Tina J. Ayers

PLANNED FLOODING AND COLORADO RIVER RIPARIAN TRADE‐OFFS DOWNSTREAM FROM GLEN CANYON DAM, ARIZONA

Regulated river restoration through planned flooding involves trade-offs be- tween aquatic and terrestrial components, between relict pre-dam and novel post-dam re- sources and processes, and between management of individual resources and ecosystem characteristics. We review the terrestrial (wetland and riparian) impacts of a 1274 m 3 /s test flood conducted by the U.S. Bureau of Reclamation in March/April 1996, which was designed to improve understanding of sediment transport and management downstream from Glen Canyon Dam in the Colorado River ecosystem. The test flood successfully restored sandbars throughout the river corridor and was timed to prevent direct impacts to species of concern. A total of 1275 endangered Kanab ambersnail (Oxyloma haydeni kan- abensis) were translocated above the flood zone at Vaseys Paradise spring, and an estimated 10.7% of the total snail habitat and 7.7% of the total snail population were lost to the flood. The test flood scoured channel margin wetlands, including potential foraging habitats of endangered Southwestern Willow Flycatcher (Empidonax traillii extimus). It also buried ground-covering riparian vegetation under .1 m of fine sand but only slightly altered woody sandbar vegetation and some return-current channel marshes. Pre-flood control ef- forts and appropriate flood timing limited recruitment of four common nonnative perennial plant species. Slight impacts on ethnobotanical resources were detected .430 km down- stream, but those plant assemblages recovered rapidly. Careful design of planned flood hydrograph shape and seasonal timing is required to mitigate terrestrial impacts during efforts to restore essential fluvial geomorphic and aquatic habitats in regulated river eco- systems.

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.

Pollen morphology in Lysipomia (Campanulaceae: Lobelioideae) and interpretation of shape artifacts

The pollen of 32 species of Lysipomia was examined by light, scanning, and transmission electron microscopy. Two pollen types occur in the genus: 3-colporate and 6-colporate. The 3-colporate condition occurs in only two species, L. laciniata and L. pumila. The remaining 30 species are 6-colporate, a condition known from only one other genus in the Campanulaceae. Surface sculpturing among the species is uniformly striate. Pollen shape was highly variable within a single individual in comparisons of pollen gathered from herbarium specimens, FAA preserved material collected in the field, and fresh pollen from cultivated individuals grown from seed. Shape may change from oblate spheroidal to subprolate as a result of drying time and temperature, and should not be used as a morphological character in systematic studies if infraspecific variation is seen. When fresh or preserved pollen is not available, rehydrated pollen should be compared to reduce the possibility of inadvertent artifact production confounding the analysis of morphological data.

Genetic and Morphometric Assessment of the Origin, Population Structure, and Taxonomic Status of Anticlea vaginata (Melanthiaceae)

Abstract Anticlea vaginata (Melanthiaceae) is a rare and endemic plant species restricted to hanging gardens in low-elevation desert regions of the Colorado Plateau. Its more widespread congener, A. elegans, is morphologically similar, but occurs in montane forests that encompass and extend beyond the natural range of A. vaginata. Here, we use morphometric and genetic analyses to investigate the biogeographic origin, population structure, and taxonomic classification of A. vaginata relative to A. elegans. Our results demonstrate that A. vaginata is closely related to and morphologically indistinguishable from A. elegans and likely represents remnant populations of A. elegans derived from a Pleistocene vicariance event. We conclude that A. vaginata warrants treatment as Anticlea elegans subsp. vaginata, since it exhibits a similar level of differentiation from A. elegans subsp. elegans as subsp. glaucus. Since A. vaginata occupies an ecologically unique niche, exhibits a distinct flowering period and harbors unique alleles, we suggest separate conservation management in order to protect this subspecies and its fragile habitat, which is currently threatened by climate change and the potential for groundwater development.

Three new species of Lysipomia (Lobeliaceae) endemic to the páramos of southern Ecuador

Three species of Lysipomia, L. caespitosa, L. cylindrocarpa, and L. speciosa, from a tropical alpine ecosystem called páramo, are described, illustrated, and compared to related species. Notes on morphology, especially floral presentation, and general ecology are given where possible. A published key to the species in Ecuador is revised to include the three new species plus L. multiflora McVaugh, a species previously known only from northern Peru.