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Floral Ontogeny and Vasculature in Xyridaceae, with Particular Reference to Staminodes and Stylar Appendages Author(S): M

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Floral Ontogeny and Vasculature in Xyridaceae, with Particular Reference to Staminodes and Stylar Appendages Author(S): M Floral ontogeny and vasculature in Xyridaceae, with particular reference to staminodes and stylar appendages Author(s): M. Graça Sajo, Aline Oriani, Vera L. Scatena and Paula J. Rudall Source: Plant Systematics and Evolution, Vol. 303, No. 9 (November 2017), pp. 1293-1310 Published by: Springer Stable URL: https://www.jstor.org/stable/44853796 Accessed: 21-06-2021 11:00 UTC JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at https://about.jstor.org/terms Springer is collaborating with JSTOR to digitize, preserve and extend access to Plant Systematics and Evolution This content downloaded from 86.59.13.237 on Mon, 21 Jun 2021 11:00:03 UTC All use subject to https://about.jstor.org/terms Plant Syst Evol (2017) 303:1293-1310 CrossMark DOI 10. 1007/s00606-0 17- 1438-3 VOZ ORIGINAL ARTICLE Floral ontogeny and vasculature in Xyridaceae, with particular reference to staminodes and stylar appendages M. Graça Sajo1 • Aline Oriani1 • Vera L. Scatena1 • Paula J. Rudall2© Received: 29 March 2017 /Accepted: 24 June 2017 /Published online: 7 July 2017 © The Author(s) This2017. article is an open access publication Abstract We provide a detailed comparative evolutionary study historyof of the xyrid clade; this transition floral ontogeny and vasculature in Xyridaceae, occurred includ-either once followed by a reversal to fertile sta- ing Xyris , Abolboda and Orectanthe. We mensevaluate in Eriocauloideae these and staminodes in some Xyri- data in the context of a recent well-resolved daceae, phylogenetic or twice independently within both Xyridaceae and analysis of Poales to compare floral structures Eriocaulaceae. within the xyrid clade (Xyridaceae and Eriocaulaceae). Xyrids are relatively diverse in both flower structure Keywordsand anatomy; Rower anatomy • Poales • Stamens • Stylar many species incorporate diverse and unusual appendages floral • Vasculature structures such as staminodes and stylar appendages. Xyridaceae possess three generally epipetalous stamens in a single whorl; the "missing" stamen whorl Introduction is either entirely absent or transformed into staminodes. Fertile stamens each receive a single vascular bundle The commelinid diverged monocot family Xyridaceae consists of from the median petal bundle. In Xyris , the fivestamen genera bundle that are mostly neotropical, though the largest diverges at the flower base, but it diverges atgenus upper Xyris flower extends to other regions of the southern levels in both Abolboda and Orectanthe. hemisphereIn species (Table of 1). The family is highly recognizable in Abolboda that possess staminodes, staminode morphological vasculature terms, often possessing ensiform leaves and is closely associated with the lateral vasculature imbricate-bracted of each spikes with showy flowers with only petal. Despite the likely sister-group relationship three antepetalous between fertile stamens (Krai 1998), in contrast Eriocaulaceae and Xyridaceae, our character with optimization the typical six-staminate condition that predominates indicates that the stylar appendages that characterize in most other some monocots (e.g. Remizowa et al. 2010). In Xyridaceae (except Xyris and Achlyphila ) are many non-homol- other monocot families with only three stamens, such ogous with those of some Eriocaulaceae. asOn Iridaceae the other and Mayacaceae, the stamens are opposite the hand, it remains equivocal whether the loss outer of tepals; a fertile i.e. antesepalous (e.g. Carvalho et al. 2009; outer androecial whorl occurred more than once Ronse during de Craene the 2010; Oriani and Scatena 2012). Xyri- daceae are currently circumscribed into two subfamilies: Xyridoideae, with a single widespread genus Xyris L. (ca Handling Editor: Louis P. Ronse De Craene. 360 species), and Abolbodoideae, with four genera ende- mic to South America (Table 1). The family Xyridaceae E Paula J. Rudall belongs in the order Poales and is probably most closely [email protected] related to Eriocaulaceae (e.g. Davis et al. 2004; Givnish 1 Instituto de Biociências, Universidade Estadual Paulista et al. 2010; Barrett et al. 2015). Xyridaceae flowers are (UNESP), Rio Claro, SP CEP 13506-900, Brazil hermaphrodite, but Eriocaulaceae flowers are generally 2 Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, unisexual by late abortion of the stamens or carpels, UK resulting in female or male flowers, respectively (Stützel Ô Springer This content downloaded from 86.59.13.237 on Mon, 21 Jun 2021 11:00:03 UTC All use subject to https://about.jstor.org/terms 1294 M. G. Sajo et al. Table 1 Genera of Xyridaceae Genus No. of species Distribution Abolboda Kunth ca. 22 Central and South America Achlyphila Maguire and Wurdack 1 Endemic to Guiana Highlands Aratitiyopea Steyerm. and P.E.Berry 1 Rocky outcrops in NW Brazil, SW Venezuela and SE Colombia Orectanthe Maguire 2 Endemic to Guiana Highlands Xyris L. ca. 360 Mostly in Neotropics: Central and South America, Africa, Australia Data from Dahlgren et al. (1985), Kral (1998) and Campbell (2005) 1998). Both Xyridaceae and Eriocaulaceae of - green collectively sepals and colourful petals (Rudall and Sajo 1999; termed the "xyrids" - are unusual in possessing Oriani onlyand Scatena three 2012). The absence of septal nectaries in fertile stamens, at least in some species (in both Eriocaulaceae, Xyridaceae and Eriocaulaceae has prompted some the number of androecial whorls is reduced authors in only to speculate one of that nectaries have evolved de novo in the two subfamilies, Paepalanthoideae; some xyrids Eriocaulaceae from ancestors that lacked septal nectaries (Linder are also dimerous). Another commelinid family and Rudall with 2005). only Both Abolbodoideae (Xyridaceae) and three stamens, Mayacaceae (with a single genus, Paepalanthoideae Mayaca (Eriocaulaceae) ), have flowers with stylar was formerly included among the xyrids (e.g.appendages Linder that and produce nectar, which is used as a resource Rudall 2005), but a recent molecular analysis by insect(Bouchenak- pollinators (Rosa and Scatena 2007; Oriani et al. Khelladi et al. 2014) has tentatively placed Maya- 2009; Oriani and Scatena 2011, 2012). caceae closer to Rapateaceae (Fig. 1). In this paper, we examine the floral vasculature of six A colourful, showy perianth and nectar production from species of Xyridaceae, representing three genera: Abol- septal nectaries are both likely ancestral conditions for boda, Orectanthe and Xyris (Fig. 2). The other two genera monocots (Remizo wa et al. 2010). Many Poales are char- ( Achlyphila and Aratitiyopea) are both monotypie and are acterized by flowers that differ from "typical" monocot relatively rare; for these two species, we rely on earlier flowers in several traits that are sometimes associated with descriptions. Floral vasculature of Abolboda , Achlyphila wind pollination. These features include petals that are non- and Orectanthe was described by Carlquist (1960), and showy and often reduced in size and/or number, absence offlowers of Aratitiyopea were described by Campbell and nectaries and a reduced number of stamens (Linder and Stevenson (2007). Oriani and Scatena (2012) also provided Rudall 2005). Species of Xyridaceae are biotically polli- information on floral anatomy of Abolboda and Orectan- nated but lack septal nectaries; their flowers are relatively the. Our primary goals are to elucidate familial systematics showy, possessing a double (heterochlamydeous) perianth and the possible origin of the staminodial whorl that is present in Xyris and some species of Abolboda (Table 2). A Cyperaceae staminodial whorl is a relatively unusual feature in -J Mmmmmm Juncaceae monocots with hermaphrodite flowers (e.g. Walker-Larsen Thurniaceae cyperids and Harder 2000). Sajo et al. (1997) and Remizowa et al. L-lllllljrB,,,,,,'~ Rapateaceae (2012) reported an unusual pattern of perianth vasculature Mayacaceae in Xyris species: each sepal receives a single bundle and p- " Poaceae each petal receives three bundles, in contrast with many rLl other heterochlamydeous angiosperms, in which the sepals ' rļ Joinvilleaceae Flagellariaceae typically receive three bundles each, whereas the petals p f- Restionaceae receive a single major vascular bundle at the base (Eames Centrolepidaceae restiids 1931; Puri 1951; Endress 2001, 2005). - b LJ Im_ Restionaceae Vasculature has been used in combination with other Anarthriaceae characters to infer that the petals are broadly homologous __J Eriocaulaceae | xypjçjg with staminodes (i.e. sterile stamens) because the stamens ft--- Xyridaceae I also typically receive a single vascular bundle; in contrast, p- Bromeliaceae | , ,. , p- I _ . , bromeliads ,. , sepals are considered to be derived from leaf-like bracts I- mm _ Typhaceae . I (reviewed by Remizowa et al. 2012). The unusual vascu- Fig. 1 Tree diagram summarizing relationships between families latureof of Xyris, together with the absence of staminodes Poales, as suggested by molecular analysis in Bouchenak-Khelladi and presence of a floral tube in Orectanthe, Aratitiyopea et al. (2014) Ô Springer This content downloaded from 86.59.13.237 on Mon, 21 Jun 2021 11:00:03 UTC All use subject to https://about.jstor.org/terms
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