BackgroundVarious expansions or contractions of inverted repeats (IRs) in chloroplast genomes led to fluxes in the IR-LSC (large single copy) junctions. Previous studies revealed that some monocot IRs contain a trnH-rps19 gene cluster, and it has been speculated that this may be an evidence of a duplication event prior to the divergence of monocot lineages. Therefore, we compared the organizations of genes flanking two IR-LSC junctions in 123 angiosperm representatives to uncover the evolutionary dynamics of IR-LSC junctions in basal angiosperms and monocots.ResultsThe organizations of genes flanking IR-LSC junctions in angiosperms can be classified into three types. Generally each IR of monocots contains a trnH-rps19 gene cluster near the IR-LSC junctions, which differs from those in non-monocot angiosperms. Moreover, IRs expanded more progressively in monocots than in non-monocot angiosperms. IR-LSC junctions commonly occurred at polyA tract or A-rich regions in angiosperms. Our RT-PCR assays indicate that in monocot IRA the trnH-rps19 gene cluster is regulated by two opposing promoters, S10A and psbA.ConclusionTwo hypotheses are proposed to account for the evolution of IR expansions in monocots. Based on our observations, the inclusion of a trnH-rps19 cluster in majority of monocot IRs could be reasonably explained by the hypothesis that a DSB event first occurred at IRB and led to the expansion of IRs to trnH, followed by a successive DSB event within IRA and lead to the expansion of IRs to rps19 or to rpl22 so far. This implies that the duplication of trnH-rps19 gene cluster was prior to the diversification of extant monocot lineages. The duplicated trnH genes in the IRB of most monocots and non-monocot angiosperms have distinct fates, which are likely regulated by different expression levels of S10A and S10B promoters. Further study is needed to unravel the evolutionary significance of IR expansion in more recently diverged monocots.

Generic delimitation of Cyathocalyx and Drepananthus has been controversial, with some authors recognizing them as distinct genera, and others recognizing a more broadly defined Cyathocalyx, inclusive of Drepananthus. Some doubt also exists regarding the relationships between these taxa and Cananga. Molecular phylogenetic analyses are presented based on combined psbA‐trnH spacer, trnL‐F, matK and rbcL sequences. Results indicate that Cananga, Cyathocalyx s.str. and Drepananthus form three generally well‐supported clades, although with inadequate resolution of relationships among the three clades. Morphological variation is re‐evaluated, and the narrower delimitation of Cyathocalyx proposed, necessitating 21 new nomenclatural combinations following the recognition of Drepananthus as a distinct genus. Divergence times are estimated using an uncorrelated lognormal distributed (UCLD) relaxed molecular clock. Historical biogeographical analysis suggests that the ambavioid lineage originated in Africa, with subsequent dispersal into Asia. Alternative hypotheses for this dispersal, involving rafting on the Indian tectonic plate versus migration via the extensive boreotropical forests associated with the Eocene thermal maximum, are evaluated, and the latter route identified as the most consistent with the divergence age estimates and the geological and palaeoclimatic data.