A global perspective on Campanulaceae: Biogeographic, genomic, and floral evolution.

PREMISE OF THE STUDY: The Campanulaceae are a diverse clade of flowering plants encompassing more than 2300 species in myriad habitats from tropical rainforests to arctic tundra. A robust, multigene phylogeny, including all major lineages, is presented to provide a broad, evolutionary perspective of this cosmopolitan clade. METHODS: We used a phylogenetic framework, in combination with divergence dating, ancestral range estimation, chromosome modeling, and morphological character reconstruction analyses to infer phylogenetic placement and timing of major biogeographic, genomic, and morphological changes in the history of the group and provide insights into the diversification of this clade across six continents. KEY RESULTS: Ancestral range estimation supports an out-of-Africa diversification following the Cretaceous-Tertiary extinction event. Chromosomal modeling, with corroboration from the distribution of synonymous substitutions among gene duplicates, provides evidence for as many as 20 genome-wide duplication events before large radiations. Morphological reconstructions support the hypothesis that switches in floral symmetry and anther dehiscence were important in the evolution of secondary pollen presentation mechanisms. CONCLUSIONS: This study provides a broad, phylogenetic perspective on the evolution of the Campanulaceae clade. The remarkable habitat diversity and cosmopolitan distribution of this lineage appears to be the result of a complex history of genome duplications and numerous long-distance dispersal events. We failed to find evidence for an ancestral polyploidy event for this clade, and our analyses indicate an ancestral base number of nine for the group. This study will serve as a framework for future studies in diverse areas of research in Campanulaceae.

Pollinator-mediated gene flow fosters genetic variability in a narrow alpine endemic, Abronia alpina (Nyctaginaceae).

PREMISE OF THE STUDY: For rare and endemic plants that exist in small, isolated habitats, natural selection is expected to favor self-compatibility, which can result in low genetic diversity due to inbreeding and genetic drift. Using Abronia alpina, a rare alpine endemic of the California Floristic Province, we demonstrate that there are exceptions to these predictions. METHODS: We present the results of both a pollination experiment and a genetic study using AFLPs (amplified fragment length polymorphisms). Using controlled hand-pollination and pollinator observations, we examined the breeding system, pollination ecology, and mechanism for self-incompatibility in A. alpina. KEY RESULTS: Abronia alpina exhibits an allogamous mating system with probable self-incompatibility resulting from limited growth of pollen tubes originating from self-pollination. Only xenogamous crosses and open-pollinated controls produced seed, and only xenogamous crosses produced pollen tubes that reached the ovary. The molecular study shows that A. alpina has substantial genetic diversity for a rare, endemic species, evidenced by the high percentage of polymorphic loci and average expected heterozygosity. Gene flow among subpopulations, as inferred from AFLP markers, appears to be substantial, although the Kern River is an important physical barrier. CONCLUSIONS: Our results indicate that A. alpina is dependent on insects for both seed production and the maintenance of genetic diversity. This finding suggests that pollinators may be crucial to the long-term adaptive potential of rare, endemic plants and that conservation of rare endemics is, in part, dependent on community-level interactions such as plant-pollinator mutualisms.

Nectar spur evolution in the Mexican lobelias (Campanulaceae: Lobelioideae).

Phylogenetic studies are often hampered by the independent evolution of characters that may potentially obscure relationships. The adaptive significance of the nectar spur and its evolution within the Mexican lobeliads (Campanulaceae) is considered here. The taxonomic delimitations of Heterotoma from the Mexican species within the genera Lobelia and Calcaratolobelia were tested. Independent molecular data were gathered to determine whether the Mexican spurred lobeliads should be treated as distinct genera. The internal transcribed spacer (ITS) region from 18-26S nuclear rDNA and chloroplast DNA from the 3' trnK intron were sequenced from 14 representative species. Our data suggest that Heterotoma, as originally conceived, is a good evolutionary unit within Lobelia and that the presence of a nectar spur is an important morphological character that can be used in defining phylogenetic position. This study also suggests that morphological changes associated with hummingbird pollination have evolved more than once in the Mexican lobeliads, from small blue-flowered, insect-pollinated relatives.