Background In the large angiosperm order Lamiales, a diverse array of highly specialized life strategies such as carnivory, parasitism, epiphytism, and desiccation tolerance occur, and some lineages possess drastically accelerated DNA substitutional rates or miniaturized genomes. Gesneriaceae plus Calceolariaceae, which is also supported by morphological heroes. Plantaginaceae (incl. Gratioleae) and Scrophulariaceae are well separated in the backbone grade; Lamiaceae and Verbenaceae appear in distant clades, while the recently explained Linderniaceae are confirmed to become monophyletic and in an isolated position. Conclusions Confidence about deep nodes of CD86 the Lamiales tree is an important step towards understanding the evolutionary diversification of a major clade of flowering vegetation. The degree of resolution acquired here now provides a first opportunity to discuss the development of morphological and biochemical qualities in Lamiales. The multiple self-employed evolution of the carnivorous syndrome, once in Lentibulariaceae and a second time in Byblidaceae, is definitely strongly supported by all analyses and topological checks. The development of selected morphological characters such as flower symmetry is definitely discussed. The XL184 addition of further sequence data from introns and spacers keeps promise to eventually obtain a fully resolved plastid tree of Lamiales. Background With more than 23,000 varieties in at least 23 family members [1], Lamiales (eudicots/asterids) are one of the largest orders of flowering vegetation, with associates found all over the world. The highest diversity is contributed by herbaceous vegetation with mono-symmetric blossoms. Some users are economically important, such as Lamiaceae (pot-herbs like mint, sage, oregano or basil), Oleaceae (olives), Pedaliaceae (sesame), Verbenaceae (timber, medicinal) Plantaginaceae (medicines like digitalis, ornamentals) and Scrophulariaceae (ornamentals). The order consists of lineages with highly specialized existence forms and qualities of particular medical interest. So far, their comparative study has been limited by the lack of a powerful phylogenetic platform for Lamiales. Desiccation-tolerant users (so-called “resurrection vegetation”, see Number ?Number1a)1a) of the recently described family Linderniaceae [2] are a focus of molecular and evolutionary studies [3,2]. XL184 Great metabolic and genomic shifts are exhibited by parasitic vegetation. With Orobanchaceae, Lamiales harbor the largest quantity of parasitic angiosperms (Number ?(Figure1b).1b). The family comprises both hemi- and holoparasites [4], with some varieties causing serious damage in agriculture [5]. Chloroplast genomes of users of Orobanchaceae display gene order rearrangements, high evolutionary rates and gene deficits, potentially as a consequence of parasitism with this family. One line of current study in the family concentrates on progressive plastid development under increasingly peaceful practical constraints [Wicke et al., in prep]. Number 1 Example taxa from Lamiales, showing associates of desiccation-tolerant, parasitic, and carnivorous lineages, as well as users from family members regularly referred to in the text. a: the desiccation-tolerant Craterostigma pumilum XL184 from Linderniaceae; … Carnivory in Lamiales Lentibulariaceae, probably the most species-rich family of carnivorous vegetation (ca. 350 spp.) belongs to Lamiales (Number 1c, d). This family is unique for a variety of reasons: traps of Utricularia (bladderworts) are regarded as a complex changes of leaves [6,7], and the typical angiosperm body strategy is definitely strongly relaxed in users of this genus [8-10]. Utricularia and its sister genus, Genlisea (the corkscrew vegetation), are the only carnivorous angiosperms known to feed on protozoa [11]. They have the smallest holoploid genome sizes among angiosperms, with some nuclear genomes as small as 63 Mbp or less [12], and show the highest relative DNA substitution rates for some of the investigated chloroplast genome areas [13,14]. Pinguicula (butterworts), the third genus of Lentibulariaceae, is definitely far less intense in genome size, substitution rate and morphology, and exhibits glandular leaves that function as adhesive (“flypaper”) traps (Number 1c, d). Apart from Lentibulariaceae, the monogeneric Australian family Byblidaceae (Number ?(Figure1e)1e) also attracts and catches insects with simple flypaper traps similar in function to the people of Pinguicula. The carnivorous syndrome of Byblis was questioned by some authors, as the vegetation were considered to lack their personal digestive enzymes and have not been demonstrated to be able to take up released nutrients, thus being rated as merely “protocarnivorous” [15]. However, a recent study [16] recognized phosphatase activity, therefore repairing the rank of carnivory to Byblis. Morphological links -.