Structure des communautés de Phanaeini (Coleoptera : Scarabaeidae) de Guyane : étude par échantillonnage massif au piège d’interception

Summary

Seasonality and community structure of Phanaeini (Coleoptera: Scarabaeidae) in French Guiana : study by mass sampling using large flight interception traps. Phanaeini is a neotropical tribe of scarabs mostly dung or carrion feeders, generally of large size. This paper analyses data collected with large window flight interception traps set on nine forest sites in French Guiana with a primary goal of biotic inventory. The study deals with three main questions: 1) What is the spatial structure of communities and are there some species indicators of secondary forests and others of pristine conditions? 2) What is the temporal structure of the community and is there a stable pattern of seasonality between years and sites? 3) Between close species, is there a temporal asynchrony? Our samples contain more than 9,700 identified specimens, nine complete year series on four sites including a four years continuous survey near Cayenne. We found a significantly lower diversity (Shannon and Simpson index) on the most fragmented and hunted sites. A more equitable repartition of species and a relative abundance of the larger species appear typical of undisturbed sites. Other observations reinforce the hypothesis that there is a fast and huge modification in Phanaeini community structure on the most accessible and disturbed sites. Seasonal pattern shows an abundance peak at the beginning of the rainy season (December or January), a medium abundance during the rainy season with sometimes secondary peaks and a low to very low activity during the dry season. The pattern is rather consistent between years but changes with sampling site. It is however different from the results of other studies using pitfall baited traps in Amazonian and Guyanese forests. These studies show much less clear temporal pattern or no seasonal change. There is no obvious pattern of niche sharing by phenological differences between species. The methodological differences of sampling between baited traps and window flight traps are eventually discussed. Due to its passive way of collecting, interception trap is considered as quite relevant for studying flight activity spatio-temporal patterns of Scarabaeinae. This aspect may explain some differences in seasonality patterns compared to other studies.     

The mitogenomes of three beetles (Coleoptera: Polyphaga: Cucujiformia): New gene rearrangement and phylogeny

The mitochondrial genome (mitogenome) has been extensively used for studying phylogenetic relationships at different taxonomic levels. Several molecular analyses have been performed, but the phylogenetic relationships among infraorders in Polyphaga have not been well resolved. In this work, three nearly complete mitogenomes of Coleoptera, Sitophilus oryzae, Oryzaephilus surinamensis and Callosobruchus chinensis, were determined. The O. surinamensis and S. oryzae mitogenomes harbor gene content typical of other Polyphaga mitogenomes, while a gene rearrangement (trnQ) was found in the C. chinensis mitogenome. The mitogenomes of these three Coleoptera species each consist of approximately 13 protein-coding genes, 22 tRNA genes, two rRNA genes and one A + T-rich region. Phylogenetic analysis within Polyphaga was carried out based on mitochondrial data. The phylogenetic results within Polyphaga support the basal position of Cyphon sp., which belonged to Scirtoidea, Elateriformia. Within Cucujiformia, monophyletic Curculionoidea, Chrysomeloidea and Tenebrionoidea were confirmed.     

The mitochondrial genome of Gatzara jezoensis (Neuroptera: Myrmeleontidae) and phylogenetic analysis of Neuroptera

Phylogenetic relationship within Neuroptera is controversial, particularly for the various hypotheses based on both morphological and molecular evidence. In the present study, we determined the complete mitochondrial genome (mitogenome) of Gatzara jezoensis, which is the second representative of the tribe Dendroleontini. The G. jezoensis mitogenome contained the conserved set of 37 mitochondrial genes and a putative control region, with a conserved gene arrangement which was similar to that of most sequenced neuropteran mitogenomes. All transfer RNAs exhibited the canonical cloverleaf secondary structure, except for trnS^(AGN). The control region contained two conserved elements (ploy-T stretch and ATGGTTCAAYAAAATAAYYCYCTC motif) and abundant microsatellite-like elements. The phylogenetic analysis of sequenced neuropteran mitogenomes using the concatenated protein-coding genes (PCGs) and ribosomal genes recovered the monophyly of Myrmeleontidae, which revealed this dataset could generate the more robust phylogeny of Neuroptera than that of 13 PCGs dataset.     

Chromosomal diversification of diploid number,heterochromatin and rDNAs in two species of Phanaeus beetles (Scarabaeidae,Scarabaeinae)

The genus Phanaeus is included in the tribe Phanaeini, one of the most diverse tribes within the subfamily Scarabaeinae in terms of chromosomal characteristics. However, so far the species of this genus were not studied with differential cytogenetic techniques, limiting any inference of the probable mechanisms responsible for this diversity. In this work, several techniques were applied with the aim of cytogenetically characterizing two Phanaeus species. The karyotype found for Phanaeus (Notiophanaeus) chalcomelas was 2n = 12, neo-XY, and that of P. (N.) splendidulus was 2n = 20, Xy_p, considered primitive for the family Scarabaeidae. The chromosomes of both species showed a high amount of constitutive heterochromatin (CH), with blocks rich in base pairs GC (CMA₃⁺). Moreover, in P. (N.) chalcomelas the marks revealed by C-banding and fluorochrome staining were different in size, showing CH variability. Sites of 18S ribosomal DNA (rDNA) were identified in one autosomal pair of P. (N.) chalcomelas and in five autosomal pairs of P. (N.) splendidulus. On the other hand, only one autosomal pair exhibited 5S rDNA sequences in these species. The results suggest that the karyotype differentiation of the Phanaeus species studied here involved pericentric inversions and centric fusions, as well as mechanisms related to amplification and dispersion of CH and rDNA sequences.     

Mitochondrial genome of the yellow catfish Pelteobagrus fulvidraco and insights into Bagridae phylogenetics

The mitochondrial genome (mitogenome) can provide important information for understanding phylogenetic analysis and molecular evolution. Herein, we amplified the complete mitogenome sequence of Pelteobagrus fulvidraco. The mitogenome was 16,526 bp in length and included 13 protein-coding genes (PCGs), 22 transfer RNA genes, two ribosomal RNA genes and a non-coding control region (D-loop). Both the organization and location of genes in the mitogenome were consistent with those from Siluriformes fishes previously published in GenBank. The phylogenetic relationships based on Bayesian inference (BI) and Maximum likelihood (ML) methods showed that P. fulvidraco has close relationships with Pelteobagrus eupogon and Tachysurus intermedius, suggesting that P. fulvidraco belongs to Tachysurus. This study provides evidence that Tachysurus, Pseudobagrus and Leiocassis do not form monophyly, but that these three genera form a monophyletic group. Our results provide reference for further phylogenetic research of the Bagridae species.     

Insufficient resolving power of mitogenome data in deciphering deep phylogeny of Holometabola

The mitochondrial genome (mitogenome) is one of the most widely used markers for phylogenetic analysis. Compared with whole-genome data, mitogenome data are less expensive to obtain and easier to manipulate. However, compositional bias and accelerated evolutionary rate reduce the effectiveness of the mitogenome in determining insect phylogeny. This study shows that mitogenome data are not suitable to reconstruct deep holometabolan evolution, even with a most comprehensive data coding scheme and the more realistic CAT model. For the deep levels of divergence within Holometabola, protein-coding genes only retain weak phylogenetic signals, leading to peculiar interordinal relationships. Consensus relationships in the Holometabola phylogeny, such as the monophyly of Holometabola, the most basal position of Hymenoptera, and the sister group relationship between the Strepsiptera and Coleoptera were rarely resolved in our analyses. The relationships of the holometabolan groups as inferred by mitogenomes are highly vulnerable to gene types, data coding regimes, model choice, and optimality criteria, and no consistent alternative hypothesis of Holometabola's relationships is supported. Thus, we suggest that the slowly evolving nuclear genes or genome-scale approaches may be better options for resolving deep-level phylogeny of Holometabola.     

The complete mitochondrial genome sequence of the little egret (Egretta garzetta)

Many phylogenetic questions in the Ciconiiformes remain unresolved and complete mitogenome data are urgently needed for further molecular investigation. In this work, we determined the complete mitogenome sequence of the little egret (Egretta garzetta). The genome was 17,361 bp in length and the gene organization was typical of other avian mtDNA. In protein-coding genes (PCGs), a C insertion was found in ND3, and COIII and ND4 terminated with incomplete stop codons (T). tRNA-Val and tRNA-Ser (AGY) were unable to fold into canonical cloverleaf secondary structures because they had lost the DHU arms. Long repetitive sequences consisting of five types of tandem repeats were found at the 3′ end of Domain III in the control region. A phylogenetic analysis of 11 species of Ciconiiformes was done using complete mitogenome data and 12 PCGs. The tree topologies obtained with these two strategies were identical, which strongly confirmed the monophyly of Ardeidae, Threskiorothidae and Ciconiidae. The phylogenetic analysis also revealed that Egretta was more closely related to Ardea than to Nycticorax in the Ardeidae, and Platalea was more closely related to Threskiornis than to Nipponia in the Threskiornithidae. These findings contribute to our understanding of the phylogenetic relationships of Ciconiiformes based on complete mitogenome data.     

The complete mitochondrial genome of <Emphasis Type="Italic">Damora sagana</Emphasis> and phylogenetic analyses of the family Nymphalidae

The monotypic genus Damora (Nymphalidae, Heliconiinae) contains a single species, Damora sagana, which is widely distributed across southern China. Herein, its complete mitogenome was sequenced to further understand lepidopteran mitogenome characteristics, reconstruct the nymphalid family phylogeny, and infer the subdivision of Heliconiinae species. The circular mitogenome was 15,151 bp long, abundant in A and T, and comprised of 13 protein-coding genes (PCGs), 22 transfer RNA (tRNA) genes, 2 ribosomal RNA (rRNA) genes, and one control region with a gene arrangement typical of lepidopteran mitogenomes. ATN codons initiated all PCGs, except cytochrome c oxidase subunit 1 (COX1), which was initiated by a CGA sequence as has been observed in other lepidopterans. Three PCGs (COX1, COX2 and ND4) employed a single T termination signal, whereas others had the typical complete termination codon (TAA). All tRNA genes were folded into the typical cloverleaf structure except for tRNA-Ser (AGN). The A+T-rich region included the conserved motif ‘ATAGA’ followed by a 17 bp poly-T stretch, which was also observed in tribe Argynnini mitogenomes. A phylogenetic tree was constructed via multiple methods using the 13 PCGs data of D. sagana and other available mitogenomes of nymphalid species. All three phylogenetic trees yielded the same topology. These results were consistent with those from previous studies of most major nymphalid groups, except those regarding tribe subdivision in certain subfamilies such as Argynnini + (Acraeini + Heliconiini) for Heliconiine. Furthermore, our analyses identified that the genus Cethosia was grouped with the genus Acraea composing the tribe Acraeini with strong support.     

Molecular phylogeny and systematics of the tribe Chorisporeae (Brassicaceae)

Sequence data from nuclear (ITS) and chloroplast (trnL-F) regions for 89 accessions representing 56 out of 64 species from all five genera of the tribe Chorisporeae (plus Dontostemon tibeticus) have been studied to test the monophyly of the tribe and its component genera, clarify its boundaries, and elucidate its phylogenetic position in the family. Both data sets showed strong support for the monophyly of the Chorisporeae as currently delimited, though the position of its tentative member D. tibeticus was not resolved by ITS. Parrya and Pseudoclausia are poly- and paraphyletic with regard to each other, and Chorispora is either polyphyletic or at least paraphyletic (comprising Diptychocarpus) within a weakly supported monophyletic clade. The incongruence in branching pattern among the markers was most likely caused by hybridization and possibly influenced by incomplete lineage sorting. The present results suggest uniting Pseudoclausia, Clausia podlechii, and Achoriphragma with Parrya and transferring P. beketovii and P. saposhnikovii to Leiospora (Euclidieae). We also obtained support for splitting Chorispora into two geographically defined groups, one of which is closer to Diptychocarpus. Both data sets revealed a close relationship of the Chorisporeae to Dontostemoneae, while ITS also indicated affinity to Hesperideae. Therefore, the position of Chorisporeae needs further verification.     

Phylogenetic relationships,subdivision, and biogeography of the cyprinid tribe Labeonini (sensu ) (Teleostei: Cypriniformes), with comments on the implications of lips and associated structures in the labeonin classification

The Labeonini (sensu Rainboth, 1991) is a tribe of the subfamily Cyprininae, the largest subfamily of Cypriniformes. With around 400 species in 34 genera, this tribe is widely distributed in the freshwaters of tropical Africa and Asia. Most species are adapted to fast-flowing streams and rivers, and exhibit unique morphological modifications associated with their lips and other structures around the mouth. The monophyly of this tribe has been tested and generally accepted in previous morphological and molecular studies. The major objectives of this study were to reconstruct the phylogenetic relationships within the tribe Labeonini, test its monophyly and explore the taxonomic subdivisions, intrarelationships and biogeography of the group. The value of the morphological characters associated with the lips and other associated structures in the taxonomic classification of labeonins was also discussed. Nucleotide sequences (3867 bp) of four unlinked nuclear loci were obtained from 51 species in 18 Labeonini genera from throughout the range of the tribe. Maximum parsimony, partitioned maximum likelihood and partitioned Bayesian analyses were used for phylogenetic inference from combined and separate gene data sets. Based on our results, the monophyly of Labeonini was well supported. Two major clades could be recovered within the tribe. Three subclades could further be recognized from the first clade. These clades/subclades are not consistent with groupings of any of previous workers using either morphological or molecular characters for phylogenetic inference. Only five currently recognized genera in this analysis are monophyletic. The similarity between some lips and associated structures (e.g. suctorial discs) of labeonins may due to convergence or parallelism instead of common ancestry. Labeonins of Southeast Asia, India and China are closely related to each other; the multiple clades of African taxa do not form a single monophyletic group, indicating multiple, independent dispersal events of labeonins into Africa from Asia.     

Analysis of phylogenetic relationships of Brassicaceae species based on Chs sequences

Sequences of nuclear chalcone synthase gene (Chs) were analyzed for species of the Brassicaceae family to reconstruct phylogenetic relationships. The phylogeny for 106 species of 60 genera was reconstructed, and assigned to 24 tribes, using maximum parsimony, maximum likelihood, and neighbor-joining methods. Most of the tribes can be assigned to the major lineages (Lineages I–III) suggested by Beilstein et al. (2006). The tribe Camelineae was not monophyletic. Conringia planisiliqua together with Orychophragmus violaceus would not be recognized as a new tribe proposed by the previous studies, and C. planisiliqua should be a member of tribe Isatideae. The genera delimitation and monophyly of the expanded Solms-laubachia were also confirmed by our data. Furthermore, one parent of inter-tribal allopolyploid Pachycladon appeared to be most closely associated with Crucihimalaya, Transberingia and tribes Boechereae and Halimolobeae, another parent was proved to be in tribe Smelowskieae.     

Complete mitochondrial genome of the darkling beetle Gonocephalum outreyi (Coleoptera: Tenebrionidae) with phylogenetic implications

The complete mitochondrial genome (mitogenome) of Gonocephalum outreyi was determined by using next-generation sequencing approach. The full length of this mitogenome is 15,836?bp, which consists of 37 typical metazoan mitochondrial genes with an identical genome organization to ancestral insects. The majority of the protein-coding genes begin with the codon ATN, except for cox1 and cox2 with AAT and AAA, respectively. To elucidate the phylogenetic position of G. outreyi, we used various sequence coding schemes for protein-coding genes and the combined nucleotide sequences of all mitochondrial genes for tree building under the Bayesian and Maximum Likelihood inferences. The phylogenetic results consistently supported G. outreyi as a member of the family Tenebrionidae. The monophyly of both Tenebrionoidea and Tenebrionidae were strongly supported. The Scraptiidae and Melandryidae were recovered to be non-monophyletic in regards to the Osphya. Within Tenebrionidae, the subfamilies Diaperinae and Tenebrioninae were found to be non-monophyletic.     

Characterization of the complete mitochondrial genome of Hydrotaea spinigera (Diptera: Muscidae) with phylogenetic implications

Hydrotaea spinigera Stein is a muscid species with high forensic importance. To further supplement the genome-level features of related species, the complete mitochondrial genome of H. spinigera is amplified, sequenced, annotated, analyzed, and compared with 13 other species of the family Muscidae. This mitogenome is 15,517?bp in length, with a standard set of 13 protein-coding genes (PCGs), 22 transfer RNA (tRNA) genes, two ribosomal RNA (rRNA) genes, and a typical non-coding control region. The gene organization, base composition, and secondary structures of tRNA genes of this mitogenome are similar to other dipteran species. Phylogenetic analyses based on all the mitochondrial PCGs and rRNA genes for 14 muscid and one outgroup taxa are performed using Bayesian inference (BI) and maximum likelihood (ML) analysis. The inferred trees indicate that the Muscidae, the subfamily Muscinae, and the tribes Azeliini and Reinwardtiini are monophyletic, whereas the monophyly of the subfamily Azeliinae and the tribe Muscini is not supported.     

Complete mitochondrial genome sequence of bighead croaker Collichthys niveatus (Perciformes, Sciaenidae): a mitogenomic perspective on the phylogenetic relationships of Pseudosciaeniae

The monophyly and phylogenetic relationships of Pseudosciaeniae have long been controversial. Here we describe the mitochondrial genome (mitogenome) sequence of Collichthys niveatus. It is a circular double-stranded DNA molecule of 16,450 base pairs (bp) in length with a standard set of 22 transfer RNA genes (tRNAs), 2 ribosomal RNA genes (rRNAs), 13 protein-coding genes as well as a non-coding control region. The mitogenome of C. niveatus shared common features with those of other bony fishes in terms of gene arrangement, base composition, and tRNA structures. The C. niveatus mitogenome exhibited pronounced strand-specific asymmetry in nucleotide composition, which was also reflected in the codon usage of genes oriented in opposite directions. Contrary to the typical structure of the control region, the central conserved blocks (CSB-D, -E, and -F) could not be detected in C. niveatus mitogenome. Phylogenetic analysis based on whole mitogenome sequences provided strong support for the monophyly of Pseudosciaeniae, and sister-group relationships of C. niveatus + Collichthys lucidus and Larimichthys crocea + Larimichthys polyactis, which was consistent with the traditional taxonomy. Unexpected divergence was found in two C. niveatus mitogenomes and several hypotheses were proposed to explain this observation including misidentification and introgressive hybridization between C. niveatus and L. polyactis, and polyphyletic origin of C. niveatus. We considered species misidentification to be the main hypothesis. However, additional data is essential to test these proposed hypotheses.     

Characterization of the mitochondrial genome and phylogeny of the black arowana (Osteoglossum ferreirai)

In this study, we sequenced and assembled the mitochondrial (mt) genome of Osteoglossum ferreirai to re-assess the phylogenetic relationship of the family Osteoglossidae. We determined that the mitogenome of O. ferreirai contains the entire set of 37 mt genes, and the nucleotide composition and gene arrangement were similar to those of other bonytongues. Our phylogenetic analyses exhibited monophyly of the family Osteoglossidae with high bootstrap support, which is in agreement with the currently accepted phylogenetic viewpoint that is based on both morphological and molecular approaches. These findings provide additional informative data for the further study of phylogenetic relationships and help to elucidate a key component of the species radiation process within the family Osteoglossidae.     

Phylogenetic position of the genus Ferula (Apiaceae) and its placement in tribe Scandiceae as inferred from nrDNA ITS sequence variation

Recent molecular systematic investigations suggested that Ferula, an umbellifer genus of about 170 species, is polyphyletic, with its members placed in the apioid superclade and within tribe Scandiceae. We analyzed ITS sequence variation from 134 accessions of Apiaceae, including 83 accessions (74 species) of Ferula to ascertain the phylogenetic position of the genus within the family. Phylogenetic analyses of these data using maximum parsimony, Bayesian, and neighbor-joining methods support the monophyly of Ferula upon the addition of Dorema and Leutea (as Ferula sensu lato) and its placement in tribe Scandiceae. Ferula sensu is closely allied with other major lineages of Scandiceae, corresponding to subtribes Scandicinae, Daucinae, and Torilidinae. Therefore, we recognize the Ferula clade as subtribe Ferulinae. Another addition to tribe Scandiceae is a clade composed of genera Glaucosciadium and Mozaffariania. The three accessions of Ferula misplaced in the apioid superclade represent a species of Silaum.     

The Chloroplast Genome of Hyoscyamus niger and a Phylogenetic Study of the Tribe Hyoscyameae (Solanaceae)

The tribe Hyoscyameae (Solanaceae) is restricted to Eurasia and includes the genera Archihyoscyamus, Anisodus, Atropa, Atropanthe, Hyoscyamus, Physochlaina, Przewalskia and Scopolia. Even though the monophyly of Hyoscyameae is strongly supported, the relationships of the taxa within the tribe remain unclear. Chloroplast markers have been widely used to elucidate plant relationships at low taxonomic levels. Identification of variable chloroplast intergenic regions has been developed based on comparative genomics of chloroplast genomes, but these regions have a narrow phylogenetic utility. In this study, we present the chloroplast genome sequence of Hyoscyamus niger and make comparisons to other solanaceous plastid genomes in terms of gene order, gene and intron content, editing sites, origins of replication, repeats, and hypothetical open reading frames. We developed and sequenced three variable plastid markers from eight species to elucidate relationships within the tribe Hyoscyameae. The presence of a horizontally transferred intron in the mitochondrial cox1 gene of some species of the tribe is considered here a likely synapomorphy uniting five genera of the Hyoscyameae. Alternatively, the cox1 intron could be a homoplasious character acquired twice within the tribe. A homoplasious inversion in the intergenic plastid spacer trnC-psbM was recognized as a source of bias and removed from the data set used in the phylogenetic analyses. Almost 12 kb of plastid sequence data were not sufficient to completely resolve relationships among genera of Hyoscyameae but some clades were identified. Two alternative hypotheses of the evolution of the genera within the tribe are proposed.     

Complete mitochondrial genome sequence of Bonasa sewerzowi(Galliformes: Phasianidae) and phylogenetic analysis

Bonasa sewerzowi, the smallest and most southerly distributed grouse species in the world, is a bird endemic to China. The population of B. sewerzowi had shown a declining trend, which made it to be the endangered species in the China Red Data Book and Category I of nationally protected animals. So far, however, most studies about this species were mainly focused on the morphological and ecological aspects. In order to further study the feature of B. sewerzowi, the complete mitochondrial genome(mitogenome) of B. sewerzowi was sequenced by Illumina Hiseq 2000 high-throughput sequencing. Then, we focused on comparative genomics of two Bonasa species to find their characteristics. Finally, phylogenetic position of Bonasa was made based on the mitogenome dataset. Our results revealed that:(1) the mitogenome of B. sewerzowi, consisting of 16 658 bp, displayed typical genome organization and gene order found in other previously determined Galliformes mitogenomes;(2) the structure and composition of mitogenomes were similar between B. sewerzowi and B. bonasia;(3) the monophyly of Bonasa was well supported, which had a closer phylogenetic relationship with Meleagris gallopavo.     

Phylogenetic Systematics of Peccaries (Tayassuidae: Artiodactyla) and a Classification of South American Tayassuids

Tayassuidae is a family of pig-like Artiodactyla restricted to the New World. Despite its rich fossil history, they have received less attention from a taxonomic and phylogenetic perspective when compared to the Old World pigs, Suidae. In this study, we performed a computer assisted phylogenetic analysis using morphological and molecular data including fossil and extant Tayassuidae, using parsimony and Bayesian approaches. We recovered the monophyly of the family Tayassuidae, confirming previous proposals, as well as the monophyly of the subfamilies Hesperhyinae and Tayassuinae, and the genus Platygonus, which we placed in a new taxon of tribe level. The three living peccaries and a number of fossil species belong to a new, tribe level, monophyletic group. The genus Catagonus comes out as paraphyletic, leading us to propose to restrict the generic name to the type species, C. metropolitanus, and a new taxonomic arrangement for the remaining species previously included in it, revalidating the genera Brasiliochoerus and Parachoerus, and describing a new genus, Protherohyus, gen. nov.     

Speciation and evolution of eco‐climatic ranges in the intertropical African dung beetle genus,Diastellopalpus van Lansberge

We investigate how late Cenozoic orogenics and climatic change might have influenced the history of taxon diversification and current species ranges of an endemic, Afrotropical, insect genus. Diastellopalpus van Lansberge is a near basally‐derived taxon in the dung beetle tribe Onthophagini (Coleoptera: Scarabaeidae: Scarabaeinae) that has diversified into 32 known species primarily centred on intertropical forests. Basal dichotomies in both published and re‐analysed phylogenies divide the species into clades that are geographically centred either to the east or west of the south‐east highlands that underwent uplift from the Miocene. There is broad climatic overlap between many of the species but clear separation along a minimum spanning tree in ordinal space where they are divided into taxa with either lowland or highland centres of distribution. Observed spatial distributions of six defined species groups mostly differ from predicted climatic ranges, presumably as a result of historical constraints on species dispersal. A trend from dominance of montane or wet lowland forest associations in species lineages derived from more basal nodes (Groups A–C) to dominance of drier upland forest and moist woodland associations in species lineages derived from a more terminal node (Groups D–F) is perhaps linked to the stepped trend to cooler, dryer climate in the late Cenozoic. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 407–423.