Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology Official Journal of the Societa Botanica Italiana ISSN: 1126-3504 (Print) 1724-5575 (Online) Journal homepage: http://www.tandfonline.com/loi/tplb20 A new infrafamilial taxonomic setting for Liliaceae, with a key to genera and tribes L. Peruzzi To cite this article: L. Peruzzi (2016): A new infrafamilial taxonomic setting for Liliaceae, with a key to genera and tribes, Plant Biosystems - An International Journal Dealing with all Aspects of Plant Biology, DOI: 10.1080/11263504.2015.1115435 To link to this article: http://dx.doi.org/10.1080/11263504.2015.1115435 Published online: 11 Jan 2016. Submit your article to this journal Article views: 29 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tplb20 Download by: [La Trobe University] Date: 22 February 2016, At: 03:53 Plant Biosystems, 2016 http:/dx.doi.org/10.1080/11263504.2015.1115435 A new infrafamilial taxonomic setting for Liliaceae, with a key to genera and tribes L. PERUZZI* Dipartimento di Biologia, Università di Pisa, Pisa, Italy Abstract Based on currently available morphological, karyological, and phylogenetic information, a taxonomic setting of Liliaceae is provided. The family is here organized in 6 tribes, for a total of 15 genera and about 900 species. An identification key for tribes and genera is also provided. Keywords: Calochorteae, Liliales, Lilieae, Medeoleae, monocots, Streptopeae, taxonomy, Tricyrtideae, Tulipeae Luke 12: 27 Consider the lilies, how they grow: they 2003, 2009). The genera belonging to families with- neither toil nor spin; but I tell you, not even Solomon in in Asparagales are marked by the presence of septal all his glory clothed himself like one of these nectaries at the base of ovary, by seeds dark brown to blackish, rich in phytomelan, with cells of the outer coat collapsed (Dahlgren et al. 1985). By contrast, Introduction according to the same authors, the genera belonging to families within Liliales show nectaries on tepals, The family Liliaceae was subjected to several – even seeds light brown, lacking phytomelan, with cells of dramatic – circumscription changes over the last 2–3 the outer coat intact. Notably exceptions to this rule decades (see, Peruzzi et al. 2009 for a short review). are Iridaceae and Orchidaceae, currently placed in However, a general agreement concerning the in- Asparagales on phylogenetic grouds (Chase et al. cluded taxonomic groups and its sister relationship 1995; APG 1998, 2003, 2009), but placed with- with Smilacaceae, within Liliales, has been reached in Liliales by Dahlgren et al. (1985), based on the in the last 20 years (Chase et al. 1995; APG 1998, above-cited character states. However, Iridaceae and 2003, 2009; Patterson & Givnish 2002; Fay et al. Orchidaceae have three to two–one stamens, respec- 2006; Leitch et al. 2007; Peruzzi et al. 2009; Kim tively, and both have inferior ovary (Goldblatt et al. et al. 2013; Petersen et al. 2013). In particular, the 1998; Vogel 1998), while Liliales have mostly superi- Downloaded by [La Trobe University] at 03:53 22 February 2016 most important papers providing a strong phylogenetic or ovary or, if inferior, have six stamens (Vogel 1998). backbone for the family are those of Patterson and Liliaceae are easy to distinguish from the remain- Givnish (2002), Fay et al. (2006), Kim et al. (2013), ing nine families presently recognized in Liliales and Petersen et al. (2013); the first authors focused (APG 2009), thanks to several evident morphological on Liliaceae, the others with Liliales but with a large features. They share the absence of leaf base sheath- sampling of genera and families. ing with Philesiaceae, Rhipogonaceae, and Smi- Traditional angiosperm classifications used to in- lacaceae, but Liliaceae are never climbers or shrubs clude most of the monocots bearing showy flowers as the three above-cited families, or even as Alstro- with six tepals, six stamens, and a trilocular superior emeriaceae (tribe Luzuriageae) and Petermanniace- ovary in Liliaceae (e.g. Cronquist 1981). Since the ae (Conran 1998; Conran & Clifford 1998a, 1998b, seminal studies of Dahlgren et al. (1985), the genera 1998c). In addition, Liliaceae are never parasitic formerly classified under Liliaceae have been distrib- plants as Corsiaceae (Neinhuis & Ibisch 1998); they uted among many smaller families organized in two have not inferior ovary as Alstroemeriaceae (tribe Al- main orders: Asparagales and Liliales (APG 1998, stroemerieae), Corsiaceae, and Campynemataceae Correspondence: Lorenzo Peruzzi, Dipartimento di Biologia, Unita di Botanica, Universita di Pisa, via Derna 1, 56126 Pisa, Italy. Email: lorenzo.peruzzi@ unipi.it © 2016 Societa Botanica Italiana 2 L. Peruzzi (Bayer 1998; Kubitzki 1998; Neinhuis & Ibisch 1. Tribe Streptopeae Baker, J. Linn. Soc., Bot. 1998); they do not show resupinate leaves as most 14: 522. 1875. (Figure 1A–C) Alstroemeriaceae (Bayer 1998; Conran & Clifford 1998a); they lack cells with raphide crystals, occur- = Scoliopeae Baker, J. Linn. Soc., Bot. 14: 509. ring in Melanthiaceae (Tamura 1998c, 1998d) and, 1875. finally, show anthers extrorse, and not mostly int- = Scoliopinae S.Watson, Proc. Amer. Acad. Arts 14: rorse as in Colchicaceae (Nordenstam 1998). 222. 1879. Liliaceae as currently circumscribed include sev- = Streptopoideae Mabberley ex Reveal, Phytoneu- eral plants of relevant economic importance for their ron 2012–37: 219. 2012. [23 Apr 2012] remarkable beauty, such as Calochortus (American lily – C. nuttallii Torr. is the official flower of Utah, Included genera: Prosartes D.Don (6 species), USA and its bulbs are also edible), Tricyrtis, Fritillar- Scoliopus Torr. (2 species), Streptopus Michx. (7 ia, Tulipa, Lilium, one of the oldest ornamental plants species) of the Western world being indeed L. candidum L. Plants herbaceous, never climbers, rhizomatous; Some species of Calochortus, Cardiocrinum, Fritil- fruit a berry or a capsule (in Scoliopus); monosporic laria, Gagea (incl. Lloydia), Lilium, Notholirion, and embryo-sac formation of Polygonum-type, nucel- Tulipa are edible or used as drugs, especially in Asia la with long basal part, placentation axile (Tamura (Tamura 1998a, 1998b). 1998b); chromosomes generally short 1–5 μm (but However, a critical, integrative, and supported 5–13 μm in Scoliopus); genome size 4–15(30) pg, infrafamiliar taxonomic setting is still missing. This x = 7–9: x = 8(9) in Prosartes, x = 8–9 in Scoliopus, is exactly the aim of this paper, which takes into ac- x = (7)8 in Streptopus (Peruzzi et al. 2009). count the morphological, karyological, and phyloge- The tribe Streptopeae represents one of the ear- netic information available in the literature. liest branching lineages within Liliaceae, for which some authors suggest a sister relationship with Calo- chortus + Tricyrtis (Patterson & Givnish 2002), others Taxonomic setting with Tricyrtis (Kim et al. 2013) or with Calochortus Family Liliaceae Juss., Gen. Pl.: 48. 1789. nom. cons. (Fay et al. 2006) only. Despite this, the established phylogenetic relationships among the three genera = Tulipaceae Batsch, Dispos. Gen. Pl. Jenens.: 48. are the following (Patterson & Givnish 2002; Fay 1786. et al. 2006; Kim et al. 2013): ((Prosartes, Scoliopus), = Erythroniaceae Martynov, Tekhno-Bot. Slovar: Streptopus). 238. 1820. 2. Tribe Calochorteae Melchior, Pflanzenfam. = Calochortaceae Dumort., Anal. Fam. Pl.: 53. ed 12, 2: 520. 1964. (Figure 1D) 1829. = Fritillariaceae Salisb., Gen. Pl.: 56. 1866. = Calochortoideae Thorne & Reveal ex Reveal, Phy- = Medeolaceae. S.Watson in Takht., Sist. Magnoli- toneuron 2012–37: 216. 2012. [23 Apr 2012] of.: 291, 31. 1987. Included genera: Calochortus Pursh. (ca. 65 = Scoliopaceae Takht., Bot. Zhurn. Moscow & Len- species) Downloaded by [La Trobe University] at 03:53 22 February 2016 ingrad. 81(2): 86. 1996. Plants herbaceous, never climbers, bulbous; fruit = Tricyrtidaceae Takht., Divers. Classif. Fl. Pl.: 482. a septicide capsule; monosporic embryo-sac forma- 1997. nom. cons. tion of Polygonum-type, nucella with long basal part, placentation axile (Tamura 1998b); chromosomes Plants herbaceous, never climbers or shrubs, nev- generally short, 1–6(13) μm; genome size 4–15(30) er parasitic, bulbous or rhizomatous; leaf sheaths ab- pg, x = 6–10 (Peruzzi et al. 2009). sent, leaves never resupinate; flowers with two whorls The tribe Calochorteae represents a lineage cer- of 3 + 3 tepals, two whorls of 3 + 3 stamens (rare- tainly distinct from others, but of uncertain phy- ly reduced to one), with extrorse anthers, syncarpic logenetic placement: sister to Tricyrtis for Patterson superior ovary made up of three carpels; presence and Givnish (2002) and Petersen et al. (2013), sister of nectaries on tepals; hermaphrodite (rarely male to Streptopeae for Fay et al. (2006), or even sister sterile, but relatively often female sterile) flowers; to Medeoleae + Lilieae + Tulipeae for Kim et al. fruit a capsule (loculicide, septicide) or a berry; (2013). Concerning the phylogenetic relationships seeds light brown, lacking phytomelan, with cells of among species within Calochortus, a detailed cpDNA the outer coat intact; cells lacking raphide crystals; phylogeny is available (Patterson & Givnish 2004). monosporic or tetrasporic embryo-sac formation (Tamura 1998a, 1998b; Peruzzi et al. 2012; Zhang 3. Tribe Tricyrtideae [Trycirteae] K.Krause in En- et