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The earliest record of the genus Cola (Malvaceae sensu lato: Sterculioideae) from the Late Oligocene (28–27 Ma) of Ethiopia and leaf characteristics within the genus

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Abstract

A fossil leaf compression from the Late Oligocene (28–27 Ma) of northwestern Ethiopia is the earliest record of the African endemic moist tropical forest genus Cola (Malvaceae sensu lato: Sterculioideae). Based on leaf and epidermal morphology, the fossil is considered to be very similar to two extant Guineo-Congolian species but differences warrant designation of a new species. This study also includes a review of the fossil record of Cola, a comprehensive summary of leaf characteristics within several extant species of Cola, Octolobus, and Pterygota, and a brief discussion of the paleogeographic implications of the fossil species affinity and occurrence in Ethiopia.

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Acknowledgment

We would like to thank the Authority for Research and Conservation of Cultural Heritage, the Ministry of Culture and Tourism, Ethiopia, and especially Ato Jara for permission to conduct our continuing research in northwestern Ethiopia, and the Director Mamitu Yilga and staff of the National Museum, Addis Ababa, and the Gondar ARCCH and Chilga Ministry of Culture and Sports Affairs for logistical support. We thank the Missouri Botanical Garden, the Royal Botanic Gardens at Kew, and the collectors of the herbaria specimens examined for assistance and access to their collections. We are grateful to the National Museums of Kenya, the Baringo Paleontological Research Project, the East African Herbarium, and Christine Kabuye for their collaborative support of work in Kenya. This project was funded by grants from the National Science Foundation (EAR-0001259, EAR-0240251, and EAR-0617306), the National Geographic Society, and the Dallas Paleontological Society. Tillehun Selassie, Misege Birara, Habtewold Habtemichael, Mesfin Mekonnen, and Drs. Ambachew Kebede and Aklilou Asfaw provided valuable field assistance. We thank Dr. Martin Cheek for generously sharing information about Cola and other sterculioids, for providing advice, and for supplying cuticle specimens from the Royal Botanic Gardens at Kew. We are grateful to Dr. Thomas Denk for images of the Cameroon sterculioid fossils housed at the Swedish Museum of Natural History. We also appreciatively acknowledge help from Yohannes Desta, Yeshiwass Sitotaw, Gebremeskel Ayele, Elias Addissu, and Teshome Yohannes at Chilga and laboratory assistance from Kathryn Larson.

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Authors and Affiliations

  1. Fort Worth Museum of Science and History, 1600 Gendy Street, Fort Worth, TX, 76107-4062, USA

    Aaron David Pan

  2. Roy M. Huffington Department of Earth Sciences, Southern Methodist University, 750395, Dallas, TX, 75275-0395, USA

    Bonnie F. Jacobs

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  1. Aaron David Pan
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Correspondence to Aaron David Pan.

Appendix

Appendix

Leaf and epidermal characteristics and states

  1. 1.

    Leaf type: (0) simple, (1) palmately compound, (2) pinnately compound

  2. 2.

    Leaf(let) shape: (0) elliptic, (1) obovate, (2) ovate, (3) oblong, (4) special (needles, awls, etc.)

  3. 3.

    Leaf(let) attachment: (0) petiolate [petiolulate for compound-leaved species], (1) subsessile/sessile

  4. 4.

    Leaf(let) texture: (0) chartaceous (1) semicoriaceous, (2) coriaceous

  5. 5.

    Leaf(let) margin type: (0) entire, (1) lobed, (2) toothed

  6. 6.

    Leaf(let) base angle: (0) acute, (1) obtuse, (2) wide obtuse

  7. 7.

    Leaf(let) base shape: (0) cuneate/concave/decurrent, (1) convex/rounded, (2) truncate, (3) subcordate/cordate/lobate

  8. 8.

    Leaf(let) apex morphology: (0) acute, (1) subacuminate, (2) acuminate, (3) obtuse/rounded/slightly emarginate, (4) deltate/cuspidate, (5) emarginate

  9. 9.

    Primary venation: (0) pinnate, (1) tri-nerved at the base, (2) palmately veined

  10. 10.

    Number of secondary vein pairs: (0) 0–5; (1) 6–10; (2) 11–15; (3) >16

  11. 11.

    Secondary vein categories: (0) brochidodromous, (1) eucamptodromous, (2) festooned brochidodromous, (3) cladodromous, (4) reticulodromous, (5) toothed-craspedodromous, (6) toothed-semicraspedodromous, (7) toothed-festooned semicraspedodromous

  12. 12.

    Tertiary venation: (0) random reticulate, (1) opposite percurrent, (2) alternate percurrent, (3) mixed opposite/alternate percurrent

  13. 13.

    Abaxial epidermal cell shape: (0) isodiameteric, (1) rectangular

  14. 14.

    Abaxial cell size: indicated by length (first set of numbers) and width ranges, rounded to the nearest 0.5 µm

  15. 15.

    Abaxial epidermal cell arrangement: (0) random (a combination of any of the following cellular arrangements), (1) nonrandom–tetragonal, (2) nonrandom–pentagonal, (3) nonrandom–hexagonal, (4) nonrandom–polyagonal (>6-sided cells), (5) nonrandom–linear

  16. 16.

    Abaxial anticlinal cell wall pattern: (0) straight, (1) rounded, (2) undulate

  17. 17.

    Abaxial anticlinal cell wall shape of undulation: (0) U, (1) V, (2) Ω

  18. 18.

    Abaxial epidermal cell surface ornamentation: (0) absent, (1) stippled, (2) striate

  19. 19.

    Abaxial stoma: (0) absent, (1) present

  20. 20.

    Abaxial stomatal complex type: (0) polycytic types (anomocyptic, cyclocytic), (1) anisocytic, (2) paracytic types (paracytic, brachyparacytic, amphibrachyparacytic, hemiparacytic), (3) tetracytic types (staurocytic, anomotetracytic, paratetracytic, brachyparatetracytic)

  21. 21.

    Abaxial stomatal complex chains or clusters: (0) absent, (1) present

  22. 22.

    Abaxial stomatal ornamentation: (0) absent, (1) striate, (2) papillate, (3) thickened areas on periclinal wall

  23. 23.

    Abaxial guard cells: (0) level, (1) sunken, (2) raised

  24. 24.

    Number of abaxial trichomes types: (0) glabrous/absent (no trichomes present), (1) one, (2) two, (3) three, (4) four, (5) five or more

  25. 25.

    Abaxial hair types present: (0) unicellular simple, (1) unicellular glandular, (2) multicellular glandular, (3) stellate/peltate (usually only the polygonal base is preserved),

  26. 26.

    Abaxial trichome dominance: (0) glandular trichomes predominate [≥70% glandular], (1) no trichome type predominates (29–69%), (2) stellate/peltate trichomes predominate [≥70% stellate/peltate], (3) simple hairs predominate (≥70% simple)

  27. 27.

    Abaxial glandular hair pairs: (0) absent, (1) present

  28. 28.

    Adaxial epidermal cell shape: (0) isodiameteric, (1) rectangular

  29. 29.

    Adaxial cell size: indicated by length (first set of numbers) and width ranges, rounded to the nearest 0.5 µm

  30. 30.

    Adaxial epidermal cell arrangement: (0) random (a combination of any of the following cellular arrangements), (1) nonrandom–tetragonal, (2) nonrandom–pentagonal, (3) nonrandom–hexagonal, (4) nonrandom–polyagonal (>6-sided cells), (5) nonrandom–linear

  31. 31.

    Adaxial anticlinal cell wall pattern: (0) straight, (1) rounded, (2) undulate

  32. 32.

    Adaxial anticlinal cell wall shape of undulation: (0) U, (1) V, (2) Ω

  33. 33.

    Adaxial epidermal cell surface ornamentation: (0) absent, (1) stippled, (2) striate

  34. 34.

    Adaxial stoma: (0) absent, (1) present

  35. 35.

    Adaxial stomatal complex type: (0) polycytic types (anomocyptic, cyclocytic), (1) anisocytic, (2) paracytic types (paracytic, brachyparacytic, amphibrachyparacytic, hemiparacytic), (3) tetracytic types (staurocytic, anomotetracytic, paratetracytic, brachyparatetracytic)

  36. 36.

    Abaxial stomatal complex chains or clusters: (0) absent, (1) present

  37. 37.

    Adaxial stomatal ornamentation: (0) absent, (1) striate, (2) papillate, (3) thickened areas on periclinal wall

  38. 38.

    Adaxial guard cells: (0) level, (1) sunken, (2) raised

  39. 39.

    Number of adaxial trichomes types: (0) glabrous/absent (no trichomes present), (1) one, (2) two, (3) three, (4) four, (5) five or more

  40. 40.

    Adaxial hair types present: (0) unicellar simple, (1) unicellular glandular, (2) multicellular glandular, (3) stellate/peltate (usually only the polygonal base is preserved)

  41. 41.

    Adaxial trichome dominance: (0) glandular trichomes predominate (≥70% glandular), (1) no trichome type predominates (29–69%), (2) stellate/peltate trichomes predominate (≥70% stellate/peltate), (3) simple hairs predominate (≥70% simple)

  42. 42.

    Adaxial glandular hair pairs: (0) absent, (1) present

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Pan, A.D., Jacobs, B.F. The earliest record of the genus Cola (Malvaceae sensu lato: Sterculioideae) from the Late Oligocene (28–27 Ma) of Ethiopia and leaf characteristics within the genus. Plant Syst Evol 283, 247–262 (2009). https://doi.org/10.1007/s00606-009-0225-1

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