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A New Genus in Polygonaceae Based on Conventional Maximum Parsimony and Three-Taxon Statement Analyses of a Comprehensive Morphological Dataset

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A New Genus in Polygonaceae Based on Conventional Maximum Parsimony and Three-Taxon Statement Analyses of a Comprehensive Morphological Dataset Phytotaxa 314 (2): 151–194 ISSN 1179-3155 (print edition) http://www.mapress.com/j/pt/ PHYTOTAXA Copyright © 2017 Magnolia Press Article ISSN 1179-3163 (online edition) https://doi.org/10.11646/phytotaxa.314.2.1 Persepolium (Polygoneae): A new genus in Polygonaceae based on conventional Maximum Parsimony and Three-taxon statement analyses of a comprehensive morphological dataset OLGA V. YURTSEVA1*, ELENA E. SEVEROVA1 & EVGENY V. MAVRODIEV2 1Department of Higher Plants, Faculty of Biology, M.V. Lomonosov Moscow State University, 1–12, Leninskie Gory, 119234, Moscow, Russia; e-mail: [email protected] 2Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, U.S.A. *Author for correspondence Abstract Recent molecular studies revealed the polyphyletic nature of the broadly defined genus Polygonum. This paper includes a standard maximum parsimony (MP) and three-taxon statement analyses (3TA) as well as a taxonomic revision of Polygonum sect. Avicularia subsect. Spinescentia (Polygonaceae) as compared with two closely related taxa: genus Atraphaxis s.s., and genus Bactria. In total, 27 characters, including life history, habit, morphology of the shoots, leaf blades, ocreas, perianth, achene, ultrasculpture of perianth and achene surfaces, as well as pollen morphology, were analyzed, illustrated, and dis- cussed in detail. Both the standard MP and 3TA confirm that monophyletic Polygonum subsect. Spinescentia is sister to the narrowly defined Atraphaxis. The genus Persepolium (Polygonum subsect. Spinescentia), with the circumscription of five species, is established as new to science as a result of this study. Possible transformations of perianth and thyrse morphology are discussed within the framework of the Principle of variable proportions by Troll in connection with a shift of pollination mode in the group of taxa studied. Key words: Standard cladistic analysis, three-taxon statement analysis, taxonomy, achene surface sculpture, ocrea morphol- ogy, perianth morphology, pollen, sporoderm ornamentation, tepal micromorphology Introduction Recent molecular studies (Tavakkoli et al. 2015, Yurtseva et al. 2016a) have revealed the polyphyletic nature of the broadly defined genus Polygonum Linnaeus (1753: 359). This paper includes a standard Maximum Parsimony (MP) and Three-taxon statement analyses (3TA), as well as a taxonomic revision of Polygonum sect. Avicularia subsection Spinescentia (Polygonaceae) as compared with two closely related taxa: Atraphaxis Linnaeus (1753: 333), and Bactria Yurtseva & Mavrodiev (Yurtseva et al. 2016a: 42). Polygonum section Avicularia subsection Spinescentia Boissier (1879: 1027) comprises several endemics of West and South Iran: P. aridum Boiss. & Hausskn. in Boissier (1879: 1042), P. dumosum Boissier (1846: 83), P. khajeh-jamalii Khosravi & Poormahdi (2008: 477), P. salicornioides Jaubert & Spach (1844–46: tab. 123), and P. spinosum Gross (1913: 340). They are all erect dwarf caespitose undershrubs, or shrubs with abandantly branched shortly puberulent annual shoots, linear-elliptical or oblong-lanceolate leaf blades, short bifid truncate-tubular ocreas, terminal thyrses of several cymes of 1–2 (3) flowers, a campanulate or urceolate perianth with equal-sized segments, which is totally densely minutely puberulent outside. Recent molecular phylogenetic analyses (Tavakkoli et al. 2015) discovered the sister position of Polygonum subsection Spinescentia to Atraphaxis, as well as the sister relationship of the clade (P. subsection Spinescentia plus Atraphaxis) and A. ovczinnikovii (Czukavina 1962: 62) Yurtseva (Yurtseva et al. 2014: 763). As a result, Tavakkoli et al. (2015) transferred Polygonum section Avicularia subsect. Spinescentia and Atraphaxis section Ovczinnikovia Yurtseva ex Tavakkoli (Tavakkoli et al. 2015: 1167) into the genus Atraphaxis, treating this taxon with a broad circumscription, perhaps broader than has ever been done before. According to Tavakkoli et al. (2015), Polygonum subsection Spinescentia is treated as Atraphaxis section Polygonoides S.Tavakkoli, Kaz.Osaloo & Mozaff. in Tavakkoli et al. (2015: 1167). Accepted by Alexander Sennikov: 16 Jun. 2017; published: 25 Jul. 2017 151 Tavakkoli et al. (2015) also mentioned that aside from the five listed species of Polygonum, A. section Polygonoides should also include Polygonum botuliforme Mozaffarian (1988: 62), which is a rare and poorly known species from Central Iran. Both Tavakkoli et al. (2015) and Yurtseva et al. (2016a, b) reported conflicting positions for P. botuliforme in plastid vs. ITS-based topologies. As already mentioned (Yurtseva et al. 2016a, b), this issue requires further reappraisal and, at the moment, P. botuliforme is excluded from consideration here. Our results using Maximum Likelihood (ML) and Bayesian analyses (BI) based on ITS nrDNA regions as well as combined plastid regions (trnL (UAA) intron, trnL-F IGS, and rpl32-trnL (UAG) IGS) of Atraphaxis s.s. and related taxa showed that the name “A. ovczinnikovii” matches two taxonomic entities accepted at the specific rank (Yurtseva et al. 2016a). We also found that these two species form a strongly supported sister clade of (Polygonum subsection Spinescentia plus Atraphaxis s.s.) (combined plastid regions), that, however, appear paraphyletic on the phylogenetic trees based on ITS nrDNA regions. Both species are dwarf shrubs resembling some shrubby species of Polygonum by the campanulate perianth with five equal-sized segments, but have remarkable papillae at the segment margins and distinct sporoderm ornamentation. The genus Bactria (Yurtseva et al. 2016a: 42) has been proposed, which includes these two dwarf shrubs: B. ovczinnikovii (Czukav.) Yurtseva & Mavrodiev (Yurtseva et al. 2016a: 43) from Pamir and B. lazkovii Yurtseva & Mavrodiev (Yurtseva et al. 2016a: 43) from Tian-Shan. Our results using ML and BI based on ITS nrDNA regions and combined plastid regions confirmed Polygonum subsection Spinescentia as a strongly supported sister to Atraphaxis s.s. (Yurtseva et al. 2016a). However, we suggest that the proposed circumscription of Atraphaxis including Polygonum subsection Spinescentia (Tavakkoli et al. 2015) is questionable from a morphological standpoint due to the loss of morphological identity of Atraphaxis so circumscribed. The majority of species in the narrowly defined Atraphaxis (Yurtseva et al. 2016a) are shrubs or undershrubs with a rather specialized perianth characterized by accrescent inner segments and a long filiform tube. Several rare endemics from Central Asia differ in their perianth with equal-sized segments, but correspond to Atraphaxis s.s. in many other features (Yurtseva et al. 2010, Yurtseva et al. 2012a, b, 2016a, b, Schuster et al. 2011b). Atraphaxis toktogulica (Lazkov) T.M.Schust. & Reveal, A. atraphaxiformis (Botsch.) T.M.Schust. & Reveal, A. ariana (Grigorj.) T.M.Schust. & Reveal (Schuster et al. 2011b: 1663), and A. tortuosa Losinsk. (Losina-Losinskaja 1927: 44) are nested individually within the clade Atraphaxis in plastid and ITS-based phylogenetic trees (Yurtseva et al. 2016a, b) and are particularly important for comparison with the members of Polygonum subsection Spinescentia when investigating convergent features in these sister groups. A comprehensive comparative morphological analysis of Polygonum subsection Spinescentia and closely related Atraphaxis s.s., as well as of the recently described genus Bactria, is the main goal of this study. This analysis is critical in order to evaluate the results of the previous molecular phylogenetic analyses of the complex, as well as to clarify the rank of Polygonum subsection Spinescentia. Materials & Methods Data sources The morphological study involved eight type and verified specimens of Polygonum aridum, P. dumosum, P. salicornioides, and P. spinosum from P. subsection Spinescentia, and more than 400 specimens of 15 species of Atraphaxis representing all its sections and morphological groups, as well as five specimens of Bactria ovczinnikovii, and the holotype of B. lazkovii. The specimens are stored in the Herbaria of V.L. Komarov Botanical Institute RAS, St. Petersburg, Russia (LE); Botanical Department of St. Petersburg University, St. Petersburg, Russia (LECB); M.V. Lomonosov Moscow State University, Moscow, Russia (MW). High-resolution images of the type specimens of P. aridum, P. dumosum, P. salicornioides, and P. spinosum (B—http://ww2.bgbm.org/herbarium/, G-BOIS, LINN—http:// linnean-online.org/linnaean_herbarium.html, M, and P—https://science.mnhn.fr/taxon/genus), were also carefully examined. All of the taxa included in the morphological analysis are listed in Appendix 1. Additionally to the certain number of the herbarium specimens and images (summarized above), we studied and compared morphological characters of Atraphaxis and related taxa using the comprehensive observations from the literature. For Atraphaxis we used the data from Ronse De Craene & Akeroyd (1988), Ronse De Craene & Smets (1991), Hong (1995), Hong et al. (1998), Hong et al. (2005), Yurtseva et al. (2012a, 2014, 2016a, b). For Polygonum subsection Spinescentia, we used the data from Jaubert & Spach (1844–1846), Boissier (1846, 1879), Meisner (1857), 152 • Phytotaxa 314 (2) © 2017 Magnolia Press YURTSEVA ET AL. Gross (1913), Mozaffarian (2012), Khosravi & Poormahdi (2008), Tavakkoli et al. (2015), and Yurtseva et al. (2016a). Due to the complete lack of the available material of Polygonum khajeh-jamalii, the data from Khosravi & Poormahdi (2008) was obtained for
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