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Vernonieae: Asteraceae) III: Restoration of the Genus Strobocalyx and the New Genus Tarlmounia

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Vernonieae: Asteraceae) III: Restoration of the Genus Strobocalyx and the New Genus Tarlmounia PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 121(1):19–33. 2008. Studies on the Gymnantheminae (Vernonieae: Asteraceae) III: restoration of the genus Strobocalyx and the new genus Tarlmounia Harold Robinson*, Sterling C. Keeley, John J. Skvarla, and Raymund Chan (HR) Department of Botany, NHB 166, National Museum of Natural History, P.O. Box 37012, Smithsonian Institution, Washington, D.C. 20013-7012, e-mail: [email protected]; (SK) Department of Botany, University of Hawaii, Manoa, 3190 Maile Way, #101, Honolulu, Hawaii, 96822-2279, e-mail: [email protected]; (JS) Department of Botany and Microbiology, and Oklahoma Biological Survey, University of Oklahoma, Norman, Oklahoma. 73018-6131, e-mail: [email protected]; (RC) Department of Botany, NHB 166, National Museum of Natural History, P.O. Box 37012, Smithsonian Institution, Washington, D.C. 20013-7012, e-mail: [email protected] Abstract.—The genus Strobocalyx is resurrected from the synonymy of Gymnanthemum and Vernonia, and the genus Tarlmounia is described. Both of these Asian and Malaysian genera have distinct stylar nodes, blunt stylar hairs and specialized tricolporate, echinate pollen. Strobocalyx is typified by the Asian and Indonesian S. arborea, and combinations are provided for the 6 other East Asian and Malaysian species: Vernonia bockiana, V. chunii, V. esculenta, V. solanifolia, V. sylvatica and V. vidalii. Tarlmounia contains only Vernonia elliptica. A continuing series of papers by the sensu Robinson (1999a), with its woody senior author during the last quarter habit and usually imbricated involucre century has segregated many reasonably with rather deciduous inner bracts. The phyletic units from the old, traditional, first of these latest studies resurrected the aphyletic, core genus Vernonia Schreb. Asiatic genus Monosis DC. in Wight and its relatives. The series was summa- (Robinson & Skvarla 2006) which was rized up to a point in two papers, distinct in its leaf shape and its lophate Robinson (1999a), making an initial pollen. This was only a first step in effort at reorganizing genera of the correcting the overly broad concept of Eastern Hemisphere Vernonieae, and Gymnanthemum. The present study and a Robinson (1999b), dealing in more detail simultaneous study of the somewhat with Western Hemisphere members of the similar Asian and Malaysian genus De- tribe. The work is also broadly summa- caneuropsis (Robinson & Skvarla, 2007) rized in less detail in Robinson (2007). further refine the group. These treatments were all based on The present paper deals with the phenetic studies made prior to the use of groups that include two Southeast Asian DNA sequence data. Partly because of species for which Keeley et al. (2007) have these molecular data, further evaluations obtained DNA sequences, Vernonia ar- of some larger genera have begun. The borea Buch.-Ham., a large tree (Fig. 1), present series continues to deal with the and the type of the genus Strobocalyx shrubby or arborescent Gymnanthemum (Blume in DC.) Spach., and Vernonia Cass. in the subtribe Gymnantheminae elliptica DC. in Wight, a straggling shrub or sprawling vine. The sequences indicate * Corresponding author. that the two species are rather close to 20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON each other, but that they are remote from Results the mostly African and Madagascaran genus Gymnanthemum into which V. Two genera are accepted in the present arborea Buch.-Ham. was placed by Ro- treatment, Strobocalyx for V. arborea and binson (1999a). The distinction of both its relatives, and the new genus Tarlmou- species from Gymnanthemum has been nia for V. elliptica. The primary charac- confirmed by study of certain structural teristics on which the genera are based are features, pollen, sweeping hairs of the as follows. style, the basal stylar node, and the Trichomes.—Hairs on the leaves in absence of a well-developed median shield Strobocalyx vary from the long, simple, in the involucral bracts. Revised generic uniseriate form with short basal cells and dispositions of the two species and the a long nonseptate apical cell in Strobo- current understanding of generic limits calyx arborea (Fig. 3A) and S. solanifolia are presented here. The DNA sequence and in other Strobocalyx spp. The latter data (Keeley et al. 2007) offer an extra are sometimes asymmetrical at the base. problem by placing the two Southeast These differ from the very short-stalked, Asian species in close relation to Amer- very long-armed T-shaped form in the ican members of the tribe, a result apparently closely related Tarlmouna el- requiring some discussion. liptica (Fig. 3B). This places Strobocalyx and the related Tarlmounia among the groups in the Vernonieae with simple- or Materials and Methods T-shaped trichomes, in contrast to groups Specimens examined are from the with stellate, spurred, or scale-like tri- U.S. National Herbarium in Washington, chomes such as the New World Piptocar- D.C. Examination of microscopic char- phinae or Sipolisiinae. acters involved use of Hoyer’s Solution Inflorescences.—The shape of the in- (Anderson 1954). florescence in Strobocalyx is generally Preparation of pollen for scanning corymbiform with obvious cymose devel- electron microscopy (SEM) consisted of opment at the apex. The shape is not acetolysis (Erdtman 1960) followed by the strictly pyramidal as in Monosis (Robin- osmium–thiocarbohydrazide repeat pro- son & Skvarla, 2006). The length of cedure (Chissoe et al. 1995) and finally the peduncles varies from almost none pulse sputter coating with a gold/palladi- in S. sylvatica to elongate in S. chunii. um (60/40) target (Chissoe & Skvarla The receptacles may or may not have 1996). Examination was with JEOL 880, hairs. In no case does the inflorescnce Leica 440, and Amray 1810 scanning become very dense. The corollas never electron microscopes, all equipped with have elongate slender bases as in Decan- lanthanum hexaboride (LaB6) electron europsis (Robinson & Skvarla, 2007). sources. The microscopes used were at The achenes of Strobocalyx have been the University of Oklahoma and in the traditionally described as 5-ribbed or 5- SEM laboratory of the United States angled, contrasting with the 10-ribbed National Museum of Natural History in condition in most other Gymanthemum- Washington. like Vernonieae of the Eastern Hemi- The results of the DNA study and the sphere. Careful examination shows 10 methods used are reported in Keeley et al. ribs in almost all achenes, but many of (2007). The phyletic tree included here these may be faint or totally suppressed. (Fig. 2) is simplified from the phylogenet- In Tarlmounia the inflorescences are ic tree of the Vernonieae in the latter elongate and thyrsiform, the achenes paper. are 5-ribbed with little or no evidence of VOLUME 121, NUMBER 1 21 Fig. 1. Coauthor Raymund Chan standing next to Strobocalyx arborea (Buch.-Ham.) Sch. Bip. (approx. 20 m tall) in Bukit Timah Nature Reserve, Singapore (Photo by Michele Chan, plant material for study collected by Shawn Lum). extra ribs. The pappus in all species Style.—In the present study of the examined shows some shorter outer Gymnantheminae, distinctions are em- bristles, although these are not in a highly phasized between Strobocalyx, Gym- differentiated series. nanthemum and Monosis, all placed in 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Simplified phylogenetic tree of Vernonieae derived from DNA sequence data from Keeley et al. (2007). Tree shows positions of Strobocalyx Group of Malaysia that is remote from typical Gymnanthemum of Africa with which it was previous associated. Transverse line in middle of page separates primarily Old World Vernonieae (below) from primarily New World Vernonieae (above). Note position of Old World Strobocalyx Group within the primarily New World Vernonieae. Original from which tree is derived (Keeley et al. 2007) Bayesian analysis of combined data set (ITS, ndhF, trnL-F) run for 1 million generations with model TIM + 1 + G, log-likelihood score 220,656.5. Posterior probability values given above line, bootstrap values (1000 replicates) from majority rule consensus NJ tree (minimum evolution) constructed using same model given below line. Lengths of lines altered. Gymnanthemum in Robinson (1999a). of the style, but these are few and the Two of the useful characters involve the node is, at best, scarcely developed. style. In Strobocalyx the base of the style Sweeping hairs on the upper shaft and has a distinct expanded node. This is also branches of the style are blunt in Strobo- true of Monosis. Tarlmounia has a conical calyx and Tarlmounia. In contrast, they style base with a narrow basal annulus of are pointed in Gymnanthemum and Mono- thickened cells. In contrast, typical Gym- sis, and in some other elements that have nanthemum,withG. coloratum (Willd.) H. been placed in the broad concept of Rob. & B. Kahn and G. amygdalinum Gymnanthemum. The blunt hairs can be (Del.) Sch.Bip. ex Walp., has no evident seen in a distorted form using SEM, but basal stylar node. Other species of true they are easily observed under the light Gymnanthemum, such as G. myrianthum microscope. This character is not restrict- (Hook. f.) H. Rob. and G. theophrastifo- ed to Strobocalyx and Tarlmounia, which lium (Schweinf. ex Oliv. & Hiern) H. have nonlophate pollen, but also occurs Rob., have differentiated cells at the base in numerous species of the Eastern VOLUME 121, NUMBER 1 23 Fig. 3. SEM of trichomes from abaxial surfaces of leaf blades. A, Strobocalyx arborea showing simple hairs (Tonkin, Pe´telot 6624,US);B,Tarlmounia elliptica showing short-stalked T-shaped hairs (Sri Lanka, Grierson 1013, US). Hemisphere such as Decaneuropsis cu- have ordinary Type A pollen (see discus- mingiana (Benth.) H. Rob. & Skvarla, sion below). and in most members of the Piptocarphi- Pollen.—Strobocalyx and Tarlmounia nae in the Western Hemisphere which have tricolporate, echinate, nonlophate 24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pollen with perforated tectum continuous layer (Figs.
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