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The Distribution of C3 and C4 Plants in a Successional Sequence in the Okavango Delta

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The Distribution of C3 and C4 Plants in a Successional Sequence in the Okavango Delta 400 S.-Afr.TydskT.Plantk., 1992, 58(5) Specimens examined The distribution of C3 and C4 plants in a Calonectria hederae, Hedera helix leaf, Great Britain, 1958, successional sequence in the Okavango IMI 75300 (holotype); C. hederae. Hedera helix leaf, Great Delta Britain, 1978, IMI 241261; C. hederae. Hedera helix leaves, France, 1948, G. Arnaud, IMI 39232 (lectotype); Cylindro­ carpon reteaudii, on Smithia bequaertii, Indo China, F. Karen Ellery,* W.N. Ellery* and 8.Th. Verhagent Bugnicourt, 1M! 55922 (dried culture derived from type, 'Department of Botany, University of the Witwatersrand. P.O. Herb Paris). Wits, 2050 Republic of South Africa tSchonland Research Centre, University of the Witwaters­ Culture examined rand, P.O. Wits, 2050 Republic of South Africa C. leucothoeae, from Leucothoeae axillaris leaves, Florida, Received 8 January 1992; revised 26 May 1992 U.S.A., Feb. 1988, EI-Gholl, ATCC 64824 (type culture). Carbon isotope techniques were used to determine the Acknowledgements photosynthetic pathway of a number of wetland plant The authors thank the International Mycological Institute, species. The relative cover abundances of C3 and C4 plants Kew, Surrey (lMI), and the American Type Culture Collec­ were compared along a successional sequence of the tion, Rockville, Maryland (A TCC), for placing specimens Maunachira River system of the Okavango Delta. Plants with and cultures at our disposal. a C3 photosynthetic pathway were dominant in early succes­ sional stages and C4 plants were dominant in late References successional stages. It is proposed that nutrient availability, particularly nitrogen limitation, could be a determinant of the ARNAUD. G. 1952. Mycologie Concrete: Genera. Bull. Soc. change from C3 to C4 plant dominance during succession. Mycol. Fr. 68: 181 - 223. BOESEWINKEL. H.I. 1982. Heterogeneity within Cylindrocla­ Die fotosintetiese verloop in 'n aantal moerasplante is deur dium and its teleomorphs. Trans. Br. mycol. Soc. 78: 553 - middel van koolstof-isotooptegnieke vasgestel. Die relatiewe 556. dekkingsdigthede van C3 en C4 plante in opeenvolgende BOOTH, C. 1966. The genus Cylindrocarpon. Mycol. Pap. 104: seksies van die Maunachira riviersisteem van die Okavango 1 - 56. Delta is met mekaar vergelyk. In die aanvanklike seksies was BOOTH. C. & MURRAY, 1.S. 1960. Calonectria hederae plante met 'n C3 fotosintetiese verloop dominant. terwyl C4 Arnaud. and its Cylindrocladium conidial state. Trans. Br. plante in die daaropvolgende seksies dominant was. Daar mycol. Soc. 43: 69 - 72. word voorgestel dat die beskikbaarheid van voedingstowwe, BUGNICOURT, F. 1939. Les Fusarium et Cylindrocarpon de veral stikstof, 'n rol speel in die verandering van C3 l'Indochine. Encycl. Mycol. 11: 1 - 206. dominansie na C4 dominansie. CROUS. P.W., PHILLIPS, AJ.L. & WINGFIELD. M.l. 1991. The genera Cylindrocladium and CylindrocladieUa in South Keywords: carbon isotope, C3• C4• nutrients, Okavango Africa, with special reference to forest nurseries. S. Afr. For. J. Delta, succession. 157: 69 - 85. EL-GHOLL. N.E., LEAHY, R.M. & SCHUBERT, T.S. 1989. Cylindrocladium leucothoeae sp. nov. Can. J. Bot. 67: 2529 - The C4 photosynthetic pathway has been shown to have an 2532. adaptive advantage under conditions of high temperature, FISHER, N.L.. BURGESS, L.W., TOUSSOUN, T.A. & high irradiance and in an arid environment (Osmond et al. NELSON, P.E. 1982. Carnation leaves as a substrate and for 1982; Pearcy & Ehleringer 1984). The field studies in which preserving cultures of Fusarium species. Phytopathology 72: the distribution of C3 and C4 plant species was investigated, 151 - 153. have been either along environmental gradients (Tieszen et PEERALLY, A. 1974. CMI Descriptions of Pathogenic Fungi and al. 1979; Boutton et al. 1980) or on a phyto-geographical Bacteria No. 426. basis (Vogel et al. 1978; Ellis et al. 1980; Cowling 1983; PEERALLY, A. 1991. The classification and phytopathology of Hattersley 1983; Vogel et al. 1986). No work relating Cylindrocladium species. Mycotaxon 40: 323 - 366. SHERBAKOFF, C.D. 1828. Washington palm leaf spot due to photosynthetic metabolic pathways to a successional sequence of plant communities has been conducted. Cylindrocladium macrosporum n. sp. Phytopathology 18: 219 - 225. The area of the present study was the permanently SOBERS, E.K. 1967. Morphology and pathogenicity of Cylindro­ inundated Maunachira River system situated in the north­ cladium macrosporum and C. pteridis. Phytopathology 57: eastern part of the Okavango Delta, north-western 464. Botswana. The vegetation consists of a heterogeneous mix SOBERS, E.K. 1968. Morphology and host range of Cylindro­ of wetland plant communities comprising submerged and cladium pteridis. Phytopathology 58: 1265 - 1270. floating-leaved species dominant in deep open water bodies SOBERS, E.K. & ALFIERI, S.A. 1971. Species of Cylindro­ and short, emergent plant species dominant in shallow peat cladium and their hosts in Florida and Georgia. Proc. Fla. bogs. A successional sequence for the wetland plant State Hortic. Soc. 85: 366 - 369. communities of the study area, based on phytosociological WOLF, F.A. 1926. Brown leaf spot of leather fern. 1. Elisha associations and peat stratigraphy, is described by Ellery et Mitchell Sci. Soc. 42: 55 - 62. al. (1991). S.AfrJ.Bot., 1992, 58(5) 401 Carbon isotope analysis was used as an indicator of Submerged plant assemblages rooted in deep water sites photosynthetic pathway type. Species chosen for analysis (stage A) are exposed to low levels of irradiance. The plant were important components in one or more of the succes­ communities in later successional stages (stages B - F) have sional stages and comprised mainly grasses and sedges relatively open canopies where very little shading occurs. It which dominate the landscape. Gas samples for carbon isotope analysis were produced using a modified Dumas combustion method (Macko & Parker 1983). Approximately Table 1 Photosynthetic pathways (PP) and l)13C (%0 PDB) values of a number of plant species in the Maunachira 5 mg of dry, powdered plant material was added to silica River system ampoules containing CuO, Cu and Ag previously degassed at 400°C. Ampoules were evacuated, lightly flamed, sealed and heated to 900°C. After slow cooling, seals were broken Plant species at liquid-nitrogen temperature and ampoules were again Cyperus peetinatus Yahl ~ -26.0 evacuated. CO2 gas was then released at -80°C into the inlet Eleocharis acutangula (Roxb.) Schult.b C3 of a Micromass 602C mass spectrometer. The carbon iso­ Eleocharis dulcis (Bunn.F.) Hensch. ~ -26.0 tope ratios were measured in duplicate and expressed as Ficus verruculosa Warb" C3 -26.8 813C values in permille (%0) with respect to PDB Standard§. Fuirena puhescens (poir.) Kunthb C3 Measurement precision is estimated at +0.10/00. Whilst F uirena stricta Steud. C3 -27.1 known C3 plants have 813C values around -270/00, known C4 Fuirena umbel/ata Rottb. b ~ plants have 813C values around -120/00. CAM plants have Ludwigia leptocarpa (Nutt.) Hara' C3 -26.1 mainly intermediate values (Deines 1980). Panicum parvifolium Lam. b ~ b Mean percentage cover of each of the C3 and C4 plant Phragmites australis (Cav.) Steud. C3 b species within each successional stage was determined and Sacciolepis africana C.E. Hubb. & Snowden C3 the overall contribution of each of these types to total cover Scirpus cubensis Poeppig & Kunth ex Kunthlt C3 Scleria woodii C.B.CLb C3 was estimated. Thelypteris confluens (lbunb.) Morton' C -28.2 The results of the carbon isotope analysis, as well as the 3 Typha capensis (Rohrb.) N.E.Br. ~ -28.1 photosynthetic pathway of major plant components in the Cyperus articulatus L. b C4 study area, are presented in Table 1. The analysis of Eriochrysis pal/ida Munro C4 -12.6 percentage cover of each of the C3 and C4 plant species in Imperata cylindrica (L.) Raeuschel C4 -12.8 each successional stage shows that C3 plant species are Miscanthus capensis Nees Anderss. var capensis C4 -12.3 present with a higher abundance than C4 species in the first Panicum repens L. b C4 three successional stages (Figure 1), namely A, Band C, Pycreus mundtii Neesb C4 which comprise deep-water communities dominated by sub­ Pycreus nitidus (Lam.) J. Raynal C4 -11.9 merged, floating-leaved and tall emergent plant species. The Vossia cuspidata (Roxb.) Griff.b C4 opposite was apparent in the last three successional stages Nymphaea caerulea Sav.' CAM -22.3 (D, E and F). The greatly increased relative cover of C4 , Omitted from succession analysis species in these three stages was due to the dominance of b Photosynthetic pathway determined from other studies the C4 sedge Pycreus nitidus in the floating sudd communities of Stage D and the presence of many short emergent C4 species such as Eriochrysis pallida, Vossia 100 cuspidata, lmperata cylindrica and Miscanthus capensis var. capensis in the shallow peat bog communities of stages E 90 ~ 80 and F (Ellery et al. 1991). Non-sedge and non-grass species o u present in each successional stage were not included in the OJ 70 OJ analysis, as their relatively minor contribution to total cover C'" eo would have had little effect on the observed distribution OJ ~ 50 0.. pattern of C3 and C4 plants. 2 «) The time span of the successional sequence from deep '" (4 m) water bodies colonized by submerged, floating-leaved .~en 30 and tall emergent plant species (stages A, B & C, Figure 1) W 20 to floating sudd communities (stage D), to peat bog com­ 10 munities of less than half a metre water depth dominated by o A B c o E F short, emergent wetland species (stages E & F) has been Successional stage estimated to be in the order of one to two centuries in the study area (p. Smith, Dept. of Water Affairs, Maun, Figure 1 The relative proportions of cover abundance of C3 and Botswana, pers. commun.). Throughout this time, peat has C4 grasses and sedges in each of the six recognized successional stages A - F (early - late succession).
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