Ronald L. Hartman

An ITS-Based Phylogenetic Analysis of the Perennial, Endemic Apiaceae Subfamily Apioideae of Western North America

Abstract Phylogenetic analyses of 159 DNA sequences from the nuclear rDNA internal transcribed spacer region were conducted to evaluate the monophyly of the herbaceous, perennial genera of Apiaceae subfamily Apioideae endemic to North America (north of Mexico) and to determine the relationships of those elements that currently comprise Cymopterus within the group. The results of a previous phylogenetic study were equivocal in suggesting monophyly for these perennial, endemic taxa and revealed Cymopterus to be polyphyletic, with its species closely linked with those of Aletes, Lomatium, Musineon, Oreoxis, Orogenia, Podistera, Pseudocymopterus, Pteryxia, and Tauschia. Herein, we expand sampling to include comprehensive representation of Aletes, Cymopterus, Musineon, Oreoxis, Orogenia, Podistera, Pseudocymopterus, and Pteryxia, and greater representation of Lomatium and Tauschia. We also include all members of two genera not examined previously, Glehnia and Oreonana, as well as additional outgroup genera from the Angelica clade of the apioid superclade. Our results indicate that the perennial, endemic apioid umbellifers of North America constitute a (weakly supported) monophyletic group, with Angelica and the meso-American Arracacia clade comprising two of several possible sister groups. The two subspecies of Glehnia littoralis ally with Angelica and Peucedanum japonicum; Oreonana shows affinity with several species of Cymopterus and Lomatium. The lack of resolution in the ITS trees precludes unambiguous hypotheses of relationship among these perennial, endemic umbellifers but does show that many of these genera, where resolved, are not monophyletic. Indeed, Cymopterus and Lomatium are highly polyphyletic and permeate all major clades resolved in the molecule-derived trees. Evidence from branch lengths and low sequence divergence suggests that this group of North American umbellifers underwent rapid radiation, likely during the geoclimatic events of the Late Tertiary and Quaternary.

Systematic implicationsofflavonoid patterns in Haplopappus segregates

Complex flavonoid patterns are correlated with primitive woody taxa and high base chromosome numbers among Haplopappus segregates. Variability among flavonoid complements supports the polyphyletic interpretation of the genus.

TWO NEW SPECIES OF CYMOPTERUS (UMBELLIFERAE) FROM WESTERN NORTH AMERICA

Two species ofCymopterus are described as new:Cymopterus douglassi from the Lost River and Lemhi ranges of central Idaho and the closely relatedC. williamsii from the southern half of the Big Horn Mountains, Wyoming. Both are diploid on the basex=11 and occur on calcareous or dolomitic substrates. The mature fruit ofCymopterus williamsii differ from those ofC. douglassii in the absence of a functioning carpophore and in having 1, not 3–5 oil tubes in the intervals. Both species are unusual in the genus, though not unique, in that the dorsal ribs of the mericarps are not winged.

A new species of Cymopterus (Umbelliferae) from southern Idaho

A new species,Cymopterus davisii, from alpine areas in the Albion Mountains of extreme south-central Idaho, is described. It has been confused withC. nivalis, with which it is compared. The fruit ofC. davisii are subterete to somewhat compressed laterally and, contrary to the literature, correspond closely to those ofC. nivalis and related species.

Morphology, Anatomy, and Taxonomic Relationships of Otopappus australis (Asteraceae)

A collection of Otopappus australis from Cerro Campana represents the first report for the genus from Panama. Aspects of the morphology at variance with the type description include longer and narrower leaves and narrower flowering heads. Details of stem and leaf anatomy are reported; exceptionally tall primary wood rays probably relate to the plants vine-like habit. A variety of primitive and specialized features are found in the floral anatomy. Unusual topographic sites for trichomes include pappus awns, inside the corolla tube, anther appendages, and style branches. The junior authors conclude that this species is more properly a Zexmenia rather than an Otopappus. Geographical and morphological data are uti- lized to demonstrate that 0. australis, Z. columbiana, and Z. mikanjoides are conspecific. The latter two are considered Zexmenia mikanioides var. mikanioides, and the nomen- clatural combination is made to make 0. australis a variety of Z. mikanioides. The genus Otopappus Benth. belongs to the subtribe Verbesininae of the Heliantheae (Asteraceae) and contains about 15 species as presently understood (Hartman and Stuessy, unpubl.). Most taxa are found from western Mexico to Costa Rica. The closest generic relatives are Notoptera Urb. and Salmea DC. (Blake, 1915a; Stuessy, 1977). McVaugh (1972) suggested that the generic distinctiveness of Otopap- pus and Notoptera may be specious and recent revisionary studies (Hart- man and Stuessy, in prep.) have shown that the taxa are congeneric. The genus is unified by its viny habit and winged awns of the achenes. How- ever, one species, 0. australis, is isolated morphologically and geograph- ically. This taxon was originally described by Blake (1924) as a relative of 0. verbesinioides Benth. and as the first representative of the genus from South America. Features of the achenes of 0. australis differ from other species of the genus and suggest possible affinities with other gen- era. Geographically, 0. australis is known from Venezuela and Ecuador, which are outside the known range of all other species of the genus (0. simplex Badillo was also described from Venezuela, but it does not belong in Otopappus; details of that relationship will be described elsewhere). Interest in Otopappus australis has developed independently with the senior author finding material new to Panama, and the junior authors determining specific and generic relationships. Discovery of each others work suggested that the observations and interpretations be published

A Revision of the Ipomopsis spicata Complex (Polemoniaceae)

A cladistic analysis indicates that the Ipomopsis spicata complex is monophyletic, related as a sister group to I. congesta in sect. Microgilia, and is characterized by regional differentiation most likely associated with climatic and topographic changes that occurred during the late Pliocene and Pleistocene. The cladogram provides evidence for the independent evolution of alpine, sub- alpine, and lowland races in the Rocky Mountains and the eastern perimeter of the Great Basin, correlated with a progressive change in inflorescence architecture, trichome distribution, and floral morphology. Based on morphological studies and examination of all type specimens, a taxonomic revision is proposed that summarizes morphological, geographical, and cladistic patterns within the complex. We recognize I. spicata as composed of five principal geographic races treated as: subsp. spicata, subsp. capitata, subsp. orchidacea (comb. nov.), subsp. robruthii (subsp. nov.), and subsp. tridactyla (comb. nov.). Subspecies orchidacea includes var. orchidacea and var. cephaloidea (comb. nov.), which differ in ecological distribution and inflorescence architecture but intergrade in parts of their sympatry. Subspecies robruthii is described from the Absaroka Range of northwestern Wy- oming.

Phylogenetic Analysis Reveals Multiple Cases of Morphological Parallelism and Taxonomic Polyphyly in Lomatium (Apiaceae)

Abstract The genera Lomatium and Cymopterus, along with many others, form a group that has been referred to previously as the perennial endemic Apiaceae subfamily Apioideae of western North America. This group of ecologically important and widespread species has been the target of numerous systematic studies, but the evolutionary relationships among these species remain elusive. Here we show that this confusion is due to high levels of morphological parallelism and homoplasy in the characters that have traditionally been used to define them, a result that is concordant with previous studies of the group. We explore patterns of evolution in traditionally important morphological characters using Bayesian stochastic character mapping on a phylogeny constructed from novel nrDNA and cpDNA sequence data for 96 specific and infraspecific taxa of the estimated 200 species in the group. We consider the implications of these results for taxonomic classification, the evolution of morphologies, and the utility of these morphologies to delimit small and large clades. Lomatium concinnum is newly combined as Cymopterus glomeratus var. concinnus and the new combination Cymopterus glomeratus var. greeleyorus is made.

A new species of Cymopterus (Apiaceae) from the Rocky Mountain region, U.S.A.

Cymopterus constancei is described as new from mostly western portions of Wyoming, Colorado, and New Mexico and from eastern Utah and Arizona. It has been confused withC. bulbosus A. Nelson but differs in several features of its bractlets, phenology, and habitat. The novelty is among the earliest of the native flora to bloom and is quite uniform morphologically. The number of collections encountered exceeds 200, a fourfold increase, in the past 22 years.

Revision of Otopappus (Compositae, Heliantheae)

Otopappus (Compositae, Heliantheae) is revised to contain 15 species from Mexico and Central America. Prominent among the taxonomic difficulties have been generic delimitations, especially with Notoptera. The principal differentia between the two genera, pappus with and without squamellae, is shown not to correlate with other features, and the two genera are merged. Within this complex two sections are recognized: Otopappus (10 species) and Loxosiphon (5 species) with the latter divided into series Loxosiphon (2 species) and series Hirsutus (3 species). Principal characters separating these major groups include discoid vs. radiate heads, numbers of rows of phyllaries and their length in relation to the paleae, color of corollas (white or yellow), and convex vs. conical receptacles. The genus is unified by its viny habit and conspicuously adaxially winged achenes and pappus awns. Within this context, 0. australis, 0. ferrugineus, and 0. simplex seem better referred to Zexmenia and 0. jaliscensis to Lasianthaea. Otopappus seems allied most closely to Salmea or Zexmenia. New combinations are Lasianthaea jaliscensis, Otopappus sect. Loxosiphon, 0. series Hirsutus, 0. guatemalensis, 0. hirsutus, and Zexmenia simplex.

The Identity of Aster blepharophyllus (Asteraceae: Astereae)

Recent collections of Aster blepharophyllus A. Gray, the first since its discovery in 1851, prompted a reevaluation of its identity. It is properly placed in the genus Machaeranthera as M. gypsitherma Nesom, Vorobik, & Hartman. Within that genus, it shares characters with species of sect. Blepharodon as well as those of sect. Arida (both groups sensu Hartman), but we place it tentatively in the latter group. Of particular significance is its chromosome number of n = 5 pairs, reported for the first time, which is shared with species of sect. Arida.