THE ELECTRONIC COMPARATIVE PLANT ECOLOGY THE ELECTRONIC COMPARATIVE PLANT ECOLOGY Incorporating the principal data from Comparative Plant Ecology and The Abridged Comparative Plant Ecology J.G. Hodgson, J.P. Grime, R. Hunt and K. Thompson The NERC Unit of Comparative Plant Ecology Department of Animal and Plant Sciences University of Sheffield, UK. Springer-Science+Business Media, B.V. First edition 1995 © 1995 Springer Science+Business Media Dordrecht Originally published by Chapman & Hall in 1995 ISBN 978-94-011-7652-1 ISBN 978-94-011-0559-0 (eBook) DOI 10.1007/978-94-011-0559-0 Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the prior permission in writing of the publishers, or in the case of reprographic reproduction only in accordance with the terms of the licences issued by the Copyright Licensing Agency in the UK, or in accordance with the terms of licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries concerning reproduction outside the terms stated here should be sent to the publishers at the London address printed on this page. The publisher makes no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that may be made. A catalogue record for this book is available from the British Library @ Printed on acid-free text paper, manufactured in accordance with ANSI/NISO Z39.48-1992 (Permanence of Paper). Contents Preface vi INTRODUCTION 2 EXPLANATION OF TABLES 2 2.1 Nomenclature 2 2.2 Ordering of species 2 3 FIRST GROUP: ECOLOGICAL ATTRIBUTES 3 3.1 Species 3 3.2 Habitat range 3 3.3 Soil pH 5 3.4 Floristic diversity 6 3.5 Distribution in N Europe 6 3.6 Present status 7 4 SECOND GROUP: ATTRIBUTES OF THE ESTABLISHED PHASE 7 4.1 Life history 7 4.2 Established strategy 8 4.3 Life form 8 4.4 Canopy structure 9 4.5 Canopy height 10 4.6 Lateral spread 10 4.7 Mycorrhizas 11 4.8 Leaf phenology 12 4.9 Flowering time and duration 12 4.10 Polyploidy 13 5 THIRD GROUP: ATTRIBUTES OF THE REGENERATIVE PHASE 13 5.1 Regenerative strategies 13 5.2 Seed bank 14 5.3 Agency of dispersal 15 5.4 Dispersule and germinule form 16 5.5 Dispersule weight 17 5.6 Dispersule shape 17 5.7 Germination requirements 17 5.8 Family 18 Preface The 3Y2-inch floppy disk which accompanies this booklet is a standard DSIHD 1.44MB disk formatted by Microsoft® MS-DOSTM. The disk contains five files, all called ecpe (standing for Electronic Comparative Plant Ecology). All five have identical content but differ in file type. Only one of these files need be selected by the user, according to the software which will be used locally to examine the data. Filename Software ecpe.txt ascii text only, for MS-DOSTM ecpe.csv comma-delimited entries, ditto ecpe.xls for Microsoft® Excel 5.0 ecpe.wk3* for WK3 format in Lotusl-2-3® ecpe.csm comma-delimited for Macintosh® *this file is accompanied by the usual ecpe. fm3. The Electronic Comparative Plant Ecology 1 INTRODUCTION This publication consists of a collection of standardized autecological information for the more common vascular species of the British flora. It is an amalgamated, electronic copy of the data tables which concluded both of the printed works Comparative Plant Ecology (CPE) and The Abridged Comparative Plant Ecology (ACPE). In addition to providing summaries for the 281 species which are the subjects of autecological accounts in CPE and in ACPE, the table includes data on a further 221 species, which each satisfy all of three criteria: (a) they are recorded in at least 66% of Watsonian vice-counties of the British Isles; (b) they are either native or a well-naturalized alien throughout the majority of their British range; and (c) they have an extensive distribution in inland Britain. Ecological information for these additional species (and for Carlina vulgaris, Phragmites australis and Reynoutria japonica, which were recorded from fewer than ten quadrats in UCPE's 'Survey II') is derived from unpublished UCPE data. Only one of the species tabulated, Sedum anglicum Hudson, was not recorded during fieldwork in the Sheffield area. The quantity and quality of information available varies considerably according to species. Certain species are variable in certain biological characteristics, and field distributions with respect to habitat features such as slope and aspect may be too strongly affected by geographical location to allow reliable generalizations at a national scale. Despite these problems, it has been possible to present data for 22 characteristics which, apart from caveats included in the explanatory notes, can be applied with some confidence to the species observed in the British Isles; these relate to ecology (Group 1), attributes of the established phase (Group 2) and attributes of the regenerative phase (Group 3). These Groups correspond to Tables 6.1, 6.2 and 6.3 respectively in CPE and to Tables 4.1, 4.2 and 4.3 respectively in ACPE. The significance and use of each of the 22 characteristics is discussed in both CPE and ACPE. 2 The electronic comparative plant ecology 2 EXPLANATION OF TABLES 2.1 Nomenclature Nomenclature follows that of A. R. Clapham, T. G. Tutin and E. F. Warburg's Excursion Flora oj the British Isles (3rd. edn), published by Cambridge University Press in 1981, except for the following species groupings: Betula spp. (includes B. pendula, B. pubescens and their hybrid), Festuca ovina (also includes F. tenuifolia), Festuca rubra (also includes F. nigrescens), Lolium perenne (refers only to ssp. perenne; the monocarpic, and introduced, ssp. multiflorum (Lam.) Husnot is a non persistent escapee from cultivation), Medicago sativa (excludes ssp. Jalcata (L.) Arcangeli which is native in East Anglia), Nasturtium officinale agg. (N microphyllum, N officinale and their hybrid), Poa pratensis agg. (P. angustifolia, P. pratensis sensu stricto and P. subcaerulea), Quercus agg. (Q. petraea, Q. robur and their hybrid), Rosa spp. (R. can ina group, R. rubiginosa group and R. tomentosa group), Salix cinerea agg. (s. aurita L., S. caprea, S. cinerea and hybrids), Salix fragilis agg. (s. alba L., S. fragilis L. and related hybrids) and Taraxacum agg. (Section Erythrosperma, Section Obliqua, Section Palustria, Section Spectabilia and Section Taraxacum). Two columns of symbols precede the scientific name. In the first, species which are the subject of autecological accounts in CPE and ACPE are indicated by a '+'; in the second, alien species are denoted by an asterisk. The authorities for the scientific names are cited. 2.2 Ordering of species Species lists are divided into three groups: (a) herbs, and woody species up to 1.5 m in height, (b) woody species exceeding 1.5 m in height, and (c) pteridophytes. Within each group species are arranged in alphabetical order. Ecological attributes 3 3 FIRST GROUP: ECOLOGICAL ATTRIBUTES 3.1 Species For most species only one set of data is given. However, in the case of individual trees and shrubs ecological attributes are, wherever possible, presented separately for seedlings and small saplings, juveniles, and for the more mature individuals which contribute to the canopy. 3.2 Habitat range (a) General policy A habitat key (published in CPE Fig. 2.2 and in ACPE Fig. 1.3) has been used as a basis for identifying the habitat range of each species. First, an estimate of the abundance in each of the seven primary habitat groups (wetland, skeletal, arable, pasture, spoil, wasteland and woodland) is provided. Second, the terminal habitat in which the species is most frequent is indicated. (b) Abundance in the seven primary habitats The percentage frequency of the species in each of these major groups is compared with the percentage frequency of the species in the overall survey. The symbols used are as follows: ++ very common and characteristic of the particular habitat (percentage frequency >4 times the overall value) + common within habitat (2 -4 times the overall value) widespread in the habitat (0.5 -2 times the overall value) infrequent and uncharacteristic of the habitat (0.25 -0.5 times the overall value) largely absent from habitat or confined to an uncommon variant «0.25 of the overall value). In Woodland in the UCPE 'Survey II' and in Wasteland in 'Survey III' these primary habitat groups each constitute over 25% of all records, rendering the ++ rating an impossibility. It was decided that all species in 4 The electronic comparative plant ecology which at least 80% of records fall within a single primary habitat should also be classified as ++. For species for which there are insufficient field records (Hypericum androsaemum L., Salix repens L. and Sedum anglicum Hudson) only the identity of the most characteristic major habitat is indicated. (c) Commonest terminal habitat The commonest terminal habitat for the subject species is identified as follows: AQUATp (lakes, canals, ponds and ditches), AQUATr (rivers and streams), ARABLE (arable), BRICK (bricks and mortar rubble), CANALB (canal banks, +), CINDER (cinder tips and cindery railway tracks), CLIFF (cliffs), COAL (coal-mine spoil), HEDGE (hedgerows), LEAD (lead-mine spoil), MANURE (manure and sewage spoil), MEADOW (meadows), MIREs (shaded mire), MIREu (unshaded mire), OUTCRP (rock outcrop), PASTa (pasture on acidic strata), PASTe (enclosed pasture), PASTI (pasture on limestone strata), PATH (paths), PLANTb (broadleaved plantations), PLANTc (coniferous plantations), PONDBK (banks of lakes, canals, ponds and ditches, +), QRYa (quarry spoil on acidic strata), QYRl (limestone quarry spoil), RDIRLY (road verges/railway banks, +), RDVRGE (road verge), RIVBNK (river and stream banks), RLY BNK (railway banks, +), SCREE (limestone scree), SCRUB (scrub), SNDPIT (sand and gravel pits, +), SOIL (soil heaps), WALL (walls), WASTEa (wasteland on acidic strata), WASTEd (dry sandy wasteland droughted during summer, +), WASTEc (wasteland on calcareous strata), WOODa (woodland on acidic strata), WOODI (woodland on limestone strata). Habitats suffixed + are minor habitats which, because of their infrequent occurrence, were excluded from the habitat classification of 'Survey II' (CPEFig. 2.2 and ACPE Fig. 1.3). These data are particularly useful for categorizing species of narrow ecological range (e.g. Chaenorhinum minus, associated with cinders, and Lemna minor, associated with lakes, canals, ponds and ditches). However, three grassland species (Euphrasia officinalis, Leontodon autumnalis and Rumex acetosa) and seedlings of Ulmus glabra were Ecological attributes 5 most commonly recorded from lead-mine spoil even though they are characteristic of a range of base-rich habitats. In order to prevent the impression that these species are metallophytes and largely restricted to soils contaminated with heavy metals, the habitat in which these species are second most common is substituted. Rather anomalous data were obtained for two wetland species which are persistent after drainage. Eriophorum vaginatum was recorded most commonly in pasture on limestone strata and Lotus uliginosus most commonly in wasteland on limestone. In both species the second most common habitat (unshaded mire) has been promoted to first place. Four species which do not normally reach maturity in their most common habitat (Athyrium fllix jemina, cliffs and walls, Dryopteris fllix-mas, walls, Equisetum palustre, rivers and streams, and Ulex europaeus, river banks) have been similarly treated. For each of these four species the habitat code is preceded by an asterisk to warn that the information has been modified in this manner. These examples illustrate that data concerning the most common terminal habitat could, if utilized in isolation, lead to misinterpretation. Their use in conjunction with the distribution of the species in the seven major habitats is advised. N.B. The range and diversity of major habitats differs according to region. For example, in areas where (unlike that sampled here) there is little limestone, species such as Carex panicea would be regarded as a wetland rather than as a grassland species. 3.3 Soil pH The data take the form of a numeral indicating the modal pH class for the species, followed by a letter indicating the number of pH classes in which the frequency of the species exceeds 50% of that in its modal class. Thus, 7a would indicate that the species is most frequent within the range 7.0 - 7.9 and has a range of only I pH unit. In contrast, 3d would indicate a mode within the interval pH 3.0 - 3.9 and a range of 4 pH units. Where information on soil surface pH is scarce, the estimate of modal class is preceded by a question mark. A few species exploit both strongly acidic and highly calcareous soils and thus possess a bimodal pH distribution. They are identified by a W