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December 2012 Number 1 - Utah Native Plant Society

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Calochortiana <strong>December</strong> <strong>2012</strong> <strong>Number</strong> 1<br />

Section Diphysi from other putatively related sections.<br />

The thickness or texture of the valve walls, the type and<br />

distribution of pubescence on the valves, and the degree<br />

of closure of the locules by the septum (if it is complete<br />

and fused to the funicular flange throughout or just<br />

within a portion of the body of the pod) is highly variable<br />

throughout the range of this species.<br />

The second group within this complex is characterized<br />

by having scarcely inflated (cylindrical to ventricose<br />

in shape and slightly inflated dorsally, if at all),<br />

thick papery to leathery, elliptic, narrowly oblong, to<br />

linear pods. The septum is generally incomplete in this<br />

group, either semi-bilocular (the septum partially divides<br />

the two locules) or sub-unilocular (the septum is<br />

less than half the width of the locule). Unlike the first<br />

group, the pods are either deciduous or long-persistent.<br />

These scarcely inflated taxa were first comprehensively<br />

described by Rydberg (1929) as Section Palantia<br />

Rydb. within the genus Tium Medik., based on the similarity<br />

of the pod morphology. The remaining members<br />

of Section Diphysi were split and included in the old<br />

world genus Cystium Steven. The degree to which these<br />

characteristics define a section or species is a major<br />

source of disagreement among all monographs of this<br />

complex (Barneby 1945, 1964, Isely 1998, Jones 1923,<br />

Rydberg 1929, Welsh 2007). The long-persistent pods<br />

in one taxon, A. lentiginosus var. mokiacensis (A. Gray)<br />

M.E. Jones, was the primary character that lead Barneby<br />

(1989) to retain it within the Section Preussiani M.E.<br />

Jones, a section distantly related to Section Diphysi.<br />

Vastly different interpretations of the significance of<br />

this character have led to often disparate views of the<br />

species boundaries and delimitations surrounding these<br />

taxa (Alexander 2005, Barneby 1964, 1989, Welsh<br />

2007). When the taxa with persistent pods are delimited<br />

as varieties, Astragalus lentiginosus becomes the only<br />

documented North American species of Astragalus to<br />

have forms with both persistent and deciduous pods.<br />

All other examples proposed by taxonomists have been<br />

split at the species or the sub-sectional level in recent<br />

monographs.<br />

In the most recent revision, Alexander (2008) considered<br />

all of the scarcely inflated varieties of A. lentiginosus<br />

to be a single evolutionary lineage and referred to<br />

them collectively as the Palantia, based on Rydberg's<br />

sectional name. The Palantia consists of A. lentiginosus<br />

var. bryantii (Barneby) J.A. Alexander, A. lentiginosus<br />

var. iodanthus (S. Watson) J.A. Alexander, A. lentiginosus<br />

var. maricopae Barneby, A. lentiginosus var.<br />

mokiacensis (including A. lentiginosus var. trumbullensis<br />

S.L. Welsh & Atwood), A. lentiginosus var. palans<br />

(M.E. Jones) M.E. Jones, A. lentiginosus var. pseudiodanthus<br />

(Barneby) J.A. Alexander, A. lentiginosus var.<br />

ursinus (A. Gray) Barneby, and A. lentiginosus var. wilsonii<br />

(Greene) Barneby. The name, Palantia, is not used<br />

herein in a nomenclatural sense as a sub-generic or sectional<br />

name. It is unusual in botany for a collective<br />

name to be required for clarity when referring to groups<br />

of morphologically similar varieties. However, with<br />

over 40 varieties and 4 or 5 lineages that are similar<br />

morphologically, a collective naming convention for A.<br />

lentiginosus is necessary to refer to these groups. As<br />

such, the Palantia is used as a convenient, informal<br />

name for the varietal group with scarcely inflated pods<br />

and is italicized following the common literary convention<br />

for unfamiliar Latin words.<br />

Within the Palantia, two taxa, Astragalus lentiginosus<br />

var. maricopae and A. lentiginosus var. ursinus have<br />

been historically overlooked by taxonomists and have<br />

an uncertain taxonomic status. Astragalus ursinus A.<br />

Gray (first reduced to a variety of A. lentiginosus by<br />

Barneby 1964) was the first to be described in 1878,<br />

along with A. mokiacensis A. Gray (Gray 1878). Gray<br />

was the first to propose a close relationship between<br />

these taxa and A. lentiginosus var. iodanthus (at that<br />

time, and until only recently, this taxon was delimited as<br />

a species). The affinity of the types of A. ursinus to extant<br />

populations has been controversial since the taxon<br />

was first described (Alexander 2005, 2008, Barneby<br />

1964, Jones 1923, Welsh 1978, 2007, Welsh & Atwood<br />

2001). In some recent taxonomic treatments (Barneby<br />

1989, Welsh 1993), the types have been regarded as an<br />

insignificant variant of A. lentiginosus var. palans.<br />

However, in others A. lentiginosus var. ursinus is recognized<br />

as an insignificant variant of A. lentiginosus var.<br />

mokiacensis (Alexander 2005, Welsh 2007, Welsh &<br />

Atwood 2001).<br />

Based on a combination of field surveys, morphometric<br />

analyses and chloroplast haplotype analyses, Astragalus<br />

lentiginosus var. ursinus was found to be genetically<br />

distinct from its geographically closest relative,<br />

A. lentiginosus var. mokiacensis (Alexander 2008, Alexander<br />

& Liston, in prep). In addition, Astragalus lentiginosus<br />

var. ursinus is a highly restricted limestone talus<br />

endemic (totaling less than 5,000 individuals), and is<br />

confined to a small region of the Beaver Dam Mountains<br />

in Mohave County, Arizona and Washington<br />

County, <strong>Utah</strong> (Alexander 2008).<br />

The second species, Astragalus lentiginosus var.<br />

maricopae, was first described in Barneby's (1945)<br />

monograph. It is often confused with A. lentiginosus<br />

var. yuccanus due to similar floral morphology (size and<br />

color) and has remained poorly known since it was first<br />

described. The floral and pod morphology are highly<br />

distinct when compared to the other members of the<br />

Palantia. If it were placed within any other section of<br />

the genus, it would be recognized at the species-level. It<br />

has universally been recognized as a variety of A. lentiginosus<br />

in all major monographs, but has only recently<br />

been found to be a restricted endemic (Alexander 2008).<br />

135

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