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Capsicum Phylogeny and Domestication. Brian Walsh. 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

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slide2
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)
one of the most diversely utilized plants
One of the most diversely utilized plants
  • Spice
  • Food
  • Dye
  • Medicine
  • Weapons
  • Poison
  • Drinks
  • Deterrent
  • Insecticide/Pesticide
  • Preservative
  • Snuff
  • Rituals
  • Tribute
  • Town Names
  • Punishment
domestication
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)
slide6

Agricultural Centers of the New World and the theoretical flow of speciation

View from 1000000 km above 5°18’S 63°55’W

Based on C. chacoense

domesticated characters in capsicum
Domesticated Characters in Capsicum
  • Gigantism of fruit
  • Altered fruit shape
  • Fruit color other than red
  • Retained on peduncle a maturity
  • Variable capsaicin levels

Domesticated

Wild

  • small fruit
  • Fruit round
  • Red fruit
  • Peduncle dehiscence at maturity
  • High capsaicin levels
slide10

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

slide18

Capsicum eximium

C. baccatum var. praetermissum

slide19

Capsicum cardenasii

Only species that strictly outcrosses

slide20

Flower color classification of Capsicum

  • Color categories inaccurate (too many exceptions)
  • Only a fraction of total species included
  • Doesn’t agree with molecular data
current phylogenetic questions
Current phylogenetic questions
  • Monophyly of genus
  • Resolving species relationships
slide22

Capsicum Monophyletic?

Modified from Bohs & Olmstead 1997

slide23

A

B

C

A

B

C

=

Monophyletic

A

A

C

A

A

C

=

Not Monophyletic

slide24

Walsh 2001

(chloroplast atpB-rbcL spacer data)

slide25

Walsh 2001

(chloroplast atpB-rbcL spacer data)

slide26

Walsh 2001

combined atpB-rbcL spacer and waxy data

n=12/13

slide27

Walsh 2001

(chloroplast atpB-rbcL spacer data)

capsicum monophyly
Capsicum monophyly
  • Intact for the time being
  • Odd C. ciliatum seems to be the weak link

(n=13)

mascone s missing tree
Mascone’s missing tree

Mascone 2003

Flow cytometry and Feulgen densitometry

1C-value of DNA = an equation for calculating genome size in absolute units

Genome size in Mbp

slide30

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

slide31

1C G

C. chacoense

3.35 3273

C. annuum var. annuum

3.38 3312

C. frutescens

3.40 3332

C. chinense

3.42 3352

slide32

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

slide33

1C G

C. chacoense

3.35 3273

C. annuum var. annuum

3.38 3312

C. frutescens

3.40 3332

C. chinense

3.42 3352

C. baccatum var. pendulum

3.69 3621

C. baccatum var. baccatum

3.71 3636

C. baccatum var. umbilicatum

3.76 3636

Interpreted from data – not explicitly stated

slide34

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 DNAcontentintermediate between C. baccatum and C. pubescens, but genome size supports close relationship to C. pubescens as well as isozyme and cross studies

slide35

1C G

C. chacoense

3.35 3273

C. annuum var. annuum

3.38 3312

C. frutescens

3.40 3332

C. chinense

3.42 3352

C. baccatum var. pendulum

3.69 3621

C. baccatum var. baccatum

3.71 3636

C. baccatum var. umbilicatum

3.76 3636

C. eximium

4.06 3979

C. pubescens

4.47 4381

?

C. campylopodium

5.14 5032

?

C. parvifolium/cardenasii

5.77 5655

however mascone claims his data supports his hypothesis
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
slide37

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

slide38

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

x 13 species
X=13 species
  • C. ciliatum
  • C. buforum
  • C. campylopodium
  • C. cornutum
  • C. schottianum
  • C. villosum
  • C. pereirae *
  • C. friburguense *
  • 3 unnamed Capsicum
slide41

n = 13

n = 12

slide42

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
slide54

Cherry Pepper

Bolivian rainbow

slide55

Chilaca

Fluorescent

slide58

Lemon black pepper

Little nubian

slide60

Peter Pepper

Pimenta-Doce

slide61

Poblano

Suryamukhi

slide62

Tepin

Variegata

slide63

C. Frutescens

Naga Jolokia / Tezpur

slide64

Tabasco

PI451762

slide65

C02466

Antillerna (Paris)

C. chinense

slide67

C. chacoense

PI560944 - Putapario

slide70

Capsicum eximium

praetermissum