VOLUME 58, NUMBER 3 JULY-SEPTEMBER 2011
RECONSIDERATION OF THE TAXONOMIC STATUS OF MASON’S LILAEOPSIS — A
STATE-PROTECTED RARE SPECIES IN CALIFORNIA
Peggy L. Fiedler, Esa K. Crumb, and A. Kate Kn0Xx..........ccccccceeeeeeeeeereees 131]
A COMPARISON OF THE EFFECTS OF NA2SOq4 AND NACL ON THE GROWTH OF
HELIANTHUS PARADOXUS AND HELIANTHUS ANNUUS (ASTERACEAE)
MoO. Mendez and O; W, Van AUbeH sivtecc Met accsesenstoomee eee tona eater ondees 145
THE DIVERSITY AND BIOGEOGRAPHY OF THE ALPINE FLORA OF THE SIERRA
NEVADA, CALIFORNIA
SITU TTD WV RUNG CP os ese ch ace nee eA RD wk ee RM PR Anan ounesedausewadeass 53
A NEw SPECIES OF ASTRAGALUS (FABACEAE) FROM THE WASATCH
MOUNTAINS OF UTAH
DCU TCOWE: COP DIM oh cients aiecdet as ORR See AON hens oli Sulla cbueaeenaule 185
GRIMMIA VAGINULATA, (BRYOPSIDA, GRIMMIACEAE) A NEW SPECIES FROM
THE CENTRAL COAST OF CALIFORNIA
TR CATLCLIA INCU VIIIOTA te iN rene Le SN eh 190
INTRODUCTION TO CALIFORNIA CHAPARRAL
CGHAESTODICTAIBUGV LOCK. cisirs caissad Mccre tetec ae Oe 91g siRNA ned as Fes ees wae snide 199
COP ANTEINEE GOIRINITIN reteset ite cia sts ano, aeRO Me cists cscs a wits Roe oak oes Sanne vamaeemniads 201
DEIN eee se Bites aah ee iste casa ca camnem sO taneaaenaieasoadeenuasiheeeueneliel 204
IN TERS (610 SAR te oe) ee ere eee D205
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MADRONO, Vol. 58, No. 3, pp. 131-144, 2011
| GMT HSONIay
APR 129012
LIBRARIES
RECONSIDERATION OF THE TAXONOMIC STATUS OF MASON’S
LILAEOPSIS — A STATE-PROTECTED RARE SPECIES IN CALIFORNIA
PEGGY L. FIEDLER
Natural Reserve System, UC Office of the President, Office of Research and
Graduate Studies, 1111 Franklin Street, 6" Floor, Oakland, CA 94607
peggy.fiedler@ucop.edu
ESA K. CRUMB
Wetlands and Water Resources, 818 Fifth Ave, STE 208, San Rafael, CA 94901
A. KATE KNOX
7730 19% Avenue NE, Seattle, WA 98115
ABSTRACT
Lilaeopsis masonii is a California state-listed rare species with a wide range of morphologies
observed in the field throughout its range, and in herbaria collections. This extensive variation
confounds reliable taxonomic identification, particularly for those specimens intermediate between L.
masonii and its sister taxon, L. occidentalis. To investigate the genetic basis of this morphological
variation, we examined two portions of the Lilaeopsis genome in seven species. Specifically we sought
to determine whether L. masonii is sufficiently distinct from its closely related, widespread congener to
continue to warrant specific status. DNA sequence analysis of ITS1, 5.8S, and ITS2 nuclear ribosomal
DNA revealed no differences between L. occidentalis and L. masonii California collections, and
minimal differences between these samples and L. occidentalis collected from the state of Washington,
suggesting strongly that these two species form a single clade. A combination of fragment data from
three AFLP primers yielded 274 fragments from 29 samples. Genetic Manhattan distance values
calculated from the AFLP matrix within species ranged from a low of 1.4 to a high of 6.6, reflecting
minor differences among all samples. UPGMA cluster phenograms support the results of the PCA
analysis, illustrating a cluster of L. occidentalis + masonii samples distinct from other Li/aeopsis
species. Because conservation dollars should protect unique evolutionary entities, we suggest that L.
masonii be subsumed under L. occidentalis and therefore no longer receive formal state protection.
Key Words: AFLP, Apiaceae, California endangered species Act, goldilocks conundrum, ITS,
lilaeopsis masonii, lilaeopsis occidentalis, UPGMA.
Lilaeopsis masonii Mathias & Constance (Ma-
son’s lilaeopsis) is one of 15 wetland or aquatic
species of the widespread genus Lilaeopsis Greene
within the Apiaceae. The genus Lilaeopsis 1s
comprised of perennial herbs characterized by a
horizontal stem with leaves commonly in clusters
(“‘ramets’’) borne directly from the stem, although
rarely leaves occur individually. Lilaeopsis is
notable in its morphologic simplicity—entire,
generally linear leaves; simple umbels; absence
of a carpophore; and, a strongly reduced habit
(Petersen et al. 2002; Downie et al. 2000; Downie
et al. 2008). Such simple morphology has led to
a long history of taxonomic uncertainties and
difficulty in the reconstruction of its phylogeny.
Evidence for monophyly of Li/aeopsis is strong
(Petersen et al. 2002). However, recent research
based on molecular evidence from nuclear and
chloroplast genes suggests that the genus is best
placed in the Oenantheae tribe within the
Apioideae (Downie et al. 2008) and that Lilaeop-
sis 18 sister to the clade comprising Ptilimnium,
Limnosciadium, Daucosma, Cynosciadium and
rachis-leaved species of Oxypolis, not the Mexi-
can genus Neogoezia as suggested by Petersen
et al. (2002). The New World endemics clade of
tribe Oenantheae is native to North America and
comprises a monophyletic group that appears to
be evolving much faster than any other major
clade recognized in the tribe (Hardway et al.
2004).
Early taxonomic work on the genus in
California by Hill (1927) and Mason (1957)
included mention of comparatively smaller and
narrower leaves in Lilaeopsis specimens occurring
away from the coast, in contrast to a relatively
more robust coastal form. Professor Herbert
Mason, an early expert on the wetland flora of
California, first collected a relatively smaller
Lilaeopsis from Brannan Island of the San
Francisco Bay/Sacramento-San Joaquin Delta
(Bay-Delta). He referred to the smaller form as
the “San Francisco Bay and river-mouth” form
(Mason 1957: 631). This specimen, according to
Mason (unpublished) was ‘‘definitely distinct
from the coastal L. occidentalis.’’ Western
lilaeopsis (Lilaeopsis occidentalis J. M. Coulter
& Rose) is a widespread, common species,
139 MADRONO
ranging from the Queen Charlotte Islands of
British Columbia, Canada to Marin County,
California (Affolter 1985). Considered to be a
coastal species confined to salt water or brackish
water intertidal habitats, collections of L. occi-
dentalis from inland fresh water lentic and lotic
habitats are known, but considered “‘uncharac-
teristic’ (Affolter 1985).
Lilaeopsis masonii was not described as a
distinct taxon for two decades after the smaller
form in the Bay-Delta was first observed. In
1977, Mathias and Constance formally recog-
nized the diminutive nature of a specimen
obtained from Twitchell Island in the Bay-Delta
as L. masonii (Mathias and Constance 1977).
Mathias and Constance described L. masonii as
distinct from L. occidentalis based upon the
former (rare) taxon bearing narrower, typically
shorter, and more or less terete leaves, and an
inland distribution. They honored Herbert Ma-
son’s expertise in the wetland flora of the State
with the specific epithet.
Mason’s lilaeopsis was one of the first vascular
plant species to be protected as “‘rare’”’ under the
California Endangered Species Act (CESA)
(California Fish & Game Code 882050, ef seq.).
At the time of its listing in November 1979, only
seven population occurrences were known
(CNDDB 2009). Since formal protection, the
documented extent of geographic distribution
and population abundance of L. masonii has
increased nearly three-fold, primarily as a result
of concentrated field survey efforts conducted in
the early 1990’s by Golden, Fiedler, and Zebell
(Golden and Fiedler 1991; Golden 1992; Fiedler
and Zebell 1993; Zebell and Fiedler 1996). Today,
Mason’s lilaeopsis is known to occur within 24
USGS quadrangles and seven counties (CNPS
2008), spanning across roughly 690 square miles.
One hundred eighty-six documented occurrences
are on record with the state (CNDDB 2009),
although most, but not all are extant.
A History of Taxonomic Uncertainty
Confusion over the taxonomic limits of this
rare species existed from the beginning of its
description. Two examples are relevant. First, a
long-controversial Lilaeopsis specimen collected
by Schreiber (#42266 UC, 28 June 1936) from
Chicken Ranch Beach in Marin County derives
from outside the circumscribed geographic range
of the endemic inland taxon. Leaf lengths from
this specimen range between 15 to 42 mm, a
morphological range characteristic of compara-
tively larger leaf lengths for L. masonii. However,
it is possible to key these larger leaved Chicken
Ranch Beach specimens to L. occidentalis in
every relevant flora (e.g., Hickman 1993).
Affolter (1985) examined this specimen in his
monograph of Lilaeopsis, and accepted it as L.
[Vol. 58
masonii, but noted that it was a geographical
outlier for the rare, Bay-Delta endemic species.
Today, CNPS (Tibor 2001) acknowledges that
this specimen is likely to be L. occidentalis, not
the rare L. masonii, but provides no explanation.
Several attempts to relocate this Lilaeopsis
material at Chicken Ranch Beach by the authors
have failed as the population appears to be
extirpated, thereby making an independent spe-
cies corroboration impossible.
It is important to note that (1) numerous
collections of L. occidentalis from the beaches of
Marin and Sonoma counties exist, (2) leaf lengths
range by an order of magnitude or more within
and between adjacent populations of L. occiden-
talis, (3) the number and clarity of internal
crosswalls considered important diagnostic char-
acters are more likely a function of relative plant
size, exposure, or both, and (4) inland collections
of the common species are known from the state
of Washington (e.g., UC 1594452; 4 September
1962). Also noteworthy, L. masonii has never
again been collected on the Pacific coast of North
America beyond Schreiber’s Marin Co. collection
in 1936.
Further, Affolter (1985) remarked that leaves
from a collection of L. masonii (derived from
Sherman Island immediately down river of
Twitchell and Brannan islands) cultivated for
his greenhouse comparisons were “‘remarkably
longer than any of the herbarium material”
(Affolter 1985:70). He suggested the observed
overall larger and more robust greenhouse
material was evidence of how difficult it is to
understand vegetative plasticity from herbarium
material alone. However, the relatively robust
response of Mason’s lilaeopsis to the mild
conditions of a greenhouse suggests strongly that
strict morphological distinctions between the two
taxa are problematic.
Additional morphological characters further
support the assertion that L. masonii is not
distinctly different from L. occidentalis. Affolter
(1985:70) noted that the “two taxa are similar in
several respects,”’ including similar (1) leaf shapes
(linear), (2) rhizome branching architecture, (3)
fruit shapes, (4) fruit cell types, (5) fruit venation
patterns, (6) habitats, and they have (7) overlap-
ping geographic distributions. Despite all these
similarities, Affolter (1985:71) supported their
separate specific status, primarily because ““when
grown under a common-garden environment in
the greenhouse, the two species retained the
vegetative characteristics that distinguish them
in the field.”
Subsequent laboratory studies conducted by
the principal author and her students (Golden
and Fiedler 1991; Golden 1992; Fiedler and
Zebell 1993; Zebell and Fiedler 1996) have
provided little clarity. Most importantly, no
nucleotide variation was found among nine
2011]
populations of L. masonii or between L. occiden-
talis and L. masonii when the 204 nucleotides of
the ITS2 nuclear genome were ascertained
(Fiedler and Zebell 1993). Fiedler and her
colleagues thus concluded tentatively that the
rare species was most likely an inland ecotype not
clearly distinct from its widespread congener.
Field Observations
Decades of field observations of Lilaeopsis
throughout the Bay-Delta, Suisun Marsh, and
Napa River ecosystems do not reinforce many of
the conclusions offered by Hill (1927), Mason
(1957, unpublished), Mathias and Constance
(1977), and Affolter (1985) supporting the
recognition of two distinct taxa. Rather, the few
vegetative characteristics that typify this genus
are highly variable both within and between
populations throughout this region. Occurrences
of L. masonii in the lower Napa River, studied
since 2001 (WSP 2007, unpublished; Blasland,
Bouck, & L, Inc. unpublished; Entrix unpub-
lished; L.C. Lee & Associates unpublished; Still-
water Sciences and Fiedler unpublished) include
a full spectrum of individual ramet sizes. Often,
both large and small forms of Lilaeopsis species,
easily identifiable to the two different species, can
be found growing in the same location. Often the
plant stature/leaf length size gradient runs per-
pendicular to the shoreline, where the small
‘“masonii’ form (approx.1.5—4.5 cm in height)
grows relatively close to the water’s edge, while
increasing larger and more robust “‘occidentalis”’
(approx. =11 cm in height) can be found further
from the water. “Intermediate” or medium-sized
Lilaeopsis material (approx. >6.25 and <11 cm
in height) is common throughout this shoreline/
river bank habitat and geographic range, and
keys to either (or both) the rare or the common
species. We call this phenomenon—.e., range in
size of a critical morphological character, with
significant overlap between taxa—the ‘‘Goldi-
locks Conundrum”’ to highlight the problem that
the intermediate-sized material is not “‘just right,”’
but rather, highly problematic.
To resolve our conundrum and determine
whether L. masonii is a discrete species distinct
from L. occidentalis, we initiated a genetic
analysis of seven species of this genus. We
hypothesized that there were no significant
differences between diagnostic portions of the
genome selected for this study of the two species,
L. masonii and L. occidentalis. Based on these
analyses, we then explored whether L. masonii
watrants continued recognition as a distinct
species or rather, should be subsumed under the
widespread and common L. occidentalis. If no
significant differences were shown to exist be-
tween diagnostic portions of the L. occidentalis
and L. masonii genomes, then L. masonii should
FIEDLER ET AL.: RECONSIDERATION OF MASON’S LILAEOPSIS 133
be subsumed within L. occidentalis, and contin-
ued protection under the California Endanger-
ed Species Act for L. masonii should be
reconsidered.
Fallon (2007) noted that genetic information is
being used increasingly to resolve taxonomic
issues for protection at the federal level under
the U.S. Endangered Species Act of 1973 (ESA).
She conducted a review of listing decisions made
by the U.S. Fish & Wildlife Service and the
National Marine Fisheries of species, subspecies,
or distinct population segments (DPSs) proposed
for protection under the ESA. Fallon determined
that the listing fate of a DPS based upon data
from more than one genetic marker resulted in a
higher probability of protection than candidate
taxon or population segment whose discreteness
was determined by a single genetic marker. With
the cautionary tale of Fallon’s findings in mind,
we examined the ITS region of the nuclear
genome and, to corroborate our ITS findings,
conducted an amplified fragment length poly-
morphism (AFLP) analysis on a similar suite of
taxa.
We chose the ITS region in large part because
Hardway et al. (2004) found evidence for
particularly rapid evolution in the Oenanthe clade
that includes Lilaeopsis when compared to the
rest of the taxa. Sequence divergences in this
clade averaged 6—7 times higher (approx. 17%)
than between species in Oenanthe (approx. 2.8%)
or Cicuta (approx. 2.4%) (Hardway et al. 2004).
AFLP analysis was selected as a secondary
marker system based on the increasing popularity
of this form of DNA fingerprinting as a
complementary system in phylogenetic studies
(Holland et al. 2008). Additionally, AFLP
fingerprinting offers a reliable, robust, and
genomically comprehensive method of genetic
analysis for taxa lacking complex nuclear and
organellar markers (Vos et al. 1995).
MATERIALS AND METHODS
Field Collection
Lilaeopsis masonii specimens were collected in
the spring of 2007 from locations along the Napa
River and in the Sacramento/San Joaquin Delta.
Lilaeopsis occidentalis was collected from Bodega
Head, California, and Mason and Lawrence lakes
in Washington State (Fig. 1). Leaf material to be
used in DNA extraction was preserved in silica gel
at the time of collection. Vouchers were deposited
at the herbarium at San Francisco State Univer-
sity (SFSU) (Table 1). Material for L. brasiliensis
(Glaz.) Affolter and L. mauritiana G. Petersen & J.
Affolter was obtained from a commercial aquarium
supplier (freshwateraquariumplants.com). The
dataset is composed of 35 nrDNA ITS sequences
representing seven taxa, including three sequences
MADRONO
[Vol. 58
Washington Samples
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‘Hill Slough/Grizzly Island
Liberty Island
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134
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this study.
from Genbank (L. carolinensis J. M. Coulter &
Rose, L. novae-zelandiae (Gand.) A. W. Hill, and
L. occidentalis), two specimens from the aquarium
trade labeled as L. brasiliensis and L. mauritiana,
and two specimens of L. schaffneriana (Schltdl.) J.
M. Coulter & Rose subsp. recurva (A. W. Hill)
Affolter courtesy of the Desert Botanical Garden
staff. Comprehensive sampling was conducted for
L. masonii and L. occidentalis as the purpose of
this study was to resolve the taxonomic classifi-
cation for these two species. A detailed systematic
O \3sr5 75 150 Miles
Map of the geographic locations of Lilaeopsis masonii and L. occidentalis specimens collected for
study for Lilaeopsis is in preparation (S. Downie,
Univ. of Dlinois, Urbana-Champaign, personal
communication).
ITS Methods
DNA from leaf tissue of five of the seven
species was extracted using the Qiagen DNeasy
Plant Mini Kit (QIAGEN, Inc., Valencia, CA),
following the manufacturer’s protocol with slight
modifications. Lilaeopsis carolinensis and _ L.
135
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‘SASATVNV dIHV GONV SI] Od SUAAWOAN NOISSAOOY ANVENAD GNV NOILVNYOAN] NOLLOATIOD
DUDIJIANDUL *
SISUAI]ISDAG *
DUDIAIUL{DYIS °
SI]DJUIAPIIIO *
IDIPUDJIZ-IDAOU *
1UuOSDUL "JT
SISUIUTJOADI “T
so1sodsg
‘| alav bs
136
TABLE 2. AFLP PRIMER AND ADAPTER SEQUENCES.
Primer
AdlEcoRI
Ad1Msel
prampEcoRI
prampMsel
FAM-EcoRI
HEX-EcoRI
MselI + CAA
Msel + CAT
MselI + CAG
novae-zealandiae were excluded from this analysis
due to technical difficulties with the DNA
extraction from the leaf material. Dilutions of
the genomic DNA extract of 1:10 in ultrapure
water were used in PCR reactions. The contigu-
ous ITSI, 5.88, and ITS2 regions of nuclear
ribosomal DNA were PCR-amplified using the
primers ITSLEU (Baum et al. 1998) and ITS4
(White et al. 1990) in final reaction volumes of
25 ul. Positive amplifications were purified using
the MO BIO UltraClean PCR Clean-up DNA
Purification Kit (MO BIO Laboratories, Inc.,
Solana Beach, CA). Internal primers ITS2 and
ITS3 (White et al. 1990) were used in addition to
ITSLEU and ITS4 in cycle-sequencing reactions
in order to extend fragments and clarify ambigu-
ities. Fragments were sequenced with the BigDye
3.1 kit (Applied Biosystems, Foster City, CA)
following the manufacturer’s protocols, and
visualized using the ABI PRISM 3100 Genetic
Analyzer (Applied Biosystems). Sequences were
manually aligned using Sequencher 3.1.1 (Gene-
Codes Corp., Ann Arbor, MI) and MacClade
4.04 (Maddison and Maddison 2001).
AFLP Methods
AFLP fingerprinting was conducted follow-
ing a modified protocol based on the methods
described by Vos et al. (1995). DNA extracts
prepared for ITS analysis also were used for
this study undiluted. Approximate DNA con-
centrations for all samples were estimated to
contain a range of concentrations from 10 ng/ul
to 50 ng/ul using an ethidium dot test. DNA
template of each sample was digested using the
infrequent endonuclease cutter EcoR1 and the
frequent endonuclease cutter Msel. Immediately
following digestion, the entire digestion reaction
was combined with an equal volume of ligation
mix.
The resulting fragmented DNA template con-
taining “sticky ends” was diluted five-fold and
subsequently amplified by PCR using a pre-
selective primer mix. This step effectively reduces
the number of possible fragments by approxi-
mately 1/16" (Meudt and Clarke 2007). The
MADRONO
[Vol. 58
Sequence
5’ =CTCGTAGACTGCGTACG— 3”
5 =CGACGATGAGRCCTGAG— 3!
5’ =GACTGCGTACCAATTCA— 3’
5 “GATGAGTCCTGAGTAAC= 3’
5’ -GACTGCGTACCAATTCAAC- 3’
5) =GACT GCG TAGCAATTCACG=3"
5 =GATGAGTCCTGAGTAACAA— 3
5’ -GATGAGTCCTGAGTAACAT= 3’
5’ -GATGAGTCCITGAGTAACAG— 3”
pre-selective reaction condition consisted of 30
cycles of 94° for 30 sec, 56° for 1 min, and 72° for
1 min.
Three combinations of selective primer sets
were used to produce a final AFLP fingerprint
for each sample (Table 2). Each set of selective
primers consisted of a primer region matching the
known adapter sequence, as well as three selective
nucleotides on the 3’ end of the MseI primer and
three selective nucleotides plus a florescent label
on the 3’ end the of EcoRI primer. Template for
the pre-selective PCR was diluted 6-fold and then
combined with a master mix containing one set of
selective PCR primers. A step-down PCR was
used to amplify the selective fragments in a
program consisting of 13 cycles of 94° for 30 sec,
65° for 30 sec (—0.7° per cycle), and 72° for 1 min,
followed by 24 cycles of 94° for 30 sec, 56° for
30 sec, and 72° for 10 sec.
The final selective PCR product fragments
containing a fluorescently labeled EcoRI end and
unlabeled Msel end were analyzed undiluted
using an ABI 3100 genetic analyzer (Applied
Biosystems). Initial fragments were sized first
using the analysis software GeneScan (Applied
Biosystems) and using the program by GeneMar-
ker© (SoftGenetics, State College, PA). After a
comparison of fragment calling using both programs,
all samples were analyzed using GeneMarker®.
Fragments were recorded for each sample in a
data matrix based on a binary system (1 for
presence, 0 for absence); a data matrix was
developed for each primer combination and then
all data was collated into a single data matrix. To
test reproducibility of results, twenty percent of
all samples selected at random for each primer
pair were re-analyzed, starting with the initial
DNA extracts. Fragment peaks that were deter-
mined to be consistently low (below 300 peak
intensity) or unpredictable were dropped from
the matrix table.
Data Analysis
Phylogenetic analyses of ITS sequences were
conducted using Phylip version 3.68 (Felsentein
2004). All characters were weighted equally,
2011]
character state transformations were treated as
unordered, and gaps were treated as missing data.
Most-parsimonious trees were obtained in Phylip
using the “‘branch-and-bound” method of exact
search implemented by the analysis unit DNA-
PENNY. Bootstrap re-sampling (1000 replicates)
was used to assess nodal support (Felsenstein
1985). Most parsimonious trees were generated
from a search of 100,000 trees and a final tree was
derived using a strict consensus tree method
(Felsenstein 2004). Additional tree searches were
conducted in greater volumes, up to 1,000,000.
However, larger searches produced the same final
tree, thus a smaller tree search was selected to
reduce run-time during bootstrapping. Several
combinations of Lilaeopsis species outgroups
were explored before selecting L. novae-zelandiae
as the outgroup. This selection was based on
indications of a potential sister group relationship
between L. novae-zelandiae and L. occidentalis,
which was supported by ITS phylogenetic anal-
ysis of this genus within the Apiaceae tribe
described in Downie et al. (2008). Genetic
distances to determine branch lengths were
calculated in Phylip using the Jukes-Cantor
method implemented in DNADIST and a
Fitch-Margoliash (FITCH) search.
AFLP phylogenetic analysis was performed us-
ing the program Phylip version 3.68 (Felsenstein
2004). A genetic distance matrix was created
using the techniques described by Nei and Lei
(1979) as implemented by RestDist in Phylip
(Felsenstein 2004). The output matrix was then
input into NEIGHBOR using the UPGMA
method of cluster analysis (Felsenstein 2004).
Using this approach, an output tree was con-
structed by successive clustering using an aver-
age-linkage method of clustering. The output file
was then plotted as both a rooted and unrooted
tree. A search for the most parsimonious trees
was implemented first using the branch-and-
bound algorithm of DOLPENNY (100,000 trees
searched) in Phylip following bootstrap analysis
using 100 replicates. A 50% majority-rule con-
sensus tree was then generated to condense the
results into a final tree, which is presented here.
Previous studies of Lilaeopsis using AFLP
analysis have not previously been reported. As
such, outgroup selection for AFLP parsimony
analysis was determined following variable,
preliminary analysis replicates. Lilaeopsis schaff-
neriana was selected as this species is the closest
geographically to both ZL. masonii and _ L.
occidentalis. Further, L. schaffneriana also dem-
onstrated sufficient genetic differences to be used
as an appropriate outgroup.
To further visualize potential multi-dimen-
sional correlation of AFLP data based on
genetic similarities, an additional genetic dis-
tance matrix was derived using Manhattan
distance (StatisiXL; www.statistixl.com). These
FIEDLER ET AL.: RECONSIDERATION OF MASON’S LILAEOPSIS [37
data were then analyzed using a _ principal
coordinates analysis (PCA) using the Microsoft
Excel© add-in program GenAIEx 6.2 (Peakall
and Smouse 2006). To determine whether a
measurable degree of genetic dissimilarity
among L. occidentalis (WA and CA samples)
and L. masonii (CA samples) could be attributed
to geographic distance, an additional test of
molecular variance based on geographic origin as
measured by Global Position System (GPS) also
was tested. A two-way analysis of variance was
assessed for collections of L. occidentalis and L.
masonii using an analysis of molecular variance
(AMOVA) with GenAIEx 6.2 (Peakall and
Smouse 2006). Significance was assessed using
99 permutations.
RESULTS
DNA sequence analysis of the ITS regions
ITS1, 5.88, and ITS2 of nuclear ribosomal DNA,
based on a most parsimonious search of 64,631
trees using a branch-and-bound method, revealed
no differences between California samples of L.
occidentalis and L. masonii samples, including a
GenBank accession for L. occidentalis (100 of 100
trees) (Fig. 2). Within the L. occidentalis\L.
masonii clade, samples collected from Washing-
ton clustered separately. Distance values between
Washington and California samples were low for
single collections from Lawrence Lake and
Mason’s Lake (0.1 and 0.3%, respectively).
However, a second sample from Lawrence Lake
exhibited higher distance values (1.5%), which
may be due to missing data. Distance values for
California samples of L. masonii and L. occiden-
talis were 0% across all samples. Comparatively,
distance values between the additional species
used for this study ranged from 1.2—8%. Distance
based analysis of ITS sequences found an
identical tree structure as the strict consensus
tree inferred from most parsimonious results
implemented by DNAPENNY. Bootstrap esti-
mates from 1000 replicate analyses yielded 100%
nodal support for all branches. Branch placement
and relationship of Lilaeopsis species used in this
study are consistent with results of a previous ITS
phylogenetic analyses by Downie et al. (2008),
though that study excluded L. masoniti.
The three AFLP primer combinations gener-
ated 274 unique fragments among 29 samples,
with only 21 fragments shared or monomorphic
between the five species used in this study.
Although a small sample size was used, specifi-
cally for L. brasiliensis, L. schaffneriana, and L.
mauritiana, the large number of shared fragments
is potentially indicative of low genetic diversity
within this genus, which is consistent with species
exhibiting high morphologic plasticity (Linhart
and Grant 1996). The number of total diagnostic
bands from the three markers combined data set
138 MADRONO
t—{ 0.003 substitutions/base
L. brasiliensis2
L. brasiliensis?
L. mauritiana
L. occidentalis (Bodega Bay, CA)
1A-B04a, Napa
L. occidentalis (Mason Lake, WA)
L. occidentalis (Lawrence Lake2, WA)
L. occidentalis (Lawrence Lake,1 WA
L. occidentalis (GenBank)
3 Mile Slough, CA
Grizzly Island, CA
Brannan Island, CA
1A-H07b, Napa
1A-29a, Napa
Collinsville, CA
2W-B02a, Napa
2w-B02b, Napa
2E-A04, Napa
McAvoy Harboy, CA
Reach 3, Napa
2E-B07a, Napa
2E-A07, Napa
1A-B04b, Napa
Liberty Island, CA
1A-H07a, Napa
2W-B02a, Napa
1A-001, Napa
1A-29b, Napa
Maxwell Bridge, Napa
Twitchell Island, CA
1B-S07, Napa
L. carolinensis (GenBank)
L.schaffneriana?
L.schaffneriana2
L.novae-zelandiae (GenBank)
Fic. 2.
Strict consensus tree derived from ITS sequence data using the branch and bound method implemented
by DOLPENNY. Branch values are Bootstraps. Upper left tree illustrates distance values, branch lengths are
proportional to the number of nucleotide substitutions per base.
varied between species, ranging from a low of 98
bands for one sample of L. schaffneriana, to a
high of 135 bands for the Mason Lake, WA
sample of L. occidentalis, with a mean number of
AFLP bands equaling 123 (SD = 9.5) (Table 3).
Within species genetic distance values (Man-
hattan distance calculated from the AFLP
matrix) ranged from a low of 1.4 for L. masonii
(Napa River site #1A-H07, large and small
forms), to a high of 6.6 for L. masonii (Napa
site 1A-BO4 and Twitchell Island collections).
Between species values ranged from a low of 4.1
for L. occidentalis (Mason Lake, WA) and L.
masonti (Napa River #2W-B02), to high of 11.9
for L. schaffneriana and L. occidentalis (Lake
Lawrence, WA collection).
Principal coordinate analysis (PCA) illustrated
an overlapping association between samples of L.
occidentalis and L. masonii (Fig. 3), but a clear
differentiation between the L. occidentalis/L.
masonii cluster and all other Lilaeopsis species
examined in this study, i.e., L. schaffneriana, L.
mauritiana, and L. brasiliensis. The small separa-
tion observed between the L. occidentalis and L.
masonii data may be attributed to geographic
distance. Results of AMOVA analysis derived
2011] FIEDLER ET AL.: RECONSIDERATION OF MASON’S LILAEOPSIS 139
TABLE 3. AFLP FRAGMENT NUMBERS FOR EACH SPECIES.
Mean or Total No. of Fragments
Species CAA CAG CAT Total
L. masonii 33.0 46.1 21.9 125.1
L. occidentalis a paw 50 22:9 128.8
L. schaffneriana subsp. recurva 47.5 34.5 16 99.5
L. brasiliensis a2 39 21 114
L. mauritiana 43 38 16 99
All Fragments 123
from grouping samples based on geographic
location indicated that approximately 73% of
genetic variation was distributed between groups
and thus 23% among groups, supporting the
conclusion that most observed genetic variation 1s
due to geographic distance (P = 0.01).
The UPGMA cluster phenograms provides
additional support for the results of the PCA
analysis, illustrating a combined grouping of L.
occidentalis (CA and WA) and L. masonii
samples (Fig. 4). Within the L. occidentalis and
L. masonii clade, samples collected from Wash-
ington clustered separately from samples col-
lected within California, corroborative of results
of AMOVA indicating that variation within this
clade is due in large part to geographic distance.
Samples of L. brasiliensis and L. mauritiana
cluster separately but are sister to the L.
occidentalis/L. masonii clade; L. schaffneriana
samples also cluster separately but are sister to
all other specimens/species used for this study.
The most parsimonious tree from the maximum
parsimony analysis supports a single L. occiden-
talis/L. masonii clade; however, additional reso-
(gm
> L. occidentalis (CA)
Lioccidentalis (WA)
Coord.2
:
|
|
| : L.masonii
lution within this taxon is less certain of the
specific placement of Lilaeopsis samples, based
on geographic location (Fig. 5).
DISCUSSION
Taxonomic implications. Within the last two
decades, the use of genetic techniques to distinguish
discrete evolutionary units has become common
place in systematic biology. Use of genetic data in
the protection of endangered species when morpho-
logical (or other character) information is either
unreliable or 1mpossible is just one reason why this
approach to species identification and delimitation
is SO Important (Avise 2003). Thus, sole reliance on
morphological, geographic, reproductive behavior
or some combination of non-genetic characters to
delimit taxa is no longer defensible when diagnostic
genetic information is available and can be readily
assessed. In the case of Lilaeopsis masonii and L.
occidentalis, neither ITS sequence nor AFLP
fragment length data support the recognition of
the L. masonii as a distinct evolutionary entity.
L.schaffneriana
L. mauritiana
L. brasilensis
Coord.1
FIG. 3.
Principal coordinates analysis (PCA) of AFLP fragment data matrix. Codes for Napa collections, e.g.,
1A-H07, indicate different collection locations and dates along the Napa River specific to the Napa River Flood
Protection Project.
140 MADRONO
39.8
49.5
59
[Vol. 58
9.5 Lawrence Lake earns
Lawrence Lake 2 Li/aeopsis
128 occidentalis
: Lawrence Lake 3 (Washington)
9.5 Mason Lake 1
Mason Lake 2
1A-29a
1A-B04a Ey
3. 5 A-29b L. masonii
(Napa, CA)
1A-H07a
4A-HO7b
3S Maxwell Bridge
2W-B02a
2W-B02b
McAvoy Harbor
3 Mile Slough
Brannan Island
L. masonii
Liberty Island (California)
Collinsville |
Grizzly Island
4{A-001
L. masonii
Reach 3 (Napa, CA)
8.3 2E-B07
9.2 __L. occidentalis
=e Bodega Head — atifornia)
13. L. masonii
1A-B04b (Napa, CA)
14 : L. masonii
Twitchell Island — (California)
Lilaeopsis brasiliensis
Lilaeopsis mauritiana
2.9 Lilaeopsis schaffneriana1
Lilaeopsis schaffneriana2
Fic. 4. UPGMA Cluster Phenogram (rooted and unrooted trees) from AFLP data matrix of five species of
Lilaeopsis. Numbers are distance values. Codes for Napa collections, e.g., 1A-H07, indicate different collection
locations and dates along the Napa River specific to the Napa River Flood Protection Project.
The morphological and geographic information
to support two distinct taxa is weak, ambiguous,
and unreliable at best.
Fallon’s (2007) arguments regarding the im-
portance of using genetic information to resolve
taxonomic issues for species protection is borne
out in our study. While her review focused solely
on vertebrates, and on only those species,
infraspecific or population segments proposed
for listing, not those already listed, our results
add further emphasis for use of molecular
techniques in conservation efforts. We concur
that multiple genetic markers are essential for a
thorough assessment of taxonomic or population
unit (or at any appropriate level) when consider-
ing of formal protection. We further suggest that
use of best available science such as existing or
generating new genetic information is equally
valid for the periodic reviews of listed species
required of both the federal and state agencies.
Further and relevant to L. masonii, use of genetic
data is likely to be essential during a de-listing
review process.
Based upon several lines of evidence, including
decades of fieldwork throughout the range of L.
masonii, observations from the most recent
monograph (Affolter 1985), and our molecular
genetic analyses, we urge that this rare taxon no
longer be recognized as a separate taxonomic
entity. Rather, L. masonii should be subsumed
within the larger, much more widespread, com-
mon, and equally variable species, L. occidentalis.
2011]
96/100
100/100
71/100
96/100
57/100
58/100
FIEDLER ET AL.: RECONSIDERATION OF MASON’S LILAEOPSIS 14]
Reach 3, Napa
A1-001, Napa
L. occidentalis (ML1, WA)
L. occidentalis (ML2, WA)
Maxwell Bridge
L. occidentalis (LL3, WA)
1A-29b, Napa
1A-B04a, Napa
1A-29a, Napa
2W-B02a, Napa
3 Mile Slough, CA
McAvoy Harbor, CA
L. occidentalis (Bodega Hd,CA)
L. occidentalis (LL1, WA)
1A-H07a, Napa
1A-H07/b, Napa
Brannan Island, CA
Grizzly Island, CA
2W-B02b, Napa
Collinsville, CA
1A-B04b, Napa
L. occidentalis (LL2, WA)
2E-B0/, Napa
Twitchell Island, CA
Liberty Island, CA
L. brasiliensis
L. mauritiana
L. schaffneriana1
L. schaffneriana2
FIG. 5.
Most parsimonious tree from AFLP data matrix of five species of Lilaeopsis. Codes for Napa collections,
e.g., LA-H07, indicate different collection locations and dates along the Napa River specific to the Napa River
Flood Protection Project.
Significant morphological, but limited genetic,
variation exists both within and among popula-
tions of Lilaeopsis throughout the Pacific Coast
of North America, from the Queen Charlotte
Islands in British Columbia to the inland islands
of the Sacramento-San Joaquin Delta in the
Great Valley of California. Importantly, this
variation does not follow any consistent environ-
mental gradient for either taxon. As such, one
intrinsically variable species, not two, of west
coast Lilaeopsis should be recognized in relevant
floras, including those for North America.
Important additional circumstantial support
comes from the very wide geographic ranges,
some amphitropical, of the great majority of
other species of Lilaeopsis, including L. chinensis
Kuntze, L. carolinensis, L. shaffneriana, L.
macloviana A. W. Hill, and L. novae-zelandiae,
among others. A large geographic range is not
surprising for all these species, given their
vigorous vegetative reproduction by easily frag-
mented rhizomes and their restriction to aquatic
habitats, many with bi- or multi-directional flow
vectors (e.g., Napa River, Sacramento River,
Pacific Ocean).
Field observations suggest a possible explana-
tion for the inter- and intra-populational varia-
tion individual ramet size for both western North
American Lilaeopsis. Periods of rapid spring
growth occur during the spring tides and
snowmelt from the Sierra Nevada, when temper-
atures warm sufficiently to encourage an increase
in photosynthetic activity. This increased vegeta-
tive growth occurs when floodwaters from the
142 MADRONO
Sacramento, San Joaquin and Napa Rivers are at
their height of volume and rate of flow. Thus, the
comparatively high kinetic energy of flowing
water during the spring run-off, coupled with
this species’ preference for river banks and shores
characterized by high light and open exposure,
combine to restrict vegetative growth to a
comparatively shorter plant less vulnerable to
being dislodged from its habitat. Relatively taller
Lilaeopsis ramets are invariably found compara-
tively further from the shoreline in shadier and
relatively lower energy microhabitats than com-
paratively shorter stature ramets. Aquaria enthu-
siasts who work with various species of Lilaeopsis
have dubbed the short stature coupled with dense
growth phenomenon the “lawn effect” (http://
www. freshwateraquariumplants.com).
Conversely, observations of Lilaeopsis species
submerged in (low energy) water reveal individual
leaves grow comparatively longer. Affolter’s
(1985) greenhouse experiments and observations
that demonstrated that for least eight of the 13
species Lilaeopsis studied (including L. occidenta-
lis), material grown in submerged pots had larger
and wider leaves, more septae, and wider rhizome
diameter. In his monograph of the thirteen
species known in 1985, increased periods of
inundation result in a suite of morphological
changes, including an increase in leaf length and
increases in both peduncle and pedicel lengths
(Affolter 1985). Lastly, the rejection of leaf length
as a key diagnostic character distinguishing two
otherwise very similar taxa has precedent in
Affolter’s lumping of all Andean, Fuegian, and
Patagonian material into a single species, L.
macloviana, synonomizing thirteen previously
described taxa.
Regulatory implications. Neither CESA nor the
federal ESA, as amended, protects any vascular
plant distinct population segment as does the
ESA for specific vertebrate populations. While an
argument can be made that this is a form of taxon
chauvinism, plant species are not protected below
the infraspecific level. Such a comparison is
important, because some vertebrate species that
were listed relatively soon after the ESA was
passed have since been determined not genetically
distinct from common widespread relatives, but
they continue to be formally protected because of
the DPS provisions. For example, the San
Francisco garter snake (Thamnophis sirtalis tetra-
taenia), a highly restricted taxon in central coastal
California, was determined, through an exami-
nation of the clade’s mtDNA (Janzen et al. 2002),
to be a member of a California clade of the
widespread common garter snake. These authors
concluded that morphologically based subspecies
designations of 7. sirtalis in western North
America were invalid because they did not reflect
reciprocal monophyly of mtDNA _ sequences.
[Vol. 58
Extrapolating Janzen et al.’s (2002) logic to our
genetic work with Lilaeopsis, the parallel conclu-
sion that the specific designation of L. masonii is
invalid is compelling. Because neither the CESA
nor the ESA include DPS provisions for plant
species, L. masonii no longer warrants protection
as a State “rare” species and the allocation of
limited recovery resources. Given the widespread
nature of Lilaeopsis occidentalis + L. masonii, and
the large number of projects (both existing and
proposed) requiring mitigation and monitoring of
the rare L. masonii, a timely review of our
findings is essential. Conservation dollars are few,
and they should be applied to truly rare,
threatened, and real discrete species.
Finally, Pavlik (2003) recently examined the
role of state- and federally-listed species protect-
ing the ecosystems in which they are found. Of
relevance is the notion that some _ protected
species provide a “regulatory umbrella” for other
species that are unlisted, but are rare, in decline,
or otherwise of conservation concern. Lilaeopsis
masonii has long served to restrict, prevent, or
slow the conversion, degradation or destruction
of wetlands throughout the Sacramento-San
Joaquin Delta, Suisun Marsh, and San Francisco
Bay ecosystems, thereby protecting associated
but unlisted species of conservation concern.
While a suite of other protected wetland plant
taxa exist in these ecosystems (e.g., Cirsium
hydrophilum Jeps. var. hydrophilum [Suisun this-
tle], Cordylanthus mollis A. Gray ssp. mollis [soft
bird’s beak], etc.), there are many more that are
rare, in decline, and not listed (e.g., Cicuta
maculata Lam. var. bolanderi (S. Watson) G. A.
Mulligan [Bolander’s water hemlock], Plantago
elongate Pursh [slender plantain], Lycopus asper
Greene) (see Baye et al. 2000). Thus we acknowl-
edge that a delisting of Mason’s lilaeopsis may
further expedite wetland habitat loss in central
California. Nonetheless, conservation in the
twenty-first century demands the use of best
available science, despite the unintended conse-
quences that may occur. Ultimately, government
agencies charged with the protection of our
biodiversity must redouble their efforts to em-
brace new scientific results that affect listed
species, commit to diligent review of listed and
candidate species, and disseminate accurate and
up-to-date information. Similarly, conservation-
ists should redouble their efforts to provide the
best available science for decision-making. The
time to embrace current molecular genetic
techniques in routine conservation decision-mak-
ing has come.
ACKNOWLEDGMENTS
The authors would like to thank the Napa County
Flood Control and Water Conservation District,
particularly R. Thomasser, for funding this work;
L. Garrison for assistance in the field and laboratory;
2011]
A. Fullerton for collection of two Lilaeopsis from
Washington state, and J. Johnson for collection of
L. schaffneriana subsp. recurva from Scotia Canyon in
Arizona. We also wish to thank S. L. Krauss and F.
Cipriano for guidance in the ITS and AFLP analyses, and
the Conservation Genetics Laboratory, San Francisco
State University for access to its facility. Lastly, we would
like to thank J. Affolter, S. R. Downie, and B. J. Grewell
for reading the manuscript and for their helpful comments.
Four anonymous reviewers also provided comments that
improved this manuscript.
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BAYE, P. R., P. M. FABER, AND B. GREWELL. 2000.
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MADRONO, Vol. 58, No. 3, pp. 145—152, 2011
A COMPARISON OF THE EFFECTS OF NA>;SO, AND NACL ON THE GROWTH
OF HELIANTHUS PARADOXUS AND HELIANTHUS ANNUUS (ASTERACEAE)
M. O. MENDEZ! AND O. W. VAN AUKEN
Department of Biology, University of Texas at San Antonio, San Antonio, TX 78249
monica.mendez@tamiu.edu
ABSTRACT
Helianthus paradoxus Heiser (Asteraceae, puzzle sunflower), is a federally threatened hybrid species
found in salt marshes of west Texas and New Mexico. Helianthus annuus L. (Asteraceae, common
sunflower) is one of the parent species and is found throughout North America, but it is not present in
the inland salt marshes where H. paradoxus is found. Helianthus paradoxus has previously been
described as a halophyte, but its tolerance to Na »SOx4, one of the major salts found in its habitat, has
not been investigated. However, salinity has been identified as a major abiotic factor influencing the
limited distribution of H. paradoxus populations. In this greenhouse study, the effects of elevated
concentrations of NasSO,4 and NaCl, at equal ionic strengths (0.00, 0.09, 0.17, 0.34, and 0.51), on the
survival and dry mass of both H. paradoxus and H. annuus were examined. In the three-way factorial
experiment, the effects on dry mass observed were dependent on the species, the type of salt and the
ionic strength of the salt. Helianthus paradoxus produced more dry mass than AH. annuus in both salt
treatments; however, NaCl was more inhibitory of dry mass production for both species with plants
unable to survive the highest salt treatments. While dry mass of H. annuus decreased with increasing
ionic strengths of both salts, dry mass of H. paradoxus increased by 38 to 72% in low to moderate
ionic strengths of NazSO, relative to the nonsaline treatment. Both species were less tolerant of NaCl
than Na,SO, with A. paradoxus seeming to have moderate and high tolerance to elevated Cl” and
SO,’ ionic strength, respectively, while H. annuus had low to moderate tolerance. Greater dry mass
production in Na»,SOxq, along with tolerance to both salts, suggests that low to moderate sulfate soil
salinity will enhance the dry mass production of H. paradoxus.
Key Words: Halophyte, Helianthus annuus, helianthus paradoxus, ionic strength, NaCl, Na>»SOxg, salt
tolerance, sunflower.
Helianthus paradoxus Heiser (Asteraceae, puz-
zle sunflower) is a federally threatened species
with limited distribution in salt marshes in west
Texas and New Mexico (Correll and Johnston
1979; Poole and Diamond 1993; McDonald
1999). Hybridization studies (Heiser 1958, 1965;
Abbott 1992) and molecular analysis (Rieseberg
et al. 1990; Rieseberg 1991; Rieseberg et al. 1991)
have determined that H. paradoxus is a stabilized
hybrid species between H. annuus L. (Asteraceae,
common sunflower) and H. petiolaris Nutt.
(plains sunflower). Although H. paradoxus shares
several morphological and ecophysiological traits
with its parental species (Rosenthal et al. 2002), it
has diverged and is genetically isolated from its
progenitors and considered a separate species.
Helianthus anuus is common throughout North
America and grows in disturbed, heavy clay soils
that are moist in the spring and dry out by mid-
summer. Helianthus petiolaris is found in dry,
sandy soils in western North America, while H.
paradoxus grows in heavy, waterlogged, saline
soils (Van Auken and Bush 1998).
‘Current address: Department of Biology & Chem-
istry, Texas A&M International University, 5201
University Boulevard, Laredo, TX 78041, USA.
Ecological and ecophysiological studies of H.
paradoxus have determined that this homoploid
hybrid species is salt tolerant, unlike its parental
species. Helianthus paradoxus 1s restricted to
inland salt marshes with salt levels of approxi-
mately 10 g kg ' (Poole and Diamond 1993; U.S.
Fish and Wildlife Service 2005; Grunstra and
Van Auken 2007a, b), while H. annuus and H.
petiolaris can be found in low saline soils
(<0.02 g kg™' soil sodium, Welch and Rieseberg
2002). Helianthus paradoxus is a better compet-
itor than its progenitors in saline soils (Bush and
Van Auken 2004). The west Texas and New
Mexico salt marshes were key habitats in
isolating hybrids (Abbott 1992). The parental
species are glycophytes and cannot survive the
same habitat as H. paradoxus, where other salt
tolerant plants are generally present (Poole and
Diamond 1993: Lexer et al. 2003: U.S. Fish and
Wildlife Service 2005). Additionally, H. para-
doxus is capable of sequestering higher sodium
and sulfur concentrations and produces greater
leaf succulence compared to its parental species
while maintaining significantly greater fitness
when grown in elevated NaCl concentrations
(Welch and Rieseberg 2002) or in field-like highly
saline soil conditions (Karrenberg et al. 2006).
In the largest known population of H. para-
doxus, at the Diamond-Y Spring Preserve near
146
Fort Stockton, Texas, the distribution of JH.
paradoxus is mainly affected by soil salinity and
soil moisture gradients when biotic factors are
not considered (Bush and Van Auken 1997; Van
Auken and Bush 1998; Bush 2006a, b). This large
desert spring and associated salt marsh has had
1.44 to 2.70 million H. paradoxus plants (Van
Auken and Bush 1998), depending on environ-
mental conditions. In addition, there are several
rare and federally endangered invertebrates
found in the marsh (McDonald 1999). Helianthus
paradoxus plants consistently establish parallel to
the drainage of Leon Creek but their proximity
depends on seasonal climatic conditions influencing
soil salinity levels and soil water content, both of
which decrease dramatically upland from the lowest
point in the salt marsh (Van Auken and Bush 1998;
Grunstra and Van Auken 2007a, b). When annual
rainfall is high, H. paradoxus can be found further
from the drainage compared to drier years when
plants are located closer to the drainage. The
population of H. paradoxus at the Diamond-Y
Spring Preserve seems to be dependent on low to
intermediate salinity levels and intermediate
moisture levels; however, salinity appears to be
the major abiotic factor affecting the local
distribution of H. paradoxus (Bush 2006a, b).
Previous studies have identified H. paradoxus
as a salt tolerant species with characteristics of
halophytes (Welch and Rieseberg 2002; Bush and
Van Auken 2004). However, these studies were
done with NaCl, one of the major salt compo-
nents of the marsh, but not Na»,SO, which is also
found at high concentrations in soils of its salt
marsh habitat (Boghici 1997). In addition, the
effects of a limited range of salinity levels have
been examined. Chloride is generally more toxic
than SO,°- and even Na‘ at lower concentrations
(Manchanda et al. 1982; Marschner 1995; Frank-
lin and Zwiazek 2004; Munns and Tester 2008);
however, SO, salinity can be more growth
inhibitive than Cl” for some halophytes (Warne
et al. 1990; Egan and Ungar 1998). At the same
time, separating specific ion effects is difficult and
differential effects of Cl” and SO,’ salinity on H.
paradoxus are still unknown; consequently, the
influence of these anions on the ecological
isolation and distribution of H. paradoxus has
not yet been elucidated.
Ionic strength rather than salt concentration
was used as a treatment variable in the present
study. It is used as a normalization procedure and
was required because of different numbers of ions
present in equal molar solutions of NaCl (2 ions)
and Na>SO, (3 ions). Therefore, it was important
to compare the two salts at concentrations that
allowed osmotic potential of the corresponding
treatments to be equal. Concentrations of the two
salt treatments were based on equal ionic
strengths calculated using procedures in Barrow
(1966).
MADRONO
[Vol. 58
TABLE 1. IONIC STRENGTHS OF EACH TREATMENT
AND CORRESPONDING SALT CONCENTRATIONS IN
G:-KG ' EXAMINED IN THE STUDY.
NaCl Na»SOx,4
Ionic strength (g kg~')
0.00 0 0
0.09 eS) 4.1
0.17 10 8.1
0.34 20 16.2
ee | 30 24.3
The objective of this study was to examine the
survival, growth and the salt tolerance of ZH.
paradoxus in elevated levels of both Na»SO, and
NaCl, at equal ionic strengths. Both of these salts
are major contributors of the soil salts of the
Diamond-Y Spring Preserve. Our experiment
also included the more salt tolerant of its parental
species, H. annuus, a known glycophyte.
MATERIALS AND METHODS
Helianthus paradoxus seeds were collected from
the Nature Conservancy’s Diamond-Y Spring
Preserve near Fort Stockton, Texas (31°00.54’N,
102°55.49’W) and stored dry at 25°C until used.
Helianthus annuus seeds were purchased from
Native American Seed Farm (Junction, Texas
76849) and stored dry at 4°C. Seeds of both
species were cold stratified in Ziploc® storage
bags lined with paper towels, wet with deionized
water, and maintained at 4°C in the dark for
4 weeks (H. paradoxus) or 8 weeks (H. annuus).
To prevent osmotic shock, groups of seeds
(approximately 400 seeds for each treatment)
were germinated on paper toweling saturated
with deionized water (0.00 osmotic strength) in
plastic storage bags at 25°C. Germinated seed-
lings were transferred to equal or increasing levels
of the appropriate salt (NaCl and Na ,SOy, at
ionic strengths of 0.00, 0.05, 0.09, 0.13, 0.17, 0.26,
0.34, 0.43, and 0.51) every 2 d until placed in the
final salt (NaCl or Na>»SO,) and ionic strength to
be tested (0.00, 0.09, 0.17, 0.34, and 0.51).
Seedlings were kept in the appropriate solution
for a total of 18 days prior to transplanting to
pots in the greenhouse containing the corre-
sponding treatment (Table 1).
For each species, five plants per pot with five
replicate pots per treatment (25 plants per species
per treatment) were grown for 62 d. Plastic pots
(15 cm diameter < 15 cm height) lined with a
Ziploc® storage bag (to retain water, salts, and
nutrients) were filled with 1.4 kg of air-dried,
sieved (5.8 mm mesh screen) soil. The soil was the
upper 10 cm of a low nutrient Patrick-series
Mollisol (clayey-over-sandy, carbonatic-thermic,
typic calciustoll), collected from northern Bexar
Co., Texas (Taylor et al. 1966; Van Auken and
Bush 1998). The soil was friable, allowing root
2011]
TABLE 2.
MENDEZ AND VAN AUKEN: SALT TOLERANCE OF TWO HELIANTHUS SPECIES
147
THREE-WAY ANOVA RESULTS FOR THE TOTAL DRY MASS (G) OF TWO SPECIES (HELIANTHUS
PARADOXUS AND H. ANNUUS) GROWN IN TWO SALTS (NACL AND NA>SQOq) AND AT FIVE IONIC STRENGTHS (0.00,
0.09, 0.17, 0.34, AND 0.51). All main effects and interactions were significant. “ Significant interactions are
designated (* = P < 0.05; ** = P < 0.01; *** = P < 0.001; **** = P < 0.0001).
OQ.
mh
Source*
Species
Salt
Strength
Species < salt***
Species < strength**
Salt X strength****
Species X salt < strength*
Error
Total
Oo HH Hee
Nome.<)
extraction and recovery (Bush and Van Auken
2004). Appropriate amounts of anhydrous salts
(Na>SO,q, or NaCl) and a single nutrient applica-
tion (0.05 g of P from Na,HPO, - 7H,O, 0.07 g N
from NH4NOs3, 0.07 g of K from KCl, and 0.03 g
of S from MgSO, 7H,O; Tiedemann and
Klemmedson 1986) were added to each pot and
thoroughly mixed. Before seedlings were planted,
400 ml of deionized water was added to each pot.
Thereafter, soil moisture was maintained at
approximately field capacity with distilled water.
This study was conducted in a fiberglass green-
house in which the daytime temperatures ranged
from approximately 26° to 38°C and light levels
were approximately 36% of full sunlight with a
mean photosynthetically active photon flux den-
sities of 562 + 135 umol m * s_' measured with a
Li-Cor® LI-188 integrating quantum sensor.
Plant survivorship was assessed at 62 d for
each treatment combination (species X< salt xX
ionic strength). Percent survivorship is based on
the mean number of erect and green plants out of
five plants per replicate pot per treatment.
Shoot and root dry mass were measured at the
end of the experiment. For shoot dry mass, plants
from each pot were clipped at the soil surface and
placed in a pre-weighed paper bag. For the ash-
free root dry mass, all of the soil and particulate
matter from each pot was carefully washed off
the roots. Roots were then wrapped in pre-
weighed aluminum foil. All plant material was
dried in a forced air oven at 90°C to a constant
mass. After drying, roots were ashed in a
programmable muffle furnace (Fisher Scientific
Isotemp®, Fisher Scientific Research, Pittsburgh,
PA) at 625°C for 3 hours to obtain the ash-free
root dry mass (Bohm 1979). Total dry mass was
also calculated by combining the shoot dry mass
and ash-free root dry mass per pot.
All dry mass data was analyzed using SAS
Statistical software (SAS Institute 1999) with each
pot as the unit of replication and P = 0.05 as the
criterion for significance. Any plant mortalities
because of a treatment effect were taken into
account by including a dry mass of zero. To
SS F Pp
4.11 35.69 <0.0001
14.75 128.17 <0.0001
5.00 43.44 <0.0001
Lo? 12.05 0.0008
0.35 4.78 0.0016
1.67 14.50 <0.0001
0.40 3-5) 0.0108
921
57.05
determine the effects of salt type and ionic
strength on the dry mass of each species, a
three-way ANOVA including interactions was
employed. When significance was detected with
the overall ANOVA, two-way and one-way
ANOVAs were used followed by Duncan’s
Multiple Range Test to examine significant
differences between all possible combinations of
salt type and salt concentration for each species
separately. Mean mortality was calculated, but
was not analyzed statistically.
RESULTS
All H. paradoxus and H. annuus plants died in
the highest ionic strengths of NaCl examined
(0.34 and 0.51). However, both species demon-
strated 100% survivorship in the no-salt treat-
ment. For all Na»,SO, treatments, 100% of H.
paradoxus plants survived, while 100% of the H.
annuus plants survived in each of the Na»,SO,
treatments except at the 0.51 ionic strength, the
highest Na»SO, concentration tested. Plant sur-
vivorship for H. paradoxus in NaCl treatments
was 100% at the 0.09 and 0.17 ionic strengths,
while 80% and 40% of H. annuus plants survived
in these same ionic strengths, respectively.
Three three-way ANOVAs were used to
analyze shoot, root, and total dry mass; however,
only total dry mass data will be presented.
Results were similar for mean shoot and root
dry mass (analysis not shown). The three-way
ANOVA (Table 2) demonstrated a significant
overall species, salt, and ionic strength effect on
total dry mass. In addition, the three two-way
interactions were significant as was the three-way
interaction. To demonstrate more clearly the
experimental results, two of the two-way interac-
tions will be presented first. Overall, H. para-
doxus produced more dry mass than H. annuus in
both salts (Fig. la, species < salt interaction).
For both species, more dry mass was produced in
the Na>»SO, treatment than in the NaCl treat-
ment. In addition, ionic strength was significant
with more dry mass in the lower treatments;
&
(oe)
]
a)
GH. paradoxus
[_] H. annuus
Total (Shoot + Root) dry mass (g pot’)
Salt
FIG. 1.
MADRONO
[Vol. 58
H. paradoxus
H. annuus
Total (Shoot + Root) dry mass (g pot*')
0.17 0.34 0.51
lonic Strength
Two-way interaction plots of total dry mass (bars) as the response variable for (a) salt (NaCl and Na,SOx,)
by species (Helianthus paradoxus and Helianthus annuus) and (b) ionic strength (0.00—0.51) by species. A three-way
ANOVA determined there were significant differences (P < 0.0001) between salt treatments and ionic strengths for
each species. Different letters indicate significant differences between means (Duncan’s Multiple Range Test). Lines
above the bars represent + one SD (standard deviation).
however, dry mass of H. annuus was significantly
lower at an ionic strength of 0.09 (Fig. 1b, species
xX 1onic strength interaction). The salt xX ionic
strength interaction has not been presented
because the results can be seen within the three
way interaction figure. The significant three-way
interaction indicated that dry mass was depen-
dent on species, salt type, and ionic strength
(Table 2, Fig. 2a, b). Helianthus paradoxus dry
mass was higher in Na,SO4 compared to NaCl
(Fig. 2a), with the greatest dry mass at the mid
ionic strengths (Fig. 2a). On the other hand, H.
annuus dry mass was lower than AH. paradoxus,
but H. annuus did produce more dry mass in
NasSO, compared to NaCl (Fig. 2b). In addi-
tion, as the ionic strength increased dry mass of
H. annuus decreased.
In comparison to the Na»,SO, treatments, dry
mass of both species was significantly reduced in
NaCl at elevated ionic strengths (Duncan’s
Multiple Range Test, P < 0.0001, Fig. 2a, b).
In the no-salt treatment, growth of H. annuus was
elevated compared to AH. paradoxus (not signifi-
cantly). However, for all levels of salt addition,
total dry mass of H. paradoxus was greater than
H. annuus, yet the differences were salt depen-
dent. For both species, mean total dry mass was
elevated in Na>,SOy, at an ionic strength of 0.09
compared to NaCl. Dry mass of H. paradoxus
was 32% greater than dry mass of H. annuus in
this treatment. In the 0.17 to 0.51 ionic strengths
of Na>SOxq, mean total dry mass of H. paradoxus
was 2- to 12-fold greater than H. annuus. Both
species produced less dry mass in NaCl treat-
ments; however, H. annuus dry mass was reduced
most by NaCl. Total dry mass of H. paradoxus
was 7-fold greater than H. annuus in the 0.09
NaCl treatment. This difference between species
in NaCl increased at the 0.17 and 0.34 ionic
strengths of NaCl where dry mass of dH.
paradoxus was 15-fold greater than H. annuus.
DISCUSSION
Salt tolerance is the ability of a species to grow
and adjust to the presence of a specific ion (ionic
effect) or to adapt to the general effects of low
water potentials (osmotic effect) (Ungar 1991). In
this study, a potential ionic effect and an osmotic
effect were investigated using NaCl and Na,SO,4
salts at increasing ionic strengths. Both salts are
found in HA. paradoxus habitats at various
concentrations (Boghici 1997; Van Auken and
Bush 1998; Lexer et al. 2003). To differentiate
between the effects of the two salts and the Cl™
and SO,” anions, equal ionic strengths were used
in the separate salt treatments (Barrow 1966).
Due to greater salt tolerance of H. annuus in
NaCl and mixed salt environments compared to
H. petiolaris (both purported parental species)
(Ashraf and Tufail 1995; Welch and Rieseberg
2002; Bush and Van Auken 2004; Karrenberg
et al. 2006; DiCaterina et al. 2007), H. annuus was
used as a comparative species for salt effects on
growth of H. paradoxus.
As in previous studies, this investigation
demonstrated that H. paradoxus was more salt
tolerant than one of its parental species, H.
annuus, and even produced slightly more dry
mass in low saline soils compared to H. annuus
(Figs. 1b, 2a, and 2b). Helianthus paradoxus
produced 70% more dry mass than H. annuus
over all treatments (data not shown) and
consistently produced more dry mass in elevated
salinity soils (Fig. 2a, b), especially in the sulfate
treatments. Results of this greenhouse study are
consistent with observations that H. paradoxus
has greater fitness in saline conditions than H.
2011]
nad
°
a) H. paradoxus
N
n
N
°
Total (Shoot + Root) dry mass (gP°)
Oo on
lonic Strength
mmm Na2so4 C— NaCl
b) H. annuus
No
oi So
~
ra)
=
So
Total (Shoot + Root) dry mass (gP°)
oO —_
an an
=
o
lonic strength
Fic. 2. Three-way interaction plots of total dry mass
(bars) as the response variable for (a) Helianthus
paradoxus and (b) Helianthus annuus. Salts were
Na 2SO, (black bar) and NaCl (gray bar) at ionic
strengths of 0.00—0.51. A three-way ANOVA deter-
mined there were significant differences (P < 0.0001)
between salt treatments and ionic strengths for each
species. Different letters indicate significant differences
between means (Duncan’s Multiple Range Test) within
a species. Lines above the bars represent + one SD
(standard deviation).
annuus (Welch and Rieseberg 2002; Bush and
Van Auken 2004; Karrenberg et al. 2006), and
that H. annuus is not expected to be found in
areas with elevated soil salinity where H. para-
doxus is able to grow and outcompete H. annuus
and probably other non-halophytic species (Ab-
bot 2003; Van Auken and Bush 2006).
Both Helianthus spp. exhibited a specific ionic
growth inhibition at elevated levels of Cl,
compared to SQ,’ , and differences in salt
tolerance between species were evident. For both
species, NaCl caused plant mortality at ionic
strengths of 0.34 and 0.51, yet H. paradoxus was
MENDEZ AND VAN AUKEN: SALT TOLERANCE OF TWO HELIANTHUS SPECIES 149
more tolerant than H. annuus to low to moderate
NaCl concentrations (Fig. 2). Greater dry mass
production and survivorship of H. paradoxus
plants compared to H. annuus in NaCl treatments
was also observed by Welch and Rieseberg (2002)
in corresponding NaCl treatments (100 and
200 mmol LL! [~6 and 12 g kg '] NaCl). Plant
survivorship of H. annuus indicated that low
Na»SO, levels seem to be less inhibitive than
NaCl but significant dry mass reduction was still
observed above 0.09 ionic strength treatments.
Dry mass of H. paradoxus, on the other hand,
was enhanced by low to moderate levels of
Na>»SO,4 with 38 to 72% greater dry mass, relative
to the no-salt treatment. This corresponds to field
observations where H. paradoxus was most
abundant in soil salinities (mainly Na, K, Ca,
and Mg chlorides with less SO4) ranging from 5
to 12 g kg '' in the Diamond-Y Spring Preserve
(Boghici 1997; Van Auken and Bush 1998; Bush
2006b; Grunstra and Van Auken 2007a, b).
Although previous studies did not differentiate
between the effects of NaCl and Na>,SO, (Welch
and Rieseberg 2002; Bush and Van Auken 2004;
Karrenberg et al. 2006; Van Auken and Bush
2006), those studies support the salt tolerance of
H. paradoxus to low concentrations of NaCl.
Results consistently demonstrated that biomass
of H. annuus was statistically reduced by NaCl
alone (Welch and Rieseberg 2002) or by low
levels of mixed salts (Na‘*, Cl, and SO,”
included; Bush and Van Auken 2004; Karrenberg
et al. 2006), while H. paradoxus demonstrated
lower productivity in NaCl alone and greater
growth and productivity in the presence of SO4*-
as observed in the present study. Greater salt
tolerance of H. paradoxus to NaCl, as compared
to H. annuus, has been attributed to significantly
greater fitness along with Na* accumulation, leaf
succulence, and water use efficiency (Welch and
Rieseberg 2002). Because the salts were at equal
ionic strengths and Na* seems to serve as an
osmoticum for H. paradoxus (Welch and Riese-
berg 2002), it can be assumed that Cl is causing
reduced productivity in comparison to SO,’ .
Molar concentrations of Cl” in the soil water of
the Diamond-Y Spring Preserve are approxi-
mately 1.5 times that of molar concentrations of
SO, ; therefore, Cl- has the potential of inhib-
iting growth of H. paradoxus in its salt marsh
habitat (Boghici 1997). However, these data in
conjunction with previous salt studies (Welch and
Rieseberg 2002; Bush and Van Auken 2004;
Karrenberg et al. 2006) indicate that the presence
of soil sulfate may have played an important
role in the selection for and adaptation of
H. paradoxus to the Na-Cl -SO,°~ rich environ-
ments. Further, poor tolerance to chloride and
sulfate by H. annuus has limited its establishment
and therefore, fitness in the H. paradoxus salt
marsh habitat.
150
It should be noted that in order to maintain
equal osmotic potentials between treatments,
molar concentrations of Na* in NaCl treatments
were 3.0 times that of Na»,SO, treatments.
Nevertheless, the molar equivalents of Na™ at
0.09 and 0.17 ionic strengths were between that of
the 0.34 and 0.51 ionic strengths of NasSO4
where dry mass production of H. paradoxus was
not inhibited. Since an inert osmotic medium
such as polyethylene glycol was not examined, an
osmotic effect in combination with an ionic effect
cannot be excluded (Katembe et al. 1998; Munns
and Tester 2008).
In the limited studies comparing phytotoxicity
of both salts, greater toxicity to NaCl compared
to Na,SO,4 has been demonstrated for other salt
tolerant glycophytes and halophytes (Manchanda
et al. 1982: Curtin et al. 1993; Franklin and
Zwiazek 2004; Pagter et al. 2009). Chloride is
more toxic to plants than sulfate possibly due to
synergistic phytotoxicity effects with Na‘, differ-
ential inhibition of enzyme activity, reduction in
plant productivity, and imbalance of nutritional
status (Greenway and Munns 1980; Manchanda
et al. 1982; Curtin et al. 1993; Wang et al. 1997;
Veira Dos Santos and Caldeira 1999; Franklin
and Zwiazek 2004). Ion toxicity is dependent on
whether the plant possesses adaptations to
tolerate the osmotic stress and to exclude and/
or compartmentalize the ion. Although not yet
documented, it is possible that H. paradoxus
accumulated Cl” along with Na* and may even be
more sensitive to Cl’) compared to SO,?” due to
poor compartmentalization into vacuoles (Green-
way and Munns 1980; Flowers et al. 1986; Munns
1993; Rajakaruna et al. 2003). Chloride may be
considered more toxic sometimes because of poor
salt tolerance response and thus, high accumula-
tion of Cl” over Na’, or in this case SO,” . Toxic
cytoplasmic Cl” concentrations have not yet been
determined but are assumed to be equal to or
slightly lower than Nat (Flowers et al. 1986;
Greenway and Munns 1980; Munns and Tester
2008).
Sulfate may be required for salt tolerance in H.
paradoxus. Tissue ion concentrations were not
examined in this study, but previous work
suggested that (as in other halophytes) SO,’ ,
along with Na‘, may be an important vacuolar
osmoticum in plant tissue (Greenway and Munns
1980; Karrenberg et al. 2006; Johnston 2006).
Leaf sodium and sulfur concentrations were
shown to be inversely related to calcium,
magnesium, and potassium concentrations but
positively related to biomass and succulence in H.
paradoxus (Karrenberg et al. 2006). Interestingly,
several studies have found a correlation between
plants inhabiting waterlogged sulfate rich soils
and the presence of vacuole stored flavonoid
sulfates which may serve to detoxify excess
sulfates alone or in combination with sodium
MADRONO
[Vol. 58
(Harborne 1975; Tomas-Barberan et al. 1987;
Rajakaruna et al. 2003). A possible role of
sulphur in the salt tolerance of halophytes also
includes the production of methylated sulfonium
compounds that accumulate in the cytosol as
osmotically compatible organic solutes for com-
partmentalization of Na* and Cl” in vacuoles.
These organic solutes also serve to detoxify
sulfides in salt marsh plants (Flowers and Colmer
2008). Flavonoid sulfates or methylated sulfoni-
um compounds have not been measured in tissues
of H. paradoxus; however, NasSO, tolerance of
this species together with Na* and SO,?~ accu-
mulation suggests a possible detoxification mech-
anism not yet identified.
Ecological isolation of H. paradoxus within the
inland salt marshes of west Texas and New
Mexico may in part depend on the elevated levels
of soil salinity found in soils of these habitats
(Boghici 1997; McDonald 1999; Van Auken and
Bush 1998, 2006; Abbott 2003). Halophytes are
limited to saline environments because of an
advantageous adaptation to excess salts and a
reduction of competitive ability in non-saline
environments (Ungar 1991). Distribution of ZH.
paradoxus plants appears to be dependent on low
to moderate soil salinity levels at the Diamond-Y
Spring Preserve where Na*, Cl” and SO,’ are the
prevalent salts from groundwater discharge,
while Ca**, Mg**, and HCO3° are secondary ions
(Boghici 1997; Van Auken and Bush 1998; Bush
2006b; Grunstra and Van Auken 2007a, b). In the
present study, H. paradoxus outperformed H.
annuus in both soil salt treatments, partially
explaining the inability of H. annuus to survive
the salt marsh habitat of H. paradoxus (Lexer
et al. 2003; Bush and Van Auken 2004). The
presence of salts excluded H. annuus from
colonizing the salt marsh (Abbott 2003; Lexer
et al. 2003), while the salt tolerance of JH.
paradoxus to both NaCl and NazSOy, has allowed
this species to establish in the Diamond-Y Spring
Preserve and other salt marsh environments in
west Texas and New Mexico. Further, tolerance
to high concentrations of NasSO4, as demon-
strated by significantly greater productivity rela-
tive to non-saline conditions, suggests that H.
paradoxus may experience a physiological stress
response without elevated Na»SO,, which is
necessary for optimum growth and _ perhaps
necessary for salt tolerance (Munns and Tester
2008). The establishment of H. paradoxus in the
Diamond-Y Spring Preserve has been promoted
by the selection for transgressive phenotypes
promoting salt tolerance (sodium exclusion,
calcium uptake, and leaf succulence), as demon-
strated by H. paradoxus, along with the presence
of specific ions (Na*, Ca**, and SO,’~) in the salt
marsh habitat (Abbott 2003; Lexer et al. 2003;
Karrenberg et al. 2006). In addition, both
parental species are poor competitors in field-like
2011]
elevated soil salinity conditions, while H. para-
doxus iS a poor competitor against H. annuus in
nonsaline conditions (Bush and Van Auken
2004).
The potential for hybridization is still present
and environmental conditions isolating hybrid
Helianthus spp. from parental genotypes appears
to have been an important factor in their adaptive
evolution of greater fitness in their respective
habitats (Lexer et al. 2003; Whitney et al. 2010;
Donovan et al. 2010). Populations of H. annuus
are found in disturbed pockets of isolated deep,
nonsaline soil. Helianthus petiolaris, the other
reported parent species to H. paradoxus is also
found in some isolated, low saline, sandy soils.
However, environmental factors such as the soil
conditions required for the isolation and survival
of some of the H. annuus X H. petiolaris hybrids
was and continues to be the saline soils sur-
rounding the isolated springs in this area of west
Texas and New Mexico (Abbott 2003; Lexer
et al. 2003). The unique salt tolerance of H.
paradoxus compared to its parental species along
with establishment in the Na‘t-Cl -SO,° domi-
nated salt marshes will continue to promote the
ecological isolation of H. paradoxus.
ACKNOWLEDGMENTS
Some support for this project was provided by the
Nature Conservancy of Texas, the University of Texas
at San Antonio, the Texas Department of Parks and
Wildlife, the U.S. Fish and Wildlife Service, and Dr.
Loyce and Mr. William Collenback. We especially
thank Mr. John Karges of the Nature Conservancy of
Texas for help and advice during various stages of this
project.
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MADRONO, Vol. 58, No. 3, pp. 153-184, 2011
THE DIVERSITY AND BIOGEOGRAPHY OF THE ALPINE FLORA OF THE
SIERRA NEVADA, CALIFORNIA
PHILIP W. RUNDEL
Department of Ecology and Evolutionary Biology and the Institute of the Environment and
Sustainability, University of California, Los Angeles, CA 90095
rundel@biology.ucla.edu
ABSTRACT
The alpine zone of the Sierra Nevada of California, defined as non-forested areas at or above
3500 m, includes 385 species (409 taxa) of native vascular plants. Were the alpine boundary defined as
at or above 3300 m, the alpine flora would grow to 536 species (570 taxa). There are 97 species that
reach elevations of 4000 m and 27 species that reach to 4200 m. Over half of the alpine species occur in
just six families, led by the Asteraceae (55 species, 59 taxa), Poaceae (39 species, 47 taxa), Brassicaceae
(34 species), and Cyperaceae (31 species). The largest genus present is Carex with 29 species, and 18
more species would be added by lowering the alpine boundary to 3300 m. Next in size are Draba (14
species) and Lupinus (11 species, 16 taxa). Life forms of the flora are heavily dominated by broad-
leaved erect perennials (50%), followed in importance by graminoid perennials (21%) and mats and
cushions (11%). Annuals and woody shrubs each account for about 6% of the flora. Only nine species
are obligate alpine taxa with a range restricted to elevations of 3500 m or above. An additional 67
species (17% of the flora) occur in both subalpine and alpine habitats but not lower. More than a
quarter of the alpine species have elevational ranges that extend as low as foothill habitats defined as
occurring below 1200 m. In terms of biogeographic affinities, the broad relationships of the flora
include the cordillera of western North America (35%), Intermountain region of the Great Basin
(20%), Sierra/Cascade axis (16%), and widespread distributions (14%). There are 36 species in the
alpine flora endemic to the Sierra Nevada, and another 31 species that are Californian endemics.
Key Words: Alpine, arctic-alpine flora, cushion plant, Sierra Nevada, treeline.
How large and diverse is the alpine flora of the
Sierra Nevada in California and what are its
biogeographic relationships? There has been a
long history of floristic and ecological studies of
the alpine region of the Sierra Nevada addressing
this and related issues, but a clear answer to the
question has not been achieved. Unlike the
majority of alpine regions in the northern
hemisphere that share extensive elements of a
circumboreal arctic-alpine flora, the Sierra Ne-
vada has developed a unique component to its
alpine flora under the influence of mediterranean-
climate conditions with relatively dry summers
added to other alpine environmental factors of
stress. Also significant in the evolution of this
alpine flora has been the relative isolation of the
range from other high mountain floras of the
western United States. Moreover, the Sierra
Nevada possesses a complex mosaic of substrate,
glacial history, and soil variation superimposed
over broad patterns of climatic and topographic
heterogeneity.
Interest in the alpine flora dates back to early
descriptions by Coville (1893) and Harshberger
(1911), who recognized the distinctiveness of the
Sierran alpine flora. Hall and Grinnell (1919)
gave a very brief description of the alpine zone in
the context of a broader description of California
life zones, and provided a short list of character-
istic species. More significant, however, have
been five studies over the past 80 years that have
provided an analysis of the diversity and floristic
affinities of the high elevation flora of the Sierra
Nevada. The earliest of these was the work of
Smiley (1921), whose definition of the boreal
region of the Sierra Nevada comprised the
Canadian, Hudsonian, and Arctic-Alpine zones
as characterized in the Merriam system of life
zones (Daubenmire 1938). These life zones
roughly correspond to the upper montane,
subalpine and alpine zones under current con-
cepts (Fites-Kaufman et al. 2007). Smiley’s work
was followed by the classic investigation of
Sharsmith (1940), and in more recent decades
with analyses by Chabot and Billings (1972),
Major and Taylor (1977), and Stebbins (1982).
Early speculations on the origin of the Sierran
alpine flora were contributed by Went (1948,
1953). Beyond these broad floristic surveys, there
have been numerous studies of the floristics and
vegetation of regional areas of subalpine and
alpine vegetation in the Sierra Nevada (Howell
1944, 1951; Klikoff 1965; Pemble 1970; Taylor
1976b; Major and Taylor 1977; Tatum 1979;
Benedict and Major 1982; Burke 1982; Ratliff
1982: Benedict 1983; Porter 1983: Constantine-
Shull 2000; Sawyer and Keeler-Wolf 2007).
None of the existing literature has provided a
satisfactory answer to the fundamental question.
How many species are there in the alpine flora of
154 MADRONO
the Sierra Nevada? The objective of this paper is
to present a broad overview of the alpine flora of
the Sierra Nevada by providing a detailed and
updated analysis of the floristic richness, ecolog-
ical diversity, and biogeographic relationships of
the species present within the alpine zone. The
paper takes a conservative approach following
Sharsmith (1940) by defining the alpine zone
using a lower elevational limit of 3500 m.
Climatic treeline typically occurs from 3300-—
3500 m in the central and southern Sierra Nevada
where the great majority of alpine habitat in
California is located (Fig. 1). Although the
northern Sierra Nevada lacks high elevation
areas, it nevertheless has a good representation
of alpine species that reach above 3500 m in the
central or southern areas of the range. To provide
a broader context examining the significance of
elevation in the definition of the alpine zone,
analyses have been made for all species occurring
at or above 3300 m within California.
Beyond an intrinsic interest in the evolution of
biodiversity of alpine biota, there are very
significant reasons to support Sierran alpine
studies that can serve as baseline studies for
important early warning systems of potential
environmental impacts of climate change. Cli-
mate change models for California suggest that
there will be significant effects on environmental
conditions of subalpine and alpine habitats of the
Sierra Nevada (Hayoe et al. 2004; Shafer et al.
2001), and historical data on vertebrate distribu-
tion demonstrates that these effects are ongoing
today in influencing the distributions of verte-
brate species (Moritz et al. 2008; Tingley et al.
2009).
MATERIALS AND METHODS
The Jepson Manual, 2nd Edition (Baldwin et al.
2012) was used to identify California species with
an elevational distribution up to or above 3300 m
within the state, and which occurred in the Sierra
Nevada. This reference is the sole source and
reference for binomials used in this article.
Species at or above 3500 m in California were
considered to comprise the alpine flora. The
upper and lower elevational ranges of each of
these species were recorded, along with their
biogeographic distribution and occurrence within
the geographic regions of California (Hickman
1993). These geographic regions included records
of species presence in the montane and higher
elevations of the northern, central, and southern
subregions of the Sierra Nevada, as well as the
high Cascade Range, the Klamath/Siskiyou
mountains, Transverse and Peninsular ranges of
southern California, and ranges east of the Sierra
Nevada including the Sweetwater and White-
Inyo mountains (Fig. 1). The elevational limits
and geographical ranges listed in Baldwin et al.
[Vol. 58
(2012) are specimen-based records and thus
considered reliable. Only native species were
included in this analysis, however, alien species
recorded as occurring at high elevations in the
Sierra Nevada are very few. Poa pratensis L. is
recorded as reaching 3500 m and Taraxacum
officinale F. H. Wigg. reaches 3300 m.
Each taxon occurring at elevations of 3300 m
or above was categorized into a series of growth
forms, based on a modified scheme of Raunkiaer
(1934). These categories were broad-leaved her-
baceous perennials (tussocks, rosettes, and bien-
nials), graminoid perennials, mats and cushion
plants, geophytes, aquatics, annuals, subshrubs,
woody shrubs (deciduous and evergreen), and
trees.
The lower elevational limit of occurrence in
California was used to separate alpine species into
categories of lowest elevational zone of occur-
rence on the following basis: 1) foothill habitats of
woodland and chaparral— <1199 m; 2) lower
montane habitats dominated by mixed conifer
and yellow pine forests— 1200-1999 m; 3) upper
montane habitats of red fir and lodgepole pine
forests— 2000-2699 m; 4) subalpine habitats of
open conifer stands near treeline— 2700-3499 m;
and 5) alpine habitats— >3500 m. Because
elevational boundaries of these major vegetation
zones change with latitude, as well as locally with
slope exposure, these elevational ranges represent
averaged boundaries across the west slope of the
central and southern Sierra Nevada.
The biogeographic range of each alpine
species was classified into one of six categories.
These were: 1) widespread species present in
many habitats or regions across North America
and/or throughout the world; 2) cordilleran
species broadly distributed in mountain regions
of the western United States; 3) Sierra/Cascade
species with a Pacific Northwest distribution; 4)
Intermountain Region species present in the
Great Basin; 5) species endemic to the Sierra
Nevada; and 6) species endemic to California,
broadly defined to include adjacent Great Basin
ranges extending into western Nevada (_.e.,
Sweetwater, Wassuk, and White-Inyo moun-
tains) and southern Oregon. Dividing species
into such simple biogeographic categories is
inherently arbitrary for some species, and
expanded field studies in the future may well
change these classifications and alter the list of
Sierran endemics based on new records or
taxonomic revisions.
RESULTS
The Geography of California Alpine Habitats
The elevational contour interval of 3500 m is
highly irregular in the Sierra Nevada, as it defines
a relatively continuous area along the crest of the
2011] RUNDEL: SIERRA NEVADA ALPINE FLORA iS)
FiG. 1. Topographic map of California showing the major areas of mountain systems. W/I = White-Inyo
Mountains, T = Tehachapi Mountains, SG = San Gabriel Mountains, SB = San Bernardino Mountains, and P =
Peninsular Ranges.
central and southern crest of the range extending
from northern Tuolumne and Mono counties in
the area of Leavitt Peak (3527 m) near Sonoran
Pass and south across Yosemite National Park
where the highest peak is Mount Lyell (3999 m;
Fig. 2). Further south this belt of alpine habitat
continues into Kings Canyon and Sequoia
National parks where there are extensive areas
of alpine habitat with ten peaks that reach above
4000 m. Mount Whitney at 4421 m is the highest
156 MADRONO [Vol. 58
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FiG. 2. Geographic distribution of high elevations of the Sierra Nevada and adjacent ranges. The solid line marks
a rough position of the 3000 m contour. Adapted from Hovanitz (1940).
point in the contiguous United States. The end of the continuous chain of glaciated peaks in
southern limit of this extensive and virtually the Sierra Nevada. To the south, the alpine zone
contiguous alpine zone occurs at Cirque Peak reappears on Olancha Peak (3698 m; Fig. 2), the
(3932 m) in Sequoia National Park at the southern southernmost glaciated summit of the range lying
2011]
on the Tulare-Inyo county line (Howell 1951;
Tatum 1979). Two major breaks with subalpine
elevations but not true alpine provide the only
major discontinuity for this primary Sierran alpine
region. These are Tioga Pass in Yosemite National
Park (3031 m) and Mammoth Pass (Muinaret
Summit) (2824 m), which is the route for
California Highway 203.
The crest of the Sierra Nevada lies at lower
elevations to the north of the Tioga Pass area,
with only scattered areas of good alpine habitat
present. A notable ecological change occurs
north of this pass where volcanic substrates
replace the granites of the central and southern
Sierra Nevada. Fragmented communities of
alpine species are present at elevations below
3500 m, particularly along exposed ridgelines
and on steep north-facing slopes that were
heavily glaciated. However, there are no eleva-
tions in the northern Sierra Nevada that reach
the 3500 m limit used here to define the alpine
zone. Alpine habitats are weakly developed in
Alpine Co. (with Sonora Peak reaching 3493 m)
and eastern El Dorado Co. (with Freel Peak
reaching 3318 m), extending to their northern
limit on Mount Rose (3285 m) in the Carson
Range east of Lake Tahoe in Nevada (Fig. 2).
Nevertheless, there are scattered communities of
alpine-like habitat existing at upper elevations
in the northern Sierra Nevada, positioned above
local edaphically-controlled treelines, and the
alpine flora is well represented (Smiley 1915).
Despite the floristic relationships of high eleva-
tion Sierran species all along the range, Stebbins
and Major (1965) linked the Sierra Nevada
north of Lake Tahoe with the Cascade Range
rather than with the region of the central and
southern Sierra Nevada on the basis of the
dominance of volcanic substrates.
To the north of the Sierra Nevada, Mount
Shasta in the southern Cascade Range reaches an
elevation of 4322 m, while Lassen Peak is lower at
3187 m. The highest peaks in the Klamath
Mountains of northwestern California and
adjacent Oregon are Mount Eddy (2750 m) in
Siskiyou Co., Thompson Peak (2744 m) in
Trinity Co., and Mount Ashland (2296 m) in
Jackson Co., Oregon. These high peaks contain
areas with permanent or long-lasting snowfields
on north-facing slopes with associated alpine
species (Howell 1944; Major and Taylor 1977).
There are several high mountain ranges to the
east of the Sierra Nevada at the western margin
of the Great Basin. The Sweetwater Mountains,
located just 33 km east of the Sierra Nevada,
reach 3552 m on Mount Patterson (Hunter and
Johnson 1983). The Wassuk Range in west-
central Nevada lie 48 km east of the Sweetwater
Mountains and 88 km north of the White
Mountains, reaching 3427 m on Mount Grant
(Bell and Johnson 1980). The White Mountains
RUNDEL: SIERRA NEVADA ALPINE FLORA 157
have an extensive alpine area and reach to 4344 m
on White Mountain Peak, the third highest peak
in California (Rundel et al. 2008). To the south,
Mount Waucoba forms the high point at 3390 in
the Inyo Mountains. The Panamint Mountains
lying east of the White-Inyo Mountains reach a
maximum elevation of 3366 m on Telescope
Peak. Further south, the Spring Mountains in
southwestern Nevada divide the Pahrump Valley
and Amargosa River basins from the Las Vegas
Valley watershed and define part of the south-
western boundary of the Great Basin. The highest
point is Charleston Peak at 3633 m.
High elevations are also present in the Trans-
verse and Peninsular ranges of southern Califor-
nia (Fig. 1) where a subset of Sierran alpine
species 1s present in weakly developed alpine-like
communities (Hall 1902; Parish 1917; Horton
1960; Hanes 1976; Major and Taylor 1977;
Meyers 1978). Mount San Gorgonio in the San
Bernardino Mountains reaches 3506 m, while
other high points are Mount San Jacinto in the
San Jacinto Mountains at 3302 m and Mount
Baldy in the San Gabriel Mountains at 3068 m.
Alpine species are present in both xeric and mesic
habitats at high elevation, but alpine communi-
ties, defined as extended areas dominated by
assemblages of alpine species, are only poorly
developed.
The alpine zone of the Sierra Nevada experi-
ences mediterranean-type climate conditions that
differ significantly from those that characterize
the Rocky Mountains and most of the continental
alpine habitats of the world where summer
rainfall predominates. The fraction of annual
precipitation that falls as winter snow in the Sierra
Nevada is about 95% at upper treeline (Stephen-
son 1998). Deep snow packs and cool temperature
at higher elevations mean that snowmelt extends
into the spring, but the length and magnitude of
the summer drought period experienced by plants
is significant. Patterns of rainfall decline gradually
from north to south in the Sierra Nevada, and
summer drought decreases as elevation increases
because of both increased levels of precipitation
and cooler temperatures with lower evaporative
demand at higher elevations (Stephenson 1998;
Urban et al. 2000).
Winter mean monthly low temperatures are
moderate in the Sierra Nevada compared to the
Rocky Mountains, and soils only rarely freeze to
even moderate depth. While, the mean minimum
temperature above treeline is below freezing for ten
months of the year, with nighttime lows that
typically reach only —3 to —6°C, although extremes
can reach temperatures of —15°C or lower on the
high peaks. Nevertheless, these moderate low
temperatures as well as other limiting factors for
survival at high elevations sharply reduce the
diversity of species able to tolerate such conditions
(Korner 2003).
158 MADRONO
Number of species/taxa
3100 3300 3500
3700
[Vol. 58
» Taxa
WB Species
3900 4100 4300 4500
Elevation (m)
Fic. 3.
Floristic Richness
The alpine flora of the Sierra Nevada, defined
as species reaching 3500 m or more at their upper
limit of distribution, comprises 385 vascular plant
species. The species total includes 10 ferns and
fern relatives (2.6%), five conifers (1.3%), 85
monocots (22.1%), and 285 eudicots (74.0%).
With the inclusion of an additional 24 named
varieties and subspecies, the total number of
alpine taxa is 409.
Of course, the predetermined elevational
boundary has a very strong influence on the size
of the flora (Fig. 3). If the alpine flora were
defined as those species reaching to 3400 m, then
76 additional species would be added for a total
of 460 species (488 taxa). Were the limit defined
as 3300 m, there would be a flora of 536 species
(570 taxa), with the relative proportions of
monocots and eudicots virtually unchanged and
the addition of five ferns and one conifer.
There are 97 species (101 taxa) with an
elevational range that extends as high as 4000 m,
an elevation reached by only the highest Sierran
peaks (Fig. 3). This number declines to 27 species
that reach 4200 m in elevation. These 27 high
elevation species do not display dominance by a
few families as is the case with the full alpine flora
but are rather spread among 15 different families
(Appendix 1). Three species have been recorded
as reaching to 4400 m. These are Epilobium
anagallidifolium Lam. (Onagraceae), Saxifraga
hyperborea R. Br. (Saxifragaceae), and Erigeron
vagus Payson (Asteraceae). Additional taxa that
occur up to or above 4300 m are Erigeron
compositus Pursh (Asteraceae), Boechera lemmo-
nii (S. Watson) W. A. Weber (Brassicaceae),
Cerastium beeringianum Cham. & Schltdl. (Car-
yophyllaceae), Calyptridium umbellatum (Torr.)
Greene (Montiaceae), Festuca brachyphylla
Schult. & Schult. subsp. breviculmis Fred., Poa
keckii Soreng. and P. lettermannii Vasey (Poa-
Elevational distribution of species in the high mountain flora of the Sierra Nevada.
ceae), Phlox pulvinata (Wherry) Cronquist (Po-
lemoniaceae), Ranunculus eschscholtzii Schltdl.
var. oxynotus (A. Gray) Jeps. (Ranunculaceae),
and Potentilla pseudosericea Rydb. and Sorbus
californica Greene (Rosaceae).
There are six families that contribute 20 or
more taxa to the alpine flora. The largest of these
is the Asteraceae with 55 species (59 taxa),
followed in size by the Poaceae (39 species, 47
taxa), Brassicaceae (34 species), Cyperaceae (31
species), Rosaceae (21 species, 23 taxa), and
Fabaceae (18 species, 27 taxa). These six families
together comprise 52% of the alpine flora.
At the generic level, Carex stands out prom-
inently with 29 species in the alpine flora, with an
additional 18 species present at elevations be-
tween 3300 and 3500 m. Next in order of size are
Draba (Brassicaceae, 14 species), and Lupinus
(Fabaceae, 11 species, 16 taxa). There are 10
species of Boechera (Brassicaceae) and nine
species each of Epilobium (Onagraceae), Eriogo-
num (Polygonaceae), and Potentilla (Rosaceae).
There are three genera with eight species—
Penstemon (Plantaginaceae), Poa (Poaceae), and
Salix (Salicaceae).
Growth Form Distribution
Herbaceous perennial growth forms, broadly
defined, comprise the great majority of taxa
reaching to or above 3500 m in the Sierra |
Nevada. This growth form with all of its |
subgroups includes 343 taxa, or 84% of the 409 |
taxa that comprise the flora. These herbaceous |
perennials can be broken down into subgroups of
erect herbaceous perennials, perennial grami-
noids, prostrate mats and cushion plants, bienni- |
als, and geophytes. The largest numbers of |
herbaceous perennials form the category of erect ,
herbaceous perennials, with 186 species (206 taxa;
Fig. 4). The most important families for the erect ,
herbaceous perennials are the Asteraceae, Brassi- |
2011]
200
160
120
80
Number of taxa
erect mat /
annual perennial
cushion graminoid geophyte
RUNDEL: SIERRA NEVADA ALPINE FLORA lest)
aquatic shrub suburb tree
Fic. 4. Growth form distribution of alpine taxa in the alpine flora of the Sierra Nevada.
caceae, Fabaceae, Rosaceae, Polygonaceae, and
Onagraceae. Among the erect herbaceous peren-
nials are four species that are reported to have the
potential to survive as facultative annuals.
Although these have not been studies in detail,
it is expected that these species have biennial or
short-lived perennial life histories in the alpine
zone, and they are included here in the totals for
erect herbaceous perennials. Three of these are
members of the Brassicaceae, each representing a
different genus. Among these facultative annuals,
only Androsace septentrionalis L. (Primulaceae)
with an elevation range of 2700-3600 m can be
considered as a subalpine and alpine specialist.
There are several additional groups classified
broadly as erect herbaceous perennials. The 10
species of ferns and fern relatives included here
within the alpine flora represent four families
(Pteridaceae, Ophioglossaceae, Woodsiaceae, and
Selaginellaceae). Four of these species reach
elevation at or above 4000 m—Botrychium lineare
W. H. Wagner, B. paradoxum W. H. Wagner,
Cystopteris fragilis (L.) Bernh., and Selaginella
watsonii Underw. Lowering the characterization
of the alpine zone lower limit to 3300 m would
add five additional fern species (Appendix 2).
Also classified as erect perennials are seven
species of hemiparasites, all members of the
Orobanchaceae, with four species (five taxa) of
Castilleja and three species of Pedicularis. Six
more species from this family would be added by
lowering the alpine boundary to 3300 m, includ-
ing five more species of Castilleja.
Next in diversity among the herbaceous
perennials is the subgroup of graminoids (Cyper-
aceae, Juncaceae, Juncaginaceae, and Poaceae)
with 83 species (85 taxa, Fig. 4). All of the
members of these four families within the alpine
flora are perennials, with Agrostis, Bromus,
Carex, Elymus, Juncus, Luzula, Poa, and Stipa
forming genera with five or more taxa (Appendix
1). These perennial graminoids include one
species of Cy grass, Muhlenbergia richardsonis
Rydb. (Sage and Sage 2002). Two other C,
members of this genus, the perennial M. montana
Hitche. and the annual M. filiformis Rydb., just
miss inclusion, reaching to elevations of 3420 m
and 3350 m, respectively. Lowering the alpine
boundary to 3300 m would add significantly to
the diversity of graminoid perennials, with 43
additional taxa present (Appendix 2).
Prostrate mats and cushion forms of growth
are common in some of the herbaceous perenni-
als of the Sierran alpine flora (Fig. 4). These
species are low in stature and form a heteroge-
neous group that shares the characteristic of a
prostrate growth form with either a central
taproot or multiple points of rooting through
layering. Mats and cushions often form an
ecologically significant component of plant cover
on exposed ridges and fellfield. There are 46
species classified here as mats or cushions, with
19 of these high subalpine and alpine specialists
not occurring below 2700 m elevation The growth
form characteristics of mats and cushions may be
genetic in some cases but in others is environ-
mentally induced, with mat forms of growth only
occurring at higher elevations (personal observa-
tions). Alpine mat and cushion species are well
represented in the Asteraceae with 13 species
(notably taxa of Antennaria and Erigeron),
Polygonaceae (Eriogonum) with eight species,
160 MADRONO
50
x +
os 30 O
- ¢
& OC
20
E A
oh
2 AA
10
0
[Vol. 58
@ annuals
Cishrubs
a subshrubs
>
® cushion/mat
3100 3300 3500 3700 3900 4100 4300 4500
Elevation (m)
FIG. 5.
Sierra Nevada.
Brassicaceae (Draba and Anelsonia) with s1x
species, and Fabaceae (Astragalus, Lupinus,
Oxytropis, and Trifolium) with five species. Also
notable in their ecological dominance are mats of
Caryophyllaceae (Cerastium, Eremogone, Min-
uartia) and Polemoniaceae (Ph/ox).
The alpine flora includes just four species of
geophytes, which represent the Alliaceae (A//ium
obtusum Lemmon var. obtusum), Liliaceae (Ca-
lochortus leichtlinii Hook. f.), Melanthiaceae
(Veratrum californicum Durand var. californi-
cum), and Themidaceae (Triteleia dudleyi Hoo-
ver). The highest elevation species among these 1s
C. leichtlinii, which reaches up to 4000 m. None
of these geophytes can be considered to be high
elevation specialists as all reach lower elevational
limits of 1200—1500 m in California. There are six
species of geophytes that just miss reaching the
lower alpine limit, as defined here, but occur at or
above 3300 m (Appendix 2). These include A//ium
validum S. Watson (Alliaceae), /ris missouriensis
Nutt. (Iridaceae), Lilium kelleyanum Lemmon
(Liliaceae) and three Orchidaceae (Platanthera
dilatata (Pursh) Lindl. ex L. C. Beck var.
leucostachys (Lind.) Luer, P. sparsiflora Schlitr.,
and Spiranthes romanzoffiana Cham.
Only a single species of aquatic plant, Calli-
triche palustris L. (Plantaginaceae), reaches the
alpine zone of the Sierra Nevada. This is perhaps
not surprising given the relatively small area of
oligotrophic lakes that are present above 3500 m.
Potamogeton robbinsii Oakes (Potamogetona-
ceae) and Limosella acaulis Sessé & Moc.
Elevational distribution of taxa of annuals, shrubs, subshrubs, and mats/cushions in the alpine flora of the
(Scrophulariaceaeae) have a range that extends
as high as 3300 m, and a number of aquatic
species including /soetes (Isoetaceae) reach eleva-
tions of 3000 m.
Plants with an annual life history comprise a
small but significant component of the alpine
flora of the Sierra Nevada, with 24 species (26
taxa) reaching to elevations of 3500 m (Fig. 4).
The annual species occurring at the highest
elevation is Gayophytum decipiens F. H. Lewis
& Szweyk. (Onagraceae), which ranges up to
4200 m. Five other species of annuals reach
4000 m in elevation—Gentianopsis holopetala (A.
Gray) Iltis (Gentianaceae), Phacelia hastata
Douglas ex Lehm. subsp. compacta (Brand)
Heckard (Boraginaceae), Mimulus suksdorfii A.
Gray (Phrymaceae), Gayophytum racemosum
Torr. & A. Gray (Onagraceae), and Collinsia
torreyi A. Gray var. wrightii (S. Watson) I. M.
Johnst. (Plantaginaceae). The number of annual
species present increases sharply below the limit
set here for inclusion in the alpine flora. Including
the above taxa, there are 33 annual species (36
taxa) with a range reaching to or above 3400 m
and 38 species (41 taxa) occurring at or above
3300 m (Fig. 5).
Most of the annuals reaching into the alpine
zone are species with wide elevational ranges that
extend down to lower foothill habitats. Only 13
of the alpine annual species have ranges limited
to elevations at or above 1200 m, a distribution
that would indicate adaptation to montane and
higher elevation habitats. Five annual taxa can be
2011]
considered as subalpine and alpine specialists
having a lower elevation limit of 2700 m or above
and/or a median elevational range above 3000 m.
These species, none of which ranges as high as
4000 m or above, are Comastoma_ tenellum
(Rottb.) Toyok. (Gentianaceae), Cryptantha cir-
cumscissa (Hook. & Arn.) I. M. Johnst.var.
rosulata J. T. Howell (Boraginaceae), Strep-
tanthus gracilis Eastw. (Brassicaceae), and Lepto-
siphon oblanceolatus (Brand) J. M. Porter & L. A.
Johnson and Gymnosteris parvula A. Heller
(Polemoniaceae). Just missing this criteria, but
certainly also a high elevation specialist, is
Phacelia orogenes Brand (Boraginaceae). Four
of these six, with Comastoma tenellum and
Gymnosteris parvula as exceptions, are Sierra
Nevada endemics.
The most important family in contributing to
the annual flora of high elevations is the
Boraginaceae, with 11 species (12 taxa) repre-
senting five genera. Next in importance are the
Polemoniaceae with five species (comprising five
genera), and the Onagraceae with four species
(five taxa) representing just a single genus. There
are four genera that contribute three or more
species to the annual flora. These are Gayophy-
tum (Onagraceae, four species, five taxa), Phace-
lia (Boraginaceae, four species), Cryptantha
(Boraginaceae, three species, four taxa), and
Mimulus (Phrymaceae, three species).
Subshrubs, defined as semi-woody species that
maintain living perennial tissue in winter above
the ground surface, include 13 species occurring
at elevations of 3500 m or above (Fig. 4). The
Asteraceae contribute more than 60% of the
alpine flora of subshrubs, with eight species. Four
species of Ericameria (Asteraceae) and three
species each of Penstemon (Plantaginaceae), and
one Monardella (Lamiaceae) form subshrubs that
reach alpine elevations. Five species are consid-
ered to be subalpine and alpine specialists based
on a lower elevational limit of 2700 m or a mean
elevational range above 3000 m. Four of these are
members of the Asteraceae—Sphaeromeria cana
(D. C. Eaton) A. Heller, Ericameria parryi (A.
Gray) G. L. Nesom & G. I. Baird var.
monocephala (A. Nelson & P. B. Kenn.) G. L.
Nesom & G. I Baird, E. bloomer (A. gray) J. F.
Macbr., and Chrysothamnus viscidiflorus (Hook.)
Nutt. var. viscidiflorus. The two latter species
have very broad elevational occurrence from
800-4000 m.
There are 23 species of woody shrubs that
extend into the alpine zone of the Sierra Nevada
(Fig. 4). Just four families account for the
majority of the high elevation shrubs. The largest
of these is the Salicaceae (eight species of Salix),
followed by the Ericaceae (five species, each in a
different genus), Grossulariaceae (three species of
Ribes), and Rosaceae (three species, each in a
different genus). The highest elevation reached is
RUNDEL: SIERRA NEVADA ALPINE FLORA 16]
reported for Sorbus californica at 4300 m. How-
ever, this elevation record appears to not be
supported by specimen records in the Consortium
of California Herbaria (ucjeps.berkeley.edu/
consortium), and therefore needs confirmation.
There are six additional shrub species that reach
elevations of 4000 m—Salix orestera C. K.
Schneid., S. planifolia Pursh, S. petrophila Rydb.,
Gaultheria humifusa (Graham) Rydb., Holodiscus
discolor (Pursh) Maxim. var. microphyllus (Rydb)
Jeps., and Ribes montigenum McClatchie. Only
three shrub species can be considered as subal-
pine and alpine specialists based on a lower
elevational limit at or above 2700 m or median
range of occurrence above 3000 m. These are
Salix planifolia, S. brachycarpa Nutt. var. bra-
chycarpa, and S. nivalis Hook. Three more shrub
species just miss this definition of high elevation
specialist. Arctostaphylos uva-ursi (L.) Spreng.
has an elevational range of 2400-3300 m, while
Jamesia americana Torr. & A. Gray (Hydrangea-
ceae) and Ribes cereum Douglas var. inebrians
(Lindl.) C. L. Hitche. are alpine species that
extend down to lower elevations of 2070 m and
2100 m, respectively. Including the above species,
there are a total of 39 shrub species that occur at
elevations of 3300 m or above in the Sierra
Nevada. This group includes two more species of
Salix, one additional Ribes, five Ericaceae, three
Rosaceae, two species of Caprifoliaceae, and a
scattered diversity of species from other families
(Appendix 2).
Five species of coniferous trees in the Pinaceae
have scattered populations that extend well above
typical treeline elevation on favorable sites. The
treeline pines, Pinus albicaulis Engelm., P. flexilis
E. James and P. balfouriana Grev. & Balf., all
have local populations that reach as high as
3700 m in elevation in the Sierra Nevada, while P.
contorta Loudon subsp. murrayana (Grev. &
Balf.) Critchf. and Tsuga mertensiana (Bong.)
Carriere reach 3500 m. Just missing the elevation
of the alpine zone are scattered trees of Pinus
monticola Douglas ex D. Don that reach up to
3400 m.
Elevational Amplitude
Separating alpine taxa into categories of
elevational ranges over which they occur provides
some insight into their ecological amplitude and
thus a crude measure of potential niche breadth.
There are nine obligate alpine taxa in the Sierra
Nevada restricted in occurrence to elevations
at or above 3500 m. These are Boechera depaupe-
rata (A. Nelson & P. B. Kenn.) Windham &
Al-Shehbaz (Brassicaceae), Botrychium para-
doxum and B. tunux Stensvold & Farrar (Ophio-
glossaceae), Carex incurviformis Mack. (Cyper-
aceae), Draba sierra Sharsm. (Brassicaceae),
Eriogonum wrightii Torr. ex Benth. var. olanchense
162
140
120
100
80
60
Number of taxa
40
20
foothill lower montane
FIG. 6.
MADRONO
upper montane
[Vol. 58
subalpine alpine
Lower elevational zone of occurrence for taxa in the alpine flora of the Sierra Nevada. The elevational
limits of these zones are 0—1190 m for the foothill zone, 1200-1990 m for the lower montane zone, 2000—2699 m for
the upper montane zone, 2700-3490 m for the subalpine zone, and 3500 m and above for the alpine zone.
(J. T. Howell) Reveal (Polygonaceae), Minuartia
stricta (Sw.) Hiern (Caryophyllaceae), Phlox
dispersa Sharsm. (Polemoniaceae), and Poa let-
termanii (Poaceae).
There an additional 67 alpine species (70 taxa,
17.2% of all taxa) with ranges restricted to the
elevations of subalpine and alpine habitats at or
above 2700 m (Fig. 6). Examining the floristic
composition of all 76 species (79 taxa) with a
range restricted to subalpine and alpine eleva-
tions, just four families comprise more than half
of these. These are the Asteraceae (11 species),
Brassicaceae (nine species), Rosaceae (eight
species, nine taxa), and Poaceae (eight species).
Three additional species would be added to the
obligate subalpine and alpine flora if the eleva-
tional limit were reduced to 3300 m. These are
Astragalus ravenii Barneby (Fabaceae), Carex
tiogana D. W. Taylor & J. D. Mastrog. (Cyper-
aceae), and Chaenactis douglasii (Hook.) Hook.
& Arn. var. alpina A. Gray (Asteraceae).
Looking at the level of all alpine taxa, 22.4%
have a lower elevational limit in the upper
montane zone (2000-2699 m) and a further
31.0% have a lower limit in the lower montane
zone (1200-1999 m). Finally 27.5% of the alpine
taxa have a broad elevational amplitude of
occurrence extending upward from the foothill
zone below 1200 m up into the alpine (Fig. 6).
Plotting the elevational amplitude of all of the
alpine taxa shows a peak at about 2300 m, with
relatively fewer species exhibiting very broad or
very narrow elevational amplitudes (Fig. 7).
Nevertheless, there are many taxa with surprising
broad ranges of elevational occurrence. There are
77 species that have an elevational amplitude of
3000 m or more, and six species that have 4000 m
or more of amplitude in California. These latter
species, each in a different family, are Callitriche
palustris (Plantaginaceae), Calyptridium umbella-
tum (Montiaceae), Cystopteris fragilis (Woodsia-
ceae), Draba cana Rydb. (Brassicaceae), Epilo-
bium ciliatum Raf. subsp. ciliatum (Onagraceae),
and Erysimum capitatum (Hook.) Greene var.
capitatum (Brassicaceae). Were the elevational
definition of the alpine zone lowered to 3300 m, a
large number of species with broad elevational
amplitudes would be added to the flora. There
are 42 species in this group of added taxa that
have 3000 m or more of elevational amplitude in
their range of occurrence.
Biogeography and Endemism
Within the Sierra Nevada itself, the distribu-
tions of the high elevation flora are relatively well
spread between the northern, central and south-
ern subregions of the Sierra Nevada. Assessing
species reaching an elevational boundary of
3300 m, 70% of the 567 taxa occur in all three
subregions. The northern subregion has 76% of
the alpine flora present, while the central and
southern Sierra Nevada have 90% and 88% of the
alpine flora present, respectively. A number of
alpine species have their southern limit of
distribution in the central Sierra Nevada. These
include Carex whitneyi Olney (Cyperaceae),
Podistera nevadensis (A. Gray) S. Watson (Apia-
ceae), Claytonia megarhiza (A. Gray) S. Waston
(Montiaceae), Thalictrum alpinum L. (Ranuncu-
laceae), Galium grayanum Ehrend. var. grayanum
(Rubiaceae), and Salix nivalis (Salicaceae).
|
2011] RUNDEL: SIERRA NEVADA ALPINE FLORA 163
60
50
40
fae)
&
xe)
5 30
Q
=
=)
Zz
20
10
0
0 400 800 1200 1600 2100 2400 2800 3200 3600 4000 4400
Elevational amplitude (m)
FIG. 7. Elevational amplitude of alpine taxa in the Sierra Nevada. These values are based on the range expressed
from upper and lower limits of elevational distribution in California as presented in Baldwin et al. (2012).
The broader biogeographic relationships of the
alpine flora at or above 3500 m indicate its
diverse origins (Fig. 8). Widespread species dis-
tributed across North America and beyond as
boreal or arctic-alpine taxa comprise 13.6% of
the flora (Table 1). The largest group of taxa
(34.3%) shows patterns of distribution as cordil-
leran species widespread in mountain regions of
the western United States. Next in importance
160
140
120
100
Number of taxa
cord S-C
wide
FIG. 8.
are taxa with a range in the Intermountain
Region of the Great Basin, comprising 20.5% of
taxa. A group consisting of 15.8% of the taxa has
ranges extending along the Sierra Nevada axis to
the Cascade Range and often on to the Pacific
Northwest.
The alpine flora of the Sierra Nevada includes
36 endemic taxa restricted in their distribution to
the Sierra Nevada (Table 2). These endemic taxa
end end-cal int
Biogeographic relationships of the alpine flora of the Sierra Nevada. WIDE = widespread taxa present in
many habitats or regions across North America and/or throughout the world; CORD =
cordilleran taxa
widespread in mountain regions of the western United States; S-C = Sierra/Cascade taxa with a Pacific Northwest
distribution; END = taxa endemic to the Sierra Nevada; and END-CAL = taxa endemic to California, as broadly
defined; INT = intermountain taxa present in the Great Basin. See text for discussion.
164
TABLE 1.
SIERRA NEVADA.
Biogeographic range category
Widespread taxa distributed
across North America and
beyond as circumboreal or
arctic-alpine taxa
MADRONO
[Vol. 58
EXAMPLES OF THE BROADER BIOGEOGRAPHIC RELATIONSHIPS OF THE ALPINE FLORA OF THE
Examplar taxa
Anemone drummondii (Ranunculaceae), Carex capitata (Cyperaceae), Crepis
nana (Asteraceae), Cystopteris fragilis (Woodsiaceae), Deschampsia cespitosa
(Poaceae), Oxyria digyna (Polygonaceae), Phleum alpinum (Poaceae),
Rhodiola integrifolium (Crassulaceae), Salix nivalis and S. petrophila
(Salicaceae), Sibbaldia procumbens (Rosaceae), Trisetum spicatum (Poaceae)
Cordilleran taxa widespread in
mountain regions of the
western United States
Antennaria media (Asteraceae), Carex phaeocephala (Cyperaceae), Erigeron
vagus (Asteraceae), Gentiana calycosa (Gentianaceae). Lewisia pygmaea
(Montiaceae), Phlox condensata (Polemoniaceae), Poa glauca subsp. rupicola
(Poaceae), Ribes cereum (Grossulariaceae), Stipa pinetorum (Poaceae)
Intermountain Region taxa
distributed across the Great
Basin
Cryptantha nubigena (Boraginaceae), Cymopterus cinerarius (Apiaceae),
Eriogonum incanum (Polygonaceae), Ivesia shockleyi (Rosaceae), Jamesia
americana (Hydrangeaceae), Podistera nevadensis (Apiaceae), Selaginella
watsonii (Selaginellaceae), Trifolium monanthum subsp. monanthum (Fabaceae)
Taxa extending from the Pacific
Northwest and Cascade
Range
Anelsonia eurycarpa (Brassicaceae), Astragalus whitneyi (Fabaceae), Boechera
howellii (Brassicaceae), Carex breweri (Cyperaceae), Eriogonum lobbii
(Polygonaceae), Gentiana newberryi (Gentianaceae), Potentilla breweri and P.
flabellifolia (Rosaceae), Ranunculus alismifolius var. alismellus
(Ranunculaceae), Senecio fremontii var.occidentalis (Asteraceae), Silene
sargentii (Caryophyllaceae)
are heavily weighted toward subalpine and alpine
specialists, with 29 of these restricted in distribu-
tion to elevations of 2700 m or above, or with
median elevational range above 3000 m. Two
generic lineages are prominent among these
endemics, with five taxa each of Draba and
Eriogonum. Five of the alpine endemics are
annual species—Orochaenactis thysanocarpha
(A. Gray) Coville (Asteraceae), Cryptantha cir-
cumscissa var. rosulata (Boraginaceae), Strep-
tanthus gracilis (Brassicaceae), and Leptosiphon
oblanceolatus (Polemoniaceae).
The southern Sierra Nevada is the most
significant subregion for endemics, with 15 of
the 36 endemic taxa (42%) are restricted in
distribution to the area from the Kings River
drainage south that includes Kings Canyon and
Sequoia National parks (Table 2). These are
Draba cruciata Payson, D. longisquamosa O. E.
Schulz, D. sharsmithii Rollins & R. A. Price,
Eriogonum polypodum Small, E. spergulinum A.
Gray var. pretense (S. Stokes) J. T. Howell, E.
wrightii var. olanchense, Galium hypotrichium A.
Gray subsp. subalpinum (Hilend & J. T. Howell)
Ehrend., Leptosiphon oblanceolatus, Monardella
beneolens Shevock, Ertter & Jokerst, Oreonana
clementis (M. E. Jones) Jeps., Orthochaenactis
thysanocarpha, Phlox dispersa, Pinus balfouriana
var. austina R. J. Mastrog. & J. D. Mastrog.,
Streptanthus gracilis, and Trifolium kingie S.
Waston subsp. dedeckerae (J. M. Gillett) D.
Heller.
Another 13 taxa are restricted to the central
and southern Sierra Nevada. Five of the endem-
ics are present across the northern, central and
southern subregions of the Sierra Nevada, and
three endemic taxa are restricted in occurrence to
the central Sierra Nevada. These are Draba
sierrae, Eriogonum ovalifolium Nutt. var. caeles-
tinum Reveal, and Lupinus gracilentus Greene
(Table 2). Although the absence of peaks above
3500 m in the Sierra Nevada north of Yosemite
National Park explains the lack of endemics
restricted to this subregion, the scattered lower
elevation alpine communities of the northern
Sierra Nevada retain moderately high richness of
species.
In addition to the members of the alpine flora
that are endemic to the Sierra Nevada, there are
an additional 31 alpine taxa that are Californian
endemics, allowing for a broad interpretation of
the floristic region to include the westernmost
ranges of the Great Basin lying close to the Sierra
Nevada and the southern Cascade Range in
Oregon. Many of the Californian endemics have
ranges that extend to the Sweetwater and/or —
White Inyo mountains, while others extend into |
the high Transverse and Peninsular Ranges of |
southern California and a small number extend ©
into the southern Cascade Range. The pattern of |
dominant endemism centered in the southern
Sierra Nevada is not seen among these taxa.
Twelve of these occur throughout the Sierra
Nevada and an additional 12 are restricted to the ©
central and southern areas of the range, while |
only four species are limited to the southern
Sierra Nevada.
If the lower limit of the alpine zone were |
dropped to 3300 m, 11 additional Sierra Nevada
endemics would be added (Table 2). Four of |
these are restricted to the southern Sierra Nevada |
(Astragalus ravenii, Boechera pygmaea (Rollins) |
Al-Shehbaz, Castilleja praeterita Heckard & ©
Bacig., and Phacelia orogenes), with three each —
present in the central and southern Sierra Nevada |
and in all three regions. One species is restricted
2011]
to the central Sierra Nevada. Five additional
Californian endemics would be added if the lower
alpine limit was dropped to 3300 m (Table 2).
DISCUSSION
Defining the Alpine Zone of California
Critically defining what species should be
included in an alpine flora is an imperfect task
given the lack of a simple operational definition, as
discussed below. The high elevation areas of the
Sierra Nevada broadly classified as subalpine and
alpine, or upper Hudsonian and Arctic-Alpine in
the Merriam life zone classification (Daubenmire
1938), would roughly include those areas lying
above about 3000 m elevation (Fig. 2). Such
subalpine and alpine habitats cover extensive areas
of the central and southern Sierra Nevada, but
only scattered areas of the northern Sierra Nevada
lying north of Sonora Pass. The higher elevation
area of this northern Sierran region, however,
supports mosaics of subalpine forest, shrublands,
and low alpine-like vegetation (Smiley 1915, 1921).
A simple definition of alpine habitat is that
area occurring above treeline, with the caveat
that most alpine species are not obligate in
inhabiting habitats above treeline and typically
occur to varying degrees at lower elevations
(Packer 1974). While this approach sounds
logical, timberline itself can be highly variable
even in a local area depending on slope exposure,
erosional history, parent material, disturbance
history, and local microclimate (Billings 2000).
Sharsmith (1940) recognized the alpine flora as
a distinct subdivision of the overall California
flora, characterized by its geographic range,
growth forms, species composition, and constancy
with which the alpine association of species is
maintained. Although he described the alpine zone
as reaching its lower limit at an average elevation
of 3500 m, the limit used in this paper, nowhere in
his dissertation is there a clear statement of criteria
for his inclusion of species. He stated,
While certain species are absolute indicators of
the alpine flora, it is the particular association
of species which gives it its characteristic.
Although many species occur elsewhere, this
special assemblage is not met until the alpine
area 1s reached. Everywhere the flora presents
the impression of unity, an impression rein-
forced by increased field experience.
The combined subalpine and alpine flora of the
Sierra Nevada as defined by Smiley (1921)
included 633 species, with 41 listed as indicators
of the Arctic-Alpine zone. He considered 158
Species to be Sierra Nevada endemics and
another 20 species to be restricted in distribution
to the Sierra Nevada and the southern California
mountains. Sharsmith (1940) included 189 species
RUNDEL: SIERRA NEVADA ALPINE FLORA 165
in his alpine flora of the Sierra Nevada, with 31
of these considered to be endemic. This flora was
composed of 183 herbaceous perennials and six
annual species, but did not include any woody
species. A similar estimate was made by Stebbins
(1982) who stated that there were 207 species in
the Sierra Nevada alpine flora. Finally, a much
higher estimate came from Major and Bamberg
(1967) who used the species descriptions in Munz
(1959) to estimate a Sierran alpine flora of about
600 species, a number similar to that reported
here for taxa reaching 3300 m or above.
Growth Forms
The relative dominance of a herbaceous
perennial growth forms present in the alpine
flora of the Sierra Nevada is typical of other
alpine floras worldwide and does not change
dramatically in a gradient from the Rocky
Mountains west across the Great Basin (Billings
1978, 2000; Rundel et al. 2008). Herbaceous
perennials have the characteristic of maintaining
large proportions of total biomass belowground
where they play an important role in carbohy-
drate storage over the winter months (Mooney
and Billings 1960; Billings 1974). The herbaceous
perennials include species with a variety of
ecological forms and life history strategies of
carbon allocation to belowground and above-
ground vegetative, and reproductive tissues
(Rundel et al. 2005), and many of these are
relatively long-lived plants surviving for decades
(Billings 1974; Pollak 1991).
As in other alpine regions, perennial grami-
noids in alpine habitats of the Sierra Nevada
commonly dominate plant communities of wet
meadows that dry earlier than fellfield commu-
nities. In contrast, fellfield habitats exhibit a
mixed dominance of broad-leaved erect perenni-
als, perennial graminoids, and mats and cushions
(Rundel et al. 2005). Mat and cushion growth
forms of herbaceous perennials are widespread in
the high elevation Sierra Nevada, where the 46
taxa listed here represent 12% of the alpine flora.
These are most prominent ecologically on wind-
swept rocky slopes or other exposed areas that
remain snow-free during the winter.
Because of limiting stress factors of short and
severe growing conditions, annual plants are
generally rare in the typical circumboreal arctic-
alpine floras of the Northern Hemisphere,
comprising no more than 1—2% of the flora
(Billings 2000). Although not abundant, annuals,
nevertheless, are more common in alpine flora of
the Sierra Nevada and White Mountains where
they comprise about 6—-8% of the floras (Jackson
and Bliss 1982; Jackson 1985; Rundel et al. 2008).
The species richness of alpine annual taxa,
however, drops rapidly at elevations above 3300
in the Sierra Nevada (Fig. 5). Went (1948, 1953)
166 MADRONO [Vol. 58
TABLE 2. ENDEMIC TAXA OF THE ALPINE FLORA OF THE SIERRA NEVADA, CALIFORNIA WITH THEIR UPPER
RANGE OF OCCURRENCE AND GROWTH FORMS. For range: n = northern Sierra Nevada, c = central Sierra
Nevada, and s = southern Sierra Nevada. For growth forms: A = annual, G = geophyte, P = erect herbaceous
perennial, P-G = perennial graminoid, P-MAT = mat or cushion, SS = subshrub, T = tree. Species names follow
Baldwin et al. (2012).
Endemic group Family Range Growth form
Sierra Nevada endemics >3500 m
Aquilegia pubescens Ranunculaceae n,c,s P
Calamagrostis muiriana Poaceae C,S P-G
Carex congdonii Cyperaceae C,s P-G
Cryptantha circumscissa var. rosulata Boraginaceae n,Cc,s A
Dodecatheon subalpinum Primulaceae C,S P
Draba cruciata Brassicaceae S P
Draba lemmonii Brassicaceae n,C,s P
Draba longisquamosa Brassicaceae S P
Draba sharsmithii Brassicaceae S P
Draba sierrae Brassicaceae Cc P-MAT
Eriogonum nudum var. scapigerum Polygonaceae C,S P
Eriogonum ovalifolium var. caelestinum Polygonaceae Cc P-MAT
Eriogonum polypodum Polygonaceae S P-MAT
Eriogonum spergulinum var. pratense Polygonaceae S P
Eriogonum wrightii var olanchense Polygonaceae S P-MAT
Galium hypotrichium subsp. subalpinum Rubiaceae S P
Hazardia whitneyi var. whitneyi Asteraceae n,c,s SS
Ivesia muirti Rosaceae C,S P
Ivesia pygmaea Rosaceae CS P
Leptosiphon oblanceolatus Polemoniaceae S A
Lewisia disepala Montiaceae c,S P
Lupinus covillei Fabaceae C,S P
Lupinus gracilentus Fabaceae Cc P
Luzula orestera Juncaceae n,c,$S P-G
Monardella beneolens Lamiaceae S SS
Oreonana clementis Apiaceae S P
Oreostemma peirsonii Asteraceae C,S P
Orochaenactis thysanocarpha Asteraceae S A
Phlox dispersa Polemoniaceae S P-MAT
Pinus balfouriana var. austrina Pinaceae S T
Poa stebbinsii Poaceae c,S P-G
Polemonium eximium Polemoniaceae c,S P
Stipa kingii Poaceae C,S P-G
Streptanthus gracilis Brassicaceae S A
Trichophorum clementis Cyperaceae C,S P-G
Trifolium kingii subsp. dedeckerae Fabaceae s P
Sierra Nevada endemics 3300-3499 m
Astragalus ravenii Fabaceae S P
Boechera pygmaea Brassicaceae S P
Castilleja praeterita Orobanchaceae S FP
Erigeron elmeri Asteraceae C,S P
Hulsea vestita subsp. vestita Asteraceae C;s P
Ipomopsis aggregata subsp. bridgesii Polemoniaceae C,Ss P
Lilium kelleyanum Liliaceae c,s G
Lomatium torreyi Apiaceae n,c,s P
Phacelia eisenti Boraginaceae c,s A
Phacelia orogenes Boraginaceae S A
Trifolium monanthum subsp. tenerum Fabaceae n,c,s P
Californian endemics >3500
Astragalus kentrophyta var. danaus Fabaceae C,S P-MAT
Carex mariposana Cyperaceae n,c,s P-G
Castilleja nana Orobanchaceae n,c,s P
Chaenactis alpigena Asteraceae n,c,s P-MAT
Delphinium polycladon Ranunculaceae n,c,s P
Draba breweri Brassicaceae n,c,s P
Draba subumbellata Brassicaceae S P-MAT
Eriogonum gracilipes Polygonaceae C,S P-MAT
2011]
TABLE 2.
Endemic group
Galium hypotrichium subsp. hypotrichium
Hulsea vestita subsp. pygmaea
Ivesia lycopodioides subsp. lycopodioides
Ivesia lycopodioides subsp. scandularis
Ivesia santolinoides
Lewisia glandulosa
Lupinus breweri var. breweri
Lupinus breweri var. bryoides
Lupinus latifolius var. parishii
Lupinus lepidus var. ramosus
Lupinus padre-crowleyi
Lupinus pratensis var. pratensis
Phyllodoce breweri
Poa keckii
Potentilla pseudosericea
Potentilla wheeleri
Primula suffrutescens
Ranunculus eschscholtzii var. oxynotus
Tonestus peirsonii
Triteleia dudleyi
Viola pinetorum subsp. grisea
Viola purpurea subsp. mesophyta
Californian endemics 3300-3499 m
Eriogonum latens
Frasera puberulenta
Hordeum brachyantherum subsp. californicum
Penstemon caesius
Plagiobothrys torreyi var. diffusus
suggested that many of the high elevation annuals
in the Sierra Nevada were related to desert
species.
Severe winter conditions typically limit the
occurrence of woody plants above treeline, with
prostrate mats and cushions as prominent excep-
tions. The upright growth form of woody shrubs
and krummbholtz tree species exposes their tissues
to extreme conditions of temperature and wind
exposure (Korner 2003). This impact on shrub
occurrence can be seen in Fig. 5 where shrub
richness in the Sierra Nevada drops sharply with
increasing elevation above 3300 m, similar to the
pattern for annual species. Much of the alpine
flora of woody species comes from species of Salix
and members of the Ericaceae, groups which
favor moist habitats with some level of protection.
Biogeography and Endemism
The alpine flora of mountain ranges on the
western margin of the Great Basin of California
and western Nevada exhibit very strong relation-
ships to that of the Sierra Nevada (Rundel et al.
2008). The Sweetwater Mountains supports a
flora of 173 species in 16 km? of alpine habitat,
with 94% of this flora common to the Sierra
Nevada (Hunter and Johnson 1983). The Wassuk
Range has an alpine flora of 70 species in just
2.6 km’ of alpine habitat (Bell and Johnson
1980). Again, this flora is has stronger floristic
RUNDEL: SIERRA NEVADA ALPINE FLORA
167
CONTINUED.
Family Range Growth form
Rubiaceae C,S P
Asteraceae C,S P
Rosaceae ee P
Rosaceae C,S P
Rosaceae n,C,s P
Montiaceae C.S P
Fabaceae n,c,s P-MAT
Fabaceae S P-MAT
Fabaceae C,S P
Fabaceae C,S P
Fabaceae S P-MAT
Fabaceae Gs P
Ericaceae n,c,S S
Poaceae n.c.S P-G
Rosaceae C,S P
Rosaceae S P
Primulaceae n.c.s Pr
Ranunculaceae N:c.s P
Asteraceae Cc Pp
Themidaceae C,S G
Violaceae n,c,s P
Violaceae n,c,s P
Polygonaceae C,S Fr
Gentianaceae C,S P
Poaceae n,c,S P-G
Plantaginaceae S SS
Boraginaceae Ie.s A
relationships to the Sierra Nevada than the
Rocky Mountains.
As with the Sweetwater Mountains and Was-
suk Ranges, the flora of the White Mountains
exhibits much stronger floristic relationships to
the Sierra Nevada than to the Rocky Mountains.
About 90% of the species in the alpine flora of
the White Mountains are also found in the Sierra
Nevada (Rundel et al. 2008), compared with only
58% that occur in the ranges of the central Rocky
Mountains (Scott 1995). These values are signif-
icantly higher for both ranges than earlier
estimates made on incomplete data (Lloyd and
Mitchell 1973).
Mountain ranges in the central Great Basin
generally show strong floristic linkages to the
Rocky Mountains and weaker links to the Sierra
Nevada (Billings 1978). Loope (1969) reported
189 alpine species from the Ruby Mountains in
northeastern Nevada, with this flora heavily
linked to the Rocky Mountains. The isolated
San Francisco Mountains in Arizona with only
5.2 km’ of alpine habitat has 80 species, and
likewise shows strong floristic relationships to the
Rocky Mountains despite its separation of about
200 km (Schaak 1983).
The level of endemism in the alpine Sierra
Nevada flora is a relatively small part of the
overall endemism for the montane and higher
parts of the range. Based on current information,
there are 205 taxa endemic to what The Jepson
168
Manual (Hickman 1993) classifies as the north-
ern, central, and southern high Sierra Nevada,
i.e., the montane, subalpine and alpine zones
above foothill habitats (R. Moe, Univ. of
California, Berkeley, personal communication).
The 36 Sierran endemics present in the alpine
flora would thus comprise 18% of the endemic
flora of the higher Sierra Nevada.
The unique California component of the alpine
flora of the Sierra Nevada is considerably greater
if one considers the endemic component of 31
species in the alpine flora that are not uniquely
limited to the Sierra Nevada but are Californian
endemics as defined earlier. Combining the
endemic taxa with Sierran and Californian limits
of distribution, the total of 66 taxa represents
16% of the alpine flora. This is a relatively high
figure compared to other alpine ranges in
continental North America and Europe, and
reflects the environmental stress conditions asso-
ciated with the summer-dry mediterranean-type
climate present in the Sierra Nevada.
Stebbins (1982) analyzed the flora of the high
Sierra Nevada, defined similarly to that of Smiley
(1921) as the upper montane to alpine zones, and
identified 119 endemic species, 13.5% of the total
flora. He further noted that another 60% of the
flora extended beyond the Sierra Nevada only as
far as southern California, western Nevada, and
southern Oregon.
Raven and Axelrod (1978) briefly discussed the
diversity and evolution of the subalpine and
alpine flora of the Sierra Nevada, listing 68
endemics for this region. Their table of endemics,
however, is outdated by more recent information
on distribution patterns and species concepts.
Shevock (1996) gave a figure of 405 endemic taxa
of vascular plants for the entire Sierra Nevada.
The 36 alpine endemics reported here would
comprise 9% of this total. Of the three geograph-
ical subregions (northern, central, and southern)
of the entire range, the southern Sierra Nevada is
the richest in endemics, rare species, and total
floristic composition (Shevock 1996), a finding
similar to that reported here.
The Evolution of the Sierran Alpine Flora
A detailed assessment of the biogeographic and
evolutionary origin of the alpine flora of the
Sierra Nevada is beyond the scope of this review.
Broad interpretations of biogeographic relation-
ships within alpine lineages have been discussed
by previous authors (e.g., Smiley 1921; Sharsmith
1940; Chabot and Billings 1972; Taylor 1977;
Major and Taylor 1977; Raven and Axelrod
1978; Stebbins 1982) but recent phylogenetic
studies have made many of these earlier interpre-
tations subject to re-evaluation.
Evidence for a north to south route of
colonization of high mountain areas of the Sierra
MADRONO
[Vol. 58
Nevada comes from a pattern of decreasing
presence of Rocky Mountain floristic elements
and an increasing number of endemics alpine
species as one moves from the northern to
southern crest of the range (Chabot and Billings
1972; Raven and Axelrod 1978). The southern
limit of a number of alpine species on Mount
Lassen suggests the possibility that some of these
and other Cascade Range species may well have
been present in the Sierra Nevada in the late
Pliocene or early Pleistocene. Although the
species composition of lower and middle eleva-
tion conifer forests of Lassen National Park are
strongly related to that of the Sierra Nevada, the
summits of the highest peaks in Lassen support
an alpine flora that exhibits stronger floristic
links to Mount Shasta and the Cascade Range to
the north (Gillett et al. 1995). Alpine species with
disjunct patterns of distribution from Mount
Lassen to the Cascade Range volcanoes include
Cardamine_ bellidifolia L. (Brassicaceae), Carex
illota L. H. Bailey (Cyperaceae), Collomia larsenii
(A. Gray) Payson (Polemoniaceae), Draba aure-
ola S. Watson (Brassicaceae), Erigeron elegantu-
lus Greene and E. nivalis Nutt. (Asteraceae),
Hulsea nana A. Gray (Asteraceae), Polemonium
pulcherrimum Hook. var. pilosum (Greenm.)
Brand (Polemoniaceae), and Silene suksdorfii B.
L. Rob. (Caryophyllaceae). The Klamath Moun-
tains also mark the southern distribution limit of
a number of boreal species that do not occur in
the high elevations of the Sierra Nevada (Howell
1944).
Alpine and subalpine species characteristic of
wet meadows and other moist sites typically have
broad geographic ranges but become increasing
habitat specific moving to the south in the Sierra
Nevada as precipitation decreases (Kimball et al.
2004; Moore et al. 2007). The relative isolation of
the Sierra Nevada from northern ranges and the
summer-dry have clearly acted as a filter to
exclude some widespread circumpolar arctic-
alpine species such as Dryas integrifolia Vahl
(Rosaceae) and Silene acaulis L. (Caryophylla-
ceae) which do not occur anywhere in California. ©
Species growing in xeric rocky habitats show |
higher levels of endemism and smaller range size —
due to isolation and divergence from ancestral |
populations distributed in wetter habitats to the |
north. |
More controversial, however, is the origin of |
disjunct Rocky Mountain species present in the —
central and southern Sierra Nevada, often
growing in azonal soil conditions. There is both
geological and paleobotanical evidence to suggest —
that the mean elevation of the Great Basin was as |
much as 1500 m higher in the Miocene and that
the current Basin and Range topography is the |
result of subsidence rather than uplift (Wernicke ©
et al. 1988; Wolfe et al. 1997). The presence of |
higher elevations in the Great Basin during the ©
2011]
Pleistocene could possibly have provided step-
ping stones to allow the dispersal of alpine
organisms from the east (Major and Bamberg
1967; Taylor 1976a). Molecular evidence indi-
cates that at least one lineage of butterflies
entered the Sierra Nevada by this route (Nice
and Shapiro 2001). However, other authors feel
that the majority of these disjunct plant species
reached the Sierra Nevada by the same dominant
route from the Western Cordillera via the
Cascade Range and south (Chabot and Billings
1972).
Modes of speciation in the development of the
endemic alpine flora of the Sierra Nevada are
clearly complex. Polyploidy and associated apo-
mixis are widely recognized as major factors in
plant evolution, and these factors have had a
relatively recent impact on speciation in produc-
ing stable self-propagating lineages (Soltis et al.
2009). In the alpine region of the Sierra Nevada,
as in other alpine regions, diploid lineages of
polyploid complexes often occupy unglaciated
areas and resist introgression due hypothetically
to a significantly higher seed set. However,
asexual apomictic populations are more wide-
spread than their sexual relatives in glaciated
areas. Sexual and asexual polyploids may become
distinct stabilized species through hybrid origin.
Reproductive isolation and stability of tetra-
ploids within their respective distribution as well
as the value of uniparental reproduction provide
the advantages of apomixis. Many important
genera in the alpine flora of the Sierra Nevada are
notable for the presence of apomixis, with
Boechera (Schranz et al. 2005; Dobes et al.
2007), Draba (Jordon-Thaden and Koch 2008),
and Antennaria (Bayer and Stebbins 1987) as
examples. Additional speciose genera in the
Sierra Nevada known to have complex apomictic
populations include Arnica and Crepis (Aster-
aceae; Noyes 2007), Poa and Calamagrostis
(Poaceae), and Potentilla (Rosaceae) (Asker and
Jerling 1992).
Other modes of alpine speciation have also
been described for the Sierra Nevada. Some
speciation, for example, has hypothetically come
from lowland arid-adapted taxa colonizing the
glaciated terrain of the range at the end of the
Pleistocene (Went 1948, 1953). Speciation has
also been shown to be the result of population
disjunction and reproductive isolation (Chase
and Raven 1975).
Although the Transverse and Peninsular rang-
es are well separated from the higher elevations of
the Sierra Nevada, more than one third of the
Sierran alpine flora has a range of distribution
that extends to these southern California ranges.
While some of these species occur at lower
elevations, others are typically subalpine and
alpine species that must have crossed the Mojave
Desert during the cold conditions of the Pleisto-
RUNDEL: SIERRA NEVADA ALPINE FLORA 169
cene. This latter group includes Androsace
septentrionalis (Primulaceae), Hulsea vestita A.
Gray subsp. pygmaea (A. Gray) Wilken (Aster-
aceae), Oxyria digyna (L.) Hill (Polygonaceae),
and Podistera nevadensis (Apiaceae).
There are lessons to be learned from recent
studies of the patterns of diversification in the
European alpine flora. These strongly demon-
strate that speciation have been promoted by
diverse ecological, evolutionary, and life history
traits related to population structure, phyloge-
netic relationships, breeding system, dispersal
syndromes, ecophysiological ranges of habitat
requirements, and competitive abilities (Comes
and Kadereit 1998; Taberlet et al. 1998: Hewitt
2000; Gugerli and Holderegger 2001; Vargas
2003). The complex and dynamic climatic and
geological history of the Sierra Nevada operating
on such traits suggests that there have been a
range of different colonization and extinction
histories that are species specific. Much more
work on the comparative phylogeography of
alpine plants in the Sierra Nevada will be
necessary before we understand all of the factors
responsible for present distributions and predom-
inant modes of speciation in the alpine flora of
the range.
Research Needs
There is little doubt that the stability of the
ecotone between alpine and treeline ecosystems in
the Sierra Nevada and other high mountain
regions has been and continues to be a function
of complex interactions, with multiple drivers
operating across diverse scales of time and space.
This ecotone has been highly dynamic in the past
and given the importance of temperature in
controlling the elevation of treeline and higher
alpine ecosystems, this ecotone and associated
species are likely to be particularly sensitive to
climate change in the future (Lloyd and Graum-
lich 1997; Graumlich et al. 2005; Grabherr et al.
2010). Beyond treeline studies, the expansion of
woody shrub species into alpine habitats has been
shown to also be a sensitive indicator of potential
climate change, with significant feedbacks on
microclimate and soil ecosystems (Hallinger et al.
2010), as well as species facilitation (Callaway et
al. 2002). The potential sensitivity of alpine
ecosystems to climate change has been the
stimulus for establishing the worldwide research
program Global Observation Research Initiative
in Alpine Environments (GLORIA, http://www.
gloria.ac.at) with the aim of providing long-term
observations on the state and dynamics of alpine
biota.
ACKNOWLEDGMENTS
I thank Bruce Baldwin and Richard Moe for their
help in accessing the Jepson Manual database. Peggy
170
Moore and Sylvia Haultain kindly shared species lists
for Yosemite and Sequoia/Kings Canyon National
Parks, respectively. This work is a contribution from
the UCLA La Kretz Center for California Conserva-
tion Science.
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2011] RUNDEL: SIERRA NEVADA ALPINE FLORA 173
APPENDIX 1. Annotated checklist of the alpine flora of the Sierra Nevada, including all taxa reaching an elevation
of 3500 m. Lower and upper elevations limits are those for all of California and taken from Baldwin et al. (2012).
Growth form abbreviations are: P = erect broad-leaved perennial; G = geophytes; P-G = graminoid perennial;
P-MAT = mat or cushion; A = annual; Q = aquatic perennial; SS = subshrub; S = woody shrub; and T = tree.
Biogeographic relationships are abbreviated as follows: WIDE = widespread taxa present in many habitats or
regions across North America and/or throughout the world; CORD = cordilleran taxa widespread in mountain
regions of the western North America; S-C = Sierra/Cascade taxa with a Pacific Northwest distribution; INT =
intermountain taxa present in the Great Basin; END = taxa endemic to the Sierra Nevada; and END-CAL = taxa
endemic to California, as broadly defined in the text. Species names follow Baldwin et al. (2012).
Specific or Lower Upper Growth Biogeographic
Higher level taxon infraspecific taxon elevation (m) elevation (m) form relationship
PTERIDOPHYTA
Ophioglossaceae Botrychium crenulatum 1500 3600 P CORD
Ophioglossaceae Botrychium lineare 2500 4000 P CORD
Ophioglossaceae Botrychium paradoxum 4000 4200 P CORD
Ophioglossaceae Botrychium simplex var. 1500 3800 Pp WIDE
compositum
Ophioglossaceae Botrychium tunux 3600 3600 led WIDE
Pteridaceae Pellaea breweri 1500 3700 P INT
Selaginellaceae Selaginella watsonii 1350 4100 P INT
Woodsiaceae Athyrium distentifolium var. 1700 3700 P WIDE
americanum
Woodsiaceae Cystopteris fragilis 50 4100 i CORD
Woodsiaceae Woodsia scopulina 1300 3500 P WIDE
CONIFERAE
Pinaceae Pinus albicaulis 2135 3700 T CORD
Pinaceae Pinus balfouriana var. 2700 3700 i END
austrina
Pinaceae Pinus contorta subsp. 1525 3500 fi S-C
murrayana
Pinaceae Pinus flexilis 2600 3700 T CORD
Pinaceae Tsuga mertensiana 1200 3500 T S-C
MONOCOTY LEDONAE
Alliaceae Allium obtusum var. obtusum 1500 3500 G INT
Cyperaceae Carex albonigra 3000 4200 P-G CORD
Cyperaceae Carex breweri 2000 3900 P-G S-C
Cyperaceae Carex capitata 1200 3900 P-G WIDE
Cyperaceae Carex congdonii 2600 3900 P-G END
Cyperaceae Carex deflexa var. boottii 0 3800 P-G CORD
Cyperaceae Carex douglasii 300 3800 P-G CORD
Cyperaceae Carex filifolia var. erostrata 1500 3700 P-G CORD
Cyperaceae Carex haydeniana 2400 4200 P-G CORD
Cyperaceae Carex helleri 2400 4100 P-G S-C
Cyperaceae Carex heteroneura 1300 4000 P-G INT
Cyperaceae Carex hoodii 650 3600 P-G CORD
Cyperaceae Carex incurviformis 3700 4000 P-G CORD
Cyperaceae Carex jonesii 900 3500 P-G CORD
Cyperaceae Carex lenticularis var. 0 3600 P-G CORD
lipocarpa
Cyperaceae Carex leporinella 1900 4000 P-G CORD
Cyperaceae Carex mariposana 750 3600 P-G END-CAL
Cyperaceae Carex multicostata 1900 3500 P-G CORD
Cyperaceae Carex nigricans 1900 3700 P-G CORD
Cyperaceae Carex phaeocephala 2500 4000 P-G CORD
Cyperaceae Carex praeceptorium 2200 3500 P-G CORD
Cyperaceae Carex proposita 3000 4100 P-G S-C
Cyperaceae Carex rossii 0 3800 P-G CORD
Cyperaceae Carex scirpoidea var. 2800 3700 P-G CORD
pseudoscirpoidea
Cyperaceae Carex specifica 1200 3500 P-G INT
Cyperaceae Carex spectabilis 1800 3700 P-G CORD
Cyperaceae Carex straminiformis 1700 4100 P-G S-C
Cyperaceae Carex subfusca 700 3800 P-G INT
Cyperaceae Carex subnigricans 2600 3800 P-G CORD
Cyperaceae Carex tahoensis 3200 3700 P-G CORD
174
Higher level taxon
Cyperaceae
Cyperaceae
Cyperaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaceae
Juncaginaceae
Liliaceae
Melanthiaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
MADRONO
APPENDIX |. _CONTINUED.
Specific or Lower Upper
infraspecific taxon elevation (m) elevation (m)
Carex vernacula 1800 4000
Eleocharis quinqueflora 40 3600
Trichophorum clementis 2400 3600
Juncus bryoides 600 3600
Juncus drummondii 200 3500
Juncus mertensianus 1200 3500
Juncus mexicanus 0 3800
Juncus orthophyllus 1200 3500
Juncus parryi 2000 3800
Luzula divaricata 2100 3700
Luzula orestera 2700 3600
Luzula spicata 2900 3700
Luzula subcongesta 2000 3500
Triglochin palustris 2400 3500
Calochortus leichtlinii 1300 4000
Veratrum californicum var. 0 3500
californicum
Agrostis idahoensis 0 3500
Agrostis pallens 200 3500
Agrostis scabra 100 3500
Agrostis thurberiana 1300 3500
Agrostis variabilis 1600 4000
Alopecurus aequalis var 50 3500
aequalis
Bromus carinatus var. 0 3500
carinatus
Bromus carinatus var. 0 3500
marginatus
Bromus orcuttianus 560 3500
Bromus porteri 550 3500
Bromus richardsonii 1200 3600
Calamagrostis muiriana 2480 3900
Calamagrostis purpurascens 1300 4000
Deschampsia cespitosa 0 3820
subsp. cespitosa
Elymus elymoides subsp. 295 4200
californicus
Elymus multisetus 0 3800
Elymus scribneri 2900 4200
Elymus sierrae 1800 3530
Festuca brachyphylla subsp. 2800 4300
breviculmis
Festuca minutiflora 2850 4050
Hordeum jubatum var. 20 3500
jubatum
Koeleria macrantha 0 3840
Muhlenbergia richardsonis 1220 3670
Phleum alpinum 0 3700
Poa abbreviata subsp. 3300 3660
pattersonit
Poa cusickii subsp. epilis 2400 3600
Poa cusickii subsp. 2100 3500
purpurascens
Poa glauca subsp. rupicola 3300 4100
Poa keckii 3300 4340
Poa lettermanii 3500 4300
Poa secunda subsp. secunda 0 3900
Poa stebbinsii 2700 3700
Poa wheeleri 1300 3800
Stipa hymenoides 60 3500
Stipa kingii 2000 3650
Stipa nelsonii subsp. dorei 450 3500
Stipa occidentalis subsp. 1200 3500
pubescens
Growth
form
P-G
ror rg Rg tg ig bg hg by
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an)
ac lacldeclixcMaciacline| inc). \a-Sa°imia-|ta- MMM sc]ga°)iin-iin> MMMa-e>/ai tae) xc) og gela- ac) x-/bn-/ae) o7[la-]s Mela~ | lange ae) ec aclac exc >
QQOQQQQQHQ0Q QAO AQAOQ AM AANAM ©Q AMAH0OO0H 0A 0 AHAMO0O0
[Vol. 58
Biogeographic
relationship
CORD
WIDE
END
CORD
CORD
CORD
WIDE
CORD
CORD
S-C
END
WIDE
S-C
WIDE
S-C
CORD
CORD
CORD
WIDE
CORD
CORD
WIDE
S-C
S-C
S-C
CORD
CORD
END
WIDE
WIDE
CORD
CORD
CORD
INT
WIDE
CORD
WIDE
WIDE
CORD
WIDE
CORD
CORD
S-C
CORD
END-CAL
CORD
CORD
END
CORD
CORD
END
CORD
CORD
2011]
Higher level taxon
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Themidaceae
EUDICOTS
Adoxaceae
Apiaceae
Apiaceae
Apiaceae
Apiaceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
RUNDEL: SIERRA NEVADA ALPINE FLORA
APPENDIX 1.
Specific or
infraspecific taxon
Stipa pinetorum
Stipa webberi
Torreyochloa erecta
Torreyochloa pallida var.
pauciflora
Trisetum spicatum
Triteleia dudleyi
Sambucus racemosa vat.
melanocarpa
Cymopterus cinerarius
Oreonana clementis
Podistera nevadensis
Sphenosciadium capitellatum
Achillea millefolium
Ageratina occidentalis
Agoseris aurantiaca var.
aurantiaca
Agoseris monticola
Antennaria media
Antennaria pulchella
Antennaria rosea subsp.
confinis
Antennaria rosea subsp.
rosea
Antennaria umbrinella
Arnica chamissonis
Arnica lanceolata subsp.
prima
Arnica longifolia
Arnica mollis
Arnica ovata
Artemisia arbuscula subsp.
arbuscula
Artemisia ludoviciana subsp.
incompta
Artemisia norvegica subsp.
saxatilis
Artemisia spiciformis
Chaenactis alpigena
Chaenactis douglasii var.
douglasii
Chrysothamnus viscidiflorus
var. viscidiflorus
Cirsium arizonicum var.
arizonicum
Cirsium occidentale var.
venustum
Cirsium scariosum var.
americanum
Crepis nana
Ericameria bloomeri
Ericameria discoidea
Ericameria nauseosa var.
Speciosa
Ericameria parryi var.
monocephala
Ericameria suffruticosa
Erigeron algidus
Erigeron compositus
Erigeron lonchophyllus
Erigeron pygmaeus
Erigeron vagus
Lower
elevation (m)
2000
1450
2000
0
1370
1200
1800
2100
1500
3000
CONTINUED.
Upper
elevation (m)
3900
3500
3500
3500
3900
3500
3600
3500
4000
4000
3500
3650
3700
3500
3800
3900
3700
3700
3700
3900
3500
3500
3500
3500
3600
3800
3500
3800
3700
3900
3500
4000
3500
3600
3500
4000
4000
3800
3500
3700
3800
3700
4300
3550
4100
4400
Growth
form
PPP
@uaane
eo
26
N
wy Oy oC oy
uv
> >
HH
175
Biogeographic
relationship
INT
INT
INT
CORD
WIDE
END-CAL
CORD
INT
END
INT
CORD
WIDE
CORD
CORD
S-C
CORD
INT
WIDE
CORD
WIDE
WIDE
CORD
CORD
CORD
CORD
CORD
INT
WIDE
CORD
END-CAL
CORD
INT
INT
INT
CORD
CORD
INT
INT
INT
INT
INT
INT
WIDE
WIDE
S-C
INT
176
Higher level taxon
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Boraginaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
Brassicaceae
APPENDIX |.
Specific or
infraspecific taxon
Eriophyllum lanatum var.
integrifolium
Hazardia whitneyi var.
whitneyi
Hieracium nudicaule
Hieracium triste
Hulsea algida
Hulsea vestita subsp.
pygmaea
Hymenoxys hoopesii
Oreostemma alpigenum var.
andersonii
Oreostemma peirsonii
Orochaenactis
thysanocarpha
Packera cana
Packera werneriifolia
Pyrrocoma apargioides
Raillardella argentea
Raillardella scaposa
Senecio fremontii var.
occidentalis
Senecio integerrimus var.
major
Senecio pattersonensis
Senecio scorzonella
Senecio spartioides
Solidago multiradiata
Sphaeromeria cana
Stenotis acaulis
Tonestus peirsonii
Cryptantha circumscissa vat.
CIFCUMSCISSA
Cryptantha circumscissa vat.
rosulata
Cryptantha glomeriflora
Cryptantha humilis
Cryptantha nubigena
Hackelia micrantha
Hackelia sharsmithii
Nama densum
Phacelia hastata subsp.
compacta
Phacelia mutabilis
Phacelia ramosissima
Anelsonia eurycarpa
Boechera depauperata
Boechera howellii
Boechera inyoensis
Boechera covillei
Boechera inyoensis
Boechera lemmonii
Boechera lyallii
Boechera paupercula
Boechera repanda
Boechera tiehmii
Cardamine cordifolia
Descurainia incana
Draba albertina
Draba breweri
Draba cana
Draba cruciata
Draba densifolia
Draba lemmonii
MADRONO
CONTINUED.
Lower Upper
elevation (m) elevation (m)
1400 3500
1200 3500
1800 3500
1650 3550
3000 4000
3200 3900
1500 3650
1200 3500
3000 3800
1600 3800
1200 3500
3000 3650
2200 3800
1800 3900
2000 3500
2800 4000
100 3600
3000 3700
1600 3500
1000 3500
1250 3950
1800 4000
1800 3600
2900 3700
150 3650
2950 3650
1800 3750
1700 3600
2400 3900
1200 3500
3150 3700
880 3560
1500 4000
900 3500
0 3800
1600 4100
3650 3900
1500 3800
1200 3500
2200 3500
1200 3500
2000 4350
2000 3900
2500 3700
1400 3600
3000 3600
600 3600
100 3500
900 3700
3100 4100
0 4100
2500 3963
1900 3650
3050 4000
Growth
form
iy
eee Aan Dae lla
>
=
-
TUVUU DUE UU US UT >>Duv~TU> D> Sve AvA vy 4
a
ti
>
v
yuu
>
=
[Vol. 58
Biogeographic
relationship
CORD
END
2011] RUNDEL: SIERRA NEVADA ALPINE FLORA Li?
APPENDIX |. CONTINUED.
Specific or Lower Upper Growth Biogeographic
Higher level taxon infraspecific taxon elevation (m) — elevation (m) form relationship
Brassicaceae Draba lonchocarpa 2800 4000 P WIDE
Brassicaceae Draba longisquamosa 3000 3900 P END
Brassicaceae Draba novolympica 1500 3700 P-MAT CORD
Brassicaceae Draba oligosperma 2000 3900 P-MAT CORD
Brassicaceae Draba praealta 2500 4100 P WIDE
Brassicaceae Draba sharsmithii 3300 3800 P END
Brassicaceae Draba sierrae 3500 4114 P-MAT END
Brassicaceae Draba subumbellata 3300 4100 P-MAT END-CAL
Brassicaceae Erysimum capitatum var. 0 4000 P WIDE
capitatum
Brassicaceae Erysimum perenne 2000 4000 P S-C
Brassicaceae Lepidium densiflorum 0 3500 Pp-A WIDE
Brassicaceae Rorippa curvipes 100 3500 P-A CORD
Brassicaceae Rorippa curvisiliqua 0 3500 A CORD
Brassicaceae Streptanthus gracilis 2600 3600 A END
Brassicaceae Streptanthus tortuosus 200 4100 P S-C
Caryophyllaceae Cerastium beeringianum 2900 4300 P-MAT WIDE
Caryophyllaceae Ereomogone kingii var. 2100 4050 P-MAT S-C
glabrescens
Caryophyllaceae Minuartia nuttallii var. 2600 3800 P-MAT S-C
gracilis
Caryophyllaceae Minuartia obtusiloba 3150 3700 P-MAT CORD
Caryophyllaceae Minuartia rubella 2400 3800 P CORD
Caryophyllaceae Minuartia stricta 3500 3900 Pp CORD
Caryophyllaceae Sagina saginoides 1000 3800 P WIDE
Caryophyllaceae Silene bernardina 1350 3600 P CORD
Caryophyllaceae Silene sargentii 2400 3800 Pp S-C
Caryophyllaceae Stellaria calycantha 1700 3800 Pp WIDE
Chenopodiaceae Chenopodium atrovirens 300 3500 A CORD
Chenopodiaceae Monolepis nuttalliana 0 3700 A CORD
Crassulaceae Rhodiola integrifolia 1800 4000 Pr WIDE
Crassulaceae Sedum obtusatum subsp. 1200 3700 P S-C
obtusatum
Ericaceae Cassiope mertensiana 1800 3505 S CORD
Ericaceae Gaultheria humifusa 1350 4000 S CORD
Ericaceae Kalmia polifolia subsp. 1000 3500 S CORD
microphylla
Ericaceae Phyllodoce breweri 1200 3500 S END-CAL
Ericaceae Pterospora andromedea 60 3700 P WIDE
Ericaceae Rhododendron columbianum 0 3630 S CORD
Fabaceae Astragalus kentrophyta var. 2900 4000 P-MAT END-CAL
danaus
Fabaceae Astragalus kentrophyta var. 2700 3600 Er CORD
tegetarius
Fabaceae Astragalus lentiginosus var. 1250 3700 P INT
ineptus
Fabaceae Astragalus platytropus 2350 3500 P INT
Fabaceae Astragalus purshii var. 1500 3650 P INT
lectulus
Fabaceae Astragalus whitneyi var. 1550 3500 P S-C
whitneyi
Fabaceae Lupinus adsurgens 1000 3500 P S-C
Fabaceae Lupinus angustiflorus 1000 3500 P INT
Fabaceae Lupinus argenteus var. 1500 3500 P INT
meionanthus
Fabaceae Lupinus argenteus var. 2500 3500 P INT
montigenus
Fabaceae Lupinus breweri var. breweri 1000 4000 P-MAT END-CAL
Fabaceae Lupinus breweri var. 2500 4000 P-MAT END-CAL
bryoides
Fabaceae Lupinus breweri var. 2000 3500 P-MAT INT
grandiflorus
Fabaceae Lupinus covillei 2500 3500 P END
Fabaceae Lupinus gracilentus 2500 3500 P END
178 MADRONO [Vol. 58
APPENDIX |. CONTINUED.
Specific or Lower Upper Growth Biogeographic
Higher level taxon infraspecific taxon elevation (m) elevation (m) form relationship
Fabaceae Lupinus latifolius var. 1000 3500 iP S-C
columbianus
Fabaceae Lupinus latifolius var. 0 3500 ig END-CAL
parishii
Fabaceae Lupinus lepidus var. lobbii 2000 3500 P S-C
Fabaceae Lupinus lepidus var. ramosus 3000 4000 P END-CAL
Fabaceae Lupinus obtusilobus 2500 3500 P S-C
Fabaceae Lupinus padre-crowleyi 2500 4000 P-MAT END-CAL
Fabaceae Lupinus pratensis var. 1000 3500 P END-CAL
pratensis
Fabaceae Oxytropis borealis var. 3300 3900 P INT
australis
Fabaceae Oxytropis borealis var. 3300 3900 P CORD
viscida
Fabaceae Oxytropis parryi 3100 3800 P-MAT INT
Fabaceae Trifolium kingii subsp. 2100 3500 P END
dedeckerae
Fabaceae Trifolium monanthum subsp. 1700 3900 P-MAT INT
monanthum
Gentianaceae Comastoma tenellum 3200 3900 A WIDE
Gentianaceae Gentiana calycosa 1300 3900 P CORD
Gentianaceae Gentiana newberryi var. 1500 4000 P. S-C
tiogana
Gentianaceae Gentianella amarella subsp. 1500 3500 A WIDE
acuta
Gentianaceae Gentianopsis holopetala 1800 4000 A S-C
Grossulariaceae Ribes cereum var. inebrians 2100 3850 S INT
Grossulariaceae Ribes montigenum 800 4000 S CORD
Grossulariaceae Ribes velutinum 700 3500 S CORD
Hydrangeaceae Jamesia americana 2070 3700 S INT
Lamiaceae Monardella beneolens 2500 3600 SS END
Lamiaceae Monardella linoides subsp. 1000 3500 SS INT
sierrae
Lamiaceae Monardella odoratissima 1000 3500 SS INT
subsp. glauca
Linaceae Linum lewisii 400 3657 P INT
Montiaceae Calyptridium monospermum 300 3970 P INT
Montiaceae Calyptridium roseum 1500 3800 A CORD
Montiaceae Calyptridium umbellatum 240 4300 Pp CORD
Montiaceae Claytonia nevadensis 2200 3500 P S-C
Montiaceae Lewisia disepala 1300 3500 P END
Montiaceae Lewisia glandulosa 3000 4000 P END-CAL
Montiaceae Lewisia nevadensis 609 3596 P S-C
Montiaceae Lewisia pygmaea 1700 4020 P CORD
Montiaceae Lewisia triphylla 1300 3500 P CORD
Montiaceae Montia chamissoi 1100 3700 P S-C
Onagraceae Epilobium anagallidifolium 1500 4500 Pp WIDE
Onagraceae Epilobium ciliatum subsp. 0 4000 P CORD
ciliatum
Onagraceae Epilobium ciliatum subsp. 0 3500 P WIDE
glandulosum
Onagraceae Epilobium clavatum 1200 4200 P CORD
Onagraceae Epilobium glaberrimum 1200 3800 P CORD
subsp. fastigiatum
Onagraceae Epilobium hallianum 100 3700 P CORD
Onagraceae Epilobium hornemannii 1200 3900 P WIDE
subsp. hornemannii
Onagraceae Epilobium obcordatum 1700 4000 P S-C
Onagraceae Epilobium oregonense 1200 3500 P CORD
Onagraceae Epilobium saximontanum 1400 3500 P CORD
Onagraceae Gayophytum decipiens 1800 4200 A INT
Onagraceae Gayophytum diffusum subsp. 800 3700 A S-C
diffusum
2011]
Higher level taxon
Onagraceae
Onagraceae
Onagraceae
Orobanchaceae
Orobanchaceae
Orobanchaceae
Orobanchaceae
Orobanchaceae
Orobanchaceae
Orobanchaceae
Orobanchaceae
Parnassiaceae
Phrymaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Plantaginaceae
Polemoniaceae
Polemoniaceae
Polemoniaceae
Polemoniaceae
Polemoniaceae
Polemoniaceae
Polemoniaceae
Polemoniaceae
Polemoniaceae
Polemoniaceae
Polemoniaceae
Polygonaceae
Polygonaceae
Polygonaceae
Polygonaceae
Polygonaceae
Polygonaceae
Polygonaceae
Polygonaceae
Polygonaceae
Polygonaceae
Polygonaceae
RUNDEL: SIERRA NEVADA ALPINE FLORA
APPENDIX 1. CONTINUED.
Specific or Lower Upper
infraspecific taxon elevation (m) elevation (m)
Gayophytum diffusum subsp. 800 3700
parviflorum
Gayophytum racemosum 1000 4000
Gayophytum ramosissimum 500 3500
Castilleja applegatei subsp. 1900 3600
pallida
Castilleja applegatei subsp. 300 3600
pinetorum
Castilleja lemmonii 1550 3700
Castilleja miniata subsp. 1500 3500
miniata
Castilleja nana 2400 4200
Pedicularis attollens 1200 4000
Pedicularis groenlandica 1000 3600
Pedicularis semibarbata 1500 3500
Parnassia palustris 0 3600
Mimulus suksdorfii 1100 4000
Callitriche palustris 0 4000
Collinsia parviflora 800 3500
Collinsia torreyi var. wrightii 800 4000
Penstemon davidsonii 2000 3750
Penstemon heterodoxus var. 2700 3900
heterodoxus
Penstemon newberryi var. 1000 3700
newberryi
Penstemon procerus var. 2100 3600
formosus
Penstemon roezlii 300 3500
Penstemon rostriflorus 500 3500
Penstemon rydbergii var. 1000 3600
oreocharis
Penstemon speciosus 850 3800
Veronica wormskjoldii 1500 3500
Collomia linearis 600 3650
Gymnosteris parvula 2400 3700
Ipomopsis congesta subsp. 1500 3700
montana
Leptosiphon oblanceolatus 2800 3700
Linanthus pungens 1700 4000
Phlox condensata 2000 4000
Phlox diffusa 1100 3600
Phlox dispersa 3600 4200
Phlox pulvinata 3300 4300
Polemonium eximium 3000 4200
Polemonium pulcherrimum 2400 3700
var. pulcherrimum
Eriogonum gracilipes 2900 3900
Eriogonum incanum 2100 4000
Eriogonum lobbii 1600 3800
Eriogonum nudum var. 2800 3800
scapigerum
Eriogonum ovalifolium var. 3000 3600
caelestinum
Eriogonum ovalifolium var. 1700 4200
nivale
Eriogonum polypodum 2800 3500
Eriogonum rosense var. 2300 4000
rosense
Eriogonum spergulinum var. 1300 3500
pratense
Eriogonum umbellatum var. 3000 3600
covillei
Eriogonum wrightii var. 3500 3600
olanchense
Growth
form
A
tT > >
>PrOruwu yw ys UT CU
P-MAT
P-MAT
P-MAT
P-MAT
~
Su> Uppy VAL
-
179
Biogeographic
relationship
INT
CORD
INT
INT
INT
S-C
INT
END-CAL
S-C
S-C
S-C
WIDE
INT
WIDE
WIDE
INT
INT
S-C
INT
INT
INT
INT
INT
INT
WIDE
WIDE
CORD
S-C
END
CORD
CORD
CORD
END
CORD
END
S-C
END-CAL
INT
S-C
END
END
INT
END
INT
END
END-CAL
END
180 MADRONO [Vol. 58
APPENDIX |. CONTINUED.
Specific or Lower Upper Growth Biogeographic
Higher level taxon infraspecific taxon elevation (m) elevation (m) form relationship
Polygonaceae Oxyria digyna 1800 4000 P WIDE
Polygonaceae Rumex californicus 0 3500 P CORD
Polygonaceae Rumex paucifolius 1500 4000 P S-C
Polygonaceae Rumex salicifolius 0 3500 P INT
Polygonaceae Rumex utahensis 1000 3500 P CORD
Primulaceae Androsace septentrionalis 2700 3600 P-A WIDE
Primulaceae Dodecatheon redolens 2400 3600 P INT
Primulaceae Dodecatheon subalpinum 2100 4000 P END
Primulaceae Primula suffrutescens 2000 4200 P END-CAL
Pteridaceae Pellaea breweri 1500 3700 P INT
Ranunculaceae Aconitum columbianum 300 3500 | CORD
subsp. columbianum
Ranunculaceae Aquilegia pubescens 2600 3650 P END
Ranunculaceae Delphinium polycladon 2200 3600 P END-CAL
Ranunculaceae Ranunculus alismifolius var. 1400 3600 P S-C
alismellus
Ranunculaceae Ranunculus eschscholtzii var. 2200 3600 P S-C
eschscholtzii
Ranunculaceae Ranunculus eschscholtzii var. 2700 4300 P END-CAL
oxynotus
Ranunculaceae Ranunculus glaberrimus 1200 3600 P CORD
Ranunculaceae Thalictrum alpinum 2900 3700 P WIDE
Ranunculaceae Thalictrum sparsiflorum 1400 3500 P CORD
Rosaceae Dasiphora fruticosa 2000 3600 S WIDE
Rosaceae Drymocaulis lactea var. 1800 3700 P INT
lactea
Rosaceae Drymocaulis pseudorupestris 3200 3900 P CORD
var. crumiana
Rosaceae Drymocaulis pseudorupestris 2300 3500 leg CORD
var. saxicola
Rosaceae Holodiscus discolor var. 1159 4000 S CORD
microphyllus
Rosaceae Ivesia gordonii var. 1800 3500 P INT
ursinorum
Rosaceae Ivesia lycopodioides subsp. 3000 4000 P END-CAL
lycopodioides
Rosaceae Ivesia lycopodioides subsp. 3000 4115 P END-CAL
scandularis
Rosaceae Ivesia muirii 2900 4000 P END
Rosaceae Ivesia pygmaea 2700 4000 P END
Rosaceae Ivesia santolinoides 1500 3600 P END-CAL
Rosaceae Ivesia shockleyi 2700 4000 P-MAT INT
Rosaceae Potentilla breweri 1500 3700 P S-C
Rosaceae Potentilla flabellifolia 1700 3700 P S-C
Rosaceae Potentilla bruceae 1200 3700 P INT
Rosaceae Potentilla glaucophylla var. 2600 3500 P WIDE
glaucopylla
Rosaceae Potentilla gracilis var. 800 3500 P INT
fastigiata
Rosaceae Potentilla jepsonii 2700 3800 P INT
Rosaceae Potentilla pensylvanica 2700 3800 P WIDE
Rosaceae Potentilla pseudosericea 3200 4300 P END-CAL
Rosaceae Potentilla wheeleri 1800 3500 P END-CAL
Rosaceae Sibbaldia procumbens 1820 3700 | a WIDE
Rosaceae Sorbus californica 1200 4300 S INT
Rubiaceae Galium bifolium 1500 3700 A CORD
Rubiaceae Galium grayanum var. 1830 3500 P S-C
grayanum
Rubiaceae Galium hypotrichium subsp. 3000 4200 P END-CAL
hypotrichium
Rubiaceae Galium hypotrichium subsp. 2650 3880 P END
subalpinum
Salicaceae Salix brachycarpa var. 3200 3500 S CORD
brachycarpa
2011] RUNDEL: SIERRA NEVADA ALPINE FLORA 18]
APPENDIX |. CONTINUED.
Specific or Lower Upper Growth Biogeographic
Higher level taxon infraspecific taxon elevation (m) elevation (m) form relationship
Salicaceae Salix eastwoodiae 1600 3800 S CORD
Salicaceae Salix geyeriana 1450 3600 S CORD
Salicaceae Salix lemmonii 1400 3500 S CORD
Salicaceae Salix nivalis 3100 3500 S CORD
Salicaceae Salix orestera 1100 4000 S S-C
Salicaceae Salix petrophila 1670 4000 S CORD
Salicaceae Salix planifolia 2500 4000 S WIDE
Saxifragaceae Heuchera rubescens 1000 4000 P CORD
Saxifragaceae Lithophragma glabrum 0 3750 P CORD
Saxifragaceae Micranthes aprica 1600 3600 P CORD
Saxifragaceae Micranthes bryophora 1600 3500 P CORD
Saxifragaceae Micranthes nidifica 1000 3500 lg CORD
Saxifragaceae Micranthes tolmiei 1980 3596 eg CORD
Saxifragaceae Pectiantia breweri 1500 3500 P S-C
Saxifragaceae Saxifraga hyperborea 3000 4500 P WIDE
Valerianaceae Valeriana californica 1500 3700 P INT
Violaceae Viola adunca 0 3570 P WIDE
Violaceae Viola bakeri 900 3800 P INT
Violaceae Viola macloskeyi 609 3600 P WIDE
Violaceae Viola pinetorum subsp. 1981 3700 P END-CAL
grisea
Violaceae Viola purpurea subsp. 1400 3598 le END-CAL
mesophyta
APPENDIX 2. Annotated checklist of the flora of the Sierra Nevada with an upper elevational limit of 3300—
3499 m. Abbreviations as in Appendix 1. Species names follow Baldwin et al. (2012).
Higher level Specific or Lower Upper Growth Biogeographic
taxon infraspecific taxon elevation (m) elevation (m) form relationship
PTERIDOPHYTA
Aspleniaceae Asplenium septentrionale 2500 3350 P WIDE
Ophioglossaceae Botrychium lunaria 2300 3400 Pp WIDE
Pteridaceae Adiantum aleuticum 0 3400 | WIDE
Pteridaceae Aspidotis densa 100 3400 Pp CORD
Pteridaceae Cryptogramma acrostichoides 1400 3400 P INT
CONIFERAE
Pinaceae Pinus monticola 150 3400 T S-C
MONOCOTS
Alliaceae Allium validum 1200 3400 G CORD
Cyperaceae Carex abrupta 1200 3450 P-G S-C
Cyperaceae Carex aurea 1100 3300 P-G WIDE
Cyperaceae Carex buxbaumii 0 3300 P-G WIDE
Cyperaceae Carex davyi 1400 3300 P-G S-C
Cyperaceae Carex disperma 1100 3400 P-G WIDE
Cyperaceae Carex fissuricola 1500 3300 P-G CORD
Cyperaceae Carex fracta 250 3300 P-G S-C
Cyperaceae Carex illota 2100 3400 P-G CORD
Cyperaceae Carex integra 800 3400 P-G S-C
Cyperaceae Carex microptera 1500 3400 P-G CORD
Cyperaceae Carex pellita 60 3300 P-G WIDE
Cyperaceae Carex petasata 600 3400 P-G CORD
Cyperaceae Carex preslii 1800 3400 P-G CORD
Cyperaceae Carex simulata 0 3300 P-G CORD
Cyperaceae Carex tiogana 3100 3350 P-G S-C
Cyperaceae Carex utriculata 0 3400 P-G WIDE
Cyperaceae Carex vesicaria 0 3300 P-G WIDE
Cyperaceae Carex whitneyi 1200 3400 P-G S-C
182 MADRONO [Vol. 58
APPENDIX 2. CONTINUED.
Higher level Specific or Lower Upper Growth Biogeographic
taxon infraspecific taxon elevation (m) elevation (m) form relationship
Cyperaceae Eleocharis acicularis var. acicularis 0 3300 P-G WIDE
Cyperaceae Eleocharis acicularis var. gracilescens 0 3300 P-G WIDE
Cyperaceae Eleocharis suksdorfiana 0 3400 P-G CORD
Cyperaceae Eriophorum criniger 2000 3350 P-G S-C
Iridaceae Tris missouriensis 900 3400 G CORD
Juncaceae Juncus hemiendytus var. abjectus 1400 3400 P-G INT
Juncaceae Juncus nevadensis subsp. nevadensis 1200 3300 P-G CORD
Juncaceae Luzula parviflora var. parviflora 1000 3300 P-G WIDE
Liliaceae Lilium kelleyanum 2200 3300 G END
Orchidaceae Platanthera dilatata var. leucostachys 0 3400 G CORD
Orchidaceae Platanthera sparsiflora 100 3400 G CORD
Orchidaceae Spiranthes romanzoffiana 0 3300 P WIDE
Poaceae Agrostis humilis 1500 3350 P-G CORD
Poaceae Bromus suksdorfii 1250 3300 P-G S-C
Poaceae Calamagrostis canadensis var. 1500 3400 P-G WIDE
canadensis
Poaceae Calamagrostis canadensis vat. 1500 3400 P-G WIDE
langsdorfii
Poaceae Calamagrostis stricta subsp. inexpansa 0 3400 P-G WIDE
Poaceae Calamagrostis stricta subsp. stricta 1500 3350 P-G WIDE
Poaceae Danthonia intermedia var. intermedia 1460 3450 P-G WIDE
Poaceae Elymus trachycaulus subsp. 0 3400 P-G WIDE
trachycaulus
Poaceae Hordeum brachyantherum subsp. 0 3400 P-G CORD
brachyantherum
Poaceae Hordeum brachyantherum subsp. 0 3400 P-G END-CAL
californicum
Poaceae Melica bulbosa 0 3400 P-G CORD
Poaceae Melica stricta 1200 3350 P-G INT
Poaceae Muhlenbergia filiformis 150 3350 A CORD
Poaceae Muhlenbergia montana 1640 3420 P-G WIDE
Poaceae Stipa nevadensis 1000 3450 P-G CORD
Poaceae Stipa occidentalis subsp. californica 150 3450 P-G CORD
Poaceae Stipa occidentalis subsp. occidentalis 1200 3450 P-G CORD
Poaceae Trisetum wolfii 1740 3300 P-G CORD
EUDICOTS
Apiaceae Angelica lineariloba 1700 3300 P END-CAL
Apiaceae Ligusticum grayi 1000 3300 P CORD
Apiaceae Lomatium torreyi 1100 3300 P END
Apiaceae Perideridia parishii subsp. latifolia 2000 3400 P S-C
Asteraceae Agoseris parviflora 1400 3400 P CORD
Asteraceae Artemisia cana subsp. bolanderi 1200 3300 S S-C
Asteraceae Artemisia dracunculus 0 3400 ig INT
Asteraceae Chaenactis douglasii var. alpina 3000 3400 P-MAT CORD
Asteraceae Crepis acuminata 1000 3300 P CORD
Asteraceae Crepis intermedia 800 3300 CORD
Asteraceae Ericameria parryi var. aspera 1900 3300 SS INT
Asteraceae Erigeron barbellulatus 2100 3300 P END-CAL
Asteraceae Erigeron clokeyi var. pinzliae 2200 3400 P INT
Asteraceae Erigeron coulteri 1900 3400 P CORD
Asteraceae Erigeron elmeri 1300 3300 P END
Asteraceae Erigeron glacialis var. glacialis 1300 3400 E CORD
Asteraceae Erigeron tener 2300 3400 P-MAT CORD
Asteraceae Helenium bigelovii 0 3400 P S-C
Asteraceae Hieracium albiflorum 0 3300 P WIDE
Asteraceae Hieracium horridum 1350 3300 P S-C
Asteraceae Hulsea vestita subsp. vestita 2400 3350 P END
Asteraceae Microseris nutans 1000 3400 P CORD
Asteraceae Nothocalais alpestris 1300 3400 P S-C
Asteraceae Packera pauciflora 1800 3300 P WIDE
Asteraceae Senecio triangularis 100 3300 P CORD
Asteraceae Tetradymia canescens 1000 3400 S CORD
Asteraceae Tonestus eximius 1800 3300 P CORD
Asteraceae Wyethia mollis 900 3400 P S-C
2011) RUNDEL: SIERRA NEVADA ALPINE FLORA 183
APPENDIX 2. CONTINUED.
Higher level Specific or Lower Upper Growth Biogeographic
taxon infraspecific taxon elevation (m) elevation (m) form relationship
Boraginaceae Cryptanrtha watsonii 1250 3300 A CORD
Boraginaceae Cryptantha confertiflora 1050 3350 P INT
Boraginaceae Lappula redowskii 1300 3300 A WIDE
Boraginaceae Mertensia ciliata 1300 3380 P S-C
Boraginaceae Phacelia bicolor 700 3400 A INT
Boraginaceae Phacelia eisenii 1300 3400 A END
Boraginaceae Phacelia orogenes 2060 3400 A END
Boraginaceae Plagiobothrys hispidulus 1200 3400 A CORD
Boraginaceae Plagiobothrys torreyi var. diffusus 1200 3400 A END-CAL
Brassicaceae Barbarea orthoceras 0 3400 B/P WIDE
Brassicaceae Boechera calderi 2050 3350 P CORD
Brassicaceae Boechera davidsonii 1200 3400 PB S-C
Brassicaceae Boechera pygmaea 2600 3400 P END
Brassicaceae Boechera stricta 1800 3400 P CORD
Brassicaceae Cardamine oligosperma 50 3300 A/B CORD
Brassicaceae Descurainia californica 1700 3400 A/B CORD
Brassicaceae Draba asterophora 2600 3300 P INT
Brassicaceae Physaria occidentalis 600 3350 PE INT
Caprifoliaceae Lonicera conjugialis 140 3300 S S-C
Caprifoliaceae Symphicarpos rotundifolius var. parishii 1100 3300 S INT
Caryophyllaceae Ereomogone congesta var. 1200 3300 P INT
subfrutescens
Ericaceae Arctostaphylos patula 750 3350 S CORD
Ericaceae Arctostaphylos uva-ursi 2400 3300 S WIDE
Ericaceae Vaccinium caespitosum 0 3400 S WIDE
Ericaceae Vaccinium uliginosum subsp. 0 3400 S WIDE
occidentale
Fabaceae Astragalus bolanderi 1400 3300 P INT
Fabaceae Astragalus raventii 3400 3450 P END
Fabaceae Trifolium monanthum subsp. tenerum 1600 3300 P END
Fagaceae Chrysolepis sempervirens 700 3300 S S-C
Gentianaceae Frasera puberulenta 1700 3400 P END-CAL
Gentianaceae Gentianopsis simplex 1200 3400 P INT
Grossulariaceae Ribes inerme var. inerme 1200 3300 S CORD
Lamiaceae Monardella breweri subsp. lanceolata 0 3400 A INT
Montiaceae Claytonia megarhiza 2600 3300 P CORD
Montiaceae Lewisia leana 1300 3350 P S-C
Onagraceae Chamerion angustifolium subsp. 0 3300 P WIDE
circumvagum
Orobanchaceae Castilleja arachnoidea 1300 3300 P INT
Orobanchaceae Castilleja linariifolia 1000 3350 P INT
Orobanchaceae Castilleja peirsonii 1500 3400 P S-C
Orobanchaceae Castilleja pilosa 1200 3400 P INT
Orobanchaceae Castilleja praeterita 2200 3400 ley END
Orobanchaceae Orobanche fasciculata 0 3300 P WIDE
Papaveraceae Dicentra uniflora 1000 3300 Pp. CORD
Phrymaceae Mimulus breweri 1200 3400 A CORD
Phrymaceae Mimulus nanus var. mephiticus 1520 3445 A S-C
Phrymaceae Mimulus tilingii 1400 3400 P CORD
Plantaginaceae Penstemon caesius 1800 3400 SS END-CAL
Plantaginaceae Veronica americana 0 3300 P WIDE
Polemoniaceae Ipomopsis aggregata subsp. bridgesii 1800 3300 P END
Polemoniaceae Microsteris gracilis 0 3300 A WIDE
Polemoniaceae Navarretia breweri 1000 3300 A CORD
Polemoniaceae Polemonium occidentale subsp. 900 3300 P WIDE
occidentale
Polygonaceae Eriogonum latens 2600 3400 P END-CAL
Polygonaceae Eriogonum microthecum var. alpinum 2500 3300 SS END-CAL
Polygonaceae Eriogonum microthecum var. ambiguum 1100 3300 SS INT
Polygonaceae Eriogonum saxatile 800 3400 P-MAT INT
Polygonaceae Eriogonum spergulinum vat. 1300 3400 A INT
reddingianum
Polygonaceae Eriogonum wrightii var. subscaposum 200 3400 P-MAT INT
184
Higher level
taxon
Polygonaceae
Polygonaceae
Potamogetonaceae
Primulaceae
Ranunculaceae
Ranunculaceae
Ranunculaceae
Ranunculaceae
Rhamnaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Rosaceae
Salicaceae
Salicaceae
Scrophulariaceae
MADRONO
APPENDIX 2.
Specific or
infraspecific taxon
Polygonum polygaloides subsp.
kelloggii
Polygonum shastense
Potamogeton robbinsii
Dodecatheon alpinum
Anemone drummondii
Aquilegia formosa
Caltha leptosepala var. biflora
Delphinium nuttallianum
Ceanothus cordulatus
Amelanchier utahensis
Fragaria virginiana
Geum macrophyllum var. perincisum
Horkelia fusca subsp. parviflora
Ivesia gordonii var. alpicola
Ivesia saxosa
Rosa woodsii subsp. gratissima
Spiraea splendens
Salix jepsonii
Salix scouleriana
Limosella acaulis
CONTINUED.
Lower Upper
elevation (m) elevation (m)
1500 3300
2100 3400
1600 3300
1700 3400
1200 3350
0 3300
900 3300
300 3300
365 3365
200 3400
1200 3300
1000 3300
1400 3300
2100 3300
900 3300
800 3400
548 3400
1000 3400
l 3400
) 3300
[Vol. 58
Growth Biogeographic
form
P
NNnNNANDODIadDAWMN DI VIdIVOV
a
Q
relationship
CORD
S-C
WIDE
CORD
S-C
S-C
CORD
CORD
INT
INT
WIDE
WIDE
CORD
INT
INT
INT
S-C
S-C
CORD
WIDE
MADRONO, Vol. 58, No. 3, pp. 185-189, 2011
A NEW SPECIES OF ASTRAGALUS (FABACEAE) FROM THE WASATCH
MOUNTAINS OF UTAH
BETH LOWE CORBIN
Owyhee Field Office, Bureau of Land Management, 20 First Ave. W, Marsing, ID 83639
ecorbin@blm.gov
ABSTRACT
Astragalus kelseyae B.L. Corbin, sp. nov. is described as a new species from the central Wasatch
Mountains, where it is known from only one small occurrence in Weber County, Utah. It grows in
shale talus within the Gambel oak and bigtooth maple shrubland.
Key Words: Astragalus, Fabaceae, rare plant, Utah, Wasatch Mountains.
A new species of Astragalus is described from a
single population in the foothills of the Wasatch
Mountains in Weber County, Utah. This distinc-
tive milkvetch’s fruit shape and texture resemble
some forms of A. Jentiginosus Douglas ex
Hooker, but plants differ in having a branching,
subterranean caudex, a smaller fruit beak, and
generally larger (and fewer) flowers and fruit. Its
humistrate growth form and fruit shape resemble
A. amblytropis Barneby, but it has firmer fruits
and larger flowers. Its large pods are similar to A.
megacarpus (Nutt.) A. Gray, and flowers and
leaflets are similar to A. beckwithii Torr. & A.
Gray, but it differs from both species by having
bilocular fruit, shorter leaves, and larger flowers.
It differs significantly from each of those species
by having dolabriform (malpighian) hairs.
TAXONOMIC TREATMENT
Astragalus kelseyae B.L. Corbin, sp. nov. (Fig. 1)
— Type: USA, Utah, Weber Co., talus slope
above Ogden, TON, RIW, Sec 26, SLM,
41°14'N, 111°55'W, 1625 m (5340 ft) elevation,
28 May 2010, Beth Lowe Corbin 1292 (holo-
type: UT; isotypes: NY, CIC, UVSC, to be
distributed).
Planta similis A. /entiginosus Douglas ex
Hooker var. negundo S.L. Welsh & N.D. Atwood
et A. amblytropis Barneby in legumina biloculares
et e var. negundo in caudices subteranneis
elongatis et e ambo in legumina cartilagineis nec
chartaceis et pubescentiis dolabriformis differt.
Perennial herb from branched, subterranean,
woody caudex branches. Above-ground stems
10-20 cm long, prostrate (Fig. 2). Leaves humi-
strate, 3.05.2 cm long, 1.5—3.0 cm wide, with (5)
7-11 (13) leaflets, the terminal jointed. Leaflets
widely elliptical, 8-15 mm long, 6-12 mm wide,
tips rounded to obtuse, more or less alternate on
the rachis. Leaves and stems silvery green with
appressed dolabriform hairs about 0.5—0.8 mm
long. Stipules free, triangular, 4-5 mm _ long.
Inflorescence 2—7 flowered, congested, not much
elongating in fruit. Peduncle 13—25 mm; flowering
axis 5-10 mm. Calyx 13—16 mm long, 3.5—4.5 mm
wide, with narrow teeth 3-4 mm long; calyx
pinkish, with light and dark hairs. Petals white,
with keel tip slightly purple. Banner 22—26 mm
long, 9-11 mm wide, bent midway at about a 120°
angle. Wings 19-21 mm long, narrow, slightly
cupped. Keel 18-19 mm long. Fruit firm,
cartilaginous, fleshy, inflated, bilocular, humi-
strate, sessile, with narrow and shallow dorsal
and ventral grooves, not curved, sometimes red
mottled. Fresh pod 35-53 mm long, 18—30 mm
thick, 10-17 mm tall (dorsiventrally compressed),
with a small beak 3-6 mm long, and _ short,
dolabriform hairs. Dehiscence through the beak,
after separation.
Paratypes (topotypes): 19 May 2009 Beth Lowe
Corbin 1235 (BRY), 4 September 2011 Beth Lowe
Corbin 1523 (UTC — to be distributed).
DISTRIBUTION AND HABITAT
Astragalus kelseyae grows on the lower, west/
southwest-facing slope of the central Wasatch
Mountains, on talus openings within Quercus
gambelii Nutt. (Gambel oak) and Acer grand-
identatum Nutt. (bigtooth maple) shrublands.
The talus consists of fine-textured Ophir shale
on about 50-60% slope, at about 1625 m
elevation. This habitat is just above the old
shoreline of the Pleistocene Lake Bonneville.
Precipitation is about 51—64 cm (20-25 inches)
per year. The site 1s within the Uinta-Wasatch-
Cache National Forest. Astragalus kelseyae 1s
known only from the type locality, where about
150-200 plants were seen in a localized area of
about 0.1 ha. It was first found in 2009, and
revisited in 2010 and 2011. A popular. hiking trail
bisects the occurrence, and additional undesig-
nated trails occur within the habitat.
The geology of the Wasatch Mountains is a
complicated mix of sedimentary, metamorphic
(such as quartzite), and igneous deposits (Yonkee
and Lowe 2004). Ophir shale is a Paleozoic era
186 MADRONO
[Vol. 58
Fic. 1.
sedimentary deposit. Bands of Ophir shale and
other similar talus types occur in scattered
locations across the Wasatch Front, and addi-
tional occurrences of Astragalus kelseyae may be
found in the future.
The talus microsite is very open, with low cover
by associates, including Scutellaria angustifolia
Astragalus kelseyae habit, flower detail, pod cross section, and leaf hair detail.
Pursh subsp. micrantha Olmstead, Asclepias
asperula (Decne.) Woodson, Apocynum androsae- —
mifolium L., Epilobium canum (Greene) P.H. |
Raven subsp. garrettii (A. Nelson) P.H. Raven, |
Hedysarum boreale Nutt., Erysimum capitatum |
(Douglas) Greene, Eriogonum umbellatum Torr., |
Pseudoroegneria spicata (Pursh) A. Love, Ame- |
2011]
CORBIN: A NEW ASTRAGALUS FROM UTAH 187
FIG. 2.
Astragalus kelseyae in its shale talus habitat.
lanchier utahensis Koehne, and Phacelia hastata
Douglas ex Lehm. Although no weeds occur
directly with the Astragalus, several weedy species
occur in the vicinity, including J/satis tinctoria L.,
Linaria dalmatica (L.) Mill., Euphorbia myrsinites
L., and Bromus tectorum L., and pose a threat to
this species.
RELATIONSHIPS
Astragalus kelseyae appears to have similarities
to A. lentiginosus, A. amblytropis, A. megacarpus,
and A. beckwithii, but differs significantly from
each (S. Welsh, Brigham Young Univ., personal
communication). Its pod resembles A. lentigino-
sus var. negundo with a large, bilocular fruit, but
differs in having a less prominent beak, generally
wider fruit, fewer flowers, a branched, subterra-
nean caudex, and dolabriform hairs. It is similar
to A. amblytropis in having a subterranean
caudex, humistrate stems, leaves, and fruit, and
somewhat similar fruit shape, but differs in
having firmer pods, larger flowers, free stipules,
-and dolabriform hairs. It superficially appears
similar to A. megacarpus or A. oophorus S.
_Watson with its large pods, and A. beckwithii
-with its white flowers, but differs from these
species in the section Megacarpi by both its
bilocular fruit and dolabriform hairs. Thus, A.
kelseyae differs from each of these species by the
presence of dolabriform hairs, and other charac-
teristics as shown in Table 1. The new species’
resemblance to A. /entiginosus is likely due to
independent parallel evolution, rather than a
close relationship; the evolution of a fruit septum
(and dolabriform hairs) does not necessarily
imply shared ancestry with other species with
these characters (J. Alexander, Utah Valley
Univ., personal communication). Dolabriform
hairs have apparently arisen independently in
several sections within genus, but none of the
other species with dolabriform hairs have large,
bilocular pods, relatively few, large flowers, and
prostrate stems. The combination of characteris-
tics present in A. ke/sevyae appears unique.
Astragalus lentiginosus, with its plethora of
varieties, 1s widespread, including Weber Co.,
Utah; var. negundo is known from Box Elder Co.
(which is adjacent to the north side of Weber Co.)
and Millard and Tooele counties (southwest of
Weber Co.), Utah, so its range somewhat
overlaps that of A. kelseyae (Welsh 2007).
Astragalus amblytropis is limited to the vicinity
of Challis, in Custer and Lemhi counties, Idaho,
some 290 miles north of A. ke/seyvae (Welsh 2007).
Astragalus megacarpus 1s known from Wyoming
MORPHOLOGICAL COMPARISON OF ASTRAGALUS KELSEYAE AND SIMILAR SPECIES. Measurements (except A. kelseyae) come from Barneby 1989, Welsh on
TABLE 1.
88 MADRONO [Vol. 58
(at least 50 miles east of this occurrence), central
26 od 6 ae
Uo) of) oO :
Go ee eee rey: sone Utah (about 120 miles south), Nevada, and
Gos Sf 8S 8s Oc ; eee:
as 2 O ee 5 Colorado, while A. beckwithii occurs in Weber
a 0° EAE Aas 3 = :
= Z = 0 © 5 Seiten fi aise Co. and generally to the west in Utah, and in
fae) : Ic
a 5 ee ee eo Idaho, Nevada, Oregon, Washington, and British
S| 2S8i 582 B57 FEL] Columbia (Albee et al. 1988; Welsh et al. 2008;
— oe > iol Swe OW
aI(PEOEREES o8y Sure Shultz et al. 2010; USDA 2010). Astragalus
Gs owns — = .
Seleaceec o fF Tess, = oophorus occurs mostly in the southern Great
— qc ba — > ° . 5
se Pore ee. Wee, geet ae Basin, east into western Colorado, but is also
. far} mo
oe i? wa = recorded from northwest Utah (Box Elder and
fan} on .
; Tooele counties) (Welsh et al. 2008; USDA 2010).
Bs 9 ae
a ae Soe Fes & Thus, the location of A. kelseyae is at the east edge
= 9 ae +6 ON rs wn ey <a, 1) es
= POA. EF 5SgSVN Ee tie of the ranges of A. lentiginosus var. negundo and
= OnbK- MEAOEB S| aa serie
= BOO es Fe 7 bie Ons oo A. beckwithii, close to the northern edge of the
~x SA eae ee ee Sea gs
= Sane a 5S§ Seat R eth 2 ranges of A. megacarpus and A. oophorus, and
a 3 Po ae a7 3 = 5 cs ee disjunct from A. amblytropis.
9) Loma q) Om — = . 3
N| x eED SSR ES Ss ad eat oe Astragalus kelseyae is named in honor of Ann
a eCSaCRSSEOSSSTEH! Kelsey, plant collect for the Uni
2 aaa sé co sie a elsey, plant collection manager for the Univer-
ov ee ° ‘Oo ~ sity of Utah, with whom I’ve spent many pleasant
a = a ee f field days in the mountains and deserts of Utah.
Om (o) i) Oe 5 ‘. *
an a a0 a oe This plant is of conservation concern because
= Mey As a 6 © 4 aE:
Al Ve cag & 6 : SS 4 wo only one occurrence with few individuals is
= ee | Nese? sat x 3
3 22 wen |S . + 2 65 = known, and because potential threats exist from
~ Seed ss erigs — QO ra Oo . . 5
Bo Be +84 ott Fe 8 trails and invasive weeds.
iN¥) nv ~S & : roe . i) & va) 5
= oO mM .e PO tH CO Om
: 5 On se s6 a _&
x x 8 3 = oh vy A& > Oo7r& KEY TO SIMILAR SPECIES
cee: ome Zo S86 pee (adapted from Barneby 1989 and Welsh 2007)
On = OQ, mM WH
1. Hairs dolabriform; pods fully bilocular ...A. kelseyae
o 8 = fon) é : P : :
i oh. - — 1’. Hairs basifixed; pods unilocular or bilocular
7 Sea 6... Be oA 5 ns 2. Pods unilocular
™ > ~ sce
S 5 c os c S So 2 bby 3. Pod 1.5—3.7 cm long, valves leathery. . .
= BS %Rh560 FS pee eae ye ee eee eee A. beckwithii |
S = 5 « a..8 & 5 a = 3’. Pod (2) 3-6 (7.5) cm long, valves papery
= v . A oS Ss wee. = Oe 4. Plants subacaulescent, stems 1-—
i ees na Bas Z eG of b eC TONS yiext cen eee A, megacarpus
meas 8 7 = 38 a ce SE S 4’. Plants caulescent, stems mostly 5—
@ : o ee oe oe at 20 cmon? ta.e foe ee A. oophorus
ere “a a = 2’. Pods bilocular
eo = Z . 4 .
Sp - a a 5. Stipules connate; caudex deeply sub-
: Ces : oe ae .
= =z 6yaea E LORRAINE Ato sre ane ee A. amblytropis
> a OO Aa & bh : :
i f2 ¢ a s & = a. 5’. Stipules free; caudex superficial (near
SS Oo aie SB aT £2 SOil Surlace): = 4..°7 eee ee A. lentiginosus
aS AN a : ms 2 ia 2 do
5 ns nm Pos Ot N 0) = op
80 ne | a | “oO Q = 7 5° oa
= on On ase oO. a Res ACK
Se a ;S a0, Ff oO Bo CKNOWLEDGMENTS
aS Dn Ymadiad K Oh fice ee ; :
; A 5 fs S B ears a2 St mI = I thank Stanley Welsh (Brigham Young University,
= @ Go = FREL SOO mo BRY) for detailed information, the Latin diagnosis, and
O79 a 5 mn 2 access to the herbarium. I also thank Stephen Clark
(Weber State University, WSCO), Don Mansfield
Prt y . .
a ~< (College of Idaho, CIC), Michael Piep (Utah State
S ai i= re NaTi-=| University, UTC) and Ann Kelsey (University of Utah,
= | qaq59S FS 3 oak .
ew) SxS, 2eB = = ty ip SO ob UT) for access to collections. Jason Alexander (Utah
~_ 2 ot fw ao o & < Soh is a S Valley University) provided helpful comments and
S c 5 = ; HAE 5 2 o = oo ffi 7 insights. Dan Scott produced the illustration, which was
s NJBPU Sm we gunk. 6 5- funded by the Intermountain Region of the U.S. Forest
= S2H2 sees aud adaat 5 ear
= SERB F RSA SBSNS EL KMS Service, thanks to Teresa Prendusi, who also reviewed a
co) = G c - s : 7
= Coe c Z Sayer draft of this article, as did Don Mansfield. This paper was _|
ao) also improved by reviews from Stanley Welsh and Leila |
= Shultz; however, any mistakes are all mine.
lan} (cD) > 3
oc} 18 =
S| 88 : 3
al | 9 5 on LITERATURE CITED
ia) Nn 0) =— @
= S| 86 = 2 ogee
= =|3 0 S 2 = 3 ALBEE, B. J., L. M. SHULTZ, AND S. GOODRICH. 1988. Atlas
D UIAn [Ly [Ly = 4
of the vascular plants of Utah. Utah Museum of |
2011]
Natural History, University of Utah, Salt Lake City,
Li
BARNEBY, R. C. 1989. Fabaceae. Pp. 12-267 in A.
Cronquist, A. H. Holmgren, N. H. Holmgren, J. L.
Reveal, and P. K. Holmgren (eds.), Intermountain
flora, Vol. 3, pt. B: Fabales. The New York
Botanical Garden, Bronx, NY.
SHULTZ, L. M., R. D. RAMSEY, W. LINDQUIST, AND C.
GARRARD. 2010. Digital atlas of the vascular
plants of Utah. Utah State University, Logan,
UT. Website http://earth.gis.usu.edu/plants/ [ac-
cessed 01 September 2011].
CORBIN: A NEW ASTRAGALUS FROM UTAH 189
USDA, NRCS. 2010, The PLANTS Database. Na-
tional Plant Data Center, Baton Rouge, LA.W-
ebsite http://plants.used.gov [accessed 01 Oct 2010].
WELSH, S. L. 2007. North American species of
Astragalus Linnaeus (Leguminosae): a taxonomic
revision. Brigham Young University, Provo, UT.
, N. D. ATwoop, S. GOODRICH, AND L. C.
HIGGINS. 2008. A Utah flora, fourth edition,
revised. Brigham Young University, Provo, UT.
YONKEE, A. AND M. Lowe. 2004. Geologic map of the
Ogden 7.5-minute quadrangle, Weber and Davis Coun-
ties, Utah. Utah Geological Survey, Salt Lake City, UT.
MADRONO, Vol. 58, No. 3, pp. 190-198, 2011
GRIMMIA VAGINULATA, (BRYOPSIDA, GRIMMIACEAE) A NEW SPECIES
FROM THE CENTRAL COAST OF CALIFORNIA
KENNETH KELLMAN
California Academy of Sciences, 55 Music Concourse Drive, San Francisco, CA 94118
kkellman@sbcglobal.net
ABSTRACT
A new California endemic species, Grimmia vaginulata, is described and illustrated. It is
characterized by its very small size, julaceous habit, immersed capsule on a straight, centrally
inserted seta, very large annulus, keeled unistratose leaves, autoicous sexuality, and a large cylindrical
ochrea atop the vaginula that sheathes the seta to the base of the capsule. The differentiation and
ecology of the new species is discussed.
Key Words: California, endemic moss, Grimmia vaginulata, Grimmiaceae, new species, ochrea,
vaginula.
In the summer of 2008, prior to the annual
meeting of the American Bryological and Li-
chenological Society, Dale Vitt asked to see
Orthotrichum kellmanii D. H. Norris, Shevock
& Goffinet in the field. During that short trip
Dale noticed tiny little plants growing on the
same sandstone boulders that supported the
Orthotrichum. These plants were so_ scattered
and insignificant that they appeared to be
immature stems of a small Grimmia, and not
worth collecting. That impression proved incor-
rect. Upon microscopic examination, it was
immediately obvious that these plants represent-
ed a remarkable plant that was unlike any other
Grimmia in North America.
Chief among the distinguishing characteristics
was a transparent extension of the vaginula that
extended to the base of the capsule. This
extension, known as an ochrea, 1s not present in
any North American Grimmia, and it is not
mentioned in any North and Central American
Grimmia literature (Flowers 1973; Crum and
Anderson 1981; Crum 1994; Hastings and
Greven 2007; Munoz 1999; Munoz and Allen
2002). To the casual student of the genus, this
novel character suggested the possibility of a new
genus. That hypothesis was supported by the
scattered growth habit. A wider examination of
Grimmia was undertaken resulting in the expan-
sion of the North American parameters of the
genus.
TAXONOMY
Grimmia vaginulata Kellman, sp. nov. (Figs. 1A,
IC, 2, 3, 4, 5)—Type: USA, California, Santa
Cruz Co., Big Basin Redwoods State Park, on
dry vertical walls of calcareous sandstone
boulders eroded from the sandstone bedrock
in open chaparral, above the Basin Trail and
below China Grade Road ca. 1.5 mi. beyond
the northern intersection with SR 236, elev.
685 m, 37°12'40"N, 122°12'42’”W, 13 Aug 2008.
Kellman, Vitt, & Shevock 5869 (holotype: CAS).
Species Grimmia capillata De Notaris affinis,
sed differt ab statura breviore, foliis statu madido
vel sicco persistenter julaceis, ochrea cylindracea
longiore usque ad prope basem capsulae exten-
denti differt.
Plants up to 2 mm tall, scattered to very loosely
tufted (Fig. 5), simple, or with a short, tightly
appressed branch; green when young, tan or
white in age; julaceous wet or dry. Lower and
perigonial leaves short ovate, 1:1, muticous,
increasing in length upwards. Upper stem and
perichaetial leaves (Fig. 2B) obovate or elliptical,
soft when moist, weakly keeled concave, uni-
stratose throughout, with at least some portion of
the leaf margin hyaline, most common at the base
and often extending to midleaf or slightly
beyond; 1—1.65 mm long X 0.4—-0.75 mm wide
(without the awn); the keel is even less pro-
nounced in transverse section, appearing more
convex, even tubular, especially on non-sporo-
phytic plants (Fig. 4); apex acute or occasionally
slightly acuminate. Basal juxtacostal cells short
rectangular with thin to moderately thick,
straight; relatively uniform across the base, but
outer basal cells often somewhat narrower in | to
3 marginal rows. Distal and medial cells decreas-
ing in length gradually from the base to the apex,
1—1.5(2.5):1, 12-16(26) um long X 7—13 um wide;
rectangular, triangular, or irregularly polygonal,
thick-walled, often with some portion of the
lumen rounded creating small trigones, not or
weakly sinuose; in transverse section plane to
slightly bulging. Margins weakly recurved on one
or both sides; unistratose. Costa narrow, to 40 um
wide at the base, broadening toward the apex;
excurrent in a hyaline, weakly toothed awn
shorter than the lamina, decurrent at most 1—2
cells down the margin, and those often projecting
2011] KELLMAN: A NEW SPECIES OF GRIMMIA 19]
Fic. 1. A. Grimmia vaginulata with leaves stripped exposing the sporophyte. (O) ochrea (V) entire vaginula. Scale
bar = 500 um (from the type); B. G. capillata with leaves stripped exposing the sporophyte. (O) ochrea. Scale bar =
500 um (Handel-Mazetti 1778, FH: syntype of G. mesopotamica); C. Detail of typical upper leaf apex of G.
vaginulata Scale bar = 80 um; D. Detail of typical upper leaf apex of G. capillata Scale bar = 175 um.
as short, blunt teeth; costa in transverse section at
midleaf with two homogenous rows of cells, 2
cells wide adaxially. Gonioautoicous or cladau-
toicous. Calyptra irregularly crenate at the base,
conical or campanulate, hyaline at the extreme
-apex and at the base, naked and smooth, just
covering the operculum. Vaginula (Fig. 1A),
measured with ochrea, 0.5—0.6 mm long, epider-
mal cells irregular, about 2—3:1, with very thin,
straight walls; distally with a cylindrical ochrea
surrounding but not connate to the seta and
flaring just below the base of the capsule, 340—
430 um long. Seta straight, attached to the center
of the capsule, 0.5 mm long. Capsule immersed
192 MADRONO
G. capilla ta
G. vaginulata
Fic, 2.
A. Sporophytic plants of Grimmia vaginulata and G. capillata showing size difference. Marks are | mm
apart; B. G. vaginulata, two upper and perichaetial leaves. Scale bar = 500 um.
with only the operculum exposed, irregularly
wrinkled when dry, slightly wrinkled when wet,
obloid, 0.9 mm long X 0.6 mm wide, abruptly
contracted to the seta. Annulus (Fig. 3) of 3 rows
of differentiated, transparent, thick-walled en-
larged and elongated cells; remaining on the urn
after dehiscence, but gradually falling off in
sections. Operculum mammilose to low conical,
crenulate to erose at the base. Exothecial cells
irregularly rectangular to hexagonal, thin-walled,
almost transparent when mature, easily revealing
the stalked theca within. Stomata present. Peri-
stome (Fig. 3) of 16 orange-red cribrose-dissected
teeth, irregularly divided nearly to the base into
3-4 strongly spiculose filaments, ca. 185 um long.
Spores smooth, 10—13 um in diameter.
The specific epithet refers to the persistent
cylindrical ochrea atop the vaginula, which
extends nearly to the base of the capsule.
IDENTIFICATION AND
TAXONOMIC RELATIONSHIPS
Grimmia vaginulata can be distinguished from
all other North American Grimmia by the
following combination of characters: very small
plants that are julaceous wet or dry, upper leaves
with hyaline margins, dissected peristome, and a
conspicuous and persistent ochrea sheathing the
entire seta to just below the base of the capsule.
Grimmia vaginulata resembles G. anodon Bruch
& Schimp. and G. plagiopodia Hedw. in leaf
shape. However, G. vaginulata remains julaceous
wet or dry, and is a much smaller and narrower
plant. The straight, centrally inserted seta of G.
vaginulata contrasts with the sigmoid, eccentri-
cally inserted seta that characterizes both G.
anodon and G. plagiopodia. The extremely dis-
sected peristome resembles G. orbicularis Bruch
and G. moxleyi R.S. Williams. Grimmia orbicu-
laris shares the unistratose margins, and the
autoicous sexuality with G. vaginulata, but the
awns of G. orbicularis are evenly distributed
along the stem. Grimmia moxleyi has awns
restricted to the upper leaves like G. vaginulata,
but the leaf margins are bistratose. Furthermore,
both G. orbicularis and G. moxleyi have exserted
capsules on arcuate setae. Again, both are much
larger than G. vaginulata.
Both gametophytically, and sporophytically, G.
vaginulata appears most closely related to G.
capillata De Not., a species scattered around the
Mediterranean Sea (Greven 2003). They share the
keeled, entirely unistratose lamina with the
margins at least somewhat recurved on one or >
two sides, a costa that broadens in the distal half |
of the leaf, perichaetial leaves with proximal
hyaline areas, and autoicous sexuality. Sporophy-
tically, the two taxa are very close. Lastly, both
prefer calcareous substrates. Again, G. vaginulata
2011]
FIG. 3.
is amuch smaller plant that stays tightly julaceous
wet or dry. My examination of 16 specimens of G.
capillata and G. mesopotamica Schiffn. (a syno-
nym of G. capillata fide Munoz and Pando 2000;
Greven 2003) from FU, MUB, and NY show
plant size in G. capillata averages around 7 mm,
dwarfing G. vaginulata. (Fig. 2A) The dry leaves
of G. capillata are erect to erect-patent and are
usually strongly keeled. Thus the costae project
from the rest of the leaves giving the dry plants a
more tumid and textured appearance. When
moist, the leaves of G. capillata are at least
erect-patent, and are relatively easy to dissect
from the stem. The tiny and tightly appressed
leaves of G. vaginulata are very difficult to strip
for examination without tearing. Leaf size is
another point of separation, with the upper stem
leaves of G. capillata measuring 2—-3 mm long
(Cortini Pedrotti 2001; Ignatova and Munoz
2004), while the much smaller G. vaginulata has
leaves with a maximum length of 1.65 mm. In G.
capillata, the more or less rounded apex (Fig. 1D)
is the most commonly hyaline portion of the leaf,
while in G. vaginulata, the margins are hyaline,
and the apex is acute. There is some small portion
at the base of the awn that is hyaline, but at most
it is one to two cells down the margin (Fig. 1C).
KELLMAN: A NEW SPECIES OF GRIMMIA 193
Detail of annulus and peristome of Grimmia vaginulata. Scale bar = 120 um.
Fic. 4. Drawings of leaf cross sections, Grimmia
vaginulata; a) leaf from sterile plant taken at midleaf; b,
c, d) upper leaf from sporophytic plant. Scale bar =
100 um.
194
MADRONO
FIG. 5.
Although evidence is admittedly sparse, with the
only known population being the type, the growth
habit of G. vaginulata adds another basis for
separation. In G. vaginulata, the plants are at best
loosely associated (Fig. 5), not forming a contin-
uous turf or clump. In G. capillata, the plants
form dense green mats of sterile plants inter-
spersed with a few fertile plants (Cortini Pedrotti
2001; Greven 2003; Heyn and Herrnstadt 2004;
Ignatova and Munoz 2004). The base of the
capsule in G. capillata is less abruptly contracted
to the seta than is seen in G. vaginulata. Lastly, in
G. capillata the ochrea, while pronounced, extends
only half way up the seta, not to the base of the
capsule as in G. vaginulata. (Figs. 1A, B)
Grimmia pseudoanodon Deguchi, described in
1987 from Peru (Deguchi 1987), is another small
autoicous plant (ca 5 mm tall) with immersed
sporophytes with a centrally attached, straight
seta, and like G. vaginulata, the base of its capsule
is abruptly contracted. Examination of an isotype
from NY, showed several important differences
from G. vaginulata. First, G. pseudoanodon grows
in cushions, contrasting the scattered habit of G.
Grimmia vaginulata. Photograph of growth habit on sandstone.
vaginulata. G. pseudoanodon has no peristome, its
cauline leaves are not reduced down the stem,
being linear to lanceolate and bearing awns to the
base of the plant. They are patent to spreading
when moist, and are easy to separate from the
stem. Similar to G. capillata, the apex of the leaf»
is hyaline for 15-20 cells, and the awn is often
\
decurrent down the margins of the leaf. Most |
leaves of G. pseudoanodon show bistratose |
margins. The vaginula of G. pseudoanodon 1s
extended, demonstrated by the aborted archego- —
nia scattered throughout its length, however, the
ochrea is at most represented by a few hyaline
flaps at the apex of the vaginula, far from the
tube sheathing the seta of G. vaginulata. Inter- |
estingly, slight pressure expels the seta from the |
end of the vaginula, leaving a conical stub at the |
base of the seta.
GENERIC TAXONOMY
It is clear that this new plant belongs in the
Grimmiaceae. The only other familial possibility
is Ptychomitriaceae, and that is ruled out by the
2011]
unlobed calyptra. Within Grimmuiuaceae, other
genera can easily be dismissed as well. The large
annulus, and the operculum falling independent
of the columnella deny placement in Schistidium.
Although the cribrose peristome resembles those
of both Coscinodon and Jaffueliobryum, and the
autoicous sexuality, unistratose lamina and large
annulus further support placement in Jaffuelio-
bryum, both genera require a pleated and
somewhat sheathing calyptra (Churchill 1987;
Hastings 2007; Spence 2007). The calyptra of G.
vaginulata is so short that it does not even split or
tear during capsule development, sitting instead
just atop the operculum. Racomitrium (sensu lato)
is ruled out by the lack of sinuosity in the cell
walls of the lamina or the vaginula (Deguchi
1978).
Not only can we suggest placement in Grimmia
by elimination, but the dramatic characters
displayed by G. vaginulata fit well within the
limits of variation of plants already in that genus.
Deguchi (1978) describes the “‘Affinis type”
annulus as “‘well differentiated, composed of (2)
3-4 rows of cells, which are thick walled, but
transparent, and becoming increasingly larger
from the lower to the upper rows. ... Upper rows
of cells of the annulus are also removed when the
lid falls, but their lower rows usually remain
attached to the orifice of the urn, disjoining little
by little in the course of time.” This is a perfect
description of the annulus in G. vaginulata. He
attributes this “‘Affinis type’ annulus to “G.
affinis, G. anomala, G. apiculata, G. brachydic-
tyon, G. curvata, G. olympica, and G. pilifera.”
Munoz (1999) assigns this annulus type to “G.
involucrata, G. longirostris, G. poecilostoma, and
G. trichophylla’ in the introductory section, and
14 other taxa in the species descriptions.
The ochrea is a structure that is poorly
understood and has received little taxonomic
discussion. Deguchi (1978) makes it clear that the
ochrea is only a section of the vaginula, but
Magill (1990) defines ochrea as “‘vaginula, or
upper part of vaginula’’. The haploid vaginula
includes cells originating from both the archego-
nium and the upper part of the stem (Deguchi
1978). Deguchi’s proof of this contention is the
“occurrence of aborted archegonia at the basal
part of the vaginule” (Fig. 1A).
Maier (2002) discusses the vaginula and the
ochrea and describes Grimmia donniana Sm. ex
Spruce with ‘“‘ochrea broad’’, with most other
Species with ochrea ‘“‘short” or “‘small’”’. Some
species such as G. orbicularis (a close relative of
the aforementioned G. moxleyi) she describes
with ‘“‘vaginula 0.8 mm, with ochrea.” Unfortu-
nately, she does not provide illustrations to
inform that aspect of her excellent descriptions,
nor does she describe how she measures the
vaginula. Delgadillo (Universidad Nacional
Autonoma de Mexico, personal communication)
KELLMAN: A NEW SPECIES OF GRIMMIA Is
wrote “I believe the ochrea is a fairly common
feature of the neotropical species of Grimmia. It
can be demonstrated in G. americana, G.
donniana and G. elongata. It is particularly
evident in G. anodon.”’ But more convincing
examples can be found in the persistent ochreae
of G. capillata and G. involucrata Cardot.
Being clear then that this new plant properly
belongs in Grimmia, it is necessary to at least
explore its relationships within the genus. The
history of the genus Grimmia, s./, 1s a tortured and
complex story of confused concepts, alternating
periods of splitting and lumping, and regional
authors describing numerous plants that have
been later synonymized in worldwide treatments
(Munoz and Pando 2000; Greven 2003). Deguchi
(1978) and Ochyra et al. (2003) both provide a
good summary of this nomenclaturally difficult
subject. Although Hernandez-Maqueda et al.
(2008) casts some doubt on Coscinodon, most
modern authorities agree on the separation of
Jaffueliobryum, Schistidium, and Coscinodon from
Grimmia, but there is disagreement whether
Grimmia itself requires further subdivision.
Ochyra et al. (2003) proposed to split Grimmia
into five genera, with each genus corresponding in
concept with the subgenera reluctantly suggested
by Hastings and Greven (2007). Both systems are
rooted in the work of Hagen (1909) and are
summarized in Table 1.
Grimmia vaginulata can be ruled out of
Gasterogrimmia based on the straight, centrally
attached seta. Litoneuron requires 2-stratose,
concave leaves with the costa not prominently
projecting abaxially: Grimmia vaginulata meets
none of these criteria. An expanded Guembelia
that included unistratose leaves would fit all
characters of G. vaginulata, but if one were to
accept the Ochyra et al. (2003) concept of
Dryptodon, which includes plants with short
straight setae and plicate or wrinkled capsules,
then G. vaginulata would belong there. In fact,
Ochyra et al. (2003) places G. capillata in
Dryptodon.
The difficulty is compounded, however, by
statements by various authors suggesting that
pairs of plants within Grimmia s./. are closely
related, but subsequent authors treat each
member of the pair in separate subgenera or
genera. For example, G. capillata is often paired
with G. crinita Brid. (Greven 2003; Ignatova and
Munoz 2004) and in fact De Notaris (1836, 1838),
who first described G. capillata in 1836, reduced it
to a variety of G. crinita in 1838, and Maier
(2010) reduced G. capillata into synonymy with
G. crinita. In this pair, G. crinita is universally
placed in Gasterogrimmia (Grimmia sensu Ochyra
et al. 2003) but as stated above G. capillata is
placed in Dryptodon. These cross subgeneric or
generic affinities, as well as the taxa that defy
placement within any system, illustrate the
196
TABLE 1.
Hastings and Greven (2007)
Gasterogrimmia
Guembelia
Litoneuron
Rhabdogrimmia
MADRONO
Ochyra et al. (2003)
Grimmia
Orthogrimmia
Guembelia
Dryptodon
[Vol. 58
COMPARISON OF TAXONOMIC TREATMENTS OF GRIMMIA.
Description
Capsules immersed, smooth, seta
sigmoid and eccentrically attached
to the capsule, leaves 1- or 2-stratose
that are concave or concave-keeled.
Capsules immersed to exserted,
smooth, seta straight and centrally
attached to the capsule, leaves
mostly 2-stratose and keeled.
Capsules exserted or emergent,
smooth, seta straight and centrally
attached to the capsule, leaves 2-
stratose that are concave with the
costa not prominent dorsally.
Capsules emergent to exserted, ribbed,
seta arcuate and centrally attached
to the capsule, leaves mostly 1-
stratose (2-stratose at the margins)
and keeled.
complexity of Grimmia s./. It is likely that this will
only be sorted out by a massive study combining
worldwide genetic and morphological data.
The very dramatic and persistent ochrea in G.
vaginulata opens the question of the importance
of the ochrea in Grimmia taxonomy. Only Maier
(2002) and Deguchi (1978) discuss this character.
Deguchi briefly mentions the ochrea in his
introduction, and then does not use it in his
species descriptions. It is ignored in Crum (1994),
Munoz (1999), Cortini Pedrotti (2001), Greven
(2003), Ignatova and Munoz (2004), and Hastings
and Greven (2007). Maier (2002) lists G. donniana
with ochrea “‘broad’’, and G. elatior with ochrea
“distinct”. Many other species were described
with ochrea “‘small”’ or the character was ignored
altogether. A quick examination of five specimens
in CAS annotated by R. Hastings as G. donniana
displayed three with no ochrea, one very juvenile
capsule with an ochrea already disintegrating, and
one specimen with a persistent ochrea about as
long as broad at the base of a mature sporophyte.
Thus, it seems that this character by itself does not
seem to be reliable enough to be useful for
separating G. donniana, one of the few taxa that
Maier thought had a distinctive ochrea; it is likely,
therefore, that the character is not useful beyond
the G. capillata—vaginulata group. Nonetheless,
further study of such taxa as G. involucrata may
show that the ochrea could be an additional
character useful for identification.
ECOLOGY AND DISTRIBUTION
Grimmia vaginulata has thus far only been
found on vertical or underhanging surfaces on
calcareous sandstone boulders that have eroded
out of less calcareous sandstone bedrock of the
Butano Formation. These rocks are scattered
through a variably dense chaparral comprised of
Adenostoma fasciculatum Hook. & Arn., Arcto-
staphylos tomentosa (Pursh.) Lindl., Ceanothus
cuneatus (Hook.) Nutt., C. papillosus Torr. & A.
Gray, and Eriodictyon californicum Hook. &
Arn.. The climate is Mediterranean with moder-
ate but rainy winters, and hot rainless summers
(Kellman 2003). The elevation is slightly below
700 meters, placing the site above all but the
thickest summer maritime fog. The lack of
summer fog keeps these sites very hot and dry
until the winter storms arrive.
One of the more peculiar aspects of this very
interesting plant is the substrate. With the
exception of one recent collection from marble
rock (Kellman, Shevock & Lodder 6133 [CAS]) the
boulders are also the sole known substrate of ©
Orthotrichum kellmanii, another rare coastal
California endemic (Norris et al. 2004). The
bedrock was deposited in the Upper Eocene, and
is part of the Butano Sandstone Formation
(Brabb 1989), but the geologic history of the
boulders is not known. Presumably, the rocks
were formed and eroded into their rounded shape
some time prior to the Eocene deposits that |
formed the bedrock. In Santa Cruz Co., the |
Butano Formation is exposed along the western
slopes of the highest mountains on the eastern ©
side of the county (Brabb 1989), but the boulders
are only present in a small patch in Big Basin |
Redwoods State Park. There is also a small field |
of the boulders in southern San Mateo Co., also |
in Butano State Park. Recently, this same |
formation with exposed boulders has been |
found in Monterey Co. within the Ventana |
Wilderness of the Los Padres National Forest. |
Orthotrichum kellmanii was collected here al- |
though much of the area remains to be surveyed.
The new population greatly expanded its range |
to the south. Although access to these sandstone |
outcrops was difficult due to hiking through
2011]
dense stands of chaparral, the area subsequently
burned in the summer of 2008. This area 1s now
among the highest priority sites to survey for G.
vaginulata—and to document the extent of Ortho-
trichum kellmanii in this area.
Associated bryophytes growing on and around
the boulders include Amphidium californicum
(Hampe ex Mill. Hal.) Broth., Antitrichia cali-
fornica Sull. in Lesq., Gemmabryum californicum
(Sull.) Spence, Didymodon vinealis (Brid.) R.H.
Zander, Tortula muralis Hedw., Grimmia torenti
Hastings, G. pulvinata (Hedw.) J. E. Smith,
Gymnostomum calcareum Nees & Hornsch., G.
viridulum Brid., Orthotrichum kellmanii, and
Cephaloziella divaricata (Sm.) Warnst.
CONSERVATION
At the type location, G. vaginulata has been
found on only four out of several hundred
boulders, and these fit within a circle of ten
meters. It is imperative that no further collections
should be made until more populations are
found. Additional surveys need to be conducted
throughout the highly restricted habitat within
the Butano Formation within State Park Lands
and a federally designated wilderness area. Even
if additional occurrences are discovered, Grimmia
vaginulata will almost certainly remain a very
narrowly restricted endemic that could be ad-
versely impacted by stochastic events.
Even though the chaparral is a fire adapted
ecosystem (Schoenherr 1992), the extreme rarity
of G. vaginulata leaves open the possibility of fire
killing the entire known population. In March of
2009, the author visited the Bonny Doon Ecolog-
ical Preserve in Santa Cruz Co. Nine months
before, in June of 2008, a very hot fire raced
through the preserve, where sandstone rocks are
scattered through a chaparral very similar to that
found at the type location of G. vaginulata. Even
in places where the brush was minimal, virtually
all bryophytes were killed on the rocks, especially
those bryophytes growing on the walls and
underhanging surfaces of the rocks. A few mosses
and liverworts survived on the tops of a few rocks.
Apparently the heat of the fire was trapped under
the rocks and then bathed the vertical surfaces,
killing all plant life. It is conceivable that thick
cushions, or gemmiform plants with densely
imbricate leaves could insulate at least some part
of a dry moss plant from the heat, but the loose
colonies and cylindrical plant form of G. vaginu-
lata offers no such protection.
To date, G. vaginulata is known only from the
type and in general, taxonomic novelties should
not be based on a single specimen, particularly in
variable genera such as Grimmia. However, this
plant’s combination of characters, unique to
Grimmia in North America, along with the
conservation implications of its rarity, demand a
KELLMAN: A NEW SPECIES OF GRIMMIA
LOT
full description. Of course, it is always possible
that further collections from new locations could
alter the concept of G. vaginulata. Of all the
characters discussed above, the most likely
features to change would be the size and the
growth form. It is possible that other populations
of G.vaginulata could be composed of larger
plants. Arguing against that possibility is the fact
that both sterile and fruiting plants exhibit the
same size and growth form in the type population.
It seems more likely that G. vaginulata could be
found growing in turfs or small cushions.
However, turf and cushion formation require
the survival of the originating stems. There are no
subapical innovations in the type population
which suggest that this species may be short lived
and that individuals die after sexual reproduction.
ACKNOWLEDGMENTS
I am indebted to Jim Shevock and Dan Norris for
teaching me how to do the research necessary to write a
paper of this type. They may or may not have been
aware that they were teaching me these methods, but an
interested student learns as much by observation as by
lecture and discussion. I honor their example. I would
also like to thank Roxanne Hastings for teaching me the
basic understanding of Grimmia, and for a thorough
review of this paper resulting in many positive changes. I
thank Henk Greven for his review of this paper, and the
suggestion to compare Grimmia vaginulata with G.
pseudoanodon. David Toren deserves credit for pointing
out the illustration of Grimmia mesopotamica in the
Bryophyte Flora of Israel that put me onto the
similarities of G. vaginulata and G. capillata. Thanks
also to Eva Maier, and Claudio Delgadillo, for their
insights on the vaginula and ochrea. I also appreciate
Claudio Delgadillo and Jesus Munoz for their helpful
comments on an earlier version of this paper. Steve
Lodder was invaluable in providing the beautiful
photographs. I thank Patricia Eckel for providing the
Latin description. I would like to thank the curators of
MUB, FH, and NY for the loan of specimens of
Grimmia capillata, Grimmia mesopotamica, and Grimmia
pseudoanodon, and Deb Trock at CAS who arranged for
the loans.
LITERATURE CITED
BRABB, E. E. 1989. Geologic map of Santa Cruz
County, California. Miscellaneous Investigation
Series. U.S. Geological Survey, Denver, CO.
CHURCHILL, S. P. 1987. Systematic and biogeography
of Jaffueliobryum (Grimmiaceae). Memoirs of the
New York Botanical Garden 45:691—708.
CORTINI PEDROTTI, C. 2001. Flora dei muschi d’Italia:
Sphagnopsida, Andreaeopsida, Bryopsida (1
Parte). Antonio Delfino Editore, Rome, Italy.
CruM, H. 1994. Grimmiales. 386-415. in A. J. Sharp,
H. Crum, and P. M. Eckel (eds.), The moss flora of
Mexico. Memoirs of the New York Botanical
Garden 69:1—1113.
AND L. ANDERSON. 1981. Mosses of Eastern
North America. Columbia University Press, New
York, NY.
DE NOTARIS, G. 1836. Memorie della Reale Accademia
delle Scienze di Torino 39:248.
198 MADRONO
. 1838. Syllabus Muscorum in Italia et in Insulis
Circumstantibus Hucusque Cognitorum, Turin,
Italy.
DEGUCHI, H. 1978. A revision of the genera Grimmia,
Schistidium and Coscinodon (Musci) of Japan.
Journal of Science of the Hiroshima University
Series B, Div. 2 (Botany) 16:121—256.
. 1987. Studies on some Peruvian species of the
Grimmiaceae. Pp. 19-74 in H. Inoue (ed.), Studies
on cryptogams in Southern Peru. Tokai University
Press, Tokyo, Japan.
FLOWERS, S. 1973. Mosses: Utah and the West.
Brigham Young University Press, Provo, UT.
GREVEN, H. C. 2003. Grimmias of the world. Bachuys,
Leyden, The Netherlands.
HAGEN, I. 1909. Forarbejder tilen Norsk lovmosflora.
IX. Grimmiaceae. Det Kongelige Norske Videns-
kabers Selskabs Skrifter 1909(5):1—94.
HASTINGS, R. I. 2007. Coscinodon. Pp. 258-262 in
Flora of North America Editorial Committee
(eds.), Flora of North America North of Mexico,
Vol. 27. Oxford University Press, New York, NY
, H. C. GREVEN. 2007. Grimmia. Pp. 225-258 in
Flora of North America Editorial Committee
(eds.), Flora of North America North of Mexico,
Vol. 27. Oxford University Press, New York, NY.
HERNANDEZ-MAQUEDA, R., D. QUANDT, O. WERNER,
AND J. MUNOZ. 2008. Phylogeny and classification
of the Grimmiaceae/Ptychomitriaceae complex
(Bryophyta) inferred from cpDNA. Molecular
Phylogenetics and Evolution 46:863—877.
HEYN, C. C. AND I. HERRNSTADT. 2004. The
bryophyte flora of Israel and adjacent regions.
The Israel Academy of Sciences and Humanities,
Jerusalem, Israel.
IGNATOVA, E. A. AND J. MUNOZ. 2004. The genus
Grimmia in Russia. Arctoa 13:101—182.
[Vol. 58
KELLMAN, K. M. 2003. A catalog of the mosses of
Santa Cruz County, California. Madrofio
50:61-82.
MAGILL, R. E. 1990. Glossarium Polyglottum Bryolo-
giae, a multilingual glossary for bryology. Missouri
Botanical Garden, Saint Louis, MO.
MAIER, E. 2002. The genus Grimmia (Musci, Grimmia-
ceae) in the Himalaya. Candollea 57:143—238.
. 2010. The genus Grimmia Hedw. (Grimmia-
ceae, Bryophyta) - a morphological-anatomical
study. Boissiera 63:1—377.
MuNoz, J. 1999. A revision of Grimmia (Musci,
Grimmiaceae) in the Americas. 1: Latin America.
Annals of the Missouri Botanical Garden
86:118—191.
AND F. PANDO. 2000. A world synopsis of the
genus Grimmia (Musci, Grimmiaceae). Missouri
Botanical Garden Press, St. Louis, MO.
AND B. ALLEN. 2002. Grimmia. Pp. 221—231 in
B. Allen (ed.), Moss flora of Central America,
Part 2. Encalyptaceae—Orthotrichaceae. Missouri
Botanical Garden Press, Saint Louis, MO.
Norris, D. H., J. R. SHEVOCK, AND B. GOFFINET.
2004. Orthotrichum kellmanii (Bryopsida, Ortho-
trichaceae), a remarkable new species from the
central coast of California. The Bryologist
107:209-214. _
OCHYRA, R., J. ZARNOWIEC, AND H. BEDNAREK-
OCHYRA. 2003. Census catalog of Polish mosses.
Polish Academy of Sciences, Krakow, Poland.
SCHOENHERR, A. A. 1992. A natural history of
California. University of California Press, Berkeley
and Los Angeles, CA.
SPENCE, J. R. 2007. Jaffueliobryum. Pp. 262-264 in
Flora of North America Editorial Committee
(eds.), Flora of North America North of Mexico,
Vol. 27. Oxford University Press, New York, NY.
MADRONO, Vol. 58, No. 3, pp. 199-200, 2011
REVIEW
Introduction to California Chaparral. By RONALD
D. QUINN AND STERLING C. KEELEY. 2006.
University of California Press, Berkeley, CA.
344 pp. ISBN 9780520219731, $55.00, hardcover;
ISBN 9780520245662, $21.95, paperback.
In Introduction to California Chaparral, au-
thors Ronald D. Quinn and Sterling C. Keeley
condense the beauty and diversity of California’s
most prominent habitat into a beginners guide
suitable for curling up with on the couch or toting
along on a hike. The diminutive tome attempts to
familiarize the casual observer not only with that
superficial ““‘bluish-green blanket gently covering
the hills,’ but also with the dynamic interplay of
organisms and underlying ecological processes
beneath that oft go largely unseen. It is a monu-
mental task to be sure. To accomplish this task,
the authors guide us through the climatic forces
that shape the system; the dominant plant, animal
and insect species that inhabit it; and other forces
such as fire, flooding and urban encroachment
that affect it. In addition to describing the
chaparral world that modern day adventurers
might observe, Quinn and Keeley take us briefly
through the history of the chaparral to provide a
backdrop of the forces that created such diversity.
In distilling such broad and complex subject
matter, the authors only stumble in perhaps the
most complex of chaparral arenas: wildfire.
Dedicating roughly 1/6 of the book specifically
to fire and regularly referring to it throughout the
rest, the authors succeed in divulging most of the
issues surrounding fire in chaparral. However,
their attempt at presenting those issues together
in a cohesive and coherent whole falls short and
proves dangerously misleading. Notwithstanding,
the authors bring hundreds of years of scientific
discovery, field observation and a love of Cali-
fornia’s defining ecosystem into an accessible and
enjoyable guide for the masses.
This attractive collection consists of 302 pages
of text, color photos, black and white diagrams,
plus a glossary and reference section for those
who wish to dig deeper. All this is printed on
thick, sturdy pages that will no doubt survive the
jostles of the many hikes it is sure to inspire.
Quinn and Keeley, Professors of biological
science and botany, respectively, draw on a wide
range of sources and personal research in com-
piling this guide. They use these sources to paint a
colorful mural of wood-rat nests, heat-seeking
fire beetles, iridescent hummingbirds and an
impressive array of plants that have managed to
survive eons of yearly drought and occasional
wildfire. The descriptions are presented at an
accessible level of detail for its intended audience.
Plant and animal common names are provided in
conjunction with their scientific equivalents.
Throughout the guide, the particular adaptations
that help chaparral plants and animals survive
this difficult environment, 1.e., the very features
that make them uniquely chaparralian, are
highlighted. Reference especially is often given
to a particular species’ relationship to fire. The
authors provide not only colorful descriptions
but interesting facts that keep even the most
hardened urbanite engaged.
Unfortunately, colorful language occasionally
gives way to literary excess. The authors too often
employ personification, especially with respect to
plant characteristics, that can undermine the
integrity of the data presented.
An example on the innocuous end of the
spectrum involves a species of monkey flower,
Mimulus aurantiacus. The authors describe the
unique stigma that assumes a closed position
when pollinated or even just touched by an insect
or animal. The authors then claim that in so doing
the plants ‘“‘advertise their status” as pollinated
or not, presumably giving pollinators the plant
kingdom’s equivalent of a wedding ring to would-
be suitors. Does a closed stigma make it more
likely that pollinators will move to another flower,
or is there some other evolutionary incentive for
the stigma closing? It’s a question the authors
don’t address, nor is the conclusion supported
with cited research. In this case, it’s an admittedly
harmless conclusion.
The authors’ phrasing becomes critically
dangerous with regard to fire. The first example
comes in the punctuating sentence of the
introductory paragraphs to the fire section. It
reads: “In short, where there is chaparral, there is
fire.” While this may technically be true, it is
misleading. Where there is chaparral there is
going to be fire. But where there are grasslands
there is fire. Where there are conifer forests there
is fire. In other words, the presence of fire in
chaparral is not due to something inherent in the
plant community. Rather, where there is chapar-
ral there is a Mediterranean climate and where
there is a Mediterranean climate (or any other
dry climate with enough moisture to sustain
vegetative growth) and an ignition source there
will be fire, regardless of vegetation type.
The phrase implies that chaparral fosters a fire
environment rather than the more accurate
paradigm that chaparral species exist because
they are able to survive infrequent fires. The
result of the authors’ approach is a_ subtle
perpetuation of the oft repeated and poorly
200
applied concept that chaparral needs to burn. As
the authors themselves state, certain species do
produce seeds that require fire cues to germinate.
On the other hand, we know chaparral stands
existing for more than a century without fire are
perfectly healthy systems and if chaparral stands
burn too frequently they can be converted to
non-native grasslands. Extrapolating this dichot-
omy is a tightrope that the authors have difficulty
walking.
The most prominent example comes in their
‘Living with the Chaparral” chapter. In it, the
authors describe the merits of prescribed fire and
clearing fuel breaks, offering them as effective
means to reduce mature chaparral and thus
reduce the hazards fire presents to humans.
Young chaparral, they suggest, burns less intense-
ly and cuts off the fuel supply, extinguishing fires.
Later, not as a counterpoint but as a discrete
section, they explain that extreme weather condi-
tions drive fires right through fuel breaks and
fuel-age mosaics presumably negating the value of
such “‘barriers”’.
Further on, the intrinsic watershed value of
mature chaparral stands is described in detail. The
authors explain that ““mature chaparral acts as a
sponge”’ retaining rainfall in the soil and aquifers
beneath. Without a mature chaparral and root
system, water skims off the soil surface creating
disastrous debris flows and leaving no water
behind for the ecosystem. That the previously
advocated prescribed burning and man-made fuel
breaks destroy this important ““sponge’’ mecha-
nism is a conclusion never fully reconciled by the
authors.
In truth, land management with respect to
wildfire, watershed values, and protecting lives
and property is very much a balancing act that
few, if any, have yet to master. It is perhaps unfair
MADRONO
[Vol. 58
then to expect the authors to provide solutions to
this dilemma. Rather than offer a cohesive
recommendation, the authors drop all the issues
in our lap and leave us to sort them out. It is a safe
approach. Nevertheless, since this work appears
to imply that mature chaparral is a threat to
humans rather than the other way around, it will
likely draw ire from some ecologists and conser-
vationists. And in that sense it is perplexing to
find the text demonizing an ecosystem because it
can be inconvenient to humans. While perhaps
not packaged in the most ideal manner, the
authors at least conclude the wildfire section with
a plea for humans to take responsibility for their
urban planning practices to avoid further losses.
Muddled fire issues aside, this 1s an ““/ntroduc-
tion to California Chaparral” and in that sense it
serves its purpose well. Quinn and Keeley’s work
is a celebration of a unique and teeming habitat.
To the reader unfamiliar with that amorphous
blanket of shrubs covering the hillsides outside
their car windows, Introduction to California
Chaparral provides a comprehensive glimpse into
that complex habitat. Where the authors partic-
ularly shine is in capturing the sheer diversity and
dynamic drama found within the often impene-
trable chaparral canopy. Were the reader to carry
this guide out into the field with them they would
likely recognize many more twitters, rustles and
floral displays than ever before. Ultimately,
creating that connection to the natural environ-
ment is the greatest gift any environmental author
can bestow upon a reader. In Introduction to
California Chaparral Quinn and Keeley are likely
to succeed in doing so many times over.
—CHRISTOPHER BLAYLOCK, California Chaparral Insti-
tute, P.O. Box 545, Escondido, CA 92033; chris@
blaylock.org.
MADRONO, Vol. 58, No. 3, p. 201, 2011
NOTEWORTHY COLLECTION
CALIFORNIA
PORPHYRA SUBORBICULATA Kjellman_ 1897:10—13
(BANGIACEAE).—Marin Co., epizoic on barnacles
attached to pilings in the upper intertidal at Nick’s
Cove, Tomales Bay, 38°11'57.42”N, 122°55'16.40"W,
thalli sterile and fertile, 12 May 2011, J. R. Hughey s.n.
(UC 1966687, UC 1966688) and 20 July 2011, J. R.
Hughey s.n. (UC 1966689).
Previous knowledge. Porphyra suborbiculata is natu-
rally distributed throughout Asia (type locality: Goto-
retto, Nagasaki Prefecture, Japan (Silva et al. 1996) and
reportedly also occurs naturally in the Indian Ocean,
Australia, and New Zealand (Broom et al. 2002).
However, based on recent molecular evidence some P.
suborbiculata from Australia and New Zealand, as well
as North America (Mexico and the Atlantic coast of
North America) represent introduced populations
(Broom et al. 2002; Klein et al. 2003; Neefus et al.
2008). The occurrence of this seaweed in areas
associated with increased shipping activity suggest that
its numerous introductions were the result of hitchhik-
ings on the hulls of seagoing vessels (Broom et al. 2002).
In the eastern Pacific, P. suborbiculata was first
collected in 1985 from Baja California, Mexico
(Aguilar-Rosas and Aguilar-Rosas 2003). Although P.
suborbiculata shows some variation in morphology, it is
generally characterized as being a relatively small (0.5—
4 cm in diameter), monostromatic alga, with brownish
red to pink, or bronze and violet colored blades that are
ovate to cordate or reniform in shape (Broom et al.
2002; Aguilar-Rosas and Aguilar-Rosas 2003; Neefus et
al. 2008).
Significance. First report of P. suborbiculata in
California. The gametophytic thalli collected from
Tomales Bay are in agreement with descriptions and
illustrations of this species. Specimens from California
are cordate at the base, and ovate to deeply reniform in
shape. The margins are slightly ruffled and appear
dentate to the unaided eye. Under microscopic exam-
ination thalli show the diagnostic marginal teeth. The
blade color is reddish-brown in spring and steel greenish
in summer, and fronds are more or less equal in width
and height, measuring to 1.8 cm. Spot checks for this
alga at Marshall and Marconi failed to yield additional
specimens. Identification of this invasive species was
confirmed using rbcL (GenBank JN413680) and ITS-1
(GenBank JN413679) DNA _ sequences. The rbcL
sequence was identical to two sequences from Japan
(Kanagawa, Yokosuka, Sajima and Yamaguchi, Shi-
monoseki, Tsunoshima) and differed from others in the
database by 2 or more bp. The ITS-1 sequence was
identical to fifteen other sequences deposited in
GenBank representing populations from around the
world. Since the rbcL sequence generated for P.
suborbiculata from Tomales Bay matches specimens
from Japan and the ITS-1 sequence is identical to other
invasive populations of this species from Australia, Baja
California, New Zealand, and the western Atlantic, it is
concluded that the population from California is also
the result of an introductory event. In Tomales Bay, P.
suborbiculata joins a list of five other non-native algae:
Caulacanthus ustulatus (Turner) Kutzing, Codium
fragile subsp. tomentosoides (van Goor) P. C. Silva,
Gelidium vagum Okamura, Lomentaria hakodatensis
Yendo, and Sargassum muticum (Yendo) Fensholt (C.
K. Kjeldsen, Sonoma State University, unpublished
data; Hughey 1995; Hughey et al. 1996). The maricul-
ture of oysters in Tomales Bay began around 1907
(Barrett 1963), and oysters are the likely vector for the
introduction of P. suborbiculata in the bay.
—JEFFERY R. HUGHEY, Department of Science and
Engineering, Hartnell College, Salinas, CA 93901.
jhughey@hartnell.edu.
LITERATURE CITED
AGUILAR-ROSAS, R. AND L. E. AGUILAR-ROSAS. 2003.
El género Porphyra (Bangiales, Rhodophyta) en la
costa Pacifico de México. I. Porphyra suborbiculata
Kjellman. Hidrobioldgica 13:51—S6.
BARRETT, E. M. 1963. Fish Bulletin 123. The California
oyster industry. Scripps Institution of Oceanogra-
phy Library, UC San Diego, San Diego, CA,
Website http://escholarship.org/uc/item/1870g57m
[accessed 10 September 2011].
BRooM, J. E., W. A. NELSON, C. YARISH, W. A.
JONES, R. AGUILAR ROSAS, AND L. E. AGUILAR
ROSAS. 2002. A reassessment of the taxonomic
status of Porphyra suborbiculata, Porphyra caroli-
nensis and Porphyra lilliputiana (Bangiales, Rho-
dophyta) based on molecular and morphological
data. European Journal of Phycology 37:227—235.
HUGHEY, J. R. 1995. A systematic study of Chondra-
canthus Kutzing (Rhodophyceae) with a contribu-
tion to the marine flora of Tomales Bay, California.
M.A. thesis, Sonoma State University, Rohnert
Park, CA.
, C. K. KJELDSEN, P. C. SILVA, R. L. MOE, AND
T. C. DECEW. 1996. Noteworthy Collections.
Madrono 43:432.
KLEIN, A. S., A. C. MATHIESON, C. D. NEEFUS, D. F.
CAIN, H. A. TAYLOR, B. W. TEASDALE, A. L.
WEST, E. J. HEHRE, J. BRODIE, C. YARISH, AND
A. L. WALLACE. 2003. Identification of north-
western Atlantic Porphyra (Bangiaceae, Bangiales)
based on sequence variation in nuclear SSU and
plastid rbcL genes. Phycologia 42:109—122.
NEEFUS, C. D., A. C. MATHIESON, T. L. BRAY, AND C.
YARISH. 2008. The distribution, morphology, and
ecology of three introduced Asiatic species of
Porphyra (Bangiales, Rhodophyta) in the north-
western Atlantic. Journal of Phycology 44:1399—
1414.
SILVA, P. C., P. W. BASSON, AND R. L. MOE. 1996.
Catalogue of the benthic marine algae of the Indian
Ocean. University of California Publications in
Botany, Berkeley, CA.
MADRONO, Vol. 58, No. 3, pp. 202-203, 2011
NOTEWORTHY COLLECTION
CALIFORNIA
SEQUOIADENDRON GIGANTEUM (Lindl.) J. Buchholz
(CUPRESSACEAE) (giant sequoia, big tree, or Sierra
redwood).—Riverside Co., northwestern San Jacinto
Mts., northeast of HW 243 about 18 highway km NW
of Idyllwild, in the southern unit of the San Bernardino
National Forest, on the northwestern flank of Black
Mt. along both sides adjacent to and well back from the
Black Mountain Trail, from upper Hall Canyon,
through the sloping plateau or saddle near the westerly
ridge, and into the final switchback ascent to the
summit (elev. 2369 m). Species seen from 2036 to 2236 m
elev. in mixed-conifer forest (Lower Montane Conifer-
ous Forest) on sandy granitic soil; associates include:
Abies concolor, Pinus coulteri, P. ponderosa var.
pacifica, and Quercus chrysolepis dominant at lower
elevations; P. jeffreyi and P. lambertiana dominant at
higher elevations; Calocedrus decurrens, P. flexilis, and
Ribes nevadense throughout most of area; 1 May 2009.
Collections (vouchers to be deposited at RSA, UC,
and UCR). Tree DBH 65 cm, sterile, just left (as
ascending) of trail on slope by large trunk lying across
trail, N33 49.540, W116 45.977, 2036 m elev., R. Schmid
& M. Schmid 2009-1; tree with abundant male cones,
right (as ascending) of trail on slope covered with
Pteridium aquilinum and near drainage (Indian Creek)
with Alnus rhombifolia, N33 49.579, W116 45.891,
2066.5 m elev., R. Schmid & M. Schmid 2009-2; mature
seed cones with opened cone scales and dislodged seeds,
plus mix of sandy granitic soil and sparse duff, along
Black Mountain Trail above drainage (Indian Creek)
and below sloping plateau or saddle, ca. 2070 m elev.,
R. Schmid & M. Schmid 2009-4; tree DBH 20 cm, with
medium-sized female cones, right (as ascending) side of
trail in flat open area, N33 49.607, W116 45.854, 2093 m
elev., R. Schmid & M. Schmid 2009-5; tree ca. 5.5 m
high, with many, very large, immature female cones,
just left (as ascending) of trail in flat open area, N33
49.713, W116 45.773; 2144 m elev., R. Schmid & M.
Schmid 2009-6.
In addition to these collections made on 1 May 2009,
we did a GPS census in the vicinity of the Black
Mountain Trail, starting in upper Hall Canyon. The
census revealed both in the canyon and upslope beyond
it at least 157 individuals from 2036 to 2236 m elev. (ca.
0.7 km linear distance, ca. 0.1 km’), plus an outlier
sapling 450 m distant at 2361 m elev. near the summit of
Black Mt. (2369 m). Our set of plots involved four
groups at progressively higher elevations: (1) at the
head of Indian Creek in the drainage (including
vouchers 2009-1 and 2009-2); (2) on the slope coming
out of the drainage (including voucher 2009-4); (3) at
the sloping plateau or saddle (including vouchers 2009-
5 and 2009-6); (4) on the northwest-facing slope closer
to the summit. This species alien to southern California
is regenerating prolifically on Black Mt., as revealed by
multiple age classes, from seedlings and _ saplings
(juveniles) about 20-60 cm tall to trees over 6 m tall,
about 40 years old, and reproductively mature.
Previous knowledge. The monotypic California en-
demic Sequoiadendron giganteum is native to the
western slope of the Sierra Nevada, where it occurs in
isolated groves in mixed-conifer forest (Lower Montane
Coniferous Forest) between 825 and 2700 m elev. The
67 groves are mostly of very restricted extent and/or
threatened. They occur in a narrow strip measuring
about 395 km long (northwest-southeast) and 19-21 km
at the widest points (east-west) and extending over
seven counties from southern Placer Co. southeasterly
to southern Tulare Co. (Flint 2002; Willard 2000).
This species has been extensively planted in Califor-
nia as an ornamental, as part of afforestation attempts,
and as reforestation efforts, especially post-fire re-
vegetation ones. For example, Burns and Sauer (1992)
noted that 22,900 seedlings of S. giganteum were
planted in southern California in the San Gabriel
Mts. alone, but neither this species nor 44 other alien
conifer species planted there as part of afforestation
projects “‘are invading adjacent natural habitats” (p. 49).
However, Rogers (1986: p. 33) wrote: “On the San
Bernardino [National Forest] about 5000 to 10,000
seedlings are planted each year, and at least one
instance of natural regeneration from some of the early
plantings has been observed.’’ Rogers’s “‘one instance”’
may well be the introduction of S. giganteum after the
Aug. 1974 fire in Hall Canyon (Keeler-Wolf 1990;
Cheng 2004) and its subsequent naturalization on Black
Mt., which is the subject of the present preliminary
report. Our extensive analysis of the print and Internet
literature for the floristics and ecology of southern
California suggests S. giganteum is possibly also
naturalized in the San Gabriel Mts. of Los Angeles
Co. and southwestern San Bernardino Co., and in the
San Bernardino Mts. of San Bernardino Co.
Significance. First report and collections for River-
side Co. and the San Jacinto Mts. (see the database of
the Consortium of California Herbaria, http://ucjeps
.berkeley.edu/consortium). More importantly, first doc-
umented record for naturalization of this Sierra- Nevada
endemic species in montane southern California. State
and regional floras for California should acknowledge
in their keys and descriptions such naturalizations.
Details of these findings will be published elsewhere
(Schmid and Schmid in press).
—RUDOLF SCHMID, Department of Integrative
Biology, University of California, Berkeley, CA
94720-3140; MENA SCHMID, Somerville, MA 02144.
schmid@berkeley.edu.
LITERATURE CITED
BURNS, C. AND J. SAUER. 1992. Resistance by natural
vegetation in the San Gabriel Mountains of
California to invasion by introduced conifers.
Global Ecology and Biogeography Letters 2:46—51.
CHENG, S. (tech. ed). 2004. Forest Service Research
Natural Areas in California. General Technical Re-
port PSW-GRT-188. USDA Forest Service, Pacific
Southwest Research Station, Albany, CA. Website
http://www. fs.fed.us/psw/publications/documents/
psw_gtr188/ [accessed 20 June 2011].
FLINT, W. D. 2002. To find the biggest tree, revised ed.
Sequoia Natural History Association, Three Riv-
ers, CA.
2011]
KEELER-WOLF T. 1990. Ecological surveys of Forest
Service Research Natural Areas in California. Gen-
eral Technical Report PSW(GTR)-125. USDA Forest
Service, Pacific Southwest Research Station, Berkeley,
CA. Website http://www.fs.fed.us/psw/publications/
documents/psw_gtr125/ [accessed 20 June 2011].
ROGERS, R. R. 1986. Management of giant sequoia in
the national forests of the Sierra Nevada,
California. Pp. 32-36 in C. P. Weatherspoon,
Y. R. Iwamoto, and D. D. Piirto (tech. coordi-
nators). Proceedings of the workshop on man-
agement of giant sequoia, May 24-25, 1985,
NOTEWORTHY COLLECTION 203
Reedley, California. General Technical Report
PSW(GTR)-95. USDA Forest Service, Pacific
Southwest Forest and Range Experimental Sta-
tion, Berkeley, CA. Website http://www. fs.fed.us/
psw/publications/documents/psw_gtr095/_— [ac-
cessed 20 June 2011].
SCHMID, R. AND M. SCHMID. In Press. Naturalization
of Sequoiadendron giganteum (Cupressaceae) in
montane southern California. Aliso 30.
WILLARD, D. 2000. A guide to the sequoia groves of
California. Yosemite Association, Yosemite Na-
tional Park, CA.
MADRONO, Vol. 58, No. 3, p. 204, 2011
NOTEWORTHY COLLECTION
COLORADO
DRABA WEBERI R.A. Price & Rollins (BRASSICA-
CEAE).—Park Co., Middle Fork of the South Platte,
near Magnolia Mill, Pike National Forest, in mossy
depressions on north side of granite boulder, 10 plants
in flower and fruit, at 39.36112°N, 106.09013°W,
3,398 m elev., 20 July 2010, Bernadette Kuhn 7927a
and 7927b, along with Gina Glenne, Sheila Lamb,
Alicia Langton, Ellen Mayo, Steve Olson, and Jeff
Sprovkin (MO). Dr. Ihsan Al-Shehbaz verified the
identification.
Previous knowledge. Global distribution is limited to
the type locality in the Monte Cristo Creek drainage,
below the Upper Blue Lake Reservoir Dam, on the west
side of Continental Divide, Summit County, Colorado.
As described in the original publication (Price and
Rollins 1991) the holotype (Price #464) was collected
from “‘crevices of rocks beside cascades, amid rocks at
edge of stream’. The type locality was described by
Price as containing “‘perhaps 100 individuals”, and in
2009, 81 individuals were reported present at the site
(Colorado Natural Heritage Program 2011). All indi-
viduals documented are on private land owned and
operated by Colorado Springs Utility, though the site is
surrounded by National Forest System lands (Decker
2006).
Significance. Second documented occurrence. For
twenty years botanists have been searching for another
occurrence outside the species’ type locality. Our
collection, located 2.8 km to the south, expands the
known distribution of D. weberi to another county
(Park), and to the east side of the Continental Divide.
In addition, it confirms the long-suspected presence of
D. weberi on National Forest System land. This finding,
along with the recent addition of D. weberi to the
USDA Forest Service Sensitive List, may present new
opportunities for the management and conservation of
this extremely rare Colorado endemic (USDA Forest
Service 2011).
—BERNADETTE KUHN, Colorado Natural Heritage
Program, 254 General Services Building, 1474 Campus
Delivery, Fort Collins, CO 80523-1474. bernadette.
kuhn@colostate.edu.
LITERATURE CITED
COLORADO NATURAL HERITAGE PROGRAM. 2011,
Biodiversity Tracking and Conservation System
(BIOTICS). Colorado State University, Fort Col-
lins, CO. Website http://www.cnhp.colostate.edu/
exchange/request.asp [accessed 01 May 2011].
DECKER, K. 2006. Draba weberi Price & Rollins
(Weber’s draba): a technical conservation assess-
ment. USDA Forest Service, Rocky Mountain
Region, Golden, CO. Website http://www. fs.fed.us/
r2/projects/scp/assessments/drabaweberi.pdf [ac-
cessed 1 May 2011].
PRICE, R. A. AND R. C. ROLLINS. 1991. New taxa of
Draba (Brassicaceae) from California, Nevada, and
Colorado. Harvard Papers in Botany 3:75—77.
USDA FOoRrRESsT SERVICE. 2011. Supplement No. 2600-
2011-1. Chapter 2670—Threatened, endangered,
and sensitive plants and animals. USFS Forest
Service Manual, FSM 2600-Wildlife, Fish, and
Sensitive Plant Habitat Management. USDA
Forest Service, Rocky Mountain Region (Region
2), Denver, CO. Website http://www.fs.fed.us/im/
directives/field/r2/fsm/2600/fsm_r2_2670_201 1-1.
docx [6 January 2012].
MADRONO, Vol. 58, No. 3, p. 205, 2011
NOTEWORTHY COLLECTION
MEXICO
ANEMONE TUBEROSA Rydb. (RANUNCULA-
CEAE).—Baja California, Municipio of Ensenada,
Ejido Nativos del Valle heading W to Santo Tomas,
31.42561°N, 116.34858°W (WGS 84), 434 m/1428 ft, 26
March 2010, Sula Vanderplank, Sean Lahmeyer, Ben
Wilder and Karen Zimmerman 100326-29 (RSA), Less
than 100 plants observed growing with Zigadenus sp. on
N-facing side of a steep limestone outcrop. Most plants
in full flower.
Previous knowledge. The general habitat of this
species is given by Dutton et al. (1997), in the Flora
of North America, as from rocky slopes and stream
sides 800-2500 m. It is known from desert regions of the
southwestern USA (CA, NM, NV, TX, UT) and NW
Mexico (Baja California, Sonora), and has been well
documented within the Sonoran and Mojave deserts:
Wiggins (1980) reports it from the western edge of the
Colorado Desert and the eastern Mojave Desert;
Wilken (1993) gives the range in California as eastern
Desert Mountains; 900-1900 m; Munz (1974) reports
elevations of 3000—5,000 ft (914-1520 m), in Joshua
Tree Woodland and Pinyon-Juniper Woodland, west-
ern edge of the Colorado Desert, eastern Mojave
Desert. The westernmost known collections for this
taxon in California are from the southern Cuyamaca
Mtns., near the southwestern end of Poser Mtn, March
19 1995, Jeri Hirshberg 253 (RSA) 60 km from the
Pacific coast; and the Laguna Mountains, below Desert
View, 06 April 1939, A. J. Stover 245 (SD), 80 km from
the Pacific coast (CCH 2011). In Arizona, Kearney and
Pebbles (1951) report elevations of 2,500—S000 ft (760—
1520 m). In Mexico, the two Baja California collections
closest to the Pacific Ocean are both 50 km inland (30—
45 km from the Gulf of California to the east) in the
central desert, close to the narrowest part of the
peninsula: summit of volcanic hill at top of Jaraguay
Grade, 24 Feb 1973, Reid Moran 20248 (SD), elev.
875 m, (the lowest documented elevation for this species
in the state); Sierra San Borja, summit of Cerro la
Chona, 19 Mar 1966, Reid Moran 12782 (SD), 1450 m
(San Diego Natural History Museum 2011). There are a
small number of California specimens from ~400 m
elev., all from a small area within the Whipple
Mountains, on the border with Arizona (CCH).
Significance. First collection of this species west of the
peninsular ranges and inside the California Floristic
Province. This collection represents a range extension of
ca. 70 km west from the peninsular ranges, and is the
most coastal collection documented to date, being just
15 km (9 mi) from the Pacific Ocean. The elevation is the
lowest recorded in Baja California, and is near the
lowest elevation recorded for the species anywhere.
Further investigation of the foothills surrounding this
site may help determine if this small population is indeed
isolated from the core range, perhaps as a result of the
unusual habitat provided by the limestone outcrop.
—SULA VANDERPLANK, Deptartment of Botany &
Plant Sciences, University of California, Riverside, CA,
92507; SEAN LAHMEYER, The Huntington Library, Art
Collections, and Botanical Gardens, 1151 Oxford
Road, San Marino, CA 91108. sula.vanderplank@
gmail.com.
LITERATURE CITED
CONSORTIUM CALIFORNIA HERBARIA (CCH). 2011.
University of California, Berkeley, CA, Website
http://ucjeps.berkeley.edu/consortium/ [accessed 3
March 2011].
DUTTON, B. E., C. S. KEENER AND B. A. FORD. 1997.
Anemone. Pp. 139-158 in Flora of North America
Editorial Committee (eds.), Flora of North America
North of Mexico, Vol. 3. Oxford University Press,
New York, NY.
KEARNEY, T. H. AND R. H. PEEBLES. 1951. Arizona
flora. University of California Press, Berkeley, CA.
Munz, P. A 1974. A flora of southern California.
University of California Press, Berkeley, CA.
SAN DIEGO NATURAL HISTORY MUSEUM, 2011. The
flora of Baja California. San Diego Natural
History Museum, San Diego, CA. Website http://
bajaflora.org/ [accessed 3 March 2011].
WIGGINS, I. 1980. The flora of Baja California. Stanford
University Press, Stanford, CA.
WILKEN, D. H. 1993. Anemone. Pp. 912-913 in J. C.
Hickman (ed.), The Jepson manual: higher plants
of California. University of California Press,
Berkeley, CA.
Volume 58, Number 3, pages 131-206, published 28 March 2012
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