tion of the soil seem critical for most succulents, with each species having additional requirements such as altitude and aspect. By far the most favoured soils for succulent Asclepiads seem to be those derived from red granite with a high mica content, and as these are not very widespread in the south west where most succulents grow; this further limits their distribution. Ceropegias are also sensitive to waterlogging and, being palatable to livestock, must also have some protection from grazing, such as hiding among distasteful or thorny shrubs.

With experience, a glance can identify those areas which may harbour succulents, and because some species are so exacting in their requirements, can predict the particular species as well. Many ceropegias, especially those that grow on the Tihama plain, will not tolerate competition from plants such as grasses, Pentatropis nivalis or Sarcostemma species. However, species growing at higher altitudes are not only tolerant of more shade, but will also grow happily among thick bushes of Euphorbia schimperi, E. cactus and the tangled stems of Cissus rotundifolius.

One of the joys of Ceropegia-hunting in Saudi Arabia is the knowledge that in the very next unexplored wadi there may be a new population or even a new species. Unfortunately many of these potential sites may have already been destroyed.

REFERENCES

Bruyns, P. V. (1988). Studies in the Flora of Arabia XXIV: The genus Ceropegia in Arabia. Notes Roy. Bot. Card. Edinb. 45: 287-326.

Chaudhary, S. A. & Lavranos, J. J. (1985). Studies in the Flora of Arabia XVI: Two new species of Ceropegia from.Saudi Arabia. Notes Roy. Bot. Card. Edinb. 42: 315-319.

Collenette, I . S. (1985). An illustrated guide to the JIowers of Saudi Arabia. Scorpion Publishing Ltd., Buckhurst Hill.

Field, D. V. & Collenette, I. S. (1984). Ceropegia superba (Asclepia- daceae), a new species from Arabia. Kew Bull. 39: 639-642.

Huber, H. (1957). Revisionder Gattung Ceropegia. Mem. SOC. Brot. 12: 138.

A CLOSE LOOK AT A CRETAN DAPHNE

Nicholas Turland

Until now, two species of Daphne were known to occur on Crete: D. oleoides Schreber and D. sericea Vahl. However, in Michel Gan- doger’s Flora Cretica (1916), there is a record for D. gnidioides Jaub. &

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Spach, .based on a collection by Cousturier at Patsianos in the Sfakia district in the south-west of the island. This record was never confirmed, and no further sites were found before the publication of Flora Europaea vol. 2 in 1968, so the species-distribution given in that work omits mention of Crete. This article confirms the occurrence of D. gnidioides on the island, and provides comparisons with D. oleoides and D. gnidium L., both ofwhich are widespread in the Mediterran- ean area, although the latter is unknown on Crete. A detailed description of D. gnidioides is given, along with notes on the Cretan localities found to date.

The story begins in April 1984 when I was participating in a botanical holiday based at Plakias on the south coast of western Crete, led by Lance Chilton. Among the plants we found was a Daphne species growing on Akrotiri Kakomouri at the southern end of Plakias bay. Initially, the exact identity of this plant eluded us, but when a few terminal flower-clusters were noticed on one of the bushes, we found that it keyed out to D. oleoides in Flora Europaea. In spite of this provisional determination, Lance Chilton noted that the plants seemed closer to D. gnidium in general appearance, and their size (up to 2 m tall) was unusually large for D. oleoides.

Three years passed, during which time I led a number of botanical holidays myself at Plakias and continued to assume the Daphne to be D. oleoides. No further observations were made, except that the flowering period appeared, curiously, to be September to November ( D . oleoides is summer-flowering) . The flowers seen in late April 1984 were presumably out-of-season, since not even the first visible stages of bud-development were observed in subsequent springs (as late as mid-May), nor was evidence found in autumn of plants having flowered during the summer. However, in May 1987, while climbing Mount Kedhros in south-western central Crete, I found what was undoubtedly true D. oleoides growing at an altitude of approximately 1,500 m. These plants were obviously very different from those near Phkias (even though they were without flowers) and so I began to suspect the latter to be something hitherto unknown on Crete, since it clearly did not belong to the remaining Cretan taxon, D. sericea. The main differences between the Plakias plant and D. oleoides are outlined in Table 1.

In late October 1987 I took a close look at the plants growing on Akrotiri Kakomouri and collected flowering and fruiting herbarium specimens. Almost all of the several tens of individuals present were

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TABLE 1

Plakias plants Daphne oleoides

to c. 2 m high

flowering September to November

perianth-lobes broad, obtuse

altitude c. 5 1 2 0 m

to c. 50 cm high

flowering between April and August, depending on altitude

perianth-lobes narrow, acuminate

altitude (450-) 1,400 m upwards

in full flower with up to 18 creamy white flowers in each terminal head. Sometimes there were also juicy, orange-red berries co- existent with the flowers. The plants gave the immediate impression of being very much like D.gnidium but without the paniculate inflorescences characteristic of this species. I sent material to Mr J. R. Press at the British Museum .(Natural History), who in turn sent it on to Kew, where Brian Mathew determined it as D . gnidi- oides, ruling out D. gnidium for the reasons outlined in Table 2.

In late October 1988 I collected further herbarium material from the four populations then known to me (all within 8 km of Plakias, see following notes on distribution). Thespecimens weresubsequent- ly taken to Kew, where Brian Mathew and I compared them with the D. gnidioides already present there - all Turkish, except for two

TABLE 2

Daphne gnidioides Daphne gnidium

stems comparatively sparsely leafy; leaves erecto-patent to patent

leaves, on average, comparatively shortinrelationtowidth (ratio3.5

inflorescence adense terminal head (occasionally with axillary flower- clusters below), not paniculate

flowers comparatively large: peri- anth-tubes up to 7 mm long; peri- anth-lobes up to5 mmlongand 4.5 mmwide

stems moredensely leafy; leaves mostly erecto-patent

leaves, on average, longer in rela- tiontowidth (ratio4to 12.1 : 1)

inflorescence paniculate

to9.1: 1)

flowers smaller: perianth-tubes up to 5 mm long; perianth-lobes up to c. 3 mm long and 2 mm wide

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collections from the East Aegean islands. There was a small degree of variation among my Cretan plants, with few-flowered, relatively long- and acute-leaved specimens from one population and com- paratively many-flowered, short- and obtuse-leaved ones from the other three. However, there was even more variation (including forms comparable with the two just described) among the Kew specimens. We concluded that although the Cretan plants were

Map to show the distribution ofD. gnidioides.

referable to D. gnidioides, they bore two apparently constant features which appeared to render them different from Turkish plants. Firstly, flowering season: material from Turkey had been collected with flowers in May, June, July, August and October, whereas, as mentioned earlier, Cretan ones flowered almost exclusively between September and November. One could speculate that the Turkish plants also have their main flowering period during the autumn, the spring and summer collections having been taken from unusual individuals which happened to be bearing out-of-season flowers at the time of the collectors’ visits. Secondly, altitude: Turkish plants had been collected from sea-level to 1,200 m, whereas I had never seen Cretan ones above c. 120 m (and know the Plakias region of

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south-western Crete in considerable detail). The possibility of higher altitude populations being found on Crete cannot be ruled out, however.

The treatment of D. gnidioides in Flora Europaea is rather mislead- ing, probably as a result of the species having been so little known and seldom collected in the years prior to 1968. Faced with a number of specimens, one would be almost certain to find features which disagreed with the description. The account in Flora of Turkey, however, is more accurate.

The currently known distribution on Crete is as follows: Eparchy of Aghios Vasilios, 1.5 km south-west of Kato Rodhak-

ino, headland at western end of Koraka beach; rocky, limestone phrygana; associated with Ceratonia siliqua L. and Pistachia lentiscus L.; altitude c. 5-20 m.

Ep. Ag. Vasilios, 1 km north of Plakias, by Kotsifos stream; olive groves and scrub over stony, calcareous stream-deposits; associated with Callicotome villosa (Poiret) Link and Ceratonia siliqua; altitude c. 15 m.

Ep. Ag. Vasilios, southern end of Plakiasa bay, Akrotiri Kakom- ouri; sand dunes, grazed, rocky, limestone phrygana and crevices in limestone cliffs; altitude c. 5-120 m.

Ep. Ag. Vasilios, 3 km south-east of Plakias, headland between Damnoni and Amoudi beaches; grazed, rocky, limestone phrygana and crevices in limestone cliffs; altitude c. 5-20 m.

All four populations occur within a small area of south-western Crete. Despite having visited seemingly suitable habitats in many other parts of the island I have yet to find the plant outside this range. Gandoger's collection from Mt. Trocharis, a 2,401 m peak in the White Mountains of western Crete (Rechinger, 1943), is almost certainly referable to D. oleoides because of the high altitude.

CULTIVATION. The Alpine Garden Society handbook Daphne, the Genus in the Wild and in Cultivation (Brickell & Mathew, 1976) gives the impression of an insignificant species of little value to horticulture but possibly of interest to collectors. However, the information included was based upon existing literature, which in turn was probably based upon dried herbarium specimens rather than living material. Certainly the more floriferous Cretan plants are most attractive at anthesis, and have the added distinction of a somewhat sickly-sweet fragrance. They definitely rival D. gnidium, which is occasionally cultivated. I collected two seeds from Akrotiri Kakom-

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Daphne gnidioides A, habit; B, flowering branch; C, flower; D, half-flower; E, stamens; F, fruit. Drawn by Christine Grey-Wilson.

ouri in autumn 1986, both germinated and one plant is still alive, now growing in my glasshouse border and measuring 60 cm tall (December 1990). The degree of hardiness is still unknown but, in view of the low altitudes of the Cretan populations, I should think a sunny, sheltered site with a light, well-drained soil would be necessary if the plant is to be tried outdoors in a cool-temperate climate. I know that D. gnidium will grow outside in southern England (B. Mathew, pers. comm.), so Cretan D. gnidioides may well prove similarly tolerant, although I doubt if cold regions of Britain would be suitable, or indedd comparable areas further afield. Seedlings and young plants would probably be more susceptible to cold than established ones, so raising in a frost-free environment during the early years would be advisable. Plants grown from seed collected at the upper altitudinal limit ofthe species in Turkey might prove less vulnerable, however.

Daphne gnidioides Jaub. & Spach, Ill. P1. Orient. 4: 4, t.304 (1850); Meissn. in DC., Prodr. 14: 536 (1857)) Boiss., F1. Or. 4: 1049 (1879); Rech. fil., FI. Aegaea 392-393 (1943); Webb & Ferguson in Tutin et al. (eds.), F1.

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Europaea 2: 256-258 (1968); Brickell & Mathew, Daphne 107-108 (1976); Tan in Davis (ed.), F1. Turkey 7: 521-526 (1982). Type: Jaubert & Spach cite 3 syntypes 'mountains of Leros, Olivier 6' Bruguiire (P); old ruins of Samos, D'Urville (P) and dry coastal areas of Lydia and Caria, Olivier 6' Bruguiire ( P) . ' D. oleoides sensu d'Urv. in Mtm. SOC. Linn. Paris 1: 299 (1822) non

Schreber, Ic. Descr. P1. 1: 13, t.7 (1766).

The following description is based upon (1) living plants both in the wild and in cultivation; (2) my herbarium specimens from Crete and (3) herbarium material a t K and BM.

DESCRIPTION. Much-branched shrub, up to c. 3.5 m high. Twigs slender, usually straight and erect, rounded in cross-section, brown, reddish-tinged (or green) and white-puberulent when young. Bud-scales triangular-ovate, usually less than 1.5 mm long, margin white-puberulent. Leaves alternate, patent to erecto-patent, carried on twigs up to 2 years old; blade normally mid-green above, minutely pale-dotted beneath (particularly conspicuous when dry), glaucous and odorous when young, oblanceolate to narrowly elliptic, gradually tapering towards the base, obtuse-rounded to acute at the apex, mucronate, (lo-) 15-35(-45) mm long, (2-)2.5-6.5(-9) mm wide, leathery, glabrous (or with a few hairs when very young); mucro sharply pointed, often slightly recurved, up to 1.5 mm long; margin translucent, cartilaginous, entire, often slightly revolute; mid-vein impressed above and raised beneath throughout its length; lateral veins f obscure; petiole short, articulate at node, indistinct from blade, often twisted through up to 90" so that blade is held vertically. (N.B. a few leaves immediately below inflores- cence rarely subglabrous, the uppermost occasionally reduced in size, sometimes greatly so (down to c. 4.5 mm long and 0.5 mm wide), when sparsely white-pubescent and bract-like.) Injorescence terminal, a dense rounded head of 1 to c. 18 sickly-sweet-scented flowers (occasionally also axillary, with clusters of 1 -few flowers below the terminal head), appearing sometime between April and November. Bracts few, falling off early, no longer than much-reduced uppermost leaves; outer lanceolate, acute, with white-pubescent margin; inner becoming ovate, obtuse and densely white-pubescent on outer surface except along mid-vein. (There appear to be no clear-cut discontinuities between much-reduced uppermost leaves, outer bracts and inner bracts.) Pedicels very short and erect at bud-stage, often progressively elongating to a maximum of c. 7 mm and becoming patent to pendent as flowers (and subsequently fruits) develop, densely white-pubescent. Perianth hypogynous, actinomorphic, waxy-textured, sub- persistent; tube whitish, darkening to yellowish and then brown with age, narrowly ovoid, up to 7 mm long and 2.5 mm wide, 8-veined, densely white- pubescent outside: lobes 4, petaloid, patent to slightly recurved, concolorous with tube, very occasionally pinkish-tinged, oblong- to ovate-orbicular, sometimes very shortly cuspidate, up to 5 mm long and 4.5 mm wide, densely white-pubescent on reverse. Stamens 8, in 2 whorls, attached to inner surface of perianth-tube; filaments white, 0.5-0.75 mm long, glabrous;

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anthers basifixed, yellow, 0.75-1.25 mm long, 0.3-0.5 mm wide, glabrous, the upper whorl barely exserted from perianth- tube. Hypogynousdiscannular. Ovary concealed by but free from perianth-tube, greenish to brownish*, ovoid, c. 2.5 mm long and 1.5 mm across, densely silvery-sericeous. Style brownish*, c. 0.5 mm long, glabrous. Stigma brownish*, k hemispherical with style attached to centre of flattened underside, 0.75-1 mm across, densely pale-papillose on upper surface. Fruit a berry, enclosed by perianth-tube until ripe, orange-red, ovoid, c. 9 mm long and 7 mm wide, fleshy and juicy, sparsely sericeous, often borne simultaneously with flowers and buds in the same inflorescence. Seeds 1 per fruit, pyriform, c. 5 mm long and 3 mm wide; surface grey-brown with pale patches, somewhat shiny with a dark brown basal scar and light brown very low lateral ridge.

DISTRIBUTION. Greece (Crete, East Aegean islands of Samos, Leros, Kalimnos, Kos, Nisiros and Tilos, Megiste (Kastellorizo) island, ? Asti- palaia, ? Evvoia, ? Skiathos); S and SW Turkey (Izmir, Mugla, Denizli, Burdur, Antalya, Iqel and Maras vilayets); altitude c. 5-120 m (Crete), sea- level to c. 1,400 m (Turkey).

HERBARIUM SPECIMENS SEEN. TURKEY: Mugla: Sogut to Bozburun, s.1.-50 m, 1965, Davis41 167 (K); CalDag, Sslope, 35 kmNofFethiye, 1968, Lambert &? T h o 9 556 (K); Sandras Dag, at Agla, 1947, Davis 13619 (K); Fethiye to Kalkan, near Xanthus, 1962, Dudley 35 17 1 (K); Yilani Dag, Hikmet Birand30 (K) . Antalya: Finike, 1,200 m, 1960, Khan, Prance &? Ratclijjfe 255 (K) ; 1958, P. B. Smith 259 (K); c. 20 kin W of Kizilcadag on Fethiye road, 1970, Fraser Jenkins 2142 (BM). Iqel: Anamur, 1872, Peionin 89 (K, BM). E. AEGEAN IS: Kos, Mt. Dhikion, above Asfendiou, 1965, Davis 40474 (K); Kos, between Pili and Kardamena, 1935, Rechinger 7890 (BM); Kos, Mt. Dikaio, c. 600 m, 1935, Rechinger 8027 (BM); Kos, Aucher-Eloy 2514 (K, BM). Samos, near Tigani, c. 200 m, 1934, Rechinger3621 (BM). A specimen without collector’s name (?Forbes) was labelled ‘ruins ofcnidus, Caria, 1841’ (K).

My own Cretan material is from: Koraka beach headland, Plakias olive groves, Akrotiri Kakomouri and Damnoni-Amoudi beach headland. All specimens were collected in October 1988 and are now at K except for the October 1987 specimens from Akrotiri Kakomouri which are at BM.

ACKNOWLEDGMENTS. I should like to thank Lance Chilton, Brian Mathew and Bob Press for their help and information which have been invaluable to me in the writing of this article.

REFERENCES

Alden, B. (1986). Daphne L. In Strid, A. (ed.) Mountain Flora of Greece 1:

Boissier, P. E. (1879). Flora Orientalis 4: 1049. Basel, Geneva & Lyon.

*Colours may differ on fresh material. White indumentum may become

59 1-592. Cambridge University Press.

brownish on or after drying.

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Brickell, C. D. & Mathew, B. (1976). Daphne, the Genus in the Wild and in

Gandoger, M. ( 19 16). Flora Cretica 9 1. Paris. Jaubert, C. & Spach, E. (1850-53). Illustrationes Plantarum Orientalium 4: 4,

Meissner, C. F. (1857).\Daphne L. In de Candolle, A. P. Prodromus Sys-

Rechinger, K. H. (1943). Flora Aegaea 391-393. V‘ ienna. Schreber, J. C. D. von (1766). Icones et Descriptiones Plantarum 1: 13, t.7.

Halle. Tan, Kit (1982). Daphne L. In Davis,’P. H. (ed.), Flora of Turkey 7: 521-

526. Edinburgh University Press. d’urville, J. S. C. D. (1822). Enumeratio plantarum quas in insulis Archi-

pelagi aut littoribus Ponti-Euxini, armis 1819 et 1820, collegit atquae detexit J. Dumort d’Urville. Mim. SOC. Linn. Paris 1: 299.

Webb, D. A. & Ferguson, I. K. (1968). Daphne L., in Tutin, T. G. et al. (eds.) Flora Europaea 2: 256-258. Cambridge University Press.

Cultivation 107-108. Alpine Garden Society, Surrey.

t.304. Paris.

[ematis naturalis Regni vegetabilis 14: 536. Paris.

A SUPPLEMENTARY NOTE O N EUPHORBIA SCHILLING11 AND A NEW VARIETY

Alan Radcliffe-Smith

Euphorbia schillingii, which was first described in The Kew Magazine 4: 112, pl. 80 (1987), was discovered in central Nepal by Tony Schilling who subsequently introduced it into cultivation. In the 14 years in which it has been cultivated it has proved to be an easy, adaptable and attractive species, which is now available commercially. Unlike its cousins, E. wallichii and E. sikkimense, E. schillingii forms discrete clumps and is non-invasive. For this reason it is a better plant for the smaller garden.

Since its discovery a distinct variant ofE. schillingii has been found, also in central Nepal, but further west than the type plant. It is here described as var. praecox. This variety is less garden-worthy than its close relations on account of the less vividly yellow-green flowering heads. I t flowers at least a month earlier than typical E. schillingii however, and produces fruits which are slightly more warty, from early mid-July onwards. Living material was brought back in 1979 by Stephen Wright, and grown on at Wakehurst Place, where it thrives on a moisture-retentive peaty soil on a slope in the ‘Himalayan Glade’. Euphorbia schillingii A. Radcliffe-Smith in Kew Mag. 4: 112, pl. 80 (1987). Type: Nepal, Schilling 2060 (holotype K).

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