Pre-Columbian land use in the ring

Pre-Columbian land use in the ring-ditch
region of the Bolivian Amazon
Article
Accepted Version
Accepted manuscript
Carson, J., Watling, J., Mayle, F., Whitney, B. S., Iriarte, J.,
Prumers , H. and Soto, J. D. (2015) Pre-Columbian land use in
the ring-ditch region of the Bolivian Amazon. The Holocene,
25 (8). pp. 1285-1300. ISSN 0959-6836 doi:
10.1177/0959683615581204 Available at
http://centaur.reading.ac.uk/40180/
It is advisable to refer to the publisher’s version if you intend to cite from the
work.
Published version at: http://hol.sagepub.com/content/early/2015/04/29/0959683615581204.full.pdf+html
To link to this article DOI: http://dx.doi.org/10.1177/0959683615581204
Publisher: Sage Publications
All outputs in CentAUR are protected by Intellectual Property Rights law,
including copyright law. Copyright and IPR is retained by the creators or other
copyright holders. Terms and conditions for use of this material are defined in
the End User Agreement .
www.reading.ac.uk/centaur
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Central Archive at the University of Reading
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Page 15 of 62
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Page 17 of 62
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A61
1
HOLOCENE
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http://mc.manuscriptcentral.com/holocene
A71
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Page 19 of 62
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A81
1
HOLOCENE
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http://mc.manuscriptcentral.com/holocene
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Page 21 of 62
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HOLOCENE
http://mc.manuscriptcentral.com/holocene
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http://mc.manuscriptcentral.com/holocene
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Page 23 of 62
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94+5 CE9D234 FAF5 B3245 25 >F5 2%%245 %F%>FAB5 25 34%+5 F,FAAFF45
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HOLOCENE
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HOLOCENE
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FAF5B5439>F+5453B59F5)FA2+5E+FAE5C97(F75!2F,FAE53F53)23FBB525
)2B?82495 =9,F5 0%FA945 +F%2A9)35 29)BF5 )A2)2BFB5 395 B+5 2A+5
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HOLOCENE
+BF9BFB5 24+5 39,F5 >FF45 B)AF9+5 A9)+5 ,95 F:F4B,F5 49,F5 A9+F5 A24FB5
E'F4F,94E5C98/E5C99K5'2>4BE5C9/FE532453F54FF+52A5+AF5249535
4A2)F94B75F59>94+24%F4P+F4F59523FA5B243FA450%9D24945BFBE5B435
9B5 3F5 A4?+35 ,9FB5 45 3F5 -))FA5 ;44E5 24AAF+5 45 3F5 F44A5 9FA5
4A2)F945 9AA,9E5 94+5 B5 9BA>F+5 25 +BF9BF?+A,F45 +F%2A9)35 29)BF5
E!F"F4>FAFAE566F755
http://mc.manuscriptcentral.com/holocene
3B1
1
HOLOCENE
45 3F5 AB5 4B94FE5 3F5 %45 25 BF5 +F4F5 95 I*5 24+5 9))F9A5 25
B4))2A5 95 )2)49245 29)BF5 4"F+5 35 4A2)F945 24975 !2F,FAE5 2BFA5
4B)F245 25 3F5 AF2A+5 AF,F9B5 395 >4A445 >F945 25 +F4F5 A2%5 J/665 #5
94+5 9B5 9F+5 >5 2AFB5 F:)94B24E5 335 9B5 2%)FF5 >5 J(665 #75 53B5
B4FBB5395BF5+F4F59A244+5I*5>F945>F2AF5AB54A2)F94524975F5
%4B5 AF95 3F5 +945 25 3B5 AF2A+5 9424BE5 9B5 3F5 9F5 %2+F5 45 3B5 4))FA5
BF245 25 3F5 2AF5 39B5 95 +F5 FAA2A5 A94F5 EBFF5 64AF5 &F753F45 F5 9"F5 425
Fo
92445 3F5 4FA)29F+5 9(U5 FAA2A5 A94F5 25 3F5 +9FBE5 3B5 )9FB5 3F5 2AFB5
F:)94B245 9A244+5 I*5 B2%F3FAF5 >FFF45 &66?/65 AB5 #E5 %F9445 395
rP
92AFB9245 25 3F5 BF5 24+5 39,F5 24AAF+5 4)5 25 95 F44A5 >F2AF5 2A5 9FA5
4A2)F94524975!2F,FAE5,F453953F5+945253B532AD245B5>244+F+5>5
25A9+29A>245+9FBE53F549AA2524+F4F54FA,952453F52FA524BA9445
ee
+9F5 *.&(5 E9(U5 9F5 A94F5 25 /76?8/65 95 #FE5 94+5 3F5 4F9A5 BF+%F49245
A9F5 3A243245 3F5 2)5 C665 %5 25 3F5 2AFE5 9B5 4+9F+5 >5 24A5 9F5 %2+F5
rR
E64AF5&FE5 9F)453F54FA)29F+59F5252AFB5F:)94B245J(665#5BFF%B5
AF9B249>F75 F5 9B25 42F5 395 3F5 +945 25 3F5 39A295 +F4F5 95 I*E5 3F45
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3F5 4FA)29F+5 9(U5 24+F4F5 A94F5 B5 9"F45 425 9244E5 B5 >FFF45 ((6?
/965 95 #75 53B5 B5 )9FB5 3F5 >F4445 25 3F5 >4A445 +F4F5 >F2AF5 C&95
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5B3FA5 AFF45 )4>B3F+5 AF2A+B5 25 )AF?824%>945 24)9245 95
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F9A32A"5 BFB524F%)2A9A5 35395 25 I*E5 29F+5453F5 F4A95 BC3DE74 F24
EE75E34F5F597E56CE56C&FE539,F5+9F+5BF5+F4FP94+54BF5394F525
B32A5 >F2AF5 3F5 9AA,95 25 4A2)F94B75 'F4F5 95 3FBF5 BFB5 9B5 F,+F45
94BF+5>592AB523FA53945B+52A+5+BF9BF75 5B59B252A3542453953F5
244FB5 A9+29A>245 +9FB5 2>94F+5 B25 9A5 A2%5 A45 +35 EF2)3F5
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http://mc.manuscriptcentral.com/holocene
3A1
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Page 33 of 62
)AF?C&95 0'5 E944942%9E5 6C6F75 B4A5 )99F2F2295 AF2A+E5 24)F+5 35
3F5+945259A39F22959FAB595OO5>5#A$%FAB5F5975E66/FE59525335
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3B5BF5B5%2AF5"F755
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94+53F5+F4F545943A2)2F459,52>BFA,F+595I*5B5425B4))2AF+5A2%5
24A5+99E5F:394FB5>FFF454A2)F94B594+549,F50%9D2494B54F,FA3FFBB5
Fo
"F5 24A>4F+5 25 394FB5 45 94+5 4BF5 BA9F5 245 BFB5 "F5 OO5 45 3F5
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E'F4F,94E5 66CF75 53FBF5 4F5 22B5 %9+F5 B9B3?94+?>4A45 9A44AF5 95
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0A44A9594+594+?4BF5BA9F5
45 24A5 )2F45 949BFBE5 29ABF5 A924B5 FAF5 B944F+5 2A5 2%%245 4F45
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%9424E3DE8427#C2D8348A94DFE494+5%9DF5E123453674I7FE5952533539,F5
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53FBF5 B)FFB5 9B25 39,F5 +942B5 )323BE5 9B5 +25 9AA2A225 E393D834
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HOLOCENE
39#DFD3234 I4+7F5 94+5 AF45 E(3C38234 3CCE#34 I7F75 45 >235 3F5 )2F45 94+5
)3235 AF2A+BE5 14 53675 9B5 3F5 245 4F45 AF2,FAF+4 045 949BB5 25
)323B594+5B9A35A94B5A2%5B24F522594+5)2FA5AF%94B5AF2,FAF+5A2%5
3F5 *'#5 A45 +35 9B25 244+5 %9DF5 25>F595 4>N424B5 4F45 E'"945 F5975
6CF75 B4A5 +995 24A%5 395 %9DF5 9B5 "F5 945 %)2A945 B9)F5 A2)5 25 3F5
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9B5 C7(65 AB5 #75 53F5 335 39A295 F,FB5 9BB29F+5 35 3F5 )AF?824%>945
http://mc.manuscriptcentral.com/holocene
321
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HOLOCENE
24)9245 B4FB5 395 3F5 94+5 4BF5 BA9F5 4,2,F+5 3F9,5 >4A445 25 3F5
829934 9524 B9,944935 9A244+5 3F5 BF75 !2F,FAE5 23FA5 B24AFB5 25 39A295 9AF5
9B25"FE5B4359B54)4B5A2%5F,FA+95AF54BF5A2%53F9A3B7553F5B24AFB525
4F52A53B5>4A44524+539,F544+F+5B9,944935AFFBE5AFFB5A2%53F59FA5
2AFB5 9A244+5 I*5 94+5 3F5 4F9A>5 945 9AR45 .,FAE5 94+5 9FA5 J6665 AB5 #E5
AFFB5A2%53F5F,FAAFF45A942AFB53959%F525B4AA244+53F5OO5BF52245
AF2495A942AFB5F:)94B24759A2B2)539A295B5+F)2BF+545959"F5>9B45
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>5 ,9A24B5 )A2FBBFBE5 %F9445 395 )F9"B5 94+5 A243B5 45 3F5 39A295 AF2A+5
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)2F4595JC666595A5#75094E53B5+BAF)945>FFF453F525)A2:FB5%95
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244B245 B5 B4))2AF+5 >5 3F5 9))9AF45 F9A94F5 25 9FA5 2AFB5 +4A45 3F5
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Page 34 of 62
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http://mc.manuscriptcentral.com/holocene
331
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Page 35 of 62
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9A44AF5 45 3B5 AF245 B5 %942E5 335 B5 F4FA95 24B+FAF+5 25 >F5 95
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HOLOCENE
@FE5 667F5 94+5 3F5 >9BB5 2A5 3F5 +F,F2)%F45 25 9AFE5 BF+F49AE5
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http://mc.manuscriptcentral.com/holocene
341
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HOLOCENE
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Page 36 of 62
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http://mc.manuscriptcentral.com/holocene
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Page 37 of 62
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HOLOCENE
53B52524ABF5+2FB5425A4F52453F5)2BB>53952AFB5AFB24AFB5FAF5
%949F+595F95OB975 4AF9BFB545F242%5B)FFB5%95425>F5BFF45453F5
)99F25 AF2A+5 >F94BF5 3FA5 )2F45 %2A)3225 +2FB5 425 925 3F%5 25 >F5
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0FA49,FE5 3F5 )249245 BA9F5 25 B2%F5 F242%5 9:95 %95 %F945 395
3F5 9AF5 44+FAAF)AFBF4F+5 45 3F5 )2F45 AF2A+5 E4B35 94+5 .,FA9E5 66CF5 94+5
94425>F59)4AF+54595B94+9A+566?(665)2F45A94524475'FB)F53BE524F5
http://mc.manuscriptcentral.com/holocene
361
1
HOLOCENE
253F5%2B54A445242%FB52524A54,FB924B595I*5B53F5F,+F4F5395
)F2)F5)AF+9F53F5 9AA,95 252AFB52453B5BFE594+53953FAF5 9B595245 94+5
244424B5 24)9245EB)94445JC(665F9ABF75 5B53FAF2AF5335"F5395
3F52AFB52453B5BF539B54F,FA5>FF453245B2%F5+FAFF52534%945%)97555
53F5 +FA5 BF5 9A244+5 I*5 9B5 %9494F+5 9B5 945 2)F45 943A2)2F45
94+B9)F5 445J/66?(665 AB5#E59FA5335 92AFB924522"5 )9F75F5945
244+FE53FAF2AFE5395%435253F52AFB5425B4AA244+453F5%2+FA45,9F5
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9B5 FB9>B3F+5 45 3F5 9B5 (665 F9AB75 B45 95 AF2495 B9F5 9B2E5 >F24+5 3F5
D24F5 25 4F4B,F5 34%945 94+5 4BF5 9A244+5 3F5 24)9245 BFE5 3F5 F,FAAFF45
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2AFB5 9B5 FB9>B3F+5 AF9,F5 AFF4E5 F:)94+45 A2%5 J6665 AB5 #5 45
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395 3FAF5 9B5 B2%F5 AFF4FA9245 25 3F5 9FA5 2AFB5 A2%5 J(665 AB5 #75
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BFF%F4B5 A93FA5 3945 +F2AFB924E5 9B5 B4FBF+5 >5 3F5 )99F2F2295
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=F,F5F597E56CCF7551
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http://mc.manuscriptcentral.com/holocene
371
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Page 39 of 62
56D78B6DE
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HOLOCENE
.93FA5 3945 F:)FAF445 BF5 9>94+24%F45 9FA5 4A2)F945 2495 45
0'5 C&9E5 3F5 F95 OB95 BF5 2444F+5 25 >F5 24)F+5 3A2435 25 4F9A5 3F5
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25 3F5 2AFB5 395 F:BB5 9A244+5 3F5 BF5 2+975 53B5 +F%24BA9FB5 39E5 A93FA5
http://mc.manuscriptcentral.com/holocene
381
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HOLOCENE
3945>F4594F45E7F75%F4495459FFE53F52AFB52,FA453B5BF52+95B5a5
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http://mc.manuscriptcentral.com/holocene
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9A39F2>294952A"59AAF+524595OO75F59B25394"512B5944F59AA2B5
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5
5
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5
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1
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HOLOCENE
5
5
5
5
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http://mc.manuscriptcentral.com/holocene
4B1
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HOLOCENE
$3323D3EE
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http://mc.manuscriptcentral.com/holocene
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HOLOCENE
839,F51E5824+5.E54FA?I94+945!8E5F5975E667F50BBFBB45F,+F4F52A595)FA,9B,F5
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http://mc.manuscriptcentral.com/holocene
421
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http://mc.manuscriptcentral.com/holocene
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HOLOCENE
*2B45E59F56E559F5=E5F5975E669F5'FAF49245>FFF45=F2A2)95
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http://mc.manuscriptcentral.com/holocene
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HOLOCENE
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http://mc.manuscriptcentral.com/holocene
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HOLOCENE
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http://mc.manuscriptcentral.com/holocene
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Page 62 of 62
http://mc.manuscriptcentral.com/holocene
Page 63 of 62
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HOLOCENE
http://mc.manuscriptcentral.com/holocene