Table Of ContentMadrono, Vol. 46, No. 4, pp. 205-207, 1999
NOTES
Observations on the Morphology and Geo- leaves, flagless1 upper cauline leaves, and the pa-
graphic Range of Antennaria—dioica (L.) Gaertn. pery laminae of the phyllaries are olivaceous to
(Asteraceae: Gnaphalieae). Randall J. Bayer, brown or black in color. The fact that the leaves of
CSIRO, Plant Industry, Australian National Her- all ten flowering stalks mounted on the herbarium
barium, GPO Box 1600, Canberra, ACT, 2601, sheet lack flags distinguishes them from A. alpina
Australia. (L.) Gaertn., which possesses prominent flags. The
length of the staminate florets is greater than 3.5
Antennaria dioica (L.) Gaertn. (Asteraceae: mm, which separates these specimens from the
Gnaphalieae) is widely distributed across Eurasia closely related diploid species, A. pulchella.mEm.
from the British Isles toJapan and its range extends Greene, which has florets shorter than 3.0
east into North America only in the western most (Bayer 1990). The observation that the adaxial sur-
Aleutian Islands (Bayer, 1993; Bayer and Stebbins, faces of the leaves are pubescent and the capituli
1993). It is characterized by having glabrous ad- are relatively small and the laminae of the phyllar-
axial leafsurfaces andphyllaries with pink orwhite ies are dark colored, distinguishes them from A.
laminae. The circumscription ofA. dioica in North marginata, E. Greene which has leaves that are gla-
Americahas long been a topic ofdebate, asA. mar- brous adaxially and has relatively large heads with
ginata (syn. = A. dioica var. marginata (Greene) white tipped phyllaries.
Jeps.) ofthe southwestern United States bears are- The three specimens (Fig. 1, arrows), referred to
markable similarity to A. dioica. The two species as A. dioica by Chmielewski (1998), appear to be
those with pink flecking in the lamina of some of
differ in that A. dioica has the upper flowering
stems lacking glandularhairs and the stolon surface the phyllaries, although the entire sheet, i.e., all ten
misarpgubiensacteantt,hebuutppneortfdleonwseerliyngwosotlelmy,iswuhseuraelalysbiensAe.t ksipe(cFiimge.ns1,).isAanfnouorttahtesdpaesciA.medniowiictahbypicnhkmifleelcekwisn-g
with purple glandular hairs and the stolon surface in the lamina is in a fragment folder on the sheet.
is densely woolly obscuring the surface of the sto- tItheapspaemaerscltohnaet,alals othfeitrhelseeavsesp,ecsitmeemnshecigahmte, pfhryolm-
lon. The species are also strongly allopatric, there-
lary color, and stage of maturity seem to be iden-
fsopreeciietss(eBeamyserbeasntdtoStreebcbiongsn,iz1e99t3h)e.tNwoo aosthdeirstsipnec-t tical (Fig. 1, arrows). I would determine these three
(four counting the specimen in the fragment folder)
cies in North America is so similar to A. dioica as specimens as either A. media or later generation
to pose a difficulty in determination. hybrids betweenA. media andA. rosea Greene, not
Recently, Chmielewski (1998) reported a large
A. dioica.
range extension forA. dioica from the Aleutian Is- The basal leaves of the specimens are heavily
lands to the Sierra Nevada, a distance of ca. 5,000 pubescent on both the abaxial and adaxial surfaces
km. Antennaria dioica (sensu Bayer and Stebbins, and resemble the other eight specimens ofA. media
1993) has not previously been reported for Califor- on the sheet. Antennaria dioica usually has gla-
nia (Stebbins and Bayer, 1993). After examining brous adaxial leaf surface, less often are these sur-
the herbarium specimen upon which the range ex- faces subglabrous (Tutin et al. 1976; Hulten 1968;
tension is based (California, Sierra Nevada, Bayer and Stebbins 1993). As for the phyllary col-
"Among granite fell-fields & scattered Pinus albi- or, my examination ofthe material leads me to con-
caulis near Margaret Lakes, east of Bishop Pass, clude that these three staminate plants do not bear
no. Inyo Co. Elev. 10,800 ft., 7-19-62, Betty H. "pink bracted involucres" (c.f. Chmielewski 1998).
Johnson #692," CAS 898202!), I came to the con- The base of the papery lamina is olivaceous-black,
clusion that this collection represents atypical in- the tips of some are white, whereas others are red-
dividuals ofA. media Greene (= A. alpina var. me- dish-pink orwhite withreddish-pink flecks. The en-
dia (Greene) Jeps.). tire involucre of these three plants gives the im-
Chmielewski (1998) states that "The single col- pression of being white spotted with reddish-pink.
lection ofAntennaria dioica that is the basis ofthis Typical A. dioica has heads in which laminae are
report consists ofthree staminate shoots . . .", how- entirely white, pink or rose. Phyllary color in A.
ever, also on the sheet are seven other staminate dioica is a sex-linked trait; the laminae of the sta-
flowering stalks and their associated basal leaves minate plants usually being white, whereas those of
and stolons.
These seven specimens are part ofthe same col-
lection and in my view represent typical staminate the1 tFilpasgsofatrheefplahty,lllairneiaers,nsoctartiooubse,ctoinpsfutsheadt awriethsiomridlianrartyo
plants ofAntennaria dioica. They have strongly pu- subulate or blunt leaf tips that are essentially green and
bescent basal leaves, narrowly spathulate basal herbaceous.
206 MADRONO
[Vol. 46
AustralianNationalHerbarium(CANBJ
Antennaria mediaE.L.Greene
mr. RandallJ.Bayer November1999
Antennaria dioica (L.) Gaertner
J.G. Chmielewski (SLRO)
X 2 3 4 5 6 7 8 9 10 11 12 1998
CALIFORNIA ACADEMY OF SCIENCES
PlantsofCalifornia 5ie.r?aK/«i-v«Mi&.
Fig. 1. Partial view of herbarium sheet (CAS #898202) showing staminate plants ofAntennaria media; reputed
specimens ofA. dioica indicated with arrows. Scale as indicated on figure.
1999] BAYER: OBSERVATIONS ON ANTENNARIA DIOCIA 207
the pistillate plants are normally pink or rose (von some pinkish or rose flecking in the laminae ofthe
Ubisch 1930), therefore the situation we see in phyllaries. The reddish-pink phyllary color may be
these specimens is the opposite of the usual phyl- due to introgressivehybridizationbetweenA. media
lary color in staminate A. dioica. Typical A. dioica and A. rosea. I maintain that the distribution ofA.
donotusuallyhave alarge darkolivaceous orblack dioica is mainly Eurasian, with a few populations
spot at thejuncture ofthe base ofthe phyllary and in the Aleutian Islands of Alaska. Therefore, An-
the papery lamina, as do the three (four) specimens tennaria dioica is not a part ofthe flora ofCalifor-
in question. I believe that the pink flecking in these nia, and the only North American populations are
specimens is simply the expression of a rare char- in the Aleutian Islands.
acter, notnormally seen inA. media. The samerose
flecking is also seen in the phyllaries ofsome spec- Acknowledgments
imens ofA. umbrinella Rydb. (see discussionunder I thank the curator of CAS for loan of the herbarium
A. umbrinella in Bayer and Stebbins 1993), whose specimen and to Laurie Adams, Greg Chandler, Lyn Cra-
laminae are normally light to dark brown. I also ven, Don Les, Rogier de Kok, and Michael Trebby for
have made the same observation inA. anaphaloides critical reviews of this manuscript. I am grateful to Neil
Rydb and A. pulcherrima (Hook.) Greene (Bayer Bagnall for measuring and scoring pollen grains for this
study.
personal observation). It is also possible that the
specimens in question represent later generation Literature Cited
backcross hybrids between A. media and A. rosea,
E. Greene, the pink or red phyllary characterbeing Alexander, M. R 1980. Aversatilestainforpollen,fungi,
yeast and bacteria. Stain Technology 55:13-18.
a typical trait ofA. rosea subsp rosea. Bayer, R. J. 1990. A systematic study ofAntennaria me-
The protoplasts of pollen grains from a typical dia, A. pulchella, andA. scabra (Asteraceae: Inuleae)
Scandanavian A. dioica (taken from CANB 11884 of the Sierra Nevada and Inyo Mountains. Madrono
and stained with Alexander's stain (Alexander 37:171-183.
1980)) are an average of 21.3 u- in diameter and . 1993. A taxonomic revision ofthe genus Anten-
87.5% of the grains stained viable. The dark-phyl- naria (Asteraceae: Inuleae: Gnaphaliinae) of Alaska
laried typicalA. media specimens on the sheet, i.e., and Yukon Territory, northwestern North America.
Arctic and Alpine Research 25:150-159.
tarheoseanwiatvheoruatgteheofpi2n7ki.s2h|fxleicnkidnigaminettehre pwhiytlhlar9i0es%, foratnhedgGe.nuLs.ASntteebnbninasr.ia19(9A3s.terAacseyaneo:psIinsulewaiet:hGkneay-s
viability. The size difference in the protoplasts is phaliinae) for North America. Canadian Journal of
typical of diploid (A. dioica) vs. tetraploid (A. me- Botany 71:1589-1604.
dia) pollen (Bayer unpublished). If the three spec- Chmielewski, J. G. 1998. Antennaria dioica (Asteraceae:
imens in question wereA. dioica one would expect Inuleae): Addition to the vascular flora ofCalifornia.
the pollen to be around 21 |x in diameter and to be Madrono 45:271-272.
rtheeasootnhaebrlyplvainatbsleo,ntthhues sdhiesetti.ngUunifsohritnugnattheelym,ftrhoemy Hultteonr,iesE:.A19m6a8n.uaFlloorfatohfeAvlaasscuklaaranpdlanntesi.gShtbaonrfionrgdtUenrrii--
versity Press, Stanford, CA.
are sterile, the anthers are empty, notproducing any Stebbins, G. L. and R. J. Bayer. 1993. Antennaria. in J.
identifiable pollen even though the flowering stalks Hickman (ed.), The Jepson Manual of Higher Plants
are of mature height and the phyllaries are mature. of California. University of California Press, Berke-
This is highly supportive of the idea that these ley.
plants may represent hybrids betweenA. media and Tutin, T. G., V. H. Heywood, N. A. Burges, D. M.
A. rosea, which grow in sympatry throughout the Moore, D. H. Valentine, S. M. Walters, and D. A.
Sierra Nevada. Webb. 1976. Flora Europaea, Vol. 4. Plantagmaceae
In summary, I believe the three specimens (four tsiotyCoPmrepsoss,iMtealebo(uarnnde.Rubiaceae). Cambridge Univer-
including the one in the fragment folder) in ques- von Ubisch, G. 1930. Geschlechtsverteilung und sekun-
tion on CAS 898202 {Johnson #692) to be some- dare Geschlechtsmerkmale bei Antennaria dioica
what atypical staminate plants of A. media with (Gaertn.). Biol. Zbl. 50:532-540.
