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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 sub- species, 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 particu- larly 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 stud- ies 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 1995. This species is named in gratitude of two sisters, Göta Curtis’s Botanical Magazine 2014 vol. 31 (3): pp. 214–222 214 © The Board of Trustees of the Royal Botanic Gardens, Kew 2014.

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Page 1: 791. FRITILLARIA SORORUM

791. FRITILLARIA SORORUMLiliaceae

Martyn Rix and Kit Strange

Summary. Fritillaria sororum J. Persson & K. Persson from the Taurosmountains in southern Turkey is illustrated; its relationship with otherspecies in the area is discussed.

The genus Fritillaria consists of around 150 species and sub-species, found all round the northern hemisphere, with theexception of eastern North America. DNA studies, primarilymade at Kew, have shown that the genus has evolved in twovery distinct branches or clades (Day et al., 2014); one cladeconsists of the majority of the Eurasian species, extending fromwestern Europe to Japan: the other clade consists of 20 speciesin North America, extending south into California and east toNorth Dakota, with three species in central and eastern Siberiaand one (Fritillaria camschatcensis) on both sides of the northernPacific. Within these clades there are five areas of particu-larly active evolution and clusters of species: in California; inMediterranean Greece and Turkey; in Anatolia and the Zagrosmountains; in central Asia from Uzbekistan to western Xinjiang,and in the eastern Himalaya in southwestern China. DNA stud-ies show the species in each area to be related, and specificationappears to be driven in response different pollinators. Even inthe only distantly related Californian and Turkish species thereare remarkably similar-looking pairs of species, F. pudica andF. carica for example, and F. purdyi and F. crassifolia. Notably theCalifornian, humming-bird pollinated, red-flowered F. recurvahas no similar-looking counterpart in Eurasia.

The species illustrated here, Fritillaria sororum, is found ina small area of southern Turkey, in the foothills of the Taurosmountains, an area very rich in Fritillaria species (Plate 791).It was first collected in fruit by Jimmy and Karin Persson at theend of April 1987 and described in The New Plantsman in 1998from plants which flowered in Göteborg Botanical Garden in1995. This species is named in gratitude of two sisters, Göta

Curtis’s Botanical Magazine 2014 vol. 31 (3): pp. 214–222214 © The Board of Trustees of the Royal Botanic Gardens, Kew 2014.

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Plate 791 Fritillaria sororum joanna langhorne

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and Svea Blomqvist, who bequeathed half their estate to theBotanical Garden, Göteborg.

Fritillaria sororum is easily recognised by its rounded andbell-shaped, tessellated and stippled flowers and particularly bythe uppermost leaves, which are solitary, very long and modifiedinto tendrils. The large dark ovate nectary, 7–11 mm long, andthe very different shapes of the inner and outer perianth seg-ments, suggest that this species is related to F. acmopetala Boiss.,which is one of the most widespread of the eastern Mediter-ranean species, found from southwestern Turkey near Kalkan,along the coast, eastwards to the Lebanon and in Cyprus. It isdistinct in its marked but not tessellated flowers, and the verywide, apiculate and recurved inner perianth segments, whichoften form a triangular entrance to the perianth. This speciesis found commonly, though not exclusively, in cornfields oron abandoned terraces, and it may be that in the past it wasdistributed as a cornfield weed.

An unusual tessellated form of Fritillaria acmopetala was col-lected by Peter Davis and Oleg Polunin (D. 26044, E!, K!); it hasa more narrowly campanulate flower, faint tessellation all over,and the upper leaves do not show any sign of tendrils, so appearsdistinct from F. sororum.

The solitary uppermost leaves of Fritillaria sororum are partic-ularly distinct; leaves modified into tendrils are not found inany other Turkish species, though they are found in numerousspecies in central Asia and China, as well as in F. ruthenica whichextends from eastern Europe (Ukraine) to eastern Kazakhstan(Fig. 1). The tendrils of F. sororum are different from those ofother species; they are solitary, long and narrowly strap-like; inother species the tendril-forming leaves are in whorls, and havea very narrow, coiling tip, and though their function is the same,the two types probably evolved independently. This concurs withthe tendency mentioned above, for very similar characters toappear in widely separated branches of the genus. When stud-ied in the wild, none of the upper leaves of F. sororum were seento be coiling around supports, and in many cases the leaf tipshad been eaten by adult lily beetles. However, the photograph inThe New Plantsman (Persson & Persson, 1998) clearly shows the

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Fig. 1. Fritillaria sororum, showing coiling leaf tips, LCD 2007–1838, photographed at Kew.

leaf of one plant coiled around the stem of another. F. sororumis also unusual in having smooth or minutely papillose stamenfilaments. In most Mediterranean species stamen filaments aredistinctly papillose (see Fig. 2).

In spite of these distinctions, and taking into account theirown extensive fieldwork, Teksen & Aytaç (2011) includedF. sororum as a synonym under F. acmopetala … ‘The F. acmopetalasubsp. acmopetala specimens collected in the field have manyindividuals 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 theuppermost leaves often with a tendril-like apex. This diagnosticcharacter does not distinguish F. sororum from F. acmopetalasubsp. acmopetala. The topotype specimens of F. sororum donot have tendril leaves (M. Teksen 2019, 2021, 2108). All thecharacteristics 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.

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Fig. 2. Fritillaria sororum. A, habit, × 1/7; B, bulb, × 2/3; C, leaf tip from front, × 10; D, leaftip from side, × 10; E, section through leaf tip, × 10; F, l/s of outer tepal, × 11∕2; G, outertepal from front, × 11∕2; H, inner tepal from front, × 11∕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, × 11∕2; P, stigma and style, × 41∕2; Q, capsule from side, × 1; R, capsule fromabove, × 1; S, seed from side, × 5. All figures drawn by Joanna Langhorne from Kew LCDacc. no. 2007-1838.

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samples of F. sororum. However it must be admitted that thegroupings indicated by DNA studies on Turkish and easternMediterranean Fritillaria species do not often coincide withgroupings based on floral morphology. More testing of samplesof known wild origin might clarify some of these anomalies.

Another species of Fritillaria in southern Turkey is somewhatsimilar to F. sororum and in the phylogenetic tree published inDay et al. (2014), is indicated to be more closely related. Thisis F. whittallii Baker, which also has a tessellated, green andbrown flower and narrow leaves, but has green rather thanglaucous leaves and large distinct tessellations on the perianthsegments, 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 character-istic species of Cedrus libani forest on limestone in Antalya andIsparta vilayet, (Byfield et al., 2010), well-separated from the twoknown populations of F. sororum.

Because Fritillaria sororum is so rare that its exact locality isnot mentioned in the original description, it is not given here.The valley where it grows is very isolated, the rock a confusedmixture of schist and limestone. When we visited the typelocality in early April 2002, flowering was well advanced. Therewere a reasonable number of non-flowering leaves, but onlyfive 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, as well as Dry-opteris and various grasses. We failed to find the second, higheraltitude locality. Another, as yet unnamed, species with smallflowers was found nearby.

Pollination has not been recorded in the wild, but is probablyby queen wasps.

Cultivation. Fritillaria sororum is cultivated in the collectionat Göteborg and bulbs from the type collection are also at Kew,grown from seed from Göteborg. Seeds germinate readily inabout 3 to 4 months from a winter sowing, generally appear-ing any time from May to July, the bulbs becoming mature in

220 © The Board of Trustees of the Royal Botanic Gardens, Kew 2014.

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three or four years. At Kew the bulbs are grown with a selec-tion of like-minded fritillaries from the Mediterranean region,planted in clay pots and plunged in a free standing, raised sandbed. It is east facing and has a glass roof as a cover. The bulbsnever get too hot as the frame affords plenty of shade and theplunge gets sprayed over in the summer, so that it never driesout completely. As with most species in Mediterranean climates,F. sororum is found growing inside deciduous bushes, which bothprotects the stems against grazing, and keeps the bulbs cool, soit is important that they should not become too hot nor dry outtoo much when dormant.

In September they are re-potted into a loam-based compostwith added grit and a little bit of coir to make it slightly moistureretentive. The bulbs are set on a small layer of river-washed sandto keep them snug. To start the bulbs in autumn, both pots andplunge are drenched once very thoroughly, and then up to thepoint the bulbs appear, only the plunge is watered when it feelsdry. Only when the shoots and leaves are above ground are theywatered freely. F. sororum, is generally helped along in the grow-ing season with a few pea-sticks, to support its tall stems, and itcan coil onto the pea-sticks with its tendrils. It can get very tall, inone year the plants reaching over 1m. They grow fast for abouttwo months until they go dormant again in early May. In theabsence of a number of herbarium specimens, the descriptionbelow 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 brookravine in Pinus brutia forest, on limestone, coll. 30. 04. 1987 in fruit, cult.20. 05. 1995, J. Persson & K. Persson 87–99 (holo: GB).

Description. Bulb to 3 cm across. Stem 25–45(−100) cm, glaucous,smooth. Leaves 6–10, all alternate, glaucous, linear, shallowly canalicu-late, 12–16 cm long, 7–10 mm broad, the uppermost around 12 cm longand tendril-like at the tip, sometimes coiling around a support. Flow-ers 1, rarely 2, broadly bell-shaped, rounded at the angle, glaucous out-side, marked and tessellated, especially on the inside, with dark choco-late brown, often darker towards base and apex, without fascia; scent notrecorded. Perianth segments 3–4 cm, the inner 1.5–2.3 cm wide, broadlyoblanceolate to obovate, with a slightly reflexed, apiculate and papilloseapex, the outer narrowly ovate, 1–1.7 cm wide. Nectary 7–11× 3.5–5 mm,

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narrowly ovate, brownish-black, 4–5 mm above the tepal base. Filaments9–10 mm, slender, smooth or minutely papillose. Style 12–15 mm, thebranches 5–8 mm, slender, smooth, the branches recurved. Capsule cylin-drical, not winged (Fig. 2). Chromosome number: 2n= 24.

Distribution and habitat. Turkey: Içel; growing in grassy places inQuercus scrub and Pinus brutia forest, 500–950 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 FritillariaL. (Liliaceae). Molecular Phylogenetics and Evolution 80: 11–19.

Persson, J. & Persson, K. (1998). Fritillaria sororum (Liliaceae): a newspecies from Turkey. The New Plantsman 5(2): 114–116.

Teksen, M. & Aytaç, Z. (2011). The revision of the genus Fritillaria L.(Liliaceae) in the Mediterranean region (Turkey). Turkish Journal ofBotany 35: 447–478.

222 © The Board of Trustees of the Royal Botanic Gardens, Kew 2014.