NOMENCLATURAL HISTORY AND TAXONOMIC STATUS OF ECHINACEA ANGUSTIFOLIA, E. PALLIDA, AND E. TENNESSEENSIS (ASTERACEAE)

8 Pages·1993·4.6 MB·English

by  Jerry M Baskin

AND
NOMENCLATURAL TAXONOMIC
HISTORY
STATUS OF ECHINACEA ANGUSTIFOLIA,
AND
TENNESSEENSIS (ASTERACEAE)
PALLIDA,
E. E.
SNYDER CAROL BASKIN
JERRY M. BASKIN, KRISTIN M. and
C.
School of Biological Sciences
University Kentucky
of
KY
40^06-0223
Lexington,
ABSTRACT
A
names
problem exists in the literature concerning the application of correct to Echhiacea
We
DC,
and
angnstifolia E. pallida (Nutt.) Nutt., and £. temmseensh (Beadle) Small. review clarify
the past confusion concerning the nomenclatural history of these three taxa from 1753 to 1991-
The contrasting views of A. Cronquisr and R.L. McGregor on the taxonomic status of/:.
angustifolm, E.pallicl-a, and E. tennesseensis^TQ briefly discussed. Based on various types ofevidence,
McGregors
these three taxa should be recognized as distinct species; thus, view is accepted.
RESUMEN
nombres
Existe un problema en la bibliograffa respecto a la aphcacion de los correctos a
DC,
Echinacea angnstifolia £. palltda (Nutr.) Nucc, y E. tenmssemsis (Beadle) Small. Se revisa
y
confusion pasada relativa a historia nomenclatural de estos tres taxa desde 1753 a
clarifica la la
McGregor estatustaxonomico
991. Los diferentespuntosde vista de A. Cronquisty R.L. sobre el
1
En
de E. angmtifolia, E. pallida, y E. tenncsseensis se discuten brevemente. base a evidencias de varios
como punto
tipos, estos tres taxa deben ser reconocidos especies distintas. Asi pues, se acepta el
de de McGregor.
vista
INTRODUCTION
In a recent study of the comparative ecology of the three closely-related species,
DC,
and
Echinacea angnstifolia E. pallida (Nutt,) Nutt., E. tennesseensis (Beadle)
we
Small (Snyder encountered considerable confusion in the literature
1991),
names For
concerning the application of the correct to these three species.
example, several Latin names have been applied to each of these taxa, including
the same name to different taxa and different names to the same taxon. Thus, it
whose
was evident that a knowledge of the nomenclatural history of these species,
geographical distribution well documented (McGregor 1968c), was required
is
summary
The
to accurately interpret the literature. purpose here is to present a
( 1 )
and
of the history of the nomenclature of £. angnstifolia, E. pallida, E. tennesseensis,
McGregor
and compare the views of A. Cronquist and R.L. on the taxonomic
(2)
status of the three taxa.
NOMENCLATURAL
HISTORY
A
summary from 1753
of the nomenclatural history of the three Echinacea- taxa
m
The which
to 1991 presented Table material in the table for references are
is 1
.
597-604.
SiDA 1993
15(4):
11
598 SiDA 1993
15(4):
TahiJ: I. SLimmary of the nonu-nchuural histor)- of Ec/j/^uuea cnigi/snjitliii^ II. pidlnhi, and /:. fcnfiesxecf/^/L
1753 LinnacLis dcscrihct! RinlhcckiLi p/nf/in\-i " * Rudhrkui pnypinwi L., the species of the L^cniis
t)'p{.'
1790 Neckcr descrilxxl genus Bviiiinoia (no species mentioned) Bvii/ourhi Necker.
tiie *
1791 iNLoench described clie genus Echifhicea based on Linnaeus' Rinlhijck'ui puvpiin.i *" VAJ)i}iiicva
Moench.
S34 T. Nurrail 8 V1) described Rihlkckut paUuL Kudkcku} pallida Nurc,
( >
I 1
'^})G A, DeCandolle described tour species of /:t/vJA/rcv/, one of which was E. iitii^NSfifolia " ^ EibiULneu
I
DCl
folia
Lni\\nst'i
8 T. Nutiall (1811) transferred Riidkxkia pallnla Nutt. to ELh'niaaia > Echinacea pallida (Nutt.)
I 1
Nurr., and also placed E^chitiacea augusttjoliii DC^ in synonym)' wirli E.piill'nLi, often written as
/:c/'/>A7av//j<7///i/./ Nutt. (e.g., Small L933; I'ernald 19^0; Cronquisr 1952, 1955; Gleason and
Cmnquisc
1963, 1991 Steyermark 1963; Hartley 1966; Radford 196S; Barker 1969;
et al.
;
Hitchcock and Croiiquist 1973; Swink an^l Wilhehii 1979; Molilenbrock 1986).
S1
1 T. Nuttall ( 8 1 ) Llescrllxxl Echinacea saiigidniCii ^ Echifh/cca Sinigaiiica Nutt.
I
1
8-1 1-94 Tiunking that E. i/i/gi/stijolia DC7 and E. pc/llida{N\nx.)'H\nT. were synon\'ms, authors of many
1
name
works
floristic [uiblished (.ku'ing this period use(.l the E. angustipolia for /:, pallidii [e.g.,
Torrey and Gray 842; Darby 1857; Woods 1872;Drendel 1887; Chapman 887; Warson and
I I
Coulter 1889, also eds. 2-5 of Gray's manual(1856-1867)].
189 Bnrton (189 made the combination EviUincna pallida (Nutt,) Britton, whicli incluckxl
1)
I
Rihlhcckia palluLi Nutt. [^ Echifiacca piillicLi {'H\\tt.)^utti] and E. angustifolia DC. as synonyms.
898 Bruton and Brown 89S) mcluck-d Rudheckui pallida Nucr. [- E. pallnla (Nutt.) Nutt.] and E.
1 (
I
under name
a}igu.\ii[i)lia the Ryyainicrla pallida (Nutt.) Britton
.
Hunking DC,
1898 that the long-rayed E. pidl/da (Nutt.) Nucc. was E. and unfamiliar
cuigiistijidiii
with the true /:. a7igiistij(dia DC. (Fernald 9<*t'), Beadle 898) ck^scribcd B}\naicria tcnncsscaisis
I ( 1
"** Braf/Jieria fetniesseefisis Beadle, which previously had been inclntkd in E.-angNstifnlhi DC7 (e.g.,
One
Gattinger 887). wonders Beadle would have described E. teiint'sseaisis if Britton (1 89-1)
1 if
made
had the nomenclatural blunder Fernald 1900).
[lot (cf.
1900
Fernald (1900) recognix.cd that E. pallida and angi/stifolia were different species, placed
/:. /:.
synonymy with and ins\^nonymy
.r<'///^i;///m^Nutr. in E^ra imeria Umu'sscois s^v:\(\\v witl
/:./?.////;/:/,
i ^
DC.
E.
angiistifolia
1900 Hl Her describes! Byaioieria aiigustipdia froni Tc^xas, the descrij:)tion of which marched (and a
is
synonym oQ E. august ijidia DC. *" Bra/niiriaaugNslijolia I leller, which often has been designated
as Byaiincria augNslipdia {r>C) leller (e.g., Britton 9(H Rydberg 906; Robinson and Fernald
\ 1
1
;
l908.Shimek
1011; Blake 1928; Stevens 1950).
1901 Britton (I9OI) recognized that he had iiicUided the names of two disrinet tuxa undi^r Byaiineria
Brown
pallida (Britton 1891, Britton and 1898), and thus used the names Byamicyia pallida
(Nutt.) Britton and B. august ijolia (DC.) Heller to refer ro these raxa. These names continued
ro be used In editions 2 and of Britton manual (Britton 1905, 1907).
^ s
1901 Gattinger 901), apparently following Britton and Brown (1898), used ihe name Byaiiueria
(
I
pallida (Nutt.) Britton for Bya/auria tcuuessamis Beadle, which previous to Beadle's (1898)
new
description of this species had been assumed to be the true Echinacea {Byaiiucyia)aug//sfifolia
DC.
of the western prairies and plains Gattinger 1887),
(e.g.,
1903 Small (1903) recognizet.1 Bca/nicyia pallida (Nutt.) Britton, />. augiistifidiii (I'^C.) Heller, and /3.
Beadle
temiisseifisis as distinct species.
I9O8 hi the seventh edition olGra)-'s Manual, Robinson and Fernald 908) recognized that E. pallida
(1
was
a separate species from Echinacea {Byainnria) angiistifidia ^ Byaiifuria a>igusfifolia (DC.)
Heller (including Tennesse^e material, i.e., B. lefiuesseensis) and B. pallnLi (Ntitt.) Britton.
1913 Britton and Brown (1913) recognized Echinacd/ pallid/ (Nutt.) Britton and E. august DC.
iftdia
as separate species.
1913 Small (1913) again recognized Byauneyia pallida (Nutt.) Brirton, B. august (DC.) Heller,
ifolia
and B. tenmsseensis Beadle as distinct species.
1929 Blake (1929) accepted Echinacea rather than Byauneyia, and made Brauncyia Beadle
tennesseensis
DC. DC.
a variety of Z:. augustijolia » E. augustijulia var. tennesseensis (Beadle) Blake.
and
Snyder
Baskin, Baskin, Echinacea 599
L
Table continued
1930 Johns (I93O) recognized Braunerla pdllida (Nutc.) Britton and Braumrui angHstifolia (DC.)
Heller separate
as taxa.
1932 Rydberg (1932) recognized Echinacea pallida (Nutt.) Britton and E. angustifolia DC. as distinct
species.
made
1933 Small (1933) the combination Echinacea tennesseensis (Beadle) Small based on Bva/inerta
Beadle.
tennesseensis
1942 Steyermark (1942) described Echinacea pall Nutt. forma albida Steyerm., which McGregor
ixia
(1968c) put in synonymy with pallida (Nutt.) Nutt. However, this white form may have been
E..
described from plants of E. nmidata McGregor. McGregor (1968c) has dots for £. nmulata, but
not for E. pallida, in the counties in southeastern Missouri where Steyermark 963) has symbols
( 1
for forma albida. According to R.L. McGregor (personal communication), '\ white-rayed
. .
We
lorms are found in E. angifstifolia, E. pallida^ E. siniulata^ and E.purpiirear have seen a white-
rayed form of E. simulata in Trigg County in western Kentucky and of/;, tennesseensis in Wilson
County, Tennessee.
950 In the eighth edition of Gray's manual, Fcrnald (1950) used the DC.
n'<xn\(^^Echinaceaangi/.stifolia
1
and E. pallida Nutt. The range given for E. angHstijolia does not include Tennessee, and no
mention made of
£.
is tennesseensis.
1952 Cronquist( 1952) did not presumably because he did not think that
!istZ:.^/«'^wj"f//f>/iV?separately,
the taxon occurred within the Britton and Brown manual range (R.L. McGregor, personal
communication). However, under E. atror/d?ens Nutt., he made the comment "Similar to E.
folia"
angiisti
—^
1955 A. Cronquist (1955) reduced £. angustifolia to a variety o^E. pallida E, pallida (Nutt.) Nutt,
Cronq. Hitchcock and Cronquist Cronquist and Gleason
var. angustifolia (1973), (1980),
iX^^^.)
and Cronquist (1991) maintained this classification.
On
1958 page 550 in the appendix of the second (1958) and subsequent printings ofTheNetv Britton
and Brown Illustrated flora, Cronquist (1958) stated that, 'The Great Plains phase of E.
"
pallida. .may be recognized as var. angustifolia (DC.) Cron
.
1959 The question on Echinacea vs. Brauneria was settled at the ninth International Botanical Congress
in Montreal, when Necker's 1790 name Branneria v^rs considered illegitimate. As Echinacea was
the validly published name, took priority over Brai/neria and became the genus name.
first it
E
1961 W.J. Dress (1961) maintained E.angnstijolia and pallida as separate species. However, he did
.
not recognize E. tennesseensis, as lie stated that "... it [E. angi/stifolia] also occurs in Tennessee.
1963 In the appendix added to the third printing o{ Handbook ofNorth Dakota Plants (Stevens 1963),
DC.
Brauneria angustifolia (DC.) Heller changed to Echinacea angustifolia
is
1968 McGregor 968a) describetl Echinacea speciosa McGregor. This name was applied to plants of
(1
KY
MO,
AR,
E. pallida (Nutt.) Nutt. in northcentral southeast southern IL, and wcstcentral
with yellow pollen and = in contrast to E. pallida (Nutt.) Nutt, which has white pollen
;/ 1 1,
and ;;= 22. However, since tlic binomial speciosa already had been used [synonym oi Echinacea
/:.
purpurea (L.) Mocnch], McGregor 968b) used the speci epithet simulata to describe the new
fie
( 1
species -^ E. simulata McGregor. According to Smith 978, 988), difficult to determine
it is
( 1
1
pollen color In herbarium specimens, and thus fresh pollen needed to distinguish these two
is
species in the field, since otherwise they are difficult to separate taxonomically (Smith 1973).
1968 McGregor (1968a) described a new variety of angustifdia DC. -* E. angustifolia DC. var.
Z:.
McGregor; becoming DC.
strigosa typical variety E. angustifolia var. angustifolia.
1970 McGregor's (1970) treatment o^ Fuhinacea species in the Texas flora sensu McGregor 1968c.
is
1980 A. Cronquist (1980) did not recognize E. simulata McGregor as a distinct taxon from E. pallida
(Nutt.) Nutt. var. pallida, or E. angustijolia var. strigosa as a distinct taxon from E. pallida var.
angustifolia; suggested that £, sanguinea Nutt. should be treated as a variety diE. pallida; and
considered E. tennesseensis (Beadle) Blake "an eastern outlier" o{ E. pallula (Nutt.) Nutt. var.
(DC.) Cronq.
angustijolia
1981 In a letter (dated 27 February 1981 to M. Baskin, R.L. McGregor states, ^'Iknow angustifolia
) J.
my
cjuite well and based on observations just cannot merge the two [E. angustifolia and E.
1
together Cronquist must be done."
tennesseensis} as insists
600 SiDA 1993
15(4):
Tahu; concinucd
1.
McGregor
9^6 McGregor's 98(^0 trcatnient o{Ecbhuia'a species of the Great Plains scnsu 986c.
is
1 ( 1 1
1987-90 Bauer and associates demonstrated that E. angiistijidhi and [hdlida can be distinguished by
/:.
Wagner
cnfferencesinthechemicalconsriruenrsin the roots (Bauer and 1987; Baiter cral., 1988;
Bauer and Remitter L989), but that tiie alkamides and caffeic acid derivatives in E. angustifuliu
and icnfiesseensis "... appear to be very similar" (Bauer et al. 1990),
11.
Thomas
989 Gandhi and 989) reduced sangimiea to a variety of/:. puUiihi -* E. pdU'ida (Nutt.)
1 ( /:.
1
Nntt. van sanguines (Nutt.) Gandhi Thomas.
(Sc
1990 Hatch et al. (1990) used the combinacion llchithtcdi pcdlidii (Nutr.) Nutt. van slrigosa (R.L.
McGreg.) Gandhi, based on DC. R.L. McGregor.
iingi/stifoUci var. slrigosa
/:.
99 Cronquist (Gleason dud Cronquist 1991) continued to recognize aNg//s/i/o//a as a variety of/i.
1
1
pcilluL, and suggested that E. .uw/zA/A/ perhaps shoidd be recognized as another variety o^hillicLr
"An
le states that, eastern outlier of van angustiful'u!^ in the cedar glades of c. Tenn., has been
I
called tennesseefjsis"
/T.
1993 In a comparative study of the [X)pulation genetics of/:, angnst'ifolid van ungustifidia and
/:.
tcnmsscmm, Baskauf 993) concluded that, "£, laincsseen.ds and ctngusi'ifidla are genetically
/:.
(
1
»
distinct taxa.
McGregor
not cited was taken from Fernalcl 900), Sharp (1935), Dress (196
( ),
1 1
(1968c) and/or Foster (1984).
AND MCGREGOR
VIEWS OF GRGNQUIST
more
Authors of recent systematic works that include one or of the three
who
Echinacm species are divided between those follow the tcixonomic views of
Cronquist 198())-e.g.,Scoggan(1979), Gandhi and Thom^is(1989),and
1955,
(
—
who McGregor Van Bruggen Mohlenbrock
those follow (1968c) (1976),
e.g.,
Dorn
^itskievych and Turner and Taylor and Taylor
(1986), (1988), (1990),
A
McGregor
199 !) comparison of the views of and Croncjuist on the taxonomic
(
and
presented Table
status of E. angNstifol'ui^ E. E. terinesseensh in 2.
pciIlicL^i, is
Whereas Oonquist
one with
recognizes species, E. pall'uLi, three varieties,
McGregor
recognizes five species and one additional variety, E. angiistifol'ia var.
strlgosa. Tlius, Croncjuist does not recognize either £. simulata (But see next to the
item Table
last in E. augastiforia var. strigosa, or E. teinmseensis as distinct taxa.
1
.)?
The
demonstration chemical and
ol differences in the roots of E. august alia E.
if
pallida by Bauer and associates (cited in Table lend support to McGregor's
1)
And
968c) view two although no
that these taxa are distinct species. differences
( 1
were detected in the root chemistry of £. angustijolta and E. tennesseemh (Bauer et
I99O), Baskauf (1993) convincingly demonstrated by use of allozyme
al.
two
electrophoresis that these entities are genetically distinct taxa. Intraspecific
genetic identities differed signihcantly from interspecific identity (Baskauf
We
conclude and should be
1993)- that E. angustifolia, E. palluL/, E. teunesseensis
McGregor
recognized as distinct species (sensu 1968c), based on a combination
ofdifferences found in comparative studies on the morphology, anatomy, chromo-
some number, geographical distribution (McGregor 1968c), root chemistry
(Bauer and associates, see Table and genetics (Bciskauf 1993).
1 ),
SnYDBR and
BaSKIN, BASKIN, Echinacea 601
Table 2. The two current: views on the caxonomic sracus of Echinacea ang/Lstifolia^ E. pallnl^jy and E
tennesseemis.
A. Cronquist(1955, 1980)
1.
Echinacea
pallida (Nucc.) Nucc. van pallida
McGregor
E. smulata (But see next to last item in Table
1
.)
E. pallida (Nutc.) Nucr. van angustifolia (DC.) Cronq.
DC.
£. angustifolia var. angustifolia
DC. McGregor
E. angustifolia var. strigosa
E. tennesseensis (Beadle) Small - "...an eastern outlier of var. angustifolia in the cedar barrens of c
Tenn. ." (Cronquisr 19^0) (Also see next to last item in Tible
I .)
.
E. pallida (Nutt) Nutt. sanguinea (Nutt.) Gandhi and Thomas [Varietal status of sangianea
var.
/:.
was suggested by Cronquist (1980), and formal combination made by K. Gandhi and R. D.
Thomas
(1989).}
McGregor
R.L. (1968a)
2,
Echinacea pallida (Nutt.) Nutt.
simulata McGregor
E.
sanguinea Nutt.
E.
DC.
E. angustifolia var. angiiscifoHa
angustifoha DC. strigosa McGregor
E. var.
E, tennesseensis (Beadle) Small
ACKNOWLEDGMENT
We
Tom
Wayne
thank Bloom, Kentucky Nature Commission,
State Preserves
Chester, Austin Peay State University (Tennessee), and Barney Lipscomb,
Botanical Research Institute ofTexas, for providing us with some of the hterature
cited in this paper.
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