 7I 7D ;N7CFB; ?D IJ;F  M7I J7A;D 8O %;IKI =K?HH;"KJ?;HH;P EL;H 7 L;=;J7J?ED FBEJ KI?D= 7 CKBJ?IF;9JH7B '/0( 97C;H7 CEKDJ;: ED 7D $DIF?H;
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=;IJ;: 8O J>; 2EHB: (;J;EHEBE=?97B *H=7D?P7J?ED 2(* MMMMCE
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79J;H?I; J>; FBEJI JEFE=H7F>O 7I ?J >7I 8;;D I>EMD J>7J JEFE=H7F>O
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7L7?B78?B?JO /;HH7?D IBEF; M7I 97B9KB7J;: KI?D= 7 >?=>H;IEBKJ?ED :?=
?J7B ;B;L7J?ED CE:;B Q C F?N;B I?P; 7J J>; ;GK7JEH <HEC J>; .>KJ
JB; /EFE=H7F>O (?II?ED !7HH ;J 7B
UNCORRECTED PROOF
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Table 3
.F;9JH7B H;CEJ; I;DI?D= ;DL?HEDC;DJ7B 7D: IE?B H;B7J;: L7H?78B;I KI;: :KH?D= J>; CE:
;BB?D= FHEJE9EB BB 9B?C7J?9 L7H?78B;I 8KJ IBEF; M;H; 97B9KB7J;: KI?D= 7 9B?C7JEBE=O E<
 O;7HI  BB IE?B L7H?78B;I M;H; 97B9KB7J;: <EH J>; JEF 9C IE?B B7O;H .;D
J?D;B 87D: M7L;B;D=J>I DC 7H; =?L;D ?D F7H;DJ>;I?I 7<J;H J>; 87D: D7C;
/OF; 17H?78B; ;I9H?FJ?ED -;<;H;D9;I
-. 
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M?J>IF7J?7B
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
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7DDK7B
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J>;"BE87B DL?HEDC;DJ7B
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:7J787I;
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.7D: CEKDJE<I7D:
M;?=>J
.BEF; /;HH7?DIBEF;
 CH;IEBKJ?ED
!7HH;J7B
 J CEIJ I?J;I IE?B :7J7 M;H; I7CFB;: BE97BBO 7D: 7D7BOI;: JE 7
IJ7D:7H:?I;: FHEJE9EB ?D B78I ?D ;?J>;H $)+ (7D7KI H7P?B EH J>; 0D?
L;HI?JO E< ';;:I 0& <EBBEM?D= J>; -$)!*- IE?B FHEJE9EB ,K;I7:7 ;J
7B  !HEC J>;I; :7J7 M; IKCC7H?I;: FBEJ B;L;B IE?B :7J7 7L;H7=;:
EL;H J>; `HIJ  9C <EH J;NJKH; .7D: 7D: B7O 97J?ED ;N9>7D=; 97
F79?JO ;  7D: F##* F# +BEJ B;L;B J;NJKH; :7J7 M;H; DEJ 7L7?B
78B; <EH FBEJI ?D KIJH7B?7 7D: J>; )31 FBEJI 7D: M;H; J>KI :;H?L;:
<HEC J>; .E?B"H?:I :7J7I;J 7J  C F?N;B IF7J?7B H;IEBKJ?ED <EH J>EI;
FBEJI EDBO #;D=B ;J 7B 
 <:=/?7;5 1<::B;7AF 93C39 A?/7A 27@A?70BA7<;@ /1?<@@ ?357<;@
2; J;IJ;: ?< 7D: JE M>7J ;NJ;DJ J>; 9ECCKD?JOB;L;B JH7?J :?IJH?8K
J?EDI :?<<;H;: 7CED= H;=?EDI 2; IGK7H;HEEJ JH7DI<EHC;: J>; JH7?J L7BK;
JE ?CFHEL; DEHC7B?JO 7D: 7FFB?;: 7D 7D7BOI?I E< L7H?7D9; )*1 2;
J>;D 7FFB?;: 7 /KA;OI #ED;IJ .?=D?<?97DJ ?<<;H;D9; /KA;O #. J;IJ JE
?DL;IJ?=7J; J>; I?=D?<?97D9; E< J>; :?<<;H;D9;I 8;JM;;D J>; C;7DI E< J>;
9ECCKD?JO M;?=>J;: C;7D 2( JH7?J L7BK;I 7CED= BE97J?EDI />;
)*1 7D: /KA;O J;IJ M;H; 97HH?;: EKJ KI?D= J>; IJ7JIF79A7=; <EH -
 &39/A7;5 =7E3993C39 A?/7A 1<:=<@7A7<; A< @=31A?/9 ?3T31A/;13
3;C7?<;:3;A /;2 @<79 1<;27A7<;@
2; CE:;BB;: J>; 9ECCKD?JO M;?=>J;: C;7D 2( E< ;79> JH7?J 7J
J>; F?N;BB;L;B  e C 7I 7 <KD9J?ED E< J>; .;DJ?D;B H;CEJ; I;DI
?D= ;DL?HEDC;DJ7B 7D: IE?B 9EL7H?7J;I /78B;  KI?D= 7 IF7J?7BL;HI?ED
E< J>; C79>?D; B;7HD?D= -7D:EC !EH;IJ -! 7B=EH?J>C H;?C7D 
D7C;: ";E=H7F>?9 -7D:EC !EH;IJI "-! ";EH=7DEI ;J 7B  -!
?I 7 DEDF7H7C;JH?9 7B=EH?J>C J>7J >7I 8;;D I>EMD JE 8; HE8KIJ JE EL;H
`JJ?D= 7D: L7H?78B; ?DFKJI J>7DAI JE J>; 87==?D= FHE9;II 7D: ?JI H7D
:EC <;7JKH; I;B;9J?ED #7IJ?; ;J 7B  (EH;EL;H ?J >7I 8;;D ;N
J;DI?L;BO KI;: JE CE:;B 7D: FH;:?9J ;9EBE=?97B 7D: H;CEJ; I;DI?D= :7J7
M?J>?D 7D: 79HEII ;9EIOIJ;CI ;= ID;H ;J 7B  17D :;H +B7I ;J
7B  $D 9EDJH7IJ JE -! "-! :?I7==H;=7J;I J>; KD:;HBO?D= :7J7 ?D
=;E=H7F>?9 IF79; ?D J>?I 97I; 87I;: ED J>; IF7J?7B 9EEH:?D7J;I E< J>;
.;DJ?D;B F?N;BI 8K?B:?D= =BE87B 7D: BE97B IK8CE:;BI FBEJ B;L;B C7A
?D= J>; CE:;BB?D= <H7C;MEHA J>KI IF7J?7BBO ;NFB?9?J />; ;NFB?9?J ?D9BK
I?ED E< J>; IF7J?7B 9ECFED;DJ 34 F?N;B BE97J?ED ?D J>; CE:;BI M>?9>
7H; I;GK;DJ?7BBO `JJ;: M?J> :?<<;H;DJ I;JI E< J>; JH7?D?D= :7J7 J>; 87=
=?D= FHE9;II C7O 9EDJH?8KJ; JE J>; E8I;HL;: H;:K9;: IF7J?7B 7KJE9EHH;
B7J?ED E< "-! ?D 9ECF7H?IED JE J>; 9ECCED -! ";EH=7DEI ;J 7B 
$D J>; "-! 7 =BE87B CE:;B ?I 8K?BJ 7I ?D EJ>;H -! 7FFB?97J?EDI #EM
;L;H "-! 7BIE =;D;H7J;I 7 BE97B -! <EH ;79> BE97J?ED M>?9> ?D9BK:;I 7
IF;9?`;: DKC8;H E< D;7H8O E8I;HL7J?EDI >;H; :;`D;: 8O 7BB F?N;BI ?D
J>; L;=;J7J?ED FBEJ CEIJBO  >7 /78B;  97BB;: D;?=>8EKH>EE: E8
J7?D?D= ?D J>?I M7O C;JH?9I E< BE97B 7D: =BE87B CE:;B FH;:?9J?L; FEM;H
7D: L7H?78B; ?CFEHJ7D9; !EH CE:;B FH;:?9J?EDI 7 <KI?ED 8;JM;;D J>;
=BE87B CE:;B J>7J KI;I CEH; :7J7 7D: BE97B CE:;BI M?J> BEM 8?7I 97D
8; 7FFB?;: M;?=>J?D= J>; 9EDJH?8KJ?EDI E< J>; =BE87B 7D: BE97B CE:;BI
87I;: ED J>; F7H7C;J;HI J>7J ?D9H;7I; J>; FH;:?9J?L; 799KH79O 7D: :;
9H;7I; J>; CE:;BI -EEJ (;7D .GK7H; HHEH -(.  2; KI;: J>; IF7J?7B
"-! JE `J 7 =BE87B CE:;B <EH ;79> <KD9J?ED7B JH7?J 7D: 7BIE `J 7 IF;9?`9
CE:;B <EH ;79> H;=?ED KIJH7B?7 H7P?B ./ H7P?B )3 "78ED ">7D7
(7B7OI?7 7D: +;HK KI?D= J>; .F7J?7B(' F79A7=; ?D -
2; F;H<EHC;: 7D ;NJ;DI?L; I;J E< CE:;B EFJ?C?P7J?ED 7D: H;=KB7H
?P7J?ED FHE9;:KH;I JE H;:K9; EL;H`JJ?D= !EH J>; 2( CE:;BI M; I;
B;9J;: J>; DKC8;H E< JH;;I JE `J 8O <EB: 9HEIIL7B?:7J?ED 7D7BOI?I M?J>
DKC8;H E< JH;;I H7D=?D= 8;JM;;D  7D:  7D: J>; DKC8;H E< L7H?
78B;I H7D:ECBO I7CFB;: 7I 97D:?:7J;I 7J ;79> IFB?J CJHO H7D=?D= 8;
JM;;D  7D:  KI?D= ?D J>; `D7B CE:;B J>; 9EC8?D7J?ED E< J;HCI J>7J
=;D;H7J;: J>; BEM;IJ -(.  BB 9EL7H?7J;I ?D9BK:;: ?D J>; CE:;BI >7:
F7?HM?I; +;7HIED 9EHH;B7J?ED 9E;<`9?;DJI ? /78B;  !EH J>; `D7B
=BE87B 7D: BE97B CE:;BI M; KI;:  E< J>; :7J7 <EH CE:;B `JJ?D= 7D:
J>; H;C7?D?D=  <EH CE:;B ;L7BK7J?ED 17H?78B; ?CFEHJ7D9; <EH ;79>
CE:;B M7I 9ECFKJ;: 7I J>; :;9H;7I; ?D DE:; ?CFKH?J?;I <HEC IFB?JJ?D=
ED J>; L7H?78B; 7L;H7=;: EL;H 7BB JH;;I 7D: :;H?L;: <HEC J>; *KJ E< 7=
** ;HHEH />;D J>; H;IKBJ?D= ?CFEHJ7D9; M7I IJ7D:7H:?I;: JE 7 
I97B; <EH 9ECF7H?IED FKHFEI;I
2; 97HH?;: EKJ 7BB 7D7BOI;I IJ7J;: 78EL; M?J> J>; <KBB I;J E< JH;; ?D
:?L?:K7BI FH;I;DJ ?D ;79> L;=;J7J?ED FBEJ M?J> <KD9J?ED7B JH7?JI 7IIKC
?D= J>7J J>; 9EDJH?8KJ?ED E< IC7BB ?D:?L?:K7BI JE J>; JH7?J 2( L7BK;
7D: J>KI JE J>; 9ECCKD?JO H;a;9J7D9; 7J J>; F?N;BB;L;B MEKB: 8; C?D
?C7B =?L;D J>; M;?=>J?D= <79JEH KI;: ?; J>; ?D:?L?:K7BI 9HEMD 7H;7
#EM;L;H JE KD:;HF?D J>?I M; 97HH?;: EKJ 7BB 7D7BOI;I ED JME IC7BB;H
:7J7I;JI ED; M>;H; J>; J> 7D: EJ>;H M>;H; J>; J> F;H9;DJ?B; E< J>;
IC7BB;IJ JH;;I F;H H;=?ED M;H; H;CEL;: BB 7D7BOI;I M;H; 97HH?;: EKJ
?D J>; - IJ7J?IJ?97B ;DL?HEDC;DJ M?J> J>; 97H;JJ?:OL;HI;7D: .F7
J?7B('F79A7=;I
3. Results
 */?7/A7<; 7; A?/7A 1<:=<@7A7<; /1?<@@ A?<=71/9 4<?3@A@
(EIJ B;7< <KD9J?ED7B JH7?JI ;N>?8?J;: I?=D?<?97DJ :?<<;H;D9;I 79HEII J>;
JHEF?9I !?=  ?D9BK:?D= M?:; JH7?J H7D=; L7H?7J?ED M?J>?D J>; I7C; H;
=?ED !?= . M?J> B;7< <H;I> C7II 7D: B;7< J>?9AD;II 8;?D= ED 7L;H7=;
B;II L7H?78B; 7CED= BE97J?EDI /78B; .
UNCORRECTED PROOF
5B7??3BA7L??3G3A/9 &3:<A3'3;@7;5<4;C7?<;:3;A EEE EEEE EEEEEE
Fig. 2. ECF7H?IED E< JH7?J :?IJH?8KJ?EDI 79HEII JHEF?97B H;=?EDI />; 8ENFBEJI 7H; 87I;:
ED J>; F?N;BB;L;B  e  C 9ECCKD?JO JH7?J L7BK;I <EH ;79> JH7?J 7D: H;=?ED ; 
<EH KIJH7B?7  <EH H7P?B)3  <EH H7P?B./  <EH "78ED  <EH ">7D7
 <EH (7B7OI?7 7D:  <EH +;HK #EH?PEDJ7B B?D;I ?D ;79> 8ENFBEJ I>EM J>; C;
:?7D L7BK; 7D: L;HJ?97B B?D;I 7H; J>; M>?IA;HI J>7J ;NJ;D: JE J>; B7H=;IJ L7BK; EH DEJ
<KHJ>;H J>7D  J?C;I J>; ?DJ;HGK7HJ?B; H7D=; !EH IEC; BE97
J?EDI ?D<EHC7J?ED <EH 7BB JH7?JI M7I DEJ 7L7?B78B; !EH <KBB :;J7?BI ?D I?=D?<?97DJ :?<<;H;D9;I
?D C;7D JH7?J L7BK;I 7CED= BE97J?EDI I;; /78B; . H7P?B )3 )EL7 37L7DJ?D7 H7P?B ./
.7DJ7H;C
';7< 9>;C?IJHO 7D: F>EJEIODJ>;J?9 97F79?JO C7N 7D: I7J E<J;D
I>EM;: I?=D?<?97DJ :?<<;H;D9;I 7CED= BE97J?EDI /78B; . H?;H BE97
J?EDI 7I ?D )EL7 37L7DJ?D7 H7P?B )3 :?IFB7O;: JH7?J 7:7FJ7J?EDI JE
I;7IED7B H7?D<7BB 7D: J;CF;H7JKH; M?J> ED 7L;H7=; J>?9A;H 7D: IC7BB;H
 ]  CC 7D:  ]  9CH;IF;9J?L;BO B;7L;I 7J J>; 9EC
CKD?JO B;L;B M?J> IEC; E< J>; >?=>;IJ 9ECCKD?JOB;L;B B;7< D?JHE=;D
9ED9;DJH7J?ED  ]  7D: >?=>;IJ F>EJEIODJ>;J?9 97F79?JO C;7D
C7N   ]  μCEB CI 7D: I7J   ]  μCEB CI $D
9EDJH7IJ M;JJ;H H;=?EDI IK9> 7I (7B7OI?7 :?IFB7O;: ED 7L;H7=; IEC;
E< J>; 8?==;IJ  ]  9C 7D: J>?DD;IJ  ]  CC B;7L;I
M?J> >?=> B;7< M7J;H 9EDJ;DJ  ]  />; +;HKL?7D 7BJ?JK:?D7B JH7D
I;9J I>EM;: B7H=; L7H?7J?ED ?D 9ECCKD?JOB;L;B JH7?JI L7BK;I M>?9>
E<J;D EL;HB7FF;: M?J> JH7?J L7BK;I <HEC 7BB EJ>;H I7CFB;: BE97J?EDI
79HEII J>; JHEF?9I !?=  !EH CEIJ DKJH?;DJI B;7< DKJH?;DJ 9ED9;DJH7
J?ED M7I E<J;D >?=>;IJ ?D <EH;IJI <EKD: ?D ">7D7 ;= &  ] 
7D: (=   ]  7D: (7B7OI?7 &   ]  7D:
(=   ]  KIJH7B?7D <EH;IJI I>EM;: ED 7L;H7=; IEC; E< J>;
BEM;IJ 9ECCKD?JOB;L;B )  ]  7D: + ]  B;7<
9ED9;DJH7J?EDI
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J>; J> 7D: J> F;H9;DJ?B; E< J>; IC7BB;IJ JH;;I F;H H;=?ED M;H; H;
CEL;: =7L; I?C?B7H H;IKBJI <EH J>; =BE87B & 7D: & H;
IF;9J?L;BO /78B; . 7D: BE97B & 7D: & H;IF;9J?L;BO
/78B; . CE:;BI E< FB7DJ JH7?J :?IJH?8KJ?EDI />;H;<EH; ?D J>; <EBBEM?D=
M; EDBO FH;I;DJ J>; H;IKBJI <EH J>; CE:;BI 97HH?;: EKJ M?J> J>; <KBB L;=
;J7J?ED :7J7I;J
/>; 799KH79O E< J>; F7DJHEF?97B FH;:?9J?ED E< <KD9J?ED7B JH7?JI H7D=;:
8;JM;;D 7 C?D?CKC E< &  <EH B;7< <H;I> C7II 7D: 7 C7N ?CKC
E< &  <EH B;7< 97H8ED 9EDJ;DJ  87I;: ED J>; EKJE<I7CFB;:
J;IJ?D= :7J7 79HEII J>; JHEF?9I /78B;  />; FH;:?9J?L; 799KH79?;I E<
B;7< 9>;C?IJHO 7D: F>EJEIODJ>;J?9 JH7?JI M;H; E<J;D >?=>;H J>7D <EH CEH
F>EBE=?97B 7D: IJHK9JKH7B JH7?JI IK9> 7I B;7< :HO C7II &  7D:
B;7< 7H;7 & !?=  J J>; F7DJHEF?97B B;L;B J>; >?=>;IJ FH;
:?9J?ED 799KH79O M7I E8J7?D;: <EH B;7< J>?9AD;II &  <EH CEH
F>EBE=?97B 7D: IJHK9JKH7B JH7?JI <EH B;7< 7 7 &  7D: B;7< &
& &  <EH J>; 9>;C?97B JH7?JI EJ>;H J>7D 97H8ED ';7< ) 7D: +
9ED9;DJH7J?EDI M;H; 7BIE FH;:?9J;: M?J> >?=> 799KH79O &  ';7<
F>EJEIODJ>;J?9 97F79?JO JH7?JI C7N 7D: I7J I>EM;: IEC; E< J>; >?=>
;IJ FH;:?9J?ED 799KH79?;I H7D=?D= <HEC &  JE  H;IF;9J?L;BO
(E:;B IF7J?7B FH;:?9J?EDI <EH I;L;H7B JH7?JI 7D: BE97J?EDI 7H; I>EMD ?D
!?=  7D: EJ>;HI 97D 8; I;;D ?D !?= .!?= .
(E:;BI 8K?BJ <EH ;79> JHEF?97B H;=?ED 7D: JH7?J KD9EL;H;: C7HA;:
:?<<;H;D9;I ?D FH;:?9J?ED 799KH79O 7CED= J>;C !?=  /78B;  7D:
/78B; . ';7< 7H;7 FH;:?9J?ED 799KH79O H7D=;: <HEC &  H7P?B
./ JE  KIJH7B?7
Table 4
.J7J?IJ?97B H;IKBJI ED J>; J;IJ :7J7  E< <KBB :7J7I;J <EH J>; =BE87B JH7?J :?IJH?8KJ?ED
CE:;BI />; FH;:?9J?ED 799KH79O ?I I>EMD 8O J>; -I9EH;
/OF; /H7?J ( -(. -
(EHF>EBE=?97B7D:IJHK9JKH7B H;79C   
HOC7II=   
!H;I>C7II=   
.'C=   
/>?9AD;IICC   
>;C?IJHO '2   
   
7   
&   
(=   
)   
+   
+>EJEIODJ>;J?9 C7NμCEB CI   
I7JμCEB CI   
(  (;7D 8IEBKJ; HHEH -(.  -EEJ C;7D IGK7H; ;HHEH
UNCORRECTED PROOF
5B7??3BA7L??3G3A/9 &3:<A3'3;@7;5<4;C7?<;:3;A EEE EEEE EEEEEE
Fig. 3. (E:;B FH;:?9J?EDI JE J>;  J;IJ :7J7 <HEC J>; =;D;H7B CE:;B M>?9> M7I `JJ;: M?J>  E< J>; JH7?J :7J7 <HEC 79HEII J>; JHEF?9I "H;O :EJI 7H; J>; E8I;HL;: 7=7?DIJ FH;:?9J;:
JH7?J L7BK;I E< J>; F?N;BB;L;B  e C 9ECCKD?JO M;?=>J;: C;7D JH7?JI <HEC J>; J;IJ :7J7I;J />; 8B79A B?D; I>EMI J>;  H;B7J?EDI>?F 8;JM;;D E8I;HL;: 7D: FH;:?9J;: L7BK;I (E:;B
FH;:?9J?ED 799KH79O ?I I>EMD ?D J>; JEF B;<J !KBB CE:;B H;IKBJI 7H; I>EMD ?D /78B; 
UNCORRECTED PROOF
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Fig. 4. .F7J?7B FH;:?9J?EDI E< JH7?J :?IJH?8KJ?EDI <EH 7 I;B;9J;: IK8I;J E< FB7DJ JH7?JI 7D: BE97J?EDI />; C7F C?::B; I>EMI J>; BE97J?EDI E< L;=;J7J?ED FBEJI J>7J M;H; KI;: :KH?D= J>;
CE:;BB?D= <H7C;MEHA />; IF7J?7B FH;:?9J?EDI JEF 7D: 8EJJEC HEMI M;H; E8J7?D;: KI?D= J>; =;D;H7B CE:;BI /78B;  <EH ;79> E< J>; JH7?JI 7D: BE97J?EDI 7J 7  e C F?N;B H;IEBKJ?ED
/>; 7FFHEN?C7J;: BE97J?ED E< ;79> L;=;J7J?ED FBEJ KI;: ?I I>EMD 7I 7 M>?J; IGK7H; M?J>?D ;79> IF7J?7B FH;:?9J?ED C7F <EH L?IK7B?I7J?ED FKHFEI;I M>?J; IGK7H;I 7H; DEJ I97B;: JE J>; FBEJI
H;7B I?P; .F7J?7B FH;:?9J?EDI <EH EJ>;H JH7?JI 97D 8; <EKD: ?D !?= ..
7D: J>7J E< IF;9?`9 B;7< 7H;7 .' H7D=;: <HEC &  <EH (7B7OI?7
JE  <EH H7P?B )3 /78B; . />; BE97B CE:;BI I>EM;: 7 >?=>;H
799KH79O <EH FH;:?9J?D= BE97B B;L;B B;7< 9>;C?97B DKJH?;DJI KF JE
&  ;IF;9?7BBO <EH + 7 7D: ) 9ED9;DJH7J?EDI ?D 9ECF7H?IED JE
CEHF>EBE=?97B ;= B;7< 7H;7 7D: .' JH7?JI /78B;  !?=  /H7?JI H;
B7J;: JE F>EJEIODJ>;J?9 97F79?JO I>EM;: 7D EL;H7BB 8;JJ;H FH;:?9J?ED 79
9KH79O J>7D B;7< 7H;7 7D: .' M?J> FH;:?9J?ED L7BK;I H7D=?D= 8;JM;;D
 +;HK JE  ">7D7 <EH C7N 7D: KF JE  <EH I7J H7P?B
)3 !?=  *D 7L;H7=; J>; >?=>;IJ FH;:?9J?ED 799KH79O 79HEII H;=?EDI
<EH 7 =?L;D JH7?J M;H; H;79>;: <EH B;7< + 9ED9;DJH7J?ED &  7D:
C7N & 7D: J>; BE97J?EDI M?J> J>; >?=>;IJ 7L;H7=; FH;:?9
J?ED 799KH79O 79HEII JH7?JI M;H; J>; )EL7 37L7DJ?D7 I7L7DD7 H7P?B
)3 &  7D: J>; +;HK ;B;L7J?ED =H7:?;DJ &  /78B; 
8EJ> I?J;I ;D9ECF7II?D= IJHED= =H7:?;DJI ?D L;=;J7J?ED CEHF>EBE=O 7D:
IJHK9JKH;
 :=<?A/;13 <4 @=31A?/9 ?3:<A3 @3;@7;5 197:/A71 /;2 @<79 2/A/ 4<?
:/==7;5 A?/7A 27@A?70BA7<;@
2; ?D9BK:;: .;DJ?D;B 87D: :;H?L;: H;a;9J7D9; L7BK;I L;=;J7J?ED
?D:?9;I J>;?H 97DEFO J;NJKH; F7H7C;J;HI 9B?C7J?9 7D: IE?B L7H?78B;I
?D J>; =;D;H7B JH7?J CE:;BI JE FH;:?9J 9ECCKD?JO B;L;B JH7?JI 7J J>;
F?N;BB;L;B /78B;  />; ?CFEHJ7D9; E< J>;I; L7H?78B;I <EH FH;:?9J?D=
JH7?JI :;F;D:;: ED J>; IF;9?`9 JH7?J 8;?D= 7::H;II;: !?=  $D J>;
=BE87B CE:;B J>; H;CEJ; I;DI?D= J;NJKH; F7H7C;J;HI M;H; J>; `HIJ EH
I;9ED: C7@EH 9EDJH?8KJEH <EH FH;:?9J?D= D?D; E< J>; <KD9J?ED7B JH7?JI
79HEII J>; JHEF?9I !?=  7D: !?= . -7M IF;9JH7B L7H?78B;I M;H;
J>; I;9ED: CEIJ ?CFEHJ7DJ =HEKF <EH FH;:?9J?D= <EKH E< IK9> <KD9J?ED7B
JH7?JI 8KJ E<J;D BEM;H ?D ?CFEHJ7D9; J>7D J>; J;NJKH7B F7H7C;J;HI $D
J>; =BE87B CE:;B IE?B 7D: J;HH7?D <79JEHI M;H; ED 7L;H7=; IEC; E< J>;
CEIJ ?CFEHJ7DJ <EH FH;:?9J?D= F>EJEIODJ>;J?9 JH7?JI 7D: <EB?7H + 9ED9;D
JH7J?ED *D 7L;H7=; 9B?C7J?9 L7H?78B;I M;H; ?CFEHJ7DJ <EH FH;:?9J?D= 
EKJ E<  <KD9J?ED7B JH7?JI 8KJ J>;?H 9EDJH?8KJ?ED M7I BEM;H <EH FH;:?9J
?D= B;7< :HO 7D: <H;I> C7II 7D: B;7< M7J;H 9EDJ;DJ !?=  #EM;L;H ?J
?I ;L?:;DJ J>7J 7 9EC8?D7J?ED E< J;NJKH7B IF;9JH7B 9B?C7J?9 7D: IE?B ?D
<EHC7J?ED ?I H;GK?H;: JE E8J7?D J>; 8;IJ =;D;H7B CE:;B FH;:?9J?EDI 79HEII
<KD9J?ED7B JH7?JI 7D: DE I?D=B; L7H?78B; 7FF;7HI 7I J>; CEIJ ?CFEHJ7DJ
79HEII 7BB JH7?JI !?= .
/>; BE97B CE:;BI FHEL?:;: 7 I?J;IF;9?`9 L?;M E< J>; CEIJ ?CFEHJ7DJ
H;CEJ; I;DI?D= :;H?L;: L7H?78B;I ;DL?HEDC;DJ7B 7D: IE?B 9ED:?J?EDI <EH
:;H?L?D= 9ECCKD?JO B;L;B JH7?JI 9ECFEI?J?ED !?= . .;DJ?D;B H;
CEJ; I;DI?D= H;B7J;: L7H?78B;I M;H; CEH; ?CFEHJ7DJ <EH :;J;9J?D= B;7<
CEHF>EBE=O 7D: DKJH?;DJ L7BK;I J>7D ;DL?HEDC;DJ7B 7D: IE?B H;B7J;:
L7H?78B;I ?D  E< J>; JH7?J CE:;BI ?D  EKJ E<  FEII?8B; JH7?JI 8O
H;=?ED 9EC8?D7J?EDI ?=>JOED; F;H9;DJ E< J>; J?C;  BE97J?ED 8O
JH7?J 9EC8?D7J?EDI J>; 97DEFO J;NJKH; F7H7C;J;HI M;H; CEH; ?CFEHJ7DJ
J>7D J>; H7M IF;9JH7B H;a;9J7D9; <79JEHI $D  7D:  E< J>; FEI
I?8B; JH7?J 7D: H;=?ED 9EC8?D7J?EDI 9B?C7J?9 EH IE?BJEFE=H7F>O H;B7J;:
L7H?78B;I H;IF;9J?L;BO M;H; J>; CEIJ ?CFEHJ7DJ <EH :;J;9J?D= 9ECCKD?JO
JH7?JI !?= .
4. Discussion
/E J>; 8;IJ E< EKH ADEMB;:=; J>?I ?I J>; `HIJ IJK:O ;L7BK7J?D= J>;
78?B?JO E< .;DJ?D;B I7J;BB?J; :7J7 JE C7F FB7DJ <KD9J?ED7B JH7?JI 79HEII
JHEF?97B ;9EIOIJ;CI /HEF?97B <EH;IJ JH7?J C7FF?D= ?I <KD:7C;DJ7B <EH KD
:;HIJ7D:?D= E< FB7DJ H;IFEDI;I JE =BE87B 9>7D=; 7D: DEJ78BO J>; FB7DJ
<KD9J?ED7B JH7?JI M; FH;:?9J ?D J>?I IJK:O 7H; H;B;L7DJ JE FB7DJ IF;9?;I
H;IFEDI;I JE 7 9>7D=?D= ;DL?HEDC;DJ EJ> ;J 7B  )KD;I ;J 7B
 .EK:P?BELIA7?7 ;J 7B  =K?HH;"KJ?THH;P ;J 7B  2;
>7L; :;CEDIJH7J;: J>7J 799KH7J; F?N;BB;L;B  e  C FH;:?9J?EDI E<
JHEF?97B <EH;IJ <KD9J?ED7B JH7?J :?IJH?8KJ?EDI 79HEII J>; JHEF?9I 97D 8;
=;D;H7J;: 8O C7A?D= KI; E< ;NJ;DI?L; ?DI?JK 9EBB;9J;: FB7DJ <KD9J?ED7B
JH7?JI =;EBE97J;: 97DEFO IJHK9JKH; L;=;J7J?ED 9;DIKI;I 7D: >?=> IF;9
JH7B 7D: IF7J?7B H;IEBKJ?ED H;CEJ; I;DI?D= :7J7 <HEC J>; .;DJ?D;B I7J;B
B?J;I
 (?<=71/9 4<?3@A A?/7A 27@A?70BA7<;@
+B7DJ <KD9J?ED7B JH7?JI 7H; 9>7H79J;H?IJ?9I J>7J 7?: IF;9?;I JE J>H?L; ?D
J>;?H ;DL?HEDC;DJ EH 7:7FJ JE D;M 9ED:?J?EDI "?L;D IK9> 7:7FJ7J?EDI
JE IF;9?`9 ;DL?HEDC;DJI ?J C?=>J 8; ;NF;9J;: J>7J JH7?J L7H?7J?ED MEKB:
8; >?=>;H ?D H;=?EDI J>7J ;D9ECF7II CEH; L7H?;: ;DL?HEDC;DJ7B 9ED
:?J?EDI  DGK?IJ ;J 7B  DL?HEDC;DJ7B 7:7FJ7J?ED ?I ;N;CFB?`;:
8O J>; IJHED= L7H?7J?ED ?D L7BK;I <EH CEIJ JH7?JI ?D +;HK 7D: (7B7OI?7
$D +;HK J>; :7J7 H;FH;I;DJ 7 9B?C7J?9 7D: 7BJ?JK:?D7B =H7:?;DJ H7D=?D=
<HEC J>; BEMB7D: C7PED ?D J>; /7C8EF7J7 )7J?ED7B +7HA 7J 7D ;B;
L7J?ED E<  C7IB JE FBEJI ?D 9@7D79E 7J 78EL;  C7IB $D
(7B7OI?7 J>; L;=;J7J?ED FBEJI 7H; :?IJH?8KJ;: 79HEII 7 B7D:KI; =H7:?;DJ
H7D=?D= <HEC KD:?IJKH8;: JE >;7L?BO BE==;: <EH;IJI EJ> ;J 7B 
DL?HEDC;DJ7B 7:7FJ7J?ED C7O 8; 7BIE I>EMD 8O J>; E8I;HL;: :?<<;H
;D9;I ?D JH7?J :?IJH?8KJ?EDI 8;JM;;D :?<<;H;DJ H;=?EDI 79HEII J>; JHEF?9I
;= KIJH7B?7 7D: "78ED I;; 7BIE !?=  />; F?N;B87I;: 9ECCKD?JO
JH7?J L7BK;I ?D J>; +;HKL?7D JH7DI;9J E<J;D ;NJ;D: 79HEII CK9> E< J>;
H7D=; ?D JH7?J L7BK;I E8I;HL;: ?D EJ>;H BE97J?EDI !?=  2; :;J;9J;:
7D EL;H7BB I?=D?<?97DJ :?<<;H;D9; 7CED= BE97J?EDI ?D J;HCI E< CEHF>E
BE=?97B 9>;C?97B 7D: F>EJEIODJ>;J?9 JH7?JI /78B; . />?I M?:; L7H?7
J?ED ?D JH7?JI IK==;IJI J>; FH;I;D9; E< BE97B 8?EJ?9 7D: 78?EJ?9 9EDJHEBI E<
JH7?J :?IJH?8KJ?EDI 7D: FB7DJ IF;9?;I 7:7FJ7J?EDI J>7J C7O :?<<;H 7CED=
JHEF?97B H;=?EDI .K9> :?<<;H;D9;I ?D JH7?J 9ECFEI?J?ED >?=>B?=>J J>; ?C
FEHJ7D9; 7D: J>; 9>7BB;D=; E< I7CFB?D= 7I <KBBO 7I FEII?8B; J>; <KD9
J?ED7B JH7?J :?L;HI?JO 79HEII :?<<;H;DJ JHEF?97B <EH;IJI />?I ?I E< F?LEJ7B
?CFEHJ7D9; M>;D 9ECF7H?D= <EH;IJ H;IFEDI;I JE 9>7D=?D= ;DL?HEDC;DJI
79HEII CKBJ?FB; H;=?EDI 2; J>KI IK==;IJ J>7J <KHJ>;H `;B: JH7?J IKHL;O
97CF7?=DI 79HEII J>; JHEF?9I 7H; D;;:;: JE ?CFHEL; F7DJHEF?97B JH7?J
FH;:?9J?EDI I <EH J>; BE97B 8?EJ?9 7D: 78?EJ?9 9EDJHEBI E< JH7?J :?IJH?
8KJ?EDI <EH ?DIJ7D9; ?J ?I M?:;BO ADEMD J>7J <H?97D JHEF?97B <EH;IJI
7H; ?D =;D;H7B B;II IF;9?;I :?L;HI; J>7D J>;?H I?7D 7D: .EKJ> C;H?
97D 9EKDJ;HF7HJI 8KJ J>7J J>;O >7L; IEC; E< J>; >?=>;IJ 8?EC7II 97H8ED
UNCORRECTED PROOF
5B7??3BA7L??3G3A/9 &3:<A3'3;@7;5<4;C7?<;:3;A EEE EEEE EEEEEE
Fig. 5. (E:;BI FH;:?9J?EDI JE J>;  J;IJ :7J7 <HEC J>; H;=?ED7B CE:;BI `JJ;: M?J>  E< J>; JH7?J :7J7 <HEC ;79> H;=?ED 79HEII J>; JHEF?9I 79> 9EBEKH H;FH;I;DJI 7D ?D:?L?:K7B H;=?ED7B
CE:;B 7D: J>; 9EBEKH;: IOC8EBI 7H; J>; E8I;HL;: 7=7?DIJ FH;:?9J;: JH7?J L7BK;I E< J>; F?N;BB;L;B  e C 9ECCKD?JO M;?=>J;: C;7D JH7?JI <HEC J>; J;IJ :7J7I;J />; 8B79A B?D; I>EMI
J>;  H;B7J?EDI>?F 8;JM;;D E8I;HL;: 7D: FH;:?9J;: L7BK;I (E:;B FH;:?9J?ED 799KH79O ?I I>EMD ?D /78B;  !KBB CE:;B H;IKBJI 7H; I>EMD ?D /78B; . H7P?B )3 )EL7 37L7DJ?D7 H7P?B
./ .7DJ7H;C

UNCORRECTED PROOF
5B7??3BA7L??3G3A/9 &3:<A3'3;@7;5<4;C7?<;:3;A EEE EEEE EEEEEE
Table 5
+H;:?9J?ED 799KH79O - ED J>; J;IJ?D= :7J7 7CED= H;=?EDI I>7:;: -;=?ED C;7D -9EBKCD 7D: <KD9J?ED7B JH7?JI I>7:;: /H7?J C;7D -HEM )EJ I>7:;: L7BK;I ?D J>; J78B; I>EM J>;
FH;:?9J?ED 799KH79O - ED J>; J;IJ :7J7 F;H H;=?ED 7D: JH7?J
 DE :7J7 7L7?B78B; H7P?B )3 )EL7 37L7DJ?D7 H7P?B ./ .7DJ7H;C
IJEH7=; 97F79?JO F;H KD?J 7H;7 .KBB?L7D ;J 7B  /HEF?97B <EH;IJI
?D 2;IJ <H?97 7H; ?D =;D;H7B :H?;H ?D 9ECF7H?IED JE C7PED?7D JHEF?
97B <EH;IJI +7HC;DJ?;H ;J 7B  7D: IEC; <H?97D H;=?EDI IK9> 7I
"78ED >7L; ;NF;H?;D9;: ?D9H;7I;I ?D J;CF;H7JKH; 7D: :;9H;7I;I ?D FH;
9?F?J7J?ED EL;H J>; B7IJ  O;7HI KI> ;J 7B  />KI IK9> 9>7D=;I
?D 9B?C7J?9 9ED:?J?EDI 7I J>EI; E8I;HL;: ?D 2;IJ <H?97D JHEF?97B <EH;IJ
C7O 7BIE KD:;HB?; L7H?7J?EDI ?D IF;9?;I 9ECFEI?J?ED 7D: J>; BE97BBO E8
I;HL;: <KD9J?ED7B JH7?J FEEB 7I I>EMD ?D J>?I IJK:O $J ?I 7BIE MEHJ> DEJ
?D= J>7J JME 97L;7JI E< J>; 9ECCKD?JOM;?=>J;: C;7D JH7?J 7FFHE79>
C7O 799EKDJ <EH F7HJ E< J>; KD;NFB7?D;: JH7?J L7H?7J?ED !?HIJ ?J C7A;I
J>; 7IIKCFJ?ED E< 7 KD?GK; <KD9J?ED7B EFJ?CKC ?D 7 =?L;D ;DL?HEDC;DJ
M>?B; CKBJ?FB; EFJ?C7B IJH7J;=?;I FEJ;DJ?7BBO 9EHH;IFED:?D= JE 9ED
JH7IJ?D= JH7?J L7BK;I 9EKB: 9E;N?IJ '7K=>B?D ;J 7B  .;9ED:BO ?J
:E;I DEJ 799EKDJ <EH J>; :OD7C?9 D7JKH; E< 9ECCKD?J?;I IE J>7J 7 9EC
CKD?JO M;?=>J;: C;7D 7J 7 =?L;D J?C; FE?DJ C?=>J DEJ ;D9ECF7II J>;
EFJ?CKC 7J ;GK?B?8H?KC '7K=>B?D ;J 7B 
(EHF>EBE=?97B 7D: IJHK9JKH7B JH7?JI IK9> 7I B;7< 7H;7 <H;I> 7D: :HO
C7II B;7< J>?9AD;II .' 7D: '2 H;FH;I;DJ JH7:;EcI 8;JM;;D ;D
;H=O 79GK?I?J?ED 9EDIKCFJ?ED 7D: IKHL?L7B 7D: <EHC 7 C7?D F7HJ E<
J>; =BE87B IF;9JHKC E< FB7DJ <KD9J?ED?D= V7P ;J 7B  ;I?:;I ?D
L;IJ?=7J?D= J>; FH;:?9J78?B?JO E< IK9> FB7DJ IJHK9JKH7B JH7?JI M; <KHJ>;H
7D7BOI;: J>; FEJ;DJ?7B <EH FH;:?9J?D= B;7< 9>;C?IJHO  & (= 7 )
+ 7D: F>EJEIODJ>;I?I H;B7J;: JH7?JI C7N 7D: I7J (7FF?D= 9>;C?97B
7D: F>EJEIODJ>;J?9 JH7?JI 7J 7 F7DJHEF?97B I97B; >7I J>; FEJ;DJ?7B <EH ?D
9H;7I?D= EKH KD:;HIJ7D:?D= E< >EM F>EJEIODJ>;J?9 97F79?JO I>?<JI 79HEII
JHEF?97B H;=?EDI 7D: ED FEII?8B; ?CF79JI E< 7 9>7D=?D= ;DL?HEDC;DJ ED
JHEF?97B <EH;IJI FHE:K9J?L?JO "K7D ;J 7B  (K;BB;H ;J 7B 
 '3;A7;39 ?3:<A3 @3;@7;5 4<? :/==7;5 1<::B;7AF 93C39 A?/7A
27@A?70BA7<;@ /1?<@@ A63 A?<=71@
$D J>;?H F?ED;;H?D= MEHA M?J> >OF;HIF;9JH7B ?C7=;HO 7D: I?CKB7J;:
CKBJ?IF;9JH7B .;DJ?D;B :7J7 EL;H ">7D7 '7KH?D ;J 7B  :;CED
IJH7J;: J>7J .;DJ?D;B ?C7=;HO 9EKB: 8; KI;: JE :?I9H?C?D7J; JHEF?97B
<EH;IJ JOF;I 7D: C7F FB7DJ <KD9J?ED7B JOF;I />; 7KJ>EHI 7H=K;: J>7J
J>; <KBB 87D: I;J 7D: L;=;J7J?ED ?D:?9;I :;H?L;: <HEC J>; .;DJ?D;B
MEKB: 8; 7:L7DJ7=;EKI <EH 799KH7J;BO C7FF?D= FB7DJ <KD9J?ED7B =K?B:I
?D J>; JHEF?9I O KI?D= <KD9J?ED7B JH7?J :7J7 9EBB;9J;: ?D I?JK 79HEII
JHEF?97B <EH;IJI 7D: CE:;BB?D= 7J >?=> IF7J?7B H;IEBKJ?ED F?N;BB;L;B
M; I>EM J>7J CEIJ E< EKH =BE87B JH7?J :?IJH?8KJ?ED CE:;BI FH;I;DJ 7
>?=> FH;:?9J?L; FEM;H <EH CEIJ JH7?JI 7D7BOI;: M?J> FH;:?9J?ED 799K
H79O ED J>; J;IJ?D= :7J7I;JI 8;?D= >?=>;IJ <EH FH;:?9J?D= B;7< 9>;C
?97B 7D: F>EJEIODJ>;J?9 97F79?JO JH7?JI #EM;L;H M; 7BIE I>EM J>7J
J>; BE97B B;L;B JH7?J CE:;BI FHE:K9;: B;II 799KH7J; FH;:?9J?EDI J>7D
J>; =BE87B CE:;BI FHE878BO 7I 7 H;IKBJ E< J>; D7HHEM;H H7D=; E< ?D
FB7DJ JH7?J L7BK;I <EKD: 7J J>; BE97B H;=?ED ?D 9ECF7H?IED JE 79HEII
J>; H;=?EDI IEC;J>?D= 7BIE I>EMD 8O 27BB?I ;J 7B  />; FH;
:?9J?ED 799KH79O E8J7?D;: <HEC EKH CE:;BI KI?D= .;DJ?D;B CKBJ?IF;9
JH7B :7J7 ?I I?C?B7H 7D: ?D IEC; 97I;I >?=>;H J>7D J>7J I>EMD 8O H;
9;DJ IJK:?;I J>7J C7A; KI; E< >OF;HIF;9JH7B ?C7=;HO 7D: EJ>;H CKB
J?IF;9JH7B I;DIEHI JE C7F <KD9J?ED7B JH7?JI (7HJ?D ;J 7B  ID;H
;J 7B  ID;H ;J 7B  !EH ?DIJ7D9; EKH FH;:?9J?EDI ED J;IJ
:7J7 <EH B;7< D?JHE=;D F>EIF>EHKI 7D: 97H8ED 7H; 9ECF7H78B; EH >?=>;H
J>7D J>EI; E8J7?D;: 8O EJ>;H ?DDEL7J?L; IJK:?;I ?D (7B7OI?7 & 
 7D:  H;IF;9J?L;BO (7HJ?D ;J 7B  +;HK &  
7D:  ID;H ;J 7B  7D: J;CF;H7J; <EH;IJI & 
  )KD;I ;J 7B  7D: 9BEI;BO H;B7J;: JE J>EI; E8J7?D;:
8O 27BB?I ;J 7B  M?J> EJ>;H CKBJ?IF;9JH7B I;DIEH <EH D?JHE=;D 7D:
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?D :;J;HC?D?D= ;9EIOIJ;C FHE:K9J?L?JO 7D: <KD9J?ED?D= ;= 97H8ED 97F
JKH; V7P ;J 7B  7HCED7 ;J 7B  (EH;EL;H IF7J?7BBO ;N
FB?9?J CE:;BI E< <KD9J?ED7B JH7?JI I>?<J 79HEII J>; JHEF?9I 97D >;BF :;9?
F>;H >EM ;9EIOIJ;C <KD9J?ED?D= L7H?;I ;L;D 7CED= JHEF?97B 7H;7I FHE
L?:?D= 7 9EIJ;<<;9J?L; F7J>M7O JE ?:;DJ?<O?D= H;=?EDI E< >?=> 9EDI;HL7
J?ED L7BK; 7D: >;D9; 7?: ?D J>; 9H;7J?ED E< BE97BBO 7:;GK7J; 8?E:?L;H
I?JO 9EDI;HL7J?ED FEB?9?;I *L;H7BB EKH `D:?D=I 7H; E< H;B;L7D9; <EH ?D
<EHC?D= 8?E:?L;HI?JO CED?JEH?D= FEB?9?;I KD:;H ;9EIOIJ;C 9>7D=; 7I M;
I>EM J>7J 799KH7J; FH;:?9J?EDI E< H;B;L7DJ FB7DJ <KD9J?ED7B JH7?JI 97D 8;
E8J7?D;: ?D >?=> 8?E:?L;HI?JO 7H;7I IK9> 7I J>; JHEF?9I *KH 7FFHE79>
J>KI <79?B?J7J;I JH79A?D= FEII?8B; I>?<JI ?D JH7?J :?IJH?8KJ?EDI 7D: 9ECFEI?
J?ED 79HEII B7H=; IF7J?7B ;NJ;DJI 7I 7 H;IFEDI; JE ;DL?HEDC;DJ7B 9>7D=;I
KI?D= J>; .;DJ?D;B I7J;BB?J;I
5. Conclusions
/HEF?97B <EH;IJ ;9EIOIJ;CI 7H; M?JD;II?D= 7 H7F?: JH7DI<EHC7J?ED 7I
7 H;IKBJ E< 9>7D=?D= ;DL?HEDC;DJ7B 9ED:?J?EDI 7D: :?H;9J >KC7D ?C
F79JI ';M?I ;J 7B  /7K8;HJ ;J 7B  =K?HH;"KJ?THH;P ;J 7B
 #EM;L;H M; 97DDEJ 7:;GK7J;BO KD:;HIJ7D: EH I?CKB7J; JHEF?
97B ;9EIOIJ;C H;IFEDI;I JE ;DL?HEDC;DJ7B 9>7D=;I 87I;: IEB;BO ED 9KH
H;DJ ;9EIOIJ;C CE:;B 7FFHE79>;I 7I J>;I; 7H; KD78B; JE 97FJKH; J>;
>?=> :?L;HI?JO E< FB7DJ ;9EIOIJ;C <KD9J?EDI ?D J>; IF;9?;IH?9> JHEF
?9I );=B;9J E< <KD9J?ED7B 8?E:?L;HI?JO 97D EL;HI?CFB?<O J>; I?CKB7J;:
H;IFEDI; E< 7D ;9EIOIJ;C JE 7D ;DL?HEDC;DJ7B :?IJKH87D9; #;H; M;
I>EM J>; >?=> L7H?7J?ED ?D <KD9J?ED7B JH7?JI J>7J ;N?IJI 7CED= JHEF
?97B H;=?EDI M>?9> >?DJI 7J J>; :?<<;H;DJ 97F78?B?J?;I E< IK9> <EH;IJI
JE H;IFED: JE 7 9>7D=?D= ;DL?HEDC;DJ 2; :;CEDIJH7J; EFFEHJKD?
J?;I <EH C;7IKH?D= J>; :?IJH?8KJ?ED E< A;O <KD9J?ED7B JH7?JI 79HEII JHEF
?97B <EH;IJ ;9EIOIJ;CI 7J J>; F?N;BB;L;B KI?D= J>; .;DJ?D;B I7J;BB?J;I
M>?9> ?< :ED; 79HEII J?C; 9EKB: H;L;7B 7H;7I M>;H; <KD9J?ED7B I>?<JI
>7L; E99KHH;: 7D: B?A;BO M>;H; 8?E:?L;HI?JO 9EDI;HL7J?ED7C;B?EH7
J?ED C;7IKH;I 7H; D;;:;: BJ>EK=> J>; .;DJ?D;B I7J;BB?J;I I>EM >?=>
FHEC?I; <EH J>?I ;D:;7LEKH EKH 7FFHE79> ?I B?C?J;: 8O J>; I>EHJ J?C;
?DJ;HL7B I?D9; J>;O M;H; B7KD9>;: ?;  7D: J>; BEM;H IF;9JH7B H;I
EBKJ?ED E< .;DJ?D;B ?C7=;HO ?D 9ECF7H?IED JE J>7J :;H?L;: <HEC >OF;H
IF;9JH7B I;DIEHI (;J>E:I 7D: :7J7 FHE:K9JI 7H; D;;:;: JE JH79A 9>7D=;I
?D <KD9J?ED7B 9ECFEI?J?ED ?D <EH;IJ ;9EIOIJ;CI 79HEII J?C; 7D: IF79;
2; :;CEDIJH7J; 7 D;M 7FFHE79> JE :;L;BEF 7 H7F?: CED?JEH?D= JEEB
<EH 97FJKH?D= J>; ;<<;9JI E< 7 9>7D=?D= ;DL?HEDC;DJ 79HEII J>; JHEF?9I
/>?I D;M JEEB >7I J>; FEJ;DJ?7B JE 9EDJH?8KJ; JE 7 CEH; HE8KIJ 7D: ;L
?:;D9;87I;: FEB?9OC7A?D= <EH 9EDI;HL7J?ED E< JHEF?97B <EH;IJ ;9EIOI
J;CI
Authorship contribution statement
%" 9ED9;?L;: J>; IJK:O :;I?=D;: 7D: 97HH?;: EKJ J>; 7D7BOI?I 7D:
MHEJ; J>; `HIJ :H7<J E< J>; F7F;H 4( 9ED9;?L;: 7D: ?CFB;C;DJ;: J>;
" ( );JMEHA E8J7?D;: <KD:?D= <EH CEIJ E< J>; " ( JH7?JI `;B: 97C
F7?=DI 7D: 9ECC;DJ;: ED ;7HB?;H L;HI?EDI E< J>; C7DKI9H?FJ .- 7:
L?I;: ED IJ7J?IJ?97B 7D: H;CEJ; I;DI?D= 7D7BOI?I 7D: 9ECC;DJ;: ED ;7H
B?;H L;HI?EDI E< J>; C7DKI9H?FJ BB 9E7KJ>EHI F7HJ?9?F7J;: ?D EH 9EEH:?
D7J;: L;=;J7J?ED JH7?J :7J7 7D:EH IE?B :7J7 9EBB;9J?ED EH FHE9;II;: `;B:
:7J7 />; 7KJ>EHI D7C;: 8;JM;;D . 7D: '%/2 7H; B?IJ;: 7BF>7
8;J?97BBO BB 9E7KJ>EHI 9ECC;DJ;: ED 7D: 7FFHEL;: J>; C7DKI9H?FJ
Declaration of Competing Interest
/>; 7KJ>EHI :;9B7H; J>7J J>;O >7L; DE ADEMD 9ECF;J?D= `D7D9?7B
?DJ;H;IJI EH F;HIED7B H;B7J?EDI>?FI J>7J 9EKB: >7L; 7FF;7H;: JE ?DaK;D9;
J>; MEHA H;FEHJ;: ?D J>?I F7F;H
Acknowledgements
/>?I MEHA ?I 7 FHE:K9J E< J>; "BE87B 9EIOIJ;CI (ED?JEH?D= " (
D;JMEHA =;CJHEF?97B<EH;IJIEN79KA %" M7I <KD:;: 8O J>; )7J
KH7B DL?HEDC;DJ -;I;7H9> EKD9?B ) - ) / 7D: ) 
. 7D: J>; );J>;HB7D:I *H=7D?P7J?ED <EH .9?;DJ?`9 -;I;7H9>
)2* KD:;H J>; -K8?9ED FHE=H7CC; M?J> FHE@;9J DKC8;H
'2 />; JH7?JI `;B: 97CF7?=D M7I <KD:;: 8O 7 =H7DJ JE
4( <HEC J>; KHEF;7D -;I;7H9> EKD9?B :L7D9;: "H7DJ
" (/-$/  KD:;H J>; KHEF;7D 0D?EDI .;L;DJ> !H7C;MEHA
+HE=H7CC; !+ M?J> 7::?J?ED7B IKFFEHJ <HEC ) -
"H7DJ )  7D: ) % <EH JH7?JI MEHA ?D +;HK ) -
"H7DJ *!*- ) & <EH JH7?JI MEHA ?D .7DJ7H;C ) -
"H7DJ '$ ) & <EH FBEJ 7D: JH7?JI MEHA ?D (7B7OI?7 7D:
- :L7D9;: "H7DJ /!*- .  JE +>?BB?FI <EH JH7?JI MEHA ?D
KIJH7B?7 +BEJ I;JKF ?D ">7D7 7D: "78ED M;H; <KD:;: 8O 7 ) - "H7DJ
) $ 7D: 8O J>; -EO7B .E9?;JO';L;H>KBC; <H?97 7F79?JO
K?B:?D= +HE=H7CC; />; (7B7OI?7 97CF7?=D M7I 7BIE <KD:;: 8O ) -
"H7DJ ) & +BEJ ?DL;DJEH?;I ?D +;HK M;H; IKFFEHJ;: 8O <KD:
?D= <HEC J>; 0. )7J?ED7B .9?;D9; !EKD:7J?ED 'ED=/;HC -;I;7H9> ?D
DL?HEDC;DJ7B ?EBE=O FHE=H7C '/-     7D: J>; "EH
:ED 7D: ;JJO (EEH; !EKD:7J?ED D:;IC7PED +HE=H7C +BEJI ?DL;D
JEH?;I ?D )EL7 37L7DJ?D7 H7P?B M;H; IKFFEHJ;: 8O J>; )7J?ED7B EKD
9?B <EH .9?;DJ?`9 7D: /;9>DEBE=?97B ;L;BEFC;DJ )+G 'ED= /;HC
9EBE=?97B -;I;7H9> +HE=H7C + ' +HE9  7D: J>;
!EKD:7J?ED E< -;I;7H9> .KFFEHJ E< (7JE "HEIIE !+ (/ +HE@;9J
-;!BEH +HE9  KH?D= :7J7 9EBB;9J?ED $* M7I IKFFEHJ;:
8O 7 (7H?; KH?; !;BBEMI>?F !++ *+' $ ! " ( JH7?J
:7J7 ?D "78ED M7I 9EBB;9J;: KD:;H 7KJ>EH?I7J?ED JE 4( 7D: IKFFEHJ;:
8O J>; "78ED )7J?ED7B +7HAI =;D9O  M7I <KD:;: 8O J>; !ED:7
J?ED 2?;D;HDIF79> 2& 79ADEMB;:=;I IKFFEHJ <HEC J>; !79KBJO
-;I;7H9> BKIJ;H "BE87B 9EBE=OE< J>; 0D?L;HI?JO E< CIJ;H:7C (.
M7I <KD:;: 8O 7 =H7DJ <HEC J>; (?D?IJHO E< :K97J?ED 4EKJ> 7D: .FEHJI
E< J>; P;9> -;FK8B?9 $)/ -/-).! - '// 4( ?I IKFFEHJ;:
8O J>; %79AIED !EKD:7J?ED 2; J>7DA J>; JME 7DEDOCEKI H;L?;M;HI 7D:
IIE9?7J; :?JEH " #;D;8HO <EH J>;?H ?DI?=>J<KB 9ECC;DJI J>7J >;BF;:
?CFHEL;: J>?I C7DKI9H?FJ
UNCORRECTED PROOF
5B7??3BA7L??3G3A/9 &3:<A3'3;@7;5<4;C7?<;:3;A EEE EEEE EEEEEE
Appendix A. Supplementary data
.KFFB;C;DJ7HO :7J7 JE J>?I 7HJ?9B; 97D 8; <EKD: EDB?D; 7J >JJFI
:E?EH=@HI;
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87JPE=BEK %/ E8HEMIA? .5 +7HAI . #;=;M?I9> &  /;HH7B?C7J; 7
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 .9? :7J7  
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7B  H?;H JHEF?97B <EH;IJI 7H; IKI9;FJ?8B; JE <KD9J?ED7B 9>7D=;I ?D H;IFEDI; JE 7
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... Relative accuracy is widely used to evaluate the performance of impervious surface indices, such as RF-based [57][58][59], SVM-based [17,60,61], and CART [62] indices. In this study, these algorithms had high omission errors and misclassifications due to the limited quantity and quality of the self-constructing samples. ...
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... With ongoing technological advances, satellite remote sensing observations are poised to move beyond the monitoring of land cover change and quantify plant functional diversity across regional landscapes [33,35]. Indeed, a growing number of studies demonstrate the retrieval and analysis of individual canopy and leaf traits in discrete plots and locations through satellite remote sensing [36][37][38][39][40][41][42]. Yet, the step from using internally consistent trait estimates to deriving functional diversity patterns from satellite remote sensing is undertaken less often and remains limited to spatially discrete observations as opposed to the potential to facilitate spatially continuous 'wall-to-wall' inference [42][43][44][45]. ...
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Global biodiversity losses erode the functioning of our vital ecosystems. Functional diversity is increasingly recognized as a critical link between biodiversity and ecosystem functioning. Satellite earth observation was proposed to address the current absence of information on large-scale continuous patterns of plant functional diversity. This study demonstrates the inference and spatial mapping of functional diversity metrics through satellite remote sensing over a large key biodiversity region (Sabah, Malaysian Borneo, ~53,000 km2) and compares the derived estimates across a land-use gradient as an initial qualitative assessment to test the potential merits of the approach. Functional traits (leaf water content, chlorophyll-a and -b, and leaf area index) were estimated from Sentinel-2 spectral reflectance using a pre-trained neural network on radiative transfer modeling simulations. Multivariate functional diversity metrics were calculated, including functional richness, divergence, and evenness. Spatial patterns of functional diversity were related to land-use data distinguishing intact forest, logged forest, and oil palm plantations. Spatial patterns of satellite remotely sensed functional diversity are significantly related to differences in land use. Intact forests, as well as logged forests, featured consistently higher functional diversity compared to oil palm plantations. Differences were profound for functional divergence, whereas functional richness exhibited relatively large variances within land-use classes. By linking large-scale patterns of functional diversity as derived from satellite remote sensing to land-use information, this study indicated initial responsiveness to broad human disturbance gradients over large geographical and spatially contiguous extents. Despite uncertainties about the accuracy of the spatial patterns, this study provides a coherent early application of satellite-derived functional diversity toward further validation of its responsiveness across ecological gradients.
... Methods or techniques that help to obtain functional data on a large scale can contribute to the understanding of the ecosystem. Therefore, mappings based on multispectral and hyperspectral images have sought to fill this information gap [40][41][42][43][44]. Satellite observations to study the phenology of tropical forests using vegetation indices as a proxy for photosynthetic activity and leaf renewal (leaf flush or leaf flushing), combined with in situ data, can help to minimize the uncertainties raised and improve the biophysical understanding of forest dynamics in wet and dry periods. ...
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... To gain a better understanding of how the ecosystem reacts to environmental pressures, it is important to combine information on the extent and fragmentation of mangroves with biophysical variables such as forest density, leaf area index, chlorophyll content, and other functional attributes and vegetation characteristics (Pham et al. 2019). Mapping of biophysical variables that inform us about vegetation characteristics of mangroves would allow us to assess the ecosystems' health, phenology, functional attributes, and diversity thereof (Aguirre-gutiérrez et al. 2021;Lausch et al. 2016). However, such detailed indicators of ecosystem integrity are still rarely measured in mangroves at large spatial scales (Younes Cárdenas, Joyce, and Maier 2017). ...
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Mangrove forests provide vital ecosystem services. The increasing threats to mangrove forest extent and fragmentation can be monitored from space. Accurate spatially explicit quantification of key vegetation characteristics of mangroves, such as leaf area index (LAI), would further advance our monitoring efforts to assess ecosystem health and functioning. Here, we investigated the potential of radiative transfer models (RTM), combined with active learning (AL), to estimate LAI from Sentinel-2 spectral reflectance in the mangrove-dominated region of Ngoc Hien, Vietnam. We validated the retrieval of LAI estimates against in-situ measurements based on hemispherical photography and compared against red-edge NDVI and the Sentinel Application Platform (SNAP) biophysical processor. Our results highlight the performance of physics-based machine learning using Gaussian processes regression (GPR) in combination with AL for the estimation of mangrove LAI. Our AL-driven hybrid GPR model substantially outperformed SNAP (R2 = 0.77 and 0.44 respectively) as well as the red-edge NDVI approach. Comparing two canopy RTMs, the highest accuracy was achieved by PROSAIL (RMSE = 0.13 m2.m−2, NRMSE = 9.57%, MAE = 0.1 m2.m−2). The successful retrieval of mangrove LAI from Sentinel-2 can overcome extensive reliance on scarce in-situ measurements for training seen in other approaches and present a more scalable applicability by relying on the universal principles of physics in combination with uncertainty estimates. AL-based GPR models using RTM simulations allow us to adapt the genericity of RTMs to the peculiarities of distinct ecosystems such as mangrove forests with limited ancillary data. These findings bode potential for retrieving a wider range of vegetation variables to quantify large-scale mangrove ecosystem dynamics in space and time.
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