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Home , Burseraceae, Dipentodon, Huerteales, Kirkia, Kirkiaceae, Melia (plant)

The Families and Genera of Vascular 10

Flowering Plants.

Sapindales, , Myrtaceae

Bearbeitet von Klaus Kubitzki

1. Auflage 2011. Buch. x, 436 S. Hardcover ISBN 978 3 642 14396 0 Format (B x L): 19,3 x 26 cm Gewicht: 1142 g

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K. KUBITZKI

CONSPECTUS OF F AMILIES drupaceous with resinous mesocarp; with oily and starchy ; endotegmen lignified, 1. Herbs or low-growing 2 usually thickened; embryo curved, with fleshy – Erect shrubs or (some herbaceous) cotyledons.n ¼ 7–12, 14–16, 21. 81/c. 800. Pantropi- 4 cal, also temperate Anacardiaceae 2. Perennial herbs; nectary glands 5, at base of antese- 6. dehiscing with 4 or 8 one-seeded mericarps palous ; carpels with distinct stylodia arising from a central column; flowers isomerous, 4-merous; from base of ovarioles; solitary, pendulous, testa thin; endosperm 0; trees with alternate, imparipin- epitropous; embryo sac tetrasporic, 16-celled; n ¼ 5. nate ; ellagic acid present. 1/6. , 1/4 or 5. E Mediterranean to C Biebersteiniaceae Tetradiclis – Fruit not dehiscing from central column 7 – Low shrubs of saline habitats, rarely ( ) 7. Pericycle containing a cylinder of sclerenchyma annual herbs; intrastaminal nectary disk annular or (, Guindilia, and some Acereae excepted); angular; with simple style; ovules 1 or several plants containing in idioblasts but no bitter per carpel, epitropous or apotropous; embryo sac, as nortriterpenoids; leaves alternate or less often (Acer- far as known, of Polygonum type 3 eae, Hippocastaneae) opposite; flowers actinomorphic 3. 1 per carpel, apotropous; fruit drupaceous; n ¼ or obliquely zygomorphic; disk extrastaminal or less 12, 30. 1/5–8. Old World, Australia often intrastaminal, annular (in Xanthoceras,with horn-like appendages) or unilateral; – Ovules several to many per carpel, epitropous; fruit n ¼ sometimes (Hippocastaneae, ) with a loculicidal or a ; 7, 12, 13. 3/7–8. scale-like appendage concealing nectary; ovules E and S Europe to Middle Asia, 1 or 2 per carpel or rarely more, usually apotropous. Tetradiclidaceae n ¼ 10–16, 20. 141/c. 1,900. Pantropical, with some 4. Plants usually strongly resinous, with vertical temperate genera s.l. canals in the bark and also with resin ducts in the phloem of the larger veins of the leaves and some- – Pericycle without a cylinder of sclerenchyma; pro- times in wood rays; producing biflavonyls 5 ducing bitter nortriterpenoids (limonoids or quassi- – Plants resinous or not, but without resin ducts in the noids) 8 bark, rays, and veins; biflavonyls 0 6 8. Leaves pellucid-punctate and secretory schizogenous 5. Ovules 2 in each , epitropous, collateral or cavities scattered through the parenchymatous tissue (Beiselia) superposed; nodes mostly 5-lacunar 5-trace; (not in all Cneoroideae); flowers mostly actinomorphic flowers actinomorphic and obdiplostemonous, or and obdiplostemonous, sometimes stamens in one cycle and antesepalous; nectary disk intrastaminal; with the antesepalous whorl reduced; gynoe- þ cium of (2)3–5(9–13 in Beiselia) connate carpels; style carpels (2)4–5 , more or less connate proximally and simple with 2–3-lobed or capitate ; usually held together by the joined stylodia, less often with 1–5 one-seeded pyrenes or pseudocap- completely connate; ovules (1)2–many in each locule, sules releasing pyrenes; endotesta lignified; seeds usually epitropous; fruits follicles, drupes, , or samaras; producing limonoids, canthin-6-ones, and exalbuminous, with hemicellulosic reserves; embryo n ¼ þ minute, with folded, usually palmately lobed cotyle- alkaloids of different types. 7–11, 18 .154/c. dons. n ¼ 11, 13, 23. 19/640. Pantropical 1,800. Pantropical and temperate – Leaves not pellucid-punctate 9 – Ovule solitary in each locule, apotropous, more 9. Stamen filaments not appendaged, usually connate rarely epitropous; nodes mostly 3-lacunar 3-trace; into a staminal tube with anthers in one or two flowers often monosymmetric, obdiplostemonous or whorls, less often filaments distinct; nodes mostly with (1)5–10þ stamens; of 4–12 distinct 5-lacunar 5-trace; ovary (1)2–6(–20)-carpellate, syn- carpels of which usually only one is fertile, or of carpous; style simple; ovules 1–2 or more per carpel, (2)3(–5) connate carpels; stylodia distinct or more usually epitropous; seeds often sarcotestal or arillate; or less connate into a simple style; fruit usually coat exotegmic with fibres or pachychalazal; 2 K. Kubitzki

producing limonoids. n ¼ 8(–180). 50/c. 575. Pan- other Malvids tropical, some temperate – Stamen filaments distinct, usually with scaly append- age; nodes 3-lacunar; carpels (1)2–5, distinct or Biebersteiniaceae basally or ventrally connate; stylodia distinct, conglu- Nitrariaceae tinate or connate into a common style; ovule 1 per carpel, epitropous; seeds not fleshy; seed coat usually Tetradiclidaceae nondescript, pachychalazal in and ; producing bitter quassinoids, limonoids, and canthin-6- Sapindaceae ones. n ¼ 10–13. 22/100. Pantropical, some temperate Kirkiaceae Nineteenth century botanists, such as Bentham Anacardiaceae (in Bentham and Hooker 1862) and Engler (e.g., Burseraceae 1931), tended to treat Sapindales and Rutales (the latter sometimes as ) as distinct Rutaceae orders, a concept followed by Takhtajan (2009) Simaroubaceae to the present day; however, a wider ordinal Meliaceae concept with Rutales included in Sapindales, Fig. 1. Phylogenetic relationships of Sapindales families, as Terebinthales (Wettstein 1901)orSapindales based on rbcL sequence data from Muellner et al. (2007) (Cronquist 1968), is now broadly supported and Sheahan and Chase (1996) and accepted. Gene sequence studies (Sheahan and Chase 1996;Gadeketal.1996; Muellner et al. 2007, among others) have contributed to shaping the present concept of the and the Burseraceae/Anacardiaceae clade, with which provided support for its monophyly, with it shares important similarities in floral structure increasing indications for and Brassi- (Bachelier and Endress 2008). Burseraceae are tra- cales and the little known as close ditionally distinguished from Anacardiaceae by relatives of Sapindales (Worberg et al. 2009). having two collateral ovules (except for Beiselia The multigene analysis of Wang et al. (2009) in which the two ovules are superposed) that are has recovered the strongly supported relation- epitropous, in contrast to all other Sapindales. ship [ [Sapin- Bachelier and Endress (2009) report, however, dales [Huerteales [ þ Malvales]]]]. that in the earliest developmental stages the ovules Insights from morphology and molecular work, in Burseraceae appear apotropous. Thus, the ratio- particularly a two-gene analysis with a broader nale for the use of ovule curvature as a criterion sampling of Sapindales (Muellner et al. 2007), for ordinal distinction becomes questionable. suggest the topology presented here (Fig. 1), The close relationship between Burseraceae in which, however, the precise relationship and Anacardiaceae is well established both by between Simaroubaceae and Meliaceae remains anatomical (Takhtajan 2009), floral morphologi- weakly supported. cal (Bachelier and Endress 2009), and molecular The androecium is often (basically?) obdi- evidence. Sapindaceae are treated here to include plostemonous (with the carpels in antepetalous and Hippocastanaceae, in a return to position), and the two stamen whorls sometimes the practice of several nineteenth century authors (in Burseraceae, Rutaceae, and Sapindaceae) (for historical aspects, see the treatment) appear in a single cycle (meta-obdiplostemony, and in conformity with the results of recent Lam 1931), or one cycle is missing. The herba- molecular studies (e.g., Harrington et al. 2005; ceous and shrubby, early diverging Nitrariaceae, Buerki et al. 2009), which have also brought to Tetradiclidaceae, and Biebersteiniaceae are still light the peculiar position of Xanthoceras as little known but exhibit variation in ovule curva- a basal branch of Sapindaceae. Rutaceae, - tion and in seed and fruit structure, obviously ceae, and Simaroubaceae share the possession in adaptation to the challenges of their saline of unusual bitter compounds, the limonoids or semiarid habitats. , formerly included and quassinoids, which are based on degraded in Simaroubaceae, is now recognised as sister to triterpenes, the nortriterpenoids. The simplest Introduction to Sapindales 3 limonoids are found in Rutaceae, and occur in Buerki, S., Forest, F., Acevedo-Rodrı´guez, P., Callmander, increasing complexity in Meliaceae and in - M.W., Nylander, J.A.A., Harrington, M., Sanmartin, I., Kupfer,€ P., Alvarez, N. 2009. Plastid and nuclear ceae/Cneoroideae. Cneoroideae comprise genera DNA markers reveal intricate relationships at sub- that until recently had been treated as belonging familial and tribal levels in the soapberry family to either Rutaceae or Simaroubaceae, or had been (Sapindaceae). Mol. Phylogen. Evol. 51: 238–258. separated into small satellite families, but the Cronquist, A. 1968. The evolution and classification of flowering plants. London: Nelson. presence of triterpenoid bitter compounds and Engler, A. 1931. , Rutaceae. In: Engler, A., particularly the results of gene sequence studies Prantl, K. (eds.) Die naturlichen€ Pflanzenfamilien, have yielded strong arguments for combining 2nd edn, vol. 19a, pp. 184–359. Leipzig: W. Engelmann. them with the Rutaceae. The peculiar apocarpy Gadek, P., Fernando, E.S., Quinn, C.J., Hoot, S.B., Terrazas, T., Sheahan, M.C., Chase, M.W. 1996. of Rutaceae and Simaroubaceae, thought by some Sapindales: molecular delimitation and infraordinal to be inherited directly from groups. Am. J. Bot. 83: 802–811. or , has been revealed to be a phy- Harrington, M.G., Edwards, K.J., Johnson, S.A., Chase, M.W., Gadek, P.A. 2005. Phylogenetic inference in logenetically secondary condition, as is evidenced Sapindaceae lato using plastid matK and rbcL by the peculiar postgenital connation of the DNA sequences. Syst. Bot. 30: 366–382. stylodia that hold together the carpels in the Kubitzki, K., Krutzsch, W. 1996. Origins of East flowering stage (Ramp 1988). and Southeast Asian diversity, pp. 56–70. In: Floristic characteristics and and diversity of East Sapindales are an ancient lineage with a Asian plants. Beijing: Higher Education record dating back to the Cretaceous. At least Press, and Berlin: Springer. from the onward, Meliaceae, Rutaceae, Lam, H.J. 1931. Beitr€age zur Morphologie der dreiz€ahli- Sapindaceae, Anacardiaceae, and Burseraceae are gen Burseraceae-Canarieae. Ann. Jard. Bot. Buiten- zorg 42: 25–56, t. v–vii. represented by reliable in the northern Lam, H.J. 1932. Beitr€age zur Morphologie der - hemisphere, particularly in North America and ceae, insbesondere der Canarieae. Ann. Jard. Bot. Europe; Simaroubaceae follow in the early Buitenzorg 42: 97–226, t. xi–xvi. (for documentation, see family treatments in this Muellner, A.N., Vassiliades, D.D., Renner, S.S. 2007. Placing Biebersteiniaceae, a herbaceous clade of volume). It is likely that the early evolution of Sapindales, in a temporal and geographic context. Sapindales took place in North America, and that Pl. Syst. Evol. 266: 233–252. in the Eocene they dispersed eastward through Ramp, E. 1988. Struktur, Funktion und systematische the warm-temperate belt north of the Sea of Bedeutung des Gynoeciums bei den Rutaceae und Simaroubaceae. Inaug.-Diss., Philos. Fak. II, Univ. Tethys (often erroneously called “paratropical”, Zurich:€ ADAG. see Kubitzki and Krutzsch 1996), and from there Sheahan, M.C., Chase, M.W. 1996. A phylogenetic analysis invaded and diversified in tropical regions. of R.Br. based on morphological, anatomical and rbcL DNA sequence data. Bot. J. Linn. Soc. 122: 279–300. Takhtajan, A. 2009. Flowering plants, 2nd edn. Dordrecht: Springer. Wang, H., Moore, M.J., Soltis, P.S., Bell, C.D., References Brockington, S.F., Alexandre, R., Davis, C.D., Latvis, M., Manchester, S.R., Soltis, D.E. 2009. Bachelier, J.B., Endress, P.K. 2008. Floral structure of Rosid radiation and the rapid rise of angiosperm- Kirkia (Kirkiaceae) and its position in Sapindales. dominated forests. Proc. Natl. Acad. Sci. USA 106: Ann. Bot. 102: 539–550. 3853–3858. Bachelier, J.E., Endress, P.K. 2009. Comparative floral Wettstein, R. v. 1901. Handbuch der systematischen morphology and anatomy of Anacardiaceae and Botanik, vol. 1. Leipzig and Wien: F. Deuticke. Burseraceae (Sapindales), with a special focus on Worberg, A., Alford, M.H., Quandt, D., Borsch, T. 2009. gynoecium structure and evolution. Bot. J. Linn. Huerteales sister to Brassicales plus Malvales, Soc. 159: 499–571. and newly circumscribed to include , Bentham, G., Hooker, J.D. 1862. Genera Plantarum, vol. I , , , and . (1). London: Reeve. Taxon 58: 468–478. http://www.springer.com/978-3-642-14396-0