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The End of Pseudodracontium N.E. Br 40 AROIDEANA, Vol. 35 The End of Pseudodracontium N.E. Br. Wilbert L. A. Hetterscheid Von Gimborn Arboretum Velperengh 13 3941 BZ Doorn, The Netherlands [email protected] (*31)318418187 Cyrille Claudel Biocentre Klein Flottbek and Botanical Garden Hamburg University Ohnhorststrasse 18 22609 Hamburg, Germany ABSTRACT soon be transferred to Amorphophallus, this new species, clearly a member of The aroid genus Pseudodracontium N.E. ‘‘Pseudodracontium’’, was published as a Br. is reduced to Amorphophallus Bl. ex species of Amorphophallus. Decne. New names and a new key to taxa Pseudodracontium has always been of the former Pseudodracontium are pre- thought to be very closely related to sented. A discussion on its phylogenetic Amorphophallus and both genera solely position within Amorphophallus is given. make up the tribe Thomsonieae. Thomso- nieae is now regarded to be the sister group KEY WORDS of Caladieae (incl. Zomicarpeae; Cabrera et al., 2008; Cusimano et al., 2011). Amorphophallus, Pseudodracontium,tax- onomy, Thomsonieae, Araceae, phylogeny. PSEUDODRACONTIUM AND AMORPHOPHALLUS INTRODUCTION Hetterscheid (1994) already suggested that The genus Pseudodracontium was es- Pseudodracontium may well be a part of tablished by N.E. Brown in 1882. He Amorphophallus instead of its sister genus. introduced one new species, P. anomalum This was based on morphological observa- N.E. Br., and transferred Amorphophallus tions which resulted in a small suite of lacourii Lind. & Andre´toPseudodracon- apomorphies for Amorphophallus + Pseudo- tium. The first mentioned species was dracontium, whereas without Pseudodra- chosen as the lectotype species by Nicolson contium there was no single character to be (1967). Subsequent years saw the addition found for a monophyletic Amorphophallus of several new names in Pseudodracon- (Hetterscheid, 1994; Serebryanyi, 1995). tium. These were dealt with taxonomically The monophyly of Pseudodracontium in a revision of the genus by Serebryanyi itself was never disputed and strongly (1995) which left a total of 6 accepted supported by a suite of morphological species, 4 of which were newly introduced (near-) autapomorphies (Serebryanyi, 1995; in that revision. After the revision only one Van der Ham & Van Heuven, 2001; Van new name was published in relation to der Ham et al., 1998, 2000, 2005). The most Pseudodracontium, namely Amorphophal- obvious ones are: unilocular thecae, slender lus glaucophyllus Hett. & Serebryanyi filaments (fully free or more or less connate (Hetterscheid, 2006): Because it was al- to form a slender column, Fig. 1b), appendix ready then strongly suspected by the with recognizable staminodial structure authors that Pseudodracontium would (Fig. 1c), pollen with ‘‘polar caps’’ (smooth Aroideana arod-35-00-05.3d 3/4/12 10:58:43 40 W. L. A. HETTERSCHEID, C. CLAUDEL, 2012 41 Plate 1. 1a. Amorphophallus lacourii group. Habit. 1b. Amorphophallus lacourii group. Male flowers. 1c. Amorphophallus pseudoharmandii Appendix. 1d. Amorphophallus macrophyllus Pollen with polar caps. or irregularly structured polar areas, erably less complex than the lateral ones, fig. 1d). and there is a general morphocline from We have always been puzzled by the more complex to rather simply divided leaf retention of seemingly juvenile leaf archi- laminas (or the reverse). In some speci- tecture in many adult plants of Pseudodra- mens the leaf remains fully juvenile with contium taxa (fig 1a). In almost all taxa the only three, very large, leaflets in adult central main segment of the leaf is consid- plants, as is seen in seedlings, also in many Aroideana arod-35-00-05.3d 3/4/12 10:58:44 41 42 AROIDEANA, Vol. 35 other Amorphophallus species. In nearly all Molecular analyses of Amorphophallus + seedlings ever observed by us (several Pseudodracontium published to date hundred) the first seedling leaf lacks the (Grob et al., 2002, 2004; Sedayu et al., central segment (leaflet) entirely. 2010) strongly support the sister-group The polar caps of the pollen grains appear relationship of Pseudodracontium to the to be a retained non-adult phase of pollen A. longituberosus group within Amorpho- formation. Looking at the inflorescence, we phallus and thus confirms the earlier observe a few characters that may be seen as morphology-based hypotheses. More re- ‘‘plesiomorphic’’ for Amorphophallus at cent molecular analysis (Randomized Ax- large, like the appendix wall with fully elerated Maximum Likelihood [RAxML]) by developed staminodes and the long, free or the second author using the markers ITS1, partly fused filaments. We hypothesize that FLint2, rbcL and matK and applied to ca. the morphogenic pathways of several phe- 130 species of Amorphophallus (incl. Pseu- notypic characters in Pseudodracontium is dodracontium) again strongly supports the deregulated. It may well be that the chaotic aforementioned grouping of Pseudodra- molecular and morphological patterns dis- contium in Amorphophallus (Claudel et al., play an evolutionary phase Pseudodracon- in prep., Hetterscheid & Claudel, in prep.) tium is going through whereby (past and but expands it with a small strongly ongoing) hybridisation (see also paragraph supported clade of 3 (possibly only 2) 3.1) may well have been the onset of the species (A. saraburiensis Gagn., A. scuta- deregulation. Heterochrony seems to be part tus Hett. & T.C. Chapman and A. tenuistylis of this deregulation, leading to adult plants Hett.). This group of the Amorphophallus with partly juvenile morphologies. species mentioned + Pseudodracontium is Pseudodracontium as a group within part of a larger and strongly supported Amorphophallus is considered to be closely clade of the ‘‘Continental Asia-II’’-clade of related to A. longituberosus (Engl.) Engl. & Sedayu et al. (2010). Gehrm. and its immediate allied species A. In the light of all evidence discussed albispathus Hett., A. coudercii (Bogn.) above it is here decided to reduce the Bogn. and A. tenuispadix Hett. This group genus Pseudodracontium to the synonymy turned up again and again in morphology- of Amorphophallus. The necessary new based attempts to unravel the phylogeny of combinations and one new name are Amorphophallus. In these analyses Pseudo- presented below. The tribe Thomsonieae dracontium species were always associat- will become monotypic. ed with the aforementioned group as a sister group. Even in chemical analyses of THE ‘‘SPECIES’’-PROBLEM IN the scent of species in this alliance (Kite & Hetterscheid, 1997; Kite et al., 1998), traces ‘‘PSEUDODRACONTIUM’’ AND A of a compound (4-methoxyphenetyl alco- PROVISIONAL NEW KEY TO THE TAXA hol, or ‘‘anise oil’’) unique to it, were found Species Number in Pseudodracontium in P. lacourii (Lind. & Andre´) N.E. Br. and P. fallax Serebryanyi. This seemed to Serebryanyi (1995) recognized 6 species strengthen the suggested relationship. The in Pseudodracontium and provided an trouble with all full-morphology-based identification key. In subsequent years, phylogeny reconstructions of Amorpho- many more collections of Pseudodracon- phallus was that no statistically relevant tium species were made by many botanists support for this grouping could be found, and studied by the first author morpholog- not even applying the Implied Weighing ically. It turned out that many species- procedure of the TNT program of Goloboff defining character combinations proposed et al. (2004; Hovenkamp & Hetterscheid, by Serebryanyi can no longer be used. 2008). The species group of A. longituber- Many character combinations that were osus reappeared in all analyses but never used to define particular species prove to associated with Pseudodracontium. be more widespread and recombined with Aroideana arod-35-00-05.3d 3/4/12 10:58:51 42 W. L. A. HETTERSCHEID, C. CLAUDEL, 2012 43 other characters to form an ever more Amorphophallus kuznetsovii (Sereb- complex of combinations blurring almost ryanyi) Hett. & C. Claudel, comb. all conventional species borders in the nov. Basionym: Pseudodracontium ‘‘genus’’. kuznetsovii Serebryanyi. Blumea Molecular analyses by the second author 40(1)(1995): 226, fig. 2. 1995 - Type: (Claudel at al., in prep.) also show huge from a plant cultivivated in Hortus overlap in characters between ‘‘taxa’’ as Botanicus Leiden, the Netherlands, 15 well as suggesting extensive hybridization July 1992, Hetterscheid H.AM.165-T (particularly in ITS1 analyses). (holotypus, MHA, spiritcoll.) - orig. As a result there is a strong conviction coll. Vietnam, Xuen Moc Reserve, that the number of species of Pseudodra- easternmost part of the Dong Nai contium is overstated. In the forthcoming province, Kuznetsov s.n., 1991. Amorphophallus treatment in the Flora of Amorphophallus lanceolatus (Sereb- Thailand (Hetterscheid, in press) a key to ryanyi) Hett. & C. Claudel, comb. the taxa of ‘‘Pseudodracontium’’ is present- nov. Basionym: Pseudodracontium ed, which is duplicated here (see below) lanceolatum Serebryanyi, Blumea using the new combinations and new name 40(1)(1995): 230, fig 4a, b. - Type: in Amorphophallus. from a plant cultivivated in Hortus Botanicus Leiden, the Netherlands, 15 Five New Combinations and One New July 1992, Hetterscheid H.AM.179-T Name in Amorphophallus (holotypus, MHA, spiritcoll.) - orig. coll. Vietnam, Xuen Moc Reserve, As a result of the transfer of all Pseudo- dracontium species to Amorphophallus,five easternmost part of the Dong Nai new nomenclatural combinations and one province, Kuznetsov s.n., 1991. new name in Amorphophallus are presented Amorphophallus latifolius (Serebrya-
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