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The Anatomy and Morphology of Tmesipteris The Anatomy and Morphology of Tmesipteris. BY M. G. SYKES. Girton College, Cambridge; Bathurst Student, Newnkam College, Cambridge. With Plates VII and Vm and thirteen FigureB in the Text. HE material on which the following investigation is based was kindly T sent by Professor Thomas, of Auckland in New Zealand, at the request of Professor Seward ; it comprised two forms, differing only slightly in appearance and structure, but separated by Dangeard 1 as two species: T. tannensis (Fig. I) and T. lanceolata (Fig. II). My thanks are due to Professor Seward, both for liis kindness in obtaining the material for me and for his helpful interest in my work and useful suggestions during its progress. I. HABITAT AND DISTRIBUTION. Tmesipteris is found living as an epiphyte on tree-ferns in New Zealand, Australia, and Polynesia.8 Each plant consists of an aerial portion and a rhizome, or subterranean region, but has no roots. In my specimens the aerial part varied from three to eight inches in length. It is very difficult to extricate any but small pieces of the rhizome from the tree-fern roots with which it intertwines.3 I therefore received only small broken posterior portions of the rhizome and short lengths of the anterior region, the latter being attached to the aerial shoots.* Our knowledge of Tmesipteris is based entirely on the adult plant: spores have never been germinated, so nothing is known of the gameto- phyte. It seems probable that the plant has a saprophytic mode of life, and the occurrence of a fungus8 growing in the cortical cells of the rhizome supports this suggestion. ... 1 Dangeard, Le Botaniste, ii, p. 316. ' Baker, Fern Allies, 297-30. 3 Dangeard, Le Botaniste, ii, p. 168. * Jennings and Hall, 1891, PI. I, Fig. 4. 1 Dangeard, Le Botaniste, ii, pp. 233-30, PI. IX, Figs. 15, 16. [Annals of.BoUny, Vol. XXIL No. LXXXV. January, 1908.] Downloaded from https://academic.oup.com/aob/article-abstract/os-22/1/63/201535 by Queen Mary University of London user on 24 March 2018 6\ Sykes.—T/ie Anatomy and Morphology of Tmesipleris. II. EXTERNAL FEATURES. (x) Vegetative regions. The underground portion of Tmesipteris runs more or less horizontally and forks at intervals. Generally one of the branches thus formed remains small, while the other continues as the main axis. Rhizome apices were very rare in my material; they are easily distinguished by their smoothness, while the rest of the rhizome is covered FIGURE I. 7*. ianntnsis. FIGURE II. T. laiucolata (f nat. tize). with soft hairs. One apex was divided into three lobes, but investigation of the vascular structure revealed no branching of the stele, and I can offer no explanation of this unusual case. In every specimen examined the anterior end of the rhizome is directly transformed into a single aerial shoot, but Dangeard describes plants Downloaded from https://academic.oup.com/aob/article-abstract/os-22/1/63/201535 by Queen Mary University of London user on 24 March 2018 Sykes.— The Anatomy and Morphology of Tmesipteris. 65 in which several aerial stems arose from one rhizome.1 Usually the winding, hairy, and irregular rhizome passes by a more or less straight and glabrous transition region into the upright stem. The surface of the upper part of the transition region becomes gradually ridged owing to the fact that the leaf, bases are decurrent. At this stage a number of small scale- leaves, varying from two to six, according to the size of the plant, and reminding one of the leaves of Psilotum, are given off. Every transition is found between these and normal leaves. The latter in T. tannensis are from a half to three-quarters of an inch long, while in T. lanceolata they may reach one and a quarter inches in length. In the former species the largest leaves are found about half way up the stem, and decrease in size again towards the apex of the plant. Each leaf is sessile and ovate— lanceolate, has a mucronate tip, and is adnate to the stem by its base and part of one side. In T. lanceolata the largest leaves are found lower down on the stem than in T. tannensis; they have no mucronate tip, but taper gradually to a point, and they are lanceolate in shape. It is doubtful whether these two kinds of leaves are sufficiently distinct to warrant in themselves the distinction of the forms bearing them as two species; especially as in some cases2 both kinds have been found on the same plant. There appears to be no other important difference between the so-called species. The leaves are arranged spirally in two to five irregular rows, and are vertically placed, like the phyllodes of Acacia. (a) Fertile region. In T. tannensis the first branches are formed at the region of maximum development of the leaves, in T. lanceolata rather above this region. There may be a leaf here and there among the branches, but generally the purely foliar regions of the stem are sharply marked off from the branch-bearing regions, somewhat in the same manner as was noted by Bower in Lycopodium Selago? In several cases these regions were found to alternate, six to eight sterile leaves being placed between two fertile regions. No sterile branches such as those described by Bertrand4 and Dangeard 8 were borne on my specimens, but every branch was fertile and generally bore two leaves and a bisporangiate synangium. In some plants growth had stopped, and in these cases the main axis ended in a fertile branch or sporophyll, bearing as usual two leaves and a synangium. Presumably the apex is here employed in the formation of the terminal synangium. In one case the last organ borne by the mature plant was a leaf. Other specimens were still growing at the time 1 Dangeaid, Le Bot., li, p. 168. 3 Vanghan JenniDgs and Hall, 1891. * Bower, 1894, p. 353. * Bertrand, Figs. 203, 243-6. » Dangeard, Le Bot., ii,.p. 181, PI. XII, Fig. 5, and PL XIV, Fig. 8. F Downloaded from https://academic.oup.com/aob/article-abstract/os-22/1/63/201535 by Queen Mary University of London user on 24 March 2018 66 Sykes.— The Anatomy and Morphology of Tmesipleris. of preservation, and all these bore at their apex numerous fertile branches in all stages of development (Text-fig. II, p. 64). Such plants do not appear to have been among Bertrand's or Dangeard's material, but are figured by Jennings.1 Each ' sporophyll' consists of a short axis at the apex of which two leaves and a synangium are situated. The synangium is borne on a short pedicel or sporangiophore, which may, however, in unusual cases become much elongated, and even attain a considerable length-2 The leaves are exactly similar to those on the main axis. The synangia are generally bisporangiate, consisting of two lobes, each dehiscing longitudinally, and separated from each other by a plate of sterile tissue. A few abnormal fertile branches are described below. III. SHORT SUMMARY OF PREVIOUS WORK ON TMESIPTERIS. The most exhaustive account of Tmesipteris is found in Bertrand's monograph3, which deals very fully with both the anatomy and the morphology. Dangeard4 has given a more concise description of the genus, of which he recognizes several species, differing chiefly in the shape of their leaves and synangia. This author was the first to investigate the rhizome, and he studied to some extent the fungus5 found in its cells, and considered by him to be a member of the Chytridiaceae. Jennings and Hall6 have described the anatomy of Tmesipteris, but few new facts are to be gleaned from their account. The results of the above authors may be shortly summarized as follows: Tmesipteris is a rootless plant,' the underground portion absorbing moisture by means of rhizoids, containing a fungus and having a single concentric stele, which has usually two exarch protoxylem groups. In no case has an endodermis been seen either in the rhizome or in the aerial stem, and the minute structure of the phloem is as yet uninvestigated. The aerial portion of the plant bears leaves and sterile and fertile branches. The branches each bear a pair of leaves, and in the case of the fertile branches each has also a two-lobed spore-producing organ, formed on the end of a short pedicel arising at the point of divergence of the two leaves. The aerial stem also is monostelic, but here the phloem forms a continuous ring, while the xylem is arranged in several mesarch groups. Bertrand states that the leaves and branches each receive a single trace with a single protoxylem group, but Dangeard describes the formation of diarch steles to supply the branches. Each leaf-trace arises from one of the xylem groups of the stem by division at or near the place of origin, but each 1 Jennings and Hall, 1891, PI. I, Fig. II. ' Thomas, 1902, p. 343. * Bertrand, Recherches sur 1M Te'mripte'rid^es. 1 DflDgeard, Le Botanisle, ii, pp. 163-319, PI. IX-XV. ' Dangeard, Le Botaniste, ii, p. 222-30. * Jennings and Hall, 1891. Downloaded from https://academic.oup.com/aob/article-abstract/os-22/1/63/201535 by Queen Mary University of London user on 24 March 2018 Sykes.—The Anatomy and Morphology of Ttnesipteris. 67 branch trace is formed by the passing out of a whole group from the stem, its place being subsequently taken in the stem by. a group formed by division from one of the others. The various authors agree that in each fertile branch three bundles are finally found, one of which supplies the spore-producing organ, while the other two form the traces of the two leaves.
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