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Capsicum Phylogeny and Domestication
Brian Walsh
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Subfamily: Solanoideae Tribe: Capsicinae (since 1999)
Family: Solanaceae Subfamily: Solanoideae Tribe: Capsicinae (since 1999) Estimated 25 species (my count =37) Native to South and Central America Current distribution is global One of the most utilized spice in the world Part of human diet for 9000 years All diploid Typically self-fertilize Most n=12 (some n=13)
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One of the most diversely utilized plants
Spice Food Dye Medicine Weapons Poison Drinks Deterrent Insecticide/Pesticide Preservative Snuff Rituals Tribute Town Names Punishment
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Domestication Davenport (1970) proposes all five domesticated species arose from a single ancestor (C. frutescens L.) (Lumper) (based on morphology) Heiser (1971) proposes each domesticated species arose from its own unique wild ancestor (Splitter) (hybridization studies) Pickersgill (1972) agrees with Heiser - different cultivated species show a combination of characters suggesting domestication by different groups of people living in different areas. Eshbaugh (1983) argues at least three distinct domesticated lineages based on morphology, plant breeding and enzyme analysis (C. chacoense is plesiomorphic)
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Agricultural Centers of the New World and the theoretical flow of speciation
View from km above 5°18’S 63°55’W Based on C. chacoense
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Domesticated Characters in Capsicum
Wild Gigantism of fruit Altered fruit shape Fruit color other than red Retained on peduncle a maturity Variable capsaicin levels small fruit Fruit round Red fruit Peduncle dehiscence at maturity High capsaicin levels
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Capsicum annuum
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Eshbaugh’s breakdown into flower color categories (why?)
Capsicum annuum complex C. annuum var. annuum C. annuum var. avicular C. frutescens white flowers C. chinense C. chacoense Capsicum baccatum group C. baccatum var. praetermissum white flowers with yellow spots C. baccatum var. pendulum C. baccatum var. baccatum Capsicum eximium complex C. pubescens C. eximium purple flowers C. cardenasii Total species=8
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C. frutescens Tabasco
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C. baccatum var. praetermissum
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Capsicum eximium
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Capsicum eximium C. baccatum var. praetermissum
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Only species that strictly outcrosses
Capsicum cardenasii Only species that strictly outcrosses
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Flower color classification of Capsicum
Color categories inaccurate (too many exceptions) Only a fraction of total species included Doesn’t agree with molecular data
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Current phylogenetic questions
Monophyly of genus Resolving species relationships
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Capsicum Monophyletic?
Modified from Bohs & Olmstead 1997
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A B C A B C = Monophyletic A A C A A C = Not Monophyletic
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Walsh 2001 (chloroplast atpB-rbcL spacer data)
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Walsh 2001 (chloroplast atpB-rbcL spacer data)
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Walsh 2001 combined atpB-rbcL spacer and waxy data n=12/13
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Walsh 2001 (chloroplast atpB-rbcL spacer data)
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Capsicum monophyly Intact for the time being
Odd C. ciliatum seems to be the weak link (n=13)
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Mascone’s missing tree
Flow cytometry and Feulgen densitometry 1C-value of DNA = an equation for calculating genome size in absolute units Genome size in Mbp
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Visualization of Mascone’s results
Capsicum anuum complex C. annuum var. annuum C. annuum var. avicular C. frutescens white flowers C. chinense C. chacoense United by similar DNA amount and little heterochromatin
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1C G C. chacoense C. annuum var. annuum C. frutescens C. chinense
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Capsicum baccatum group
C. baccatum var. praetermissum white flowers with yellow spots C. baccatum var. pendulum C. baccatum var. baccatum C. baccatum var. umbilicatum New variety described in 1998 Domesticated status unknown Similar DNA amounts – not surprising
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C. baccatum var. pendulum 3.69 3621
1C G C. chacoense C. annuum var. annuum C. frutescens C. chinense C. baccatum var. pendulum C. baccatum var. baccatum C. baccatum var. umbilicatum Interpreted from data – not explicitly stated
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Capsicum eximium complex
C. pubescens C. eximium purple flowers C. cardenasii C. parvifolium synonym with C. cardenasii? (but description is off – tree?) C. parvifolium only compared to C. campylopodium points out distantly related and nothing more C. campylopodium groups with C. pubescens – large genome size, and large amounts of heterochromatin forming complex patterns (tone implies trepidation) C. eximium has DNA content intermediate between C. baccatum and C. pubescens, but genome size supports close relationship to C. pubescens as well as isozyme and cross studies
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? ? 1C G C. chacoense 3.35 3273 C. annuum var. annuum 3.38 3312
C. annuum var. annuum C. frutescens C. chinense C. baccatum var. pendulum C. baccatum var. baccatum C. baccatum var. umbilicatum C. eximium C. pubescens ? C. campylopodium ? C. parvifolium/cardenasii
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However, Mascone claims his data supports his hypothesis
Increase in DNA amount is evolutionary advancement C. chacoense most plesiomorphic (ancestral) C. campylopodium most advanced n=13 in more recently evolved
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Mascone tree rooted ? Least DNA C. chacoense C. annuum var. annuum
C. frutescens C. chinense C. baccatum var. pendulum C. baccatum var. baccatum C. baccatum var. umbilicatum C. eximium C. pubescens Most DNA C. campylopodium x=13 ? C. parvifolium/cardenasii
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Mascone molecular ? ? Least DNA C. chacoense C. annuum var. annuum
C. frutescens C. chinense C. baccatum var. pendulum C. baccatum var. baccatum C. baccatum var. umbilicatum C. eximium C. pubescens Most DNA C. campylopodium x=13 ? ? C. parvifolium/cardenasii ancestral
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X=13 species C. ciliatum C. buforum C. campylopodium C. cornutum
C. schottianum C. villosum C. pereirae * C. friburguense * 3 unnamed Capsicum
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Capsicum flexuosum
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n = 13 n = 12
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Conclusion Capsicum needs more research
Appears to be an x=12/x=13 split in distant past research hindered by incomplete sampling Difficult to study domestication when phylogeny unresolved 6 new species identified in last 8 years
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Naga Morich
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Following Photos unused in presentation
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Cherrapeño
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Cherry Pepper Bolivian rainbow
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Chilaca Fluorescent
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Fredrik from Amalfi
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Jalapeño - Biker Billy
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Lemon black pepper Little nubian
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O'odham Indian Piquin
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Peter Pepper Pimenta-Doce
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Poblano Suryamukhi
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Tepin Variegata
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C. Frutescens Naga Jolokia / Tezpur
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Tabasco PI451762
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C02466 Antillerna (Paris) C. chinense
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Capsicum galapagoense
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C. chacoense PI Putapario
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C. Pubescens
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Capsicum eximium praetermissum
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C. ciliatum
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Capsicum Cardenasii
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C. tovarii
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