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FERN GAZETTE � .A ,,\ R; tf � THE FERN GAZETTE � .A ,,\ r;/ Edited by lAoCrabbe & Bo5oParris THE BRITISH PTERIDOLOGICAL SOCIETY 1/J 1\� Volume 13 Part 4 _______..;;._..____;;...___", 1988_ The Bri tish Pteridological Society THE FERN GAZETTE VOLUME 13 PART 4 1988 CONTENTS Page MAIN ARTICLES A chromosome count from Azolla filiculoides (Azollaceae: Pteridophyta) - You-Xing Lin & Anne 5/eep 193 Cytological and anatomical Observations on Tm esipteris (Tmesipteridaceae: Pteridophyta) species from New Caledonia - A. F. Braithwaite 199 Shoot temperature measurements of montane Cyathea (Cyatheacea: Pteridophyta) species in Papua New Guinea - M.J. Ea mshaw, T. C. Gunn & J.R. Croft 209 Pteridophytes of Zarate, a forest on the western side of the Peruvian Andes - 8/anca Le6n & Niels Valencia 217 A field survey of Pteridium aquilinum (Dennstaedtiaceae: Pteridophyta) mycorrhizas - H.M. Jones & E. Sheffield 225 Adaptive strategies of Marsi/ea (Marsileaceae: Pteridophyta) in the Lake Chad basin of N.E. Nigeria - Jan Komas 231 SHORT NOTES Dryopteris villarii (Dryopteridaceae: Pteridophyt&) a new high-mountain species in the Carpathians - Halina Piekos-Mirkawa & Zbigniew Mirek 24 5 Lygodium japonicum in Singapore - Y.C. Wee & L.L. Chua 24 6 Dryopteris x gomerica (Dryopteridaceae: Pteridophyta) new for Europe - R. Viane 24 7 REVIEWS 250 OBITURY: lrene Manton - Anne Sleep 253 (THE FERN GAZETTE Volume 13 Part 3 was pub/ished on 28th Ju/y 1988) Published by THE BRITISH PTERIDOLOGICAL SOCIETY, c/o Department of - Botany, British Museum (Natural History), London SW7 580 ISSN 0308-0838 Metlee Printers Ltd, Old Station Road, Loughton, Essex FERN GAZ. 1 3(4) 1 988 193 A CHROMOSO. ME COUNT FROM AZOLLA FILICULOIDES (AZOLLACEAE: PTERIDOPHYTA)* YOU XING LIN** and ANNE SLEEP Department of Pure and Applied Biology, The University, Leeds, LS2 9JT, England ABSTRACT Azol/a filiculoides has been investigated cytologically for the first time. Counts from microsporangia establish its chromosome number as n = 22 . INTRODUCTION To date there have been very few cytological investigations of Azolla (the Water Fern). The minute chromosomes present technical problems which are compounded by the delicate nature of the roots and a scarcity of fertile material at the right stage. The extremely small chromosome size also causes considerable difficulty in thß establishment of unequivocal counts. The cytology of Azolla has thus lagged behind research into other aspects of its biology. Recently much interest has focussed on this genus because of its nitrogen-fixing ability; this arises from the symbiotic re lationship between Azolla and the endophytic cyanobacterium (or blue-green alga), Anabaena azollae, which is invariably present in the air chambers in the upper Iobes of its leaves. The consequent economic importance of Azolla has long been recognized in the Far East, where, particularly in China and Vietnam, A. pinnata R.Br. has been used for centuries as a green manure for increasing the fertility of paddyfields (lumpkin 1985). During the last ten years various other species of Azolla have been introduced into China for the purposes of breeding and other research, and the arrival in Leeds of Y. X. Lin presented an opportunity to count one of the most vigorous of these introduced Azolla species, a particularly strong-growing form identified as A. filiculoides Lam. Azolla shows much variation in the morphology of its vegetative parts, and the same species can Iook quite different when growing under different environmental conditions. This phenotypic plasticity has led to problems over identification; the boundaries between species are unclear and in general the taxonomy of the genus is confused. MATERIAL The material used in this study came originally from the Forstbotanischer Garten Eberswalde der Humboldt, at the University of Berlin in East Germany. This culture was sent to the botanic garden of the Botanical lnstitute, Chinese Academy of Sciences, Beijing, in 1978, and was established in cultivation there. ln 1986 material of the same collection was successfully brought into'"cultivation at Leeds. Sporangia were produced in the autumn of 1986 and again in 1987. Material of botanic-garden origin may sometimes be misidentified, especially in view of the problems outlined above. lt was therefore thought desirable to check the identification of our culture as A. filiculoides, and to this end various morphological features were studied. Characters for the identification of A. filicu/oides are summarized by Ashton & Walmsley (1984) and by Ruiz de Clavijo et al. (1 984). Fertile material of mature, well-grown A. filiculoides, one of two American species introduced and now widespread in Europe, is said to be very distinctive (the other, A. caroliniana Willd., is much more difficult to identify positively and its taxonomic status may be doubtful, see Derrick, Jermy & Paul 1987) • Dedicated to Prof. T. Reichstein on the occasion of his 90th birthday (2 0.7.1987) in recognition of his friendship and encouragement. **On leave from the Institute of Botany, Chinese Academy of Sciences, Beijing, People's Republic of China. 194 FERN GAZETIE: VOLUME 13 PART 4 (1 988) PLATE 1 I. Morphology of our cytologically attested Azolla material (A. filiculoides): SEM photographs of leaves. Fig. 1. Upper epidermis showing lhe wlde margi n of 3-4 rows of large, hyaline cells and the central Iamina bearing stomata and unicellular trichomes or papillae. Fig. 2. Enlarged view of the unicellular trlchomes. AZOLlA FILICULOIDES 195 Our material is a strong-growing plant, cons1stmg, when mature, of both a decumbent, free-floating system as weil as erect branches which can extend out of the water by as much as 5cm. When the emergent branches are well-developed the horizontal system tends to die back, forming a tangled brown mat on the water surface. The ascending growth habit arises particularly when the plant is growing in a limited space or under dry conditions. Lin (1980) has observed that on these upright branches the colourless and usually submerged lower Iobes of the leaves also become green and develop Anabaena-containing cavities. The flattened, leafy stems of the horizontal system are elongate, and the branches distant one from another. ln young plants the branching is dichotomous; on adult material unequal development at branch-points Ieads to a false dichotomy and branching of apparently irregular arrangement. The tips of the emergent branches tend to roll inwards to give a distinctive crisped appearance mentioned by many authors. Roots arise in no ·set pattern or position but acropetally at intervals along the underside of the floating stems. The ovate, imbricate, closely appressed branch-leaves agree with the description of Svenson (1 944); they are alternate, greyish-green in colour, with subacute to obtuse tips and a broad, membranaus margin (�s described by Lawalree in Tutin 1964) formed of 2-3 rows of large, hyaline cells (see Figs. 1 & 2). Unicellular trichomes (Lumpkin & Plucknett 1980, Pieterse, de Lange & van Vliet 1977) are borne on the surface of the epidermis (see Figs. 1 & 2). Sporocarps develop, usually in pairs, in the axils of leaves close to the branch apices. Either two microsporocarps (very rarely, two megasporocarps) or a microsporocarp and accompanying megasporocarp occur together. Within each microsporocarp there are numerous microsporangia, each containing 4-6 massulae. The massulae bear barbed glochidia which are clearly aseptate (see Fig. 3). Fowler and Stennett-Willson (1 978) attach great importance to the structure of the megaspore wa ll in classification; although not studied in detail the megaspore sculpturing of our material agrees weil with that illustrated by them. On the basis of these Observations we have no hesitation in confirming the identification of our plants as A. filiculoides Lam. Voucher herbarium specimens of our material will be deposited in the national herbaria at Kew and the British Museum (Natural History), as weil as in the herbaria of the Botanical lnstitute, Chinese Academy of Sciences, Beijing and of the International Rice Research Institute, Manila, Philippines. METHODS Young microsporangia were fixed in a solution of 3 parts absolute alcohol: 1 part glacial acetic acid, sta ined in acetocarmine and sq uashed according to the technique described by Manton (1 950). Preparations were made permanent by McCiintock's method (1 929). Suitable cells were photographed in phase-contrast on a Zeiss Ultraphot II microscope. Stereoscan photographs of leaves and glochidia were taken from material fixed in 1% osmium tetroxide, dehydrated through an ascending acetone series and critical-point dried before mounting and coating with gold. The specimens were examined on a CamScan Series 111 scanning electron microscope. CYTOLOGY Several cells in metaphase I yielded counts of n = 22 bivalents. The cell illustrated in Fig. 4 (with accompanying explanatory diagram in Fig. 5) is at late metaphase and some of the bivalents are already beginning to separate prior to anaphase I. One prominent pair is very much I arger than the rest. Root-tip preparations by ourselves from the same material gave counts of 2n = c.44, which are in accordance with the meiotic observations. 196 FERN GAZETIE:VOLUME 13 PART 4 (1988) 5 I � PLATE 2 Fig. 3. SEM photograph of surface of massula showin�Jthe aseptate glochidia. II. Cytology. melosis from microsporanglum of A. filiculotdes. Flg. 4. Late metaphase I showing 22 bivalents. One chromosome (arrowed) is much larger than the rest. Some bivalents are already beginning to separate. Magnlfication: x 2,400. Fig. 5. Explanatory diagram to fig. 4. n • 22. Magnification: x 3,800. AZOLLA FILICULOIDES 197 DISCUSSION Löve, Löve and Pichi-Sermolli (1977) give base numbers for Azo/la of x "' 11, x "' 12. The latter number is based on two early reports, the first by de Litardiere (1921 ), who obtained a count of 2n "' 48 from material of A. caroliniana. He used sectioned material, from which it is not always easy to establish a precise count. However, Tschermak-Woess and Dolezai-Janisch (1 959), in a paper on the cytology of Cyrtomium fa lcatum, state that they found 2n "' 48 in material of A.
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