Two decades of research [1][2][3][4] have not established whether tropical insect herbivores are dominated by specialists or generalists. This impedes our understanding of species coexistence in diverse rainforest communities. Host specificity and species richness of tropical insects are also key parameters in mapping global patterns of biodiversity 1,4,5 . Here we analyse data for over 900 herbivorous species feeding on 51 plant species in New Guinea and show that most herbivorous species feed on several closely related plant species. Because species-rich genera are dominant in tropical floras, monophagous herbivores are probably rare in tropical forests. Furthermore, even between phylogenetically distant hosts, herbivore communities typically shared a third of their species. These results do not support the classical view that the coexistence of herbivorous species in the tropics is a consequence of finely divided plant resources; non-equilibrium models of tropical diversity 6 should instead be considered. Low host specificity of tropical herbivores reduces global estimates of arthropod diversity from 31 million (ref. 1) to 4-6 million species. This finding agrees with estimates based on taxonomic collections, reconciling an order of magnitude discrepancy between extrapolations of global diversity based on ecological samples of tropical communities with those based on sampling regional faunas 7,8 .Host specificity is difficult to measure, and the limitations of existing studies include sampling only certain taxonomic groups rather than entire guilds, or sampling limited numbers of host plant species and lineages. Studies are often of insufficient duration, producing samples too small for quantitative analysis, or insects are sampled destructively, which precludes feeding experiments and the study of immature stages. Further, previous studies 2 failed to consider the phylogenetic relationships of host plants by using measures of host specificity that relied on counts of higher plant taxa (for example, genera or families). This approach can be misleading when taxonomic ranks are not commensurate with plant lineages. We examined the impacts of sampling bias and phylogenetic effects on estimates of host specificity by analysing the largest available data set of its kind. The leaf-chewing insect community on 51 plant species was characterized by using a sample

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Rubiaceae are one of the largest families of plants, with ;13,000 species. In this study, we have estimated the phylogeny for 534 Rubiaceae taxa from 329 genera with 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 taxa was used with 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 90.4 Ma. The estimated lineage divergence times for the subfamilies are 84.4 (86.6) Ma for Rubioideae, 73.1 Ma for Ixoroideae, and 73.1 Ma for Cinchonoideae. The estimated lineage divergence times for the tribes vary between 86.6 and 14.2 Ma. Classification, relationships, geographical distribution, and age estimates are presented and discussed for all tribes.

A phylogenetic study of plastid DNA sequences (ndhF, trnL/F, and rps16) in Lamiales is presented. In particular, the inclusiveness of Scrophulariaceae sensu APG II is elaborated. Scrophulariaceae in this sense are mainly a southern hemisphere group, which includes Hemimerideae (including Alonsoa, with a few South Americanspecies), Myoporeae, the Central American Leucophylleae (including Capraria), Androya, Aptosimeae, Buddlejeae, Teedieae (including Oftia, Dermatobotrys, and Freylinia), Manuleeae, and chiefly Northern temperate Scrophularieae (including Verbascum and Oreosolen). Camptoloma and Phygelius group with Buddlejeae and Teedieae, but without being well resolved to any of these two groups. Antherothamnus isstrongly supported as sister taxon to Scrophularieae. African Stilbaceae are shown to include Bowkerieae and Charadrophila. There is moderate support for a clade of putative Asian origin and including Phrymaceae,Paulownia, Rehmannia, Mazus, Lancea, and chiefly parasitic Orobanchaceae, to which Brandisia is shown to belong. A novel, strongly supported, clade of taxa earlier assigned to Scrophulariaceae was found. The clade includes Stemodiopsis, Torenia, Micranthemum and probably Picria and has unclear relationships to the restof Lamiales. This clade possibly represents the tribe Lindernieae, diagnosed by geniculate anterior filaments, usually with a basal swelling.

We present a phylogenetic dating of asterids, based on a 111-taxon tree representing all major groups and orders and 83 of the 102 families of asterids, with an underlying data set comprising six chloroplast DNA markers totaling 9914 positions. Phylogenetic dating was done with semiparametric rate smoothing by penalized likelihood. Confidence intervals were calculated by bootstrapping. Six reference fossils were used for calibration. To explore the effects of various sources of error, we repeated the analyses with alternative dating methods (nonparametric rate smoothing and the Langley-Fitch clock-based method), alternative tree topologies, reduced taxon sampling (22 of the 111 taxa deleted), partitioning the data into three genes and three noncoding regions, and calibrating with single reference fossils. The analyses with alternative topologies, reduced taxon sampling, and coding versus noncoding sequences all yielded small or in some cases no deviations. The choice of method influenced the age estimates of a few nodes considerably. Calibration with reference fossils is a critical issue, and use of single reference fossils yielded different results depending on the fossil. The bootstrap confidence intervals were generally small. Our results show that asterids and their major subgroups euasterids, campanulids, and lamiids diversified during the Early Cretaceous. Cornales, Ericales, and Aquifoliales also have crown node ages from the Early Cretaceous. Dipsacales and Solanales are from the Mid-Cretaceous, the other orders of core campanulids and core lamiids from the Late Cretaceous. The considerable diversity exhibited by asterids almost from their first appearance in the fossil record also supports an origin and first phase of diversification in the Early Cretaceous.

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• Premise of the study: The pantropical, species-rich Psychotrieae and Palicoureeae are sister tribes of mostly drupe-bearing and nonbacterial leaf-nodulating species with problematic generic limits. This problem is more complicated in Psychotrieae due to the paraphyly of the genus Psychotria, the lack of diagnostic characters for some major lineages, and the poor sampling from some biodiversity hotspots. Schizocarps and bacterial leaf nodules have been used for recognizing formal groups in Psychotrieae, but their evolution and taxonomic value have not been studied using a robust phylogeny of the tribe.• Methods: We analyzed 287 samples from the entire ranges of the tribes, with particular emphasis on the Western Indian Ocean region, with the Bayesian Markov chain Monte Carlo method.• Key results: All allied Psychotria genera investigated are nested within a paraphyletic Psychotria. Schizocarps evolved independently two times within Psychotria, and one reversal back to the drupaceous condition is inferred. The Malagasy leaf-nodulated Psychotrieae (except Apomuria bullata) and the Comorian non-leaf-nodulated Psychotria conocarpa are nested within the (African) leaf-nodulated clade. Within Palicoureeae, Chassalia is paraphyletic with respect to Geophila sensu stricto, and the Malagasy Geophila gerrardii and the African Hymenocoleus are closely related.• Conclusions: A widely circumscribed Psychotria encompassing the entire Psychotrieae is supported. Within Psychotria, two separate origins of schizocarps from drupes, one reversal back to the drupaceous condition, and two independent origins of the Malagasy leaf-nodulated species are inferred. A new genus Puffia is described to accommodate Geophila gerrardii, and a narrow circumscription of Chassalia is adopted. Thirty-two new combinations, two lectotypifications, and 25 new names are presented.