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

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

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 ^ O O <N <N — Tj- OrN ^<ONN *fi^ NO0r0 <^N OmwO^ rON O^OO<NOiO/^ O"/^ ' 5 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 O ^ OOOOOW^U^TtsOO «0 <N OS '^ On 00 r^ -^ — O ^ O • • « « t ft OO -^ r^ 00 -^ -^ en <N r-i 3 M , J 4> ^ O — 4> ^H Tt Ov O^ -^ <^ v^^ ON li-l a uo <^ fN 00 -^ rs r- \0 <N — ^ O 1 t « • 4 V * 00 w^ r<i ^3 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 m — V 00 »o 00 NO 00 OON O — — 00 <N On NO 00 ; m O ^ T3 ON <N rn <N »n w^ f*^ r-* (N 03 oo oo ON NO OO oo V Q oo 00 r^ OC NO NO c O o O o O O o # o rs <^J <3 +1 +1 +1 +1 +1 +1 +1 +i +1 +1 +1 +1 +1 +1 — O Tj-<^^00W^Ot^<^ 3 NO rO <N ^OvO'^r^^'^OO — On OO w-i ^ ^ o o\ 00 -^ -^ -^ -^ lO -^ r^ >/^ o — — m 2 <Nsoo^NOTfTt *> ON At* sOONTtsOsor^NOr^Jsow^ O — r^ 13 I: n •a +i +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 "I O O rn 00 f^ 00 00 O O O r*^ f -a ,L' c ON NO rO w^ O^ (N m O '}i P3 »ri so tN r^ >0 */^ »y^ > O o i c ON 00 O^ p NO NO <N ON <N r^ r^ 00 w^ o o o o « * « * +1 + +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 o 1 w^ oo (N r^ O<^ "O rr r^ rsj Tj- <N 00 00 rj 00 tN <N f*^ «i^ "^^ m — 00 rt *^ y^ <^ rr r*^ CO oo' B B C/5 E CO E H c E B c E E E E. E E E s E E o E E -J E U < CuO >- s= -J X X 03 < < u < < o a. 9 a. EI a- U z > Z a. o:: o c/) en D < O < < < < z z ^ z o O u Z z a. O) ca cQ 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. . . .

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