Seed morphology of the paleotropical tribe Paropsieae (Passifloraceae, Malpighiales), and paleobotanical implications

  • Flavien Lagrange CR2P – Centre de Recherche en Paléontologie – Paris, MNHN – Sorbonne Université – CNRS, 43 Rue Buffon, 75005 Paris, France
  • Camila Martínez Área de Sistemas Naturales y Sostenibilidad, Universidad EAFIT, Medellín, Colombia
  • Cédric Del Rio CR2P – Centre de Recherche en Paléontologie – Paris, MNHN – Sorbonne Université – CNRS, 43 Rue Buffon, 75005 Paris, France
Keywords: seeds, taxonomy, anatomy, paleobotany, botany


The fossil record of the diverse subfamily Passifloroideae (>750 species and 17 genera) is relatively poor. Despite the distinctiveness of its leaves (glandular and often emarginate), most of the fossils from this group have been described from seeds. Fossil seeds have been recovered from Europe, and North and South America. A lack of information on seed morphology for all the genera and tribes of this subfamily has prevented a tribe-level identification of the fossils and a better understanding of their biogeographic patterns. The Passifloroideae is divided into three tribes: Passifloreae with 10 genera, Paropsieae with six genera and the monotypic Jongkindieae. This study provides new descriptions for 15 species from 5 genera from the mostly Afrotropical tribe Paropsieae based on herbarium material, and introduces an online seed database and a key for 100 species of Passifloroideae compiled from literature and direct observations. Our study shows a low morphological diversity among the seeds of Paropsieae in comparison to a much larger diversity within Passifloreae. Some rare morphologies are only present in Passifloreae and can be used to assign seeds to this tribe. Within the Paropsieae, Androsiphonia has seed that are very distinct from those in the other genera in the tribe and also from the rest of the subfamily. The genus Paropsia exhibits two main morphotypes, while the genera Barteria, Paropsiopsis and Smeathmannia have very similar seeds with a highly conserved morphology. These results suggest that living or fossil Paropsieae cannot be identified confidently based solely on seed characters.


Angiosperm Phylogeny Group 1998. An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Garden 85 (4): 531–53.

Angiosperm Phylogeny Group 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141 (4): 399–436.

Angiosperm Phylogeny Group 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161 (2): 105–121.

Angiosperm Phylogeny Group 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181 (1): 1–20.

Benedict J.C. 2015. A new technique to prepare hard fruits and seeds for anatomical studies. Applications in Plant Sciences 3 (10): Art. 1500075.

Bonilla Morales M.M., Aguirre Morales A.C. & Agudelo Varela O.M. 2015. Morfología de Passiflora: una guía para la descripción de sus especies. Revista de Investigación Agraria y Ambiental 6: 91–109.

Breteler F.J. 1999. Barteria Hook f. (Passifloraceae) revised. Adansonia 21 (2): 307–318.

Breteler F.J. 2003. Novitates Gabonenses 48. A new species of Paropsia (Passifloraceae) from Gabon. Adansonia 25 (2): 247–249.

Breteler F.J., Breman F.C., Lei D. & Bakker F.T. 2022. Wrong flowers? The evolutionary puzzle of Jongkindia (Passifloraceae s.l.), a new monotypic genus and tribe from Liberia, West Africa. Webbia 77: 229–246.

D’Apolito C., Jaramillo C. & Harrington G. 2021. Miocene palynology of the Solimões Formation (Well 1-AS-105-AM), western Brazilian Amazonia. Smithsonian Contributions to Paleobiology 105: 1–134.

de Vos J. & Breteler F.J. 2009. A revision of the African genera Paropsiopsis and Smeathmannia (Passifloraceae – Paropsieae), including a new species of Paropsiopsis from Cameroon. Edinburgh Journal of Botany 66 (1): 27–49.

de Wilde W.J.J.O. 1971. The systematic position of tribe Paropsieae, in particular the genus Ancistrothyrsus, and a key to the genera of Passifloraceae. Blumea: Biodiversity, Evolution and Biogeography of Plants 19 (1): 99–104.

de Wilde W.J.J.O. 1972. Passifloraceae. Flora Malesiana – Series 1, Spermatophyta 7: 405–434.

Espinoza T.E.B., Jørgensen P.M. & MacDougal J.M. 2018. A taxonomic revision of Passiflora sect. Xerogona (Passifloraceae) using principal component analysis. Annals of the Missouri Botanical Garden 103: 258–313.

Feuillet C. 2020. Ancistrothyrsus scopae (Passifloraceae), a new species from Amazonian Brazil and Guyana, with keys to the genus and species. Phytotaxa 438 (3): 207–212.

Feuillet C. & MacDougal J.M. 2007. Passifloraceae. In: Kubitzki K. (ed.) The Families and Genera of Vascular Plants, Vol. 9, Flowering Plants. Eudicots: 270–281. Springer, Berlin, Heidelberg.

Gilbert L.E. & MacDougal J.M. 2000. Passiflora microstipula, a new species of Passifloraceae from Southeast Mexico. Lundellia 3: 1–5.

Hermsen E. 2021. Review of the fossil record of Passiflora, with a description of new seeds from the Pliocene Gray Fossil Site, Tennessee, USA. International Journal of Plant Sciences 182 (6): 533–550.

Hermsen E. 2023. Pliocene seeds of Passiflora subgenus Decaloba (Gray Fossil Site, Tennessee) and the impact of the fossil record on understanding the diversification and biogeography of Passiflora. American Journal of Botany 110 (3): e16137.

Hutchinson J. & Daziel J.M. 1952. Flora of West tropical Africa, Vol. 1. Crown Agents for Overseas Governments and Administrations, London.

Jørgensen P.M., Lawesson J.E. & Holm-Nielsen L.B. 1984. A guide to collecting passionflowers. Annals of the Missouri Botanical Garden 71: 1172–1174.

Jørgensen P.M., Holm-Nielsen L.B. & Lawesson J.E. 1987. New species of Passiflora subg. Plectostemma and subg. Tacsonia (Passifloraceae). Nordic Journal of Botany 7: 127–133.

Jørgensen P. & Vásquez R. 2009. A revision of Passiflora sections Insignes and Inkea (Passifloraceae). Anales del Jardín Botánico de Madrid 66: 35–53.

Kerner A., Bouquin S., Portier R. & Vignes Lebbe R. 2021. The 8 years of existence of Xper3: State of the art and future developments of the platform. Biodiversity Information Science and Standards 5: e74250.

Kiew R., Chung R.C.K., Saw L.G. & Soepadmo E. 2018. Flora of peninsular Malaysia. Series II: Seed plants. Volume 7. Malayan Forest Records 49: 1–321.

Krosnick S.E., Ford A.J. & Freudenstein J.V. 2009. Taxonomic revision of Passiflora subgenus Tetrapathea including the monotypic genera Hollrungia and Tetrapathea (Passifloraceae), and a new species of Passiflora. Systematic Botany 34 (2): 375–385.

Krosnick S.E., Porter-Utley K.E., MacDougal J.M., Jørgensen P.M. & McDade L.A. 2013. New insights into the evolution of Passiflora subgenus Decaloba (Passifloraceae): phylogenetic relationships and morphological synapomorphies. Systematic Botany 38 (3): 692–713.

Lozada-Pérez L. & Gutiérrez J.F.R. 2016. A new species of Passiflora L. (Passifloraceae) from Guerrero and Oaxaca, México. Phytotaxa 263 (3): 3 June 2016.

Maas P.J.M., Baas P., Christenhusz M.J.M., Clarkson J.J., Koek-Noorman J., Mennega A.M.W., Tokuoka T., Van Der Bank M., Van Der Ham R.W.J.M., Van Marle E.-J., Westra L.Y.TH. & Chase M.W. 2019. “Unknown yellow”: Pibiria, a new genus of Passifloraceae with a mixture of features found in Passifloroideae and Turneroideae. Botanical Journal of the Linnean Society 189 (4): 397–407.

MacDougal J.M. 1994. Revision of Passiflora Subgenus Decaloba Section Pseudodysosmia (Passifloraceae). Systematic Botany Monographs 41: 1–146.

MacDougal J.M. 2001. Two new species of passionflower (Passiflora, Passifloraceae) from Southwestern Mexico. Novon 11 (1): 69–75.

Martínez C. 2017. Passifloraceae seeds from the late Eocene of Colombia. American Journal of Botany 104 (12): 1857–1866.

Mezzonato-Pires A.C., Mendonça C.B.F., Milward-de-Azevedo M.A. & Gonçalves-Esteves V. 2017. The taxonomic significance of seed morphology in the Passiflora subgenus Astrophea (Passifloraceae). Acta Botanica Brasilica 31 (1): 68–83.

Mezzonato-Pires A.C., de Souza Freitas G.H.G., Mendonça C.B.F. & Gonçalves-Esteves V. 2022. The systematic value of pollen morphology in the tribe Paropsieae (Passifloraceae sensu stricto). Review of Palaeobotany and Palynology 299: Art. 104606.

Morton J.F. 1987. Passion fruit. In: Morton J.F. (ed.) Fruits of Warm Climates: 320–328. Florida Flair Book, Miami, FL.

Ngumbau V., Nyange M., Dai C., Zhong Z-X., Wei N., Malombe I., Hu G.-W. & Wang Q.-F. 2017. Adenia angulosa (Passifloraceae), a new species from coastal forests of Kenya and Tanzania. Phytotaxa 313 (1): 137–142.

Olivier D. 1871. Flora of Tropical Africa, Vol II. L. Reeve and co., London.

Palazzesi L., Barreda V.D., Cuitiño J.I., Guler M.V., Tellería M.C. & Ventura Santos R. 2014. Fossil pollen records indicate that Patagonian desertification was not solely a consequence of Andean uplift. Nature Communications 5: Art. 4558.

Pérez-Cortéz S. 2007. Atlas morfológico de Semillas en Especies del Género Passiflora L. Presentes en Venezuela. Fundación Instituto Botánico de Venezuela, Caracas, Venezuela.

Pérez-Cortéz S., Tillett S. & Escala M. 2002. Estudio Morfológico de la semilla de 51 especies del género Passiflora L. Acta Botánica Venezuelica 25 (1): 67–96.

Pérez-Cortéz S., Escala M. & Tillett S. 2005. Anatomia de la cubierta seminal en ocho especies de Passiflora L., subgénero Passiflora. Acta Botánica Venezuélica 28: 337–348.

Peréz-Cortéz S., Escala M. & Tillett S. 2009. Morfoanatomia de la cubierta seminal en siete especies de Passiflora L., subgénero Passiflora (Passifloraceae). Hoehnea 36: 131–137.

Perrier de la Bathie H. 1945. Passifloracées. In: Humbert H. (ed.) Flore de Madagascar et des Comores. Tananarive Imprimerie officielle, Madagascar.

POWO 2023. Plants of the world online. Facilitated by the Royal Botanic Gardens, Kew. Available from [accessed 3 May 2024].

Rasband W.S. 2016. ImageJ. US National Institutes of Health, Bethesda, MD. Available from [accessed 3 May 2024].

Schemelzer H.H. & Gurib-Fakim A. 2008. Plant Resources of Tropical Africa 11 (1): Medicinal Plants 1. PROTA Foundation, Wageningen, Netherlands.

Sleumer H. 1970. Le genre Paropsia Noronha ex Thouars (Passifloraceae). Bulletin du Jardin botanique National de Belgique / Bulletin van de National Plantentuin van België 40 (1): 49–75.

Stapff O. 1904. Contribution to the flora of Liberia. Botanical Journal of the Linnean Society 61: 79–115.

Stevens P.F. 2001 onwards. Angiosperm Phylogeny Website. Ver. 14, July 2017. Available from [accessed 3 May 2024].

Tokuoka T. 2012. Molecular phylogenetic analysis of Passifloraceae sensu lato (Malpighiales) based on plastid and nuclear DNA sequences. Journal of Plant Research 125: 489–497.

Ulmer T. & MacDougal J.M. 2004. Passiflora: Passionflowers of the World. Timber Press, Portland, USA

Ung V., Dubus G., Zaragüeta-Bagils R. & Vignes-Lebbe R. 2010. Xper2: introducing e-taxonomy. Bioinformatics 26 (5): 703–704.

Vanderplank J. & Ochoa J. 2020. Passiflora insoliti. Curtis’s Botanical Magazine 37 (1): 131–138.

Vanderplank J. & Zappi D. 2011. Passiflora cristalina, a striking new species of Passiflora (Passifloraceae) from Mato Grosso, Brazil. Kew Bulletin 66: 149–153.

How to Cite
Lagrange, F., Martínez, C., & Del Rio, C. (2024). Seed morphology of the paleotropical tribe Paropsieae (Passifloraceae, Malpighiales), and paleobotanical implications. European Journal of Taxonomy, 943(1), 1–23.
Research article