December 2012 Number 1 - Utah Native Plant Society

Utah Native Plant Society
Table 2. List of 24 morphological characters and
their abbreviations used in the 153 specimen
Palantia PCoA. Two were constant (C: calyxd, calyxo)
and not used in the PCoA, parsimony or cluster
analysis. One (O: calyxs) was not used due to the finding
that it was variable on the same plant (and within
most calyces). For a list of character states for the characters
below, see Appendix 2. Characters were coded as
multistate continuous variation (R), binary state (B), or
multistate (M).
1. Adaxial leaflet pubescence (Leafad) M
2. Abaxial leaflet pubescence (Leafab) M
3. Leaf and stem hair length (leafh) R
4. Leaflet number (leafn) M
5. Inflorescence length in flower (inflw) R
6. Calyx tube length (calyxl) R
7. Calyx pubescence density (calyxd) M, C
8. Calyx teeth shape (calyxs) M, O
9. Calyx teeth orientation (calyxo) M, C
10. Keel length (keell) R
11. Keel color (keelc) M
12. Banner color (bannc) M
13. Inflorescence length in fruit (infr) R
14. Pod pedicel orientation (podpo) M
15. Pod length X width ratio (podr) R
16. Pod deciduous or persistent (poddp) B
17. Pod shape, longitudinal section (podsl) M
18. Pod shape, cross section (podsc) M
19. Pod orientation on raceme (podro) M
20. Pod orientation, degree of pod incurve (podpi) R
21. Pod inflation (podin) M
22. Pod valve texture (podt) M
23. Pod pubescence (podpu) M
24. Pod valve color (podvc) M
scarcely inflated, unilocular, leathery, deciduous pods),
and not to members of Section Inflati (4-6 base pair divergence;
Alexander, unpublished data, Wojciechowski
et al. 1993, 1999). The ITS sequence data suggest that a
deciduous, unilocular, leathery, scarcely inflated pod is
the putative ancestral state in this complex. Based, in
part, on these data, members of Section Argophylli were
used as outgroups for chloroplast haplotype analyses in
Knaus (2008). Taxa in other putatively closely related
sections (Section Monoenses Barneby, Section Cystiella
Barneby, Section Circumdati (M.E. Jones) Barneby, or
Section Platytropides Barneby; all of which have taxa
with inflated pods) have not been fully investigated in
molecular analyses. The selection of any member of
Section Argophylli as an outgroup automatically polarizes
the ancestral state of the group as a unilocular or
partially bilocular, scarcely inflated, deciduous pod. In
addition, haplotypes within the Argophylli sampled by
Knaus (2008) were found to be nearly twenty steps
more distant from the haplotypes examined in the
Palantia. Finding that A. lentiginosus var. iodanthus and
A. lentiginosus var. pseudiodanthus (both of which have
been universally delimited as species until Alexander
2009), are not highly genetically or morphologically
distinct from A. lentiginosus is a problem for outgroup
selection in this study, primarily with the parsimony
analysis and the genetic analyses (Alexander & Liston,
in prep). The terminal taxa in this study are also not recognized
at the species level, which violates assumptions
in parsimony analyses. As a result, robust phylogenetic
conclusions cannot be made with these data.
Instead, the cladistic analyses were used to investigate
patterns of character state changes within the
Palantia. Astragalus lentiginosus var. iodanthus, and A.
lentiginosus var. pseudiodanthus were selected as outgroups
based on their relatively greater genetic and morphologic
distance from the Palantia based on the results
from Alexander (2008), Knaus (2008), and Alexander &
Liston (in prep). Even if the inflated varieties related to
A. lentiginosus var. yuccanus were used as outgroups
for the Palantia, the morphological trends discussed
herein would not change (see Figures 5-8) since the major
clades of the Palantia would still be split in two
clades. More thorough molecular analyses of species
potentially closely related to Section Diphysi (with both
inflated and scarcely inflated pods) are needed before a
robust molecular and morphological phylogenetic study,
with a satisfactory outgroup, can be attempted.
Despite this, conclusions of the taxonomic status of
Astragalus lentiginosus var. maricopae and A. lentiginosus
var. ursinus can be made. The results of the PCoA
analyses and the genetic analyses (Alexander & Liston,
in prep) indicate that species level delimitations for
many of the Palantia have much weaker support than
previously thought by Alexander (2005). Astragalus
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