Ecology, chorology and commonness of the Trifolium taxa in Greece

Feddes Repertorium 121 (2010) 1–2, 66–80 DOI: 10.1002/fedr.200911129 Weinheim, Mai 2010

© 2010 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 0014-8962/10/1-205-0066

1 Technological Educational Institute of Kavala, Department of Forestry and Natural Environment, Drama, Greece 2 Technological Educational Institute of Larisa, School of Forestry and Management of Natural Environment, Karditsa, Greece

G. FOTIADIS1; M. S. VRAHNAKIS

2; TH. MEROU1 & K. VIDAKIS

1

Ecology, chorology and commonness of the Trifolium taxa in Greece

With one Map and 6 Figures

Abst rac t

The taxa of the genus Trifolium L. are consid-ered valuable genetic plant resources world-wide. The study of their chorological and flo-ristic characteristics is of great importance. This paper aims at evaluating the taxa of the genus found in the flora of Greece in terms of their ecological, chorological and commonness characteristics. A floristic catalogue was pro-duced based on published, but not genus-specific, floristic data collected at various parts of the country and field surveys carried out by the authors in years 2004 and 2005. Informa-tion on the localities of the taxa, either found in the field surveys or scattered into floristic re-ports, was gathered. The present analysis was based on several research results and biblio-graphic information. It was found that the Greek flora contains 113 taxa (95 species, 17 subspecies, one hybrid) out of a total of ap-proximately 255 species reported worldwide; six of them being Greek endemics. The taxa are distributed into two subgenera of the genus; subgenus Chronosemium including 14 species and subgenus Trifolium that includes 98 taxa (without counting one hybrid) distributed in five out of eight sections: Trifolium (56), Vesi-caria (18), Trifoliastrum (15), Trichocephalum (7) and Paramesus (2). The majority of the taxa are mediterranean elements (38%), annuals (76%), therophytes (76%), and of scapose growth type (78%). It was also found that the centre of distribution of Trifolium is located in Anatolia (Turkey) and Greece, so pointing out this part of east Mediterranean as an important

“gene-pool” region. The most common Tri-folium species in Greece are T. repens, T. campestre, T. fragiferum, T. physodes and T. arvense.

Introduction

The taxa of the genus Trifolium L. (clovers) are important floristic elements almost found in all natural ecosystems of the world. The knowl-edge of indigenous in each country Trifolium taxa is important not only for their rational exploitation in terms of rural economy but also for sustaining ecosystems’ integrity. These taxa are preeminent elements for atmospheric nitro-gen fixation, contribute to the improvement of nutritive value of forages for grazing animals, facilitate the root expansion of other species through their fibrous, tap-like roots, increase soil water retention capacity, and protect soils from mineral leaching (KNIGHT & WATSON 1977; TAYLOR 1985; LOI et al. 2000). Conse-quently, they constitute valuable natural re-sources, for which ecological, biotic and choro-logical attributes must be adequately defined prior to any exploitation or restoration effort. Also, many Trifolium taxa, especially those from the Mediterranean centre of diversity, are invasive species to other places of the globe, especially in the south hemisphere (e.g. in New Zealand, GRAVUER 2004; and in Australia, EMMS et al. 2005). The knowledge of their ecology and distribution may help improve their rational exploitation or in conducting studies for invasion prevention of the taxa.

G. FOTIADIS et al.: Ecology, chorology and commonness of Trifolium taxa 67

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The first historical, taxa-specific description of the genus Trifolium on a worldwide basis was carried out by ZOHARY & HELLER (1984) in their monumental book “The Genus Tri-folium”, while GILLETT & TAYLOR (2001) report the distributional characteristics of Tri-folium around the globe. The first, completed identification key, accompanied by European taxa description, was produced by TUTIN et al. (1968) in Flora Europaea (vol. 2). The first catalogue of the floristic elements of the Leguminosae family (Trifolium genus in-cluded) for the Mediterranean countries was presented in the Med-Checklist of GREUTER et al. (1989, vol. 4). According to that cata-logue, the flora of Greece contains 102 Tri-folium taxa. On the other hand, the Mountain Flora of Greece (STRID 1986, vol. 1) reports 26 taxa, but in altitudes over 1000 m. a.s.l. The aim of this study is to present choro-logical and ecological information and to infer about the commonness of the genus Trifolium in Greece by pointing out the great diversity of the Trifolium taxa. For this purpose, additional data obtained from posterior bibliographic and field work aiming at complementing the existing knowledge on Trifolium taxa found in Greece was used. A floristic catalogue is given by re-porting the floristic regions where these taxa are present, and their biotic and ecological charac-teristics. The proximity of Greece to the Tri-folium distributional centre is additionally dis-cussed. Finally, the ecological adaptation of Trifolium is investigated and the most widely adapted taxa are highlighted.

Method of research

Several floras and standard bibliographic sources were used to identify the Trifolium taxa found in Greece, namely: Flora Europaea, vol. 2 (TUTIN et al. 1968); Mountain Flora of Greece, vol. 1 (STRID 1986); Flora of Turkey and the East Aegean Islands, vol. 3 (DAVIS 1969); Med-Checklist, vol. 4 (GREU-TER et al. 1989), and information from the Med-Checklist, Notulae 16, 18, 20, 21, 23 (GREUTER & RAUS 1995, 2001a, b, 2002, 2005). In addition, research papers reporting the results of field surveys were reviewed, namely: ATHANASIADIS & THEO-DOROPOULOS (1990), ATHANASIADIS et al. (1993), ATHANASIADIS & DROSOS (1990), BABALONAS (1984, 1981), BERGMEIER & MATTHÄS (1995), BERGMEIER et al. (2001), CHRISTODOULAKIS et al.

(1990), GANIATSAS (1963), GEORGIADIS et al. (1990), DIMITRELLOS & CHRISTODOULAKIS (1995), DIMOPOULOS (1993), ELEFTHERIADOU (1992), FOTIADIS & ATHANASIADIS (2008), FOTIADIS et al. (2009), GREUTER et al. (1983), HANLIDOU & KOKKINI (1997), HANSEN & NIELSEN (1993), KARAGIANNAKIDOU & KOKKINI (1987), KARAGIAN-NAKIDOU & RAUS (2001), KARETSOS (2002), KAROUSOU et al. (2008), KOUMPLI-SOVANTZI (1983), MINOGLOU (2000), PANITSA (1997), PAPADIMITRIOU & ISPIKOUDIS (2008), PAPANICOLAOU (1985), PAVLIDIS (1976, 1982, 1985), RAUS (1996), SARI- KA et al. (2005), STRID (1978), TSIRIPIDIS & ATHA-NASIADIS (2003) and VLACHOS et al. (2002). Fur-thermore, field data were collected in years 2004 and 2005 in surveys carried out in several regions of Greece (Lakonia, Etoloakarnania, Thesprotia, Ioannina, Kiklades and Northern Greece). Collected plant material was taxonomically identified by using Flora Europaea vol. 2 (TUTIN et al. 1968), Mountain Flora of Greece vol. 1 (STRID 1986), Flora of Turkey and the East Aegean Islands vol. 3 (DAVIS 1969), Flora d’Italia (PIGNATTI 1982) and “The Genus Trifolium” (ZOHARY & HELLER 1984). Information for Trifolium taxa were sorted out and a floristic catalogue was build. The catalogue includes taxa allocated to subgenus and sections in alphabetical order according to ELLISON et al. (2006). Taxa nomenclature followed STRID (1986), GREUTER et al. (1989) and HENDRYCH (1993). The name of the authors of plant names are written ac-cording to BRUMMITT & POWELL (1992). Data analysis included the determination of the major a) ecological and b) chorological character-istics of the Trifolium taxa, while in addition c) commonness of each taxon was determined. Specifi-cally, ecological analysis included the classification of the taxa according to their (i) life duration, (ii) life form, (iii) growth type growth type according to RAUNKIAER’s (1910) system as modified by ELLEN-BERG (1956). Classification was based on GARCKE (1972), PIGNATTI (1982), DAVIS (1969), STRID (1986) and OBERDORFER (1990). Moreover, taxa were classified according to their (iv) altitudinal expansion range and (v) type of habitat. Altitudinal expansion range was determined as the difference between the lower and the highest altitude of taxon’s reference. For example, an altitudinal expansion range of 300 m for a given taxon may be resulted from a difference of 1500 m (highest altitude) minus 1200 m (lower altitude), while the same range for another taxon may derive from other altitudinal differences (e.g. 1000 m minus 700 m). Taxa were allocated into eight types of habitats according to their dominant appearance (grasslands, shrublands, open woodlands, forests, cultivated fields, roadsides, urban areas, and coasts). Chorological analysis was performed by means of the chorological spectrum

68 Feddes Repert., Weinheim 121 (2010) 1–2


following Oberdorfer’s system of geographical partition of Europe into major floristic units (OBER-DORFER 1990) in order to locate the distribution centre of each taxon and to define its level of endem-ism. Information for species distribution was drawn from GARCKE (1972), PIGNATTI (1982), DAVIS (1969), GREUTER et al. (1989), GREUTER & RAUS (1995, 2001a, b, 2002, 2005), STRID (1986) and OBERDORFER (1990). More over, chorological analy-sis included the allocation of taxa into the phyto-geographical areas of Greece as described by STRID & TAN (1997). Finally, results on the commonness of the Trifolium taxa were produced by graphically combining the relative location of each taxon into the two dimensional space of its altitudinal expansion rate with the number of phytogeographical areas of Greece where each taxon is found. Wide-spread Trifolium taxa were those that occur frequently in high numbers of phytogeographical areas and have wide altitudinal expansion rates. On the opposite side, narrow-spread taxa were those extending over a narrow area in terms of number of phytogeographi-cal areas and altitudinal expansion rate. The rest of the taxa were classified as having intermediate commonness.

Results and discussion

Genus Trifolium is one of the large genera in the legumes’ family with ca. 255 species (ZO-HARY & HELLER 1984; GILLET & TAYLOR 2001; ELLISON et al. 2006). Of these species, 95 are naturally found in Greece and, according to existing records, there are almost 113 Tri-folium taxa, including one hybrid (T. hybridum subsp. anatolicum × elegans). In the past, taxa nomenclature varied considerably; there were taxa with over 25 synonyms (e.g. T. arvense, T. pratense), while at least eight genera were used to describe Trifolium species (Amarenus, Amo-

ria, Calycomorphum, Chrysaspis, Falcatula, Micrantheum, Mistyllus, Paramesus) (ZOHARY & HELLER 1984). A recent study allocated two subgenera to the genus Trifolium (ELLISON et al. 2006); the subgenus Chronosemium com-prising approximately 20 Mediterranean spe-cies, 14 of them present in Greece, and the subgenus Trifolium comprising the remaining species, with 81 of them present in Greece. According to the same study, 81 species are distributed into five out of a total of eight sec-tions, namely Paramesus (2 species), Tricho-cephalum (4 species), Trifoliastrum (9 species), Trifolium (51 species) and Vesicaria (15 spe-cies). The other three sections, the Central-European section of Glycyrrhizum, the Euro-Siberian section of Lupinaster, and the Ameri-can section of Involucratum are not present in Greece.

Ecological characteristics

The life duration spectrum revealed that the majority of the taxa are annuals, followed by perennials, while one (T. velenovskyi) is bien-nial (Fig. 1). Therophytes dominate over hemicryptophytes and geophytes. The majority of the taxa is of the scapose growth type fol-lowed by caespitose, reptans, and rhizomatous. All reptans taxa are perennials, and all rhizo-matous taxa are geophytes. The majority of species are found in areas of low altitude (below 1000 m), while the number of species reduces with increasing altitude; only four species are recorded in altitudes over 2500 m (T. pratense, T. repens, T. parnassi, T. pallescens) (Fig. 2).

Fig. 1 Life duration (a), life form (b) and growth type (c) spectra of Trifolium species in Greece (n = 95) A – annuals; B – biennials; P – perennials; T – therophytes; H – hemicryptophytes; G – geophytes; Scap – scapose; caesp – caespitose; rept – reptans: rhiz – rhizomatous



Fig. 2 Altitudinal distribution of Trifolium species (n = 95) 31 species out of 95 are found in closed or open forest communities (Fig. 3), of which only five (T. ochroleucon, T. heldreichianum, T. pseudomedium, T. diffusum, T. pignantii) are distributed in a wide altitudinal range [(s.l.–)300–1800(–2300) m]. Phytosociological studies have shown that six perennial taxa (T. pseudomedium, T. patulum, T. alpestre, T. ochroleucum, T. physodes, T. medium subsp. medium) are typical elements of Mediterranean forests (and shrublands) (MUCINA 1997; TSIRIPIDIS 2001; FOTIADIS & ATHANASIADIS 2008). Μany typical forest taxa are perennials (e.g. T. ochroleucon, T. pannonicum, T. held-reichianum, T. patulum, T. pignantii, T. pseu-domedium, T. hybridum subsp. elegans) in contrast to the dominant annual life duration pattern of the whole genus. Consequently, these species have the potential to be introduced into

forests or silvopastoral systems for improving their forage quality potential. The majority of taxa (48) are found in open places; 16 of them in an altitude of 0–1000 m. Ten species are commonly found in a variety of habitats (more than six habitats).

Chorological characteristics and endemism

Chorological spectrum revealed the dominance of Mediterranean elements, followed from sub-Mediterranean, and Balkanic/sub-Balcanic (Fig. 4). There are six endemic taxa [T. praetermissum, T. aurantiacum (uncertain status), T. dolopium, T. parnassi, T. barbeyi, T. ottonis], mostly found on limestone, on is-lands and sub-alpine grasslands. The number of Eurasiatic elements is small (seven), with one

Fig. 3 Habitats of Trifolium species(n = 95)



(T. spadiceum) northern, and three (T. badium, T. pallescens, T. noricum subsp. praetutianum) alpine. The presence of T. glomeratum is im-pressive, since apart from the Mediterranean basin it is also found in South Africa and Chile (ZOHARY & HELLER 1984). Chorologically, the east Mediterranean taxa (omed/osmed) (20 in number) dominate over west Mediterra-nean/Atlantic (wmed/atl/subatl) ones (five in number) (Fig. 5). This is in accordance with ZOHARY & HELLER (1984) who argued that Anatolia (Asia Minor) is the preponderant centre of distribution for Trifolium. There is strong evidence that Anatolia holds the greater number of endemic Trifolium species (eight in number) followed by Greece (six) and Syria (three) (DAVIS 1969; JORDANOV et al. 1976; PIGNATTI 1982; ZOHARY & HELLER 1984; GREUTER et al. 1989; GREUTER & RAUS 1995, 2001a, b, 2002, 2005). It seems that a great number of endemic species is accumulated between the Balkans and Anatolia, since the

Fig. 5 Percentage distribution of west- and east-Mediterranean elements (n = 25) wmed – west- Mediterranean, atl – atlantic, omed – east- Mediterranean, osmed – east-sub- Mediterranean

endemics of a) the east Mediterranean (Anato-lia, Cyprus, Syria, Israel), b) the Balkans and Anatolia, and c) the Italian peninsula and the Balkans are proportionally distributed accor-ding to the rate 29:33:27. In addition, the maximum number of Trifolium species is found in the Balkans and Anatolia area, since the species proportion of West Mediterranean Europe (Italy included): Balkans:Asian Mediterranean:African Mediterranean is 77:116:119:59, while the species proportion of Balkans:Anatolia :Asian Mediterranean is 116:105:73. These numbers and proportions clearly indicate that the distribution centre of Trifolium is in Anatolia, but slightly shifted to the Balkans, and more specifically to Greece. This is further confirmed by examining the chorological distribution of Trifolium in the Greek territory (Map 1); north-east Greece (NE) hosts the highest number of species, while the region of the east Aegean islands (EAe) is found in the fourth position.

Commonness

Most of the species are present in only one phytogeographical region of Greece (Fig. 6), while five species (T. campestre, T. repens, T. arvense, T. fragiferum, T. physodes) are present almost in the whole country. Most of the species appear in an altitudinal expansion range of 1000 m, while those recorded in a range over 2000 m are 15 (T. tomentosum, T. physodes, T. fragiferum, T. pratense, T. pignantii, T. heldreichianum, T. ochroleucon, T. arvense, T. alpestre, T. repens, T. hybridum,

Fig. 4 Chorological spectrum of Trifolium taxafound in Greece (n = 95) med – Mediterranean; smed – sub-Mediterranean; euras – eurasiatic; kont –continental; no – northern; alp – alpine; balc-subalc – balkanic/sub-balkanic; hell –Greece endemics

G. FOTIADIS et al.: Seed morphology, chorology and commonness of Trifolium taxa 71


Map 1 Distribution of Trifolium species in the 13 phytogeographical regions of Greece KK – Kriti; Pe – Peloponnisos; StE – Sterea Ellada; SPi – south Pindos; NPi – north Pindos; EC – east central; NC – north central; NE – north east; Iol – Ionian islands; WaE – West Aegean islands; NAe – North Aegean islands; EAe – east Aegean islands; Kik – Kiklades Phytogeographical regions of Greece according to STRID & TAN (1997)

T. uniflorum, T. patens, T. campestre, T. aureum); of those, T. pignantii and T. held-reichianum have wide altidutinal distributions but narrow chorological ones (Fig. 6). The latter are species mostly found under forest canopies where the climatic conditions are rather stable and mild, a fact that propably determines their large altidutinal expansion range. By combining (with graphical means) the relative location of each Trifolium species into

the two dimensional space of altitudinal expan-sion rate with the number of phytogeographical areas of Greece where each species is found, three distinct groups of Trifolium species are produced according to their commonness (Fig. 6; groups separated by gray line). The first group includes the narrow-spread Trifoli-um species, i.e. those extending over a narrow area in terms of number of phytogeographical areas and altitudinal expansion rates. There are 29 narrow-spread species; four of them



Fig. 6 Number of phytogeographicalareas (1–13) and altitudinalexpansion range (1–27 viz. from100 to 2700 m) where Trifoliumspecies are distributed



(T. cinctum, T. obscurum, T. pilczii, T. scu-tatum) appear in a single phytogeographical region and up to 100 m of altitudinal expansion range, while the rest are found up to three phy-togeographical regions and up to 1200 m of altitudinal expansion range. Four out of six Greek endemic species (T. praetermissum, T. dolopium, T. barbeyi, T. ottonis) are included in this group. On the contrary, 15 species are found in more than seven phytogeographical regions and in altitudinal expansion range over 1400 m, thus belonging to the wide-spread group. Some characteristic wide-spread taxa are T. arvense, T. physodes, T. fragiferum, T. campestre and T. repens found in at least 12 phytogeographical regions and in altitudinal expansion ranges of over 2000 m. The wide commonness of the perennial T. repens and other species justify and support their cultiva-tion in many areas of the globe (BAKER & WILLIAMS 1987; TAYLOR & QUESENBERRY 1996), while in other areas of the world they are the most important in terms of invasion, like in New Zealand (GRAVUER 2004). The rest of the species were classified as having inter-mediate commonness.

Conclusions

The present research maintains that the current number of Trifolium taxa found in Greece is 113. They are distributed into two subgenera of the genus; subgenus Chronosemium includes 14 species and subgenus Trifolium includes 98 taxa (without counting one hybrid); the latter are further distributed in five out of eight sections: Trifolium (56), Vesicaria (18), Trifolias-trum (15), Trichocephalum (7) and Paramesus (2). The majority of the taxa are Mediterranean elements (38%), annuals (76%), therophytes (76%), and of scapose growth type (78%). Six species are Greek endemics [T. praetermissum, T. aurantiacum (uncertain status), T. dolopium, T. parnassi, T. barbeyi, T. ottonis]. The greatest number of taxa is found in north-east Greece (NE) and the endemic elements found in Greece indicate that the distribution centre of Trifolium is in Anatolia, but slightly shifted to the Bal-kans, and more specifically to Greece. The most common Trifolium species in Greece are T. repens, T. campestre, T. fragiferum, T. phy-

sodes and T. arvense. Trifolium species are fur-ther classified as those having wide common-ness (29 species), narrow commonness (15 species), and intermediate commonness (the remaining 51 species).

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Adresses of the authors

G. Fot iad is , Th. Merou, K. Vidakis , Techno-logical Educational Institute of Kavala, Department of Forestry and Natural Environment Management, 66100, Drama, Greece; e-mail: [email protected]

M. S. Vrahnakis , Technological Educational Institute of Larisa, School of Forestry and Manage-ment of Natural Environment, Annex of Karditsa, 43100, Karditsa, Greece. E-mail: [email protected]

Manuscript received: November 26th, 2009.

Αnnex

Floristic catalogue

Subgenus Chronosemium

Trifolium aurantiacum BOISS. & SPRUNER: A/ Tscap/ hell (?)/ 200–1900 m/ Pe, StE, NPi, NC, NE, WAe, EAe/ shrublands, forests and rocky places. – Even though ELLISON et al. (2006) did not include the species in the subgenus Chronosemium, according to ZOHARY & HELLER (1984) the species belongs to section Chronosemium which became a subgenus. According to the Med-Checklist (GREUTER et al. 1989), it is a Greek endemic and it is not detectable in the East Aegean islands. Pro-pably, it is misidentified or confused with other Trifolium taxa (T. boissieri, T. grandiflorum or even T. campestre). According to LASSEN (GREUTER & RAUS 1999), the single record from the Cretan area (KK) is in fact, a misiden-tified specimen of T. campestre. Trifolium aureum POLLICH: A(Β)/ T(Η)scap/ gemabkont(-smed)/ s.l.–2000 m/ Pe, StE, SPi, EC, NC, NE, Kik/ fields, grasslands, shrublands, open woodlands, forests, coasts, roadsides, wet places and rocky places. Trifolium badium SCHREBER: P/ Hscap/ alp/ 700–2100 m/ NPi, NC/ grasslands, open Fagus forests and wet places but mostly in subalpine calcareous grasslands. Trifolium boissieri GUSS.: A/ Tscap/ omed/ s.l.–1250 m/ KK, Pe, StE, NE, WAe, EAe, Kik/



olive fields, shrublands (mainly maquis and phrygana), forests and rocky places. Trifolium campestre SCHREBER: A/ Tscap/ smed-subatl/ s.l.–2200 m/ KK, Pe, StE, SPi, NPi, EC, NC, NE, IoI, WAe, NAe, EAe, Kik/ almost everywhere, but mostly in grass-lands. Trifolium dolopium GIBELLI & BELLI: A/ Tscap/ hell/ ~300 m/ NE/ slopes. Stenotopic endemic restricted in an area of the Athos pen-insula. – Even though ELLISON et al. (2006) did not include the species in the subgenus Chronosemium, according to ZOHARY & HELLER (1984) the species belongs to section Chronosemium which became a subgenus. Trifolium dubium SIBTH.: A/ Tscap/ subatl(-smed)/ 100–1400 m/ EC, NC, NE/ grasslands, urban areas, roadsides and sandy places. Trifolium filiforme L.: A/ Tscap/ med-atl/ s.l.–1800 m/ KK, SPi, EC, NC, NE, EAe, Kik/ fields, Fagus forests, wet, damp places and sandy places. – Even though ELLISON et al. (2006) did not include the species in the subge-nus Chronosemium, according to ZOHARY & HELLER (1984) the species belongs to section Chronosemium which became a subgenus. Trifolium grandiflorum SCHREBER: A/ Tscap/ omed/ s.l.–1600 m/ KK, Pe, StE, SPi, NPi, EC, NC, NE, WAe, EAe, Kik/ shrublands, forests, urban areas and rocky places. Trifolium patens SCHREBER: A/ Trept/ smed-med/ s.l.–2200 m/ KK, Pe, StE, SPi, EC, NC, NE, IoI, WAe, EAe/ fields, grasslands, shrublands, open woodlands, forests, roadsides, wet and rocky places but mostly in nutrient-rich grasslands. Trifolium praetermissum GREUTER et al.: P(A)/ H(T)scap/ hell/ 100–1000 m/ KK/ calca-reous soils. Stenotopic endemic restricted in two areas east of Iraklio (islands of Crete and Karpathos). Trifolium sebastianii SAVI: A/ Tscap/ os-med/ 300–1300 m/ NPi, NE/ open woodlands. – Even though ELLISON et al. (2006) did not include the species in the subgenus Chrono-semium, according to ZOHARY & HELLER (1984) the species belongs to section Chrono-semium which became a subgenus. Trifolium spadiceum L.: A(P)/ T(H)scap/ nokont-pralp/ 1300–1600 m/ NC, NE/ open woodlands and forests.

Trifolium velenovskyi VANDAS: Β/ Hscap/ balc (Al-Ju-Bu)/ 900–1900 m/ StE, NC, NE/ grasslands, shrublands, open woodlands and forests. – Even though ELLISON et al. (2006) did not include the species in the subgenus Chronosemium, according to ZOHARY & HELLER (1984) the species belongs to section Chronosemium which became a subgenus. The species has also been recorded on the island of Zakinthos (IoI), but its occurrence needs con-firmation.

Subgenus Trifolium

Section Paramesus

Trifolium glanduliferum BOISS.: A/ Tscap/ omed/ 600–1000 m/ EAe/ fields, grasslands, shrublands (mainly maquis) and Pinus forests. Trifolium strictum L.: A/ Tscap/ smed/ s.l.–1550 m/ StE, SPi, EC, NE/ grasslands, open woodlands.

Section Τrichocephalum

Trifolium globosum L.: A/ Tscap(rept)/ omed/ grasslands, amongst shrubs and sandy places/ s.l.–900 m. StE, NE, WAe, EAe, Kik. Trifolium pauciflorum DUM.-URVILLE: A/ Tscap/ omed/ s.l.–1200 m/ EAe/ fields, grass-lands and stony calcareous places. Trifolium pilulare BOISS.: A/ Tscap/ οmed/ s.l.–800 m/ EAe/ grasslands. Trifolium subterraneum L.: A/ Trept/ smed/ s.l.–1250 m/ KK, Pe, StE, SPi, EC, NC, NE, IoI, WAe, NAe, EAe, Kik/ fields, grass-lands, shrublands, open woodlands, urban areas and wet places. Three subspecies recognized in Greece: a. the Mediterranean subsp. oxaloides NYMAN, b. the submediterranean subsp. subter-raneum, and c. the balkan endemic subsp. yan-ninicum KATZN. & F.MORLEY. GREUTER et al. (1989) do not mention the existance of the subspecies subterraneum in the Mediterranean region, even though, according to ZOHARY & HELLER (1984), the subspecies is native only in Greece.

Section Trifoliastrum

Trifolium retusum L.: A/ Tscap/ euras/ 100–1100 m/ NC, NE/ grasslands, shrublands, for-ests, coasts and roadsides but mostly in inland salt-marches, salt-pans and salt-steppes. – The



species not mentioned for Greece by GREUTER et al. (1989), although ZOHARY & HELLER (1984) mentions the existance of the species in Greece and RAUS et al. (GREUTER et al. 2002) give the information above cited. Trifolium suffocatum L.: A/ Tscap/ med/ s.l.–1500 m/ KK, StE, EC, NE, WAe, EAe, Kik/ grasslands and waste places but mostly in short-lived, trampled ruderal plant communities. Trifolium nigrescens VIV.: A/ Tscap/ med-smed/ s.l.–1800 m/ KK, Pe, StE, SPi, EC, NC, NE, WAe, NAe, EAe, Kik/ fields, grasslands, shrublands, roadsides, wet and waste places, sandy places and rocky places but mostly in annual, herb-rich ruderal plant communities. Two subspecies recognized in Greece: subsp. nigrescens and subsp. petrisavii (G.C.CLEMENTI) HOLMBOE. Trifolium pilczii ADAMOVIĆ: P/ Hcaesp/ balc (Al-Ju)/ 2100–2200 m/ NP/ calcareous grasslands. Trifolium uniflorum L.: P/ Hcaesp/ med/ s.l. – 2400 m/ KK, Pe, StE, EC, NE, WAe, NAe, EAe, Kik/ fields, grasslands, shrublands (mainly maquis and phrygana), coasts, urban areas, roadsides and rocky places. Trifolium repens L.: P/ H(Ch)rept/ euras-subozean-smed/ s.l.–2700 m/ KK, Pe, StE, SPi, NPi, EC, NC, NE, IoI, WAe, NAe, EAe/ al-most everywhere but mostly in nutrient-rich and wet grasslands. Four subspecies recognized in Greece: a. the continental subsp. orbelicum (VELEN.) PAWL., b. the mediterranean subsp. orphanideum (BOISS.) D.E.COOMBE, c. the sub-mediterranean subsp. prostratum NYMAN, and d. the eurasiatic subsp. repens. – GREUTER et al. (1989) do not mention the existance of the subspecies repens in the Mediterranean region, even though, according to ZOHARY & HELLER (1984), the subspecies is native in the Medi- terranean countries. Trifolium pallescens SCHREBER: P/ Ηcaesp/ alp-pralp/ 1600–2800 m/ NC, NE/ grasslands but mostly in screes, gravel river banks and related habitats. – According to STRID & PAPANICOLAOU (1981), the occur-rence of the species in Vermio and Voras (NC) needs confirmation, but according to STRID (1986) the material from Tzena (NC) looks like the material from Olympus (NC) of T. palles-cens and that of Voras resembles that from the Alps of T. pallescens.

Trifolium glomeratum L.: A/ Tscap/ smed (also in S. Africa and Chile)/ s.l. – 1500 m/ KK, Pe, StE, SPi, EC, NC, NE, WAe, EAe, Kik/ fallow fields, grasslands, shrublands, for-ests, urban areas, roadsides, riverbanks and rocky places but mostly in annual herb-rich ruderal communities. Trifolium parnassi BOISS. & SPRUNER: P/ Hcaesp/ hell/ 1500–2700 m/ Pe, StE, SPi, NPi, NC/ grasslands and rocky places but mostly in siliceous alpine and boreal grasslands.

Section Τrifolium

Trifolium affine C.PRESL: A/ Tscap/ balc (bu-tu)-an/ s.l.–300 m/ NE, NAe, EAe/ grasslands and dry places. Trifolium alpestre L.: P/ Hscap/ ge-mabkont-smed/ 200–2300 m/ Pe, StE, SPi, NPi, EC, NC, NE, NAe/ grasslands, shrublands, open woodlands, forests and roadside but mostly in herb-rich heliophilous and (sub)-thermophilous fringe plant communities of temperate forests. Trifolium angustifolium L.: A/ Tscap/ smed-subatl/ s.l.–1100 m/ KK, Pe, StE, SPi, NPi, EC, NC, NE, IoI, WAe, NAe, EAe, Kik/ fields, grasslands, amongst scrubs, open wood-lands, urban areas and rocky places but mostly in terrestrial plant communities dominated by annual low-growing herbs and grasses. Trifolium apertum BORDOV: A/ Tscap/ osmed/ s.l.–1000 m/ IoI/ shrublands, meadows and clay hills. Trifolium arvense L.: A/ Tscap/ euras-subozean-smed/ s.l.–2000 m/ KK, Pe, StE, SPi, NPi, EC, NC, NE, IoI, NAe, WAe, EAe, Kik/ fields, grasslands, open woodlands, urban ar-eas, roadsides and rocky places, but mostly in grasslands and related short-lived, herb-rich plant communities of sandy and skeletal, pre-dominantly nutrient-poor and drought-stressed soils. Trifolium barbeyi GIBELLI & BELLI: A/ Tscap/ hell/ s.l.–500 m/ KK/ dry grasslands. Trifolium bocconei SAVI: Α/ Tscap/ med/ s.l.–800 m/ KK, Pe, NC, NE, EAe, Kik/ grass-lands, amongst scrubs and roadsides. Trifolium caudatum BOISS.: P/ Hcaesp/ hell-an/ ~400m/ EAe/ pastures. – Even though ELLI-SON et al. (2006) did nοt include the species in the section Trifolium, according to ZOHARY &



HELLER (1984) the species belongs to section Trifolium. Trifolium cherleri L.: A/ Tscap/ smed/ s.l.–1800 m/ KK, Pe, StE, EC, NC, NE, WAe, EAe, Kik/ grasslands, shrublands, urban areas and rocky places but mostly in terrestrial plant communities dominated by annual low-growing herbs and grasses. Trifolium cinctum DC.: A/ Tscap/ balc (Al-Ju)/ ~90 m/ IoI/ wet grasslands. – Even though ELLISON et al. (2006) did nοt include the spe-cies in the section Trifolium, according to ZO-HARY & HELLER (1984) the species belongs to section Trifolium. Trifolium clypeatum L.: A/ Tscap/ omed/ s.l.–700 m/ KK, Pe, EAe, Kik/ fields and road-sides. Trifolium congestum GUSS.: A/ Tscap/ med/ s.l.–600 m/ EAe/ dump open places. – Even though ELLISON et al. (2006) did nοt include the species in the section Trifolium, according to ZOHARY & HELLER (1984) the species be-longs to section Trifolium. Trifolium constantinopolitanum SER.: A/ Tscap/ οmed/ s.l.–700 m/ NE/ fields, roadsides, river banks and damp places. Trifolium dalmaticum VIS.: A/ Tscap/ balc (Al-Ju-Bu)/ 100–1600 m/ Pe, StE, SPi, NPi, EC, NC, NE, IoI/ grasslands, shrublands, open woodlands, forests and disturbed places but mostly in terrestrial plant communities domi-nated by annual low-growing herbs and grasses. Trifolium dasyurum C.PRESL: A/ Tscap/ οmed/ s.l.–1300 m/ KK, Pe, StE, IoI, EAe, Kik/ fields, grasslands, stony and calcareous soils. Trifolium desvauxii BOISS. & BLANCHE: A/ Tscap/ omed/ s.l.–100 m/ EAe. – Even though ELLISON et al. (2006) did nοt include the spe-cies in the section Trifolium, according to ZO-HARY & HELLER (1984) the species belongs to section Trifolium. Trifolium diffusum EHRH.: Α/ Tscap/ smed/ 200–1800 m/ StE, SPi, NC, NE/ shrub-lands (mainly maquis), open woodlands and forests. Trifolium echinatum M.BIEB.: A/ Tscap/ osmed/ 100–750 m/ NPi, EC, NC, NE, WAe, EAe/ open woodlands, urban areas and sandy places. Trifolium heldreichianum (GIBELLI & BELLI) HAUSSKN.: P/ Hcaesp/ balc (Al-Ju-Bu)-

Αn/ 100–2100 m/ StE, SPi, NPi, NC, NE/ grasslands, shrublands, open woodlands and forests. Trifolium hirtum ALL.: Α(B)/ Tscap/ smed/ s.l.–1400 m/ KK, Pe, StE, SPi, NPi, EC, NC, NE, WAe, NAe, EAe, Kik/ grasslands, shrub-lands, open woodlands, forests, urban areas, roadsides. Trifolium incarnatum L.: A(B)/ Tscap-(Hscap)/ med-atl/ 500–1100 m/ KK, Pe, NPi, NE, WAe/ fields, grasslands, open woodlands, forests, roadsides and rocky places. Two subspecies recognized in Greece: a. subsp. incarnatum – GREUTER et al. (1989) do not mention the existance of this subspecies in the Mediterranean region, even though, according to ZOHARY & HELLER (1984), the subspecies (as var. incarnatum) is widely cultivated and naturalized in the Mediterranean region – and b. subsp. molinerii (HORNEM.) SYME – Accor-ding to GREUTER et al. (1989), the occurrence of the subspecies in Greece needs confirmation. According to LASSEN (GREUTER & RAUS 1999), the only record of the subspecies from the EAe (island of Rhodos) is based on mixed material of T. infamia-ponertii and T. stellatum. Trifolium infamia-ponertii GREUTER: Α/ Tscap/ med-omed/ s.l.–1150 m/ KK, Pe, StE, WAe, EAe, Kik/ grasslands and mostly in ter-restrial plant communities dominated by annual low-growing herbs and grasses. – Even though ELLISON et al. (2006) did nοt include the spe-cies in the section Trifolium, according to ZO-HARY & HELLER (1984) the species belongs to section Trifolium. Trifolium lappaceum L.: A/ Tscap/ smed-med/ s.l.–1100 m/ KK, Pe, StE, EC, NC, NE, IoI, WAe, EAe, Kik/ fields, grasslands, amongst scrub, roadsides, rocky and wet places. Trifolium latinum SEBASTIANI: A/ Tscap/ balc (Bu)–It-an/ 100–600 m/ Pe, StE, SPi, EC/ shrublands (mainly maquis) and forests. Trifolium leucanthum M.BIEB.: A/ Tscap/ smed/ 200–1600 m/ KK, StE, NC, NE, EAe/ grasslands, shrublands (mainly phrygana), open woodlands and mountain slopes. Trifolium ligusticum LOISEL.: A/ Tscap/ wmed/ s.l.–800 m/ KK, Pe, WAe/ fields and rocky places. Trifolium lucanicum GUSS.: A/ Tscap/ med-smed/ 50–1600 m/ KK, Pe, StE, NE/ dry



and rocky places, but mostly in terrestrial plant communities dominated by annual low-growing herbs and grasses of Thero-Brachypodietea. Trifolium medium L. subsp. medium: P/ Grhiz/ eurossubozean (-smed)/ 600–1500 m/ NPi, NC, NE/ forests and shrublands, but mostly in herb-rich heliophilous and (sub)-thermophilous fringe communities of temperate woodlands. – Despite the several records of the subspecies, GREUTER et al. (1989) do not men-tion its existence in the Mediterranean region. Additionally, according to ZOHARY & HELLER (1984), the subspecies is native in the Mediter-ranean countries but they do not mention Gree-ce. Trifolium noricum subsp. praetutianum (SAVI) ARCANG.: P/ Hcaesp/ oalp/ 1400–2400 m/ Pe, StE, SPi, NPi, NC, NE/ rocky places but mostly in subalpine calcareous grasslands. Trifolium obscurum SAVI: A/ Tscap/wmed/ ~40 m/ EAe/ sandy soils damp in winter. Trifolium ochroleucon HUDS.: P/ Hcaesp/ smed-subatl/ 50–2100 m/ Pe, StE, EC, SPi, NPi, NC, NE, WAe, EAe/ grasslands, shrub-lands, open woodlands, forests and roadsides but mostly in steppes, rocky steppes and conti-nental sandy grasslands of Festuco-Brometea. Trifolium ottonis BOISS.: P/ Hscap/ hell/ 2050–2350 m/ Pe, StE/ grasslands. – Even though ELLISON et al. (2006) did nοt include the species in the section Trifolium, according to ZOHARY & HELLER (1984) the species be-longs to section Trifolium. Trifolium pallidum WALDST. & KIT.: A-B/ T(H)scap/ med-smed/ s.l.–900 m/ KK, Pe, StE, EC, NC, NE, WAe, EAe, Kik/ grasslands, shrublands (mainly phrygana), forests, road-sides and rocky places. Trifolium pamphylicum BOISS. & HELDR.: A/ Tscap/ omed/ s.l.–100 m/ KK/ grasslands and amongst scrubs. – Even though ELLISON et al. (2006) did nοt include the species in the section Trifolium, according to ZOHARY & HELLER (1984) the species belongs to section Trifolium. Trifolium pannonicum JACQ. subsp. pan-nonicum: P/ Hcaesp/ οsmed/ 1000–2000 m/ StE, NC/ open woodlands and wet places. Trifolium patulum TAUSCH: P/ Hcaesp/ balc (Al-Ju)-Ιt/ 450–1500 m/ Pe, StE, SPi, NPi, NC, NE, WAe/ shrublands and forests.

Trifolium phleoides WILLD.: A/ Tscap/ smed/ 200–1600 m/ StE, SPi, NC, NE/ fields, grasslands, open woodlands and rivulets. Trifolium pignantii FAUCHÉ & CHAUB.: P/ Grhiz/ balc (Al-Ju-Bu)/ 50–2300 m/ Pe, StE, SPi, NPi, EC, NC, NE, WAe/ grasslands and mostly forests. Trifolium pratense L.: P/ Hscap/ euras-subozean(-smed)/ s.l.–2500 m/ Pe, StE, SPi, EC, NC, NE, IoI, NAe, WAe/ almost every-where but mostly in nutrient-rich and wet grasslands. Trifolium pseudomedium HAUSSKN.: P/ Grhiz/ balc (Ju-Bu)/ 100–1800 m/ StE, SPi, NPi, EC, NC, NE/ grasslands, shrublands, open woodlands and mostly forests. Trifolium purpureum LOISEL.: A/ Tscap/ med/ s.l.–1200 m/ StE, NPi, EC, NC, NE, NAe, EAe, Kik/ fields, grasslands, urban areas, road-sides and limestones slopes. Trifolium. scabrum L.: A/ Trept-scap/ med-smed (-subatl)/ s.l.–1600 m/ KK, Pe, StE, EC, NC, NE, WAe, EAe, Kik/ fields, grass-lands, shrublands (mainly phrygana and ma-quis), coasts, urban areas, roadsides, rocky, calcareous, sandy places but mostly in terres-trial plant communities dominated by annual low-growing herbs and grasses and in grass-lands and related short-lived, herb-rich plant communities of sandy and skeletal, predomi-nantly nutrient-poor and drought-stressed soils. Trifolium scutatum BOISS.: A/ Tscap/ οmed/ s.l.–500m/ EAe/ grasslands and shrub-lands. Trifolium squamosum L.: A/ Tscap/ ge-mabkont-smed/ s.l.–1000 m/ KK, IoI, Pe, EC, NC, EAe/ fields, grasslands and forests. Trifolium squarrosum L.: A/ Tscap/ smed/ s.l.–600 m/ NE, IoI/ fields, grasslands and urban areas. Trifolium stellatum L.: A/ Tscap/ med-smed/ s.l.–1500 m/ KK, Pe, StE, EC, NC, NE, IoI, WAe, EAe, Kik/ fields, grasslands, shrub-lands, open woodlands, urban areas, roadsides and rocky places but mostly in terrestrial plant communities dominated by annual low-growing herbs and grasses. Trifolium striatum L.: A(B)/ T(H)scap/ subatl-smed/ 400–1600 m/ Pe, StE, SPi, NC, NE/ grasslands, shrublands, forests, roadsides and disturbed places but mostly in annual,



herb-rich ruderal communities of Stellarietea mediae. Trifolium sylvaticum GÉRARD: A/ Tscap/ med/ 200–1600 m/ KK, StE, NE, EAe/ hillsides and montane valleys. Trifolium tenuifolium TEN.: A/ Tscap/ balc (Al-Ju-Bu-Tu)-Ιt-Αn/ s.l.–1100 m/ KK, Pe, StE, EC, NC, NE, WAe, Kik/ grasslands, shrublands, forests, dry, rocky places, valleys and hillsides. – Even though ELLISON et al. (2006) did nοt include the species in the section Trifolium, according to ZOHARY & HELLER (1984) the species belongs to section Trifolium. Trifolium trichopterum PANČIĆ: A/ Tscap/ balc (Al-Ju-Bu-Tu)/ 1100–1400 m/ StE, NC, NE/ forests, dry and stony hillsides. Trifolium xanthinum FREYN: Α/ Tscap/ hell-an/ 1000–1500 m/ Pe, StE/ ?. – Even though ELLISON et al. (2006) did nοt include the species in the section Trifolium, according to ZOHARY & HELLER (1984) the species be-longs to section Trifolium.

Section Vesicastrum

Trifolium argutum BANKS & SOL.: A/ Tscap/ omed/ 200–500 m/ EAe, Kik/ fields, shrublands and roadsides. Trifolium balansae BOISS.: A/ Tscap/ balc(Ju-Bu-Tu)-An/ ?/ StE, EAe, Kik/ fields. Trifolium michelianum SAVI: A/ Tscap/ wmed/ s.l.–1800 m/ KK, Pe, StE, EC, NC, EAe/ grasslands, sandy and damp places. – According to LASSEN (GREUTER & RAUS 1999), the occurrence of species in Crete (KK) needs confirmation. Trifolium mutabile PORT.: A/ Tscap/ med/ s.l.–600 m/ Pe, StE, NC, NE/ field and grass-lands. Trifolium ornithopodioides L.: A/ Tscap/ smed-subatl/ s.l.–600 m/ KK/ slopes. Trifolium setiferum ΒOISS.: A/ Tscap/ balc (Al-Ju-Bu-Tu)-It-An/ 100–600 m/ NE/ amongst scrubs. Trifolium spumosum L.: A/ Tscap/ med/ s.l.–1000 m/ KK, Pe, StE, NE, WAe, EAe, Kik/ fields, grasslands, roadsides and damp places. Trifolium vesiculosum SAVI: A/ Tscap/ smed/ s.l.–700 m/ KK, Pe, StE, SPi, EC, NC,

NE, IoI/ grasslands, shrublands and open woodlands. Trifolium clusii GORDON & GREN.: A/ Tscap/ med/ ?/ EAe, Kik/ mostly in annual, herb-rich ruderal communities. Trifolium fragiferum L.: P/ Hrept/ med-smed(-gemabkont)/ s.l.–2200 m/ KK, Pe, StE, EC, SPi, NPi, NC, NE, IoI, WAe, NAe, EAe, Kik/ grasslands, open woodlands, urban areas, damp open and sandy places. Trifolium hybridum L.: P/ Hcaesp/ ge-mabkont(-osmed)/ s.l.–2400 m/ KK, Pe, StE, SPi, NPi, EC, NC, NE/ roadsides, grasslands, open woodlands, urban areas, roadsides and wet places but mostly in nutrient-rich grass-lands. Three subspecies recognized in Greece: a. the balkan endemic subsp. anatolicum (BOISS.) HOSSAIN, b. the Mediterranean subsp. elegans (SAVI) ASCH. & GRAEBN., and c. the Mediterranean subsp. hybridum – GREUTER et al. (1989) do not mention the existence of the subspecies in the Mediterranean region, even though, according to ZOHARY & HELLER (1984), the subspecies is native only in the Mediterranean countries. Additionally, STRID (1986) found in SPi and NPi a hybrid of subsp. anatolicum × subsp. elegans. Trifolium physodes M.BIEB.: P/ Hscap/ omed/ s.l.–2100 m/ KK, Pe, StE, SPi, NPi, EC, NC, NE, IoI, WAe, NAe, EAe, Kik/ grasslands, shrublands, open woodlands, forests, coasts and wet meadows but mostly in annual, herb-rich ruderal plant communities. Trifolium resupinatum L.: A(B)/ T(H)rept/ med-smed/ s.l.–1500 m/ KK, Pe, StE, SPi, EC, NC, NE, IoI, WAe, EAe, Kik/ fields, grass-lands, open woodlands, roadsides, wet and disturbed places but mostly in nutrient-rich and wet grasslands. Trifolium tomentosum L.: A/ Trept/ smed/ s.l.–1300 (2200) m/ KK, Pe, StE, EC, NE, IoI, WAe, EAe, Kik/ fields, grasslands, shrublands, urban areas and roadsides but mostly in annual, herb-rich ruderal plant communities. Trifolium bullatum BOISS. & HAUSSKN.: A/ Tscap/ omed/ s.l.– 200 m/ EAe/ in a small rocky island.

Funded by The present work is part of the research project “Impact of Climate on the Ecophysiology, SeedDormancy and Viability of Self-Reseeding Legumes” (2005–2007), co-financed by the European

Ecology, chorology and commonness of the Trifolium taxa in Greece

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