Flora of North America species comparison

Varieties 7 (7 in the flora).

The variation within Dodecatheon pulchellum is substantial and, for the most part, each of the entities recognized here seems distinct although nearly all break down in one or more features; most seem to have intergraded with other entities in the past. Variety pulchellum is the most widespread and remains, even as treated here, quite variable. The depauperate, often uniflorous, high-elevation form, var. watsonii, is included in var. pulchellum; there is no difference except in overall size even on the East Humboldt and Ruby mountains of northeastern Nevada, the type location of var. watsonii. Widely disjunct populations assigned to var. monanthum remain problematic. The plants of northwestern California and southwestern Oregon differ only slightly from those found elsewhere in Oregon, including the Blue Mountains, where the type of var. monanthum was obtained. The southern Utah expression, while similar morphologically, displays a biogeographic pattern that is unique. A better understanding of the variation between the western and eastern phases of var. monanthum is needed.

The coastal var. macrocarpum has consistent morphological differences and a higher ploidy level (2n = 88, 132) compared with the more inland var. pulchellum (2n = 44). Whether or not the ploidy level difference is consistent remains to be shown.

The arid forms of Dodecatheon pulchellum exhibit remarkable morphological differences that require recognition (J. L. Reveal 2005). Hanging garden plants in Utah are recognized as var. zionense, following N. H. Holmgren (2005). Some populations assigned to this variety may ultimately prove to be merely large-leaved plants of either var. pulchellum or the Utah phase of var. monanthum. Zion shootingstar may owe both its large leaves and its glandular-puberulent pedicels and calyces to hybridization with D. redolens sometime in its evolutionary past, even though the latter taxon is no longer close geographically. Variety shoshonense, usually growing in moist, alkaline meadows, is found mainly in the northern Mojave Desert and the Intermountain West. The color pattern associated with the stamens differs from most other varieties of the species, suggesting a fundamental change associated with pollination and likely a closer relationship to var. cusickii (which also has yellow pollen sacs) than to var. pulchellum.

This taxon inadvertently was named Dodecatheon puberulum (Nuttall) Nuttall three years before the establishment of Exinia pulchella. To avoid nomenclatural disruptions, the basionym D. meadia var. puberulum Nuttall has been proposed for rejection (J. L. Reveal and K. N. Gandhi 2008).

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Genera 20, species ca. 600 (5 genera, 52 species in the flora).

The largest genera in Primulaceae are Primula (ca. 500 species) and Androsace (ca. 100 species). No genera are endemic to the flora area; Dodecatheon and Douglasia have relatively few representatives elsewhere (northeastern Asia).

The family contains ornamental taxa, especially in Dodecatheon and Primula. Primula can cause dermatitis. Some taxa are pollinated by insects; selfing also occurs. Seeds are dispersed by gravity, water, wind, or ants (Primula; B. Ståhl and A. A. Anderberg 2004).

As typically described (e.g., A. Cronquist 1981; V. H. Heywood 1978), Primulaceae were clearly polyphyletic, closely related to Myrsinaceae and Theophrastaceae. M. Källersjö et al. (2000) and B. Ståhl and A. A. Anderberg (2004) removed the nonrosette terrestrial members from Primulaceae in the broad sense and placed them in the Myrsinaceae, which are further distinguished by leaves and calyx often dotted with yellow or dark streaks, flowers with relatively shorter corolla tubes, seeds immersed in placentae, and wood devoid of rays or with multiseriate rays only. Maesa, consisting entirely of trees found in the Eastern Hemisphere tropics, also has semi-inferior ovaries, pedicels with two bracts, and wood with both uniseriate and multiseriate rays; it, too, was removed from Primulaceae/Myrsinaceae and placed in its own family (Källersjö et al.). The families Primulaceae in the narrow sense, Myrsinaceae, Theophrastaceae (including Samolaceae), and Maesaceae then form a monophyletic clade within Ericales (P. F. Stevens, http://www.mobot.org/MOBOT/research/APweb/), sharing some features, most notably flowers with sympetalous corollas, stamens in a single series and opposite the petals, free-central placentation, bitegmic, tenuinucellate ovules, and plants generally with tannins and saponins.

Additional evidence (L. Martins et al. 2003) indicates that Androsace and Primula may not be monophyletic; more work is needed to resolve these issues. The work of M. Källersjö et al. (2000) showed that Douglasia should remain separate from Androsace, and Dodecatheon should remain separate from Primula, although Dodecatheon clearly is derived from Primula subg. Auriculastrum. Alternative views suggesting more inclusive concepts of Primula and Androsace have been offered by I. Trift et al. (2002), A. R. Mast et al. (2004), and G. M. Schneeweiss et al. (2004). The phylogenetic position of Cyclamen, a scapose taxon currently included in Myrsinaceae, has not been resolved. Our understanding of Primulaceae is still in flux, and future taxonomic realignments at the familial and generic levels are to be expected.

(Discussion copyrighted by Flora of North America; reprinted with permission.)