LIMODORUM ABORTIVUM: Orchidaceae

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c. 2.7 mm long including the stout, 1.4 mm long filaments; anther flaps c. 1.3 mm long. Gynoecium urn-shaped, c. 2.3 mm long and 1.5 mm wide. Fruit sausage-shaped to obovoid, c. 7 mm long, bluish-black. Seeds not seen.

DISTRIBUTION. China: Sichuan, Hubei and Guizhou Provinces; 700-1200 m.

ACKNOWLEDGMENTS. The writer wishes to thank Mr John Bond VMH (Savill Garden, Windsor) for supplying material for illustration, and Mr Martin Gardner and Ms Sabina Knees (Royal Botanic Garden, Edinburgh) for their helpful discussions and assistance in the preparation of these notes.

REFERENCES

Ahrendt, L. W. A. (1961). Berberis and Mahonia. Journal of the Linnean Society, Botany 57: 1410.

Andrews, S. & Nelson, C. (1986). Augustine Henry’s plants in Kew Gardens. Kew Magazine 3: 136-140.

Lancaster, R. (1989). Trauels in China. Antique Collectors’ Club. Wood- bridge, Suffolk.

Loconte, H. & Estes, J. R. (1989). Phylogenetic Systematics of Berberidaceae and Ranunculales (Magnoliidae) . Systematic Botany 14: 565-579.

Rehder, A. (1929). Ligneous plants described by LCveillt. Journal of the Arnold Arboretum 10: 190.

230 and 23 1. LIMODORUM ABORTIVUM Orchidaceae

Jeffrey Wood

The majority of saprophytic orchids are generally rather small, often -pallid or dull brownish in colour and usually well camouflaged against the leaf-litter of their forest floor habitat. Limodorum abortiuum (L.) Sw., by contrast, is a tall, striking plant with brightly coloured flowers that cannot fail to impress all who encounter a colony in bloom in the dappled sunlight of a woodland setting.

The generic name is thought to be derived from the Greek leimon, a meadow, and doron, a gift, referring possibly to a meadow habitat, although it is more typical of woodland and scrub. Another explanation is that it is a literary error by the herbalist Dodonaeus, who applied to this genus the name haemodoron which was used by

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Theophrastus for a red-flowered parasitic plant of unknown identity. Limodorum abortivum and L. trabutianum Batt. are the sole members of

the genus. Limodorum trabutianum is found only in south-west Europe and western North Africa, extending northwards to the French Atlantic coast and eastwards to the Italian island ofpantelleria offthe Tunisian coast. I t is a less attractive plant and looks somewhat like a peloric form of L. abortivum. It is distinguished by the narrower spathulate lip which is not constricted near the base, and the shorter spur only 1-4 mm long. It may often be found growing alongside L. abortivum and some would argue that it should perhaps be given varietal status only.

Plants of Limodorum abortivum sometimes occur singly, but more often form colonies, the emergent inflorescences reminding one of asparagus shoots. Some individuals may be very robust, attaining 80 cm in height, while others may be quite slender and graceful in habit. In suitable locations it is not unusual to find several groups of flowering stems. Ifwinter and early spring rainfall, particularly in the Mediterranean part of the range, should be inadequate, plants suffer and stems are often little more than 20 cm tall. Such plants appear dingy and have flowers that barely open and are quite unlike the forms illustrated here.

Some populations are more floriferous than others. Plants occur- ring in rough grassy scrub at 600 m in the southern limestone foothills ofthe CCvennes in southern France, for example, are short in stature, but have dense floriferous spikes of wide-opening lilac-pink flowers (see Plate 231, right). These are found between low bushes of the legume Dorycnium pentaphyllum subsp. gracile, associated with orchids such as Anacamptis pyramidalis, Cephalanthera rubra, Himantoglossum hircinum, Ophrys aplfra and 0. scolopax.

The flowers of Limodorum are thought to be exclusively self- pollinating and vary somewhat in shade and intensity of colour. The usual coloration is reddish-violet to violet with some yellow on the lip. Plate 230 shows a specimen found growing in acidic soil on the beautiful Massif des Maures between Toulon and St Tropez in the Provenqal department of Var in southern France. Some populations in Cyprus and Turkey (Anatolia) have dark rose-red flowers and have beendistinguished asvar. rubrumH. Sund. (seeplate 231, left). Plants with paler violet as well as white flowers occasionally occur. Plants from Turkey which have somewhat smaller flowers have been called var. anatolicum C. Koch.

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Limodorum abortivum

Plate 230

CHRISTINE HART-DAVIES

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Limodorum abortivum grows in a variety of habitats and, although preferring calcareous soils, is sometimes found on some igneous formations. I t grows most frequently in scrubby places, along woodland margins and on shady banks under pine trees. In Cyprus, it is often found under Pinus brutia and ascends to 1,750 m in the Troodos Mountains. Populations occur as high as 2,300 m in Turkey and south-west Iran where they are frequently found growing under oak, beech and hornbeam.

CULTIVATION (by Sandra Bell). Attempts have been made to bring Limodorum abortivum into cultivation, but none has been successful as far as I am aware. This is hardly surprising, given the plant’s unusual mode of nutrition. Limodorum is ‘usually described as saprophytic, deriving nutrients from the breakdown of organic matter in the soil. Its cluster of short, fleshy roots is heavily impregnated with a mycorrhizal fungus and it has been suggested (E. & 0. Danesch, 1984: 132-3) that this makes the breakdown products available to the orchid in a way similar to that known to occur in the bird’s nest orchid Neottia nidus-avis. Neottia, however, grows in soils rich in organic matter whilst Limodorum grows in soils conspicuously lacking in leaf-mould which suggests that it may not be so much a saprophyte but an epiparasite like the two Australian underground orchids, Rhizanthella gardneri and R. slateri. R. gardneri has been cultivated successfully in Australia, where, in the wild, it only occurs within the root-spread of the tree species Melaleuca uncinata (Myrtaceae) . At the University of Adelaide researchers inoculated roots of M. uncinata with a mycorrhizal fungus which had been isolated from roots of Rhizanthella gardneri, then scattered seeds of the underground orchid around the tree. Germination was achieved and the plants flowered within 18 months (Dixon et al.,

It may well be shown in future that Limodorum abortivum can be grown following a similar method with perhaps a species of pine as the host tree. In the meantime it cannot be recommended as a subject for cultivation and must be admired in the wild.

1990).

Limodorum abortivum (L.) Sw. in Nov. Act. Reg. SOC. Sci. Upsal. 6: 80 ( 1 799). Orchis abortiva L., Sp. Pl., 943 (1753). Type: ‘in Gallia, Helvetia, Anglia,

Serapiasabortiva (L.) Scop., F1. Cam. (edn. 2) 11: 205 (1772). Epipactisabortiva (L.) All., F1. Pedem. 11: 151 (1785).

I talia’.

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Neottia abortiva (L.) Claim., Mant., 264 (181 1 ) . Centrosisabortiva (L.) Sw., Summ. Veg. Sc., 32 (1814).

Jonorchis abortiva (L.) Beck, F1. Nied.-Oest., 215 (1890). Lequeetia abortiva (L.) Bubani, F1. Pyr., 58 (1901). Limodorum abortivum (L.) Sw. var. anatolicum C. Koch in Linnaea 19: 12

(1847). Type: unlocated. L. abortivum (L.) Sw. var. rubrum H. Sund., Europ. Medit. Orch. (edn. 2),

201 (1975), nom. nud.

DESCRIPTION. Terrestrial, saprophytic or possibly epiparasitic herb (20-)40-75(-80) cm high. Rhizome short, stout, up to 1 cm thick, with

Limodorum abortivum. A, dorsal sepal, X 3; B, lateral sepal, X 3; C, petal, X 3; D, lip, column and spur, X 3; E, column, X 6; F, lip with spur, X 3. Drawn by Sarah Thomas.

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Limodorum abortivum

Plate 231

CHRISTINE HART-DAVIES

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numerous thick, fleshy roots. Stem violet, violet-red or bluish green, erect, robust, thick, rigid, bearing numerous violet, violet-red or bluish violet, ovate, obtuse, rarely acute sheaths possessing very little or no chlorophyll. Green leaves absent. Injorescence a 4-25-flowered, elongate spike-like raceme up to 35 cm long, lax to dense; floral bracts bluish green usually flushed with violet, oblong-elliptic, acute or acuminate, concave, membranous, up to 4 cm long, the lowermost exceeding the ovary, the uppermost a little shorter than the ovary. Sepals pale to dark violet, flushed greyish violet on the exterior, rarely rose-pink or white, erect or slightly spreading; dorsal sepal ovate to obovate, obtuse, strongly concave, sparsely papillose on the undersurface, erect or curved forward, forming a hood over the column, 20-25 mm long, 7-1 1 mm wide; lateral sepals obliquely ovate to ob- lanceolate, acute or subacuminate, flat or slightly concave, glabrous or only very sparsely papillose on the undersurface, 16-20 mm long, 5-7 mm wide. Petals pale violet towards the apex, whitish towards the base, narrowly elliptic to linear, acute, glabrous, spreading, 16-19 mm long, 3-4 mm wide. Lip violet, fading to dingy yellow with age, rarely rose-pink or white, adnate to the base of the column, distinctly constricted between the hypochile and epichile, 1&15(-17) mm long, 7-12 mm wide; hypochile whitish or pale violet, deltoid, porrect, parallel to the column; epichile bright violet with darker violet longitudinal nerves and often a yellow flush at the centre, entire, triangular-ovate to oblong, obtuse, sometimes obscurely 2-lobed at the apex, curved, margin slightly crenulate-undulate, turned upwards, particularly towards the apex, glabrous; spur pale violet or whitish, cylindrical, slender, obtuse, downturned, c. 15(-17) mm long, seldom shorter, rarely absent. Column erect, slender, 13-17 mm long; anther oblong, 5 mm long, 3 mm wide; ovary shortly pedicellate, subclavate- cylindrical, not twisted, glabrous, 1.5-3 cm long; pedicel 1 cm long, much enlarged after anthesis. Ripe capsule up to 4.5 cm long.

DISTRIBUTION. Central, southern and south-east Europe, northwest to Belgium and Germany, east to Turkey, Cyprus, Syria, Lebanon, Crimea, Caucasus, Iraq and western and northern Iran, south to North Africa.

HABITAT. Dry, grassy places, scrub, open coniferous and mixed woodland, woodland margins, on calcareous and igneous soils; sea level to 2,300 m.

BIBLIOGRAPHY

Camus, E. G. & A. (1928). Iconographie des Orchidies d’Europe et du Bassin

Danesch, E. & 0. (1984). Les Orchidies de Suisse. Editions Silva, Zurich. Dixon, K. W., Pate, J. S. & Kuo, J. (1990). The Western Australian fully

subterranean orchid, Rhizanthella gardneri. In Arditti, J. (ed.), Orchid Biology, Reviews and Perspectives V: 37-62. Timber Press, Portland, Oregon.

Renz, J. & Taubenheim, G. (1984). Orchidaceae. In Davis, P. H., (ed.), Flora of Turkey and the East Aegean Islands 8: 45&627. Edinburgh Univ. Press.

Miditerranien. Paul Lechevalier, Paris.

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Ruckbrodt, U. & D., & Hansen, K. & R.-B. (1992). Bemerkungen zu den in der Tiirkei vorkommenden Orchideenarten und ihrer Verbreitung. Ber. Arbeitskr. Heim. Orch. 9( 1): 4-103.

Wood, J. J. (1985). Orchidaceae. In Meikle, R. D., Flora of Cyprus 2: 1491-1558. Bentham-Moxon Trust, Kew.

TWO CULTIVATED SPECIES O F BOUVARDIA

Daniela Zappi

The genus Bouvardia in the Rubiaceae was described by Salisbury in homage to Dr Charles Bouvard, Superintendent of the Jardin des Plantes, Paris. Placed in the tribe Hedyotideae by Bremekamp (1952), together with Houstonia, Hindsia and Mannetia, it comprises small shrubs or climbers with showy, tubular, tetramerous flowers and many, winged seeds per capsule. The genus was revised by Blackwell ( 1968), who accepted 3 1 species, ranging from the south- western United States to northern Central America. However, its main centre ofdiversity is in Mexico, at elevations above 500 metres. The habitats of Bouvardia vary from semi-desert to relatively mesic ravines covered by oak and pine forest.

Blackwell ( 1968) accepted the classification proposed by Schlechtendal (1854), dividing the genus in three subgenera: subgenus Bougardiastrum, characterized by opposite leaves and many-flowered inflorescences with tubular, variously coloured corollas, and including B . Zaevis and 14 other species; subgenus Bouvardioides, comprising 8 species with opposite leaves and mostly solitary flowers with white, salverform corollas; and subgenus Bouvardia, with 3-6 verticillate leaves at each node, where B. terniflia is placed alongside 7 other species of ‘typical bouvardias’.

First introduced to Britain in 1794 by Sir Joseph Banks, B . ternifolia has already been illustrated twice before in Curtis’s Botanical Magazine, but not under its correct name. The first time was in 1817, as B . triphylla Salisbury (t. 1854), based probably on the same introduction as the type of B . terniflia, with somewhat orange flowers and, later, as a new species, B . splendens Graham ( 1840). This time it was drawn by W. H. Fitch and based on a much more attractive and vigorous specimen of unknown origin obtained from the Horticultural Society’s Chiswick Garden by James McNab in 1838. On this occasion, the plant received the following exaltation: ‘There is scarcely any thing in cultivation more brilliant than a large

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