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Studies on Indian Hymenophyllaceae

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Studies on Indian Hymenophyllaceae STUDIES ON INDIAN HYMENOPHYLLACEAE Part III. Contributions to Our Knowledge ofMeringium edentulum (v.d.B. Copeland) BY A. R. RAO AND (Mlss) USHA SHARMA (Botany Department, Lucknow University) Received December 12, 1962 [Communicated by Prof. L. N. Rao, M•SC., Ph.D. (Lond.), r.ms., r.A.sc.] CONTENTS PAGE ° INTRODUCTION ...... .. 300 2. MATERIAL AND METHODS .... •. 301 3. GENERAL MORPHOLOGY ...... •. 301 4. ANATOMY- (a) Root I m .. 303 (b) Rhizome t 4 .. 303 (c) Leaf • ° .. 304 5. SUMMARY • ° 304 6. ~NCES • • O t 305 7. EXPLANATION OF PLATE 9 305 8. EXPLANATION OF TEXT-FIGURES • . 306 1. INTRODUCTION The morphological studies of two genera of Hymenophyllaceae, Crepidomanes latealatum (v.d.B) Copel. Sharma, 1960 and Hymenophyllum simonsianum. Hook. (Sharma, in press) have already been worked out. The present paper deals with the vegetative parts of a North Indian species of Meringium, M. edentulum (v.d.B.)Copel. The developmental stages of the sporangium could not be followed as adequate fertile material was not available for study. Observations on these will be published in duo course. 3OO Studies on Indian Hymenophyllaceae-- III 301 The plant comes under Hymenophyllum Linn. (Hooker and Baker, 1868, p. 68; Beddome, 1883, p. 34; Bower, 1908, p. 576; 1926, p. 250, Christensen, 1938, p. 351). According to vanden Bosch it has also been referred to Leptocionium edentulum v.d.B. (Copeland, 1938, p. 40). A comparison can also be made with H. serrulatum of Holttum (1954, p. 75) which according to Copeland is Meringium meyenianum. The material investigated resembles M. meyenianurn in every respect except that the margi- nal teeth are lacking and the margin is wavy. Such specimens have been called M. edentulum by Copeland (1947, p. 39). Meringium edentulum (v.d.B.) Copel. occurs in Assam, Luzon and Borneo. 2. MATERIAL AND METHODS The present material was collected from Khasi and Jayantia hills in Assam in November 1959 and preserved in 70.%o alcohol. Microtome sections were cut 8 to 15/~ in thickness. Free-hand sections were also found to be useful. Haemotoxylin-orange G and safranin- fast-green combinations gave good results. Phloroglucin was used to stain the lignified tissues (Foster, 1942, p. 141). Schulze's fluid for clearing up of the tissues and a 10~o solution of glyce- rine in water for softening hard tissues, like the rhizome and petiole, were also used. 3. GENERAL MORPHOLOGY The plants are epiphytic and are 10 to 17 cm. long (Plate XIV, Photo 1). The creeping rhizome, which is irregularly branched and black in colour, measures 0-3 ram. in thickness. The thin roots come out irregularly mostly from the lowr side of the rhizome (Text-Fig. 1 and Plate XIV, Photo 1) and are covered all round by unicellular hairs (Plate XIV, Photo 2). The fronds are pinnately lobed, measure 10 to 17 cm. in length and 3 to 4 cm. in width; pinnae measuring commonly 0" 5 to 1 cm. long, are very deeply cleft; clefts narrow, pinnules closely placed (Text-Fig. 2 and Photos 1 and 3). Basal pinnae are widely spaced and reduced in size (Photo 1). The rachis is thicker than the rhizome, measuring 0.4 mm. in thickness. The lamina is one cell thick, translucent and as stated earlier deeply cleft. The venation is dichotomous with free veinlets (Text-Fig. 2; Plate XIV, Photo 3). Young leaves show circinate vernation as in other ferns (Text-Fig. 1). Dermal appendages are one to three cells long hairs (Text-Figs. 3-5) found on the surface of the rhizome but absent on the leaf, They are rather 302 A. R. RAO A~,rD (MIss) USHA SHARMA ~ Y Y ~.-_ r. c. , 4 $.~. teo~. l ~ _ per Pxy. i -- ph 5 2 Tmcr-l~os. 1-9 Studies on Indian Hymenophyllaceae--III 303 scanty and not in abundance as in Hymenophyllum simonsianum (Sharma, in press). The sorus is terminal on the lowest anterior pinnules of the upper pinnae (Plate XIV, Photos 1 and 3 and Text-Fig. 2). Theindusium is 1.5 mm. long, cup- shaped and bivalved in its upper part. The base of the indusium is triangularly cup-shaped. In the upper part the triangular flaps are slightly longer than the basal hollow part. Copeland (1947, p. 8) has already pointed out that the receptacle is included when young and excluded in later stages. The pinnule supply divides into three (Text-Fig. 2) at the base of the receptacle. The central one supplies the receptable itself and the other two lateral ones enter the two valves along the line of their fusion (Text-Fig. 2). 4. ANATOMY (a) Root.--Text-Figure 6 represents a transverse section of a fairly old root, whose outer layers have been worn out. A single-layered epidermis with thick walls is succeeded by the cortex (C.) made up of two to three layers of thick-walled cells. The innermost layer (s.L) of this tissue has very highly stratified cell walls and narrow lumen. But here and there a thin-walled parenchymatous cell with unstratified walls may also be seen like a passage cell (p.c.). A single-layered endodermis (end.) and pericycle (per.) are also clear. The phloem (ph.) is very thin and one cell thick. The exarch xylem is diarch or triarch. (b) Rhizome.pPhoto 4 is a transverse section of the rhizome showing only the inner layers of the cortex. The cortex contains five to six layers of thick- walled cells, with narrow lumen and highly stratified walls. The stele is essentially slightly dorsiventral, concentric, protostelic with mesarch xylem. The endodermis (end.) is single-layered. The pericycle (per.) is two to three layered, made up of parenchymatous cells. According to Boodle (1900, p. 459), all the layers in between the endodermis and phloem are taken as pericycle. A central group of more or less thirty tracheids are surrounded by one to two layered phloem made up of narrow cells (ph.), separated from the xylem by one to two layered parenchyma. The protoxylem (p.xy.) is at one point and is mesarch. The xylem can roughly be compared with the I type of xylem found in Meringium subtype of Hymenophyllum type as reported by Nozu (1950, p. 75). The stele though similar to that of Hymenophyllum simonsianum differs from it in being mesarch. 304 A.R. RAO AND (MIss) USI-IA SHASMA Leaf or root traces arise from the main stele of the rhizome (Plate XIV, Photo 4) without any gaps. (c) Leaf--Text-Figure 7 is a transverse section of the petiole in the basal region. The structure is exactly similar to the rhizome in all the details. A single-layered epidermis is succeeded by a cortex six to seven cells deep and made up of thick-walled cells. The stele is a concentric one (Text-Fig. 7) as in the rhizome with a single-layered endodermis (end.) and several layered pericycle (per.). The xylem (xy.) is plate-like and the protoxylem is exarch. A little higher up the stele becomes collateral and remains so till the apex. The venation of the lamina is fundamentally dichotomous and the pinna supply is by the abstriction of the main stele, alternately right and left. Text-Figure 8 is a transverse section of the leaf in the apical region. Con- necting the upper and lower epidermises are a single layer of cells constituting the lamina (/), which is throughout one cell thick except round the midrib and veinlets. The cells of the lamina are heavily loaded with chloroplasts. Plate XIV, Photo 5 shows them in surface view. The large cortex seen at the earlier levels of the petiole is now reduced to a single layer. The stele is very much reduced in size and is collateral in form. The endodermis and peficycle are one-layered. The xylem is made up of six to seven tracheids and four or five small cells of the phloem, separated from the xylem by one or two cells of conjunctive parenchyma. Text-Figure 9 shows a lateral vein cut transversely. The stele is further reduced with a central mass made up of four to five tracheids (tr.) surrounded by parenchymatous tissue. 5. SUMMARY The paper deals with the morphological studies of a North Indian species of Meringium, M. edentulum (v.d.B) Copel. collected from Khasi and Jayantia hills of Assam. The numerous roots, coming out irregularly from the rhizome and densely covered by unicellular hairs, are usually diarch, rarely triarch also. The rhizome is black in colour, creeping and has a mesarch protostele. The leaves are once pinnate with a free dichotomous venation. Dermal appendages are one to three celled hairs and are present only on the rhizome and not on the leaf. The sorus is terminal on the lowest pinnules of the upper pinnae. The indusium is cup-shaped in its lower half and is bivalved in the upper half. The receptacle is included at first but becomes excluded later on. Studies on Indian Hymenophyllaceae~IlI 305 6. REmu~NC~S Beddome, R.H. .. Handbook to the Ferns of British India, Thacker, Spink and Co., Calcutta, 1883. Boodle, L.A. • • "Comparative anatomy of the Hymenophyllaceae, Schi2eaeceae and Gleicheniaceae. 1. On the anatomy of Hymeno- phyllaceae," Ann. Bot., 1900, 14, 455-96. Bower, F.O. .. The Origin of a Land Flora, Macmillan & Co., London, 1908~ .. The Ferns (Filicales), The University Press, Cambridge, 1926, 2. Copeland, E.B. .. "Genera Hymenophyllacearum," The Phillipp. J. Sci., 1938, 67, 1-110. .. Genera Felicum, Waltham, Mass., U.S,A., 1947. Christensen, C. .. "'Filicinae," in Verdoorn, Fr. Ed., Manual of Pteridology, Martinus Nijhoff, The Hague, 1938, 522-50.
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