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Forest Ecology A.G. Van der Valk Editor Forest Ecology Recent Advances in Plant Ecology Previously published in Plant Ecology Volume 201, Issue 1, 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: 2009927489 DOI: 10.1007/978-90-481-2795-5 ISBN: 978-90-481-2794-8 e-ISBN: 978-90-481-2795-5 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 Quantitative classification and carbon density of the forest vegetation in Lüliang Mountains of China X. Zhang, M. Wang & X. Liang . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1–9 Effects of introduced ungulates on forest understory communities in northern Patagonia are modified by timing and severity of stand mortality M.A. Relva, C.L. Westerholm & T. Kitzberger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11–22 Tree species richness and composition 15 years after strip clear-cutting in the Peruvian Amazon X.J. Rondon, D.L. Gorchov & F. Cornejo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23–37 Changing relationships between tree growth and climate in Northwest China Y. Zhang, M. Wilmking & X. Gou . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39–50 Does leaf-level nutrient-use efficiency explain Nothofagus-dominance of some tropical rain forests in New Caledonia? A. Chatain, J. Read & T. Jaffré . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51–66 Dendroecological study of a subalpine fir (Abies fargesii) forest in the Qinling Mountains, China H. Dang, M. Jiang, Y. Zhang, G. Dang & Q. Zhang . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67–75 Aconceptual model of sprouting responses in relation to fire damage: an example with cork oak (Quercus suberL.) trees in Southern Portugal F. Moreira, F. Catry, I. Duarte, V. Acácio & J.S. Silva . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77–85 Non-woody life-form contribution to vascular plant species richness in a tropical American forest R. Linares-Palomino, V. Cardona, E.I. Hennig, I. Hensen, D. Hoffmann, J. Lendzion, D. Soto, S.K. Herzog & M. Kessler . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87–99 Relationships between spatial configuration of tropical forest patches and woody plant diversity in northeastern Puerto Rico I.T. Galanes & J.R. Thomlinson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101–113 Vascular diversity patterns of forest ecosystem before and after a 43-year interval under changing climate conditions in the Changbaishan Nature Reserve, northeastern China W. Sang & F. Bai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115–130 Gap-scale disturbance processes in secondary hardwood stands on the Cumberland Plateau, Tennessee, USA J.L. Hart & H.D. Grissino-Mayer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131–146 Plurality of tree species responses to drought perturbation in Bornean tropical rain forest D.M. Newbery & M. Lingenfelder . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147–167 Red spruce forest regeneration dynamics across a gradient from Acadian forest to old field in Greenwich, Prince Edward Island National Park, Canada N. Cavallin & L. Vasseur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169–180 Distance- and density-dependent seedling mortality caused by several diseases in eight tree species co-occurring in a temperate forest M. Yamazaki, S. Iwamoto & K. Seiwa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181–196 Response of native Hawaiian woody species to lava-ignited wildfires in tropical forests and shrub- lands A. Ainsworth & J. Boone Kauffman . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197–209 Evaluating different harvest intensities over understory plant diversity and pine seedlings, in a Pinus pinasterAit. natural stand of Spain J. González-Alday, C. Martínez-Ruiz & F. Bravo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211–220 Land-use history affects understorey plant species distributions in a large temperate-forest complex, Denmark J.-C. Svenning, K.H. Baktoft & H. Balslev . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221–234 Short-term responses of the understory to the removal of plant functional groups in the cold-temperate deciduous forest A. Lenière & G. Houle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235–245 Host trait preferences and distribution of vascular epiphytes in a warm-temperate forest A. Hirata, T. Kamijo & S. Saito . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247–254 Seed bank composition and above-ground vegetation in response to grazing in sub-Mediterranean oak forests (NWGreece) E. Chaideftou, C.A. Thanos, E. Bergmeier, A. Kallimanis & P. Dimopoulos . . . . . . . . . . . . . . . . . . 255–265 On the detection of dynamic responses in a drought-perturbed tropical rainforest in Borneo M. Lingenfelder & D.M. Newbery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267–290 Changes in tree and liana communities along a successional gradient in a tropical dry forest in south-eastern Brazil B.G. Madeira, M.M. Espírito-Santo, S. D’Ângelo Neto, Y.R.F. Nunes, G. Arturo Sánchez Azofeifa, G. Wilson Fernandes & M. Quesada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291–304 Woody plant composition of forest layers: the importance of environmental conditions and spatial configuration M. Gonzalez, M. Deconchat & G. Balent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 305–318 The importance of clonal growth to the recovery of Gaultheria procumbens L. (Ericaceae) after forest disturbance F.M. Moola & L. Vasseur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 319–337 Species richness and resilience of forest communities: combined effects of short-term disturbance and long-term pollution M.R. Trubina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 339–350 Hurricane disturbance in a temperate deciduous forest: patch dynamics, tree mortality, and coarse woody detritus R.T. Busing, R.D. White, M.E. Harmon & P.S. White . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 351–363 Quantitative classification and carbon density of the forest vegetation in Lu¨liang Mountains of China Xianping Zhang Æ Mengben Wang Æ Xiaoming Liang OriginallypublishedinthejournalPlantEcology,Volume201,No.1,1–9. DOI:10.1007/s11258-008-9507-x(cid:1)SpringerScience+BusinessMediaB.V.2008 Abstract Forestsplayamajorroleinglobalcarbon the 9 forest formations was 32.09 Mg ha-1 in 2000 (C) cycle, and the carbon density (CD) could reflect and33.86 Mg ha-1in2005.Form.Piceameyerihad its ecological function of C sequestration. Study on thehighestCD(56.48 Mg ha-1),andForm.Quercus theCDofdifferentforesttypesonacommunityscale liaotungensis ? Acer mono had the lowest CD is crucial to characterize in depth the capacity of (16.14 Mg ha-1). Pre-mature forests and mature forest C sequestration. In this study, based on the forestswere very important stages inC sequestration forest inventory data of 168 field plots in the study among four age classes in these formations. Forest area (E 111(cid:2)300–113(cid:2)500, N 37(cid:2)300–39(cid:2)400), the densities, average age of forest stand, and elevation forest vegetation was classified by using quantitative hadpositiverelationshipswithforestCD,whileslope method (TWINSPAN); the living biomass of trees location had negative correlation with forest CD. was estimated using the volume-derived method; the CD of different forest types was estimated from the Keywords TWINSPAN (cid:1) Carbon density (cid:1) biomassoftheirtreespecies;andtheeffectsofbiotic Volume-derived method (cid:1) Forest vegetation (cid:1) and abiotic factors on CD were studied using a China multiple linear regression analysis. The results show that the forest vegetation in this region could be classifiedinto9forestformations.TheaverageCDof Introduction Forests play a major role in global carbon (C) cycle X.Zhang(cid:1)M.Wang(&) (Dixon et al. 1994; Wang 1999) because they store InstituteofLoessPlateau,ShanxiUniversity, 80% of the global aboveground C of the vegetation 580WuchengRoad,Taiyuan030006, and about 40% of the soil C and interact with People’sRepublicofChina e-mail:[email protected] atmospheric processes through the absorption and respiration of CO (Brown et al. 1999; Houghton 2 X.Zhang et al. 2001a, b; Goodale and Apps 2002). Enhancing ShanxiForestryVocationalTechnologicalCollege, C sequestration by increasing forestland area has Taiyuan030009,People’sRepublicofChina been suggested as an effective measure to mitigate X.Liang elevated atmospheric carbon dioxide (CO ) concen- 2 GuandiMountainState-OwnedForestManagement trationandhencecontributetowardthepreventionof BureauofShanxiProvince,Jiaocheng,Lishi032104, People’sRepublicofChina global warming (Watson 2000). Recent researches A.G.VanderValk(ed.),ForestEcology.DOI:10.1007/978-90-481-2795-5_1 1 2 A.G.VanderValk(ed.) focus mainly on carbon storage of forest ecosystem Methods on landscape or regional scale (Fang et al. 2001; Hiura 2005; Zhao and Zhou 2006). Many studies Study region have shown that the C sequestration abilities of different forests change considerably, which can be The study was conducted in the middle-north of well explained by their CD values (Wei et al. 2007; Lu¨liang Mountains (E 111(cid:2)300–113(cid:2)500, N 37(cid:2)300 HuandLiu2006).MeanwhiletheCstorageofforests –39(cid:2)400) with its peak (Xiaowen Mountain) 2831 m maychangesubstantiallywithforestecosystemsona abovesealevel(asl).Thetemperateterrestrialclimate community scale. This type of moderate-scale ischaracterizedbyawarmsummer,acoldwinter,and research into the C storage of forests, however, has a short growing season (90–130 days) with a mean been rarely conducted. annual precipitation of 330–650 mm and a mean Many methods have been used to estimate the annual temperature of 8.5(cid:2)C (min. monthly mean of biomass of forest vegetation (Houghton et al. 2001a, -7.6(cid:2)CinJanuaryandmax.monthlymeanof22.5(cid:2)C b). Among them, the volume-derived method has in July). The soils from mountain top to foot are been commonly used (Brown and Lugo 1984; Fang mountain meadow soil, mountain brown soil, moun- et al. 1996; Fang and Wang 2001). Forest volume tain alfisol cinnamon soil, and mountain cinnamon production reflects the effects of the influencing soil (The Editing Committeeof Shanxi Forest1984). factors, such as the forest type, age, density, soil There are two national natural reserves in this condition, and location. The forest CD estimated region with Luya Mountain National Nature Reserve from forest biomass will also indicate these effects. in the north and Pangquangou National Nature Zhou et al. (2002) and Zhao and Zhou (2005) Reserve in the south, in which Crossoptlon mant- improved the volume-derived method by hyperbolic churicum (an endangered bird species), Larix function, but the method has not been used to principis-rupprechtii forest, and Picea spp. (P. mey- estimate forest CD on the moderate scale. eri and P. wilsonii) forest are the key protective The Lu¨liang Mountains is located in the eastern targets. part of the Loess Plateau in China, where soil and Based on the system of national vegetation water losses are serious. To improve ecological regionalization, this area was classified into the environmentthere, theChinesegovernment hasbeen warm-temperate deciduous broad-leaved forest zone. increasing forestland by carrying out ‘‘The Three- With the elevation rising, vegetation zone are, North Forest Shelterbelt Program,’’ ‘‘The Natural respectively, deciduous broad-leaved forest, needle- Forest Protection Project,’’ and ‘‘The Conversion of broad-leavedmixedforest,cold-temperateconiferous Cropland to Forest Program’’ since 1970s. Previous forest, and subalpine scrub-meadow. studies on the forest vegetation in this region focus mainly on the qualitative description of its distribu- Data collection tionpattern(TheEditingCommitteeofShanxiForest 1984). The objectives of this study were (1) to The forest inventory data from a total of 168 field classify the forest vegetation on Lu¨liang Mountains plotsin2000and2005wereusedinthisstudy.These usingquantitativeclassificationmethod(TWINSPAN) permanent plots (each with an area of 0.0667 ha) (Zhang et al. 2003; Zhang 2004); (2) to estimate the were established systematically based on the grid of CD of different forest types through biomass based 4 km 9 4 km across the forestland of 2698.85 km2 on the modified volume-derived method (Zhou et al. in 1980s under the project of the forest survey of the 2002) and to clarify the distribution pattern of forest Ministry of Forestry of P. R. China (1982), in which CDinthisregion;and(3)toquantifythecontribution the data, such as tree species, diameter at breath ofbioticandabioticfactors(includingaverageforest height of 1.3 m (DBH), the average height of the age, density, soil thickness, elevation, aspect, and forest stand, and the average age of the forest stand slope) to forest CD based on a multiple linear had been recorded along with the data of location, regression analysis. The results would provide basic elevation, aspect, slope degree, slope location, and data for further study of forest C storage pattern in soil depth. For trees with C5 cm DBH, the values of this region. their DBH were included in the inventory. ForestEcology 3 TWINSPAN classification Table1 Parameters of biomass calculation for dominant speciesinthisstudy A total of 26 tree species had been recorded in the Species Parametersinequation 168 plots. The importance values (IV) for every tree a b n R2 species in each plot were calculated using the following formula: Larixprincipis-rupprechtii 0.94 0.0026 34 0.94 Pinustabulaeformis 0.32 0.0085 32 0.86 IV¼ ðRelative densityþRelative dominance Piceameyeri 0.56 0.0035 26 0.85 þRelative frequencyÞ=300 Platycladusorientalis 1.125 0.0002 21 0.97 where relative density is the ratio of the individual Pinusarmandii 0.542 0.0077 17 0.73 number for a tree species over the total number for Populusdavidiana 0.587 0.0071 21 0.92 all tree species in a plot, relative dominance is the Betulaplatyphylla 0.975 0.001 14 0.91 ratio of the sum of the basal area for a tree species Quercusliaotungensis 0.824 0.0007 48 0.92 over the total basal area of all tree species in a plot, and the relative frequency is the percentage of the plot number containing a tree species over the total CD¼B(cid:2)C ð2Þ c plot number (168) in this inventory. Based on the whereBisthetotallivingbiomassoftreespeciesina matrix of IVs of 26 9 168 (species 9 plots), the plot; C is the average carbon content of dry matter, forest vegetation can be classified into different C which is assumed to be 0.5, though it varies slightly formations using the two-way indicator-species for different vegetation (Johnson and Sharpe 1983; analysis (TWINSPAN) (Hill 1979). Zhao and Zhou 2006). Effects of influencing factors Estimation of biomass and CD The qualitative data of the aspect and slope location Thevolumeproductionofanindividualtreecouldbe were first transformed into quantitative data to obtained in the volume table (Science and Technol- quantify their effects on forest CD. According to ogy Department of Shanxi Forestry Bureau 1986) the regulations of the forest resources inventory by according to its DBH. The volume of a species (V) the Ministry of Forestry (1982), the aspect data were was the sum ofits individual tree’s volume ina plot. transformedtoeightclassesstartingfromnorth(from The total living biomass (B) (Mg ha-1) of a species 338(cid:2)to360(cid:2)plusfrom0(cid:2)to22(cid:2)),turningclockwise, in a plot was calculated as: and taking every 45(cid:2) as a class: 1 (338(cid:2)–22(cid:2), north V aspect), 2 (23(cid:2)–67(cid:2), northeast aspect), 3 (68(cid:2)–112(cid:2), B¼ ð1Þ aþbV east aspect), 4 (113(cid:2)–157(cid:2), southeast aspect), 5 where V represents the total volume (m3 ha-1) of a (158(cid:2)–202(cid:2), south aspect), 6(203(cid:2)–247(cid:2),southwest), speciesinaplot,a(0.32–1.125)andb(0.0002–0.001) 7 (248(cid:2)–292(cid:2), west aspect), and 8 (293(cid:2)–337(cid:2), are constants (Zhou et al. 2002). The constants for northwest aspect). The slope locations in the moun- most ofthe tree species inthis study were developed tains were transformed to 6 grades: 1 (the ridge), 2 by Zhao and Zhou in 2006 (Table 1). (the upper part), 3 (the middle part), 4 (the lower In regard to companion tree species in this study, part), 5 (the valley), and 6 (the flat). theirbiomassestimationwasbasedontheparameters A multiple linear regression model was used to of above known species according to their morpho- analyze the effects of biotic and abiotic factors on logical similarity, i.e., Pinus bungeana is referred to forest CD, assuming a significant effect if the theparametersofPinusarmandii;Ulmuspumillaand probability level (P) is\0.05: TiliachinensistothoseofQuercusliaotungensis;and Y^¼aþb1X1þb2X2þb3X3þ...þbkXkh ð3Þ Acer mono and the rest of broad-leaved species to those of Populus davidiana. whereaisaconstant,b ,b ,b ,andb areregression 1 2 3 k Forest CD (Mg ha-1) was calculated as: coefficients. Y^represents CD and X , X , X , X , X , 1 2 3 4 5 4 A.G.VanderValk(ed.) X6, and X7 represent forest density (X1), average age 168 plots (X ), elevation (X ), slope location (X ), aspect (X ), 2 3 4 5 slope degree (X ), and soil depth (X ) in each plot, 6 7 respectively. Here forest density is the individual number of all tree species per area in a plot, and 2nd level forest age isthe average age ofdominanttrees inthe 3rd level plot. 4th level Results 1 2 3 4 5 6 7 8 9 (12) (20) (17) (24) (35)(26) (11) (5) (18) Forest formations from TWINSPAN Fig.1 DendrogramderivedfromTWINSPANanalysis.Note: 1.Form.Larixprincipis-rupprechtii;2.Form.Piceameyeri;3. According to the 4th level results of TWINSPAN Form.Betulaplatyphylla;4.Form.Populusdavidiana;5.Form. classification, the 168 plots were classified into 9 Pinustabulaeformis;6.Form.Pinustabulaeformis?Quercus formations (Table 2), which were named according liaotungensis;7.Form.Quercusliaotungensis;8.Form.Pinus bungeana?Platycladus orientalis, and 9. Form. Quercus to Chinese Vegetation Classification system (Wu liaotungensis?Acer mono. The number of plots for each 1980). The dendrogram derived from TWINSPAN formationisshownbetweenthebrackets analysis is shown in Fig. 1. The basic characteristics of species composition, structure along with its common companion species were Picea meyeri environment for each formation are described as and P. wilsonii in the tree layer. follows: 2. Form. Picea meyeri (Form. 2): P. meyeri forest 1. Form. Larix principis-rupprechtii (Form. 1 for belongedtocold-temperateevergreenconiferous short, the same thereafter): L. principis- forest. Its ecological amplitude was relatively rupprechtii was the dominant tree species of narrow with a range of vertical distribution from the cold-temperate coniferous forest in north 1860 mto2520 m.BetulaplatyphyllaandPicea China. It grew relatively faster with fine timber. wilsonii appeared commonly in this forest. Therefore it was a very important silvicultural 3. Form.Betulaplatyphylla(Form.3):B. platyphy- tree species at middle-high mountains in this lla was one of main tree species in this region region. This type of forest distributed vertically and occupied the land at moderate elevation from 1610 m to 2445 m above sea level, and (1700–2200 m). In the tree layer, Populus Table2 Thestructurecharacteristicsof9forestformationsandtheirenvironmentalfactors Form Density(No./ha) Age(Year) Coverage(%) Slopelocation Elevation(m) Slope((cid:2)) Aspect Soildepth(cm) 1 849.3±121.8 40.0±5.4 54±8.7 2.7±0.1 1610–2445 19.1±1.1 4.1±.6 56.4±5.1 2 869.6±179.1 55.4±4.8 62±8.3 2.3±0.2 1860–2520 19.6±2.2 4.7±0.6 50.6±5.9 3 774.3±57.8 45.5±5.3 45±4.1 2.6±0.2 1700–2200 21.6±1.9 4.2±0.8 48.7±3.3 4 1071.9±124.4 31.6±2.6 41±6.3 3.5±0.2 1350–1997 23.0±1.6 4.1±0.6 49.2±6.2 5 770.9±139.7 54.7±2.6 49±5.7 2.9±0.2 1360–2010 23.9±2.2 2.9±0.5 41.0±4.1 6 756.2±87.7 60.9±3.7 46±4.2 2.6±0.2 1235–1820 29.4±2.3 3.7±0.4 34.2±4.1 7 731.3±154.7 56.8±6.2 46±7.4 3.0±0.3 1452–2010 25.9±2.1 3.4±0.8 53.2±3.7 8 1589.2±616.2 53.8±3.8 41±2.5 2.6±0.5 1250–1270 26.6±3.5 3.6±0.7 34.0±7.1 9 910.3±136.8 51.3±4.6 51±7.3 3.4±0.2 1350–1660 23.2±2.5 4.8±0.5 39.4±4.4 Note:1.Form.Larixprincipis-rupprechtii;2.Form.Piceameyeri;3.Form.BetulaPlatyphylla;4.Form.Populusdavidiana;5.Form. Pinus tabulaeformis; 6. Form. Pinus tabulaeformis?Quercus liaotungensis; 7. Form. Quercus liaotungensis; 8. Form. Pinus bungeana?Platycladusorientalis;9.Form.Quercusliaotungensis?Acermono

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