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Flowering Plants. Eudicots The Families and Genera of Vascular Plants 8 Flowering Plants. Eudicots Asterales Bearbeitet von Joachim W. Kadereit, Charles Jeffrey 1. Auflage 2006. Buch. xi, 636 S. Hardcover ISBN 978 3 540 31050 1 Format (B x L): 20,3 x 27,6 cm Weitere Fachgebiete > Chemie, Biowissenschaften, Agrarwissenschaften > Biowissenschaften allgemein > Ökologie Zu Inhaltsverzeichnis schnell und portofrei erhältlich bei Die Online-Fachbuchhandlung beck-shop.de ist spezialisiert auf Fachbücher, insbesondere Recht, Steuern und Wirtschaft. Im Sortiment finden Sie alle Medien (Bücher, Zeitschriften, CDs, eBooks, etc.) aller Verlage. Ergänzt wird das Programm durch Services wie Neuerscheinungsdienst oder Zusammenstellungen von Büchern zu Sonderpreisen. Der Shop führt mehr als 8 Millionen Produkte. Asterales: Introduction and Conspectus J.W. Kadereit Asterales (incl. Campanulales of many authors), 1999; Cronquist 1981: Celastrales; Thorne 1992: with Alseuosmiaceae, Argophyllaceae, Composi- Theales; Takhtajan 1997: Icacinales) and Rous- tae (= Asteraceae), Calyceraceae, Campanulaceae seaceae (Lundberg 2001; Takhtajan 1997: Brexi- (incl. Cyphiaceae, Lobeliaceae, Nemacladaceae), ales), and partly also to Menyanthaceae (Downie Carpodetaceae (included in Rousseaceae by and Palmer 1992; Olmstead et al. 1992; Cronquist APG II 2003), Goodeniaceae, Menyanthaceae, 1981: Solanales; Thorne 1992: Campanulales; Pentaphragmataceae, Phellinaceae, Rousseaceae Takhtajan 1997: Menyanthales) and Stylidiaceae and Stylidiaceae (incl. Donatiaceae), contain about (Cronquist 1981: Campanulales; Thorne 1992: Sax- 26,300 species in c. 1,720 genera. The large major- ifragales; Takhtajan 1997: Stylidiales). Further sam- ity of species and genera belong to Compositae pling may identify other taxa from distant corners and Campanulaceae. The order is well supported of the traditional angiosperm system which should in all major molecular phylogenetic analyses be included in the order. On the other hand, Sphe- (APG II 2003), and is part of the Euasterids II or nocleaceae, as a family often associated with Aster- Campanulids sensu Bremer et al. (2002). ales/Campanulales (e.g. Lammers 1992), do not Phylogenetic structure within Campanulids belong here but rather in Solanales (APG II 2003). (also containing Apiales, Aquifoliales, Dipsacales Members of Asterales are mostly herbaceous and several families of uncertain ordinal place- and in most cases have alternate leaves without ment; APG II 2003) is not sufficiently well resolved stipules. Flowers are very rarely solitary but mostly to identify the sister group of Asterales. It appears aggregated in sometimes axillary but more com- to be evident, however, that of all representatives monly terminal inflorescences which are capitulate of the Campanulids, Aquifoliales are least closely and involucrate in most of the closely related Good- related to Asterales (Savolainen et al. 2000a, b; eniaceae, Calyceraceae and Compositae, and also Soltis et al. 2000; Albach et al. 2001; Bremer et al. in some Campanulaceae. The mostly zoophilous 2001, 2002). flowers typically are tetracyclic and pentamerous Although several of the constituent families but variation of organ number per whorl is of the order had been recognized to be closely known from several families. Flower symmetry is related to one another long ago (for discussion, see actinomorphic or zygomorphic with bilabiate or Lammers 1992), the recognition of the relationship unilabiate flowers – actinomorphic and zygomor- of others to Asterales (Lundberg and Bremer phic flowers are both found in the capitula of many 2003) is the result mainly (but not only) of recent Compositae – and resupination of flowers is known molecular phylogenetic work. This applies partic- from Campanulaceae-Lobelioideae and some Sty- ularly to Alseuosmiaceae (Backlund and Bremer lidiaceae. The sepals are commonly fused (not in 1997; Gustafsson and Bremer 1997; Kårehed Alseuosmiaceae and some Menyanthaceae), and et al. 1999; Cronquist 1981: Rosales; Thorne 1992: in Compositae the calyx commonly is replaced by Saxifragales; Takhtajan 1997: Hydrangeales), Argo- a pappus of variable structure assisting in fruit dis- phyllaceae (Kapil and Bhatnagar 1992; Gustafsson persal. Petals are free only in Carpodetaceae, Phelli- et al. 1996; Kårehed et al. 1999; Olmstead et al. naceae and some Argophyllaceae, Pentaphragmat- 2000; Cronquist 1981: Rosales; Takhtajan 1997: aceae and Stylidiaceae (Donatia). The androecium Hydrangeales), Carpodetaceae (Gustafsson and normally is isomerous with calyx and corolla, and Bremer 1997; Lundberg 2001; Takhtajan 1997: Hy- the stamens alternate with the petals. Reduction of drangeales), Phellinaceae (Backlund and Bremer stamen number is largely limited to Stylidiaceae. 1997; Gustafsson and Bremer 1997; Kårehed et al. Stamens can be inserted on the corolla or not, and 2 J.W. Kadereit anthers are mostly tetrasporangiate, basifixed and Following Lundberg and Bremer (2003), valvate commonly introrse. Pollen grains are mostly tri- corolla aestivation and the absence of apotracheal colporate, but both colpate or porate pollen grains wood parenchyma can be identified as synapo- withanincreasednumberofaperturesareknown. morphic. Both these characters, however, are not Carpodetus (Carpodetaceae) and Lechenaultia unique for the order and are variable within it. (Goodeniaceae) are unusual in having pollen Previously identified synapomorphies, such as sec- tetrads. The pluri- to unilocular ovary is commonly ondary pollen presentation (which is present in the inferior (or semi-inferior) but superior ovaries form of different mechanisms and is likely to have are found in some Carpodetaceae, some Goode- arisen more than once; see above) and the pres- niaceae, some Campanulaceae, and in Menyan- ence of inulin, are characteristic only of subgroups thaceae, Phellinaceae and Rousseaceae. Ovules of Asterales. usually are anatropous (hemi- to campylotropous Relationships within the order are clear and in Phellinaceae), unitegmic and tenuinucellate well supported in some parts but not in others and, where known, endosperm formation is mostly (Lundberg and Bremer 2003). One well-supported cellular, but nuclear in some Compositae. Fruits clade identified in several analyses (Chase et al. are commonly capsules or achenes (= cypselae), 1993; Morgan and Soltis 1993; Cosner et al. 1994; rarely berries or drupes. Inulin is found in several Gustafsson and Bremer 1995; Olmstead et al. 2000; families (Calyceraceae, Campanulaceae, Compo- Soltis et al. 2000; Bremer et al. 2001; Lundberg and sitae, Goodeniaceae, Menyanthaceae and Stylidi- Bremer 2003) consists of Menyanthaceae, Good- aceae), and iridoids or seco-iridoids are common, eniaceae, Calyceraceae and Compositae (MGCA but absent from Campanulaceae and Composi- clade; Fig. 1). This clade is characterized by the tae, and apparently also from Alseuosmiaceae, presence of petal lateral veins (Gustafsson 1995), Phellinaceae and Rousseaceae. the loss of micropylar endosperm haustoria (Cos- A tight integration of stamens and style is ner et al. 1994), and a thick and multilayered (> 10 found in several families. In most Stylidiaceae, cells) integument (Inoue and Tobe 1999). Within the two stamens are fused with the style to form this clade, the sister-group relationship between a pressure-sensitive gynostemium. In Calycer- Calyceraceae and Compositae is supported by sev- aceae, Campanulaceae, Compositae and Goodeni- eral potential synapomorphies in wood anatomical aceae, the interaction of style and either fused or (Carlquist and De Vore 1998), inflorescence, flower free anthers results in various forms of secondary and fruit morphological and anatomical (Hansen pollen presentation (Carolin 1960; Leins and Erbar 1992; Gustafsson 1995), and pollen (Hansen 1990, 2003; Erbar and Leins 1995). Erbar and Leins 1992) characters. Goodeniaceae are sister to (1995) classified these as (1) brushing or pump these two families, and the clade consisting of mechanism in Compositae and Campanulaceae- Goodeniaceae/Calyceraceae/Compositae may be Lobelioideae (pollen is removed from an anther supported by pollen grains with a prominent layer tube by the elongating style which is hairy or not), with branched columellae and secondary pollen (2) deposition (or rarely brushing) mechanism presentation involving fused anthers (Lundberg in Campanulaceae-Campanuloideae (pollen from and Bremer 2003). Lundberg and Bremer (2003) free anthers is deposited on hairs on the outside of suggested that Stylidiaceae incl. Donatiaceae, the style, these hairs can invaginate or not), (3) cup astronglysupportedcladeintheirstudy,aresister and cup/brushing mechanism in Goodeniaceae to the MGCA clade. A close relationship between (pollen is deposited in a cup-like outgrowth below Donatiaceae and Stylidiaceae, however, was not the stigma, the indusium; in addition to this cup, found in other analyses (Albach et al. 2001; Bremer hairs can be present on the style) and (4) deposition et al. 2002), and neither Donatiaceae nor Stylidi- mechanism of Goodeniaceae (deposition of pollen aceaeweresistertotheMGCAcladeinthesetwo grains on top of the style). Detailed summaries analyses. Instead, Stylidiaceae were sister to Cam- of character distribution in Asterales have been panulaceae (Albach et al. 2001; Bremer et al. 2002), provided by Lammers (1992; excl. Alseuosmiaceae, and Donatiaceae sister to Alseuosmiaceae/Argo- Argophyllaceae, Carpodetaceae, Phellinaceae, phyllaceae/Phellinaceae (Bremer et al. 2002) or Rousseaceae) and, covering the entire
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