SUBSPECIFIC VARIATION IN THE WIDESPREAD BURL-FORMING ARCTOSTAPHYLOS GLANDULOSA

The genus Arctostaphylos consists mostly of chaparral shrubs known by the common name manzanita, and one of the widest ranging of these is A. glandulosa Eastw., distributed from Baja California to Oregon. Particularly in the southern half of its range it exhibits complex patterns of morphological variation that have long presented taxonomic challenges. Phenetic analysis of morphological traits from over 1400 individuals from throughout the range were used to examine intra- and inter-population patterns of variation. Multivariate ordination and hierarchical cluster analysis were used to determine phenetic patterns linked with ecological and geographical distributions. These analyses suggest the hypothesis that this species comprises two lineages with a common origin but divergent in the presence or absence of glandularity: A. glandulosa Eastw. subsp. glandulosa, characterized by branchlets with long glandular hairs, scabrous or pubescent leaves, and nascent inflorescences with mostly foliaceous bracts; and A. glandulosa Eastw. subsp. cushingiana (Eastw.) Keeley, Vasey and Parker comb. nov., with non-glandular tomentose branchlets, glabrate or pubescent leaves and either foliaceous or short deltoid bracts. Populations dominated by one or the other of these morphotypes occur throughout the range and tend to be separated by elevation or distance from the coast, although mixed populations occur where these taxa come together. Two other glandular subspecies are named here. One is A. glandulosa Eastw. subsp. leucophylla Keeley, Vasey and Parker, subsp. nov., with intensely glaucous leaves and commonly with foliaceous bracts. A second glandular subspecies is A. glandulosa Eastw. subsp. atumescens Keeley, Vasey & Parker, subsp. nov., a narrowly distributed Baja California endemic similar to the nominate subspecies except that it lacks a basal burl and does not resprout after fire. Of the non-glandular tomentose taxa, in addition to A. glandulosa subsp cushingiana, several others are also recognized. One is A. glandulosa Eastw. subsp. crassifolia (Jepson) Wells, a well established coastal San Diego endemic recognized by darker and thicker leaves and smaller and flatter fruits. Another is a newly described taxon A. glandulosa Eastw. subsp. erecta Keeley, Vasey & Parker, subsp. nov., an endemic to northern Baja California recognized by the erect nascent inflorescenses. Two others have glabrate leaves and highly reduced deltoid often marcescent bracts; A. glandulosa subsp. adamsii (Munz) Wells, which has intensely glaucous leaves and is distributed from interior Riverside Co. south, and A. glandulosa Eastw. subsp. gabrielensis (Wells) Keeley, Vasey and Parker comb. nov., which has bright lustrous green leaves and greater fusion of nutlets, and is distributed from the interior San Gabriel Mountains of Los Angeles Co. north to the Sierra Madre Mountains of Santa Barbara Co. All non-glandular plants with long setose or villous hairs are A. glandulosa Eastw. subsp. mollis (Adams) Wells. This taxon includes plants with foliaceous as well as reduced bracts and is distributed throughout the Transverse Ranges from Santa Barbara to San Bernardino counties, with some outlying populations further south. This taxon shows a marked tendency for reduced stomatal densities on the upper leaf surface in the westernmost populations. Although all of the A. glandulosa taxa described here are known from allopatric populations, intergradations of these closely related taxa occur and thus some populations reflect a mixture of traits and can not be assigned a unique name of practical value.