Anacamptis Rich.

First published in De Orchid. Eur.: 25 (1817)
This genus is accepted
The native range of this genus is Europe, Medit. to Central Asia.


General Description

Rootstock tuberous, tubers void, entire. Stem glabrous, leafy. Leaves basal and cauline, unspotted. Inflorescence densely to laxly few to many-flowered, shortly racemose to elongate; floral bracts membranous, sometimes leafy, slightly shorter than, equalling or exceeding ovary. Sepals and petals usually slightly concave, glabrous, dorsal sepal and petals connivent to form a helmet or hood (galea), or free but forming a loose hood (in A. papilionacea), lateral sepals spreading. Labellum spurred, entire and flabellate, shallowly three-lobed or distinctly three-lobed, side lobes (when present) rounded, oblong-rhombic, obliquely triangular or linear, mid-lobe (when present) entire or bilobed, glabrous, papillose or occasionally hirsute, sometimes with two longitudinal basal calli, spur filiform or broad and thick, entrance sometimes with crests and angular in profile, descending, horizontal or ascending, sometimes nectariferous. Column short; rostellum small; pollinia two, attached to a single viscidium or attached to separate viscidia, placed in a simple bursicle. Ovary cylindrical, subsessile or sessile, twisted, glabrous. (JW).


Members of the European genus Anacamptis are terrestrial orchids many of which typically grow on shallow, well-drained, calcareous soils such as chalk and limestone. These include the Mediterranean species, A. boryi, A. collina, A. papilionacea, A. sancta, and the more widespread species A. pyramidalis which, in Britain, is a characteristic orchid of the chalk downlands in southern England and also calcareous sand dunes (Summerhayes 1951; Davies et al. 1988; Delforge 1995; Rasmussen 1995). Anacamptis morio has the widest ecological range of the group (Summerhayes 1951; Delforge 1995). Although it prefers calcareous soils, it often grows on neutral to slightly acid substrates. In parts of Britain it flourishes in sand dume slacks (Lang 1989), and it is particularly common on heavy marl and clay soils if they are not too wet (Summerhayes 1951; Sanford 1991; Wells et al. 1998). ln contrast, A.Iaxiflora and A. coriphora, which tend to grow on slightly acid to alkaline soils, are more often found where the soils are permanently wet, such as in bogs, marshes, seepage zones, and other areas prone to flooding (Davies et al. 1988; Delforge 1995).
Anacamptis grows at a range of elevations throughout its distribution. Anacamptis sancta, for example, often occurs near the coast and always below 900 m, whereas A. pyramidalis and A. coriophora can grow at elevations over 2000 m (Davies et al. 1988; Delforge 1995). Members of the genus grow in a variety of grassland habitats including dry, nutrient poor pastures and meadows, old established dunes, marshy moorland, roadside verges and railway embankments, as well as in open shrub and woodland such as maquis, open pine woods, and olive groves in the Mediterranean region (Pettersson 1976; Davies et al. 1988; Delforge 1995). The variety of habitats within which individual Anacamptis species are able to grow depends on their level of shade tolerance. Anacamptis morio, for example, normally only occurs in open habitats in full sunlight. Thus, in Britain it typically grows in species-rich grasslands such as old hay meadows (i.e. Cynorus cristatus-Centaurea nigra grassland, MG5, in Rodwell's National Vegetation Classification, 1992) and in pastures where the turf is short and may be grazed. In these habitats, A. morio is often associated with cowslips (Primula veris Mill.; Primulaceae) and other species characteristic of undisturbed grassland such as adder's tongue (Ophioglossum vulgatum L.; Ophioglossaceae) (Summerhayes 1951; Lang 1989; Sanford 1991) . Summerhayes (1951) noted that where the vegetation is taller A. morio is absent and that it rarely occurs in shaded habitats such as woodlands. Anacamptis pyramidalis is more shade-tolerant. In Britain it is a component of the Hieracium sub-community of Bromus-Brachypodium grassland, CG5 (Rodwell 1992). The sward of its typical chalk grass land habitat in southern England is composed of relatively tall grasses. Anacamptis pyramidalis also grows among marram grass (Ammophila arenaria Link) and lyme grass (Elymus spp.) on sand dunes (Lang 1989; Allan et al. 1993), but it is less common in short grassland where other orchid calcicoles are often found (Summerhayes 1951). It can withstand partial shade since it sometimes occurs among low shrub of hawthorn and blackthorn or in some beech, oak, and other woodlands where their canopies are relatively open (Summerhayes 1951; Davies et al. 1988). Other members of the genus that occur in open woodlands or shrubland in the Mediterranean include A. champagneuxii, A. sancta, A. collina, A. papilionacea, and A. boryi (Davies et al. 1988; Delforge 1995).
The peak flowering period of Anacamptis is from March to June, although this varies between species due to the elevation and latitude of individual populations. The Mediterranean species, A. collina, is early flowering, commencing in December in some localities and with successive cohorts coming into flower until April (Delforge 1995). In contrast, A. sancta and A. coriophora, also growing in the Mediterranean, flower later, with anthesis taking place between April and July (Davies et al. 1988; Delforge 1995). In the south of its range in the Algarve in Portugal, Anacamptis morio flowers in March, several weeks earlier than the same species in me north of its range in Scotland where it flowers in May (Neiland 1994). The widespread species A. pyramidalis flowers early in the summer across western Europe, e.g. from mid -June until the end of July in Britain, which is several weeks later than many other orchids growing in the same habitat in England (Lang 1989). This characteristic of flowering some time after co-occurring orchid species is also noted for the eastern Mediterranean species, A. sancta, which flowers in April (Davies et al. 1988).
Anacamptis orchids are usually 'winter-green' species. Normally each plant produces a rosette of leaves in the autumn that remains green and functional throughout the winter and spring and senesces early, often during me flowering period. Following fruit-set, the orchids perennate as underground tubers. This is of adaptive advantage for growth in a Mediterranean climate where Anacamptis is thus able to survive the summer drought. The tubers are replaced annually, and under favourable conditions more man one tuber may develop from me buds at the base of the aerial stem. Formation of extra daughter tubers from the moth plant is the means by which vegetative propagation can be achieved in this and other terrestrial genera such as Dactylorhiza (Rasmussen 1995).
Fruit formation in Anacamptis species is normally quite low. Average fruit-set figures of 21.1 %, 29.0%, and 33 .5% have been recorded from A. morio, A. collina, and A. pyramidalis populations in Europe (Neiland and Wilcock 1998). A higher proportion of flowers of A. papilionacea examined by Vogel (1972) in Germany set fruit (50%), but this population was unusual in that it was pollinated during regular patrolling flights of territorial bees. Average seed set per capsule varies among species in me genus. Salisbury (1942) counted about 2000 seeds in a single capsule of A. pyramidalis and estimated that the average output of seeds from an individual plant could be 35 000 seeds. The number of seeds formed per capsule of A. morio ranges from 2000 to 4000, and seed outputs per plant of between 20 000 and 32 000 seeds have been calculated using field observations of average numbers of capsules per plant of A. morio (Salisbury 1942; Neiland 1994).
Germination of Anacamptis seeds normally takes place in me spring after which a protocorm is formed. In some species, e.g. A. morio and A. papilionacea, this rapidly develops and produces its first root in the autumn and the first foliage leaf in the following winter or spring after which time a tuber develops at the base of the stem. Growth of the protocorm is slower in A. pyramidalis, and it may take 1-3 years before the first tuber and foliage leaves appear, depending on growth conditions at field sites (Rasmussen 1995). The formation of the inflorescence is delayed still further. Wells (1981) stated that flowering first takes place four to five years after germination in A. morio and seven to eight years after germination in A. pyramidalis. The protocorm and rhizome of Anacamptis species are highly mycotrophic, but the endophytes involved have yet to be identified (Rasmussen 1995). Seeds of some species have been germinated symbiotically with fungal strains, including seeds of A. laxiflora with Ceratobasidium corrigerum (Muir 1987, 1989), A. papilionacea with Epulorhiza sp. (Andersen 1990, cited in Rasmussen 1995), and A. morio with a Ceratobasidium-like isolate obtained from the Royal Botanic Gardens, Kew (McKendrick 1996a). It is now possible to propagate a variety of Anacamptis species a symbiotically on a range of nutrient media (e.g. Malmgren 1993). The time to reach maturity may be quite short; Frosch (1980) reported that A. morio plants initially cultured by asymbiotic methods reached flowering within 23 months. Symbiotic techniques have been successfully used at Kew to provide seedlings of several Anacamptis species (including A. laxiflora and A. morio) for re-introduction to field sites (Muir 1989) as a means of conserving these species, the numbers of which have been declining in England in recent years.
Some members of the genus are abundant locally, and under favourable conditions large populations of flowering plants may be formed (Summerhayes 1951) that can be long-lived. Anacamptis pyramidalis, for example, is known to have persisted in high numbers for over 100 years at one locality on the east coast of Scotland (Allan et al. 1993) and at another site in Kent in southern England (Lang 1989). As with many congeneric orchids, A. pyramidalis is able to colonize new localities rapidly by means of seed dispersal. For example, Nothdurft (1995) described the appearance, presumably due to wind dispersal, f a flowering plant of A. pyramidalis in a garden in Germany 40 km from the nearest population. Anacamptis laxiflora in the Mediterranean is also a typical weedy species and quickly invades abandoned fields (Pettersson 1976). Populations of Anacamptis species have been described as fluctuating in numbers from year to year. Anacamptis morio, for example, is thought to be short-lived and monocarpic (Summerhayes 1951). However, in a recent long-term demographic study by Wells and his co-workers it has been shown that, after reaching a critical size, A. morio plants can flower for several years in succession (for example up to 17 out of the 18 years of the study in a few cases). In this population, a large proportion of plants flowered each year (usually >40%). The probability of an individual flowering increases with leaf number, but there seem to be no clear relation hip between flowering and climatic factors, which could explain variations in the flowering performance of the population from year to year (Wells et al. 1998).
In addition to the more common and widespread species, Anacamptis includes some species with Iocalized distributions, e.g. the eastern Mediterranean species, A. sancta, A. coriopbora, and the southern Greek endemic A. boryi (Delforge 1995). Many of these localized species are also declining in numbers. Anacamptis morio in Britain was cited as one of the commonest of British orchids by Summerhayes (1951), but within three decades it had undergone a dramatic decline and is now considered to be a threatened species (Lang 1989). The major reason for the disappearance of Anacamptis species has been alteration of their grassland habitat. Seedlings can be physiologically tolerant of some shade (e.g. A. morio, McKendrick 1996b), but in the field the plants can readily become shaded-out if taller plants become dominant in the habitat. For example, on the Swedish island of Oland, Lind (1992) showed that populations of A. pyramidalis growing at sites that were not kept open by grazing were decreasing because of the invasion of taller grasses and shrubs. Changes in the vegetation composition due to re-seeding old pastures, drainage or fertilization can also have a detrimental effect on Anacamptis populations (Sanford 1991; McKendrick 1996). Agricultural improvement of ancient herb-rich meadows by the application of fertilizer has been used to explain the recent dramatic decline of A. morio in England. High levels of inorganic phosphorus can be toxic to the orchids; even low level may improve hay yield but lead to the demise of the orchid, which suffers increased competition from the surrounding plants (Silvertown et al. 1994; McKendrick 1996a). (RN).


A dozen species distributed in northern, central, and southern Europe, western Asia and south to North Africa. Anacamptis collina and A . pyramidalis extend east to Iran, and A. pseudolaxiflora extends east to Afghanistan and south to Yemen. (JW).




None recorded other than horticultural. (JW).



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