The limit for successful DNA extraction was tested by amplification and sequencing of an over 200-year-old herbarium specimen collected by Adam Afzelius, a student of Carl Linnaeus. We amplified and sequenced a 800-bp region between 16S ribosomal DNA and the 3′ part of the trnI gene (16S-trnI) in the chloroplast genomeof Phaulopsis talbotii S. Moore (Acanthaceae). To test the replicability and to control for contamination the procedure was performed in sealed vials and with negative PCR controls. The procedure was also repeated in a separate laboratory. In addition, the chloroplast rpl16 intron was successfully amplified and sequenced and the rps16 intron amplified. Sequences of taxa closely related to Acanthaceae were found to be most similar to the produced sequences. The results suggest that molecular investigations of other 18th century botanical collections are feasible and that molecular methods could be employed for comparative studies to extant plant collections. An important application would be to identify descendants or clones of Linnaean lectotypes by comparing DNA from these with potentially remnant plants from Linnaeus’ cultivations.

Phylogenctic analyses of taxa that have been included in the tribe Paederieae are performed on the basis of a molecular dataset of plastid DNA sequences (the rbcL gene, rps16 intron, and the regions trnT-F). The paraphyly of Paederieae as currently circumscribed is confirmed. A clade comprising Paederia, Spermadietyon, Leptodermis, and Serissa is retrieved and the circumscription of Paederieae is proposed to be restricted to this clade. Paederia is sister to a clade with Spermadiclyon, Leptodermis, and Serissa. Paederia is subdivided into an Asian and an African clade, and within the African clade seven species endemic to Madagascar are sister to P. pospischilii in north-eastern Africa. The position of Kelloggia as sister to Rubieae is confirmed and a position of Saprosma in the Spermacoceae alliance is supported. The genera Putoria, Gaillonia (with segregates), Plocama, and Aitchisonia form a clade that is sister to Theligoneae/Ke//oggia/Rubieae. The name Putoricae is resurrected for the members of this clade. A synopsis of Putorieae is provided, where all species are placed in an expanded Plocama with 34 species. Thirty-one new combinations in Plocama are proposed. Plocama pendula on the Canary Islands is strongly supported as sister to a clade with P. crocyllis (former Crocyllis) in southern Africa and P. yemenensis and P. tinctoria (former Gaillonia) in southern Arabia/Horn of Africa.

Background: The Philippine archipelago is globally one of the most important model island systems for studying evolutionary processes. However, most plant species on this archipelago have not yet been studied in sufficient detail. The main aim of this study is to unravel the evolutionary history and biogeographic relationships of the Philippine members of the pantropical genus Ixora. Results: The complex plastid and nuclear divergence patterns in Philippine Ixora, documented using tree and network approaches, reveal a highly dynamic evolution in Ixora, involving several phases of radiation and recolonization. Philippine Ixora comprises at least five lineages, of which one is most closely related to species from Wallacea, and the remaining four to species from Asia. Conclusions: Our study highlights the importance of Philippine species for understanding phytogeographic patterns in the Indomalayan-Australasian eco-region. The overall genetic differentiation, as well as the incongruence between genealogies based on the biparentally inherited nucleome and the maternally inherited plastome in Ixora, reflect the complex tectonic history of the Philippine archipelago. The Ixora lineage related to Wallacean species supports the delimitation of different ecozones along Huxley's line, because it is absent from Palawan. The remaining four lineages are all allied with Asian taxa, reflecting several waves of colonization. Close relationships between some widespread Philippine species and locally adapted narrow endemics suggest that the widespread, genetically diverse species act as pools for the formation of new species in a process of ongoing speciation. Our results suggest that the species concepts of some of the more widespread taxa need to be revised.

Rubiaceae is one of the five largest families of flowering plants with over 13,000 species. We have seen a tremendousincrease in our understanding of the phylogeny of the family through studies on molecular data during the 15-year period from1991 to 2005; some new relationships are completely unexpected and different from traditional classification. At the end of2005, ca. 50 phylogenetic reconstructions from the family had been published based on more than 4400 sequences. Moststudies are based on ITS and rbcL sequences, but 13 different markers have been used. Most sequences available in GenBank(as of 2005) are from rps16, trn(T)L-F, rbcL, and ITS. We can now see a framework of the family phylogeny with support forthree subfamilies and over 43 tribes; subfamily Cinchonoideae (Chiococceae, Cinchoneae, Guettardeae, Hamelieae, Hillieae,Hymenodictyeae, Isertieae, Naucleeae, Rondeletieae), subfamily Ixoroideae (Alberteae, Bertiereae, Coffeeae, Condamineeae,Cremasporeae, Gardenieae, Ixoreae, Mussaendeae, Octotropideae, Pavetteae, Posoquerieae, Retiniphylleae, Sabiceeae,Sipaneeae, Vanguerieae), and subfamily Rubioideae (Anthospermeae, Argostemmateae, Coussareeae, Craterispermeae,Danaideae, Gaertnereae, Knoxieae, Lasiantheae, Morindeae, Ophiorrhizeae, Paederieae, Psychotrieae, Putorieae, Rubieae,Schradereae, Spermacoceae, Theligoneae, Urophylleae), and tribe Coptosapelteae, which is placed outside the threesubfamilies. Two of these tribes, Gardenieae and Morindeae, are paraphyletic/polyphyletic. Only about half of the tribes havebeen the focus of specific investigations. However, we have seen increased interest in using Rubiaceae phylogenies for studiesof ecology, evolution, and biogeography, e.g., and also for morphological and anatomical investigations. Evolution of fruittraits, flower types, and myrmecophytism has been investigated, and biogeographic patterns for specific taxa in Africa, theCaribbean, and the Pacific have been studied. In addition, distribution of pollen types, chemical substances, and woodcharacteristics have been compared with molecular phylogenies.Key words:

Carl Linnaeus brought order to the knowledge of plants and animals by arranging all known species in encyclopaedic

works. He proposed a system of plants, the sexual system, based on the number and arrangement of male and female

organs. His artificial sexual system has since long been replaced by ‘natural’ or phylogenetic systems but there has never

been a comprehensive comparison of the sexual system with modern plant classification. The currently most often used

classification of flowering plants is the APG-system. It is based on comprehensive phylogenies of flowering plants,

reconstructed by analyses of DNA data. The APG-system covers all flowering plants which are classified in 453 families

and these are classified in 45 orders. Most of the species were not known at time of Linnaeus. Families and orders in the

APG-system are arranged in larger informal groups representing major branches in the flowering plant phylogenetic tree.

Three such groups are the monocots, the rosids, and the asterids.

I have examined all genera published in Species plantarum (1753) and classified them according to order and major

groups in the APG-system. All classes except one, number 15 Tetradynamia, comprises groups of unrelated plants. Not

surprisingly, the sexual system does not display what we know today about plant relationships. As is evident from this

analysis, there is little correspondence between the sexual system and the APG-system. This does not mean that the sexual

system has been useless or misleading. When it was introduced, it formed the basis for much intensified research and

increased knowledge of plants.

Schistidium Brid. is a bryophyte genus mainly distributed in temperate to arctic zones. This revision is based on field, herbarium, and literature studies. There are descriptions of the genus and the 12 species and discussions of morphology, relationships, history, nomenclature, habitat, and distribution, and keys to accepted taxa. About 300 synonyms are listed and four new combinations are made. Many lectotypes and a few neotypes are selected.

This study is published in four parts, the first paper deals with species with large spores from the northern hemisphere and the second with the remaining species in the genus. The third paper is the general part and the fourth is a proposal to reject two erroneously used names.

A revised and updated classification for the families of flowering plants is provided. Many recent studies have yielded increasingly detailed evidence for the positions of formerly unplaced families, resulting in a number of newly adopted orders, including Amborellales, Berberidopsidales, Bruniales, Buxales, Chloranthales, Escalloniales, Huerteales, Nymphaeales, Paracryphiales, Petrosaviales, Picramniales, Trochodendrales, Vitales and Zygophyllales. A number of previously unplaced genera and families are included here in orders, greatly reducing the number of unplaced taxa; these include Hydatellaceae (Nymphaeales), Haptanthaceae (Buxales), Peridiscaceae

(Saxifragales), Huaceae (Oxalidales), Centroplacaceae and Rafflesiaceae (both Malpighiales), Aphloiaceae, Geissolomataceae and Strasburgeriaceae (all Crossosomatales), Picramniaceae (Picramniales), Dipentodontaceae and Gerrardinaceae (both Huerteales), Cytinaceae (Malvales), Balanophoraceae (Santalales), Mitrastemonaceae (Ericales) and Boraginaceae (now at least known to be a member of lamiid clade). Newly segregated families for genera previously understood to be in other APG-recognized families include Petermanniaceae (Liliales), Calophyllaceae (Malpighiales), Capparaceae and Cleomaceae (both Brassicales), Schoepfiaceae (Santalales), Anacampserotaceae, Limeaceae, Lophiocarpaceae, Montiaceae and Talinaceae (all Caryophyllales) and Linderniaceaeand Thomandersiaceae (both Lamiales). Use of bracketed families is abandoned because of its unpopularity, and in most cases the broader circumscriptions are retained; these include Amaryllidaceae, Asparagaceace and Xanthorrheaceae (all Asparagales), Passifloraceae (Malpighiales), Primulaceae (Ericales) and several other smaller families. Separate papers in this same volume deal with a new linear order for APG, subfamilial namesthat can be used for more accurate communication in Amaryllidaceae s.l., Asparagaceace s.l. and Xanthorrheaceae s.l. (all Asparagales) and a formal supraordinal classification for the flowering plants. 

Rubiaceae are one of the largest families of plants, with ;13,000 species. In this study, we have estimated the phylogeny for 534Rubiaceae taxa from329 generawith up to five different chloroplast regions by Bayesian analysis. It resulted in a highly resolved tree with many strongly supported nodes. There is strong support for the three subfamilies (Cinchonoideae, Ixoroideae, Rubioideae) and most of the 44 included tribes. A scaled-down data set of 173 Rubiaceae taxawas usedwith a Bayesian approach to estimate divergence times for clades classified as tribes and subfamilies. Four fossils were used as minimum age priors, one inside each subfamily and one for Rubiaceae as a whole (Faramea-type pollen, Scyphiphora pollen, Cephalanthus pusillus fruits, and Paleorubiaceophyllum eocenicum leaves). The estimated lineage (stem) divergence time for Rubiaceae is 90.4Ma. The estimated lineage divergence times for the subfamilies are 84.4 (86.6)Ma for Rubioideae, 73.1Ma for Ixoroideae, and 73.1Ma for Cinchonoideae.The estimated lineage divergence times for the tribes vary between 86.6 and 14.2Ma. Classification, relationships, geographical distribution, and age estimates are presented and discussed for all tribes.

Five new combinations are made in the genus Bremeria: B. arachnocarpa, B. eriantha, B. scabrella, B. landia var. holosericea, and B. landia var. stadmanii. Bremeria gerrardii is conspecific with Gaertnera phanerophlebia, and thus excluded from Bremeria. Lectotypes are designated for Mussaenda erectiloba var. scabrella, M. stadmanii, and M. trichophlebia.

This is the first phylogenetic study focused on the Pavetteae, one of the most species-rich and morphologically diverse tribes within the coffee family (Rubiaceae). Fifteen of the 17 currently recognized genera, represented by 85 taxa, were sequenced for rps16, trnT-F and ITS and analysed using Bayesian inference and maximum likelihood methods. The monophyly of the Pavetteae is confirmed. Four major lineages are identified, but their phylogenetic relationships are not fully resolved. The continental African genera Rutidea, Nichallea and Tennantia, the Madagascan genera Homollea and Robbrechtia, and the paleotropical genus Pavetta are monophyletic. Other genera are paraphyletic in their current circumscriptions and the following changes are made: Homolliella is placed in synonymy with Paracephaelis, and Coleactina and Dictyandra with Leptactina, resulting in four new combinations. The large paleotropical genus Tarenna is shown not to be monophyletic. In the future, the name Tarenna should not be used for continental African species. Most of these could be transferred to the hitherto monospecific genus Cladoceras, but other species might constitute altogether new genera. The relationship between the monophyletic Asian-Pacific and Madagascan Tarenna species remains unclear. The phylogeny of the Madagascan genera of the Pavetteae is largely unresolved and the largest Madagascar-centred genus Coptosperma was not recovered as monophyletic. The low resolution for the Madagascan taxa can be considered as an indication of rapid radiation. Further molecular and morphological studies are necessary to clarify the phylogeny of the Pavetteae, especially regarding the African Tarenna species and the Madagascan genera of the tribe.

According to current generic delimitations, supported by wood anatomy, palynology, and recent molecular phylogenies, Antirhea, traditionally treated as a Pantropical genus, is instead restricted to the Paleotropics. At the same time, the Neotropical species traditionally positioned in Antirhea should be positioned in Stenostomum. Following these definitions, Stenostomum is a genus with most of the species present in the Caribbean Region, and two species present in the Guianas, namely S. acreanum and S. guianensis, distributed throughout the Amazon Basin. In addition, according to these new generic delimitations, two species previously positioned in Guettarda and Antirhea need to be transferred to Chomelia, as C. triflora and C. ulei.

Driven by limited resources and a sense of urgency, the prioritization of species for conservation has been a persistent concern in conservation science. Gymnosperms (comprising ginkgo, conifers, cycads, and gnetophytes) are one of the most threatened groups of living organisms, with 40% of the species at high risk of extinction, about twice as many as the most recent estimates for all plants (i.e. 21.4%). This high proportion of species facing extinction highlights the urgent action required to secure their future through an objective prioritization approach. The Evolutionary Distinct and Globally Endangered (EDGE) method rapidly ranks species based on their evolutionary distinctiveness and the extinction risks they face. EDGE is applied to gymnosperms using a phylogenetic tree comprising DNA sequence data for 85% of gymnosperm species (923 out of 1090 species), to which the 167 missing species were added, and IUCN Red List assessments available for 92% of species. The effect of different extinction probability transformations and the handling of IUCN data deficient species on the resulting rankings is investigated. Although top entries in our ranking comprise species that were expected to score well (e.g. Wollemia nobilis, Ginkgo biloba), many were unexpected (e.g. Araucaria araucana). These results highlight the necessity of using approaches that integrate evolutionary information in conservation science.

Alchemilla (the lady’s mantles) is a well known but inconspicuous group in the Rosaceae, notable for its ornamental leaves and pharmaceutical properties. The systematics of Alchemilla has remained poorly understood, most likely due to confusion resulting from apomixis, polyploidisation and hybridisation, which are frequently observed in the group, and which have led to the description of a large number of (micro-) species. A molecular phylogeny of the genus, including all sections of Alchemilla and Lachemilla as well as five representatives of Aphanes, based on the analysis of the chloroplast trnL–trnF and the nuclear ITS regions is presented here. Gene phylogenies reconstructed from the nuclear and chloroplast sequence data were largely congruent. Limited conflict between the data partitions was observed with respect to a small number of taxa. This is likely to be the result of hybridisation/introgression or incomplete lineage sorting. Four distinct clades were resolved, corresponding to major geographical division and life forms: Eurasian Alchemilla, annual Aphanes, South American Lachemilla and African Alchemilla. We argue for a wider circumscription of the genus Alchemilla, including Lachemilla and Aphanes, based on the morphology and the phylogenetic relationships between the different clades.

Argostemmateae (Rubiaceae, subfam. Rubioideae) are a mostly tropical Asian group of about 200 species currently classified in four morphologically distinct genera (Argostemma, Mouretia, Mycetia, Neohymenopogon). The monophyly of the tribe and Mycetia is strongly supported by molecular data, however, the tropical Asian genus Myrioneuron, traditionally associated with Mycetia based on its berry fruits, has not previously been investigated. The Bornean and Sulawesian genus Cyanoneuron, described based on the species of Myrioneuron with drupaceous fruits, had not been sequenced. Therefore, the phylogenetic positions of Cyanoneuron and Myrioneuron within Rubiaceae and their generic status have yet to be assessed with molecular data. These genera have tentatively been placed in tribe Spermacoceae (Rubioideae). We reconstructed a robust phylogeny of Rubioideae with sequence data from five plastid regions of 176 accessions and using the Bayesian Markov chain Monte Carlo and parsimony methods. Once the positions of Cyanoneuron and Myrioneuron were revealed, a robust phylogeny of the Spermacoceae alliance was reconstructed with the combined plastid and nuclear data (nrETS, nrITS) from 61 accessions to reassess its tribal limits. Mycetia and Myrioneuron are non-monophyletic and intermixed, and formed a well-supported clade diagnosed by berry fruits. We formally transfer Myrioneuron to Mycetia (older name), and present nine new combinations in the latter genus. Cyanoneuron was resolved with high support as monophyletic, and appears to be closely related to the Chinese monogeneric tribes Foonchewieae and Dunnieae. A new tribe Cyanoneuroneae is described to accommodate Cyanoneuron. This tribe is morphologically distinct from related tribes by its stipules apically divided into multiple linear segments, condensed-cymose inflorescences and drupe-like fruits with numerous small seeds. A new key to the genera of Argostemmateae is provided.

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.

We performed molecular phylogenetic analyses of the Randia clade of the tribe Gardenieae using both plastid and nuclear DNA data. In the phylogenetic hypotheses presented, the African genera Calochone, Euclinia, Macrosphyra, Oligocodon, Pleiocoryne, and Preussiodora are resolved as a monophyletic group. Support is also found for a clade of the Neotropical genera Casasia, Randia, Rosenbergiodendron, Sphinctanthus, and Tocoyena. This Neotropical clade is resolved as sister group to the African clade in analyses of combined plastid and nuclear data. The genus Euclinia appears polyphyletic. The species described from Madagascar represent an independent lineage, the position of which is moreover found to be incongruent between datasets. Plastid and ribosomal DNA data support a sister-group relationship to the mainland African clade, whereas the low-copy nuclear gene Xdh supports a closer relationship to the Neotropical genera. The phylogenetic reconstructions also indicate that Casasia and Randia are not monophyletic as presently circumscribed. A taxonomic proposal is made for the recognition of the Malagasy taxon at generic level as Melanoxerus.

Subfamily Ixoroideae is one of three major lineages in Rubiaceae, with approximately 4000 species. Previous molecular phylogenetic studies have indicated that many genera and tribes previously placed in other subfamilies are better considered as part of Ixoroideae. However, the internal resolution and clade support have generally been low, and several genera found to be nested in the subfamily do not appear to be associated with any described tribe. In order to resolve the phylogeny and assess the tribal delimitations in the expanded Ixoroideae, phylogenetic reconstructions were performed using Bayesian and parsimony analyses of six plastid DNA regions and a broad sampling of genera from all tribes of the subfamily. In the inferred phylogenetic hypotheses, the tribal relationships were mostly well supported, with Ixoroideae consisting of the Coffeeae and the Vanguerieae alliances as sister groups and a grade comprising Condamineeae, Henriquezieae, Posoquerieae, Retiniphylleae, Sipaneeae and the genus Steenisia. A revised tribal classification, including the description of five new tribes, Airospermeae, Augusteae, Scyphiphoreae, Steenisieae and Trailliaedoxeae, is provided.

Extensive efforts have been made to resolve the phylogeny of the large coffee family (Rubiaceae) based on molecular data. As a result, several small tribes have been described, but the phylogenies and generic delimitations for many of these groups remain unclear. This study focuses on the small tribe Danaideae that belongs to subfamily Rubioideae and whose generic limits have not previously been addressed with molecular data. It is the sole rubiaceous tribe distributed almost entirely in the Western Indian Ocean region, with the exception of the East African Danais xanthorrhoea. The tribe consists of three genera: Danais, Payera (including the monotypic genus Coursiana), and Schismatoclada. We present the first molecular phylogenetic study of Danaideae including representatives from all three genera and using Bayesian and maximum parsimony methods and sequence data from nuclear DNA (nrITS) and chloroplast DNA (petD, psbA-trnH, rpl32-trnL(UAG), rps16). Our main objectives were to rigorously test the monophyly of Danaideae as currently circumscribed and assess phylogenetic relationships within the tribe. The findings of this study shed light on the colonization history of the tribe. Our analyses reaffirm the monophyly of Danaideae and Danais but reveal the paraphyly of Payera and Schismatoclada. The close relationship between the three Danaideae genera and Coursiana is supported. However, we found very little support for the inclusion of the latter genus in Payera as proposed earlier. The tribe is resolved in two morphologically distinct major lineages, the highly supported Damns clade with lianescent habit (= Danais sensu Buchner & Puff) and the Payera-Schismatoclada clade with arborescent habit. The Malagasy and Mauritian specimens of Danais fragrans are not closely related, and we restrict D. fragrans to the Mauritian taxa and resurrect Danais lyallii Baker to accommodate the Malagasy D. fragrans. According to our analysis. Madagascar is the origin of all species of Danaideae occurring in the Comoro archipelago, East Africa, and Mauritius. The Mauritian and East African Danais each is the result of a single colonization event, while there were at least two independent colonization events to the Comoros.

The tribe Knoxieae consists of genera formerly included in (at least) four tribes of the Rubiaceae (the coffee family). Apart from Knoxia itself, which is mainly Indomalesian, the group is from Africa and Madagascar. Most genera are herbaceous or shrubby, but small trees are also present. They generally have fimbriate, colleter-tipped stipules and five-merous flowers with unequal calyx lobes. We present a molecular phylogeny of the group (based on two chloroplast markers, rps16 and trnT-F, and the nuclear ITS) and discuss the morphological support for the resulting groups and the taxonomic consequences of the phylogeny. Two hundred and sixty-one new sequences from 90 species have been obtained. Of the 19 genera of the Knoxieae, all but three (two monotypic and one with two species) are represented. The genera Calanda, Chlorochorion, Neopentanisia, and Paraknoxia are merged with Pentanisia. Pentas is shown to be not monophyletic and is split into the new genera Phyllopentas, Dolichopentas, and Rhodopentas. We also suggest the Pentas subgenera Megapentas and Chamaepentadoides be included in Chamaepentas. The genus Carphalea is also shown to be not monophyletic and for the African members of the genus (section Dirichletia) the name Dirichletia should be used. Placopoda is suggested to be merged with Dirichletia. Triainolepis is enlarged to include Paratriaina and Thyridocalyx. The tribal position of Lathraeocarpa is discussed. Twenty-nine new combinations for species and seven for infraspecific taxa are made. A key to the genera of Knoxieae is provided.

Potential events of allopolyploidy may be indicated by incongruences between separate phylogenies based on plastid and nuclear gene sequences. We sequenced two plastid regions and two nuclear ribosomal regions for 34 ingroup taxa in Fragariinae (Rosaceae), and six outgroup taxa. We found five well supported incongruences that might indicate allopolyploidy events. The incongruences involved Aphanes arvensis, Potentilla miyabei, Potentilla cuneata, Fragaria vesca/moschata, and the Drymocallis clade. We evaluated the strength of conflict and conclude that allopolyploidy may be hypothesised in the four first cases. Phylogenies were estimated using Bayesian inference and analyses were evaluated using convergence diagnostics. Taxonomic implications are discussed for genera such as Alchemilla, Sibbaldianthe, Chamaerhodos, Drymocallis and Fragaria, and for the monospecific Sibbaldiopsis and Potaninia that are nested inside other genera. Two orphan Potentilla species, P. miyabei and P. cuneata are placed in Fragariinae. However, due to unresolved topological incongruences they are not reclassified in any genus.

The Hymenodicteae-Naudeeae clade is a predominantly Paleotropical group with 220 species in 28 genera. The phylogertetic relationships and generic limits within Naucleeae have previously been assessed using combined molecular-morphological data, however the status of some genera remains questionable. The evolutionary relationships within Hymenodictyeae have never been investigated before. We performed phylogenetic analyses of the Hymenodictyeae-Naucleeae clade using nuclear [nrETS; nrITS] and chloroplast [ndhF; rbcL; rps16; trnT-F] data and a large sampling of both tribes. Our study supports the monophyly of the tribes, all subtribes of Naucleeae (Adininae, Breoniinae, Cephalanthinae, Corynantheinae, Mitragyninae, Naucleinae, and Uncariinae), and the Hymenodictyeae genera Hymenodictyon and Paracorynanthe. In Naucleeae, the monotypic genera Adinauclea, Metadina, and Pertusadina are nested within Adina, Mitragyna within Fleroya, Ludekia, Myrmeconauclea, and Ochreinauclea within Neonauclea, and Burttdavya and Sarcocephalus within Nauclea. Corynanthe and Pausinystalia are mutually paraphyletic. We provisionally maintain the current generic status of Neonauclea and its allied genera, pending further study. In sum, we recognize 17 genera in Naucleeae: Adina s. l., Breonadia, Breonia, Cephalanthus, Corynanthe s. l., Diyaminauclea, Gyrostipula, Janotia, Khasiaclunea, Ludekia, Mitragyna s. l., Myrmeconauclea, Nauclea s. l., Neolamarckia, Neonauclea, Ochreinauclea, and Uncaria. Five new combinations were made: Adina euryncha, Adina malaccensis, Corynanthe lane-poolei subsp. iturense, Corynanthe talbotii, and Nauclea nyasica.

The Hymenodictyeae-Naucleeae clade is a predominantly Paleotropical group with 220 species in 28 genera. The phylogenetic relationships and generic limits within Naucleeae have previously been assessed using combined molecular-morphological data, however the status of some genera remains questionable. The evolutionary relationships within Hymenodictyeae have never been investigated before. We performed phylogenetic analyses of the Hymenodictyeae-Naucleeae clade using nuclear [nrETS; nrITS] and chloroplast [ndhF; rbcL; rps16; trnT-F] data and a large sampling of both tribes. Our study supports the monophyly of the tribes, all subtribes of Naucleeae (Adininae, Breoniinae, Cephalanthinae, Corynantheinae, Mitragyninae, Naucleinae, and Uncariinae), and the Hymenodictyeae genera Hymenodictyon and Paracorynanthe. In Naucleeae, the monotypic genera Adinauclea, Metadina, and Pertusadina are nested within Adina, Mitragyna within Fleroya, Ludekia, Myrmeconauclea, and Ochreinauclea within Neonauclea, and Burttdavya and Sarcocephalus within Nauclea. Corynanthe and Pausinystalia are mutually paraphyletic. We provisionally maintain the current generic status of Neonauclea and its allied genera, pending further study. In sum, we recognize 17 genera in Naucleeae: Adina s.l., Breonadia, Breonia, Cephalanthus, Corynanthe s.l., Diyaminauclea, Gyrostipula, Janotia, Khasiaclunea, Ludekia, Mitragyna s.l., Myrmeconauclea, Nauclea s.l., Neolamarckia, Neonauclea, Ochreinauclea, and Uncaria. Five new combinations were made: Adina euryncha, Adina malaccensis, Corynanthe lane-poolei subsp. iturense, Corynanthe talbotii, and Nauclea nyasica.

This study focuses on the subfamily Cinchonoideae s.s. utilizing information from six DNA markers and 206 taxa. The nine tribes (i.e. Cinchoneae, Chiococceae s.l., Guettardeae s.s., Hamelieae, Hillieae, Hymenodictyeae, Isertieae, Naucleeae s.l., and Rondeletieae s.s.) are resolved in four major lineages, all strongly supported and relationships between them are resolved. The tropical American Cinchoneae and Isertieae constitute the first diverging lineage within the subfamily, followed by the predominantly paleotropical Naucleeae and Hymenodictyeae. The remaining two lineages primarily include neotropical taxa: Rondeletieae and Guettardeae are sister clades in the first, while the second comprises Chiococceae, Hamelieae, and Hillieae. Additionally, taxonomic placement of several genera, not previously included in molecular analyses, were confirmed: Acunaeanthus belongs in Rondeletieae, Ottoschmidtia in Guettardeae, Nernstia in Chiococceae, Pinarophyllon, Plocaniophyllon, and Syringantha in Hamelieae, and Balmea in Hillieae. Colleteria, of previously unknown taxonomic position, is resolved as sister to Chione.

The Rubiaceae is the fifth largest plant family and is found on all continents, mostly in (sub-)tropical regions. Despite a large representation of Rubiaceae in the Paleotropics, the subfamily Cinchonoideae has its primary distribution in the Neotropics. Within the Cinchonoideae only two tribes, Naucleeae and Hymenodictyeae, have Paleotropical centers of distribution. In this study, we used information from five chloroplast DNA markers and fossil data to investigate when the subfamily was established in the Neotropics as well as major subsequent dispersal events within and out of the Neotropics and within the Paleotropics. Our results show that the ancestor of Cinchonoideae and its sister, Ixoroideae, was present in South America during the Late Cretaceous. Dispersal to Central America was estimated to occur during the Early Paleocene, and subsequent dispersals to the Caribbean islands occurred during the Oligocene-Miocene. The ancestor of Naucleeae and Hymenodictyeae dispersed to the Paleotropics no later than the Eocene, and the findings of fossils of Cephalanthus in Europe and western Siberia support its presence in the boreotropical forests. Long-distance, trans-Pacific dispersal during the Miocene was indicated for the remaining Paleotropical Cinchonoideae.

In the Comoro Islands, Rubiaceae are one of the largest families of flowering plants, represented by c. twenty genera and seventy species. Among these genera, Coptosperma (Pavetteae, Ixoroideae) is represented by two widely distributed species, also occurring in Madagascar and

continental Africa, and a third species new to science. Coptosperma mitochondrioides Mouly & De Block possesses large stipules of the “bec de canard” type and ribbed fruits. While this type of stipule is typical for many Malagasy Coptosperma species, ribbed fruits are very rare within the genus. Within the Comoros archipelago C. mitochondrioides is restricted to Mayotte, but the species is also known from northern Madagascar. The new species is described and illustrated and a key to the species of Coptosperma occurring in the Comoros is provided.

In this study we investigate the large and diverse Rubiaceae-Gardenieae and closely related tribes Bertiereae, Coffeeae, Cremasporeae, Octotropideae, and Pavetteae. Some of the tribes or groups have been shown to be monophyletic and strongly supported, but the phylogeny of this large complex is still far from being satisfactorily elucidated particularly for Gardenieae, both in terms of intertribal relationships as well as tribal delimitations. We reconstruct the phylogeny of the complex using an extensive sampling of 108 genera and five plastid DNA regions. Phylogenetic relationships demonstrate that Gardenieae sensu Andreasen & Bremer is polyphyletic, as Burchellia, Didymosalpinx, Monosalpinx, and Mantalania are closer to Octotropideae-Cremasporeae. In addition, Pavetteae and the investigated members of Aulacocalyceae are nested in a supported but partially unresolved Gardenieae-Pavetteae clade. Within this clade, several strongly supported groups are resolved: an Aidia group, an Alibertia group, a Gardenia group, Pavetteae including Pelagodendron, a Porterandia group, a Randia group, a Rothmannia group (including Aulacocalyx and Heinsenia), a Sherbournia group, and the two isolated genera Massularia and Schumanniophyton. The latter genus presented a high rate of genetic substitutions, which resulted in perturbations of the phylogenetic reconstruction. A revised tribal circumscription is given for Gardenieae, the Alibertia group is recognized at tribal level as an emended Cordiereae, and a new tribe, Sherbournieae, is described to accommodate the members of the Sherbournia group.

Ixora clarae and I. elisae (Rubiaceae) are two endemic species described from New Caledonia. Ixora clarae is easily recognised by its (1-)3-flowered inflorescences, made up by thick and large flowers embedded in a calyculus. It is known from a few specimens collected on the nearest mountains by Nouméa, and is considered as endangered (EN). Ixora elisae is remarkable for its very enlarged campanulate calyx-limb. The latter one, from the Isle of Pines and Lifou, known by just two herbarium collections, is probably on its way to disappear in nature. Consequently, it is proposed to a critically endangered (CR) classification.

A new species of Paederia L. (Rubiaceae, Rubioideae, Paederieae), P. ntiti Mouly & Puff, endemic to the Comoros (Grande Comore, Anjouan and Mayotte) and occurring in natural elevated forests is described and illustrated. Its conservation status is evaluated and the species is rated 'Endangered' (EN). A key to the Paederia taxa of the Comorian archipelago is provided. Also included are notes on the three other species of rubiaceous climbers and lianas known from the Comoros: Paederia bojeriana (A. Rich. ex DC.) Drake, Danais comorensis Drake and Uncaria africana G. Don.

We performed phylogenetic analyses of DNA sequences of three chloroplast markers: rbcL, rps16, and trnT-F, to rigorously test the monophyly of competing circumscriptions of the tribe Ixoreae. Several genera traditionally or currently associated with the type genus Ixora L. were included in the analyses. Ixora as currently circumscribed appears paraphyletic, as many other genera are nested within it with strong support: Captaincookia N. Halle´, Doricera Verdc., Hitoa Nadeaud, Myonima Comm. ex Juss., Sideroxyloides Jacq., Thouarsiora Homolle ex Are`nes, and Versteegia Valeton. Further, Aleisanthia Ridl., Aleisanthiopsis Tange, and Greenea Wight & Arn. are more closely related to Ixora and allies than the monotypic genus Scyphiphora C. F. Gaertn. Consequently, Ixoreae fide Andreasen and Bremer (2000) is not monophyletic without an exclusion of Scyphiphora. Ixoreae fide Robbrecht and Manen (2006) is not monophyletic unless Captaincookia and Doricera are included. The monophyly of a morphologically heterogeneous Ixoreae alliance consisting of Ixora and its relatives Aleisanthia, Aleisanthiopsis, and Greenea is, however, strongly supported. In order to recognize monophyletic and morphologically consistent groups, we adopt a narrow circumscription of Ixoreae (including Bemsetia Raf., Captaincookia, Charpentiera Vieill., Doricera, Hitoa, Ixora, Myonima, Pancheria Montrouz., Sideroxyloides, Thouarsiora, Tsiangia But, H. H. Hsue & P. T. Li, and Versteegia), and two new tribes are erected for Aleisanthia + Aleisanthiopsis and Greenea, respectively. The Indo-Malesian Aleisanthieae and the pantropical Ixoreae s. str. are sister groups, and the Southeast Asian Greeneeae is sister to the Ixoreae–Aleisanthieae clade.

Species-rich genera often have various conflicting circumscriptions from independent regional fl ora treatments. Testing the monophyly of these groups of plants is an important step toward the establishment of a phylogenetic classifi cation. The genus Ixora of the tribe Ixoreae in the subfamily Ixoroideae (coffee family or Rubiaceae) is a species-rich pantropical genus of ca. 500 species. Phylogenetic analyses of Ixoreae based on combined sequence data from one nuclear (nrETS) and two chloroplast ( rps16 and trnT-F ) markers reveal the paraphyly of Ixora as presently delimited and also show that the tribe can be subdivided into three major

clades: the Mascarene/neotropical/Malagasy/African clade, the Pacifi c clade, and the Asian clade. Given the lack of morphological synapomorphies supporting the different Ixora clades and the morphological consistency of the ingroup taxa, we propose a broad circumscription of Ixora including all its satellite genera: Captaincookia , Doricera , Hitoa , Myonima , Sideroxyloides , Thouarsiora, and Versteegia . The current infrageneric classifi cation of Ixora is not supported. The different Ixora subclades represent geographical units. Nuclear and chloroplast tree topologies were partially incongruent, indicating at least four potential natural hybridization events. Other confl icting positions for the cultivated species are most likely due to anthropogenic hybridization.

Hedyotis and related genera (here called the Hedyotis-Oldenlandia complex) are highly debated groups in the Rubiaceae family with no consensus to date on their generic delimitations. The present study focuses on Asian-Pacific taxa from these groups and aims at resolving taxonomic inconsistencies by describing monophyletic genera within the complex. The generic circumscriptions presented in our study are based on the phylogenetic trees of nuclear (ITS, ETS) and plastid (petD, rps16) sequence data inferred using Bayesian and maximum likelihood methods. Morphological key features of the group such as habit, fruit type, seed form, and pollen type are studied and compared with the phylogeny to characterize the clades. Based on these results, the Asian-Pacific members are placed in 14 monophyletic groups across the Hedyotis-Oldenlandia complex. Of these, we accept and circumscribe 13 monophyletic genera: Debia, Dentella, Dimetia, Edrastima, Exallage, Hedyotis, Involucrella, Kadua, Kohautia, Leptopetalum, Neanotis, Oldenlandia, and Scleromitrion. Two of these, Debia and Involucrella, are here described as new genera.