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of Triassic Equisetites sp. and<br />

Schizoneura paradoxa, Jurassic Equisetum<br />

columnare and living Equisetum has<br />

allowed the recognition of an evolutionary<br />

series confirming that Calamites and<br />

Equisetum are closely related and belong<br />

to the same lineage. Such a relationship<br />

was previously proposed by Good<br />

[Good, C.W., 1975. Pennsylvanian-age<br />

calamitean cones, elater-bearing spores,<br />

and associated vegetative organs. Palaeontogr.<br />

Abt. B 153, 28–99.] on the basis<br />

of morphological and anatomical similarities.<br />

Indeed, this study clearly shows<br />

that the ultrastructure of the spores of<br />

Triassic and Jurassic Equisetales is intermediate<br />

between that of Carboniferous<br />

Calamites and modern Equisetum,<br />

demonstrating that Equisetum evolved<br />

from Calamites. The present study illustrates<br />

how Calamites spores of the<br />

Calamospora type progressively<br />

evolved into the quite different spores of<br />

living Equisetum. Indeed, it would not<br />

have been possible to understand how<br />

such a transition occurred without this<br />

TEM study. Similarly, it underscores the<br />

proclivity of the Equisetales to develop<br />

unusual structures in both their spores<br />

and their reproductive organs. Moreover,<br />

it demonstrates that the ultrastructural<br />

features of equisetalean spores changed<br />

greatly during their long evolutionary<br />

history, whereas those of other Pteridophytes<br />

groups, such as ferns and lycopsids,<br />

remained unchanged. However, the<br />

most striking outcome of this comparative<br />

TEM study is the demonstration that<br />

spores of the Triassic horsetails show<br />

many ultrastructural similarities with<br />

spores of the Ophioglossaceae, a living<br />

family of primitive ferns. According to<br />

Lugardon and Brousmiche Delcambre<br />

[Lugardon, B., Brousmiche-Delcambre,<br />

C., 1994. Exospore ultrastructure in<br />

Carboniferous sphenopsids. In: Kurmann,<br />

M.H., Doyle, J.A. (Eds.), Ultrastructure<br />

of fossil spores and pollen, pp.<br />

53–66, Royal Botanic Gardens, Kew.],<br />

who made the same observations for the<br />

spores of Calamites, these similarities<br />

indicate that horsetails and ferns are<br />

closely related and have a common origin.<br />

These results are in agreement with<br />

those of a recent DNA analysis which<br />

shows that the horsetails and the ferns<br />

form a monophyletic group of plants.<br />

Moreover, they support and are in<br />

agreement with the phylogenetic analysis<br />

using cladistic principles of Stein et<br />

al. [Stein, W.E., Wight, D.C., Beck,<br />

C.B., 1984. Possible alternatives for the<br />

origin of Sphenopsida. Syst. Bot. 9 (1),<br />

102–118.] which suggests that the<br />

sphenophytes and the ferns as a whole<br />

are descended from a common ancestor,<br />

the Devonian complex Cladoxylopsida<br />

including the Hyeniales. The fact that<br />

the Hyeniales combine sphenophyte and<br />

fern-like features would explain why the<br />

spores of the ferns and those of the ancient<br />

sphenophytes have the same ultrastructural<br />

features. Moreover, this study<br />

indicates that spore ultrastructure retains<br />

ancestral features for a longer time than<br />

the other parts of the plants which<br />

evolve and thus change more rapidly.<br />

This study provides a further demonstration<br />

that spore ultrastructure, particularly<br />

of those preserved in situ, is of great<br />

value in researching the evolutionary<br />

and phylogenetic relationships of plants.<br />

2010010064<br />

现 生 和 化 石 买 麻 藤 孢 子 外 壁 超 微 结 构<br />

和 形 态 的 对 比 = Comparative pollen<br />

morphology and ultrastructure of modern<br />

and fossil gnetophytes. ( 英 文 ). Tekleva<br />

M V; Krassilov V A. Review of Palaeobotany<br />

and Palynology, 2009,<br />

156(1-2): 130-138 2 图 版 .<br />

A considerable disparity of pollen<br />

characters in the modern Gnetales (including<br />

the inaperturate and monosulcate<br />

germination types, the psilate, polyplicate<br />

and spinulose sculptural types)<br />

stands in sharp contrast with their ultrastructural<br />

uniformity. In all of the ultrustructurally<br />

studied living species of<br />

Ephedra, Welwitschia and Gnetum, the<br />

26

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