Teratophyllum sect. Teratophyllum
Content
Distribution
Asia-Tropical, Ceylon present, Lesser Sunda Is absent, Mergui present, Peninsular Thailand present
Mergui and Peninsular Thailand, Ceylon; throughout Malesia, except Lesser Sunda Is.; 9 spp.
Ecology
Confined to primary forest. Prothalli grow on exposed tree-roots or bases of buttresses; young sporophytes develop a slender rhizome which usually grows upwards and may branch (in T. ludens it produces long trailing branches which pass along the ground from one tree to another) bearing successively larger and more complex bathyphylls, the earlier ones often asymmetric, with lamina more fully developed on lower side, with a more or less abrupt transition to acrophylls at c. 2 m above ground level. Fertile fronds produced at 3-5 m or more above ground, seasonally (probably in response to short periods of drier weather, which in many parts of Malesia are of irregular occurrence). A copious root- system develops on the lower parts of the climbing stem, spreading in the soil, and I believe this to provide most of the water needed by the plant. Karsten, believing Teratophyllum plants to lack such a soil- penetrating root-system, suggested that the chief function of bathyphylls is water-absorption (). Bathyphylls can absorb some water, but not enough to be of much importance; and they do not always lie in close contact with the bark of the supporting tree, as described by Karsten (see ).
Taxonomy
The two species native in Java were first described by Blume (1828) in the genus Lomaria; he saw bathyphylls and thought they were fertile fronds. John Smith, when enumerating Cuming's Philippine ferns (1841) established a new genus Stenochlaena, and stated, on the evidence of Cuming's specimens, that Stenochlaena scandens (now known as S. palustris (Burm.f.) Bedd.), the type-species of Stenochlaena, sometimes produced abnormal bipinnate fronds. These fronds were bathyphylls of Teratophyllum aculeatum, which were associated with normal sterile and fertile fronds of Stenochlaena palustris by Cuming under his n. 347, as shown by specimens in John Smith's herbarium (BM), one annotated by him. This idea persisted with Smith and is repeated in his last book (); it was also accepted by Hooker () and by Beddome (Handb.) who so named a specimen of T. aculeatum bearing bathyphylls collected by Wallich in Penang in 1822. Wallich had given the name Davallia achillaeifolia to this specimen, and the name was formally published, with a figure, by Hooker (), with doubts expressed as to the affinity of the fern and a reference to its resemblance to Lomaria aculeata Bl. Baker named similar ferns from Moulmein Lindsaea parishii in 1867. Even as late as 1929 Copeland wrote of "the polymorphism of the fronds of immature plants" of Stenochlaena palustris (). Sporeling plants of S. palustris are very rare in Malaya, though mature plants are abundant and frequently fertile. I have however raised young plants from spores. Their first fronds were simple, then simply pinnate, showing from the first the leaflet-form and venation of Stenochlaena, quite unlike Teratophyllum. I can confidently assert that no true Stenochlaena produces fronds which could be confused with the bathyphylls of Teratophyllum. It should be noted that young plants of Asplenium epiphyticum Copel. also produce bathyphylls. These were confused with those of species of Teratophyllum by Christ, who concluded that "Stenochlaena" was an acrostichoid derivative of Asplenioid origin; Copeland accepted this derivation as "absolutely clear" in 1929. (See ; also his earlier composite fig. 96, p. 40, under Lomariopsis sorbifolia, which included fronds of Lomariopsis, Teratophyllum and Asplenium, in Farnkr. der Erde, a figure copied without question by ).
In his monograph of the acrostichoid ferns (1845) Fee did not mention Stenochlaena palustris. He included the species of Teratophyllum sect. Teratophyllum known to him in his new genus Lomariopsis; the species were L. spinescens (Lomaria aculeata Bl.), L. leptocarpa (based on Cuming 132 from Luzon) and L. ludens (based on a Wallich specimen from Singapore). Of these, the first was known only from bathy- phylls and sterile acrophylls, the second from sterile and fertile acrophylls, the third only from bathyphylls. Hooker later included all these, and all true Lomariopsis specimens known to him, in Acrostichum sorbifolium L. (), a species now regarded as confined to tropical America.
Mettenius was the first to recognize Teratophyllum as a genus (1869; posthumous work edited by Kuhn); he included in it two species, one in each of the sections here recognized (he included Stenochlaena, as a section, in Lomariopsis). He included all specimens of sect. Teratophyllum in the species T. aculeatum.
Underwood clearly distinguished for the first time between Stenochlaena, Lomariopsis and Teratophyllum, ranking all as sections of Stenochlaena (). But he failed to recognize the great differences between Stenochlaena proper and the other sections, as he did not examine spores, scales, or vascular anatomy, and failed to notice the "glands" at the bases of pinnae (including basal reduced ones) in Stenochlaena; he also failed to notice that Teratophyllum differs from Lomariopsis in having all pinnae jointed to the rachis, including the apparently terminal one. Underwood did not recognize the close relationship between sect. Teratophyllum and sect. Polyseriatae, remarking only that the latter (as Arthrobotrya J. Sm.) appeared to be a valid distinct genus.
The present account is based on that of , with some additional material, especially that recently collected by M. G. Price in the Philippines.
In his monograph of the acrostichoid ferns (1845) Fee did not mention Stenochlaena palustris. He included the species of Teratophyllum sect. Teratophyllum known to him in his new genus Lomariopsis; the species were L. spinescens (Lomaria aculeata Bl.), L. leptocarpa (based on Cuming 132 from Luzon) and L. ludens (based on a Wallich specimen from Singapore). Of these, the first was known only from bathy- phylls and sterile acrophylls, the second from sterile and fertile acrophylls, the third only from bathyphylls. Hooker later included all these, and all true Lomariopsis specimens known to him, in Acrostichum sorbifolium L. (), a species now regarded as confined to tropical America.
Mettenius was the first to recognize Teratophyllum as a genus (1869; posthumous work edited by Kuhn); he included in it two species, one in each of the sections here recognized (he included Stenochlaena, as a section, in Lomariopsis). He included all specimens of sect. Teratophyllum in the species T. aculeatum.
Underwood clearly distinguished for the first time between Stenochlaena, Lomariopsis and Teratophyllum, ranking all as sections of Stenochlaena (). But he failed to recognize the great differences between Stenochlaena proper and the other sections, as he did not examine spores, scales, or vascular anatomy, and failed to notice the "glands" at the bases of pinnae (including basal reduced ones) in Stenochlaena; he also failed to notice that Teratophyllum differs from Lomariopsis in having all pinnae jointed to the rachis, including the apparently terminal one. Underwood did not recognize the close relationship between sect. Teratophyllum and sect. Polyseriatae, remarking only that the latter (as Arthrobotrya J. Sm.) appeared to be a valid distinct genus.
The present account is based on that of , with some additional material, especially that recently collected by M. G. Price in the Philippines.
Cytology
The only Observation is from roots of a plant of T. ludens in cultivation at Kew, showing 2n = 82.
Citation
Mett. sensu Holttum 1954 – In: Rev. Fl. Mal.: 470
Bedd. 1883: Handb. Ferns Br. India: 423
J. Sm. 1875: Hist. Fil.: 139
Mett. sensu Holttum 1960: Fern Fl. Philip.: 269
Copel. 1947: Gen. Fil.: 117
1917: Handb.: 428
J. Sm. 1841 – In: Hook., J. Bot. 4: 149
J. Sm. 1875: Hist. Fil.: 312
Backer & Posth. 1939: Varenfl. Java: 151
Christ 1897: Farnkr. Erde: 39
Presl 1851: Epim. Bot.: 165
C. Chr. 1934: Ind. Fil.: 186
Mett. sensu Holttum 1937 – In: Gard. Bull. S. S.: 142
v.A.v.R. 1908: Handb.: 722