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One of 132 equally most-parsimonious trees resulting from phylogenetic analysis of cpDNA restriction site data for taxa of Saxifragac s.s. using an outgroup consisting of Ribes aureum, R. sanguineum, and Francoa sonchifolia (not including autapomorphies; length = 409 steps, consistency index = 0.760, retention index = 0.931). This tree is identical to the 50% majority rule consensus tree (including compatible groupings) constructed from the shortest trees. In those instances where multiple populations were analyzed for a species (Table 1) and identical restriction site maps were obtained, only one entry is given (e.g., Darmera pelt ata). Nodes that did not occur in all shortest trees are marked by black triangles. Monophyletic groupings occurring in some shortest trees but not in this one are: 1) Darmera group-Saxifraga group, 2) Rodgersia sambucifolia-R. pinnata-R. aesculifolia, 3) Boykinia major 11450, 11452, 2084-B. occidentalis 11386, and 4) Boykinia major 11398, 2077-B. occidentalis 11636- B. intermedia. Numbers above each branch indicate the number of restriction site mutations; bootstrap and decay values are indicated as in Fig. 1. Well-supported groups are named and indicated by brackets (Lept. -Tan = Leptarrhena-Tanakaea).  

One of 132 equally most-parsimonious trees resulting from phylogenetic analysis of cpDNA restriction site data for taxa of Saxifragac s.s. using an outgroup consisting of Ribes aureum, R. sanguineum, and Francoa sonchifolia (not including autapomorphies; length = 409 steps, consistency index = 0.760, retention index = 0.931). This tree is identical to the 50% majority rule consensus tree (including compatible groupings) constructed from the shortest trees. In those instances where multiple populations were analyzed for a species (Table 1) and identical restriction site maps were obtained, only one entry is given (e.g., Darmera pelt ata). Nodes that did not occur in all shortest trees are marked by black triangles. Monophyletic groupings occurring in some shortest trees but not in this one are: 1) Darmera group-Saxifraga group, 2) Rodgersia sambucifolia-R. pinnata-R. aesculifolia, 3) Boykinia major 11450, 11452, 2084-B. occidentalis 11386, and 4) Boykinia major 11398, 2077-B. occidentalis 11636- B. intermedia. Numbers above each branch indicate the number of restriction site mutations; bootstrap and decay values are indicated as in Fig. 1. Well-supported groups are named and indicated by brackets (Lept. -Tan = Leptarrhena-Tanakaea).  

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... Boykinia has B. aconitifolia Nutt. in eastern North America, B. lycoctonifolia (Maxim.) Engl. in eastern Asia, and at least five species in western North America (Gornall & Bohm, 1985;Soltis et al., 1993): B. intermedia (Piper) G.N. Jones, B. major A. Gray, B. occidentalis Torrey & Gray, B. rotundifolia Parry, and B. richardsonii (Hook.) ...
... Within the North American clade, the ITS tree placed B. rotundifolia from western North America sister to a clade containing B. aconitifolia from eastern North America and the remaining western North American species, whereas the cpDNA tree shows a trichotomy among B. aconitifolia, B. rotundifolia, and the clade of the remaining species. Phylogenetic analyses of the Boykinia group as well as of Saxifragaceae s. s. (Soltis et al., 1993(Soltis et al., , 1995(Soltis et al., , 1996 indicate that Boykinia is sister to Bolandra A. Gray and Suksdorfia A. Gray, both from western North America. The center of origin for Boykinia is most likely western North America. ...
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This review shows a close biogeographic connection between eastern Asia and western North America from the late Cretaceous to the late Neogene in major lineages of vascular plants (flowering plants, gymnosperms, ferns and lycophytes). Of the eastern Asian–North American disjuncts, conifers exhibit a high proportion of disjuncts between eastern Asia and western North America. Several lineages of ferns also show a recent disjunct pattern in the two areas. In flowering plants, the pattern is commonly shown in temperate elements between northeastern Asia and northwestern North America, as well as elements of the relict boreotropical and Neogene mesophytic and coniferous floras. The many cases of intercontinental biogeographic disjunctions between eastern Asia and western North America in plants supported by recent phylogenetic analyses highlight the importance of the Bering land bridge and/or the plant migrations across the Beringian region from the late Cretaceous to the late Neogene, especially during the Miocene. The Beringian region has permitted the filtering and migration of certain plant taxa since the Pliocene after the opening of the Bering Strait, as many conspecific taxa or closely related species occur on both sides of Beringia.
... Willdenowia 46 -2016 Saxifragaceae (J. W. Kadereit) The non-monophyly of Saxifraga L., first shown by Soltis & al. (1993), has been confirmed in several studies (for discussion see Fernández Prieto & al. 2013;Tkach & al. 2015). In particular, a group of 70 -90 species from North America and Eurasia is only very distantly related to the remainder of Saxifraga and has to be treated as the genus Micranthes Haw. ...
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The use of DNA sequence data in plant systematics has brought us closer than ever to formulating well-founded hypotheses about phylogenetic relationships, and phylogenetic research keeps on revealing that plant genera as traditionally circumscribed often are not monophyletic. Here, we assess the monophyly of genera documented in Rothmaler's "Exkursionsflora von Deutschland" (Gefäßpflanzen: Grundband, 19th Ed.; Jäger 2005). Using a survey of the phylogenetic literature, we discuss which classifications would be consistent with the phylogenetic relationships found and could be followed, provided monophyly is accepted as the primary criterion for circumscribing taxa. We indicate whether and which names are available when changes in generic assignment are made (but do not present a comprehensive review of the nomenclatural aspects of such names). Among the 840 genera examined, we identified c. 140 where data quality is sufficiently high to conclude that they are not monophyletic, and an additional c. 20 where monophyly is questionable but where data quality is not yet sufficient to reach convincing conclusions. While it is still fiercely debated how a phylogenetic tree should be translated into a classification, our results could serve as a guide to the likely consequences of systematic research for the taxonomy of the German flora and the floras of neighbouring countries.
... In addition, two species of Chrysosple- 83 nium (C. valdivicum, C. macranthum) occur in Tierra del Fuego 84 (Hara, 1957), and several species of Saxifraga are also found in 85 the Andes (Webb and Gornall, 1989).DeChaine et al., 2013; Johnson and Soltis, 1995; Johnson et al., 108 1994; Kuzoff et al., 1999; Morgan and Soltis, 1993; Prieto et al., 109 2013; Soltis and Soltis, 1997; Soltis et al., 1993; Soltis et al., 110 2001a; Soltis et al., 2001b; Soltis et al., 1996; Vargas, 2000; Xiang 111 et al., 2012; Zhu et al., 2013) Brochmann et al., 1998; 116 Godsoe et al., 2013; Ness et al., 1989; Segraves and Thompson, 117 1999; Soltis and Doyle, 1987; Soltis and 118 Rieseberg, 1986; Soltis and Soltis, 1989; Soltis and Soltis, 1999; 119 Soltis and Soltis, 1988; Soltis and Soltis, 1989; Soltis et al., 1993; 120 Soltis et al., 1992; Soltis et al., 1991; Soltis et al., 1990; Soltis 121 et al., 1989a; Soltis, 1984; Wolf et al., 1990), 122 biogeography/phylogeography (Oliver et al., 2006; Reisch, 2008; 123 Szövényi et al., 2009; Westergaard et al., 2010), and evolution 124 and diversification (Holten et al., 2006; Kuzoff et al., 2001; Kuzoff 125 et al., 1999; Okuyama et al., 2005; Soltis et al., 2001a; Steen 126 et al., 2000; Vargas et al., 1999; Winkler et al., 2012) Increasing amounts of sequence data and wider taxon sampling 147 have yielded better resolution, support, and evolutionary insights 148 throughout the tree of life (Heath et al., 2008; Moore et al., 2007; 149 Parfrey et al., 2010; Soltis et al., 2011), Ribes (Grossulariaceae), and Liquidambar (Altingiaceae) 178 were selected as outgroups based on previous studies (Fishbein 179 et al., 2001; Jian et al., 2008; Soltis et al., 2013; Soltis et al., 2001a). ...
... In addition, two species of Chrysosple- 83 nium (C. valdivicum, C. macranthum) occur in Tierra del Fuego 84 (Hara, 1957), and several species of Saxifraga are also found in 85 the Andes (Webb and Gornall, 1989).DeChaine et al., 2013; Johnson and Soltis, 1995; Johnson et al., 108 1994; Kuzoff et al., 1999; Morgan and Soltis, 1993; Prieto et al., 109 2013; Soltis and Soltis, 1997; Soltis et al., 1993; Soltis et al., 110 2001a; Soltis et al., 2001b; Soltis et al., 1996; Vargas, 2000; Xiang 111 et al., 2012; Zhu et al., 2013) Brochmann et al., 1998; 116 Godsoe et al., 2013; Ness et al., 1989; Segraves and Thompson, 117 1999; Soltis and Doyle, 1987; Soltis and 118 Rieseberg, 1986; Soltis and Soltis, 1989; Soltis and Soltis, 1999; 119 Soltis and Soltis, 1988; Soltis and Soltis, 1989; Soltis et al., 1993; 120 Soltis et al., 1992; Soltis et al., 1991; Soltis et al., 1990; Soltis 121 et al., 1989a; Soltis, 1984; Wolf et al., 1990), 122 biogeography/phylogeography (Oliver et al., 2006; Reisch, 2008; 123 Szövényi et al., 2009; Westergaard et al., 2010), and evolution 124 and diversification (Holten et al., 2006; Kuzoff et al., 2001; Kuzoff 125 et al., 1999; Okuyama et al., 2005; Soltis et al., 2001a; Steen 126 et al., 2000; Vargas et al., 1999; Winkler et al., 2012) Increasing amounts of sequence data and wider taxon sampling 147 have yielded better resolution, support, and evolutionary insights 148 throughout the tree of life (Heath et al., 2008; Moore et al., 2007; 149 Parfrey et al., 2010; Soltis et al., 2011), Ribes (Grossulariaceae), and Liquidambar (Altingiaceae) 178 were selected as outgroups based on previous studies (Fishbein 179 et al., 2001; Jian et al., 2008; Soltis et al., 2013; Soltis et al., 2001a). ...
... Molecular phylogenies in plants are traditionally based on chloroplast DNA (cpDNA) sequence variation (Despres et al., 2003). The low evolutionary rate of these sequences limits the power of cpDNA for the assignment at the genus or species level (Soltis 1993). When both cpDNA and ITS sequencing fail to resolve phylogenies, the AFLP approach has the potential to solve such difficulties, particularly among closely related species, or at the intra-specific level (Koopman et al., 2001). ...
... Don) Engl. Saxifraga L. is the largest genus in the family Saxifragaceae (Soltis et al. 2001) and consists of about 400 species (Webb 1993;Soltis et al. 1993Soltis et al. , 1996Healy and Gillespie 2004). Engler (1930) reported that Saxifragaceae are morphologically diverse assemblage of annual, biennial and perennial herbs, shrubs, trees and vines. ...
... It comprises 17 subfamilies (Schulze-Menz 1964). In recent attempts to clarify the circumscription and relationships of the morphologically diverse members of Saxifragaceae using molecular systematic tools (18S rDNA and rbcL), Soltis and Soltis (1997) and Soltis et al. (1993Soltis et al. ( , 2001 concluded that Saxifragaceae is a polyphyletic family of the Saxifragales clade, that encompasses the subfamilies of Rosidae, Dilleniidae and Hamamelidae. Soltis and Soltis (1997) suggested that many of the families (Cunoniaceae, Droseraceae, Cephalotaceae, Gunneraceae, Rosaceae and Greyiaceae) traditionally considered close relatives of Saxifragaceae are only distantly related to this narrowly defined family. ...
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Two varieties of Saxifraga diversifolia, diversifolia and parnassifolia, collected from the moist alpine slopes around Gauri Kund, 3930 m (Manimahesh Hills, Himachal Pradesh, India), are studied for detailed male meiosis, chiasma frequency and pollen fertility. Both varieties exist at diploid level (based on x = 8), and show a meiotic chromosome count of n = 8 at diakinesis and metaphase-I, and regular chromosome distributions at anaphases-I/II. The meiotic chromosome count of n = 8 ascertained here represents a new aneuploid cytotype, supplementing the earlier report of a diploid cytotype with 2n = 20 from the north-west Himalayas in India and the Nepal Himalayas. Of the two varieties, var. diversifolia showed the presence of multiple chromosomal associations and univalent chromosomes at diakinesis and metaphase-I of meiosis-I. On the other hand, var. parnassifolia does not have multivalent formation but showed only 2–4 univalent chromosomes at diakinesis and metaphase-I. Occurrence of univalents in pollen mother cells of var. diversifolia and parnassifolia reduced the chiasma frequency significantly and also caused some pollen sterility (7–8%). The paper herein discusses for the first time the occurrence of structural heterozygosity and univalent chromosomes and their apparent affect on chiasma frequency and pollen fertility in S. diversifolia.
... This systematic treatment, supported by numerous authors in the last years (e.g. Brouillet and Elvander 2007b;Brouillet and Gornall 2007;Gornall et al. 2000;McGregor 2008;Wells and Elvander 2007), is based on phylogenetic analyses of chloroplast DNA (cpDNA) restriction sites (Soltis et al. 1993), sequences of the cpDNA regions such as matK, rbcL, trnL-trnF and psbA-trnH Soltis 1994, 1995;Soltis et al. 1996Soltis et al. , 2001, and nuclear ribosomal sequences from the internal transcribed spacer (ITS) regions (Johnson and Soltis 1998). ...
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Recent phylogenetic studies have shown that Saxifraga, as currently understood, must be divided into two genera: Saxifraga L. sensu stricto and Micranthes Haw. To better understand the evolutionary history of these two genera, we performed phylogenetic analyses inferred from the nuclear ribosomal sequences from the internal transcribed spacer and the sequences of the plastid DNA (rbcL). Our molecular data confirmed the monophyly of the genus Micranthes and the consistency of the existing systematic treatments based on morphological criteria. Moreover, Micranthes species native from the Iberian Peninsula (i.e. M. clusii, M. lepismigena and M. stellaris) should be included into Micranthes sect. Arabisa.
... The subtribe was later raised to the tribal level as Astilbeae of subfamily Saxifragoideae (Schulze-Menz, 1964). Recent molecular studies did not support the traditional classification of some genera and tribes, especially concerning tribe Astilbeae (Soltis et al., 1993Soltis et al., , 2001). Cytology and flavonoid chemistry also suggested that Astilbeae is not monophyletic (Bohm & Bhat, 1985;Soltis, 1986). ...
... Cytology and flavonoid chemistry also suggested that Astilbeae is not monophyletic (Bohm & Bhat, 1985;Soltis, 1986). Rodgersia and Astilboides have been shown to be closely related to Darmera Voss, Mukdenia Koidzumi, Oresitrophe Bunge and Bergenia Moench of tribe Saxifrageae (Soltis, 1986;Gornall, 1989;Soltis et al., 1993Soltis et al., , 2001). All these genera share the base chromosome number of x = 17 or 15 (Hamel, 1949;Soltis, 1986;Funamoto et al., 2000). ...
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Phylogenetic analyses were conducted for Astilbe (Saxifragaceae), an Asian/eastern North American disjunct genus, using sequences of nuclear ribosomal internal transcribed spacer (ITS) and plastid matK, trnL-trnF and psbA-trnH regions. The monophyly of Astilbe is well supported by both ITS and plastid sequences. Topological incongruence was detected between the plastid and the ITS trees, particularly concerning the placement of the single North American species, A. biternata, which may be most probably explained by its origin involving hybridization and/or allopolyploidy with plastid capture. In Astilbe, all species with hermaphroditic flowers constitute a well-supported clade; dioecious species form a basal grade to the hermaphroditic clade. Astilbe was estimated to have split with Saxifragopsis from western North America at 20.69 Ma (95% HPD: 12.14–30.22 Ma) in the early Miocene. This intercontinental disjunction between Astilbe and Saxifragopsis most likely occurred via the Bering land bridge. The major clade of Astilbe (all species of the genus excluding A. platyphylla) was inferred to have a continental Asian origin. At least three subsequent migrations or dispersals were hypothesized to explain the expansion of Astilbe into North America, Japan and tropical Asian islands. The intercontinental disjunct lineage in Astilbe invokes a hybridization event either in eastern Asia or in North America. This disjunction in Astilbe may be explained by a Beringian migration around 3.54 Ma (95% high posterior density: 1.29–6.18 Ma) in the late Tertiary, although long-distance dispersal from eastern Asia to North America is also likely. The biogeographical connection between continental Asia, Taiwan, the Philippines and other tropical Asian islands in Astilbe provides evidence for the close floristic affinity between temperate or alpine south-western China and tropical Asia. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, ●●, ●●–●●.
... Familial circumscriptions and nomenclature are based on the treatment of the Saxifragaceae by Soltis et al. (2001a) and Soltis (2007). Outgroup taxa were selected based on the well supported placement of Iteaceae, Grossulariaceae, and Pterostemonaceae as closest relatives to Saxifragaceae (Morgan and Soltis, 1993; Soltis et al., 1993 Soltis et al., , 2001a Johnson and Soltis, 1995 ). Therefore, one species from Pterostemonaceae (Pterostemon rotundifolius Ramírez), two species of Iteaceae (Itea virginica L., I. yunnanensis Franch.), and five species of Grossulariaceae (Ribes glaciale Wall., R. maximowiczianum Kom., R. moupinense Franch. ...
... In total, the study material consisted of 74 accessions representing 71 species, 25 of which were newly sequenced as part of this study. Other sequences came from previous studies (Soltis et al., 1991aSoltis et al., , 1993Soltis et al., , 1996aSoltis et al., ,b, 2001a Soltis and Kuzoff, 1995; Soltis, 1995, 1998; Kuzoff et al., 1998 Kuzoff et al., , 1999 Conti et al., 1999; Fishbein et al., 2001; Senters and Soltis, 2003; Okuyama et al., 2005 Okuyama et al., , 2008 Okuyama and Kato, 2009).Table 1 Voucher information and GenBank accession numbers for taxa used in this study. Vouch specimens are deposited in the following herbaria: K = Royalo Botanic Gardens, Kew, England; KUN = Kunming Institute of Botany, China; OS = Ohio State University, United States; UC = University of California, United States; WA = University of Warsaw, Poland; WS = Washington State University, United States; UBC = University of British Columbia, Canada; WTU = University of Washington, United States. ...
... The Saxifragaceae are a morphologically highly diverse group, variability of characters such as carpels, sepals, petals, stamens, is generally considered to be of taxonomic importance. Morphological characters supporting the clades found in the molecular analyses are as yet unknown or limited, although some morphological trends, which corroborate the relationships based on DNA sequence date, are becoming apparent (Nakazawa et al., 1997; Soltis et al., 1993 Soltis et al., , 2001a). While there are some broad morphological trends within clades it remains difficult to identify morphological synapomorphies. ...
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Saniculiphyllum, a monotypic genus distributed in Southwest China, was thought to be extinct before our recent rediscovery. The taxonomic position of this genus has been enigmatic ever since its publication. It was originally treated as the only member of a distinct tribe Saniculiphylleae in the family Saxifragaceae. Some proposed a new family, Saniculophyllaceae, to accommodate this genus, although its affinities are clearly with members of Saxifragaceae. Here we analyzed six DNA regions, the nuclear ribosomal ITS and 26S rDNA and the plastid rbcL, matK, trnL-trnF, psbA-trnH genes, spacers, and intron to explore the phylogenetic position of Saniculiphyllum within Saxifragaceae. The combined nuclear and chloroplast dataset includes 63 ingroup species, representing all genera but Hieronymusia in the family. Results from likelihood, parsimony and Bayesian phylogenetic methods corroborate earlier results. Two clades of Saxifragaceae, the Heucheroid and Saxifragoid clades, were recovered. The topologies obtained from different analyses confirm the placement of Saniculiphyllum in Saxifragaceae, but our analyses reveal that Saniculiphyllum is embedded within the large Heucheroid clade. However, the closest relatives of Saniculiphyllum within the Heucheroid clade remain unclear. Combined with morphological data, our results suggest that Saniculiphyllum should best be regarded as a highly distinctive lineage within the Heucheroid clade of Saxifragaceae. Morphological novelties and conservation status of Saniculiphyllum are also presented.
... When molecular sequences were not available for the species surveyed for morphology, sequences from close relatives were used instead (names and GenBank accession numbers provided). Numbers in parenthesis indicate the references for molecular data: (1) Zurawski et al. (1986), (2) Albert et al. (1992), (3) Fernando et al. (1993), (4) Soltis et al. (1993), (5) Morgan et al. (1994), (6) Ka¨ss and Wink (1996), (7) Sheahan and Chase (1996), (8) Doyle et al. (1997), (9) Swensen et al. (1998), (10) Thulin et al. (1998), (11) Savolainen et al. (2000), (12) Yokoyama et al. (2000), (13) Kajita et al. (2001), (14) Kita and Kato (2001), (15) Lia et al. (2001), (16) Kamiya et al. (2002), (17) fig. 1) for Atroxima and Carpolobia; Rudall and Bateman (2002) for orchids]. The opposite disposition of the flowers of Polygalaceae and Leguminosae fig. 1) is not the result of resupination as coded here, but by an opposite arrangement of the floral organs 4. Hypanthium (0) absent; (1) present. ...
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Relationships between the four families placed in the angiosperm order Fabales (Leguminosae, Polygalaceae, Quillajaceae, Surianaceae) were hitherto poorly resolved. We combine published molecular data for the chloroplast regions matK and rbcL with 66 morphological characters surveyed for 73 ingroup and two outgroup species, and use Parsimony and Bayesian approaches to explore matrices with different missing data. All combined analyses using Parsimony recovered the topology Polygalaceae (Leguminosae (Quillajaceae + Surianaceae)). Bayesian analyses with matched morphological and molecular sampling recover the same topology, but analyses based on other data recover a different Bayesian topology: ((Polygalaceae + Leguminosae) (Quillajaceae + Surianaceae)). We explore the evolution of floral characters in the context of the more consistent topology: Polygalaceae (Leguminosae (Quillajaceae + Surianaceae)). This reveals synapomorphies for (Leguminosae (Quillajaceae + Surianaceae)) as the presence of free filaments and marginal/ventral placentation, for (Quillajaceae + Surianaceae) as pentamery and apocarpy, and for Leguminosae the presence of an abaxial median sepal and unicarpellate gynoecium. An octamerous androecium is synapomorphic for Polygalaceae. The development of papilionate flowers, and the evolutionary context in which these phenotypes appeared in Leguminosae and Polygalaceae, shows that the morphologies are convergent rather than synapomorphic within Fabales.
... This approach has proved to be very powerful at the family level through the sequencing of coding regions such as rbcL[16]. However, low evolutionary rate of these sequences limits the power of cpDNA for the assignment at the genus or species level[17]. As a consequence, the relationships among closely related taxa have been inferred using non-coding sequences[18]. ...
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