Three New Subspecies of Monardella (Lamiaceae) from Southern California, U.S.A

2015·28.5 MB·English

by  Mark A Elvin

New
Three Subspecies Monardella (Lamiaceae) from Southern
of
U.S.A.
California,
Mark
Elvin
A.
UCLA
Herbarium (LA), Department of Ecology and Evolutionary Biology, University of California
Los Angeles, Los Angeles, 90095, U.S.A. [email protected]
California
Andrew
Sanders
C.
Herbarium, Department Botany and
of Plant Sciences, University of California Riverside,
Riverside, California 92521-0124, U.S.A. [email protected]
Riehard Burgess
A.
Thousand 2100 Thousand Oaks Thousand
Oaks, Boulevard, Oaks, 91362, U.S.A.
City of California
[email protected]
Barbara
Hellenthal
J.
Greene-Nieuwland Herbarium (NDG/ND), Department
of Biological Sciences, University of Notre
Dame, Dame,
Notre Indiana 46556, U.S.A. [email protected]
eomplex synonymy M.
Abstract. Continuing work on the flora of Califor- Benth. results in the of
Elmer M.
The Manual: franciscana subsp.
nia after the publieation of Jepson to villosa villosa.
Vascular Plants of California has led to the diseovery Monardella villosa subsp. subserrata (Greene) Epling
new upland and
of three Monardella Benth. (Lamiaeeae) subspe- reeognized as the interior taxon in this
is
& A
eies: M. australis Abrams subsp. gabrielensis Elvin speeies. leetotype is designated for M. purpurea
Howell and M. Elmer
M. reeognized both franciscana
A. C. Sanders, australis subsp. occidentalis Elvin, for
&
and M.
R. A. Burgess A. Sanders, and M. sinuata Elvin neglecta Greene.
C.
& &
lUCN
Key Red
A. C. Sanders subsp. gerryi Elvin, A. C. Sanders words: California, Lamia-
List,
R. A. Burgess. Monardella australis subsp. gabri- eeae, Monardella, San Gabriel Mountains, South
and San
elensis oeeurs in the eentral western Gabriel Coast Ranges, Transverse Ranges.
Mountains Transverse Ranges, and M.
of the australis
subsp. occidentalis oeeurs on Pine Mountain Ridge in Monardella Benth. (Lamiaeeae) oeeurs throughout
the Western Transverse Ranges and the San Rafael western North Ameriea from northern Mexieo
to
Mountains of the adjaeent southern portion of the southern Canada with 59 taxa distributed among 35
South Coast Ranges. The two new subspeeies of M. speeies. California the eenter of diversity for
is
australis are most similar the autonymie subspe- Monardella with 52 taxa distributed among 30
to
and
eies, but both differ in their braet leaf speeies (Sanders et ah, 2012). Reeent work for The
morphology, pubeseenee on the stems and ealyees, Jepson Manual: Vascular Plants California
of
and Monardella
stature, distribution. sinuata subsp. (Baldwin ah, 2012) has stimulated eontinuing
et
Work
gerryi oeeurs in the Las Posas and Camarillo Hills in taxonomie investigation of Monardella. on a
&
the eoastal plain of Ventura County in the South flora of Ventura County, California (Burgess
The new
Coast Region. subspeeies most similar to Munro, in prep.), and other ongoing loeal projeets
is
M. Most
sinuata subsp. sinuata but differs in its stem, leaf, has led to further diseoveries in Monardella. of
braet, and glomerule size and in pubeseenee, the taxa in this genus have very limited distributions,
its
new
nutlets, and eeologieal setting. Monardella neglecta whieh also the ease for the three subspeeies
is
new
Greene rank subspeeies deseribed
transferred a as a of herein.
to
is
M. purpurea Howell, as M. purpurea subsp. neglecta Taxonomically important characters Monardel-
in
&
The main
(Greene) Elvin A. C. Sanders. a diminutive, morphologieal eharaeters that have
It is la.
glabrous, serpentine endemie on the Marin and been used over time distinguish between taxa in
to
A
Tiburon peninsulas in northern California. taxo- the genus Monardella inelude plant habit and
leaf,
nomie assessment names M. and morphology
of within the villosa braet, infloreseenee (Gray, 1876,
Novon 416-431. Published on 9 January 2015. 10.3417/2014002
23:
doi:
Number
Volume 417
4
23, Elvin et
al.
2014
Monardella (Lamiaceae) from Southern
California
1886; Abrams, 1912a, 1912b, 1951; Epling, 1925, in elevation in the Peninsular and Transverse ranges
Munz,
1939; Jepson, 1925, 1943; 1935, 1959, 1974; of southern California.
While used
these eharae-
Jokerst, 1993). historieally
Monardella
la. australis subsp. australis. Eigure
ters ean be highly variable leaf morphology,
(e.g.,
lA.
stem length) within a single taxon, they ean be
still
when
used
helpful for identifieation, espeeially in
Monardella
subsp. oeeurs
australis australis in
tandem. Other
eharaeters ealyx length, braet
(e.g.,
mid-montane
subalpine roeky
ehaparral,
to forest,
some m
morphology) ean be variable in taxa but are
1450-3300
San
openings, in elevation in the
at
more More we have
eonsistent in others. reeently,
Jaeinto and San Bernardino mountains, in Riverside
found pubeseenee be
of stem, ealyx)
(i.e., leaf, to and San Bernardino
eounties, respeetively, in
among
most used
the stable of eharaeters
to
southern California. distinguished from the other
It is
between and
distinguish speeies subspeeies in
M. subspeeies by long ealyees
australis (greater
its
&
Monardella (Elvin Sanders, 2003, 2009; Sanders mm) and (12-18 mm)
than 8.5 long, purple braets
ah, 2012; Elvin ah, 2013). exeeed
et et that the ealyees.
We
have eaeh
notieed that plant strueture
(e.g.,
lb. Mouardella australis subsp. ciuerea (Abrams) A.
stem, ealyx, braet) in Monardella generally has
leaf,
&
Novon
Sanders 323. 2009.
own pubeseenee, and eaeh ean have one C. Elvin, 19(3):
distinet
its
Monardella Abrams, Muhlenbergia
cinerea
8:
several different types of triehomes that are
to
TYPE:
1912. U.S.A. Los Angeles
33. California:
intermixed on these Eor example, M.
struetures.
San Antonio 9000
Co., Mt. [Mt. Baldy], or
ft.
robisonii Epling has one type of glandular triehome
24 Abrams 1928
more,
July 1901, R.
L.
and
two types of non-glandular triehomes
all
MO
(holotype, DS; isotypes, photo of holotype
many
on
intermixed the stems. Unfortunately, of the
OSC
sheet, not seen). Eigure IB.
pubeseenee some
different types of are minute, with
mm
0.01-0.03 Triehomes
small
as as in length. of
Monardella australis subsp. cinerea oeeurs in mid-
lOX
this size eannot be seen reliably with a standard
montane and endemie
subalpine the
to forest to
is
hand but they ean be distinguished with the use
lens,
upper (1800-3100 m) San
elevations in the Gabriel
The
of a mieroseope. triehomes do not lend
Mountains Los Angeles County,
in California.
It is
themselves to easy identifieation in the field, but distinguished from the other M. australis subspeeies
they do provide aeeurate
for identifieation. by short stems (3-15 em), very short internodes
its
As
between
noted below, there are differenees the mm), (5-10 mm),
(3-8[10] short leaves the grayish
new many
and
subspeeies subspeeies hue and mm).
their sister in of herbage, short ealyees
(5-7[8]
its its
morphology,
of the standard, old eharaeters leaf
(e.g.,
Mouardella
many l australis subsp. gabrieleusis Elvin
stem length); however, in instanees there c.
is
&
TYPE:
A. Sanders, subsp. nov. U.S.A.
C.
overlap between taxa the edge of the eharaeter
at
Los Angeles San
Gabriel Mtns.,
California: Co.,
The new
ranges leaf length). three subspeeies
(e.g.,
NE
2051 27
faee of Paeifieo Mtn., m., Sep.
presented below unequivoeally from
are distinet their
&
2011, D. E. Bramlet, E. Kentner Zylstra
sister subspeeies in pubeseenee eharaeters J.
(e.g.,
4972 UCR; MO,
LA,
JEPS,
(holotype, isotypes,
stem,
braet, ealyx).
SBBG, VEWO).
US,
Eigures 1C,
2.
Monardella Abrams, Muhlenbergia
australis
1. 8:
34. 1912. Monardella odoratissima Benth. The new
Diagnosis. subspeeies similar Monardella
to
is
Abrams
(Abrams) Ann. australis subsp. australis, but differs in having
subsp. Epling, Missouri
australis
green stramineous braets purple or rose-tinged in
to (vs.
Card. 70-71. 1925. Monardella
Bot. 12:
mm
12-18
subsp. australis) that are shorter vs.
([5J6.5-9
odoratissima Benth. (Abrams)
var. australis mm
mm) 4—8
and mm),
narrower (2—4.5 shorter ealyees
vs.
mm
TYPE:
437. 1943.
Jeps., El. Calif. [Jepson] 3: (7—8 8.5—10 mm), and laeking non-glandular
in
vs.
open ealyx triehomes.
U.S.A. Riverside
California: Co., forests of
Tamaraek Valley, July 1901, H. M. Hall 2486
em
Caespitose perennials subshrubs, (6)15-25
to
UC
MO,
DS;
(holotype, isotypes, not seen).
matted; rhizomatous; stems
tufted visibly
to
tall,
Monardella mat-forming
australis is a eaespitose, woody base, pubeseenee dense, with 2 types of
at
mm
rhizomatous perennial subshrub distinguished by stem 0.02-0.04 and
to triehomes, glandular,
(1) (2)
mm
X
mm.
having long non-glandular triehomes, 0.3-0. 6 non-glandular, 0.3-0. 6 Leaves
(6)10-19(21)
mm
mm,
minute 0.02-0.04
with glandular triehomes, narrowly widely
2)3. 5-5(6) to elliptie, sessile,
(2.
m
1350-3300
beneath. oeeurs in montane areas pale or grayish green; with 2 types of triehomes on
at
It
418 Novon
—
Figure A-F. Glomemles and bracts of Monardella Benth. subspecies in southern California. A. Monardella australis
1.
— —
&
Abrams
subsp. australis. B. Monardella australis subsp. cinerea A. C. Sanders Elvin. C. Monardella australis subsp.
— —
& &
gabrielensis Elvin A. C. Sanders. D. Monardella australis jokerstii Elvin A. C. Sanders. E. Monardella australis
—
& &
subsp. oeeidentalis Elvin, R. A. Burgess A. C. Sanders. E. Monardella linoides A. Gray subsp. ereeta (Abrams) Elvin A.
Abrams 1928
C. Sanders. A, photographed from MeClintoek (LA-62613); B, from (holotype, DS); C, from D. E. Bramlet, E.
s.n. L.
&
&
Kentner Zylstra 4972 (holotype, UCR); D, from M. A. Elvin K. VinZant 5085 (isotype, IRVC); E, from R. A. Burgess, M. A.
J.
& E E
Elvin Simpson 9700 (holotype, UCR); E, from Abrams 2861 (isotype, US). All images taken and prepared by Michael R.
Tiffany.
mm
adaxial blade surface, glandular, 0.02-0.04 less than or equaling the calyces. Flowers with the
(1)
mm
mm,
7-8
and 0.4-0.6
(dense) non-glandular, (sparse calyx green, rarely purple or rose-tinged
to
(2)
dense); type of trichome on abaxial blade surface, the calyx with 2 types of trichomes,
1 distally,
(1)
mm mm mm
0.02-0.04 1-0.2 and
glandular, (sparse dense); leaves glandular, glandular, 0.4
to 0.
(2)
subsessile to decurrent, base acute to obtuse, apex (sparse), but non-glandular trichomes absent; corolla
mm,
12-14
acute obtuse, rounded. Inflorescence a terminal white cream with fine purple markings
to to
10- make
glomerule on primary stems, solitary; glomemles that the corolla appear lavender in general
mm
X
mm, 14-17 mm, 14-16
18 wide; bracts (5)6.5-9 2-4.5 green stigma exserted; stamens
to color;
mm,
Fmit
stramineous, occasionally purple-tinged distally, exserted. a nutlet, tan to light brown,
X
mm.
narrowly narrowly lanceolate, apices acute. mottled dark brown, oblong, 2-2. 3 0.8-0. 9
elliptic to 2.
1
Number
Volume 419
4
23, Elvin et
al.
2014
Monardella (Lamiaceae) from Southern
California
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Abrams
Figure Monardella australis subsp. gabrielensis Elvin A. C. Sanders. Holotype, D. E. Bramlet E. Kentner
2.
J.
4972
(UCR).
Zylstra
and decomposed
Monardella
Distribution habitat. australis granite/grandiorite soils or granitic
m
1600-2200
endemic
subsp. gabrielensis the north central outcrops in elevation. Associated
to at
is
San Gabriel Mountains in the Transverse Ranges of dominants in these areas include Cupressaceae such
southern occurs openings montane Calocedrus decurrens Pinaceae such
California. in in as (Torr.) Florin,
It
coniferous forest, montane coniferous woodland, and as Pinus Balf. and P. lambertiana Douglas,
jeffreyi
canyon oak woodland sometimes has and Fabaceae such Quercus Liebm.
live that a as chrysolepis
montane chaparral understory. The new subspecies Other commonly associated species include Poaceae
most commonly occurs in either colluvium of such as Bromus tectorum L. and Poa secunda Presl;
J.
Novon
420
Asteraceae such as Ericameria nauseosa (Pall.) G. L. Gabriel Mountains are distinct from those in the San
&
Nesom
Eriophyllum Bernardino Mountains.
G. Baird,
confertiflorum
1.
(DC.) A. Gray, and Hulsea A. Gray subsp. Monardella australis subsp. gabrielensis most
vestita is
M.
gabrielensis Wilken; Onagraceae such as Gayophy- similar to australis subsp. australis but differs
&
tum A. Gray; Orobanchaceae such from in having glomerule bracts that are shorter
diffusum Torr.
it
mm mm
12-18
subspecies
as Cordylanthus nevinii A. Gray; Polygonaceae such ([5]6.5-9 vs. in australis),
mm
4-8
Eriogonum nudum Douglas ex Benth. and narrower (2-4.5 vs. mm), and green versus
as E.
and
purple having
umhellatum and Rubiaceae such Galium or rose-tinged, in bracts that are
Torr.; as
&
Hilend Howell. less than or equal to the calyces versus noticeably
jepsonii T.
J.
them The
exceeding lA,
calyces are shorter
(Fig. C).
mm
lUGN
Red
Monardella
List category. australis in M. australis subsp. gabrielensis (7-8 8.5-
vs.
mm)
subsp. gabrielensis has a restricted distribution in the 10 mm), with shorter 1-0.2 glandular
(0.
north-central San Gabriel Mountains. Using historical trichomes and sparse, longer glandular trichomes
to
new mm;
and current records, the subspecies has an 0.4 these differ from M. australis subsp. australis
mm)
extent of occurrence (EOO) of approximately 148.6 and lack minute (0.04-0.06 glandular
its
mm)
km^ (minimum convex polygon) and an area of trichomes and longer (0.3-0. 6 non-glandular
occupancy (AGO) approximately 8 km^ (sum trichomes. Occasionally the sparse glandular
of of
tri-
mm
chomes on M.
occupied have been 0.4 long the calyces of australis
grid squares). All collections
made can and
within the boundaries of the Angeles National subsp. gabrielensis lose their glandular tips
The
Department which appear be non-glandular. two subspecies have
Forest (U.S. of Agriculture), to
is
managed many may some separate ranges, with the San Gabriel Mountains the
This provide
for uses.
and San
may range for subspecies gabrielensis the
protection, but also result in threats the
to
it
Bernardino and San Jacinto mountains the range
depending on Given for
species, the land use. the
too,
subspecies however, appears be
there a
australis; to
uncertainty regarding climate change, the extremely
zone of introgression with several specimens that
and and
limited distribution fragmentation, the
represent putative intermediates between the two
burn
increase in megafires that large areas
(fires
subspecies, example, Swinney 9256 (UCR,
for R. G.
100,000 and have considerable on
acres] effects
[e.g.,
VFWO)
11330
and Swinney (UCR).
R.
G.
humans)
southern assessed here
in California, as
is
it
Monardella
australis subsp. gabrielensis similar
CR, Endangered (CR is
or Critically B2ab[i,iii,iv,v]),
and M.
occurs adjacent australis subsp. cinerea
lUCN to to
according
to criteria (2001).
can be from by
but distinguished longer
corollas
its
it
mm mm
9-11
(12-14 and
subspecies
Phenology. Plants of Monardella australis subsp. vs. in cinerea)
mm)
absence minute
have been documented between of (0.06 glandular calyx
gabrielensis flower
to
mm)
trichomes and long 6 non-glandular
mid-July and mid-September. (0.3-0.
The
calyx trichomes. two subspecies occur
in
The
Etymology.
Latin epithet gabrielensis refers and
different ecological settings different eleva-
at
m
to the San Gabriel Mountains, in which this tions (1600-2200 1800-3100 m). While M.
vs.
subspecies’ entire range occurs. australis subsp. cinerea also endemic the San
is to
Gabriel Mountains, appears not overlap with
to
it
Taxonomic Monardella
subsp.
australis
affinities. howev-
subspecies geographic
gabrielensis in ranges;
gabrielensis closely related the three previously
to
is one specimen 953
there least Dressier
er, is at (R. L.
M.
described subspecies of australis (subspecies
between
[LA]) that represents a putative intermediate
new
and and M.
australis, cinerea, jokerstii) to the More
the two subspecies. introgressant individuals
subspecies described below
australis in habit,
its are expected be found.
to
new
and
general morphology. This subspe-
vestiture, Monardella
australis subsp. gabrielensis distin-
is
cies overlaps in several gross morphological characters guished from M. by
subsp. calyx
australis jokerstii
its
mm
with the other subspecies of M. australis, such as stem pubescence (lacking the minute glandular 0.06
mm
and
height leaf size, indicating their close relationship. trichomes and long non-glandular 0.3-0. 6
However, M. can be
australis subsp. gabrielensis trichomes seen subspecies morphol-
in leaf
jokerstii),
by and
distinguished several morphological characters ogy (blades entire vs. serrate), inflorescence branch-
occupies a separate geographic range. Prince (2009) ing pattern glomerules an open com-
(solitary vs.
conducted pound cyme number
a population genetics analysis of several of glomerules), the of glomerules
Monardella taxa from southern California, and her per main stem (one one and longer
vs. to five),
mm
10-11
M. San (12-14 mm). Monardella
results indicate that australis plants in the corollas
vs.
Number
Volume
4 421
23, Elvin et
al.
2014
Monardella (Lamiaceae) from Southern
California
&
and M. Simpson 9700
australis subsp. gabrielensis occurs west north- Burgess, A. Elvin
L.
UCR; MO,
west of Mount San Antonio (the highest peak in the (holotype, isotype, CAS, K, LA,
SBBG, VFWO).
San Gabriel Mountains 3069 m), while M. australis OBI, Figures IE,
at 3.
and
subsp. jokerstii occurs south east of
it.
San Bernardino Mountains Monardella
In the
The new
Diagnosis. subspeeies similar Monardella
to
is
australis subsp. australis (Fig. lA) and M. linoides australis Abrams subsp. australis, but differs in having
mm mm
& wider leaves (4-1 1 vs. 2.5-5 in subsp. australis),
A. Gray subsp. erecta (Abrams) Elvin A. C.
mm
12—18
(7—12 mm),
shorter braets shorter ealyees (7—
vs.
Sanders IF) appear interbreed extensively,
(Fig. to
mm
8.5-10 mm), and nonglandular stem
8.5 short
vs.
based on numerous hybrid specimens. This mm
putative
1—0.2
triehomes
0. long.
San
pattern continues in the Gabriel Mountains,
cm
8-22
Caespitose perennials subshrubs,
to
number tall,
where specimens
there are a of that exhibit
woody
matted; rhizomatous; stems
tufted visibly
to at
pubescence, and
intermediate states in leaf bract size
pubescence stem
base, dense, with 3 types of
and and between M.
shape, general habit
australis
mm,
triehomes, glandular, 0.02-0.04 non-
and M. (1) (2)
subsp. gabrielensis linoides subsp. erecta
mm,
1-0.2 and 0.3-
glandular, non-glandular,
0.
1886
specimens [RSA], (3)
0. Mistretta R.
G.
(e.g.,
mm
X
mm. 11-17 4-11 mm,
Leaves
0.6
elliptic to
Swinney 5589 [RSA, UCR], Swinney 5646
R.
G.
ovate, sessile, pale or grayish green; with 2 types of
[RSA, UCR], Swinney 5656 [RSA, UCR]).
on 0.02-0.04
triehomes adaxial blades, glandular,
(1)
mm
mm;
Paratypes. U.S.A. California: Los Angeles Co., R. (dense), and non-glandular, 0.4-0.6 with
(2)
6419 4959
Bacigalupi (JEPS not seen, LA), D. Bramlet
on
3 types of triehomes abaxial blade surfaces,
(1)
Ewan
8417
(GH,
OBI), T. (LA), Sep. 1917, F. Grinnell mm
/.
0.02-0.04
glandular, (dense), non-glandular,
(2)
(RSA-40I600), 29 Aug. (RSA-40I603), 29
1917,
s.n. s.n.
mm, mm;
1-0.2 and non-glandular, 0.4-0. 6
0.
Aug. (POM-8I30), 23 Aug. Hasse (3)
1917, 1893, H. E.
s.n. s.n.
1824 1924 leaves subsessile decurrent, bases acute obtuse
(NY), 0. Mistretta (RSA), (RSA), F. W. Peirson to to
UC
1070 (RSA), T. S. Ross 5594 (RSA, not seen), 5779 (truncate) decurrent, apices widely acute obtuse.
to
UC
(RSA, Swinney 9071 (RSA, UCR), 9210
not seen), R.
G.
Inflorescence a terminal glomerule on primary stems,
VFWO), 38267
(UCR, Thorne (RSA), Wheeler
R. F.
L. C. mm
14-19
glomerules wide; glomerule
solitary;
10058 (RSA), 10308 (RSA), 23 (RSA-
July 1968,
s.n.
mm
X
mm,
7-12
bracts 5)4-5. 5 stramineous
to
638957), 7 Aug. 1968, (RSA-609405). (3.
s.n.
purple, widely greater than or equaling the
elliptic,
&
mm,
Monardella 7-8.5
l australis subsp. jokerstii Elvin A. calyces. Flowers with the calyx purple;
d.
Novon
C. Sanders, 19: 323-324, 2009. calyx with 3 types of triehomes, glandular, 0.2
fig 4.
(1)
mm
mm,
TYPE:
U.S.A. San Bernardino S glandular, 0.4 and non-
California: Co., (sparse),
(2) (3)
W
mm; mm,
Cucamonga Day 13-16
face of Peak, just of Creek, glandular, 0.3-0. 6 corolla white to
&
make
25 Aug. 2006, M. A. Elvin K. VinZant 5085 cream, occasionally with purple markings that
UCSB; ARIZ, BRY, CAS, appear pale lavender general stigma 16-
(holotype, isotypes, in color;
it
mm, mm,
CHSC, GH, IRVC, MO, NY, 19 stamens 15-18
JEPS, K, OBI, exserted; exserted. Fruit
F,
RSA, SBBG, UCR,
SD, US). Figure ID. a nutlet, tan to light brown, mottled dark brown,
X
mm.
2.25-2.35 8-0.9
oblong,
0.
Monardella
subsp. occurs
australis jokerstii in
and
Monardella
colluvium on steep scree or talus slopes or stony Distribution habitat. australis
an
benches on canyon bottoms montane and subsp. occidentalis is extremely rare plant with
in forest
m V
known
(160)1350-1750 Mon- only one extant occurrence, which in entura
chaparral in elevation. is
at
been Western
endemic San County. has collected in the
ardella australis suhsp. jokerstii is to the It
Ranges and
Gabriel Mountains San Bernardino County, Transverse the southeastern edge of the
in
Ranges
Outer South Coast Santa Barbara County.
where in
California, occurs in the vicinity of
it
The Ventura County were on Pine
Cucamonga Peak (Mount San and plants in collected
Antonio) the
Mountain Mount and
Ridge, south of Pinos Frazier
western portion of Lytle Creek. distinguished
It is
The
Mountain. Santa Barbara County were
plants in
from the other M. australis subspecies by serrate
its
collected in the San Rafael Mountains near Big Pine
and branching
leaves inflorescence.
Specimens have been taken from
[Mountain].
dry,
m
Monardella 1890
le. australis subsp. occidentalis Elvin, exposed rocky slopes elevations between
at
&
R. A. Burgess A. C. Sanders, subsp. nov. and 2433 m. occurs as an understory species in
It
TYPE:
U.S.A. California: Ventura Co., Pine mixed conifer woodlands and montane chaparral
Mountain loam
Ridge, steep slope with clay soils associated with Pinaceae such as Abies concolor
&
derived from a sandstone base in a rocky/scree (Gordon Glend.) Hildebr., Pinus P.
jeffreyi,
&
colluvium matrix, 2130 m, 20 June 2014, R. A. lambertiana, P. ponderosa P. Lawson C. Lawson;
Novon
422
&
Figure Monardella australis Abrams subsp. occidentalis Elvin, R. A. Burgess A. C. Sanders. Holotype, R. A. Burgess, M.
3.
& L
9700
Elvin Simpson (UCR).
A.
moha- Raven
Asteraceae such as Ericameria nauseosa var. (Greene) P. H. subsp. latifolium (Hook.) P. H.
&
Nesom
vensis (Greene) G. L. G. Baird; and Raven, and Polemoniaceae such as Eriastrum densi-
1.
Rhamnaceae Mason
such as Ceanothus cordulatus Kellogg folium (Benth.) H. subsp. austromontanum
(T.
and Frangula californica (Eschsch.) A. Gray subsp. T. Craig) H. Mason. Monardella australis subsp.
&
tomentella (Benth.) Kartesz Gandhi. Associated occidentalis occurs in colluvium on a sandy clay loam
herbaceous and suffruticose perennials of the under- derived from weathered sandstone, sandy loam,
soil
story include Asteraceae such as Chaenactis santoli- and very gravelly sandy loam. This edaphic condition
canum
noides Greene, Onagraceae such as Epilobium unique within M. australis because other M.
is all
Number
Volume 423
4
23, Elvin et
al.
2014
Monardella (Lamiaceae) from Southern
California
mm
5
(VFWO)
&
Office
Hartadum Ventura Fish Wildflfe
of
USA
of V«ilwi County, Cslifomuk,
PLmiis
&
Lamtsccaf
A.C. Sanders
fUvin
Sfomtntnllu
CanumHo km of CamanllQ
aulhcm Coasul; nilJs; «0 411.1
SW N <.44nS7,6'N.
4 km of Min.tpaik (air), .iffieiry Road.
scrub
2J6m.775ft, Constnl saac
|ir“i&'16,rw, t.lOOm). tlev
:
sw
from sandstone bedrock. Assoc,
derived
sanih- soiSs
ilb
Mtimrdella simiaia.
ruron (W(fo™j™.t. fforieirofirteotJ,
£rwgtiifuiv
^^^n•r(lm:a cnfifi>rni(ii.
rremuiiJ Ciiii/ontiirtl.
O.
tilKF^tfs. Opuitlilt orrfflAj. Uttonttif--.
lii'tcuSaium., E.
em 50-100
Skflfcc annual lO-JS cm tall. S-15 iviste; fls purptc: ca-
Dutgess
Rk'k
lA
i' VK wo)
vfwCt FwiliBe
irju^d fl 1
lata 1
& &
Figure Monardella sinuata Elvin A. C. Sanders subsp. gerryi Elvin, A. C. Sanders R. A. Burgess. Holotype, M. A. Elvin
4.
&
R. A. Burgess 7121 (UCR), with pedicel and cuplike structure in inset box at top right.
australis subspecies occur on soils derived from in Ventura County, from the same These two
all site.
km
mountain 35
granite. areas are adjacent islands less than
known
With
apart. all collections from only two sites
lUCN
Red
Monardella
List category. australis
AOO
we known
on adjacent mountains, estimate the
subsp. occidentalis appears be exceedingly rare
to
km^
be than 2 km^, with each occurrence.
to less 1 for
and has an extremely There have
limited distribution.
Because have been no M.
been only one from Big Pine there other australis subsp.
five collections,
EOO
be
[Mountain] in the San Rafael Mountains in Santa occidentalis collections, the is likely to small,
Barbara County and four from Pine Mountain Ridge and we estimate be approximately 27 km^, using
to
it
424 Novon
minimum
a convex polygon encompassing the two trichomes on the calyces (lacking in subspecies
occurrences, but only 2 km^, using the occupied gabrielensis).
1
Much
km^ method. between Monardella
grid of the area these australis subsp. occidentalis differs
two occurrences (and on the two mountains) consists from M. australis subsp. cinerea in larger leaves
its
mm mm
X X
4-11 5-10 2-5
(11-17
below do appear
of areas the conifer forests that not vs. in
be suitable habitat. Given extremely limited subspecies cinerea), green vestiture cinerous),
to its (vs.
its
cm
3-15
and (8-22
distribution, the separation of the two occurrences its slightly taller plants vs. cm). It
mm
and
from each and uncertainty regarding climate has two types of glandular trichomes, 0.2
other,
(1)
mm
on
change, M. subsp. assessed 0.4 (sparse) the calyces that are both
australis occidentalis
is
(2)
here CR, Endangered (CR longer than the short glandular calyx trichomes (0.1-
as or Critically
mm)
lUCN
observed subspecies Monardella
according 0.2 in cinerea.
(2001)
B2ab[i,ii,iii,v]c[iii,iv]), to
much
and Of australis subsp. occidentalis has corollas that are
categories criteria. note that of this
is
mm mm
9-11
(13-16
area does not have easy access, and the region has longer vs. for subspecies
been more cinerea).
not intensely botanized; therefore,
may Monardella
subsp.
occurrences be discovered with intensive australis occidentalis is distin-
guished from M. by
australis subsp. jokerstii leaf
fieldwork. its
more
blade margins
that are or less entire serrate
(vs.
Monar
Phenology.
Plants of della australis subsp.
in subspecies by inflorescences as
jokerstii),
its
have been documented between
occidentalis in flower
glomerules one glomerules an
solitary to five in
(vs.
June and but blooms
early early July, likely later
it
compound 13-16
open cyme), by longer
corollas
its
summer
and
into the possibly into early mm
fall.
10-11 mm), and by (8-22
smaller habit
(vs. its
cm
5-35
cm).
The vs.
Etymology.
Latin epithet occidentalis refers
Monardella subsp. oblonga (Greene)
linoides
to the distribution of this subspecies in relation to the
Abrams
occurs in the general region of Pine
M.
other australis subspecies; the westernmost,
is
it
A
Mountain Ridge and Monardella
the north
(to east).
or occidental, of the subspecies.
five
specimen nearby on Sewart Mountain
collected
Taxonomic Monardella
subsp.
australis
affinities. 9433 [VFWO])
{Burgess represents a putative
M.
from
occidentalis differs australis subsp. australis in
specimen between M.
introgressant australis subsp.
mm mm
wider leaves (4-11 2.5-5 in subspecies
vs.
its Some
and M.
subsp. oblonga.
occidentalis linoides
mm
12-18
mm),
(7-12
shorter bracts
australis), vs.
introgression between these two taxa expected
is
mm
and 8.5-10 mm).
shorter calyces (7-8.5
vs. It is
given the observed introgression between M. australis
mm)
by 1-0.2
distinguished short non-glandular
its (0. and M. San San
linoides in the Bernardino,
s.l. s.l.
trichomes on the stems (lacking in subspecies
and San mountains where
Jacinto, Gabriel in areas
mm)
and by 1-0.2 &
non-glandular
australis) short
its (0.
those species occur in close proximity (Elvin
on Monardella
trichomes the abaxial leaf surfaces.
Sanders,
2009).
subsp. lacks the minute (0.04-
australis occidentalis
mm) Paratypes. U.S.A. California: Santa Barbara Co., R.
0.06 glandular calyx trichomes found
in
Hoffmann 644 (CAS, SBBG); Ventura Burgess
Co., R. A.
new
subspecies Rather, the subspecies has
australis.
8921 (PH, UCR), 9479 (CIC, GH, JEPS, NDG, NY, RM,
mm)
short (0.2-0.4 glandular calyx trichomes that are WTU), 3124
US, Chandler (SBBG), 8 June
E. R. 1966,
s.n.
A
not found in subspecies australis, and has longer (SBBG-84125), M. Elvin 7447 (MSC, OSC, UTC,
it
mm VFWO).
(13-16 10-13 mm).
corollas
vs.
Monardella
subsp.
australis occidentalis differs
&
Monardella Novon
sinuata Elvin A. Sanders,
2. C.
from M. wider
australis subsp. gabrielensis in leaf
its
TYPE:
338-340, 2009. U.S.A.
19: 10.
mm mm fig.
blades (4-11 subspe-
vs. [2.2]3.5-5[6] in W
Santa Barbara Burton Mesa,
California: Co.,
and wider glomerule
cies gabrielensis) bracts
its
N
Santa Lucia Canyon, Lakes Canyon, 15
of of
mm
2-4.5 mm), which exceed
versus
or
([3.5J4-5.5
&
Lum
May
M.
2006, A. Elvin, Ballard
L. L.
equal the calyces versus being less than calyx
4904 UCSB; IRVC,
JEPS,
(holotype, isotypes,
by 1-0.2
lengths. distinguished the short
It is (0.
MO, RSA, UCR,
US).
mm)
non-glandular trichomes on stems
the (lacking
in subspecies gabrielensis) and by the short 1-0.2 Monardella sinuata occurs sand dunes and
in
(0.
mm)
non-glandular trichomes on abaxial leaf openings scrub and oak woodlands
in coastal
at
300
surfaces (lacking in subspecies gabrielensis). elevations below m. Monardella sinuata occurs
It
mm)
lacks short 1-0.2 glandular trichomes on from Marin County in northern California Ventura
to
(0.
the calyces (that are present in subspecies gabri- County in southern California. distinguished by
It is
mm)
elensis) and has longer (0.3-0. 6 non-glandular undulating leaf margins and annular habit.
its
Number
Volume 425
4
23, Elvin et
al.
2014
Monardella (Lamiaceae) from Southern
California
Monardella
2a. sinuata subsp. sinuata. glabrous very sparsely pubescent; with 2 types of
to
mm,
triehomes, glandular, 0.02-0.04 and non-
(1) (2)
Monardella on
sinuata subsp. sinuata occurs
mm;
0.3-0.6
glandular, calyx teeth with type of
1
sandy dunes and sandy
soils in relictual areas in
mm;
11-14
trichome, non-glandular, 0.3-0. 6
corolla
scrub and oak woodland
coastal habitats elevations
at
mm,
upper
2 corolla lobes gland-tipped, purple;
300
below m. Monardella
sinuata subsp. sinuata
is
mm, mm,
13-17 13-17
stigma stamens
exserted;
found in coastal areas from Morro Bay in San Luis
Emit brown brown
exserted. a nutlet, light with dark
Obispo County south sand dunes in the Purisima
to
X
mm.
and 6
spots streaks, oblong, 1.5-1. 0.8
Santa Barbara County,
Hills in California.
It is
and
distinguished from M. sinuata subsp. nigrescens Elvin Distribution habitat. Monardella sinuata
&
A. C. Sanders by its taller, yet more gracile, habit subsp. gerryi is endemic to the Las Posas and
and by lacking the darkened stems and bract veins Camarillo west of the Santa Monica Mountains
hills
and apices of subspecies nigrescens. and southeast of the Santa Clara River in Ventura
between
County, California. occurs elevations
at
It
m m
Monardella
2b. sinuata subsp. gerryi Elvin, A. C.
150 and 243 openings scmb.
in in coastal sage
&
TYPE:
Sanders R. A. Burgess, subsp. nov.
Associated species in the microhabitats include a
Ventura
U.S.A. Camarillo
California: Co., Hills, number uncommon
unique and
of taxa typically
May
236 m, 24 M.
sandstone 2013, A.
soils,
found in coastal maritime scrub or other arid
&
UCR;
7121
Elvin Burgess
R. A. (holotype,
environments: Euphorbiaceae such Croton
as cal-
MO, VEWO).
RSA,
JEPS, US, Eigure
isotypes,
ornicus Mull. Arg., Euphorbia polycarpa Benth., and
if
4.
Watson; Polygonaceae such
Stillingia linearfolia
S.
Mucronea
Polemoniaceae such
arnica Benth.;
as
calif
The new
Diagnosis. subspeeies similar Monardel- Mason
is to Eriastrum densfolium H.
as (Benth.) subsp.
&
la sinuata Elvin A. C. Sanders subsp. sinuata, but differs
elongatum H. Mason; and Rosaceae such
(Benth.) as
mm mm
7-8
in having the ealyx shorter (5.5-6 in
vs.
Horkelia cuneata Lindl. subsp. puberula (Rydb.) D.
0.02-0.04
subspeeies glandular ealyx triehomes
sinuata),
mm common
D. Keck. Other associates of the dominant
long (laeking in subspeeies sinuata), sparse eonoideus
glands along the stems (laeking in subspeeies. sinuata), and plant community of the area, coastal sage scmb,
mm
mm).
longer nutlets (1.5- 1.6 1.1
vs. ea.
include Asteraceae such Artemisia
as californica
Eabaceae such Acmispon
Less., as glaber (Vogel)
cm
Annual, 7-30(43) simple
erect, gracile, to
tall,
Lamiaceae such Greene,
Brouillet, as Salvia mellifera
±
moderately branched, glabrous very sparsely
to
and Polygonaceae such Eriogonum
fasciculatum
as
pubescent, stem with type of trichome, non-
1
Monardella
Benth. sinuata subsp. gerryi occurs in
mm,
1-0.2 stems stramineous
glandular,
0. retrorse,
sandy Las Posas Eormation
in the of Pleistocene
soils
to tan, with eonoideus glands sparsely present.
The
Age. soils in this unit are derived from weakly
X
10-35 2-6 mm,
Leaves
with length-to-width
some
sand
consolidated sandstone, with gravelly
±
(L:W) ratios of 5-7:1, blades narrowly elliptic,
and
units, are highly susceptible to landsliding (Tan
glabrous very sparsely pubescent, with type of
to 1
ah, 2004).
et
on 1-0.2
trichome adaxial surface, non-glandular,
0.
mm
lUCN
Red
Monardella
with type of trichome on abaxial surface, List category. sinuata
(rare), 1
mm
and been
non-glandular, 1-0.2 leaves subses- subsp. gerryi exceedingly rare has only
0. (sparse), is
sile to decurrent (rarely cuneate), margins generally collected three times since botanists started to
document
slightly undulate, occasionally strongly undulate, the flora of southern California in the
known
M.
bases acute, apices acute. Inflorescence solitary to 1800s. Currently, sinuata subsp. gerryi is
an open compound cyme; flowers in terminal only from one small extant occurrence in a remnant
glomerules; glomerules 1 to 7 per plant, glomerules patch of fragmented habitat in the Las Posas and
mm
on main stem 10-17 wide, glomerules on Camarillo Of the three M. sinuata subsp. gerryi
axillary hills.
mm
made
branches, 7-12 wide; glomerule bracts 7-9(12) collections, two were over 40 years ago and are
X We
mm;
3.5-6 with 1 type of trichome, non-glandular, considered to be extirpated. estimate that the Las
mm, may
1-0.2 and with 2 types of non- Posas and Camarillo one time have
0. cilia, hills at
(1)
mm
glandular, 1-0.2 and non-glandular, 0.3- supported up 15.64 km^ of suitable habitat before
0. to
(2)
mm
human
modern The
0.5 (sparse), bracts elliptic to widely lanceolate, habitation in the area. Las Posas
±
veins green, translucent between veins, apices and Camarillo have experienced considerable
hills
and
acute to acuminate, purple-tinged or not, less than or residential agricultural development, particularly
mm;
equaling calyces. Elowers with pedicels 1-1.5 in the last 10 to 20 years, which has resulted in a
±
mm,
calyx 5.5-6 rapidly caducous after anthesis, considerable loss of habitat and increased flagmen-