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Angiosperm Phylogeny Flowering Plant Systematics woody, vessels lacking; dioecious; flw T5–8, A∞, G5–8; 1 ovule/carpel; embryo sac 9-nucleate; 1 species (New Caledonia) MBORELLALES Amborellaceae A G A N R aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A3–∞; embryo sac 4-nucleate seeds operculate, perisperm; mucilage; alkaloids (no benzylisoquinolines) NYMPHAEALES Cabombaceae Hydatellaceae Nymphaeaceae I A D woody, vessels solitary; flw T>10, ∞A , G ca.9; embryo sac 4-nucleate tiglic acid, aromatic terpenoids Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae T E AUSTROBAILEYALES A lvs opposite, interpetiolar stipules; nodes swollen E flw small T0–3, A1–5, G1, 1 apical ovule/carpel CHLORANTHALES Chloranthaceae A woody; pollen uniporate R aromatic terpenoids CANELLALES Canellaceae Winteraceae L nodes trilacunar ± herbaceous; lvs two-ranked, leaf base sheathing Y single adaxial prophyll; swollen nodes IPERALES Aristolochiaceae Hydnoraceae Piperaceae Saururaceae P sesquiterpenes woody; lvs opposite; flw with hypanthium, staminodes frequent Calycanthaceae Hernandiaceae Monimiaceae (pellucid dots) (pellucid ethereal oils in spherical idioblasts idioblasts spherical in oils ethereal A often valvate anthers; carpels with 1 ovule; embryo large LAURALES Gomortegaceae Lauraceae Siparunaceae AGNOLIIDS N M woody; pith septate; lvs two-ranked; ovules with obturator Annonaceae Eupomatiaceae Magnoliaceae G features as in endosperm ruminate MAGNOLIALES “Early Angiosperms” Degeneriaceae Himantandraceae Myristicaceae I infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules O ovules atropous, with epidermal perisperm and copious endosperm; idioblasts with ethereal oils ACORALES Acoraceae Alismataceae (incl. Limnocharitaceae) Juncaginaceae Ruppiaceae S mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green LISMATALES Aponogetonaceae Butomaceae Posidoniaceae Scheuchzeriaceae P A Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae embryo sac 8-nucleate sac embryo E alkaloids; benzylisoquinoline stem with ring of bundles Ca oxalate fr a follicle; East Asia Petrosaviaceae R MONOCOTS raphides PETROSAVIALES endosperm M atactostele (scattered bundles) often twining vines; lvs often reticulate nuclear Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae no secondary thickening helobial ovary often inferior, style short, branched; steroid sapogenins/alkaloids DIOSCOREALES S mostly herbaceous nectaries when present pollen monosulcate septal some woody (with terminally tufted leaves) woody; vessels absent sieve tube plastids with flw spadix ANDANALES Cyclanthaceae Pandanaceae Velloziaceae eustele; nodes protein crystals P sieve tube plastids sympodial branching often geophytes (bulbs, tubers, rhizomes) with starch grains vascular bundles in stem scattered flw tepals sometimes spotted, nectaries at tepals; many seeds, seeds coat (testa) cellular Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae lvs simple, persistent, entire lvs parallel-veined, entire phytomelan lacking; fructans in stems, chelidonic acid, steroid saponins LILIALES Colchicaceae Liliaceae Petermanniaceae Smilacaceae flw parts free, strobilar, no glandular teeth perfect, P parts varying, flw pentacyclic, P 3-merous, A opp. P often geophytes; anomalous sec. growth often in threes, filaments narrow capsule or berry Amaryllidaceae (incl. Agapanthaceae, Alliaceae) Hypoxidaceae Iridaceae weakly differentiated, anthers broadly attached, seed coat obliterated or with phytomelan (incl. Agavaceae, Hyacinthaceae, Ruscaceae) stamen with broad filaments septal nectary; single cotyledon ASPARAGALES Asparagaceae Lanariaceae Orchidaceae pollen monosulcate adventitious roots Tecophilaeaceae Xanthorrhoeaceae (incl. Asphodelaceae, Hemerocallidaceae) G apocarpous (style short in most) monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate nectaries absent intense primary growth, large apical meristem embryo very small infl with spathe;alkaloids Arecaceae aromatic terpenoids ARECALES mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae lvs grassy; flw often anemophilous, minute, chaffy, without nectaries POALES Cyperaceae Juncaceae Rapateaceae Typhaceae (incl. Sparganiaceae) flw often irregular; few fertile stamens infl thyrsus of scorpioid cymes COMMELINIDS phenylphenalenones Commelinaceae Haemodoraceae Pontederiaceae COMMELINALES UV-fluorescing cell walls rhizomatous, large-leafed herbs; pseudostem: central infl (ferulic/coumaric acids) flw irregular/monosymmetric, septal nectaries silicic acid in leaves G inferior, A often strongly modified/reduced Cannaceae Heliconiaceae Marantaceae Strelitziaceae cuticular waxes often in rodlets seeds arillate; silicic acid INGIBERALES aggregated into scallops Z Costaceae Lowiaceae Musaceae Zingiberaceae aquatic, herbaceous; lvs whorled, no pellucid dots; vessels lacking; monoecious flw T9–10, A∞, G1, 1 apical ovule/carpel, pollen inaperturate, pollen tube branched; hydrophilous CERATOPHYLLALES Ceratophyllaceae lvs often divided; flw whorled, P single or multiple whorls Berberidaceae Eupteleaceae Menispermaceae G apocarpous/paracarpous, superior; berberines Ranunculaceae RANUNCULALES Circaeasteraceae Lardizabalaceae Papaveraceae woody; flw 4–5-merous, nectary: disk, A epipetalous, G syncarpous pollen colporate; endosperm helobial; triterpenoids Sabiaceae SABIALES mostly woody; flw tepals often 4-merous A epitepalous, connectives sometimes with apical appendage PROTEALES Nelumbonaceae Platanaceae Proteaceae axial woody; vessels lacking; idioblasts in cortex; flw tepals missing, A∞ E nectary G>5 laterally connate with abaxial nectaries; fr aggregate of follicles U common TROCHODENDRALES Trochodendraceae D mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic flw tepals ± uniform or missing;pregnan pseudoalkaloids I BUXALES Buxaceae Didymelaceae C unisexual, dioecious; lvs toothed, sec. veins palmate flw tepals small to lacking O ellagic acid UNNERALES Gunneraceae Myrothamnaceae T G mostly woody; lvs veins proceed to apex & teeth; flw K5–∞, persisting S mostly A∞, G mostly slightly connate seeds often with aril; fr often both ventrally/dorsally dehiscent Dilleniaceae pollen tricolpate DILLENIALES flw K/C/P opp A lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent benzylisoquinolines absent benzylisoquinolines filaments rather narrow fr mostly dry, dehiscent Altingiaceae Daphniphyllaceae Hamamelidaceae anthers basifixed myricetin, flavonols Paeoniaceae nodes trilacunar SAXIFRAGALES Cercidiphyllaceae Grossulariaceae stomata anomocytic often tendrillar vines; lvs often divided and with glandular teeth Crassulaceae Haloragaceae Saxifragaceae ethereal oils absent A epipetalous, 2 ovules per carpel; raphides, pearl glands berries stipules VITALES Vitaceae nodes cork origin deep-seated 3:3 endosperm lacking resinous, lignans/neolignans, harman alkaloids ZYGOPHYLLALES Krameriaceae Zygophyllaceae R flw small, G often 3-merous, nectary: intrastaminal disk seeds often with aril (red-orange) O Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae) Lepidobotryaceae S infl cymose CELASTRALES I lvs often compound, pulvini (sleep movement) ellagic acid tannins Brunelliaceae Connaraceae Elaeocarpaceae D flw A5 or multiple, branched style common Oxalidaceae S mucilage cells; oxalates OXALIDALES Cephalotaceae Cunoniaceae Huaceae F Achariaceae Euphorbiaceae Rafflesiaceae Ochnaceae Podostemaceae lvs margins toothed Chrysobalanaceae Hypericaceae Passifloraceae Rhizophoraceae A flw G often tricarpellate MALPIGHIALES B Clusiaceae Linaceae Phyllanthaceae Salicaceae flw often “papilionaceous”: wing, standard, keel, mostly G1 Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae I mostly A10; fr a pod; symbiosis with root nodule bacteria D diverse alkaloids, NP amino acids, lectins (in Fabaceae) FABALES Fabaceae Polygalaceae Quillajaceae Surianaceae C E S U lvs mostly simple with stipules Barbeyaceae Elaeagnaceae Rosaceae O N fix flw K valvate (and hypanthium) persisting R D carpels with 1 ovule, stigma dry; dihydroflavonols ROSALES Cannabaceae Moraceae Ulmaceae E I embryo large Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae) C endosperm scanty lvs mostly alternate flw often unisexual, G mostly inferior Anisophyllaceae Coriariaceae Cucurbitaceae O parietal placentation; cucurbitacins Tetramelaceae CUCURBITALES Begoniaceae Corynocarpaceae Datiscaceae T lvs undivided; flw small, unisexual S anemophilous, thus T reduced or lacking, G mostly inferior infl spikes or catkins; fr 1-seeded, mostly nuts Betulaceae Fagaceae Myricaceae Rhoipteleaceae flw pentacyclic ectomycorrhiza; tannins, dihydroflavonols FAGALES Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae parts alternating lvs with glandular teeth G connate flw A obdiplostemonous, nectary on filament K + C (free) Francoaceae Geraniaceae Ledocarpaceae Melianthaceae (A polyandrous) stems jointed at nodes; ethereal oils, ellagic acid GERANIALES pollen tricolporate lvs opposite, colleters (glandular hair on adaxial surface of petiole base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae) trihydroxyl-flavonoids stipules small (if any); cork deep seated flw K valvate, persisting, A incurved in bud, ovary inferior; ovules many Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae) endosperm scanty; scaly bark; flavonols, myricetin
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    ANGIOSPERM PHYLOGENY GROUP (APG) SYSTEM The Angiosperm Phylogeny Group, or APG, refers to an informal international group of systematic botanists who came together to try to establish a consensus view of the taxonomy of flowering plants (angiosperms) that would reflect new knowledge about their relationships based upon phylogenetic studies. As of 2010, three incremental versions of a classification system have resulted from this collaboration (published in 1998, 2003 and 2009). An important motivation for the group was what they viewed as deficiencies in prior angiosperm classifications, which were not based on monophyletic groups (i.e. groups consisting of all the descendants of a common ancestor). APG publications are increasingly influential, with a number of major herbaria changing the arrangement of their collections to match the latest APG system. Angiosperm classification and the APG Until detailed genetic evidence became available, the classification of flowering plants (also known as angiosperms, Angiospermae, Anthophyta or Magnoliophyta) was based on their morphology (particularly that of the flower) and their biochemistry (what kinds of chemical compound they contained or produced). Classification systems were typically produced by an individual botanist or by a small group. The result was a large number of such systems (see List of systems of plant taxonomy). Different systems and their updates tended to be favoured in different countries; e.g. the Engler system in continental Europe; the Bentham & Hooker system in Britain (particularly influential because it was used by Kew); the Takhtajan system in the former Soviet Union and countries within its sphere of influence; and the Cronquist system in the United States.
  • Functional Integration of Floral Plant Traits: Shape and Symmetry, Optical Signal, Reward and Reproduction in the Angiosperm Flower

    Functional Integration of Floral Plant Traits: Shape and Symmetry, Optical Signal, Reward and Reproduction in the Angiosperm Flower

    Functional Integration of Floral Plant Traits: Shape and Symmetry, Optical Signal, Reward and Reproduction in the Angiosperm Flower Dissertation zur Erlangung des Doktorgrades (Dr. rer. nat.) der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn vorgelegt von Andreas Wilhelm Mues aus Kirchhellen Bonn, den 20. Januar 2020 1 2 Angefertigt mit Genehmigung der Mathematisch-Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn Erstgutachter: Prof. Dr. Maximilian Weigend, Universität Bonn Zweitgutachter: Prof. Dr. Eberhard Fischer, Universität Koblenz Tag der Promotion: 30. April 2020 Erscheinungsjahr: 2020 3 4 Acknowledgements I thank Prof. Dr. Maximilian Weigend, supervisor, for his guidance and support, and for giving me the opportunity to study the holistic subject of floral functional integration and plant-animal interaction. I am grateful for the experience and for the research agendas he entrusted to me: Working with the extensive Living Collections of Bonn Botanical Gardens was an honour, and I have learned a lot. I thank Prof. Dr. Eberhard Fisher, for agreeing to be my second supervisor, his advice and our shared passion for the plant world. I would like to thank many people of the Nees Institute and Bonn Botanical Gardens who contributed to this work and who gave me good memories of my years of study: I thank Lisabeth Hoff, Tianjun Liu, Luisa Sophie Nicolin and Simon Brauwers for their contribution in collecting shares of the raw data together with me, and for being eager students – especially counting pollen and ovule numbers and measuring nectar reward was a test of patience sometimes, and we have counted and measured a lot … Thank you! Special thanks go to Gardeners of the Bonn Botanical Gardens, for their constant support throughout the years, their love for the plant world in general and their commitment and care for the Living Collection: Klaus Mahlberg (Streptocarpus), Birgit Emde (carnivorous plants), Klaus Bahr (Geraniales), Bernd Reinken and Klaus Michael Neumann.