Summary• The family Araceae (3790 species, 117 genera) has one of the oldest fossil records among angiosperms. Ecologically, members of this family range from free-floating aquatics (Pistia and Lemna) to tropical epiphytes. Here, we infer some of the macroevolutionary processes that have led to the worldwide range of this family and test how the inclusion of fossil (formerly occupied) geographical ranges affects biogeographical reconstructions.• Using a complete genus-level phylogeny from plastid sequences and outgroups representing the 13 other Alismatales families, we estimate divergence times by applying different clock models and reconstruct range shifts under different models of past continental connectivity, with or without the incorporation of fossil locations.• Araceae began to diversify in the Early Cretaceous (when the breakup of Pangea was in its final stages), and all eight subfamilies existed before the K ⁄ T boundary. Early lineages persist in Laurasia, with several relatively recent entries into Africa, South America, South-East Asia and Australia.• Water-associated habitats appear to be ancestral in the family, and DNA substitution rates are especially high in free-floating Araceae. Past distributions inferred when fossils are included differ in nontrivial ways from those without fossils. Our complete genus-level time-scale for the Araceae may prove to be useful for ecological and physiological studies.

This study focuses on reconstructing the time‐calibrated phylogeny of the nine families comprising the order Sapindales, representing a diverse and economically important group of eudicots including citrus, mahogany, tree‐of‐heaven, cashew, mango, pistachio, frankincense, myrrh, lychee, rambutan, maple, and buckeye. We sampled three molecular markers, plastid genes rbcL and atpB, and the trnL‐trnLF spacer region, and covered one‐third of the generic diversity of Sapindales. All three markers produced congruent phylogenies using maximum likelihood and Bayesian methods for a set of taxa that included outgroups, i.e., members of the closely related orders Brassicales and Malvales, and the more distantly related Crossosomatales, Ranunculales, and Ceratophyllales. All results confirmed the current delimitation of the families within Sapindales, and the monophyly of the order. Concerning inter‐familial relationships, Biebersteiniaceae and Nitrariaceae formed a basal grade (or sister clade) to the rest of Sapindales with moderate support. The sister relationship of Kirkiaceae to Anacardiaceae and Burseraceae was strongly supported. The clade combining Anacardiaceae and Burseraceae as well as the clade combining Meliaceae, Simaroubaceae, and Rutaceae each received strong support. The sister relationship between Meliaceae and Simaroubaceae was moderately supported. The position of Sapindaceae could not be resolved with confidence. The Sapindales separated from their sister clade, comprising Brassicales and Malvales, in the Early Cretaceous at ca. 112 Ma, and diversified into the nine families from ca. 105 Ma until ca. 87 Ma during Early to Late Cretaceous times. Biebersteiniaceae and Nitrariaceae have the longest stem lineages observed in Sapindales, possibly indicating that extinction may have had a greater role in shaping their extant diversity than elsewhere within the order. Divergence within the larger families (Anacardiaceae, Burseraceae, Meliaceae, Rutaceae, Sapindaceae, Simaroubaceae) started during the Late Cretaceous, extending into the Paleogene and Neogene.

Alocasia comprises over 113 species of rainforest understorey plants in Southeast Asia, the Malesian region, and Australia. Several species, including giant taro, Alocasia macrorrhizos, and Chinese taro, Alocasia cucullata, are important food plants or ornamentals. We investigated the biogeography of this genus using plastid and nuclear DNA sequences (5200 nucleotides) from 78 accessions representing 71 species, plus 25 species representing 16 genera of the Pistia clade to which Alocasia belongs. Divergence times were inferred under strict and relaxed clock models, and ancestral areas with Bayesian and maximum likelihood approaches. Alocasia is monophyletic and sister to Colocasiagigantea from the SE Asian mainland, whereas the type species of Colocasia groups with Steudnera and Remusatia, requiring taxonomic realignments. Nuclear and plastid trees show topological conflict, with the nuclear tree reflecting morphological similarities, the plastid tree species' geographic proximity, suggesting chloroplast capture. The ancestor of Alocasia diverged from its mainland sister group c. 24 million years ago, and Borneo then played a central role in the expansion of Alocasia: 11-13 of 18-19 inferred dispersal events originated on Borneo. The Philippines were reached from Borneo 4-5 times in the Late Miocene and Early Pliocene, and the Asian mainland 6-7 times in the Pliocene. Domesticated giant taro originated on the Philippines, Chinese taro on the Asian mainland.

The mustard family (Brassicaceae) is a scientifically and economically important family, containing the model plant Arabidopsis thaliana and numerous crop species that feed billions worldwide. Despite its relevance, most published family phylogenies are incompletely sampled, generally contain massive polytomies, and/or show incongruent topologies between datasets. Here, we present the most complete Brassicaceae genus-level family phylogenies to date (Brassicaceae Tree of Life, or BrassiToL) based on nuclear (>1,000 genes, almost all 349 genera and 53 tribes) and plastome (60 genes, 79% of the genera, all tribes) data. We found cytonuclear discordance between nuclear and plastome-derived phylogenies, which is likely a result of rampant hybridisation among closely and more distantly related species, and highlight rogue taxa. To evaluate the impact of this rampant hybridisation on the nuclear phylogeny reconstruction, we performed four different sampling routines that increasingly removed variable data and likely paralogs. Our resulting cleaned subset of 297 nuclear genes revealed high support for the tribes, while support for the main lineages remained relatively low. Calibration based on the 20 most clock-like nuclear genes suggests a late Eocene to late Oligocene icehouse origin of the family. Finally, we propose five new or re-established tribes, including the recognition of Arabidopsideae, a monotypic tribe to accommodate Arabidopsis. With a worldwide community of thousands of researchers working on this family, our new, densely sampled family phylogeny will be an indispensable tool to further highlight Brassicaceae as an excellent model family for studies on biodiversity and plant biology.

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Heterotrophic plants provide intriguing examples of reductive evolution. This is especially evident in the reduction of their plastid genomes, which can potentially proceed toward complete genome loss. Several milestones at the beginning of this path of degradation have been described; however, little is known about the latest stages of plastome reduction. Here we analyze a diversity of plastid genomes in a set of closely related non-photosynthetic plants. We demonstrate how a gradual loss of genes shapes the miniaturized plastomes of these plants. The subject of our study, the genus Thismia, represents the mycoheterotrophic monocot family Thismiaceae, a group that may have experienced a very ancient (60–80 mya) transition to heterotrophy. In all 18 species examined, the plastome is reduced to 14–18 kb and is highly AT-biased. The most complete observed gene set includes accD, seven ribosomal protein genes, three rRNA, and two tRNA genes. Different clades of Thismia have undergone further gene loss (complete absence or pseudogenization) compared to this set: in particular, we report two independent losses of rps2 and rps18.

Plastid genomes exhibit different levels of variability in their sequences, depending on the respective kinds of genomic regions. Genes are usually more conserved while noncoding introns and spacers evolve at a faster pace. While a set of about thirty maximum variable noncoding genomic regions has been suggested to provide universally promising phylogenetic markers throughout angiosperms, applications often require several regions to be sequenced for many individuals. Our project aims to illuminate evolutionary relationships and species-limits in the genus Pyrus (Rosaceae)—a typical case with very low genetic distances between taxa. In this study, we have sequenced the plastid genome of Pyrus spinosa and aligned it to the already available P. pyrifolia sequence. The overall p-distance of the two Pyrus genomes was 0.00145. The intergenic spacers between ndhC–trnV, trnR–atpA, ndhF–rpl32, psbM–trnD, and trnQ–rps16 were the most variable regions, also comprising the highest total numbers of substitutions, indels and inversions (potentially informative characters). Our comparative analysis of further plastid genome pairs with similar low p-distances from Oenothera (representing another rosid), Olea (asterids) and Cymbidium (monocots) showed in each case a different ranking of genomic regions in terms of variability and potentially informative characters. Only two intergenic spacers (ndhF–rpl32 and trnK–rps16) were consistently found among the 30 top-ranked regions. We have mapped the occurrence of substitutions and microstructural mutations in the four genome pairs. High AT content in specific sequence elements seems to foster frequent mutations. We conclude that the variability among the fastest evolving plastid genomic regions is lineage-specific and thus cannot be precisely predicted across angiosperms. The often lineage-specific occurrence of stem-loop elements in the sequences of introns and spacers also governs lineage-specific mutations. Sequencing whole plastid genomes to find markers for evolutionary analyses is therefore particularly useful when overall genetic distances are low.

Reticulation events caused by hybridization are common and important sources of novelty in angiosperm evolution (Wood et al., 2009;Palfalvi et al., 2020). The detection, investigation, and representation of hybridization remains a challenge in phylogenomics (Kellogg, 2016;Mallet et al., 2016;Spooner et al., 2020). The combination of divergent genomes in hybrids (herein used for any organism that contains divergent genomes due to a hybridization event, e.g., many polyploids) introduces conflicting phylogenetic signal and can lead to topologically incorrect or poorly resolved phylogenetic trees (McDade, 1992;Soltis et al., 2008). However, the advancement of target capture data and universal probe kits such as Angiosperms353 (Johnson et al., 2018) provides an opportunity to gain insight into historical reticulations in angiosperm evolution and reduce phylogenetic conflict, if ortholog (or homeolog in polyploids) gene variants can be identified and separated (phased).Previously, inclusion of phased gene variants in phylogenetic studies has been used to confirm hybrid status of organisms (Sang and Zhang, 1999), determine the origin of polyploids (Popp and Oxelman, 2001), reveal parental lineages (Triplett et al., 2012;Estep et al., 2014), enable reconstruction of past reticulations (Estep et al., 2014;Brassac and Blattner, 2015), and date ancient hybridization events (Marcussen et al., 2015). Using Sanger sequencing, single-gene studies generated sequences for each variant separately using cloning (Sang and Zhang, 1999;Popp and Oxelman, 2001), and multi-gene studies linked these gene variants using their phylogenetic association in single-gene