791. FRITILLARIA SORORUM Liliaceae. Martyn Rix and Kit Strange

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1 791. FRITILLARIA SORORUM Liliaceae Martyn Rix and Kit Strange Summary. Fritillaria sororum J. Persson & K. Persson from the Tauros mountains in southern Turkey is illustrated; its relationship with other species in the area is discussed. The genus Fritillaria consists of around 150 species and subspecies, found all round the northern hemisphere, with the exception of eastern North America. DNA studies, primarily made at Kew, have shown that the genus has evolved in two very distinct branches or clades (Day et al., 2014); one clade consists of the majority of the Eurasian species, extending from western Europe to Japan: the other clade consists of 20 species in North America, extending south into California and east to North Dakota, with three species in central and eastern Siberia and one (Fritillaria camschatcensis) on both sides of the northern Pacific. Within these clades there are five areas of particularly active evolution and clusters of species: in California; in Mediterranean Greece and Turkey; in Anatolia and the Zagros mountains; in central Asia from Uzbekistan to western Xinjiang, and in the eastern Himalaya in southwestern China. DNA studies show the species in each area to be related, and specification appears to be driven in response different pollinators. Even in the only distantly related Californian and Turkish species there are remarkably similar-looking pairs of species, F. pudica and F. carica for example, and F. purdyi and F. crassifolia. Notably the Californian, humming-bird pollinated, red-flowered F. recurva has no similar-looking counterpart in Eurasia. The species illustrated here, Fritillaria sororum, is found in a small area of southern Turkey, in the foothills of the Tauros mountains, an area very rich in Fritillaria species (Plate 791). It was first collected in fruit by Jimmy and Karin Persson at the end of April 1987 and described in The New Plantsman in 1998 from plants which flowered in Göteborg Botanical Garden in This species is named in gratitude of two sisters, Göta Curtis s Botanical Magazine 2014 vol. 31 (3): pp The Board of Trustees of the Royal Botanic Gardens, Kew 2014.

2 Plate 791 Fritillaria sororum joanna langhorne

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4 and Svea Blomqvist, who bequeathed half their estate to the Botanical Garden, Göteborg. Fritillaria sororum is easily recognised by its rounded and bell-shaped, tessellated and stippled flowers and particularly by the uppermost leaves, which are solitary, very long and modified into tendrils. The large dark ovate nectary, 7 11 mm long, and the very different shapes of the inner and outer perianth segments, suggest that this species is related to F. acmopetala Boiss., which is one of the most widespread of the eastern Mediterranean species, found from southwestern Turkey near Kalkan, along the coast, eastwards to the Lebanon and in Cyprus. It is distinct in its marked but not tessellated flowers, and the very wide, apiculate and recurved inner perianth segments, which often form a triangular entrance to the perianth. This species is found commonly, though not exclusively, in cornfields or on abandoned terraces, and it may be that in the past it was distributed as a cornfield weed. An unusual tessellated form of Fritillaria acmopetala was collected by Peter Davis and Oleg Polunin (D , E!, K!); it has a more narrowly campanulate flower, faint tessellation all over, and the upper leaves do not show any sign of tendrils, so appears distinct from F. sororum. The solitary uppermost leaves of Fritillaria sororum are particularly distinct; leaves modified into tendrils are not found in any other Turkish species, though they are found in numerous species in central Asia and China, as well as in F. ruthenica which extends from eastern Europe (Ukraine) to eastern Kazakhstan (Fig. 1). The tendrils of F. sororum are different from those of other species; they are solitary, long and narrowly strap-like; in other species the tendril-forming leaves are in whorls, and have a very narrow, coiling tip, and though their function is the same, the two types probably evolved independently. This concurs with the tendency mentioned above, for very similar characters to appear in widely separated branches of the genus. When studied in the wild, none of the upper leaves of F. sororum were seen to be coiling around supports, and in many cases the leaf tips had been eaten by adult lily beetles. However, the photograph in The New Plantsman (Persson & Persson, 1998) clearly shows the The Board of Trustees of the Royal Botanic Gardens, Kew

5 Fig. 1. Fritillaria sororum, showing coiling leaf tips, LCD , photographed at Kew. leaf of one plant coiled around the stem of another. F. sororum is also unusual in having smooth or minutely papillose stamen filaments. In most Mediterranean species stamen filaments are distinctly papillose (see Fig. 2). In spite of these distinctions, and taking into account their own extensive fieldwork, Tekşen & Aytaç (2011) included F. sororum as a synonym under F. acmopetala The F. acmopetala subsp. acmopetala specimens collected in the field have many individuals with tessellated perigon and smooth filaments. These flower features are very well-matched with F. sororum. F. sororum differs from F. acmopetala subsp. acmopetala by the uppermost leaves often with a tendril-like apex. This diagnostic character does not distinguish F. sororum from F. acmopetala subsp. acmopetala. The topotype specimens of F. sororum do not have tendril leaves (M. Tekşen 2019, 2021, 2108). All the characteristics fall within the variation of F. acmopetala subsp. acmopetala. In the phylogenetic tree published by Day et al. (2014), Fritillaria acmopetala subsp. acmopetala is well-separated from two 218 The Board of Trustees of the Royal Botanic Gardens, Kew 2014.

6 Fig. 2. Fritillaria sororum. A, habit, 1 / 7 ;B,bulb, 2 / 3 ; C, leaf tip from front, 10; D, leaf tip from side, 10; E, section through leaf tip, 10; F, l/s of outer tepal, 1 1 2;G,outer tepal from front, 1 1 2; H, inner tepal from front, 1 1 2; J, tip of tepal from front, 4; K, stamen from back, 3; L, tip of anther from front, 6; M, base of filament, 3; N, stigma, style and ovary, 1 1 2; P, stigma and style, 4 1 2; Q, capsule from side, 1; R, capsule from above, 1; S, seed from side, 5. All figures drawn by Joanna Langhorne from Kew LCD acc. no The Board of Trustees of the Royal Botanic Gardens, Kew

7 samples of F. sororum. However it must be admitted that the groupings indicated by DNA studies on Turkish and eastern Mediterranean Fritillaria species do not often coincide with groupings based on floral morphology. More testing of samples of known wild origin might clarify some of these anomalies. Another species of Fritillaria in southern Turkey is somewhat similar to F. sororum and in the phylogenetic tree published in Day et al. (2014), is indicated to be more closely related. This is F. whittallii Baker, which also has a tessellated, green and brown flower and narrow leaves, but has green rather than glaucous leaves and large distinct tessellations on the perianth segments, which are acute and almost equal. The flower of F. whittallii is also more angular, with square rather than rounded shoulders, and a much smaller nectary around 3 mm across. It also has papillose stamen filaments. F. whittallii is a characteristic species of Cedrus libani forest on limestone in Antalya and Isparta vilayet, (Byfield et al., 2010), well-separated from the two known populations of F. sororum. Because Fritillaria sororum is so rare that its exact locality is not mentioned in the original description, it is not given here. The valley where it grows is very isolated, the rock a confused mixture of schist and limestone. When we visited the type locality in early April 2002, flowering was well advanced. There were a reasonable number of non-flowering leaves, but only five flowering plants were seen in spite of careful searching. Associated species include Daphne sericea, Coronilla emerus subsp. emeroides, Cistus creticus, Paliurus spina-christi, Styrax officinalis, Quercus coccifera and Q. infectoria subsp. boissieri, aswellasdryopteris and various grasses. We failed to find the second, higher altitude locality. Another, as yet unnamed, species with small flowers was found nearby. Pollination has not been recorded in the wild, but is probably by queen wasps. Cultivation. Fritillaria sororum is cultivated in the collection at Göteborg and bulbs from the type collection are also at Kew, grown from seed from Göteborg. Seeds germinate readily in about 3 to 4 months from a winter sowing, generally appearing any time from May to July, the bulbs becoming mature in 220 The Board of Trustees of the Royal Botanic Gardens, Kew 2014.

8 three or four years. At Kew the bulbs are grown with a selection of like-minded fritillaries from the Mediterranean region, planted in clay pots and plunged in a free standing, raised sand bed. It is east facing and has a glass roof as a cover. The bulbs never get too hot as the frame affords plenty of shade and the plunge gets sprayed over in the summer, so that it never dries out completely. As with most species in Mediterranean climates, F. sororum is found growing inside deciduous bushes, which both protects the stems against grazing, and keeps the bulbs cool, so it is important that they should not become too hot nor dry out too much when dormant. In September they are re-potted into a loam-based compost with added grit and a little bit of coir to make it slightly moisture retentive. The bulbs are set on a small layer of river-washed sand to keep them snug. To start the bulbs in autumn, both pots and plunge are drenched once very thoroughly, and then up to the point the bulbs appear, only the plunge is watered when it feels dry. Only when the shoots and leaves are above ground are they watered freely. F. sororum, is generally helped along in the growing season with a few pea-sticks, to support its tall stems, and it can coil onto the pea-sticks with its tendrils. It can get very tall, in one year the plants reaching over 1m. They grow fast for about two months until they go dormant again in early May. In the absence of a number of herbarium specimens, the description below is adapted from Persson & Persson (1998). Fritillaria sororum J. Persson & K. Persson, The New Plantsman 5(2): 114 (1998). Type: Turkey: Içel; 500 m, meadow fragments adjoining brook ravine in Pinus brutia forest, on limestone, coll in fruit, cult , J. Persson & K. Persson (holo: GB). Description. Bulb to 3 cm across. Stem 25 45( 100) cm, glaucous, smooth. Leaves 6 10, all alternate, glaucous, linear, shallowly canaliculate, cm long, 7 10 mm broad, the uppermost around 12 cm long and tendril-like at the tip, sometimes coiling around a support. Flowers 1, rarely 2, broadly bell-shaped, rounded at the angle, glaucous outside, marked and tessellated, especially on the inside, with dark chocolate brown, often darker towards base and apex, without fascia; scent not recorded. Perianth segments 3 4 cm, the inner cm wide, broadly oblanceolate to obovate, with a slightly reflexed, apiculate and papillose apex, the outer narrowly ovate, cm wide. Nectary mm, The Board of Trustees of the Royal Botanic Gardens, Kew

9 narrowly ovate, brownish-black, 4 5 mm above the tepal base. Filaments 9 10 mm, slender, smooth or minutely papillose. Style mm, the branches 5 8 mm, slender, smooth, the branches recurved. Capsule cylindrical, not winged (Fig. 2). Chromosome number: 2n = 24. Distribution and habitat. Turkey: Içel; growing in grassy places in Quercus scrub and Pinus brutia forest, m, flowering in early April. REFERENCES Byfield, A., Atay, S. & Özhatay, N. (2010). Important Plant Areas in Turkey: 122 Key Turkish Botanical Sites. WWF Türkiye, Istanbul. Day, P.D., Berger, M., Hill, L., Fay, M.F., Leitch, A.R. & Kelly, L.J. (2014). Evolutionary relationships in the medicinally important genus Fritillaria L. (Liliaceae). Molecular Phylogenetics and Evolution 80: Persson, J. & Persson, K. (1998). Fritillaria sororum (Liliaceae): a new species from Turkey. The New Plantsman 5(2): Tekşen, M. & Aytaç, Z. (2011). The revision of the genus Fritillaria L. (Liliaceae) in the Mediterranean region (Turkey). Turkish Journal of Botany 35: The Board of Trustees of the Royal Botanic Gardens, Kew 2014.