A NEW HEDGE-NETTLE (STACHYS: LAMIACEAE) FROM THE INTERIOR HIGHLANDS OF THE UNITED STATES, AND KEYS TO THE SOUTHEASTERN SPECIES

2008·7.3 MB·English

by  John B Nelson

NEW
A
HEDGE-NETTLE
LAMIACEAE)
(STACHYS:
FROM
THE INTERIOR HIGHLANDS OF THE UNITED
STATES,
AND
TO THE SOUTHEASTERN
KEYS
SPECIES
John
Nelson
B.
Department ofBiological Sciences
University ofSouth Carolina
Columbia, South Carolina 29208,
U.S.A.
ZUSAMMENFASSUNG
Stachys is one of the larger genera within the Lamiaceae, comprising nearly 300 species, and nearly cosmo-
from
politan, absent Australasia. Its centers of species diversity include warm-temperate portions of western
Asia and the Mediterranean, southern Africa, and portions of North and South America (Hcim 2008).
et
al.
The genus in North America, north of Mexico, contains about 45 species, particularly concentrated west of
Rocky Mountains and
the (especially the Pacific states) in the east, along the Atlantic seaboard states and
in the Appalachian region. Southeastern species occur in a wide variety of habitats, most often in mesic
sites, and at a broad range of elevations. The North and South American species were studied in by
detail
Epling (1934); Mulligan and Munro (1989) provide a comprehensive assessment of the North American taxa
north of Mexico. Ongoing studies in the genus show that an additional species should be recognized.
Stachys
iltisii Nelson, sp. nov. (Figs. 1-3). Type: UNITED STATES. Arkansas. Logan Co.: crowded population on rocky
J.
m
Potentially robust perennial herbs to 1 tall or taller from vigorous, pale, fragrant rhizomes, often in dense
commonly
colonies; steins erect, occasionally branched, older plants branched; stem sides and angles
densely pubescent with mixture and
a of glandular eglandular hairs, the glandular hairs capitate, the eg-
mm
landular hairs with 3 or more cells, to 3 long, spreading and reflexed; leaves spreading, somewhat lax,
densely pubescent on both surfaces with glandular and eglandular hairs, the latter mostly appressed on
upper surface, and erect on lower surface, the lower surface equipped as well with scattered
sessile (or
musky
nearly sessile) capitate glands; the crushed leaves with a walnut-spice-fish scent; mid-stem leaves
4-6 cm cm
commonly
broad, 10-17(-19) somewhat narrowed
long, lyrate basally, featuring a waist, with
distally convex margins; inflorescences commonly elongated maturity the lowest and
at (fruiting), oldest
several verticils usually slightly separated, the more distal and younger verticils more or less crowded to-
gether, thus nearly compact; cymules each with 6-8-flowers, the thus 12-16-flowered, mostly
verticils
(turbinate)-hemispheric to nearly spherical; open flowers are visited by a variety of insects, including hon-
and
small bumblebees, and wasps; calyces campanulate, densely pubescent with glandular eglan-
eybees,
mm
mm
dular tube (3.2-)3.8-4 long, the lobes less that one-half the length of the tube, 1.5-2 long,
hairs,
mm
minutely corollas pink, 11-11.5 long from base to tip of galea, galeae frequently
deltoid, apiculate;
mm
90° more with
notched or indented apically; the lower lip generally declined at full anthesis, so age, 6
prominent pink-purple blotches and spots on adaxial surface; corolla tube prominently sac-
long, featured
toward base on lower internally glabrous, but with prominently slanting (oblique) annulus, this
cate side,
mm mm
copiously pubescent with bulbous trichomes; mericarps 1.5 wide, 1.5-1.6 long, dark brown,
soft,
and minutely verrucose.
irregularly
-eFlore Co.: M. Hopkins
A new
Nelson, species of Stachys from the Interior Highlands of the US
endemic
s to the Interior Highlands of Arkai
made
is that by E. Palmer in 1920, from Izard County, Arkansas, which
represents the eastern-most
loca-
J.
tion for this plant. Its first recognition in the literature was made by Epling who considered
(1934), ad-
it,
known
ditionally by then from collections in Oklahoma, as a thin-leaved and unnamed variant of Stachys
nuttallii Shuttlew. ex Bentham. Fernald (1950), despite considerable disagreement with Epling's assessments
on
Stachys in general concurred
(1943), that the plants of the Interior Highlands belong in as did
S. nuttallii,
Hugh
Gleason (WIS) by way on
(1952). litis of annotation the original label of his #5374 (BRIT) from
AR
Magazine Mountain, Logan County, on 9 Jun 1955, suggested that these plants rightly constitute
a
recognizable entity within S. nuttallii, offering "ssp. nov. arkansanus" for Nelson and Fairey (1979) es-
it.
sentially concurred with Epling in recognizing these plants as distinctive within the complex,
S. nuttallii
but maintained them being and new
as a part of a distinctive taxon, they
S. eplingii; treated the plants of the
Interior Highlands as forming the western (and disjunct) end of the range of S. eplingii (the eastern distribu-
tion consisting of primarily Appalachian portions of West Virginia, Virginia, Maryland, Colum-
District of
and
North The somewhat
bia, historically, Carolina). conservative approach to the taxonomic status of these
plants taken by Nelson and
Fairey at the time is evident within the protologue of their description of
S.
eplingii: they list (1979, p. 493) three specimens from Arkansas as paratypes for (Demaree 17743
S. eplingii
from Logan
County; was
Miller 187 "Smith" used for this collection, but in from Montgomery
... error...
County; litis 5175 from Polk County). Their listing these paratypes for a different taxon has no bearing on
the matter hand, and
at these three collections presently serve as paratypes
for (see above).
S.
iltisii
This taxonomic position was maintained by Nelson and by Mulligan and Munro whose
(1981),
(1989;
formulation most
the recent treatment North American
is for all the species of by Gleason and
Stachys),
Cronquist and by Smith
(1991), finally (1994).
Recent field investigation in Arkansas and Oklahoma (during one of the hotter and stormier, lightning-
summers
filled on record, 2007) and study of considerably more herbarium material since 1979 reveals that
the opinion expressed by Epling and was
litis legitimate, at least in the sense that the plants of the Interior
Highlands
are distinctive within the S. nuttallii "complex." There no question that these plants share many
is
features with S. eplingii, and the two entities are surely related. Substantial differences between these two
species are apparent, and based on more
recent collections, it is obvious that the two taxa can be consis-
tently separated, both from fresh material, and from herbarium specimens. Both
species bear short-stalked,
capitate glands on the abaxial leaf surface, with unicellular, spheroidal heads, 25-30 urn in diameter. The
lower
leaf surface of sparingly glandular, while and
S. is that of densely prominently
iltisii S. eplingii is
atomiferous-glandular (Fig. 1), to the point of having a somewhat shiny, golden surface (with magnification)
below
the non-glandular pubescence, as well as often leaving something of a yellowish, slightly oily stain
on newspapers when The
pressed. leaf blades of are longer, broader, and somewhat thinner
S. iltisii in texture
than
those
of S. eplingii. Additionally, the leaf blades of are frequently somewhat narrowed
S. iltisii in the
lower third, thus somewhat lyrate, and widest toward the middle. Stachys eplingii has more ovate leaves,
usually widest in the lower third. In S. iltisii, the apex of the early blooming stem is relatively delicate and
somewhat
flexuous, while The
that in commonly
S. eplingii is stiffer. fruiting inflorescence of
S. iltisii is
compact, 1.5-2cm between
the lowest whereas commonly
verticils, that of S. eplingii exhibits more space,
(2-)3-4cm between
the lowest
verticils.
Stachys eplingii is found in boggy places within the Appalachians. most common from West
habitat
Its
Virginia into northern Virginia includes wet, saturated meadows. Stachys is generally found in upland
iltisii
commonly
forested habitats, on thin soil of rocky places, in both the Ozark Plateau and the Ouachita Moun-
on
tains, thus both sides of the Arkansas River Valley, though probably more widely
distributed in the
Ouachitas. Although
substantial differences in geology and soil types exist between the Ozark Plateau and
the Ouachitas, sites occupied by do not appear be unusual and
5. iltisii to or particularly special, in
fact,
Journal of the Botanical Research Institute of Texas 2(2)
OKLAHOMA
make where frequently grows with weedy plants such as
open tend excellent habitat for
roadsides to it, it
Common woody disturbed include
Daucus and Ambrosia associates, in less sites
Coronilla varia, pusilla, trifida.
and albidum.
Ostrya virginiana, Sassafras
Acer rubrum, Robinia pseudo-acacia, Juglans nigra,
Quercus
alba,
grew with Toxicodendron radicans, Parthenocissus
during
Nearly every population located this field effort
and Rubus
occidentalis.
quinquefolia,
An endemism Highlands not the purpose of this paper. This
overview of biological in the Interior is
and discussed
number and animal species,
been amply addressed considerable of plants is
subject has for a
summary. Zollner (2005) provide
Robison and Allen (1995) provide a useful et al.
in the recent literature;
Highlands species plus 12 subspecies
an enumeration of the vascular plant taxa endemic to the Interior (24
between
more widely distributed species). Vicarious distributions of plant taxa the Interior
or varieties of
Highlands and the Appalachians have been long recognized: Steyermark (1934) lists 87 vascular taxa of
examples include the distributions
Ozark occur in the southern Appalachians. "Classic"
the Plateau that also
The
and perhaps Halesia Carolina (Little 1970).
Robinia pseudo-acacia,
of Magnolia tripetala, Cladrastis lutea,
some
presumes prob-
two
common notion regarding disjunction of the same species, or of related species,
&
by evolutionary divergence.
interconnection in the past (Hardin Cooper 1967), followed potential
able
forming species "pairs" in the
been demonstrated species
and subsequent speciation has for fish
Vicariance
&
such
Burr and various examples exist for vascular plants,
Ozarks and the Appalachians (Strange 1997),
commonality
and such potential
as Castanea ozarkensis and C. pumila. In the case of Stachys iltisii S. eplingii,
km
now Approximately 1000 (640
no longer and the two entities are fully separated.
of distribution exists,
Based upon the isolation of the plants within the
mi) currently separate their nearest points of distribution.
on
based
and upon the ease of separating plants
Highlands from other related hedge-nettles,
Interior all
simple morphology, Stachys is here recognized and described at the level of species.
iltisii
Personnel within the Arkansas Natural Heritage Commission (2005) list Stachys iltisii as S. eplingii (thus
&
rank "INV" (Inventory Element, "of conser-
and have assigned as
following Nelson Fairey 1979), its state
The Oklahoma Natural Heritage Inventory (2003) similarly maintains S. iltisii as S. eplingii,
vation concern").
Arkansas and
At 40 known populations are verified for 13 counties in
tracking as a rare species. least
it
on
Arkansas (Howard and Scott Counties) but not
Oklahoma. Reports of additional county occurrences in
known
with range
specimens seen from them, consistent the (Fig. 2).
is
comes from
United
Reference to this species as an element of the flora of the Southeastern States its
recognized by botanists as a portion of the
delimited area generally
presence in Arkansas, a politically
living plants (population of Nelson 26653,
(/: L
many
Although Oklahoma not Southeastern eastern portions do indeed harbor
"Southeast." a state, its
is
"Southeastern" species.
"Ozark may be an appropriate common name. This a handsome species, often robust
hedge-nettle" is
moving Due tendency toward
and conspicuous (Fig. and fairly easy to spot from a vehicle. to their a
3),
somewhat compact inflorescence, especially in the sun, the plants do resemble Teucrium canadense... but
commonly confused with "Canada germander," frequently causing field-trip
then, various Stachys species are
new
Etymology.— unofficial differentiation of this plant as a taxon deserves recognition. Its occur-
litis'
rence in Oklahoma as well as Arkansas, however, makes the choice of the epithet "arkansana" (Stachys as a
Hugh
noun requires a feminine ending) unwarranted. As a way of commemorating litis' interest in the
and genus and well acknowledging his tremendous
vegetation of the Interior Highlands, in the Stachys, as
named
contributions to botany, this taxon is thus in his honor.
common, and
Except that Stachys and S. eplingii are (were?) never really that there is a relatively
iitisii
low number of historic collections of each, it is somewhat remarkable that Epling (1934) did not consider
formally recognizing them in 1934. On the other hand, Stachys in the Southeastern United States has com-
monly been considered troublesome genus. current taxonomy fairly stable, although several of the
a Its is
characters stem pubescence, petiole length, and calyx lobe dimensions) useful in separating taxa are
(e.g.
:
variously continuous. The considerable variation in some species groups suggests future usefulness to be
attained from cytological and breeding system studies.
taxonomy demands To this end, two keys are provided for distinguish-
Nevertheless, practical results.
primary
ing Stachys from the other Southeastern species. The first uses relative petiole length as a
iitisii
distinction, the second using shape of the calyx lobes. The different structuring of these keys is provided to
may
emphasize different characters; with a particular plant or specimen in hand, the user find that one or
the other key provides the easier route to identification.
prominently exserted from cah widespread.
Perennial; corolla /x,
middle portion of the stem at least 1/5 as long as the leaf
?
blade).
Calyx tubes glandular.
3.
mm),
4. Leaf blade margins deiTtate or nearly s o; stem a gles copiously pubescent with long (to 3
mo
Leaf blade margins crenate; stern angles glabrate or pubescent with mostly short, retrorse
4. stly
Leaf blades <3 cm wide; plants perni ciously weedy, commonly in dense patches, scarcely to .75
5.
minatinq rhiznmes S.I
m
commonly
Leaf blades commonly >3 cm wide; lants not weedy, never in dense patches, tall
5. p 1
rounded!
of greater;
margins IN-OH-WV-KY-VA S
rrpnulafp;
Leaf blades ovate, tP-
6.
Leaf blades
6. elliptic
Calyx not glandular, or ver /slightly so.
3.
Calyx glabrous to sparsely pubescent; petioles well-developed, es|
.
Calyx variously hairy, but at least hispidulous, frequently strongly hispid; petioles short or long.
.
Petioles usually well developed; stem angles abundantly pubescent with spreading hairs; blade
8.
commonly clingmanii
margins dentate S.
stem
Petioles short to nearly absent;
8.
i
margins but not dentate
serrate,
5mm)
Dies short (the midstem leaves with petioles less than or absent.
1
»w species of Stachys from the Interior Highlands of the US
Leaf blades abundantly pubescent below
with appressed
h
.
surface
felty or velvety; corolla white, pink, or purplish.
Stems angles and abundantly wr
sides pubescent,
soft
white; adventive
rare (SC)
Stem angles and sides abundantly pubesce
1 1
.
retrorse hairs, corolla purple;
r
nulate
hyssopifolia
S.
.
mm
1 2. Leaf blades broader, 5-8 wide; plants frequently hispidulous or at least moderately pube-
cent, or abundantly hairy; blade margins crenulate to serrulate aspera
S.
Leaves ovate to elliptic, widest near the center or toward the apex, oblong; leaf margins crenate to
.
sharply serrate for nearly the entire length.
Stem commonly
sides pubescent above.
3.
1
5. Leaves generally subsessile, the petioles thick, not longer than 6mm; blades densely pubes-
cent, frequently soft-pilose to scaberulous; leaf margins serrate; introduction from farther
15mm
15. Leaves generally petioled up to long, although commonly and
shorter, relatively
more
-Itoid; flowers 8 or per verticil; leaf margins crenate-crenulate; native with
s
ution,OK-AR,WV-MD.
surface densely and
f sessile- stipitate-glandular, blades ovate, widest lower
in
adjacent upper separated
verticils (NC)
af;
:
f surface glandular, but not densely so; blades elliptic-ovate, widest at middle;
'. Fruiting calyx lobes lanceolate, hispidulous to hispid, about half as long as the calyx tube hispida
S.
'. Fruiting calyx lobes deltoid or triangular, shorter than half the length of the calyx tube.
18. Leaf margins sharply serrate, nearly always with sharp teeth; bracts gradually reduced upward
from lowest flowering node, leaf blades elliptic, rounded to truncate at base; Blue Ridge
Leaf margins more
8. serrate to crenate; teeth often rounded; bracts abruptly reduced u
1
from lowest flowering node;
leaf
fc
VA-NC-TN-SC-GA
SOUTHEASTERN
STACHYS: KEY
#2
Fruiting calyx lobes deltoid to broadly mostly <1/2 tube
triangular, length.
.
Stem
2. sides pubescent; foliage glandular, often with musky scent when crushed.
Petioles short to absent, blades mostly rounded
3. to truncate-cordate.
Lower
4. leaf surface densely glandular, with atomiferous/short-stipitate golden glands; blades
prevail-
WV-VA-MD
ingly ovate, broadest toward base; (NC)
Lower
4. surface glandular, but not densely; blades prevailing somei
elliptic, base,
broadest toward midblade;AR-OK
commonly
3. Petioles well developed, blades cordate at the base.
5. Top of blooming stem frequently flexuous, somewhat blades ovate-rounded, IN-OH-WV-KY-VA-
lax;
Leaf blade margins entire to crenate; pic
9.
margins with
Leaf blade serrulate at leas
9.
Stems or sparingly branched; k
0. strict
1
VA
merely SEUS, adventive
o felty; rare in in fi
pubescent,
Blades t
.
from widespread.
prominently exserted calyx;
rennial; corolla
commonly glandular on stems, leaves, and within inflorescence.
Plants
.
segmented, rhizomes; blades oblong; [weedy,
producing tuber-like
Plant thick,
18.
s
mostly of coastal plain
iizomatous, but not producing thick, tuberous thickening of rhizomes; blades
8.
on and within inflorescence.
not glandular stems, leaves,
Plants
.
abundantly pubescent with eglandular hairs; fruiting calyx lobes
19. Calyx usually stiff,
jpiculate
_
ACKNOWLEDGMENTS
Commission sharing
am Arkansas Natural Heritage for
Theo Witsell and other personnel of the
grateful to
I
from Oklahoma Natural Heritage Inventory, for similar
and personnel the
information from their also
files,
US (Ouachita and Ozark National Forest units) for providing col-
thank the National Forest Service
help.
I
were provided through the W.T.
Funding support work and for publication costs
lection permits. for field
Endowment Moore Herbarium, USC. Kathleen Ross (Department of Languages, Litera-
Batson the A.C.
for
Johannes Stratmann (Department of Biological
and USC) provided the Latin diagnosis.
Cultures,
tures,
USC) and Hartmut Hilger (Systematische Botanik und Pflanzengeographie, Frei Universitat, Ber-
Sciences,
USC)
provided, usual, his
helped with the abstract. Clint Cook (Department of Biological Sciences, as
lin)
Zack Murrell (BOON), Bert Pittman (South
and within paper.
constant help with the graphics figures this
on
and Alan Weakley (NCU) graciously provided reviews the
Carolina Department of Natural Resources),
who
am George Johnson the herbarium (APCR) of Arkansas Tech University,
especially grateful to at
I
me new
me work, and in introducing to a
was so accommodating in helping with the field aspect of this
pressing technique.
field
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Arkansas Natural Commission. 2005. State species of special concern Plants. available from Depart-
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of Arkansas Heritage, 1500 Tower Building, 323 Center Street, Little Rock, AR 72201. http://www.
url:
naturalheritage.com/program/rare-species/. Accessed
July 2008.
Epling, C.C. 1934. Preliminary revision of American Stachys. Feddes Repert. Spec. Nov. Regni Veg. 80:1-75.
Ml.
Fernald, 1943. Virginian botanizing under Rhodora 45:357-516.
restrictions.
Ml
Fernald, 1950. Gray's manual of botany, 8th American Book Company, New
Edition. York.
New
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Bot.
1
New
York.
Gleason, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and Adjacent
New New
Canada.
York
Ed. Bot. Gard., York.
2.
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Hardin, J.W. and A.W. Cooper. 967. Mountain disjuncts the eastern Piedmont of North
in Carolina. Elisha Mitch-
1 J.
39-1
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83:1 50.
ell
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Lcim, A., C. Menderes, and Dadandi. 2008. Stachys marashica (Lamiaceae), a species from Turkey. Ann.
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In:
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University, Blacksburg. Pp. 249-290.
Taxonomy
Mulligan G.A. and D.B. Munro. 989. of North American species of Stachys (Labiatae) found north of
1
Mexico. Naturaliste Canad. 16:35-51.
1
Nelson, 981 Stachys (Labiatae) southeastern United Sida 04-1
J.B. in States. 23.
1 9:1
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name
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1
species. Brittonia 31:491-494.
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1 in
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