Abstract
Complementary to our previous project on the molecular phylogeny of Camphorosmeae, the leaf anatomy of ca. 35 species including all non-Australian and selected Australian species was studied by use of light microscopy. Nine anatomical leaf types were described, compared to previous classifications, and discussed with regard to their putative evolution on the background of phylogenetic trees. Particular emphasis was given to the relationships between the C3 and C4 leaf types: Chenolea type (C3), Eokochia type (C3), Neokochia type (C3), Sedobassia type (C3/C4 intermediate), Bassia prostrata type (C4), B. muricata type (C4), B. eriantha type, B. lasiantha type (C4), Camphorosma type (C4). The main results and conclusions were: (1) Two unusual new C3 leaf types: Chenolea with microfenestrate chlorenchyma, Eokochia with unique complex vascular bundles; (2) Sedobassia interpreted as anatomically C3/C4 intermediate by kranz-like bundle sheath cells is the first C3/C4 intermediate in Camphorosmeae and found in a derived position; (3) Neokochia type detected as the likely starting point for all four C4 leaf types and for the C3/C4 intermediate; (4) hypodermis of C4 types originated from outermost chlorenchyma layer of C3 types and lost multiple times during further evolution; (5) atriplicoid Bassia. lasiantha type without water storage tissue evolved from kochioid B. muricata type; (6) two independent gains of C4 photosynthesis, one in Bassia and one in Camphorosma; (7) depending on the lineage, leaf architecture remains comparatively stable (Australian Camphorosmeae) or shows an unexpected plasticity (Bassia scoparia group).
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Notes
Genus Austrobassia Ulbr. had once ca. 70 ssp. It is now discarded, with most species included in Sclerolaena and Maireana.
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Acknowledgments
We are indebted to many colleagues and institutions who supplied herbarium specimens, seeds for cultivation, wet-conserved material or photographs, e.g., Maria Lomonosova (Novosibirsk), Valentin Golub (Togliatti), Antonina Butnik (Almaty), Norbert Juergens (Hamburg), Alexander Sukhorukov (Moscow), Steffi Ickert-Bond (Fairbanks), M. Schnittler (Greifswald), Manolo Gil González (Lanzarote) and José María Escolano (Zaragoza). We are also most thankful to Sabine Bringmann (Kassel) for the careful cultivation of plants, Irene Diebel for scrupulous laboratory work, Dagmar Müller (Kassel) and Doris Franke (Mainz) for support given to the preparation of plates. Willibald Stichler (Neuherberg) and Gerald E. Edwards (Pullman) provided some new carbon isotope data. With gratitude we acknowledge the information given by Elena Voznesenskaya (St. Petersburg), Gerry Edwards (Pullmann) and Rowan Sage (Toronto) about unpublished results concerning the photosynthesis in Sedobassia. Finally we thank Elena Voznesenskaya and an anonymous reviewer for useful critical comments and Ian C. Hedge (Edinburgh) for linguistic improvement of the manuscript. Parts of the project were financially supported by the German Science Foundation (DFG, grant WE 1830/2-1) and different travel grants (mostly by DFG) to HF.
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Freitag, H., Kadereit, G. C3 and C4 leaf anatomy types in Camphorosmeae (Camphorosmoideae, Chenopodiaceae). Plant Syst Evol 300, 665–687 (2014). https://doi.org/10.1007/s00606-013-0912-9
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DOI: https://doi.org/10.1007/s00606-013-0912-9