Recuerdo de Ramón Margalef - Encuentros en la Biología

Primavera 2014
!!
!
>"2%"(+*#+"#>.1?)#!.(&.,"9#
F. Xavier Niell
Catedrático de Ecología de la Universidad de Málaga
fxn@uma.es !
!
!"#$%&'"("#%)*#+"#,*$#"+'-*("$#+"#,.#("/'$-.#0%"#1"#.2%"(+"#+"#0%'3)#-.)4
-*#.5(")+67#8,#"+'-*(#0%"#1"#,*#5'+"#9%"#%)*#+"#1'$#.,%1)*$#1:$#+"$-.2.4
+*$;#<*=#"$#=.#5(*9"$*(#=#"$#+"#,*$#5(*9"$*("$#%)'/"($'-.('*$#0%"#5'")$.7##
>.1?)# !.(&.,"9# "(.# %)# 5")$.+*(;# 2*)# "$-.# .@'(1.2'?)# 2%15,*# +*$# <'-*$A# .2'"(-*# +"# 1*+*#
"B.2-*#.,#2.,'@'2.(,*;#"(.#%)#5")$.+*(;#=#+*=#$.-'$9.22'?)#.#$%#1"1*('.#=#,"#(')+*#.$6#<*1").C";#
5%"$-*#0%"#",#/.,*(#+",#5")$.1'")-*#"(.#,*#0%"#1:$#"$-'1.D.7##
8$-"# ")$.=*# )*# ("2*)$-(%'(:# ,.# /'+.# +",# E(*9"$*(# # !.(&.,"9F# "$-:# =.# 5%D,'2.+.7# !'# +'$2%($*#
+.(:#/%",-.$#.,("+"+*(#+"#$%$#2%.,'+.+"$#2*1*#5")$.+*(#=#,*#0%"#+"#+'2<.$#2%.,'+.+"$#$"#+"$-',?#
2'")-6@'2.#=#"+%2.-'/.1")-"7#
!
7
8,#5,.2"(#+"#C%&.(#2*)#",#2*)*2'1'")-*#
!.(&.,"9#%$.D.#5.(.#.2%+'(#.#,.#G.2%,-.+#",#-(.)$5*(-"#5HD,'2*#+"#I.(2",*).;#+"26.#0%"#.$6#,"#
+.D.#%)#-'"15*#5.(.#5")$.(#.)-"$#+"#,,"&.(777#J%#$")-'+*#+"#%$*#=#.5(*/"2<.1'")-*#+",#-'"15*#
("0%"(6.#+"#1*1")-*$#+"#$*,"+.+#=#+"#.5.(")-"#.%$")2'.#0%"#1%2<*$#("2*(+.1*$#2*1*#'1.4
&")#$%=.#.,#/"(,"#")$'1'$1.+*##2%.)+*#5.$:D.1*$##+",.)-"#+"#$%$#+"$5.2<*$7#
!*$-(.D.# $%# 5("9"(")2'.# 5*(# ,*$# 13-*+*$# 2*)$-(%2-'/*$# +"# "B5,*(.2'?)# +"# ,*$# +.-*$;# 5*(#
"C"15,*#'(#')2*(5*(:)+*,*$#%)*#.#%)*;#)*#.#%).#-.D,.#8B2",#5.(.#5.$.(,*$#.#%)#5(*&(.1.##&(:@'2*#
KLM.,"'+.&(.5<#%#*-(*$NO;#$')*#.#&(:@'2.$#0%"#'D.#2*)$-(%=")+*##5%)-*#.#5%)-*#$*D("#5.5",#1','4
1"-(.+*7# P.+.# /.,*(# )%"/*# 0%"# ')2*(5*(.D.# .# ,.# &(:@'2.;# -(.6.# %).# <'5?-"$'$# 0%"# <.26.# 5.(.# $'#
$*D("#,.#5*$'2'?)#"$5"(.+.#+",#)%"/*#+.-*#")#(",.2'?)#.#,*$#+"1:$7#!'")-(.$#3$-"#"(.#%)#13-*+*#
2*)$-(%2-'/*#"#')-"(.2-'/*;#,*$#2*15%-.+*("$#,"#+.D.)#%).#$*,%2'?)#'(('-.)-"1")-"##$HD'-.#=#5,.4
).;#,.#"B5,*(.2'?)#)*#$"#5")$.D.;#-")'.#0%"#+"+%2'($"#+",#"B.1")#Q.#5*$-"('*('R;#0%"#"(.#1%2<*#
1")*$#+'):1'2*#0%"#",#/"(#).2"(;#5*2*#.#5*2*;#",#2*15*(-.1'")-*#+",#$'$-"1.777#!.(&.,"9#$')-"4
-'S.D.#$%$#$")-'1'")-*$#+"#1*+*#$*(5(")+")-"1")-"#"$2%"-*#=#")#("9"(")2'.#.#,*#0%"#<"#2*1")4
-.+*#+"26.#0%"#,*$#*(+").+*("$#Q)*#,"#.=%+.D.)#.#5")$.(R7##
J%#2(6-'2.#.#,*$#*(+").+*("$#"(.#%).#<'53(D*,"#.,&*#9(6/*,.;#5%"$-*#0%"#4.#5"$.(#+"#$%#'(*)6.#
$*D("#,.$#1:0%').$4#"(.#%)#")-")+'+*#")#1*+",.+*7#!"#("@'"(*#.#0%"#2*15(")+6.#,*$#1*+",*$#=#
$.D6.#,.#9*(1.#0%"#-")6.)#0%"#-")"(#")#2.+.#2.$*;#$"#")-%$'.$1.D.#QC%&.)+*R#2*)#",,*$;#5.(.1"4
-('S:)+*,*$#=#$'1%,:)+*,*$#2*)#$%#1.0%')'-.#+"#2.,2%,.(;#0%"#1.)"C.D.#+%(.)-"#$%$#/'.C"$#5.(.#
1.-.(#",#-'"15*7#T"#"$-.#1.)"(.#"B5,*(?#",#2*15*(-.1'")-*#2.?-'2*#+"#$'$-"1.$#0%"#5(*D.D,"4
1")-"#)*#,*#$*)#2*1*#,.$#')-"(.22'*)"$#+"5("+.+*(45("$.7#
8,#.$5"2-*#,H+'2*#+*1').D.#")#&(.)#1.)"(.#,.#5(*$5"22'?)#+"##,.#).-%(.,"S.#0%"#<.2'.#!.(&.4
,"97#J"#")-%$'.$1.D.#=#$"#$*(5(")+6.#.#1"+'+.#0%"#'D.#5(*$5"2-.)+*#,*$#("$%,-.+*$#0%"#'D.#')4
2*(5*(.)+*#.#$%$#2*)2,%$'*)"$##=#$"#',%$'*).D.#2%.)+*#2*)@'(1.D.#%).#("&,.#*#%).#,"=7#T%(.)-"#
1%2<.$#"B2%($'*)"$#2*)#3,;#("2%"(+*#0%"#)*#+"C.D.#+"#1*C.(#%)#5"+.2'-*#1%=#5"0%"U*#K!.(&.4
,"9#<'S*#,.#&%"((.#2'/',O#+"#5.5",#')+'2.+*(#+"#5V#")#-*+*#2%.)-*#2<.(0%'-*#./'$-.D.7#W,#$.2.(,*#
+",#.&%.#1*$-(.D.#",#2*,*(#1*(.+*;#*(&%,,*$*#+",#1'$1*;#")#,*$#2<.(2*$#+*)+"#<.D6.#9*-*$6)-"4
$'$;#2%.)+*#,*#')"$5"(.+*#"(.#0%"#$.,'"$"#+"#*-(*#2*,*(7#8)#%).#*2.$'?)#2%.)+*#=*#=.#1"#<.D6.#
<"2<*#1.=*(;#=#)*#.)-"$;#,"#5("&%)-3A#XLY#0%3#"$5"(.D.;#")#%)#2<.(2*#,,")*#+"#.,&.$;#.#5,").#,%S#
+",#!"+'-"((:)"*N#LZ%"#$.,'"(.#(*C*NX#!"#2*)-"$-?#0%"#')$'$-'")+*#")#",#5.(.+'&1.#$"#")2*)-(.4
D.)#,.$#.)*1.,6.$;#,*#1'$1*#<.D6.#+'2<*#.,&%)*$#$'&,*$#.)-"$#['*(+.)*#I(%)*;#.%)0%"#.#3$-"#,*#
0%"1.(*)777#
!.(&.,"9#1.)"C.D.#",#2*)*2'1'")-*#")#$")-'+*#"$-('2-*;#$.D6.#,*#0%"#$"#-")'.#0%"#$.D"(;#=#$%#
2*)*2'1'")-*#"(.#.15,'*#=#5(:2-'2*;#1.)"C.D.#1*+",*$#+'9"(")2'.,"$#*#.(("&,.D.#,.#("$'$-")2'.#
+"#%)#"$5"2-(*9*-?1"-(*;#-")6.#+"$+"##-.,")-*#.#<.D','+.+#1.)%.,7#W)+.(#2*)#",#5*(#,.#).-%(.,"S.#
5.(.#%)#C*/")#,'2")2'.+*#"(.#+"2*(.S*).+*(;#,*#2*)*26.#=#,*#"B5,'2.D.#-*+*;#=#2%.)+*#+'&*#-*+*;#
5.(.#0%3#/.1*$#.#")-(.(#")#+"-.,,"$777#1"#("@'"(*#.#-*+*7#
Vol.7 ¦ Nº 147
"
T'$9(%-.D.#2*)#,.#*(-*+*B'.;#5"(*#"(.#<"-"(*+*B*;#2*15.-'D','S.D.#.14
D.$#2*$.$#+"1*$-(.)+*#0%"#,.#1'B-%(.#)*#$*,*#"$#5*$'D,";#$')*#0%"#-.14
D'3)#"$#$.).;#5")$.D.#2*)-')%.1")-"#")#1*+*$#+'9"(")-"$#+"#"B5,'2.(#%)#
1'$1*#5(*2"$*;#2*)#,*#2%.,#.D(6.#,.#5*$'D','+.+#+"#,.#9*(1%,.2'?)#+"#<'5?4
-"$'$#.,-"().-'/.$#=#$%#2*15(*D.2'?)7#>"%)6.#$*,/")-"1")-";#.#$%#1.)"(.;#
.#G"="(.D")+#=#.#E*55"(#=#.#,.#$?,'+.#"+%2.2'?)#M(.%$'$-.#+"#,.#\)$-'-%4
2'?)#]'D("#+"#8)$"U.)S.;#5*(0%"#)*(1.,1")-"#"(.#2.5.S#+"#'1.&').(#,.#
$*,%2'?)#0%"#$"#+.(6.#5.(.#2.+.#<'5?-"$'$777#Y#$'#)*#$"#+.D.;#",,*#2*)$-'-%6.;#
5("2'$.1")-";#,.#D.$"#5.(.#9.,$.(,.7#
!
!
8,#5(*9"$*(#!.(&.,"9#
8
8,#5(*9"$*(#>.1?)#!.(&.,"9#+.D.#2,.$"$7#J%$#2,.$"$;#+"26.1*$#$%$#.,%1)*$;#"(.)#+"#Q#W(-"#=#
8)$.=*R# 2*1*# ",# 2')"# +"# ,.# 35*2.;# ",# +"# # I"(&1.));# M%(*$.^.;# W)-*)'*)';# G.$$D')+"(;# [*+.(+#
]*$"=;#=#*-(*$;#5"(*#)*#"(.#G",,')'.)*777#]*$#.,%1)*$#+"26.1*$#0%"#,.$#2,.$"$#Q"$-.D.)#1%=#D'")#
+"#-32)'2.#5"(*#)*#$"#")-")+6.#",#.(&%1")-*R777#8$-.#2('-'2.;#0%"#<"#*6+*#1:$#/"2"$#2%.)+*#=*#<"#
"B5,'2.+*#"2*,*&6.#5*$-"('*(1")-";#1"#5.("2"#'15(*5'.#+"#5"($*).$#0%"#0%'$'"(.)# #.5(")+"(#.#
5")$.(7#8)#,.$#2,.$"$#<.D6.#%).#+*2-(').#+"#9*)+*;#)*#+"1.$'.+*#.15,'.;#2*)-')%.1")-"#$"#"/'4
+")2'.D.#$%#/.,'+"S;#")#2.+.#2'(2%)$-.)2'.;#5.(.#2.+.#$'$-"1.;#=#<.$-.# #")#2.$*$#.5.(")-"1")-"#
+'9"(")-"$7#8,#')2*)/")'")-"#"$#0%"#,.$#"B2%($'*)"$#5*(#",#.15,'*#"$5.2'*#+"#2*)*2'1'")-*$#)*#$"#
./'$.D.)#=#",#-(.D.C*#+"#")$.1D,.(#2.+.#2.$*#=#*(+").(,*#+*)+"#,"#2*(("$5*)+6.;#")#",#$")*##+"#
,.#_"*(6.;#"(.#",#+",#"$-%+'.)-"7##
Vol.7 ¦ Nº 147
Primavera 2014
!
P("6.#-.)-*#")#,.#2.,'+.+#')-","2-%.,#+"#,.$#5"($*).$#0%"#.D.)+*).D.# #,*$#
"B:1")"$#=#+"C.D.#$*,*$# .# ,*$# "$-%+'.)-"$F# .,&%)*$# 2*5'.D.);# 5"(*# )*# ,"#
'15*(-.D.;#3,#D%$2.D.#,.#+'9"(")2'.#+"#,*$#0%"#.5*(-.D.)#$%#'+".#*#$%#/'4
$'?)#5"($*).,#$*D("#",#-"1.#+"$'&).+*;#1:$#0%"#,.#("5(*+%22'?)#1"1*4
(6$-'2.# +"# ,*$# 2*)-")'+*$7# Z%"(6.# X2(*1.-*&(.@'.(X# ")# QD.)+.$R# .# ,*$# +'$4
-')-*$# "$-%+'.)-"$;# "$-.D.# ')-"("$.+*# 5*(# ,.# 2(".-'/'+.+7# W,&%).$# /"2"$#
")2*)-(.D.#+"$/'.2'*)"$#.#,*#9("2%")-";#,*#2%.,;#1"#2*)$-.;#,"#.,"&(.D.7#
P*)$'+"(.D.#0%"#")$"U.(#.#)'/",#%)'/"($'-.('*#"(.#-(.)$1'-'(#5(*D,"1.$#=#
1*+*$#+"#5")$.1'")-*#=#.(&%1")-.2'?)7#J%$#.$'&).-%(.$;#")1.(2.+.$#+"#5,")*#")#,.#I'*,*&6.#
&")"(.,;#2*1*#)*#"(.)#*D,'&.-*('.$;#)*#"(.)#"$2*&'+.$#5*(#1%2<*$#.,%1)*$7#8)#,.$#2,.$"$#-"?('4
2.$#"B'$-6.#%)#2*)$-.)-";#=#5.(.#.,&%)*$#+"$2*)2"(-.)-";#2(%2"#+",#2*)*2'1'")-*#+"#%)#,.+*#.,#
*-(*#+"#,.#1.-"('.;#6D.1*$##+"#"B-("1*#.#"B-("1*7#J%$#5(:2-'2.$;#&")"(*$.$#")#",#%$*#+"#-*+*#,*#
0%"# <.D6.# .# +'$5*$'2'?);# $"# .9(*)-.D.)# 2*)# %).# $"('"+.+# (*-%)+.# 2*1*# $'# 2*)$-'-%="(.)# %)#
-"1.#+"#-(.D.C*#(".,7#V.26.#5"(2'D'(#",#+"-.,,"#+",#"B5"('1")-*#2*1*#$")-'1'")-*#+"#,.#1:B'1.#
'15*(-.)2'.#")#,.#.+0%'$'2'?)#+"#+.-*$7#
`*#+.D.#+"1.$'.+.##'15*(-.)2'.#.#,*$#2*)-")'+*$;#5*+6.#5("$2')+'(#+"#-"1.$#5%"$-*#0%"#,.$#
1.-"('.$#"(.)#.,&*#1:$#0%"#,.#$%1.#+"#2.+.#%)*#+"#,*$#-"1.$;#0%"#$'"15("#"(.)#+'$2%-'+*$#2*)#
%).#"B<'D'2'?)#+"#2(6-'2.#5"($*).,7##
!
9
a(&.)'S.(# =# E(*&(.1.(# 9(")-"# .# [")"(.(# P.5.2'+.+# +"# W2-%.4
2'?)#\)+"5")+'")-"###
# # >"@,"B'*).D.# -.1D'3)# $*D("# ,.# ')%-','+.+# +"# +"-.,,.(# ,*$# 5,.)"$# +"# "$-%+'*;# %)# -"1.# 1%=#
.2-%.,# =# ("2%((")-"# ")# ,*$# H,-'1*$# &*D'"()*$# +",# "$-.+*7# J"# +"2.)-.D.# 5*(# 5,.)"$# +"# "$-%+'*#
0%"# ")$"U.(.)# ,.$# ,"="$# 9%)+.1")-.,"$# +",# %)'/"($*;# =# +"# ,.# D'*,*&6.# ")9.-'S.)+*# $%$# 2*')2'4
+")2'.$;#$*D("#-*+*#-"(1*+'):1'2.$;#0%"#$"#5("*2%5.(.)#1:$#+"#*('")-.(#,.$#2.5.2'+.+"$#+"#
,*$#"$-%+'.)-"$#0%"#,*$#2*)*2'1'")-*$#$%1.+*$#%)*#-(.$#*-(*7#8)#"$-"#.$5"2-*#"$-*=#+"#.2%"(+*#
2*)#3,;#$*D("#-*+*#2%.)+*#+"26.#0%"#+"$5%3$#+"#0%"#,*$#5*,6-'2*$#<%D'"(.)#2.1D'.+*#-.)-.$#/"4
2"$#,*$#5,.)"$#+"#"$-%+'*;#",#("$%,-.+*#"(.#",#1'$1*A#$.,6.)#5*2.$#5"($*).$#1%=#5*(#")2'1.#+"#
,.#1"+'.;#.,&%).$#+"$-.2.+.$;#=#%).#&(.)#1.$.#5*(#+"D.C*#+"#,.#1"+'.7#8$#+"2'(;#0%"#,.#("$5%"$4
-.#+",#2*,"2-'/*#<%1.)*;#.9*(-%).+.1")-";#$"#1.)-")6.#')2,%$*#")#,.$#5"*("$#2'(2%)$-.)2'.$##=#
0%"#,.#"+%2.2'?)#)*# #1"C*(.D.#1:$#0%"#,"/"1")-"#2%.)+*#,.#"+%2.2'?)#-")6.#-*+*$#,*$#')&("4
+'")-"$#5.(.#2*)$'+"(.($"#9./*("2'+.7#8,#")*(1"#"$9%"(S*#+"#,*$#5"+.&*&*$#5*(#",#2*)-(*,#+"#,.#
")$"U.)S.#)*#"$-.#+.)+*;#b)'#,*$#+.(:c;#("$%,-.+*$#+'&)*$#+"#1"("2"(#%)#2*1")-.('*7#d)#e8>\4
G\PW;# =# $%$# X1.('.2<'$X;# <%D'"(.)# 5"(-%(D.+*# ",# -.,.)-"# =# ",# <%1*(# +",# E(*9"$*(# !.(&.,"97# ].#
2.,'+.+# )*# "$# "$*;# )'# $"# 1'+"# 5*(# ,.$# ")2%"$-.$;# $"# 1'+"# 5*(# ,.# 2.5.2'+.+# +"# ,*$# "$-%+'.)-"$#
5.(.#$%5"(.(#,.$#1.-"('.$#$"('.$7#
!.(&.,"9#"(.#1%=#"$-'1.+*#")-("#.,&%)*$#=#("2<.S.+*#5*(#*-(*$;#)*#$3#"B.2-.1")-"#")#0%3#
5(*5*(2'?)7#P*)#$"&%('+.+#.#3,#)*#,"#'15*(-.D.#+"1.$'.+*;#$%#3-'2.#-(.$2")+6.#.#$%$#)%,.$#.)4
$'.$# +"# 5*5%,.('+.+7# E("+'2.D.# 2*)# ",# "C"15,*A# 1:$# 0%"# 5("*2%5.($"# 5*(# ",# 13-*+*# +"# ")$"4
U.)S.;#$"#5("*2%5.D.#5*(#-(.)$1'-'(#%).#5.%-.# #+"#2*15*(-.1'")-*#(.2'*).,'S.+*(7#Y*#1'$1*#
,"#("2%"(+*#1:$#5*(#,.#1.)"(.#+"#-(.D.C.(#")#,.#("@,"B'?)#+"#,.#`.-%(.,"S.#0%"#5*(#$%$#")$"4
U.)S.$#2*)2("-.$7#]*#0%"#$"#.5(")+6.#+"#3,#"(.#2*)$'$-")-";#%$.D,"#=#5"($*).,'S.D,";#%)*#,*#5*4
+6.#2*)/"(-'(#")#%).#5.%-.#5(*5'.;##5"(1'-6.#2(".(#",#1*+",*#1")-.,#0%"#+'"(.#.##")-")+"(#2.+.#
5(*D,"1.;#*#$".#$"#')2'-.D.#.#,.#'1.&').2'?)#"$5.2'.,;#.#,.#2*)$-(%22'?)#+"f#b$?,'+*$#2.$-',,*$#")#
",#.'("c#T"#,*#0%"#)*#2.D"#+%+.#"$#0%"#,.#'15(*)-.#0%"#+"C.D.#$"#$"5.(.D.#+"##,*#2*1H)7#
J'#",#,"2-*(#"$-:#')-"("$.+*#")#,.#5.(-"#1*(D*$.#+"#,.#-(.="2-*('.#/'-.,#+",#E(*9"$*(#!.(&.,"9#
0%"#,".#*-(.$#("2")$'*)"$A#=*#("2%"(+*#=#"$-'1*#",#('"&*#0%"#2*(('?#2%.)+*#,"#,,"/.(*)#.#2*1D.4
-'(#")#,.#+'/'$'?)#g\#2*)#",#[")"(.,#]6$-"(#")#,.#D.-.,,.#+",##8D(*;#+%(.)-"#"$.#&%"((.#0%"#5(*/*4
2.(*)#,*$#0%"#,.#5(*/*2.(*)F#"$-%+'?#-.(+";#-"(1')?#,.#2.(("(.#0%"#2%($?#")#",#$"&%)+*#0%')4
0%")'*#+"#,*$#/"')-"#.#,*$#-("')-.;#")#-("$#.U*$;#,,")*#+"#!.-(62%,.$#+"#V*)*(;#9%"#",#5('1"(#2.4
-"+(:-'2*#+"#82*,*&6.#+",#8$-.+*#=#1%2<*$#"-2777#
!
!
Vol.7 ¦ Nº 147
10
W#1';#@',*$?@'2.1")-";#!.(&.,"9#1"#5.("2"#%)#:2(.-.#0%"#+"$-',?#",#1:B'1*#/.,*(#+"#,.#.2(.2'.A#
,.#,'D"(-.+#+"#5")$.1'")-*#=#,.#')+"5")+")2'.#5"($*).,7#\)-"(2.,*#%).#"/*2.2'?)#$%=.#"$2('-.#")#
%).#*D(.#1%=#("2*)*2'+.A#!"#$%&'()*+#+#K!.(&.,"9#hiijO#0%"#"B-(.'&*#+"#,.#"B2",")-"#("/'$'?)#+"#
G(.)2"$2#E"-"($#KE"-"($#klhlO#A#
!"#$%&#'(%(#)*+'%(,-(%+#.+/$.%/+#0#123%4%0*5/%)#/('%&#$/%(,-.%0-62/$'%-.7%8//0%&#$/%*.+0*./7%(#%8-.9
(-'23%8//0*.1%-.7%*.')*$-:#.%(,-.%(#%$*1#$3%+#.'*'(/.+2%-.7%/;/.%$/')#.'*<*0*(2=%4.%&2%;*/6%#.%/.;*$#.9
&/.(-0%)$#<0/&'3%4%8//0%&#$/%->$-+(/7%<2%(,/%#$*1*.%#8%(,/%($#?<0/'%(,-.%<2%(,/*$%'#0?:#.'3%-(%0/-'(%*.%
(,/%6-2%(,/%)$#<0/&%*'%?'?-002%8-+/7%-(%)$/'/.(=@%
T"$-.2*#,.#H,-'1.#9(.$"A#")#",,.#<.D,.#+",#1*+*#+"#")9*2.(#,.$#2*$.$#")#",#5("$")-";#,*$#-?5'2*$#
,"#1*,"$-.D.);#"(.#$"('.1")-"#<*$-',#.#,.$#1*+.$#+"#,.#')/"$-'&.2'?)7#`*#<%D'"(.#5"(1'-'+*#0%"#
%)#-(.D.C*#$*D("#,.#')@,%")2'.#+"#,.#_"15"(.-%(.#=#",#de#$"#-'-%,.(.#1:$#0%"#')@,%")2'.#+"#+'2<.$#
/.('.D,"$#")#,*#0%"#9%"(.;#$')#)"2"$'+.+#"/*2.(#.,#2.1D'*#2,'1:-'2*7#e'/'?#,.#5('1"(.#35*2.#+"#,.$#
.5,'2.2'*)"$#=#+"#,.#%-','+.+#+"#,.#')/"$-'&.2'?)7#8)#(".,'+.+#2*)$'+"(.D.#0%"#",#2*)*2'1'")-*#")#
&")"(.,#2*)$-'-%="#%)#5.-('1*)'*;#",#1:$#/.,'*$*#0%"#"B'$-"#5.(.#",#<*1D(";#5*(#$6#1'$1*;#$')#0%"#
-")&.# 0%"# $"(/'(# 5.(.# ).+.7# ],"&?# .# +"2'(# Q`*# <.=# 2'")2'.# D:$'2.# )'# .5,'2.+.A# <.=# D%").# =# 1.,.#
2'")2'.R7#].$#2*)$'&).$#+"#,.#35*2.#"(.)#-.)#5"$.+.$#2*1*#,.$#+"#,*$#5*,6-'2*$#.2-%.,"$7#
E.(.# .)'1.(# .# ,*$# +"$.)'1.+*$# 0%'"(*# ("2*(+.(# %).# H,-'1.# .)32+*-.A# W# >.1?)# !.(&.,"9# ,"#
("2<.S.(*)#%)#5(*="2-*#+"#')/"$-'&.2'?)#")#,.$#5('1"(.$#2*)/*2.-*('.$#")#0%"#9%)2'*)?#%)#$'$4
-"1.#+"#@').)2'.2'?)#+"#5(*="2-*$#5.("2'+*#.,#.2-%.,1")-"#/'&")-"7#T"$5%3$#+"#,.#,?&'2.#$*(5("$.#
=#.,&*#+"$.$*$"&.+*#5*(##)*#$"(#5(*9"-.#")#$%#-'"((.;#')+.&?#")#,.$#+*$#')$-'-%2'*)"$#0%"#5%+'"(*)#
"/.,%.(# =# +"$2.,'@'2.(# $%# 5(*="2-*;# 5%"$-*# 0%"# "(.)# ,.$# H)'2.$# 0%"# <.D6.# ")# ",# 8$-.+*# ")# .0%",#
-'"15*;#5.(.#")-")+"(#0%3#(.S*)"$#<%D*#5.(.#",#("2<.S*777#]*$#2'")-6@'2*$#+"#,.$#+*$#$"#.2%$.D.)#
1%-%.1")-"7# P%.)+*;# 1:$# -.(+";# ,*# 2*)-.D.;# ("2%((6.# .# ,.# '(*)6.# =# 1%(1%(.D.# 2*)# %).# $*)('$.#
562.(.A#X5*(#5('1"(.#/"S#"$#5*$'D,"#0%"#-*+*$#+'C"(.)##,.#/"(+.+X777###
P'")-6@'2.1")-"# 1"# 0%"+.# ",# ("2%"(+*# +"# $%# -(.D.C*# "$-(",,.;# 0%"# 5.(.# 1'# 1*+*# +"# /"(# "$# ,.#
.5,'2.2'?)#+"#,.#_"*(6.#+"#,.#\)9*(1.2'?)#.#,.#I'*,*&6.7#8$#.)-*,?&'2*#5*(#$%#('&*(#"#'1.&').2'?)#.#
,.#/"S7#8$-"#-(.D.C*#"),.S.#2*)#$%#2*)$-.)-"#5("*2%5.2'?)#5.(.#.).,'S.(#",#$'&)'@'2.+*#+"#,.#/.('.4
D','+.+#).-%(.,#+"#,*$#*(&.)'$1*$#=#,*$#"2*$'$-"1.$#+.)+*#5*(#$")-.+*#",#2.(:2-"(#5(*D.D',6$-'2*#
+"#2%.,0%'"(#.@'(1.2'?)#0%"#$"#<'2'"(.#$*D("#",#2*15*(-.1'")-*#+"#,.#`.-%(.,"S.7#8$-"##5,.)*#.H)#
)*#$"#<.#.,2.)S.+*#")#,.#")$"U.)S.#%)'/"($'-.('.#.2-%.,#+*)+"#$?,*#$"#")$"U.#+"-"(1')'$1*#$')#
')-(*+%2'(# .# ,*$# .,%1)*$# ")# ,.# 2*)$'+"(.2'?)# +"# ,.# ')2"-'+%1D("# +"# 2%.,0%'"(# .@'(1.2'?);# +"#
2%.,0%'"(# ,"=;# +"# 2%.,0%'"(# 5(')2'5'*7# ]"# 2%5*# ,.# *('&').,'+.+# =# -.1D'3)# ,.# /.,")-6.# +"# ')2,%'(# .,#
<*1D("#2*1*#mR5'"S.#)*(1.,R#+",#9%)2'*).1'")-*#+"#,*$#"2*$'$-"1.$7#8(.#%)#5.$*#<.2'.#%).#&,*4
D.,'S.2'?)#+",#%$*#+"#,.#")"(&6.7#J"#.5*=.D.#$*D("#%).#')&")-"#=#2*-'+'.).#2.5.2'+.+#5.(.#*D$"(4
/.(#1%"$-(.$#+"#.&%.$#2*)-')")-.,"$#=#1.(').$7#!.(&.,"9#5")$.D.#"$-.$#1%"$-(.$;#0%"#,"#')$5'(.4
D.)#5(')2'5'*$#&")"(.,"$#+"#9%)2'*).1'")-*#=#*(&.)'S.2'?)#9%"(.)#+"#,*#0%"#9%"(.)7#`*#+"$5("4
2'.D.#,*#5"0%"U*;#2*15(")+6.#0%"#%)#2<.(2*#"(.#%)#"2*$'$-"1.#=#0%"#")#%)#"2*$'$-"1.#.15,'*#
5*+6.#("%)'(#.#+'/"($*$#2<.(2*$7#J%#2*)-('D%2'?)#.,#2*)*2'1'")-*#+"#,*$#*(&.)'$1*$#.2%:-'2*$#")#
&")"(.,#"$#%).#D.$"#5.-('1*)'.,#K+'(6.#=*O#+"#2*)*2'1'")-*#5.(.#,.#]'1)*,*&6.#+"#8$5.U.7##
P*1*#1."$-(*#+"$-.2.(3#$'"15(";#5*(#")2'1.#+"#,*#0%"#)*$##")$"U?;#,*#0%"#+"#3,#.5(")+'1*$#=#
-*+./6.#5*+"1*$#.5(")+"(7##
!
Referencias:
! Margalef, R. 1997. Our Biosphere. Excellence in Ecology 10. Ecology Institute. Oldendorf/Luhe.
! Peters, F. 2010. Ramón Margalef, the curiosity driven life of a self-taught Naturalist. Limnologia and Oceanography Bulletin
19 (1): 2-15.
Vol.7 ¦ Nº 147