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A Morphometric Study of Species Delimitation in Sternbergia Lutea (Alliaceae, Amaryllidoideae) and Its Allies S

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A Morphometric Study of Species Delimitation in Sternbergia Lutea (Alliaceae, Amaryllidoideae) and Its Allies S Botanical Journal of the Linnean Society, 2008, 158, 460–469. With 5 figures A morphometric study of species delimitation in Sternbergia lutea (Alliaceae, Amaryllidoideae) and its allies S. sicula and S. greuteriana EWAN GAGE1,2 and PAUL WILKIN2* FLS 1Plant Science Laboratories, The University of Reading, Whiteknights, Reading, Berkshire RG6 6AS, UK 2Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, UK Received 22 October 2007; accepted for publication 13 May 2008 The morphological chararacters used to differentiate the species Sternbergia lutea (L.) Ker Gawl. ex Spreng., Sternbergia sicula Tineo ex Guss. and Sternbergia greuteriana Kamari & R.Artelari were found not to possess discrete or consistently different states during an attempt to produce an electronic multi-access key to the genus. Thus, variation in floral and leaf morphology in the three species was further explored to re-evaluate taxon limits using herbarium specimens and statistical methods, including principal components analysis (PCA) and elliptic Fourier analysis (EFA). This confirmed that variation was continuous between the three species. Sternbergia sicula and S. greuteriana are sunk into S. lutea and a revised description provided. It is suggested that cultivar status is the most appropriate rank for the cultivated forms of the S. lutea complex. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158, 460–469. ADDITIONAL KEYWORDS: Amaryllidaceae – CITES – conservation status – geographical distribution – Mediterranean – systematics. INTRODUCTION although S. lutea (L.) Ker Gawl. ex Spreng. is a notable exception. It was first described as Amaryllis Sternbergia L. (Alliaceae, Amaryllidoideae) is a genus lutea by Linnaeus (1753), before being reassigned to of seven (Mathew, 1983) to nine (Govaerts et al., 2007) Sternbergia by Sprengel (1825). This was followed by species of geophytes that are mainly distributed the description of S. sicula Tineo ex Guss. in 1845. around the Mediterranean basin. Species diversity is These two species are similar in their macromorphol- highest in the eastern Mediterranean. Some species ogy, and Webb (1978) suggested that subspecific rank are valued in horticulture and have become natural- would be more appropriate because of the high level ized elsewhere. The genus can readily be differenti- of morphological overlap between them. This led to ated from other Eurasian Amaryllidoideae through their treatment as S. lutea subsp. lutea and S. lutea possession of a solitary, goblet-shaped flower with an subsp. sicula (Tineo) D.A.Webb in Flora Europaea extended perianth tube (Mathew, 1983). Most species (Webb, 1980). However, this treatment has not been produce yellow flowers in the autumn, although in S. universally accepted; S. sicula and S. lutea were vernalis (Mill.) Gorer & J.H.Harvey and S. candida retained as species by Mathew (1984) in The Flora of B.Mathew & T.Baytop anthesis occurs in spring, with Turkey and by Davis & Mathew in the CITES bulb the latter possessing a creamy–white perianth. checklist (Davis et al., 1999). In their paper, in which The majority of species within the genus are a third closely related species was described (S. greu- defined by discrete morphological characters, teriana Kamari & R.Artelari), Kamari & Artelari (1990) assigned specific status to S. sicula and sepa- *Corresponding author. E-mail: [email protected] rated it from S. lutea on the basis of vegetative and 460 © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158, 460–469 STERNBERGIA LUTEA COMPLEX MORPHOMETRICS 461 floral morphological characters. Sternbergia greuteri- and S. greuteriana, giving ranges of 10–17 mm and ana is an endemic of Crete and the Eastern Aegean 15–32 mm, respectively. However, Kamari & Artelari Islands and was said by Kamari & Artelari to show a (1990) reported that although some differences strong similarity to S. lutea. Sternbergia sicula is occur in filament length between the three species, distributed throughout the Aegean Islands, Crete, intraspecific variation renders it unusable as a diag- Cyclades, Greece, Italy, Sicily and Turkey. Sternbergia nostic character. Pasche & Kerndorff (2002) separated lutea has been recorded through the Mediterranean S. lutea from S. sicula on the basis of leaf width and from Spain to Iran and the Caucasus, although this colour and perianth segment dimensions. distribution has probably been artificially extended When these classifications were evaluated and by its horticultural use (Mathew, 1983). A fourth checked against specimens from K, BM and RNG taxon has also been described, S. minoica Ravenna. (see http://sweetgum.nybg.org/ih/ for herbarium acro- However, this is thought to be a form of S. sicula in nyms) with the aim of writing an electronic multi- which the scape fails to develop and thus the ovary access key, it was discovered that none of the remains subterranean, an occurrence which has been characters used by previous authors enabled a key to observed in several individuals growing in popula- these three species to be produced. This suggested tions near the type locality in Crete. that re-examination of species limits was necessary. Different systematists have employed a number of Other than the studies cited above, recent taxonomic characters to separate the three species. Mathew treatments of Sternbergia in, for example, Turkey (1983) and Kamari & Artelari (1990) used leaf width (Mathew & Baytop, 1984), Spain (Morales & Castillo, as a key character, with the latter authors also 2004), the former USSR (Artjushenko, 1970) and the placing importance upon the cross-sectional shape Aegean Islands (Kamari & Artelari, 1990) have each and colour of fresh leaf material. Polunin & Huxley considered a country in isolation. Even the paper of (1965) suggested that S. sicula and S. lutea could Artelari & Kamari (1991) did not sample taxa outside separated on the basis of the presence or absence of Greece. The work of Pasche & Kerndorff (2002) was glandular cilia on the leaf margins, although Kamari orientated towards plant material in cultivation. & Artelari (1990) reported that no significant differ- Thus, it appeared that the morphological characters ence could be found in this character between either of S. sicula, S. lutea and S. greuteriana had not been species or S. greuteriana. In his account of Sternber- evaluated across the full extent of their geographical gia for Flora Europaea, Webb (1980) separated distributions since Webb (1978) and there has been no S. lutea subsp. lutea from subsp. sicula through attempt to employ modern computational methods to the former having 4 to 15 mm wide leaves and 7 explore variation in a group of species with complex to 15 mm wide perianth segments, with the leaf variation patterns. margins usually being entire or sometimes obscurely A number of conflicting chromosome counts have crenulated and the latter having 3 to 5 mm wide been published for S. sicula and S. lutea: Moore leaves and 4 to 8 mm wide perianth segments and (1982) cited counts of 2n = 18 and 2n = 22 for the two leaf margins with small but discernible crenulae. species, respectively, whereas Löve & Löve (1974) Webb also stated that there was a difference in the listed counts of 12, 16, 18, 20, 22, 24 and 33 for diploid chromosome number of the two subspecies S. lutea. Kamari & Artelari (1990) and Artelari & (see below). Kamari (1991) suggested a basic chromosome number Kamari & Artelari (1990) and Artelari & Kamari of x = 11, with 2n = 22 in all three species, although (1991) placed strong emphasis on the shape and they commented that S. lutea can also be found to dimensions of the perianth segments as diagnostic exhibit a count of 2n = 3x = 33. As a result of such characters. They suggested that the perianth seg- variability in chromosome number, the extent to ments of S. sicula were acute and oblanceolate and which karyological data can be applied as a system- contrasted this with the obtuse and obovate forms atic research tool is limited until carefully gathered, found in both S. lutea and S. greuteriana. Sternbergia population-level data are available. Molecular sys- greuteriana and S. lutea were distinguished from tematic methods have been applied to Sternbergia in each other by leaf and perianth segment dimensions, two studies to date. Açik et al. (1997) reported that, with S. lutea having significantly larger dimensions whereas a number of other members of Sternbergia in each case (for example, they gave leaf width ranges were separable on the basis of random amplified of 2–5(-6) mm in S. greuteriana and 7–12 mm in S. polymorphic DNA–polymerase chain reaction (RAPD- lutea). They stated that S. greuteriana more closely PCR) results, they were unable to separate S. sicula resembles S. lutea than S. sicula in both morphology and S. lutea from each other. Meerow et al. (2006) and karyology (Kamari & Artelari, 1990). As part of suggested that S. lutea and S. sicula could not be their recent survey of the genus, Pasche & Kerndorff differentiated, but that S. colchiciflora Waldst. & Kit. (2002) employed filament length to separate S. sicula was a distinct taxon with significant sequence diver- © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158, 460–469 462 E. GAGE and P. WILKIN gence from S. lutea, S. sicula and S. greuteriana.The MATERIAL AND METHODS sequences used in that study are available from GenBank. The material held at BM, K and RNG of S. sicula, S. The shape and dimensions of perianth segments lutea and S. greuteriana was examined and specimens are among the main characters that have been used in good condition exhibiting all necessary organs were in delimiting species in this complex, e.g. by Kamari scored (see Appendix). An isotype of S. sicula is held at & Artelari (1990). Although these characters are well K(Tineo s.n.) and a digital image of the holotype of S. preserved in herbarium material, unlike the presence lutea (LINN 416/1) was examined.
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