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Evolution and biodiversity of the Ixoroideae (Rubiaceae)
Stockholm University, Faculty of Science, Department of Botany.
2010 (English)Doctoral thesis, comprehensive summary (Other academic)
Abstract [en]

The phylogenetic relationships within subfamily Ixoroideae of the coffee family are investigated by phylogenetic reconstruction of molecular data, including regions of the chloroplast DNA (matK, ndhF, rbcL, rps16, trnH-psbA, trnS-G, and trnT-F), and the nuclear ribosomal DNA (ITS). The evolution of morphological characters within the group are inferred, with focus on characters used in classification. Ixoroideae have primarily been characterized by secondary pollen presentation, contorted corolla aestivation, and fleshy fruits. Secondary pollen presentation appears synapomorhic of a clade comprising the Ixoroideae crown group together with Retiniphyllum, whereas contorted corolla aestivation has evolved earlier and is synapomorphic for the crown group, Retiniphyllum, and Steenisia. Capsules likely represent a plesiomorphy from which various dry or fleshy indehiscent fruits have evolved independently in different clades. Reductions in seed number have also occured in many clades, none of which shows a secondary increase in the number of seeds.

Within Ixoroideae, the phylogeny and tribal delimitations of Alberteae and Condamineeae are studied in more detail. The former appears restricted to Alberta, Nematostylis, and Razafimandimbisonia, a new genus described here. The Condamineeae are a diverse tribe largely unresolved in previous molecular phylogenetic studies. Our results support a synonymization of both Calycophylleae and Hippotideae, because these are nested within the Condamineeae. Ancestral state reconstructions indicate that intrapetiolar stipules, poricidal anthers, and protogyny, otherwise uncommon characters in Rubiaceae, all have evolved more than once in the Condamineeae.

The rare genera Jackiopsis, Glionnetia, and Trailliaedoxa previously not included in molecular phylogenetic analyses, are all found nested within the Ixoroideae, and their systematic positions are discussed. The genera Bathysa, Calycophyllum, Elaeagia, and Rustia do not appear monophyletic. Consequently, resurrections of the names Holtonia, Schizocalyx, and Semaphyllanthe, and synonymizations of Phitopis (as Schizocalyx) and Tresanthera (as Rustia) are proposed. Also proposed are five new tribal names for clades that are not associated with any previously described tribes in the phylogenetic hypotheses presented.

Place, publisher, year, edition, pages
Stockholm: Department of Botany, Stockholm University , 2010. , 28 p.
Keyword [en]
Alberteae, character evolution, classification, Condamineeae, fruit evolution, Ixoroideae, phylogenetics, pollen presentation, Rubiaceae, systematics, taxonomy
National Category
Biological Systematics
Research subject
Plant Systematics
Identifiers
URN: urn:nbn:se:su:diva-48850ISBN: 978-91-7447-198-4 (print)OAI: oai:DiVA.org:su-48850DiVA: diva2:376046
Public defence
2011-01-14, föreläsningssalen, Botanicum, Lilla Frescativägen 5, Stockholm, 10:00 (English)
Opponent
Supervisors
Note
At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: Manuscript. Paper 4: Manuscript.Available from: 2010-12-22 Created: 2010-12-09 Last updated: 2010-12-14Bibliographically approved
List of papers
1. Molecular phylogenetic analysis of the tribe Alberteae (Rubiaceae), with description of a new genus, Razafimandimbisonia
Open this publication in new window or tab >>Molecular phylogenetic analysis of the tribe Alberteae (Rubiaceae), with description of a new genus, Razafimandimbisonia
2009 (English)In: Taxon, ISSN 0040-0262, E-ISSN 1996-8175, Vol. 58, no 3, 757-768 p.Article in journal (Refereed) Published
Abstract [en]

The tribe Alberteae, presently classified in the subfamily Ixoroideae (Rubiaceae), has historically been an artificial grouping of genera. In the present study, phylogenetic analyses of the chloroplast DNA markers rbcL, ndhF, trnS-G, trnT-F and trnH-psbA as well as the ITS region of the nuclear ribosomal DNA, are done to assess the delimitation of Alberteae. The resulting phylogenetic hypothesis is highly resolved, with most clades strongly supported. The genus Alberta is found to be paraphyletic as presently circumscribed. As a consequence, we propose the new genus Razafimandimbisonia Kainul. & B. Bremer to accommodate the Malagasy species. The newly delimited Alberta is distinguished by having two calycophylls that expand after anthesis as well as awl-shaped stigma lobes. Razafimandimbisonia is distinguished from the remaining Alberteae by having dehiscent fruits and anthers without basal appendages. We demonstrate that the genera Airosperma, Boholia and Crossopteryx are not associated with Alberteae, as has previously been suggested. Alberteae is considered restricted to the genus Alberta endemic to Southeast Africa, and the two Malagasy endemic genera Nematostylis and Razafimandimbisonia.

Place, publisher, year, edition, pages
Wien: International Association for Plant Taxonomy, 2009
Keyword
Airosperma, Alberta, Boholia, cpDNA, Crossopteryx, Madagascar, Nematostylis, Rubiaceae
National Category
Biological Systematics
Research subject
Systematic Botany
Identifiers
urn:nbn:se:su:diva-35237 (URN)000269774900006 ()
Available from: 2010-01-15 Created: 2010-01-15 Last updated: 2017-12-12
2. Molecular systematics and morphological character evolution of the Condamineeae (Rubiaceae)
Open this publication in new window or tab >>Molecular systematics and morphological character evolution of the Condamineeae (Rubiaceae)
2010 (English)In: American Journal of Botany, ISSN 0002-9122, E-ISSN 1537-2197, Vol. 97, no 12, 1961-1981 p.Article in journal (Refereed) Published
Abstract [en]

• Premise of the study: The Condamineeae have in previous molecular studies been shown to be part of an early-divergent cladewithin the subfamily Ixoroideae, together with the tribes Calycophylleae, and Hippotideae, and genera of the former Cinchoneae and Rondeletieae. Generic relationships within this clade have, however, remained largely unresolved

.• Methods: In this study, the systematics of the Condamineeae was further examined by phylogenetic reconstruction of six cpDNA regions and one nrDNA region using parsimony and Bayesian Markov chain Monte Carlo inference. Morphological character evolution within the tribe was assessed by ancestral state reconstruction using likelihood optimization of characters onto Bayesian trees.

• Key results: Calycophylleae appears polyphyletic. “Hippotideae” is monophyletic but nested within the Condamineeae. The phylogenetic hypotheses presented support a resurrection of the genera Holtonia, Schizocalyx, and Semaphyllanthe. Furthermore, Bathysa is found to be polyphyletic, Tresanthera is found nested within Rustia, and the taxonomically disputed genus Dialypetalanthus is here shown to be sister to a BothriosporaWittmackanthus clade. Morphological ancestral state reconstructions indicate that protogyny have evolved at least two times within the tribe and that indehiscent fruits, loculicidal fruit dehiscence, and intrapetiolar stipules have evolved independently several times. The occurrence of calycophylls (leaf-like calyx lobes), poricidal anthers, and winged seeds also appear homoplastic within the tribe.

• Conclusions : A diagnosis and delimitation of the tribe Condamineeae is presented, with taxonomic proposals to synonymize Tresanthera and to transfer several species of Bathysa as well as Phitopis to a resurrected Schizocalyx.

Keyword
character evolution, Condamineeae, Dialypetalanthus, Hippotideae, Ixoroideae, Rubiaceae, seed morphology, systematics
National Category
Biological Systematics
Research subject
Plant Systematics
Identifiers
urn:nbn:se:su:diva-48797 (URN)10.3732/ajb.1000090 (DOI)000284760800016 ()
Funder
Knut and Alice Wallenberg Foundation
Available from: 2010-12-09 Created: 2010-12-09 Last updated: 2017-12-11Bibliographically approved
3. Molecular support for a basal grade of morphologically peculiar, monotypic genera in the species-rich Vanguerieae alliance (Rubiaceae): its systematic and conservation implications
Open this publication in new window or tab >>Molecular support for a basal grade of morphologically peculiar, monotypic genera in the species-rich Vanguerieae alliance (Rubiaceae): its systematic and conservation implications
Show others...
(English)Manuscript (preprint) (Other academic)
Abstract [en]

The species-rich coffee family (Rubiaceae) contains 211 monotypic genera, representing 34.5% of the total number of rubiaceous genera. We rigorously assessed the generic status and phylogenetic positions of the enigmatic, monotypic genera, the Seychellean Glionnetia, the Southeast Asian Jackiopsis, and the Chinese Trailliaedoxa, in Rubiaceae using sequence data from four plastid markers (ndhF, rbcL, rps16, and trnT-F). The present study provides molecular support for isolated phylogenetic positions of these genera in the subfamily Ixoroideae, and reveals the presence of a basal grade of morphologically odd, monotypic genera (Crossopteryx, Jackiopsis, Scyphiphora, Trailliaedoxa, and Glionnetia, respectively) in the species-rich Vanguerieae alliance. We consider these phylogenetically isolated genera potential candidates for a high conservation priority, as they appear to be sole representatives of their respective lineages and therefore carry unique genetic information and evolutionary history. The monogeneric tribe Jackieae as defined by Ridsdale is supported.

Keyword
Conservation, Crossopteryx, Glionnetia, Jackiopsis, Rubiaceae, Scyphiphora, systematics, Trailliaedoxa
National Category
Biological Systematics
Research subject
Plant Systematics
Identifiers
urn:nbn:se:su:diva-48800 (URN)
Funder
Knut and Alice Wallenberg Foundation
Note
Manuscript part of second authors Thesis in Systematic Botany.Available from: 2010-12-09 Created: 2010-12-09 Last updated: 2010-12-14Bibliographically approved
4. Phylogeny and evolution of secondary pollen presentation, corolla aestivation patterns, fruit types, and seed number in the Ixoroideae (Rubiaceae)
Open this publication in new window or tab >>Phylogeny and evolution of secondary pollen presentation, corolla aestivation patterns, fruit types, and seed number in the Ixoroideae (Rubiaceae)
(English)Manuscript (preprint) (Other academic)
Abstract [en]

Phylogenetic reconstructions using Bayesian and parsimony analyses of six chloroplast DNA region were performed in order to investigate character evolution and tribal relationships within the subfamily Ixoroideae (Rubiaceae). In the inferred phylogenetic hypotheses, the tribal relationships were mostly well supported, with the subfamily comprised of a crown group of two major sister clades termed the Coffeeae-alliance and Vanguerieae-alliance respectively, and a basal grade comprising Condamineeae, Henriquezieae, Posoquerieae, Retiniphyleae, Sipaneeae, and Steenisia. Five new tribes are here recognized (Airospermeae, Augusteae, Scyphiphoreae, Steenisieae and Trailliaedoxeae). Secondary pollen presentation, corolla aestivation patterns,fruit types, and number of ovules are all characters that have been considered of great importance in the classification of Rubiaceae. Ancestral state reconstructions of these characters using likelihood optimization indicate that secondary pollen presentation is synapomorphic for a clade comprising the Ixoroideae crown group and Retiniphyllum, whereas left-contorted corolla aestivation is synapomorphic for a clade comprising the crown group, Retiniphyllum, and Steenisia. Capsular fruits with numerous seeds are plesiomorphic in Ixoroideae, from which dry or fleshy indehiscent fruits have evolved numerous times independently. Reductions in seed number appears to have occurred within several lineages, none of which show a secondary increase in the number of seeds.

Keyword
character evolution, cpDNA, pollen presentation, Rubiaceae, seed number, systematics
National Category
Biological Systematics
Research subject
Plant Systematics
Identifiers
urn:nbn:se:su:diva-48798 (URN)
Funder
Knut and Alice Wallenberg Foundation
Note
Manuscript part of the first authors Thesis in Systematic Botany.Available from: 2010-12-09 Created: 2010-12-09 Last updated: 2010-12-14Bibliographically approved

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