A Palaeogenomic Window Into the History of Wine

A Palaeogenomic Window into the History of Wine

Tom Gilbert

What shapes biodiversity? 1) Management practices

Insect community

Fungal community Insect community

Fungal community Fungal diversity 2) Domestication 2) Domestication Key Questions:

Where? Key Questions:

Where?

When? Key Questions:

Where? Why?

When?

Key Questions:

Where?

When? How many times? Key Questions:

Where?

When? How many times? How did we distribute it? Key Questions:

Where? How do we continue to When? shape it? How many times? How did we distribute it? Wild grapes Domesticated grapes Vitits vinifera ssp. sylvestris Vitis vinifera ssp. vinifera Wild grapes Domesticated grapes Vitits vinifera ssp. sylvestris Vitis vinifera ssp. vinifera

Wine grapes Table grapes Vegetative propagation

Grape varieties = identical clones Myles et al. PNAS 2010, 108:9 Myles et al. PNAS 2010, 108:9 8000 year history

Romans

Ancient Egypt Areni-1, Armenia

Romans

Ancient Egypt Romans Rome 100 BCE

Egyptian tomb painting depicting grape cultivation, around 1400 BCE. Image the Metropolitan Museum of Art. Ancient Egypt Romans

Ancient Egypt

Questions

Ancient grapes/cultivars Modern cultivars (Roman grapes) Questions

Ancient grapes/cultivars Modern cultivars (Roman grapes)

How do the ancient grapes used by ancient civilizations relate to modern grapes? Questions

Ancient grapes/cultivars Modern cultivars (Roman grapes)

How do the ancient grapes used by ancient civilizations relate to modern grapes?

How were they cultivating these grapes? Questions

Ancient grapes/cultivars Modern cultivars (Roman grapes)

How do the ancient grapes used by ancient civilizations relate to modern grapes?

How were they cultivating these grapes?

Were they transporting grapes from other regions or cultivating local wild grapes?

Ancient grape seeds

Horbourg-Wihr Órleans n=1 n=3

28 archaeological seeds

Iron Age (~Bay 2,480 of Biscayyo), n=1 France

Colletiére, Charavines Roman (2,200–1,700 yo), n=19 n=1

Late medieval (1,300-900 yo), n=8 Magalas Lattes (trading center) n=3 n=1 Nimes n=9 La Lesse-Espagnac Sauvian Marseille n=5 Roumeges, Poussan n=2 Mont Ferrier, Tourbes n=3 • aDNA extraction (archaeological seeds)

• Illumina library

• Enrichment of 10,000 selected SNPs

• High-throughput sequencing

• High quality genotypes for 21 samples

Copenhagen clean lab for aDNA processing • aDNA extraction (archaeological seeds)

• Illumina library

• Enrichment of 10,000 selected SNPs

• High-throughput sequencing

• High quality genotypes for 21 samples

Ancient grape seed Copenhagen clean lab for aDNA processing GrapeReSeq panel

~789 modern domesticated varieties

112 wild grapes

~10,000 polymorphic sites

Lacou et al. PLoS ONE 2018 Wild grapes Wine grapes: West and Central Europe Medieval grapes Iberian Peninsula and Roman grapes Northwest Africa Table grapes Iron Age grapes Balkans and East Europe TABLE WILD

WINE

Lacou et al. 2018

Wild grapes Wine grapes: West and Central Europe Medieval grapes Iberian Peninsula and Roman grapes Northwest Africa Table grapes Iron Age grapes Balkans and East Europe Ancient grapes are closer to domesticated grapes

Grape varieties = ‘identical’ clones

Myles et al. PNAS 2010, 108:9 19 genetically identical clusters = 19 clones = 19 ‘varieties’

Myles et al. PNAS 2010, 108:9 Distribution of ancient grape cultivars

Horbourg-Wihr Órleans n=2 n=1

Bay of Biscay France Colletiére, Charavines n=1

Magalas Lattes n=1 n=3 Nimes n=9 La Lesse-Espagnac Sauvian Roumeges, Poussan n=5 n=2 Mont Ferrier, Tourbes n=3 Mediterranean Sea Distribution of ancient grape cultivars

Horbourg-Wihr Órleans n=2 Same variety n=1 cultivated in multiple regions Bay of Biscay France Colletiére, Charavines n=1

Magalas Lattes n=1 n=3 Nimes n=9 La Lesse-Espagnac Sauvian Roumeges, Poussan n=5 n=2 Mont Ferrier, Tourbes n=3 Mediterranean Sea Distribution of ancient grape cultivars

Horbourg-Wihr Órleans n=2 Same variety n=1 cultivated in multiple regions Thus evidenceBay of Biscay of France Colletiére, Charavines cultivation by cuttings n=1 during Roman and Magalas Lattes n=1 medieval period n=3 Nimes n=9 La Lesse-Espagnac Sauvian Roumeges, Poussan n=5 n=2 Mont Ferrier, Tourbes n=3 Mediterranean Sea Distribution of ancient grape cultivars

Horbourg-Wihr Órleans n=2 Multiple varieties in n=1 single locations

Bay of Biscay France Colletiére, Charavines n=1

Magalas Lattes n=1 n=3 Nimes n=9 La Lesse-Espagnac Sauvian Roumeges, Poussan n=5 n=2 Mont Ferrier, Tourbes n=3 Mediterranean Sea Assigning identity Ancient grapes Parent−offspring relationship Full siblings or similar Late Iron age (480 BCE) Ancient grapes Parent−offspring relationship Full siblings or similar

Late Iron age (480 BCE)

Roman period (200 BCE – 300 CE) Ancient grapes Parent−offspring relationship Full siblings or similar

Late Iron age (480 BCE)

Roman period (200 BCE – 300 CE)

Medieval (790 – 1020 CE) Ancient grapes Parent−offspring relationship Full siblings or similar

Late Iron age (480 BCE)

Roman period (200 BCE – 300 CE)

Medieval (790 – 1020 CE)

Late medieval (1125 CE) Ancient grapes Parent−offspring relationship Full siblings or similar

Late Iron age (480 BCE)

Roman period (200 BCE – 300 CE)

Medieval (790 – 1020 CE)

Late medieval (1125 CE) >1000 years of clonal Savignan/Traminer Savignan/Traminer Earliest appearance is Southern France

Horbourg-Wihr Órleans n=2 n=1

Bay of Biscay France Colletiére, Charavines n=1

Magalas Lattes n=1 n=3 Nimes n=9 La Lesse-Espagnac Sauvian Roumeges, Poussan n=5 n=2 Mont Ferrier, Tourbes n=3 Mediterranean Sea Conclusions

Multiple varieties were cultivated by ancient Romans Conclusions

Multiple varieties were cultivated by ancient Romans Transportation and propagation by cuttings Conclusions

Multiple varieties were cultivated by ancient Romans Transportation and propagation by cuttings Savagnin has been propagated for over 1,000 years Perspectives

• Reintroduction of now discarded cultivars? Perspectives

• Reintroduction of now discarded cultivars? • Recreation of lost cultivars? Perspectives

• Reintroduction of now discarded cultivars? • Recreation of lost cultivars? • Local adaptation? • Maybe they were discarded for a reason… Perspectives

• Reintroduction of now discarded cultivars? • Recreation of lost cultivars? • Local adaptation? • Maybe they were discarded for a reason… • Recreation of lost wine styles? • Couple with ancient microbe analyses Acknowledgments

Roberto Bacillieri Jazmin Ramos Laurent Bouby Thierry Lacombe

Nathan Wales Shyam Gopalakrisnan J. Alfredo Samaniego

Víctor Moreno …Mayar AK Runge Filipe G. Vieira Acknowledgments

• Hentie Swiegers

• Lars Hestbjerg Hansen, Lea Ellegaard-Jensen