Unraveling Our Human History: Habitats of Early Hominid Evolution

!
!
!
!
!
!
!
!
!
!
!
"#$%&'()#*!+,$!-,.%#!-)/01$23!-%4)0%0/!15!6%$(2!-1.)#)7!6&1(,0)1#!
8#%(2/)/!15!09'!:;<=!>%(,'/!15!?(%#0!@'%5!A%B'/!)#!%!C%$)#'!D'7).'#0!=1$'!
!"#$%&'(#)%*+(&
'#,(-)&./#0(0&'123(44#5&6%7&89:;&
!"#$%&'#()*+,*-+.(/%&"*012#,+&"#13*4%51&(6%"(*+,*71&(/*-8#%"8%'3*9+'*:";%2%'3*0:
EF!84/0$%G0!
!
<#7&#=-"14(-,%)7&>#%41)#0?&01*/&%0&2(@#5%"(03?&%)#&A(5#"7&)#B%)5#5&%0&
%5%@4(=#&)#0@-,0#0&4-&*"(3%4#C&&.-&2#44#)&1,5#)04%,5&A/%4&*"(3%4#&*-,5(4(-,0&5)-=#&
4/#0#&%5%@4%4(-,0?&@%"#-#,=()-,3#,4%"&)#*-,04)1*4(-,0&*%,&2#&@)-51*#5&10(,B&
4#))#04)(%"&@"%,4&"#%>&A%$#0&@)#0#)=#5&(,&3%)(,#&0#5(3#,40C&&D"%,4&"#%>&A%$#0&%)#&
4)%,0@-)4#5&27&A(,5&%,5&5#@-0(4#5&(,&3%)(,#&0#5(3#,40?&*-,4%(,(,B&%,&(0-4-@(*&
>(,B#)@)(,4?&4/#&*%)2-,&(0-4-@(*&*-3@-0(4(-,&E!:;FG?&4/%4&)#*-)50&A/(*/&
@/-4-07,4/#4(*&@%4/A%7&EF;&-)&FHG&A%0&5-3(,%,4&%4&4/%4&@%)4(*1"%)&4(3#C&&I7&
-20#)=(,B&-0*(""%4(-,0&(,&4/#&!:;F&=%"1#0?&4/#&0/(>4&2#4A##,&>-)#04#5&*%,-@(#0&4-&-@#,&
0%=%,,%/&*%,&2#&-20#)=#5?&%""-A(,B&10&4-&)#*-,04)1*4&/%2(4%40&>-)&#%)"7&/13%,&
#=-"14(-,C&&6%)(,#&0#5(3#,40&A#)#&%,%"7J#5&>)-3&K##@&'#%&K)(""(,B&D)-L#*4&'(4#&8;:&
(,&4/#&M1">&->&!5#,?&*-))#0@-,5(,B&4-&;CNOHC8&6%C&&./(0&%B#&)%,B#&(0&->&@%)4(*1"%)&
(,4#)#04&2#*%10#&(4&#,*%@01"%4#0&0-3#&->&4/#&#%)"(#04&2(@#5%"&/-3(,(50C&&
!@@)-$(3%4#"7&:99&0%3@"#0&A#)#&@)-*#00#5?&>()04&(0-"%4(,B&4/#&@"%,4&"#%>&A%$#0&27&
*-,51*4(,B&%**#"#)%4#5&0-"=#,4&#$4)%*4(-,&%,5&*-"13,&*/#3(04)7?&4/#,&10(,B&3%00&
0@#*4)-3#4)7?&B%0&*/)-3%4-B)%@/7?&%,5&(0-4-@(*&)%4(-&%,%"70#0&4-&5#4#)3(,#&!:;F&
=%"1#0C&&P#01"40&0/-A#5&%,&-@#,(,B&(,&4/#&"%,50*%@#&2#4A##,&HC8&%,5&HC9&6%?&
(,5(*%4#5&27&%&HCQR&0/(>4&4-&"#00&5#@"#4#5&!:;F&=%"1#0?&%,5&4/#,&%&;CSR&)#41),&4-&
3-)#&5#@"#4#5&!:;F&=%"1#0&>)-3&HC9&4-&;CN&6%?&(,5(*%4(,B&%&F;&5-3(,%,*#C&&T,&
*-3@%)(0-,?&4/#&-=#)%""&)%,B#&->&=%)(%2("(47&2#4A##,&@1)#&F;&%,5&@1)#&FH&#*-0704#30&
(0&:URC&&./(0&04157&(0&@%)4&->&%&"%)B#)&#>>-)4&4-&)#*-,04)1*4&4/#&/(04-)7&->&
@%"#-#,=()-,3#,40&0@%,,(,B&4/#&"%04&:8&3(""(-,&7#%)0C&
&
8&
EEF!E#0$17,G0)1#!
!
V,5#)04%,5(,B&4/#&*"(3%4#&->&
,-)4/#%04&!>)(*%&51)(,B&4/#&"%4#&
F#,-J-(*&(0&*)1*(%"&(,&1,5#)04%,5(,B&
/13%,&#=-"14(-,C&&!>)(*%,&*"(3%4#&(0&
(,>"1#,*#5&27&4#*4-,(*&*/%,B#0?&B"-2%"&
*"(3%4#&4)%,0(4(-,0?&%,5&"-*%"&=%)(%4(-,0&
(,&-)2(4%"&>-)*(,B&E6%0"(,&#4&%"C?&899SGC&&
./#0#&*-,5(4(-,0&=%)(#5&B)#%4"7&(,&4/#&
"%4#&F#,-J-(*?&*%10(,B&#$4)#3#&
-0*(""%4(-,0&(,&,-)4/#%04&!>)(*%,&
*"(3%4#&4/%4&3%7&/%=#&0#)=#5&%0&4/#&
*%4%"704&>-)&#=-"14(-,%)7&*/%,B#0&
%3-,B04&#%)"7&/-3(,(5&0@#*(#0&
E5#6#,-*%"?&899HGC&&6%)(,#&0#5(3#,40&
(,5(*%4#&4/%4&51)(,B&4/#&"%4#&W#-B#,#?&
H)*,$'!;3&&F-3@%)(0-,&->&*/)-,-"-B(*%"&
01**#00(-,&->&4/#&/-3(,(5&)#*-)5&(,&!>)(*%&A(4/&
@%04&=#B#4%4(-,&*/%,B#0&%,5&B"%*(%"&*7*"#0&
EI-,,#>(""#?&89:9GC&
0124)-@(*%"&!>)(*%,&*"(3%4#&#$@#)(#,*#5&
>)#X1#,4&-0*(""%4(-,0&2#4A##,&A#4&%,5&5)7&*-,5(4(-,0?&*-))#0@-,5(,B&4-&/(B/O
"%4(415#&B"%*(%"&*7*"#0&EY(B1)#&:GC&&./#0#&*"(3%4#&-0*(""%4(-,0&%)#&)#"%4#5&4-&+#7&
L1,*41)#0&(,&#%)"7&/13%,&#=-"14(-,&(,*"15(,B?&0@#*(%4(-,&*/%,B#0?&4/#&#3#)B#,*#&->&
4/#&B#,10&<+6+?&%,5&4/#&5#=#"-@3#,4&->&%5%@4(=#&>#%41)#0&01*/&%0&2(@#5%"(03&
E6%0"(,&#4&%"C?&899SGC&&
!
&
;&
!
I)55'$'#G'!)#!?9101/2#09'0)G!?%09J%2/&&
&P#*-,04)1*4(,B&4/#&4)%,0(4(-,0&2#4A##,&>-)#04&%,5&B)%00"%,5&=#B#4%4(-,&
*-=#)&(,&,-)4/#%04&!>)(*%&51)(,B&4/#&"%4#&W#-B#,#&(0&(3@-)4%,4&(,&1,5#)04%,5(,B&4/#&
/%2(4%40&->&#%)"7&/13%,&0@#*(#0?&2#*%10#&=#B#4%4(-,&5#4#)3(,#0&>--5&)#0-1)*#0&
E'@-,/#(3#)&#4&%"C?&899QG?&/%2(4%40?&%,5&#*-"-B(*%"&,(*/#0&EI-,,#>(""#?&89:9GC&&./#0#&
4)%,0(4(-,0&2#4A##,&>-)#040&%,5&B)%00"%,50&)#@)#0#,4&%&0/(>4&(,&5-3(,%,*#&->&@"%,40&
14("(J(,B&F;&%,5&FH&@/-4-07,4/#4(*&@%4/A%70&EY#%+(,0&#4&%"C?&899UGC&&F;?&4/#&3-)#&
@)(3(4(=#&@%4/A%7?&10#0&4/#&F%"=(,&F7*"#&%"-,#&>-)&*%)2-,&>($%4(-,&EY#%+(,0&#4&%"C?&
89:;GC&&./#&FH&Z[%4*/O'"%*+\&@%4/A%7&>()04&%@@#%)#5&(,&4/#&]"(B-*#,#&214&>"-1)(0/#5&
"%4#)&(,&4/#&6(-*#,#&EY#%+(,0&#4&%"C?&89:;GC&&V,"(+#&4/#&F;&@%4/A%7?&4/#&FH&@%4/A%7&
10#0&4A-&0#@%)%4#&#,J73#0?&-,#&>-)&4/#&1@4%+#&->&*%)2-,&5(-$(5#&%,5&%,-4/#)&>-)&
@/-4-07,4/#0(0&E^%""#)&%,5&_#A(0?&:`S`GC&&./(0&1,(X1#&%5%@4%4(-,&#,%2"#0&FH&@"%,40&
4-&10#&A%4#)&3-)#&#>>(*(#,4"7?&%,5&*%))7&-14&@/-4-07,4/#0(0&(,&#$4)#3#&/#%4&%,5&"(B/4&
*-,5(4(-,0&E^%""#)&%,5&_#A(0?&:`S`GC&&./#)#>-)#?&F;&@"%,40&4/)(=#&(,&*--"?&A#4&
*-,5(4(-,0&*-))#0@-,5(,B&4-&%&>-)#04#5&#,=()-,3#,4?&A/("#&FH&@"%,40&4/)(=#&(,&/-4?&
5)7&*-,5(4(-,0&01*/&%0&B)%00"%,50C&
D'('G0)1#!15!?$1B2!
!
F;&%,5&FH&@/-4-07,4/#4(*&@%4/A%70&/%=#&5(04(,*4&04%2"#&*%)2-,&(0-4-@(*&
*-3@-0(4(-,0?&!:;F?&A/(*/&%)#&)#*-)5#5&(,&@"%,4&"#%>&A%$#0&E]a_#%)7?&:`N:GC&&D"%,4&
"#%>&A%$#0&%)#&)#%5("7&%2)%5#5&>)-3&/-04&@"%,40?&%,5&4)%,0@-)4#5&27&A(,5&4-&3%)(,#&
0#5(3#,40?&A/#)#&4/#7&@)#0#)=#&4/#&(0-4-@(*&*-3@-0(4(-,&->&4/#&=#B#4%4(-,&0-1)*#&
E6#7#)0&%,5&b%5(#?&:``;c&Y)##3%,&%,5&F-"%)100-?&899:c&'*/#>100&#4&%"C?&899;GC&&
&
H&
V0(,B&4/#&!:;F&->&21"+&-)B%,(*&*%)2-,&(0&(,01>>(*(#,4&>-)&@%"#-=#B#4%4(-,&
)#*-,04)1*4(-,?&2#*%10#&4/#)#&3%7&2#&31"4(@"#&*%)2-,&0-1)*#0&->&=%)7(,B&(0-4-@(*&
*-3@-0(4(-,0&ED#%0-,&%,5&bB"(,4-,?&8999GC&&&
&
_-,B&*/%(,&,O%"+%,-(*&%*(50?&0@#*(>(*%""7&F8Q?&F8N?&%,5&F;9?&%)#&4/#&3-04&)#"(%2"#&
@)-$(#0&>-)&4#))#04)(%"&=#B#4%4(-,&EY#%+(,0&#4&%"?&899SGC&&./#0#&,O%"+%,-(*&%*(50&/%=#&
3-"#*1"%)&5(04)(214(-,0&*/%)%*4#)(04(*&->&%&@"%,4&"#%>&A%$&0-1)*#&%,5&/%=#&*%)2-,&
(0-4-@(*&=%"1#0&A(4/(,&4/#&)%,B#&->&*1))#,4&F;&%,5&FH&#,5&3#32#)0&EY#%+(,0&#4&%"C?&
899UGC&&!:;F&=%"1#0&>-)&F;&4)##0&%,5&0/)120&47@(*%""7&)%,B#&2#4A##,&O;:CH&%,5&&
O8HCQR?&A/("#&FH&B)%00#0&)%,B#&2#4A##,&O:HC:&4-&O::CUR&EF#)"(,B&#4&%"C?&:``8GC&&Y-)&
4/(0&4/#0(0&F;9&,O%"+%,-(*&%*(50&A#)#&10#5?&2#*%10#&4/#7&0/-A&4/#&"%)B#04&57,%3(*&
)%,B#&(,&(0-4-@(*&=%"1#0&%,5&%)#&4/-1B/4&4-&2#&5#)(=#5&#$*"10(=#"7&>)-3&4#))#04)(%"&
@"%,40&Ed-"+3%,&#4&%"C?&:`N9c&6#7#)0?&:``Sc&[1B/#,&#4&%"C?&899HGC&&F;9&(0&*-,0(5#)#5&%&
2#44#)&2(-3%)+#)&4/%,&4/#&0/-)4#)&*/%(,O"#,B4/&/-3-"-B1#0?&2#*%10#&)%5(-*%)2-,&
%B#0&(,*)#%0#0&A(4/&(,*)#%0(,B&*/%(,&"#,B4/&51#&4-&%&/(B/#)&)#0(04%,*#&4-&
5#B)%5%4(-,&E<10*/?&89:9GC&&
8KK(2)#*!09'!?$1B2!
&
T,&4/(0&4/#0(0&T&(3@"#3#,4&4/(0&*-3@-1,5O0@#*(>(*&(0-4-@(*&%,%"70(0&%@@)-%*/&
10(,B&@"%,4&"#%>&A%$#0&4-&%,%"7J#&3%)(,#&0#5(3#,40&>)-3&K'KD&'(4#&8;:&(,&4/#&M1">&
->&!5#,&EY(B1)#&8GC&&V,"(+#&4#))#04)(%"&)#*-)50?&A/(*/&4#,5&4-&2#&>)%B3#,4#5&%,5&
"(3(4#5&(,&4(3#&%,5&0@%*#?&3%)(,#&0#5(3#,40&@)-=(5#&%&*-,4(,1-10&)#*-)5&4/%4&(0&
0@%4(%""7&%,5&4#3@-)%""7&#$4#,0(=#C&&K1#&4-&4/#&"-*%4(-,&->&'(4#&8;:?&5-A,A(,5&>)-3&
,-)4/#%04&!>)(*%?&A(,52"-A,&@"%,4&"#%>&A%$#0&@)-=(5#&%&*-,4(,1-10&)#*-)5&->&
=#B#4%4(-,&>-)&4/#&)#B(-,C&&D)#=(-10&0415(#0&10(,B&4/#&0%3#&3#4/-5-"-B(*%"&%@@)-%*/&
&
U&
EY#%+(,0&#4&%"C?&899UG?&/%=#&>-1,5&#=(5#,*#&>-)&0(B,(>(*%,4&#,=()-,3#,4%"&
>"1*41%4(-,0&%4&4/(0&0(4#C&&!&"-,B#)&)#*-)5?&0@%,,(,B&:8&3(""(-,&7#%)0?&/%0&%"0-&2##,&
@)-51*#5&>-)&4/(0&0(4#&EY#%+(,0&#4&%"C?&89:;GC&&[#)#&T&@)#0#,4&*%)2-,&(0-4-@(*&5%4%&>-)&
'(4#&8;:?&0@%,,(,B&;CN&4-&HC8&6%?&4/%4&)#@)#0#,40&%&03%""&@-)4(-,&->&4/#&"-,B#)&:8&
3(""(-,&7#%)&)#*-)5C&&./(0&)#*-)5&#,%2"#5&3#&4-&%,%"7J#&4/#&>"1*41%4(-,0&2#4A##,&F;&
%,5&FH&5-3(,%4#5&"%,50*%@#0&51)(,B&%,&#$*(4(,B&@#)(-5&->&/13%,&#=-"14(-,&A/#)#&
+#7&#=-"14(-,%)7&>#%41)#0?&01*/&%0&2(@#5%"(03?&A#)#&2#B(,,(,B&4-&5#=#"-@C&&&T&%"0-&
*-3@%)#&4/(0&3%)(,#&)#*-)5&A(4/&@#5-B#,(*&*%)2-,%4#&5%4%&4-&>1)4/#)&#$%3(,#&4/#&
#,=()-,3#,4%"&0/(>40&-20#)=#5&(,&4/#&3%)(,#&)#*-)5C&&&
EEEF!D)0'!@1G%0)1#
H)*,$'!L3&&8f&_-*%4(-,&->&K'KD&'(4#&8;:&(,&)#"%4(-,&4-&@)#0#,4O5%7&=#B#4%4(-,&J-,#0&%,5&%=#)%B#&
A(,5&0@##50&51)(,B&4/#&0-14/A#04&3-,0--,&E6%7Og1"7GC&&Mf&_-*%4(-,&->&+#7&/-3(,(,&>-00("&0(4#0C&&
EY(B1)#&>)-3&Y#%+(,0&#4&%"C?&899UCG&
&
./#&3%)(,#&0#5(3#,4&*-)#&A%0&-24%(,#5&>)-3&K##@&'#%&K)(""(,B&D)-L#*4&
EK'KDG&'(4#&8;:?&"-*%4#5&(,&4/#&*#,4#)&->&4/#&M1">&->&!5#,&E::CN`eW?&HNC8Ueb?&8:8U&3&
A%4#)&5#@4/?&K'KD&_#B&8HG&EY(B1)#&8!GC&&'(4#&8;:&(0&4/#&*"-0#04&D"(-OD"#(04-*#,#&
3%)(,#&0#5(3#,4%)7&)#*-)5&4-&4/#&b%04&!>)(*%,&P(>4&'704#3C&&./(0&0(4#&*%@41)#0&@"%,4&
&
Q&
"#%>&A%$#0&4/%4&%)#&4)%,0@-)4#5&27&A(,5&%,5&5#@-0(4#5&(,&3%)(,#&0#5(3#,40?&4/10&
@)-=(5(,B&%&*-,4(,1-10&4#))#04)(%"&%)*/(=#&->&,-)4/#%04&!>)(*%&EY#%+(,0&#4&%"C?&89:;GC&&
K1#&4-&4/#&@)-$(3(47&->&'(4#&8;:&4-&3%,7&+,-A,&/-3(,(,&>-00("&0(4#0?&-14"(,#5&(,&
Y(B1)#&8I?&(4&(0&#$@#*4#5&4/%4&4/#&0(4#&A(""&*%@41)#&4/#&#,=()-,3#,4%"&=%)(%2("(47&
#$@#)(#,*#5&27&4/#0#&#%)"7&/13%,&0@#*(#0C&&&!
E>F!8*'!C17'(!
!"#$%&'#(
!"#$)%*+
./#&%B#&3-5#"&EY(B1)#&;G&
4544$
4544$
7544$
9544$
;544$
<544$
64544$
67544$
A%0&*-,04)1*4#5&10(,B&%,&
644544$
)#*-)5&EI)-A,&#4&%"C?&:`NUc&
I1+)7?&:`SHc&Y#%+(,0&#4&%"C?&899Sc&
,#-./$)0123+
1@5%4#5&4#@/)%&%,5&,%,->-00("&
Y#%+(,0&#4&%"C?&89:;c&P%>>(&#4&%"C?&
744544$
844544$
944544$
:44544$
899Qc&P-4/?&:`SHc&'%),%O
;44544$
^-L*(*+(&#4&%"C?&:`NUGC&&./#&
85=:$
6=:544$
!"#$)%*+
2#(,B&10#5&4-&5%4#&0%3@"#0&
95:4$
95=:$
:544$
:57:$
:5:4$
6>:544$
,#-./$)0123+
@-(,40?&A(4/&"(,#%)&(,4#)@-"%4(-,&
957:$
6<:544$
0#5(3#,40&*-))#0@-,5(,B&4-&;CNO
HC8&6%&*-,4%(,&;&%B#&*-,4)-"&
9544$
74:544$
76:544$
77:544$
78:544$
79:544$
2#4A##,&4/#0#&@-(,40C&&^/("#&4/#&
7::544$
*1))#,4&%B#&3-5#"&/%0&0#=#)%"&"%)B#&B%@0&
2#4A##,&%B#&*-,4)-"&@-(,40?&4/#&
H)*,$'!<3&&!B#&3-5#"&>-)&K'KD&'(4#&8;:&2%0#5&
-,&"(,#%)&(,4#)@-"%4(-,&2#4A##,&5%4#5&4#@/)%&
%,5&,%,->-00("0&E5#4%("#5&(,&.%2"#&:GC&
0#5(3#,4%4(-,&)%4#0&%@@#%)&4-&/%=#&
2##,&>%()"7&*-,04%,4?&@)-=(5(,B&*-,>(5#,*#&>-)&4/(0&3-5#"C&&&&&
&
S&
&
.#@/)%&A%0&10#5&%0&4/#&@)(3%)7&3#%,0&->&5%4(,B&4/(0&*-)#?&2#*%10#&'(4#&8;:&
(0&,#%)&3%,7&+,-A,&b%04&!>)(*%,&#)1@4(-,&0(4#0&51)(,B&4/#&D"(-*#,#&%,5&D"#(04-*#,#C&&
.#@/)%&>)-3&#$@"-0(=#?&/(B/O=-"13#&#)1@4(-,0&*%,&2#&4)%,0@-)4#5&27&4/#&
4)-@-0@/#)#&%,5&04)%4-0@/#)#&1@&4-&8S99&+3&>)-3&4/#&0-1)*#C&&./#0#&4#@/)%&*%,&2#&
)#*-=#)#5&%,5&B#-*/#3(*%""7&*-))#"%4#5&4-&+,-A,&#)1@4(-,0&4-&@)-=(5#&5()#*4&
*/)-,-04)%4(B)%@/(*&4(#0&(,5#@#,5#,4&->&5%4(,B&1,*#)4%(,4(#0&EY#%+(,0&#4&%"C?&899SGC&&&
>F!C'0917/!
!
T,&4/(0&4/#0(0&T&@)#0#,4&%&@"%,4&"#%>&
A%$&)#*-)5&->&=#B#4%4(-,&*/%,B#&>)-3&
3%)(,#&0#5(3#,40&*-""#*4#5&%4&K'KD&EK##@&
'#%&K)(""(,B&D)-L#*4G&'(4#&8;:&(,&4/#&M1">&->&
!5#,&EY(B1)#&8GC&&Y-)&4/(0&@)-L#*4&3-"#*1"%)O
"#=#"&2(-3%)+#)&(0-4-@(*&5%4%&A%0&3#%01)#5&
10(,B&@"%,4&"#%>&A%$#0&(,&4/#&>-)3&->&>%447&
%*(5&3#4/7"&#04#)0&EY!6b0GC&&Y%447&%*(50?&
0@#*(>(*%""7&,O%"+%,-(*&%*(50?&A#)#&(0-"%4#5&
>)-3&4/#&21"+&0#5(3#,4&0%3@"#&%,5&%,%"7J#5&
10(,B&4/#&*/#3(*%"&@)-*#51)#&-14"(,#5&(,&
Y(B1)#&H&%,5&5#0*)(2#5&2#"-AC&&&&&&&
6%)(,#&0#5(3#,4&*-)#0&A#)#&
*-,4(,1-10"7&0%3@"#5&%,5&(,4#B)%4#5&#=#)7&
:9&*3?&*-))#0@-,5(,B&4-&h;?999&7#%)0C&&Y-)&
4/(0&@)-L#*4?&h:99&0%3@"#0&A#)#&%,%"7J#5&
&
H)*,$'!N3&&Y"-A&*/%)4&5#4%("(,B&4/#&
*/#3(*%"&%,5&%,%"74(*%"&3#4/-50&
#$#*14#5&4-&)#4)(#=#&!:;F&=%"1#0&>)-3&
3%)(,#&0#5(3#,40&
N&
*-))#0@-,5(,B&4-&5#@4/0&->&:NUCUO:``CH&320>?&%,5&%B#0&->&;CNNQOHC:U8&6%C&&&&
./)-1B/-14&4/#&#$@#)(3#,4%"&@)-*#51)#&0%3@"#0&A#)#&04-)#5&(,&4/#&>)##J#)&%4&&
O89eFC&&&
'%3@"#0&A#)#&>)##J#O5)(#5?&10(,B&%&d()4(0&8+&1,(4?&%,5&/-3-B#,(J#5&A(4/&%&
3-)4%)&%,5&@#04"#C&&./#&"(@(5&@-)4(-,&A%0&4/#,&#$4)%*4#5&>)-3&#%*/&0%3@"#&10(,B&%,&
!**#"#)%4#5&'-"=#,4&b$4)%*4(-,&0704#3&E!'b&;U9i?&K(-,#$GC&&b%*/&!'b&*%@01"#&A%0&
@%*+#5&A(4/&:9O:U&B&->&5)7?&/-3-B#,(J#5&0%3@"#&%,5&>(""#5&A(4/&*-32104#5&0%,5C&&
'%3@"#0&A#)#&#$4)%*4#5&A(4/&`9j&5(*/"-)-3#4/%,#&4-&:9j&3#4/%,-"&%4&/(B/&
4#3@#)%41)#&E:99eFG&%,5&@)#001)#&E:U99&@0(GC&&_(@(5&#$4)%*40&A#)#&4/#,&0#@%)%4#5&
27&*-"13,&*/)-3%4-B)%@/7&E*-"13,f&U&*3&$&H9&33&D%04#1)&@(@#44#?&W[8&0#@)%&21"+&
@%*+(,B?&Q9kG&#"14(,B&A(4/&8f:&5(*/"-)-3#4/%,#&A(4/&(0-@)-@%,-"?&>-""-A#5&27&Hj&
>-)3(*&%*(5&(,&5(#4/7"&#4/#)C&&b"14(-,0&7(#"5#5&,#14)%"&%,5&%*(5&>)%*4(-,0?&
)#0@#*4(=#"7C&&&
!*(5&>)%*4(-,0&A#)#&4/#,&3#4/7"%4#5&4-&*-))#0@-,5(,B&Y!6b'&A(4/&HCSU&3_&
3#4/%,-"&El:;F3#4/%,-"&m&OHSC8U&n&9C:9?&,m;G&%,5&9C8U&3_&/75)-*/"-)(*&%*(5&EH9j&
*-,*#,4)%4#5G&%4&S9eF&>-)&:8&/-1)0C&&[75)-"7J#5&@)-51*40&A#)#&4/#,&4)#%4#5&A(4/&&
:&3_&3(""(o&A%4#)C&&_(@(50&A#)#&@%)4(4(-,#5&(,4-&4/#&"#00&5#,0#&/#$%,#&@-)4(-,?&
A/(*/&A%0&*-""#*4#5&%,5&5)(#5&27&@%00(,B&(4&4/)-1B/&%&*-"13,&->&%,/75)-10&0-5(13&
01">%4#C&&_(@(50&A#)#&>1)4/#)&@1)(>(#5&27&*-"13,&*/)-3%4-B)%@/7&E*-"13,f&U&*3&$&H9&
33&D%04#1)&@(@#44#?&Uj&A%4#)O5#%*4(=%4#5&0("(*%OB#"?&:99O899&3#0/G&#"14(,B&A(4/&
/#$%,#&%,5&5(*/"-)-3#4/%,#C&&&b"14(-,0&7(#"5#5&,-,O@-"%)&>)%*4(-,0&%,5&@-"%)&Y!6b&
>)%*4(-,0&)#0@#*4(=#"7C&&&
&
`&
!:;F&5%4%&%)#&)#@-)4#5&>-)&4/#&F;9&,O%"+%,-(*&%*(5&@-)4(-,&->&4/#&Y!6b0C&&F;9&
A%0&X1%,4(>(#5&10(,B&%&MFp6'&EM%0&F/)-3%4-B)%@/7O6%00&'@#*4)-3#4)7G?&4-&
5#4#)3(,#&4/#&%@@)-@)(%4#&%3-1,4&4-&(,L#*4&(,4-&4/#&MFOTP6'&EM%0&
F/)-3%4-B)%@/7OT0-4-@#&P%4(-&6%00&'@#*4)-3#4)7G&>-)&(0-4-@(*&%,%"70(0&EY#%+(,0&#4&
%"C?&899UGC&&'%3@"#0&A#)#&%,%"7J#5&27&[%,,%/&_(557&10(,B&4/#&3#4/-50&5#0*)(2#5&(,&
Y#%+(,0&#4&%"C?&89:;&%,5&Y#%+(,0?&89:;C&&T&@)-*#00#5&4/#&)#01"40&->&4/#0#&MFOTP6'&
%,%"70#0&>-)&4/(0&4/#0(0&10(,B&T0-5%4&0->4A%)#C&&
>EF!O'/,(0/!
!
]>&4/#&h:99&0%3@"#0&4/%4&A#)#&@)-*#00#5&10(,B&4/#&*/#3(*%"&@)-*#51)#&
-14"(,#5&(,&Y(B1)#&U?&NQ&0%3@"#0&@)-51*#5&)#"(%2"#&!:;F&5%4%C&&./#0#&0%3@"#0&/%5&%&
3#%,&=%"1#&->&O8SCUR?&%&04%,5%)5&5#=(%4(-,&->&:C:8?&%,5&%&)#@"(*%4#&@)#*(0(-,&->&
9C:RC&&F;9&@#%+0&A#)#&5#4#)3(,#5&)#"(%2"#&2%0#5&-,&)#B1"%)&@#%+&0/%@#?&%20#,*#&->&
@#%+&*-#"14(-,?&%,5&%3@"(415#&B)#%4#)&4/%,&U99&3dC&
&
&!:;F&=%"1#0&>)-3&4/(0&@-)4(-,&->&4/#&0#5(3#,4&*-)#&)%,B#&2#4A##,&O8`CNR&
%,5&O8HCNRC&&./#&3-04&,#B%4(=#&=%"1#0&%)#&)#@-)4#5&%4&;C`&%,5&HC:&6%?&%,5&4/#&3-04&
@-0(4(=#&=%"1#0&%4&HC9&6%C&&T0-4-@(*&=%"1#0&=%)(#5&01204%,4(%""7?&/-A#=#)?&)#01"40&
(,5(*%4#&%&0/(>4&4-A%)50&"#00&5#@"#4#5&!:;F&=%"1#0?&%,5&%&)#41),&4-&3-)#&5#@"#4#5&
=%"1#0C&&./#)#>-)#&4/#&)#01"40&A(""&2#&*-,0(5#)#5&(,&8&@/%0#0f&%&0/(>4&4-A%)50&"#00&
5#@"#4#5&=%"1#0&EHC8OHC9&6%G&%,5&%&)#41),&4-&3-)#&5#@"#4#5&=%"1#0&EHC9O;CN&6%GC&&&
&
&
&
:9&
&&&&&&&&&&&&&
H)*,$'!P3&&6%)(,#&)#*-)5&->&=#B#4%4(-,&>-)&,-)4/#%04&!>)(*%&0@%,,(,B&h;CNOHC8&6%C&&!))-A0&(,5(*%4#&F;&
%,5&FH&#,5&3#32#)0C&&F-)#&)#*-=#)7&(0&)#@)#0#,4#5&27&4/#&2%)&%5L%*#,4&4-&4/#&7O%$(0C&&K1#&4-&@--)&
*-)#&)#*-=#)7&51)(,B&4/#&5)(""(,B&@)-*#00&0#5(3#,40&A#)#&,-4&)#*-=#)#5&>-)&HC9U&4-&HC:;&6%C&
T,&4/#&)%,B#&HC8OHC9&6%?&4/#&3(,(313&!:;F&=%"1#&(0&O8`CHR&%,5&4/#&
3%$(313&=%"1#&(0&O8HCNRC&&K1)(,B&4/(0&@#)(-5&4/#)#&(0&%&HCQR&0/(>4&4-A%)50&"#00&
5#@"#4#5&=%"1#0C&&./#)#&(0&%&B%@&(,&4/#&5%4%&0@%,,(,B&hHC9UOHC:;&6%&51#&4-&@--)&
0%3@"#&)#*-=#)7&51)(,B&4/#&5)(""(,B&@)-*#00?&*/#3(*%"&@)#@%)%4(-,?&-)&%,%"74(*%"&
@)-*#51)#C&&&
&
T,&4/#&)%,B#&HC9O;CN&6%?&%&)#41),&4-A%)50&3-)#&5#@"#4#5&!:;F&=%"1#0&(0&
-20#)=#5C&&K1)(,B&4/(0&@#)(-5&4/#)#&(0&%&3%$(313&(0-4-@(*&=%"1#&->&O8HCNR&%,5&%&
3(,(313&->&O8`CHR?&3%)+(,B&%&O;CSR&0/(>4C&&&&&
&
::&
>EEF!I)/G,//)1#!
E#0'$K$'0%0)1#!15!=%$41#!E/101K'!O'G1$7!
!
./#&5%4%&(,&4/(0&4/#0(0&)#=#%"0&0(B,(>(*%,4&#,=()-,3#,4%"&=%)(%2("(47&2#4A##,&
;CNOHC8&6%?&(,5(*%4(,B&4/%4&,-)4/#%04&!>)(*%,&#,=()-,3#,4%"&=%)(%2("(47&A%0&
01204%,4(%"&@)(-)&4-&4/#&W-)4/#),&[#3(0@/#)#&B"%*(%4(-,&E_(0(#*+(&%,5&P%73-?&899UGC&&
P#01"40&(,5(*%4#&%&A(5#&)%,B#&->&*%)2-,&(0-4-@(*&=%"1#0&51)(,B&4/(0&4(3#&@#)(-5?&
(,5(*%4(,B&%&"%,50*%@#&3($#5&A(4/&>-)#04&%,5&B)%00"%,5C&&d%)(%2("(47&A%0&01204%,4(%"?&
)#=#%"(,B&%&@#)(-5&->&FH&#$@%,0(-,&%,5&%&0120#X1#,4&)#41),&4-&F;&5-3(,%,*#C&&./#)#&
(0&%&HCQR&0/(>4&4-A%)5&3-)#&#,)(*/#5&!:;F&=%"1#0&EHC8OHC9&6%G&(,5(*%4(,B&%&0/(>4&
>)-3&F;&4-&FH&*-,5(4(-,0&E>-)#04&4-&B)%00"%,5GC&&6-)#&#,)(*/#5&*%)2-,&(0-4-@(*&=%"1#0&
(,5(*%4#&A%4#)&04)#00&%,5&%&3-)#&-@#,&#,=()-,3#,4?&A/(*/&%)#&*-,5(4(-,0&4/%4&>%=-)&
FH&#$@%,0(-,&%,5&*%10#&F;&@"%,40&4-&5(0@"%7&3-)#&#,)(*/#5&!:;F&=%"1#0&EF#)"(,B&#4&%"C?&
:``SGC&&./#0#&*-,5(4(-,0&%@@#%)&4-&>"1*41%4#&4/)-1B/-14&4/(0&)#*-)5?&%,5&2#4A##,&
HC9&4-&;CN&6%&4/#)#&(0&%&;CSR&0/(>4&4-&3-)#&5#@"#4#5&=%"1#0?&(,5(*%4(,B&%&)#41),&4-&F;&
5-3(,%,*#C&
&
./(0&5%4%&(0&@%)4&->&%&"-,B#)?&:8&3(""(-,&7#%)&)#*-)5?&%,5&(0&#32#55#5&A(4/(,&
4/#&5%4%0#4&@)#@%)#5&27?&Y#%+(,0&#4&%"C?&89:;?&10(,B&0(3("%)&3#4/-50&EY(B1)#&QGC&&./(0&
"-,B#)&)#*-)5&@)-=(5#0&#=(5#,*#&>-)&4A-&@#)(-50&->&FH&#$@%,0(-,?&::O`&6%&%,5&HC;O
:CH&6%C&&./#&5%4%&>-)&4/(0&4/#0(0&>%""0&A(4/(,&4/(0&0#*-,5&@#)(-5&->&FH&#$@%,0(-,C&&]=#)&
4/#&@%04&:8&3(""(-,&7#%)0&4/#)#&(0&%&"-,B&4#)3&4)#,5&4-A%)5&"#00&5#@"#4#5?&3-)#&FH&
=%"1#0?&/-A#=#)?&4/#&@-)4(-,&->&4/(0&)#*-)5&*-=#)#5&(,&4/(0&4/#0(0&=%)(#0&>)-3&4/(0&
-=#)%""&4)#,5C&&K1)(,B&4/(0&@#)(-5?&%&FH&#$@%,0(-,&(0&-20#)=#5?&/-A#=#)?&(4&(0&>-""-A#5&
27&%&)#41),&4-&F;&5-3(,%,*#C&&K1)(,B&4/(0&0120#X1#,4&FH&*-,4)%*4(-,?&!:;F&=%"1#0&
&
:8&
)#%*/&0-3#&->&4/#&3-04&5#@"#4#5&=%"1#0&->&4/#&#,4()#&:8&3(""(-,&7#%)&)#*-)5C&&./(0&
)#41),&4-&F;&=%"1#0&5#0@(4#&4/#&-=#)%""&4)#,5&4-A%)50&FH&(0&%&0(B,(>(*%,4&>(,5?&(3@"7(,B&
4/%4&#,=()-,3#,4%"&*/%,B#&(,&,-)4/#%04&!>)(*%&A%0&,-4&%&"(,#%)&@)-*#00?&214&)%4/#)&
-**1))#5&%0&%&0#)(#0&->&-0*(""%4(,B&#=#,40&4/%4&4)#,5#5&4-A%)5&4/#&-=#)%""&FH&
5-3(,%4#5&"%,50*%@#&-20#)=#5&4-5%7C&&&&&
=1.K%$)/1#!01!D1)(!=%$41#%0'!O'G1$7!
!
H)*,$'!Q3&&F%)2-,&(0-4-@(*&#=(5#,*#&>-)&,-)4/#%04&!>)(*%,&=#B#4%4(-,C&&!:;F&=%"1#0&%)#&)#@-)4#5&
>-)&@"%,4&"#%>&A%$#0&E"#>4G&%,5&@#5-B#,(*&*%)2-,&E)(B/4GC&&!B#&*-,4)-"&@-(,40&>-)&K'KD&8;:&%)#&
0/-A,&%0&5(%3-,50&%5L%*#,4&4-&4/#&7O%$(0C&&EY(B1)#&>)-3&Y#%+(,0&#4&%"C?&89:;G&&
!
!,&#*-0704#3?&01*/&%0&,-)4/#%04&!>)(*%?&4/%4&(0&*-3@)(0#5&->&%&3($41)#&->&F;&
%,5&FH&@"%,40&/%0&%&5(04(,*4&*%)2-,&(0-4-@(*&0(B,%41)#&4/%4&(0&)#*-)5#5&(,&2-4/&4/#&
%2-=#&B)-1,5&%,5&2#"-A&B)-1,5&2(-3%00C&&g104&%0&4/(0&*%)2-,&(0-4-@(*&0(B,%41)#&(0&
&
:;&
)#*-)5#5&(,&@"%,4&"#%>&A%$#0?&(4&(0&%"0-&)#*-)5#5&(,&0-("&*%)2-,%4#0&q8U&*3&2#"-A&4/#&
01)>%*#&EF#)"(,B?&:`NH&%,5&F#)"(,B&#4&%"C?&89::GC&&D"%,40&)#0@()#&*%)2-,&%0&F]8&(,&4/#&
0-("?&%,5&4/(0&)#0@()#5&F]8&/%0&%&0(3("%)&*%)2-,&(0-4-@(*&=%"1#&%0&4/#&%2-=#&B)-1,5&
*%)2-,C&&I#*%10#&4/#&*%)2-,&(0-4-@(*&*-3@-0(4(-,&->&0-("&*%)2-,%4#0&(0&*-,4)-""#5&27&
4/#&*-3@-0(4(-,&->&4/#&)#0@()#5&F]8?&0-("&*%)2-,%4#0&*%,&0#)=#&%0&(3@-)4%,4&
@%"#-*"(3%4#&%,5&@%"#-=#B#4%4(-,&(,5(*%4-)0&EF#)"(,B&#4&%"C?&89::?&]5#&#4&%"C?&:`N9?&
%,5&K#(,#0&#4&%"C?&:`N9GC&&!&,#4&(0-4-@(*&#,)(*/3#,4&->&:HO:QR&(0&-20#)=#5&(,&0-("&
*%)2-,%4#0?&)#"%4(=#&4-&0-("O)#0@()#5&F]8?&51#&4-&@)-*#00#0&01*/&%0&5(>>10(-,?&(0-4-@#&
>)%*4(-,%4(-,?&%,5&%43-0@/#)(*&3($(,B&EF#)"(,B&#4&%"C?&89::GC&&&&
&
!&*-3@%)(0-,&->&4/#&3%)(,#&%)*/(=#&->&@"%,4&"#%>&A%$#0&%,5&4/#&"-*%"&)#*-)5&
>)-3&4#))#04)(%"&0(4#0&@)-=(5#0&>1)4/#)&*-,4#$4&>-)&4/#&3%)(,#&5%4%&EY(B1)#&QGC&&
F%)2-,&(0-4-@(*&%,%"70#0&->&0-("&*%)2-,%4#0&*-3@("#5&>)-3&0#=#)%"&0415(#0&01@@-)4&
4/#&-=#)%""&4)#,5&5(0@"%7#5&27&4/#&@"%,4&"#%>&A%$#0c&(0-4-@(*&#,)(*/3#,4&%,5&FH&
#$@%,0(-,&>)-3&:8&6%&4-&@)#0#,4?&A(4/&0(B,(>(*%,4&*"(3%4#&=%)(%2("(47&2#B(,,(,B&%0&
#%)"7&%0&HC;&6%&EY#%+(,0&#4&%"C?&89:;GC&&./#&0-("&*%)2-,%4#&5%4%&%"0-&01@@-)40&4/#&
>(,5(,B0&->&4/(0&4/#0(0?&5(0@"%7(,B&%&FH&#$@%,0(-,&%,5&0120#X1#,4&*-,4)%*4(-,&>)-3&
;CN&4-&HC8&6%C&&&
&
D)(-)&4-&h;CN&6%&*%)2-,%4#&5%4%&(0&>%()"7&5(0@#)0#5&%,5&)%,B#0&>)-3&hO:9O&&
OURC&&!4&h;C`&6%&%&*"104#)&->&5%4%&@-(,40&(,5(*%4#&%&4)#,5&4-A%)50&3-)#&#,)(*/#5&
=%"1#0?&(,5(*%4(=#&->&%&FH&#$@%,0(-,?&A(4/&!:;F&=%"1#0&@#%+(,B&%)-1,5&9RC&&./(0&@#%+&
(,&*%)2-,&(0-4-@(*&=%"1#0&*-(,*(5#0&,(*#"7&A(4/&4/#&FH&#$@%,0(-,&*%@41)#5&(,&4/#&
@"%,4&"#%>&A%$&5%4%C&&!&)#41),&4-&3-)#&5#@"#4#5?&F;?&=%"1#0&27&;CN&3%&(0&%"0-&)#*-)5#5&
(,&4/#&0-("&*%)2-,%4#0C&&!:;F&=%"1#0&(,&0-("&*%)2-,%4#0&)#41),&4-&"#=#"0&0(3("%)&4-&4/#&
&
:H&
@#)(-5&2#>-)#&FH&#$@%,0(-,&EhONRG?&0-3#&->&4/#&3-04&5#@"#4#5&=%"1#0&->&4/#&:8&
3(""(-,&7#%)&)#*-)5?&5#0@(4#&-=#)%""&FH&#$@%,0(-,C&&'-("&*%)2-,%4#&5%4%&%"0-&0/-A0&%&
0(3("%)&)%,B#&(,&(0-4-@(*&=%"1#0&51)(,B&4/(0&@#)(-5&EhNRG?&*-3@%)#5&4-&4/#&@"%,4&
"#%>&A%$&)#*-)5&EURG?&@)-=(5(,B&>1)4/#)&*-,>(5#,*#&(,&4/#&5%4%C&&&&
&&&&&&!
!
>EEEF!=1#G(,/)1#/!
!
6-"#*1"%)O"#=#"&2(-3%)+#)&(0-4-@(*&5%4%&@)-=(5#0&=%"1%2"#&(,0(B/40&(,4-&4/#&
,%41)#&->&,-)4/#%04&!>)(*%,&*"(3%4#&=%)(%2("(47C&&T,&4/(0&4/#0(0&T&@)#0#,4&*%)2-,&
(0-4-@(*&5%4%&>-)&4/#&@-)4(-,&->&4/#&0#5(3#,4&*-)#&*-))#0@-,5(,B&4-&;CNOHC8&6%C&&./(0&
)#*-)5&(0&*-,0(04#,4&A(4/&0-("&*%)2-,%4#&5%4%&%,5&(0&@%)4&->&%&"%)B#)&#>>-)4&4-&
)#*-,04)1*4&,-)4/#%04&!>)(*%,&*"(3%4#&-=#)&4/#&@%04&:8&3(""(-,&7#%)0&EY#%+(,0&#4&%"C?&
89:;GC&&!:;F&=%"1#0&->&"#%>&A%$&2(-3%)+#)0&(,5(*%4#&%&/(B/"7&=%)(%2"#&"%,50*%@#&
*-3@-0#5&->&%&3($41)#&->&>-)#04&%,5&B)%00"%,5C&&T0-4-@(*&5%4%&%"0-&(,5(*%4#5&%,&
-@#,(,B&(,&4/#&"%,50*%@#&2#4A##,&HC8&%,5&HC9&6%?&>-""-A#5&27&%&)#41),&4-&F;&
*-,5(4(-,0&>)-3&HC9&4-&;CN&6%C&&./(0&FH&#$@%,0(-,&%,5&0120#X1#,4&*-,4)%*4(-,&
5#=(%4#0&>)-3&4/#&-=#)%""&4)#,5&4-A%)50&FH&5-3(,%4#5&"%,50*%@#0&-=#)&4/#&@%04&:8&
3(""(-,&7#%)0C&&./#&)#01"40&->&4/(0&4/#0(0&>1)4/#)&01@@-)4&4/#&,-4(-,&4/%4&,-)4/#%04&
!>)(*%,&*"(3%4#&A%0&/(B/"7&=%)(%2"#&%,5&A%0&@1,*41%4#5&27&%2)1@4&0/(>40&(,&*"(3%4#?&
)%4/#)&4/%,&-**1))(,B&%0&%&B)%51%"?&1,(5()#*4(-,%"&@)-*#00C!
&
8GR#1J('7*'.'#0/!
!
./(0&4/#0(0&10#5&0%3@"#0&@)-=(5#5&27&4/#&T,4#B)%4#5&]*#%,&K)(""(,B&D)-B)%3C&&
Y1,5(,B&A%0&%A%)5#5&4-&!'&27&^('b&E^-3#,&(,&'*(#,*#&%,5&b,B(,##)(,BG&%,5&4/#&
V,(=#)0(47&->&'-14/#),&F%"(>-),(%&D)-=-04a0&P#0#%)*/&Y1,5C&&T&A-1"5&"(+#&4-&4/%,+&
&
:U&
D)->#00-)&'%)%/&Y#%+(,0?&6(B1#"&P(,*-,?&%,5&[%,,%/&_(557&>-)&4/#()&A-,5#)>1"&
3#,4-)0/(@&-=#)&4/#&@%04&4/)##&7#%)0C&&&
O'5'$'#G'/!
Bonnefille, R. (2010). Cenozoic vegetation, climate changes and hominid evolution in
tropical Africa. Global and Planetary Change. Vol. 72, pp. 390-411.
Brown, F. H., Sarna-Wojcicki, A. M., Meyer, C. E., and Haileab, B., 1992, Correlation of
Pliocene and Pleistocene tephra layers between the Turkana Basin of East Africa
and the Gulf of Aden: Quaternary International, v. 13–14, p. 55–67.
Bukry, D., 1974, Coccolith zonation of cores from the western Indian Ocean and the Gulf
of Aden, Deep Sea Drilling Project Leg 24: Government Printing Office, 24, 995–
996.
Cerling, T.E., Levin, N.E., Passey, B.H. (2011). Stable Isotope Ecology in the OmoTurkana Basin. Evolutionary Anthropology, v. 20, p. 228-237.
Cerling, T.E., Harris, J.M., MacFadden, B.J., Leakey, M.G., Quade, J., Eisenmann, V.,
and Erleringer, J.R. (1997). Global vegetation change through the
Miocene/Pliocene boundary. Nature, v. 389, p. 153-158.
Cerling, T.E. (1984). The stable isotopic composition of modern soil carbonate and its
relationship to climate. Earth and Planetary Science Letters, v. 71, p. 229-240.
Deines, P. (1980). The isotopic composition of reduced organic carbon in: Handbook of
Environmental Isotope Geochemistry, 1. The Terrestrial Environment, A.P. Fritz
and J.Ch. Fontes. Eds., pp.329-406, Elsevier, Amsterdam.
deMenocal, P. (2004). African climate change and faunal evolution during the PliocenePleistocene. Earth Planet. Sci. Lett. 220, 3-24.
Feakins, S.J., Brown, F. H., deMenocal, P.B. (2007). Plio-Pleistocene Microtephra in
DSDP Site 231, Gulf of Aden. Journal of African Earth Sciences, v. 48, p. 341352.
Feakins, S.J., deMenocal, P.B., Eglinton, T.I. (2005). Biomarker records of Late
Neogene Changes in East African Vegetation. Geology, v. 33, p. 977-980.
Feakins, S.J., Levin, N.E., Liddy, H.M., Sieracki, A., Eglinton, T.I., Bonnefille, R.
(2013). Northeast African vegetation change over 12 million years. Geology, p.
doi:10.1130/G33845.1.
&
:Q&
Feakins, S.J. (2013). Pollen-corrected leaf wax D/H reconstructions of northeast
African hydrological changes during the late Miocene. Palaeogeography,
Palaeoclimatology and Palaeoecology, p. doi:10.1016/j.palaeo.2013.01.004.
Freeman, K., Colarusso, L. (2001). Molecular and isotopic records of C4 grassland
expansion in the late Miocene. Geochimica et Cosmochimica, v. 65, p. 14391454.
Hughen, K., Eglinton, T., Xu, L., Makou, M. (2004). Abrupt tropical vegetation response
to rapid climate changes. Science, v. 304, p. 1955-1959.
Kusch, S., Rethemeyer, J., Schefuß, E., Mollenhauer, G. (2010). Controls on the age of
vascular plant biomarkers in Black Sea sediments. Geochimica et Cosmochimica,
v. 74, p.7031-7047.
Lisiecki, L.E., Raymo, M.E. (2005). A Pliocene-Pleistocene stack of 57 globally
distributed benthic !18O records, Paleoceanography, 20, PA 1003,
doi:10.1029/2004PA001071.
Maslin, M.A., Christensen, B. (2007). Tectonics, orbital forcing, global climate change,
and human evolution in Africa: introduction to the African paleoclimate special
volume. Journal of Human Evolution. v. 53, p. 443-464.
Meyers, P. (1997). Organic geochemical proxies for paleoceanographic, paleolimnologic,
and paleoclimatic processes. Organic Geochemistry, v. 27, p. 213-250.
Meyers, P., Eadie, B. (1993). Sources, degradation, and recycling of organic matter
associated with sinking particles in Lake Michigan. Organic Geochemistry, v. 20,
p. 47-56.
Ode, D.J., Tieszen, L.L., Lerman, J.C. (1980). The seasonal contribution of C3 and C4
plant species to primary production in a mixed prairie. Ecology, v. 61, p. 13041311.
O’Leary, M. (1981). Carbon isotope fractionation in plants. Phytochemistry, v. 20,
p. 553-567.
Pearson, A., Eglinton, T. (2000). The origin of n-alkanes in Santa Monica Basin surface
sediment: a model based on compound specific !14C and !13C data. Organic
Geochemistry, v. 31, p. 1103-1116.
&
:S&
Raffi, I., Backman, J., Fornaciari, E., Palike, H., Rio, D., Lourens, L., and Hilgen, F.,
2006, A review of calcareous nannofossil astrobiochronology encompassing the
past 25 million years: Quaternary Science Reviews, v. 25, no. 23-24, p. 3113–
3137.
Sarna-Wojcicki, A. M., Meyer, C. E., Roth, P. H., and Brown, F. H., 1985, Ages of tuff
beds at East African early hominid sites and sediments in the Gulf of Aden:
Nature, v. 313, p. 306–308.
Schefuss, E., Ratmeyer, V., Stuut, J.B.W., Jansen, J.H.F., Damste, J.S.S. (2003). Carbon
isotope analyses of n-alkanes in dust from the lower atmosphere over the central
eastern Atlantic. Geochimica et Cosmochimica, v. 67, p. 1757-1767.
Sponheimer, M., Loudon, J.E., Codron, D., Howells, M.E., Pruetz, J.D., Lee-Thorp, J.A.
(2006). Do “savanna” chimpanzees consume C4 resources? Journal of Human
Evolution. v. 51, p. 128-133.
Volkman, J., Johns, R., Gillan, F., Perry, G. (1980). Microbial lipids of an intertidal
sediment – 1. Fatty acids and hydrocarbons. Geochimica et Cosmochimica, v. 44,
p. 1133-1143.
Waller, S.S., lewis, J.K. (1979). Occurrence of C3 and C4 Photosynthetic Pathways in
North American Grasses. Journal of Range Management. v. 32, no. 1, p. 12-28.&
&
&
:N&
8KK'#7)B!8F!I%0%!S%4('/!
.%2"#&:f&!B#&6-5#"&>-)&K'KD&'(4#&8;:&
K%413&
K#@4/&E320>G&
!B#&E6%G&
!
!
K%4(,B&3#4/-5&
P#>#)#,*#&
!
!
H9!)p;`!)&
S'K9$%!
'("2-&.1>>&
<-+(0#"#(&.1>>&
IO.1"1&I-)&.1>>&
_-+-*/-4&.1>>&
_-3-B-"&.1>>&
A%$*1(1!S,55!
C1)0)!S,55!
;UC8U&
::NCQ9&
:Q9CN9&
:QNCSQ&
:S9CSU&
;TUFTV!
;WWF<P!
9CSU&n&9C98&
8CH;&n&9C9U&
;CH9&n&9C9;&
;CUN&n&9C9:&
;CQ9&
<FWV!
<FUL!Y!VFVN!
(,4#)@-"%4(-,&
H9!)p;`!)&
-)2(4%""7&41,#5&
(,4#)@-"%4(-,&
)#0'$K1(%0)1#!
NV8$Z<U8$!
I&
I&
'^?&I&
Y&
I&
DAX!M!
DAX!M!
[%#151//)(/!
-=*"%+1>#%'*<?*
-=*1>#%'*<?*
!"#$%&'&(#)*#
4=*@.#"@.%&16.'*9?*
4=*/161(.'*<?*
4=*"%+/161(.'*9?*
4=*/161(.'*9?*
0=*6#+5%21;#8.'*<?*
&
:S:C`N&
:S:C`N&
LPWF<<!
8QQCUU&
HSUC:9&
HNQC89&
U8:CNU&
U8;C8N&
&
;CQU&n&9C9:&
;CQU&n&9C9:&
PF<P!Y!VFVL!
UCUH&n&9C9H&
`CQ`&n&9C:8&
`CQ`&n&9C::&
:9CUU&n&9C:8&
::C98&n&9C:9&
&
-)2(4%""7&41,#5&
-)2(4%""7&41,#5&
1$4)0%((2!0,#'7!
-)2(4%""7&41,#5&
-)2(4%""7&41,#5&
-)2(4%""7&41,#5&
-)2(4%""7&41,#5&
-)2(4%""7&41,#5&
&
P-?&I1?&P&
P-?&I1?&P&
O1X!M,X!O!
P-?&I1?&P&
P-?&I1?&P&
P-?&I1?&P&
P-?&I1?&P&
P-?&I1?&P&
!
W-4#f&I-"5#5&)-A0&(,5(*%4#&%B#&*-))#"%4(-,&@-(,40&*-=#)#5&(,&4/#&0@%,&->&4/(0&4/#0(0C&
P#>#)#,*#0f&I&m&I)-A,&#4&%"C?&:``8c&'^&m&'%),%O^-L*(*+(&#4&%"C?&:`NUc&Y&m&Y#%+(,0&&
#4&%"C?&899Sc&P-&m&P-4/?&:`SHc&I1&m&I1+)7?&:`SHc&P&m&P%>>(&#4&%"C?&899Qc&%,5&&
)#>#)#,*#0&4/#)#(,C&_]&m&"-A#04&-**1))#,*#c&[]&m&/(B/#04&-**1))#,*#C&&
E.%2"#&>)-3&Y#%+(,0&#4&%"C?&89:;G&
&
:`&
.%2"#&8f&_#%>&^%$&F%)2-,&T0-4-@(*&F-3@-0(4(-,&
&
K#@4/&E320>G&
!B#&E6%G&
l:;F&F;9&ERG&
:NUCU&
:NUCQ&
:NUCS&
:NUCN&
:NUC`&
:NQ&
:NQC:&
:NQC;&
:NQCU&
:NQCQ&
:NQCS&
:NQCS&
:NQCN&
:NQC`&
:NN&
:NN&
:NNC:&
:NNC8&
:NNC;&
:NNCH&
:NNCU&
:NNCQ&
:NNCS&
:NNC`&
:N`&
:N`C8&
:N`C;&
:N`CH&
:N`CU&
:N`CU&
:N`CQ&
:N`CQ&
:N`CS&
:N`CN&
:N`C`&
:`9&
:`9C:&
:`9C8&
:`9C;&
:`9CU&
:`9CQ&
:`9CQ&
:`9CS&
:`9CN&
:`9C`&
;CNNQ&
;CNNS&
;CNNN&
;CN`9&
;CN`:&
;CN`;&
;CN`H&
;CN`S&
;CN``&
;C`9:&
;C`98&
;C`98&
;C`9H&
;C`9U&
;C`89&
;C`89&
;C`88&
;C`8;&
;C`8H&
;C`8Q&
;C`;9&
;C`;8&
;C`;H&
;C`;N&
;C`H9&
;C`HH&
;C`HQ&
;C`HN&
;C`U9&
;C`U9&
;C`U8&
;C`U8&
;C`UH&
;C`US&
;C`U`&
;C`Q:&
;C`Q;&
;C`QU&
;C`QS&
;C`S:&
;C`S;&
;C`S;&
;C`SU&
;C`SS&
;C`S`&
O8NCH&
O8NC`&
O8NCN&
O8NCQ&
O8NCS&
O8SCS&
O8NCU&
O8NCS&
O8SC`&
O8`C:&
O8NC:&
O8NC:&
O8SCS&
O8NC9&
O8SCS&
O8SCS&
O8QC`&
O8SC;&
O8SCN&
O8`C;&
O8SCN&
O8NC9&
O8NC:&
O8NC;&
O8SC`&
O8SCN&
O8NCH&
O8SCQ&
O8SC8&
O8SC8&
O8`C8&
O8`C8&
O8`CN&
O8`CQ&
O8NCH&
O8NCU&
O8QC`&
O8NC:&
O8SCN&
O8SCH&
O8QC:&
O8QC:&
O8SCN&
O8NCH&
O8`C8&
89&
&
K#@4/&E320>G&
!B#&E6%G&
l:;F&F;9&ERG&
:`:&
:`:C:&
:`:C8&
:`:CH&
:`:CH&
:`:CU&
:`:CU&
:`:CQ&
:`:CS&
:`:CS&
:`:CN&
:`8&
:`8C:&
:`8C8&
:`8C8&
:`8C;&
:`8CH&
:`8CU&
:`8CQ&
:`8CS&
:`8CS&
:`8CN&
:`8C`&
:`;&
:`;C:&
:`;C:&
:`;C;&
:`;CH&
:`;CU&
:`;CQ&
:`;CQ&
:`;CS&
:`;CN&
:`;C`&
:`H&
:`HC:&
:`HC:&
:`HC8&
:`HC;&
:`HCH&
:`HCU&
:`HCQ&
:`HCQ&
:`HCS&
:`HCN&
:`NC;&
:`NCH&
;C`N:&
;C`N;&
;C`NU&
;C`N`&
;C`N`&
;C``:&
;C``:&
;C``;&
;C``U&
;C``U&
;C``S&
HC99:&
HC99;&
HC99U&
HC99U&
HC99N&
HC9:9&
HC9:8&
HC9:H&
HC9:Q&
HC9:Q&
HC9:N&
HC989&
HC988&
HC98H&
HC98H&
HC98N&
HC9;9&
HC9;8&
HC9;H&
HC9;H&
HC9;Q&
HC9;N&
HC9H9&
HC9H8&
HC9HH&
HC9HH&
HC9HQ&
HC9HN&
HC9U9&
HC9U8&
HC9UH&
HC9UH&
HC9UQ&
HC9U`&
HC:;9&
HC:;8&
O8`C;&
O8SC;&
O8NCH&
O8QC:&
O8QC:&
O8QC:&
O8SC8&
O8QCU&
O8UC8&
O8UC8&
O8UCQ&
O8UCH&
O8SC9&
O8SCH&
O8SCH&
O8QC`&
O8SCS&
O8`C9&
O8QCU&
O8SC`&
O8SC`&
O8SC;&
O8SC9&
O8NC8&
O8SC;&
O8SC;&
O8HC`&
O8UCQ&
O8UCH&
O8QCH&
O8QCH&
O8SCS&
O8UC8&
O8NCQ&
O8UC`&
O8QC8&
O8QC8&
O8UC8&
O8QCN&
O8SCH&
O8SC9&
O8NC8&
O8NC8&
O8NC;&
O8SCU&
O8QCU&
O8SC9&
8:&
K#@4/&E320>G&
!B#&E6%G&
l:;F&F;9&ERG&
:`NCH&
:`NCQ&
:`NCS&
:`NCN&
:`NCN&
:`NC`&
:``&
:``C:&
:``C;&
:``CH&
:``CH&
HC:;8&
HC:;Q&
HC:;N&
HC:H9&
HC:H9&
HC:H8&
HC:HH&
HC:HQ&
HC:U9&
HC:U8&
HC:U8&
O8SC9&
O8NCQ&
O8SCU&
O8NCH&
O8NCH&
O8`CH&
O8QCS&
O8SCU&
O8SC`&
O8QC;&
O8QC;&
&
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
!
&
88&