(Stachys: Lamiaceae) from the - Botanical Research Institute of Texas
(Stachys: Lamiaceae) from the - Botanical Research Institute of Texas
(Stachys: Lamiaceae) from the - Botanical Research Institute of Texas
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A new hedge-nettle (staChYs: lAmiAceAe) <strong>from</strong> <strong>the</strong><br />
mid-AtlAntic Piedmont And coAStAl PlAin <strong>of</strong> <strong>the</strong> united StAteS<br />
Gary P. Fleming John B. Nelson John F. Townsend<br />
Virginia Dept. <strong>of</strong> Conservation and A.C. Moore Herbarium Virginia Dept. <strong>of</strong> Conservation and<br />
Recreation, Division <strong>of</strong> Nat. Heritage Dept. Biological Sciences Recreation, Division <strong>of</strong> Nat. Heritage<br />
217 Governor St., 3rd Floor University <strong>of</strong> South Carolina 217 Governor St., 3rd Floor<br />
Richmond, Virginia 23219, U.S.A. Columbia, South Carolina 29208 U.S.A. Richmond, Virginia 23219, U.S.A.<br />
gary.fleming@dcr.virginia.gov nelson@sc.edu john.townsend@dcr.virginia.gov<br />
abstract<br />
A new, geographically restricted “hedge-nettle,” <strong>Stachys</strong> mat<strong>the</strong>wsii (lamiaceae) is described <strong>from</strong> <strong>the</strong> region that includes sou<strong>the</strong>astern<br />
and south-central Virginia and north-central north carolina. this taxon resembles stachys latidens Small ex Britton, a species <strong>of</strong><br />
higher-elevation forests <strong>of</strong> <strong>the</strong> Blue ridge mountains, but has consistent morphological differences in stem pubescence and has distinctly<br />
different habitat requirements. two keys are provided for distinguishing hedge-nettles <strong>of</strong> <strong>the</strong> sou<strong>the</strong>astern united States.<br />
zusammenFassung<br />
Zur Beschreibung des neuen Ziests, <strong>Stachys</strong> mat<strong>the</strong>wsii (lamiaceae), lässt sich sagen, dass seine geographische Verbreitung auf den<br />
südöstlichen und den mittleren Süden Virginias und den mittleren norden north carolinas begrenzt ist. die neue Art ähnelt stachys<br />
latidens Small in Britton, einer Pflanze aus den höher gelegenen “Blue ridge mountains,” aber sie hat konsistente morphologische<br />
unterschiede in der Stammpubeszenz und hat völlig andere habitat-Ansprüche. Zwei Schlüssel dienen zur Bestimmung der Ziest-Arten<br />
aus den südöstlichen uSA.<br />
resumen<br />
Se describe una nueva “ortiga muerta,” <strong>Stachys</strong> mat<strong>the</strong>wsii (lamiaceae) restringida geográficamente a la región que incluye el sureste<br />
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<br />
bosques a mayor elevación de las montañas del Blue ridge, pero tiene diferencias morfológicas consistentes en la pubescencia del tallo<br />
y distintos requerimientos de hábitat. Se aportan dos claves para diferenciar las xxx del sureste de los estados unidos.<br />
introduction<br />
field work and herbarium studies in north carolina and Virginia has led to <strong>the</strong> circumscription <strong>of</strong> a new<br />
species in <strong>the</strong> genus stachys, a large group <strong>of</strong> mints widely recognized for its taxonomic complexity. changing<br />
taxonomic interpretations in <strong>the</strong> past have confounded efforts to classify this plant properly, but enough has<br />
now been learned to recognize this species’ unique place within <strong>the</strong> genus. A history <strong>of</strong> taxonomic opinions<br />
regarding stachys mat<strong>the</strong>wsii is presented, as is a discussion <strong>of</strong> habitat and geography, particularly its primary<br />
range in <strong>the</strong> Piedmont and its somewhat disjunct coastal Plain populations in Virginia.<br />
<strong>Stachys</strong> mat<strong>the</strong>wsii g.P. fleming, J.B. nelson, & J.f. townsend, sp. nov. (Figs. 1–5). type: united StAteS: north<br />
carolina. montgomery co.: fairly shady woodland on e side <strong>of</strong> Yadkin river, just downstream <strong>from</strong> falls dam, uwharrie national<br />
forest, 11 Jun 1979, John B. nelson 1152 with J. mat<strong>the</strong>ws (holotype: uSch; isotypes: fSu, mo, ncu, uncc[2]).<br />
differt a stachys latidens ob trichomata longa expansaque ad caulis angulas. differt a stachys hispida ob calycis lobos breviores et minus deltatos<br />
pro parte. differt a stachys cordata ob folia parviora, minus cordata, et petiolos breviores. differt a stachys nuttallii ob caulis facies leves.<br />
Perennial herbs <strong>from</strong> vigorous, pale, fragrant rhizomes; flowering stems erect, 0.8–1.05m tall, sparingly<br />
branched, even within inflorescence, or branched toward base on older plants; <strong>the</strong> sides glabrous as high as<br />
<strong>the</strong> lowest verticil, fertile internodes with scattered short glandular hairs, <strong>the</strong> angles prominently and consistently<br />
pubescent with stiffish, spreading (or retrorse) hairs, <strong>the</strong>se eglandular, to 3mm long; nodes bearded<br />
with hairs as those on <strong>the</strong> stem angles; <strong>the</strong> overall pubescence is dense at <strong>the</strong> base <strong>of</strong> <strong>the</strong> plant, becoming<br />
progressively less so upwards; leaves consistently petioled, <strong>the</strong> petioles 2–5mm long, blades mostly rounded<br />
at base or somewhat cordate, <strong>the</strong> blades 5–10 cm long, 2–5cm broad, broadly elliptical to lance-ovate, acute<br />
J. Bot. Res. Inst. <strong>Texas</strong> 5(1): 9 – 18. 2011
10 Journal <strong>of</strong> <strong>the</strong> <strong>Botanical</strong> <strong>Research</strong> <strong>Institute</strong> <strong>of</strong> <strong>Texas</strong> 5(1)<br />
at apex, <strong>the</strong> margins crenate (crenulate-serrulate), leaves spreading-ascending, upper surface prominently<br />
bullate, abundantly pubescent with appressed and erect non-glandular, pustulate-based hairs, <strong>the</strong> lower<br />
surface glabrate to s<strong>of</strong>tly pubescent with erect, eglandular hairs; inflorescences <strong>of</strong> interrupted verticils;<br />
bracts abruptly reduced upward <strong>from</strong> lowest fertile nodes; cymules each with 3 flowers, <strong>the</strong> verticils thus<br />
6-flowered; fruiting calyces campanulate, <strong>the</strong> tube densely short-s<strong>of</strong>t pubescent, and with longer, stiffish<br />
hairs, especially along <strong>the</strong> veins, and slightly atomiferous-glandular, tubes mostly 4mm (3.5–4.5mm) long;<br />
<strong>the</strong> lobes deltoid, narrowed apically and terminated with minute apiculum, <strong>the</strong> lobes less than one-half <strong>the</strong><br />
length <strong>of</strong> <strong>the</strong> tube, 1.5–1.8(–2.4)mm long; corollas pink, corolla tube prominently saccate toward base on<br />
lower side, tube internally glabrous, but with prominently slanting (oblique) annulus, copiously pubescent<br />
with s<strong>of</strong>t, bulbous trichomes; tube 11–14mm <strong>from</strong> base to apex <strong>of</strong> galea; galeae abundantly pubescent with<br />
stalked glands and s<strong>of</strong>t eglandular hairs; lower lip with prominent dark pink (nearly purple) blotches on<br />
<strong>the</strong> median portion <strong>of</strong> <strong>the</strong> lower lip; declined to 90° <strong>from</strong> <strong>the</strong> tube, 6.5–8mm long, <strong>the</strong> middle lobe <strong>of</strong> lower<br />
lip narrowly ovate, slightly notched at apex; mericarps dark brown, minutely pebbled, 1.4–1.5mm wide,<br />
1.8–2mm long.<br />
Additional collections examined: UNITED STATES. VIRGINIA. Amelia Co.: local in wet meadows and old fields, 5 Jul 1938, J.B.<br />
Lewis 1592 (fSu). Brunswick Co.: along wooded stream, Jun 1942, J.B. Lewis 3588 (VPi). Charlotte Co.: young floodplain forest along<br />
cargills creek, 3.5 mi e <strong>of</strong> dryburg, 14 Aug 2008, Gary P. Fleming 15453 with John F. townsend (VPi). Halifax Co.: moist swale in powerline<br />
clearing 2.2 mi Sw <strong>of</strong> Public fork, 14 Jul 1995, , Gary P. Fleming 10804 (gmuf, uSch, VPi, willi); same location, 1 Jul 1997,<br />
Gary P. Fleming 13418 (unc, uS); same location, 14 Aug 2008, John F. townsend 4061 with Gary P. Fleming (VPi); thinly wooded toe slope<br />
along Staunton river, 0.6 mi. south <strong>of</strong> uS 360 bridge and 2.3 mi. west-southwest <strong>of</strong> Public fork, 26 may 2010, Gary P. Fleming 15470<br />
(VPi). Pittsylvania Co.: dry woodland edges (rich), 22 Jun 1963, W. hathaway s.n. (lYn); sandy bottomland, county road #608 near<br />
Pigg river, 25 Jun 1966, W. hathaway s.n. (lYn); below bridge over Staunton river on [uS] hwy 29 across <strong>from</strong> Altavista, 29 Jun 1966,<br />
G. ramsey s.n. et al. (lYn); old jeep trail in woods on west side <strong>of</strong> uS 29 bypass on south side <strong>of</strong> Staunton river, 2 river mi. upstream<br />
<strong>from</strong> Alta Vista, 30 Jul 1994, John B. nelson 16019 (gh, uSch). Surry Co.: rich alluvial woods and thickets back <strong>of</strong> sand-beach <strong>of</strong> James<br />
river, below Sunken meadow Beach, 14 Jun 1938, m.L. Fernald and B. Long 8446 (gh); thicket back <strong>of</strong> sand-beach <strong>of</strong> cobham Bay, James<br />
river, northwest <strong>of</strong> chippokes, 25 Aug 1940, m.L. Fernald and B. Long 12788 (gh); sandy river bank, sou<strong>the</strong>ast <strong>of</strong> claremont, 25 Jun<br />
1950, B. mikula 5514 (fArm, willi). NORTH CAROLINA. Durham Co.: wooded pasture, nor<strong>the</strong>ast <strong>of</strong> durham on neuse river at<br />
co. line (Sw <strong>of</strong> cozart), 21 Aug 1965, harry e. ahles 59795 (ncu). Granville Co.: disturbed right <strong>of</strong> way, former Seaboard coastline<br />
rr bed 0.2 mi Sw <strong>of</strong> Sr 1726, 28 Jun 2006, Dale e. suiter 2505 et al. (ncu). Montgomery Co.: same location as type, 21 Jun 1978, J.<br />
mat<strong>the</strong>ws et al. s.n. (uncc); 16 Jun 1998, h. selvig s.n. (uSch); 23 Jun 1988, J. nelson 6733 (uSch).<br />
discussion<br />
<strong>the</strong> earliest known collections <strong>of</strong> stachys mat<strong>the</strong>wsii were made by fernald and long in 1938 and 1940,<br />
during <strong>the</strong>ir exploration <strong>of</strong> unusual, shell-rich, calcareous bluffs and shores along <strong>the</strong> James river in <strong>the</strong><br />
sou<strong>the</strong>astern Virginia coastal Plain (fernald 1938, 1941). fernald initially determined <strong>the</strong>se collections<br />
as “stachys nuttallii Shuttlew.,” but later changed his diagnosis to “stachys clingmanii Small,” (fernald 1943,<br />
1950). confusion in <strong>the</strong> interpretation <strong>of</strong> o<strong>the</strong>r historical collections <strong>of</strong> s. mat<strong>the</strong>wsii is also evident, and is<br />
perhaps best epitomized by J.B. lewis’ 1942 collection <strong>from</strong> Brunswick county, Virginia. this specimen was<br />
originally labeled s. tenuifolia willd., but was subsequently annotated between 1972 and 2005 as s. clingmanii<br />
Small, s. nuttallii Shuttlew. ex Benth., s. eplingii J.B.nelson, and finally, s. hispida Pursh.<br />
James mat<strong>the</strong>ws’ (uncc) discovery <strong>of</strong> this plant in montgomery county, north carolina arose <strong>from</strong><br />
an inventory <strong>of</strong> unusual habitats and rare species within <strong>the</strong> uwharrie mountains region. on June 21, 1978,<br />
mat<strong>the</strong>ws collected two specimens <strong>of</strong> <strong>the</strong> hedge-nettle in question during a search for helenium brevifolium<br />
(nutt.) wood, along <strong>the</strong> east side <strong>of</strong> <strong>the</strong> Yadkin river just below <strong>the</strong> falls dam. <strong>the</strong> plants were considered<br />
unusual and tentatively identified as stachys riddellii house (= s. cordata riddell). nelson (<strong>the</strong>n at fSu) was<br />
alerted to <strong>the</strong> presence <strong>of</strong> this unusual hedge-nettle by mat<strong>the</strong>ws in 1979, and driven to <strong>the</strong> population for<br />
some observations. he later made trips to <strong>the</strong> same spot, again accompanied by mat<strong>the</strong>ws, in 1988 and<br />
1998. Between 1994 and 2008, new populations and/or herbarium specimens <strong>from</strong> four additional counties<br />
in south-central Virginia were located by nelson, fleming, and townsend.<br />
conflicting determinations <strong>of</strong> historical specimens <strong>of</strong> stachys mat<strong>the</strong>wsii can be partly explained by <strong>the</strong><br />
incomplete and evolving treatments <strong>of</strong> stachys during <strong>the</strong> mid-20th century, and partly by <strong>the</strong> lack <strong>of</strong> any
Fleming et al., A new species <strong>of</strong> <strong>Stachys</strong> <strong>from</strong> <strong>the</strong> sou<strong>the</strong>astern United States 11<br />
available key or description that would account for <strong>the</strong> particular combination <strong>of</strong> characters that are diagnostic<br />
<strong>of</strong> <strong>the</strong> taxon. in recent decades, interpretation <strong>of</strong> <strong>the</strong> genus has become more stable, with circumscriptions<br />
<strong>of</strong> native sou<strong>the</strong>astern species generally based on combinations <strong>of</strong> three character-groups: relative petiole<br />
length, shape and relative length <strong>of</strong> <strong>the</strong> calyx lobes, and distribution and type <strong>of</strong> stem pubescence (nelson<br />
1981; mulligan & munro 1989). stachys mat<strong>the</strong>wsii is characterized by <strong>the</strong> combination <strong>of</strong> short petioles,<br />
short triangular-apiculate calyx lobes, and copious pubescence restricted to <strong>the</strong> stem angles (figs. 2, 3, and<br />
5). this taxon was not clearly recognized in any treatment until that by nelson (1981), who maintained<br />
it within s. nuttallii (= s. cordata) as a short-petioled variant, disjunct <strong>from</strong> <strong>the</strong> rest <strong>of</strong> <strong>the</strong> range. current<br />
treatments <strong>of</strong> <strong>the</strong> nutallii-cordata group, however, distinguish its members not only by <strong>the</strong>ir long-petioled<br />
leaves but also by <strong>the</strong> presence <strong>of</strong> hairs on <strong>the</strong> stem sides and <strong>the</strong>ir musky-scented, atomiferous-glandular<br />
foliage (nelson 2008), all characters absent in s. mat<strong>the</strong>wsii. contemporary interpretation <strong>of</strong> s. clingmanii<br />
as a longer-petioled, high-elevation plant <strong>of</strong> <strong>the</strong> sou<strong>the</strong>rn Appalachians has rendered fernald’s view <strong>of</strong> <strong>the</strong><br />
Surry county hedge-nettles both taxonomically erroneous and phytogeographically improbable (nelson<br />
1981; gleason & cronquist 1991).<br />
<strong>the</strong> treatment by mulligan and munro (1989) would place <strong>the</strong> collections <strong>of</strong> this taxon within s. latidens<br />
Small. Annotations <strong>of</strong> <strong>the</strong> holotype and one paratype <strong>of</strong> stachys mat<strong>the</strong>wsii (nelson 6733), supplied by g.A.<br />
mulligan in 1990, read “stachys latidens Small, very close to”; <strong>the</strong> holotype additionally bears mulligan’s<br />
note “angles long erect hairs.” like s. mat<strong>the</strong>wsii, s. latidens has short petioles and short deltoid calyx lobes.<br />
however, it is a relatively smooth plant with only scattered, pustulate, short-deflexed hairs on <strong>the</strong> stem<br />
angles. Additionally, <strong>the</strong> bearded nodes and abundant, prominently spreading, multicellular hairs <strong>of</strong> s.<br />
mat<strong>the</strong>wsii will consistently separate it <strong>from</strong> s. latidens. moreover, <strong>the</strong> two species have allopatric distributions<br />
and disparate ecological affiliations, with s. latidens a plant <strong>of</strong> higher-elevation forests and clearings in<br />
<strong>the</strong> Blue ridge and s. mat<strong>the</strong>wsii a plant <strong>of</strong> low-elevation clearings and edges in <strong>the</strong> Piedmont and coastal<br />
Plain. this geographic and ecological separation <strong>of</strong> s. mat<strong>the</strong>wsii and s. latidens would seem to preclude any<br />
biological interaction between <strong>the</strong> two taxa at present. o<strong>the</strong>r species, such as s. hispida and s. tenuifolia,<br />
are occasionally found sympatrically with s. mat<strong>the</strong>wsii, but no intermediates have been seen to suggest<br />
any current gene exchange among members <strong>of</strong> <strong>the</strong> group. <strong>the</strong> type <strong>of</strong> multicellular pubescence seen in s.<br />
mat<strong>the</strong>wsii could possibly indicate some past kinship with s. hispida, although <strong>the</strong> distribution and density<br />
<strong>of</strong> <strong>the</strong>se pustular-based, multicellular hairs is distinctly different in <strong>the</strong> two taxa. Specifically, s. hispida is<br />
more or less uniformly hispid with a moderate to somewhat dense concentration <strong>of</strong> hairs along <strong>the</strong> stem,<br />
whereas those <strong>of</strong> s. mat<strong>the</strong>wsii are only moderately dense above but rapidly become very densely concentrated<br />
in <strong>the</strong> proximal 3–4 nodes and internodes. without a molecular framework for placing <strong>the</strong> various species<br />
<strong>of</strong> stachys, such morphological and ecological evidence will remain <strong>the</strong> most practical tools for recognizing<br />
new taxa in this complex genus.<br />
<strong>the</strong> population at <strong>the</strong> type locality is near <strong>the</strong> margin <strong>of</strong> <strong>the</strong> Yadkin river, at <strong>the</strong> narrows just below<br />
(and within sight <strong>of</strong>) <strong>the</strong> falls dam, east <strong>of</strong> <strong>the</strong> town <strong>of</strong> Badin. <strong>the</strong> plants grow in considerable shade on<br />
sandy, loamy ground, with <strong>the</strong> population best developed on a somewhat sandy ridge. <strong>the</strong> site is probably<br />
best considered an occasionally flooded-scoured river levee complex. Associated species include<br />
Liriodendron tulipifera l., Pinus echinata mill., Liquidambar styraciflua l., Quercus nigra l., robinia pseudoacacia<br />
l., albizia julibrissin durazz., sassafras albidum (nutt.) nees, alnus serrulata (Aiton) willd., amorpha fruticosa<br />
l., Par<strong>the</strong>nocissus quinquefolia (l.) Planch., apios americana medik., Vitis rotundifolia michx., and asplenium<br />
platyneuron (l.) Britton, Sterns, & Poggenb., all <strong>of</strong> which are locally common and hardly indicating special<br />
or unusual habitats.<br />
Subsequent study <strong>of</strong> populations and collections <strong>from</strong> Virginia demonstrate a wider distribution (fig.<br />
4). most Virginia populations occur ei<strong>the</strong>r along streams or in low, seasonally damp swales. habitats could<br />
generally be considered mundane, except for those <strong>of</strong> <strong>the</strong> Surry county populations, which occupy calcareous<br />
open swales and forest edges along shell-laden shores <strong>of</strong> <strong>the</strong> James river. repeated observations <strong>of</strong> populations<br />
in halifax and charlotte counties indicate that s. mat<strong>the</strong>wsii is quite shade-intolerant and not very
12 Journal <strong>of</strong> <strong>the</strong> <strong>Botanical</strong> <strong>Research</strong> <strong>Institute</strong> <strong>of</strong> <strong>Texas</strong> 5(1)<br />
Fig. 1. <strong>Stachys</strong> mat<strong>the</strong>wsii habit.<br />
Fig. 2. Middle stem, leaves, and petioles <strong>of</strong> <strong>Stachys</strong> mat<strong>the</strong>wsii. Fig. 3. Flowering and fruiting calyces <strong>of</strong> <strong>Stachys</strong> mat<strong>the</strong>wsii.
Fleming et al., A new species <strong>of</strong> <strong>Stachys</strong> <strong>from</strong> <strong>the</strong> sou<strong>the</strong>astern United States 13<br />
Fig. 4. County distribution <strong>of</strong> <strong>Stachys</strong> mat<strong>the</strong>wsii specimens in Virginia and North Carolina.<br />
competitive with taller, ruderal perennials. Plants growing in a formerly cleared floodplain now invaded by<br />
young trees have become progressively shade-stressed and do not flower. A periodically mowed powerline<br />
swale with short to medium-height herbaceous vegetation appears to represent optimal habitat for <strong>the</strong> species,<br />
supporting a population with hundreds <strong>of</strong> flowering plants. Perhaps <strong>the</strong> combination <strong>of</strong> shade-intolerance<br />
and poor competitive abilities, as well as low levels <strong>of</strong> botanical inventory in <strong>the</strong> Piedmont, accounts for <strong>the</strong><br />
relatively low number <strong>of</strong> known populations, despite an abundance <strong>of</strong> seemingly suitable habitat in <strong>the</strong> region.<br />
Notes on <strong>the</strong> Distribution <strong>of</strong> <strong>Stachys</strong> mat<strong>the</strong>wsii<br />
<strong>the</strong>re is precedence for <strong>the</strong> somewhat disjunct distribution <strong>of</strong> populations <strong>of</strong> stachys mat<strong>the</strong>wsii between <strong>the</strong><br />
north carolina - Virginia Piedmont and calcareous habitats <strong>of</strong> <strong>the</strong> Virginia coastal Plain. Previous investigations<br />
by ware and ware (1992) and macdonald (2000) point to numerous species <strong>of</strong> primarily Piedmont<br />
and mountain distribution with outlying populations in or adjacent to coastal ravine complexes underlain<br />
by <strong>the</strong> calcareous Yorktown and eastover formations. <strong>the</strong>se Pliocene deposits, due to <strong>the</strong>ir highly erodible<br />
nature, have allowed for <strong>the</strong> creation <strong>of</strong> relatively dramatic ravine complexes within <strong>the</strong> o<strong>the</strong>rwise slightly<br />
undulating and acidic landscape <strong>of</strong> <strong>the</strong> tidewater region. it is widely assumed that <strong>the</strong> disjuncts present in<br />
or near such ravines are relics <strong>of</strong> early holocene climates, when species now confined to montane habitats<br />
were more broadly distributed in Virginia (delcourt & delcourt 1986). while stachys mat<strong>the</strong>wsii is not confined<br />
to <strong>the</strong>se steep - sided erosional features, <strong>the</strong> presence <strong>of</strong> populations immediately adjacent to <strong>the</strong>m<br />
on shelly substrates (and nowhere else in <strong>the</strong> region) certainly points to relictual distribution as a possible<br />
factor involved in <strong>the</strong>ir presence.
14 Journal <strong>of</strong> <strong>the</strong> <strong>Botanical</strong> <strong>Research</strong> <strong>Institute</strong> <strong>of</strong> <strong>Texas</strong> 5(1)<br />
Fig. 5. <strong>Stachys</strong> mat<strong>the</strong>wsii G.P. Fleming, J.B. Nelson, and J.F. Townsend. A. Habit, upper stem. B. Base <strong>of</strong> stem showing bearded nodes and densely hairy<br />
internodes. C. Calyx, S. mat<strong>the</strong>wsii. D. Calyx, S. hispida (for comparison). E. Seed, dorsal view. F. Seed, ventral view (<strong>from</strong> G.P. Fleming 13418, NCU).<br />
Illustration by Lara Call Gastinger.<br />
numerous examples <strong>of</strong> this mountain/Piedmont–coastal Plain disjunction type are described by ware<br />
and ware (1992) and macdonald (2000), but <strong>the</strong> species scutellaria incana Biehler and stewartia ovata (cav.)<br />
weath. provide <strong>the</strong> best analogs for <strong>the</strong> distribution <strong>of</strong> stachys mat<strong>the</strong>wsii in Virginia. each is found almost<br />
exclusively within <strong>the</strong> sou<strong>the</strong>rn Piedmont <strong>of</strong> Virginia but have outlying populations within <strong>the</strong> coastal ravine<br />
systems discussed here.<br />
stachys mat<strong>the</strong>wsii also belongs to a small group <strong>of</strong> species that have distributions centered in <strong>the</strong> sou<strong>the</strong>astern<br />
Piedmont, sometimes extending into <strong>the</strong> adjacent inner coastal Plain or to scattered disjunctions<br />
in <strong>the</strong> mountains. five <strong>of</strong> <strong>the</strong>se species are apparently endemic to Virginia and north carolina: Cardamine<br />
micran<strong>the</strong>ra rollins, eupatorium saltuense fernald, hexastylis lewisii (fernald) Blomq. & oosting, Isoetes vir-
Fleming et al., A new species <strong>of</strong> <strong>Stachys</strong> <strong>from</strong> <strong>the</strong> sou<strong>the</strong>astern United States 15<br />
ginica n. Pfeiff.and Phemeranthus piedmontana S. ware (see ware, this issue). Additional species that occupy<br />
a somewhat larger range, occurring primarily <strong>from</strong> sou<strong>the</strong>rn maryland or Virginia to South carolina or<br />
nor<strong>the</strong>astern georgia, include Clematis ochroleuca Aiton, heuchera caroliniana (rosend., Butters & lakela) e.f.<br />
wells, hypericum lloydii (Svens.) w.P. Adams, acmispon helleri (Britton) A.A. heller, Phacelia covillei S. wats.<br />
ex A. gray, rhus michauxii Sarg., solidago pinetorum Small, and symphyotrichum grandiflorum (l.) g.l. nesom.<br />
Several granite flatrock endemics that also fit this pattern include Cyperus granitophilus mcVaugh, Diamorpha<br />
smallii (Britton ex Small), Isoetes piedmontana (n. Pfeiff.) c.f. reed, and Portulaca smallii (P. wilson).<br />
Etymology<br />
in presenting this new taxon, we have selected <strong>the</strong> epi<strong>the</strong>t “mat<strong>the</strong>wsii” as a way <strong>of</strong> honoring James f. mat<strong>the</strong>ws,<br />
who initiated this process with his early collections, and who alerted nelson to <strong>the</strong> potential issue. Jim’s<br />
record as an outstanding botanist, teacher, and mentor is continued through his dedication to and support<br />
for <strong>the</strong> uncc herbarium; we are honored to recognize him in this way.<br />
keys to sou<strong>the</strong>astern stachys<br />
Building on <strong>the</strong> work <strong>of</strong> nelson (2008), two field keys are provided to aid identification <strong>of</strong> stachys mat<strong>the</strong>wsii<br />
and <strong>the</strong> integration <strong>of</strong> this taxon into contemporary treatments <strong>of</strong> sou<strong>the</strong>astern stachys. <strong>the</strong> first key uses<br />
relative petiole length as <strong>the</strong> primarily character, while <strong>the</strong> second uses shape and relative length <strong>of</strong> <strong>the</strong><br />
calyx lobes. <strong>the</strong> user may find that one key has greater utility with a given specimen, and that <strong>the</strong> use <strong>of</strong><br />
two approaches is helpful in identifying difficult specimens.<br />
key 1<br />
1. Annual; corolla scarcely exserted <strong>from</strong> calyx ____________________________________________________ S. agraria<br />
1. Perennial: corolla prominently exserted <strong>from</strong> calyx; widespread.<br />
2. Petioles obvious, those <strong>of</strong> <strong>the</strong> mid-stem leaves generally 1∕5 as long as <strong>the</strong> leaf blades.<br />
3. Calyx tubes glandular.<br />
4. Leaf blade margins dentate or nearly so; stem angles copiously pubescent with long (to 3 mm),<br />
spreading hairs ________________________________________________________________ S. clingmanii<br />
4. Leaf blade margins mostly crenate; stem angles glabrate or pubescent with mostly short, retrorse<br />
hairs.<br />
5. Leaf blades < 3 cm wide; plants weedy, scarcely 0.75 m tall, with white, rounded tubers terminating<br />
rhizomes _________________________________________________________________ S. floridana<br />
5. Leaf blades commonly > 3 cm wide; plants not weedy, commonly 1 m or more tall; rounded<br />
tubers absent.<br />
6. Leaf blades ovate, <strong>the</strong> bases cordate; margins crenulate _______________________________S. cordata<br />
6. Leaf blades elliptic-oblong, <strong>the</strong> bases rounded to slightly cordate; margins crenate to serrate<br />
__________________________________________________________________________ S. nuttallii<br />
3. Calyx not glandular, or very slightly so.<br />
7. Calyx glabrous to sparsely pubescent; petioles well-developed, especially in shade forms<br />
______________________________________________________________________________ S. tenuifolia<br />
7. Calyx variously hairy, but at least hispidulous, frequently strongly hispid; petioles short or long.<br />
8. Petioles usually well developed; stem angles abundantly pubescent with spreading hairs; blade<br />
margins dentate _____________________________________________________________ S. clingmanii<br />
8. Petioles short to nearly absent; stem angles commonly pubescent, with retrorse hairs; blade<br />
margins serrate _________________________________________________________________ S. hispida<br />
2. Petioles short, those <strong>of</strong> <strong>the</strong> mid-stem leaves < 1∕5 as long as <strong>the</strong> blades or absent.<br />
9. Leaves linear-lanceolate to narrowly lanceolate, usually widest at or near <strong>the</strong> base; leaf margins entire<br />
to crenulate, rarely serrulate, and <strong>the</strong>n mostly towards <strong>the</strong> apex.<br />
10. Upper stem sides prominently pubescent; leaf blades abundantly pubescent below with appressed<br />
hairs, thus closely tomentose, <strong>the</strong> lower surface felty or velvety; corolla purplish; rarely adventive<br />
______________________________________________________________________________S. palustris<br />
10. Upper stem sides glabrous; leaf blades variously pubescent or glabrate, but never felty nor velvety;<br />
corolla pink; native species<br />
11. Leaf blades narrow, 3–6 mm wide; plants generally glabrous to moderately pubescent; blade<br />
margins entire to obscurely crenulate __________________________________________ S. hyssopifolia
16 Journal <strong>of</strong> <strong>the</strong> <strong>Botanical</strong> <strong>Research</strong> <strong>Institute</strong> <strong>of</strong> <strong>Texas</strong> 5(1)<br />
11. Leaf blades broader, 5–8 mm wide; plants frequently hispidulous or abundantly pubescent;<br />
blade margins crenulate to serrulate _______________________________________________ S. aspera<br />
9. Leaves ovate to elliptic or oblong, widest near <strong>the</strong> center or toward <strong>the</strong> apex; leaf margins crenate to<br />
sharply serrate for nearly <strong>the</strong> entire length.<br />
12. Stem sides commonly pubescent above.<br />
13. Calyx lobes lanceolate; flowers 6 per verticil; leaf margins serrate-crenate<br />
14. Leaves generally subsessile, <strong>the</strong> petioles thick, not longer than 6 mm; blades densely<br />
pubescent, frequently s<strong>of</strong>t-pilose or velvety; leaf margins serrate; VA and WV only ______ S. arenicola<br />
14. Leaves generally petioled up to 15 mm long, although shorter, and relatively slender; blades<br />
pubescent on both surfaces, but never pilose or velvety; leaf margins crenate- serrulate;<br />
KY-TN-GA-AL-SC __________________________________________________________ S. nuttallii<br />
13. Calyx lobes deltoid; flowers 8 or more per verticil; leaf margins crenate-crenulate.<br />
15. Lower leaf surface densely sessile- and stipitate-glandular; blades ovate, widest below <strong>the</strong><br />
middle; adjacent upper verticils separate in fruit __________________________________S. eplingii<br />
15. Lower leaf surface glandular, but not densely so; blades elliptic-ovate, widest at <strong>the</strong> middle;<br />
adjacent upper verticils commonly crowded in fruit _________________________________ S. iltisii<br />
12. Stem sides glabrous, <strong>the</strong> angles pubescent.<br />
16. Fruiting calyx lobes lanceolate or deltoid-acuminate, more than half as long as <strong>the</strong> calyx tube.<br />
17. Calyx hispidulous to strongly hispid with long, stiff hairs; stem angles moderately to<br />
densely pubescent; leaves subsessile, with very short petioles; bracts abruptly reduced<br />
upward <strong>from</strong> lowest flowering node ____________________________________________ S. hispida<br />
17. Calyx sparsely to moderately pubescent with short s<strong>of</strong>t hairs; stem angles moderately<br />
pubescent to nearly glabrous; leaves with petioles 1 to 3 cm long; bracts gradually reduced<br />
upward <strong>from</strong> lowest flowering node ________________________________________ S. subcordata<br />
16. Fruiting calyx lobes triangular-deltoid and abruptly apiculate, less than half as long as <strong>the</strong> calyx<br />
tube.<br />
18. Stem nodes not bearded (sometimes obscurely fine- hairy); stem angles with scattered,<br />
pustulate, short deflexed hairs; plants <strong>of</strong> high-elevation montane habitats _____________ S. latidens<br />
18. Stem nodes bearded; lower stem angles copiously pubescent with 3-celled spreading hairs;<br />
plants <strong>of</strong> low-elevation Piedmont and Coastal Plain habitats _____________________ S. mat<strong>the</strong>wsii<br />
key 2<br />
1. Fruiting calyx lobes deltoid to broadly triangular, less than half as long as <strong>the</strong> calyx tube.<br />
2. Stem sides pubescent; foliage glandular, <strong>of</strong>ten musky-scented when crushed.<br />
3. Petioles short to absent, blades mostly rounded to truncate-cordate.<br />
4. Lower leaf surface densely glandular with atomiferous/short- stipitate golden glands; blades<br />
generally ovate, broadest toward <strong>the</strong> base ______________________________________________S. eplingii<br />
4. Lower leaf surface glandular, but not densely so; blades generally elliptic, somewhat lyrate toward<br />
base, broadest near <strong>the</strong> middle _________________________________________________________ S. iltisii<br />
3. Petioles well developed, blades cordate at base.<br />
5. Top <strong>of</strong> blooming stem frequently flexuous, somewhat lax; blades ovate-rounded ________________S. cordata<br />
5. Top <strong>of</strong> blooming stem stiffish, not lax; blades oblong-elliptic _______________________________ S. nuttallii<br />
2. Stem sides glabrous; foliage not glandular (occasionally with a few scattered glands) or musky-scented.<br />
6. Stem nodes not bearded (sometimes obscurely fine-hairy); stem angles with scattered, pustulate, short<br />
deflexed hairs; plants <strong>of</strong> high-elevation montane habitats ___________________________________ S. latidens<br />
6. Stem nodes bearded; lower stem angles copiously pubescent with 3-celled spreading hairs; plants <strong>of</strong><br />
low-elevation Piedmont and Coastal Plain habitats ______________________________________ S. mat<strong>the</strong>wsii<br />
1. Fruiting calxy lobes lanceolate, narrowly triangular, deltoid-acuminate, or nearly subulate, more than half as<br />
long as <strong>the</strong> calyx tube.<br />
7. Leaf blades linear to lanceolate.<br />
8. Leaf blade margins entire to crenate; plants generally glabrous ____________________________ S. hyssopifolia<br />
8. Leaf blade margins serrulate with at least a few teeth; plants glabrous or pubescent.<br />
9. Stems strict or sparingly branched; leaves sessile or barely petioled, <strong>the</strong> blades crenate to serrate<br />
with shallow teeth _________________________________________________________________ S. aspera<br />
9. Stems frequently lax and branched <strong>from</strong> <strong>the</strong> upper nodes; leaves obviously petioled, <strong>the</strong> blades<br />
sharply too<strong>the</strong>d _________________________________________________________________ S. tenuifolia<br />
7. Leaf blades wider, rounded, oblong to elliptic.
Fleming et al., A new species <strong>of</strong> <strong>Stachys</strong> <strong>from</strong> <strong>the</strong> sou<strong>the</strong>astern United States 17<br />
10. Petioles poorly developed or absent, < 1 ∕5 as long as <strong>the</strong> blade.<br />
11. Upper stem sides glabrous.<br />
12. Calyx hispidulous to strongly hispid with long, stiff hairs; stem angles moderately to densely<br />
pubescent; leaves subsessile, with very short petioles; bracts abruptly reduced upward <strong>from</strong><br />
lowest flowering node ________________________________________________________ S. hispida<br />
12. Calyx sparsely to moderately pubescent with short s<strong>of</strong>t hairs; stem angles moderately pubescent<br />
to nearly glabrous; leaves with petioles 1 to 3 cm long; bracts gradually reduced upward<br />
<strong>from</strong> lowest flowering node ________________________________________________ S. subcordata<br />
11. Upper stem sides variously pubescent or glandular-pubescent.<br />
13. Corollas purple; rare adventive _________________________________________________S. palustris<br />
13. Corollas pink; native species.<br />
14. Blades copiously pubescent, usually velvety; VA and WV only _____________________ S. arenicola<br />
14. Blades pubescent but never velvety; KY-TN-GA-AL-SC ___________________________ S. nuttallii<br />
10. Petioles obvious, those <strong>of</strong> <strong>the</strong> mid stem > 1 ∕5 <strong>the</strong> length <strong>of</strong> <strong>the</strong> blade.<br />
15. Leaf blades dentate or sharply dentate; stem angles abundantly pubescent with long (to 3 mm),<br />
spreading or somewhat retrorse hairs _____________________________________________ S. clingmanii<br />
15. Leaf margins crenate to serrulate, but never dentate; stem angles glabrous or pubescent (if <strong>the</strong><br />
latter, <strong>the</strong>n with scattered, stiffish, retrorse hairs).<br />
16. Annual; corolla scarcely exserted <strong>from</strong> calyx ________________________________________ S. agraria<br />
16. Perennial; corolla prominently exserted <strong>from</strong> calyx.<br />
17. Plants commonly glandular on stems, leaves, and within inflorescence.<br />
18. Rhizomes terminated by white, rounded tubers; blades oblong; weedy _________ S. floridana<br />
18. Rhizomes without tubers; blades elliptic, cordate; not weedy __________________ S. nuttallii<br />
17. Plants not glandular on stems, leaves, or within inflorescence.<br />
19. Calyx usually hispidulous to hispid with stiff, eglandular hairs; fruiting calyx lobes<br />
straight and conspicuously apiculate ______________________________________ S. hispida<br />
19. Calyx usually glabrous or sparingly pubescent; fruiting calyx lobes usually recurved<br />
and only weakly apiculate ____________________________________________ S. tenuifolia<br />
acknowledgments<br />
we acknowledge <strong>the</strong> kind cooperation <strong>of</strong> <strong>the</strong> herbarium curators who have made loans <strong>of</strong> <strong>the</strong>ir material<br />
available to us. we especially thank randee humpreys and personnel <strong>of</strong> lewis ginter <strong>Botanical</strong> garden for<br />
allowing long-term access to <strong>the</strong> Virginia commonwealth university herbarium, greg Plunkett, pr<strong>of</strong>essor at<br />
Vcu, for helping arrange and process loans and for enabling a critical study <strong>of</strong> specimens in this workspace,<br />
and artist lara call gastinger for preparation <strong>of</strong> <strong>the</strong> botanical illustration. <strong>the</strong> South carolina nongame &<br />
heritage trust Program provided travel assistance for field work (nelson 1988) at <strong>the</strong> site <strong>of</strong> <strong>the</strong> type collection.<br />
Substantial support for field work by nelson was provided by a grant <strong>from</strong> <strong>the</strong> Barbara J. harvill fund<br />
for floristic research in Virginia. holly Selvig provided preliminary morphological studies as an independent<br />
study through a howard hughes grant at <strong>the</strong> university <strong>of</strong> South carolina, and assisted with field observations.<br />
grant godden and an anonymous reviewer provided helpful reviews. ursula engelbrecht (uSc) and<br />
Boris Schlumperger (herrenhausen gardens, hanover, germany) considerably improved <strong>the</strong> german abstract.<br />
reFerences<br />
delCourt, H.R. and P.A. delCourt. 1986. Late-Quaternary vegetational history <strong>of</strong> <strong>the</strong> central Atlantic states. In: J.<br />
McDonald and S.O. Bird, eds. The Quaternary <strong>of</strong> Virginia, Virginia Commonwealth Division <strong>of</strong> Mineral Resources,<br />
Charlottesville. Pp. 23–35.<br />
fernald, M.L. 1938. Noteworthy plants <strong>of</strong> sou<strong>the</strong>astern Virginia. Rhodora 40:364–491.<br />
fernald, M.L. 1941. Ano<strong>the</strong>r century <strong>of</strong> additions to <strong>the</strong> Virginia flora. Rhodora 43:485–665.<br />
fernald, M.L. 1943. Virginian botanizing under restrictions. Rhodora 45:357–516.<br />
fernald, M.L. 1950. Gray’s manual <strong>of</strong> botany, 8th edition. American Book Company, New York.<br />
gleason, H.A. and A. CronQuist. 1991. Manual <strong>of</strong> vascular plants <strong>of</strong> nor<strong>the</strong>astern United States and adjacent Canada.<br />
Ed. 2. New York Bot. Gard., New York.
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MaCdonald, L.E. 2000. Plant species <strong>of</strong> <strong>the</strong> Virginia Coastal Plain flora that are disjunct <strong>from</strong> <strong>the</strong> mountains: <strong>the</strong>ir<br />
distribution, abundance and substrate selectivity. Unpublished M.S. Thesis, The College <strong>of</strong> William and Mary,<br />
Williamsburg, VA.<br />
Mulligan, G.A. and D.B. Munro. 1989. Taxonomy <strong>of</strong> species <strong>of</strong> North American <strong>Stachys</strong> (Labiatae) found north <strong>of</strong><br />
Mexico. Naturaliste Canad. 116:35–51.<br />
nelson, J.B. 1981. <strong>Stachys</strong> (Labiatae) in sou<strong>the</strong>astern United States. Sida 9:104–123.<br />
nelson, J.B. 2008. A new hedge-nettle (<strong>Stachys</strong>: <strong>Lamiaceae</strong>) <strong>from</strong> <strong>the</strong> Interior Highlands <strong>of</strong> <strong>the</strong> United States, and<br />
keys to <strong>the</strong> sou<strong>the</strong>astern species. J. Bot. Res. Inst. <strong>Texas</strong> 2:761–769.<br />
Ware, D.M.E. and S. Ware. 1992. An Acer barbatum-rich ravine forest community in <strong>the</strong> Virginia Coastal Plain.<br />
Castanea 57:110–122.