The marattialean ferns (Order Marattiales, Class Polypodiopsida) comprise six genera and around 150 modern species found throughout tropical and subtropical regions (Encyclopaedia Britannica). They also have a significant fossil record.
Several phylogenetic analyses of the Marattiales have been published in recent years (e.g.Rothwell et al, 2018; Lehtonen et al, 2020; May et al, 2021), but they do not show good agreement. The current author is not qualified to evaluate the relative merits of these studies, but the results of the work by Lehtonen et al (2020) are highlighted here because they are based on a more recent and more complete molecular dataset than that used by Rothwell et al (2018) and present an older (more consistent with the fossil data) crown-group clade than that derived by May et al (2021). The following time tree is based on Lehtonen et al (2020):
Several phylogenetic analyses of the Marattiales have been published in recent years (e.g.Rothwell et al, 2018; Lehtonen et al, 2020; May et al, 2021), but they do not show good agreement. The current author is not qualified to evaluate the relative merits of these studies, but the results of the work by Lehtonen et al (2020) are highlighted here because they are based on a more recent and more complete molecular dataset than that used by Rothwell et al (2018) and present an older (more consistent with the fossil data) crown-group clade than that derived by May et al (2021). The following time tree is based on Lehtonen et al (2020):
Figure 1. Time tree of the stem-Marattiales
The presence of several polytomies in the above tree indicates that the phylogeny is poorly resolved.
While there are literature references (e.g. Liu et al, 2000) to the presence of marattialean fossils in the Early Carboniferous, no specific names seem to be available. The oldest well-documented marattialeans are of Late Carboniferous (Middle Pennsylvanian) age (Millay, 1982). The oldest stem-Marattiales represented on Figure 1 (Acitheca polymorpha, Eoangiopteris andrewsii and several species of Scolecopteris) are of this age. Acitheca polymorpha and a species of Scolecopteris are illustrated below, together with other members of the stem group for which images are available in the public domain (click on image for a larger view):
While there are literature references (e.g. Liu et al, 2000) to the presence of marattialean fossils in the Early Carboniferous, no specific names seem to be available. The oldest well-documented marattialeans are of Late Carboniferous (Middle Pennsylvanian) age (Millay, 1982). The oldest stem-Marattiales represented on Figure 1 (Acitheca polymorpha, Eoangiopteris andrewsii and several species of Scolecopteris) are of this age. Acitheca polymorpha and a species of Scolecopteris are illustrated below, together with other members of the stem group for which images are available in the public domain (click on image for a larger view):
Figure 2. Images of stem-Marattiales
The above images are ordered (approximately, given the unresolved nature of the phylogenetic tree shown in Figure 1) from most basal to most crownward, but no obvious trends can be observed.
The oldest known member of the crown-Marattiales is Marattiopsis crenulatus, described from Late Triassic (Rhaetian) sediments in Scania, Sweden (Lundblad, 1950; Rothwell et al, 2018). No public-domain image is available.
Taking into account the ghost lineage of at least 31 million years on the marattialean stem line (shown as a blue bar in Figure 1), the stem-to-crown transition of the Marattiales appears to have lasted between 138 and 158 million years, from the Early Carboniferous to the Late Triassic (Figure 1).
The oldest known member of the crown-Marattiales is Marattiopsis crenulatus, described from Late Triassic (Rhaetian) sediments in Scania, Sweden (Lundblad, 1950; Rothwell et al, 2018). No public-domain image is available.
Taking into account the ghost lineage of at least 31 million years on the marattialean stem line (shown as a blue bar in Figure 1), the stem-to-crown transition of the Marattiales appears to have lasted between 138 and 158 million years, from the Early Carboniferous to the Late Triassic (Figure 1).
References
Lehtonen, S., Poczai, P., Sablok, G., Hyvönen, J., Karger, D. N., & Flores, J. (2020). Exploring the phylogeny of the marattialean ferns. Cladistics, 36(6), 569-593.
Liu, Z. H., Hilton, J., & Li, C. S. (2000). Review on the origin, evolution and phylogeny of Marattiales. Chinese Bulletin of Botany, 17(专辑), 39.
Lundblad, A. B. (1950). Studies in the Rhaeto Liassic Floras of Sweden: Pteridophyta, Pteridaspermae, and Cycadophyta from the Mining District of Nw Scania. I. Almquist and Wiksell.
May, M. R., Contreras, D. L., Sundue, M. A., Nagalingum, N. S., Looy, C. V., & Rothfels, C. J. (2021). Inferring the total-evidence timescale of marattialean fern evolution in the face of model sensitivity. Systematic biology, 70(6), 1232-1255.
Millay, M. A. (1982). Studies of Paleozoic marattialeans: an early Pennsylvanian species of the fertile fern Scolecopteris. American Journal of Botany, 69(5), 728-733.
Rothwell, G. W., Millay, M. A., & Stockey, R. A. (2018). Resolving the overall pattern of marattialean fern phylogeny. American journal of botany, 105(8), 1304-1314.
Simpson, M. G. (2019). Evolution and diversity of vascular plants. Plant systematics, Third Edition (Academic Press), 75-130.
Liu, Z. H., Hilton, J., & Li, C. S. (2000). Review on the origin, evolution and phylogeny of Marattiales. Chinese Bulletin of Botany, 17(专辑), 39.
Lundblad, A. B. (1950). Studies in the Rhaeto Liassic Floras of Sweden: Pteridophyta, Pteridaspermae, and Cycadophyta from the Mining District of Nw Scania. I. Almquist and Wiksell.
May, M. R., Contreras, D. L., Sundue, M. A., Nagalingum, N. S., Looy, C. V., & Rothfels, C. J. (2021). Inferring the total-evidence timescale of marattialean fern evolution in the face of model sensitivity. Systematic biology, 70(6), 1232-1255.
Millay, M. A. (1982). Studies of Paleozoic marattialeans: an early Pennsylvanian species of the fertile fern Scolecopteris. American Journal of Botany, 69(5), 728-733.
Rothwell, G. W., Millay, M. A., & Stockey, R. A. (2018). Resolving the overall pattern of marattialean fern phylogeny. American journal of botany, 105(8), 1304-1314.
Simpson, M. G. (2019). Evolution and diversity of vascular plants. Plant systematics, Third Edition (Academic Press), 75-130.
Image credits – stem-Marattiales
- Header (King fern - Angiopteris evecta – photographed in Else Kientzler Botanical Garden, Sarchi Norte, Costa Rica): Photo by David J. Stang, CC BY-SA 4.0 <https://creativecommons.org/licenses/by-sa/4.0>, via Wikimedia Commons
- Figure 2 (Radstockia kidstonii): Field Museum of Natural History under Creative Commons license CC BY-NC 4.0
- Figure 2 (Sydneia manleyi): From Open Access article Psenicka, J., Zodrow, E. L., & DAngelo, J. A. (2013). Sterile foliage of fertile Sydneia manleyi and synangial chemistry (eusporangiate fern, Late Asturian, Canada): a new subfamily Sydneideae. Sciendo, Folia vol. 47, No. 1-2.
- Figure 2 (Acitheca polymorpha): From Open Access article Dangelo, J. A., Zodrow, E. L., & Psenicka, J. (2021). 3D chemometric model simulating the Acitheca polymorpha frond: implications for reconstructing carboniferous ferns (Marattiales, Canada). Sciendo, Folia vol. 55, No. 1–2.
- Figure 2 (Pectinangium xuanweiense): From Open Access article Zhou, Y., Guo, Y., Pšenička, J., Bek, J., Yang, S. L., & Feng, Z. (2021). A new marattialean fern, Pectinangium xuanweiense sp. nov., from the Lopingian of Southwest China. Review of Palaeobotany and Palynology, 295, 104500.
- Figure 2 (Scolecopteris sp.): Daderot, CC0, via Wikimedia Commons
- Figure 2 (Zhutheca densata, fossil): From Open Access article Guo, Y., Zhou, Y., Pšenička, J., Bek, J., Yang, S. L., & Feng, Z. (2020). Reinvestigation of the marattialean Zhutheca densata (Gu et Zhi) Liu, Li et Hilton from the Lopingian of Southwest China, and its evolutionary implications. Review of Palaeobotany and Palynology, 282, 104310.
- Figure 2 (Zhutheca densata, life restoration): From Open Access article Guo, Y., Zhou, Y., Pšenička, J., Bek, J., Yang, S. L., & Feng, Z. (2020). Reinvestigation of the marattialean Zhutheca densata (Gu et Zhi) Liu, Li et Hilton from the Lopingian of Southwest China, and its evolutionary implications. Review of Palaeobotany and Palynology, 282, 104310.
- Figure 2 (Danaeopsis fecunda): From Open Access article Zhou, N., Xu, Y., Li, L., Lu, N., An, P., Popa, M. E., ... & Wang, Y. (2021). Pattern of vegetation turnover during the end-Triassic mass extinction: Trends of fern communities from South China with global context. Global and Planetary Change, 205, 103585.
- Figure 2 (Danaeopsis marantacea): Hans Hagdorn, licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International License,
- Figure 2 (Qasimia yunnanica): From Open Access article Guo, Y., Zhou, Y., Bek, J., Yang, S. L., & Feng, Z. (2021). Qasimia yunnanica sp. nov., a marattialean fern with bivalvate synangia from the Lopingian of Southwest China. Review of Palaeobotany and Palynology, 293, 104497.