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Afroalpine Plant Ecology

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Afroalpine Plant Ecology ACTA PHYTOGEOGRAPHICA SUECICA BD ID IT SVENSKA VAXTGEOGRAFISKA SALLSKAPET 49 FEATURES OF AFROALPINE PLANT ECOLOGY Avec un resume en frant;ais BY OLOV HEDBERG UPPSALA 1964 A L M Q V I S T & W I K S E L L S B 0 K T R Y C K �- R I A B SVENSKA VAXTGEOG,RAFISKA SALLSI(APET SOCIETAS PHYTOGEOGRAPHICA SUECANA Adress: Vaxtbiologiska Institutionen, Villavagen 14, Uppsala, Sverige Styrelse (council): prof. Hugo Sjors, ordf. (chairman); lab. Magnus Fries, v. ordf. (vice chairman); fil.lic. Bengt M. P. Larsson, sekr.; fil. kand. Folke Bjorkback, skattm. (treasur�r); doe. Mats Wrern, red. (editor); fil. lic. Gunnar Wassen, v. sekr.; doe. Olov Hedberg; prof. John Axel Nannfeldt; doe. Nils Quennerstedt. Sallskapet har till andama.I att vacka och underhalla The object of the Society is to promote investigation intresse for vaxtgeografien i vidstracktaste mening, in flora and vegetation, their history and their eco­ att framja utforskandet_av flora och vegetation i Sve­ logical background. Through publication of mono­ rige och andra lander och att havda geobotanikens graphs, and other activities, the Society tries to praktiska och vetenskapliga betydelse. stimulate geobotanical research and its application Sallskapet anordnar sammankomster och exkur­ to practical and scientific problems. Membership is sioner och utger publikationsserien Acta phytogeo­ open to all persons, in Sweden and e]sewhere, who graphica suecica. Varje for sallskapets. syften och have a personal interest in the advancement of verksamhet intresserad person kan vinna intrade phytogeography. efter anmalan hos sekreteraren. Foreningar, bibliotek, The Society issues the Acta Phytogeographica laroanstalter och andra institutioner kunna inga som Suecica. Individual members and subscribers (so­ abonnenter. Arsavgift 24 kr. (18:- :£or studerande); cieties, institutes, libraries, etc.) receive the Acta standigt medlemskap 400 kr. for annual dues of 24 Skr plus postage. There are Medlemmar och abonnenter erhalla Acta mot additional fees in years when more than one volume postforskott pa arsavgift jamte porto och expedi­ are issued. Permanent membership 400 Skr. For tionskostnad. Da flera band utges under samma ar, membership please apply to the Secretary. uttages en tillaggsavgift for extra band. Postgiro 7 504 7. Back volumes may be obtained by members and Medlemmar och abonnenter kunna erhalla aldre subscribers at reduced cost (see list of publications, publikationer till reducerat pris (parentespriset i prices in brackets). To receive the Acta by exchange listan). for other publications, please apply to the Institute Librarian (address as above). SVENSKA VAXTSOCIOLOGISKA SALLSKAPETS HANDLINGAR I. H. OsvALD, Die Vegetation des Hochmoores Komosse. 6. TB. C. E. FRIES, Die Rolle des Gesteinsgrundes bei der 1923. 40: - (27: -}. Verbreitung der Gebirgspflanzen in Skandinavien. 2. G. E. Du RIETZ, Gotlii.ndische Vegetationsstudien. 1925. 3: - (2: -). 1925. 12: - (8: -}. 7. H. OsvALD, Zur Vegetation der ozeanischen Hochmoore 3. G. E. Du RIETZ & J. A. NANNFELDT, Ryggmossen und in Norwegen. 1925. 15:- (10: -). Stigsbo Rodmosse, die letzten lebenden Hochmoore 8. G. E. Du RIETZ, Die regionale Gliederung der skandi­ der Gegend von Uppsa.la.. 1925. 6: - (4: -). na.vischen Vegetation. 1925. 15: - (10: -) . 4. G. E. Du RIETZ, Zur Kenntnis der flechtenreichen 9. G. SAMUELSSON, Untersuchungen iiber die hohere Zwergstra.uchheiden im kontinenta.len Siidnorwegen. Wasserflora von Dalarne. 1925. 6:- (4: -). 1925. 12: - (8: -). 10. TB. C. E. FRIEs, En vaxtsociologisk huvudfraga. (A 5. TB:. C. E. FRIEs, Okologische Un.d phii.nologische Beob­ principal problem of phytosociology.) 1926. 0: 50 (0: 25). a.chtungen bei Abisko in den Jahren 1917-1919. I. 1925. 15: - (10: -). ACTA PHYTOGEOGRAPHICA SUECICA 1. E. Al.MQUIST, Upplands vegetation och flora. (Vegeta­ their distribution in. Northwestern Europe.) 1932. 12: - tion and flora of Uppland.) 1929. 40:- (27: -). (8: -). 2. S. THUNMARK, Der See Fiolen und seine Vegetation. 5. H. OsvALD, Vegetation of the Pacific coast bogs of 1931. 21:- (14: -). North America. 1933. 12:- (8: -). 3. G. E. Du RIETZ, Life-forms of terrestrial flowering 6. G. SAMUELSSON, Die Verbreitung der hoheren Wasser­ plants. I. 1931. 15:- (10: -). pflanzen in Nordeuropa. 1934. 21: - (14: -). 4. B. LINDQUIST, Om den vildvaxa.nde skogsalmens raser 7. G. DEGELIUS, Das ozea.nische Element der Strauch­ och deras utbredning i Nordvii.steuropa. (Summary: und Laubflechtenflora von Skandinavien. 1935. 36: ;_ The races of spontaneous Ulmus glabra. Huds. and (24: -). ACTA PHYTOGEOGRAPHICA SUECICA 49 FEATURES OF AFROALPINE PLANT ECOLOGY avec un resume en fran�aise BY OLOV HEDBERG UPPSALA 1964 Almqvist & Wiksells Boktryckeri AB Illustrations printed with contribution from Langmanska Kulturfonden Contents Frontispiece Lobelia keniensis 59 Preface 5 Carduus keniensis 61 Introduction 7 Conclusion on "Giant rosette plants" 62 Frost heaving resistance 64 I. AFROALPINE ENVIRONMENTS 8 Xeromorphy 70 Macro climates 8 Reduced length of internodes 73 Atmospheric pressure 8 AFROALPINE LIFE-FORMS Wind conditions and cloudiness 8 Ill. 80 Precipitation . 10 Introduction 80 Insolation and (outward) long wave Current life-form systems 80 radiation . 15 Giant rosette plants . 84 Temperature conditions 16 Tussock grasses (and sedges) 86 Relative humidity . 19 Acaulescent rosette plants 87 Number of humid months 19 Cushion plants 88 Conclusion . 20 Sclerophyllous shrubs (and dwarf-shrubs) 89 Snow and glaciers 20 Other phanerogamic life-forms 90 Microclimates. 22 Solifluction floaters 92 Soils. 26 IV. WATER ECONOMY IN THE AFROALPINE Soil water 28 FLORA 93 Solifluction 29 V. PRINCIPAL AFROALPINE PLANT COMMU- Fire 35 NITIES 96 Biotic influences 3 7 Introduction 96 Dendrosenecio woodlands 96 Il. ADAPTIVE TRENDS IN THE .AFROALPlNE FLORA 41 Helichrysum scrub I 04 Introduction 41 Alchemilla scrub 110 Protection against radiation 42 Tussock grassland 114 Temperature insulation 45 Carex bogs and related communities 118 Big grass tussocks 45 Other plant communities 128 Giant leaf rosettes . 49 Vicarious plant communities 129 Senecio keniodendron 49 RESUME. 131 Senecio brassica . 52 EXPLANATION TO TABLES 7--10 132 Other Dendrosenecios; comparison with REFERENCES. 133 Espeletia . 52 Lobelia telekii 56 INDEX TO LATIN NAMES 140 Acta Phytogeogr. Suec. 49 Upper photo: Two leaf rosettes of Lobelia keniensis in night position with densely folded leaves, covered by hoar frost (cf. p. 59 and Fig. 50). The big grass tussocks are formed by Festuca pilger·i. Photo 0. Hedberg, Aug. 1948. Lower photo: Leaf rosette of Lobelia lceniensis in day position with patent leaves, between which appears the surface of the "water reservoir" described in the text (p. 59). Photo 0. Hedb rg, Aug. 1948. (Printed in Svensk Naturvetenskap, 1963.) Preface The author's first notions about afroalpine plant elaboration had to be done in spare time between ecology were received through a lecture by Pro­ other projects and has therefore required a number fessor Carl Troll given in Uppsala in the autumn of of years. In the course of this work much valuable 194 7. The impulses received on that occasion were assistance has been received from various sources, released through a minor mishap less than a year only part of which can be acknowledged here. later, when I spent a few weeks exploring the flora First of all I am indebted to Dr. Ake Holm, Upp­ on the upper parts of Mt Kenya. A bad knee forced sala, the initiator and leader of the Swedish East me to remain for a few days more or less stationary Mrica expedition in 1948, for the opportunity to around my camp in the upper part of Teleki Valley. join the expedition as well as for much help and These days were spent in making some ecological cheerful comradeship during the field work. My observations, and a few analyses of the local participation in this expedition was rendered vegetation. This vegetation, growing at an altitude economicaJly feasible by a grant from the Swedish of more than 4000 metres not far from the equator, Natural Science Research Council. The completion is exposed to a peculiar and rigorous climate with of the present book was achieved at the Institute small seasonal variations but large diurnal changes. of Systematic Botany of the University of Uppsala. Heavy frost occurs almost every night, whereas Most of the meteorological data included here were the insolation in daytime is very intense. I soon obtained from the Meteorological Institute, and discovered that the weird Giant Lobelias and Giant much of the ecological literature was consulted at Senecios growing around my camp behaved in a the Institute of Plant Ecology, both of the same very peculiar manner at night. On closer study these University. At various stages of my work I have plants proved to display some quite remarkable profited from discussions with numerous colleagues adaptations to the inhospitable afroalpine climate, and friends, notably G. E. Du Rietz, J. B. Gillett, with "summer every day and winter every night". J. Leonard, J. A. Nannfeldt, S. Nilsson, H. Rufelt, Also several other plants were found to possess R. Santesson, and H. Sjors. For multifarious help peculiar adaptations to the perils of the afro­ and encouragement, including fast and sagacious alpine environment. Similar conditions obtain, of proof-reading, I am indebted to my wife, Mrs. course, on the other high East Mrican mountains Inga Hedberg. Some of the photographs reproduced as well, and the following account embodies also here were taken by Dr. A. Holm.
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