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Abstract

An updated suprageneric classification is provided for Apocynaceae to bring the family into better agreement with recent morphological and molecular, mainly phylogenetic-based, results. A total of 366 genera are recognized and placed within five subfamilies, 25 tribes and 49 subtribes. In Apocynaceae s. str., one new tribe (Amsonieae) and two new subtribes (Tonduziinae and Vincinae) are described in Rauvolfioideae, and one new tribe (Rhabdadenieae) and nine new subtribes (Amphineuriinae, Beaumontiinae, Chonemorphinae, Galactophorinae, Papuechitinae, Peltastinae, Pentalinoninae, Prestoniinae and Urceolinae) are described or validated in Apocynoideae. Within Asclepiadoideae, one new tribe (Eustegieae) and three subtribes (Diplolepinae, Pentacyphinae and Tassadiinae) are described or validated.
Accepted by Piero Delprete: 21 Dec. 2013; published: 14 Feb. 2014 175
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Copyright © 2014 Magnolia Press
Phytotaxa 159 (3): 175–194
www.mapress.com/phytotaxa/Article
http://dx.doi.org/10.11646/phytotaxa.159.3.2
An updated classification for Apocynaceae
MARY E. ENDRESS1, SIGRID LIEDE-SCHUMANN2 & ULRICH MEVE2
1Institute of Systematic Botany, University of Zurich, Zollikerstrasse 107, 8008 Zürich, Switzerland. mendress@systbot.unizh.ch
2 Department of Plant Systematics, University of Bayreuth, 95440 Bayreuth, Germany sigrid.liede@uni-bayreuth.de,
ulrich.meve@uni-bayreuth.de
Abstract
An updated suprageneric classification is provided for Apocynaceae to bring the family into better agreement with recent
morphological and molecular, mainly phylogenetic-based, results. A total of 366 genera are recognized and placed
within five subfamilies, 25 tribes and 49 subtribes. In Apocynaceae s. str., one new tribe (Amsonieae) and two new
subtribes (Tonduziinae and Vincinae) are described in Rauvolfioideae, and one new tribe (Rhabdadenieae) and nine new
subtribes (Amphineuriinae, Beaumontiinae, Chonemorphinae, Galactophorinae, Papuechitinae, Peltastinae,
Pentalinoninae, Prestoniinae and Urceolinae) are described or validated in Apocynoideae. Within Asclepiadoideae, one
new tribe (Eustegieae) and three subtribes (Diplolepinae, Pentacyphinae and Tassadiinae) are described or validated.
Introduction
Robert Brown (1810) chose to split the asclepiads out of Jussieu’s (1789) Apocineae, and recognized them as a
separate family, Asclepiadaceae, for practical reasons. For nearly 200 years thereafter most traditional treatments
recognized the two separate but closely related families, although it has long been known that for a number of
morphological characters, more highly evolved taxa of Apocynaceae are more similar to Asclepiadaceae than they
are to the less complex members of their own family. From the beginning, application of molecular phylogenetics
in the group indicated that traditional Asclepiadaceae is not monophyletic because the included taxa of subfamily
Periplocoideae came out as sister to those of Apocynoideae, rather than with the other Asclepiadaceae (Sennblad &
Bremer 1996), and thus that Asclepiadaceae should be included in Apocynaceae, which, in turn, renders
Apocynoideae paraphyletic. And in both Endress et al. (1996) and in Sennblad & Bremer (1996), Rauvolfioideae
were found to be strongly paraphyletic, with all other subfamilies nested therein. In addition, these first two
molecular-based studies brought to light some hitherto unexpected inconsistencies with the then-current
classification of Leeuwenberg (1994) at the tribal level. For example, within Apocynaceae-Rauvolfioideae, taxa
with fleshy, syncarpous berries had traditionally been considered to be “primitive” and were thus placed together at
the base of classifications as the tribe Carisseae, whereas taxa with dry, dehiscent fruits were considered as
“advanced”. However, in both Endress et al. (1996) and Sennblad & Bremer (1996), the taxon that came out at the
base of the family was Alstonia, a genus with dry, dehiscent fruits. Furthermore, both Carisseae and Plumerieae
(Rauvolfioideae) were found to be grossly paraphyletic, and there were indications that some tribes in
Apocynoideae were also likely not monophyletic. This was corroborated for Leeuwenberg’s (1994) Wrightieae in a
study on the tribe by Sennblad et al. (1998), in which the included genera were dispersed among four clades;
furthermore, their results hinted that essentially all other tribes of Apocynoideae, as circumscribed by Leeuwenberg
(1994) were also paraphyletic, though the sample size was too small to address this question with confidence.
In a first step at bringing the taxonomy of the group into better agreement with the available morphological and
molecular evidence, Endress & Bruyns (2000) published a classification, reuniting the two families and accepting
five subfamilies: the two traditionally recognized in Apocynaceae (Rauvolfioideae and Apocynoideae) and the
three from Asclepiadaceae (Periplocoideae, Secamonoideae and Asclepiadoideae), respectively. The two smaller
subfamilies, Periplocoideae and Secamonoideae, were not further divided. Among the three larger subfamilies, 17
tribes were recognized: nine in Rauvolfioideae, five in Apocynoideae and three in Asclepiadoideae. Subtribes were
ENDRESS ET AL.176 Phytotaxa 159 (3) © 2014 Magnolia Press
not included. Within Rauvolfioideae, three additional tribes (Willughbeieae, Hunterieae and Melodineae) were
added to accommodate the disperate lineages that had been included in Leeuwenberg’s (1994) Carisseae, and two
new tribes (Alstonieae and Vinceae), to house the deviating genera within his Plumerieae. Within Apocynoideae,
fewer changes could be made, as resolution was still too poor in several parts of the tree; but an additional tribe,
Malouetieae, was split out of Wrightieae, and another new tribe, Mesechiteae, was recognized.
Between 2000–2007, several studies came out—a number of them based on DNA and broad sampling—that
had far-reaching implications for the systematics of the family and clarified some previously unresolved regions of
the family tree (Liede 2001, Meve & Liede 2001, 2002, 2004a, 2004b, Potgieter & Albert 2001, Liede & Täuber
2000, 2002, Liede & Meve 2001, 2002, Liede & Kunze 2002, Liede et al. 2002a, b, Liede-Schumann et al. 2005,
Rapini et al. 2003, 2004, 2006, 2007, Simões et al. 2004, 2006, 2007, Verhoeven et al. 2003, Venter et al., 2006,
Meve & Liede-Schumann 2007, Endress et al. 2007b, Goyder et al. 2007, Ionta & Judd 2007, Lahaye et al. 2007,
Livshultz et al. 2007, Wanntorp & Forster 2007). Of particular significance was a series of morphology-based
papers focusing on pollen, which revealed the unexpected variation present within tetrads and pollinia of
Periplocoideae (Venter &Verhoeven 2001, Verhoeven & Venter 1998, 2001), or the peculiar pollen of Alyxieae,
which proved to be the most important morphological character supporting the indicated polyphyly of the tribe
sensu Leeuwenberg (1994) and subsequent exclusion of several of the genera (Van der Ham et al. 2001). Despite
greatly increased sample size in molecular-based phylogenetic studies, the closest ancestor of Periplocoideae
remained uncertain. However, this became a moot point as far as the monophyly of the former Asclepiadaceae is
concerned, since at least three independent studies showed that three genera from Africa (Baissea, Oncinotis and
Motandra), which were validated as a separate subtribe, Baisseinae, by De Kruif (1983) but not recognized by
Leeuwenberg (1994), are strongly supported as sister to Secamonoideae and Asclepiadoideae (Potgieter & Albert
2001, Livshultz et al. 2007, Lahaye et al. 2007), corroborating the earlier result based on much a smaller sample by
Sennblad et al. (1998).
The results of these studies were incorporated into a revised classification (Endress et al. 2007a), transferring a
number of genera to new positions in accordance with the new results and correcting other shortcomings in the
previous classification (Endress & Bruyns 2000). In Rauvolfioideae one additional tribe, Aspidospermeae, was
recognized; in Apocynoideae, Baisseinae was raised to tribal rank and two additional tribes (Nerieae and
Odontadenieae) were recognized. Within Asclepiadoideae, 12 subtribes were recognized (four in Ceropegieae, and
eight in Asclepiadeae). Within Rauvolfioideae and Apocynoideae recognition of subtribes was not feasible at the
time, since major clades in several regions of more basal parts of the tree were still unresolved or only weakly
supported.
In the years since the last classification outline was published (Endress et al. 2007a), further publications have
come out, which have brought greater resolution in several places in the phylogeny, allowing us to better
understand the relationships among genera, in general, and to place with more confidence a number of genera
whose positions were previously uncertain. These include papers focused on wood anatomy (Lens et al. 2008,
2009), floral morphology (Wanntorp & Kunze 2009), pollen morphology (Van der Weide & Van der Ham 2012), as
well as mainly molecular-based studies and their taxonomic consequences (Goyder 2009, Livshultz 2010; Simões
et al. 2010, Fishbein et al. 2011, Wanntorp et al. 2011, Meve & Liede-Schumann 2012, Liede-Schumann & Meve
2013, Liede-Schumann et al. 2012, in press, Middleton & Livshultz 2012, Silva et al. 2012, Wanntorp & Kunze
2009, Wanntorp et al. 2011). In addition, the treatment of Apocynaceae s. str. for the Flora Malesiana (Middleton
2007) and Flora of Peninsular Malaysia (Middleton 2011) came out during this time. Furthermore, David
Middleton kindly provided us with the as yet unpublished treatment for the Flora of Cambodia, Laos and Vietnam
(Middleton in press), which together, made available a wealth of new morphological information on some hitherto
poorly known genera, allowing these to be placed with more confidence.
In this paper, we provide an updated, lineage-based classification for the Apocynaceae, focusing in particular
on suprageneric ranks. Subtribes are recognized within Rauvolfioideae and Apocynoideae here for the first time
since phylogenetic methods have been applied to systematics in the family. The great majority are based on earlier
names originally published by the indefatigable apocynologist Marcel Pichon with only a French description.
Leeuwenberg (1994) already validated one tribe and a few subtribes that were among the many originally
published by Pichon (Pichon 1948a, 1949, 1950a, 1950b). Here, one new tribe (Amsonieae) and two new subtribes
(Tonduziinae and Vincinae) within Vinceae are described in Rauvolfioideae, all of them monogeneric. In
Apocynoideae one new monogeneric tribe (Rhabdadenieae) is described, and nine subtribes, published originally
Phytotaxa 159 (3) © 2014 Magnolia Press 177
AN UPDATED CLASSIFICATION FOR APOCYNACEAE
without a Latin diagnosis by Pichon (1947, 1948a, 1948b, 1949, 1950a, 1950b, 1952) are validated:
Amphineuriinae, Beaumontiinae, Chonemorphinae, Galactophorinae, Papuechitinae, Peltastinae, Pentalinoninae,
Prestoniinae and Urceolinae. Although we have used Pichon’s names, the composition of the subtribes (and in
many cases also the tribes) differs significantly compared to his and other earlier classifications. Therefore, a table
is provided comparing the generic composition of tribes and subtribes of Rauvolfioideae and Apocynoideae
recognized here with those of Pichon and Leeuwenberg (Table 1). Within Asclepiadoideae, one new tribe
(Eustegieae) is recognized and three subtribes (Diplolepinae, Pentacyphinae and Tassadiinae) are described or
validated within Asclepiadeae.
UPDATED CLASSIFICATION OF APOCYNACEAE
I. RAUVOLFIOIDEAE Kostel., Allg. Med.-Pharm. Fl. 3: 1054 (1834).
(79 genera) [Including Carissoideae Endl., Gen. Pl.: 578 (1838); Ophioxyloideae Endl., Gen. Pl.: 579 (1838);
Allamandoideae Endl., Ench. Bot.: 294 (1841); Plumerioideae Luerss., Handb. Syst. Bot. 2: 1061 (1882);
Cerberoideae Pichon , Notul. Syst. (Paris) 13: 212 (1948); Tabernaemontanoideae Stapf ex Boiteau & Sastre ,
Adansonia, sér. 2, 15: 240 (1975); Ervatamioideae Y. Tsiang & P.T. Li, Fl. Reipubl. Popul. Sin. 63: 98 (1977).]
1. Tribe Aspidospermateae Miers, Apocyn. S. Amer. 7 (1878).—T: Aspidosperma Mart. & Zucc. (1824). (6
genera)
1. Aspidosperma Mart. & Zucc., Flora 7 (Beil. 4): 135 (1924), nom. cons.
2. Geissospermum Allemão, Pl. Nov. Brasil. : 707 (1846).
3. Haplophyton A. DC. in DC. & A. DC., Prodr. 8: 412 (1844).
4. Microplumeria Baill., Bull. Mens. Soc. Linn. Paris 1: 749 (1899).
5. Strempeliopsis Benth. in Benth. & Hook.f., Gen. 2: 702 (1876).
6. Va l l es i a Ruiz & Pav., Fl. Peruv. Chil. Prodr.: 28 (1794).
2. Tribe Alstonieae G. Don, Gen. Hist. 4: 70, 86 (1837-1838).—T: Alstonia R. Br., nom. cons. (1810). (2 genera)
7. Alstonia R. Br., Asclepiadeae: 64 (1810), nom. cons.
8. Dyera Hook. F., J. Linn. Soc. Bot. 19: 293 (1882).
3. Tribe Vinceae Duby, Bot. Gall. 1: 324 (1828).—T: Vinca L. (1753). (9 genera) [Including Rauvolfieae Bartl.,
Ord. Nat. Pl. 205 (1830; Kopsieae G. Don, Gen. Hist. 4: 71, 100 (1837-1838); Ophioxyleae Spach, Hist. Nat. Vég.
8: 501, 511 (1839).]
3.1. Subtribe Kopsiinae Leeuwenb., Wageningen Agric. Univ. Pap. 94(3): 57. (1994).—T: Kopsia Blume, nom.
cons. (1823).
9. Kopsia Blume, Cat. Buitenzorg: 12 (1823), nom. cons.
3.2. Subtribe Ochrosiinae Pichon ex Boiteau in A. Aubrév. & J.-F. Leroy, Fl. Nouv.-Calédonie & Dépend. 10: 8
(1981).—T: Ochrosia Juss (1789).
10. Ochrosia Juss., Gen.: 144 (1789).
3.3 Subtribe Tonduziinae M.E. Endress, subtrib. nov.—T: Tonduzia Pittier (1908). (Supported by Potgieter &
Albert 2001 and Simões et al. unpubl. results).
—Trees or shrubs with white latex. Leaves alternate or whorled. Flowers small; calycine colleters absent; corolla
salverform; lobe aestivation sinistrorse; anthers free from the style-head, wholly fertile, included; nectaries poorly
developed, annular or two lobes alternating with the carpels; ovary apocarpous; style filiform; style-head with basal
collar. Fruit a pair of long, slender follicles; seeds numerous, compressed, elliptic, covered with long hairs or with
clumps of shorter hairs marginally.
11. Laxoplumeria Markgr., Notizbl. Bot. Gart. Berlin-Dahlem 9: 981 (1926).
12. Tonduzia Pittier, Contr. U.S. Natl. Herb. 12: 103 (1908).
ENDRESS ET AL.178 Phytotaxa 159 (3) © 2014 Magnolia Press
3.4. Subtribe Vincinae M.E. Endress, subtrib. nov.T: Vinc a L. (1853). (Supported by Simões et. al., unpubl.
results).
– Perennial herbs, often procumbent, with white latex. Leaves opposite. Flowers showy; calycine colleters absent;
corolla infundibuliform, usually blue-violet; lobe aestivation sinistrorse; anthers free from the style-head,
specialized with thick, geniculate filaments and spathulate apical appendage; nectary two lobes alternating with the
carpels; ovary apocarpous; style gradually thickened toward the apex; style-head with basal collar. Fruit a pair of
slender follicles; seeds naked, several, not compressed, oblong with a hilar groove, testa warty.
13. Vinca L., Sp. Pl. 1: 109 (1853).
3.5 Subtribe Catharanthinae Pichon ex Boiteau in A. Aubrév. & J.-F. Leroy, Fl. Nouv.-Calédonie & Dépend. 10:
8 (1981).—T: Catharanthus G. Don (1837).
14. Catharanthus G. Don, Gen. Hist. 4(1): 95 (1837).
15. Kamettia Kostel., Allg. Med. Pharm. Fl. 3: 1062 (1834).
16. Petchia Livera, Ann. Roy. Bot. Gard. (Peradeniya) 10: 140 (1926).
3.6. Subtribe Rauvolfiinae Benth. & Hook.f., Gen. Pl. 2: 684 (1876). –T: Rauvolfia L. (1753).
17. Rauvolfia L., Sp., Pl. 1: 208 (1753).
4. Tribe Willughbeieae A. DC. in DC. & A. DC., Prodr. 8: 318 (1844). –T: Willughbeia Roxb., nom. cons. (1820).
(18 genera)
4.1. Subtribe Leuconotidinae Pichon ex Leeuwenb., Wageningen Agric. Univ. Pap. 94(3): 53 (1994).—T:
Leuconotis Jack (1823).
18. Bousigonia Pierre, Bull. Mens. Soc. Linn. Paris, sér. 2, 1: 35 (1898).
19. Cyclocotyla Stapf, Bull. Misc. Inform. 1908: 259 (1908).
20. Leuconotis Jack, Trans. Linn. Soc. 14: 121 (1823).
4.2. Subtribe Willughbeiinae A. DC. in DC. & A. DC., Prodr. 8: 319 (1844).—T: Willughbeia Roxb. (1820), nom.
cons.
21. Willughbeia Roxb., Pl. Coromandel 3: 77, t. 280 (1820), nom. cons.
4.3. Subtribe Lacmelleinae Pichon ex Leeuwenb.,Wageningen Agric. Univ. Pap. 94(3): 54 (1994)
(Lacmelliinae).—T: Lacmellea Karst. (1857). [Including Couminae Pichon ex Leeuweb., Wageningen Agric. Univ.
Pap. 94(3): 54 (1994).]
22. Couma Aubl., Hist. Pl. Guiane Suppl.: 39, t. 392 (1775).
23. Hancornia Gomes, Observ. Bot.-Med. Nonnullis Bras. Pl. 2: 1, t. 1 (1803).
24. Lacmellea H. Karst., Linnaea 28: 449 (1857).
25. Parahancornia Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 242 (1922).
4.4. Subtribe Landolphiinae K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 122, 127 (1895).—T:
Landolphia P. Beauv. (1806).
26. Ancylobothrys Pierre, Bull. Mens. Soc. Linn. Paris, sér. 2, 1: 91 (1898).
27. Chamaeclitandra (Stapf) Pichon, Mém. Inst. Franç. Afrique Noire 35: 202 (1953).
28. Clitandra Benth. in Hook., Niger Fl.: 445 (1849).
29. Cylindropsis Pierre, Bull. Mens. Soc. Linn. Paris, sér. 2, 1: 38 (1898).
30. Dictyophleba Pierre, Bull. Mens. Soc. Linn. Paris, sér. 2, 1: 92 (1898).
31. Landolphia P. Beauv., Fl. Oware Bénin 1: 54 (1804), nom. cons.
32. Orthopichonia H. Huber, Kew Bull. 15: 437 (1962).
33. Pacouria Aubl., Hist. Pl. Guiane: 268, t. 105 (1775).
34. Saba (Pichon) Pichon, Mém. Inst. Franç. Afrique Noire 35: 302 (1953).
35. Vahadenia Stapf in Dyer, Fl. Trop. Africa 4(1): 29 (1902).
Phytotaxa 159 (3) © 2014 Magnolia Press 179
AN UPDATED CLASSIFICATION FOR APOCYNACEAE
5. Tribe Tabernaemontaneae G. Don, Gen. Hist. 4: 70, 87 (1838).—T: Tabernaemontana L. (1753). (15 genera)
[Including Macoubeeae Boiteau & Sastre, Adansonia sér. 2, 15: 244 (1975); Ambelanieae Pichon ex Boiteau, L.
Allorge & Sastre, Adansonia, sér. 2, 18: 276 (1978); Voacangeae Boiteau & Sastre, Adansonia, sér. 2, 15: 246
(1975); Woytkowskieae Boiteau & Sastre, Adansonia, sér. 2, 15: 244 (1975); Ervatamiinae Pichon ex Boiteau in A.
Aubrév. & J.-F.Leroy, Fl. Nouv.-Calédonie & Dépend. 10: 9 (1981).]
5.1. Subtribe Ambelaniinae A.O. Simões & M.E. Endress, Taxon 59: 786 (2010).—T: Ambelania Aubl. (1775).
36. Ambelania Aubl., Hist. Pl. Guiane: 263, t. 104 (1775).
37. Macoubea Aubl., Hist. Pl. Guiane Suppl.: 17, t. 378 (1775).
38. Molongum Pichon, Mém. Mus. Natl. Hist. Nat. 24: 167 (1948).
39. Mucoa Zarucchi, Agric. Univ. Wageningen Pap. 87(1): 40 (1988).
40. Neocouma Pierre, Bull. Mens. Soc. Linn. Paris, sér. 2, 1: 33 (1898).
41. Rhigospira Miers, Apocyn. S. Amer.: 67, t. 10A (1878).
42. Spongiosperma Zarucchi, Agric. Univ. Wageningen Pap. 87 (1): 40 (1988).
5.2. Subtribe Tabernaemontaninae A. DC. in DC. & A. DC., Prodr. 8: 349 (1844).—T: Tabernaemontana L.
(1753). [Including Callichiliinae L. Allorge in L. Allorge & H. Couderc, Bull. Mus. Natl. Hist. Nat., B, Adansonia,
sér. 4, 5: 235 (1983); Sarcopharyngiinae L. Allorge, Bull. Mus. Natl. Hist. Nat., B, Adansonia, sér. 4, 5(2): 235
(1983).]
43. Callichilia Stapf in Dyer, Fl. Trop. Afr. 4(1): 130 (1902).
44. Calocrater K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 175 (1895).
45. Carvalhoa K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 189 (1895).
46. Crioceras Pierre, Bull. Mens. Soc. Linn. Paris 1: 1311 (1897).
47. Schizozygia Baill., Bull. Mens. Soc. Linn. Paris 1: 782 (1888).
48. Tabernaemontana L., Sp. Pl. 1: 210 (1753).
49. Tabernanthe Baill., Bull. Mens. Soc. Linn. Paris 1: 782 (1888).
50. Voacanga Thouars, Gen. Nov. Madag.: 10 (1806).
6. Tribe Melodineae G. Don, Gen. Hist. 4: 71, 101 (1837-1838).T: Melodinus J.R. Forst. & G. Forst. (1775). (5
genera)
51. Diplorhynchus Welw. ex Ficalho & Hiern., Trans. Linn. Soc. London, Bot. ser. 2,2: 22, t. 5 (1881).
52. Craspidospermum Bojer ex A. DC. in DC. & A. DC., Prodr. 8: 323 (1844).
53. Melodinus J.R. Forst. & G. Forst., Charact. Gen. Pl.: 37, t. 19 (1776).
54. Pycnobotrya Benth. in Benth. & Hook.f., Gen. 2: 715 (1876).
55. Stephanostegia Baill., Bull. Mens. Soc. Linn. Paris 1: 748 (1888).
7. Tribe Hunterieae Miers, Apocyn. S. Amer. 6 (1878).—T: Hunteria Roxb. (1824). (4 genera) [Including
Pleiocarpeae K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 122, 133 (1895).]
56. Gonioma E. Mey., Comment. Pl. Africae Austr: 188 (1938).
57. Hunteria Roxb., Fl. Indica 2: 531 (1824).
58. Picralima Pierre, Bull. Mens. Soc. Linn. Paris 2: 1278 (1896).
59. Pleiocarpa Benth. in Benth. & Hook.f., Gen. 2: 699 (1876).
8. Tribe Amsonieae M.E. Endress, trib. nov.—T: Amsonia Walter (1788). (1 genus). (Supported by Potgieter &
Albert 2001 and Endress et al. 2007b).
– Subshrubs or perennial herbs with white latex. Leaves alternate. Flowers medium-sized, relatively showy;
calycine colleters absent; corolla salverform, often blue; lobe aestivation sinistrorse; anthers free from the style-
head, wholly fertile, included; necaries absent, adnate or indistinct; ovary apocarpous; ovules numerous; style-head
with basal collar. Fruit a pair of thin-walled follicles; seeds naked, not compressed, oblong to ovoid with slanted
ends; testa rugulose or corky.
60. Amsonia Walter, Fl. Carol.: 98 (1788).
ENDRESS ET AL.180 Phytotaxa 159 (3) © 2014 Magnolia Press
9. Tribe Alyxieae G. Don, Gen. Syst. 4: 71, 96 (1837-1838).—T: Alyxia Banks ex R. Br. (1810), nom. cons. (7
genera)
9.1. Subtribe Condylocarpinae Pichon ex Leeuwenb., Wageningen Agric. Univ. Pap. 94(3): 56 (1994).—T:
Condylocarpon Desf. (1822). [Including Chilocarpeae Pichon ex Leeuwenb., Wageningen Agric. Univ. Pap. 94(3):
54 (1994).]
61. Chilocarpus Blume, Cat. Buitenzorg 12: 251 (1895).
62. Condylocarpon Desf., Mém. Mus. Hist. Nat. 8: 119 (1822).
63. Plectaneia Thouars, Gen. Nov. Madag.: 11 (1806).
9.2. Subtribe Alyxiinae A. DC. in DC. & A. DC., Prodr. 8: 345 (1844).—T: Alyxia Banks ex R. Br. (1810), nom.
cons.
64. Alyxia Banks ex R. Br., Prodr.: 469 (1810), nom. cons.
65. Lepinia Decne., Ann. Sci. Nat. Bot., sér. 3, 12: 194, t. 9 (1849).
66. Lepiniopsis Valeton, Ann. Jard. Bot. Buitenzorg 12: 251 (1895).
67. Pteralyxia K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 151 (1895).
10. Tribe Plumerieae E. Mey., Comm. Pl. Afr. Austr. 2: 188 (1838).—T: Plumeria L. (1753). (10 genera)
[Including Allamandeae G. Don, Gen. Hist. 4: 71, 102 (1837-1838); Skytantheae Pichon, Notul. Syst. (Paris) 13:
217 (1948); Cerbereae Leeuwenb., Wageningen Agric. Univ. Pap. 94(3): 57 (1994) Thevetieae Miers, Apocyn. S.
Amer.: 6 (1878).]
10.1. Subtribe Allamandinae A. DC. in DC. & A. DC., Prodr. 8: 318 (1844).—T: Allamanda L. (1771).
68. Allamanda L., Mant.: 146, 214 (1771).
10.2. Subtribe Plumeriinae Pichon ex Leeuwenb., Wageningen Agric. Univ. Pap. 94(3): 53 (1994).—T: Plumeria
L. (1753).
69. Himatanthus Willd. ex. Schult. in Roem. & Schult., Syst. Veg. 5: 221 (1819).
70. Mortoniella Woodson, Ann. Missouri Bot. Gard. 26: 257 (1939).
71. Plumeria L., Sp. Pl. 1: 209 (1753).
10.3. Subtribe Thevetiinae A. DC. in DC. & A. DC., Prodr. 8: 342 (1844).—T: Thevetia L. (1758), nom. cons.
[Including Cerberinae Benth. & Hook.f., Gen. Pl. 2: 685 (1876); Camerariinae Pichon, Notul. Syst. (Paris) 13: 219
(1948).]
72. Anechites Griseb., Fl. Brit. W. Indian Isl.: 410 (1861).
73. Cameraria L., Sp. Pl. 1: 210 (1753).
74. Cerbera L., Sp. Pl. 1: 208 (1753).
75. Cerberiopsis Viell. ex Pancher & Sébert in Sébert, Not. Bois. Nouv. Caledonie: 187 (1874).
76. Skytanthus Meyen, Reise 1: 376 (1834).
77. Thevetia L., Opera Varia: 212 (1758), nom. cons.
11. Tribe Carisseae Dumort, Anal. Fam. Pl.: 26 (1829).—T: Carissa L. (1767). (2 genera) [Including Carandieae
G. Don, Gen. Hist. 4: 71, 103 (1837-1838); Arduineae Baill., Hist. Pl. 10: 165, 172. (Sep-Oct 1889).]
78. Acokanthera G. Don, Gen. Hist. 4(2): 485 (1838).
79. Carissa L., Mant.: 52 (1767).
II. APOCYNOIDEAE Burnett, Outlines Bot. 1012, 1095, 1104 (1835). (82 genera) [Including Echitoideae
Luerss., Handb. Syst. Bot. 2: 1062 (1882).]
1. Tribe Wrightieae G. Don, Gen. Syst. 4: 70, 85 (1837-1838).—T: Wrightia R. Br. (1810). (3 genera)
80. Pleioceras Baill., Bull Mens. Soc. Linn., Paris 1: 759 (1888).
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AN UPDATED CLASSIFICATION FOR APOCYNACEAE
81. Stephanostema K. Schum., Bot. Jahrb. Syst. 34: 325 (1904).
82. Wrightia R. Br., Prodr.: 467 (1810).
2. Tribe Nerieae Baill., Hist. Pl. 10: 166, 198 (1889).—T: Nerium L. (1753). (6 genera)
2.1. Subtribe Neriinae Benth. & Hook.f., Gen. Pl. 2: 688 (1876).—T: Nerium L. (1753).
83. Adenium Roem. & Schult., Syst. Veg. 4: 35 (1819).
84. Nerium L., Sp. Pl. 1: 209 (1753).
2.2. Subtribe Alafiinae Pichon ex Leeuwenb., Wageningen Agric. Univ. Pap. 94(3): 59 (1994).—T: Alafia Thours
(1806).
85. Alafia Thouars, Gen. Nov. Madag.: 11 (1806).
86. Isonema R. Br., Asclepiadeae: 52 (1810).
87. Farquharia Stapf, Bull. Misc. Inform. 1912: 278 (1912).
88. Strophanthus DC., Bull. Sci. Soc. Philom. Paris 3: 122 (1802).
3. Tribe Malouetieae Müll. Arg., in Mart., Fl. Brasil. 6(1): 5, 89 (1860).T: Malouetia A. DC. (1844). (13
genera) [Including Robbieae Miers, Apocyn. S. Amer.: 8. (Mai-Jun 1878).]
3.1. Subtribe Galactophorinae Pichon ex M.E. Endress, subtrib. nov. ≡ Pichon, Mém. Mus. Nat. Hist. Nat. sér.
2B, Bot. 1: 38 (1950). Validated here and description amended.—T: Galactophora Woodson (1932). (Supported by
Simoes et. al. 2004, Livshultz et. al. 2007 and Livshultz 2010).
—Subshrubs or perennial herbs with glandular hairs and translucent or white latex. Leaves opposite, rarely
whorled. Flowers large and showy; calycine colleters absent; corolla infundibuliform, with 5 longitudinal ridges
outside; lobe aestivation dextrorse; anthers with lignified guide-rails, sagittate basally, weakly attached to style-
head, included; nectaries five, coalesced basally; ovary apocarpous; style filiform; style-head with five fleshy
projections at base. Fruit a long-stipitate pair of terete follicles; seeds numerous, glabrous; coma micropylar,
sessile.
89. Galactophora Woodson, Ann. Missouri Bot. Gard. 19: 49 (1932).
3.2. Subtribe Pachypodiinae Pichon ex Leeuwenb., Wageningen Agric. Univ. Pap. 94(3): 58 (1994).—T:
Pachypodium Lindl. (1830).
90. Neobracea Britton in Britton & Millsp., Bahama Fl.: 335 (1920).
91. Pachypodium Lindl., Edwards' Bot. Reg. 16: t. 1321 (1830).
3.3. Subtribe Malouetiinae Pichon, Mém. Mus. Nat. Hist. Nat. sér. 2B, Bot. 1: 45 (1950).—T: Malouetia A. DC.
(1844). [Including Holarrheneae Y. Tsiang & P.T. Li, Fl. Reipubl. Popularis Sin. 63: 117 (1977).]
92. Allowoodsonia Markgr., Gard. Bull. Straits Settlem. 22: 23 (1967).
93. Carruthersia Seem., Fl. Vitiensis: 155 (1866).
94. Eucorymbia Stapf, Hook. Icon. Pl. 28: t. 2764 (1903).
95. Funtumia Stapf, Proc. Linn. Soc. 1899: 2 (1899).
96. Holarrhena R. Br., Asclepiadeae: 51 (1810).
97. Kibatalia G. Don, Gen. Hist. 4(1): 86 (1837).
98. Malouetia A. DC. in DC. & A. DC., Prodr. 8: 378 (1844).
99. Malouetiella Pichon, Bull. Jard. Bot. Etat 22: 131 (1952).
100. Mascarenhasia A. DC. in DC. & A. DC., Prodr. 8: 487 (1844).
101. Spirolobium Baill., Bull. Mens. Soc. Linn. Paris 1: 773 (1889), nom. cons.
4. Tribe Rhabdadenieae Pichon ex M.E. Endress, trib. et stat. nov. Pichon, Mém. Mus. Nat. Hist. Nat. sér. 2B,
Bot. 1: 37 (1950) (Rhabdadeniinae). Validated here and description amended.—T: Rhabdadenia Müll. Arg. (1860).
(1 genus). (Supported by Livshultz et al. 2007 and Livshultz 2010).
Slender woody lianas or perennial herbs with white latex. Leaves opposite. Flowers large and showy; calycine
ENDRESS ET AL.182 Phytotaxa 159 (3) © 2014 Magnolia Press
colleters absent; corolla infundibuliform; lobe aestivation dextrorse; anthers with truncate, lignified guide-rails
fused to filaments, firmly attached to style-head, included; nectaries five separate or coalesced basally; ovary
apocarpous; style filiform; style-head cylindrical, conspicuously pilose apically, with a large membranous basal
collar. Fruit a pair of narrowly fusiform follicles; seeds numerous, narrowly boat-shaped, glabrous; coma
micropylar, long-rostrate.
102. Rhabdadenia Müll. Arg. in Mart., Fl. Brasil. 6(1): 173 (1860).
5. Tribe Echiteae Bartl., Ord. Nat. Pl.: 204 (1830).—T: Echites P. Browne (1756). (19 genera) [Including
Parsonsieae A. DC. in DC. & A. DC., Prodr. 8: 399 (1844); Prestonieae Miers, Apocyn. S. Amer.: 9 (1878).]
5.1. Subtribe Pentalinoninae Pichon ex M.E. Endress, subtrib. nov. ≡ Pichon, Mém. Mus. Nat. Hist. Nat. sér. 2B,
Bot. 1: 35 (1950) (Urechitinae). Validated here and description amended.—T: Pentalinon Voigt (1845). (Supported
by Livshultz et al. 2007 and Livshultz 2010).
—Woody lianas, subshrubs or perennial herbs, often with twining stems; latex white. Leaves opposite. Flowers
large and showy; calycine colleters several in a ring or fewer and alternisepalous; corolla infundibuliform, tubular-
campanulate or salverform; lobe aestivation dextrorse; anthers with lignified guide-rails, slender and abruptly
incurved basally and firmly attached to style-head, included; nectaries two separate or five coalesced basally; ovary
apocarpous; style filiform; style-head fusiform with a narrow membranous basal collar. Fruit a pair of slender,
terete to fusiform follicles; seeds numerous, elliptic, attenuate to rostrate, with a coma at micropylar end.
103. Angadenia Miers, Apocyn. South Amer.: 173 (1878).
104. Pentalinon Voigt, Hortus Suburb. Calcutta: 523 (1845).
105. Salpinctes Woodson, Bull. Torrey Bot. Club 58: 453 (1931).
5.2. Subtribe Peltastinae Pichon ex M.E. Endress, subtrib. nov. ≡ Pichon, Mém. Mus. Nat. Hist. Nat. sér. 2B,
Bot. 1: 40 (1950). Validated here and description amended.—T: Peltastes Woodson (1932). (Supported by
Livshultz et. al. 2007 and Livshultz 2010).
—Woody lianas, most parts of the plant usually ferruginous-tomentulose to ferruginous-hispid; latex translucent or
rarely white. Leaves opposite. Flowers showy; calyx markedly foliaceous, colleters opposite and solitary, but
deeply dissected or several in a ring; corolla infundibuliform, tubular-campanulate or salverform; lobe aestivation
dextrorse; anthers with lignified guide-rails, slender and acuminate or rarely auriculate basally, firmly attached to
the style-head, included; nectaries five, separate or coalesced at the base; ovary apocarpous; style filiform; style-
head fusiform with a narrow membranous basal collar. Fruit a pair of stout, fusiform follicles, often remaining
fused at the tips; seeds numerous, elliptic, long-attenuate to rostrate, with a coma at the micropylar end.
106. Asketanthera Woodson, Ann. Missouri Bot. Gard. 19: 46 (1932).
107. Macropharynx Rusby, Mem. New York Bot. Gard. 7: 327, t. 6 (1927).
108. Peltastes Woodson, Ann. Missouri Bot. Gard. 19: 375 (1932).
109. Temnadenia Miers, Apocyn. South Amer.: 207 (1878).
5.3. Subtribe Echitinae Kitt., Taschenb. Fl. Deutschl., ed. 2, 1: 449 (1843).—T: Echites P. Browne (1756).
119. Bahiella J. F. Morales, Sida 22: 342 (2006).
111. Echites P. Browne, Civ. Nat. Hist. Jamaica: 182 (1756).
112. Fernaldia Woodson, Ann. Missouri Bot. Gard. 19: 48 (1932).
5.4. Subtribe Prestoniinae Pichon ex M.E. Endress, subtrib. nov. ≡ Pichon, Mém. Mus. Nat. Hist. Nat. sér. 2B,
Bot. 1: 19 (1950). Validated here and description amended.—T: Prestonia R. Br. (1810), nom. cons. (Supported by
Livshultz et. al. 2007 and Livshultz 2010).
—Woody lianas or rarely small trees; latex usually translucent, sometimes white. Leaves opposite. Flowers
medium-sized, showy; calycine colleters opposite, solitary, but mostly deeply dissected; corolla usually
salverform, rarely sub-infundibuliform; tube orifice constricted by a thickened annulus, often also five free corona
lobes behind anthers; lobe aestivation dextrorse; anthers with lignified guide-rails, acute to acuminate basally and
firmly attached to style-head, included; nectaries five, separate or coalesced basally; ovary apocarpous; style
filiform; style-head fusiform with a narrow membranous basal collar. Fruit a pair of slender, terete to fusiform
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follicles (these very rarely postgenitally fused); seeds numerous, narrowly boat-shaped, with a sessile coma at the
micropylar end.
113. Hylaea J. F. Morales, Novon 9: 83 (1999).
114. Laubertia A. DC. in DC. & A. DC., Prodr. 8: 486 (1844).
115. Prestonia R. Br., Asclepiadeae: 58 (1810), nom. cons.
116. Rhodocalyx Müll. Arg. in Mart., Fl. Brasil. 6(1): 172 (1860).
5.5. Subtribe Parsonsiinae Benth. & Hook.f., Gen. Pl. 2: 687 (1876).—T: Parsonsia R. Br. (1810), nom. cons.
117. Artia Guillaumin, Bull. Soc. Bot. France 88: 380 (1941).
118. Ecua Middleton, Blumea 41: 33 (1996).
119. Parsonsia R. Br., Prodr.: 465 (1810), nom. cons.
120. Thenardia Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. 3 (4th ed.): 209 (1819).
121. Thoreauea J. K. Williams, Lundellia 5: 47 (2002).
6. Tribe Odontadenieae Miers, Apocyn. S. Amer.: 8 (1878).—T: Odontadenia Benth. (1841). (7 genera)
[Including Stipecomeae Miers, Apocyn. S. Amer.: 9 (1878).]
122. Cycladenia Benth., Pl. Hartw.: 322 (1849).
123. Elytropus Müll. Arg., Bot. Zeitung (Berlin) 18: 21 (1860).
124. Odontadenia Benth., J. Bot. (Hooker) 3: 242 (1841).
125. Pinochia B. F. Hansen & M. E. Endress, Edinburgh J. Bot. 64: 270 (2007).
126. Stipecoma Müll. Arg. in Mart., Fl. Brasil. 6(1): 175 (1860).
127. Secondatia A. DC. in DC. & A. DC., Prodr. 8: 445 (1844).
128. Thyrsanthella (Baill.) Pichon, Bull. Mus. Nat. Hist. Natl., 2. sér., 20: 192 (1948).
7. Tribe Mesechiteae Miers, Apocyn. S. Amer.: 10 (1878).—T: Mesechites Müll.-Arg. (1860). (5 genera)
[Including Macrosiphonieae Miers, Apocyn. S. Amer.: 9 (1878).]
129. Allomarkgrafia Woodson, Ann. Missouri Bot. Gard. 19: 45 (1932).
130. Forsteronia G. Mey., Prim. Fl. Esseq.: 133 (1818).
131. Mandevilla Lindl., Edwards' Bot. Reg. 3: t. 7 (1840).
132. Mesechites Müll. Arg. in Mart., Fl. Brasil. 6(1): 150 (1860).
133. Tintinnabularia Woodson, Ann. Missouri Bot. Gard. 23: 387, t. 7 (1936).
8. Tribe Apocyneae Rchb., Fl. Germ. Excurs. 1(3): 410, 429 (1831).—T: Apocynum L. (1753). (24 genera)
[Including Epigyneae Y. Tsiang & P.T. Li, Fl. Reipubl. Popularis Sin. 63: 244 (1977).]
8.1. Subtribe Beaumontiinae Pichon ex M.E. Endress, subtrib. nov. ≡ Pichon, Mém. Mus. Nat. Hist. Nat. sér. 2B,
Bot. 1: 59 (1950). Validated here and description amended.—T: Beaumontia Wall. (1824). (Supported by Livshultz
et. al. 2007, Livshultz 2010 and Middleton & Livshultz 2012).
—Woody lianas with white latex. Leaves opposite. Flowers showy, sometimes large; calycine colleters several in a
ring, few and alternisepalous or absent; corolla infundibuliform to tubular-campanulate; lobe aestivation dextrorse;
anthers with lignified guide-rails, sagittate basally and firmly attached to style-head, exserted or included but
exposed at the base of the expanded throat; nectaries united, sometimes only basally, into a five-lobed disc; ovary
postgenitally (weakly) syncarpous; style slender, cylindrical; style-head broadest at middle, conical apically, ovoid
basally, without a collar. Fruit single, of two mericarps weakly connate into a fat “double follicle”; seeds numerous,
elliptic, compressed, glabrous or pubescent; coma micropylar, sessile or shortly beaked.
134. Beaumontia Wall., Tent. Fl. Napal.: 14 (1824).
135. Parepigynum Tsiang & P.T. Li, Acta Phytotax. Sin. 11: 394 (1973).
136. Vallaris Burm. f., Fl. Indica: 51 (1768).
8.2. Subtribe Papuechitinae Pichon ex M.E. Endress, subtrib. nov. ≡ Pichon, Mém. Mus. Nat. Hist. Nat. sér. 2B,
Bot. 1: 93 (1950). Validated here and description amended.—T: Papuechites Markgr., Nova Guinea 14: 287
(1927). (Supported by Livshultz et. al. 2007, Livshultz 2010 and Middleton & Livshultz 2012).
ENDRESS ET AL.184 Phytotaxa 159 (3) © 2014 Magnolia Press
—Woody lianas with white latex. Leaves opposite. Flowers inconspicuous; calycine colleters few, alternisepalous;
corolla salverform, small; lobe aestivation dextrorse; anthers with lignified guide-rails, sagittate basally and firmly
attached to style-head, included; nectaries five, separate or united into a five-lobed disc; ovary apocarpous; style
cylindrical, short; style-head broadest at middle, conical apically, ovoid basally, without a collar. Fruit a pair of
rather stout follicles; seeds numerous, narrowly ovate to elliptic, compressed, glabrous; coma micropylar, long-
rostrate.
137. Anodendron A. DC. in DC. & A. DC., Prodr. 8: 443 (1844).
138. Ixodonerium Pit., in Lecomte, Fl. Gén. Indo-Chine 3: 1228 (1933).
139. Papuechites Markgr., Nova Guinea 14: 287 (1927).
8.3. Subtribe Amphineuriinae Pichon ex M.E. Endress, subtrib. nov. ≡ Pichon, Mém. Mus. Nat. Hist. Nat. sér.
2B, Bot. 1: 55 (1950). Validated here and description amended.—T: Amphineurion (A. DC.) Pichon (1948).
(Supported by Livshultz et. al. 2007, Livshultz 2010 and Middleton & Livshultz 2012).
—Woody lianas with white latex. Leaves opposite. Flowers medium-sized; calycine colleters few, alternisepalous,
or several spread in a ring; corolla salverform, smallish to medium-sized, white; lobe aestivation dextrorse; anthers
with lignified guide-rails, often with a tuft of hairs apically, sagittate basally and firmly attached to style-head,
included; nectary disc irregularly two to five-lobed; ovary apocarpous; style slender, cylindrical; style-head
broadest at middle, elongate conical apically, narrow cylindrical basally, with or without collar. Fruit a pair of
slender follicles; seeds numerous, linear-oblong, compressed, glabrous; coma micropylar, sessile.
140. Amphineurion (A. DC.) Pichon, Bull. Soc. Bot. Fr. 95: 215 (1948).
141. Sindechites Oliv., Hook. Icon. Pl. 18: t. 1772 (1888).
142. Streptoechites D.J. Middleton & Livsh., Adansonia, sér. 3, 34: 370 (2012).
8.4. Subtribe Apocyinae Pichon ex Leeuwenb., Wageningen Agric. Univ. Pap. 94(3): 59 (1994).—T: Apocynum
L. (1753).
143. Apocynum L., Sp. Pl. 1: 213 (1753).
144. Cleghornia Wight, Icon. Pl. Ind. Orient. (Wight) 4(2): 5, tt. 1310, 1312 (1848).
8.5. Subtribe Urceolinae Pichon ex M.E. Endress, subtrib. nov. ≡ Pichon, Mém. Mus. Nat. Hist. Nat. sér. 2B, Bot.
1: 81 (1950). Validated here and description amended.—T: Urceola Roxb (1799), nom. cons. (Supported by
Livshultz et. al. 2007, Livshultz 2010 and Middleton & Livshultz 2012).
—Woody lianas with white latex. Leaves opposite, usually with domatia. Flowers small, inconspicuous; calycine
colleters several, spreading in a ring or few, alternisepalous, sometimes absent; corolla salverform, urceolate or
campanulate; lobe aestivation dextrorse (sinistrorse in Parameria); anthers with lignified guide-rails, sagittate
basally and firmly attached to style-head, included; nectary of five separate lobes or these fused into a five- or
irregularly-lobed disc; ovary apocarpous; style short; style-head broadest at the middle, conical apically, ovoid
basally, without a basal collar. Fruit a pair of slender to stout follicles, sometimes weakly moniliform; seeds
numerous, ellipsoid, compressed, hirsute; coma micropylar, sessile, with crown of shorter hairs basally.
145. Aganonerion Pierre ex Spire, Contr. Apoc.: 43 (1905).
146. Parameria Benth. in Benth. & Hook.f., Gen. 2: 715 (1876).
147. Urceola Roxb., Asiat. Res. 5: 169 (1799), nom. cons.
8.6. Subtribe Chonemorphinae Pichon ex M.E. Endress, subtrib. nov. Pichon, Mém. Mus. Nat. Hist. Nat. sér.
2B, Bot. 1: 28 (1950). Validated here and description amended.—T: Chonemorpha G. Don (1837). (Supported by
Livshultz et. al. 2007 and Middleton & Livshultz 2012).
—Woody lianas with white latex. Leaves opposite. Flowers inconspicuous to showy; calycine colleters several,
spreading in a ring; corolla salverform or tubular-campanulate; lobes asymmetric to falcate, aestivation dextrorse;
anthers subsessle, with lignified guide-rails, sagittate basally and firmly attached to style-head, included or
exserted; nectary of five separate lobes or a five-lobed disc; ovary apocarpous; style slender, cylindrical; style-head
broadest at middle, conical to ovoid apically, cup-shaped basally, without a basal collar. Fruit a pair of long
fusiform follicles; seeds numerous, long and narrow, glabrous; coma micropylar, sessile.
148. Chonemorpha G. Don, Gen. Hist. 4(1): 76 (1837).
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149. Trachelospermum Lem., Jard. Fleur. 1: t. 61 (1851), nom. cons.
150. Vallariopsis Woodson, Philipp. J. Sci. 60: 288 (1936).
8.7. Subtribe Ichnocarpinae Benth. & Hook.f., Gen. Pl. 2: 688 (1876).—T: Ichnocarpus R. Br., nom. cons.
(1810).
151. Aganosma (Blume) G. Don, Gen. Hist. 4(1): 77 (1837).
152. Amalocalyx Pierre, Bull. Mens. Soc. Linn. Paris, sér. 2, 1: 28 (1898).
153. Baharuia Middleton, Blumea 40: 445 (1995).
154. Epigynum Wight, Icon. Pl. Ind. Orient. (Wight) 4(2): 4 (1848).
155. Ichnocarpus R. Br., Asclepiadeae: 50 (1810), nom. cons.
156. Micrechites Miq., Fl. Ind. Bat. 2: 456 (1857).
157. Pottsia Hook. & Arn., Bot. Beechey's Voyage: 198 (1837).
9. Tribe Baisseeae M.E. Endress, Ann. Missouri Bot. Gard. 94: 10 (2007).—T: Baissea A. DC. (1844). (4 genera)
158. Baissea A. DC. in DC. & A. DC., Prodr. 8: 424 (1844).
159. Dewevrella De Wild., Mission Laurent 1: 548 et 2: tt. 414, 415 (1907).
160. Motandra A. DC. in DC. & A. DC., Prodr. 8: 423 (1844).
161. Oncinotis Benth. in Hook., Niger Fl.: 451 (1849).
III. SUBFAMILY PERIPLOCOIDEAE Endl., Gen. Pl.: 587 (1838).—T: Periploca L. (33 genera) [Including
Cryptolepideae Venter, Taxon 46: 713 (1997); Gymnanthereae Y. Tsiang & P.T. Li, Reipubl. Popularis Sin. 63: 260
(1977).]
162. Atherandra Decne., Prodr. 8: 497 (1844).
163. Baroniella Costantin & Gallaud, Ann. Sci. Nat. Bot. 9, 6: 354 (1907).
164. Baseonema Schltr. & Rendle, J. Bot. 34: 97 (1896).
165. Batesanthus N.E. Br., Hook. Icon. Pl. 5: ad t. 2500 (1896).
166. Buckollia Venter & R.L. Verh., S. Afr. J. Bot. 60: 97 (1994).
167. Camptocarpus Decne. in DC. & A. DC., Prodr. 8: 493 (1844). nom. cons.
168. Cryptolepis R. Br., Asclepiadeae: 58 (1810).
169. Cryptostegia R. Br., Bot. Reg. 5: t. 435 (1820).
170. Decalepis Wight & Arn. in Wight, Contr. Bot. India: 64 (1834).
171. Ectadium E. Mey., Comm. Pl. Afr. Austr.: 188 (1838).
172. Epistemma D.V. Field & J.B. Hall, Kew Bull. 37: 117 (1982).
173. Finlaysonia Wall., Pl. Asiat. Rar. 2: 48, t. 162 (1831).
174. Gymnanthera R. Br., Asclepiadeae: 47 (1810).
175. Hemidesmus R. Br., Asclepiadeae: 45 (1810).
176. Ischnolepis Jum. & H. Perrier, Rev. Gén. Bot. 21: 53 (1909).
177. Kappia Venter, A.P. Dold & R.L. Verh., S. Afr. J. Bot. 72: 530 (2006).
178. Maclaudia Venter & R. L. Verh., Bot. J. Linn. Soc. 115: 58 (1994).
179. Mondia Skeels, U.S.D.A. Bur. Pl. Industr. Bull. 223: 45 (1911).
180. Myriopteron Griff., Calcutta J. Nat. Hist. 4: 385 (1844).
181. Parquetina Baill., Bull. Mens. Soc. Linn. Paris 2: 806 (1889).
182. Pentopetia Decne. in DC. & A. DC., Prodr. 8: 500 (1844).
183. Periploca L., Sp. Pl.: 211 (1753).
184. Petopentia Bullock, Kew Bull. 1954: 362 (1954).
185. Phyllanthera Blume, Bijdr.: 1048 (1827).
186. Raphionacme Harv., London J. Bot. 1: 22 (1842).
187. Sacleuxia Baill., Hist. Pl. 10: 265 (1890).
188. Sarcorrhiza Bullock, Hooker's Icon. Pl. 36: ad t. 3585 (1962).
189. Schlechterella K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. Index 2–4: 462 (1899).
190. Stomatostemma N.E. Br. in Dyer, Fl. Trop. Afr. 4(1): 252 (1902).
ENDRESS ET AL.186 Phytotaxa 159 (3) © 2014 Magnolia Press
191. Streptocaulon Wight & Arn. in Wight, Contr. Bot. India: 64 (1834).
192. Tacazzea Decne. in DC. & A. DC., Prodr. 8: 492 (1844).
193. Telectadium Baill., Bull. Mens. Soc. Linn. Paris 2: 801 (1889).
194. Zygostelma Benth. in Benth. & Hook.f., Gen. Pl. 2: 740 (1876).
IV. SUBFAMLY SECAMONOIDEAE Endl., Gen. Pl.: 589 (1838).T: Secamone R. Br. (1810). (8 genera)
[Including Secamoneae G. Don, Gen. Hist. 4: 109, 159 (1837–1838); Toxocarpeae Y. Tsiang & P.T. Li, Fl. Reipubl.
Popularis Sin. 63: 280 (1977).]
195. Calyptranthera Klack., Novon 6: 27 (1996).
196. Genianthus Hook. f., Fl. Brit. Ind. 4: 15 (1883).
197. Goniostemma Wight, Contr. Bot. India: 62 (1834).
198. Pervillaea Decne. in DC. & A. DC., Prodr. 8: 613 (1844).
199. Secamone R. Br., Prodr.: 464 (1810).
200. Secamonopsis Jum., Compt. Rend. Hebd. Séances Acad. Sci. 147: 689 (1908).
201. Toxocarpus Wight & Arn. in Wight, Contr. Bot. India: 61 (1834).
202. Trichosandra Decne. in DC. & A. DC., Prodr. 8: 625 (1844).
V. SUBFAMILY ASCLEPIADOIDEAE R. Br. ex Burnett, Outlines Bot.: 1012, 1095, 1103 (1835).—T:
Asclepias L. (1753). (164 genera) [Including Cynanchoideae Luerss., Handb. Syst. Bot. 2: 1064 (1882);
Marsdenioideae Luerss., Handb. Syst. Bot. 2: 1066 (1882).]
1. Tribe Fockeeae H. Kunze, Meve & Liede, Taxon 43: 373 (1994).—T: Fockea Endl. (1839). (2 genera)
203. Cibirhiza Bruyns, Notes Roy. Bot. Gard. Edinb. 45: 51 (1988).
204. Fockea Endl. in Endl. & Fenzl, Nov. Stirp. Dec.: 17 (1839).
2. Tribe Eustegieae Rchb. ex Liede & Meve, trib. nov.Rchb., Deut. Bot. Herb.-Buch: 129 (1841). Validated
here and description amended —T: Eustegia R. Br. (1810). (2 genera) (Supported by Rapini et al. 2007).
—Herbs with clear latex. Leaves opposite to alternate, palmately lobed or (at least some) with one or two spreading
lateral teeth a little above the base; inflorescences extra-axillary and terminal; corolla rotate, glabrous; corona
gynostegial, consisting of a very basally fused ring of simple staminal and deeply tripartite interstaminal parts with
an additional inner ring of smaller staminal lobes; gynostegium (sub-)sessile.
205. Emicocarpus K. Schum. & Schltr., Bot. Jahrb. Syst. 29, Beibl. 66: 21 (1900).
206. Eustegia R. Br., Asclepiadeae: 40 (1810).
3. Tribe Marsdenieae Benth., Fl. Austral. 4: 325, 333 (1868).—T: Marsdenia R. Br. (1810), nom. cons. (26
genera)
207. Anatropanthus Schltr., Bot. Jahrb. Syst. 40, Beibl. 92: 18 (1908).
208. Anisopus N. E. Br., Bull. Misc. Inform. 1895: 259 (1895).
209. Asterostemma Decne., Ann. Sci. Nat. Bot., II, 9: 371, t.10, f. D (1838).
210. Campestigma Pierre ex Costantin in Lecomte, Fl. Indo-Chine 4: 117 (1912).
211. Cathetostemma Blume, Rumphia 4: 30 (1849).
212. Cionura Griseb., Spicil. Fl. Rumel. 2: 69 (1844).
213. Cosmostigma Wight, Contr. Bot. India: 41 (1834).
214. Dischidia R. Br., Prodr.: 461 (1810).
215. Dolichopetalum Tsiang, Acta Bot. Sin. 15: 137 (1973).
216. Gongronema (Endl.) Decne. in DC. & A. DC., Prodr. 8: 624 (1844).
217. Gunnessia P.I. Forst., Austrobaileya 3: 282 (1990).
218. Gymnema R. Br., Prodr.: 461 (1810).
219. Heynella Backer, Blumea 6: 381 (1950).
220. Hoya R. Br., Prodr.: 459 (1810).
221. Jasminanthes Blume, Ann. Mus. Bot. Lugduno-Batavum 1: 148 (1850).
222. Lygisma Hook. f., Hooker's Icon. Pl. 15: ad t. 1423 (1883).
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223. Marsdenia R. Br., Prodr.: 460 (1810), nom. cons. (1810).
224. Oreosparte Schltr., Beih. Bot. Centralbl., Abt. 2, 34: 16 (1916).
225. Pycnorhachis Benth. in Benth. & Hook.f., Gen. Pl. 2: 737, 776 (1876).
226. Rhyssolobium E. Mey., Comm. Pl. Afr. Austr.: 217 (1838).
227. Sarcolobus R. Br., Asclepiadeae: 23 (1810).
228. Stephanotis Thouars, Gen. Nov. Madagasc.: 11 (1806).
229. Stigmatorhynchus Schltr., Bot. Jahrb. Syst. 51: 141 (1913).
230. Tel o s m a Coville, Contr. U.S. Natl. Herb. 9: 384 (1905).
231. Treutlera Hook. f., Hooker's Icon. Pl. 15: t.1425 (1883).
232. Wattakaka Hassk., Flora 40: 99 (1857).
4. Tribe Ceropegieae Decne. ex Orb., Dict. Univ. Hist. Nat. 3: 339 (1843). T: Ceropegia L. (1753). (47 genera)
4.1. Subtribe Heterostemminae Meve & Liede, Taxon 53: 71 (2004).—T: Heterostemma Wight & Arn. (1834).
233. Heterostemma Wight & Arn. in Wight, Contr. Bot. India: 42 (1834).
4.2. Subtribe Leptadeniinae Meve & Liede, Taxon 53: 70 (2004).—T: Leptadenia R. Br. (1810).
234. Conomitra Fenzl in Endl. & Fenzl, Nov. Stirp. Dec.: 65 (1839).
235. Leptadenia R. Br., Asclepiadeae: 23 (1810).
236. Orthanthera Wight, Contr. Bot. India: 48 (1834).
237. Pentasachme Wall. ex Wight, Contr. Bot. India: 60 (1834).
4.3. Subtribe Anisotominae Meve & Liede, Taxon 53: 70 (2004).—T: Anisotoma Fenzl (1844).
238. Anisotoma Fenzl, Linnaea 17: 330 (1844).
239. Emplectanthus N.E. Br. in Dyer, Fl. Cap. 4(1): 771 (1908).
240. Neoschumannia Schltr., Bot. Jahrb. Syst. 38: 38 (1905).
241. Riocreuxia Decne. in A. DC, Prodr. 8: 640 (1844).
242. Sisyranthus E. Mey., Comm. Pl. Afr. Austr.: 197 (1838)
4.4. Subtribe Stapeliinae G. Don, Gen. Hist. 4: 106, 109 (1837-1838).—T: Stapelia L., nom. cons. (1753).
[Including Ceropegiinae Decne. ex Miq., Fl. Ned. Ind. 2: 501. 20 Aug 1857]
243. Anomalluma Plowes, Cact. Succ. J. (Los Angeles) 65: 167 (1993).
244. Apteranthes J.C. Mikan, Nova Acta Acad. Caes. Leop.-Carol. Nat. 17: 594, t. 41. (1835).
245. Australluma Plowes, Haseltonia 3: 54 (1995).
246. Baynesia Bruyns, Novon 10: 354 (2000).
247. Boucerosia Wight & Arn. in Wight, Contr. Bot. India: 34 (1834).
248. Brachystelma Sims, Bot. Mag. 49: ad t. 2343 (1822).
249. Caralluma R. Br., Asclepiadeae: 14 (1810).
250. Caudanthera Plowes, Haseltonia 3: 58 (1995).
251. Ceropegia L., Sp. Pl.: 211 (1753).
252. Desmidorchis Ehrenb., Abh. Königl. Akad. Wiss. Berlin 1829: 31, 39 (1832).
253. Duvalia Haw., Syn. Pl. Succ.: 44 (1812).
254. Duvaliandra M.G. Gilbert, Cact. Succ. J. Gr. Brit. 42: 101 (1980).
255. Echidnopsis Hook. f., Bot. Mag.: ad t. 5930 (1871).
256. Edithcolea N.E. Br., Bull. Misc. Inform. Kew 1895: 220 (1895).
257. Hoodia Sweet ex Decne. Prodr. 8: 664 (1844).
258. Huernia R. Br., Asclepiadeae: 11 (1810)
259. Larryleachia Plowes, Excelsa 17: 5 (1996).
260. Lavrania Plowes, Cact. Succ. J. (Los Angeles) 58: 122 (1986).
261. Monolluma Plowes, Haseltonia 3: 64 (1995).
262. Notechidnopsis Lavranos & Bleck, Cact. Succ. J. (Los Angeles) 57: 255 (1985).
263. Ophionella Bruyns, Cact. Succ. J. Gr. Brit. 43: 70 (1981).
ENDRESS ET AL.188 Phytotaxa 159 (3) © 2014 Magnolia Press
264. Orbea Haw., Syn. Pl. Succ.: 37 (1812).
265. Orbeanthus L.C. Leach, Excelsa Tax. Ser. 1: 71 (1978).
266. Pectinaria Haw., Suppl. Pl. Succ.: 14 (1819), nom. cons., non Pectinaria Bernh. (1800), Apiaceae, nom. rej.
267. Piaranthus R. Br., Asclepiadeae: 12 (1810).
268. Pseudolithos P.R.O. Bally, Candollea 20: 41 (1965).
269. Quaqua N.E. Br., Gard. Chron., II, 12: 8 (1879).
270. Rhytidocaulon P.R.O. Bally, Candollea 18: 335 (1962).
271. Richtersveldia Meve & Liede, Pl. Syst. Evol. 234: 204 (2002).
272. Socotrella Bruyns & A.G. Miller, Novon 12: 330 (2002).
273. Stapelia L., Sp. Pl.: 217 (1753), nom. cons.
274. Stapelianthus Choux ex White & Sloane, Stapelieae: 71 (1933).
275. Stapeliopsis Pillans, S. African Gard. 18: 32 (1928).
276. Tavaresia Welw., Ann. Cons. Ultramarino, ser. 1: 79 (1854).
277. Tridentea Haw., Syn. Pl. Succ.: 34 (1812).
278. Tromotriche Haw., Syn. Pl. Succ.: 36 (1812).
279. White-sloanea Chiov., Malpighia 34: 541 (1937).
5. Tribe Asclepiadeae Duby, Bot. Gall. 1: 323 (1828).—T: Asclepias L. (1753). (87 genera) [Including
Cynancheae Endl., Gen. Pl.: 589 (1838); Gonolobeae G. Don, Gen. Hist. 4: 107, 136 (1837-1838); Oxypetaleae
Orb., Dict. Univ. Hist. Nat. 9: 370 (1847), Tylophoreae Engl., Syllabus, ed. 2: 174 (1898); Orthosieae Malme, Ark.
Bot. 21A: 7. 31 (1927); Vincetoxiceae Small, Man. S.E. Fl.: 1064 (1933).]
5.1. Subtribe Astephaninae Endl. ex Meisn., Pl. Vasc. Gen.: Tab. Diagn. 257 et 266, Comm.: 174 (1840).—T:
Astephanus R. Br. (3 genera)
280. Astephanus R. Br., Asclepiadeae: 43 (1810).
281. Microloma R. Br., Asclepiadeae: 42 (1810).
282. Oncinema Arn., Edinburgh New Philos. J. 17: 261 (1834).
5.2. Subtribe Asclepiadinae Decne. ex Miq., Fl. Ned. Ind. 2: 484 (1857) (Asclepiadeae).—T: Asclepias L.
(1753). (25 genera) [Including Pergularieae Endl., Gen. Pl.: 595 (1838); Calotropidinae Endl. ex Meisn., Pl. Vasc.
Gen.: Tab. Diagn. 257, 268, Comm. 175 (1840).]
283. Aidomene Stopp, Bot. Jahrb. Syst. 87: 21 (1967).
284. Asclepias L., Sp. Pl.: 214 (1753).
285. Aspidoglossum E. Mey., Comm. Pl. Afr. Austr.: 200 (1838).
286. Aspidonepsis Nicholas & Goyder, Bothalia 22: 24 (1992).
287. Calciphila Liede & Meve, Novon 16: 369 (2006).
288. Calotropis R. Br., Asclepiadeae: 28 (1810).
289. Cordylogyne E. Mey., Comm. Pl. Afr. Austr.: 218 (1838).
290. Fanninia Harv., Gen. S. Afr. Pl., ed. 2: 235 (1868).
291. Glossostelma Schltr., J. Bot. 33: 321 (1895).
292. Gomphocarpus R. Br., Asclepiadeae: 26 (1810) .
293. Kanahia R. Br., Asclepiadeae: 28 (1810).
294. Margaretta Oliv., Trans. Linn. Soc. London 29: 111 (1875).
295. Miraglossum Kupicha, Kew Bull. 38: 625 (1984).
296. Odontostelma Rendle, J. Bot. 32: 161, t. 344 (1894).
297. Oxystelma R. Br., Prodr.: 462 (1810).
298. Pachycarpus E. Mey., Comm. Pl. Afr. Austr.: 209 (1838).
299. Parapodium E. Mey., Comm. Pl. Afr. Austr.: 221 (1838).
300. Pergularia L., Syst. Nat., ed. 12, 2: 135, 191 & Mant. Pl.: 8, 53 (1767).
301. Schizoglossum E. Mey., Comm. Pl. Afr. Austr.: 218 (1838).
302. Solenostemma Hayne, Getreue Darstell. Gew. 9: ad t. 38 (1825).
303. Stathmostelma K. Schum., Bot. Jahrb. Syst. 17: 129 (1893).
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304. Stenostelma Schltr., Bot. Jahrb. Syst. 18, Beibl. 45: 6 (1894).
305. Trachycalymma (K. Schum.) Bullock, Kew. Bull. 1953: 348 (1953).
306. Woo d i a Schltr., Bot. Jahrb. Syst. 18. Beibl. 45: 30 (1894).
307. Xysmalobium R. Br., Asclepiadeae: 27 (1810).
5.3. Subtribe Cynanchinae K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 209, 245 (1895).—T:
Cynanchum L. (1753). (13 genera) [Including Sarcostemminae Decne. ex Miq., Fl. Ned. Ind. 2: 480 (1857);
Glossonematinae K. Schum in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 209, 225 (1895).
308. Adelostemma Hook. f., Hooker's Icon. Pl. 15: 22, t. 1427 (1883).
309. Cynanchum L., Sp. Pl.: 212 (1753).
310. Glossonema Decne., Ann. Sci. Nat. Bot., II, 9: 335 (1838).
311. Graphistemma (Champ. ex Benth.) Champ. ex Benth. in Benth. & Hook.f., Gen. Pl. 2: 760 (1876).
312. Holostemma R. Br., Asclepiadeae: 31 (1810).
313. Mahawoa Schltr., Beih. Bot. Centralbl. 34(2): 2 (1916).
314. Metaplexis R. Br., Asclepiadeae: 37 (1810).
315. Odontanthera Wight, Madras J. Lit. Sci. 7: 143 (1838).
316. Pentarrhinum E. Mey., Comm. Pl. Afr. Austr.: 199 (1838).
317. Raphistemma Wall., Pl. Asiat. Rar. 2: 50, t. 163 (1831).
318. Schizostephanus Hochst. ex Benth. in Benth. & Hook.f., Gen. Pl. 2: 762 (1876).
319. Seshagiria Ansari & Hemadri, Indian Forester 97: 126 (1971).
320. Sichuania M.G. Gilbert & P.T. Li, Novon 5: 12 (1995).
5.4. Subtribe Tylophorinae K. Schum. in Engl. & Prantl, Nat. Pflanzenfam. 4(2): 209 (1895).—T: Tylophora R.
Br. (1810). (4 genera)
321. Merrillanthus Chun & Tsiang, Sunyatsenia 6: 105, f. 5, t. 17–18 (1941).
322. Pentastelma Tsiang & P.T. Li in Chun & al., Fl. Hainan. 3: 577 (1974).
323. Pentatropis R. Br. ex Wight & Arn. in Wight, Contr. Bot. India: 52 (1834).
324. Vincetoxicum Wolf, Gen. Pl.: 130 (1776).
5.5. Subtribe Pentacyphinae Liede & Meve, subtrib. nov.T: Pentacyphus Schltr. (1906).
(Supported by Liede-Schumann et al. 2005). (1 genus)
Weakly twining subshrubs with white latex; leaves opposite; inflorescences sciadioidal; flowers
nodding; corolla campanulate, adnate to gynostegium; gynostegial corona of free staminal lobes with
additional peg-shaped part along filament tube; pollinia medifixed or (sub-)basally inserted on
corpusculum; mericarps solitary, fusiform.
325. Pentacyphus Schltr., Bot. Jahrb. Syst. 37: 605, f. 3 (1906).
5.6. Subtribe Diplolepinae Liede & Meve, subtrib. nov.T: Diplolepis R. Br. (1810).
(Supported by Liede-Schumann et al. 2005 and by Hechem et al. 2011). (1 genus)
– Suffrutescent twiners, shrublets or shrubs, with white latex; leaves opposite; inflorescences sciadioidal; corolla
rotate, campanulate or urceolate; corona gynostegial, sometimes vestigial or absent; gynostegium sessile; pollinia
ovoid to oblongoid, laterally attached to horizontal, flattened caudicles; style-head often rostrate; mericarps
solitary, usually narrowly obclavate; seeds usually ovate, winged.
326. Diplolepis R. Br., Asclepiadeae: 30 (1810).
5.7. Subtribe Orthosiinae Liede & Rapini, Syst. Bot. 30: 191 (2005).—T: Orthosia Decne. (1844). (4 genera)
327. Jobinia E. Fourn. in Mart., Fl. Bras. 6(4): 327, t. 97 (1885).
328. Monsanima Liede & Meve, Ann. Missouri Bot. Gard. 99: 66 (2013).
329. Orthosia Decne. in DC. & A. DC., Prodr. 8: 526 (1844).
330. Scyphostelma Baill., Hist. Pl. 10: 252 (1890).
ENDRESS ET AL.190 Phytotaxa 159 (3) © 2014 Magnolia Press
5.8. Subtribe Metastelmatinae Endl. ex Meisn., Pl. Vasc. Gen.: Tab. Diagn. 257, 267, Comm.: 174 (1840).—T:
Metastelma R. Br. (1810). [Including Ditassinae Endl. ex Meisn., Pl. Vasc. Gen.: Tab. Diagn. 257, 267, Comm. 174
(1840).] (12 genera)
331. Barjonia Decne. in DC. & A. DC., Prodr. 8: 512 (1844).
332. Blepharodon Decne. in DC. & A. DC., Prodr. 8: 603 (1844).
333. Ditassa R. Br., Asclepiadeae: 38 (1810).
334. Hemipogon Decne. in DC. & A. DC., Prodr. 8: 509 (1844).
335. Hypolobus E. Fourn. in Mart., Fl. Bras. 6(4): 311 (1885).
336. Metastelma R. Br., Asclepiadeae: 41 (1810).
337. Minaria T.U.P. Konno & Rapini, Taxon 55: 424 (2006).
338. Nautonia Decne. in DC. & A. DC., Prodr. 8: 509 (1844).
339. Nephradenia Decne. in DC. & A. DC., Prodr. 8: 604 (1844).
340. Peplonia Decne. in DC. & A. DC., Prodr. 8: 545 (1844).
341. Petalostelma E. Fourn. in Mart., Fl. Bras. 6(4): 328, t. 98. (1885).
342. Rhyssostelma Decne. in DC. & A. DC., Prodr. 8: 590 (1844).
5.9. Subtribe Tassadiinae Liede & Meve, subtrib. nov.T: Tassadia Decne. (1844).
(Supported by Liede-Schumann et al. 2005, and Liede-Schumann et al. in press). (2 genera)
– Suffrutescent twiners with white latex; leaves opposite; inflorescences usually axillary, paired, with one
inflorescence shorter (indicating derivation from side branches), often multi-flowered, thyrsoidal; corolla with
trichomes in tube; lobes frequently with trichomes; gynostegial corona a ring of connate staminal and interstaminal
segments; gynostegium usually (sub-)sessile; caudicles cylindrical; mericarps paired, narrowly oblong; seeds
oblong.
343. Stenomeria Turcz., Bull. Soc. Imp. Naturalistes Moscou 25: 312 (1852).
344. Tassadia Decne. in DC. & A. DC., Prodr. 8: 579 (1844).
5.10. Subtribe Oxypetalinae E. Fourn. in Martius, Fl. Bras. 6(4): 193 (1878).—T: Oxypetalum R. Br. (1810).
[Including Araujiinae E. Fourn. in Mart., Fl. Bras. 6(4): 193 (1878); Mitostigmatinae E. Fourn. in Mart., Fl. Bras.
6(4): 193 (1878).] (7 genera)
345. Araujia Brot., Trans. Linn. Soc. London 12: 62. t. 4–5 (1817).
346. Funastrum E. Fourn., Ann. Sci. Nat. Bot.4(14): 388 (1882).
347. Morrenia Lindl., Edwards's Bot. Reg. 24, Misc.: 71 (1838).
348. Oxypetalum R. Br., Asclepiadeae: 30 (1810).
349. Philibertia Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. 3 (4th ed.): 195 & ed. fol.: 152 (1818, publ.
1819).
350. Twe edia Hook. & Arn., J. Bot. (Hooker) 1: 291 (1834).
351. Widgrenia Malme, Kongl. Svenska Vetenskapsakad. Handl. II, 34(7): 69, t. 2, f. 4 (1900).
5.11. Subtribe Gonolobinae G. Don ex Liede, Taxon 46: 238 (1997).—T: Gonolobus Michx. (1803). (15 genera)
352. Fischeria DC., Cat. Horti Pl. Monsp.: 112 (1813).
353. Gonolobus Michx., Fl. Bor.-Amer. 1: 119 (1803).
354. Gyrostelma E. Fourn. in Mart., Fl. Bras. 6(4): 302 (1885).
355. Ibatia Decne. in DC. & A. DC., Prodr. 8: 599 (1844).
356. Lachnostoma Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. 3 (4th ed.): 198 (1818, publ. 1819).
357. Macroscepis Kunth in Humb., Bonpl. & Kunth, Nov. Gen. Sp. 3 (4th ed.): 201 et ed. fol.: 156 (1818, publ.
1819).
358. Matelea Aubl., Hist. Pl. Guiane: 277, t. 109, f. 1 (1775).
359. Phaeostemma E. Fourn. in Mart., Fl. Bras. 6(4): 311–312, pl. 91 (1885).
360. Pherotrichis Decne., Ann. Sci. Nat. Bot., II, 9: 322 (1838).
361. Pseudolachnostoma Morillo, Pittieria 36: 44 (2012).
362. Rhytidostemma Morillo, Pittieria 37: 127 (2013).
363. Rojasia Malme, Ark. Bot. 4(14): 10, t. 2, f. 4 (1905).
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364. Schubertia Mart., Nov. Gen. Sp. Pl. 1: 55 (1824), nom. cons., non Schubertia Mirb. (1812), nom. rej.,
Taxodiaceae
365. Stelmagonum Baill., Hist. Pl. 10: 287 (1890).
366. Tylodontia Griseb., Cat. Pl. Cub.: 175 (1866).
Acknowledgements
We are grateful to the reviewers for their keen eye and incisive comments, which greatly improved the paper. In
particular, we would express our sincere thanks James L. Reveal (BH) for his sage advice and for patiently
explaining several of the nomenclatural conundrums we encountered when putting together this paper.
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ENDRESS ET AL.192 Phytotaxa 159 (3) © 2014 Magnolia Press
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Sequence data for the rbcL gene from twenty-four taxa of the families Apocynaceae and Asclepiadaceae were cladistically analysed in order to evaluate the existing familial and subfamilial classification. The taxa sampled represent all described subfamilies and a majority of the described tribes. The cladistic analysis shows that the Asclepiadaceae are nested within the Apocynaceae. An amalgamation of the two families is therefore recommended. The subfamilial classification is also in need of revision: the subfamilies Plumerioideae and Apocynoideae of the current classifications are paraphyletic, as are many of the tribes. Potential subfamily candidates and characters traditionally used in the classification are discussed.
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A revised phylogeny of Tylophorinae, based on 77 species of all genera presently attributed to the subtribe (Biondia, Blyttia, Diplostigma, Goydera, Pentatropis, Pleurostelma, Rhyncharrhena, Tylophora, Vincetoxicum) is presented. Five chloroplast markers (trnT-L, trnL-F, and psbA-trnH intergenic spacers as well as the trnL and trnG introns) and two nuclear markers (ITS, ETS) were used, totalling in 3809 base pairs. Tylophorinae are strongly monophyletic and consist of several well-defined clades. Backbone resolution between these clades is low, indicating a rapid early radiation. Pentatropis is sister to all remaining taxa, albeit with weak support; all other taxa form one big clade not corresponding to previous generic delimitations. Therefore, all genera except for Pentatropis are subsumed under Vincetoxicum. The early branching clades of Vincetoxicum s.l. all grow in Africa, where the group originated ca. 18 Ma ago, correlating with the closure of the Tethys Ocean. The first round of differentiation took place between 12 and 15 Ma ago, resulting in Tylophorinae being distributed over almost all of their present range except for South Africa, Arabia and Europe. Two unrelated lineages of temperate, mostly erect plants hitherto named “Vincetoxicum” have arisen from more tropical lineages, hitherto named “Tylophora ”. One clade of African species is nested inside the Australasian assemblage; this clade contains exclusively polyploid species and its re-immigration to Africa took place in the Messinian, at ca. 6.8 Ma. The European Vincetoxicum species split from erect steppe plants of temperate Asia at ca. 4.5 Ma, coinciding with the uplift of the Tibetan plateau. This group contains two species that are presently aggressively spreading in North America after chance introductions.
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The nuclear gene phytochrome A (PHYA) from 71 species of crown clade Apocynaceae (subfamilies Asclepiadoideae, Secamonoideae, Periplocoideae, and four lineages of Apocynoideae) and outgroups is used to (1) test the chloroplast phylogeny that places the African tribe Baisseeae, with solitary pollen grains, as sister to the pollinia‐bearing milkweeds (Secamonoideae plus Asclepiadoideae); (2) resolve the position of tetrad‐bearing Periplocoideae, the proposed milkweed sister group based on morphology; (3) place the enigmatic Dewevrella; and (4) clarify relationships of the three other primary crown clade lineages: Rhabdadenia, New World clade, and Asian clade. Separate analyses of PHYA and chloroplast sequences agree in placing an African monad‐bearing clade (Baisseeae plus Dewevrella) as the sister group of the milkweeds. Combined PHYA and chloroplast datasets under parsimony and maximum likelihood reject Periplocoideae as the milkweed sister group with statistical significance. Rhabdadenia is placed as sister to the rest of the crown clade, consistent with aspects of its floral morphology and wood anatomy, but without statistical support. Other relationships among the primary crown clade Apocynaceae lineages are also not supported statistically, but power analysis indicates that four to eight times as many characters (26,376–52,752 aligned positions) will be sufficient for a robust estimate.
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Based on morphological, karyological and DNA sequence data the circumscription of Lavrania Plowes, Larryleachia Plowes and Notechidnopsis Lavranos and Bleck is reconsidered. The data presented point to an intermediate position of Notechidnopsis between the more distantly related Larryleachia and Lavrania. Keeping the three genera distinct is the best reflection of the complex relationships within this group.
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Both the tribe Tabernaemontaneae and the genus Tabernaemontana have been much d iscussed concerning delimitation and classification. In the present paper, the pollen morphology of Tabernaemontaneae s.l. is studied with light and scanning electron microscopy (SEM), and compared with the results of a phylogenetic analysis of molecular data. No pollen morphological synapomorphy was found supporting the monophyly of the tribe Tabernaemontaneae s.l. The monophyly of Tabernaemontaneae s.str. is supported by the occurrence of distinctly lalongate endoapertures that might be fused to form an endocingulum. Although the endocingulum does occur in some Old World species, it is interpreted as a synapomorphy of the New World species, which make up a monophyletic group according to molecular data. The inclusion of the genus Stemmadenia in the New World clade within Tabernaemontana is fully supported by pollen morphology. The Old World Tabernaemontana clade is supported by the occurrence of densely perforate pollen with long colpi.
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Analysis of the trnT-trnL spacer, the trnL intron and the trnL-trnF spacer of 65 species of Cynanchum, 33 members of the Metastelminae, and seven representatives of other Asclepiadeae shows that Cynanchum in its present circumscription is polyphyletic. Both the genus Cynanchum and the subtribe Metastelminae s.l. split along the Old World - New World geographic division. The only exception to this division is Cynanchum subgenus Mellichampia, which belongs to Cynanchum s. str. For the Old World Cynanchum relatives, the subtribe Cynanchinae is re-established. The remaining New World sections of Cynanchum are not monophyletic. The South American members of section Cynanchum join the morphologically aberrant Grisebachiella hieronymi. Section Macbridea is associated with Funastrum. In the Old World, the succulent genera Folotsia, Karimbolea, Platykeleba, and Sarcostemma are monophyletic with stem-succulent Cynanchum and nested within a clade of exclusively Malagasy Cynanchum species.