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A Taxonomic Revision of Timonius Subgen. Pseudobobea (Valeton) S

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A Taxonomic Revision of Timonius Subgen. Pseudobobea (Valeton) S MEP Candollea 65-2_. 18.11.10 13:28 Page217 A taxonomic revision of Timonius subgen. Pseudobobea (Valeton) S. P. Darwin (Rubiaceae) Steven P. Darwin Abstract Résumé DARWIN, S. P. (2010). A taxonomic revision of Timonius subgen. DARWIN, S. P. (2010). Révision taxonomique de Timonius subgen. Pseu- Pseudobobea (Valeton) S. P. Darwin (Rubiaceae). Candollea 65: 217-240. dobobea (Valeton) S. P. Darwin (Rubiaceae). Candollea 65: 217-240. In English, English and French abstracts. En anglais, résumés anglais et français. Twenty-one species, distributed from Sulawesi (Indonesia) to Vingt et une espèces, réparties de Sulawesi (Indonésie) à l’Ile Rossel Island (Louisiade Archipelago) through the Moluccas Rossel (archipel de la Louisiade) en passant par les Moluques (Indonesia) and Papua New Guinea, are assigned to Timonius (Indonésie) et la Papouasie-Nouvelle Guinée, ont été assignées subgen. Pseudobobea (Valeton) S. P. Darwin, a new taxon based à Timonius subgen. Pseudobobea (Valeton) S. P. Darwin, un on Timonius sect. Pseudobobea Valeton (Rubiaceae, Guet- nouveau taxon basé sur Timonius sect. Pseudobobea Valeton tardeae). Nine new species are described: Timonius akunensis (Rubiaceae, Guettardeae). Neuf nouvelles espèces sont décrites: S. P. Darwin, Timonius bracteoides S. P. Darwin, Timonius Timonius akunensis S. P. Darwin, Timonius bracteoides S. P. carrii S. P. Darwin, Timonius clathratus S. P. Darwin, Timonius Darwin, Timonius carrii S. P. Darwin, Timonius clathratus leptocalyx S. P. Darwin, Timonius moluccanus S. P. Darwin, S. P. Darwin, Timonius leptocalyx S. P. Darwin, Timonius Timonius reflexus S. P. Darwin, Timonius repertus S. P. Darwin, moluccanus S. P. Darwin, Timonius reflexus S. P. Darwin, and Timonius rosselensis S. P. Darwin. Species of the new sub- Timonius repertus S. P. Darwin et Timonius rosselensis S. P. genus are distinctive in having usually vertically oriented fruit Darwin. Les espèces du nouveau sous-genre se distinguent pyrenes, reflexed calyx lobes, and leaf blades with corivate vena- par leurs pyrènes de fruits habituellement verticales, leurs tion. Lectotypes are designed for the names Timonius argenteus lobes du calice recurvés, et leurs feuilles à nervation corivée. Valeton, Timonius merokensis Wernham, Timonius minahassae Un lectotype est désigné pour les noms Timonius argenteus Koord., Timonius papuanus Merr., Timonius schefferi Valeton, Valeton, Timonius merokensis Wernham, Timonius mina- Timonius schumannii Koord., and Timonius versteegii Valeton. hassae Koord., Timonius papuanus Merr., Timonius schefferi Neotypes are designated for the names Timonius branderhorstii Valeton, Timonius schumannii Koord. et Timonius versteegii Valeton, and Timonius grandifolius Valeton. A taxonomic Valeton. Un néotype est désigné pour les noms Timonius key to all species is provided as well as information on their branderhorstii Valeton et Timonius grandifolius Valeton. Une ecology and distribution. clé de détermination et des informations sur l’écologie et la distribution de chaque espèce sont fournies. Key-words RUBIACEAE – GUETTARDEAE – Timonius – Papuasia New Guinea – Indonesia – Systematics – Taxonomy Address of the author: Department of Ecology and Evolutionary Biology, Tulane University, New Orleans, Louisiana 70118, U.S.A. Email: [email protected] Submitted on June 2, 2009. Accepted on August 31, 2010. Edited by P. Bungener ISSN: 0373-2967 Candollea 65(2): 217-240 (2010) © CONSERVATOIRE ET JARDIN BOTANIQUES DE GENÈVE 2010 MEP Candollea 65-2_. 18.11.10 13:28 Page218 218 – Candollea 65, 2010 Introduction A recent study by ROVA & al. (2009), based on ITS, rps16, and trnL-F sequence data, suggests that Hodgkinsonia F. Muell. Timonius DC., with perhaps as many as 200 species, is the (two species in tropical Australia) may be sister to Timonius largest of fourteen genera assignable to Rubiaceae tribe Guet- (represented in their study only by T. nitidus (DC.) Fern.-Vill.). tardeae (BREMER & ERIKSSON, 2009), and one of the twenty Hodgkinsonia morphology does suggests a close relationship largest genera of the family (DAVIS & al., 2009). Timonius is with Timonius: dioecy, valvate corolla aestivation, short sta- strictly paleotropical in distribution, extending from Malesia men filaments and dorsifixed anthers, branched stigmas, and westward to Sri Lanka and the Seychelles, northward to the drupaceous (though only bilocular) fruits (BREMER, 1992). Philippines, Micronesia, and Taiwan, and southeastward to A phylogenetic study by MOYNIHAN & WATSON (2001), tropical Australia and the South Pacific as far as the Tuamotu also based on ITS sequence data, suggested that the neotropical Islands. Centers of species diversity are the Malay Peninsula genus Neolaugeria Nicolson may share a more recent common (WONG, 1988), Borneo (PUFF & WONG, 1993), the Philippines ancestor with Timonius than with any New World Guettardeae. (MERRILL, 1923), and Papuasia (Sulawesi eastward to the However, support for a Neolaugeria-Timonius sister relation- Solomon Islands). ship was admittedly weak, and the subsequent analysis by Several detailed descriptions of Timonius have been pub- ACHILLE & al. (2006) placed Neolaugeria in a basal position lished (VALETON, 1909, 1927; DARWIN, 1993, 1994), but the relative to a clade that included the entire Paleotropical Dio ecious salient generic features may be summarized as follows: Clade. Affinity between Timonius and neotropical Guettardeae Dioecious woody plants, sometimes epiphytes or hemiepi- likewise was suggested by BREMER & ERIKSSON (2009). In their phytes, sometimes myrmecophytes. Leaves often with unusual molecular analysis of representative Rubiaceae, two species venation (e.g., paxillate, corivate), and with crypt- or pit-like of Timonius, along with Guettarda crispiflora Vahl (ϵ Tourne- domatia in the abaxial axils of lateral nerves. Inflorescences fortiopsis crispiflora (Vahl) Borhidi), were united in a strongly axillary, cymose, often conspicuously bracteate, several- to supported clade (Bayesian analysis). many-flowered with the flowers secund on the inflorescence arms, or (pistillate plants) reduced to a solitary flower. Calyces toothed, lobed, or truncate, and persistent in fruit. Corollas Infrageneric classification of Timonius infundibular to salverform, the lobes valvate (their edges The above summary of published phylogenies indicates that pressed together and interlocking) and adaxially invested a much more wide-ranging molecular analysis of Guettardeae with a thickened callus. Stamens (or staminodia) as many genera and species is needed in order to resolve the phyloge- as the corolla lobes and inserted on the corolla tube, the fila- netic relationships of Timonius. While published studies do not ments short, the anthers included. Ovary several- to many- contradict a hypothesis of Timonius monophyly, none has inves- loculed, the styles of pistillate flowers with several usually tigated more than a handful of Timonius species. It is unlikely unequal stigmatic arms, the pistillodium of staminate flowers that future analyses will be able to sample even a majority of unbranched or bifid. Fruits drupaceous and containing several the 200 or so species of this genus. Thus, the circumscription of to many free pyrenes, the ovules solitary and pendulous in each infrageneric taxa based on presumed synapomorphies (e.g., pyrene. unusual leaf architecture; reduction of flower number in pistil- The evolutionary affinities of Timonius are only now being late inflorescences; multiplication of ovary locules and pyrenes; elucidated. A phylogeny of selected species of Guettarda L., valvate, calyptrate, or other specialized stipule aestivation) published by ACHILLE & al. (2006) and based on nrDNA may usefully guide future taxon sampling for the purpose ITS sequence data, included five Timonius species. While of reconstructing phylogeny and developing hypotheses of Guettarda was shown to be polyphyletic, the monophyly of character evolution and adaptation. The carving out of a few Timonius was strongly supported in a Bayesian analysis. In well-characterized species-groups (formally named and ranked, that study, the Timonius clade was sister to a clade comprised or not) will leave a number of unassignable species which, of five “Guettarda” species (all from New Caledonia) and presumably, have retained relatively more plesiomorphic traits, two species of “Antirhea” (from Fiji and New Caledonia); and may prove to occupy basal positions in the phylogenetic the sister taxon to the “Antirhea”-“Guettarda”-Timonius clade history of the genus. was Antirhea borbonica J. F. Gmel. (Mascarene Islands Thus far, two subgenera of Timonius have been described: and Madagascar). Interestingly, all of the above taxa are subgen. Timonius, with eight species in Java, the Moluccas, dioecious, and were designated by ACHILLE & al. (2006) New Guinea, tropical Australia, and the Solomon Islands the “Paleotropical Dioecious Clade” of Guettardeae. A close (DARWIN, 1993); and subgen. Abbottia S. P. Darwin, with 29 relationship between Timonius and the Hawaiian genus Bobea species distributed from eastern Borneo through Papuasia to Gaudich., as was suggested by CHAW & DARWIN (1992), was the Philippines, tropical Australia, Fiji, and Samoa (DARWIN, not supported. 1994). Another (unranked) species group allied with Timonius MEP Candollea 65-2_. 18.11.10 13:28 Page219 A taxonomic revision of Timonius subgen. Pseudobobea (Valeton) S. P. Darwin (Rubiaceae) – 219 flavescens (Jack) Baker was described from Papuasia, but Size and distribution.
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