Inge Groeninckx

Phylogeny of the herbaceous tribe Spermacoceae (Rubiaceae) based on plastid DNA data

Abstract In its current circumscription, the herbaceous tribe Spermacoceae s.l. (Rubiaceae, Rubioideae) unites the former tribes Spermacoceae s. str., Manettieae, and the Hedyotis–Oldenlandia group. Within Spermacoceae, and particularly within the Hedyotis–Oldenlandia group, the generic delimitations are problematic. Up until now, molecular studies have focused on specific taxonomic problems within the tribe. This study is the first to address phylogenetic relationships within Spermacoceae from a tribal perspective. Sequences of three plastid markers (atpB-rbcL, rps16, and trnL-trnF) were analyzed separately as well as combined using parsimony and Bayesian approaches. Our results support the expanded tribe Spermacoceae as monophyletic. The former tribe Spermacoceae s. str. forms a monophyletic clade nested within the Hedyotis–Oldenlandia group. Several genera formerly recognized within the Hedyotis–Oldenlandia group are supported as monophyletic (Amphiasma Bremek., Arcytophyllum Willd. ex Schult. & Schult. f., Dentella J. R. Forst. & G. Forst., Kadua Cham. & Schltdl., and Phylohydrax Puff), while others appear to be paraphyletic (e.g., Agathisanthemum Klotzsch), biphyletic (Kohautia Cham. & Schltdl.), or polyphyletic (Hedyotis L. and Oldenlandia L. sensu Bremekamp). Morphological investigations of the taxa are ongoing in order to find support for the many new clades and relationships detected. This study provides a phylogenetic hypothesis with broad sampling across the major lineages of Spermacoceae that can be used to guide future species-level and generic studies.

Evolutionary history of the Afro-Madagascan Ixora species (Rubiaceae): species diversification and distribution of key morphological traits inferred from dated molecular phylogenetic trees

BACKGROUND AND AIMS Previous work on the pantropical genus Ixora has revealed an Afro-Madagascan clade, but as yet no study has focused in detail on the evolutionary history and morphological trends in this group. Here the evolutionary history of Afro-Madagascan Ixora spp. (a clade of approx. 80 taxa) is investigated and the phylogenetic trees compared with several key morphological traits in taxa occurring in Madagascar. METHODS Phylogenetic relationships of Afro-Madagascan Ixora are assessed using sequence data from four plastid regions (petD, rps16, rpoB-trnC and trnL-trnF) and nuclear ribosomal external transcribed spacer (ETS) and internal transcribed spacer (ITS) regions. The phylogenetic distribution of key morphological characters is assessed. Bayesian inference (implemented in BEAST) is used to estimate the temporal origin of Ixora based on fossil evidence. KEY RESULTS Two separate lineages of Madagascan taxa are recovered, one of which is nested in a group of East African taxa. Divergence in Ixora is estimated to have commenced during the mid Miocene, with extensive cladogenesis occurring in the Afro-Madagascan clade during the Pliocene onwards. CONCLUSIONS Both lineages of Madagascan Ixora exhibit morphological innovations that are rare throughout the rest of the genus, including a trend towards pauciflorous inflorescences and a trend towards extreme corolla tube length, suggesting that the same ecological and selective pressures are acting upon taxa from both Madagascan lineages. Novel ecological opportunities resulting from climate-induced habitat fragmentation and corolla tube length diversification are likely to have facilitated species radiation on Madagascar.

A new species of Colletoecema (Rubiaceae) from southern Cameroon with a discussion of relationships among basal Rubioideae

Colletoecema magna, a new species from the Ngovayang Massif (southern Cameroon) is described and illustrated. A comparative morphological study illustrates the similar placentation and fruit anatomy of the novelty and Colletoecema dewevrei, the only other species of the genus. Colletoecema magna essentially differs from C. dewevrei by its sessile flowers and fruits, the corolla tube that is densely hairy above the insertion point of the stamens and the anthers that are included. Further characters that separate the novelty are its larger leaves, more condensed inflorescences, and larger fruits. Its position within Colletoecema is corroborated by atpB-rbcL and rbcL chloroplast sequences. The relationships among the basal lineages of the subfamily Rubioideae, to which Colletoecema belongs, are briefly addressed. Based on our present knowledge, a paleotropical or tropical African origin of the Rubioideae is hypothesized.

Dispersing towards Madagascar: Biogeography and evolution of the Madagascan endemics of the Spermacoceae tribe (Rubiaceae).

Despite the close proximity of the African mainland, dispersal of plant lineages towards Madagascar remains intriguing. The composition of the Madagascan flora is rather mixed and shows besides African representatives, also floral elements of India, Southeast Asia, Australia, and the Neotropics. Due to its proportionally large number of Madagascan endemics, the taxonomically troublesome Spermacoceae tribe is an interesting group to investigate the origin and evolution of the herbaceous Rubiaceae endemic to Madagascar. The phylogenetic position of these endemics were inferred using four plastid gene markers. Age estimates were obtained by expanding the Spermacoceae dataset with representatives of all Rubiaceae tribes. This allowed incorporation of multiple fossil-based calibration points from the Rubiaceae fossil record. Despite the high morphological diversity of the endemic herbaceous Spermacoceae on Madagascar, only two colonization events gave rise to their current diversity. The first clade contains Lathraeocarpa, Phylohydrax and Gomphocalyx, whereas the second Madagascan clade includes the endemic genera Astiella, Phialiphora, Thamnoldenlandia and Amphistemon. The tribe Spermacoceae is estimated to have a Late Eocene origin, and diversified during Oligocene and Miocene. The two Madagascan clades of the tribe originated in the Oligocene and radiated in the Miocene. The origin of the Madagascan Spermacoceae cannot be explained by Gondwanan vicariance but only by means of Cenozoic long distance dispersal events. Interestingly, not only colonization from Africa occurred but also long distance dispersal from the Neotropics shaped the current diversity of the Spermacoceae tribe on Madagascar.

A New Species of Hedythyrsus (Rubiaceae) with Comments on the Generic Delimitations within African Spermacoceae s. l.

Abstract A new Hedythyrsus species from Katanga (D. R. Congo), Hedythyrsus katangensis , is described and illustrated based on classic herbarium taxonomy. The generic placement of the new species was determined by comparing it to African genera of the Hedyotis-Oldenlandia complex of the Spermacoceae s. l. that share the terminal position of the inflorescences and the capsular fruits opening by four valves, i.e. Agathisanthemum, Dibrachionostylus, Hedythyrsus, Mitrasacmopsis, and Pseudonesohedyotis. The new species differs from the known Hedythyrsus species in being geoxylic herbs instead of shrubs, and in having bifid, not multifid, stipules. Within Hedythyrsus katangensis two varieties are recognized. Variety ternifolius is distinguished from the type variety by the position of the leaves (often ternate in var. ternifolius, always opposite in var. katangensis) and the pubescence of stems, ovary, and calyx (scabrous to shortly hairy in var. ternifolius, glabrous in var. katangensis). Following the IUCN red list criteria the species can be classified as near threatened.