- Email: [email protected]
The biological flora of Central Europe — aims and concept
Flora (2000) 195, 116-122 http://www.urbanfischer.de/joumals!flora ©by Urban & Fischer Verlag
The biological flora of Central Europe - aims and concept DIETHART MATTHIES 1 and PETER POSCHLOD2 1
2
Institut fiir Umweltwissenschaften, Universitat Ziirich, Winterthurerstr. 190, CH-8057 ZUrich, Schweiz. Current address: Fachgebiet Pflanzenokologie, Fachbereich Biologie, Philipps-Universitat, D-35032 Marburg. Tel.: +49-6421-2822085, Fax: +49-6421-2822057 E-mail : matthies@ mailer. uni-marburg.de Fachbereich Biologie, Fachgebiet Naturschutz (II), Philipps-Universitat, D-35032 Marburg
Accepted: April30, 1999
Summary Information on the biology of individual species is important in many fields, but standard floras contain very little information on the ecology and life-history of plants. We call for collaboration in a new series: "The biological flora of Central Europe". Contributions to the series should summarize the available knowledge of the biology of a plant occurring in Central Europe. The concept of the "Biological Flora of Central Europe" and a standard format for contributions is presented. Key words: Flora, ecology, species biology.
I. Introduction Information on the biology of plant species is very important in many fields. Understanding the life-cycle of a species is a prerequisite for understanding its population dynamics. Population processes like the decline of threatened plant species (SCHMID & MATTHIES 1994, FISCHER & MATTHIES 1998) or the rapid expansion of the range and abundance of non-native plants (SUKOPP & KOWARIK 1986, DI CASTRI et al. 1990) cannot be understood if only abiotic factors are analysed (MATTHIES et al. 1995, POSCHOLD et al. 1998). Data on the population biology of a species, its dispersal capabilities, the persistence of the seed bank, its breeding system and pollinators make it possible to predict more accurately the response of a plant population to human interventions and to suggest management measures. Similarly, the management of weeds in agricultural systems or of agressive exotic species in nature reserves requires a comprehensive knowledge of the biology of the respective species. Because of the population structure of vegetation, an understanding of the population biology of species is also frequently necessary to understand the dynamics of vegetation (WHITE 1985). Moreover, many problems encountered in botanical research (unsatisfactory germination, weak growth, uncertainty about the time of first flowering) 116
FLORA (2000) 195
could be avoided if autecological and population-biological data were more easily available. The Central European standard floras contain very little information on the biology of species (e.g. HEss et al. 1967, 0BERDORFER 1994, ROTHMALER 1990-99, SCHMEIL-FITSCHEN 1993, ADLER et al. 1994; for an assessment ofHEGI 1908ff, 1935ff, 1981 ff, see below). Anyone interested in a certain species has to collect the required data from many different sources, which is very laborious and time-consuming. Moreover, there are frequently large gaps in our knowledge of individual species. Many sources that are particularly valuable (e.g. Diploma theses) have not been published and are thus not generally accessible. To reduce the obvious deficits in the information available on the biology of plants, we call for collaboration in a new series : "The Biological Flora of Central Europe". This project originated within the framework of the "Working Group on Plant Population Biology" of the German Ecological Society. The "Biological Flora" is planned as a series of papers in the journal "Flora" that summarize the available information on individual species. In the following we will first give a brief summary of earlier attempts to compile the available information on the biology of plants growing in Central Europe, and discuss the relationship between "The Biological Flora 0367-2530/001195/02-116
$ 12.00/0
of Central Europe" and other biological floras. We then present the concept of the "Biological Flora of Central Europe" and a standard format for contributions. Selected works of reference which contain useful information on the biology of plant species are given in an appendix.
II. Other biological floras One of the oldest floras containing information on the biology of some species is the "Flora von Stuttgart" (KIRCHNER 1888). In particular the reproductive biology of plants is well covered. Later, this flora formed the core of the ambitious "Lebensgeschichte der Bltitenpflanzen Mitteleuropas" (KIRCHNER et al. 1908 to 1940), which, however, remained unfinished. A very useful series that is still continued is the "Biological Flora of the British Isles", which is published in the "Journal of Ecology" (BRITISH EcoLOGICAL SociETY 1941). This British series is one of the best sources of information on individual plant species and in many respects the model for the "Biological Flora of Central Europe". More recent series include the "Biological Flora of the Moscow Region" (RABOTNOV 1974ff.), and the "Biology of Canadian Weeds" (CAVERS & MULLIGAN 1972). A detailed description of the various biological floras (including electronic data bases), their history, aims and concepts, can be found elsewhere (PoscHLOD et al. 1996). A lot of biological information can also be found in the volumes of the second and third editions of the "Illustrierte Flora von Mitteleuropa" (HEGI 1908ff, 1935ff, 1981ff). However, the main emphasis of this work is still on taxonomy and morphology, and important biological data (e.g. breeding system, seed mass, germination requirements, longevity of seeds in soil) are frequently still missing in recent volumes. Moreover, for several families only the first edition from the beginning of the century is available, which contains hardly any information on the biology of species. Because the "Illustrierte Flora von Mitteleuropa" is published as a series of books, new volumes can only be added once a large number of taxonomically related species has been revised. This strongly delays the publication of new information. In contrast, the "Biological Flora of Central Europe" is planned as a series of separate papers. This makes it possible to publish contributions on individual species as they become available. We especially hope to encourage botanists to write contributions, who have intensively studied individual species (e.g. for their diploma or PhD-thesis), and know them well. According to our experience there exists a lot of knowledge on the biology of plants that is currently inaccessible.
III. Relationship of the "Biological Flora of Central Europe" to similar projects elsewhere The model for the project "Biological Flora of Central Europe" is the "Biological Flora of the British Isles", whose structure we have largely followed. The "Biological Flora of Central Europe" is intended to be complementary to the British series. Many Central European plant species do not occur in the British Isles and the British series has made only slow progress. We think that Central European botanists, who have perhaps shied away from writing a contribution to the British series, can make important contributions to a better knowledge of the European flora. Contributions to the "Biological Flora" should be written in English. The question of the language has been discussed by the members of the "Working group on Plant Population Biology" and a clear majority was in favour of English. We are aware that this may present a hurdle for some who would like to contribute to the series, but we hope that potential problems can be overcome.
IV. Procedure of compiling contributions to the "Biological Flora of Central Europe"
IV.l. Basis The basis of a contribution to the "Biological Flora" is a thorough analysis of available information in the literature. In appendix 1 of this paper a list of standard works of reference is given, which are useful sources of information on certain aspects of the biology of plants. A lot of the older literature on the biology of central European plants is listed in JAGER & MOLLER-URI (1982). A literature search in the "Biological Abstracts" is strongly recommended. In case of species that are important weeds the "Weed Abstracts" often contain additional information. The analysis of the literature should be complemented by specific studies to fill important gaps in the knowledge of a species. One of the aims of the "Biological Flora" is indeed to encourage studies on the biology of plants. In order to make it possible to assess the reliability of information, the sources of all information should be given in form of references. This is also useful for further studies of the literature. Data for which no reference is given are assumed to be based on observations by the author. Contributions on all species that occur in Central Europe (including the Alps) are welcome (see EHRENDORFER 1973, WISSKIRCHEN & HAEUPLER 1998). All contributions should conform to a standard format (see FLORA (2000) 195
117
below). If no information is available on a major point, this should be stated explicitly. The standard format provides a framework for contributions, but leaves the authors a lot of freedom. We have deliberately avoided to give very detailed and inflexible instructions, because otherwise potential contributors might be deterred if no information is available on many points. It is also possible for individual contributions to have a different emphasis. In general, species for which an account in the "Biological Flora of the British Isles", or the "Biology of Canadian Weeds" is available, should have a low priority. However, some of the early contributions to the British series are very brief, and those species would certainly merit a new account. A list of Central European species, for which accounts exist in one or more biological floras is given in PoscHLOD et al. (1996). Especially welcome are accounts on species that are rare, endangered or invasive. To avoid that several authors work on the same species without knowing of each other, authors who intend to write a contribution should inform the editors of "Flora". Submitted contributions will be reviewed as usual to ensure scientific quality and compliance with the standard format.
IV.2. Format of a contribution to the "Biological Flora of Central Europe". Summary
2. Distribution and habitat requirements 2.1. Geographical distribution. Distribution and abundance in Central Europe and other areas (distribution maps for Germany/Central Europe/world). Altitudinal limits (minimum - maximum) of the species. Possible factors responsible for distribution limits. Historical data on distribution. Is the species indigenous or introduced (when?). For maps of global distribution write to Prof. E. J. JAGER Institut fi.ir Geobotanik, Neuwerk 21, D-06108 Halle. Recent maps of the distribution of a species in Germany are available from the "Zentralstelle fi.ir die Floristische Kartierung der Bundesrepublik Deutschland", Spezielle Botanik, Ruhr-Universitat Bochum, Postfach 102146, D-44780 Bochum. 2.2. Habitat. Habitat requirements (light, temperature, precipitation, soil moisture, sensitivity against frost, wind etc.). Exposition. Soil types or aquatic habitats in which the species is found growing, parent material, soil profile, hydrological data, form of humus, pH at different depths, CaC0 3 content, other chemical characteristics of the soil (e.g. C/N, potassium, phosphate, total nitrogen, ammonium, nitrate, salinity) or water. 2.3. Communities. Brief description of vegetation types in which the species occurs, list of typical accompanying species; if the species is a character species, name of syntaxon according to 0BERDORFER (1977-1992) or PoTT (1992).
A brief abstract should state that in the paper information on the biology of a species is given and identify fields (morphology, ecology, physiology etc.) which receive special emphasis.
2.4. Response to abiotic factors. Performance in various habitats: growth rates, productivity, production of flowers and fruits. Response to unusual weather conditions (effects of late frost, drought etc.).
I. Taxonomy and morphology
2.5. Abundance. Abundance of species in its habitats (dominant, frequent, rare).
1.1. Taxonomy (EHRENDORFER 1973, WISSKIRCHEN & HAEUPLER 1998), Name of species, if necessary, subspecies and important synonyms. Family to which species belongs. Common hybrids.
3. Life cycle and biology
1.2. Morphology. Concise descriptions of species morphology, including below-ground parts. Growth form; typical, minimal and maximal size. Variability in height, number of leaves, etc. Branching, existence of clonal growth. Structure of inflorescence. Number of shoot generations per year. Form and growth of below-ground structures, rooting depth and extension. Primary root system and/or adventitious roots. Microscopical and anatomical information (e.g. distribution and density of stomata). Other available morphological data, especially if those are not covered by conventional floras. 118
FLORA (2000) 195
3.1. Life cycle. Life form according to Raunkiaer (therophyte, hemikryptophyte, geophyte, chamaephyte, phanerophyte, helophyte, hydrophyte). Form and frequency of vegetative propagation. Longevity. Frequency of reproduction during the life-cycle, e.g. obligate annual, mostly annual, obligate biennial, monocarpic perennial, polycarpic perennial. Do plants appear every year above ground (orchids, gentians!)? For perennial species : Does the plant flower every year, or at what intervals? Relative importance of reproduction by seed and vegetative propagation. Patterns of mortality in the life cycle, mortality of seedlings and adults (for
example, present survivorship curves). Patterns of growth (e.g. leaf demography, leaf area index, relative growth rate, root turnover, root/shoot ratio, seasonal allocation patterns). 3.2. Spatial distribution of plants within populations. Does the species occur as solitary individuals or in patches? Analysis of spatial patterns and their causes. Typical densities, evidence for density-dependent mortality or reproduction. 3.3. Phenology. Development of shoots, leaves, main period of growth, deciduous, evergreen, time of flowering and fruiting. When are the diaspores dispersed? 3.4. Reproduction. Relation to age of plant. Distribution of sexes (is the species hermaphroditic, monoeceous, dioeceous, gynodioeceous ?). Is species proterandrous or proterogynous? Does apomixis, vivipary, cleistogamy occur? Is self pollination possible or common? Incompatibility system. Important pollinators and their behaviour on the plant. Number of pollen grains and ovules per flower. Seed I ovule ratio. Fruit set (% flowers developing into fruits), seed set (% ovules developing into seeds). Is there evidence for pollen limitation of reproduction? Typical values and range for number of fruits per plant, seeds per fruit, seeds per plant. Mean mass of diaspores (if possible, samples from different populations, air dried) with standard deviation and sample size. Size of seeds or diaspores. What are the diaspores (seeds, fruits, vegetative diaspores)? How are they dispersed (wind, water, ants, birds, other animals)? Special morphological or other adaptations to dispersal. Fall velocity, observed or experimentally obtained dispersal distances. 3.5. Germination. Type of dormancy (see HARPER 1957, BASKIN & BASKIN 1998) and conditions which break dormancy (e.g. scarification, stratification). Experimental data on the effects of environmental conditions on germination (e.g. temperature, temperature fluctuations, light, light quality, nitrate). It is important to state when seeds or diaspores were collected and how they were stored prior to the experiments. Description of germination. Influence of storage conditions on viability of seeds or diaspores. Time of germination in nature, rate of germination. Size of diaspore bank in different habitats (e.g. % of diaspores produced, number per m 2), survival of diaspores in the soil (e.g. time after which 50% of diaspores are dead). Special microhabitats that are required for germination and successful establishment of seedlings. Morphology of seedlings (drawings of several stages).
3.6. Response to competition and management. Competitive ability. Results of competition experiments. Response to management measures used in agriculture or forestry (logging, coppicing, mowing, grazing by livestock, ploughing, burning, herbicide treatment). Response to other human influences (trampling, air pollution, picking etc.). Possible management measures to control the species (weeds or invasive species) or increase population size and density (rare or threatened species). 3. 7. Herbivores and pathogens. Animals that feed on the species. Fungi or parasitic flowering plants for which the species is a host. Bacterial or viral diseases. Which parts of the plants are attacked or eaten? Variation among seasons or different habitats. Typical extent of damage, compensatory growth, importance for population dynamics. 3.8. Mycorrhiza. Presence or absence of mycorrhiza, type of mycorrhiza. 3.9. Physiological data. Rates of net photosynthesis under different conditions (light compensation point, optimal temperature), type of photosynthesis (C 3 , C 4 , CAM), water potential at wilting point, requirements for the initiation of flowers. 3.10. Biochemical data. Production of special chemical compounds, toxicity of plant parts. Importance for the ecology of the species (protection against animals, allelopathic effects on other plants). 3.11. Genetic data. Chromosome number (preferably from Central European specimens). Occurrence of polyploid plants. Distribution of genetic variation within and among populations. Heterozygosity, mean number of alleles per locus, total allelic diversity. Variability within species, common mutations. 3.12. Hybrids. If natural hybrids are known, is the fertility of hybrids lower than that of parents? Differences in ecology (lower I higher performance, habitat requirements), occurence of introgression. 3.13. Status of the species. Is the species spreading or declining? For threatened species: Causes of threat, protection, threat categories for various regions, number of individuals in populations. Reference to more detailed published studies. Importance of species as food or medicinal plant.
Acknowledgements We thank E. JAGER, R. RunsHAUSER and B. ScHMID for helpful comments on the manuscript. D. MATTHIES was supported by grants 5001-35231 and 5001-44626 of the Swiss National Science Foundation. FLORA (2000) 195
119
References ADLER, W., OSWALD, K. & FISCHER, R. (1994): Exkursionsflora von Osterreich. Stuttgart. BASKIN, C. C., & BASKIN, J. M. (1998): Seeds: Ecology, biogeography, and evolution of dormancy and germination. San Diego. BRITISH EcoLOGICAL SociETY (1941): Biological Flora of the British Isles. J. Ecol. 29: 356-357. CAVERS, P. B., & MULLIGAN, G. A. (1972): The biology of Canadian weeds. Canadian J. Plant. Sci. 52: 651-654. DI CASTRI, F., HANSEN, A. J., & DEBUSSCHE, M. (1990): Biological invasions in Europe and the Mediterranean basin. Dordrecht. EHRENDORFER, F. (Hrsg.) (1973): Liste der GefaBpflanzen Mitteleuropas. 2nd ed. Stuttgart. FiscHER, M. & MATTHIES, D. (1998). The effect of population size on performance in the rare plant Gentianella germanica J. Ecol. 86: 195-204. HEGI, G. (1908ff): Illustrierte Flora von Mitteleuropa. Miinchen. (1935ff): 2nd ed. Miinchen, Berlin. (198lff): 3rd ed. Berlin. HEss, H. E., LANDOLT, E., & HIRZEL, R. (1967): Flora der Schweiz. Birkhauser, Basel. JAGER, E. & MULLER-URI, C. (1981-1982): Wuchsform und Lebensgeschichte der GefaBpflanzen Zentraleuropas Bibliographie. Teil 1 : Quellenverzeichnis; Teil 2: Artenverzeichnis (4 Biinde). Halle, Universitiits- und Landesbibliothek Sachsen-Anhalt. KIRCHNER, 0. ( 1888): Flora von Stuttgart und Umgebung mit besonderer Beriicksichtigung der pflanzenbiologischen Verhiiltnisse. Stuttgart. - LoEW, E., & SCHROETER, C. (1908-1940): Lebensgeschichte der Bliitenpflanzen Mitteleuropas. Stuttgart. MATTHIES, D., SCHMID, B., & SCHMID-HEMPEL, P. (1995): The importance of population processes for the maintenance of biological diversity. Gaia 4: 199-209. MEUSEL, H., JAGER, E. & WEINERT, E. (1965-1992): Vergleichende Chorologie der zentral europiiischen Flora. (1965): Vol. 1. MEUSEL, H., JAGER, E. WEINERT, E. & RAUSCHERT, S. (1978): Vol. 2. MEUSEL, H. & JAGER, E. (1992): Vol. 3. Jena. 0BERDORFER, E. (ed.) (1977-1992): Siiddeutsche Pflanzengesellschaften. Part I- IV. Stuttgart. - ( 1994) : Pflanzensoziologische Exkursionsflora. 7th ed. Stuttgart. PosCHOLD, P., KIEFER, S., TRANKLE, U., FISCHER, S., & BONN, S. (1998): Plant species richness in calcareous grasslands as affected by dispersability in space and time. Appl. Veg. Sci. 1: 75-90. -, MATTHIES, D., JORDAN, S., & MENGEL, C. (1996): The biological flora of Central Europe - an ecological bibliography. Bull. Geobot. Inst. ETH 62: 89-108. RABOTNOV, T. A. (ed.) (1974ff): Biologiceskaja flora Moskovskoj oblasti (Biological flora of the Moscow area). (1974): Vol. 1. (1975): Vol. 2. (1976): Vol. 3. (1978): Vol.4. (1980): Vol. 5. (1980): Vol. 6. (1983): Vol. 7. PAVLOV, V. N., RABOTNOV, T. A. & TICHOMIROV, V. N. (1990): Vol. 8. PAVLOV, V. N. & TICHOMIROV, V. N. (eds.) 120
FLORA (2000) 195
(1993): Vol. 9. (1995): Vol. 10. (1995): Vol. 11. (1996): Vol. 12. (1997): Vol. 13. Mosvow. RoTHMALER, W. (1990-1999): Exkursionsflora von Deutschland. 4 Biinde. Edited by Schubert, R. et al. Jena. SCHMEIL-FITSCHEN ( 1993) : Flora von Deutschland und angrenzender Lander. 89th ed., edited by SENGHAS, K. & SEYBOLD, S. Heidelberg. SCHMID, B. & MATTHIES, D. (1994): Populationsbiologische Grundlagen des Artenschutzes. Naturwissenschaften 81: 283-292. SuKOPP, H. & KowARIK, I. (1986): Okologische Folgen der Einfiihrung neuer Pflanzenarten. Gentechnologie 10: 111-135. WHITE, J. (ed.) (1985): The population structure of vegetation. Dordrecht. WISSKIRCHEN, R. & HAEUPLER, H. (1998): Standardliste der Fam- und Bliitenpflanzen Deutschlands. Stuttgart.
Appendix. Selected works of reference on aspects of the biology of plants ADLER, W., OswALD, K. & FISCHER, R. (1994): Exkursionsflora von Osterreich. Stuttgart. ANDERBERG, A. L. (1994): Atlas of seeds and small fruits of Northwest-European plant species. Part 4: Resedaceae Umbelliferae. Stockholm. BARNES, H. F. (1946-1969): Gall midges of economic importance. 8 vol. London. BASKIN, C. C. & BASKIN, J. M. (1998): Seeds: Ecology, biogeography, and evolution of dormancy and germination. San Diego. BENKERT, D., MEUSEL, H., FUKAREK, F. & KORSCH, H. (eds.) (1996): Verbreitungsatlas der Fam- und Bliitenpflanzen Ostdeutschlands. Jena. BERGGREN, G. (1969): Atlas of seeds and small fruits of Northwest-European plant species. Part 2: Cyperaceae. Stockholm. - (1981): Atlas of seeds and small fruits of Northwest-European plant species. Part 3: Salicaceae- Cruciferae. Stockholm. Bergmann, A. ( 1954): Die GroBschmetterlinge Mitteldeutschlands. Jena. BoRNER, C. ( 1952): Europae centralis Aphides. Die Blattlause Mitteleuropas. Mitt. Thiir. Bot. Ges. Beih. 3. BONN, S. & POSCHLOD, P. (1998): Ausbreitungsbiologie der Pflanzen Mitteleuropas. Wiesbaden. BRANDENBURGER, W. (1963): Vademecum zum Sammeln parasitischer Pilze. Stuttgart. BRENCHLEY, W. E. & WARNIGTON, K. (1930): The weed seed population of arable soil. 1. Numerical estimation of viable seeds and observations on their natural dormancy. J. Ecol. 18: 235-272. BuHR, H. (1964-1965): Bestimmungstabellen der Gallen (Zoo- und Phytocecidien) an Pflanzen Mittel- und Nordeuropas. 2 vol. Jena. BURGESS, N. A., CHATER, A. 0. & EDMONDSON, J. R. (1993): Flora Europaea. 2nd ed. Cambridge. CARTER, D. J. (1987): Raupen und Schmetterlinge Europas und ihre Futterpflanzen. Hamburg.
CSAPODY, V. (1968): Keimlingsbestimmungsbuch der Dikotyledonen. Budapest. DIECKMANN, L. ( 1972): Beitrage zur Insektenfauna der DDR. Coleoptera- Curculionidae: Ceutorhynchinae. Beitr. Ent. 22: 3-128. - (1972): Beitrage zur Insektenfauna der DDR. ColeopteraCurculionidae: Ceutorhynchinae. Beitr. Ent. 22: 3-128. - (1983): Beitrage zur Insektenfauna der DDR. ColeopteraCurculionidae : Tanymecinae, Leptopiinae, Cleoninae, Tanyrhynchinae, Cossoninae, Raymondionyrninae, Bogoinae, Tanysphyrinae. Beitr. Ent. 33: 257-381. DuLL, R. & KuTZELNIGG, H. (1994): Botanisch-okologisches Exkursionstaschenbuch. 5th ed. Heidelberg. ELLENBERG, H. (1996): Vegetation Mitteleuropas mit den Alpen. 5th ed. Stuttgart. -, WEBER, H. E., DuLL, R., WIRTH, v., WERNER, w. & PAuLissEN, D. (1991): Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica 18. ELLIS, M. B. & ELLIS, J.P. (1997): Microfungi on land plants. Slough. FAEGRI; K. & VANDER PIJL, L. (1979). The principles of pollination ecology. 3rd ed. Oxford. FITTER, A. H. (1978): An atlas of the wild flowers of Britain and Northern Europe. London. - & PEAT, H. J. (1994): The ecological flora database. J. Ecol. 82:415-425. FORSTER, W. & WOHLFAHRT, T. A. (1971): Die Schmetterlinge Mitteleuropas. Vol. IV. Bulen (Noctuidae). Stuttgart. FREIDENFELT, T. (1902): Studien tiber die Wurzeln krautiger Pflanzen. Flora 91: 115-208. FRANK, D. & KloTz, S. (1990): Biologisch-okologische Daten zur Flora der DDR. 2nd ed., Wiss. Beitr. Martin Luther-Univers. Halle-Wittenb. 32. FRYXELL, P. A. (1957): Mode of reproduction of higher plants. Bot. Rev. 23: 135-233. GRIME, J.P., HoDGSON, J. G. & HUNT, R. (1988): Comparative plant ecology - A functional approach to common British species. London. -, MASON, G., CuRTIS, A. V., RoMAN, J., BAND, S. R., MowFORTH, M.A. G., NEAL, A.M. & SHAW, S.C. (1981): A comparative study of germination characteristics in a local flora. J. Ecol. 69: 1017-1059. HAEUPLER, H. &. ScHONFELDER, P. (1989): Atlas der Famund Bliitenpflanzen der Bundesrepublik Deutschland. 2nd ed. Stuttgart. HAMANN, U. & WAGENITZ, G. (1977): Bibliographie zur Flora von Mitteleuropa. 2nd ed. Berlin. HANF, M. (1990): Ackerunkrauter Europas mit ihren Keimlingen und Samen. 3rd ed. Miinchen. Hannemann, H. J. (1961): Kleinschmetterlinge oder Mieralepidoptera I. Die Wickler (s.str.) (Tortricidae). In: DAHL, F. (ed.): Die Tierwelt Deutschlands und der angrenzenden Meeresteile. Jena. - (1964): Kleinschmetterlinge oder Micro Iepidoptera II. Die Wickler (s.l.) (Cochylidae und Carposinidae), die Ziinslerartige (Pyraloidea). In: DAHL, F. (ed.): Die Tierwelt Deutschlands und der angrenzenden Meeresteile. Jena. HARLEY, J. L. & HARLEY, E. L. (1987): A check-list of mycorrhiza in the British Flora. New Phytol. Suppl. lOS: 1-102.
HARPER, J. L. (1957): The ecological significance of dormancy and its importance in weed control. Proceedings of the 4th international congress on crop protection, Hamburg. pp. 415-420. HARTMANN, E., SCHULDES, H., KUBLER, R. & KONOLD, W. (1995): Neophyten. Biologie, Verbreitung und Kontrolle ausgewahlter Arten. Stuttgart. HEATH, J. et al. (1983ff): The moths and butterflies of Great Britain and Ireland. Vol. 1-10. Colchester. HEm, G. (1908ff): Illustrierte Flora von Mittel europa. Miinchen. (1935ff): 2nd ed., Miinchen, Berlin. (198lff): 3rd ed., Berlin. HEGNAUER, R. (1990): Chemotaxonomie der Pflanzen. Berlin. HEINTZE, A. (1932): Handbuch der Verbreitungsokologie der Pflanzen. Stockholm. HEJNY, S. & SLAVIK, B. (eds.) (1988-1997): Kvetena geske Socialisticke Republiky. HEJNY, S. & SLAVIK, B. (eds.) (1988): Vol. 1. HEJNY, S. & SLAVIK, B. (eds.) (1990): Vol. 2. SLAVIK, B. (ed.) (1992): Vol. 3. SLAVIK, B. (ed.) (1995): Vol. 4. SLAVIK, B. (ed.) (1997): Vol. 5. Praha. HENDEL, F. ( 1938): Agromyzidae. Vol. VI, 2 of Lindner, E. : Die Fliegen der palaearktischen Region. Stuttgart. HERING, M. (1935): Die Blattrninen Mittel- und Nord-Europas. Neubrandenburg. HESS, D. (1990): Die Bliite. 2nd ed. Stuttgart. HEss, H. E., LANDOLT, E. & HIRZEL, R. (1976): Flora der Schweiz und angrenzender Gebiete. 2nd ed. Basel. HODGSON, J. D., GRIME, J. P., HUNT, R. & THOMPSON, K. (1995): The electronic comparative plant ecology. London. HoLM, L., DoLL, J., HoLM, E. & PANCHO, J. (1997): World weeds. New York. HONARD, C. (1908-1913): Les Zoocecidies des Plantes d'Europe et du Bassin de Ia Mediterranee. (1908): Tome I, (1909): Tome II, (1913): Tome III. Paris. HuLTEN, E. (1971): Atlas of the distribution of vascular plants in North Western Europe. Stockholm. - & FRIES, M. (1986): Atlas of North European vascular plants north of the tropic of cancer II. Konigsstein. HuNDT, R. (1966): Okologisch-geobotanische Untersuchungen an Pflanzen der Mitteleuropaischen Wiesenvegetation. Jena. JAGER, E. & MuLLER-URI, C. (1981-1982): Wuchsform und Lebensgeschichte der (GefliBpflanzen Zentraleuropas Bibliographie. Part 1 : Quellenverzeichnis; Part 2: Artenverzeichnis (4 vol.). Halle, Universitats- und Landesbibliothek Sachsen-Anhalt. }ALAS, J. & SUOMNINEN, J. (1972ff): Atlas Florae Europaeae Vol 1-10. Helsinki. KALTENBACH, J. H. (1874): Pflanzenfeinde aus der Klasse der Insekten. Stuttgart. KIRCHNER, 0., LOEW, E. & SCHROETER, C. (1908ff): Lebensgeschichte der Bliitenpflanzen Mitteleuropas. Stuttgart. KNUTH P. (1889): Handbuch der Bliitenbiologie II. Leipzig. KORNECK, D., SCHNITTLER, M., & VOLLMER, I. (1996): Rote Liste der Fam- und Bliitenpflanzen (Pteridophyta et Spermatophyta) Deutschlands. Schr. R. Vegetationskunde 28: 21-187. KuGLER, H. (1970): Bliitenokologie. 2nd ed., Stuttgart. FLORA (2000) 195
121
KuTSCHERA, L. (1960): Wurzelatlas mitteleuropaischer Ackerunkrauter und Kulturpflanzen. Frankfurt. KUTSCHERA, L. & LICHTENEGGER, E. (1982-1992): Wurzelatlas mitteleuropaischer Gri.inlandpflanzen. Stuttgart. - -, SoBOTIK, M. & HAAS, D. (1997): Bewurzelung von Pflanzen in verschiedenen Lebensraumen. Stapfia 49: 1-331. LANDOLT, E. (1977): Okologische Zeigerwerte zur Schweizer Flora. Veroff. Geobot. lust. ETH, Stift. Riibel 64. LECK, M.A., PARKER, V. T. & SIMPSON, R. L. (eds) (1989): Ecology of soil seed banks. London. LovE, A. & LovE, D. (1961): Chromosome numbers of central and northwest European plant species. Opera Bot. Soc. Lund 5: 1-581. MEUSEL, H., JAGER, E. & WEINERT, E. (1965-1992): Vergleichende Chorologie der zentraleuropaischen Flora. (1965): Vol. 1. MEUSEL, H., JAGER, E. WEINERT, E. & RAUSCHERT, S. (1978): Vol. 2. MEUSEL, H. & JAGER, E. (1992): Vol. 3. Jena. MoHR, K. H. (1966): Chrysomelidae. In: FREUDE, H., HARDE, K. W. & LoHSE, G. A. (eds.): Die Kafer Mitteleuropas, Vol. 9. Krefeld. MooRE, D. M. (1982): Flora Europaea check-list and chromosome index. Cambridge. MULLER, H. (1873): Die Befruchtung der Blumen durch Insekten und gegenseitige Anpassungen beider. Leipzig. - ( 1881): Alpenblumen, ihre Befruchtung durch Insekten und ihre Anpassung an dieselben. Leipzig. MuLLER-SCHNEIDER, P. (1977): Verbreitungsbiologie (Diasporologie) der Bliitenpflanzen. Veroff. Geobot. lust. ETH, Stift. Riibel61: 1-226. - (1986): Verbreitungsbiologie der Bliitenpflanzen Graubiindens. Veroff Geobot. lust. ETH, Stift. Riibel85: 1-263. MuLLER, F. M. (1978): Seedlings of the North-Western European lowland. A flora of seedlings. The Hague, Boston. OBERDORFER, E. (1994): Pflanzensoziologische Exkursionsflora. 7th ed. Stuttgart. - (ed.) (1977-1992): Siiddeutsche Pflanzengesellschaften. Part I-IV. Stuttgart. PERRING, F. H. & WALTERS, S.M. (eds.) (1976): Atlas of the British flora. Wakefield. PERTTULA, U. (1941): Untersuchungen tiber die generative und vegetative Vermehrung der Bliitenpflanzen in der Wald-, Hainwiesen- und Hainfelsenvegetation. Ann. Acad. Scient. Fenn. Ser. A Tom. 58, No. 1. PETERSEN, A. (1988): Die Graser als Kulturpflanzen und Unkrauter auf Wiese, Weide und Acker. Berlin. PIRSON, A., ZIMMERMANN, M. H. (eds.) (1975ff.): Encyclopaedia of plant physiology; new series. Vol. 1-20. Berlin. PoTT, R. (1992): Die Pflanzengesellschaften Deutschlands. 2nd ed. Stuttgart. RABOTNOV, T. A. (ed.) (1974ff): Biologiceskaja flora Moskovskoj oblasti (Biological flora of the Moscow area). (1974): Vol. 1. (1975): Vol. 2. (1976): Vol. 3. (1978): Vol.
122
FLORA (2000) 195
4. (1980): Vol. 5. (1980): Vol. 6. (1983): Vol. 7. PAVLOV, V. N., RABOTNOV, T. A. & TICHOMIROV, V. N. (eds.) (1990): Vol. 8. PAVLOV, V. N. & TICHOMIROV, V. N. (eds.) (1993): Vol. 9. (1995): Vol. 10. (1995): Vol. 11. (1996): Vol. 12. (1997): Vol. 13. Moscow. RAUNKIAER, C. (1895-1899): De danske Blomsterplanters N aturhistorie. Kopenhagen. ROBERTS, H. A. (1981): Seed banks in soils. In: CoACKER, T. H. Advances in applied biology 6. London, 1-55. RoTHMALER, W. ( 1990-1999): Exkursionsflora von Deutschland. BASSLER, M., JAGER, E. J. & WERNER, K. (eds.) (1999): Vol. 2, 3rd ed. JAGER, E. J. & WERNER, K. (eds.) (1999): Vol. 3. ScHUBERT, R., VENT, W. & BASSLER, M. (eds.) (1990): Vol. 4. SALISBURY, E. J. (1964): Weeds and aliens. 2nd ed. London. - (1942): The reproductive capacity of plants. London. ScHNEIDER, C., SuKOPP, U. & SuKOPP, H. (1994): Biologisch-okologische Grundlagen des Schutzes geflihrdeter Segetalpflanzen. Schr. R. Vegetationskunde 26: 1-356. SCHOENICHEN, W. (1940): Biologie der geschiitzten Pflanzen Deutschlands. ScHROETER, C. (1926): Das Pflanzenleben der Alpen. 2nd ed. Ziirich. SCHUTZE, K. T. ( 1931): Die Biologie der Kleinschmetterlinge unter besonderer Beri.icksichtigung ihrer Nahrpflanzen und Erscheinungszeiten. Frankfurt. SEBALD, 0., SEYBOLD, S. & PHILIPPI, G. (1990ff): Die Famund Bliitenpflanzen Baden Wiirttembergs. Stuttgart. SERNANDER, R. (1906): Entwurf einer Monographie der europaischen Myrmekochoren. K. Sv. Vet.-Akad. Hand. 41: 1-410. SPENCER, K. A. (1990): Host specialization in the world Agromyzidae (Diptera). Dordrecht. STEWARD, A., PEARMAN, D. A. & PRESTON, C. D. (1994): Scarce plants in Britain. Peterborough. STOKOE, W. J. (1948): The caterpillars of British moths including the eggs, chrysalids and food-plants. London. THOMPSON, K., BAKKER, J.P. & BEKKER, R. M. (1997): The soil seed hanks of North West Europe: methodology, density and longevity. Cambridge. TUTIN, T. G., HEYWOOD, V. H., BURGESS, N. A., VALENTINE, D. H., WALTERS, S.M. & WEBB, D. A. (1964): Flora Europaea. Cambridge. VAN DER PIJL, L. (1982): Principles of dispersal in higher plants. 3rd ed. Berlin. VOGEL, E. F. de (1980): Seedlings of dicotyledons. Wageningen. WEEDA, E. J, WESTRA, R., WESTRA, c. & WESTRA, T. (1985-1991): Nederlandse oecologische Flora: wilde planten en hun relaties. 5 volumes. Hilversum. WELTEN, M. & SuTTER, R. (1982): Verbreitungsatlas der Fam- und Bliitenpflanzen der Schweiz. Basel. WISSKIRCHEN, R. & HAEUPLER, H. (1998): Standardliste der Fam- und Bliitenpflanzen Deutschlands. Stuttgart.
The biological flora of Central Europe - aims and concept DIETHART MATTHIES 1 and PETER POSCHLOD2 1
2
Institut fiir Umweltwissenschaften, Universitat Ziirich, Winterthurerstr. 190, CH-8057 ZUrich, Schweiz. Current address: Fachgebiet Pflanzenokologie, Fachbereich Biologie, Philipps-Universitat, D-35032 Marburg. Tel.: +49-6421-2822085, Fax: +49-6421-2822057 E-mail : matthies@ mailer. uni-marburg.de Fachbereich Biologie, Fachgebiet Naturschutz (II), Philipps-Universitat, D-35032 Marburg
Accepted: April30, 1999
Summary Information on the biology of individual species is important in many fields, but standard floras contain very little information on the ecology and life-history of plants. We call for collaboration in a new series: "The biological flora of Central Europe". Contributions to the series should summarize the available knowledge of the biology of a plant occurring in Central Europe. The concept of the "Biological Flora of Central Europe" and a standard format for contributions is presented. Key words: Flora, ecology, species biology.
I. Introduction Information on the biology of plant species is very important in many fields. Understanding the life-cycle of a species is a prerequisite for understanding its population dynamics. Population processes like the decline of threatened plant species (SCHMID & MATTHIES 1994, FISCHER & MATTHIES 1998) or the rapid expansion of the range and abundance of non-native plants (SUKOPP & KOWARIK 1986, DI CASTRI et al. 1990) cannot be understood if only abiotic factors are analysed (MATTHIES et al. 1995, POSCHOLD et al. 1998). Data on the population biology of a species, its dispersal capabilities, the persistence of the seed bank, its breeding system and pollinators make it possible to predict more accurately the response of a plant population to human interventions and to suggest management measures. Similarly, the management of weeds in agricultural systems or of agressive exotic species in nature reserves requires a comprehensive knowledge of the biology of the respective species. Because of the population structure of vegetation, an understanding of the population biology of species is also frequently necessary to understand the dynamics of vegetation (WHITE 1985). Moreover, many problems encountered in botanical research (unsatisfactory germination, weak growth, uncertainty about the time of first flowering) 116
FLORA (2000) 195
could be avoided if autecological and population-biological data were more easily available. The Central European standard floras contain very little information on the biology of species (e.g. HEss et al. 1967, 0BERDORFER 1994, ROTHMALER 1990-99, SCHMEIL-FITSCHEN 1993, ADLER et al. 1994; for an assessment ofHEGI 1908ff, 1935ff, 1981 ff, see below). Anyone interested in a certain species has to collect the required data from many different sources, which is very laborious and time-consuming. Moreover, there are frequently large gaps in our knowledge of individual species. Many sources that are particularly valuable (e.g. Diploma theses) have not been published and are thus not generally accessible. To reduce the obvious deficits in the information available on the biology of plants, we call for collaboration in a new series : "The Biological Flora of Central Europe". This project originated within the framework of the "Working Group on Plant Population Biology" of the German Ecological Society. The "Biological Flora" is planned as a series of papers in the journal "Flora" that summarize the available information on individual species. In the following we will first give a brief summary of earlier attempts to compile the available information on the biology of plants growing in Central Europe, and discuss the relationship between "The Biological Flora 0367-2530/001195/02-116
$ 12.00/0
of Central Europe" and other biological floras. We then present the concept of the "Biological Flora of Central Europe" and a standard format for contributions. Selected works of reference which contain useful information on the biology of plant species are given in an appendix.
II. Other biological floras One of the oldest floras containing information on the biology of some species is the "Flora von Stuttgart" (KIRCHNER 1888). In particular the reproductive biology of plants is well covered. Later, this flora formed the core of the ambitious "Lebensgeschichte der Bltitenpflanzen Mitteleuropas" (KIRCHNER et al. 1908 to 1940), which, however, remained unfinished. A very useful series that is still continued is the "Biological Flora of the British Isles", which is published in the "Journal of Ecology" (BRITISH EcoLOGICAL SociETY 1941). This British series is one of the best sources of information on individual plant species and in many respects the model for the "Biological Flora of Central Europe". More recent series include the "Biological Flora of the Moscow Region" (RABOTNOV 1974ff.), and the "Biology of Canadian Weeds" (CAVERS & MULLIGAN 1972). A detailed description of the various biological floras (including electronic data bases), their history, aims and concepts, can be found elsewhere (PoscHLOD et al. 1996). A lot of biological information can also be found in the volumes of the second and third editions of the "Illustrierte Flora von Mitteleuropa" (HEGI 1908ff, 1935ff, 1981ff). However, the main emphasis of this work is still on taxonomy and morphology, and important biological data (e.g. breeding system, seed mass, germination requirements, longevity of seeds in soil) are frequently still missing in recent volumes. Moreover, for several families only the first edition from the beginning of the century is available, which contains hardly any information on the biology of species. Because the "Illustrierte Flora von Mitteleuropa" is published as a series of books, new volumes can only be added once a large number of taxonomically related species has been revised. This strongly delays the publication of new information. In contrast, the "Biological Flora of Central Europe" is planned as a series of separate papers. This makes it possible to publish contributions on individual species as they become available. We especially hope to encourage botanists to write contributions, who have intensively studied individual species (e.g. for their diploma or PhD-thesis), and know them well. According to our experience there exists a lot of knowledge on the biology of plants that is currently inaccessible.
III. Relationship of the "Biological Flora of Central Europe" to similar projects elsewhere The model for the project "Biological Flora of Central Europe" is the "Biological Flora of the British Isles", whose structure we have largely followed. The "Biological Flora of Central Europe" is intended to be complementary to the British series. Many Central European plant species do not occur in the British Isles and the British series has made only slow progress. We think that Central European botanists, who have perhaps shied away from writing a contribution to the British series, can make important contributions to a better knowledge of the European flora. Contributions to the "Biological Flora" should be written in English. The question of the language has been discussed by the members of the "Working group on Plant Population Biology" and a clear majority was in favour of English. We are aware that this may present a hurdle for some who would like to contribute to the series, but we hope that potential problems can be overcome.
IV. Procedure of compiling contributions to the "Biological Flora of Central Europe"
IV.l. Basis The basis of a contribution to the "Biological Flora" is a thorough analysis of available information in the literature. In appendix 1 of this paper a list of standard works of reference is given, which are useful sources of information on certain aspects of the biology of plants. A lot of the older literature on the biology of central European plants is listed in JAGER & MOLLER-URI (1982). A literature search in the "Biological Abstracts" is strongly recommended. In case of species that are important weeds the "Weed Abstracts" often contain additional information. The analysis of the literature should be complemented by specific studies to fill important gaps in the knowledge of a species. One of the aims of the "Biological Flora" is indeed to encourage studies on the biology of plants. In order to make it possible to assess the reliability of information, the sources of all information should be given in form of references. This is also useful for further studies of the literature. Data for which no reference is given are assumed to be based on observations by the author. Contributions on all species that occur in Central Europe (including the Alps) are welcome (see EHRENDORFER 1973, WISSKIRCHEN & HAEUPLER 1998). All contributions should conform to a standard format (see FLORA (2000) 195
117
below). If no information is available on a major point, this should be stated explicitly. The standard format provides a framework for contributions, but leaves the authors a lot of freedom. We have deliberately avoided to give very detailed and inflexible instructions, because otherwise potential contributors might be deterred if no information is available on many points. It is also possible for individual contributions to have a different emphasis. In general, species for which an account in the "Biological Flora of the British Isles", or the "Biology of Canadian Weeds" is available, should have a low priority. However, some of the early contributions to the British series are very brief, and those species would certainly merit a new account. A list of Central European species, for which accounts exist in one or more biological floras is given in PoscHLOD et al. (1996). Especially welcome are accounts on species that are rare, endangered or invasive. To avoid that several authors work on the same species without knowing of each other, authors who intend to write a contribution should inform the editors of "Flora". Submitted contributions will be reviewed as usual to ensure scientific quality and compliance with the standard format.
IV.2. Format of a contribution to the "Biological Flora of Central Europe". Summary
2. Distribution and habitat requirements 2.1. Geographical distribution. Distribution and abundance in Central Europe and other areas (distribution maps for Germany/Central Europe/world). Altitudinal limits (minimum - maximum) of the species. Possible factors responsible for distribution limits. Historical data on distribution. Is the species indigenous or introduced (when?). For maps of global distribution write to Prof. E. J. JAGER Institut fi.ir Geobotanik, Neuwerk 21, D-06108 Halle. Recent maps of the distribution of a species in Germany are available from the "Zentralstelle fi.ir die Floristische Kartierung der Bundesrepublik Deutschland", Spezielle Botanik, Ruhr-Universitat Bochum, Postfach 102146, D-44780 Bochum. 2.2. Habitat. Habitat requirements (light, temperature, precipitation, soil moisture, sensitivity against frost, wind etc.). Exposition. Soil types or aquatic habitats in which the species is found growing, parent material, soil profile, hydrological data, form of humus, pH at different depths, CaC0 3 content, other chemical characteristics of the soil (e.g. C/N, potassium, phosphate, total nitrogen, ammonium, nitrate, salinity) or water. 2.3. Communities. Brief description of vegetation types in which the species occurs, list of typical accompanying species; if the species is a character species, name of syntaxon according to 0BERDORFER (1977-1992) or PoTT (1992).
A brief abstract should state that in the paper information on the biology of a species is given and identify fields (morphology, ecology, physiology etc.) which receive special emphasis.
2.4. Response to abiotic factors. Performance in various habitats: growth rates, productivity, production of flowers and fruits. Response to unusual weather conditions (effects of late frost, drought etc.).
I. Taxonomy and morphology
2.5. Abundance. Abundance of species in its habitats (dominant, frequent, rare).
1.1. Taxonomy (EHRENDORFER 1973, WISSKIRCHEN & HAEUPLER 1998), Name of species, if necessary, subspecies and important synonyms. Family to which species belongs. Common hybrids.
3. Life cycle and biology
1.2. Morphology. Concise descriptions of species morphology, including below-ground parts. Growth form; typical, minimal and maximal size. Variability in height, number of leaves, etc. Branching, existence of clonal growth. Structure of inflorescence. Number of shoot generations per year. Form and growth of below-ground structures, rooting depth and extension. Primary root system and/or adventitious roots. Microscopical and anatomical information (e.g. distribution and density of stomata). Other available morphological data, especially if those are not covered by conventional floras. 118
FLORA (2000) 195
3.1. Life cycle. Life form according to Raunkiaer (therophyte, hemikryptophyte, geophyte, chamaephyte, phanerophyte, helophyte, hydrophyte). Form and frequency of vegetative propagation. Longevity. Frequency of reproduction during the life-cycle, e.g. obligate annual, mostly annual, obligate biennial, monocarpic perennial, polycarpic perennial. Do plants appear every year above ground (orchids, gentians!)? For perennial species : Does the plant flower every year, or at what intervals? Relative importance of reproduction by seed and vegetative propagation. Patterns of mortality in the life cycle, mortality of seedlings and adults (for
example, present survivorship curves). Patterns of growth (e.g. leaf demography, leaf area index, relative growth rate, root turnover, root/shoot ratio, seasonal allocation patterns). 3.2. Spatial distribution of plants within populations. Does the species occur as solitary individuals or in patches? Analysis of spatial patterns and their causes. Typical densities, evidence for density-dependent mortality or reproduction. 3.3. Phenology. Development of shoots, leaves, main period of growth, deciduous, evergreen, time of flowering and fruiting. When are the diaspores dispersed? 3.4. Reproduction. Relation to age of plant. Distribution of sexes (is the species hermaphroditic, monoeceous, dioeceous, gynodioeceous ?). Is species proterandrous or proterogynous? Does apomixis, vivipary, cleistogamy occur? Is self pollination possible or common? Incompatibility system. Important pollinators and their behaviour on the plant. Number of pollen grains and ovules per flower. Seed I ovule ratio. Fruit set (% flowers developing into fruits), seed set (% ovules developing into seeds). Is there evidence for pollen limitation of reproduction? Typical values and range for number of fruits per plant, seeds per fruit, seeds per plant. Mean mass of diaspores (if possible, samples from different populations, air dried) with standard deviation and sample size. Size of seeds or diaspores. What are the diaspores (seeds, fruits, vegetative diaspores)? How are they dispersed (wind, water, ants, birds, other animals)? Special morphological or other adaptations to dispersal. Fall velocity, observed or experimentally obtained dispersal distances. 3.5. Germination. Type of dormancy (see HARPER 1957, BASKIN & BASKIN 1998) and conditions which break dormancy (e.g. scarification, stratification). Experimental data on the effects of environmental conditions on germination (e.g. temperature, temperature fluctuations, light, light quality, nitrate). It is important to state when seeds or diaspores were collected and how they were stored prior to the experiments. Description of germination. Influence of storage conditions on viability of seeds or diaspores. Time of germination in nature, rate of germination. Size of diaspore bank in different habitats (e.g. % of diaspores produced, number per m 2), survival of diaspores in the soil (e.g. time after which 50% of diaspores are dead). Special microhabitats that are required for germination and successful establishment of seedlings. Morphology of seedlings (drawings of several stages).
3.6. Response to competition and management. Competitive ability. Results of competition experiments. Response to management measures used in agriculture or forestry (logging, coppicing, mowing, grazing by livestock, ploughing, burning, herbicide treatment). Response to other human influences (trampling, air pollution, picking etc.). Possible management measures to control the species (weeds or invasive species) or increase population size and density (rare or threatened species). 3. 7. Herbivores and pathogens. Animals that feed on the species. Fungi or parasitic flowering plants for which the species is a host. Bacterial or viral diseases. Which parts of the plants are attacked or eaten? Variation among seasons or different habitats. Typical extent of damage, compensatory growth, importance for population dynamics. 3.8. Mycorrhiza. Presence or absence of mycorrhiza, type of mycorrhiza. 3.9. Physiological data. Rates of net photosynthesis under different conditions (light compensation point, optimal temperature), type of photosynthesis (C 3 , C 4 , CAM), water potential at wilting point, requirements for the initiation of flowers. 3.10. Biochemical data. Production of special chemical compounds, toxicity of plant parts. Importance for the ecology of the species (protection against animals, allelopathic effects on other plants). 3.11. Genetic data. Chromosome number (preferably from Central European specimens). Occurrence of polyploid plants. Distribution of genetic variation within and among populations. Heterozygosity, mean number of alleles per locus, total allelic diversity. Variability within species, common mutations. 3.12. Hybrids. If natural hybrids are known, is the fertility of hybrids lower than that of parents? Differences in ecology (lower I higher performance, habitat requirements), occurence of introgression. 3.13. Status of the species. Is the species spreading or declining? For threatened species: Causes of threat, protection, threat categories for various regions, number of individuals in populations. Reference to more detailed published studies. Importance of species as food or medicinal plant.
Acknowledgements We thank E. JAGER, R. RunsHAUSER and B. ScHMID for helpful comments on the manuscript. D. MATTHIES was supported by grants 5001-35231 and 5001-44626 of the Swiss National Science Foundation. FLORA (2000) 195
119
References ADLER, W., OSWALD, K. & FISCHER, R. (1994): Exkursionsflora von Osterreich. Stuttgart. BASKIN, C. C., & BASKIN, J. M. (1998): Seeds: Ecology, biogeography, and evolution of dormancy and germination. San Diego. BRITISH EcoLOGICAL SociETY (1941): Biological Flora of the British Isles. J. Ecol. 29: 356-357. CAVERS, P. B., & MULLIGAN, G. A. (1972): The biology of Canadian weeds. Canadian J. Plant. Sci. 52: 651-654. DI CASTRI, F., HANSEN, A. J., & DEBUSSCHE, M. (1990): Biological invasions in Europe and the Mediterranean basin. Dordrecht. EHRENDORFER, F. (Hrsg.) (1973): Liste der GefaBpflanzen Mitteleuropas. 2nd ed. Stuttgart. FiscHER, M. & MATTHIES, D. (1998). The effect of population size on performance in the rare plant Gentianella germanica J. Ecol. 86: 195-204. HEGI, G. (1908ff): Illustrierte Flora von Mitteleuropa. Miinchen. (1935ff): 2nd ed. Miinchen, Berlin. (198lff): 3rd ed. Berlin. HEss, H. E., LANDOLT, E., & HIRZEL, R. (1967): Flora der Schweiz. Birkhauser, Basel. JAGER, E. & MULLER-URI, C. (1981-1982): Wuchsform und Lebensgeschichte der GefaBpflanzen Zentraleuropas Bibliographie. Teil 1 : Quellenverzeichnis; Teil 2: Artenverzeichnis (4 Biinde). Halle, Universitiits- und Landesbibliothek Sachsen-Anhalt. KIRCHNER, 0. ( 1888): Flora von Stuttgart und Umgebung mit besonderer Beriicksichtigung der pflanzenbiologischen Verhiiltnisse. Stuttgart. - LoEW, E., & SCHROETER, C. (1908-1940): Lebensgeschichte der Bliitenpflanzen Mitteleuropas. Stuttgart. MATTHIES, D., SCHMID, B., & SCHMID-HEMPEL, P. (1995): The importance of population processes for the maintenance of biological diversity. Gaia 4: 199-209. MEUSEL, H., JAGER, E. & WEINERT, E. (1965-1992): Vergleichende Chorologie der zentral europiiischen Flora. (1965): Vol. 1. MEUSEL, H., JAGER, E. WEINERT, E. & RAUSCHERT, S. (1978): Vol. 2. MEUSEL, H. & JAGER, E. (1992): Vol. 3. Jena. 0BERDORFER, E. (ed.) (1977-1992): Siiddeutsche Pflanzengesellschaften. Part I- IV. Stuttgart. - ( 1994) : Pflanzensoziologische Exkursionsflora. 7th ed. Stuttgart. PosCHOLD, P., KIEFER, S., TRANKLE, U., FISCHER, S., & BONN, S. (1998): Plant species richness in calcareous grasslands as affected by dispersability in space and time. Appl. Veg. Sci. 1: 75-90. -, MATTHIES, D., JORDAN, S., & MENGEL, C. (1996): The biological flora of Central Europe - an ecological bibliography. Bull. Geobot. Inst. ETH 62: 89-108. RABOTNOV, T. A. (ed.) (1974ff): Biologiceskaja flora Moskovskoj oblasti (Biological flora of the Moscow area). (1974): Vol. 1. (1975): Vol. 2. (1976): Vol. 3. (1978): Vol.4. (1980): Vol. 5. (1980): Vol. 6. (1983): Vol. 7. PAVLOV, V. N., RABOTNOV, T. A. & TICHOMIROV, V. N. (1990): Vol. 8. PAVLOV, V. N. & TICHOMIROV, V. N. (eds.) 120
FLORA (2000) 195
(1993): Vol. 9. (1995): Vol. 10. (1995): Vol. 11. (1996): Vol. 12. (1997): Vol. 13. Mosvow. RoTHMALER, W. (1990-1999): Exkursionsflora von Deutschland. 4 Biinde. Edited by Schubert, R. et al. Jena. SCHMEIL-FITSCHEN ( 1993) : Flora von Deutschland und angrenzender Lander. 89th ed., edited by SENGHAS, K. & SEYBOLD, S. Heidelberg. SCHMID, B. & MATTHIES, D. (1994): Populationsbiologische Grundlagen des Artenschutzes. Naturwissenschaften 81: 283-292. SuKOPP, H. & KowARIK, I. (1986): Okologische Folgen der Einfiihrung neuer Pflanzenarten. Gentechnologie 10: 111-135. WHITE, J. (ed.) (1985): The population structure of vegetation. Dordrecht. WISSKIRCHEN, R. & HAEUPLER, H. (1998): Standardliste der Fam- und Bliitenpflanzen Deutschlands. Stuttgart.
Appendix. Selected works of reference on aspects of the biology of plants ADLER, W., OswALD, K. & FISCHER, R. (1994): Exkursionsflora von Osterreich. Stuttgart. ANDERBERG, A. L. (1994): Atlas of seeds and small fruits of Northwest-European plant species. Part 4: Resedaceae Umbelliferae. Stockholm. BARNES, H. F. (1946-1969): Gall midges of economic importance. 8 vol. London. BASKIN, C. C. & BASKIN, J. M. (1998): Seeds: Ecology, biogeography, and evolution of dormancy and germination. San Diego. BENKERT, D., MEUSEL, H., FUKAREK, F. & KORSCH, H. (eds.) (1996): Verbreitungsatlas der Fam- und Bliitenpflanzen Ostdeutschlands. Jena. BERGGREN, G. (1969): Atlas of seeds and small fruits of Northwest-European plant species. Part 2: Cyperaceae. Stockholm. - (1981): Atlas of seeds and small fruits of Northwest-European plant species. Part 3: Salicaceae- Cruciferae. Stockholm. Bergmann, A. ( 1954): Die GroBschmetterlinge Mitteldeutschlands. Jena. BoRNER, C. ( 1952): Europae centralis Aphides. Die Blattlause Mitteleuropas. Mitt. Thiir. Bot. Ges. Beih. 3. BONN, S. & POSCHLOD, P. (1998): Ausbreitungsbiologie der Pflanzen Mitteleuropas. Wiesbaden. BRANDENBURGER, W. (1963): Vademecum zum Sammeln parasitischer Pilze. Stuttgart. BRENCHLEY, W. E. & WARNIGTON, K. (1930): The weed seed population of arable soil. 1. Numerical estimation of viable seeds and observations on their natural dormancy. J. Ecol. 18: 235-272. BuHR, H. (1964-1965): Bestimmungstabellen der Gallen (Zoo- und Phytocecidien) an Pflanzen Mittel- und Nordeuropas. 2 vol. Jena. BURGESS, N. A., CHATER, A. 0. & EDMONDSON, J. R. (1993): Flora Europaea. 2nd ed. Cambridge. CARTER, D. J. (1987): Raupen und Schmetterlinge Europas und ihre Futterpflanzen. Hamburg.
CSAPODY, V. (1968): Keimlingsbestimmungsbuch der Dikotyledonen. Budapest. DIECKMANN, L. ( 1972): Beitrage zur Insektenfauna der DDR. Coleoptera- Curculionidae: Ceutorhynchinae. Beitr. Ent. 22: 3-128. - (1972): Beitrage zur Insektenfauna der DDR. ColeopteraCurculionidae: Ceutorhynchinae. Beitr. Ent. 22: 3-128. - (1983): Beitrage zur Insektenfauna der DDR. ColeopteraCurculionidae : Tanymecinae, Leptopiinae, Cleoninae, Tanyrhynchinae, Cossoninae, Raymondionyrninae, Bogoinae, Tanysphyrinae. Beitr. Ent. 33: 257-381. DuLL, R. & KuTZELNIGG, H. (1994): Botanisch-okologisches Exkursionstaschenbuch. 5th ed. Heidelberg. ELLENBERG, H. (1996): Vegetation Mitteleuropas mit den Alpen. 5th ed. Stuttgart. -, WEBER, H. E., DuLL, R., WIRTH, v., WERNER, w. & PAuLissEN, D. (1991): Zeigerwerte von Pflanzen in Mitteleuropa. Scripta Geobotanica 18. ELLIS, M. B. & ELLIS, J.P. (1997): Microfungi on land plants. Slough. FAEGRI; K. & VANDER PIJL, L. (1979). The principles of pollination ecology. 3rd ed. Oxford. FITTER, A. H. (1978): An atlas of the wild flowers of Britain and Northern Europe. London. - & PEAT, H. J. (1994): The ecological flora database. J. Ecol. 82:415-425. FORSTER, W. & WOHLFAHRT, T. A. (1971): Die Schmetterlinge Mitteleuropas. Vol. IV. Bulen (Noctuidae). Stuttgart. FREIDENFELT, T. (1902): Studien tiber die Wurzeln krautiger Pflanzen. Flora 91: 115-208. FRANK, D. & KloTz, S. (1990): Biologisch-okologische Daten zur Flora der DDR. 2nd ed., Wiss. Beitr. Martin Luther-Univers. Halle-Wittenb. 32. FRYXELL, P. A. (1957): Mode of reproduction of higher plants. Bot. Rev. 23: 135-233. GRIME, J.P., HoDGSON, J. G. & HUNT, R. (1988): Comparative plant ecology - A functional approach to common British species. London. -, MASON, G., CuRTIS, A. V., RoMAN, J., BAND, S. R., MowFORTH, M.A. G., NEAL, A.M. & SHAW, S.C. (1981): A comparative study of germination characteristics in a local flora. J. Ecol. 69: 1017-1059. HAEUPLER, H. &. ScHONFELDER, P. (1989): Atlas der Famund Bliitenpflanzen der Bundesrepublik Deutschland. 2nd ed. Stuttgart. HAMANN, U. & WAGENITZ, G. (1977): Bibliographie zur Flora von Mitteleuropa. 2nd ed. Berlin. HANF, M. (1990): Ackerunkrauter Europas mit ihren Keimlingen und Samen. 3rd ed. Miinchen. Hannemann, H. J. (1961): Kleinschmetterlinge oder Mieralepidoptera I. Die Wickler (s.str.) (Tortricidae). In: DAHL, F. (ed.): Die Tierwelt Deutschlands und der angrenzenden Meeresteile. Jena. - (1964): Kleinschmetterlinge oder Micro Iepidoptera II. Die Wickler (s.l.) (Cochylidae und Carposinidae), die Ziinslerartige (Pyraloidea). In: DAHL, F. (ed.): Die Tierwelt Deutschlands und der angrenzenden Meeresteile. Jena. HARLEY, J. L. & HARLEY, E. L. (1987): A check-list of mycorrhiza in the British Flora. New Phytol. Suppl. lOS: 1-102.
HARPER, J. L. (1957): The ecological significance of dormancy and its importance in weed control. Proceedings of the 4th international congress on crop protection, Hamburg. pp. 415-420. HARTMANN, E., SCHULDES, H., KUBLER, R. & KONOLD, W. (1995): Neophyten. Biologie, Verbreitung und Kontrolle ausgewahlter Arten. Stuttgart. HEATH, J. et al. (1983ff): The moths and butterflies of Great Britain and Ireland. Vol. 1-10. Colchester. HEm, G. (1908ff): Illustrierte Flora von Mittel europa. Miinchen. (1935ff): 2nd ed., Miinchen, Berlin. (198lff): 3rd ed., Berlin. HEGNAUER, R. (1990): Chemotaxonomie der Pflanzen. Berlin. HEINTZE, A. (1932): Handbuch der Verbreitungsokologie der Pflanzen. Stockholm. HEJNY, S. & SLAVIK, B. (eds.) (1988-1997): Kvetena geske Socialisticke Republiky. HEJNY, S. & SLAVIK, B. (eds.) (1988): Vol. 1. HEJNY, S. & SLAVIK, B. (eds.) (1990): Vol. 2. SLAVIK, B. (ed.) (1992): Vol. 3. SLAVIK, B. (ed.) (1995): Vol. 4. SLAVIK, B. (ed.) (1997): Vol. 5. Praha. HENDEL, F. ( 1938): Agromyzidae. Vol. VI, 2 of Lindner, E. : Die Fliegen der palaearktischen Region. Stuttgart. HERING, M. (1935): Die Blattrninen Mittel- und Nord-Europas. Neubrandenburg. HESS, D. (1990): Die Bliite. 2nd ed. Stuttgart. HEss, H. E., LANDOLT, E. & HIRZEL, R. (1976): Flora der Schweiz und angrenzender Gebiete. 2nd ed. Basel. HODGSON, J. D., GRIME, J. P., HUNT, R. & THOMPSON, K. (1995): The electronic comparative plant ecology. London. HoLM, L., DoLL, J., HoLM, E. & PANCHO, J. (1997): World weeds. New York. HONARD, C. (1908-1913): Les Zoocecidies des Plantes d'Europe et du Bassin de Ia Mediterranee. (1908): Tome I, (1909): Tome II, (1913): Tome III. Paris. HuLTEN, E. (1971): Atlas of the distribution of vascular plants in North Western Europe. Stockholm. - & FRIES, M. (1986): Atlas of North European vascular plants north of the tropic of cancer II. Konigsstein. HuNDT, R. (1966): Okologisch-geobotanische Untersuchungen an Pflanzen der Mitteleuropaischen Wiesenvegetation. Jena. JAGER, E. & MuLLER-URI, C. (1981-1982): Wuchsform und Lebensgeschichte der (GefliBpflanzen Zentraleuropas Bibliographie. Part 1 : Quellenverzeichnis; Part 2: Artenverzeichnis (4 vol.). Halle, Universitats- und Landesbibliothek Sachsen-Anhalt. }ALAS, J. & SUOMNINEN, J. (1972ff): Atlas Florae Europaeae Vol 1-10. Helsinki. KALTENBACH, J. H. (1874): Pflanzenfeinde aus der Klasse der Insekten. Stuttgart. KIRCHNER, 0., LOEW, E. & SCHROETER, C. (1908ff): Lebensgeschichte der Bliitenpflanzen Mitteleuropas. Stuttgart. KNUTH P. (1889): Handbuch der Bliitenbiologie II. Leipzig. KORNECK, D., SCHNITTLER, M., & VOLLMER, I. (1996): Rote Liste der Fam- und Bliitenpflanzen (Pteridophyta et Spermatophyta) Deutschlands. Schr. R. Vegetationskunde 28: 21-187. KuGLER, H. (1970): Bliitenokologie. 2nd ed., Stuttgart. FLORA (2000) 195
121
KuTSCHERA, L. (1960): Wurzelatlas mitteleuropaischer Ackerunkrauter und Kulturpflanzen. Frankfurt. KUTSCHERA, L. & LICHTENEGGER, E. (1982-1992): Wurzelatlas mitteleuropaischer Gri.inlandpflanzen. Stuttgart. - -, SoBOTIK, M. & HAAS, D. (1997): Bewurzelung von Pflanzen in verschiedenen Lebensraumen. Stapfia 49: 1-331. LANDOLT, E. (1977): Okologische Zeigerwerte zur Schweizer Flora. Veroff. Geobot. lust. ETH, Stift. Riibel 64. LECK, M.A., PARKER, V. T. & SIMPSON, R. L. (eds) (1989): Ecology of soil seed banks. London. LovE, A. & LovE, D. (1961): Chromosome numbers of central and northwest European plant species. Opera Bot. Soc. Lund 5: 1-581. MEUSEL, H., JAGER, E. & WEINERT, E. (1965-1992): Vergleichende Chorologie der zentraleuropaischen Flora. (1965): Vol. 1. MEUSEL, H., JAGER, E. WEINERT, E. & RAUSCHERT, S. (1978): Vol. 2. MEUSEL, H. & JAGER, E. (1992): Vol. 3. Jena. MoHR, K. H. (1966): Chrysomelidae. In: FREUDE, H., HARDE, K. W. & LoHSE, G. A. (eds.): Die Kafer Mitteleuropas, Vol. 9. Krefeld. MooRE, D. M. (1982): Flora Europaea check-list and chromosome index. Cambridge. MULLER, H. (1873): Die Befruchtung der Blumen durch Insekten und gegenseitige Anpassungen beider. Leipzig. - ( 1881): Alpenblumen, ihre Befruchtung durch Insekten und ihre Anpassung an dieselben. Leipzig. MuLLER-SCHNEIDER, P. (1977): Verbreitungsbiologie (Diasporologie) der Bliitenpflanzen. Veroff. Geobot. lust. ETH, Stift. Riibel61: 1-226. - (1986): Verbreitungsbiologie der Bliitenpflanzen Graubiindens. Veroff Geobot. lust. ETH, Stift. Riibel85: 1-263. MuLLER, F. M. (1978): Seedlings of the North-Western European lowland. A flora of seedlings. The Hague, Boston. OBERDORFER, E. (1994): Pflanzensoziologische Exkursionsflora. 7th ed. Stuttgart. - (ed.) (1977-1992): Siiddeutsche Pflanzengesellschaften. Part I-IV. Stuttgart. PERRING, F. H. & WALTERS, S.M. (eds.) (1976): Atlas of the British flora. Wakefield. PERTTULA, U. (1941): Untersuchungen tiber die generative und vegetative Vermehrung der Bliitenpflanzen in der Wald-, Hainwiesen- und Hainfelsenvegetation. Ann. Acad. Scient. Fenn. Ser. A Tom. 58, No. 1. PETERSEN, A. (1988): Die Graser als Kulturpflanzen und Unkrauter auf Wiese, Weide und Acker. Berlin. PIRSON, A., ZIMMERMANN, M. H. (eds.) (1975ff.): Encyclopaedia of plant physiology; new series. Vol. 1-20. Berlin. PoTT, R. (1992): Die Pflanzengesellschaften Deutschlands. 2nd ed. Stuttgart. RABOTNOV, T. A. (ed.) (1974ff): Biologiceskaja flora Moskovskoj oblasti (Biological flora of the Moscow area). (1974): Vol. 1. (1975): Vol. 2. (1976): Vol. 3. (1978): Vol.
122
FLORA (2000) 195
4. (1980): Vol. 5. (1980): Vol. 6. (1983): Vol. 7. PAVLOV, V. N., RABOTNOV, T. A. & TICHOMIROV, V. N. (eds.) (1990): Vol. 8. PAVLOV, V. N. & TICHOMIROV, V. N. (eds.) (1993): Vol. 9. (1995): Vol. 10. (1995): Vol. 11. (1996): Vol. 12. (1997): Vol. 13. Moscow. RAUNKIAER, C. (1895-1899): De danske Blomsterplanters N aturhistorie. Kopenhagen. ROBERTS, H. A. (1981): Seed banks in soils. In: CoACKER, T. H. Advances in applied biology 6. London, 1-55. RoTHMALER, W. ( 1990-1999): Exkursionsflora von Deutschland. BASSLER, M., JAGER, E. J. & WERNER, K. (eds.) (1999): Vol. 2, 3rd ed. JAGER, E. J. & WERNER, K. (eds.) (1999): Vol. 3. ScHUBERT, R., VENT, W. & BASSLER, M. (eds.) (1990): Vol. 4. SALISBURY, E. J. (1964): Weeds and aliens. 2nd ed. London. - (1942): The reproductive capacity of plants. London. ScHNEIDER, C., SuKOPP, U. & SuKOPP, H. (1994): Biologisch-okologische Grundlagen des Schutzes geflihrdeter Segetalpflanzen. Schr. R. Vegetationskunde 26: 1-356. SCHOENICHEN, W. (1940): Biologie der geschiitzten Pflanzen Deutschlands. ScHROETER, C. (1926): Das Pflanzenleben der Alpen. 2nd ed. Ziirich. SCHUTZE, K. T. ( 1931): Die Biologie der Kleinschmetterlinge unter besonderer Beri.icksichtigung ihrer Nahrpflanzen und Erscheinungszeiten. Frankfurt. SEBALD, 0., SEYBOLD, S. & PHILIPPI, G. (1990ff): Die Famund Bliitenpflanzen Baden Wiirttembergs. Stuttgart. SERNANDER, R. (1906): Entwurf einer Monographie der europaischen Myrmekochoren. K. Sv. Vet.-Akad. Hand. 41: 1-410. SPENCER, K. A. (1990): Host specialization in the world Agromyzidae (Diptera). Dordrecht. STEWARD, A., PEARMAN, D. A. & PRESTON, C. D. (1994): Scarce plants in Britain. Peterborough. STOKOE, W. J. (1948): The caterpillars of British moths including the eggs, chrysalids and food-plants. London. THOMPSON, K., BAKKER, J.P. & BEKKER, R. M. (1997): The soil seed hanks of North West Europe: methodology, density and longevity. Cambridge. TUTIN, T. G., HEYWOOD, V. H., BURGESS, N. A., VALENTINE, D. H., WALTERS, S.M. & WEBB, D. A. (1964): Flora Europaea. Cambridge. VAN DER PIJL, L. (1982): Principles of dispersal in higher plants. 3rd ed. Berlin. VOGEL, E. F. de (1980): Seedlings of dicotyledons. Wageningen. WEEDA, E. J, WESTRA, R., WESTRA, c. & WESTRA, T. (1985-1991): Nederlandse oecologische Flora: wilde planten en hun relaties. 5 volumes. Hilversum. WELTEN, M. & SuTTER, R. (1982): Verbreitungsatlas der Fam- und Bliitenpflanzen der Schweiz. Basel. WISSKIRCHEN, R. & HAEUPLER, H. (1998): Standardliste der Fam- und Bliitenpflanzen Deutschlands. Stuttgart.