Vascular Plant Flora of Estonia

Summary



Introduction

During 270 years of studying Estonian vascular plant flora, an impressive amount of material on its species composition and distribution, cultural relationships and other aspects has been collected. A larger part of this vast information is summarised in Eesti NSV Floora (Flora of Estonian SSR), published in 11 volumes over the period from 1953 to 1984, but also in plant identification guides, handbooks and in several studies.

The present publication contains much basic data necessary for the study of flora: information on the scientists who have studied flora and on their works, an Estonian floristic bibliography, the list of vascular plants with their correct modern names, the position of the species in floristic elements, cultural relations between species, species distribution and frequency in Estonia. The present publication also presents a new analysis of Estonian flora and gives a brief overview of plant protection.



Abbreviations

BF - Flora of the Baltic Countries (Vol. 1 1993, Vol. 2 1996)

BKA - West-Estonian Archipelago Biosphere Reserve

BÖI - Institute of Botany and Ecology, University of Tartu; its library

EAR - Estonian Academic Library (former Library of Estonian Academy of Science)

Eesti TA KA - Central Archive of Estonian Academy of Science

EF - Eesti NSV Floora (Flora of Estonian SSR, Vol. 1-11 1953-1984)

ELF - Estonian Fund for Nature

ELM - Estonian Museum of Natural History

EMK - Estonian Forest Management Centre

EPA - Estonian Agricultural Academy

EPMÜ - Estonian Agricultural University

EPMÜ MKI - Institute of Silviculture, Estonian Agricultural University

KM - Estonian Literary Museum

LKA - Nature protection area

LK1, LK2, LK3 - protection categories for the species under protection (see Chapter 4.10)

LUS - Estonian Naturalists' Society

LÜÖS - Livländische Gemeinnützige und Ökonomische Sozietät (Public and Economic Society of Livonia)

(O) - Plant running wild by cut branches

ÕES - Learned Estonian Society

PR0, PR1, PR2, PR3, PR4, PR5 - the danger category for the taxon, by the Red Book of Estonia

RT - Riigi Teataja (State Gazette)

(S) - Plant running wild by seeds

TBA - Tallinn Botanical Gardens

TPÜ - Tallinn Pedagogical University

TPÜ B - Chair of Biology, Tallinn Pedagogical University

TPÜ ÖI - Institute of Ecology, Tallinn Pedagogical University

TRÜ - Tartu State University

TÜ - University of Tartu, its library

TÜ GG - Library of the Institute of Geography, University of Tartu

TÜ ZHI - Institute of Zoology and Hydrobiology, University of Tartu

(V) - A plant running wild by root suckers or rhizome

ZBI - Institute of Zoology and Botany of Estonian Agricultural University, its library

ZBI B - Library of the Botany department of Institute of Zoology and Botany

(!) - Mainly after the references to the Floras from 18th and 19th centuries on Estonia, Livonia and Kurland, if the habitat situated within the present borders of Estonia is mentioned there

(?) - Used in referring to the first mentioning of the taxon if in the publication referred to the presence of the taxon is marked with a questioning mark

-> - Arrow referring to the valid name

*** - (Before the name of a taxon) A taxon without herbarium evidence



History of Estonian Flora Research (Chapter 2)

Only studies on flora and plant systematics are discussed here, and therefore the scientific works concerning plant ecology and vegetation science, as well as those on bryology and algology have not been included. Besides, a good overview of the history of Estonian vegetation science was published lately (Masing et al. 1995). The contribution of the researchers is discussed only from the viewpoint of floristic studies, leaving aside all other (and often more significant) studies of the scientist.



Chapter 2.1. describes the historiography of learning the flora of Estonia. The most noteworthy among the historical surveys is the description of the early history of vegetation studies (up to the end of tsarist times) with an extensive bibliography compiled by G. Vilbaste. It was published as two almost identical articles (Vilberg 1929c, Vilbaste 1937e). A longer period is discussed in a survey of history given in Eesti NSV taimkate (Vegetation of the Estonian SSR) written by L.-M. Laasimer (1965a. A monograph on the history of Estonian nature studies up to the year 1917 published by L. Kongo (Êîíãî 1987). The history of collecting Estonian folk plant names has been written by G. Vilbaste (1993). These works end the list of profound discussions on the history of Estonian flora studies.

The first scientific works on flora of Estonia are dated back to the first half of the 18th century, when Johann Christian Buxbaum, a scholar from the Russian Academy of Science (founded in 1725) studied the flora of Ingermanland. In his works published in 1729, 1732 and 1740, Buxbaum describes some species noticed occasionally at several places near Narva where he stayed and rested.

The first proper list of vascular plants growing in Estonia is found from the work Topographische Nachrichten von Lief- und Ehstland, published in 1777 by August Wilhelm Hupel (1737-1819). His alphabetic list of natural species of Estonia and Livonia contains about 310 names, 260 of which are considered as species of natural and semi-natural flora nowadays (M. Kask 1983).

The main reason why the flora on the territory of Estonia was so modestly described in the 18th century was the lack of local scientists as well as the lack of a common institution for naturalists here (Chapter 2.2). Although Baltic-Germans from Estonia studied in the University of Tartu (Academia Gustaviana, Academia Gustavo-Carolina 1632-1710) and in universities abroad, they graduated the universities as medical doctors, pharmacists or pastors. One of the few local naturalists with the education of a botanist was J. B. Fischer, and we must stress here the pioneering significance of his works (Fischer 1778, 1784, 1791) - these publications were the main source for studying Livonian flora during the next sixty or seventy years.

The first half of the 19th century (Chapter 2.3.1.) is characterised by only brief studies of flora: only notes of encountered species and plant identification guides were published (Germann 1805; Grindel 1803; Friebe 1805; Drümpelmann 1809-1810). We could call this a period of amateur botanists, as those who studied plants here were pastors, medical doctors, teachers, and so on, while the university professors of botany were fascinated by more distant and less studied places, such as Crimea and Altai. The most noteworthy works written by amateur botanist of that period were the studies on flora of Saaremaa written by J. W. L. von Luce (1823, 1829b).

During the years 1811-1836, C. Fr. von Ledebour (1785-1851) was the professor of science in the University of Tartu, and his main contribution was developing the University Botanical Gardens into an institution accepted on a world scale, and the founding of the Tartu school of botany. In his major work, Flora Rossica (in four volumes, published in Stuttgard in 1841-1853), Ledebour gives the first complete survey of the flora of Russian Empire. According to Flora Rossica, 1037 species of vascular plants grow in Estonia (M. Kask 1994). On the material gathered from Estonia, Ledebour described at least one new species - Viola epipsila Ledeb.

The period of 1855-1865 is especially important for the studies of Estonian flora, because this is the period when intensive description of local flora started (Chapter 2.3.2). The period coincides partly with the publication of exchange herbarium Flora Exsiccata Liv-, Esth- und Kurlands by A. G. von Bunge (1803-1890). The leading role in the research was played by the professors and students of the University of Tartu (Fr. Schmidt, Ed. Russow, L. Gruner, P. Glehn, G. Pahnsch, etc.) who were assisted by amateurs (A. von Sass, Ed. Lehmann, A. Schmidt, G. K. Girgensohn, H. A. Dietrich, etc.). The centre of research was Tartu, especially after the founding of the Estonian Naturalists' Society there in 1853.

In 1852, a plant identification guide, Beschreibung der phanerogamischen Gewächse Esth-, Liv- und Curlands, by two amateur botanists - linguist F. J. Wiedemann (1805-1887) and school teacher Fr. Ed. Weber ( 1894) was published. This book is considered one of the best handbooks on Estonian vegetation from the last century. The guide covers 1050 species of vascular plants. The descriptions of the taxa are given in more detail than in earlier Flora publications, and the locations are given for the majority of the species.

During the years 1865-1920, no major survey on the local flora was published in Estonia, but smaller articles still appeared in periodical publications. The interest of the scientists towards studying the flora of Estonia, which was relatively well studied at that time, dropped significantly. The attention of the professors of botany was turned to unstudied regions in Russia, and the flora of Estonia was again left to the amateur botanists.

One of the most famous botanists working in Tartu was J. C. Klinge (1851-1902), whose most important work on the flora of Eastern Baltic was a plant identification guide with the title Flora von Est-, Liv und Curland (Klinge 1882b). Even today, this Flora by Klinge is used as a good auxiliary material in identifying the names of taxa used in the botanical literature in the 18th and 19th centuries. Another famous work by Klinge is on dendrological flora of the eastern Baltic (Klinge 1883c). The most famous work of Klinge, however, discusses the systematics of orchids (Orchidaceae).

At the end of the 19th century, several botanists from Sweden and Russia also studied Estonian vegetation. In 1901, a Flora on Saaremaa Island was published by K. J. F. Skottsberg (1880-1963) and J. T. C. Vestergren (1875-1930), and this is the last thorough discussion published on the flora of this island. On the basis of material gathered by these two researchers, G. A. H. Dahlstedt (1856-1943) published his Flora of Hieracium's in Saaremaa (1901), and L.P.R. Matsson his overview of Rosa's in Saaremaa (Matsson 1900). Russian botanists Fr. Ruprecht, K. F. Meinshausen, I. Schmalhausen and others studied the flora of Ingermanland and Narva region mainly. Based on Ruprecht's work, Flora Ingrica (Ruprecht 1860), and on the material collected by Kurilin and himself, Meinshausen compiled his famous Flora Ingrica (1870).

During the first half of the 20th century, K. R. Kupffer (1872-1935) became the leading scientist in floristic and vegetation studies. Kupffer started studying Estonian vegetation during his student years (1886-1889) in Tartu. He published about 350 works on flora of the eastern Baltics, including the systematics of some more complicated taxa (e.g. Violaceae). Kupffer was one of the first scientists in the Baltic countries to come out with plant protection initiatives.

Among the amateur botanists from the turn of the century, one of the best known is the pharmacist R. Lehbert, whose herbarium (over 11,000 pages) is among the most representative in all the Baltic countries. He is famous for his studies on the systematics of the genus Calamagrostis.

During the imperial university in Tartu (up to 1918), the flora of Estonia was studied rather disproportionately - more detailed descriptions appeared in the places where the botanists lived, but also on the areas surrounding manors, resorts, post stations and major roads. Some regions, however, especially in central and south-western Estonia, were not studied at all. The species composition of Estonian flora was outlined by the beginning of the 20th century, but data on their distribution was scarce. The studies on more complicated - the so-called critical taxa - were often carried out by visiting foreigners, while the scientists living in Estonia preferred to study the systematics of vegetation of distant and unstudied regions.

Researches on Estonian flora during the Estonian university in Tartu could be divided in two, according to the centre co-ordinating the studies. Thus, the period of 1919-1947 could be called the period of the Institute of Botany of the University of Tartu (Chapter 2.4.1.). An important turning point here is the year 1947, when the Institute of Biology (since 1952 the Institute of Zoology and Botany) was founded by the Academy of Science of the Estonian SSR. At this time the floristic studies transformed into a side subject in the University, and the Institute of Zoology and Botany became the centre of floristic studies. It is up till now still playing the key role in this field in Estonia (Chapter 2.4.2).

Before World War II, the vast work of organising both professional and amateur botanists in systematic studies of Estonian territory was carried out by G. Vilbaste and T. Lippmaa, and the latter was also the Head of the Institute of Botany in the University of Tartu.

The scientific importance of the floristic studies by Vilbaste are the most outstanding ones of that period. As an eager herbarium maker, he collected materials from all over Estonia and published numerous articles on the distribution of rare and interesting plants or on flora of specific regions. Vilbaste wrote the first complete plant field key-book published in Estonian (Vilberg 1922b, 1925a), which stayed in active use for over twenty years. His more profound studies discuss the vegetation of alvars (doctoral thesis, Vilberg, 1929b). In addition to his main work, he also published (at his own expense) a journal Loodusvaatleja ('Naturalist'), and was the editor of the journals Loodus ('Nature', 1923-1924), Eesti Looduskaitse ('Nature Protection in Estonia', 1938), Loodushoid ja Turism ('Nature Protection and Tourism', 1939-1940), and several publications of collected articles. The list of Vilbaste's publications includes over 1200 works. Another remarkable feature of this scientist is the variety of topics he has discussed. Several works are with the inclination towards country study, folklore, ethnographies or medical herbs, such as Harjumaa. Maateadusline lugemik (Harju County: Reading Book on Geography, Vilberg 1921), Kodumaal rännates I-III (Travelling in Estonia, Vilberg 1923c, 1924i), Meie kodumaa taimi rahva käsitluses I-II (The Plants of Our Country in Folklore, Vilberg 1934f, 1935b), and his masterpiece published after his death Eesti taimenimetused (Estonian Plant Names, Vilbaste 1993). Therefore, it is entirely justified to consider Vilbaste as the founder of ethnobotanics in Estonia.

T. Lippmaa was the Head of the Institute of Botany and the Botanical Gardens of the University of Tartu in the years 1930-1943. During the 1920s, his main interest was plant physiology, but his first floristic studies on Setumaa (Lipman 1923a, b; Lippmaa 1928 a, b) and monograph on the flora and vegetation of Pärnu County (Lippmaa 1931 a, b) are also written at that time. The height of his research career as vegetation scientist was achieved in the analysis of Estonian vegetation and division of it into floristic districts (Lippmaa 1935 a, b), which is one of the most frequently used division in plant geography nowadays.

A large-scale project of mapping the vegetation of Estonia was started in 1934 under the supervision of T. Lippmaa. In addition to being valuable from the viewpoint of vegetation sciences, it was an important work also for the plant geography - the explanations to the maps contain a lot of floristic material. The mapping was finished under the supervision of L.-M. Laasimer by the year 1955.

In the second half of the 1920s, the Institute of Botany of the University of Tartu started research on unstudied areas of Estonia. The project was aimed at the systematic and thorough study of the flora and vegetation on the whole territory of Estonia. Initiated by Lippmaa, several amateur botanists started to study the systematics of many critical plant genera in Estonia (Hieracium - A. Üksip, Alchemilla - W. Reinthal, Taraxacum - B. Saarsoo, etc.).

One of the largest projects joining all botanists in Estonia at that time was the compilation of the exchange herbarium Eesti taimed ('Estonian Plants', I-IV, Lippmaa, Eichwald 1933, 1935; Eichwald 1938d, 1939a). The aim of the collection was to advance the studies on Estonian flora, improve the University's Herbarium Generale by complementing it with new plants from other countries, as well as to supply the larger educational establishments in our country with herbariums (Lippmaa 1933c). In addition to its authors, 39 professional and amateur botanists participated in compiling the collection. The majority of the work in compiling the exchange collection was carried out by Karl Eichwald (1889-1976), who was the conservator of the herbarium of the University of Tartu in 1930-1944.

The Soviet occupation that started in 1940 did not have as much influence on the unpolitical botanical studies as it had on such fields as geography and the humanities. However, the publication possibilities worsened significantly. The libraries started to destroy inadvisable literature or close such books up in special depositories. The mapping of vegetation and plant species became more difficult as the maps were subject to secrecy during the Soviet time, or they were inaccurate or intentionally deformed.

From the years of German occupation, the most noteworthy publication was a plant identification guide, Kodumaa taimestik ('Vegetation of Our Country', Enari et al. 1943), which was the next identification guide on all vascular plants after that of Vilbaste (Vilberg 1925a). Although the majority of herbariums, manuscripts and literature survived the war, the lines of the botanists thinned. T. Lippmaa, the leader of Estonian vegetation science, was killed in the war (1943). In the last year of independence, or during the years of occupation, several botanists fled from Estonia (L. Enari, P. Kaaret, E. Leppik, A. Mathiesen, B. Saarsoo, H. Salasoo, A. Tamsalu, P. W. Thomson, B. Tuiskvere, etc.).

In 1947, the Institute of Biology was founded at the Academy of Science of the Estonian SSR. During the first years of its existence, the main fields of activity were inventories of Estonian nature, starting the collections, and beginning the education of young biologists. The department of Botany of the Institute of Zoology and Botany decided to start working on the complete "Flora of Estonia", and to publish several identification guides and manuals to help people learn about plants.

The first volume of this complete work, Eesti NSV floora ("The Flora of Estonian SSR"), written by the botanists of the Institute of Zoology and Botany and the botanists of the University of Tartu, was published in 1953. The last volume appeared in 1984 and the Index in 1998. The collection includes 54 orders, 124 families, 676 genera and 2028 species (if we add also the smaller taxa, the number of taxa is 2500). Besides the native taxa, also more important cultivated plants (about 1/5 of the species) and aliens are described in this work. Every species is presented with parallel names and references to literature, and a detailed morphological description is given together with data on its habitat and distribution in Estonia, its economic importance, etc. Twenty-seven authors, mainly from the Institute of Zoology and Botany and the University of Tartu, participated in writing the Flora.

Nowadays, floristic research is mainly carried out in two institutions - the Institute of Zoology and Botany of the Estonian Agricultural University and the Institute of Botany and Ecology of Tartu University. Other institutions are engaged with either applied research, teaching botany or popularisation. At the moment, the major works of the botanists of the Institute of Zoology and Botany include the compilation of the atlas of Estonian vascular plants and the Flora of the Baltic Countries (in three volumes; 1st Vol. published in 1993, 2nd in 1996). The new identification guide of vascular plants is ready (Krall et al. 1999). The scientists participate in the compiling and revision of the Red Book of Estonia, and in compiling the lists of plants under protection. Since 1970, the botanists have contributed to the map of European Flora (Atlas Florae Europaeae, 1972-). The last volume of the Flora of the Baltic countries is still in progress.

The floristic research carried out in the Institute of Botany and Ecology of Tartu University are usually connected with the Bachelor, Master or Doctoral studies. Such floristic studies often discuss the morphological variations in several difficult plant genera (Potentilla, Alchemilla, Pilosella, etc.). Another type of floristic research carried out by the Institute concerns the primary tasks of its researchers (E. Roosaluste: protected species, flora of Saaremaa and Vormsi islands).



Herbariums (Chapter 3.1)

The largest herbarium in Estonia is situated in the Institute of Botany and Ecology of the University of Tartu, where the total number of sheets is estimated to be between 245,000 and 275,000 (of which about 45,000 are collected from Estonia). The largest collection of Estonian plants in herbarium is situated in the Institute of Zoology and Botany of the Estonian Agricultural University (110,000 sheets from Estonia and about 25,000 from foreign countries, with the herbarium of 11,000 sheets collected by G. Vilbaste stored separately). A lot of herbarium specimens collected in Estonia during the 19th and 20th centuries are stored in the Estonian Museum of Natural History (the size of this herbarium is about 62,000 sheets). Smaller herbariums are stored in the Tallinn Botanical Gardens, in protected nature areas, and in country museums. The largest private herbarium is owned by H. Aasamaa (over 30,000 sheets).



The Bibliography of Literature on Estonian Flora (Chapter 9, comments in Chapter 3.2)

This list contains 6,500 entries of literature published from 1637 to 1999. The bibliography focuses on the following sources.

1. The literature referred to in the present publication. Therefore, also occasional seed exchange lists of botanical gardens occur in this bibliography.

2. Floristic, dendrological and geobotanical publications containing data on the distribution of vascular plants in Estonia. However, the typological literature containing the description of species composition of a community with any specific locations mentioned are not included. In spite of all efforts, the list of dendrological and forestry literature is subjective.

3. General publications on the vegetation and plant systematics of the eastern Baltic, which are interesting especially from the viewpoint of phytogeographical studies. Of the publications on systematics and monographs, the works based on herbarium specimens collected from Estonia have been preferred.

4. Phytogeographical studies and atlases containing the data on species distribution in Estonia. The list focuses on publications using original data. "Secondary" floristic literature, i.e. the literature repeating the data of earlier authors without presenting anything new, has also, however, been included.

5. Various kinds of brief notes, notices, travel articles, reviews and so on containing floristic data. We must also say that the list of literature on ancient trees is not complete in this bibliography.

6. Non-botanical works containing notes on vegetation, especially on plant distribution. For example, the majority of the zoological studies contain some observations on vegetation. The list is more complete in biological publications of the last century and the beginning of this century, as the number of special flora studies was much smaller during that time.

7. Plant identification guides on Estonian vascular plants (in any language) and the lists of vascular plants.

8. Major works on plant protection in Estonia.

9. Biographies, bibliographies, and other publications on botanists, dendrologists, amateur botanists and others who have worked in Estonia; literature discussing the history of Estonian flora study.

10. Major manuscripts on the topic (graduate and diploma papers, candidate and doctoral theses, final reports of research grants, etc.; also some term papers written by students).

Data on flora is also found in mycological literature, primarily through mentioning the hosts of the micromycetes. However, as there are very good bibliographies of Estonian mycological literature (Järva, Parmasto 1980; Järva, Parmasto, Vaasma 1998), there is no need to repeat this data in the present list.

The bibliography is compiled mainly de visu. Articles published in newspapers or journals, rather than scientific ones, are occasionally represented. The complete inclusion of such articles has not been the purpose of this bibliography. In the case of older and rare literature, as well as for the majority of the manuscripts, the notes on the library where the copy used is situated is often added (sometimes with call number). If it was not possible to find the publication (the reference originates from another bibliography), it is marked with abbreviation non v. (non vidi).



Unpublished Databases (Chapter 3.3)

A database here is defined as a collection of data on any kind of carrier (paper, electronic carriers, etc.). Often the term of a database coincides with the term of manuscript, as the database is usually formed with the purpose to fulfil certain tasks (e.g. research or writing a book). The survey gives the name of the collector (person or institution), contents, and form of the database, as well as its volume and limitations for use. The databases are arranged in four groups according to their topic (entire flora of Estonia, local floras, floristic data by habitats, collections and bibliographies). The overview of the manuscripts is not complete; the complete list is given in the bibliography at the end of this publication.



The List of Estonian Vascular Plants (Chapter 7) and its Structure (Chapter 4)

The list of Estonian vascular plants (Chapter 7) tries to give as complete as possible a list of all the indigenous, naturalised, run wild and casual alien species and subspecies of vascular plants found on the territory of Estonia. The list contains also the taxa without herbarium evidence and therefore not considered as belonging to Estonian flora. Such taxa are marked with an asterisk (*)

The species concept used in this publication is wider than its traditional use in Estonian botanical literature during the last decades. Here, more attention is paid to the valid name, and the rank of the taxon has not been considered so important. Of hybrid taxa, the species with independent spread by either generative or vegetative means has been the ground for inclusion.

Within the division, the families are ordered alphabetically, and within the angiosperms, the families of monocotyledons and dicotyledons are listed separately. Within the families, the genera, species and subspecies, sometimes also the varieties (if the variety has been discussed in earlier Estonian floristic literature as an independent species) are also ordered alphabetically.

The valid Latin names are given in bold italics, and the synonyms in ordinary italics. The list also contains some valid genus names with no valid species name - in such cases no valid species of these genera are found in Estonia. The indigenous species are discussed in more detail. The data on subspecies and micro-species, hybrids, introduced or unverified taxa may be incomplete.



Scientific Names and Synonyms (Chapter 4.1)

The ordering of the synonyms was based on the following publications: Flora Europaea 1963-1981 (also the reprint of the 1st volume in 1994); Atlas Florae Europaea, parts 1-11 (1972-1998) and proof sheets of the 12th part; Wiersema et al. 1990; Brummitt 1992; Gunn et al. 1992; Hämet-Ahti et al. 1992, 1998; Greuter 1993, 1994. Also, more recent discussions published in taxonomical periodicals and recent monographs were used here. In case of contradiction, traditional names in use for decades have been favoured.

The orthography and abbreviations of the authors of the taxa's names are usually harmonised with the handbook by Brummitt and Powell (1992). The exceptions occur mainly in the names of Estonian authors (Juxip or Yuksip pro Üksip, Vilyasoo pro Viljasoo, also Nenukow pro Nenjukov, etc.).

In choosing between the synonyms, the names used in Estonia for decades have been preferred. If Eesti NSV floora and Flora Europaea give different valid names, these names are always given as a synonym. The older names which are no longer in use are given only when the name is connected with the first mentioning of the taxon in Estonia.



First Mentioning in Publications on Estonian Flora (Chapter 4.2)

First mentioning is discussed here in rather wide scope. The mentioning of the Latin name of the species (subspecies, variation, etc.) has been considered sufficient here. There has been no similar survey on Estonian flora before.

In cases where the occurrence of a taxon in Estonia was denied, but it was still found later, were not considered as first mentioning. If the later authors have proved the earlier data to be wrong, we have tried to include both sources of information. If the first mentioning has also been a new species description, the reference is marked in bold type. The problems concerning the first mentioning in literature are the following:

1. Often the names given in the literature are written so that it is not possible to identify the species clearly, especially in the cases where no author name of the taxon is given. Such names are very frequent in the works published before the Species plantarum... by K. Linné (Linnaeus 1753). The earlier publications are referred to only if we can be sure of the identification of the taxon (description or illustration is added).

2. If the name used in the first mentioning is different from the presently used one, the old name is also included. If the name has been out of use and forgotten, it is not given as a synonym after the present valid name. The names are given with the names of the authors (if known), irrespective of whether the name has been used with or without the author's name (sine auct.) in the referred publication.

3. Several problems arise from the different species concepts. If the earlier discussions of the taxon have been broad and it is not possible to harmonise all the subspecies rank with the present narrow understanding of the species (resp. subspecies), usually (but not always) two references are given: first mentioning for a larger and a smaller unit. However, if the species concept has broadened, the mentioning of one of its micro-species is understood as the first mentioning of the species.

4. In the Floras of the 18th and 19th centuries, Estonia, Livonia and Kurland are often discussed as one entity. Several authors (Fischer, Fleischer, Grindel, etc.) do not mention the territories separately and do not give much data on the specific locations the plant was found. When the specific place is mentioned in these works, the reference is marked with an exclamation mark (!). The references to the Floras of Ingermanland are referred to only if the location is in Viru County or the Narva region.

5. We could not find some of the literature referred to in the earlier works. The data of first mentioning in the works published abroad on Estonia is probably also incomplete.

6. The year of publication of the old Floras referred to here may differ from the number printed on the title page, if the work has been published in parts during several years (most famous among such publications is Flora Rossica by Ledebour).

7. Finding out the first mentioning in literature for the plants cultivated in Estonia has not been the aim of the present publication, as it would presume profound studies on the literature discussing forestry, dendrology, gardening and agriculture.



References in Flora of the Baltic Countries and Eesti NSV floora (Chapter 4.3)

These references are given for the taxa discussed in these publications. The first two volumes of Eesti NSV floora have appeared in reprint, and both references are included (e.g. 1/1 meaning 1st print of 1 volume; - the reprint of the volume, etc.). Only the two first volumes of the Flora of the Baltic Countries, have appeared; the third one on Monocotyledons, Campanulaceae and Asteraceae has not been published yet. The number of volume and the page in the Flora of the Baltic Countries stands after the family and genus names only.



Estonian Name (Chapter 4.4)

The correctness of Estonian names was checked from the database of the Committee for Estonian Plant Names. No new names are introduced in this publication, but the Committee did change the orthography for some of the names. Folkloristic or earlier standardised names are discussed in more details in the monograph by G. Vilbaste (1993) and therefore it is not considered necessary to repeat them here. The Estonian names are usually written without capitals, with the exception of persons' names used as species epithets.



Floristic elements (Chapter 4.5)

In the present publication, the floristic element is defined as a group of species whose geographical distribution area is overlapping. The grounds for defining floristic elements have always stirred up discussions, as the areas are overlapping only to a certain extent. Moreover, the grouping of the species between different floristic elements depends on the species concept - if we consider the American variety of a circumpolar species as an individual species, we must classify the former taxon as one of Eurasian or a European floristic element.

There are two common approaches to this problem: the floristic elements are numerous and various, or only a few general but inaccurate groups are used. During the last decades the Estonian floristic literature has traditionally used the North-European vascular plants' distribution atlas by Hultén (1950, reprint 1971), that defines 48 area types. The present publication is also based on the floristic elements by Hultén, but the groups defined are somewhat different, both in volume and substance, from those given in earlier analyses of Estonian flora (Laasimer 1965a; Masing 1979). In defining the floristic elements, mainly the geographical position of the mass centre was considered. If the taxon is not present on the maps by Hultén or is grouped as a species with unknown area there, other sources were used to determine the floristic element, the most important of which was the revised edition of Hultén's atlas (Hultén, Fries 1986). Disputable cases are marked with a question mark. In the case of immigrant plants or plants run wild, the origin of the taxon (natural area) is given.

Arctic and arctic-alpine element (Hultén groups 1-11): Contains taxa with arctic and circumpolar distribution (in polar regions of America and Eurasia) in arctic Europe or Asia. The species of arctic-alpine element have part of their distribution range in the mountains in warmer climate zones.

Circumpolar element (synonyms: holarctic, circumboreal; Hultén groups 16, 24, 29, 30): The taxon is found in the temperate zone of the northern hemisphere, partially extending to other vegetation zones. The total area of discontinuous circumpolar subtype (Hultén groups 17, 31-33, 41) is divided into sub-areas with vast territories in between where the taxon has not been found. The larger sub-areas of such discontinuous circumpolar taxa are usually found in Europe, North America and the Far East. Circumpolar coastal plants (Hultén groups 21, 22) are found mainly in the temperate coasts of Atlantic and Pacific Oceans, in smaller numbers also on the coasts of inland seas (Mediterranean, Black and Caspian Sea) or on the coasts of larger rivers and lakes.

Eurasian element: The taxon is found all over the temperate zone of Eurasia (Hultén groups 15, 23, 28, 38). We could separate here the continental version of the Eurasian element, where the mass centre of the taxon is situated in eastern or south-eastern Europe or the western part of Asia, while the distribution area is much smaller in western Europe and the Far East (Hultén groups 14, 37, 39, 40). The continental subtype of the Eurasian element coincides more or less with the pontine (pontiline) and ponto-sarmentous (pontosarmaatiline) elements defined by Lippmaa (1935b, c).

Euro-Siberian element: The mass centre of this floristic element is situated in the western and central Europe, and the area continues up to Siberia. This element covers the Hultén groups 25-27, 35 and 36. One could see several differences if we compare this division with the composition of Eurasian and Euro-Siberian floristic elements by Masing (1979). The present work uses the broader concept of the Euro-Siberian element.

European element [Hultén groups 12, 13, 19 (Atlantic), 20 (Subatlantic), 34, 42]: The mass centre is in western and southern Europe, but the area extends to northern and eastern Europe. Some taxa (many so-called sub-Atlantic taxa) may even extend to Central Asia in the form of a few isolated distribution areas. European coastal plants (Hultén group 18) are usually distributed on the coastal regions of the Atlantic Ocean, but also on the coasts of the Baltic Sea and the Mediterranean Sea. For some taxa, the area extends to Iceland and Spitsbergen in the north, and the Caspian Sea and the Red Sea in the south. The Atlantic group is not contrasted here with the continental European group, as the mass centre for both of these groups is situated in western Europe, where there are no significant differences between the continental and maritime regions.

Asian element: The mass centre of this floristic element lays in certain regions in Asia (e.g. South-east Asia, China, etc.), and the number of areas outside these regions is small. This element covers the Hultén groups 43 and 44.

American element (Hultén groups 45): The natural distribution area of the taxon is in North and/or South America, but it is found in other places as an introduced or immigrated taxon.

Species known in cultivated forms only (Hultén group 46): It is hard or impossible to tell the origin of the taxon, although several theories exist. This group does not correspond to the definition of a floristic element really, but, on the basis of our present knowledge, it is not possible to list these taxa under other floristic elements.

Endemic element (Hultén group 47): Endemic species of the Baltic countries and Estonia (see also Chapter 4.8).



Status of Flora (Chapter 4.6)

The status of cultivated, introduced or naturalised plants, or a species run wild is recorded here. All other species (incl. archaeophytes) are indigenous species. Archaeophytes arrived to the territory of Estonia together with the first humans, usually during the prehistoric period, but in any case not later than in the middle of the 18th century. Anthropophytes have arrived to Estonia since the times of the first settlements, while the first scientific notes on species introduction could be dated by the middle of the 18th century. Differences in comparison with the discussions in Flora Europaea or Atlas Florae Europaeae are given here.

Cultivated plants are species and subspecies of intentionally introduced and cultivated plants. If an indigenous species is cultivated in some places, it is also mentioned here.

Aliens are plants who have migrated here along with humans and grow by railway stations, ports, wool mills and elevators, on refuse dumps and other similar sites.

The taxa runs wild contain plants who have "fled" the places of cultivation, and such species are always found near fields and gardens, on refuse dumps, abandoned farmlands, by the roads, etc. Distinctions have been made between occasional running wild (the taxon is not able to reproduce and spread without the help of humans in this habitat) and permanent running wild (the species shows at least some extent of vegetative or generative reproduction without human help). The type of running wild is added mainly in the case of woody plants: by seeds (S), by root suckers or rhizome (V), or by cut branches (O). We have tried to make the list of the species running wild by seeds as complete as possible, although some of the species cultivated in few collections only, where the running wild is limited to the nearest surrounding or a compost heap have not been included. Running wild by seeds is considered to be the most important way of running wild, and therefore the ability of such plants to reproduce by cut branches or develop root suckers has not always been referred to.

Naturalised plants are introduced plants which are more or less common in natural communities without needs of human help in distribution (e.g. silvicultural plantations of foreign species with any natural second growth are not considered as naturalised species). However, the introduced species reproduce usually more successfully in anthropogenic communities. Some of the naturalised taxa have originally been aliens, but the majority is formed of cultivated plants run wild. In reality, we could also speak of naturalisation in the case of several taxa known only as weeds or ruderal plants, but which have acclimatised here as successfully as archaeophytes. To distinguish these species from the true naturalisation and migration into natural communities, such species are called acclimatised aliens or plants permanently running wild.

We must also mention the different interpretations of the term naturalisation, as the majority of the communities are nowadays more or less influenced by human activities. Several species have naturalised on our western islands where the climate is milder, while representatives of the same taxon may suffer hard from frost damage and do not reproduce naturally in eastern Estonia.



Distribution and Frequency in Estonia (Chapter 4.7)

The frequency evaluations are based on the publications or manuscripts from the last decades, on the author's observations, and on evaluations received from co-workers. The manuscripts of the Estonian plant atlas is not ready yet and therefore we have no objective grounds for evaluating the frequency of taxa. However, we may say that the evaluated frequency is good enough for the rare and common species. The locations of rarities are carefully recorded, and, similarly, in the case of a common taxon, the possibilities to make errors here are rather small. Such frequency estimations as "uncommon", "scattered" or "occasional" are defined less precisely.

The following frequency classes are used:

Unclear - (a) The presence in Estonian flora is uncertain due to missing or untrustworthy data in herbariums or references in literature ("presence unclear"); (b) The species has not been found during the last 50 years, but all the earlier sites have not yet been checked ("presence unclear"); © The taxon has been described lately, its present existence in Estonia is proved but its distribution has not yet been studied sufficiently ("distribution unclear").

Extinct - The last proved findings (as herbarium specimens or notes) dates back more than 50 years and all the former sites have been checked.

Very rare - One to three proved findings during the last 50 years and the taxon may not be permanent in the locations (especially characteristic to aliens or species running wild).

Rare - 4-10 findings during the last 50 years.

Uncommon - More locations (11-30) where the species have been found; usually the locations are in one and the same region (e.g. Saaremaa, south-eastern Estonia).

Scattered - Distributed all over Estonia, but sparse everywhere; no specific distribution locations.

Occasional - The frequency of the taxon is similar to the previous one, but the distribution is limited to certain regions in Estonia and in these regions (e.g. in Saaremaa, south-eastern Estonia, etc.) the species may be rather common.

Common - Grows in suitable locations, usually not very numerous but found all over Estonia, or in large numbers in some areas in Estonia (while there are areas where this taxon is missing).

Frequent - The taxon grows in almost every suitable location and in large numbers.

In case of aliens, and sometimes also the plants run wild, the location, year and the name of the collector (or the data known at the moment) of the first (preserved) herbarium specimens are added. The herbarised plant may be dated significantly later than the first mentioning of the species in the literature.

The frequency of occurrence is used uniformly for indigenous species and cultivated plants, species run wild, aliens and naturalised species. Therefore, one taxon may have more than one frequency evaluation, for example, an indigenous species may be "common, rarely cultivated and run wild", or a cultivated taxon may be "scattered in cultivations, rarely run wild".



Margins of the distribution area (Chapter 4.8)

When a taxon in Estonia is on the margin of a distribution area, the quarter of the area's margin in Estonia is mentioned. In some cases also the position on the area margin of the discontinued area or near the area margin is mentioned, but these cases are not included in the analysis of the flora (Chapter 4.3). The area margin is clearly shown on the distribution map of the taxon, but its explanation in words, however well composed, always leaves room for different interpretations. The margin of area is recorded for the indigenous species only. Endemic taxon is here understood as a taxon with relatively small distribution area (region, mountain area, island). In the list of Estonian vascular plants, the narrower concept of endemism has been used: endemic species are only the species distributed in Estonia and the Baltic countries. However, sometimes a taxon with much wider area is called endemic - for example, the Atlas of European Flora discusses European endemic plants. When the larger part of the area of an endemic species is situated in Estonia, its position on the area margin is not mentioned separately.



Sensitivity to Human Impact (Chapter 4.9)

Hemeraphob is a taxon which is disturbed by human activities; such plants usually disappear from the ordinary cultivated areas or show clear signs of stress there (no flowers, no seeds, etc.). Usually, the reasons can be found from rapid changes in environment caused by human activities. Some hemeraphobs are obligatory mycotrophic plants.

Hemeradiaphor is a taxon indifferent to a certain limit or human activities, but if the activities become more intensive, the plants turn into hemeraphobes of (more rarely) apophytes. Such plants are often found in communities with little human influence (coastal regions, denudations, dunes, etc.).

Apophyte is an indigenous taxon of plants preferring moderate to strong human impact and the communities changed by human activities. Some of the apophytes have been anthropophytes before.

Anthropophyte is an introduced taxon of plants (either alien or run wild from cultivation resp. naturalised) surviving only in communities significantly changed by human activities. Such plants are very rare in natural communities.

In determining the relations for the cultivated plants, mainly the thesis by L. Enari (1944) and an article by H. Rebassoo (1962a) has been used, but other sources have been helpful here as well. The thesis by L. Enari is always referred to in the list, and the apophytes and anthropophytes are given the number of the divisions by Enari. If the relations are given without references, it has a different source or originates from observations by the author.

Endangered Species and their Protection (Chapter 4.10)

The evaluation of danger (threatenedness) is added to the taxa (a) uncommon in Estonia (many of which being on their area margin here); (b) with significant decrease in their distribution during the last 50 years; © with significant decrease in their distribution in neighbouring regions (the Baltic countries) during the last 50 years. The species under protection are marked with abbreviations for the protection category, i.e. LK1, LK2, or LK3.

The 1st category of protected species is those with high scientific value (relicts, species with narrow area and on the area margin), very rare plants (1-5 locations) and clearly endangered species (Ü, Kukk 1995e: 73). There are 22 species in the first category. The list of the species and grounds for their protection were ratified in 1994 (RT1 1994, 46, 773; RT1 1998 23, 323).

The 2nd category of protected species also consists of rare and endangered species and species of scientific value, but the danger is not so direct as that of the species of the first protection category. The rarest species of the second category are nowadays usually found in nature protection areas. More common species are endemic, relict from the former climatic periods or situated on the margin of their distribution area (Ü. Kukk 1995c: 113). The list consists of 145 species, and it was ratified in 1994 (RT1 1994, 94, 1610; Rt1 1998, 23, 323).

The 3rd category of protected species covers the species quite common in Estonia but still endangered for different reasons. Many of these plants are decorative plants, herbs or edible plants. It is either difficult to protect the locations of such plants or it has not yet been necessary (Ü. Kukk 1995d: 158). The list consists of 41 species (RTl 1995, 36).

Special references have been used for species included in the Red Book of Estonia (Lilleleht 1998a: 14), using the abbreviation's PR0, PR1, PR2, PR3, PR4 and PR5. The number here signifies the protection category.

0 Extinct or probably extinct. Species with permanent natural population extinct or probably extinct in Estonia. In spite of repeated searches, the species has not been found in Estonia after 1950 (water plants after 1965), but there are reliable documents on their earlier existence.

1 Endangered. The species is in great danger of becoming extinct, with the numbers growing very small or the habitats disturbed to a critical extent, and if the dangerous situation continues, the future existence of these species is questionable.

2 Vulnerable. The species with populations decreasing due to overexploitation, or due to destruction or damage of habitats, and their distribution and numbers in Estonia are decreasing. If the situation continues, the species may become highly endangered in the future.

3 Rare. Species which occur in Estonia within restricted areas or in few habitats or very sparsely, but not belonging to the 1st or 2nd protection categories. The population could be damaged easily.

4 Care demanding. Species needing monitoring. This covers species that are still quite common but their numbers have shown a rapid decline lately and if the situation continues, the species may soon pass to the endangered species category. This category also includes species that have been in the former categories before, with their status having improved significantly.

5 Indeterminate. Species probably belonging to the categories of extinct, highly endangered, endangered or rare species, but due to insufficient data it is not possible to determine their exact status.



Analysis of the Present Flora of Estonia (Chapter 5)

The indigenous flora of Estonia includes 1441 species (incl. hybridogenic species) of vascular plants. The number also covers taxa with unclear existence in present Estonian flora (herbarium evidence proves earlier existence, but no fresh findings), and micro-species. Usually, the species of the genera Taraxacum, Hieracium and Pilosella are considered as microspecies, but we have no objective grounds for such discrimination. Existence of another 130 taxa has been published in literature, on which no proof has been found, and these thus taxa have not been included.

If we add the 97 subspecies to the 1441 indigenous species (i.e. the species are found as two or more subspecies in Estonian indigenous flora), the total number of taxa increases to 1538. The number of indigenous pteridophytes is 50, the number for gymnosperms is 4 and angiosperms 1387. These indigenous species belong to 113 families and 443 genera. A general picture of the number of species in the families is given in Table 3. The largest family is Taraxacum with 165 species, followed by Hieracium with 81 and Carex with 69 species.

There are 82 naturalized species belonging to 30 families known in Estonia (see also Chapter 4.6). The majority of naturalized species (51) are cultivated species run wild; 26 species have arrived in Estonia as casual aliens, and five species are cultivated species run wild and aliens.

The total number of species and subspecies migrated or cultured and run wild (including naturalized) is 718, and 10 of the alien species are represented by two subspecies. However, 17 of our indigenous species are also represented by one or more alien subspecies. The number of alien taxa is 373 and there are 409 taxa run wild, while 54 taxa occur both as casual aliens and cultivated plants run wild. The adventive taxa belong to 84 families, of which 24 are represented by migrated species and subspecies only (cf. the number of families consisting of indigenous taxa only is 52).



Protection of Estonian Vegetation (Chapter 6)

Dozens of good overviews have been published on the history of plant protection in Estonia (Eilart 1967a; Eichwald 1970b, 1973; R. Sander 1975a; M. Kask, Kuusk 1981; Laasimer 1986d; Ü. Kukk 1987c, 1988b, 1992b, 1994f, 1998a, c; Varep 1990; Kalda 1991; Kongo 1998a; Lilleleht 1998a, b; etc.). The Red Book of Estonia (Lilleleht 1998a) covers 309 endangered species and subspecies of vascular plants, making up 20.6 per cent of the total number of species in Estonian flora. Of these, 159 species are protected due to their rarity or endangered status, and 26 species are protected mainly due to their decorative value. The preservation of rare species or communities may sometimes be especially complicated:



1. Protecting oligotrophic and semidystrophic lakes. The health of several plant species (Lobelia dortmanna, Isoetes lacustris, I. echinospora, Sparganium gramineum, etc.) has worsened significantly during the last decades. The occurrence of water lobelia (Lobelia dortmanna) was known in 15 bodies of water, but in two of them it has died out during this century and its numbers in the majority of the locations have decreased.

2. Protecting fens. While the main danger during the last decades has been drainage of the fens, the last years have brought along the problem of overgrowth in the swampy areas - the majority of the fens have been preserved as opened landscapes due to brief or moderate grazing.

3. Disappearance of meadow communities. If there is no grazing or mowing, the alvars develop into alvar forests, coastal meadows overgrow with reed, and the flood plains overgrow with bush. The tendency is especially clear if we look at the decrease of the area under the wooded meadows, along with the decrease in the distribution of plants growing in these communities. Although the territory of wooded meadows is about 1000 times smaller than it was at the beginning of this century (T. Kukk, Kull 1997), we fortunately cannot say the same about the distribution of the plant species on the wooded meadows.

4. The number of old weeds, especially the weeds of flax cultivations, has decreased significantly. Unfortunately, the same tendency is clearly visible in other European countries as well. Protecting a specific weed species is a very complicated task. Some help may be found from leaving a special weed zone around the fields, which is ploughed every year but with no herbicides used. The best way of protecting endangered anthropophytes is to grow them in the botanical gardens.

Reintroduction and re-plantation have sometimes been advised as a means of protecting flora. Both of these methods, however, especially re-plantation, are not acceptable from the viewpoint of plant geography. We must not discuss the indigenous species separately from its location, because the fact that the individual plant is growing in the location where the environmental factors have taken it is as important as the individual plant itself. All kinds of activities aimed at the artificial distribution of rare plants must be considered equal to the distributing of a non-indigenous plant.

There are no vast open communities in Estonia which could favour the distribution of introduced species, and the only natural communities on mineral soils are heaths, limestone shores and some coastal communities. The inclusion of new species is also slowed down by Estonia's location in the northern part of the temperate zone, where the growth of vegetation is relatively slow. Introduction, therefore, is generally not seen here as dangerous as it is in the warmer regions with paleoendemic vegetation, such as Australia, and Africa, where the species of local flora tend to suffer hard through the competition with introduced species.

According to evaluations by A. Paivel (1960), about 900 introduced species of woody plants have been cultivated in Estonia at some time. About half of these species are still growing in our gardens and parks. Of these 900 species, about 55% have shown good acclimatisation and 35% adequate acclimatisation, while 10% of the plants have not acclimatised (Paivel 1960a: 70). According to H. Sander (1998, unpublished data), the total number of introduced taxa in Estonian dendrological flora is about 1700, of which 1/3 are cultivars. There are forest cultivations with at least 40 introduced tree species in Estonia.

During the last years, the herbs causing problems include the extensively running wild and in some places naturalised plants Heracleum sosnowskyi and Galega orientalis. The introduced species often show certain negative characteristics; they may become garden and field weeds which are hard to eliminate, and although they cannot be considered as a direct danger to the natural species and communities, they are dangerous or productivity-decreasing plants from the viewpoint of agriculture. Of the naturalised aliens, the most dangerous one is Galinsoga cilata which is a common and difficult weed on the fields of southern Estonia. Veronica filiformis, a weed of garden lawns and rock gardens, escaped from a private collection in Jõgeva in the 1950s.

The data on naturalised plants, their situation, distribution and other characteristics must be collected into a Black Book. This Black Book could then also serve as a media for popularising the dangers of species introduction. The species of this book should be grouped according to their dangerousness, present and potential distribution.



Tables

Table 1. Conference excursions of Baltic botanists (between1954 and 1994)

Table 2. The number of species in Estonian vascular plant flora (according to the literature data, in chronological order)

Table 3. The number of indigenous, casual alien and running-wild taxa

Table 4. The composition of families by the sensibility to human impact the taxa

Table 5. The composition of Estonian native flora by floristic elements and their frequency

Table 6. The composition of Estonian alien flora by regions of origin

Table 7. The composition of Estonian indigenous flora by floristic elements and sensibility to human impact

Table 8. The significance of species at the margins of their area

Table 9. The taxa at the margins of their area by floristic elements. The percentage value after the quarter represents the share of the taxa reaching their area margin in this floristic element. The second to last line shows the share of the floristic element in the taxa reaching their area margin, and the last column shows the share of the taxa on their area margin in the total number of taxa in the floristic element.

Table 10. The taxa at the margins of their area by frequency. The percentage value indicates the share of a frequency class in the taxa at the margins of their area in a specific quarter.

Table 11. Sensibility to human impact and species at the margins of their area. The percentage value shows the share of the group in the taxa at the margins of their area in a specific quarter.

Table 12. Frequency and sensibility to human impact. The percentage value shows the share of a frequency group in a specific type of sensibility to human impact.

Table 13. Origin of naturalized species and their frequency in Estonia

Fig. 1. The number of species at the margins of their area