(Stachys: Lamiaceae) from the - Botanical Research Institute of Texas

A new hedge-nettle (staChYs: lAmiAceAe) from the
mid-AtlAntic Piedmont And coAStAl PlAin of the united StAteS
Gary P. Fleming John B. Nelson John F. Townsend
Virginia Dept. of Conservation and A.C. Moore Herbarium Virginia Dept. of Conservation and
Recreation, Division of Nat. Heritage Dept. Biological Sciences Recreation, Division of Nat. Heritage
217 Governor St., 3rd Floor University of South Carolina 217 Governor St., 3rd Floor
Richmond, Virginia 23219, U.S.A. Columbia, South Carolina 29208 U.S.A. Richmond, Virginia 23219, U.S.A.
gary.fleming@dcr.virginia.gov nelson@sc.edu john.townsend@dcr.virginia.gov
abstract
A new, geographically restricted “hedge-nettle,” Stachys matthewsii (lamiaceae) is described from the region that includes southeastern
and south-central Virginia and north-central north carolina. this taxon resembles stachys latidens Small ex Britton, a species of
higher-elevation forests of the Blue ridge mountains, but has consistent morphological differences in stem pubescence and has distinctly
different habitat requirements. two keys are provided for distinguishing hedge-nettles of the southeastern united States.
zusammenFassung
Zur Beschreibung des neuen Ziests, Stachys matthewsii (lamiaceae), lässt sich sagen, dass seine geographische Verbreitung auf den
südöstlichen und den mittleren Süden Virginias und den mittleren norden north carolinas begrenzt ist. die neue Art ähnelt stachys
latidens Small in Britton, einer Pflanze aus den höher gelegenen “Blue ridge mountains,” aber sie hat konsistente morphologische
unterschiede in der Stammpubeszenz und hat völlig andere habitat-Ansprüche. Zwei Schlüssel dienen zur Bestimmung der Ziest-Arten
aus den südöstlichen uSA.
resumen
Se describe una nueva “ortiga muerta,” Stachys matthewsii (lamiaceae) restringida geográficamente a la región que incluye el sureste
y sur-centro de Virginia y norte-centro de carolina del norte. este taxon se parece a stachys latidens Small ex Britton, una especie de los
bosques a mayor elevación de las montañas del Blue ridge, pero tiene diferencias morfológicas consistentes en la pubescencia del tallo
y distintos requerimientos de hábitat. Se aportan dos claves para diferenciar las xxx del sureste de los estados unidos.
introduction
field work and herbarium studies in north carolina and Virginia has led to the circumscription of a new
species in the genus stachys, a large group of mints widely recognized for its taxonomic complexity. changing
taxonomic interpretations in the past have confounded efforts to classify this plant properly, but enough has
now been learned to recognize this species’ unique place within the genus. A history of taxonomic opinions
regarding stachys matthewsii is presented, as is a discussion of habitat and geography, particularly its primary
range in the Piedmont and its somewhat disjunct coastal Plain populations in Virginia.
Stachys matthewsii g.P. fleming, J.B. nelson, & J.f. townsend, sp. nov. (Figs. 1–5). type: united StAteS: north
carolina. montgomery co.: fairly shady woodland on e side of Yadkin river, just downstream from falls dam, uwharrie national
forest, 11 Jun 1979, John B. nelson 1152 with J. matthews (holotype: uSch; isotypes: fSu, mo, ncu, uncc[2]).
differt a stachys latidens ob trichomata longa expansaque ad caulis angulas. differt a stachys hispida ob calycis lobos breviores et minus deltatos
pro parte. differt a stachys cordata ob folia parviora, minus cordata, et petiolos breviores. differt a stachys nuttallii ob caulis facies leves.
Perennial herbs from vigorous, pale, fragrant rhizomes; flowering stems erect, 0.8–1.05m tall, sparingly
branched, even within inflorescence, or branched toward base on older plants; the sides glabrous as high as
the lowest verticil, fertile internodes with scattered short glandular hairs, the angles prominently and consistently
pubescent with stiffish, spreading (or retrorse) hairs, these eglandular, to 3mm long; nodes bearded
with hairs as those on the stem angles; the overall pubescence is dense at the base of the plant, becoming
progressively less so upwards; leaves consistently petioled, the petioles 2–5mm long, blades mostly rounded
at base or somewhat cordate, the blades 5–10 cm long, 2–5cm broad, broadly elliptical to lance-ovate, acute
J. Bot. Res. Inst. Texas 5(1): 9 – 18. 2011

10 Journal of the Botanical Research Institute of Texas 5(1)
at apex, the margins crenate (crenulate-serrulate), leaves spreading-ascending, upper surface prominently
bullate, abundantly pubescent with appressed and erect non-glandular, pustulate-based hairs, the lower
surface glabrate to softly pubescent with erect, eglandular hairs; inflorescences of interrupted verticils;
bracts abruptly reduced upward from lowest fertile nodes; cymules each with 3 flowers, the verticils thus
6-flowered; fruiting calyces campanulate, the tube densely short-soft pubescent, and with longer, stiffish
hairs, especially along the veins, and slightly atomiferous-glandular, tubes mostly 4mm (3.5–4.5mm) long;
the lobes deltoid, narrowed apically and terminated with minute apiculum, the lobes less than one-half the
length of the tube, 1.5–1.8(–2.4)mm long; corollas pink, corolla tube prominently saccate toward base on
lower side, tube internally glabrous, but with prominently slanting (oblique) annulus, copiously pubescent
with soft, bulbous trichomes; tube 11–14mm from base to apex of galea; galeae abundantly pubescent with
stalked glands and soft eglandular hairs; lower lip with prominent dark pink (nearly purple) blotches on
the median portion of the lower lip; declined to 90° from the tube, 6.5–8mm long, the middle lobe of lower
lip narrowly ovate, slightly notched at apex; mericarps dark brown, minutely pebbled, 1.4–1.5mm wide,
1.8–2mm long.
Additional collections examined: UNITED STATES. VIRGINIA. Amelia Co.: local in wet meadows and old fields, 5 Jul 1938, J.B.
Lewis 1592 (fSu). Brunswick Co.: along wooded stream, Jun 1942, J.B. Lewis 3588 (VPi). Charlotte Co.: young floodplain forest along
cargills creek, 3.5 mi e of dryburg, 14 Aug 2008, Gary P. Fleming 15453 with John F. townsend (VPi). Halifax Co.: moist swale in powerline
clearing 2.2 mi Sw of Public fork, 14 Jul 1995, , Gary P. Fleming 10804 (gmuf, uSch, VPi, willi); same location, 1 Jul 1997,
Gary P. Fleming 13418 (unc, uS); same location, 14 Aug 2008, John F. townsend 4061 with Gary P. Fleming (VPi); thinly wooded toe slope
along Staunton river, 0.6 mi. south of uS 360 bridge and 2.3 mi. west-southwest of Public fork, 26 may 2010, Gary P. Fleming 15470
(VPi). Pittsylvania Co.: dry woodland edges (rich), 22 Jun 1963, W. hathaway s.n. (lYn); sandy bottomland, county road #608 near
Pigg river, 25 Jun 1966, W. hathaway s.n. (lYn); below bridge over Staunton river on [uS] hwy 29 across from Altavista, 29 Jun 1966,
G. ramsey s.n. et al. (lYn); old jeep trail in woods on west side of uS 29 bypass on south side of Staunton river, 2 river mi. upstream
from Alta Vista, 30 Jul 1994, John B. nelson 16019 (gh, uSch). Surry Co.: rich alluvial woods and thickets back of sand-beach of James
river, below Sunken meadow Beach, 14 Jun 1938, m.L. Fernald and B. Long 8446 (gh); thicket back of sand-beach of cobham Bay, James
river, northwest of chippokes, 25 Aug 1940, m.L. Fernald and B. Long 12788 (gh); sandy river bank, southeast of claremont, 25 Jun
1950, B. mikula 5514 (fArm, willi). NORTH CAROLINA. Durham Co.: wooded pasture, northeast of durham on neuse river at
co. line (Sw of cozart), 21 Aug 1965, harry e. ahles 59795 (ncu). Granville Co.: disturbed right of way, former Seaboard coastline
rr bed 0.2 mi Sw of Sr 1726, 28 Jun 2006, Dale e. suiter 2505 et al. (ncu). Montgomery Co.: same location as type, 21 Jun 1978, J.
matthews et al. s.n. (uncc); 16 Jun 1998, h. selvig s.n. (uSch); 23 Jun 1988, J. nelson 6733 (uSch).
discussion
the earliest known collections of stachys matthewsii were made by fernald and long in 1938 and 1940,
during their exploration of unusual, shell-rich, calcareous bluffs and shores along the James river in the
southeastern Virginia coastal Plain (fernald 1938, 1941). fernald initially determined these collections
as “stachys nuttallii Shuttlew.,” but later changed his diagnosis to “stachys clingmanii Small,” (fernald 1943,
1950). confusion in the interpretation of other historical collections of s. matthewsii is also evident, and is
perhaps best epitomized by J.B. lewis’ 1942 collection from Brunswick county, Virginia. this specimen was
originally labeled s. tenuifolia willd., but was subsequently annotated between 1972 and 2005 as s. clingmanii
Small, s. nuttallii Shuttlew. ex Benth., s. eplingii J.B.nelson, and finally, s. hispida Pursh.
James matthews’ (uncc) discovery of this plant in montgomery county, north carolina arose from
an inventory of unusual habitats and rare species within the uwharrie mountains region. on June 21, 1978,
matthews collected two specimens of the hedge-nettle in question during a search for helenium brevifolium
(nutt.) wood, along the east side of the Yadkin river just below the falls dam. the plants were considered
unusual and tentatively identified as stachys riddellii house (= s. cordata riddell). nelson (then at fSu) was
alerted to the presence of this unusual hedge-nettle by matthews in 1979, and driven to the population for
some observations. he later made trips to the same spot, again accompanied by matthews, in 1988 and
1998. Between 1994 and 2008, new populations and/or herbarium specimens from four additional counties
in south-central Virginia were located by nelson, fleming, and townsend.
conflicting determinations of historical specimens of stachys matthewsii can be partly explained by the
incomplete and evolving treatments of stachys during the mid-20th century, and partly by the lack of any

Fleming et al., A new species of Stachys from the southeastern United States 11
available key or description that would account for the particular combination of characters that are diagnostic
of the taxon. in recent decades, interpretation of the genus has become more stable, with circumscriptions
of native southeastern species generally based on combinations of three character-groups: relative petiole
length, shape and relative length of the calyx lobes, and distribution and type of stem pubescence (nelson
1981; mulligan & munro 1989). stachys matthewsii is characterized by the combination of short petioles,
short triangular-apiculate calyx lobes, and copious pubescence restricted to the stem angles (figs. 2, 3, and
5). this taxon was not clearly recognized in any treatment until that by nelson (1981), who maintained
it within s. nuttallii (= s. cordata) as a short-petioled variant, disjunct from the rest of the range. current
treatments of the nutallii-cordata group, however, distinguish its members not only by their long-petioled
leaves but also by the presence of hairs on the stem sides and their musky-scented, atomiferous-glandular
foliage (nelson 2008), all characters absent in s. matthewsii. contemporary interpretation of s. clingmanii
as a longer-petioled, high-elevation plant of the southern Appalachians has rendered fernald’s view of the
Surry county hedge-nettles both taxonomically erroneous and phytogeographically improbable (nelson
1981; gleason & cronquist 1991).
the treatment by mulligan and munro (1989) would place the collections of this taxon within s. latidens
Small. Annotations of the holotype and one paratype of stachys matthewsii (nelson 6733), supplied by g.A.
mulligan in 1990, read “stachys latidens Small, very close to”; the holotype additionally bears mulligan’s
note “angles long erect hairs.” like s. matthewsii, s. latidens has short petioles and short deltoid calyx lobes.
however, it is a relatively smooth plant with only scattered, pustulate, short-deflexed hairs on the stem
angles. Additionally, the bearded nodes and abundant, prominently spreading, multicellular hairs of s.
matthewsii will consistently separate it from s. latidens. moreover, the two species have allopatric distributions
and disparate ecological affiliations, with s. latidens a plant of higher-elevation forests and clearings in
the Blue ridge and s. matthewsii a plant of low-elevation clearings and edges in the Piedmont and coastal
Plain. this geographic and ecological separation of s. matthewsii and s. latidens would seem to preclude any
biological interaction between the two taxa at present. other species, such as s. hispida and s. tenuifolia,
are occasionally found sympatrically with s. matthewsii, but no intermediates have been seen to suggest
any current gene exchange among members of the group. the type of multicellular pubescence seen in s.
matthewsii could possibly indicate some past kinship with s. hispida, although the distribution and density
of these pustular-based, multicellular hairs is distinctly different in the two taxa. Specifically, s. hispida is
more or less uniformly hispid with a moderate to somewhat dense concentration of hairs along the stem,
whereas those of s. matthewsii are only moderately dense above but rapidly become very densely concentrated
in the proximal 3–4 nodes and internodes. without a molecular framework for placing the various species
of stachys, such morphological and ecological evidence will remain the most practical tools for recognizing
new taxa in this complex genus.
the population at the type locality is near the margin of the Yadkin river, at the narrows just below
(and within sight of) the falls dam, east of the town of Badin. the plants grow in considerable shade on
sandy, loamy ground, with the population best developed on a somewhat sandy ridge. the site is probably
best considered an occasionally flooded-scoured river levee complex. Associated species include
Liriodendron tulipifera l., Pinus echinata mill., Liquidambar styraciflua l., Quercus nigra l., robinia pseudoacacia
l., albizia julibrissin durazz., sassafras albidum (nutt.) nees, alnus serrulata (Aiton) willd., amorpha fruticosa
l., Parthenocissus quinquefolia (l.) Planch., apios americana medik., Vitis rotundifolia michx., and asplenium
platyneuron (l.) Britton, Sterns, & Poggenb., all of which are locally common and hardly indicating special
or unusual habitats.
Subsequent study of populations and collections from Virginia demonstrate a wider distribution (fig.
4). most Virginia populations occur either along streams or in low, seasonally damp swales. habitats could
generally be considered mundane, except for those of the Surry county populations, which occupy calcareous
open swales and forest edges along shell-laden shores of the James river. repeated observations of populations
in halifax and charlotte counties indicate that s. matthewsii is quite shade-intolerant and not very

12 Journal of the Botanical Research Institute of Texas 5(1)
Fig. 1. Stachys matthewsii habit.
Fig. 2. Middle stem, leaves, and petioles of Stachys matthewsii. Fig. 3. Flowering and fruiting calyces of Stachys matthewsii.

Fleming et al., A new species of Stachys from the southeastern United States 13
Fig. 4. County distribution of Stachys matthewsii specimens in Virginia and North Carolina.
competitive with taller, ruderal perennials. Plants growing in a formerly cleared floodplain now invaded by
young trees have become progressively shade-stressed and do not flower. A periodically mowed powerline
swale with short to medium-height herbaceous vegetation appears to represent optimal habitat for the species,
supporting a population with hundreds of flowering plants. Perhaps the combination of shade-intolerance
and poor competitive abilities, as well as low levels of botanical inventory in the Piedmont, accounts for the
relatively low number of known populations, despite an abundance of seemingly suitable habitat in the region.
Notes on the Distribution of Stachys matthewsii
there is precedence for the somewhat disjunct distribution of populations of stachys matthewsii between the
north carolina - Virginia Piedmont and calcareous habitats of the Virginia coastal Plain. Previous investigations
by ware and ware (1992) and macdonald (2000) point to numerous species of primarily Piedmont
and mountain distribution with outlying populations in or adjacent to coastal ravine complexes underlain
by the calcareous Yorktown and eastover formations. these Pliocene deposits, due to their highly erodible
nature, have allowed for the creation of relatively dramatic ravine complexes within the otherwise slightly
undulating and acidic landscape of the tidewater region. it is widely assumed that the disjuncts present in
or near such ravines are relics of early holocene climates, when species now confined to montane habitats
were more broadly distributed in Virginia (delcourt & delcourt 1986). while stachys matthewsii is not confined
to these steep - sided erosional features, the presence of populations immediately adjacent to them
on shelly substrates (and nowhere else in the region) certainly points to relictual distribution as a possible
factor involved in their presence.

14 Journal of the Botanical Research Institute of Texas 5(1)
Fig. 5. Stachys matthewsii G.P. Fleming, J.B. Nelson, and J.F. Townsend. A. Habit, upper stem. B. Base of stem showing bearded nodes and densely hairy
internodes. C. Calyx, S. matthewsii. D. Calyx, S. hispida (for comparison). E. Seed, dorsal view. F. Seed, ventral view (from G.P. Fleming 13418, NCU).
Illustration by Lara Call Gastinger.
numerous examples of this mountain/Piedmont–coastal Plain disjunction type are described by ware
and ware (1992) and macdonald (2000), but the species scutellaria incana Biehler and stewartia ovata (cav.)
weath. provide the best analogs for the distribution of stachys matthewsii in Virginia. each is found almost
exclusively within the southern Piedmont of Virginia but have outlying populations within the coastal ravine
systems discussed here.
stachys matthewsii also belongs to a small group of species that have distributions centered in the southeastern
Piedmont, sometimes extending into the adjacent inner coastal Plain or to scattered disjunctions
in the mountains. five of these species are apparently endemic to Virginia and north carolina: Cardamine
micranthera rollins, eupatorium saltuense fernald, hexastylis lewisii (fernald) Blomq. & oosting, Isoetes vir-

Fleming et al., A new species of Stachys from the southeastern United States 15
ginica n. Pfeiff.and Phemeranthus piedmontana S. ware (see ware, this issue). Additional species that occupy
a somewhat larger range, occurring primarily from southern maryland or Virginia to South carolina or
northeastern georgia, include Clematis ochroleuca Aiton, heuchera caroliniana (rosend., Butters & lakela) e.f.
wells, hypericum lloydii (Svens.) w.P. Adams, acmispon helleri (Britton) A.A. heller, Phacelia covillei S. wats.
ex A. gray, rhus michauxii Sarg., solidago pinetorum Small, and symphyotrichum grandiflorum (l.) g.l. nesom.
Several granite flatrock endemics that also fit this pattern include Cyperus granitophilus mcVaugh, Diamorpha
smallii (Britton ex Small), Isoetes piedmontana (n. Pfeiff.) c.f. reed, and Portulaca smallii (P. wilson).
Etymology
in presenting this new taxon, we have selected the epithet “matthewsii” as a way of honoring James f. matthews,
who initiated this process with his early collections, and who alerted nelson to the potential issue. Jim’s
record as an outstanding botanist, teacher, and mentor is continued through his dedication to and support
for the uncc herbarium; we are honored to recognize him in this way.
keys to southeastern stachys
Building on the work of nelson (2008), two field keys are provided to aid identification of stachys matthewsii
and the integration of this taxon into contemporary treatments of southeastern stachys. the first key uses
relative petiole length as the primarily character, while the second uses shape and relative length of the
calyx lobes. the user may find that one key has greater utility with a given specimen, and that the use of
two approaches is helpful in identifying difficult specimens.
key 1
1. Annual; corolla scarcely exserted from calyx ____________________________________________________ S. agraria
1. Perennial: corolla prominently exserted from calyx; widespread.
2. Petioles obvious, those of the mid-stem leaves generally 1∕5 as long as the leaf blades.
3. Calyx tubes glandular.
4. Leaf blade margins dentate or nearly so; stem angles copiously pubescent with long (to 3 mm),
spreading hairs ________________________________________________________________ S. clingmanii
4. Leaf blade margins mostly crenate; stem angles glabrate or pubescent with mostly short, retrorse
hairs.
5. Leaf blades < 3 cm wide; plants weedy, scarcely 0.75 m tall, with white, rounded tubers terminating
rhizomes _________________________________________________________________ S. floridana
5. Leaf blades commonly > 3 cm wide; plants not weedy, commonly 1 m or more tall; rounded
tubers absent.
6. Leaf blades ovate, the bases cordate; margins crenulate _______________________________S. cordata
6. Leaf blades elliptic-oblong, the bases rounded to slightly cordate; margins crenate to serrate
__________________________________________________________________________ S. nuttallii
3. Calyx not glandular, or very slightly so.
7. Calyx glabrous to sparsely pubescent; petioles well-developed, especially in shade forms
______________________________________________________________________________ S. tenuifolia
7. Calyx variously hairy, but at least hispidulous, frequently strongly hispid; petioles short or long.
8. Petioles usually well developed; stem angles abundantly pubescent with spreading hairs; blade
margins dentate _____________________________________________________________ S. clingmanii
8. Petioles short to nearly absent; stem angles commonly pubescent, with retrorse hairs; blade
margins serrate _________________________________________________________________ S. hispida
2. Petioles short, those of the mid-stem leaves < 1∕5 as long as the blades or absent.
9. Leaves linear-lanceolate to narrowly lanceolate, usually widest at or near the base; leaf margins entire
to crenulate, rarely serrulate, and then mostly towards the apex.
10. Upper stem sides prominently pubescent; leaf blades abundantly pubescent below with appressed
hairs, thus closely tomentose, the lower surface felty or velvety; corolla purplish; rarely adventive
______________________________________________________________________________S. palustris
10. Upper stem sides glabrous; leaf blades variously pubescent or glabrate, but never felty nor velvety;
corolla pink; native species
11. Leaf blades narrow, 3–6 mm wide; plants generally glabrous to moderately pubescent; blade
margins entire to obscurely crenulate __________________________________________ S. hyssopifolia

16 Journal of the Botanical Research Institute of Texas 5(1)
11. Leaf blades broader, 5–8 mm wide; plants frequently hispidulous or abundantly pubescent;
blade margins crenulate to serrulate _______________________________________________ S. aspera
9. Leaves ovate to elliptic or oblong, widest near the center or toward the apex; leaf margins crenate to
sharply serrate for nearly the entire length.
12. Stem sides commonly pubescent above.
13. Calyx lobes lanceolate; flowers 6 per verticil; leaf margins serrate-crenate
14. Leaves generally subsessile, the petioles thick, not longer than 6 mm; blades densely
pubescent, frequently soft-pilose or velvety; leaf margins serrate; VA and WV only ______ S. arenicola
14. Leaves generally petioled up to 15 mm long, although shorter, and relatively slender; blades
pubescent on both surfaces, but never pilose or velvety; leaf margins crenate- serrulate;
KY-TN-GA-AL-SC __________________________________________________________ S. nuttallii
13. Calyx lobes deltoid; flowers 8 or more per verticil; leaf margins crenate-crenulate.
15. Lower leaf surface densely sessile- and stipitate-glandular; blades ovate, widest below the
middle; adjacent upper verticils separate in fruit __________________________________S. eplingii
15. Lower leaf surface glandular, but not densely so; blades elliptic-ovate, widest at the middle;
adjacent upper verticils commonly crowded in fruit _________________________________ S. iltisii
12. Stem sides glabrous, the angles pubescent.
16. Fruiting calyx lobes lanceolate or deltoid-acuminate, more than half as long as the calyx tube.
17. Calyx hispidulous to strongly hispid with long, stiff hairs; stem angles moderately to
densely pubescent; leaves subsessile, with very short petioles; bracts abruptly reduced
upward from lowest flowering node ____________________________________________ S. hispida
17. Calyx sparsely to moderately pubescent with short soft hairs; stem angles moderately
pubescent to nearly glabrous; leaves with petioles 1 to 3 cm long; bracts gradually reduced
upward from lowest flowering node ________________________________________ S. subcordata
16. Fruiting calyx lobes triangular-deltoid and abruptly apiculate, less than half as long as the calyx
tube.
18. Stem nodes not bearded (sometimes obscurely fine- hairy); stem angles with scattered,
pustulate, short deflexed hairs; plants of high-elevation montane habitats _____________ S. latidens
18. Stem nodes bearded; lower stem angles copiously pubescent with 3-celled spreading hairs;
plants of low-elevation Piedmont and Coastal Plain habitats _____________________ S. matthewsii
key 2
1. Fruiting calyx lobes deltoid to broadly triangular, less than half as long as the calyx tube.
2. Stem sides pubescent; foliage glandular, often musky-scented when crushed.
3. Petioles short to absent, blades mostly rounded to truncate-cordate.
4. Lower leaf surface densely glandular with atomiferous/short- stipitate golden glands; blades
generally ovate, broadest toward the base ______________________________________________S. eplingii
4. Lower leaf surface glandular, but not densely so; blades generally elliptic, somewhat lyrate toward
base, broadest near the middle _________________________________________________________ S. iltisii
3. Petioles well developed, blades cordate at base.
5. Top of blooming stem frequently flexuous, somewhat lax; blades ovate-rounded ________________S. cordata
5. Top of blooming stem stiffish, not lax; blades oblong-elliptic _______________________________ S. nuttallii
2. Stem sides glabrous; foliage not glandular (occasionally with a few scattered glands) or musky-scented.
6. Stem nodes not bearded (sometimes obscurely fine-hairy); stem angles with scattered, pustulate, short
deflexed hairs; plants of high-elevation montane habitats ___________________________________ S. latidens
6. Stem nodes bearded; lower stem angles copiously pubescent with 3-celled spreading hairs; plants of
low-elevation Piedmont and Coastal Plain habitats ______________________________________ S. matthewsii
1. Fruiting calxy lobes lanceolate, narrowly triangular, deltoid-acuminate, or nearly subulate, more than half as
long as the calyx tube.
7. Leaf blades linear to lanceolate.
8. Leaf blade margins entire to crenate; plants generally glabrous ____________________________ S. hyssopifolia
8. Leaf blade margins serrulate with at least a few teeth; plants glabrous or pubescent.
9. Stems strict or sparingly branched; leaves sessile or barely petioled, the blades crenate to serrate
with shallow teeth _________________________________________________________________ S. aspera
9. Stems frequently lax and branched from the upper nodes; leaves obviously petioled, the blades
sharply toothed _________________________________________________________________ S. tenuifolia
7. Leaf blades wider, rounded, oblong to elliptic.

Fleming et al., A new species of Stachys from the southeastern United States 17
10. Petioles poorly developed or absent, < 1 ∕5 as long as the blade.
11. Upper stem sides glabrous.
12. Calyx hispidulous to strongly hispid with long, stiff hairs; stem angles moderately to densely
pubescent; leaves subsessile, with very short petioles; bracts abruptly reduced upward from
lowest flowering node ________________________________________________________ S. hispida
12. Calyx sparsely to moderately pubescent with short soft hairs; stem angles moderately pubescent
to nearly glabrous; leaves with petioles 1 to 3 cm long; bracts gradually reduced upward
from lowest flowering node ________________________________________________ S. subcordata
11. Upper stem sides variously pubescent or glandular-pubescent.
13. Corollas purple; rare adventive _________________________________________________S. palustris
13. Corollas pink; native species.
14. Blades copiously pubescent, usually velvety; VA and WV only _____________________ S. arenicola
14. Blades pubescent but never velvety; KY-TN-GA-AL-SC ___________________________ S. nuttallii
10. Petioles obvious, those of the mid stem > 1 ∕5 the length of the blade.
15. Leaf blades dentate or sharply dentate; stem angles abundantly pubescent with long (to 3 mm),
spreading or somewhat retrorse hairs _____________________________________________ S. clingmanii
15. Leaf margins crenate to serrulate, but never dentate; stem angles glabrous or pubescent (if the
latter, then with scattered, stiffish, retrorse hairs).
16. Annual; corolla scarcely exserted from calyx ________________________________________ S. agraria
16. Perennial; corolla prominently exserted from calyx.
17. Plants commonly glandular on stems, leaves, and within inflorescence.
18. Rhizomes terminated by white, rounded tubers; blades oblong; weedy _________ S. floridana
18. Rhizomes without tubers; blades elliptic, cordate; not weedy __________________ S. nuttallii
17. Plants not glandular on stems, leaves, or within inflorescence.
19. Calyx usually hispidulous to hispid with stiff, eglandular hairs; fruiting calyx lobes
straight and conspicuously apiculate ______________________________________ S. hispida
19. Calyx usually glabrous or sparingly pubescent; fruiting calyx lobes usually recurved
and only weakly apiculate ____________________________________________ S. tenuifolia
acknowledgments
we acknowledge the kind cooperation of the herbarium curators who have made loans of their material
available to us. we especially thank randee humpreys and personnel of lewis ginter Botanical garden for
allowing long-term access to the Virginia commonwealth university herbarium, greg Plunkett, professor at
Vcu, for helping arrange and process loans and for enabling a critical study of specimens in this workspace,
and artist lara call gastinger for preparation of the botanical illustration. the South carolina nongame &
heritage trust Program provided travel assistance for field work (nelson 1988) at the site of the type collection.
Substantial support for field work by nelson was provided by a grant from the Barbara J. harvill fund
for floristic research in Virginia. holly Selvig provided preliminary morphological studies as an independent
study through a howard hughes grant at the university of South carolina, and assisted with field observations.
grant godden and an anonymous reviewer provided helpful reviews. ursula engelbrecht (uSc) and
Boris Schlumperger (herrenhausen gardens, hanover, germany) considerably improved the german abstract.
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18 Journal of the Botanical Research Institute of Texas 5(1)
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