Cat’s Eyes and Popcorn Flowers: Phylogenetic Systematics of the Genus Cryptantha s. l. (Boraginaceae)
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چکیده
Cryptantha (Boraginaceae) is a group of approximately 200 annual and perennial species, representing two-thirds of the diversity within subtribe Cryptanthinae. The genus exhibits an amphitropic distribution, occurring in temperate and desert regions of western North and South America. Fifty samples of 45 species of Cryptantha s. l., exemplars of the related genera Amsinckia, Pectocarya, and Plagiobothrys, and four outgroup taxa were sequenced for two gene regions, the nuclear ribosomal gene, ITS, and the trnLUAA intron region of the chloroplast genome. These data were used to assess phylogenetic relationships using parsimony, maximum likelihood, and Bayesian inference methods. Cryptantha s. l. was found to be polyphyletic, with its members placed among several well-supported clades. Based on these analyses, we propose resurrection of the genera Eremocarya, Greeneocharis, Johnstonella, and Oreocarya, and recognition of a newly delimited Cryptantha s. s. The related genera Amsinckia and Pectocarya were resolved as monophyletic and most closely related to various clades within Cryptantha s. l. Plagiobothryswas resolved as polyphyletic in three clades, these clades corresponding to previously named sections or groups of sections. The Cryptanthinae is supported as monophyletic. Character trait analyses support the multiple, derived evolution of perenniality, reduction in nutlet number, nutlet heteromorphism, smooth nutlet sculpturing, heterostyly, and cleistogamy. Although sampling is incomplete, this study generally supports the hypothesis of repeated, unidirectional dispersal events, from North to South America. Genera resurrected include: Eremocarya, Greeneocharis, Johnstonella, and Oreocarya. New combinations include: Greeneocharis circumscissa var. rosulata, Greeneocharis similis, Oreocarya atwoodii, Oreocarya barnebyi, Oreocarya compacta, Oreocarya crassipes, Oreocarya creutzfeldtii, Oreocarya fulvocanescens var. nitida, Oreocarya grahamii, Oreocarya hypsophila, Oreocarya johnstonii, Oreocarya ochroleuca, Oreocarya roosiorum, Oreocarya schoolcraftii, Oreocarya semiglabra, Oreocarya shackletteana, Oreocarya sobolifera, Oreocarya subcapitata, Oreocarya suffruticosa var. arenicola, Oreocarya suffruticosa var. laxa, Oreocarya suffruticosa var. pustulosa, Oreocarya suffruticosa var. setosa, Oreocarya welshii, Johnstonella angelica, Johnstonella angustifolia, Johnstonella costata, Johnstonella diplotricha, Johnstonella echinosepala, Johnstonella fastigiata, Johnstonella grayi var. cryptochaeta, Johnstonella grayi var. grayi, Johnstonella grayi var. nesiotica, Johnstonella holoptera, Johnstonella micromeres, Johnstonella parviflora, and Johnstonella pusilla. Keywords—Amsinckia, biogeography, Pectocarya, Plagiobothrys, taxonomy, trait evolution. Cryptantha Lehmann ex G. Don, commonly known as “cat’s eye” or “popcorn flower,” is a genus of the family Boraginaceae. The circumscription of this family has changed repeatedly over the years (Engler and Prantl 1897; Ferguson 1999; Gottschling et al. 2001; Heywood et al. 2007; APG III 2009), with various authors recognizing broad or narrow family concepts with different arrangements of subgroups. Here we accept the APG III (2009) system of classification recognizing a broad Boraginaceae, which may be divided into subfamilies Boraginoideae, Cordioideae, Ehretioideae, Heliotropoideae, Hydrophylloideae, and Lennoideae (Stevens 2001 onwards). Cryptantha firmly belongs within subfamily Boraginoideae, a group characterized by an inflorescence that is a circinate, scorpioid cyme (Buys and Hilger 2003), a deeply 4–lobed ovary with a gynobasic style, and a fruit that is a schizocarp of nutlets (Gottschling et al. 2001). Since the 1920s, Cryptantha has been circumscribed broadly, inclusive of several segregate genera; we refer to this genus concept as Cryptantha s. l. The genus is amphitropically distributed, with taxa in mostly temperate or desert regions of both western North America and western South America, but absent in intervening tropical regions. As currently delimited in most treatments, Cryptantha s. l. has about 200 species (see Simpson 2011). The greatest diversity, approximately 130 species, occurs in western North America, with distributions from Alaska to southern Mexico, and from the Pacific coast and east to Texas (Johnston 1925; Payson 1927; Higgins 1971, 1979). Approximately 70 species occur in western South America, in Peru, Chile, Bolivia, and east to Argentina (Reiche 1910; Johnston 1927; Schwarzer 2007; Teillier 2009; Zuloaga et al. 2008). Three species, C. albida, C. circumscissa, and C. maritima, are distributed in both North and South America. Members of the genus are strigose to hispid, annual, biennial, or perennial herbs, with simple to highly branched, generally ascending to erect stems and simple, basal to cauline, generally linear, lanceolate, or oblanceolate leaves (Johnston 1925; Payson 1927; Munz and Keck 1959, 1968; Higgins 1971; Kelley and Wilken 1993, Mabberley 2008; Simpson and Hasenstab 2009; Kelley et al. 2012). The shape, degree of fusion, vestiture, and orientation of the accrescent, fruiting calyx are often used as diagnostic features. Flowers are chasmogamous in most taxa, but may also be cleistogamous in members of subgenera Cryptantha, Krynitzkia, and Geocarya, plants of the last bearing fruits modified into lenticular structures (Grau 1983). The corolla limb ranges in size from less than one mm to two cm wide. Corollas are almost universally white (yellow in a few species) and are rotate to salverform, with five, often yellow fornices (invaginated, folded regions) surrounding the corolla throat. The number of nutlets that develop to maturity can vary from one to four; these are attached to narrowly-pyramidal receptacular tissue (the gynobase) from which a style and stigma arise. Nutlets are generally ovate to lanceolate in shape and can detach from or remain attached to the gynobase within the calyx at dispersal. Several species on both continents have fruits with nutlets that are heteromorphic in size and/or sculpturing; in these, there are often three smaller, easily detached nutlets, and one larger nutlet that is strongly adnate to the gynobase. The pericarp wall is variable in sculpturing and color. The attachment scar is generally a shallow,
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