The Families and Genera of Vascular Plants 7 Flowering Plants · Dicotyledons Lamiales (except Acanthaceae including Avicenniaceae) Bearbeitet von Joachim W. Kadereit 1. Auflage 2004. Buch. IX, 478 S. Hardcover ISBN 978 3 540 40593 1 Format (B x L): 21 x 27,9 cm Gewicht: 1433 g 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. Lamiales: Introduction and Conspectus 1 Lamiales: Introduction and Conspectus J.W. Kadereit Lamiales as presented here are a well-supported son to the latter three systems, the Lamiales as cir- clade of the Lamiids (Bremer et al. 2002) found in cumscribed here differ by the exclusion of Borag- all major, recent molecular studies of the Asterids inaceae (incl. Lennoaceae), treated as part of (APG II 2003; see also Wagenitz 1992). The Lamiales by Cronquist (1981), and by the inclusion Lamiids, comprising approximately 23,600 species of Byblidaceae (Cronquist 1981: Rosales; Thorne in 1140 genera, are herbaceous or less commonly 1992: Pittosporales; Takhtajan 1997: Aralianae- woody plants with often opposite leaves. The Byblidales), Carlemanniaceae (Cronquist 1981: flowers are sympetalous and mostly 5-merous, Dipsacales; Thorne 1992: not recognized; Takhta- often representing zygomorphic lip flowers with a jan 1997: Rubiales) and Plocospermataceae (Cron- showy corolla and with a single whorl of stamens quist 1981: Gentianales; Thorne 1992: Gentianales; (the five stamens often reduced to four or two), Takhtajan 1997: Gentianales), treated as belonging and a bicarpellate ovary with the two carpels in elsewhere by Cronquist (1981), Thorne (1992) and median position (Endress 1994). Anthers very Takhtajan (1997). There is good molecular evi- often have an outgrowth of the connective into dence for the inclusion of these three families in the anther locule, known as pollen sac placentoid Lamiales (Byblidaceae: Hedrén et al. 1995; Bremer (Hartl 1964), glandular hairs with a head with only et al. 2001; Albach et al. 2001; Carlemanniaceae: vertical cell walls are often present, and pollen Savolainen, Fay et al. 2000; Bremer et al. 2001; tubes appear to lack callose (Prósperi and Cocucci Plocospermataceae: Bremer et al. 2001), and 1979; Cocucci 1983). Both micropylar and chalazal other characters in support of this can be found. endosperm haustoria are common (Yamazaki Thus, Plocospermataceae contain cornoside as a 1974), endosperm formation is cellular, and caffeic acid derivative commonly found in the embryogeny normally is of the Onagrad type order (Jensen 1996, 2000), the carnivorous Bybli- (Polygonad in Oleaceae; Albach et al. 2001). daceae share embryological characters with the Protein bodies in the nuclei are widespread (Speta equally carnivorous Lentibulariaceae (Lang 1901; 1977, 1979; Bigazzi 1984, 1989a, b). Chemically, Conran 1996), and Carlemanniaceae have flowers typical constituents of the order are 6- and/or 8- with only two stamens which may prove to be hydroxylated flavone glycosides (Grayer et al. morphological support of their sister-group 1999), verbascosides and cornoside as charac- relationship with Oleaceae. teristic caffeic acid derivatives (Mølgaard and The exact relationships of Lamiales within Ravn 1988; Jensen 1992), shikimate-derived Lamiids are not finally resolved, but it is clear from anthraquinones (Jensen 1992), C11-decarboxy- molecular evidence (e.g. Downie and Palmer 1992; lated iridoids (Jensen 1991,1992),and stachyose or Olmstead et al. 1992, 1993; Backlund and Bremer other oligosaccharides as carbon storage com- 1997; Olmstead et al. 2000; Savolainen, Chase et al. pounds (Hegnauer 1989). As will be discussed 2000; Savolainen, Fay et al. 2000; Soltis et al. 2000; below, several of the characters listed above, Albach et al. 2001; Bremer et al. 2001; Olmstead including the zygomorphic lip flowers, originated et al. 2001) that they are part of a larger clade only within the order (Stevens 2001). containing Boraginaceae/Boraginales, Gen- In large part, Lamiales as circumscribed here tianales, Solanales, Vahliaceae. In some of these correspond to the Tubiflorae (Wettstein 1935; analyses (Olmstead et al. 2000; Savolainen, Chase Melchior 1964) or Personatae (Drude 1885– et al. 2000) they are sister to Gentianales. Inclusion 1887; Warming and Möbius 1929; Huber 1991) of Vahliaceae into Lamiales should be considered of earlier authors, and to Cronquist’s (1981) (Savolainen, Fay et al. 2000; Albach et al. 2001). Lamiales, Callitrichales, Plantaginales and Scro- Although the Lamiales clearly represent a phulariales, to Thorne’s (1992) Bignoniales, and to monophylum, their division into families in this Takhtajan’s (1997) Oleales, Scrophulariales, Lami- treatment differs from that proposed by APG II ales, Callitrichales and Hippuridales. In compari- (2003; see Table 1). 2J.W.Kadereit Table 1. Families recognized in this volume compared with aceae. Buddlejaceae, Callitrichaceae, Globulari- the APG classification aceae, Hippuridaceae, Plantaginaceae s.str. as well APG II FGVP as Myoporaceae are treated as separate families. Also, Phrymaceae here are treated as a monotypic Acanthaceae Acanthaceae family. Such treatment of Scrophulariaceae s.l. is Bignoniaceae Bignoniaceae non-phylogenetic in at least one respect. It is quite = Scrophulariaceae s.str. Buddlejaceae clear that Scrophulariaceae s.l. are paraphyletic Byblidaceae Byblidaceae = Plantaginaceae s.l. Callitrichaceae when Buddlejaceae, Callitrichaceae, Globulari- Calceolariaceae Srophulariaceae s.l. aceae, Hippuridaceae, Myoporaceae and Plantagi- Carlemanniaceae Carlemanniaceae naceae, and probably also Phrymaceae are treated Orobanchaceae Cyclocheilaceae as separate families. Buddlejaceae and Myopo- Gesneriaceae Gesneriaceae raceae group with one part of Scrophulariaceae s.l. = Plantaginaceae s.l. Globulariaceae = Plantaginaceae s.l. Hippuridaceae with good support (= Scrophulariaceae s.str.: 78%; Labiatae Labiatae Olmstead et al. 2001), and Callitrichaceae, Globu- Lentibulariaceae Lentibulariaceae lariaceae and Hippuridaceae group with another Martyniaceae Martyniaceae part of Scrophulariaceae s.l., although with only = Scrophulariaceae s.str. Myoporaceae Orobanchaceae Nesogenaceae low support (= Plantaginaceae = Veronicaceae: Oleaceae Oleaceae 48%; Olmstead et al. 2001). The recognition of a Orobanchaceae = Scrophulariaceae s.l. close relationship between most of the preceding Paulowniaceae = Scrophulariaceae s.l. families and Scrophulariaceae is by no means Pedaliaceae Pedaliaceae novel and had already been recognized (and partly Phrymaceae Phrymaceae Plantaginaceae s.l. Plantaginaceae s.str. formalized) by, for example, Hallier (1903) and Plocospermataceae Plocospermataceae Hartl (1965–1974). Following Olmstead et al. Schlegeliaceae = Scrophulariaceae s.l. (2001), recognition of Scrophulariaceae s.l. includ- Scrophulariaceae Scrophulariaceae ing Buddlejaceae, Callitrichaceae, Globulariaceae, Stilbaceae Stilbaceae Hippuridaceae, Myoporaceae and Plantaginaceae Tetrachondraceae Tetrachondraceae = Pedaliaceae Trapellaceae s.str. is not significantly less parsimonious than Ver b enaceae Verbenaceae dividing them into the above five families, pro- vided that Calceolariaceae are excluded. This latter family groups quite distant from Scrophulariaceae s.l. Considering this and the comparatively small Apart from the inclusion of Avicenniaceae in sample of genera analysed so far, treatment of Acanthaceae here (Schwarzbach and McDade Scrophulariaceae in a wide circumscription 2002), in contrast to APG II, and the inclusion of is justifiable, but not when Buddlejaceae, Trapellaceae in Pedaliaceae by APG II, also in Callitrichaceae, Globulariaceae, Hippuridaceae, contrast to the present work, the major difference Myoporaceae and Plantaginaceae (and probably between APG II and this FGVP volume lies in the Phrymaceae) are excluded, and Calceolaria treatment of Scrophulariaceae s.l. and allies. and allies are included. However, it is very likely Following the molecular evidence last summa- that eventually Calceolariaceae, Orobanchaceae, rized and expanded by Olmstead et al. (2001),APG Paulowniaceae, Plantaginaceae/Veronicaceae II divide Scrophulariaceae s.l. and allies into (incl. Callitrichaceae, Globulariaceae and Hippuri- Orobanchaceae, Paulowniaceae, Plantaginaceae daceae), Schlegeliaceae, Scrophulariaceae s.str. (incl. Callitrichaceae, Globulariaceae and Hippuri- (incl. Buddlejaceae and Myoporaceae) and an daceae; these are the Veronicaceae of Olmstead expanded Phrymaceae will have to be recognized et al. 2001), Schlegeliaceae and Scrophularia- because these lineages are, at least partly, more ceae s.str. (incl. Buddlejaceae and Myoporaceae). closely related to other families of Lamiales Calceolariaceae are recognized as an additional than to each other. Accordingly, the treatment of family by Olmstead et al. (2001), and there is Scrophulariaceae s.l. (see Fischer, this Vol.) increasing evidence that Phrymaceae, containing indicates the likely contents of the above lineages. Phryma and several genera mainly (but not only) As regards the treatment of Labiatae/