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Peer-reviewed journal article

Seed fossil record of Solanaceae revisited

Rocío DeannaiD, Aleksej Vladimirovič HvaljiD, Edoardo MartinettoiD, Sandra KnappiD, Eva‐Maria SadowskiiD, Steven R. ManchesteriD, Abel CamposiD, Vincent FernandeziD, Gloria E. BarbozaiD, Hervé SauquetiD, Ellen DeaniD, Tiina SärkineniD, Franco ChiariniiD, Gabriel BernardelloiD, Stacey D. SmithiD

Abstract

The fossil record for Solanaceae has a complex taxonomic history, with many species originally described in the family being subsequently shown to belong in other plant families. In this work, we present an in-depth analysis of the nightshade seed fossil record, which corresponds to the largest amount of fossil material for the family despite having relatively few described taxa. We re-examined 110 fossil seeds previously assigned either to one of the six taxa or unplaced within the family. All fossils were imaged and measured for 14 discrete and continuous characters, and a subset of 22 well-preserved specimens was additionally subjected to micro-CT scanning. In parallel, we scored the same set of characters for seeds from 354 extant taxa of Solanaceae. We carried out a multivariate analysis of the combined extinct and extant dataset to visualize the placement of fossil seeds within Solanaceae seed morphospace. This analysis revealed a wide spread of fossils across morphospace, including a variety of shapes, sizes and testal wall features. For one fossil taxon (Hyoscyosperma daturoides Deanna & S.D.Sm. gen. et sp. nov.), the micro-CT images confirmed the presence of an elaiosome, otherwise only known in the family from the genus Datura L. Based on these analyses, we re-circumscribe three species and describe six new taxa (four in new genera), doubling the number of fossil seed species in the family. The broad geographic and stratigraphic distribution of these diverse taxa, from Eocene North America and Europe to Pleistocene Eurasia, support the notion that the Solanaceae is likely a family of a late Cretaceous origin that attained nearly a cosmopolitan distribution by the Eocene.

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Claude Opus 5.5 · xhigh effort · 8 Nov 2026 · from the full text of version 1 (5 Jul 2025)

Morphometric and micro-CT analysis of 110 fossil and 354 extant Solanaceae seeds identifies 10 specimen-backed fossil species, including four new genera and a fossilized elaiosome, spanning Eocene to Holocene deposits.

Reconstruction of the biogeographic history and diversification timeline of nightshades (Solanaceae) has been limited by scarce, misassigned, or unrevised seed fossils. Combining 14-character morphometrics and micro-CT scanning across 110 fossil seeds and 354 extant species doubles the number of recognized fossil seed species and demonstrates a broad Northern Hemisphere distribution by the Eocene.

Key results

  1. ShownMultivariate morphospace analysis of 14 external and internal seed traits groups Solanaceae seed fossils into 17 morphological clusters within extant family diversity.

    Non-metric multidimensional scaling (NMDS, 2D stress < 0.2) of 464 specimens (110 fossils and 354 extant species across all but 3 of 99 genera) found 12 of 14 characters significantly informative (P = 0.001-0.04), identifying 10 clusters with physical fossil specimens and 7 clusters with literature-only records. · Scoring of extant and fossil seeds; Morphospace analysis; Morphospace of fossil and extant Solanaceae…

  2. ShownHigh-resolution X-ray micro-CT preserves internal embryo curvature and reveals a fossilized seed-coat elaiosome in Hyoscyosperma daturoides gen. et sp. nov.

    Among 22 fossil and 14 extant seeds scanned by micro-CT, 3 fossils preserved internal anatomy: a curved embryo in Solanoides dorofeevii (MB.Pb.1993/6802), a coiled embryo in BIN K453/175, and a hilar-chalazal elaiosome in Hyoscyosperma daturoides (holotype BIN K453/133, 27.8-0.01 Ma), a trait otherwise known in extant Solanaceae only in Datura. · X-ray computed tomography; Micro-CT of fossil and extant Solanaceae seeds; TAXONOMIC DESCRIPTIONS; Evolutionary and biogeographic implications of…

  3. ShownTaxonomic revision of the 10 specimen-supported clusters doubles the described diversity of Solanaceae seed fossils to 10 species across 9 genera.

    The revision emends 3 species (Solanum foveolatum, Solanispermum reniforme, Nephrosemen reticulatum), establishes 1 new combination (Albionites arnensis from Solanum arnense), and describes 4 new genera (Hyoscyosperma, Solanoides, Albionites, Thanatosperma) and 6 new species (Hyoscyosperma daturoides, Solanum miocenicum, Hyoscyamus undulatus, Solanoides dorofeevii, Capsicum pliocenicum, Thanatosperma minutum). · INTRODUCTION; TAXONOMIC DESCRIPTIONS; DISCUSSION

  4. ShownStratigraphic and geographic distributions show Solanaceae seeds present in both Europe and North America by the middle Eocene (~48 Ma) with peak fossil diversity at the Pliocene-Pleistocene boundary.

    Globose and flattened seed fossils appear at 48-46 Ma in the UK (Solanispermum reniforme, Albionites arnensis) and 47.8 Ma in Oregon, USA (Nephrosemen reticulatum), while Neogene radiations across Eurasia produce peak taxonomic richness around 2.58 Ma. · TAXONOMIC DESCRIPTIONS; DISCUSSION; Evolutionary and biogeographic implications of…

Methods, data and reproducibility

Scored 7 continuous and 7 discrete seed characters across 110 fossil specimens (95 physical museum specimens, 15 literature records) and 354 extant Solanaceae species, merging stereomicroscope stacks (254 composites) for ImageJ cell-wall morphometrics and scanning 22 fossils and 14 extant seeds with Zeiss Versa 520 micro-CT. Ran NMDS with Gower's dissimilarity in R (VEGAN) to delimit clusters and revise taxonomy.

  • Morphometric dataset of 14 discrete and continuous seed characters recorded in the PROTEUS database for fossil and extant Solanaceae · plant (other) (Solanaceae: 110 fossil specimens and 354 extant species) · n = 464 specimens (95 physical fossils, 15 literature records, 354 extant species)
  • High-resolution X-ray micro-computed tomography (micro-CT) volume scans of fossil and extant Solanaceae seeds · plant (other) (Solanaceae) · n = 22 fossil specimens and 14 extant species scanned by micro-CT
Methods & tools
HeliconFocus Pro, ImageJ, PROTEUS database, Volume Graphics Studio Max 3.4, VEGAN (R package), GGPLOT2 (R package), Zeiss Versa 520 micro-CT
Reagents & environment
Volume Graphics Studio Max 3.4; HeliconFocus Pro; ImageJ; R packages VEGAN (metaMDS) and GGPLOT2
Reproducibility gaps
The Open Science Framework (OSF) repository link for merged photomicrographic composites and supplemental micro-CT video is noted as to be provided after revisions.
Limitations (4)
  • Internal anatomical traits (embryo shape and elaiosome attachment) were recovered from only 3 of the 22 micro-CT-scanned fossil seeds due to variable preservation and pyritization. · stated by the authors
  • Seven of the 17 NMDS clusters consisted solely of fossils scored from incomplete literature descriptions without physical specimens examined and could not be formally revised. · stated by the authors
  • The Solanaceae fossil record lacks any representation from Africa and Oceania, large (>3.65 mm) or winged seeds, and fossils assignable to extant capsular subfamilies or tribes (Schwenckieae, Petunieae, Nicotianoideae). · stated by the authors
  • Taphonomic compression during fossilization may alter seed flatness, requiring caution when weighting lateral compression as a diagnostic character. · stated by the authors

Who should read:Read this if you study Solanaceae evolution, paleobotanical calibration of plant phylogenies, or multivariate morphometrics and micro-CT analysis of fossil seeds.

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