Comparative morphology of the leaf epidermis in Fritillaria (Liliaceae) from China QIANG WANG†, SONG-DONG ZHOU†, XIAO-YAN DENG, QI ZHENG and XING-JIN HE* College of Life Sciences, Sichuan University, Chengdu 610064, China Received 3 November 2007; accepted for publication 17 March 2008 The leaf epidermis of 16 species and one putative species of Fritillaria was examined using light microscopy (LM) and scanning electron microscopy (SEM). The results showed that the stomatal and other epidermal features were constant within species. Epidermal cells of Fritillaria under LM were usually polygonal and anticlinal cell walls were straight or curved. In a few species they were irregular, with sinuous anticlinal cell walls. The cuticular membrane of Fritillaria was usually striated, and the wax ornamentations were flaked, granular or concomitant. Based on leaf epidermal characteristics, the subdivision of Fritillaria is discussed, and the statistical t-test method was used to ascertain the significance level of the differences in the stomata of each species. All orientations of the stomatal poles in Fritillaria were the same, and this phenomenon was named ‘stomatal orientation’. The stomatal characteristics support the origin of section Fritillaria in China from two floristic elements. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160, 93–109. ADDITIONAL KEYWORDS: stomata – taxonomy – t-test – wax ornamentation. INTRODUCTION Fritillaria L. is a genus of tribe Lilieae of Liliaceae. It consists of about 130 species distributed in the tem- perate regions of the Northern Hemisphere, mainly in Central Asia and the Mediterranean region (Chen & Mordak, 2000). There are 24 species and two varieties in China, 15 species of which are endemic. All species of Fritillaria are geophytic perennials and bulbifer- ous. The bulbs are composed of a few fleshy, farina- ceous scales, often covered with a translucent tunic. In China, the bulbs of some species have been used medicinally under the name ‘bei-mu’ since ancient times. In the Pharmacopoeia of the People’s Republic of China (Editorial Board of the State Pharmacopoeia of the People’s Republic of China, 2000), eight species are prescribed in Chinese medicine, including F. cir- rhosa D.Don, F. unibracteata Hsiao & K.C.Hsia, F. delavayi Franch., F. przewalskii Maxim. ex Batalin., F. thunbergii Miq., F. usuriensis Maxim., F. walujewi Regel and F. pallidiflora Schrenk. Numerous studies of the morphology of Fritillaria have been performed. Baker (1874) studied the bulb, style, form of the nectary and capsule valves. Bakshi- Khaniki & Persson (1997) studied the nectary morphology in 31 species from south-western Asia. However, few studies have been performed on the leaf epidermal morphology in non-Chinese Fritillaria. In a review of the leaf epidermal morphology in Chinese Fritillaria, Li et al. (1993, 1994) investigated 22 Chinese species and four varieties. Zhang, Cheng & An (1998) studied leaf epidermal morphology in eight Chinese species and two varieties of Fritillaria from Gansu Province, China. Pu et al. (2001) studied the leaf epidermis in three species, one variety and one putative species. However, the descriptions of the leaf epidermis in all the reported species are not detailed, and the widely adopted terminology is not used well. Moreover, no taxonomic conclusion has been reached on the basis of the leaf epidermis. Thus, a more detailed description of the leaf epidermis of Chinese Fritillaria using the widely adopted terminology (Wilkinson, 1979; Baranova, 1987, 1992) is necessary. In this study, the leaf epidermis of 16 species and one putative species of Fritillaria from China was *Corresponding author. E-mail: hexingjin7@gmail.com †Co-first authors. Botanical Journal of the Linnean Society, 2009, 160, 93–109. With 91 figures © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160, 93–109 93 Downloaded from https://academic.oup.com/botlinnean/article-abstract/160/1/93/2418345 by guest on 08 June 2020