Table Of ContentMadrono, Vol. 56, No. 2, pp. 104-111, 2009
NOTES ON CALIFORNIA MALVACEAE INCLUDING NOMENCLATURAL
CHANGES AND ADDITIONS TO THE FLORA
Steven R. Hill
Illinois Natural History Survey, 1816 S. Oak Street, Champaign, IL 61820
[email protected]
Abstract
The writing of revised treatments for selected California Malvaceae for the upcoming second
edition of the Jepson Manual and the Flora ofNorth America series (volume 6) has made several
nomenclatural changes and explanations necessary. New combinations are made here for taxa in
Sidalcea, including Sidalcea asprella subsp. nana, Sidalcea calycosa subsp. rhizomata, Sidalcea celata,
and Sidalcea sparsifolia. Several taxa previously included within Sidalcea malviflora have been
removed from that species and re-interpreted, resulting in the resurrection and acceptance of the
names Sidcdcea asprella Greene and Sidalcea elegans Greene. Comments are presented here on the
status of Hibiscus lasiocarpos and Lavatera vs. Malva in the California flora. At least one native
species has been added to the flora, namely, Iliamna rivularis, though it may no longer occur in the
state. FourspeciesofMalvaceaehave becomenaturalized orhavebeen found aswaifsin recent years
and are added to the flora, namely, Anoda pentaschista, Lagunaria patersonia, Lavatera olbia, and
Lavatera trimestris.
Key Words: California, Hibiscus, Lagunaria, Lavatera, Malva, Malvaceae, North America, Sidalcea.
The preparation of revised treatments of identify many ofthe individuals conclusively. His
several genera of California Malvaceae—for the hand-written notes on specimens in some herbar-
upcoming revision ofthe Jepson Manual Higher ia also revealed his frustration with these plants
Plants ofCalifornia (TJMl=Hickman 1993) and (e.g., on Blankinship s.n., JEPS 2856, the type of
for the new Flora ofNorth At7ierica, Volunje 6 (in Sidalcea malviflora (DC.) A. Gray var. celata
preparation) led to the need for several nomen- Jeps.,—C. L. Hitchcock wrote: "5*. malvaeflora ssp.
clatural changes as well as an explanation for celata unless I find cause for changing my
some of the changes as compared to previous opinion", —and on JEPS 2855: "5*. malvaeflora
treatments (e.g.. Hill 1993). In addition, several ssp. celata I believe I shall call this").
taxa have been noted that were not included in A previous treatment ofthe genus in California
TJMl for the flora of California. Newly (Hill 1993) attempted to make some sense ofthe
described taxa in Sidalcea have been or shall be species, but it tended to err on the side of
published elsewhere (Hill 2008; Clifton, Buck and combining variants rather than recognizing them
Hill unpublished). to reduce the number of names. After working
several more years with these species as well as
New Combinations and Interpretations with many more both new and old collections, I
IN Sidalcea have attempted to clarify some of the problems
created by combining the variants. Admittedly,
Sidalcea A.Gray is the most species-rich ofthe the changes still have not resulted in a 'perfect'
genera of Malvaceae in California and it is a treatment by any means, but my first goal has
near-endemic there. It is also one of the most been to re-interpret some of the variants. The
perplexingofthegenera taxonomically, and while changes in interpretation are supported by both
several attempts have been made to better define morphological and geographicconsistencyaftera
the taxa (Roush 1931; Hitchcock 1957; Dimling reexamination of type and additional material.
1991; Hill 1993; Andreasen and Baldwin 2001, Second, the work of Andreasen and Baldwin
2003a, b; Andreasen 2005), some remain difficult (2001, 2003a, b) and Andreasen (2005) utilizing
to delineate. The treatment by Hitchcock (1957) new molecular phylogenetic data has helped to
attempted a synthesis using phytogeographic, clarify some ofthe relationships within the genus
morphological, and chromosomal data, and he since the 1993 treatment, and a goal was to
utilized four ranks: genus, species, subspecies, reposition and rename some taxa to incorporate
and variety in an attempt to sharply define the some of the major changes suggested by the
variants. An examination of his treatment molecular work. Among the hypotheses support-
revealed that most species, subspecies, and ed by the new data are the following: 1) the basal
varieties that he described were said to have perennials are Sidalcea hickmanii Greene, S.
transitional individuals to other taxa, and in malachroides (Hook. & Arn.) A. Gray, and S.
somegroups ofspecies it was nearly impossible to stipularisJ. T. Howell &True, 2) fourlineages, or
— —
2009] HILL: NOTES ON CALIFORNIA MALVACEAE 105
clades, ofthe remaining perennial species appear Inheriting this problem, I decided to start over
to be well-supported, the 'malviflora clade\ the and reexamine the type specimens in the group,
'oregana clade\ the ^glaucescens clade', and the keeping this new molecular data result always in
"asprella clade\ 3) within the malviflora clade, the mind. Within this group, a new and undescribed
primarily coastal subspecies of Sidalcea malvi- species of Sidalcea had also been brought to my
flora (subsp. malviflora, subsp. laciniata C. L. attention (Clifton, Buck and Hill unpubhshed),
Hitchc, subsp. patula C. L. Hitchc, subsp. and studies of this as well as the other entities
purpurea C. L. Hitchc, and subsp. rostrata within the 'asprella' and 'glaucescens' clades have
(Eastw.) Wiggins) form a very coherent and helped to resolve the problems to a certain extent.
closely related group, whereas the somewhat I decided to recognize and describe as best I could
more interior subsp. sparsifolia C. L. Hitchc. the morphologically and geographically distinct
and subsp. californica (Torr. & A. Gray) C. L. taxa that sorted out with the new data. Therefore,
Hitchc. (and subsp. dolosa C. L. Hitchc. ?) are I now propose the following nomenclatural and
somewhat divergent from those, and 4) the plants taxonomic changes within Sidalcea.
treated as S. malviflora subsp. asprella (Greene)
Sidalcea asprella Greene subsp. asprella.
C. L. Hitchc. in the 1993 revision are not in the
Bulletin ofthe California Academy ofSciences
same lineage, or clade, as the other subsp. of S. l:78.1885.--Type: USA, California, Yuba Co.,
malviflora, nor are they all necessarily very close
troelaetaedchtoothtehre,mbouutn,taiinnstesapde,ciaesre5'.mogrleauccelsocseenlsy sn.ena.r(Cleacmtpottoynpev,ilhleer,e1deJsuilgn1a8t8e4d,: EC.ALS. G1r12e1e!n)e.
Greene and the foothill species S. robusta Heller Synonym: Sidalcea malviflora ['mcdvaeflora"]
ex Roush. While several additional working (DC.) A. Gray subsp. asprella (Greene) C. L.
Hitchc, University of Washington Publica-
hypotheses can be derived from the molecular
work, these four, especially the latterthree, would tions in Biology 18:25. 1957.
seem to affect the classification and nomenclature
ofthe Cahfornia perennials the most. The inland, Edward L. Greene, in describing this species in
mostly mountain plants that had been tossed into 1885, cited two specimens, and some ofhis other
the 'dust bin' of Sidalcea malviflora subsp. remarks (p. 78) bear repeating: "On bushy
asprella had to be reassessed, and this has been hillsides of the lower Sierras, just below the
habitat of Chamaebatia\ apparently not collected
the emphasis in the recent studies.
before last season; found by Mrs. Curran in El
Hitchcock (1957) considered Sidalcea malvi-
Dorado County, and by the writer on Mr. John
fslpoercaies["mraalnvgaiefnlgorfa'r]omtoBabjea aCashifnogrlneiaw,idMeesxpirceoa,d Ramm's ranch, near Camptonville, in Yuba
County. Peculiar, at least among the perennial
north to the Willamette Valley ofOregon, and he
species, in having the leaves all of precisely the
divided it into 12 rather geographically coherent same shape, the lowest and the uppermost
subspecies, some of which were subdivided into
differing only in point of size. The rough
varieties. Eleven of the subspecies were recog-
nHiiztcehdci.noCfahOfroergnoian,bseuibnsgp.thveirognaltya e(xHcoewpetlilo)n.C.HiLl.l lpeucbteostcyepneceshioswsliktehweisseefveaetruyrecshawrealclt,eraisstdico."maTnhye
(1993) reduced the number ofCahfornia subspe- other specimens from the Sierras. However,
Hitchcock (1957) changed the circumscription
cies to eight, combining Hitchcock's subsp. of this species to include many plants with hair,
celata, elegans, and nana into the single Sierran
habit, and leaf features that did not match the
subsp. asprella partly because of the numerous
comments on transitional individuals in Hitch- ntyupmeeorrouGsresepneec'ismceonnscetphtaitovna.rOyvceornstihdeeyreaabrlsysfirncoem,
cock's 1957 revision. Over the years since, and
after the examination of many more collections, tthhiesosrpiegciineasl, acnondctehpetnhaamvee hbaesenbedcetoemremian'eddusttobibne'
it was decided that this subspecies circumscrip- for difficult Sierra plants. While some variation
tion has become far too broad to be useful, and
certainly appears to be present, the resurrection
refinement has been attempted.
and more precise application ofGreene's original
This group ofdifficult variants resides primar- name, and to a greater extent, his concept ofthe
ily in Andreasen and Baldwin's (2003a, b) species, should prove useful for current and
'asprella clade'. Regarding this 'asprella clade',
future studies.
Hitchcock's treatment and keys were generally
unusable. Plants of very different appearance Sidalcea asprella Greene subsp. nana (Jeps.)
from distant geographical areas and habitats S. R. Hill, comb. nov. Sidalcea reptans var.
would often key to the same subspecies. Andrea- nana Jeps., Flora ofCalifornia, 2:489. 1936.
sen and Baldwin (2003a, b) demonstrated that the Type: USA, California, Trinity Co., Soldier's
Hill (1993)concept ofsubsp. asprellawas actually Ridge, SE Trinity Co. (Yollo Bolly Moun-
polyphyletic, and their different samples of that tains), 24 Jul 1897, W. L. Jepson 14061
subspecies did not cluster together in the final (holotype: JEPS 2856!; isotype: JEPS 2858!).
analysis. Synonym: Sidalcea malvijlora ['malvaeflora']
—
MADRONO
106 [Vol. 56
—
subsp. nana (Jeps.) C. L. Hitchc, University of Jeps., Flora of CaUfornia 2:493. 1936. Type:
Washington Publications in Biology 18:29. USA, Cahfornia, Shasta Co., Olinda, 11 May
1957. 1911, /. W. Blankinship s.n. (holotype: JEPS
2856!; isotype: JEPS 2855!, possibly WIS!—but
Willis L. Jepson, in describing this as a variety dated 16 Apr 1911).
in 1936, considered it to be a close relative of
Sidalcea reptans Greene because of its very long The two JEPS specimens are quite different at
thin rhizomes. The specimens on the type sheet first glance - JEPS 2856 is a single stem with
are dwarfed, and all of the inflorescences are nicely spread leaves, and JEPS 2855 has 2 stems
<10 cm—long. Not all specimens are as small as and a good caudex, but the leaves are badly
the type instead, robust individuals can reach a wrinkled. The two together supply a good series
height of as much as 40 cm, yet they still share ofcharacters to define the species, however JEPS
the other important diagnostic characters. The 2856 bears a label indicating it is the type, and
morphological features match those ofS. asprella JEPS 2855 bears a label indicatingit is an isotype.
subsp. asprella well, but the subspecies is Both bear several annotation labels by the
distinctive in its long slender rooting rhizomes experts, including C. L. Hitchcock whose com-
and its fewer-flowered, often short inflorescences ments have been included above. While the lower
and occasionally few-leaved (1-3) stems that are leaves ofthe holotype have some resemblance to
decumbent-based. Otherwise the hairs through- those of the type of 5'. asprella, other characters
out and toothed leaflobes are a close match to S. do not fit that species; some of the contrasting
asprella subsp. asprella. Studies by Andreasen features include the lack of rhizomes, the
and Baldwin (2003a, b) show this plant to be, presence of stiff reflexed bristle hairs at the base
perhaps, closer to S. glaucescens than S. asprella, of the stem (a primary character for S. celata),
but more samples are needed to test this as the and the upper leaves have narrow, often entire
morphology does not support this placement lobes, whereas S. asprella, as here defined,
(e.g., S. glaucescens and the very similar S. generally has some short rhizomes, coarse stellate
miiltifida Greene do not have elongated rooting hairs at the stem base, and upper leaves that are
rhizomes of any kind). Certainly this subspecies, somewhat similar to those below, with wider
as well as the typical subspecies, can no longer be lobes that are generally toothed. Upon using
included within the more coastal S. malviflora these characters on additional specimens, I
based upon the molecular data, and so a new discovered that Sidalcea celata is a species that
name was needed. is rather narrowly distributed in dry open oak
Sidalcea calycosa subsp. rhizomata (Jeps.) woodlands mostly associated with serpentine in
Munz ex S. R. Hill, comb. nov. Sidalcea Shasta and adjacent Tehama Cos., whereas S.
rhizomata Jeps., Manual of Flowering Plants asprella appears to be widely distributed in the
of California 629. 1925.—Type: USA, Cali- central and northern Sierra Nevada range and to
fornia, Marin Co., Point Reyes Peninsula, the northwest, at the margin of more mesic
marsh near Russell's Creamery, 16 Sep 1900, coniferous woodlands, either associated with
W. L. Jepson 1174 (holotype: JEPS 2861!; serpentine and serpentine-like minerals, or not.
isotypes: JEPS 2859! MO!—not yet acces- Sidalcea elegans Greene, Cybele Columbiana
sioned with a number at the time of its 1:35. 1914.—Type: USA, Oregon, Josephine
inspection). C
Co., Eight Dollar Mountain, 12 Jun 1904,
V. Piper 6171 (holotype: US 527772!; photo-
There are two sheets of the type collection at graph of holotype at MO 940080!). Synonym:
JEPS; the sheet accessioned as JEPS 2861 has
Sidalcea malviflora ['malvaeflora'] (DC.) A.
Jepson's handwritten designation as 'type' on it, Gray subsp. elegans (Greene) C. L. Hitchc,
and includes the fertile material; the duplicate University ofWashington PubHcation in Biol-
sheet, JEPS 2859, consists ofthe sterile, creeping, ogy 18:27. 1957.
rooting rhizomes with scattered leaves also
characteristic of this subspecies. Munz (Munz
annadmeKec''kSi1d9al5c9eap. c1a3l2y;cMosuanzM.196E8.p.Jo1n2e)susseudbstph.e froSimdatlhceeoatheelregmaensmbiesrrsatohfertheeas'ialsypredlilsaticnlgaduei'shbeyd
rhizomata (Jeps.) Munz," but did not validly means ofthe relatively long, soft, simple hairs at
publish this new combination there or elsewhere. the base ofthe stem, sometimes so sparse as to be
He, perhaps, did not realize that post-1952 new nearly lacking. The stems are characteristically
combinations require direct references to the brittle and easily snapped when fresh, a character
not mentioned for other taxa in the genus (but
basionyms. Its inclusion here serves to validate
not especially useful on herbarium specimens!).
the combination.
The upper stems are sometimes glaucous, and
Si—dalcea celata (Jeps.) S. R. Hill, comb. nov. because of that feature as well as the one-sided
Sidalcea malvijlora {'malvaejlora''] var. celata inflorescences that are often slightly curved
—
2009] HILL: NOTES ON CALIFORNIA MALVACEAE 107
between the flowers, the long acuminate calyx Hitchcock (1957) divided his Sidcdcea malviflora
lobes, and the decumbent stems the species has subsp. sparsifolia further into four morphologi-
sometimes been reported as Sidalcea glciucescens, cally defined varieties that also have some
a species without the long rooting rhizomes ofS. geographic coherence. These, upon further study,
elegansand that isnot known in Oregon. Sidalcea both morphological and molecular, perhaps
elegans appears to be restricted to serpentine, and could be defined as subspecies of this newly
it is found in the Klamath Mountains of circumscribed species. I have not yet focused on
California and Oregon. Roush (1931) treated this this aspect in the current study, but some of the
taxon as a synonym ofS. asprella and Hitchcock extremes are not only quite different in appear-
(1957) stated that "Ifasprella were to be treated ance, but they are also somewhat difficult —to
as a species, ssp. eleganswould best beconsidered separate from S. malvifora subsp. cal(fornica
thereunder." In contrast, Dimling (1991) stated another mostly inland taxon that needs addition-
''Since this subspecies [S. malviflora ['malvae- al study.
floixf] subsp. elegans] is so clearly distinct from S.
malvaeflora ssp. asprella, its taxonomic identity Sidalcea diploscypha vs. Sidalcea keckii
will not be discussed further". Hill (1993) treated
it as a synonym of S. malviflora {'malvaeflora') Sidalcea keckii Wiggins has been of special
ssp. asprella and left it within that variable interest in California because it was once thought
complex. In an attempt to clarify its position to have been extirpated (Hill 1993) then, upon
and nomenclature, the name Sidalcea elegans beingrediscovered at a laterdate, it was proposed
Greene is here resurrected, because the taxon is and accepted for inclusion in the Federal Register
not a part of S. malviflora based on molecular as a Federally Endangered plant species (United
evidence, and because it appears to have several States Fish and Wildlife Service 2000). It was
features that separate it easily from S. asprella. thought to be restricted to the White River region
of southern Tulare County. In the years since it
Sidalcea sparsifolia (C. L. Hitchc.) S. R. Hill, was rediscovered, it has been sought out there
comb. nov. Sidalcea malviflora ['malvaeflora'] and elsewhere. It was known since its original
subsp. sparsifolia C. L. Hitchc, University of description to be very closely related and similar
Washington Publications in Biology 18:32. to Sidalcea diploscypha (Torr. & A.Gray) A.Gray
UL11.9Tm5HiCl7iet.8c—8shT1ocy8uop4tc!ehk:DJoS9fU5S4F3tA66.,8(0hT3oeC6ljaoolatnity,fpoCer2A:n9Si!Wa)MT,.aUyK;eri1s9no5t2y,Cpoe.Cs.,: ifhpnluaobvweeietssrcsneaunanmcnneeud—raolfuarssuihotasnbl,ilotyna,gntdhifetisseneeslpteepawcfeioarlpmaleonynrndpurihaecolgulalroasdirgdiyan,lhgcaieiirattssss
along the stem. The molecular work of Andrea-
Andreasen and Baldwin (2003b) included this sen and Baldwin (2001, 2003b) and especially
plant in their molecular studies of Sidalcea, and Andreasen (2005) utilizing new molecular phylo-
found that it grouped generally with the coastal genetic data demonstrated convincingly that the
subspecies of S. mcdviflora as well as with S. two species are distinct. It has been proposed by
covillei Greene, S. pedata A. Gray, and S. some that S. keckii is quite recognizable because
neomexicana A. Gray. They stated: "The position on the inside of the calyx there are five reddish
of S. malviflora subsp. sparsifolia basally to the spots thought not to be present in S. diploscypha.
clade (jk 76%) consisting of the other subspecies A re-investigation of the two species from both
of S. malviflora plus S. pedata and S. neomex- old and new herbarium specimens revealed that
icana, provides evidence for the paraphyly of S. both species can have these red spots (sometimes
malviflora and may justify treatment of S. reduced to narrow red lines) on the internal calyx
malviflora subsp. sparsifolia as a separate spe- surface. Specimens sorted out well using other
cies". I agree with this, not only because of the characters, to the point that it appears that
molecular data but because of its more inland Sidcdcea keckii is more wide-ranging than previ-
range and semi-desert habitats, as well as a series ously thought.
of morphological differences. It is rather similar Several morphological features can be used to
to the other species in the clade particularly in the distinguish the tw—o similar species. Regarding the
morphology ofthe fruits and ofthe pubescence of hairs on the stem one group of specimens (first
its various parts. The reduced stem leaves and identified as S. diploscypha) had, mixed with the
shortened rhizomes may be adaptations to its characteristic long perpendicular hairs, a consid-
transitional hot and dry desert environment, and erable number of glandular hairs and odd
it is the southernmost species of the genus in multicellular trichomes with green pigment (that
southern California and Baja California, along resembled short algal filaments) on both the
with its desert wetland-adapted relative S. neo- upper stems and on calyces and these were found
mexicana. also on specimens known to be S. keckii. This
Extreme variation remains problematic in this same group of specimens had one or a very few
species despite its removal from 5". malviflora. tiny bristles on the upper portion of the fruit
— ——
MADRONO
108 [Vol. 56
where many sidalceas have a smallcusp ormucro lb. Upper paired stipules and bracts each simple,
(sometimes called a 'beak') whereas S. diploscy- linear, undivided (a few divided in robust
pha has no such bristles on its fruit. The upper plants)generallyshorterthancalyx;lengthof3
leaves of S. keckii are not only lobed, typical of apical teeth of widened middle leaf lobe on
upper stem leaves essentially equal; inflores-
both species, but the lobe tips have three equal cence, calyx generally with many minute
toefetthheonuptpheerwliedaevnesedofloSb.edaippelxoswchyeprheaaasrethuesluoablelsy glandular, multicellular simple hairs . . . . S. keckii
narrow throughout, and either entire or the
Hibiscus lasiocarpos
lateral teeth are positioned far below the central
elongated tooth. The primary difference between The Cahfornia Hibiscus was treated by Hill
the two species is the presence of long, multi- (1993) as part of the widespread Hibiscus
divided bracts and stipules at the base of the
lasiocarpos Cav. in accordance with the opinions
flowers and upper leaves, respectively, of S. of Fryxell (1988) and others. The California
diploscypha vs. the smaller undivided bracts and populations (primarily in the Sacramento Valley)
stipules in the same positions in S. keckii. Upon remain quite scarce and isolated from any other
examining a large num—ber of specimens, this populations of this species, the closest of which
feature did hold up well bu—t there were always a are in northwestern Chihuahua, Mexico, and in
few that did not 'quite fit' and so a very few Dona Ana Co., New Mexico. Its scarcitycausesit
specimens with divided bracts were called S. to be ofconservation concern in California (List
keckii. As a result ofthesecharacterobservations, 2: Plants Rare, Threatened, or Endangered in
several specimens ofS. diploscypha from Colusa, California, But More Common Elsewhere). I
Fresno, Merced, Napa, Solano, and Yolo coun- have reconsidered this restricted Cahfornia plant,
ties were re-annotated as S. keckii. Most ofthese and I have decided to recognize it as Hibiscus
had some features of S'. diploscypha, and it lasiocarpos Cav. subsp. lasiocarpos var. occiden-
appeared that introgression might be playing a talis (Torr.) A.Gray. Its nomenclatural history
role. This geographic distribution suggested that follows.
specimens of 5'. keckii might also be found in
Butte and Lake counties but no specimens Hibiscus lasiocarpos Cav. var. occidentalis
examined from those counties had the definitive (Torr.) A.Gray, Proceedings of the American
assemblage ofcharacters ofthat species and so all Academy ofArts and Sciences 22:303. 1887 [4
were annotated as S. diploscypha. —Mar 1887] (as "'lasiocarpus var. occidentalis'').
Sidalcea diploscypha appears to be a species Hibiscus moscheutos L. var. occidentalis
that prefers serpentine, whereas S. keckiiis not so Torr., United States Exploring Expedition,
restricted. An examination of habitat and sub- Phanerogams. Pacific North America
strate preferences in Napa Co. where both 17(2):256. 1874.—Type: USA, Cahfornia,
substrates and species have been found nearly Sacramento Co., Sacramento Valley, s.d.,
side-by-side, showed that those on serpentine Wilkes Expedition 1364 (holotype: NY).
sorted out nicely to S. diploscypha and those in Hibiscus lasiocarpos var. californicus (Kellogg)
the adjacent sandstone-derived soils were S. L. H. Bailey, The Standard Cyclopedia of
keckii though individuals were often only a few Horticulture 1486. 1915. Hibiscus californicus
meters distant from one another (B. Ertter, [UC/ Kellogg, Proceedings of the Cahfornia Acad-
JEPS], J. Ruygt, personal communication). More emy of Sciences 4:292. 1873.—Type: USA,
molecular work on these newly interpreted California, San Joaquin Co., on island near
populations from Solano to Colusa counties Middle River bridge, San Joaquin River
may help to further unravel the relationship (Byron-Stockton Hwy) Alexander & Kellogg
between these two taxa, but, as interpreted now, 3526 (neotype: CAS; isoneotypes: UC, US)
the Federally Endangered S. keckii, while still designated by P. A. Fryxell, Systematic Botany
exceedingly uncommon, is now reported for Monographs 25:211 (1988).
seven counties, rather than just one or two, as
previously thought. The number of populations This variety continues to be included here
currently extant is still unknown. The following within Hibiscus lasiocarpos Cav. That species,
key is offered to distinguish the two species: including the Cahfornia plants, has recently been
treated by Blanchard (2008) as H. moscheutos L.
subsp. lasiocarpos (Cav.) O. J. Blanchard. At
la. Upper paired stipules (at petiole bases) and least one flora (Gleason and Cronquist 1991) has
bracts (at pedicel bases) each divided to base equated Torrey's California variety with all of
into 2 or more linear lobes nearly equal to or
longer than calyx; length of central tooth of H. lasiocarpos, calling it H. moscheutos var.
middle leaf lobe on upper stem leaves much occidentalis Torr. That concept is not accepted
longer than lateral teeth, so lobe has a single here because Torrey's type is from California
apicaltooth, orlobesentire;inflorescence,calyx and clearly represents only the isolated Cali-
generally not densely glandular .... S. diploscypha fornia population here included, following Gray's
2009] HILL: NOTES ON CALIFORNIA MALVACEAE 109
example, within H. Icisiocarpos. While there is related to Malva sylvestris L., the type species of
somevariation in pubescence in thesetwo species, Malva, than they were to Lavatera trimestris L.,
treatments such as Godfrey and Wooton (1981) the type species of Lavatera, by means of both
and Mohlenbrock (1986) use this feature in sets of data. Both genera are still accepted, but
particular to distinguish these species. Other circumscriptions have changed, and the closely
treatments of the Malvaceae (e.g.. Hill 1982) related taxacould no longer bemaintained within
and most world floras use some or many hair two separate genera. Therefore, the realignment
characters to distinguish mallow species from one of species within Lavatera had to be formalized.
another, as this family has hair types varying Nomenclature for the species of Malva included
from simple to bilateral several-rayed hairs, in TJMl remain the same.
multi-rayed stellate hairs, glandular hairs, as well Ray (1998) chose to maintain both genera with
as stellate-lepidote hairs in varying combinations the types as stated above, and he defined them
and densities that usually remain surprisingly not only by their ITS characters, but also by a
consistent within a taxon. I continue to accept series of mericarp features. Malva was distin-
Hibiscus moscheutos L. and Hibiscus lasiocarpos guished from Lavatera primarily by its mericarps
Cav. as distinct species based primarily on the that 1) are rounded in only the axial direction on
following characters: the abaxial side, 2) have lateral angles oredges, 3)
completely or nearly completely enclose the seed,
la. Upper leaf surface glabrous or sparsely 4) do not separate readily from the seed, and 5)
pubescent, darker than the densely felty- act as a dispersal unit. This group contains not
cpaupbseuslceesntgllaobwreorussurofracsepaersspeelcyialpluybewshceenntdroyn; only cosmopolitan weedy species formerly in-
sutures; involucral bracts uniformly, minutely cluded in both genera, but also several unusual
canescent Hibiscus moscJieutos L. disjunct taxa in Australia, Baja California,
lb. Upperleafsurfacedensely soft-pubescent like Mexico, and California, USA, that had formerly
the lower surface, and usually similar in been treated within the genera Lavatera or
color; capsules densely pubescent through- Saviniona Webb & Berthelot.
out; involucral bracts densely coarsely pu- Ray (1998) proposed new combinations and
bescent with both short-stellate hairs and new names for several ofthe taxa formerly placed
longer spreading simple hairs especially near
the margins Hibiscus lasiocarpos Cav. within Lavatera. Consequently, all three Lavatera
species in TJMl are now considered to belong in
According to this species concept, the Califor- Malva. Ray (1998) proposed the name Malva
nia plants fit within H. lasiocarpos. This dendromorpha M. F. Ray as a substitute for
CaUfornia variety was distinguished by Bailey Lavatera arborea L., as he thought that Malva
as having more uniformly cordate leaves and a arborea was a name already taken and unavail-
less hairy capsule than the typical variety. able. However, the name that he cited, ""Malva
Furthermore, individuals of this taxon charac- arborea St.-HiU', was neverpublished (a Sphcdma
teristically produce long starchy rhizomes from typographicum (misprint) in Index Kewensis),
which they often propagate themselves in their and so the next available name, Malva arborea
native habitat, marshes and deltaic areas subject (L.) Webb. & Berthelot (1836. Histoire Naturelle
to unreliable water levels (e.g., Sacramento, 28 des lies Canaries, pt. 2. Phytographia Canariensis
Sep 1989, C M. Richard 098928 [OAKL: 4 1:30.) based on the Linnaean name is the correct
sheets]). This variety often exceeds 2 m in height, name in Malva and Malva dendromorpha M. F.
and the capsules are globose and 2.5-3 cm, Ray becomes superfluous. Ray (1998) also
whereas the typical variety normally grows to proposed the name Malva liimaei M. F. Ray to
2 m or less (not infrequently <1 m tall) and has replace the name Lavatera cretica L., as Malva
subglobose or short-cylindic capsules 2-2.5 cm cretica Cav. had already been used for a different
long. plant. However, it was brought to my attention
(Hinsley 2009) that the name Malva pseudolava-
LavATERA vs. Malva tera Webb & Berthelot (1836. Histoire Naturelle
des lies Canaries, pt. 2. Phytographia Canariensis
The genera Lavatera L. and Malva L. have 1:29.) had been proposed as a substitute name for
undergone a significant revision since the publi- Lavatera cretica long before Ray's substitute
cation ofTJMl. Studies by Ray (1994, 1995) on name, and can be considered to be the correct
nuclear rDNA Internal Transcribed Spacer (ITS) name for the plant, making Ray's name super-
sequence data as well as morphological features fluous. For the third California Lavatera, Ray
led to the conclusion that the species of Malva (1998) proposed the new combination Malva
and Lavatera are all closely related, and that a assurgentiflora (Kellogg) M. F. Ray for the
significant number of species assigned to both indigenous species formerly called Lavatera
genera were more closely related to one another assurgentiflora Kellogg, and this is now its name
than previously thought. In particular, several in Mcdva. Further study may indicate that there
species in Lavatera were found to be more closely are two distinct subspecies within this coastal
MADRONO
110 [Vol. 56
California species as suggested by Philbrick garden, Victor Reiter's garden, 1195 Stanyon
(1980). St., San Francisco, 4 Aug 1970, T.C. Fullers.n.
Lavatera itself as currently defined (Ray 1998) (CDA 5008).
is only rarely found as an introduced plant in Lavateratrimestris L. CALIFORNIA. Santa
North America and in California in particular. Barbara Co.: edge of water, Lauro Canyon
Both L. olbia L. and L. trimestris L. have been Reservoirnear San Roque Rd., Santa Barbara,
found as waifs in the state (see below). A third 25 Jun 1975, C. F. Smith 10902 (CDA 5007;
species, Lavatera thuringiaca L., has been found RSA 535067). Cited in Smith (1976).
rarely as an escape in more northern parts of
North America. Acknowledgments
I thank Katarina Andreasen and Bruce Baldwin for
Additional Malvaceae in the Flora sharing their data on these plants and for stimulating
correspondence concerning these taxa in recent years.
Ilianma rivularis (Dougl.) Greene was not Others have also been generous with their time and
included in the flora of California in 1993 experience with these plants, including Barbara Ertter,
(Hickman 1993). Two specimens collected by Richard Halse, Stewart Robert Hinsley, Fred Hrusa,
Lawrence Janeway, Kanchi Gandhi, Jake Ruygt, and
JHousmebpohldtP.CoT.rarcecyentolny cAaumgeustotli1gh4t,. T1h9e39y,haidn tDoebtrhaanTkrocBkr,ucteo mGe.ntiBoanldawifnew.aIndwouthled eLsapwecrieanlclyeliRk.e
been overlooked for many years and there are no Heckard Endowment Fund of the Jepson Herbarium
other known records of this native species in for the fundingto visit the UC/JEPS and CAS herbaria
California, though it is much more common tostudyherbariumspecimensinsupportofmyworkon
north of the state. The vouchers for this species several Malvaceae genera for the upcoming revision of
are: CALIFORNIA. Humboldt Co.: Willow the Jepson Manual: Higher Plants of California. The
Creek Canyon, along Trinity Highway, in woods UC/JEPS and CAS staff were especially hospitable
near stream, altitude 2500 feet, 14 Aug 1939, /. during my visit and very generous with their time. A
Pis.dTorubatcfyu1l6t1h0a4t i(tMstOill1e1x9is1t8s77i!n,thMeOsta1te1,91b8u7t8!t)h.erIet cTsorunaptnpisonpruotirnttgaotwigaorrnadnisstgrrfearsteeofamurlclhtyheaacnkIdlnloipwnuolbiesldigcDeaetdpioafnorrtmofifennattnhceiosalef
is always the chance it persists. It should also studies
be sought in Lassen or Modoc counties where
there is suitable habitat. Iliamna latibracteata Literature Cited
Wwfreiirgdgoiegmoinan/s.nsdriinivlsuolnHawgrueimslvsbl.bo-ykl2ndimottswmnwCiwodfiuerdnroetmybr,Xarc4teal.dne5wdt-so6o(imdtcam.dfio1flrfoceensrmgts AndrasMeyansladtvseaetmcahnetr,aieeca.tKeaC.nnoeandl2sy0est0rea5svx.aa:toiInocmonptnlhtGierecanasectttoiiinoncsngsesre6v:xao3atf9im9opm-nlo4e1lso2ef.cfurrlaoarmre
in /. rivularis) and its dense pubescence on the and B. G. Baldwin. 2001. Unequal evolution-
undersides of the leaves (hairs sparse in /. ary rates between annual and perennial lineages of
rivularis). Iliamna bakeri (Jeps.) Wiggins is found checker mallows (Sidalcea, Malvaceae): evidence
in more inland chaparral sites in northern from 18S-26S rONA internal and external tran-
California, and it has more shallowly lobed scribed spacers. Molecular Biology and Evolution
lpeetaivoelsesantdhanshotrhteerot(h2e-r5tcwom. vs. >5 cm), stouter 18:9A3N6-D944. . 2003a. Nuclear ribosomal DNA
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checker mallows {Sidalcea, Malvaceae). Molecular
naturalized or have been found as waifs in recent Phylogenetics and Evolution 29:563-581.
years and are added to the flora. AND 2003b. Reexamination ofrelation-
.
Anoda pentaschista a. Gray. California: ships, habital evolution, and phylogeography of
checker mallows {Sidalcea, Malvaceae) based on
Imperial Co.: Collins and Flood, Bard, weed in molecular phylogenetic data. American Journal of
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Lagunariapatersonia (Andr.) G. Don. CAL- 18:4-8.
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