Paper · 2025 · The Plant Genome

Integrated chloroplast genomics and whole‐genome resequencing reveals demographic history and selection signatures of black walnuts

Hang Ye; Hengzhao Liu; Huijuan Zhou; et al.

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Open access · cc-by-nc-nd

Abstract

Elucidating the impacts of demographic history and genomic selection on species evolution is a central topic in phylogeography and evolutionary biology. Black walnuts (Juglans section Rhysocaryon) are native nut trees of the NEW WORLD, with a broad distribution ranging from southern Canada to northern Argentina. The demographic history and genomic dynamics of Rhysocaryon species remain poorly understood. Here, we employed population genomics and chloroplast data to construct a high-density map of genomic variation across 108 Rhysocaryon accessions. Despite gene introgression, these accessions were clearly delimited into four groups. Evolutionary scenarios analysis showed that the diversification of black walnuts might have occurred approximately 28.74 million years ago during the late Oligocene, with the clade comprising Juglans hindsii and Juglans californica diverging earliest. The gene introgression and hybridization analysis indicated that Juglans microcarpa might be a hybrid descendant of Juglans nigra and J. hindsii. As the climate oscillated, these ancestral populations kept diverging, laying the basis for their colonization of South America. Quaternary climatic oscillations also exerted a profound influence on black walnut population size, which exhibited sensitive fluctuations in response to alternation of glacial and interglacial periods. The selection sweeps analysis unveiled highly divergent genomic regions in the economic species J. nigra, which were associated with development, reproduction, disease resistance, and stress tolerance. The genes WRKY41 and ERF012 were identified as potential drivers of J. nigra's adaptation. Our findings illuminated the demographic history and selective signatures of black walnuts, thereby providing a genetic foundation for future breeding, conservation, and genomic studies.

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Claims 10

Method / resourceobservedoriginal resultmeasured

Claim · model-written summary

The study constructed a high-density map of genomic variation across 108 Rhysocaryon accessions.

Taxon
Rhysocaryone0
Outcome
high-density map of genomic variatione0
Value
108 Rhysocaryon accessionse0

Verbatim

e0

“construct a high-density map of genomic variation across 108 Rhysocaryon accessions.”

✓ located verbatim in the source · abstract, characters 470–554 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

The 108 Rhysocaryon accessions were delimited into four groups despite introgression.

Taxon
Rhysocaryont0
Outcome
four groupse0
Value
four groupse0
Qualifier
Despite gene introgressione0

Verbatim

e0

“Despite gene introgression, these accessions were clearly delimited into four groups.”

✓ located verbatim in the source · abstract, characters 555–640 · show in text ↑
t0

“108 Rhysocaryon accessions”

✓ located verbatim in the source · abstract, characters 527–553 · show in text ↑
Observationobservedoriginal resultsimulated

Claim · model-written summary

Black walnuts may have diversified about 28.74 million years ago in the late Oligocene.

Taxon
black walnutsJuglans nigrae0
Outcome
diversification of black walnutse0
Value
approximately 28.74 million years agoe0
Qualifier
might have occurrede0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Juglans nigra
Juglans nigra · TaxonPhoto by and (c)2007 Derek Ramsey ( Ram-Man ). Location credit to the Chanticleer Garden. · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“the diversification of black walnuts might have occurred approximately 28.74 million years ago during the late Oligocene”

✓ located verbatim in the source · abstract, characters 685–805 · show in text ↑
Observationobservedoriginal resultsimulated

Claim · model-written summary

The clade containing Juglans hindsii and Juglans californica diverged earliest.

Taxon
Juglans hindsii and Juglans californicaJuglans californicaJuglans hindsiie0
Outcome
diverging earlieste0
Direction
earlieste0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Juglans californica
Juglans californica · TaxonEugene Zelenko · CC BY-SA 4.0 · Wikimedia Commons ↗
Juglans hindsii
Juglans hindsii · Taxonsarahangulo · CC0 · Wikimedia Commons ↗

Verbatim

e0

“the clade comprising Juglans hindsii and Juglans californica diverging earliest.”

✓ located verbatim in the source · abstract, characters 812–892 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Juglans microcarpa may be a hybrid descendant of Juglans nigra and J. hindsii.

Taxon
Juglans microcarpaJuglans microcarpae0
Outcome
hybrid descendant of Juglans nigra and J. hindsiie0
Qualifier
might bee0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Juglans microcarpa
Juglans microcarpa · TaxonJerry Friedman · CC BY-SA 2.0 · Wikimedia Commons ↗

Verbatim

e0

“Juglans microcarpa might be a hybrid descendant of Juglans nigra and J. hindsii.”

✓ located verbatim in the source · abstract, characters 958–1038 · show in text ↑
Associationobservedoriginal resultsimulated

Claim · model-written summary

Ancestral black walnut populations kept diverging as climate oscillated, preceding South American colonization.

Taxon
black walnutsJuglans nigrat0
Outcome
ancestral populations kept diverginge0
Direction
diverginge0
Qualifier
climate oscillatede0
Setting
South Americae0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Juglans nigra
Juglans nigra · TaxonPhoto by and (c)2007 Derek Ramsey ( Ram-Man ). Location credit to the Chanticleer Garden. · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“As the climate oscillated, these ancestral populations kept diverging, laying the basis for their colonization of South America.”

✓ located verbatim in the source · abstract, characters 1039–1167 · show in text ↑
t0

“black walnuts”

✓ located verbatim in the source · title, characters 118–131 · show in text ↑
Associationobservedoriginal resultsimulated

Claim · model-written summary

Black walnut population size fluctuated with Quaternary glacial and interglacial periods.

Taxon
black walnutJuglans nigrae0
Outcome
black walnut population sizee0
Qualifier
alternation of glacial and interglacial periodse0
Setting
Quaternary climatic oscillationse0

Not reportedDirection

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Juglans nigra
Juglans nigra · TaxonPhoto by and (c)2007 Derek Ramsey ( Ram-Man ). Location credit to the Chanticleer Garden. · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Quaternary climatic oscillations also exerted a profound influence on black walnut population size, which exhibited sensitive fluctuations in response to alternation of glacial and interglacial periods.”

✓ located verbatim in the source · abstract, characters 1168–1370 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

J. nigra contained highly divergent genomic regions associated with development, reproduction, disease resistance and stress tolerance.

Taxon
J. nigraJuglans nigrae0
Gene
genomic regionse0
Outcome
highly divergent genomic regionse0
Qualifier
development, reproduction, disease resistance, and stress tolerancee0

Not reportedDirection

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Juglans nigra
Juglans nigra · TaxonPhoto by and (c)2007 Derek Ramsey ( Ram-Man ). Location credit to the Chanticleer Garden. · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“highly divergent genomic regions in the economic species J. nigra, which were associated with development, reproduction, disease resistance, and stress tolerance.”

✓ located verbatim in the source · abstract, characters 1410–1572 · show in text ↑
Observationauthor interpretationoriginal resultmeasured

Claim · model-written summary

WRKY41 was identified as a potential driver of J. nigra adaptation.

Taxon
J. nigraJuglans nigrae0
Gene
WRKY41e0
Outcome
potential drivers of J. nigra's adaptatione0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Juglans nigra
Juglans nigra · TaxonPhoto by and (c)2007 Derek Ramsey ( Ram-Man ). Location credit to the Chanticleer Garden. · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“The genes WRKY41 and ERF012 were identified as potential drivers of J. nigra's adaptation.”

✓ located verbatim in the source · abstract, characters 1573–1663 · show in text ↑
Observationauthor interpretationoriginal resultmeasured

Claim · model-written summary

ERF012 was identified as a potential driver of J. nigra adaptation.

Taxon
J. nigraJuglans nigrae0
Gene
ERF012e0
Outcome
potential drivers of J. nigra's adaptatione0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Juglans nigra
Juglans nigra · TaxonPhoto by and (c)2007 Derek Ramsey ( Ram-Man ). Location credit to the Chanticleer Garden. · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“The genes WRKY41 and ERF012 were identified as potential drivers of J. nigra's adaptation.”

✓ located verbatim in the source · abstract, characters 1573–1663 · show in text ↑