Herbaceous Plant Ecology A.G. Van der Valk Editor Herbaceous Plant Ecology Recent Advances in Plant Ecology Previously published in Plant Ecology Volume 201, Issue 2, 2009 123 Editor A.G. Van der Valk Iowa State University Department of Ecology, Evolution and Organismal Biology 141 Bessey Hall Ames IA50011-1020 USA Cover illustration: Cover photo image: Courtesy of Photos.com All rights reserved. Library of Congress Control Number: 2009927490 DOI: 10.1007/978-90-481-2798-6 ISBN: 978-90-481-2797-9 e-ISBN: 978-90-481-2798-6 Printed on acid-free paper. © 2009 Springer Science+Business Media, B.V. No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. springer.com Contents Plant species richness and diversity of the serpentine areas on the Witwatersrand R.A. Reddy, K. Balkwill & T. McLellan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1–17 Temporal changes in species diversity and composition in abandoned fields in a trans-Himalayan landscape, Nepal C.B. Baniya, T. Solhøy & O.R. Vetaas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19–35 Role of desert annuals in nutrient flow in arid area of Northwestern China: a nutrient reservoir and provider B.-M. Chen, G.-X. Wang & S.-L. Peng . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37–45 The effects of fire frequency and grazing on tallgrass prairie productivity and plant composition are mediated through bud bank demography H.J. Dalgleish & D.C. Hartnett . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47–56 Do plant functional types based on leaf dry matter content allow characterizing native grass species and grasslands for herbage growth pattern? M. Duru, R. Al Haj Khaled, C. Ducourtieux, J.P. Theau, F.L.F. de Quadros & P. Cruz . . . . . . . . . 57–69 Clonal growth strategies in simultaneously persistent and expanding Trifolium repenspatches L. Johansen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71–80 California native and exotic perennial grasses differ in their response to soil nitrogen, exotic annual grass density, and order of emergence J.K. Abraham, J.D. Corbin & C.M. D’Antonio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81–92 Do seed and microsite limitation interact with seed size in determining invasion patterns in flooding Pampa grasslands? L.P. Herrera & P. Laterra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93–105 Feral horses dung piles as potential invasion windows for alien plant species in natural grasslands A. Loydi & S.M. Zalba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107–116 Reproductive allocation of Carex flavareacts differently to competition and resources in a designed plant mixture of five species M. Suter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117–125 Factors affecting the establishment and growth of annual legumes in semi-arid mediterranean grasslands T.P. Merou & V.P. Papanastasis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127–136 Are irrigation and grazing effects transferred, accumulated, or counteracted during plant recruitment? L. Pérez-Camacho & S. Rebollo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137–151 Extent and spatial patterns of grass bald land cover change (1948–2000), Oregon Coast Range, USA H.S.J. Zald . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153–165 Grass (Poaceae) richness patterns across China’s nature reserves Q. Liu, X. Ge, W. Chen & J.T. Columbus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167–187 Interacting effects of grass height and herbivores on the establishment of an encroaching savanna shrub N. Hagenah, H. Munkert, K. Gerhardt & H. Olff . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189–202 Effects of competition on root–shoot allocation in Plantago lanceolata L.: adaptive plasticity or ontogenetic drift? F. Berendse & F. Möller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203–209 Rock-colonizing plants: abundance of the endemic cactus Mammillaria fraileana related to rock type in the southern Sonoran Desert B.R. Lopez, Y. Bashan, M. Bacilio & G. De la Cruz-Agüero . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211–224 Ecophysiological responses of nine floodplain meadow species to changing hydrological conditions V. Jung, L. Hoffmann & S. Muller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225–234 Tolerance of a perennial herb, Pimpinella saxifraga, to simulated flower herbivory and grazing: immediate repair of injury or postponed reproduction? A-P. Huhta, P. Rautio, K. Hellström, M. Saari & J. Tuomi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235–245 Differential herbivory tolerance of dominant and subordinate plant species along gradients of nutrient availability and competition P. Tahmasebi Kohyani, B. Bossuyt, D. Bonte & M. Hoffmann . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247–255 Responses of clonal architecture to experimental defoliation: a comparative study between ten grassland species M.-L. Benot, C. Mony, S. Puijalon, M. Mohammad-Esmaeili, J.J.M. van Alphen, J.-B. Bouzillé & A. Bonis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 257–266 Diffuse knapweed (Centaurea diffusa Lam.) seedling emergence and establishment in a Colorado grassland P.J. Meiman, E.F. Redente & M.W. Paschke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267–274 Biodiversity and tallgrass prairie decomposition: the relative importance of species identity, evenness, richness, and micro-topography T.L. Dickson & B.J. Wilsey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275–285 Herbivory and local genetic differentiation in natural populations of Arabidopsis thaliana (Brassicaceae) A. Mosleh Arany, T.J. de Jong & E. van der Meijden . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287–295 The effect of nutrient supply and light intensity on tannins and mycorrhizal colonisation in Dutch heathland ecosystems J.D. Hofland-Zijlstra & F. Berendse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297–311 Spatial and temporal dynamics of floating and drift-line seeds at a tidal freshwater marsh on the Potomac River, USA K.N. Hopfensperger & A.H. Baldwin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313–322 Estimating plant competition coefficients and predicting community dynamics from non-destructive pin-point data: a case study with Calluna vulgarisand Deschampsia flexuosa C. Damgaard, T. Riis-Nielsen & I.K. Schmidt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323–333 Disturbance by mowing affects clonal diversity: the genetic structure of Ranunculus ficaria (Ranunculuaceae) in meadows and forests C. Reisch & S. Scheitler . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 335–343 Soil amendment effects on the exotic annual grass Bromus tectorumL. and facilitation of its growth by the native perennial grass Hilaria jamesii(Torr.) Benth J. Belnap & S.K. Sherrod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345–357 Effects of fire on the vegetation of a lowland heathland in North-western Italy L. Borghesio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359–367 Plant species richness and diversity of the serpentine areas on the Witwatersrand R. A. Reddy Æ K. Balkwill Æ T. McLellan OriginallypublishedinthejournalPlantEcology,Volume201,No.2,365–381. DOI:10.1007/s11258-008-9455-5(cid:1)SpringerScience+BusinessMediaB.V.2008 Abstract Soilchemistrycanplayanimportant role serpentine soils. There was no significant difference in determining plant diversity. Serpentine soils are in alpha-diversity between the serpentine and the usually toxic to many plant taxa, which limits plant adjacent non-serpentine areas, but beta-diversity is diversitycomparedtothatonadjacentnon-serpentine higher between serpentine plots than between non- soils.Theusuallyhighconcentrationsoftoxicmetals serpentine plots. Although soil factors do affect in serpentine soils are considered to be the edaphic species richness and diversity of plants on the factors that cause low diversity and high endemism. Witwatersrand toa limitedextent,the concentrations This paper aimed primarily to determine whether of soil chemicals in serpentine soils are not suffi- there is a relationship between serpentine soil chem- cientlydifferentfromthoseinnon-serpentinesoilsto istry and species richness on the Witwatersrand and significantly influence the species richness and to compare species richness of the serpentine areas diversity of the serpentine soils. The high, but with that of adjacent non-serpentine areas as well as similar, diversity on serpentine and non-serpentine withthespeciesrichnessoftheserpentineareasinthe soils on the Witwatersrand indicates that soil factors Barberton Greenstone Belt. The alpha- and beta- donotplayasignificantroleindeterminingdiversity diversity of the Witwatersrand serpentine and non- on potentially toxic soils in the area. serpentine areas was also investigated. A secondary aimofthisstudywastodeterminewhichofthenon- Keywords Alpha-diversity (cid:1) Beta-diversity (cid:1) serpentinetaxaweremorecommonontheserpentine Modified-Whittaker plots than off the serpentine, which taxa were more common off the serpentine than on the serpentine and which taxa were equally common on and off Introduction SouthernAfricahastheworld’shighestplantspecies R.A.Reddy(&)(cid:1)K.Balkwill density at the sub-continental level (Cowling et al. C.E.MossHerbarium,SchoolofAnimal,Plantand 1989). The main reasons for this high species EnvironmentalSciences,PrivateBag3,Wits2050, SouthAfrica richnessare environmental heterogeneity, two differ- e-mail:[email protected] ent climatic regimes, i.e. summer and winter rainfall regions,recurrentclimaticfluctuationssincethemid- T.McLellan Pliocene and the number of relicts that survived in SchoolofMolecularandCellBiology,Universityofthe Witwatersrand,PrivateBag3,Wits2050,SouthAfrica pockets along the coast (Goldblatt 1978). A.G.VanderValk(ed.),HerbaceousPlantEcology.DOI:10.1007/978-90-481-2798-6_1 1 2 A.G.VanderValk(ed.) Biodiversity represents the complexity of life on Environmental factors, especially soil fertility and earth (Wilsey et al. 2005). High species diversity climate,playanimportantroleindeterminingspecies means a complex community, as a greater variety of richness(Stohlgrenetal.1999;Grace2001).Edaphic speciesmeansagreaterarrayofinteractions(Brower factorsaccountformuchvariationinspeciesdataand and Zar 1984). The stability of the ecosystem is also soil pH, and the Mg:Ca ratio plays an important role linked to diversity, since most stable communities in species distribution patterns (Esler and Cowling have large numbers of species and an even distribu- 1993). In the semi-arid areas of southern Africa, tion of individuals between species (Williams 1987; soil factors play an important role in determining Tilman 1996). Humans, by overexploitation, habitat transitions between vegetation types and plant com- modification or deliberate destruction, reduce this munities (Esler and Cowling 1993). High levels of influx(Leps2004;Sodhietal.2004).Fragmentation, local diversityhave been consideredcharacteristicof grazing, forestry and human influence are decreasing nutrient poor soils in the Cape (Goldblatt and the biological diversity of earth’s resources (Tilman Manning 2002). 1996). The official estimate by the IUCN indicates Species numbers are controlled by biological that13%oftheworld’splantspeciesareunderthreat; interactions such as edaphic conditions and compe- howeverotherscientists(PitmanandJorgensen2002) tition (Grime 1979; Bond 1983 in Cowling 1990). placethisfigurebetween22%and47%.Understand- Serpentine soils are very diverse due to various ing and predicting species diversity is therefore factors such as parent material, climate, relief and essential in identifying areas vulnerable to species biological activity (Brooks 1987). These soils gen- loss and thus providing a focus for conservation erally have high concentrations of Ni, Cr, Co and efforts (Cowling et al. 1989; Pearson 1996). Fe, high Mg:Ca ratios and low concentrations of N, Three important components of species diversity P and K. All this makes for harsh environmental are: alpha-diversity—the number of species in a conditions which in turn result in a low diversity of homogenous community (Cowling et al. 1992); plant species and unique, usually endemic flora Beta-diversity—species turnover along an environ- (Kruckeburg 1954; Proctor et al. 1980). The mental or habitat gradient (Whittaker 1972); and serpentine soils on the Witwatersrand are not as gamma-diversity—species turnover for different toxic as the serpentine soils of other areas (Reddy ecosystems along geographic gradients (Cowling et al. 2001). Thus it is expected that there will be a et al. 1992). higher diversity on the serpentine soils of this area Alpha-diversity, in the strict sense, is species relative to other serpentine areas. richness(Whittaker1972).However,speciesrichness In 1934, Raunkiaer proposed a system to describe is only one aspect of diversity (Gaston 1996) and on vegetation based on the position of buds or regener- its own is not as important in an ecosystem as the ating parts which he found to be related to climate combined effects of richness and evenness (Hooper (Rickleffs 1973). He proposed several life form and Vitousek 1997; Wilsey et al. 2005). Diversity categories the major ones being: phanerophytes— indices, such as the Shannon–Weiner and Simpson buds occur on the tips of branches, associated indices, are therefore useful because they combine with tropical climates, i.e. trees and shrubs; cha- bothspeciesrichnessandevennessintoasinglevalue maephytes—buds occur at ground level during unfa- (Magurran 1988; Do¨rgeloh 1999) which can be used vourable seasons, mainly associated with cold, dry in environmental monitoring and conservation. climates, i.e. small shrubs and herbs; hemicrypto- Beta-diversityistheextent ofspeciesreplacement phytes—dormant buds occur just beneath or at the or biotic change along environmental gradients soil surface; cryptophytes—buds are deeply buried (community turnover) and depends on habitat diver- andalsostorefood;andtherophytes—‘‘annuals’’that sity(WilsonandShmida1984).Anaccuratemeasure survive in seed form, associated mainly with deserts of beta-diversity is important because it indicates the and some grasslands. Hemicryptophytes and crypto- degree to which habitats are partitioned by species, phytes are usually associated with cold, moist i.e. the degree of patchiness. Alpha- and beta- temperate areas; most dieback to ground level diversity together measure the overall diversity of during winter and regenerate the following season an area (Peet 1974; Wilson and Shmida 1984). (Rickleffs 1973). Fire plays an important role in the HerbaceousPlantEcology 3 maintenance of the grassland biome on the Witwa- low concentrations of N, P and K usually have a tersrand (Kerfoot 1987) and thus it is expected that low diversity and a high degree of endemism cryptophytes and hemicryptophytes, with their buds (Kruckeburg1954;Proctoretal.1980).Thus,the protectedbelowthesoilsurface,willbethedominant lower concentrations of chromium and nickel or life forms on the Witwatersrand. the magnesium:calcium ratio of serpentine sites The Witwatersrand region is geologically and wouldresultinahighernumberoftolerantspecies topographically diverse (McCallum and Balkwill andfewornoendemictaxa, 1999).Thisprovidesadiversityofhabitatsandhence 3. Thesimilaritybetweentheflorasoftheserpentine thepotentialforahighlevelofplantdiversityinthis and non-serpentine soils onthe Witwatersrand is region. There is no unique serpentine flora and there greater than that between the serpentine flora of arenoserpentineendemictaxaontheWitwatersrand the Witwatersrand and that of the Barberton (Reddyetal.2001).Onserpentinesoilsthathavebeen Greenstone Belt serpentines. exposedforlongperiods,prolongedevolutionhasled to a very high floral diversity at both the genus and Study areas species level (Berazain 1992; Batianoff and Specht 1992; Williamson 1995). So, with the potential high Johannesburg is situated in an area known as the diversityofthisregionandthepossiblelongexposure Highveld of South Africa. The topography of the oftheWitwatersrandserpentinesoils,aretheserpen- regionisflatgrasslandwithirregular,undulatingrocky tine areas on the Witwatersrand as species diverse as hillsandridges.AltitudeontheWitwatersrandranges theadjacentnon-serpentineareas? between1450and1750 m.Thisregion,includingthe The aims of this paper are to: (1) compare the Witwatersrandserpentineareas(Reddyetal.2001),is relationshipbetweenspeciesrichnessofthefloraand within the grassland biome and for the most part the selected soil elements at serpentine sites and their vegetation falls into the rocky Highveld grassland adjacent non-serpentine sites, (2) determine how (Bredenkamp and van Rooyen 1996). This grassland speciesrichtheserpentineareasontheWitwatersrand type, formerly known as Bankenveld, is fire-main- are and how this richness compares to the adjacent, tained (Acocks 1988),with the vegetationadaptedto non-serpentine areas, (3) investigate the alpha- and fireandconsistingmainlyofresproutingforbs.Trees beta-diversity of the Witwatersrand serpentine soils and shrubs are confined to the rocky ridges and andthe adjacent non-serpentine soils andtocompare outcropswhichoffersomeprotectionfromfire. thebeta-diversitiesoftheallWitwatersrandserpentine Precipitationrangesbetween600and750 mmper areas with those of the sites studied in the Barberton annum and occurs mainly in summer. Temperatures Greenstone Belt and (4) determine which of the range between 12(cid:2)C (daily minimum) and 39(cid:2)C bodenvagtaxa,i.e.taxathatareindifferenttoedaphic (daily maximum) with an average of 16(cid:2)C. Winters conditions, are more common on the serpentine than are very dry and severely frosty (Bredenkamp and off the serpentine, which taxa are more common off van Rooyen 1996). Many of the trees found at the the serpentine than on the serpentine and which taxa siteshavehighlatexorresincontentsinthesap(pers. areequallycommononandofftheserpentine. obs.). This is possibly a mechanism to overcome the The following predictions are made: harsh, frosty Highveld winters and another reason 1. BecausetheserpentinesoilsontheWitwatersrand why they are confined to the less fire prone, rocky are not as toxic as the serpentine soils of other areas(BalkwillandBotanyII1993,1995).Highlatex areas,e.g.theBarbertonGreenstoneBelt(Reddy or resin contents in the sap could also act as a et al. 2001), species richness of herbaceous taxa deterrent to insect herbivory. on serpentine soils on the Witwatersrand will be Theserpentineareasonthe Witwatersrandmainly similar to species richness of herbaceous taxa of occur around the south-western and south-eastern adjacent,non-serpentineareas, margins of the granite dome found to the north of 2. Studies of other serpentine sites globally have Johannesburg,andatsomelocalitieswithinthedome. shown that serpentine soils with high concentra- These sites are situated in a densely populated urban tionsofNi,Cr,CoandFe,highMg:Caratiosand area and are subject to various forms of disturbance.