Networks of Highly Branched Stigmarian Rootlets Developed on the First Giant Trees
Networks of highly branched stigmarian rootlets developed on the first giant trees Alexander J. Hetheringtona, Christopher M. Berryb, and Liam Dolana,1 aDepartment of Plant Sciences, University of Oxford, Oxford OX1 3RB, United Kingdom; and bSchool of Earth and Ocean Sciences, Cardiff University, Cardiff CF10 3AT, United Kingdom Edited by Peter R. Crane, Yale School of Forestry and Environmental Studies, New Haven, CT, and approved February 29, 2016 (received for review July 22, 2015) Lycophyte trees, up to 50 m in height, were the tallest in the that rootlet abscission resembled foliar abscission (17, 18), obser- Carboniferous coal swamp forests. The similarity in their shoot and vations on well-preserved fossil rhizomorph apices (30–32), and root morphology led to the hypothesis that their rooting (stig- their interpretation within a phylogenetic context (31, 33) led to a marian) systems were modified leafy shoot systems, distinct from complete revival of Schimper’s (27) modified shoot hypothesis. the roots of all other plants. Each consists of a branching main axis The only living relatives of these Carboniferous giant trees are covered on all sides by lateral structures in a phyllotactic arrange- small herbaceous plants in the genus Isoetes (24, 33–35). The rooting ment; unbranched microphylls developed from shoot axes, and system of Isoetes also consists of a rhizomorph meristem that de- largely unbranched stigmarian rootlets developed from rhizomorphs velops rootlets in a regular rhizotaxy (31, 36, 37). Aside from the axes. Here, we reexamined the morphology of extinct stigmarian reduction and modification of the rhizomorph, the rooting systems systems preserved as compression fossils and in coal balls from the of Isoetes and the tree lycopsids are morphologically similar (4, 19, Carboniferous period.
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