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Fig. 6. Lignins in lycophytes. (A) Detail from Selaginella martensii frond.
(B) Ma¨ le staining of transverse sections of Selaginella stems. Red and brown colours
indicate the presence of S and G lignins, respectively. Bars ¼ 300 m m. (C) Two-
dimensional HSQC NMR spectra of acetylated cellulolytic enzyme lignins from
Selaginella whole stems, revealing the presence of syringyl lignins. C box is taken
from Weng et al. (2011) , reproduced with permission from ASPB.
derivatives have not been found in the extinct arborescent lycophyte Sigil-
laria ovata ( Logan and Thomas, 1987 ). However, lignins, and more specifi-
cally, syringyl lignins, were reported by chemical degradative methods in the
herbaceous Isoetes and Huperzia ( Towers and Gibbs, 1953; Towers and
Maass, 1965 ) and, more conclusively, by chemical depolymerizing methods
including thioacidolysis in Selaginella and Isoetes ( Espi ˜ eira et al., 2011 ).
The presence of S units in some lycopods is well known and has been
associated with the existence in the xylem of angiosperm-like vessels
( Schneider and Carlquist, 2000 ). Selaginella is a representative of the Lyco-
podiophyta and one of the earliest divergent clades of extant vascular plants
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