Table Of ContentMadrono, Vol. 48, No. 4, pp. 293-297, 2001
HARMONIA GUGGOLZIORUM (COMPOSITAE-MADIINAE), A NEW
TARWEED FROM ULTRAMAFICS OF SOUTHERN MENDOCINO COUNTY,
CALIFORNIA
Bruce G. Baldwin
Jepson Herbarium and Department of Integrative Biology,
1001 Valley Life Sciences Building #2465, University of California,
Berkeley, CA 94720-2465
Abstract
Harmonia guggolziorum is a new tarweed from ultramafic (serpentine) soils of southern Mendocino
County, California. Unlike other species ofHarmonia, H. guggolziorum combines the following morpho-
logical characteristics: primary stems usually longerthan branches ofthe subumbelliformcapitulescences,
leaves unevenly distributed but not densely congested, heads erect in bud and fruit, phyllaries irregularly
hirsute, disc florets bisexual, ray cypselae weakly arcuate, ray pappi present, and disc pappi of linear,
mm
fimbriate scales 0.6-0.8 long. Based on molecularphylogenetic data, 1 suggest that H. guggolziorum
is the only known representative ofa lineage that predates diversification ofthe other serpentine endemic
species of Harmonia (i.e., H. doris-nilesiae, H. hallii, and H. stebbinsii). The apparent phylogenetic
relationships and geographic location of H. guggolziorum lead me to hypothesize that Harmonia origi-
nated in the southern NorthCoast Ranges and has undergone more extensivediversificationonultramafics
than previously suspected for the genus or any other lineage in Madiinae.
mm
California's exceptionally rich serpentine flora is squamis papporum radiorum fimbriatis, ca. 0.5
especially well represented in the northwestern part longis; squamis papporum discorum linearibus fim-
mm
of the state (Kruckeberg 1984, Harrison et al. briatis ca. 0.6-0.8 longis.
2000), where botanical exploration has continued
to reveal previously unknown ultramafic endemics. Annual herbs. Stems erect, branched mostly in
In the tarweed genus Harmonia B. G. Baldwin distal half, slender, mostly reddish-purple, to 3 dm
[=Madia Molina sensu Keck (1959) pro parte (i.e., high, sparsely to densely hirsute proximally, dense-
the yellow-anthered, pappose annuals, with 2n — 9 ly stipitate-glandular distally, the glands dark-pur-
II); see Baldwin (1999)], two ultramafic endemics plish (or yellowish). Leaves opposite proximally,
(H. doris-nilesiae and H. stebbinsii) have been de- alternate distally, sessile, mostly cauline, unevenly
scribed from the North Coast Ranges of California distributed, mostly proximal on primary stems and
since 1980 (Nelson and Nelson 1980, 1985). Here- at bases of capitulescence branches (otherwise
in, I describe yet another species ofHarmonia from sparse or absent), ascending or usually widely
the North Coast Ranges. spreading, often with reflexed apices; blades linear
mm mm
to filiform, 5-50 long (mostly 20-25 long
Harmonia guggolziorum B. G. Baldwin, sp. nov. on primary stems), 1-3 mm wide, entire or sparsely
(Fig. 1)—TYPE: USA, California, Mendocino and shallowly toothed, slightly revolute, hirsute,
Co., on serpentine on the north side of Feliz eglandular or (mostly in capitulescence) stipitate-
Creek Road (County Road 109), 2.1 miles west glandular (especially near apices), the glands dark-
of Hopland (T13N, R12W, S23, NWl/4 of purplish (or yellowish). Capitulescences subumbel-
NEl/4), ca. 150-200 plants associated with Pla- liform, branches often 5-7 cm long (max. 15 cm
tystemon californicus. Cilia capitata, and Cryp- long) and overtopping the nearly sessile head ofthe
tantha clevelandii, 164 m, 30 April 2000, Jack primary stem. Peduncles 2-12 mm long, stipitate-
and Betty Guggolz. 1635 (holotype, JEPS; iso- glandular, the glands dark-purplish (or yellowish).
type, CAS). Heads usually erect in bud, anthesis, and fruit. In-
volucres obovoid, ca. 3-4 mm diam. (4-5 mm
Ab species ceteris Harmoniae characteribus com- diam. in pressed specimens). Phyllaries (3-) 5 (-6)
binatis differt caulibus primariis plerumque ramis (1 per ray floret), uniseriate, herbaceous, linear, 4-
mm
subumbelliformium capitulescentiarumlongioribus; 5 long, each completely enveloping a ray ova-
foliis distributis impariter, non dense congestis, ry, the free apices purplish, erect or spreading, flat
plerumque proximalibus in caulibus primariis et ad or involute, <l/5 the length of enfolded basal por-
basibus capitulescentiarum; capitulis plerumque er- tion of phyllaries; abaxial faces irregularly and of-
ectis ante, per, et post anthesin; phyllariis irregular- ten sparsely hirsute with broadly arching or some-
iter et saepe sparsim hirsutis cum pilis prope mar- what appressed hairs, often with soft, matted hairs
gines moUibus saepe implicitis; flosculis discorum near margins, ciliate, irregularly stipitate-glandular,
bisexualibus; cypselis radiorum leniter arcuatis; the glands dark-purplish (or yellowish). Rayflorets
294 MADRONO [Vol. 48
Fig. 1. Harmonia guggolziorum. (A) habit; (B) head; (C) phyllary, ray floret, palea, and disc floret (right to left); (D)
adaxial view of ray floret and associated phyllary; (E) disc floret; (F) palea; (G) ray cypsela and pappus; (H) disc
cypsela and pappus.
2001 BALDWIN: HARMONIA GUGGOLZIORUM 295
10 changes
H. nutans
H. guggolziorum
rC H. hallii Serpentine
clade
H. stebbinsii
Putative serpentine- —
endemic ancestor
H. doris-nilesiae
Fig. 2. Relationships in Hannonia based on 18S/26S nuclear ribosomal DNA sequences ofthe external and internal
transcribed spacer regions (Baldwin, unpublished). The tree is rooted with sequences from the other diploid members
ofthe "Madia" lineage (Baldwin 1996).
(3-) 5 (-6), pistillate, corollas bright yellow, tubes free or weakly fused proximally. Ray cypselae
mm
ca. 1.5 long, sparsely hirtellous, laminae broad- black, slightly laterally compressed, abaxially
mm
ly overlapping in head, flabelliform, 4-5 long, rounded, adaxially angled, clavate, weakly arcuate,
5-7 mm wide, 3-lobed to ca. half length, glabrous. ca. 3-3.5 mm long, glabrous, beakless. Ray pappi
Discflorets 8-13, bisexual, corollas bright yellow, of ca. 10-12 stramineous, linear, fimbriate scales
2-3.5 mm long, tubes much shorter than the nar- ca. 0.5 mm long. Disc cypselae black, ±terete to
mm
rowly funnelform throats, lobes 5, glabrous abaxi- clavate, straight or weakly arcuate, ca. 3-3.5
ally, densely bristly adaxially. Anthers yellow. Style long, with antrorse hairs. Disc pappi of ca. 9-11,
bglraabnrcohuess. aPcaulmeianeatneo,thpiesrpsiidsutlenotu,s.caR.ec8e,ptiancloensefplaet-, fsitmrbarmiianteeousscaolres,pur0p.l6i-s0h.,8limneamr, l±ofnlga.t (Cnhortocmriosspoedm)e,
ripheral series, linear, 4-5 mm long, herbaceous number In = 9 II [reported here from B. G. Bald-
near apices or throughout, sometimes chartaceous win 1140 (JEPS)].
proximally (especially along margins), flat or with Paratype. USA, California, Mendocino County,
margins partially enveloping a disc ovary, sparsely on serpentine hillside at junction of Feliz Creek
hirsute and densely ciliate near apices, sparsely Road (County Road 109) and County Road 110,
stipitate-glandular near apices, the glands dark-pur- west of Hopland, 8 May 2001, B. G. Baldwin 1140
plish (or yellowish), the margins of adjacent bracts (JEPS).
Key to Species of Harmonia
1. Heads usually reilexed in bud and fruit; ray pappi 0; disc pappus elements lance-attenuate, fimbrillate, 2-3.7
mm
long H. nutans (Greene) B. G. Baldwin
r Heads usually erect in bud and fruit; ray pappi present (often rudimentary); disc pappus elements subulate,
mm
linear, oblong, or quadrate, fimbriate or pulmose, 0.2-3.5 long.
2. Leaves ± evenly distributed along stems; ray cypselae gibbous (bowed out abaxially), distinctlybeaked(beaks
<1 mm long); disc florets functionally staminate
H. doris-nilesiae (T. W. Nelson & J. P. Nelson) B. G. Baldwin
2' Leaves unevenly distributed, mostly restricted to proximal stems and bases ofsubumbelliformcapitulescences;
ray cypselae weakly arcuate, beakless; at least some disc florets bisexual.
mm
3. Phyllaries pilose near margins; disc pappus elements subulate, 1.2-3.5 long, plumose
H. stebbinsii (T. W. Nelson & J. R Nelson) B. G. Baldwin
3' Phyllaries with inconspicuous, soft, often matted hairs near margins; disc pappus elements linear, oblong, or
mm
quadrate, 0.2-0.8 long, fimbriate.
4. Primary stems usually shorter than branches of the subumbelliform capitulescences; distal leaves ofprimary
mm
stem densely congested; disc pappus elements oblong or quadrate, 0.2-0.5 long
H. hallii (D. D. Keck) B. G. Baldwin
4' Primary stems usually longer than branches of the subumbelliform capitulescences; distal leaves of primary
mm
stem not densely congested; disc pappus elements linear, 0.6-0.8 long .... H. guggolziorum B. G. Baldwin
Relationships. Based on phylogenetic analyses of nilesiae, H. hallii, and H. stebbinsii from a common
DNA
18S/26S nuclear ribosomal sequences of the ancestor; H. guggolziorum is the sister group ofthe
external and internal transcribed spacers (Baldwin, lineage corresponding to H. doris-nilesiae, H. hall-
unpublished), H. guggolziorum represents a basally ii, and H. stebbinsii. Support for the hypothesis that
divergent lineage in a monophyletic group com- H. guggolziorum represents an ancient, divergent
prising all serpentine endemics in Harmonia (Fig. lineage in Harmonia rather than a recent product
2). Origin ofthe lineage represented by H. guggolz- ofhybridization comes from unique character-states
iorum apparently predates divergence of H. doris- at eight rDNA nucleotide sites in H. guggolziorum
MADRONO
296 [Vol. 48
and from four rDNA mutations shared by the other sympatry [V. Parker 757 (JEPS) and V. Parker 759
three serpentine species ofHannonia but not by H. (JEPS), at a site southwest ofDubakella Mountain,
guggolziomm, H. nutans, or any other diploid spe- Trinity Co., California].
cies of the "Madia" lineage (i.e., diploid species In summary, members of the ultramafic clade of
of Anisocarpus Nutt., Carlquistia B. G. Baldwin, Harmonia appear to be outstanding examples of
Jensia B. G. Baldwin, Kyhosia B. G. Baldwin, or serpentine neoendemics, that is, groups that
Madia Molina; see Baldwin 1996, 1999). evolved on ultramafics, rather than relicts or pa-
leoendemic taxa secondarily restricted to serpen-
Biogeographic and evolutionary history ofHar- tines (see Stebbins 1942; Kruckeberg 1954, 1984;
monia. Discovery of H. guggolziomm has allowed Stebbins and Major 1965; Raven and Axelrod
for refined perspectives on the history of edaphic 1978; Mayer and Soltis 1994a, b).
endemism and overall pattern of diversification in
Hannonia. Based on phylogenetic analyses of Rarity. Discovery of H. guggolziomm along a
rDNA sequence data (Fig. 2, Baldwinunpublished), paved, public road less than 3 miles from US High-
the four species ofultramafic endemics in Hannon- way 101 at Hopland, near a University of Califor-
ia (H. doris-nilesiae, H. guggolziomm, H. hallii, nia field station, probably reflects extreme rarity of
and H. stebbinsii) represent a well-supportedmono- the species and insufficient access by botanists to
phyletic group that is sisterto H. nutans, an endem- serpentines in the vicinity. Smith and Wheeler
ic of volcanic-ash exposures in Napa and Sonoma (1990-1991) explored some nearby ultramafic sites
counties, California. In light of the phylogenetic in Mendocino County and did not report any spe-
data, I propose a simple hypothesis to explain pat- cies referable to Harmonia in their fiora of Men-
terns of edaphic endemism in Hannonia: diver- docino County. I did not find collections ofH. gug-
gence of the ultramafic and volcanic-ash lineages golziomm at CAS, CHSC, DAV, JEPS, PUA,
from a common ancestor preadapted (preapted) to ROPA, UC, or the herbarium of the University of
"harsh" edaphic conditions, followed by extensive California Hopland Research and Extension Center
diversification on serpentines in the ultramafic lin- and am unaware of any collections of the species
eage, that is, descent of H. doris-nilesiae, H. gug- from anywhere other than the holotype and para-
golziomm, H. hallii, and H. stebbinsii from a com- type localities. Harmonia guggolziomm is probably
mon, ultramafic-endemic ancestor. Lack of diver- naturally rare, based on the paucity of documented
sity in the volcanic-ash lineage may be attributable localities for other serpentine harmonias; H. doris-
in part to the limited geographic distribution ofvol- nilesiae, H. hallii, and H. stebbinsii are all listed as
canic exposures in northwestern California com- rare or endangered (List IB) by the California Na-
pared to the wide distribution of serpentine expo- tive Plant Society (2001). Exploration for new pop-
sures in the region (Kruckeberg 1984; Fox et al. ulations of H. guggolziomm on ultramafics of
southern Mendocino County and adjacent counties
1985).
Phylogeographic considerations lead me to sug- is needed.
gpeersstalaagnednearlallophaitsritcordyivfeorrsiHfaicnantioonniaamoofngwieddeapdhiisc- BetItyamGupglgeoalszedoftoCnloavmeerdatlhies,sCpaelciifeosrnifao,r Jwahcok caonld-
"islands" (see Raven 1964; Kruckeberg 1991). lected the first specimens ofHarmonia guggolzior-
Hannonia guggolziomm and H. nutans, represent- uni and who have contributed significantly to con-
servation of California's North Coast Range flora
ed by two lineages that diverge in succession at the
base of the Hannonia rDNA tree (Fig. 2), and H. through years of dedicated effort.
hallii are allopatric or parapatric taxa that are en-
Acknowledgments
demic or largely restricted to the southern North
Coast Ranges (//. nutans extends south into the Special thanks to Jack and Betty Guggolz for sending
northern San Francisco Bay area). The two species me specimens of Harmonia giiggolziorum and other in-
of the northern North Coast Ranges and southern teresting tarweeds and for sharing their enthusiasm and
Klamath Ranges {H. doris-nilesiae and H. stebbins- knowledge about native plants ofCalifornia'sNorthCoast
ii) are apically nested in the rDNA tree among the Rluasntgreasti.onIoaflsHo.tghuagngkolLezsiloenytmR;anJdoahlnlLf.orStprroetphaerrinagndthAelailn-
southern North Coast Range lineages and therefore R. Smith for assistance with the Latin diagnosis; JLS,
are suggested to be products of dispersal from the Theodore Barkley, and Kenton L. Chambers for helpful
south. Based on the rDNA tree topology, H. doris- comments on the manuscript; JLS, Susan J. Bainbridge,
nilesiae and H. stebbinsii are not sister species and andCharles F. Quibell forfieldassistance; BridgetL.Wes-
may represent independent south-to-north dispersal sa for lab assistance; and the curators of the following
events. Alternatively, the two species may have de- herbariaforloansofspecimensofHarmonia: HSC,NDG,
scended from the same northerly-dispersed ances- and the Shasta-Trinity National Forest herbariuminRedd-
ing, California.
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to-south dispersal. Hannonia doris-nilesiae and H.
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