Abstract: The papers in this volume are the result of a meeting of fig and fig wasp biologists held in China, July 2006. Here we provide an overview of current themes in fig-fig wasp research and introduce the papers in this volume. Figs are pollinated by tiny, highly specific wasps whose larvae feed on a proportion of the flowers they pollinate. Other symbionts, including a diversity of non-pollinating wasps and vertebrate fruit eaters, exploit or otherwise depend on this obligate mutualistic interaction. Recently it was found that a substantial number of fig species harbour more than one species of pollinating wasp. This has lead to renewed interest in the question of specificity and its co-evolutionary implications. There has also been a substantial increase in the interest in fig and fig wasp biology in Asia, not least in China where our meeting was held. Directly following from this is an increased interest in dioecious figs, which are predominantly Asian. Dioecious figs offer a very different perspective on the stability of the fig-fig pollinator interaction to the better studied monoecious figs. They have also radiated into a remarkable diversity of ecological niches and, therefore, promise to reveal how fig-fig wasp interactions have responded to a variety of different ecological conditions. The study of non-pollinating fig wasp communities, which are every bit as interesting as the pollinating wasps, has been hampered by difficulties in determining their life-histories. A number of the papers presented here pioneer new methods to address these difficulties, and hence promise to prise open this field. Fig and fig wasp biology appears on the cusp of rapid developments on a number of fronts, and hence fig-fig wasp biologists can look forward with excitement to the coming years.
Abstract: Figs (Ficus, Moraceae) constitute one of the largest genera of flowering plants with ca. 750 species worldwide. While the extraordinary mutualism between figs and their pollinating wasps has received attention for decades, we are only Just beginning to reconstruct the phylogeny of both partners, a necessary framework for addressing a variety of questions concerning the evolution of mutualism. Here, we present phylogenctic analyses of 100 species of Ficus, representing all subgenera. sixteen out of nineteen sections, and two outgroups, using three nuclear markers. We explore the utility of the single copy nuclear encoded glyceraldehyde-3-phosphate dehydrogenase gene (G3pdh) for phylogeny reconstruction ill Ficus, and evaluate infrageneric relationships based on G3pdh DNA sequences in combination with the nuclear ribosomal internal and external transcribed spacers (ITS and ETS). The G3pdh gene provides limited resolution within Ficus, but increases the proportion of well-supported clades when combined with ITS and ETS. Of the six subgenera traditionally recognized based on morphology and distribution patterns, only subgenus Sycidium is supported as monophyletic. We identify fifteen clades within Ficus, but the branching order of the early lineages of Ficus and some of the internal branches are not well supported and should be considered uncertain at present.
Abstract: Plants are of immense importance in providing healthcare worldwide. With over 250 000 species of angiosperms alone, the potential for finding new medicinal plants and lead compounds for drug development is enormous. Some way of selecting plants for drug discovery programs is necessary. Phylogenies have great explanatory power and also enable a predictive perspective not offered by previous classifications of plants. Phylogenetic selection of target species is a new approach to drug discovery and the present study is the first attempt to correlate acetylcholine esterase (AChE) inhibitory activity and alkaloid distribution with a molecular phylogenetic hypothesis of Narcissus. The distribution of alkaloids with AChE inhibitory activity is significantly constrained by the phylogeny. Simultaneous evaluation of all available information of alkaloids and AChE inhibitory activity in a phylogenetic framework allowed us to discuss various strategies for selection of target species for further studies of AChE inhibitory activity
Abstract: The study of chalcid wasps that live within syconia of fig trees (Moraceae, Ficus), provides a unique opportunity to investigate the evolution of specialized communities of insects. By conducting cospeciation analyses between figs of section Galoglychia and some of their associated fig wasps, we show that, although host switches and duplication have evidently played a role in the construction of the current associations, the global picture is one of significant cospeciation throughout the evolution of these communities. Contrary to common belief, nonpollinating wasps are at least as constrained as pollinators by their host association in their diversification in this section. By adapting a randomization test in a supertree context, we further confirm that wasp phylogenies are significantly congruent with each other, and build a "wasp community" supertree that retrieves Galoglychia taxonomic subdivisions. Altogether, these results probably reflect wasp host specialization but also, to some extent, they might indicate that niche saturation within the fig prevents recurrent intrahost speciation and host switching. Finally, a comparison of ITS2 sequence divergence of cospeciating pairs of wasps suggests that the diversification of some pollinating and nonpollinating wasps of Galoglychia figs has been synchronous but that pollinating wasps exhibit a higher rate of molecular evolution.
Abstract: We conducted the first molecular phylogenetic study of Ficus section Malvanthera (Moraceae; subgenus Urostigma) based on 32 Malvanthera accessions and seven outgroups representing other sections of Ficus subgenus Urostigma. We used DNA sequences from the nuclear ribosomal internal and external transcribed spacers (ITS and ETS), and the glyceraldehyde-3-phosphate dehydrogenase (G3pdh) region. Phylogenetic analysis using maximum parsimony, maximum likelihood and Bayesian methods recovered a monophyletic section Malvanthera to the exclusion of the rubber fig, Ficus elastica. The results of the phylogenetic analyses do not conform to any previously proposed taxonomic subdivision of the section and characters used for previous classification are homoplasious. Geographic distribution, however, is highly conserved and Melanesian Malvanthera are monophyletic. A new subdivision of section Malvanthera reflecting phylogenetic relationships is presented. Section Malvanthera likely diversified during a period of isolation in Australia and subsequently colonized New Guinea. Two Australian series are consistent with a pattern of dispersal out of rainforest habitat into drier habitats accompanied by a reduction in plant height during the transition from hemi-epiphytic trees to lithophytic trees and shrubs. In contradiction with a previous study of Pleistodontes phylogeny suggesting multiple changes in pollination behaviour, reconstruction of changes in pollination behaviour on Malvanthera, suggests only one or a few gains of active pollination within the section.
Abstract: Ficus section Galoglychia (subgenus Urostigma; Moraceae) includes 72 species restricted to the African floristic region (a few extending to the Arabian Peninsula and Socotra). We present the first molecular phylogenetic analysis of the section including 56 ingroup (representing 44 species) and three outgroup taxa, to investigate its monophyly, classification and evolution. We used sequence data from the nuclear ribosomal internal and external transcribed spacers (ITS and ETS). Our results suggest that section Galoglychia is paraphyletic to the neotropical section Americana, although this is not supported by bootstrap analysis and only weakly supported by Bayesian posterior probabilities. Maximum parsimony analysis conflict with maximum likelihood and Bayesian analyses with respect to the closest relatives of section Americana in Africa. The subsections of section Galoglychia proposed by Berg [Berg, C.C., 1986. Subdivision of Ficus subg. Urostigma sect. Galoglychia (Moraceae). Proc. Kon. Ned. Akad. Wetensch., Ser. C, 89, 121-127] are generally supported. We find two major clades of section Galoglychia within Africa possibly corresponding to two main centres of diversity. One clade comprises members of subsections Platyphyllae and Chlamydodorae, which are more concentrated in Eastern Africa, and extend to Madagascar and neighbouring archipelagos (Comores, Mascarenes, Aldabra Islands and Seychelles). The other main clade includes members of subsections Caulocarpae, Cyathistipulae, Crassicostae and Galoglychia, which are concentrated in West and Central Africa.
Abstract: We present phylogenetic analyses of 37 taxa of Fritillaria (Liliaceae), 15 species of Lilium, and several outgroup taxa from Liliaceae s.s. to investigate the generic delimitation of Fritillaria in relation to Lilium as well as infrageneric relationships within Fritillaria. We used DNA sequences from the maturase-coding plastid matK gene and the trnK intron, the intron of the ribosomal protein-coding rpl16 plastid gene, and the nuclear ribosomal internal transcribed spacers (ITS). Phylogenetic analysis using maximum parsimony defined Fritillaria and Lilium (the latter including Nomocharis) as sister taxa. Fritillaria sections Fritillaria and Liliorhiza are supported in part, and some of the most enigmatic species usually included in Fritillaria (sections Petilium and Theresia and the monotypic genus Korolkowia) are closely related. The results support the new classification of Fritillaria proposed by Rix. We postulate independent origins of the underground bulbils found in Fritillaria davidii and the remainder of subgenus Liliorhiza.
Abstract: Figs (Ficus; ca 750 species) and fig wasps (Agaoninae) are obligate mutualists: all figs are pollinated by agaonines that feed exclusively on figs. This extraordinary symbiosis is the most extreme example of specialization in a plant-pollinator interaction and has fuelled much speculation about co-divergence. The hypothesis that pollinator specialization led to the parallel diversification of fig and pollinator lineages (co-divergence) has so far not been tested due to the lack of robust and comprehensive phylogenetic hypotheses for both partners. We produced and combined the most comprehensive molecular phylogenetic trees to date with fossil data to generate independent age estimates for fig and pollinator lineages, using both non-parametric rate smoothing and penalized likelihood dating methods. Molecular dating of ten pairs of interacting lineages provides an unparalleled example of plant-insect co-divergence over a geological time frame spanning at least 60 million years.
Abstract: In continuation of our investigations of the genus Plantago L. (Plantaginaceae), sixteen species were investigated with respect to water-soluble glycosides. The iridoids auroside, strictoloside and globularicisin, as well as poliumoside, 3-[(4-beta-D-glucopyranosyloxy)phenyl]propionic acid and 2-[4-(beta-D-glucopyranosyloxy)phenyl]acetic acid were isolated from Plantago for the first time. The latter compound has not previously been isolated as a natural product. Sorbitol was the main carbohydrate in all the species investigated. The distribution of iridoids correlates well with the morphological classification of Rahn and also with a recently published molecular phylogenetic study of nuclear ribosomal and plastid DNA sequences. A new chemotaxonomic finding is an abundance of iridoid glucosides present in one of the two groups within subgenus Coronopus section Coronopus, while the other group lacks iridoids, supporting a subdivision of this section. Moreover, in subgenus Albicans, 10-benzoylcatalpol is a characteristic constituent of section Gnaphaloides, while the corresponding cinnamoyl ester globularin occurs in section Lanceifolia. In biosynthetic experiments, labelled epideoxyloganic acid and deoxygeniposidic acid were incorporated into aucubin and geniposidic acid in Plantago ovata, consistent with earlier findings, but no incorporation into asperuloside was observed. The evolution of biosynthetic pathways in Plantago is discussed
Abstract: From the water-soluble part of an extract of Aragoa cundinamarcensis were isolated seven iridoid glucosides, namely aucubin, catalpol, rehmannioside D, globularin, gardoside methyl ester, epiloganin and mussaenoside. The main glycoside isolated, however, was a new caffeoyl phenylethanoid triglycoside, named aragoside, containing two beta-gluco- and one alpha-arabinopyranosyl moieties which constituted almost 5% of the dry weight of the plant. Finally, sorbitol was found to be the main carbohydrate constituent of the plant. This distinctive combination of compounds is very similar to that reported from some species of Plantago. The present findings therefore support the results from a recently published molecular phylogenetic study of plastid and nuclear ribosomal DNA sequences, where Aragoa was found to be the closest relative to Plantago so far discovered.
Abstract: transcribed spacers (ITS) of nuclear ribosomal and plastid trnL-F regions. Included are 57 Plantago species, with
two Aragoa species as the ingroup and three Veronica species as the outgroup. Phylogenetic analysis using maximum
parsimony identified five major clades, corresponding to the taxonomic groups Plantago subgenera Plantago,
Coronopus, Psyllium, Littorella and Bougueria. Aragoa is sister to genus Plantago. Plantago subgenus Littorella is
sister to the other subgenera of Plantago. The results are in general correlated with a morphological phylogenetic
study and iridoid glucoside patterns, but Plantago subgenus Albicans is paraphyletic and should be included in
Plantago subgenus Psyllium sensu lato to obtain a monophyletic clade with six sections. Plantago section Hymenopsyllium
is more closely related to section Gnaphaloides than to section Albicans. Plantago subgenus Bougueria is
sister to subgenus Psyllium s.l. section Coronopus in Plantago subgenus Coronopus is subdivided in two series. Only
some of the sections can be resolved into series. DNA variation within genus Plantago is high, a result that would
not have been predicted on the basis of morphology, which is relatively stereotyped. If we calibrate a molecular clock
based on the divergence of P. stauntoni, endemic to New Amsterdam in the southern Indian Ocean, we calculate
the time of the split between Plantago and Aragoa to be 7.1 million years ago, which is congruent with the fossil
record.
Abstract: Data for 34 species of Plantago (Plantaginaceae), including subgen. Littorella (= Littorella uniflora), have been collected with regard to their content of iridoid glucosides and caffeoyl phenylethanoid glycosides (CPGs). In the present work, 21 species were investigated for the first time and many known compounds were found together with three new iridoid glucosides. Of these, arborescoside and arborescosidic acid, both of the uncommon type with an 8,9-double bond, were present in several species, while 6-deoxymelittoside was found only in P. subulata. The known compounds deoxyloganic acid, caryoptoside and rehmannioside D were isolated from the genus for the first time. The earlier reported occurrence of sorbitol in the family was confirmed, and this compound was shown by NMR spectroscopy to be the main sugar in the three species investigated for this. The combined data show that CPGs are present in all species investigated. With regard to the iridoids, the distribution patterns showed a good correlation with the classification of Rahn. Thus, aucubin is typical for the whole genus, while bartsioside and catalpol as well as 5-substituted iridoids are each characteristic for a subgenus in the family. Finally, the close relationship between Plantago and Veronica suggested by chloroplast DNA sequence analysis. could be corroborated by the common occurrence of the rare 8,9-unsaturated iridoids in these two genera.
Abstract: Floral nectar is rich in chemicals and induces pollination1. Although it may be tainted by algae or mould, it usually lacks colouring agents. However, a few plant species in Mauritius break this rule and produce red nectar. We attempted to find a function for this coloration, but its role remains unclear.
