Circumscription and phylogeny of the Laurales: evidence from molecular and morphological data.

The order Laurales comprises a few indisputed core constituents, namely Gomortegaceae, Hernandiaceae, Lauraceae, and Monimiaceae sensu lato, and an equal number of families that have recently been included in, or excluded from, the order, namely Amborellaceae, Calycanthaceae, Chloranthaceae, Idiospermaceae, and Trimeniaceae. In addition, the circumscription of the second largest family in the order, the Monimiaceae, has been problematic. I conducted two analyses, one on 82 rbcL sequences representing all putative Laurales and major lineages of basal angiosperms to clarify the composition of the order and to determine the relationships of the controversal families, and the other on a concatenated matrix of sequences from 28 taxa and six plastid genome regions (rbcL, rpl16, trnT-trnL, trnL-trnF, atpB-rbcL, and psbA-trnH) that together yielded 898 parsimony-informative characters. Fifteen morphological characters that play a key role in the evolution and classification of Laurales were analyzed on the most parsimonious molecular trees as well as being included directly in the analysis in a total evidence approach. The resulting trees strongly support the monophyly of the core Laurales (as listed above) plus Calycanthaceae and Idiospermaceae. Trimeniaceae form a clade with Illiciaceae, Schisandraceae, and Austrobaileyaceae, whereas Amborellaceae and Chloranthaceae represent isolated clades that cannot be placed securely based on rbcL alone. Within Laurales, the deepest split is between Calycanthaceae (including Idiospermaceae) and the remaining six families, which in turn form two clades, the Siparunaceae (Atherospermataceae-Gomortegaceae) and the Hernandiaceae (Monimiaceae s.str. [sensu stricto]-Lauraceae). Monimiaceae clearly are polyphyletic as long as they include Atherospermataceae and Siparunaceae. Several morphological character state changes are congruent with the molecular tree: (1) Calycanthaceae have disulculate tectate-columellate pollen, while their sister clade has inaperturate thin-exined pollen, with the exception of Atherospermataceae, which have columellate but meridionosulcate or disulcate pollen. (2) Calycanthaceae have two ventral ovules while their sister clade has solitary ovules. Within this sister clade, the Hernandiaceae (Lauraceae-Monimiaceae) have apical ovules, while the Siparunaceae (Atherospermataceae-Gomortegaceae) are inferred to ancestrally have basal ovules, a condition lost in Gomortega, the only lauralean genus with a syncarpous ovary. (3) Calycanthaceae lack floral nectaries (except for isolated nectarogeneous fields on the inner tepals), while their sister clade ancestrally has paired nectar glands on the filaments. Filament glands were independently lost in higher Monimiaceae and in Siparunaceae concomitant with pollinator changes away from nectar-foraging flies and bees to non-nectar feeding beetles and gall midges. (4) Disporangiate stamens with anthers dehiscing by two apically hinged valves are ancestral in Siparunaceae-(AtherospermataceaeGomortegaceae) and evolved independently within Hernandiaceae and Lauraceae. Depending on the correct placement of Calycanthaceae-like fossil flowers, tetrasporangiate anthers with valvate dehiscence (with the valves laterally hinged) may be ancestral in Laurales and lost in modern Calycanthaceae and Monimiaceae.