San Diego State University
Studies in Plant Systematics




















Introduction

Plagiobothrys (popcorn flowers) is a genus of low annual or perennial herbs comprising of about 60 species. The distribution of Plagiobothrys is concentrated in Chile, southwestern United States, and northwestern Mexico (Johnston 1927 ). Plagiobothrys species tend to be found in moist areas, and several species occupy sensitive vernal pool habitats (The Jepson Manual, Hickman ed. 1993).


Description of Plagiobothrys

The plants of Plagiobothrys are low annual or perennial herbs The frequently basal leaves are opposite and often connate-perfoliate with the exception of Plagiobothrys collinus which has varied leaf arrangement. The inflorescence is a naked or bracted helicoid cyme. The calyx is often cut almost to base into lanceolate or oblong erect or connivent lobes. In some taxa, the sepal bases in the mature calyx are fused into a cup containing the fruits, and is sometimes circumscissile. Short-pedicellate or subsessile flowers are white with a short corolla tube that barely surpasses the calyx. Petal lobes are rounded, spreading and imbricate. The throat can be white or yellow, with intruded appendages. The gynobasic style is slender, and usually short. The stigma is capitate or turbinate. The gynobase is low-convex, pyramidal or "frustate" (Johnston, 1923). Each of the two locules in the ovary are divided early in development by false septa. At maturity the ovary develops into four or sometimes two nutlets each of which contains a seed. The nutlets are erect, ovate to lanceolate, smooth or roughened. The position of the nutlet attachment scar (from attachment to the gynobase) is described by Johnston (1923) as basal to medial. The scar can be on, below or at the base of a strong ventral keel (ventral ridge), or rarely the stipitate terminal elongation of the ventral keel.


















Morphology of the unit fruits, nutlets, is of extreme importance in delimiting species within the Tribe Eritrichieae. Nutlet morphology may also provide potential apomorphies that identify monophyletic groups within Eritrichieae and Plagiobothrys. Preliminary SEM images of nutlet pericarps from Plagiobothrys and exemplars of other genera within Eritrichieae reveal previously unsuspected characters. Once coded, these characters will be added to a data matrix and will be used to resolve phylogenetic and taxonomic questions.


Summary of Taxonomic and Phylogenetic Relationships

Plagiobothrys belongs to the family Boraginaceae, subfamily Boraginoideae, and Tribe Eritrichieae. Plagiobothrys has formerly been classified as six former genera: Allocarya, Plagiobothrys, Sonnea, Echidiocarya, Glyptocaryopsis, and Echinoglochin (Johnston 1923, Munz 1973). The genus in not united by a known apomorphy. It is my hypothesis that Plagiobothrys is a paraphyletic group that needs revision. My most recent analysis of Parsimony (PAUP), run with a data collected so far, supports this hypothesis. I am still in the process of collecting data for analysis, the monophyly of the genus will be retested when I have a compete data set.

Summary of Project Objectives

The goals for this thesis are: 1) to evaluate the monophyly of the genus Plagiobothrys; identify monophyletic subgroups within Plagiobothrys and evaluate the monophyly of the existing sections in the genus Plagiobothrys; and 2) to evaluate the validity of a putative new species within the genus Plagiobothrys using the phylogenetic species concept. My objectives are to: qualitatively describe nutlet pericarp ultrastructure characters from SEM images taken from 10 individuals from 11 glochidiate species, from SEM images taken from single individuals from the remaining 49 Plagiobothrys species, and from SEM images taken from single exemplars from 29 genera in the tribe Eritrichieae; and quantify four fruit morphological characters from 25 individuals from each of eleven glochidiate species in the genus Plagiobothrys. Quantified variation within and between taxonomic units will be statistically analyzed and graphed. Both qualitative and quantitative characters will be coded and entered into a data matrix using the software MacClade. The software PAUP will be used for analysis of parsimony. Character evolution will be traced on strict, semistrict, and Adam's consensus trees. Taxonomy of this genus, which has been treated as four genera in the past, will be evaluated and revised based on the phylogenetic relationships hypothesized by this research.

Publications in progress:

Dellavalle, M. & M. G. Simpson. Phylogenetic analysis of the genus Plagiobothrys (Boraginaceae), with emphasis on fruit wall ultrastructure.

Dellavalle, M. & M. G. Simpson. Fruit ultrastructure of Plagiobothrys (Boraginaceae), proposed for publication in Proceedings of the San Diego Natural History Museum.