Small herbs. Rootstock tuberous, tubers two, entire, globose or ellipsoid. Stem often rather glaucous, erect. Leaves 2-4 basal, 1-2 cauline, spotted or unspotted. lnflorescence cylindrical, sometimes conical, dense, sometimes slightly twisted; floral bracts thin, membranous. Flowers small, greenish white, straw-coloured, whitish or pink, often spotted and lined darker pink or purple. Sepals and petals connivent, forming a hood. Labellum three-lobed, mid-lobe often bilobed, spurred, without calli, often minutely papillose, spur short and conical, or longer and cylindrical. Column short, with two large lateral stigmas joined below; anther prominent; pollinia two, each with a short caudicle attached to a viscidium enclosed in a single simple bursicle. Ovary cylindrical to fusiform, sessile, twisted, glabrous. (JW).
Neotinea normally grows on alkaline oil (Davies et al. 1988; Delforge 1995). Neotinea maculala has principally been recorded from limestone areas, such as the west of Ireland and from calcareous sea sand, such as the stabilized sand-dunes on the Isle of Man (Lang 1989). In Britain, N. ustulata most frequently occurs in the chalk districts of southern England (Lang 1989). Farther north in England it has also been recorded from the Carboniferous limestone areas of Derbyshire, Yorkshire, Cumbria, northern Lincolnshire, and eastern Yorkshire (Foley 1987). ln all of these cases the soils are relatively warm and dry (Summerhayes 1985), although in some parts of its European range, N. ustulata at least can grow in wetter soils (Delforge 1995). Some members of the genus can also grow in more acidic conditions. In a few sites in Ireland N. maculata occurs on semi-peaty soil overlying acid soils, and in the French and Italian Riviera it grows in soils formed from non-calcareous rocks (Summerhayes 1985). The pH range of soils in which N. ustulata occurs has also been reported as ranging from 5.2-7.3 (Arditti 1992), although relatively high calcium levels have been recorded in another study for soils in which N. ustulata and N. tridentata naturally grow (concentrations of 534.3 mg and 426 mg/100 g soil, respectively; Moller 1985, cited in Arditti 1992).
Neotinea can grow under conditions of full sun to shade in a range of habitats encompassing grassland, scrubland (including garrigue and maquis) and woodlands, and at both high and low elevations (Delforge 1995). In Cyprus, for example, N. maculata occurs in two distinct types of habitats: in pine forests high in the mountains, which comprise the southern part of the Troodos range, and on bare chalk soils on low ground near the sea (Davies et al. 1988). At lower-elevation locations on the island the two colour forms (pink and white flowers) of the species are found growing together, whereas in the mountains only the white form occurs. Similarly in the west of Ireland, N. maculata has long been associated with the limestone pavement, grassy pastures, and roadside verges of the Burren (Summerhayes 1951), but in recent years it has also been found growing in scrubby mixed ash and hazel woodland (Doyle 1985; Lang 1989) . These woodland plants resemble more closely those of Mediterranean colonies in that their inflorescences are more robust and taller than those of the more open habitats in the Burren (Lang 1989).
Elsewhere in the British Isles, N. maculata occurs at low elevations on stabilized sand-dunes (Lang 1989), whereas in parts of its range in mainland Europe it can occur in mountain locations at up to 2000 m (Delforge 1995). Other members of the genus are also reported from high elevations. Neotinea lactea occurs up to 1800 m in some localities and is said to be particularly frequent in mountain pastures on limestone. Neotinea tridentata occurs in alpine meadow, grassy hillsides, and open woodland at up to 1600 m (Davies et al. 1988; Delforge 1995). Similarly, N. ustulata is often found in mountainous areas and can occur at elevations of up to 2400 m in alpine pastures. It is perhaps less tolerant of shade than congeneric species, being rarely found in garrigue or open woodland. In the British Isles it is usually found among the grasses of well-drained chalk downland in sheltered, sunny situations that are often southfacing, although it does sometimes occur in areas of juniper scrub where it may co-occur with Anacamptis morio (Lang 1989).
Flowering of Neotinea takes place sometime between early spring and late summer depending on the species and the location of its populations. The earliest flowering member of the genus is N. lactea, which blooms in February in North Africa (Davies et al. 1988) and reaches anthesis by April or May in other parts of it range. Neotinea maculata also flowers early, between mid- Mareh and May (Davies et al. 1988). Summerhayes (195 1) attributed it early flowering in the Burren to its 'winter-green' habit whereby the aerial shoots have already developed in the autumn preceding emergence of the inflorescence. Peak flowering is normally in May, which is before that of co-occuring orchids such as Spiranthes spiralis, Ophrys apifera, Anacamptis pyramidalis, and Epipactis atrorubens (Hoffman ex Bernh.) Besser. The flowering period of individual plants is relatively short and, a with the Isle of Man populations, Lang (1989) suggested senescence may be accelerated by seasonal high winds in these exposed area , although it may simply be the result of rapid fruit-set as seems to be the case with N. ustulata. Neotinea maculata completes anthesis and capsule formation within about one month in southern England (Lang 1989). Neotinea tridentata and N. ustulata flower relatively late, between April and August throughout their ranges (Davies et al. 1988; Delforge 1995). In Britain, most plants of N. ustulata flower in May-June in the south of England and a little later farther north (Summerhayes 1951). However, a form has been discovered recently at five sites in East Sussex and Hampshire on the south coast of England that flowers in July (Lang 1989). Individual plants can flower for at least two consecutive years (Lang 1989), but flowering of populations is highly variable and influenced by rainfall levels (Foley 1987; Lang 1989) such that in drought years no flowering may occur at all (Lang 1989). Not all populations of N. ustulata are winter-green, however, because some retain their leaves throughout the summer. Rasmussen (1995) suggested that this indicates the occurrence of different ecological races within this species.
Fruit-set in this genus is expected to below because its members are nectarless (cf. Neiland and Wilcock 1998). An observation of 11.8% of flowers setting fruit among a population of N. ustulata on bland, Sweden, in the summer of 1998 conforms to this expectation (Neiland and Wilcock, personal observation). An exception may be N. maculata, which casual observations indicate may have much higher levels of fruit and seed production. Also, N. maculata is said to be self-pollinating (Summerhayes 1951; Davies et al. 1988; Lang 1989) and may even be cleistogamous (Delforge 1995). Once seed germination has taken place, Neotinea plant remain underground as a protocorm/ mycorrhizome until the first root and aerial shoot develops, followed by formation of a tuber (Rasmussen 1995). Progression to the aerial stage is believed to occur rapidly in N. tridentata, which has been observed to produce its first leaf above ground in its second growing season (Beer 1863, cited in Rasmussen 1995). In contrast, N. ustulata is frequently quoted as having a long initial life underground of 10-15 years as a protocorm during which time it is entirely dependent on mycorrhizal nutrition (Stonjanow 1916; Zuchs and Ziegenspeck 1927, cited in Rasmusssen 1995; Summerhayes 1951; Wells 1981; Davies et al. 1988). The protocorm may be as long as 20-30 mm before roots and leaves are formed and have many constrictions along its length, which are taken to indicate each year's growth. However, Rasmussen (1995) suggested that their use may lead to inaccurate age estimates because constriction lines can also arise from contraction of the protocorm or from nodes. She stated that cultivated seedlings of N. ustulata are rarely constricted and cited work by Moller (1985) in which cultivated N. ustulata plants reached the flowering stage within three years as evidence, casting doubt on the accuracy of the long-age estimates of N. ustulata seedling stages.
Neotinea is rare throughout many parts of its distribution. For example, N. tridentata is said to be quite rare in the western limits of its range, although it is relatively common in Anatolia, Turkey (Delforge 1995). Neotinea ustulata is described as local and unpredictable in its appearance, although it may sometimes be abundant, especially in mountain areas (Summerhayes 1951; Delforge 1995). Recently new records in western Ireland of N. maculata have extended the range of this species beyond the Burren in County Clare where it was first discovered by F. M. Moore in 1864. It is now known to occur from west Donegal southward to east Cork (Shepherd and Shepherd 1985; Lang 1989), but it remains a rare orchid in Ireland as elsewhere in Europe (Delforge 1995). Its western and coastal locations indicate that it is adapted to a relatively warm oceanic climate (Summerhayes 1951), and its unusual disjunct distribution in the British Isles and Mediterranean is explained by fluctuations in climatic conditions in the past. The lrish populations of N. maculata are thought to be preglacial or inter-glacial relicts from a period of much warmer climatic conditions when this species and other Mediterranean plants still growing in the Burren would have been able to migrate northward acros a continuous land bridge from the present-day Iberian peninsula. The ameliorating conditions of the Atlantic Ocean on the local climate during and after the lce Age are thought to have allowed such species to persist in the west of Ireland when they were unable to do so in the rest of the glaciated British Isles (Summerhaycs 1951). By comparison, the other isolated populations of N. maculata in the British Isles discovered on the lsle of Man in 1969 are probably of much more recent origin. Because of their visual resemblance to the plants from the Burren (white flower and unspotted leaves), they are thought to have spread from western Ireland to Britain via seeds dispersed either by the wind or sea currents (Lang 1989) .
The British distribution of another rare member of the genus, N. ustulata, has shown a marked decline in recent years due to habitat loss and agricultural practices. For example, in a survey of northern English populations, Foley (1987) found that from 118 previously recorded sites of the species, only 15 colonies still survived. The largest of these comprised over 200 plants, but the majority comprised 35 or fewer individuals. He estimated that there were probably less than 500 flowering plants still surviving in the whole of the north of England distribution area at the time of the survey. In the southern counties of England, N. ustulata can still be found flowering in large numbers at some suitable sites on the chalk downland (Lang 1989) bur has also disappeared from some previous known localities. ln Suffolk, for example, it is estimated that it had gone from the county by 1840 (Sanford 1991). The number and extent of N. ustulata populations elsewhere in its European range is also declining, for example in southern candinavia (Petterson 1976) where it is being lost from grassland. The principal explanation for the reduction in Britain and other countries seems to be agricultural improvement of ancient pastures through the application of inorganic fertilizers and disturbance by ploughing (Foley 1987). Map records show that ploughing of prime field sites in Suffolk in the 19th century resulted in rapid loss of this species from the area (Sanford 1991), and in Derbyshire the species survives only in upland pastures, which are difficult to upgrade for agriculntral purposes (Foley 1987). Absence of grazing may also lead to competition by aggressive grass species, which rapidly out-compete this orchid. (RN).
Four species distributed from western Ireland and the Isle of Man eastward across southern Europe from Portugal and the Canary Islands to Turkey and the Lebanon, south to north Africa, east to the Caucasus, Iraq, and the Caspian region in western Asia. (JW).
