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Transcribed

Villarsia in Australia

Taxonomic revision of Villarsia (Menyanthaceae) in Australia: Molecular and morphological evidence Nicholas P. Tippery', Donald H. Les', Donald Padgett?, and Surrey W. L. Jacobs3 OF ST NE 1Department of Ecology and Evolutionary Biology, College of Liberal Arts and Sciences, University of Connecticut, Storrs, Connecticut, U.S.A.; 2Bridgewater State College, Bridgewater, Massachusetts, U.S.A.; ³National Herbarium of New South Wales, Sydney, New South Wales, Australia 1881 Figure 4. Seeds of VWarsla and related taxa. Composite of equal-magnification ight micrographs. Row 1: Nym- phoides anigua, V. violifolia, V. capitata, V. congestifiora. Row 2: V. latiola, V. exaltata, Liparophylum gunni, V. lasi- osperma. Row 3: V. capensis, V. marvingiana, Row 4: V. albifiora, V. pamassifolla, V. calthifolia, V. marchanti. Row 5: V. submersa, V. reniforms, V. umbricola var, umbricola, V. umbricola var. beaugle- holei. Discriminating seed features include vestiture Figure 2. Phylogenetic relationships in Menyan- thaceae, focusing on spe- cies currently circumscribed within Vilarsa and related taxa. The tree was con- Abstract 100 K Nymphoides (40-50 spp.) 1.0 The aquatic and wetland plant family Menyanthaceae consists of five genera that are distributed worldwide. Phylogenetic analyses using both molecular and morphological data consistently have resolved two monotypic genera with boreal distributions, Menyanthes and Nephvophylidlum, as a clade sister to the remaining genera. Two of the latter, Liparophyllum (1 sp.) and Vilarsia (18 spp.), are restricted to the southem hemisphere, and species of Nymphoides (40-50 spp.) are found worldwide. Analysis of combined molecular (nrITS, tmK/matk, rbcL) and morphological data resolved several relationships that were inconsistent with current generic circumscriptions under a criterion of monophyly, among which the most conspicuous is the paraphyly of VNarsia relative to Nympholdes. Circumscriptions of these genera have fluctuated historically, with species in Australia (where both Nymphoides and VWarsia are the most diverse) belonging entirely to one or the other genus according to different authors. Recent sampling of nearly all species of VMarsla and of the most morphologically diverse and broadly distributed species of Nymphoides revealed heretofore undescribed relationships, wherein VWasia resolved to three clades: two native to Australia and one, containing the type species V. capensis, comprising only South African taxa. Species of Nymphoides resolved to a single clade, with the exception of the anomalous species N. exigua. Revised taxonomic designations have been provided for species of Villarsia in Australia and for N. exigua, in order to reflect the phylogenetic relationships that were recovered in our study. Several distinguishing morphological characters were found to support these relationships, particularly aspects of seed shape and vestiture. 100 Nymphoides exigua 1.0 Villarsla violifolia structed using combined morphological and moleo- ular (nrITS, matK, rbcl, and tmk introns) data (analyses described in Tippery et al (2008), with the addition of two Vlarsia species (V. submersa arnd V. viofola). Topology represents the single most-parsimonious tree; node values indicate 98 100 1.0 1.0 100 Villarsia capitata 1.0 100 1.0 68 55 Villarsia congestiflora VIllarsla latifolia 100 1.0 Villarsia exaltata .07 Liparophyllum gunnil 63 parsimony bootstrap above and Bayesian posterior (tuberculate or smooth), tub- ercle density, hium location (terminal or subterminal, and the presence of a nutritive caruncle at the hilum. The top eight seeds represent the "VWarsia" 1 clade, the bottom Villarsia lasiosperma probability support below each node. The tree was rooted using sequences from Monyanthes and Nephrophylidum, prior analysis using out- group taxa from the related families Asteraceae, Caly- ceraceae, and Goodenia- ceae revealed them to be the sister clade to Liparo- phyllum, Nymphoides, and Vlarsia (Tippery et al. 2008). Taxa in the "Vilarsia" 1 and "Vilarsia" 2 clades are native to Australia, while V. capensis and V. man- ningiana are found only in South Africa. Asterisk in- dicates the type species for Vilarsia (V. capensis) Villarsia capensis 100 1.0 Villarsia manningiana 100 1.0 after VIllarsla albiflora 1.0 95 1.0 72 eight belong to the "VWarsia" 2 clade, and the two large seeds in the middle are from South Villarsia parnassifolia Table 1. Taxonomic history of the genus Villarsia. Prior to 1900, several Australian Menyanthaceae had been described variously under different genera. Abbreviations: L: Liparophyllum Hook.f.: M: Menyanthes L.; N: Nymphaides Ség. (syn. Limnanthemum S.G.Gmel.); S: Swertie L. (Gentiaraceae); V: Valarsia Vent. (syn. Renealmia Houtt.). Notes: 1. Mueller (1868) included Nymphoides as a section of Wlarsia. 2. Mueller (1875) transferred "all species of VWarsia to Wympholdes)", citing instances of other congeneric plant species among which differences were more pronounced translation and paraphrase by N.P.T.). Villarsia calthifolia African Warsia 82 species. 1.0 Villarsia marchantil 100 1.0 Villarsia submersa 100 1.0 Villarsia reniformis Discussion Villarsia umbricola N. aurantiaca 1850 N - NNN N Dalzel) Kuntze N. crenata (F.Muell.) Kurtze N. exiWlora Despite overall similarity among Vilarsia species, analysis of combined morphological and molecular data indicated that the genus as currently circumscribed comprises three clades (Fig. 2). In addition, two taxa traditionally placed outside the genus, Liparophylum gunni and Nymphaides exigua, were resolved within the same clade ("VWarsia" 1) along with several Varsia species. After more careful examination of the characters that have been used to OUTGROUPPMenyanthes (1 sp.) 1854 N - NN NN Nephrophyllidium (1 sp.) 1865 v - NNNN (F.Muell.) Kuntze N. geminata (RBr.) Kuntze N. indica LI Kuntze distinguish between Villarsia and Nymphoides (i.e., inflorescence architecture, capsule dehiscence), we hypothesized that similarities among Vilarsia taxa represent traits inherited from their shared ancestor (1.8., plesiomorphic traits) rather than features unique to a monophyletic lineage. Researchers previously have suggested that seed features are the most diagnostic characters for identifying species of Nymphoides and Villarsia (Aston 1969; Chuang and Ornduff 1992), and we determined that they are also informative for delineating phylogenetic groups. Species of the strict Vilarsia clade (including the type species V. capensis) have seeds that are orbicular and the largest of any surveyed; members of the "Villarsia" 2 clade have seeds with a 1810 VNNNNN 1753 MN-NNN N. minima (F.Muell.) Kuntze N. parvifolia Wall) Kuntze L. gunni Hook.f. N. exigua (F.Muell.) Kuntze V. capitata Nees ex Lehm. 1858 N - NNN N subterminal hilum that are smooth or densely covered with tapering tubercles; seeds of species belonging to the "Villarsia" 1 clade have a terminal hilum, are smooth or sparsely tuberculate, and may have a nutritive caruncle, a feature that is entirely absent from the other clades. 1828 vN -- -N 1847 L - NLN - Box 2. Villarsia cambodiana Hance (syn. V. thombaidalis Dop), native to Southeast Asia, represents the only Villarsia species to be described from a tropical latitude. In April 2008, material identified as V. cambodiana was obtained from Lo Go - Xa Mat National Park (Tay Ninh Province, Vietnam). Preliminary DNA sequence analysis placed the specimen soundly within the Nymphoides clade, with N. aurantiaca (Dalzel) Kuntze as its nearest relative. Thanks to L. Averyanov, Nguyen Tien Hiep, Phan Ke Loc, P. Raven, J. Regalado, and Vu Ngoc Long. Photos: L. Aver- yanov, copyright Missouri Bota- nical Garden and Vietnam 1858 N - NN NN b 1845 v - v vNV V. congestifiora 1868 v- VVNV FMuell. V. exatata (Sol. ex Sims) G.Don 1807 M-- -NV V. lasiospema 1868 v - v vN V F.Muell. V. latifolla Benth. V. vioifola 1837 v - v v NV Botanical Conservation Program, funded by the Henry Luce Foundation; used with permission. 1868 v - vvN V F.Muell. V. capensis Houtt.) Merr. V. golablattiana Omduff V. manningiana Omduff V. ablara 1777 VV ---- v v Literature Cited Aston, H. I. 1969. The genus Warsia (Menyanthaceae) in Australa. Muelaria 2 3-63. Aston, H. I. 1973. Aquatio Plants of Australa. Melbourne University Press, Cariton, Victoria. Bentham, G. and F. von Mueler. 1859. Gentianaceae. Pp. 359382 in Flora Australionsls. Vol 4. L Reeve & Co., London. 1999 V- -- Figure 3. Habit variation in Vilarsia. Clockwise trom bottom left: V. submersa is obligately submersed, with floating leaves, a lax inflorescence, and flowers that touch the water; V. congestiflora has flowers borne in a congested cluster; V. pamassiolia represents the typical VWlarsia habit, with a basal rosette of leaves and an erect, expanded inflorescence. Photos: S. Jacobs. 2001 V. Chusng T. I, and R. Omduff, 1992. Seed morphology and systematics of Menyanthaceae. American Journal of Botany 79 1396-1406. Grisebach, A. H. R. 1845. Tribus I. Manyantheae. Pp. 136--141 in A P. de Candole, ed. Prodromus Systematis Noturalis Regni Vegetsbits. Vol. 9. Sumptibus Sociorum Treuttel et Würtz, Paris Mueller, F. J. H. von. 1868. Fragmenta Phytographise Australae. Vol. 6. Auctoritate Gubern. Colonise Victorise. Ex Officina Joannis Ferres, Melbourne. 1860 v - vv NV F.Muell. V. cathifolia F.Muell. V. parmessifolia Labil.) R.Br. V. reniformis 1868 v - VVNV Box 1. Liparophyllum gunnil has remained separate from other Menyanthaceae genera because of its many anomalous morphological features, including linear keaves, solitary flowers, and the absence of carpellary glands. It occupies high-altitude lakes in Tasmania and New Zealand and is the only Menyanthaceae species native to the latter. Photos: M. 1806 sv v vNV Mueller, F. J. H. van. 1875. Fragmenta Phytographiao Australao. Vol. 9. Auctoritate Gubern. Coloniae Victoriae, Ex Officina Joannis Ferres, Melbourne. Figure 1. Villarsia flowers, Clockwise from bottom left: V. manninglana photo: R. Prunier; all ather photos S. Jacobs), V. exaltata, V. lasiosperma, V. latifola, V. congestilora, V. capitata, V. abifora, V. parmassifolia, V. cahifolia, V. marchanti, V. submersa, V. reniformis. 1810 v - v v NV Tppery. N. P., D. H. Les, D. J. Padgett, and S. W. L. Jacobs. 2008. Generic circumscription in Meryanthacese: A phylogenetic evaluation. Systomatic Botany. In preas. R.Br. IV. submersa Acknowledgments 1969 V ---- V Aston V. umbricola Aston The authors are indebted to the directors and staff of the following herbaria for providing access to specimens and type material: A, BM, BRI, CAL, CONN, FI, GH, GOET, K, MEL, MO, NSW, NT, NY, PERTH, PRE, UC, US. We also thank Helen Aston for her valuable insights on the Australian taxa 1969 V- -- Wilcox. "Villarsia" 2 VIllarsia lymphoides "Villarsia" 2 Villarsia" Taxonomic revision of Villarsia (Menyanthaceae) in Australia: Molecular and morphological evidence Nicholas P. Tippery', Donald H. Les', Donald Padgett?, and Surrey W. L. Jacobs3 OF ST NE 1Department of Ecology and Evolutionary Biology, College of Liberal Arts and Sciences, University of Connecticut, Storrs, Connecticut, U.S.A.; 2Bridgewater State College, Bridgewater, Massachusetts, U.S.A.; ³National Herbarium of New South Wales, Sydney, New South Wales, Australia 1881 Figure 4. Seeds of VWarsla and related taxa. Composite of equal-magnification ight micrographs. Row 1: Nym- phoides anigua, V. violifolia, V. capitata, V. congestifiora. Row 2: V. latiola, V. exaltata, Liparophylum gunni, V. lasi- osperma. Row 3: V. capensis, V. marvingiana, Row 4: V. albifiora, V. pamassifolla, V. calthifolia, V. marchanti. Row 5: V. submersa, V. reniforms, V. umbricola var, umbricola, V. umbricola var. beaugle- holei. Discriminating seed features include vestiture Figure 2. Phylogenetic relationships in Menyan- thaceae, focusing on spe- cies currently circumscribed within Vilarsa and related taxa. The tree was con- Abstract 100 K Nymphoides (40-50 spp.) 1.0 The aquatic and wetland plant family Menyanthaceae consists of five genera that are distributed worldwide. Phylogenetic analyses using both molecular and morphological data consistently have resolved two monotypic genera with boreal distributions, Menyanthes and Nephvophylidlum, as a clade sister to the remaining genera. Two of the latter, Liparophyllum (1 sp.) and Vilarsia (18 spp.), are restricted to the southem hemisphere, and species of Nymphoides (40-50 spp.) are found worldwide. Analysis of combined molecular (nrITS, tmK/matk, rbcL) and morphological data resolved several relationships that were inconsistent with current generic circumscriptions under a criterion of monophyly, among which the most conspicuous is the paraphyly of VNarsia relative to Nympholdes. Circumscriptions of these genera have fluctuated historically, with species in Australia (where both Nymphoides and VWarsia are the most diverse) belonging entirely to one or the other genus according to different authors. Recent sampling of nearly all species of VMarsla and of the most morphologically diverse and broadly distributed species of Nymphoides revealed heretofore undescribed relationships, wherein VWasia resolved to three clades: two native to Australia and one, containing the type species V. capensis, comprising only South African taxa. Species of Nymphoides resolved to a single clade, with the exception of the anomalous species N. exigua. Revised taxonomic designations have been provided for species of Villarsia in Australia and for N. exigua, in order to reflect the phylogenetic relationships that were recovered in our study. Several distinguishing morphological characters were found to support these relationships, particularly aspects of seed shape and vestiture. 100 Nymphoides exigua 1.0 Villarsla violifolia structed using combined morphological and moleo- ular (nrITS, matK, rbcl, and tmk introns) data (analyses described in Tippery et al (2008), with the addition of two Vlarsia species (V. submersa arnd V. viofola). Topology represents the single most-parsimonious tree; node values indicate 98 100 1.0 1.0 100 Villarsia capitata 1.0 100 1.0 68 55 Villarsia congestiflora VIllarsla latifolia 100 1.0 Villarsia exaltata .07 Liparophyllum gunnil 63 parsimony bootstrap above and Bayesian posterior (tuberculate or smooth), tub- ercle density, hium location (terminal or subterminal, and the presence of a nutritive caruncle at the hilum. The top eight seeds represent the "VWarsia" 1 clade, the bottom Villarsia lasiosperma probability support below each node. The tree was rooted using sequences from Monyanthes and Nephrophylidum, prior analysis using out- group taxa from the related families Asteraceae, Caly- ceraceae, and Goodenia- ceae revealed them to be the sister clade to Liparo- phyllum, Nymphoides, and Vlarsia (Tippery et al. 2008). Taxa in the "Vilarsia" 1 and "Vilarsia" 2 clades are native to Australia, while V. capensis and V. man- ningiana are found only in South Africa. Asterisk in- dicates the type species for Vilarsia (V. capensis) Villarsia capensis 100 1.0 Villarsia manningiana 100 1.0 after VIllarsla albiflora 1.0 95 1.0 72 eight belong to the "VWarsia" 2 clade, and the two large seeds in the middle are from South Villarsia parnassifolia Table 1. Taxonomic history of the genus Villarsia. Prior to 1900, several Australian Menyanthaceae had been described variously under different genera. Abbreviations: L: Liparophyllum Hook.f.: M: Menyanthes L.; N: Nymphaides Ség. (syn. Limnanthemum S.G.Gmel.); S: Swertie L. (Gentiaraceae); V: Valarsia Vent. (syn. Renealmia Houtt.). Notes: 1. Mueller (1868) included Nymphoides as a section of Wlarsia. 2. Mueller (1875) transferred "all species of VWarsia to Wympholdes)", citing instances of other congeneric plant species among which differences were more pronounced translation and paraphrase by N.P.T.). Villarsia calthifolia African Warsia 82 species. 1.0 Villarsia marchantil 100 1.0 Villarsia submersa 100 1.0 Villarsia reniformis Discussion Villarsia umbricola N. aurantiaca 1850 N - NNN N Dalzel) Kuntze N. crenata (F.Muell.) Kurtze N. exiWlora Despite overall similarity among Vilarsia species, analysis of combined morphological and molecular data indicated that the genus as currently circumscribed comprises three clades (Fig. 2). In addition, two taxa traditionally placed outside the genus, Liparophylum gunni and Nymphaides exigua, were resolved within the same clade ("VWarsia" 1) along with several Varsia species. After more careful examination of the characters that have been used to OUTGROUPPMenyanthes (1 sp.) 1854 N - NN NN Nephrophyllidium (1 sp.) 1865 v - NNNN (F.Muell.) Kuntze N. geminata (RBr.) Kuntze N. indica LI Kuntze distinguish between Villarsia and Nymphoides (i.e., inflorescence architecture, capsule dehiscence), we hypothesized that similarities among Vilarsia taxa represent traits inherited from their shared ancestor (1.8., plesiomorphic traits) rather than features unique to a monophyletic lineage. Researchers previously have suggested that seed features are the most diagnostic characters for identifying species of Nymphoides and Villarsia (Aston 1969; Chuang and Ornduff 1992), and we determined that they are also informative for delineating phylogenetic groups. Species of the strict Vilarsia clade (including the type species V. capensis) have seeds that are orbicular and the largest of any surveyed; members of the "Villarsia" 2 clade have seeds with a 1810 VNNNNN 1753 MN-NNN N. minima (F.Muell.) Kuntze N. parvifolia Wall) Kuntze L. gunni Hook.f. N. exigua (F.Muell.) Kuntze V. capitata Nees ex Lehm. 1858 N - NNN N subterminal hilum that are smooth or densely covered with tapering tubercles; seeds of species belonging to the "Villarsia" 1 clade have a terminal hilum, are smooth or sparsely tuberculate, and may have a nutritive caruncle, a feature that is entirely absent from the other clades. 1828 vN -- -N 1847 L - NLN - Box 2. Villarsia cambodiana Hance (syn. V. thombaidalis Dop), native to Southeast Asia, represents the only Villarsia species to be described from a tropical latitude. In April 2008, material identified as V. cambodiana was obtained from Lo Go - Xa Mat National Park (Tay Ninh Province, Vietnam). Preliminary DNA sequence analysis placed the specimen soundly within the Nymphoides clade, with N. aurantiaca (Dalzel) Kuntze as its nearest relative. Thanks to L. Averyanov, Nguyen Tien Hiep, Phan Ke Loc, P. Raven, J. Regalado, and Vu Ngoc Long. Photos: L. Aver- yanov, copyright Missouri Bota- nical Garden and Vietnam 1858 N - NN NN b 1845 v - v vNV V. congestifiora 1868 v- VVNV FMuell. V. exatata (Sol. ex Sims) G.Don 1807 M-- -NV V. lasiospema 1868 v - v vN V F.Muell. V. latifolla Benth. V. vioifola 1837 v - v v NV Botanical Conservation Program, funded by the Henry Luce Foundation; used with permission. 1868 v - vvN V F.Muell. V. capensis Houtt.) Merr. V. golablattiana Omduff V. manningiana Omduff V. ablara 1777 VV ---- v v Literature Cited Aston, H. I. 1969. The genus Warsia (Menyanthaceae) in Australa. Muelaria 2 3-63. Aston, H. I. 1973. Aquatio Plants of Australa. Melbourne University Press, Cariton, Victoria. Bentham, G. and F. von Mueler. 1859. Gentianaceae. Pp. 359382 in Flora Australionsls. Vol 4. L Reeve & Co., London. 1999 V- -- Figure 3. Habit variation in Vilarsia. Clockwise trom bottom left: V. submersa is obligately submersed, with floating leaves, a lax inflorescence, and flowers that touch the water; V. congestiflora has flowers borne in a congested cluster; V. pamassiolia represents the typical VWlarsia habit, with a basal rosette of leaves and an erect, expanded inflorescence. Photos: S. Jacobs. 2001 V. Chusng T. I, and R. Omduff, 1992. Seed morphology and systematics of Menyanthaceae. American Journal of Botany 79 1396-1406. Grisebach, A. H. R. 1845. Tribus I. Manyantheae. Pp. 136--141 in A P. de Candole, ed. Prodromus Systematis Noturalis Regni Vegetsbits. Vol. 9. Sumptibus Sociorum Treuttel et Würtz, Paris Mueller, F. J. H. von. 1868. Fragmenta Phytographise Australae. Vol. 6. Auctoritate Gubern. Colonise Victorise. Ex Officina Joannis Ferres, Melbourne. 1860 v - vv NV F.Muell. V. cathifolia F.Muell. V. parmessifolia Labil.) R.Br. V. reniformis 1868 v - VVNV Box 1. Liparophyllum gunnil has remained separate from other Menyanthaceae genera because of its many anomalous morphological features, including linear keaves, solitary flowers, and the absence of carpellary glands. It occupies high-altitude lakes in Tasmania and New Zealand and is the only Menyanthaceae species native to the latter. Photos: M. 1806 sv v vNV Mueller, F. J. H. van. 1875. Fragmenta Phytographiao Australao. Vol. 9. Auctoritate Gubern. Coloniae Victoriae, Ex Officina Joannis Ferres, Melbourne. Figure 1. Villarsia flowers, Clockwise from bottom left: V. manninglana photo: R. Prunier; all ather photos S. Jacobs), V. exaltata, V. lasiosperma, V. latifola, V. congestilora, V. capitata, V. abifora, V. parmassifolia, V. cahifolia, V. marchanti, V. submersa, V. reniformis. 1810 v - v v NV Tppery. N. P., D. H. Les, D. J. Padgett, and S. W. L. Jacobs. 2008. Generic circumscription in Meryanthacese: A phylogenetic evaluation. Systomatic Botany. In preas. R.Br. IV. submersa Acknowledgments 1969 V ---- V Aston V. umbricola Aston The authors are indebted to the directors and staff of the following herbaria for providing access to specimens and type material: A, BM, BRI, CAL, CONN, FI, GH, GOET, K, MEL, MO, NSW, NT, NY, PERTH, PRE, UC, US. We also thank Helen Aston for her valuable insights on the Australian taxa 1969 V- -- Wilcox. "Villarsia" 2 VIllarsia lymphoides "Villarsia" 2 Villarsia" Taxonomic revision of Villarsia (Menyanthaceae) in Australia: Molecular and morphological evidence Nicholas P. Tippery', Donald H. Les', Donald Padgett?, and Surrey W. L. Jacobs3 OF ST NE 1Department of Ecology and Evolutionary Biology, College of Liberal Arts and Sciences, University of Connecticut, Storrs, Connecticut, U.S.A.; 2Bridgewater State College, Bridgewater, Massachusetts, U.S.A.; ³National Herbarium of New South Wales, Sydney, New South Wales, Australia 1881 Figure 4. Seeds of VWarsla and related taxa. Composite of equal-magnification ight micrographs. Row 1: Nym- phoides anigua, V. violifolia, V. capitata, V. congestifiora. Row 2: V. latiola, V. exaltata, Liparophylum gunni, V. lasi- osperma. Row 3: V. capensis, V. marvingiana, Row 4: V. albifiora, V. pamassifolla, V. calthifolia, V. marchanti. Row 5: V. submersa, V. reniforms, V. umbricola var, umbricola, V. umbricola var. beaugle- holei. Discriminating seed features include vestiture Figure 2. Phylogenetic relationships in Menyan- thaceae, focusing on spe- cies currently circumscribed within Vilarsa and related taxa. The tree was con- Abstract 100 K Nymphoides (40-50 spp.) 1.0 The aquatic and wetland plant family Menyanthaceae consists of five genera that are distributed worldwide. Phylogenetic analyses using both molecular and morphological data consistently have resolved two monotypic genera with boreal distributions, Menyanthes and Nephvophylidlum, as a clade sister to the remaining genera. Two of the latter, Liparophyllum (1 sp.) and Vilarsia (18 spp.), are restricted to the southem hemisphere, and species of Nymphoides (40-50 spp.) are found worldwide. Analysis of combined molecular (nrITS, tmK/matk, rbcL) and morphological data resolved several relationships that were inconsistent with current generic circumscriptions under a criterion of monophyly, among which the most conspicuous is the paraphyly of VNarsia relative to Nympholdes. Circumscriptions of these genera have fluctuated historically, with species in Australia (where both Nymphoides and VWarsia are the most diverse) belonging entirely to one or the other genus according to different authors. Recent sampling of nearly all species of VMarsla and of the most morphologically diverse and broadly distributed species of Nymphoides revealed heretofore undescribed relationships, wherein VWasia resolved to three clades: two native to Australia and one, containing the type species V. capensis, comprising only South African taxa. Species of Nymphoides resolved to a single clade, with the exception of the anomalous species N. exigua. Revised taxonomic designations have been provided for species of Villarsia in Australia and for N. exigua, in order to reflect the phylogenetic relationships that were recovered in our study. Several distinguishing morphological characters were found to support these relationships, particularly aspects of seed shape and vestiture. 100 Nymphoides exigua 1.0 Villarsla violifolia structed using combined morphological and moleo- ular (nrITS, matK, rbcl, and tmk introns) data (analyses described in Tippery et al (2008), with the addition of two Vlarsia species (V. submersa arnd V. viofola). Topology represents the single most-parsimonious tree; node values indicate 98 100 1.0 1.0 100 Villarsia capitata 1.0 100 1.0 68 55 Villarsia congestiflora VIllarsla latifolia 100 1.0 Villarsia exaltata .07 Liparophyllum gunnil 63 parsimony bootstrap above and Bayesian posterior (tuberculate or smooth), tub- ercle density, hium location (terminal or subterminal, and the presence of a nutritive caruncle at the hilum. The top eight seeds represent the "VWarsia" 1 clade, the bottom Villarsia lasiosperma probability support below each node. The tree was rooted using sequences from Monyanthes and Nephrophylidum, prior analysis using out- group taxa from the related families Asteraceae, Caly- ceraceae, and Goodenia- ceae revealed them to be the sister clade to Liparo- phyllum, Nymphoides, and Vlarsia (Tippery et al. 2008). Taxa in the "Vilarsia" 1 and "Vilarsia" 2 clades are native to Australia, while V. capensis and V. man- ningiana are found only in South Africa. Asterisk in- dicates the type species for Vilarsia (V. capensis) Villarsia capensis 100 1.0 Villarsia manningiana 100 1.0 after VIllarsla albiflora 1.0 95 1.0 72 eight belong to the "VWarsia" 2 clade, and the two large seeds in the middle are from South Villarsia parnassifolia Table 1. Taxonomic history of the genus Villarsia. Prior to 1900, several Australian Menyanthaceae had been described variously under different genera. Abbreviations: L: Liparophyllum Hook.f.: M: Menyanthes L.; N: Nymphaides Ség. (syn. Limnanthemum S.G.Gmel.); S: Swertie L. (Gentiaraceae); V: Valarsia Vent. (syn. Renealmia Houtt.). Notes: 1. Mueller (1868) included Nymphoides as a section of Wlarsia. 2. Mueller (1875) transferred "all species of VWarsia to Wympholdes)", citing instances of other congeneric plant species among which differences were more pronounced translation and paraphrase by N.P.T.). Villarsia calthifolia African Warsia 82 species. 1.0 Villarsia marchantil 100 1.0 Villarsia submersa 100 1.0 Villarsia reniformis Discussion Villarsia umbricola N. aurantiaca 1850 N - NNN N Dalzel) Kuntze N. crenata (F.Muell.) Kurtze N. exiWlora Despite overall similarity among Vilarsia species, analysis of combined morphological and molecular data indicated that the genus as currently circumscribed comprises three clades (Fig. 2). In addition, two taxa traditionally placed outside the genus, Liparophylum gunni and Nymphaides exigua, were resolved within the same clade ("VWarsia" 1) along with several Varsia species. After more careful examination of the characters that have been used to OUTGROUPPMenyanthes (1 sp.) 1854 N - NN NN Nephrophyllidium (1 sp.) 1865 v - NNNN (F.Muell.) Kuntze N. geminata (RBr.) Kuntze N. indica LI Kuntze distinguish between Villarsia and Nymphoides (i.e., inflorescence architecture, capsule dehiscence), we hypothesized that similarities among Vilarsia taxa represent traits inherited from their shared ancestor (1.8., plesiomorphic traits) rather than features unique to a monophyletic lineage. Researchers previously have suggested that seed features are the most diagnostic characters for identifying species of Nymphoides and Villarsia (Aston 1969; Chuang and Ornduff 1992), and we determined that they are also informative for delineating phylogenetic groups. Species of the strict Vilarsia clade (including the type species V. capensis) have seeds that are orbicular and the largest of any surveyed; members of the "Villarsia" 2 clade have seeds with a 1810 VNNNNN 1753 MN-NNN N. minima (F.Muell.) Kuntze N. parvifolia Wall) Kuntze L. gunni Hook.f. N. exigua (F.Muell.) Kuntze V. capitata Nees ex Lehm. 1858 N - NNN N subterminal hilum that are smooth or densely covered with tapering tubercles; seeds of species belonging to the "Villarsia" 1 clade have a terminal hilum, are smooth or sparsely tuberculate, and may have a nutritive caruncle, a feature that is entirely absent from the other clades. 1828 vN -- -N 1847 L - NLN - Box 2. Villarsia cambodiana Hance (syn. V. thombaidalis Dop), native to Southeast Asia, represents the only Villarsia species to be described from a tropical latitude. In April 2008, material identified as V. cambodiana was obtained from Lo Go - Xa Mat National Park (Tay Ninh Province, Vietnam). Preliminary DNA sequence analysis placed the specimen soundly within the Nymphoides clade, with N. aurantiaca (Dalzel) Kuntze as its nearest relative. Thanks to L. Averyanov, Nguyen Tien Hiep, Phan Ke Loc, P. Raven, J. Regalado, and Vu Ngoc Long. Photos: L. Aver- yanov, copyright Missouri Bota- nical Garden and Vietnam 1858 N - NN NN b 1845 v - v vNV V. congestifiora 1868 v- VVNV FMuell. V. exatata (Sol. ex Sims) G.Don 1807 M-- -NV V. lasiospema 1868 v - v vN V F.Muell. V. latifolla Benth. V. vioifola 1837 v - v v NV Botanical Conservation Program, funded by the Henry Luce Foundation; used with permission. 1868 v - vvN V F.Muell. V. capensis Houtt.) Merr. V. golablattiana Omduff V. manningiana Omduff V. ablara 1777 VV ---- v v Literature Cited Aston, H. I. 1969. The genus Warsia (Menyanthaceae) in Australa. Muelaria 2 3-63. Aston, H. I. 1973. Aquatio Plants of Australa. Melbourne University Press, Cariton, Victoria. Bentham, G. and F. von Mueler. 1859. Gentianaceae. Pp. 359382 in Flora Australionsls. Vol 4. L Reeve & Co., London. 1999 V- -- Figure 3. Habit variation in Vilarsia. Clockwise trom bottom left: V. submersa is obligately submersed, with floating leaves, a lax inflorescence, and flowers that touch the water; V. congestiflora has flowers borne in a congested cluster; V. pamassiolia represents the typical VWlarsia habit, with a basal rosette of leaves and an erect, expanded inflorescence. Photos: S. Jacobs. 2001 V. Chusng T. I, and R. Omduff, 1992. Seed morphology and systematics of Menyanthaceae. American Journal of Botany 79 1396-1406. Grisebach, A. H. R. 1845. Tribus I. Manyantheae. Pp. 136--141 in A P. de Candole, ed. Prodromus Systematis Noturalis Regni Vegetsbits. Vol. 9. Sumptibus Sociorum Treuttel et Würtz, Paris Mueller, F. J. H. von. 1868. Fragmenta Phytographise Australae. Vol. 6. Auctoritate Gubern. Colonise Victorise. Ex Officina Joannis Ferres, Melbourne. 1860 v - vv NV F.Muell. V. cathifolia F.Muell. V. parmessifolia Labil.) R.Br. V. reniformis 1868 v - VVNV Box 1. Liparophyllum gunnil has remained separate from other Menyanthaceae genera because of its many anomalous morphological features, including linear keaves, solitary flowers, and the absence of carpellary glands. It occupies high-altitude lakes in Tasmania and New Zealand and is the only Menyanthaceae species native to the latter. Photos: M. 1806 sv v vNV Mueller, F. J. H. van. 1875. Fragmenta Phytographiao Australao. Vol. 9. Auctoritate Gubern. Coloniae Victoriae, Ex Officina Joannis Ferres, Melbourne. Figure 1. Villarsia flowers, Clockwise from bottom left: V. manninglana photo: R. Prunier; all ather photos S. Jacobs), V. exaltata, V. lasiosperma, V. latifola, V. congestilora, V. capitata, V. abifora, V. parmassifolia, V. cahifolia, V. marchanti, V. submersa, V. reniformis. 1810 v - v v NV Tppery. N. P., D. H. Les, D. J. Padgett, and S. W. L. Jacobs. 2008. Generic circumscription in Meryanthacese: A phylogenetic evaluation. Systomatic Botany. In preas. R.Br. IV. submersa Acknowledgments 1969 V ---- V Aston V. umbricola Aston The authors are indebted to the directors and staff of the following herbaria for providing access to specimens and type material: A, BM, BRI, CAL, CONN, FI, GH, GOET, K, MEL, MO, NSW, NT, NY, PERTH, PRE, UC, US. We also thank Helen Aston for her valuable insights on the Australian taxa 1969 V- -- Wilcox. "Villarsia" 2 VIllarsia lymphoides "Villarsia" 2 Villarsia" Taxonomic revision of Villarsia (Menyanthaceae) in Australia: Molecular and morphological evidence Nicholas P. Tippery', Donald H. Les', Donald Padgett?, and Surrey W. L. Jacobs3 OF ST NE 1Department of Ecology and Evolutionary Biology, College of Liberal Arts and Sciences, University of Connecticut, Storrs, Connecticut, U.S.A.; 2Bridgewater State College, Bridgewater, Massachusetts, U.S.A.; ³National Herbarium of New South Wales, Sydney, New South Wales, Australia 1881 Figure 4. Seeds of VWarsla and related taxa. Composite of equal-magnification ight micrographs. Row 1: Nym- phoides anigua, V. violifolia, V. capitata, V. congestifiora. Row 2: V. latiola, V. exaltata, Liparophylum gunni, V. lasi- osperma. Row 3: V. capensis, V. marvingiana, Row 4: V. albifiora, V. pamassifolla, V. calthifolia, V. marchanti. Row 5: V. submersa, V. reniforms, V. umbricola var, umbricola, V. umbricola var. beaugle- holei. Discriminating seed features include vestiture Figure 2. Phylogenetic relationships in Menyan- thaceae, focusing on spe- cies currently circumscribed within Vilarsa and related taxa. The tree was con- Abstract 100 K Nymphoides (40-50 spp.) 1.0 The aquatic and wetland plant family Menyanthaceae consists of five genera that are distributed worldwide. Phylogenetic analyses using both molecular and morphological data consistently have resolved two monotypic genera with boreal distributions, Menyanthes and Nephvophylidlum, as a clade sister to the remaining genera. Two of the latter, Liparophyllum (1 sp.) and Vilarsia (18 spp.), are restricted to the southem hemisphere, and species of Nymphoides (40-50 spp.) are found worldwide. Analysis of combined molecular (nrITS, tmK/matk, rbcL) and morphological data resolved several relationships that were inconsistent with current generic circumscriptions under a criterion of monophyly, among which the most conspicuous is the paraphyly of VNarsia relative to Nympholdes. Circumscriptions of these genera have fluctuated historically, with species in Australia (where both Nymphoides and VWarsia are the most diverse) belonging entirely to one or the other genus according to different authors. Recent sampling of nearly all species of VMarsla and of the most morphologically diverse and broadly distributed species of Nymphoides revealed heretofore undescribed relationships, wherein VWasia resolved to three clades: two native to Australia and one, containing the type species V. capensis, comprising only South African taxa. Species of Nymphoides resolved to a single clade, with the exception of the anomalous species N. exigua. Revised taxonomic designations have been provided for species of Villarsia in Australia and for N. exigua, in order to reflect the phylogenetic relationships that were recovered in our study. Several distinguishing morphological characters were found to support these relationships, particularly aspects of seed shape and vestiture. 100 Nymphoides exigua 1.0 Villarsla violifolia structed using combined morphological and moleo- ular (nrITS, matK, rbcl, and tmk introns) data (analyses described in Tippery et al (2008), with the addition of two Vlarsia species (V. submersa arnd V. viofola). Topology represents the single most-parsimonious tree; node values indicate 98 100 1.0 1.0 100 Villarsia capitata 1.0 100 1.0 68 55 Villarsia congestiflora VIllarsla latifolia 100 1.0 Villarsia exaltata .07 Liparophyllum gunnil 63 parsimony bootstrap above and Bayesian posterior (tuberculate or smooth), tub- ercle density, hium location (terminal or subterminal, and the presence of a nutritive caruncle at the hilum. The top eight seeds represent the "VWarsia" 1 clade, the bottom Villarsia lasiosperma probability support below each node. The tree was rooted using sequences from Monyanthes and Nephrophylidum, prior analysis using out- group taxa from the related families Asteraceae, Caly- ceraceae, and Goodenia- ceae revealed them to be the sister clade to Liparo- phyllum, Nymphoides, and Vlarsia (Tippery et al. 2008). Taxa in the "Vilarsia" 1 and "Vilarsia" 2 clades are native to Australia, while V. capensis and V. man- ningiana are found only in South Africa. Asterisk in- dicates the type species for Vilarsia (V. capensis) Villarsia capensis 100 1.0 Villarsia manningiana 100 1.0 after VIllarsla albiflora 1.0 95 1.0 72 eight belong to the "VWarsia" 2 clade, and the two large seeds in the middle are from South Villarsia parnassifolia Table 1. Taxonomic history of the genus Villarsia. Prior to 1900, several Australian Menyanthaceae had been described variously under different genera. Abbreviations: L: Liparophyllum Hook.f.: M: Menyanthes L.; N: Nymphaides Ség. (syn. Limnanthemum S.G.Gmel.); S: Swertie L. (Gentiaraceae); V: Valarsia Vent. (syn. Renealmia Houtt.). Notes: 1. Mueller (1868) included Nymphoides as a section of Wlarsia. 2. Mueller (1875) transferred "all species of VWarsia to Wympholdes)", citing instances of other congeneric plant species among which differences were more pronounced translation and paraphrase by N.P.T.). Villarsia calthifolia African Warsia 82 species. 1.0 Villarsia marchantil 100 1.0 Villarsia submersa 100 1.0 Villarsia reniformis Discussion Villarsia umbricola N. aurantiaca 1850 N - NNN N Dalzel) Kuntze N. crenata (F.Muell.) Kurtze N. exiWlora Despite overall similarity among Vilarsia species, analysis of combined morphological and molecular data indicated that the genus as currently circumscribed comprises three clades (Fig. 2). In addition, two taxa traditionally placed outside the genus, Liparophylum gunni and Nymphaides exigua, were resolved within the same clade ("VWarsia" 1) along with several Varsia species. After more careful examination of the characters that have been used to OUTGROUPPMenyanthes (1 sp.) 1854 N - NN NN Nephrophyllidium (1 sp.) 1865 v - NNNN (F.Muell.) Kuntze N. geminata (RBr.) Kuntze N. indica LI Kuntze distinguish between Villarsia and Nymphoides (i.e., inflorescence architecture, capsule dehiscence), we hypothesized that similarities among Vilarsia taxa represent traits inherited from their shared ancestor (1.8., plesiomorphic traits) rather than features unique to a monophyletic lineage. Researchers previously have suggested that seed features are the most diagnostic characters for identifying species of Nymphoides and Villarsia (Aston 1969; Chuang and Ornduff 1992), and we determined that they are also informative for delineating phylogenetic groups. Species of the strict Vilarsia clade (including the type species V. capensis) have seeds that are orbicular and the largest of any surveyed; members of the "Villarsia" 2 clade have seeds with a 1810 VNNNNN 1753 MN-NNN N. minima (F.Muell.) Kuntze N. parvifolia Wall) Kuntze L. gunni Hook.f. N. exigua (F.Muell.) Kuntze V. capitata Nees ex Lehm. 1858 N - NNN N subterminal hilum that are smooth or densely covered with tapering tubercles; seeds of species belonging to the "Villarsia" 1 clade have a terminal hilum, are smooth or sparsely tuberculate, and may have a nutritive caruncle, a feature that is entirely absent from the other clades. 1828 vN -- -N 1847 L - NLN - Box 2. Villarsia cambodiana Hance (syn. V. thombaidalis Dop), native to Southeast Asia, represents the only Villarsia species to be described from a tropical latitude. In April 2008, material identified as V. cambodiana was obtained from Lo Go - Xa Mat National Park (Tay Ninh Province, Vietnam). Preliminary DNA sequence analysis placed the specimen soundly within the Nymphoides clade, with N. aurantiaca (Dalzel) Kuntze as its nearest relative. Thanks to L. Averyanov, Nguyen Tien Hiep, Phan Ke Loc, P. Raven, J. Regalado, and Vu Ngoc Long. Photos: L. Aver- yanov, copyright Missouri Bota- nical Garden and Vietnam 1858 N - NN NN b 1845 v - v vNV V. congestifiora 1868 v- VVNV FMuell. V. exatata (Sol. ex Sims) G.Don 1807 M-- -NV V. lasiospema 1868 v - v vN V F.Muell. V. latifolla Benth. V. vioifola 1837 v - v v NV Botanical Conservation Program, funded by the Henry Luce Foundation; used with permission. 1868 v - vvN V F.Muell. V. capensis Houtt.) Merr. V. golablattiana Omduff V. manningiana Omduff V. ablara 1777 VV ---- v v Literature Cited Aston, H. I. 1969. The genus Warsia (Menyanthaceae) in Australa. Muelaria 2 3-63. Aston, H. I. 1973. Aquatio Plants of Australa. Melbourne University Press, Cariton, Victoria. Bentham, G. and F. von Mueler. 1859. Gentianaceae. Pp. 359382 in Flora Australionsls. Vol 4. L Reeve & Co., London. 1999 V- -- Figure 3. Habit variation in Vilarsia. Clockwise trom bottom left: V. submersa is obligately submersed, with floating leaves, a lax inflorescence, and flowers that touch the water; V. congestiflora has flowers borne in a congested cluster; V. pamassiolia represents the typical VWlarsia habit, with a basal rosette of leaves and an erect, expanded inflorescence. Photos: S. Jacobs. 2001 V. Chusng T. I, and R. Omduff, 1992. Seed morphology and systematics of Menyanthaceae. American Journal of Botany 79 1396-1406. Grisebach, A. H. R. 1845. Tribus I. Manyantheae. Pp. 136--141 in A P. de Candole, ed. Prodromus Systematis Noturalis Regni Vegetsbits. Vol. 9. Sumptibus Sociorum Treuttel et Würtz, Paris Mueller, F. J. H. von. 1868. Fragmenta Phytographise Australae. Vol. 6. Auctoritate Gubern. Colonise Victorise. Ex Officina Joannis Ferres, Melbourne. 1860 v - vv NV F.Muell. V. cathifolia F.Muell. V. parmessifolia Labil.) R.Br. V. reniformis 1868 v - VVNV Box 1. Liparophyllum gunnil has remained separate from other Menyanthaceae genera because of its many anomalous morphological features, including linear keaves, solitary flowers, and the absence of carpellary glands. It occupies high-altitude lakes in Tasmania and New Zealand and is the only Menyanthaceae species native to the latter. Photos: M. 1806 sv v vNV Mueller, F. J. H. van. 1875. Fragmenta Phytographiao Australao. Vol. 9. Auctoritate Gubern. Coloniae Victoriae, Ex Officina Joannis Ferres, Melbourne. Figure 1. Villarsia flowers, Clockwise from bottom left: V. manninglana photo: R. Prunier; all ather photos S. Jacobs), V. exaltata, V. lasiosperma, V. latifola, V. congestilora, V. capitata, V. abifora, V. parmassifolia, V. cahifolia, V. marchanti, V. submersa, V. reniformis. 1810 v - v v NV Tppery. N. P., D. H. Les, D. J. Padgett, and S. W. L. Jacobs. 2008. Generic circumscription in Meryanthacese: A phylogenetic evaluation. Systomatic Botany. In preas. R.Br. IV. submersa Acknowledgments 1969 V ---- V Aston V. umbricola Aston The authors are indebted to the directors and staff of the following herbaria for providing access to specimens and type material: A, BM, BRI, CAL, CONN, FI, GH, GOET, K, MEL, MO, NSW, NT, NY, PERTH, PRE, UC, US. We also thank Helen Aston for her valuable insights on the Australian taxa 1969 V- -- Wilcox. "Villarsia" 2 VIllarsia lymphoides "Villarsia" 2 Villarsia" Taxonomic revision of Villarsia (Menyanthaceae) in Australia: Molecular and morphological evidence Nicholas P. Tippery', Donald H. Les', Donald Padgett?, and Surrey W. L. Jacobs3 OF ST NE 1Department of Ecology and Evolutionary Biology, College of Liberal Arts and Sciences, University of Connecticut, Storrs, Connecticut, U.S.A.; 2Bridgewater State College, Bridgewater, Massachusetts, U.S.A.; ³National Herbarium of New South Wales, Sydney, New South Wales, Australia 1881 Figure 4. Seeds of VWarsla and related taxa. Composite of equal-magnification ight micrographs. Row 1: Nym- phoides anigua, V. violifolia, V. capitata, V. congestifiora. Row 2: V. latiola, V. exaltata, Liparophylum gunni, V. lasi- osperma. Row 3: V. capensis, V. marvingiana, Row 4: V. albifiora, V. pamassifolla, V. calthifolia, V. marchanti. Row 5: V. submersa, V. reniforms, V. umbricola var, umbricola, V. umbricola var. beaugle- holei. Discriminating seed features include vestiture Figure 2. Phylogenetic relationships in Menyan- thaceae, focusing on spe- cies currently circumscribed within Vilarsa and related taxa. The tree was con- Abstract 100 K Nymphoides (40-50 spp.) 1.0 The aquatic and wetland plant family Menyanthaceae consists of five genera that are distributed worldwide. Phylogenetic analyses using both molecular and morphological data consistently have resolved two monotypic genera with boreal distributions, Menyanthes and Nephvophylidlum, as a clade sister to the remaining genera. Two of the latter, Liparophyllum (1 sp.) and Vilarsia (18 spp.), are restricted to the southem hemisphere, and species of Nymphoides (40-50 spp.) are found worldwide. Analysis of combined molecular (nrITS, tmK/matk, rbcL) and morphological data resolved several relationships that were inconsistent with current generic circumscriptions under a criterion of monophyly, among which the most conspicuous is the paraphyly of VNarsia relative to Nympholdes. Circumscriptions of these genera have fluctuated historically, with species in Australia (where both Nymphoides and VWarsia are the most diverse) belonging entirely to one or the other genus according to different authors. Recent sampling of nearly all species of VMarsla and of the most morphologically diverse and broadly distributed species of Nymphoides revealed heretofore undescribed relationships, wherein VWasia resolved to three clades: two native to Australia and one, containing the type species V. capensis, comprising only South African taxa. Species of Nymphoides resolved to a single clade, with the exception of the anomalous species N. exigua. Revised taxonomic designations have been provided for species of Villarsia in Australia and for N. exigua, in order to reflect the phylogenetic relationships that were recovered in our study. Several distinguishing morphological characters were found to support these relationships, particularly aspects of seed shape and vestiture. 100 Nymphoides exigua 1.0 Villarsla violifolia structed using combined morphological and moleo- ular (nrITS, matK, rbcl, and tmk introns) data (analyses described in Tippery et al (2008), with the addition of two Vlarsia species (V. submersa arnd V. viofola). Topology represents the single most-parsimonious tree; node values indicate 98 100 1.0 1.0 100 Villarsia capitata 1.0 100 1.0 68 55 Villarsia congestiflora VIllarsla latifolia 100 1.0 Villarsia exaltata .07 Liparophyllum gunnil 63 parsimony bootstrap above and Bayesian posterior (tuberculate or smooth), tub- ercle density, hium location (terminal or subterminal, and the presence of a nutritive caruncle at the hilum. The top eight seeds represent the "VWarsia" 1 clade, the bottom Villarsia lasiosperma probability support below each node. The tree was rooted using sequences from Monyanthes and Nephrophylidum, prior analysis using out- group taxa from the related families Asteraceae, Caly- ceraceae, and Goodenia- ceae revealed them to be the sister clade to Liparo- phyllum, Nymphoides, and Vlarsia (Tippery et al. 2008). Taxa in the "Vilarsia" 1 and "Vilarsia" 2 clades are native to Australia, while V. capensis and V. man- ningiana are found only in South Africa. Asterisk in- dicates the type species for Vilarsia (V. capensis) Villarsia capensis 100 1.0 Villarsia manningiana 100 1.0 after VIllarsla albiflora 1.0 95 1.0 72 eight belong to the "VWarsia" 2 clade, and the two large seeds in the middle are from South Villarsia parnassifolia Table 1. Taxonomic history of the genus Villarsia. Prior to 1900, several Australian Menyanthaceae had been described variously under different genera. Abbreviations: L: Liparophyllum Hook.f.: M: Menyanthes L.; N: Nymphaides Ség. (syn. Limnanthemum S.G.Gmel.); S: Swertie L. (Gentiaraceae); V: Valarsia Vent. (syn. Renealmia Houtt.). Notes: 1. Mueller (1868) included Nymphoides as a section of Wlarsia. 2. Mueller (1875) transferred "all species of VWarsia to Wympholdes)", citing instances of other congeneric plant species among which differences were more pronounced translation and paraphrase by N.P.T.). Villarsia calthifolia African Warsia 82 species. 1.0 Villarsia marchantil 100 1.0 Villarsia submersa 100 1.0 Villarsia reniformis Discussion Villarsia umbricola N. aurantiaca 1850 N - NNN N Dalzel) Kuntze N. crenata (F.Muell.) Kurtze N. exiWlora Despite overall similarity among Vilarsia species, analysis of combined morphological and molecular data indicated that the genus as currently circumscribed comprises three clades (Fig. 2). In addition, two taxa traditionally placed outside the genus, Liparophylum gunni and Nymphaides exigua, were resolved within the same clade ("VWarsia" 1) along with several Varsia species. After more careful examination of the characters that have been used to OUTGROUPPMenyanthes (1 sp.) 1854 N - NN NN Nephrophyllidium (1 sp.) 1865 v - NNNN (F.Muell.) Kuntze N. geminata (RBr.) Kuntze N. indica LI Kuntze distinguish between Villarsia and Nymphoides (i.e., inflorescence architecture, capsule dehiscence), we hypothesized that similarities among Vilarsia taxa represent traits inherited from their shared ancestor (1.8., plesiomorphic traits) rather than features unique to a monophyletic lineage. Researchers previously have suggested that seed features are the most diagnostic characters for identifying species of Nymphoides and Villarsia (Aston 1969; Chuang and Ornduff 1992), and we determined that they are also informative for delineating phylogenetic groups. Species of the strict Vilarsia clade (including the type species V. capensis) have seeds that are orbicular and the largest of any surveyed; members of the "Villarsia" 2 clade have seeds with a 1810 VNNNNN 1753 MN-NNN N. minima (F.Muell.) Kuntze N. parvifolia Wall) Kuntze L. gunni Hook.f. N. exigua (F.Muell.) Kuntze V. capitata Nees ex Lehm. 1858 N - NNN N subterminal hilum that are smooth or densely covered with tapering tubercles; seeds of species belonging to the "Villarsia" 1 clade have a terminal hilum, are smooth or sparsely tuberculate, and may have a nutritive caruncle, a feature that is entirely absent from the other clades. 1828 vN -- -N 1847 L - NLN - Box 2. Villarsia cambodiana Hance (syn. V. thombaidalis Dop), native to Southeast Asia, represents the only Villarsia species to be described from a tropical latitude. In April 2008, material identified as V. cambodiana was obtained from Lo Go - Xa Mat National Park (Tay Ninh Province, Vietnam). Preliminary DNA sequence analysis placed the specimen soundly within the Nymphoides clade, with N. aurantiaca (Dalzel) Kuntze as its nearest relative. Thanks to L. Averyanov, Nguyen Tien Hiep, Phan Ke Loc, P. Raven, J. Regalado, and Vu Ngoc Long. Photos: L. Aver- yanov, copyright Missouri Bota- nical Garden and Vietnam 1858 N - NN NN b 1845 v - v vNV V. congestifiora 1868 v- VVNV FMuell. V. exatata (Sol. ex Sims) G.Don 1807 M-- -NV V. lasiospema 1868 v - v vN V F.Muell. V. latifolla Benth. V. vioifola 1837 v - v v NV Botanical Conservation Program, funded by the Henry Luce Foundation; used with permission. 1868 v - vvN V F.Muell. V. capensis Houtt.) Merr. V. golablattiana Omduff V. manningiana Omduff V. ablara 1777 VV ---- v v Literature Cited Aston, H. I. 1969. The genus Warsia (Menyanthaceae) in Australa. Muelaria 2 3-63. Aston, H. I. 1973. Aquatio Plants of Australa. Melbourne University Press, Cariton, Victoria. Bentham, G. and F. von Mueler. 1859. Gentianaceae. Pp. 359382 in Flora Australionsls. Vol 4. L Reeve & Co., London. 1999 V- -- Figure 3. Habit variation in Vilarsia. Clockwise trom bottom left: V. submersa is obligately submersed, with floating leaves, a lax inflorescence, and flowers that touch the water; V. congestiflora has flowers borne in a congested cluster; V. pamassiolia represents the typical VWlarsia habit, with a basal rosette of leaves and an erect, expanded inflorescence. Photos: S. Jacobs. 2001 V. Chusng T. I, and R. Omduff, 1992. Seed morphology and systematics of Menyanthaceae. American Journal of Botany 79 1396-1406. Grisebach, A. H. R. 1845. Tribus I. Manyantheae. Pp. 136--141 in A P. de Candole, ed. Prodromus Systematis Noturalis Regni Vegetsbits. Vol. 9. Sumptibus Sociorum Treuttel et Würtz, Paris Mueller, F. J. H. von. 1868. Fragmenta Phytographise Australae. Vol. 6. Auctoritate Gubern. Colonise Victorise. Ex Officina Joannis Ferres, Melbourne. 1860 v - vv NV F.Muell. V. cathifolia F.Muell. V. parmessifolia Labil.) R.Br. V. reniformis 1868 v - VVNV Box 1. Liparophyllum gunnil has remained separate from other Menyanthaceae genera because of its many anomalous morphological features, including linear keaves, solitary flowers, and the absence of carpellary glands. It occupies high-altitude lakes in Tasmania and New Zealand and is the only Menyanthaceae species native to the latter. Photos: M. 1806 sv v vNV Mueller, F. J. H. van. 1875. Fragmenta Phytographiao Australao. Vol. 9. Auctoritate Gubern. Coloniae Victoriae, Ex Officina Joannis Ferres, Melbourne. Figure 1. Villarsia flowers, Clockwise from bottom left: V. manninglana photo: R. Prunier; all ather photos S. Jacobs), V. exaltata, V. lasiosperma, V. latifola, V. congestilora, V. capitata, V. abifora, V. parmassifolia, V. cahifolia, V. marchanti, V. submersa, V. reniformis. 1810 v - v v NV Tppery. N. P., D. H. Les, D. J. Padgett, and S. W. L. Jacobs. 2008. Generic circumscription in Meryanthacese: A phylogenetic evaluation. Systomatic Botany. In preas. R.Br. IV. submersa Acknowledgments 1969 V ---- V Aston V. umbricola Aston The authors are indebted to the directors and staff of the following herbaria for providing access to specimens and type material: A, BM, BRI, CAL, CONN, FI, GH, GOET, K, MEL, MO, NSW, NT, NY, PERTH, PRE, UC, US. We also thank Helen Aston for her valuable insights on the Australian taxa 1969 V- -- Wilcox. "Villarsia" 2 VIllarsia lymphoides "Villarsia" 2 Villarsia"

Villarsia in Australia

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This Infographic displays taxonomic revision of Villarsia (Menyanthacea) in Australia: Molecular and morphological evidence.

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