A Revised Classification of the Apocynaceae S.L
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THE BOTANICAL REVIEW VOL. 66 JANUARY-MARCH2000 NO. 1 A Revised Classification of the Apocynaceae s.l. MARY E. ENDRESS Institute of Systematic Botany University of Zurich 8008 Zurich, Switzerland AND PETER V. BRUYNS Bolus Herbarium University of Cape Town Rondebosch 7700, South Africa I. AbstractYZusammen fassung .............................................. 2 II. Introduction .......................................................... 2 III. Discussion ............................................................ 3 A. Infrafamilial Classification of the Apocynaceae s.str ....................... 3 B. Recognition of the Family Periplocaceae ................................ 8 C. Infrafamilial Classification of the Asclepiadaceae s.str ..................... 15 1. Recognition of the Secamonoideae .................................. 15 2. Relationships within the Asclepiadoideae ............................. 17 D. Coronas within the Apocynaceae s.l.: Homologies and Interpretations ........ 22 IV. Conclusion: The Apocynaceae s.1 .......................................... 27 V. Taxonomic Treatment .................................................. 31 A. Key to the Subfamilies of the Apocynaceae s.1 ............................ 31 1. Rauvolfioideae Kostel ............................................. 32 a. Alstonieae G. Don ............................................. 33 b. Vinceae Duby ................................................. 34 c. Willughbeeae A. DC ............................................ 34 d. Tabernaemontaneae G. Don ..................................... 35 e. Melodineae G. Don ............................................ 36 f. Hunterieae Miers .............................................. 36 g. Plumerieae E. Mey ............................................. 36 h. Carisseae Dumort .............................................. 37 i. Alyxieae G. Don ............................................... 37 Copies of this issue [66(1)] may be purchased from the NYBG Press, The New York Botanical Garden, Bronx, NY 10458-5125, U.S.A. Please inquire as to prices. The BotanicalReview 66(1): 1-56, January-March 2000 2000 The New York Botanical Garden 1 2 THE BOTANICAL REVIEW 2. ApocynoideaeBurnett ............................................. 38 a. Wrightieae G. Don ............................................. 39 b. MalouetieaeMiill.-Arg .......................................... 39 e. Apocyneae Rchb...... -......................................... 40 d. Mesechiteae Miers ............................................. 41 e. Echiteae Bartl................................................. 41 3. PeriplocoideaeR. Br. ex Endl....................................... 42 4. SecamonoideaeEndl .............................................. 44 5. AsclepiadoideaeR. Br. ex Burnett ................................... 45 a. MarsdenieaeBenth ............................................. 46 b. Ceropegieae Orb............................................... 47 c. Asclepiadeae(R. Br.) Duby...................................... 48 VI. Acknowledgments ..................................................... 51 VII. Literature Cited ........................................................ 52 I. Abstract The Asclepiadaceae, as traditionally defined, have repeatedly been shown to be an apo- morphic derivative of the Apocynaceae. It has often been recommended that the Asclepi- adaceae be subsumed within the Apocynaceae in order to make the latter monophyletic. To date, however, no comprehensive, unified classification has been established. Here we pro- vide a unified classification for the Apocynaceae, which consists of 424 genera distributed among five subfamilies: Rauvolfioideae, Apocynoideae, Periplocoideae, Secamonoideae, and Asclepiadoideae. Keys to the subfamilies and tribes are provided, with lists of genera that (as far as we have been able to ascertain) are recognized in each tribe. Zusammenfassung Es wurde wiederholt festgestellt, dass die Asclepiadaceae in der traditionellen Um- grenzung ein apomorphes Derivat der Apocynaceae sind. Deshalb wurde oft vorgeschlagen, sie bei den Apocynaceae unterzubringen, damit die Apocynaceae eine monophyletische Gruppe darstellen. Bis jetzt wurde aber keine eingehende kombinierte Klassifikation pub- liziert. Hier stellen wir eine kombinierte Klassifikation vor for die Apocynaceae, mit 424 Gat- tungen, die in fiinf Unterfamilien gegliedert sind: Rauvolfioideae, Apocynoideae, Periplocoideae, Secamonoideae und Asclepiadoideae. Schliissel zu den Unterfamilien sowie den Triben werden gegeben. Die Gattungen, die zu jeder Tribus gehSren (soweit bekannt) werden aufgelistet. II. Introduction In 1810 Robert Brown published two seminal papers, "Prodromus Flora Novae Hollan- diae" (Brown, 1810a) and "On the Asclepiadeae" (Brown, 1810b). In these two publications Brown separated the Asclepiadeae (Asclepiadaceae) from the Apocineae (Apocynaceae) of Jussieu (1789) for the first time. In his Prodromus, which came out about a week before the second paper (Mabberley, 1985; Forster, 1991), he listed the taxa in two separate orders (families), which he called the Asclepiadeae and the Apocineae. His reasons for these changes were discussed in detail in the second paper: the main one being the presence of translators in the Asclepiadeae and their absence in the Apocyneae. Although Brown's classi- fication has been universally accepted and implemented, controversy over the delimitation of CLASSIFICATION OF APOCYNACEAE S.L. 3 the two families has never been put to rest. They are clearly more similar to each other than to the rest of the Gentianales, and in a number of characters there is a gradation from the Apocy- naceae to the Asclepiadaceae. Three suggestions for recognition and expression of this rela- tionship have appeared: as one family (Hallier, 1905; Demeter, 1922; Safwat, 1962; Stebbins, 1974; Stevens, 1976; Thome, 1976, 1992; Judd et al., 1994; Struwe et al., 1994; Takhtajan, 1997); as an order separate from the Gentianales (Tsiang, 1934; Hutchinson, 1973); and as a suborder within the Gentianales (Rosatti, 1989; Nicholas & Baijnath, 1994; Omlor, 1998). None of these possibilities seems to have met with much acceptance, so that, more than 180 years after Brown's (1810b) treatise, the group is still usually maintained as two families in the Gentianales. Nevertheless, new evidence from more detailed and extensive morpho- logical studies, as well as the rapidly growing body of molecular information (Judd et al., 1994; Endress & Albert, 1995; Sennblad & Bremer, 1996; Sennblad, 1997; Civeyrel et al., 1998; Sennblad et al., 1998; Potgieter, 1999), suggests that Brown's delimitation does not re- flect natural relationships. This new information and cladistic interpretations of it support the recognition of a single entity. Here we again assess the relationship between the Apocynaceae and the Asclepiadaceae, and we present our arguments for the recognition of a single family. A new classification is proposed, descriptions of taxa down to tribal level are provided, and the genera at present rec- ognized in each tribe are listed. The priority rule has made necessary some changes of famil- iar names for subfamilies and tribes. We have adopted the oldest known names according to J. Reveal's Web site for suprageneric names (http://www.inform.umd.edu/PBIO). Thus some familiar names must be discarded. The major changes include: Rauvolfioideae Kostel. rather than Plumerioideae K. Schum.; Vinceae Duby rather than Rauvolfieae Bartl.; and Ceropegieae Orb. rather than Stapelieae Decne. Our classification is based mostly on mor- phological evidence accumulated through our own studies. Changes in classification based on results from molecular studies have been incorporated only where cladistic support was strong, or if they were also corroborated by morphological evidence. III. Discussion A. INFRAFAMILIALCLASSIFICATION OF THE APOCYNACEAES.STR. The infrafamilial classification of the Apocynaceae s.str, has not changed much in the last 50 years. Two subfamilies are usually recognized: the Plumerioideae (= Rauvolfioideae), and the Apocynoideae. Additional subfamilies, such as Tabemaemontanoideae Stapf (1902), Apocynoideae sensu Woodson (1930), Cerberoideae Pichon (1948b), Carissoideae Endlicher (1838), have been described. However, either their delimitation is more ambiguous than that of the Rauvolfioideae and Apocynoideae, or they are at variance with our data. Consequently, they are not recognized here. The Rauvolfioideae typically have the corolla lobes sinis- trorsely contorted in bud, the anthers are mostly unspecialized and free from the style head, and there is a broad array of fruit and seed types, although the seeds are almost always eco- mose. The Apocynoideae, in contrast, are characterized by having the corolla lobes dex- trorsely contorted in bud and anthers specialized with lignified guide rails and adnate to the style head, forming a gynostegium (Fig. 1); the fruit is almost always a dry follicle with co- mose seeds. With the exception of Woodson's (1930) backward hypothesis, the Rauvolfioi- deae have usually been considered to be basal and more heterogeneous than the Apocynoideae. This view is also supported in all molecular investigations of the group and is followed here. 4 THE BOTANICAL REVIEW Fig. 1. Apocynoideaewith filamentsfused to corolla more or less at base, without any staminal foot: Apocynum androsaemifolium (M. Endress). A. Flower. B. Dissected flower with two petals removed. C. Gynostegium. D. Base of dissected flower