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El color floral, junto a otros rasgos florales, un carácter ligado al éxito reproductivo, puesto que participa en la atracción de los polinizadores. Diferentes presiones selectivas actúan sobre este rasgo, ya sean de carácter biótico como los polinizadores, o de carácter abiótico, como el estrés ambiental, llegando a promover la especiación. La gran diversidad de colores florales en la naturaleza se debe principalmente a la presencia de pigmentos. Las antocianinas son los pigmentos predominantes en las flores y se sintetizan en la ruta de biosíntesis de las antocianinas (ABP). Variaciones en los genes estructurales y reguladores que participan en esta ruta son clave para redirigir el flujo hacia la síntesis de diferentes tipos de antocianinas. En la naturaleza existen especies con polimorfismo de color floral, es decir, que presentan más de dos morfotipos diferentes. Dado que cada morfotipo podría tener diferentes presiones selectivas, los polimorfismos son una fuente de especiación. Es por ello que las especies con polimorfismo de color floral son un buen sistema para estudiar el papel del color floral en procesos de especiación. En esta tesis doctoral abordamos el estudio del polimorfismo del color floral que comparten dos especies hermanas, Lysimachia monelli y Lysimachia arvensis, con flores de morfotipo azul y morfotipo naranja, desde una perspectiva multidisciplinar. Ahondamos en la repercusión que tiene el polimorfismo en el aislamiento reproductivo y la biogeografía de L. monelli (aspectos ya estudiados anteriormente en L. arvensis), y exploramos los pigmentos principales y los genes involucrados en su síntesis en ambas especies a través del análisis del transcriptoma de los pétalos. Los resultados muestran que los linajes de color de ambas especies se encuentran en un estado avanzado de especiación. L. monelli presenta un aislamiento geográfico total, y ante un posible escenario de contacto entre ambos linajes de color, solamente las barreras postcigóticas, Flower color, along with other floral traits, is a character linked to the reproductive success, since it participates in pollinator attraction. Different selective pressures act on this trait, from biotic factors such as pollinators, to abiotic factors such as environmental stress, promoting speciation. The great diversity of flower colors in nature is mainly due to the presence of pigments. Anthocyanins are the predominant pigments in flowers and are synthesized in the anthocyanin biosynthetic pathway (ABP). Variations in the structural and regulatory genes that participate in this pathway are key to redirect the flux towards the synthesis of different types of anthocyanins. In nature there are species with floral color polymorphism, that is, they have more than two different morphotypes. Since each morphotype could have different selective pressures, polymorphisms are a source of speciation. Then, species with floral color polymorphism are good systems to study the role of flower color in speciation processes. In this PhD dissertation, we address the study of flower color polymorphism shared by two closely related species, Lysimachia monelli and Lysimachia arvensis, with blue and orange flowers colors, from a multiscale perspective. We deepen into the repercussion that polymorphism has on the reproductive isolation and biogeography of L. monelli (aspects already studied previously in L. arvensis), and we explore the main pigments and the genes involved in their synthesis in both species through the analysis of the petals’ transcriptome. The results show that the color lineages of both species are in an advanced stage of speciation. Specifically, L. monelli presents a total geographic isolation, and under a possible secondary contact between both colored lineages, only the postzygotic barriers would suppose an important isolation between them (38%). Although there is evidence of gene flow in the past, the lineages of L. monelli currently present different niches

Genetic divergence between species depends on reproductive isolation (RI) due to traits that reduce interspecific mating (prezygotic isolation) or are due to reduced hybrid fitness (postzygotic isolation). Previous research found that prezygotic barriers tend to be stronger than postzygotic barriers, but most studies are based on the evaluation of F1 hybrid fitness in early life cycle stages. We combined field and experimental data to determine the strength of 17 prezygotic and postzygotic reproductive barriers between two Lysimachia species that often co-occur and share pollinators. We assessed postzygotic barriers up to F2 hybrids and backcrosses. The two species showed near complete RI due to the cumulative effect of multiple barriers, with an uneven and asymmetric contribution to isolation. In allopatry, prezygotic barriers contributed more to reduce gene flow than postzygotic barriers, but their contributions were more similar in sympatry. The strength of postzygotic RI was up to three times lower for F1 progeny than for F2 or backcrossed progenies, and RI was only complete when late F1 stages and either F2 or backcrosses were accounted for. Our results thus suggest that the relative strength of postzygotic RI may be underestimated when its effects on late stages of the life cycle are disregarded.

Flower colour has played a decisive role as an evolutionary force in many groups of plants by driving speciation. A well-known example of colour polymorphism is found across the Mediterranean populations of Lysimachia arvensis and L. monelli, in which blue- and red-flowered plants can be found. Previous studies recognized two lineages within L. arvensis differing in flower colour, but this variation has not yet been considered in a phylogenetic context. We have reconstructed the ancestral states of flower colour across Mediterranean Lysimachia spp. aiming at understanding its phylogenetic signal using the nuclear internal transcribed spacer (ITS) sequences and three plastid markers. All blue- and red-flowered specimens were nested in different clades in the ITS tree, thus supporting that L. arvensis and L. monelli are polyphyletic, whereas low phylogenetic resolution was found in plastid markers. Monophyly was reconstructed for blue-flowered L. arvensis and L. monelli samples, and similarly for red-flowered individuals of each species: (1) blue-flowered L. arvensis was reconstructed as sister to the strictly blue-flowered L. talaverae in a monophyletic clade sister to remaining Lysimachia; (2) red-flowered L. arvensis was resolved as sister to red-flowered L. monelli in a monophyletic clade; and (3) clade 2 was sister to blue-flowered L. monelli and the strictly blue-flowered L. foemina. Our results suggest that colour lineages in L. arvensis and L. monelli constitute different species, but flower colour did not promote the separation of these lineages. We propose a new name for blue-flowered L. arvensis (L. loeflingii) and a new combination for red-flowered L. monelli (L. collina), maintaining L. arvensis for red-flowered plants and L. monelli for blue-flowered plants.

2020 Chartarrayawadee et al., published by De Gruyter 2020. The Lysimachia foenumgraecum Hance extract (LHE) was used for silver nanoparticles (AgNPs) synthesis. In this study, the herbal plant of Lysimachia foenumgraecum Hance (LH) was extracted with deionized water and we are the first to successfully use LHE as reducing and stabilizing agents for the green synthesis of AgNPs. The concentration of LHE used in this study was in the range of 0.003 to 1.0 wt%. Aqueous colloidal solutions of AgNPs reduced and stabilized by LHE show long-term stability due to the steric stabilization effect. This can be confirmed by zeta potential measurements which afforded values approximately of 0 mV, indicating the steric stability of AgNPs colloidal solutions synthesized by LHE. Furthermore, the obtained AgNPs colloidal solutions show superior antibacterial effect to gram-positive bacteria (Staphylococcus aureus) comparing to Chloramphenicol (positive control). AgNPs with LHE 0.003 wt% affords the highest antibacterial effect to S. aureus showing an inhibition zone diameter of 19.08 ± 0.67 mm; which is superior to Chloramphenicol.

Background and Aims: Herkogamy, or anther-stigma separation, is known to reduce self-pollen deposition, but little is known about the relative efficacy of different modes or conformations of herkogamy. We assessed the effectiveness of vertical versus lateral herkogamy in preventing or promoting self-pollen deposition in the annual herb Lysimachia arvensis, a plant with lineages that differ in flower colour, and in which flowers first display lateral and then vertical herkogamy. Because mating between the two lineages compromises fitness through the production of low-quality hybrid offspring, we tested the prediction that individuals sampled from sites occupied by both lineages should have flowers that promote autonomous self-pollen deposition and self-fertilization as a result of selection to reduce deleterious reproductive interference. Methods: We characterized variation in herkogamy within and among 25 pure and mixed populations of L. arvensis in its European range and assessed the effectiveness of lateral versus vertical herkogamy in avoiding self-pollen deposition. Results: Lateral herkogamy was more effective than vertical herkogamy in limiting self-pollen deposition. In the case of vertical herkogamy, only approach herkogamy was effective. Lineages showed consistent differences in herkogamy traits. In general, angles were smaller for blue than red flowers in most populations, and blue flowers showed approach herkogamy, while red flowers showed predominantly reverse herkogamy. In sympatry, the red lineage showed a reduction of both herkogamy traits while for the blue lineage only lateral herkogamy was reduced. Conclusions: Our results demonstrate that pollen deposition is affected not only by the degree but also the spatial conformation of herkogamy. They also highlight reduced herkogamy as a potential mechanism for promoting reproductive assurance under pollen limitation, as well as for avoiding reproductive interference between genetically divergent lineages

Anthocyanins are the primary pigments contributing to the variety of flower colors among angiosperms and are considered essential for survival and reproduction. Anthocyanins are members of the flavonoids, a broader class of secondary metabolites, of which there are numerous structural genes and regulators thereof. In western European populations of Lysimachia arvensis, there are blue- and orange-petaled individuals. The proportion of blue-flowered plants increases with temperature and daylength yet decreases with precipitation. Here, we performed a transcriptome analysis to characterize the coding sequences of a large group of flavonoid biosynthetic genes, examine their expression and compare our results to flavonoid biochemical analysis for blue and orange petals. Among a set of 140 structural and regulatory genes broadly representing the flavonoid biosynthetic pathway, we found 39 genes with significant differential expression including some that have previously been reported to be involved in similar flower color transitions. In particular, F3′5′H and DFR, two genes at a critical branchpoint in the ABP for determining flower color, showed differential expression. The expression results were complemented by careful examination of the SNPs that differentiate the two color types for these two critical genes. The decreased expression of F3′5′H in orange petals and differential expression of two distinct copies of DFR, which also exhibit amino acid changes in the color-determining substrate specificity region, strongly correlate with the blue to orange transition. Our biochemical analysis was consistent with the transcriptome data indicating that the shift from blue to orange petals is caused by a change from primarily malvidin to largely pelargonidin forms of anthocyanins. Overall, we have identified several flavonoid biosynthetic pathway loci likely involved in the shift in flower color in L. arvensis and even more loci that may represent the complex network of gene

Anthocyanins are the primary pigments contributing to the variety of flower colors among angiosperms and are considered essential for survival and reproduction. Anthocyanins are members of the flavonoids, a broader class of secondary metabolites, of which there are numerous structural genes and regulators thereof. In western European populations of Lysimachia arvensis, there are blue- and orange-petaled individuals. The proportion of blue-flowered plants increases with temperature and daylength yet decreases with precipitation. Here, we performed a transcriptome analysis to characterize the coding sequences of a large group of flavonoid biosynthetic genes, examine their expression and compare our results to flavonoid biochemical analysis for blue and orange petals. Among a set of 140 structural and regulatory genes broadly representing the flavonoid biosynthetic pathway, we found 39 genes with significant differential expression including some that have previously been reported to be involved in similar flower color transitions. In particular, F3′5′H and DFR, two genes at a critical branchpoint in the ABP for determining flower color, showed differential expression. The expression results were complemented by careful examination of the SNPs that differentiate the two color types for these two critical genes. The decreased expression of F3′5′H in orange petals and differential expression of two distinct copies of DFR, which also exhibit amino acid changes in the color-determining substrate specificity region, strongly correlate with the blue to orange transition. Our biochemical analysis was consistent with the transcriptome data indicating that the shift from blue to orange petals is caused by a change from primarily malvidin to largely pelargonidin forms of anthocyanins. Overall, we have identified several flavonoid biosynthetic pathway loci likely involved in the shift in flower color in L. arvensis and even more loci that may represent the complex network of gene

Flower color polymorphism, an infrequent but phylogenetically widespread condition in plants, is captivating because it can only be maintained under a few selective regimes but also because it can drive intra-morph assortative mating and promote speciation. Lysimachia arvensis is a polymorphic species with red or blue flowered morphs. In polymorphic populations, which are mostly Mediterranean, pollinators prefer blue-flowered plants to the red ones, and abiotic factors also favors blue-flowered plants. We hypothesize that the red morph is maintained in Mediterranean areas due to its selfing capacity. We assessed inbreeding depression in both color morphs in two Mediterranean populations and genetic diversity was studied via SSR microsatellites in 20 natural populations. Results showed that only 44–47% of selfed progeny of the red plants reached reproduction while about 72–91% of blue morph progeny did it. Between-morph genetic differentiation was high and the red morph had a lower genetic diversity and a higher inbreeding coefficient, mainly in the Mediterranean. Results suggest that selfing maintaining the red morph in Mediterranean areas despite its inbreeding depression. In addition, genetic differentiation between morphs suggests a low gene flow between them, suggesting reproductive isolation.

Floral colour determines pollinator behaviour, strongly affecting plant-mating systems. Lysimachia arvensis has blue- and red-flowered plants and colour inheritance remains largely unknown. A control of floral colour based on one locus, with the red allele as dominant, has been proposed. This proposal cannot explain the sporadic appearance of other floral colours in wild populations. We studied floral colour segregation in L. arvensis and assessed the possibility that pollinators can visually distinguish colour morphs by using Chittka’s hexagon model, sigmoidal model of bee discrimination and experimental studies on pollinator attendance for two years. Hand crossing between morphs originated a homogeneous F1 with salmon-coloured flowers. In the F2, blue, red, salmon morphs and other plants with intermediate colours appeared, suggesting that more than one single locus are involved in colour segregation. Results from the sigmoidal discrimination model suggest that blue, red and salmon flowers can be discriminated by pollinators. In fact, pollinators showed strong colour constancy and discriminated against the salmon morph. Our study shows that “Flower colour” is a natural marker to assess the rate of crossing between morphs. The extreme rarity of salmon flowers in wild populations and flower constancy of L. arvensis pollinators indicates assortative mating.

Uno de los caracteres que promueven la selección divergente es el polimorfismo del color que se define como la presencia en una especie de dos o más morfos de color, genéticamente determinados e interfértiles en las mismas poblaciones. En plantas, el polimorfismo del color floral es un fenómeno ampliamente distribuido, pero poco frecuente, que se debe principalmente a la presencia de antocianinas. El polimorfismo del color ha sido definido como un “carácter mágico”, es decir, un carácter codificado por genes que están sujetos a selección divergente y que afectan pleiotrópicamente al aislamiento reproductivo, por lo que puede desencadenar un proceso de especiación. Aunque para que el proceso de especiación se complete es necesario el desarrollo de un considerable número de barreras pre- y postcigóticas entre las especies incipientes. A este respecto, hay que señalar que el color floral puede no ser el carácter que desencadene un proceso de especiación, pero puede ser fundamental como mecanismo de “refuerzo” creando barreras de aislamiento entre esas especies. El objetivo principal de esta tesis doctoral es conocer si el polimorfismo de color floral puede ser el desencadenante de un proceso de especiación en Lysimachia arvensis (L.) U. Manns & Anderb, una especie anual con plantas con flores azules o rojas. Para llevar a cabo este objetivo hemos estudiado diversos aspectos ecológicos, reproductivos y moleculares de esta especie, como la segregación del color floral, el papel de los polinizadores como agentes de selección, la fenología de los morfos de color, su sistema de cruzamiento, la depresión por endogamia, la diversidad genética de las poblaciones y la filogenia del grupo de especies en las que se incluye L. arvensis. Hemos encontrado que tras el cruce entre morfos de color se origina una F1 homogénea con flores salmón y que aparece con muy poca frecuencia en las poblaciones naturales. Los espectros de reflectancia muestran que el morfo azul es el que contrasta, Colour polymorphism is defined as the presence of two or more colour morphs, genetically determined and interfertile, within populations of a species. In plants, flower colour polymorphism is a widespread phenomenom but relatively infrequent and depends mainly of the presence of anthocyanins. Colour polymorphism has been defined as a “magic trait”, that is, a character encoded by genes that are subject to divergent selection that affects pleiotropically reproductive isolation, acting as a trigger for the speciation process. An essential component of speciation is the evolution of reproductive isolation. In plants, reproductive isolation depends on the existence of a considerable number of pre- and post-zygotic barriers between the incipient species. However, sometime flower colour does not initiate the speciation process but it can strongly contribute to species separation by acting as a reinforcement mechanism. The main objective of this doctoral thesis is to know if the floral colour polymorphism can be unleashing of speciation process in Lysimachia arvensis (L.) U. Manns & Anderb., an annual forb with red or blue flowers. To do this, we have carried out ecological, reproductive and molecular studies of L. arvensis such as floral colour segregation, plant phenology, breeding system and endogamy depression, genetic diversity of populations and the phylogeny of the L. arvensis group. We have found that inter-morph cross originates a homogeneous F1 with salmon-colour flowers, very infrequent in natural populations. The reflectance spectra of these morphs show that blue flowers markedly contrast with the background of the leaves. Bees, the main pollinators of this species, hardly distinguish between salmon and red flowers. In addition, flowers have two consecutive kinds of hercogamy, lateral and vertical, a very rare phenomenon in flowering plants. Both types of hercogamy have a high degree of heritability, and differ between morphs. The red morph showed a marked late

Herkogamy, spatial separation between stigma and anthers within a flower, is important in regulating plant-mating systems. We studied phenotypic variation and heritability of herkogamy traits in Lysimachia arvensis (=Anagallis arvensis) that show both lateral and vertical herkogamy in the same flower, a rare strategy in flowering plants. Both lateral and vertical herkogamy showed continuous variation in 15 natural populations. Lateral herkogamy, measured as the angle between style and stamens, ranged from 5.6 to 66.5°; vertical herkogamy ranged from reverse to approach herkogamy. Herkogamy traits were constant within plants but variable among plants and populations. Flowers with marked lateral herkogamy showed mainly reverse herkogamy, whereas flowers with low lateral herkogamy showed mainly approach herkogamy. Both herkogamy traits showed a high degree of narrow sense heritability (h2 = 0.843 for lateral and h2 = 0.635 for vertical herkogamy). We discuss the possibility that variation in both herkogamy traits among populations of L. arvensis is a consequence of differential selective pressures under different pollination environments.

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1561531%5DMICH-V-1561531, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1561531/MICH-V-1561531/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1561394%5DMICH-V-1561394, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1561394/MICH-V-1561394/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1561664%5DMICH-V-1561664, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1561664/MICH-V-1561664/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1561430%5DMICH-V-1561430, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1561430/MICH-V-1561430/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1657157%5DMICH-V-1657157, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1657157/MICH-V-1657157/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

UPLC-DAD method was developed and validated for the quantitative determination of free flavonol aglycones (kaempferol, quercetin and myricetin) after acidic hydrolysis in six Lysimachia species. Quantitative analyses showed that the amounts of various flavonol aglycones were significantly different in Lysimachia vulgaris, Lysimachia nummularia, Lysimachia punctata, Lysimachia christinae, Lysimachia ciliata and Lysimachia clethroides. The L. clethroides sample was found to be the richest in kaempferol (25.77 ± 1.29 μg/mg extract) and quercetin (97.67 ± 4.61 μg/mg extract), while the L. nummularia sample contained the highest amount of myricetin (20.79 ± 1.00 μg/mg extract). The antioxidant capacity of hydrolysed extracts was evaluated using in vitro DPPH• (2,2-diphenyl-1-picrylhydrazyl) and ABTS•+ [2,2′-azino-bis-(3-ethylbenzothiazoline-6-sulphonic acid)] decolourisation tests. The observed radical scavenging capacities of the extracts showed a relationship with the measured flavonol aglycone content and composition. The acidic treatment resulted in an increased free radical scavenging activity compared to the untreated methanol extract.

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Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1460136%5DMICH-V-1460136, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1460136/MICH-V-1460136/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1559484%5DMICH-V-1559484, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1559484/MICH-V-1559484/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1558676%5DMICH-V-1558676, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1558676/MICH-V-1558676/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1558598%5DMICH-V-1558598, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1558598/MICH-V-1558598/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1559416%5DMICH-V-1559416, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1559416/MICH-V-1559416/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1556083%5DMICH-V-1556083, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1556083/MICH-V-1556083/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Kenmerken, ecologie, voorkomen en trend van grote wederik in Noord-Holland.

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Lysimachia mauritiana Lam. (family Primulaceae), a small short-lived herb native to India, Indian and Pacific Ocean islands, and coastal east Asia, is described as a new naturalised record from the eastern suburbs of Sydney, New South Wales, Australia. It was first recorded in 1981 near Coogee, and grows in exposed rock crevices and seepages on the seacoast, very similar to its natural habitat overseas. Lysimachia mauritiana is known to have been cultivated in the area in 1961 in a home garden, which is the likely source of this introduction; it appears to be spreading locally as a weed.

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Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1552554%5DMICH-V-1552554, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1552554/MICH-V-1552554/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1496588%5DMICH-V-1496588, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1496588/MICH-V-1496588/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1514269%5DMICH-V-1514269, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1514269/MICH-V-1514269/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1552553%5DMICH-V-1552553, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1552553/MICH-V-1552553/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

We investigated seed morphology in 34 species of the genus Lysimachia and in 14 species and two subspecies of six additional genera (Anagallis, Ardisiandra, Asterolinon, Glaux, Pelletiera, Trientalis), which have been shown to be closely related to, or are placed within Lysimachia in previous molecular studies. We studied seed shape, seed coat structure, and seed coat surface patterns. Three major types of seed shape were identified: (1) sectoroid, (2) polyhedral, and (3) coarsely rugose with concave hilar area. In addition, seeds may be keeled or winged. The outer layer of the seed coat is either sponge-like and adhering only loosely to the inner seed coat or it is thin and tightly adhering to the underlying tissue. Seed surface patterns can be divided into six main types: (1) reticulate, (2) tuberculate, (3) vesiculose, (4) colliculate, (5) undulate, or (6) poroid-alveolate. Seed surface patterns are mostly congruent with molecular phylogenetic relationships. A reticulate surface pattern is diagnostic of, e.g. Lysimachia subgenera Palladia and Hawaiian Lysimachiopsis. Mapping seed characters onto a recent phylogenetic tree, reveals that they provide potentially synapomorphic character states for various subclades of Lysimachia. Salient examples include a rugose seed shape, which turns out to be synapomorphic for the clade comprising the genus Pelletiera plus Asterolinon linum-stellatum and a sponge-like outer seed coat layer, which characterizes a clade with Lysimachia vulgaris, L. thyrsiflora, and L. terrestris, with an analogue that apparently evolved in parallel in Trientalis europaea. We also discuss possible habitat factors that may have favored the independent evolution of particular seed types such as winged seeds in various lineages.

The 16 endemic Hawaiian species of Lysimachia form a monophyletic group referred to as subgenus Lysimachiopsis (Marr and Bohm 1997). The group has radiated into various habitats and most species are endemic to a single island. Previous taxonomic treatments have revealed problems, particularly within the L. hillebrandii/L. remyi complex. We conducted phylogenetic analyses including 12 species of the subgenus Lysimachiopsis using nuclear ribosomal DNA (ETS, ITS) and chloroplast DNA (rpl16, rpl20-rps12, rps16, trnH-psbA, trnS-G) sequences in order to reconstruct phylogenetic relationships. A combined analysis using all sequence data supports three major clades and provides new insights into the evolutionary history of the group, especially within the problematic L. hillebrandii/L. remyi complex. The resulting molecular phylogeny provides support for the following clades: (1) L. hillebrandii and L. waianaeensis, (2) L. remyi ssp. remyi, L. maxima, and L. remyi ssp. subherbacea, (3) L. remyi ssp. caliginis and L. remyi ssp. kipahuluensis, and (4) L. glutinosa, L. scopulensis, and L. kalalauensis. The phylogenetic pattern is largely congruent with the biogeographical distribution of the species in the Hawaiian Islands. Our results also indicate that earlier taxonomic treatments of the group need to be partially revised in order to reflect phylogenetic relationships.

The first half of the thesis deals with the seed morphology of members of the ANITA grade at the base of the angiosperm phylogeny (Amborella, Nymphaeales, Illiciales, Trimeniaceae, and Austrobaileyaceae), especially Illiciaceae and Schisandraceae. Seed characters support a major division between the New World and Old World species of Illicium. Fossil seeds previously assigned to Illiciaceae were re-evaluated. In Schisandraceae, seed and leaf epidermal characters were added to a previously published morphological data matrix. Phylogenetic analysis using the extended data matrix shows that Kadsura and Schisandra appear to be supported as monophyletic sister taxa by a number of synapomorphies in reproductive and vegetative characters. Fossil seeds from the Eocene of North America show some similarities to the modern Schisandra glabra from North America, whereas fossils from Europe are more similar to modern Asian species. In the second half of the thesis, seed morphology of Lysimachia and closely related taxa (Anagallis, Ardisiandra, Asterolinon, Glaux, Pelletiera, Trientalis) was investigated. The phylogenetic relationships among the endemic Hawaiian species of Lysimachia was also studied, using nuclear ribosomal DNA (ETS, ITS) and chloroplast DNA (rpl16, rpl20-rps12, rps16, trnH-psbA, trnS-G) sequence data. The seeds in Lysimachia and related taxa vary in, e.g., shape, seed coat structure and surface patterns. Seed surface patterns are mostly congruent with molecular phylogenetic relationships. A reticulate surface pattern is diagnostic for, e.g., the subgenus Palladia and the Hawaiian endemic subgenus Lysimachiopsis. Mapping seed characters onto a recent molecular-based phylogenetic tree, reveals that they provide potentially synapomorphic character states for various subclades of Lysimachia. The phylogenetic analysis based on the combined data set using nuclear ribosomal DNA and chloroplast DNA data provides new insights into the relationships within the Hawaiian s

New combinations are proposed for species of Anagallis, Pelletiera and Trientalis in accordance with the results of phylogenetic analyses of the Lysimachia complex, based on molecular and morphological data. These three genera as well as Glaux and Asterolinon, for the species of which names in Lysimachia are already available, have been found to be derived, specialized groups that have evolved within Lysimachia. The present classification therefore does not reflect our current understanding of evolutionary relationships within the Lysimachia complex. Merging all the genera in Lysimachia is here considered better than splitting the latter into several smaller genera. For Anagallis crassifolia and A. filifolia new names are validated and for A. alternifolia and A. pumila the names change since their epithets have already been used in Lysimachia. Lectotypes are selected for A. filifolia, A. filiformis, A. kingaënsis, A. monelli, A. schliebenii, A. serpens and Pelletiera verna, and a neotype is designated for A. foemina.

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1482545%5DMICH-V-1482545, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1482545/MICH-V-1482545/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1480850%5DMICH-V-1480850, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1480850/MICH-V-1480850/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

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Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1480635%5DMICH-V-1480635, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1480635/MICH-V-1480635/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

This thesis focuses on Anagallis, an herbaceous genus with a distribution mainly in (sub-) tropical Africa, Madagascar or South America. A few species are found in drier climate. Anagallis is considered close to Lysimachia, separated based on mode of capsule dehiscence. Monophyly of Anagallis was tested by phylogenetic analyses of nuclear and chloroplast DNA sequence data. Character evolution within Anagallis s.l., also including Asterolinon, Pelletiera and Lysimachia nemorum and L. serpyllifolia, was traced by combining morphological and molecular data. With additional chloroplast DNA data, the previously proposed close relationship between Anagallis arvensis and A. foemina was challenged, and the biogeography of “tropical Anagallis” investigated. Both parsimony and Bayesian inference analyses show Anagallis to be non-monophyletic and nested within Lysimachia. A few Anagallis species group together with two Lysimachia species, Asterolinon, and Pelletiera in one clade, whereas all other Anagallis species group in another clade. Morphological synapomorphies of the Anagallis arvensis–Pelletiera clade are few, while the second, strictly Anagallis, clade is recognized by several morphological synapomorphies. Molecular analyses show Anagallis foemina as closer related to A. monelli than to A. arvensis, and the view of A. foemina as a subspecies of A. arvensis should be abandoned. Biogeographic analyses suggest the ancestor of “tropical Anagallis”, i.e. the strictly Anagallis clade, to have had a European distribution. Dispersal to (sub-) tropical Africa has occurred once, while two parallel dispersals from African mainland to Madagascar are suggested. Dispersal from Europe to South America may be explained through long distance dispersal over water, or via land connections. As a concluding part of the thesis, Anagallis, Asterolinon, Pelletiera, and Glaux are merged with Lysimachia based on the results of analyses of molecular and morphological data.

OBJECTIVE: To study hypouricemic effect of aqueous extract of Lysimachia christinae on hyperuricemia in mice. METHOD: The uricase inhibitor potassium oxonate was used to induce hyperuricemia in mice, and serum uric acid level was determined with the phosphotungstic acid method. RESULT: The aqueous extract of Lysimachia christinae, when administered orally to the oxonate-induced hyperuricemic mice at the doses of 5.2, 10.4 and 20.8 g.kg-1, was able to elicit dose-dependent hypouricemic effects. At these doses of the extract, the serum urate levels of the oxonate-pretreated mice showed no difference from the normal mice. In normal mice, however, oral administration of the extract at the same doses did not produce any observable hypouricemic effects. CONCLUSION: The aqueous extract of Lysimachia christinae possesses potent hypuricemic effects on models of hyperuricemia in mice pretreated with oxonate.

Lysimachia minoricensis is an endemic species of the Balearic Islands that has become extinct in the wild, but persists in botanical gardens. Attempts of re-introducing the species into its natural habitat, which consisted in temporary dry streams, have failed. Low genetic variability has been reported for the garden individuals, suggesting that a reduced potential to adapt to environmental changes could be among the reasons for its extinction. In the present study, we particularly test whether photosynthesis and photoprotection responses of this species to water stress could help explaining the lack of success of this species in its natural habitat., Plants of L. minoricensis were grown in pots in a growth chamber. Soil water depletion was imposed over 20 days by stopping irrigation. Early stomata closure was observed in response to soil water depletion while leaves desiccated progressively. Although net photosynthesis was low in irrigated plants, due to a remarkably low mesophyll conductance to CO2, substantial photosynthetic activity was kept at severe drought, where leaf relative water content was as low as 50%, suggesting that L. minoricensis is a very drought-tolerant species. In parallel with decreased photosynthesis, thermal dissipation of the excess light and photorespiration progressively increased. The former was linearly related to increased de-epoxidation of the xanthophylls cycle. Photoprotection was effective, as pre-dawn maximum photochemical efficiency was maintained higher than 0.75 through the entire experiment. Moreover, photosynthetic capacity was largely (80%) recovered only 24 h after re-watering. These results show that stomatal regulation, photosynthetic metabolism and photoprotection in L. minoricensis are well adapted to water stress, suggesting that additional factors may be responsible for its status as a wild-extinct plant.

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1102640%5DMICH-V-1102640, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1102640/MICH-V-1102640/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1485821%5DMICH-V-1485821, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1485821/MICH-V-1485821/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy

Angiosperms, http://name.umdl.umich.edu/IC-HERB00IC-X-1102481%5DMICH-V-1102481, https://quod.lib.umich.edu/cgi/i/image/api/thumb/herb00ic/1102481/MICH-V-1102481/!250,250, The University of Michigan Library provides access to these materials for educational and research purposes. Some materials may be protected by copyright. If you decide to use any of these materials, you are responsible for making your own legal assessment and securing any necessary permission. If you have questions about the collection, please contact the Herbarium professional staff: herb-dlps-help@umich.edu. If you have concerns about the inclusion of an item in this collection, please contact Library Information Technology: libraryit-info@umich.edu., https://www.lib.umich.edu/about-us/policies/copyright-policy