A SYNOPSIS OF MACHAERANTHERA (ASTERACEAE: ASTEREAE), WITH RECOGNITION OF SEGREGATE GENERA

30 Pages·2003·18 MB·English

by  David R Morgan

MACHAERANTHERA
A OF
SYNOPSIS (ASTERACEAE:
WITH GENERA
RECOGNITION OF SEGREGATE
ASTEREAE),
Morgan L Hartman
David Ronald
R.
W
Department of Biok Department of B
/estern Washington Uniiversity University of Wye
WY
lingham,WA 98225-91 USy Laramie, 82070-3
6"0,
"'""°"'"'
morgan@b,ol.wwu.i
ABSTRACT
taxonomy Machaeranthera
ding of its evolutionary history, revisions to its are proposed. is cir
scribed to include two species, and other species that have been included in Machaeranthera ar
accommodate
)rtioned into three additional genera. Dieteria revived to three of these specie:
is
new
thisma is expanded to accomodate 16 more species, and the genus Arida is proposed fo
New A
remaining. combinations for the species in Dieteria, Xanthisma, and rida are providec
e
morfologic
The genus Machaeranthera Nees (Asteraceae: Astereae) was established to ac-
many
commodate
the single species M. tanacetijolia Nees. Since inception,
its
circumscriptions Machaeranthera have been proposed, and the
different of
genus has included various parts of several other genera, with agreement
little
on composition or relationships with other taxa (see Morgan and Simp-
its its
son 1992 for a more detailed taxonomic history).
The most recent published treatment of Machaeranthera was by Hartman
who
(1990), included 36 species in the genus, excluding several taxa that are
now placed in the genera Hazardia Greene (Hartman 1990), Rayjacksonia R.L.
&
Hartman M.A. Lane (Lane Hartman Tonestus A. Nelson (Nesom
1996), 1991),
6a:
and Xylorhiza (Hartman The circumscription Machaeranthera
Nutt. 1990). of
Hartman
adopted by (1990) included species with both cyanic (white, blue, pink,
purple) and yellow and apportioned them into two subgenera and eight
rays,
We
Machaeranthera
sections. will refer to this circumscription as
s.l.
Investigations of Mcicht/eranthcras.l. using molecular evidence have helped
DNA
from (cpDNA)
to clarify its systematics. Restriction site data chloroplast
&
Home
Machaeranthera was
indicated that section PsilactisiA. Gray) Turner
&
Machaeranthera (Morgan Simpson Morgan
not a close relative of 1992;
s.l.
This section was removed from Machaeranthera by Morgan
1993). (1993),
s.l.
leaving 30 species in the genus. More recent investigations of Machaeranthera
DNA
(sensu Morgan 1993) employed sequence data from nuclear ribosomal
DNA
internal and external transcribed spacers (nrDNA) (Morgan Mor-
1997;
cpDNA
gan Comparison with
2003). of this evidence data indicated that the
evolution of Machaeranthera and relatives has been complex, involving as
its
many
The
seven occurrences molecular evidence
as of reticulate evolution. also
indicated that other genera Oonopsis Nutt., Pyrrocoma Hook., and
(e.g.,
Xanthisma Machaeranthera
DC.) are closely related to various parts of (Fig.
1).
The relationships supported by a combination of molecular, morphologi-
cytological, and chemical evidence do not support any recent circumscrip-
cal,
new
tion of Machaeranthera. Therefore, a classification needed to bring the
is
taxonomy of the genus more in line with current understanding of relation-
its
A
made
ships. case could be for including Oonopsis and Pyrrocoma in an ex-
cpDNA
panded Machaeranthera, and some
evidence provides support such
for
nrDNA
a treatment (Fig. However, data do not support this circumscription
1).
we
(Fig. Furthermore, contend that enlarging Machaeranthera through ac-
1).
would
cretion obscure the phylogenetic patterns within the complex, and would
result in a large, heterogenous, and unwieldy genus that would be difficult to
We
characterize satisfactorily. therefore propose that Machaeranthera (sensu
Morgan
1993) be divided into four genera, as described below.
KEY TO THE GENERA TREATED HERE
Ray floret corollas yellow (heads radiate) Xanthisma
1.
throughout,
at least
ifid
I
;i-ventrally flattened
r
pappus
:o strongly perennial; of
f
d near base, bases overlapping or
r

pappus
long); bristles filiform, not flattened near base (or slightly so), bases
not overlapping.
(or slightly)
4. Ray florets with prominent pappus (often eradiate in the glandular-pu-
variously pubescent with glandular and/or nonglandular hairs;mountain5,
_
W
SW
Diains, and basins of U.S.A.: Canada: extreme N Mexico Dieteria
4. Ray florets with pappus absent (5 species; eradiate in the glabrous A.
carnosa) or present; pappus present then leaves pinnatifid to bipinnatifid
if
throughout or leaves entire to toothed) plants glabrous except for bristles
(if
Mexico
Arida
&
Machaeranthera Nees, Gen. Sp. Asterearum 224. 1832. Type: Aster
lacetifolius
tar,
Kunth.
Chrysopsis subgenus Pappochroma Nutt., Acad, Nat, Sci, Phila, 7:34, 1834, Dieter ia subgenus
J.
Chrysopm
(Nutt,) Walpers, Rep, Bot. Syst, 2:587, 1843, Type: coronopijolia Nutt,
Machaeranthera (Nees) A, Gray, Proc, Amer. Acad. Arts 16:99. 1880.
&
Machaeranthera series Verae Cronquist D.D. Keck, Bnttonia 9:238. 1957. nom. (ICON:
illeg. 52.1).
cm
Herbs taprooted annuals 5-100 Stems
or biennials, erect to ascending,
tall.
much when
branched well developed, moderately densely pube& with
cent
to
glandular and nonglandular Leaves
hairs. alternate; pinnatifid or bipinnatifid,
deeply lobed or dissected, moderately to densely pubescent with glandular and
nonglandular hairs, apices, including lobes and teeth, mucronate to bristle
mm
tipped, bristles present 0.2-1 long. Heads radiate. Involucres broadly tur-
if
3-6
binate to hemispheric. Phyllaries in imbricate series, graduated in length,
appressed, spreading, or ref lexed, moderately to densely pubescent with glan-
dular and nonglandular hairs; bases indurate, apices herbaceous. Receptacles
mm
convex, naked with Ray
alveolate, flat to or scales to 0.4 long. florets pistil-
late, fertile; corollas blue, violet, or purple. Disc florets bisexual, corollas
fertile,
yellow, narrowly funnelform; tubes and limbs glabrous or glabrate, lobes 0.3-1
mm
long, glabrous to pubescent. Cypselae of ray and disc florets distinctly di-
morphic, narrowly to broadly obovate, sparsely to densely pubescent; moder-
mm
4-9
ately thick walled, with ribs per face, 2,4-3.8 long, those of ray florets
-3 rounded somewhat
sided, abaxially, of disc florets compressed laterally; pa-
tawny
ppus white
to of ray florets 1/2 to nearly equalling length of disc
florets;
bristles coarsely barbellate, in 1-3 series, dorsi-ventrally flattened and dissimi-
mm
lar in width near base, bases overlapping, mostly 4.5-6.5 long, n =
4.
The
circumscription of Machaeranthera adopted here genus
results in a
We
two
of species. apportion the remaining 28 species of Machaeranthera
Morgan
(sensu 1993) to Dietena, Xanthisma, and Arida (see below). The three
species placed in Dieteria could potentially also be included in Machaeranthera
number members
because of a of similarities. All of the two genera produce
flavonols, and all have chromosome numbers of n = obovate and nar-
fruits,
4,
row (Hartman Turner This
phyllaries that are often ref lexed 1976, 1990; 1987).
cpDNA nrDNA
was supported by evidence However,
relationship also (Fig.
1).
cpDNA
with and between
evidence conflicted the substantial conflict
(Fig.
1),
them complicates the issue by suggesting reticulate evolution in the ancestors
cpDNA nrDNA
and
Machaeranthera, evidence
of Dieteria, or both. In addition,
both supported a close relationship between Machaeranthera and Oonopsis, and
expansion of Machaeranthera to include Dieteria would also have to include
&
many
which from Machaeranthera ways (Morgan Simp-
Oonopsis, differs in
son Additional synonymy Machaeranthera included in Turner
1992). for is
(1987).
MACHAERANTHERA
KEY TO THE OE
SPECIES
long, glabrous or glabra
1
mm
pubescent Machaeranthera tagetina
lobes mostly 0.7-1 long, 2.
&
Machaeranthera (Kunth) Nees, Gen. Asterearum 225. 1832.
tanacetifolia Sp.
.
&
Aster tanacetifoUus Kunth, Nov. Gen. Sp., ed. folio, 4:74. 1820. Aster chry-
m
MEXICO:
santhemoides Willd. ex Spreng., Syst. Veg. 3:538. 1826. Type: "Colitur
horto Mexicano," Humboldt (holotype: P IDC microfiche 6209.
s.d., s.n. nv., 97.111.6;
possible ISOTYPE: B-W; photoisotype: TEX!).
Machaeranthera
Nutt., Trans. Amer. Philos. Soc. ser. 2. 7:302. 1840. coronopifolia (Nutt.) A.
GH;
souri" (probably near Fort Mandon), Jul-Aug 1811, Nuttall s.n. (isOTYPE: probable ISOTYPE:
Machaeranthera tagetina Greene, Pittonia 1899. Aster tagetinus (Greene)
4:71.
2.
ARIZONA. Cochise
Blal<e, Contr. U.S. Natl. Herb. 25:263. 1925. TYPE: U.S.A. Co.:
S.F.
Huachuca, Wilcox (holOTYPE:
near Fort 1891, US).
s.n.
&
Machaeranthera series Vanahi Cronquist D.D. Keck, Bnttonia 9:237. 1957, nom. iUeg. (ICBN:
les
24.3).
cm
10-100 Stems
Herbs taprooted annuals, biennials, or perennials, erect
tall.
much when
ascending, branched well developed, glabrous to variously pu-
to
bescent with glandular and/or nonglandular hairs. Leaves alternate; entire to
moderately densely pubescent with glandular
irregularly serrate or dentate, to
and/or nonglandular and
hairs, apices, including lobes teeth, spinescent due
indurated Heads
to apiculae, bristles absent. radiate or eradiate. Involucres tur-
campanulate,
binate, or hemispheric. Phyllaries 3-12 imbricate
in to
subimbricate graduated
series, in length, appressed, spreading, or ref lexed, gla-
brous or variously pubescent with glandular and/or nonglandular
bases
hairs;
indurate, apices herbaceous, margins
scarious. Receptacles alveolate, convex,
mm
naked Ray
or scales to 0.3 long. florets pistillate, fertile (sterile or absent in
canescens var
D. shastensisX corollas white, blue, violet, or purple. Disc florets
bisexual, fertile, corollas yellow, + funnelform, tubes and limbs glabrous or
mm
glabrate, lobes 0.2-0.7 long, glabrous to minutely pubescent. Cypselae of
and somewhat
ray disc florets dimorphic, linear obovate, glabrous moder-
to to
mm
thm
ately pubescent; walled, smooth or with 4-6 ribs per 3-6
face, long,
those of ray florets generally flattened, rounded abaxially, of disc florets flat-
tened pappus white brown
laterally; to or reddish brown, of ray florets 2/3 to
nearly equalling length somewhat
of disc florets; bristles barbellate, in 1-3 se-
most somewhat
ries, at dorsi-ventrally flattened and dissimilar in width near
mm
4-8
base, bases not to slightly overlapping, mostly long, n =
4.
Although similar Machaeranthera ways mem-
to in several (see above), the
may
bers of Dieteria be distinguished from by and
foliage fruit characters.
it
Dieteria has entire to serrate or dentate leaves with thin-walled
fruits that are
mm,
mostly longer than while Machaeranthera
3 has leaves that are deeply
mm
lobed or dissected and thick-walled mostly
fruits that are 3 or less long.
There
is a great deal of morphological variation in Dietena, which has resulted
m
number
a large of species names. The treatment adopted here follows that of
Turner
(1987) in recognizing three species containing 16 varieties, with 3 vari-
and
eties in D. asteroides, 3 in D. higelovii, 10 in D. canescens. Additional syn-
onymy
Dieteria included Turner
for in
is (1987).
OF
DIETERIA
Emory
in Report 142. 1848. Machaeranthera asteroides
,
NEW
MEXICO: land between
Pittonia 1896. Type: U.S.A. "elevated
3:63.
Emory
and the waters of the Gila," 16 Oct 1847, (holotype: NY).
s.n.
nescens var. lal ifoHa A. Gray, Pi. Wright. 2:75. 1853. Aster canescens var. latifolia
NY
GH, mo,
PH, UC,
ISOLECTO-types: US).
Mac
Aster canescensPurshvartephnxies A.GravProc.Ame r.Acad. Arts 16:99. 1880. haeranthera
t^phrodc: r tephrodes (A. Gray) S.F.Blat:e,Contr,U.S.
h 25:563. 1925. Typifica Calif.].:322. 1876, Gray mistE.kenly based
name Gray on Diplopappus rtcanusLmdl plan
the A. for
ii
name
insouthe rn California. He subsequently corrected this mistake and proposed the Aster
ma New
inAnzc and
canescen: var teph rode, for thes, plants, noting tha ttheycdso occurred
2
5
A m
M
IUthough stating that/I var tephrodes " ...was named incanus tlle Botany of
c.
Cal fo n ley's plant." Gray a]jpareni;ly did not select a type specimen
At ment lonedbyGrayinconi with
anescensv^r.tephrode lection
fo
i\
m
any dby"Capt.E.K.S:^nth" "Guadalupe Canyon, Sonora,"but
of
1
w
ha e been able'to locate irner L987) placed Aster in canus. Aster
lo't (]
nonymy
a n van tephrodes, Macha eranthera tephrod. and Aster tephrodes in syi un-
:s,
Mah
de aerantheracanescens^
KEY TO THE VAR OF DIETERIA ASTEROIDES
lETIES
Stems giabroius to glandular-pubesicent; leaves covere.dwith stiff glandular hairs
glandulosa
f^esc
Stems pubesc puberulent, or carlescent, glandular
:ent,
ing; leaves puberulent or canescent, stiffglandularhair e or lacking.
Involucres hemispheric; herbaceous phyllary apice: long-
2. 5
long; mid- stem leaves 6-20 mr wide, usually c serrate or serrulate
T.
asteroides
2 Involucres broadly turbinate (ran2ly hemispheric);!lerbace?ous phyllary apices
6mmv
2-
uminate,1-3mmlong;imid-stem leaves vide,entire to slightly
Ic.D. asteroides lagunensis
serrulate'" var.
Dieteria asteroides asteroides
la. var.
&
Morgan
lb. Dieteria asteroides var glandulosa (B.L. Turner) D.R. R.L. Hart-
man, comb. nov. Basionym: Machaeranthera asteroides (Torr) Greene var.
ARIZONA. Maricopa
glandulosa Turner, Phytologia 60:77. 1986. TYPE: U.S.A. Co.:
B.L.
U.S. highway 60, 2.6 mi E of Queen Creek tunnel, 4200 ft, 19 Sep 1975, Pinkava,
&
Keil Echto L18904 (hOLOTYPE: isotypes: ASU, CSU, NY).
LLI;
&
Morgan Hartman,
Keck)
Dieteria asteroides lagunensis (D.D. D.R. R.L.
Ic. var.
comb. nov. Basionym: Machaeranthera lagunensis D.D. Keck, Brittonia 9:238. 1957.
Machaeranthera asteroides (Torr.) Greene var lagunensis (D.D. Keck) B.L. Turner,
mi main
Phytologia 60:77. 1986. Type: U.S.A. California. San Diego Co.: 2 S of the
&
Munz
Aug EX.
recreation area, Laguna Mountains, 5200 20 1952, PA. Balls
ft,
NY
17948 (holotype: iSOTYPE: RSA).
&
Dieteria bigelovii (A. Gray) D.R. Morgan R.L. Hartman, comb, nov Basionym:
..
/\s(f r bigelovii A. Gray, Pacific Railway Survey 4:97. 1857. Machaeranthera bigelovii
New
(A. Gray) Greene, Pittonia 3:63. 1896. TYPE: U.S.A. Mexico. Bernalillo Co.: "ar-
m
royos the Sandia Mountains," 10 Oct Bigelow (holotype: GH; isotype:
1853,J,M.
n.
s.
OF
lETlES DIETERIA BIGELOVII
mm
ad as tal!;phyllaries 90-1 00, mostly 0.5-1
mm
long-acuminate, 2-5 long 2c.D.bigelo>
2a. Dieteria bigelovii var bigelovii
2b. Dieteria bigelovii var commixta (Greene) D.R. Morgan R.L. Hartman, com
(Si I
nov. Basionym: Machaeranthera commixta
Greene, Pittonia 189S
4:71.
Machaeranthera canescens (Pursh) A. Gray var commixta (Greene) Welsh
S.L.
Great Basin Naturalist 43:316. 1983. Machaeranthera bigelovii (A. Gray) Green.
Garfield Co.: Henry Mountains, Bromide Pass, 10,000 27Jul 1894, M.E.Jones 5695^
ft,
(holotype: US; ISOTYPE: POM).
&
mucronata Morgan Hartman
2c. Dieteria bigelovii var. (Greene) D.R. R.L.
comb, nov
Basionym: Machaeranthera mucronata
Greene, Pittonia 4:72. 189S
Aster adenolepis S.F. Blake, Wash. Acad. Sci. 30:471. 1940, nom. nov, non Aste;
J.
mucronatus Sheldon, 1903. Machaeranthera Gray) Greene
bigelovii (A, vai
mucronata
(Greene) B.L. Turner, Phytologia 62:238. 1987. Type: U.S.A. Arizona
Coconino Thompson
Co.: Canyon, 8500 19 Sep 1894, M.E.Jones 6065hl (lectq
ft,
US
type: [designated by Turner
1987]).
Amer
3. Dieteria canescens (Pursh) Nutt., Trans. Philos. Soc, 7:300. 1840
ser. 2,
Amer
Basionym: Aster canescens Pursh, Machaerantherc
Fl. Sept. 2:547, 1813.
canescens (Pursh) A. Gray Pi. Wright. 1:89. 1852. Type: U.S.A. North Dakota: "or
denuded banks
the of the Missouri" (vicinity of Fort Mandon), Nuttall
1811, s.n
(holotype: BM; NDG).
possible ISOTYPE:
Machaera
Pittonia 4:22. 1899. nthera ca nescens (Pursh) A. Gray var viscosa (Nutt.) Piper, Conti
:
25:563. 1925, nom. nov., non Aster laetevirens Greene (190
Clover Mountains, 26 Jul 1894, EL. Greene (holotYPE:
s.n.
Gray Nelson) Welsh, Great
(Pursh) A. var. latijolia (A. S.L.
U
Utah. Lake Big Cottonwood Canyon,
Type: S.A. Salt Co.:
GH,
1933 (HOLOTYPE: RM!; ISOTYPES: LL!, US).
KEY TO THE VARIETIES OF DIETERIA
hyllaries spreading to reflexed, rarely appressed.
glandular, frequently also
y p
leucanthemifolia
r
Ray florets sterile or abs'
and
Ray
florets pistillate
.
densely
Phyllaries a
10.
glabrous
Phyllaries
10.
1
glabrous
Dieteria
(i)
&
3b. Dieteria canescens var. incana (Lmdl.) D.R. Morgan R.L. Hartman, comb,
nov. BASIONYM: Diplopappus incanus Lindl, Bot. Reg. 1693, 1834. Dieteria
20:t.
&
Amer
incana Machaer
(Lindl.) Torr. A. Gray, Fl. N. 2:100. 1841. anthera canescens
(Pursh) Gray var incana Gray Wilkes
A. (Lindl.) A. Bot. Exp. Phan. 17(2):340. 1874.
Machacranthera
incana (Lindl) Greene, Pittoma 3:62. 1896. Type: U.S.A. OREGON:
m
"Columbia River" described from seeds grown London, 1830, Douglas (ho-
s.n.
LOTYPi-:: BM; LSOTYPE: GH; ISOTYPE, microfiche: G-DC).
3c. Dieteria canescens var. leucanthemifolia (Greene) D.R. Morgan R.L. Hart-
6a:
man, comb.
nov. Basionym: Aster leucanthemijolius Greene, Erythea
1895.
3:119.
Machacranthera
leucanthemifolia (Greene) Greene,
Pittonia 1896.
3:61.
Machacranthera
canescens (Pursh) A. Gray var leucanthemijolia (Greene)
S.L.
Welsh, Great Basin Naturalist 43:316. 1983. Type: U.S.A. Nevada. Esmeralda
Co.:
WM.
near Candelaria, 6000 Jun 1886, Shockley 268 (holotype: CAS; isotypes:
ft,
DS, US).
&
3d. Dieteria canescens Morgan Hartman,
var. sessiliflora (Nutt.) D.R. R.L.
comb, nov Amer
Basionym:
Dieteria sessiliflora Nutt., Trans. Philos. Soc, ser
2,
Machacranthera
7:301. 1840. sessiliflora (Nutt.) Greene, Pittonia 3:60. 1896.
Machacranthera
canescens (Pursh) Gray var B.L Turner
A. sessiliflora (Nutt.)
Phytologia 60:78. 1986. Type: U.S.A. Idaho(?): "Denudated plains of the Rocky Moun-
and
tains Oregon," 1836, Nuttall (HOLOTYPE: BM).
s.n.
&
3e. Dieteria canescens var shastensis Gray) Morgan Hartman,
(A. D.R.
R.L.
comb, nov
Basionym: Machacranthera Gray Amer
shastensis A. Proc. Acad. Arts
6:539. 1866. Aster shastensis (A. Gray) A. Gray,
Bot. Calif, 1:322. 1876.
Machacranthera canescens (Pursh) A. Gray var shastensis (A. Gray) B.L. Turner,
Phytologia 60:79. 1986. TYPE: U.S.A. California. Siskiyou Mt. Shasta, 9000
Co.:
ft,
WH.
1860-62, Brewer 1385
C
Dieteria canescens var
3f.
nov. Basionym: Macha^
Machacranthera
Aliso 7:65. 1969. canescens (Pursh) A. Gray var ziegleri
(I
B.L. Turner, Phytologia 60:79. 1986. Type: U.S.A. California. Riverside
Co.:
i
of Santa Rosa Mountain, 6500-7500 30 Sep LB. (holotypi
1968, Zieglers.n.
ft,
&
Morgan
subsp. glabra Gray) Hartman,
(A. D.R. R.L.
xeranthera canescens (Pursh) A. Gray var glabra A. Gray
Pi.
Machacranthera
canescens (Pursh) A. Gray subsp. glabra
(A.
hytologia
62:239. 1987.
&
ambigua Morgan
var Turner) Hartman,
D.R.
(B.L. R.L.
Basionym: Machacranthera
canescens (Pursh) Gray var ambigua
A.
;
Phytologia 60:77. 1986. TYPE: U.S.A. Arizona. Coconino
Flagstaff,
Co.: