Human/Plant Interactions in Crop Evolution: Integrating Different Sources of Data in Contemporary Ethnobotany - Emshwiller

Human/Plant Interactions in Crop Evolution: Integrating Different Sources of Data in Contemporary Ethnobotany Eve Emshwiller University of Wisconsin – Madison Botany Department [email_address] 25 September 2010 Oxalis tuberosa as an example

oca – Oxalis tuberosa octoploid - 8 x clonal - vegetatively propagated

Research on oca
Origins of domesticated “oca” Oxalis tuberosa
progenitor species of domesticate
polyploid origins
phylogeny of Oxalis species closely related to oca
Continuing evolution of the crop under human influence
humans as agents of dispersal
distribution and dynamics of oca clonal genotypes

Inka kero vessel, possibly created during early Colonial “transitional” period

Archaeological evidence of oca
Cusco**
late horizon (Inca period) Vargas, 1981
Pachacamac
late horizon Towle, 1961
Jauja valley
late horizon Hastorf, 1983
Chim ú **
? Yá covleff and Herrera, 1933
Huachichocana (Valliserrana)
270 CE Pearsall, 1992
Pacheco **
middle horizon Yá covleff and Herrera, 1933
Putaca (Tarma)
middle horizon Hastorf, 1983
Anc ón-Chillón
3800 -- 4000 bp Martins, 1976
Guitarrero Cave
9000 -- 10000 bp Smith, C.E. 1980
Map modified from Pearsall 1992 (The Origins of Plant Cultivation in South America), in Cowan & Watson (eds.), The Origins of Agriculture .

Molecular datasets
DNA sequences of nuclear loci:
ITS
ncpGS
Fragment analysis:
AFLP fingerprints
DNA sequences of plastid loci:
trnS-trnG, trnS-trnT, trnT-trnL, trnL-trnL-trnF, trnH-psbA, psbJ-petA

2 X 8 X octoploid autopolyploid diploid progenitor

8 X 2 X 2 X 2 X 2 X allopolyploid octoploid diploid progenitors

Origins of domestication and polyploidy of oca ITS -- internal transcribed spacer of nuclear ribosomal DNA Emshwiller and Doyle (1998) AJB 85: 975-985 ncpGS -- chloroplast-expressed, nuclear-encoded glutamine synthetase Emshwiller and Doyle (1999) MPE 12: 310-319 Emshwiller and Doyle (2002) AJB 89: 1042-1056 “ Oxalis tuberosa alliance” x = 8

100 86 66 64 98 60 82 57 62 O. laxa var. hispidissima O. pachyrrhiza O. spiralis O. spiralis O. spiralis O. mollissima O. medicaginea O. lotoides O. subintegra O. unduavensi s O. oulophora O. longissim a O . sp. cfr. melilotoides O. humbertii O. lucumayensis ssp . lucumayensis O. lucumayensis ssp. subiens O. paucartambensis O. cuzcensis O. herrerae O. picchensis O. herrerae O. ptychoclada var. trichocarpa O. tabaconasensis O. peduncularis var. pilosa O. petrophila O. nubigena O. peduncularis O. weberbaueri O. sp. “V” O. flagellata O. flagellata O. andina O. ortgiesii O. boliviana O. boliviana O. vulcanicola O . sp. “C” O. sp. aff. ptychoclada O . sp. “A” 99 54 “ Oxalis tuberosa alliance” x = 8 66 87 79 99 100 100 strict consensus of 88 MP trees bootstrap values using strict consensus method Emshwiller (2002) Bot Rev 68:128-152 combined ITS and ncpGS data Cytology and name of Oxalis tuberosa alliance from: de Azkue & Martínez 1990

Oxalis sp. Oxalis picchensis wild tuber-bearing taxa in the Oxalis tuberosa alliance Map modified from Emshwiller (2002) Bot Rev 68:128-152

ncpGS data chloroplast-expressed nuclear-encoded glutamine synthetase (one of 208 MP trees) spiralis mollissima medicaginea lotoides longissima subintegra unduavensis oulophora aff. melilotoides sp. (Bolivia) tuberosa lucumayensis lucumayensis humbertii sp. C flagellata nubigena petrophila sp. aff. ptychoclada sp. A cuzcensis herrerae tuberosa sp. (Bolivia) tabaconasensis ptychoclada sp.V peduncularis weberbaueri herrerae tuberosa picchensis andina 1 1 1 1 5 5 1 1 1 3 3 3 2 1 1 1 4 1 2 1 2 2 1 13 1 1 other Oxalis species (not x = 8) 1 “ Oxalis tuberosa alliance” Emshwiller and Doyle (2002) AJB 89:1042-1056 7 #s = branch lengths

8 x 2 x 2 x 2 x autoallopolyploid octoploid diploid progenitors

Oxalis tuberosa 2 n = 8 x = 64 Bolivian wild tuber-bearing 4 x ? O. picchensis wild tuber-bearing 4 x One hypothesis of origin of polyploidy in oca Based on ncpGS data BUT… Oxalis sp. 2 x Oxalis sp. 2 x Oxalis sp. 2 x

Oxalis chicligastensis Oxalis sp. Oxalis picchensis wild tuber-bearing taxa in the Oxalis tuberosa alliance Oxalis sp. Map modified from Emshwiller (2002) Bot Rev 68:128-152

Ethnotaxonomy of oca in three communities in the district of Pisac in southern Peru

P’osqo (sour cultivar used only for khaya) many wayk’u folk cultivars

Khaya = oca processed by soaking, freezing & drying Khaya = oca processed by soaking, freezing & drying

Khaya = oca processed by soaking, freezing & drying

Ki 41 03 Ki 12 07 Ki?qKy 42 07 market 01 03 Ki 14 06 Ki 41 03 Ki 25 01 Ki 45 01 pP 35 06b pP 02 04 pP 19 03 pP 06 04 pP 06 02 pP 42 01 pP? 49 04 pP 35 06 hQ?ugly 16 06 O. picchensis PQ 46 15 PQ 46 15 PQ 31 03 PQ 46 14 PQ 42 04 PQ 06 06 PQ 06 05 O. sp. s/t Bolivia O. chicligastensis Ksp?pP 02 05 Ksp 44 03 Ksp 36 01 Ksp 14 02 Ksp 43 06 Ksp 31 04 Ksp 02 08 Sn 11 01 Qwk 11 06 hQ 21 06 hQ 17 02 hQ 48 02 hQ 19 08 qKy 07 04 hQ?yellow 16 04 Sn 13 04 hQ 14 03 orange 39 12 hQ 40 01 hQ 16 09 hQ?red 16 10 hQred 16 10 Sn 23 02 pPQ 40 02 pPQ 07 01 pPQ 25 02 pPQ 15 06 pPQ 13 02 pPQ 14 04 pPQ 21 02 Mch 47 10 Mch 07 02 Mch 07 02 pCh 49 02 pCh 49 02xxx market 01 02 market 01 09 Up 41 02weak Uya 31 06 Uyu 11 05 Uyu 14 07 Uya 02 09 Up 47 06 UyaKi 19 01 UyaKi 19 01 ?? 47 04 ?? 47 04 Uya 16 03& Up 39 13b Up 39 13a Uyu 21 04 Uyu 11 05 Uyu 19 06 Uyu 19 05 Uyu 07 03 ?red 10 01 ?orange 13 03 Dam 31 08 ?yellow 16 07 ?yellow 16 07 ?red 12 05 ?red 12 05 Sn 50 02 MiH 44 01 MiH 44 01 Mi 41 01 Mi?hQ 37 05 Mi 14 08 Mi 20 03 Mi 07 05 Mi 21 07 Mi 37 02 Mi 15 07 Mi 20 03 Mi 43 01 Mi 35 04 Mi 17 03 Mi 11 03 MiH 22 06& MiH 46 05 Mi 10 02 Mi? 41 08 market 01 04 Neighbor-Joining Analysis (Nei & Li distances) khaya (P’osqo) wayk’u wild tuber-bearing AFLP data from 1 primer combination wild tuber-bearing wild tuber-bearing Emshwiller (2006) Chapter in Darwin’s Harvest (Motley & al. eds.)

6 0 5 0 4 0 3 0 3 0 2 0 1 0 0 Frecuency Number of bands amplified (>95 kb) * * * * * * * * * P’osqo & 4x wild Oxalis Wayk’u oqa AFLP data from 1 primer combination

Oxalis chicligastensis Oxalis sp. Oxalis picchensis wild tuber-bearing taxa in the Oxalis tuberosa alliance Oxalis sp. Map modified from Emshwiller (2002) Bot Rev 68:128-152

Comparing oca and 4 wild, tuber-bearing Oxalis with AFLP data
7 primer combinations
399 peaks scored
57 samples cultivated oca (10 khaya, 47 wayk’u)
16 samples wild, tuber-bearing taxa
Analyses:
Non-metric multidimensional scaling (NMDS)
Neighbor-joining (N-J)
Peak-sharing evaluation
Dr. Terra Theim

wild tuber-bearing taxa in the Oxalis tuberosa alliance Oxalis sp. 4x (Emshwiller 2002b) 4x? (Brücher 1969) ?? 6x? (Kelly Vivanco, in prep.) Oxalis sp. Oxalis chicligastensis Oxalis picchensis Map modified from Emshwiller (2002) Bot Rev 68:128-152

-0.6 -0.4 -0.2 0.0 -0.3 -0.2 -0.1 0.0 0.1 0.2 0.3 NMDS1 NMDS2 NMDS analysis of 399 AFLP markers (7 primer pairs) from cultivated oca and four wild, tuber-bearing Oxalis taxa. O. picchensis [Peru, Depts. Cusco & Apurimac] Oxalis sp. [Peru, Dept. Lima] O. chicligastensis [NW Argentina] Oxalis sp. [Bolivia] wayk’u oca khaya oca (P’osqo) 8 x 4x? Thanks to analytical help from Cécile Ané Emshwiller & al. (2009) AJB 96: 1839–1848 Kelly Vivanco, unpublished

Neighbor Joining analysis of AFLP data (7 primer pairs). O. picchensis wild tuber-bearing [Peru, Depts. Cusco & Apurimac] Oxalis sp. wild tuber-bearing [Peru, Dept. Lima] O. chicligastensis wild tuber-bearing [NW Argentina] Oxalis sp. wild tuber-bearing [Bolivia] “ Wayk’u oqa” – cultivated oca used either for cooking after sunning (or for khaya) “ P’osqo oqa” – sour ocas used only for khaya Emshwiller & al. (2009) AJB 96: 1839–1848

trnT-trnL 1 of 5 MPTs O ortgiesii ORT O boliviana EE921 O rigidicaulis boliviana AAV5415 O yungasensis EE1079 O virgata EE292 O cuzcensis EE973 O cuzcensis EE982 O cuzcensis EE1123 O cuzcensis EE606 O nubigena EE188 O nubigena EE654 O nubigena EE701 O spiralis Comas EE590 O sp nov Comas EE592 O phaeotricha EE497 O teneriensis EE605 O ptychoclada EE916 O urubambensis EE926 O sp EE979 O sp EE971 O cuzcensis EE887 O picchensis EE500 O picchensis EE1152 O chicligastensis EE1001c O chicligastensis EE1002 O chicligastensis EE1003 O chicligastensis A O chicligastensis B O sp w tubers Bolivia EE284 O sp w tubers Dept Lima EE1110 O sp w tubers Dept Lima EE1169 O sp w tubers Dept Lima EE1185 O sp w tubers Bolivia EE259 O sp w tubers Bolivia EE260 O sp w tubers Bolivia EE260C O sp w tubers Bolivia EE262 O tuberosa 50 02 O tuberosa 46 14 O tuberosa 40 02 O tuberosa 19 01 O tuberosa 11 07 O. picchensis [Peru, Depts. Cusco & Apurimac] O. sp. wild tuber-bearing [Peru, Dept. Lima] O. sp. wild tuber-bearing [Bolivia] O. tuberosa cultivated oca, both wayk’u and khaya [Peru, Dept. Cusco] O. chicligastensis [nw Argentina] [Bolivia] Oxalis tuberosa alliance Still to be confirmed… Preliminary data. Please do not cite!

wild tuber-bearing taxa in the Oxalis tuberosa alliance Oxalis sp. 4x (Emshwiller 2002b) 4x? (Brücher 1969) ?? 6x? (Kelly Vivanco, in prep.) Oxalis sp. Oxalis chicligastensis Oxalis picchensis Map modified from Emshwiller (2002) Bot Rev 68:128-152

wild tuber-bearing taxa in the Oxalis tuberosa alliance Map modified from Emshwiller (2002) Bot Rev 68:128-152 Guitarrero Cave is outside of the (present day) distributions of the likely progenitors of cultivated oca.

Research on oca
Origins of domesticated “oca” Oxalis tuberosa
progenitor species of domesticate
polyploid origins
phylogeny of Oxalis species closely related to oca
Continuing evolution of the crop under human influence
humans as agents of dispersal
distribution and dynamics of oca clonal genotypes

Clonal vs. Seed-propagated crops

Conservation of crop genetic diversity Conservation of crop genetic diversity

KHAYA WAYK'U p'osqo kusipata ushpa panti q'ellu panti yuraq panti machasqa puka p'osqo misitu hank'o q'ello q'ellu k'aytu yuraq kishwar Damaso puka chaleco chachapiya waqankillay use categories folk cultivar names sub-cultivars yana misitu tullu misitu higos misitu k'aspi misitu q'ellu misitu yana ushpa puka ushpa yuraq ushpa for processing by freezing and drying for cooking fresh after sun-sweetening Emshwiller, unpublished data (field notes from 1997)

Panti Hanq’o Q’ello Kusipata Yana Ushpa Puka Ushpa Yuraq Kishwar Misitu Q’ellu Panti P’osqo Q’ellu K’aytu Machasqa Emshwiller, unpublished data (field notes from 1997)

Sn qKy Sn Sn Oxalis chicligastensis Oxalis sp. (wild tuber-bearing of Bolivia) Oxalis picchensis 0.005 changes NJ Puka Chiliku (chaleco) Ushpa Yuraq Kishwar Misitu / Higos Puka P’osqo Machasqa Damaso “ yellow group” : Q’ellu K’aytu Hanq’o Q’ello / Waqankillay Q’ellu Panti / Señorita Puka Panti Kusipata P’osqo M M M M Yana Ushpa ( & Yana Kishwar) Puka Ushpa Yuraq Ushpa Yana Misitu Q’ellu Misitu Higos Misitu Tullu Misitu K’aspi Misitu { { * * * * * * * * * Señorita P’osqo Ushpa Kishwar Kusipata Se ñ orita Puka Panti Puka Chiliku Misitu Hanq’o Q’ellu Puka P’osqo Machasqa Damaso Q’ellu K’aytu AFLP data from 1 primer combination * * { Emshwiller (2006) Chapter in Darwin’s Harvest (Motley & al. eds.)

Somatic mutations lead to chimeric tubers, and then to clone-mates of differing pigmentation?

K. S. Zimmerer 2003 Soc. & Nat. Res. 16: 583-601 Seed Network (“seed flow”) studies via surveys of farmers &/or market vendors

? ? ? ? ? ? gifts theft sharecropping payment-in-kind barter cash sale relatives and compadres within same community within district other communities Pisac Market ? ? Different modes of exchange operate at different scales Emshwiller, unpublished data (field notes from 1997)

Long-distance dispersal

Cusco Puno Cochabamba La Paz “ Ancient trade routes” hypothesis based on CIRNMA’s market surveys image source: USGS DEM and the Cornell Andes Project

Bryan Epperson Oca collection localities 2005 & 2006

Are patterns of distribution of clonal genotypes of oca correlated with distributions of different human languages or dialects? Quechua dialect map A. Chirinos Rivera (2001) Atlas Lingüístico del Perú

– Eve Emshwiller R. Carolina Girón A. Victor Nina M. Juan Tineo C. J. Eduardo Angeles M. Policarpo Catacora C. Tulio Medina H. Field Expeditions led by Coordinators of Genetic Resources at 5 of INIA’s experiment stations

and Kelly Vivanco, Nery Amaru C., Erwin Peralta, Carlos Antonio Hernandez, Rufino Martínez Cucho, Salome Altamirano Y., Manuela Huacachi Q., Alfredo Tupayachi H., Maibé Olivera G., Manuel Concha Menacha, Jorge Luís Millones I., Javier Ríos V., Roque Alberco, Eulalia Vargas, Isaias Achata T., Julio Huaripata C., Wilbert Cruz H., Iramí Pérez Hurtado

random sampling from field (chacra)

sampling from storage

directed sample based on morphological traits of tubers

Implicacions of restricted distribucions of clones for in-situ and ex-situ conservacion…

points Geographical points used in spatial autocorrelation analyses

Moran’s I expectations for IBD N e = Wright's neighborhood size

Moran’s I: average of 66 polymorphic alleles Preliminary data. Please do not cite!

Moran’s I: 66 individual polymorphic alleles Preliminary data. Please do not cite!

Moran’s I for oca AFLP alleles 1-10 Preliminary data. Please do not cite!

Preliminary data. Please do not cite!

Moran’s I for oca clones (that have over 5 representatives in sample) These preliminary results will be examined more closely soon. Stay tuned… Preliminary data. Please do not cite!

Luya Viejo

“ Yungay” improved potatoes introduced by a middleman who buys them for sale in coastal cities.

Displacement of oca and other native crops as unintended consequence of development projects

oqa kuru oca weevil gorgojo de la oca

agricultural biodiversity fairs to encourage in-situ conservation

Cuyo cuyo

Cuyo cuyo, famous for pre-Inca terraces and oca diversity

Thank you!

Human/Plant Interactions in Crop Evolution: Integrating Different Sources of Data in Contemporary Ethnobotany - Emshwiller

by Eve Emshwiller on Oct 09, 2010

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