Antennaria pulvinata Greene: The legitimate name for A. aromatica Evert (Asteraceae: Inuleae)

16 Pages·1993·7 MB·English

by  J G Chmielewski

RHODORA,
Vol. 95, No. 883/884, pp. 261-276, 1993
ANTENNARIA
PULVINATA GREENE: THE
NAME
LEGITIMATE FOR AROMATICA EVERT
A,
(ASTERACEAE: INULEAE)
Jerry G. Chmielewski
ABSTRACT
Canonical was
variates analysis used as an analytical technique to document
morphological among Antennaha =
discontinuities individuals of media
{n
63),
= = =
A. pulvinata (n 103), A. rosea {n 64) and A. umbrinella {n 65). Evaluation
of
the defined 94%
classification criterion indicated that of the specimens were
classified correctly. The classification criterion was subsequently used to classify
=
type collections of^. aromatica (n 26) into one of the previously defined groups.
Results based on dem-
these analyses as well as previously published information
onstrate that A. aromatica and A. pulvinata are morphologically indistinguishable.
The
analyses also indicate that A. pulvinata and A. rosea exhibit morphological
integrity relative to each other and would best be treated as distinct species. The
four species, A. media, A. pulvinata, A. rosea and A. umbrinella, exhibit morpho-
A
logical integrity and should be treated as distinct. of synonymy provided
list is
for A. pulvinata.
Key
Words:
Antennaria media, A. pulvinata, A. rosea, A. umbrinella, A. aromatica
INTRODUCTION
Antennaria was
aromatica Evert originally described as a sexual
diploid Subsequent have
Cordilleran species (Evert, 1984). studies
demonstrated
morphologically,
that the species cytologically
is
and
more and
reproductively variable (Bayer Stebbins, 987; Bay-
-^.
1
er, 984, 989a), as well as more widely distributed (Chmielewski
1 1
and
Chinnappa, than proposed.
1988a; Bayer, 1989a), initially
from
Additional interest in A. aromatica stems the implication
Greene
that the species one of the progenitors of the A. rosea
is
polyploid complex, described A. rosea
that entity as
specifically,
subsp. pulvinata (Greene) R. Bayer (Bayer, 1989b).
Evert was Antennaria aromatica
(1984) able to differentiate
from
both A. media Greene and A. umbrinella Rydberg, other
members
of the complex, by copious and persistent glandu-
its
and
losity, widely cuneate-spatulate basal leaves distinctive cit-
ronella-like odor. Bayer (1989b) concluded that his subsp. pul-
vinata resembled A. aromatica, except in that lacked glandular
it
and
hairs. Although Greene described both "female"
(1898)
"male"
specimens of Bayer (1989b) concluded that
^. pulvinata,
and
his subsp. apomictic polyploid.
pulvinata
entirely pistillate,
is
261
Rhodora
262 95
[Vol.
Antennaria aromatica was considered endemic to
initially
Wyoming
Montana and of Idaho and
southwestern adjacent parts
The
and
occupying limestone talus xeric habitats (Evert, 1984).
Columbia
was from
subsequently reported Alberta, British
species
and California (Chmielewski and Chinnappa, 1988a). Bayer(1991)
specimen
rejected the latter range extension, citing misidentifi-
and
cation, as determined by either inspection or deduction, in-
by Bayer
stead supported the distribution for the species presented
(1989a).
was an
Although Antennaria aromatica originally described as
endemic, morphologically uniform, sexual diploid species, five
was more
years appropriate consider the species as
later to
it
wider ranging (Chmielewski and Chinnappa, 1988a; Bayer 1989a),
and
reproductively variable (either sexual or apomictic) cytolog-
and
ically variable (diploid, tetraploid or hexaploid) (Bayer Steb-
bins, 1987; Bayer 1989a). Reproductive biology, hybridization,
polyploidization and dioecism have contributed to morpho-
all
among
of Antennaria and thereby
logical intergradation species
magnified the present state of confusion not only in this species
but also the genus (Femald, 1950; Cronquist, 1955; Anderson,
and
1959; Hulten, 1968; Welsh, 1974; Scoggan, 1979; Porsild
Cody, Moss,
1980; 1983).
This investigation incorporates the of a phenetic study
results
of gross morphology with information published during the past
decade determine whether
to possible to distinguish consis-
it is
and among
tently confidently Antennaria aromatica, A. media,
and
A. pulvinata (including subsp. albescens Nelson), A. rosea
E.
Having
A. umbrinella. which
established those taxa are distinct,
some comments
presumed
are offered concerning the evolution-
ary history of the group.
AND
MATERIALS METHODS
Herbarium
specimens, including type material, utilized in the
phenetic study were borrowed from alta, ariz, can, colo, dao,
F, LEA, MO, MONT, MOR, MT, NY, OS, PH, RM, SLRO, UAC, UBC, UC,
WIN and ws (Holmgren
US, Specimens of each species
et 990).
al,,
1
were on
selected the basis of completeness and comparability in
terms of developmental
speci
stage
_,
mens
were
These
included. specimens a range plant size
reflect in
Chmielewski—
263
1993] ^/7/^/i/2(2r/a
Table Morphological
characters included in the phenelic study of Anten-
1.
naria aromatica, A. media, A. pulvinata, A. rosea and A. umbrinella.
Structure Character
BASALLL
Vegetative Basal leaf length (mm):
BASALLW
Basal leaf width (mm):
HEIGHT
Vegetative-Reproductive (mm);
Plant height
NOCAULL
Number
of cauline leaves:
CAULLL
CauHne (mm):
leaf length
CAULLW
Cauline width (mm):
leaf
NOCAP
Number
of
capitula:
INVOHT
(mm):
Involucre height
OUTPHYL
Outer (mm):
phyllary length
OUTPHYW
Outer width (mm):
phyllary
INNPHYL
(mm):
Inner phyllary length
INNPHYW
Inner phyllary width (mm):
PAPPUSL
Reproductive Pappus (mm):
length
COROLLAL
(mm):
Corolla length
No
involucre habit and provenance. a priori restrictions were
size,
necessary
morph
=
species. Herbarium specimens {n 295) were identified prior to
=
analysis as Antennaria aromatica {n 26 type sheets), A. media
= = =
um-
and
{n 63), A. pulvinata (n 103), A. rosea {n 64) A,
=
brinella {n Qualitative characters used to identify the spec-
65).
imens included degree of pu-
to species prior to analysis habit,
of
bescence, shape and texture of the phyllaries, exsertion
color,
on
the style and the presence or absence of papillae the achenes.
These characters were not utilized in the phenetic study. Both
pistillate and staminate plants were included in the phenetic study.
Data each
were collected for 4 quantitative characters for spec-
1
imen (Table These characters were selected primarily because
1).
< form of
they were uncorrected described the general
r .70),
(
| |
vegetative and reproductive structures, were not used directly in
were not
the classification of individuals to a priori groups, ips-
ative and were previously shown to be useful in differentiating
among
and Chinnappa, 988b,
o^Antennaria (Chmielewski
species
1
1991; Chmielewski 1990a, 1990b). Several of these char-
et
al.,
number
of
capitula, size
acters, including plant height, leaf shape,
among
of
capitula and are typically used to differentiate
floret size,
species of Antennaria. The data matrix contained no missing
values.
Rhodora
264
95
[Vol.
SAS
was (SAS
Canonical with
variates analysis facilitated the
DISCRIM
and
Institute Inc., 1989) (crossvalidate posteir options)
CANDISC
To
and procedures. meet the assumptions of multi-
measurements were
variate normality (Gilbert, 1968), length
transformed to their logarithms (base 10) and count data were
transformed to their square roots prior to initiating multivariate
was
analysis. Classificatory discriminant analysis used to classify
and
specimens oi Antennaria media, A, pulvinata, A. rosea A.
on one more were
umhrinella, defined a priori or characters that
not included in the analysis, to species. Correct classification rates
and Geisser assignment probabilities were used as indicators of
among
separation the groups. Tests for equality ofgroup centroids
The same
were performed as part of the canonical analysis. data
was used both and
set to define evaluate the classification crite-
The
was
rion. classification criterion subsequently used to classify
=
the type collections of A. aromatica {n 26) into one of the
previously defined species. Antennaria aromatica was not treated
as a separate a priori group in the discriminant analysis because
the importance of glandularity in the separation of rosea subsp.
/I.
my
pulvinata from aromatica was overempha-
A. in estimation
The
sized. copious glandularity that evident on the holotype
is
is
pronounced on some
less to lacking of the other type collections.
some
Additionally, specimens appear non-glan-
that superficially
do
dular possess glands beneath the tomentum. Thus, A. aro-
matica is not considered to be consistently glandular. Trivariate
plots of the canonical variate scores were used to facilitate vi-
of
sualization the results of the multidimensional analysis
±
Mean
(DeltaPoint was
Inc., 1992). value standard deviation
determined
for each character for each species.
RESULTS
Canonical (CVA) morpho-
variates analysis of 14 quantitative
among
logical characters was used document
to discontinuities
specimens oi Antennaria media, A, pulvinata, A. rosea and A,
umbrinella. Evaluation of the indicated
classification criterion
94%
that of the specimens were (Table
classified correctly 2).
Geisser assignment
were than
probabilities generally greater .98
for the 278 correctly assigned specimens. The low error count, in
conjunction with the high Geisser assignment probabilities, in-
Chmie\Qv/ski— Ant
1993] ennaria 265
Table summary number
2. Classification by of observations and percent
clas-
sified into species for the discriminant analysis.
Number
of Observations and Percent
Classified into Species
1
From
Species media puhinata
rosea umbrinella
media
54
8
1
85.71 12.70 .00 1.59
puhinata
6 97
5.83 94.17 .00 .00
rosea
63
1
1.56 ,00 98.44 .00
umbrinella 64
1
.00 1.54 .00 98.46
dicates that classification of additional specimens, that type
is,
collections of A. aromatica, through the use of the classification
would
criterion, yield acceptable results.
The and second discriminant functions accounted for 90.5%
first
and 17.4%
(73.1 respectively) of the total variation (eigenvalues
of and among
4.263 1.015 respectively) specimens. Ordination
of
the 295 specimens by canonical analysis presented as a series
is
of which mean
trivariate plots (Figure in canonical scores are
1)
The Mahalanobis between
indicated each distance
for species.
groups and associated F-value indicate the group centroids were
< The
significantly different (P .0001), canonical correlation of
the discriminant function or that squared as the pro-
first (.900),
portion of the variance in the function explained by the four
groups high moderate correlation with the
indicates to
(-810),
K
A
groups. strong relationship between the specific linear combi-
nation of canonical variables and the groups
variables, the is
indicated.
±
Descriptive (mean standard deviation) are used to
statistics
of
predefined groupings
identify character discontinuities for the
STEMHT, NOCAULL, CAULLL,
.Li specimens (Table Except for
3).
INNPHYW
NOCAP
BASALLL,
and which exhibited moderate
no among some remaining
to overlap or of the groups, the
all
among These
characters consistently intergraded the groups. char-
acters had the highest loadings on the canonical axis.
first
Type were through
specimens of ntennaria aromatica classified,
.4
=
the use of the classification criterion, to either /I. media (7%, n
Rhodora 95
[Vol.
266
«1
c
3
5
'c^'^'
*«
*3
5
'c>alA^'*^
Figure Ordination of Antennaria media (—.818, 1.277, —1.017); b, A.
1. a.
puhinata (-2.238, -.302, .564); A. rosea (2.908, .754, .750); and d. A. urn-
c.
brinvlla (1.476, -1.502, -.646). Group cenlroids are shown in parentheses (ca-
nonical axes 2 and 3 respectively).
1,
=
ox A. puhinata (92%, n Geisser assignment probabilities
2) 24).
were typically greater than .95 for these specimens.
DISCUSSION
noted that
In describing Antennaria aromatica. Evert (1984)
the species was morphologically uniform throughout its range.
values
Subsequent comparison of holotype average species
the to
mean product-moment
via a similarity matrix employing Pearson
that
Table indicated
correlation coefficients (^e^ Bayer, 1988a,
1)
;
Chmielev^ski—Antennaria 267
1993]
m
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O ^ OOOOOW^U^TtsOO
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ft
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Rhodora
268 [Vol. 95
"atypical of the species, consisting of really small diploids
it is
comm.
that do not occur on limestone talus" (Bayer, pers. 1989).
=
Examination of available type material {n 26) indicated that
mo^X Antennaria (Chmielewski and Chinnappa,
A. aromatica, like
1988b, 1991; Chmielewski 1990a, 1990b), indeed vari-
et is
al.,
able, exhibiting different degrees of plasticity, with respect to basal
and
leaf length and width, plant height, cauline leaf length width,
degree of pubescence, glandularity and involucre height. Anten-
naria aromatica therefore a species which illustrates that the
is
most
nomenclatural type not necessarily the typical or repre-
is
sentative element of the taxon; merely that element with
is
it
name
which the permanently associated (Lanjouw et 1966).
is al.,
Antennaria
Initially described as a diploid species (Evert, 984),
1
aromatica was subsequently reported include both tetraploid
to
and
and hexaploid individuals (Bayer, 1984, 1989a; Bayer Steb-
some
Apparently confusion with respect to the
bins, 1987). exists
CO-458
identification of the tetraploid specimen designated (Bay-
er and Stebbins, 1987), as was included with both A, aromatica
it
(toward A, media) and A. rosea (toward A. aromatica) by the
authors. This confusion notwithstanding, likely that the dip-
it is
and
loid polyploid states possessed by individuals of the species
have
contributed, at least in part, via the gigas effects of poly-
ploidization (Stebbins, 97 to the observed pattems of plasticity
1 1)
{see also Bayer, 1989b) noted above. Because so few cytological
determinations valid
are available for the species, statistically
chromosome
conclusions pertaining to the relationship between
number
and morphology
are not possible at this time.
Results based on canonical variates analysis indicate that the
degree of morphological overlap among the four taxa, Antennaria
media, and
A, pulvinata, A. rosea A. umbrinella, as defined in this
study, minimal. High assignment and Geisser assignment
is rates
me-
The
probabilities support this conclusion. conclusion that A.
and
dia, A, rosea A. umbrinella are morphologically distinct is
not a novel one as the species are generally treated as such {see
Bayer, 1988a, 1988b, 1989c, 1990a; Chmielewski and Chinnappa,
The
1988b; Chmielewski 1990b
et 1990a, for discussion).
al.,
how
question remains, does Antennaria with these
pulvinata in
fit
species?
Direct visual comparison of type material indicates that frag-
ments of (RM346356) and
the holotypes o^ Antennaria aromatica
ChmielQwski—Antennaria
1993] 269
(NDG058584)
^. pulvinata
are indistinguishable,
-9 as are the para-
type of aromatica and (CANl
A, the isotype 05666) and
(f) para-
(CAN105674)
type of
^. pulvinata.
Although two
type collections of Antennaria aromatica were
originally assigned to A, media, through the use of the
classifi-
92%
cation
criterion, the success observed with
rate in line the
is
of was
capabilities the classification criterion as defined through
it
The
the
analysis. classification criterion obtained through canon-
was 94%
ical variates initially evaluated at a success rate. Thus,
when
it is unrealistic to expect a higher classification success rate
used
it is to classify collections, such as the types of^. aromatica,
which
were not used The
to define the classification criterion.
analysis demonstrates not only that A, aromatica and A. pulvinata
are indistinguishable but also that morphological intergradation
among
species of Antennaria a facet of reality.
is
In attempting to clarify the confusion that surrounds the iden-
tity of Antennaria pulvinata, Bayer (1989a) concluded that re-
it
sembles A, aromatica closely, except that lacks glandular hairs.
it
This
character not infallible however, as glandularity, even
is
among
from
type collections of A, aromatica, ranges a dense cover
to trace amounts or even lacking. Additionally, some specimens
that appear non-glandular do possess glands beneath
superficially
tomentum. Thus
the A. aromatica not consistently copiously
is
glandular, but rather variable in this regard.
Inasmuch as Antennaria pulvinata and A, aromatica are iden-
tical, the basis for inclusion of the former in A, rosea elusive.
is
If ^. pulvinata should be included some taxonomic rank the
at in
A, rosea polyploid complex, proposed by Bayer (1989b), then
as
the of should support con-
results canonical variates analysis this
tention. Support would include lower assignment rates for both
taxa, as well as lowered Geisser assignment probabilities. Neither
of however
these scenarios true in the present study. In cir-
is
cumscribing A. Bayer (1989b) concluded that subsp, pul-
rosea,
non
and This
vinata apomictic polyploid.
entirely
is pistillate,
sequitur unacceptable however as Greene 898) described both
is (1
The
"female" and "male" specimens of ^. pulvinata. protologues
of ^. pulvinata and A, rosea, gross morphology of type material
and
the results of canonical variates analysis, all contribute to
support two morphologically
the conclusion that the taxa are
distinct. The two A. pulvinata and A, rosea, do exhibit mor-
taxa,
»
Rhodora
270
95
[Vol.
and
phological integrity (see Table should therefore be treated
3)
as distinct. As such, would be inappropriate to include the
it
former species as an infraspecific subdivision of the latter.
Although an endemic, morphologically
originally described as
more
uniform, Antennaria aromatica
sexual, diploid species, is
and
appropriately described wider ranging (Chmielewski Chin-
as
nappa, 1988a; Bayer, both sexual and apomictic (Bayer,
1991),
1989a), diploid and polyploid (Bayer, 1989a) and morphologi-
cally variable. Gross morphology of type and non-type material,
the Pearson product-moment correlation coefficients presented
by Bayer (1988a), the protologues of aromatica and A, pulvi-
yl.
nata, the of using glandularity as a diagnostic character
fallibility
and
the results of canonical variates analysis contribute, to
all
and
varying degrees, to support the conclusion that A. aromatica
The deemed
A, pulvinata are conspecific. latter specific epithet is
A
name
the legitimate for the plant and the former a synonym.
synonymy
of provided
for A. pulvinata.
list is
TAXONOMY
The key may among An-
following be used the
to differentiate
tennaria media, and
A, pulvinata, A, rosea A. umbrinella.
A. dm, dm;
Plants generally greater than 1.5-2.5
typically
tall,
1
cauline leaves long, 1.5+ cm, numerous; basal leaves typ-
cm
1.2+
ically lanceolate, occasionally spatulate-lanceolate,
immature
long; involucres nodding A. rosea
A. Plants generally compact, than dm, 0.5-1.0
less typically
1
dm;
cauline leaves shorter, <1.5 cm, fewer; basal leaves
spatulate, obovate, spatulate-obovate, cuneate spatulate to
<
immature
occasionally oblanceolate, cm; in-
typically 1.2
B
volucres not nodding
B, Stolons elongate, strongly upsurgent and usually
at tips
woody;
basal leaves tufted about stem; of involucral
tips
brown
bracts dirty white or A. umbrinella
light
B. Stolons short, not upsurgent; basal leaves not tufted about
stem; of darker than
tips involucral bracts conspicuously
C
brown,
base, green, black
C. Basal leaves with per-
typically spatulate-lanceolate
manent
woolly tomentum; narrow,
inner bracts lin-
media
ear, tips usually sharp pointed, acute A.
. . .