‘Haricot de Lima’, ‘Haba de Lima’ or ‘Lima bean’: only a toponym or reflecting an evolutive value? a name derived from the name of a place, dated ~ 1770: Carter 1949, Webster 2001 D.G. Debouck @ Cali-Quito, 23 June 2022 1/25 1753 1707-1778 [pod] moon-shaped [continuously] beautiful 1737 2/25 additional reading: Debouck 2021; Linné 1737, 1753; Westphal 1974; Yoon 2010 Phaseolus lunatus L. Phaseolus inamoenus L. G27475, from Myanmar G25694, from Ethiopia . . . what Carl von Linné might have seen • No reference specimens in the LINN and CLI Herbaria (solved by Westphal 1974, Debouck 2021) • Reports about Old World origin: A.P. de Candolle 1825, Don 1832, Dietrich 1847 • Doubts about Old World origin: Bentham 1859, Al. de Candolle 1883 photos: CIAT 2022 additional reading: Mackie 1943; van Eseltine 1931 3/25 already described in the pre-Linnean botanical literature ✓ Charles de l’Écluse (1601) acknowledged as ‘foreign’ ✓ John Gerarde (1597 [1636]) reported one pair from Brazil ✓ J. Bauhin & J.-H. Cherlero (1651) reported several from Brazil from Gerarde 1597 (same woodcut as in de l’Obel 1591?!) already referring to the striae from the seed hilum already reporting the ‘willow-leaf’ variant from de l’Écluse 1601 mentioned from Africa, Brazil and America 4/25 the Amerindian/ folk names (possibly used by 1500!?) Cuba: fríjol caballero Petén: ib limits of Tahuantisuyu Haiti: pois souche Dominican R.: haba Guerrero: comba Jalapa: furuna Chiapas: patashete Córdoba: carauta Suchitepequez: ixtapacál Carabobo: guaracaro San Salvador: chilipuca Puntarenas: kedeba Calderón & Standley 1941 Antioquia: carcha Debouck 1988 Cundinamarca: machete Nariño: torta Esquivel et al. 1992 Carchi: torta Hazlett 1986 Imbabura: cacha Martínez 1987 Cañar: pallar McBryde 1947 Cabieses 1995 Piura: pallar Cajamarca: layo Miranda 1952 Cárdenas 1989 La Libertad: pallar Montero-Rojas et al. 2013 Debouck 1989 Pittier 1926 Lima: pallar Estrella 1988 Schmeda-Hirschmann 1994 Freyre et al. 1996 Ica: pallar Parodi 1932 Cochabamba: pallar Pdte. Hayes: cugué Peralta-Idrovo et al. 2019 Chuquisaca: pallar Ramos-Nuñez 1950 Jujuy: pallar Soukup 1986 Wilhelm de Mösbach 1992 Atacama: pallar 5/25 About the name and geographic origin: at contact (1492-1500) in Hispaniola the (Lima) bean was called ‘haba’: its flat seeds remind those of Vicia faba L., then grown in the Iberian Peninsula that name persisted in the Dominican R. because of the extinction of Taino people in Cuba the translation of ‘horse bean’ ‘fríjol caballero’was preferred (same reason?) 1520: discovery of the Strait of Magellan; another route to access Peru and Chile 1532: end of Inca empire 1564: the sea route from Acapulco to Manila was opened by the Spanish galleons the coastal agricultural areas (Brazil, Chile, Peru) were fully exposed to explorers pallar was the adopted name in Spanish since 1560 given the Quechuan expansion from the Ancasmayo (Nariño, Colombia) down to the Maule (R. del Maule, Chile) Lima bean was a logical European name for this pulse in 1777 onwards, as it refers to a native bean when Lima was the capital of the viceroyalty (1542-1821) the linguistic argument helped turn down any claim of origin in Old World tropics additional reading: Andrews 1992; Carter 1949; Columbus 1492-93; Cubero 1992; de Candolle 1883; Gay 1865; Lumbreras 1974; Molina 1810; Navarrete 1560; Sturtevant 1889 6/25 The biological origin: The crop (Lima bean) = the wild ancestor modified by the domestication syndrome DGD-502, Campeche, Mexico, 1979 • reduced pod shattering • increased seed size • novel seed colors & patterns • reduced seed dormancy • reduced HCN glucosides • early flowering & monocarpy ‘ib choh’ • dwarf determinate growth habit • photoperiod insensitiveness photo: Debouck 2008 1859: Bentham reported about ‘spontaneous’ forms in the Neotropics (but no specimens) 1926: Piper reported wild forms in CUB, PRI, MEX, PNM, CLB, VNZ, BRA, PER (but no specimens) 1943: Macbride reported wild forms for eastern Peru (later on shown with ‘M’ lectin pattern) 1952: Burkart reported wild forms for Formosa, Jujuy, Salta, Argentina (but no specimens) additional reading: Debouck 2016, 2021; Erickson 1982; Lewis 1987; Maquet & Baudoin 1997; Montero-Rojas et al. 2013 Smartt 1988 7/25 One often repeated story (Mackie 1943): Carter 1949; Friedberg 1958; Westphal 1974; Baudet 1977 G25217, Arizona G27555, Dominican R. Hopi branch Carib branch G25850, Guatemala Inca branch ✓ weak because of poor and uneven sampling of wild forms ✓ infraspecific diversity created by people from a single domestication G25903, CLB, Nariño although some argued about a double domestication: photos: CIAT 2022 Allard 1960; Estrella 1988; Evans 1976; Harlan 1971; Heiser 1965; Kaplan 1965 8/25 # 3245, CRA, San José, Sn Gerardo, 12/12/12 # 2863, ECD, Chimborazo, Huigra, 17/06/90 what brought a change in the thinking about double domestication: photo: Debouck 2012 photo: Debouck 1990 Background: Standley 1937 Background: Piper 1926 Explorations (13) in CRI in: 1987 - 2017 Explorations (2) in ECD in: 1989, 1990 9/25 at least 2 groups of wild forms Sonora Tamaulipas Puerto Rico Pan-Neotropical range Revillagigedo Is.BCS. small seeds : 6-12 g Panama sea level – 2200 m Yaracuy Trinidad-&-Tobago Caldas Imbabura Pacific range Huila (not very different from Pará the one of P. debouckii) Junín Bení Cajamarca small-medium seeds: 12-20 g 320 – 2000 m Paraguay Salta “sacha viuda”: Loja “layo del zorro”: Cajamarca source: Debouck 2019 “pallar de monte”: Cajamarca 10/25 A noteworthy wild Lima bean from Boyacá, Colombia • found in 1992 NE of Bogotá • racemes and flowers similar to wild P. lunatus of S Ecuador • 100-seed weight (16.8g) within the range of Andean AI wild forms • lectin pattern M1, as the ‘Mesoamerican’ but also as in Phaseolus augusti • formed a separate cluster in AFLP analysis • classified as MII on cpDNA markers • found close to MII on 4,593 SNPs sources: Caicedo et al. 1999; Chacón-S. & Martínez C. 2017; Serrano-S. et al. 2010; Toro et al. 1993 Two genepools in the wild and two domestication events: expected ! marker references lectin polymorphisms Debouck et al. 1989; Gutiérrez-S. et al. 1995; Lioi 1996 20 and 13 isozyme loci Maquet et al. 1994, 1997 RAPDs on genomic DNA Nienhuis et al. 1995; Fofana et al. 1997 RFLPs on rRNA Jacob et al. 1995 17 isozyme loci, RFLPs on rDNA Lioi et al. 1998 RFLPs on 6 IGRs of cpDNA Fofana et al. 1999 SSRs on genomic DNA Lioi & Galasso 2002 two cpDNA probes Fofana et al. 2001 sequences of cp and nDNA Serrano-Serrano et al. 2010 sequences of cpDNA and ITS Motta-Aldana et al. 2010 ITS of rDNA Serrano-Serrano et al. 2012 2 intergenic spacers of cpDNA Andueza-Noh et al. 2013 10 microsatellite loci Martínez-Castillo et al. 2014 ✓ 2 independent domestication events in distant locations (as in common bean) ✓ possibly few original populations involved  high linamarin in wild forms ✓ parallel demonstration by use of similar markers of molecular genetics 12/25 More genepools in the wild and a third domestication event: unexpected ! Existence of a 2nd genepool in the wild Neotropical “small-seeded” Lima bean (MII) marker references sequences of cp and nDNA Serrano-Serrano et al. 2010 sequences of cpDNA and ITS Motta-Aldana et al. 2010 sequences of cpDNA Andueza-Noh et al. 2013 genome-wide 4,593 SNPs Chacón-Sánchez & Martínez-Castillo 2017 Possibility of a 3rd domestication event from wild forms in MII genepool !? marker references sequences of cpDNA Andueza-Noh et al. 2013 ✓ insufficient sampling of MII in South America to claim a 3rd event ??? Existence of a 2nd genepool in the wild Andean “large-seeded” Lima bean marker references AFLP of genomic DNA Caicedo et al. 1999 genome-wide 4,593 SNPs Chacón-Sánchez & Martínez-Castillo 2017 GBS: 12,398 SNVs analyzed Garcia et al. 2021 ✓ these wild forms from Boyacá-Cundinamarca seem not to be hybrids ! 13/25 The migration northwards to Antioquia and Cundinamarca: expected, but towards the Chaco and eastern lowlands !? Elevation m; 100-seed g; Materials Region lectin pattern G25579 BLV, Sta. Cruz 450; 111.0; A6 G25540 ARG, Salta 1100; 70.0; A3 G26102 BRA. Rio Grande Sul 830; 95.4; A3 G26068C BRA, Minas Gerais 650; 86.0; A3 Santa Cruz Minas Gerais Lima bean grown by peoples of Chaco: the Ayoreo in Paraguay: Schmeda-H. 1994 Rio Grande do Sul Salta 14/25 Bentham 1859: “Specimina Brasiliensis formae macrocarpae vidi a Pohlio in Brasilia centrali et a Spruceo prope Santarem lecta”. Elevation m; 100-seed g; Materials Region lectin pattern 70 g < 100-seed weight < 130 g (39 acces.) Ceará G26003A BRA; Ceará 180; 80.0; M1 G25997 BRA; Pernambuco 1000; 73.0; M1 G26140 BRA; Sergipe 280; 70.0; M1 ? G26034 BRA; Minas Gerais 400; 95.0; M1 G26296 BRA; Goias 1000; 88.0; M1 Pernambuco Goias Sergipe ? Minas Gerais if introduced from the Caribbean, why no such big seeds there? 15/25 wThwy odo edsi tshte iAnncdtea dn odommeesstitcaictioant liooonk aesyvmemnettrsic ?al? DGD-453; Jalisco DGD-1944; Cajamarca DGD-508; Campeche DGD-1176; Cajamarca 10 g 12 g 2250 ppm increase in seed size and weight increase in seed size and weight 2747 ppm 43 ppm 328- 4g x 113311 g g 28 g 47 ppm 3-4 x 1 2747 ppm 100-1-144 x x DGD-453; Jalisco DGD-1944; Cajamarca 2250 ppm 43 ppm DGD-508; Campeche 10 g 12 g 47 ppm DGD-1176; Cajamarca 2 cm cm 16/25 Concluding remarks • “Lima bean predominates over the frijol in the majority of the American tropics” (Sauer 1950, p. 501) • in archaeological records (5,600 ybP), it appears fully modern; no transition from the wild in contrast to maize where the transition from the wild can be tracked over 8,000 years • the reasons to domesticate a lethal legume in absence of pottery (for two millennia) remain unclear, unless it was originally domesticated for non-food purposes • on the Pacific side, from Cundinamarca to the Atacama desert, into the Chaco and from there into southern Brazil, the genepool AI had a huge pre-Columbian range • 3 puzzling facts: ➢ this huge colonization of AI = MI, in spite of a narrow range in the wild ➢ also puzzling is the colonization of AI in altitude: from sea level up to 3,100 m ➢ the increase in seed size in AI: another genetic control as compared to MI? • field work, rather than methods, in South America, key to solve several pending questions additional reading: Debouck 2016; Kaplan & Lynch 1999; Vallebueno-E. et al. 2016 17/25 References (1) Allard, R.W. 1960. 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New York, New York, USA. 341p. 24/25 Many thanks for your attention! and a big ‘Thank you’ to (since 1977): Ana L Caicedo Jorge Acosta AGCD BMZ Maria I Chacón Rodolfo Araya CENTA César Azurdía CIAT Jorge Duitama CIFP Raúl Castillo COSUDE Eliana Gaitán EU Néstor Chaves DfID Alberto Gutiérrez Hipólito de la Cruz GCDT Gov Norway Antonio Hernández George Freytag+ IBPGR ICA Jorge Hernández Paul Gepts ICTA Celia Lima Rogelio Lépiz IDIAP INIA Luís López Jenny Motta INIAP INIFAP Jorge Liñan Martha Serrano INTA José Muruaga IUCN Alba M Torres UCR Raúl Ríos USAID Joe Tohme María C Sevillano USCG USDA Orlando Toro+ Luís Valera World Bank in the lab in the countries for the support 25/25