Introduction
SET IN THE NORTH-WESTERN CORNER OF GREECE – where three countries, Greece, Albania and North Macedonia, come together around two lakes – the Prespa National Park (PNP) covers an area of 257 km². It extends altitudinally from around 850 metres at the level of the lakes – Megali Prespa and Mikri Prespa, or Great and Lesser Prespa lakes respectively – to 2,330 metres on Mount Varnous. On the Greek side, the lakes are encircled by locally extensive reedbeds and floodplains turned largely to farmland for the cultivation of beans these days. The lands surrounding the basin are characterised by considerable geological diversity, revealing gneiss and granite on the slopes of Mount Varnous and limestone on the mountains to the west and south of the lakes. Prespa is also renowned for its great biological diversity and as an internationally important site for breeding waterbirds, hosting the largest colony of Dalmatian pelicans on the planet. For more information on the area, readers are referred to Lake Prespa, North-western Greece: A Unique Balkan Wetland by Alain Crivelli and Giorgos Catsadorakis, Prespa: A Story of Man and Nature by Giorgos Catsadorakis, and the Prespa Walking Guide by Kevin Standring et al.
Ideally this botanical guidebook would also include areas of the transboundary Prespa Park in Albania and North Macedonia, but so far limited data has been collected there. Thus our study, like the first comprehensive floristic account of Prespa by Georgios Pavlides in 1985, is limited to Greek territory. The book has been jointly prepared by its three authors, with the work divided as follows:
Introduction: EB, with input from AS and GF
Maps: GF
Vegetation section: EB, with input from GF and AS
Floristic section: AS, with significant contributions by EB and some by GF
Plant photo appendix: AS (unless otherwise indicated)
References and Index: AS, with input from GF and EB
The Flora section and the descriptions of vegetation are based on more than 15,500 single plant records (both field observations and herbarium specimens), the majority of them made by the authors. The Flora section includes over 1,800 accepted vascular plant taxa (species and additional subspecies). Numerous other species reported from the area are included in the keys and descriptions (presented in smaller font) but are considered questionable by the authors and require confirmation. With few exceptions, taxonomy and nomenclature follow Dimopoulos et al. (2013; 2016), unless recent work suggests otherwise. Owing to the authors’ studies the flora of the Prespa National Park is now reasonably well understood and its species richness exceeds that of almost all other areas of a similar size in northern Greece. History, human population and land use
Archaeological records indicate that the Prespa basin has been inhabited for some six thousand years. In the mid-fourth century BC it was part of the ancient kingdom of Macedonia, until it became a part of the Roman province of Macedonia in 146 BC. The nearest cities of any size at the time were Lychnidos (now Ohrid in North Macedonia) and Herakleia Lynkestis (now Bitola, also in North Macedonia), both of which were situated on the Via Egnatia, a major Roman road that ran from Dyrrachium (now Durrës in Albania) on the Adriatic Sea to Thessalonica (now Thessaloniki in Greece) by the Aegean Sea, and onwards to Constantinople (now Istanbul in Turkey).
In the late 10th century the Bulgarian Tsar Samuel made the small island of Agios Achilleios in Lake Mikri Prespa his seat of government, where he built a monumental basilica and other places of worship. After his death in 1014 the area was re-captured by the forces of the Byzantine Emperor Basil II, eventually experiencing several further incursions and losing its overall political importance, becoming part of the Ottoman Empire in 1386. Throughout more than five centuries of Ottoman rule the region remained remote, undeveloped and largely ignored. For that reason small monastic communities flourished here, particularly on the cliffs of Lake Megali Prespa, where a number of hermitages were constructed in the rocky clefts. Alongside Turkish, languages spoken locally were SlavoMacedonian, Albanian, Vlach and Greek. Turko-Albanian families lived in the village of Lefkonas, while the local pasha’s house was in Kallithea.
Greek influence in the region was limited until the late 19th century. In November 1912 Prespa was liberated from the Ottomans by the Hellenic Army. From late 1916 until 1919 the area was under the temporary military rule of the French Armée d’Orient; an enduring mark of this French presence during the First World War is the road the army built from Koula over the pass of Agios Georgios towards Psarades and on to Korça in Albania. The present national borders were established in 1923. While many local residents emigrated during the decline of the Ottoman Empire and later on, those who remained were joined in 1923 by 88 families of Greek refugees from the Pontus, a Greek-colonised region in the eastern parts of modern-day northern Turkey by the Black Sea. With the arrival of Italian forces in April 1941 until their surrender in September 1943, Prespa was incorporated into the territories administered by Italy during World War II. It remained under foreign control until Axis troops withdrew from Greece in 1944.
During the Greek Civil War between 1946 and 1949 the Prespa region was a stronghold of the Greek Democratic Army under its commander-in-chief, Nikos Zachariadis. After the defeat of his leftist forces entire villages were deserted and a considerable number of houses were left in ruins. The area was largely abandoned, with just 1,500 people left at the con-
2. Arable fields
CEREAL CROPS OF WHEAT, winter barley, rye and – more rarely – oats are chiefly cultivated in Prespa in the wider surroundings of Vrondero and the deserted village of Pyxos. Additionally, there are a few sandy fields where rye is grown at the eastern end of the isthmus between the lakes of Megali and Mikri Prespa, as well as near the abandoned hamlet of Sfika, south of the smaller lake. Most of the small-scale cereal fields around the upland villages of eastern Prespa have long been abandoned. Wartime pressures in the 1940s – from both the Second World War and the Greek Civil War which almost immediately followed – triggered a mass exodus from Prespa’s villages and the permanent abandonment of some. All extant cereal fields are non-intensive, and are cultivated without herbicides while utilising only moderate amounts of organic fertiliser, if used at all. These conditions, together with relatively sparse sowings and the use of self-saved seed (which is imperfectly purified and contains certain weed seeds that are dependent on a farmer’s assistance), are the prerequisites for an unparalleled richness and density of wild arable plants in Prespa. Most arable plants of traditional agriculture are susceptible to herbicides and industrialised farming processes and are therefore red-listed in many countries, being either vulnerable, threatened or even extinct in much of Europe, but they can still be found scattered throughout the fields of Prespa. And in the case of some species they can even be abundant. The surviving cereal fields of Vrondero conjure images of a mechanised yet pre-industrial stage in the development of the agricultural landscape, their wildflower-filled vistas exceptional in Greece and very rare in the rest of Europe.
The fields south of Vrondero, on moderately base-rich soft silty soils derived from Pliocene marly limestone sediments, are cultivated with wheat and lucerne and sometimes
two-rowed winter barley. Lucerne, otherwise known as alfalfa, is a perennial plant that’s able to be harvested several times and is increasingly used as a winter feed instead of barley and rye. Allotments have recently been planted with Sideritis, the local mountain tea plant of which two species are native to nearby Mount Devas and Mount Triklario. The cereal fields are harvested in July, and in contrast to perennial crop fields such as lucerne and mountain tea, they contain a considerable portion of the wild arable plant species of the Central Balkans, including, among many others, Adonis aestivalis subsp. aestivalis (a rarity in Greece otherwise restricted to a few fields in the northeast of the country), Adonis flammea, Agrostemma githago, Asperula arvensis, Bifora radians (which exudes an intense scent of coriander where abundant), Bupleurum rotundifolium, Camelina microcarpa, Caucalis platycarpos, Consolida regalis subsp. paniculata, Galium spurium, Legousia speculum-veneris, Lolium temulentum, Neslia apiculata, Ranunculus arvensis, Vaccaria hispanica and Vicia pannonica subsp. striata (Vicia striata). In sites with greater soil moisture Anagallis foemina, Anthemis cotula, Euphorbia platyphyllos, Ranunculus ficaria (Ficaria verna) and Oenanthe silaifolia have also been recorded.
North of Vrondero in the Pyxos area occur Pliocene sandstones weathered to sandy and silty colluvial soils which are generally leached of nutrients and slightly more acidic than those to the south of Vrondero. Understanding this, the farmers primarily cultivate barley and rye instead of wheat and lucerne. The weed flora is distinct in having greater amounts of Apera spica-venti, Avena fatua and Centaurea cyanus (Cyanus segetum ), which can be so abundant in places that whole fields are coloured cornflower blue. Among the more significant and rare plants are two annual species of the carrot family with characteristic spinose seeds, Caucalis platycarpos and Turgenia latifolia, the pink-flowered Knautia orientalis , Melampyrum arvense (which is hemiparasitic on cereal roots and striking with its purple inflorescences), a particular agro-ecotype of another hemiparasite, the yellowflowered Rhinanthus pindicus , and the golden-headed Centaurea adamii, which is widespread in the fields around Vrondero but is a rare plant in the rest of Greece and the Balkans. Indicators of more acidic sandy soils are Teesdalia coronopifolia, Galium divaricatum and Rumex acetosella subsp. acetoselloides Galeopsis ladanum is confined to only a few fields, being uncommon in Greece and found only in the north of the country. Individuals of the extremely rare parasitic plant Phelypaea boissieri were rediscovered in the area. According to local residents it used to grow in large numbers in fields and along field edges, and was collected in spring as an ornamental plant known as katifes , a Turkish word meaning ‘velvet’, due to the velvety feel of its brilliant red flowers.
Today, the reedbeds’ phytomass largely remains in the lake’s ecosystem, increasing oxygen depletion during the decomposition process and accelerating sedimentation and the long-term development of the lake. Moreover, there is a considerably higher inflow of nutrients and pesticide-polluted soil being washed in from the bean fields in the immediate environs of the lake. The growth of Phragmites rhizomes has been impaired in the previous decade or two, potentially because of seasonally anoxic conditions at times of high water levels, or due to a combination of several factors. As a result, Typha angustifolia has become considerably more prominent in the reedbeds in recent years and now predominates in larger extents of Vromolimni than before. In contrast to Phragmites , Typha forms a habitat far less conducive to the lake’s birdlife and doesn’t generate floating islands.
The current water nutrient status of the lakes – eutrophic, that is moderately nutrientrich – is reflected by the productive reedbed vegetation as well as by the water plants. The aquatic vegetation to the lakeside of the reedbeds consists chiefly of rooting plants –such as the emergent water lilies, the white-flowered Nymphaea alba and the yellowflowered Nuphar lutea – that are sheltered from waves in suitably protected parts of Lake Mikri Prespa and have large leaves floating on the surface of the water. They root at the bottom of the lake to a depth of 1.5-2.5 metres. The water lilies are commonly
accompanied by such submerged plants as Ceratophyllum demersum, Myriophyllum spicatum and Potamogeton crispus , which can also grow in deeper water. Fragments of the aquatic plants Potamogeton perfoliatus, Myriophyllum spicatum and Vallisneria spiralis are frequently washed ashore on the much deeper Lake Megali Prespa. In more shallow waters of both lakes occur Hydrocharis morsus-ranae , Trapa natans , Potamogeton lucens, Potamogeton natans and Persicaria amphibia . Elodea canadensis , a water plant of North American origin which has established itself throughout much of Europe since the 19th century, was recorded in Greece for the first time only a few years ago at two sites in the southern part of Lake Megali Prespa. Small aquatic plants not rooting in the lake bottom but instead found floating in the sheltered waters of a few miniature lagoons and bays on both lakes include, roughly listed according to abundance, Lemna minor , Lemna polyrhiza (Spirodela polyrhiza ), Lemna trisulca, Azolla filiculoides , Salvinia natans and Lemna gibba . While Azolla filiculoides , a native of tropical America, was first found in Greece in 1982 and has long since been naturalised in both of Prespa’s lakes, a second introduced floating neophyte, Lemna minuta , native to temperate South America, was discovered in Prespa in 2017 and is expected to establish itself soon. It is an invasive plant throughout much of Southern, Western and Central Europe, but with very few records in Greece so far.
11. Juniper woodlands and shrublands 1
THERE ARE FIVE TAXA OF JUNIPER IN PRESPA. The tree-shaped Greek juniper, Juniperus excelsa, and stinking juniper, Juniperus foetidissima, form open woodlands in which the former is usually predominant. The latter species appears to be mostly rare and scattered. Prickly juniper, Juniperus oxycedrus – represented in the region, as throughout the Southern Balkans and the Eastern Mediterranean, by subsp. deltoides – is a shrub, rarely exceeding 2.5 metres. In Prespa it is a common constituent in the shrub layer of oak and juniper woods, often found together with Buxus sempervirens, Acer monspessulanum, Hippocrepis emerus subsp. emeroides and Prunus webbii. It is the host of juniper mistletoe, Arceuthobium oxycedri, a specialised hemiparasitic epiphyte rooting on juniper branches, which has leaves reduced to scales and inconspicuous sessile flowers. Common juniper, Juniperus communis subsp. communis, is pervasive in the national park only on the rough pasture lands that are widespread on Mount Triklario (Sfika) above 1,400 metres and Mount Varnous between 1,300 and 1,800 metres, where any beech and fir forests were cleared centuries ago. Further up, characterising the high montane and subalpine zone of Mount Varnous, dwarf juniper, Juniperus communis subsp. nana, forms extensive low scrub. Hybrid plants of the two subspecies – dwarf and common juniper –have been recorded on Mount Varnous (and should not be mistaken for subsp. hemisphaerica, another more southerly distributed subspecies which does not occur in Prespa), but appear to be uncommon. Dwarf juniper scrublands are further discussed in the chapter on high mountain vegetation.
Woodlands of the tree junipers – Juniperus excelsa and Juniperus foetidissima – are rare in Southeastern Europe. They are western outposts of the far more extensive juniper
woodlands in southern Anatolia and the Near East. Since they have been acknowledged in the European Union’s Habitats Directive as a priority habitat type for nature conservation they are receiving increasing attention amongst conservationists, and rightly so. For the people of Prespa, however, the juniper woods are an integral part of their cultural history and were of everyday importance to life in the past. Residents used the extremely durable juniper wood for building houses, fences, roofs and boats, as well as fish traps known as pelaizia, in which juniper branches were piled up in shallow waters in winter.
Throughout much of Prespa the tree junipers are rather shrubby, multi-stemmed and not very tall, reaching only 3-8 metres high; often they are in mixed company with wild almond, Prunus webbii, and Macedonian oak, Quercus trojana, as well as other broadleaved trees (Carpinus orientalis, Fraxinus ornus, Ostrya carpinifolia, Pistacia terebinthus). In the area of Roti near Psarades, as well as in the vicinity of Vrondero, the understorey of the low juniper shrubland is thick with Buxus sempervirens. The most impressive juniper groves occur next to the chapels of Agios Georgios (otherwise known as Ai Yioryi, between Koula and Psarades), Agios Athanasios (above Vrondero) and Theotokos (above Psarades). A few trees from a former grove still survive near Agathoto, an abandoned hamlet on the west side of Lake Mikri Prespa with very few signs of habitation remaining, where the ruined church was built of stone and juniper wood. The single-stemmed juniper trees reach 20 metres and the trunks at breast height can have diameters of 1-1.5 metres; the circumference of one tree at Agios Athanasios was
-- Lower leaves sparsely pubescent. Calyx setae 2-3 times as long as corona ...................... S. ochroleuca
Scabiosa cinerea Lam.
Perennial, woody at base. Stem 40-60 cm tall, appressedpubescent throughout. Lower leaves subentire, velutinous of short, crispate stellate hairs, greyish-green above, whitish-green beneath. Upper cauline leaves 1-pinnatisect with linear to linear-lanceolate segments. Corolla white. Calyx setae 5, dark brown to blackish. – Fl. late June to July/August.
Native. Mt Triklario: Dry, rocky calcicolous grassland, 1650 m. Recently recorded in what appears to be the only Greek locality and the southernmost for this scattered species. – Mountains of SW Europe and the NE Adriatic and W Balkan region.
Scabiosa columbaria L.
Perennial, slightly woody at base, 30-70 cm tall, shortly eglandular-pubescent throughout. Lower leaves lyrate to 1- or 2-pinnatisect. Upper leaves pinnatisect with linear segments. Phyllaries lanceolate, shorter than flowers. Corolla pale lilac to pinkish-mauve. Involucel tube c. 2.5 mm, with short white hairs at base. Corona with 20-25 veins not exceeding margin. Calyx setae 5, spreading, 3-5 times as long as corona. – Fl. July-September.
Native. Mt Varnous: Grassland over granite, 1800-2000 m. – Mainland, N Peloponnisos and Ionian Islands. –Widespread in Europe, very variable.
Scabiosa ochroleuca L.
Like S. columbaria, but corolla cream; calyx setae 2-3 times as long as corona. – Fl. late June to August. – Plate 53: 6.
Native. A single record (1973) from above the village of Platy at c. 1000 m; needs confirmation but seems feasible. – Mainland, Peloponnisos and island of Thasos. – SE Europe and SW Asia, very variable.
Scabiosa taygetea subsp. portae (Huter) Kokkini
Like S. columbaria, but lower leaves subentire to lyrate, densely white- or yellowish-lanate; terminal segment of lower leaves much wider than the lateral; corolla white or cream, sometimes tinged pale lilac. – Fl. July to early September.
Native. Mt Varnous: Alpine grassland with granitic outcrops, 1950-2150 m. – Mountains from NW Greece to Taigetos. – W Balkan Peninsula, perhaps also in Italy.
Scabiosa triniifolia Friv.
Like S. columbaria, but often biennial, subglabrous throughout; all leaves 2- or 3-pinnatisect into linear segments; corolla white to cream. – Fl. June-August.
Native. Dry, mostly quartzitic or sandy grassland, 8501200 m. – Northern mainland and island of Thasos. –Balkan Peninsula.
Scabiosa webbiana D. Don
Like S. columbaria, but stem subglabrous below; lower leaves lyrate to 1-pinnatifid with large terminal lobe;
upper cauline leaves 2-pinnatisect with linear segments, all densely lanate to pannose; corolla white to cream. – Fl. July-September.
Native. Dry, sandy or rocky calcicolous grassland, 9001350 m. – Mainland, Peloponnisos and island of Thasos. – Balkan Peninsula and W Anatolia.
DROSERACEAE
Aldrovanda vesiculosa L.
Small, free-floating aquatic plant. Leaves in whorls of 69, with setaceous segments and an orbicular lobe which can close rapidly and entrap small animals. Flowers (rarely produced in European populations) axillary on short peduncles, held just above water, ephemeral, regular, 5-merous; petals white. – Plate 53: 7.
Native. Lake Mikri Prespa is the only Greek locality for this species. It was collected by Pavlides at Fragma (southern end of the lake) in 1984, and rediscovered in this area by Raus, Strid & al. in 2011, floating in 1-2 m deep water at the outer edge of reed beds, with Nymphaea alba, Nuphar lutea, Utricularia vulgaris, U. minor, Myriophyllum spicatum, etc. Subsequently it has been found also in several other localities in Mikri Prespa. It is rare in Europe and considered critically endangered in Serbia, Bulgaria and North Macedonia. The general distribution is almost worldwide but very scattered.
ERICACEAE
Bruckenthalia spiculifolia (Salisb.) Rchb.
Dwarf shrub with many ascending, branched stems up to 20 cm. Leaves in whorls of 4-5, short-petiolate; blade linear, with strongly revolute margins. Flowers in short, dense terminal racemes, 4-merous. Bracts small; bracteoles absent. Calyx c. 1.5 mm, campanulate; lobes equalling or shorter than tube. Corolla 2.5-3 mm, campanulate, bright reddish-pink. – Fl. late June to August. – Plate 53: 8
Native. Mt Varnous: Damp alpine grassland and dwarf juniper scrub, 1800-2100 m. – Mountains of N mainland, southwards to Pieria. – Balkan Peninsula, Carpathians and N Anatolia.
Monotropa hypopitys L., s. lat.
Syn.: Hypopitys monotropa Crantz
Wholly mycotrophic perennial herb without chlorophyll, whole plant cream, sometimes tinged pink. Stem 10-20 cm, erect, simple. Leaves alternate, sessile, scale-like, ± fleshy. Flowers few in a terminal raceme, nodding at anthesis, later erect. Petals 6-12 mm, saccate at base, erect, oblong-spathulate. Capsule subglobose. – Fl. JuneAugust.
Native. Fagus and mixed oak forest, 1050-1800 m. –Mountains of mainland and Peloponnisos, southwards to Taigetos, also on the island of Kefallinia. – A widespread Euro-Siberian species.
At least some of the plants from PNP appear to represent var. glabra Roth (Hypopitys hypophegea (Wallr.) G. Don), which has entirely glabrous flowers. Taxonomy and distribution are imperfectly known.
Orthilia secunda (L.) House
Rhizomatous, laxly mat-forming herb; stems suberect, simple, 5-20 cm. Leaves basal but not in a distinct rosette, petiolate; blade ovate-elliptic, subentire. Cauline leaves reduced to small scales. Flowers 10-15 in a dense, secund, somewhat nodding raceme. Calyx c. 1 mm. Corolla 4-6 mm, greenish-white, campanulate with erect, finely lacerate lobes. Stamens 10. Style 3-5 mm, straight. – Fl. June and July.
Native. Fagus and mixed beech-fir forest, 1400-1700 m. – Mountains of N & C mainland, scattered southwards to Taigetos. – Almost circum-boreal.
Pyrola minor L.
Glabrous, rhizomatous herb. Stem erect, 10-20 cm. Basal leaves in a lax rosette, somewhat coriaceous, petiolate; blade broadly elliptic, somewhat crenulate-serrulate. Flowers in a terminal raceme, not secund. Corolla 5-7 mm in diam., globose, whitish to lilac-pink. Style 1-2 mm, straight, included, not expanded below the stigma. – Fl. June and July. – Plate 53: 9.
Native. Mt Varnous: Deciduous and mixed beech-fir forest, woodland margins and by forest tracks, 1400-1700 m. – Mountains of N mainland, southwards to c. 40°N. –A widespread, circum-boreal species.
Vaccinium L.
Deciduous shrublets or small subshrubs. Leaves alternate, simple, ovate-elliptic. Corolla broadly ellipsoid to subglobose with short lobes. Fruit a subglobose berry.
1. Twigs angled, green. Leaves serrulate ....... V. myrtillus -- Twigs terete, brownish. Leaves entire, distinctly reticulate-veined ...................................... V. uliginosum
Vaccinium myrtillus L.
Rhizomatous, deciduous subshrub forming loose mats with bushy stems 10-30 cm. Stems 3-angled, glabrous, green at least in the upper part. Leaves 8-20 mm, ovateelliptic, serrulate. Flowers solitary or paired, nodding. Corolla 4-6 mm, subglobose, with short revolute lobes, pink or occasionally greenish-white. Fruit a subglobose berry 4-8 mm, blackish with a glaucous bloom. – Fl. May and June. – Plate 54: 1.
Native. Mt Varnous: Gregarious in (sub)alpine grassland and dwarf juniper scrub over granite, 1700-2100 m. – Mountains of N mainland, southwards to c. 39°30'N. –A widespread Euro-Siberian species, very common in Scandinavia.
Vaccinium uliginosum subsp. microphyllum (Lange) Tolm. Differing from V. myrtillus in the following characters: Twigs terete, scarcely green; leaves entire, glaucous, distinctly reticulate-veined; corolla ellipsoid to urceolate, with suberect lobes, greenish-white with pinkish tinge. –Fl. May and June. – Plate 54: 2
Native. Reported once (1968) from high altitude on Mt Varnous, probably in wet grassland. – Mountains of northernmost Greece, close to the borders. – V. uliginosum is a widespread, circum-boreal species; subsp. microphyllum is a high-altitude form of uncertain distribution and status.
EUPHORBIACEAE
Euphorbia L.
Annual to perennial herbs or shrublets, with whitish latex. Leaves simple, alternate or occasionally opposite. Flowers minute, in groups within an involucre with 4-5 glands at top, the whole structure called a cyathium. Synflorescence of cyathia usually compound, with rays and raylets, subtended by ray leaves and raylet leaves. Perianth absent. Female flower solitary, pedicellate, surrounded by several male flowers. Fruit a subglobose, 3-lobed capsule; seeds 3, ellipsoid, usually with a conspicuous appendix (caruncle).
1. Leaves opposite, with stipules. Procumbent annual of sandy or ruderal habitats ............. E. prostrata
-- Leaves alternate, without stipules. Annuals or perennials of various habitats
7. Seeds sulcate, with 4 or 5 transverse grooves on each surface .............................................. E. falcata
-- Seeds reticulate-pitted, without transverse grooves .................................................... E. taurinensis
8. Leaves thin, green, linear or linear-oblong, at least 7 times as long as wide ................ E. cyparissias
-- Leaves rather thick, glaucous, oblong-elliptic to obovate, not more than 3 times as long as broad .............................................................................
9. Leaves usually broadly oblong, of firm texture but not fleshy. Seeds 1.7-2 mm .................... E. thessala
-- Leaves obovate, somewhat fleshy. Seeds 3-3.5 mm ............................................................ E. myrsinites
10. Erect, single-stemmed annuals ................................ 11
-- Perennials, usually with several ascending stems ........................................................................... 14
Both subsp. germanica (with calyx teeth half as long as tube) and subsp. heldreichii (Boiss.) Hayek (with calyx teeth 1/3 as long as tube) occur in PNP, the latter generally at higher altitude.
Stachys palustris L.
Rhizome with swollen sections. Stems erect, 40-100 cm, simple or sparingly branched, eglandular-pubescent. Cauline leaves subsessile, oblong-lanceolate, crenate. Verticillasters 4-10-flowered, the upper crowded; lower floral leaves exceeding calyx. Calyx 7-10 mm, shortly eglandular-pubescent and sometimes also with glandular hairs; teeth ± equalling tube, narrowly triangular. Corolla 12-16 mm, pale pinkish-purple with darker spots. – Fl. June-October. – Plate 79: 8
Native. Marshy habitats by the lakes and along streams, 850-1050 m. – Mainland southwards to c. 38°20'N, island of Kerkira. – A widespread, circum-boreal species.
Stachys plumosa Griseb.
Short-lived perennial. Stems ascending to erect, 20-50 cm, moderately branched, ± villous. Lower leaves shortpetiolate with broadly oblanceolate blade, the upper subsessile, narrowly elliptic to lanceolate; all shallowly serrate, sparsely pilose, green. Flowers in several, 6-14flowered verticillasters confluent above. Calyx 9-12 mm, narrowly campanulate, glandular-puberulent and sparsely pilose, green; teeth equalling tube, narrowly triangular, acuminate. Corolla 14-20 mm, white, cream or pale pinkish with small pinkish-purple spots, pale yellow when dry. – Fl. May to early August. – Plate 79: 9.
Native. Rocky slopes with dry grassland, road embankments and woodland openings, on calcareous or quartzitic ground, 850-1200 m. – Mainland especially in the north, islands of Thasos and Evvia. – Balkan Peninsula from SE Albania to Bulgaria.
Stachys recta L.
Perennial with slender rhizome. Stems ascending to suberect, branched at base, usually 20-40 cm. Cauline leaves subsessile, oblong-lanceolate, cuneate at base, crenate-serrate, sparsely eglandular-pubescent. Verticillasters 4-8-flowered, crowded into a rather short inflorescence or the lower remote. Calyx 7-9 mm, sparsely pilose, green; teeth ± equalling tube, triangular-lanceolate, acuminate. Corolla 12-15 mm, cream or yellow with small reddish spots. – Fl. June-August.
Native. Reported once from near the village of Pyli at 860 m. – Scattered on mainland and Peloponnisos, very variable. – S Europe and SW Asia.
Stachys serbica Pančić
Slender annual; stem erect, 10–25 cm tall, simple or sparingly branched, sparsely patent-pilose. Cauline leaves in few pairs, petiolate to subsessile; blade narrowly ovate, create-serrate. Verticillasters 1 or 2, 4-6-flowered; floral leaves like the upper cauline leaves. Bracteoles lanceolate, equalling or exceeding calyx tube. Calyx c. 12 mm, green, pilose; teeth almost as long as tube, narrowly
triangular-lanceolate, acuminate, slightly recurved in fruit. Corolla c. 16 mm, reddish-purple; upper lip ± straight, entire; lower lip with 3 subequal lobes. – Fl. May and June. – Plate 80: 1
Native. Scrub and open woodland over limestone, c. 900 m. – Scattered in NW & NC mainland southwards to c. 39°15'N. – Balkan Peninsula from Albania to Bulgaria and northwards to C Serbia.
Stachys sylvatica L.
With long, creeping rhizomes. Whole plant sparsely covered with patent glandular and eglandular hairs, with a characteristic, unpleasant smell. Stem erect, 30-80 cm. Leaves long-petiolate; blade 4-13 cm, ovate, cordate at base, crenate-serrate. Verticillasters distant or somewhat crowded towards top, usually 6-flowered. Calyx c. 8 mm; teeth slightly shorter than tube, narrowly triangular, erecto-patent. Corolla 14-18 mm, dark dull reddish-purple with whitish pattern on lower lip. – Fl. June-August. –Plate 80: 2.
Native. Humid, shady places in deciduous woods and ravines, 860-1500 m. – Mountains of mainland, southwards to c. 39°N. – Most of Europe, extending to N Scandinavia, naturalized overseas.
Stachys tymphaea Hausskn.
Resembling S. germanica but usually shorter (15-40 cm); blade of middle and upper cauline leaves ovate, cordate at base; verticillasters few (usually 2-5); calyx glandularand eglandular-pubescent, teeth ± recurved in fruit. – Fl. June and July. – Plate 80: 3
Native. Mt Varnous: Woodland margins, damp subalpine grassland, along mountain roads, 1400-2100 m. –Mountains of mainland, southwards to c. 38°30'N. – C & S Italy, W & C Balkan Peninsula.
Teucrium L.
Perennial herbs or subshrubs. Leaves entire or crenate to lobed. Calyx regular or slightly 2-lipped; teeth ± equal. Upper lip of corolla absent, the lower conspicuous, 5lobed. Stamens usually exserted from corolla tube.
1. With scaly stolons. Plant of wet habitats ... T. scordium
-- Without stolons. Plants of dry, usually rocky habitats .......................................................................... 2
2. Leaves deeply dentate to incised. Inflorescence of well separated verticillasters. Corolla pinkishto reddish-purple .................................... T. chamaedrys
-- Leaves subentire to crenate. Corolla usually whitish to pale yellow. Inflorescence capitate .............. 3
3. Leaves entire with revolute margins. Corolla pale yellow ............................................... T. montanum
-- Leaves crenate. Corolla usually whitish, occasionally pink .... T. polium
Teucrium chamaedrys L.
Perennial with woody, much-branched rhizome producing several ascending to erect, slender, herbaceous flowering stem 10-20(-40) cm. Middle cauline leaves shortpetiolate; blade elliptic-obovate, cuneate at base, dentate to incised. Flowers in lax, 4-8-flowered verticillasters
forming a raceme. Floral leaves like the cauline but somewhat narrower. Calyx c. 6 mm, campanulate; teeth 2/3 as long as tube, triangular, sharply acute to acuminate, purplish. Corolla 12-15 mm, pinkish- to reddish-purple. –Fl. May-August. – Plate 80: 4.
Native. Woodland margins, rocky places in dry grassland, stony and gravelly places along mountain roads, mostly on calcareous ground, 900-1600 m. –Mainland and Peloponnisos, rare on the islands and absent from the South Aegean area. – A widespread and variable species of Europe and SW Asia.
Teucrium montanum L.
Decumbent, much-branched subshrub forming mats, usually appressed-puberulent on stem, leaves and calyces. Flowering stems slender, procumbent, 5-25 cm. Leaves 10-20 mm, subsessile, oblong, entire, with revolute margins. Flowers short-pedicellate in loose terminal heads with leaf-like bracts. Calyx 6-8 mm, regular; teeth subequal, triangular, half as long as tube. Corolla 10-12 mm, pale yellow. – Fl. late May to August. – Plate 80: 5
Native. Mt Triklario: Dry calcicolous grassland with rocky outcrops, 1400-1700 m. – Mainland and Peloponnisos, rare on the Aegean Islands and absent from the south. – C & S Europe and Anatolia.
Teucrium polium subsp. capitatum (L.) Arcang.
Syn.: T. capitatum L.
Bushy subshrub with ascending stems 10-30 cm, ± greytomentose throughout, especially in the inflorescence. Leaves subsessile, linear-oblong to narrowly obovate, shallowly crenate at least towards the apex. Inflorescence a simple or usually compound head at least as wide as long. Calyx 3-5 mm. Corolla cream or pinkish, with lateral lobes of lower lip short, rounded or triangular. – Fl. June to early August. – Plate 80: 6.
Native. Rocky slopes with dry grassland and open scrub, 850-1350 m. – Throughout Greece. – T. polium s. lat. is a polyploid complex, widespread in the Mediterranean area and SW Asia; all Greek material is referable to subsp. capitatum
Teucrium scordium subsp. scordioides (Schreb.) Arcang. Rhizomatous perennial, usually with scaly stolons; whole plant patent-pilose and glandular-puberulent. Stem suberect, 20-50 cm tall, usually much-branched. Leaves sessile, ovate-oblong, coarsely crenate-dentate. Flowers in distant, c. 4-flowered verticillasters forming lax spikes. Floral leaves like the cauline but somewhat smaller, exceeding calyx. Calyx c. 4 mm, somewhat gibbous at base; teeth half as long as tube, triangular, acute. Corolla 7-10 mm, pinkish-mauve to almost white. – Fl. JuneOctober. – Plate 80: 7
Native. Wet meadows and reed beds by the lakes, 850 m. – Scattered throughout Greece. – Mediterranean region and SW Asia, replaced by subsp. scordium further north.
Thymus L.
Small perennial, aromatic herbs, woody at base, often creeping or mat-forming. Leaves simple, usually gland-
dotted and ciliate at least towards base. Flowers in verticillasters which are often crowded into a capitate inflorescence. Calyx slightly 2-lipped with the 3 upper teeth different from the 2 lower. Corolla 2-lipped. Stamens 4, exserted. – A taxonomically difficult genus with centre of diversity in the Balkan Peninsula and Anatolia. Variation, ecology and distribution of the T. praecox, T. longicaulis and T. leucotrichus groups in PNP need further study.
1. Leaf margins revolute, at least in leaves of axillary fascicles, or leaves with thickened, pseudo-revolute margins .............................................. 2
-- Leaf margins flat, not revolute ..................................... 7
2. Bracts ± similar to the leaves ....................................... 3
-- Bracts different from the leaves, at least twice as wide .......................................................................... 5
3. Leaves velutinous with additional longer hairs ....................................................... T. leucotrichus
-- Leaves velutinous without additional longer hairs, sometimes glabrous ............................................ 4
4. Leaves sessile, 0.3-0.8 mm wide ................. T. boissieri
-- Leaves short-petiolate, 1.5-3 mm wide ..... T. dolopicus
5. Leaves velutinous with additional longer hairs. Bracts often purplish .................... T. leucotrichus
-- Leaves velutinous without additional longer hairs. Bracts often greenish ..........................................
6. Leaves petiolate, 1.5-3 mm wide, with slightly revolute margins; oil dots numerous .......... T. dolopicus
-- Leaves sessile, 0.5-1.3 mm wide, with strongly revolute margins; oil dots few or absent T. parnassicus
7. Stem hairy on the angles only ................ T. pulegioides
-- Stem hairy all round or on two opposite sides ..............................................................................
8. Long creeping branches absent; flowering stems 5-40 cm from a short, woody stock, without basal fascicles of small leaves ....... T. sibthorpii
-- Long creeping branches present; flowering stems in rows, up to 10 cm, with basal fascicles of small leaves ..............................................................
9. Bracts different from the leaves, at least twice as wide ........................................................... T. striatus
-- Bracts ± similar to the leaves ..................................... 10
10. Leaves widest in the apical third .............................
-- Leaves widest at middle or in the basal third ............................................................................
11. Leaves with prominent lateral veins ........................
12. Lateral veins curving, the upper pair joining to form a marginal vein .................................
13. Calyx 4-6 mm; upper teeth 1.4-2 mm, purplish ....................................................... T. thracicus
-- Calyx 3-4 mm; upper teeth 0.8-1.2 mm, often greenish .................................................... T. longicaulis
14. Flowering stems 1-5 cm. Leaves 3-6 × 1-2 mm, exceeding internodes. Corolla 3-4.5 mm, scarcely exceeding calyx ............................ T. stojanovii
-- Flowering stems 3-10 cm. Leaves 5-13 × 1.5-3 mm, shorter than internodes. Corolla 4-6 mm,
1. Geraniaceae: Geranium sylvaticum - 2. Geranium versicolor - 3. Gesneriaceae: Ramonda serbica - 4. Globulariaceae: Globularia bisnagarica - 5. Globularia cordifolia - 6. Grossulariaceae: Ribes alpinum - 7. Ribes uva-crispa - 8. Haloragaceae: Myriophyllum spicatum - 9. Hyacinthaceae: Muscari comosum
1. Hyacinthaceae: Scilla bifolia - 2. Hydrocharitaceae: Elodea canadensis - 3. Hydrocharis morsus-ranae (with Salvinia natans) - 4. Hypericaceae: Hypericum barbatum - 5. Hypericum olympicum - 6. Hypericum perforatum7. Hypericum rumeliacum - 8. Hypericum tetrapterum - 9. Iridaceae: Crocus cancellatus
