Phytoneuron

v of the V. officinalis

Guy L. Nesom
2925 Hartwood Drive
Fort Worth, TX 76109
www. guynes om. com

ABSTRACT
Verbena madrensis Nesom sp. nov., is described from Coahuila, Nuevo Leon, and
Tamaulipas in northeastern Mexico. It is most similar to V. menthifolia and apparently also closely
related to V. officinalis and V. supina. Seven South American species and tliree from Africa and
Australia also are hypothesized to be closely related to V. officinalis. The closest relationship of V.
halei has sometimes been hypothesized to be with V. officinalis, but it apparently is closer to other
species native to North America. A key to the five species known either as waifs or naturalized in
North America is provided and typification, a description, and commentary is given for each.
KEY WORDS: Verbenaceae, Verbena madrensis sp. nov., V. menthifolia, V. halei. Verbena
officinalis. V. supina, taxonomy, USA Mexico, South America. Galapagos, Australia

In review of the taxonomy of North American Verbena toward preparation of the t
the Flora of North America volumes, attention has been given to the delimitation of V. officinalis L.
and its apparent close relatives, including V. menthifolia Berth, and V. supina L. These species, along
with others from Africa, Australia, and South America, are treated by Nesom (2010) within Verbena
sect. Verbena ser. Verbena. Verbena halei Small has been hypothesized in earlier literature to be
most closely related to V. officinalis but has been placed in a different series (Nesom 2010). In North
America, V. menthifolia and V. halei are native and V. officinalis is naturalized; V. supina lias been
recorded as a waif but does not appear to be naturalized. A previously undescribed species from
northeastern Mexico also belongs in ser. Verbena.

VERBENA MADRENSIS Nesom, sp. nov. Figs. 1, 2. Type: MEXICO. Nuevo Leon. [Mpio.]

Galeana, Hacienda Pablillo, 3 Aug 1936, M.E. Taylor 49 (holotype: TEX!; isotype: MO!)
Verbenae menthifoliae Benth. similis vestimento plerumque strigilloso, spicis iaxis elongatis,
et nuculis parvis superficiebus commissuralibus apicem attengentibus sed differt rhachidibus dense
persistente glandulosis, foliis incrassatis regulatim grosse serratis marginibus revolutis, bracteis
floralibus hispido-hirsutis alandulosi^, ct eorollis majoribus.

Plants perennial herbs, taprooted, sometimes producing basal offsets or basal branches
layering (Hinton 24412). Stems mostly 1 from the bass, erect to ascending-erect, 25-60(-120) cm,
very sparsely hirsute-strigose, eglandular or sparsely stipitate-glandular. Leaves mostly on proximal
half of stems, ovate to lanceolate or oblong-lanceolate in outline, lower and midstem 2.5-4(-5) cm x
5-17(-25) mm, margins strongly revolute, coarsely seiTate with 3-6 teeth per side, sometimes 3-lobed
or pinnatifid, '• »bes acut^ \tui> slii>hil\ smptessed a<h\tali\ hi^pid-stiigose adaxially, hispid-hirsute
abaxially mostly along the veins, eglandular. Fruiting spikes 1 or 3-7 from distal branches, open-
elongate or sometimes denser with overlapping fruits, 6-20 cm, rachis densely and persistently
stipitate-glandular; floral bracts ovate to ovate-lanceolate, slightly shorter than the calyces, hispid-
hirsute, stipitate-glandular, margins ciliate. Calyces 2.2-3 mm, hirsute to hirsutulous, densely
nsmureb strpjtaifc-gLmdi'lu lul es icimmate-Tilrionn, hispid, not connivent. Corollas blue to

purplish pink, tubes (3-)3. 5^1.5 mm, 2 mm longer than the calyces, limbs 5-9 mm in diam. Nutlets
(1.3— )1.5— 1.8 mm, longitudinally ridged, commissural faces reaching very tip, minutely papillate to
papillate-bullate, rarely bare.

Flowering May-Aug(-Oct). Thorn forest, oak, pine, oak-pine, and oak-fir woods, rocky
slopes, dry stream beds, roadsides; 500-2400 m; Mexico (Coahuila, Nuevo Leon, Tamaulipas).

Additional collections examined. MEXICO. Nuevo Leon. Chipinque, pine woodland, 1 Jul
1947, Barkley et al. 7150 (TEX); Mpio. Linares, Rio Iturbide, IRF Iturbide, 900 m, 30 Sep 1979,
Hinton et al. 17695 (FflNTON); Mpio. Aramberri, N of Aramberri, IRF Lampacitos, 995 m, 16 Jun
1990, Hinton 20329 (TEX); Mpio, Montemorelos, El Pastor, oak woods, 460 m, 19 May 1994, Hinton
24216 (TEX); Mpio. Santiago. La Nogalera, glade in oak woods, 1005 m. 16 Jun 1994, Hinton 24412
(TEX-2 sheets); Mpio. Santiago. Blanquillo, bushy hillside, 540 m, 22 Jun 1994, Hinton 24440
(TEX); Mpio Galeaiia, Hacienda ! 'jeneguilla^ on < 'erro Potosi, open, drv, rocky slopes in chaparral,
8000 ft, 11 Aug 1938, Mexican Biol. Exped. Univ. of Illinois 992 (ARIZ)' Sierra Madre Mts,
Monterrey, 18 Jul 1933, Mueller 157 (TEX); Mpio. Villa Santiago, Canon Denuncio, common in less
dense oak wood of upper canyon, 22 Jun 1935, Mueller 2010 (MO); Montemorelos, 3 Apr 1902,
Nelson 6096 (MO): 5*mi SE of Monterrey on Hwy 85 and 2 mi W of Valle Alto, thorn forest and dry
stream bed, limestone and shale, 14 Jul 1971, Parker 424 (TEX); Chipinque Mesa, 1900 ft, 2 Aug
1970, Roberts 57 (TEX); Linares, 4 Jun 1971, Seigler and Becker OS-3149 (MO, TEX); 1 mi down
road from Chipinque, Monterrey, Apr 1961, Smith 476 (TEX); Hacienda Vista Hermosa, 35 mi S of
Monterrey, ca. 2200 ft, 27 Jun 1939, White 15^7 (ARIZ). Tamaulipas. Sierra de San Carlos, Cerro
del Diente, 17 km S de San Carlos, 1100 m, bosque mesofilo de montana con Abies, Ouercus,
Staphylea, etc, 22 May 1988, Hernandez 2246 (TEX); Sierra de San Carlos, ca 5 mi S of San Carlos,
near top of slope, igneous bedrock, open woods of scattered oaks and pines, ca. 1200 m, 18 Jun 1987,
Nesom 6141 (TEX).

Verbena madrensis is recognized by its single-stemmed habit, glabrate stems with sharp,
strongly nerved angles, widely spaced and elongate leaves with coarsely serrate, revoiute margins,
loose, elongate spikes in nimcLs it tiK ipics ol tl« stems in J bntithts pcisistcith ind dtn u\
4i{Htjte i mdulai t ichi^^s huuite to lit si lulus* iul stipi f ste-^hndulat fiuial biacts aid (abet.
short corolla tubes, and nutlets with commissural faces reaching the tip. The oblong-lanceolate
cauline leaves with sharp, slightly recurving teeth are reminiscent of those of V. canescens, but the
nutlet comissural faces in V. madrensis reach all the way to the nutlet tip, like those of V. officinalis
and V. menthifolia.

Among North American species. Verbena madrensis is closely similar to V. menthifolia
Berth. Moldenke (1964) cited Mueller 157, Mueller 2010, and Taylor "49 (above, as V. madrensis)
among plants he identified as V. menthifolia. The geographic range of V. madrensis lies at the
northeastern corner of that of V. menthifolia, essentially allopatric with it, and is much smaller in
comparison. From Nuevo Leon, in the region where V. madrensis is common, I have seen only a
single collection of V. menthifolia: 32 mi S of San Roberto, along Hwy 57, near small pond, 6000 ft,
11 Jul 1963, McGregor et al. 492 (SMU). Two collections from southeastern Coahuila are V.
menthifolia: Sierra de la Concordia, 6 km NE del Ejido La Casita, 25° 14' 05" N 101° 25' 25" W,
bosque de Pinus cembroides, Ouercus laeta, Q. saltillensis, Arbutus xalapensis, y Garrya wrightii,
canyones y ladera media de exposicion oeste, 2100-2300 m, 10 Oct 1996, Encina 288 (TEX). Mpio.
Arteaga. Los Lirios->El Cercado, roadside in pine and oak woods, 1 m tall, 2405 m, 29 Jul 1995,
Hinton et al. 25422 (HTNTON-digital image!). 1 have not seen a collection of V. menthifolia from
Tamaulipas.

The two species are distinguished by the following contrasts.

1. Leaf margins regularly, coarsely serrate, revolute; inflorescence rachis densely and persistently
stipitate-glandular; floral bracts hispid-hirsute, stipitate-glandular; corolla tubes (3-)3. 5-4.5 mm, 2
mm longer than the calyces, limbs 5-9 mm in diam Verbena madrensis

1. Leaf margins deeply and irregularly toothed to lobed, not revolute or only slightly so; inflorescence
rachis eglandular or rarely sparsely stipitate-glandular and quickly becoming eglandular; floral
bracts glabrous to sparsely strigillose, eglandular to spaisj\ ^tipitatL-^Luiduldi. corolla tubes 2.5-3
mm, 0.5-1 mm longer than the calyx, limbs 1.5— 3(— 5) mm in diam Verbena menthifolia

Features in common between Verbena madrensis and V. menthifolia are a mostly strigillose
vestiture, elongate, slender spikes of remote flowers and fruits (not overlapping each other on the
spikes), relatively Minll i rollas i*sd small mJets with cotiirms^unl 1 a =s i^chint, th nut) J. tin
The European native V. officinalis also is closely similar. A close relationship between V. menthifolia
and V. officinalis was hypothesized by Perry (1933), who also included V. halei as a member of this
group. She noted that distinctions of V. menthifolia are "perhaps ... differences only of degree and
may be merely variations of V. officinalis; nevertheless, for the present it seems prefereable to retain
the name V. menthaefolia for the American representative" (p. 265). The present study confirms their
distinction. These species, along with another that is newly described in this manuscript, can be
distinguished by the following contrasts.

1. Stems mostly prostrate to decumbent-ascending: mid-cauline leaves triangular to deltate or ovate-

cuneate in outline Verbena supina

1. Stems mostly erect; mid-cauline leaves ovate to lanceolate or oblong-lanceolate in outline.

2. Leaves mostly basal and lower cauline, distal cauline become linear and entire; leaf margins

revolute; commissural faces ending below the nutlet tips Verbena halei

2. Leaves evenly distributed along stems, often largest at Mjdstem, ptoxnual ^nd distal Liuliiie
similar in morphology; leaf margins not revolute or only slightly so; commissural faces reaching
the nutlet tips.

3 . Rachis eglandular or nearly so, Verbena menthifolia

3. Rachis pcisistentlv ^hpitate-glandalai

4. Corolla tubes 2.5-3 mm, 0.5-1 mm longer than the calyces, limbs 2-3 mm in diam; leaf
margins coarsely and unevenly crenate to incised-crenate or crenate-serrate, not revolute or

only slightly so Verbena officinalis

4. Corolla tubes (3-)3.5-4.5 mm, 2 mm longer than the calyces, limbs 5-9 mm in diam; leaf
margins regularly coarsely serrate, narrowly revolute Ver bena madrensis

VERBENA MENTHIFOLIA Benth., PI. Hartw., 21. 1839. TYPE: Mexico. [Guanajuato]. Leon,
1839, [no other collection information]. Hartweg 175 (holotype: K, photo-MO!; isotype: NY,
digital image!). The protologue: "175. VERBENA (Euverbena) menthaefolia, sp. n., caule
erecto ramoso pilosiusculo. foliis obovato-cuneatis subincisis laciniis acutiusculis grosse
dentatis, spicis elongatis gracilibus paniculatis, bracteis parvis, corollis calyce triplo
longioribus, fructibus remotis."

Verbena hintonii Moldenke, Phytologia 1: 439. 1940. TYPE: Mexico. Michoacan. Distr. Zitacuaro,
Zitacuaro-Bosque, grassy bank, 28 Jun 1938, Hinton et al. 11991 (holotype: NY-digital
image!; isotypes: F-digital image!, GH, HINTON-digital image!, MICH, TEX!, US-digital
image!, WTU).

Verbena setosa Mart, & Gal., Bull. Acad. Roy. Sci. Bruxelles 11: 321. 1844. TYPE: Mexico.

Hidalgo. "Dans les forets de Moran, pres de Real del Monte, et del Sabino, pres

d'lzmiquilpan, au nord de Mexico, de 6000-7500 ft," fl. lilas, Aout 1840, H.G. Galeotti 778

(holotype: BR, photo-LL!, photo-MO!). The type of V. setosa is not currently listed in the

online account of BR types.
Verbena Carolina forma albiflora Moldenke, Phytologia 7: 430. 1961. TYPE: Mexico. Sonora.

Region of the Rio de Bavispe, Canon de El Temblor, 19 Aug 1940, S.S. White 3379

(holotype: MICH digital image'). O'Leary et al. (2010) treat this as a synonym of V.

Carolina.
Verbena comonduensis Moldenke, Phytologia 18: 343. 1969. Verbena menthifolia var. comonduensis

(Moldenke) Moldenke, Phytologia 46: 1 55. 1980. TYPE: Mexico. Baja California, Comundu,

common roadside weed for 10 miles in either direction, 19 Mar 1969, A.F. and A.R.

Moldenke. 2922 (holotype: LL!). Topotype: Comondu, moist sandy depressions on flats along

roadside, 21 Apr 197 '1, Moldenke and Moldenke 25407 (LL, TEX).

Plants perennial herbs, usually taprooted (rarely with basal offsets. Encina 288, TEX).
Stems mostly 1-3 from the base, erect, (30-)40-70 cm, sparsely strigose to hirsute-strigose along the
ridges with appressed to ascending hairs, eglandular. Leaves relatively evenly distributed, on stems,
even-sized or sometimes largest at midstem, ovate to broadly lanceolate in outline, midstem blades
2.5-4.5(-8) cm x 15-25(-40) mm, strigose-hirsute to hirsute-strigose abaxially, strigillose to hirsute-
strigose adaxially, eglandular, margins not revolute or only slightly so, deeply toothed to pinnately
lobed, sometimes evidently 3-lobed through a pair of large proximal divisions, divisions ovate-oblong
to triangular, subpetiolate. Fruiting spikes (l-)3-12 in panicles, elongate and slender, 10-20(-30)
cm, rachis eglandular to sparsely stipitaie-glandular; floral bracts ovate-lanceolate, 1.5-2 mm, shorter
than the calyces, glabrous to sparsely strigillose, eglandular, margins ciliate. Calyces 2.3-3 mm,
closely to loosely strigillose with even-length hairs, eglandular to very sparsely stipitate-glandular
(moderately ^ewlu-g! mduiai in Sonoia .ind DF) lohi,s deltate to triangular, not connivent to
subconnivent. Corollas blue to purple, tubes 2.5-3 mm, 0.5-1 mm longer than the calyx, limbs 1.5-
3(-5) mm in diam. Nutlets separating at maturity, 1.4-1.8(-2) mm, commissural faces extending
completely to the nutlet tips, minutely acicular-papillate.

Flowering Apr-Tun. Canyon bottoms, around springs and tanks, river and canal banks, wet
places in arid habitats (USA); 50-1200 m; Arizona, California, New Mexico. Texas; Mexico
(Aguascalientes, Baja California, Baja California Sur, Chihuahua, Coahuila, Distrito Federal,
Durango, Guanajuato, Hidalgo, Jalisco, Mexico, Michoacan, Morelos, Nuevo Leon, Oaxaca,
Queretaro, Puebla, San Luis Potosi, Sinaloa, Sonora, Veracruz, Zacatecas); introduced in Europe.
Attributions of V. menthifolia to Chiapas and Guatemala are based on collections discussed under V.
officinalis.

Verbena menthifolia is known to me from Texas by only a single collection. Texas. Brewster
Co. : Black Gap Refuge, Horse Canyon, 100 mi S of Alpine, infrequent perennial around old dirt tank,
limestone soil, 2000 ft, 24 Jun 1950, Warnock & Hinkley BG-108 (SMTJ). Moldenke (1964a) cited
two other collections: [Bexar Co.?]: between Kerrville and San Antonio, Jones 28294 (POM); Uvalde
Co.: W of Uvalde, 26 Apr 1931, Jones s.n. (POM) — I have not seen these but it is improbable that
they are V. menthifolia.

Peterson and David (1998, p. 23) recorded the occurrence of Verbena menthifolia in Chaves
Co., New Mexico, as "Occasional in lowlands and arroyos and on roadsides" in Bitter Lake National
Wildlife Refuge, northwest of Roswell. This report has been the basis for citation of V. menthifolia as
an element of the New Mexico flora in various accounts (Martin & Hutchins 1981; Allred 2009;
USDA NRCS 2010; Kartesz 2010). A voucher for this, however, has not been located in the present
study, and a New Mexico collection of V. menthifolia is not found in NMC, UNM or SJNM. That
species would be considerably out-of-range and out-of-habitat in Chaves County, and I speculate that

the plants observed there by Peterson and David probably were V. halei at the northwestern s

On. the other hand, a collection from New Mexico cited by Moldenke (1963) and repeated in
various other accounts as Verbena halei proves instead to be V. menthifolia. Otero Co.: sand dunes
west of AJaniogotdo, 3 May 1930, M.E. Jones 26229 (DS-digital image!). The distalraost leaves on
this plant are linear and entire (as char act eristic of V. halei), but the stem is strictly erect and sparsely
strigose, the basal leaves are deciduous, the cauline narrowly elongate and narrowly lobed, and the
spikes are relatively short and few-flowered compared to typical V. halei.

Verbena menthifolia has been collected abundantly in California (at least from San
Bernadino, San Diego, Los Angeles, and Riverside counties) and Arizona (Pima. Pinal, and Yuma

counties) and in. Mexico (states as cited above) a

Verbena menthifolia from Hidalgo, Edo. Mexico, and Distrito Federal sometimes lias stems,
leaves, and calyces with vestiture variable toward strigose-hirsute with uneven length hairs, but like
other plants of the species over the rest of its range, they are essentially eglandular and produce small
corollas. An Arizona plant identified and studied by Yuan and Olmstead (2008) as Verbena
menthifolia was heterozygous at both the PHOT1 and PHOT2 nuclear gene loci, and the gene trees
indicated that the individual was of hybrid origin, with V halei as one of the putative parental species
I have not been able to study a voucher for tiiis record, but at least it is clear that V halei does not
presently occur sympatrically with V menthifolia,

VERBENA SUPINAL., Sp. PI. I: 21. 1753. TYPE: "Habitat in Hispania." LECTOTYPE (Moldenke

1965, p. 255): "Hispania," Loefiing 16 (S9. 1, Linnaean Herbarium Stockholm digital image!).

See detailed comments by Munir (2002).
Verbena supina L. forma erecta Moldenke, Phytologia 11: 259. 1965. Verbena supina L. var. erecta

(Moldenke) Munir, J. Adelaide Bot. Gard. 20: 62. 2002. TYPE: Spain. Lieux incultes,

Algeciras, 24 Jun 1887, E. Reverchon 81 (holotype: GB, photo-NY fide Moldenke 1965,

photo-TEX! ; isotypes : BR, S).

Plants annual or weakly perennial herbs, taprooted, sometimes with adventitious roots on the
proximal, procumbent portions of the stems. Stems branching profusely from crown, procumbent to
decumbent-ascending, less commonly erect to erect-ascending, (10— )15— 35(— 50) cm, sparsely to
moderately hirsute-strigose to hirsute, hairs sometimes slightly deflexed, eglandular to sparsely
stipitate-glandular. Leaves evenly distributed and relatively even-sized, triangular to deltate or ovate-
cuneate in outline, midstem blades 2-4 cm x 1.5-2.5 cm, strigose to hirsute-strigose or hirsute,
egtandular to stipitate-glandular, margins revolute, coarsely toothed with ovate to oblong-ovate or
obovate teeth or divisions to 1-2-pinnatisect, often 3(-5)-lobed, margins \cin^ rmpses^eJ adaxudK
petioles 5-12 mm. Fruiting spikes 1 or 1-3, initially dense but elongating and fruits becoming looser
and remote, 1.5— 3(— 5) cm; rachis strigillose, minutely sessile- to short-stipitatc-ghndulai floral bracts
linear-lanceolate to oblong, 1-1.5 mm, shorter than the cafyces. Calyces 1.8-2 mm, hirsutulous-
hirtellous to hirsutulous -strigose, minutely stipitate-glandular, lobes subulate, not connivent.
Corollas blue to purplish or lilac, tubes 2-2.5 mm, 0.5-1 mm longer than the calyces, limbs 1.5-2.5
mm in diam. Nutlets separating at maturity, 1.8-2.2 mm, commissural faces extending fully to nutlet
tips, bare. In = 14.

Flowering mostly spring-summer. Dry and wet places, grasslands, disturbed sites, at a wide
range of elevation; Mediterannean Europe, Asia, Africa: naturalized in northern Europe and Eurasia,
Asia, Africa, Atlantic Islands (Canary Islands), Pacific Islands (New Zealand), Australia.

Verbena supina was recorded from Alabama by Moldenke (1965), based on waifs collected
from along Mobile Bay in 1892 and 1893. The species has not been recorded since that time from
anywhere in the USA and is not considered part of the naturalized flora of North America. Alabama.
[Mobile Co.]: Hunter's Wharf, 5 Sep 1892, Mohr s.n. (US-2 sheets); Mobile, Jul 1893, Mohr s.n.
(NY) — both Mohr collections as cited by Moldenke.

Common names for Verbena supina are trailing vervain and caipet vervain, reflecting its
characteristically prostrate habit. Other distinctive and diagnostic features are the deltate to ovate (in
outline) and relatively small leaves, relatively short spikes, loosening proximally but remaining
compact in the distal 1/3-2/3, short floral bracts, small corollas, fruits with comissural faces extending
fully to the nutlet tips, and characteristic stipitate-glandularity, at least sparsely. Munir (2002)
observed that in Verbena supina in Australia, the commissural faces of the nutlets always are smooth
and the vegetative parts are eglandular, while in other parts of the native and naturalized range, plants
sometimes are stipitate-glandular and the commissural faces papillate.

The distinctiveness of Verbena supina var. erecta, was emphasized by Munir, w r ho noted (p.
64) that "the possibility of mistaking this taxon with the typical form would be minimal." And (p. 62
and 64) 'The var. erecta is closely related to the typical variety in having almost similar shaped
leaves, inflorescence, calyx, corolla and mericarps . . . [but] . . . differs from the typical form of the
species in having its stems usually erect or suberect from the beginning, usually robust and purplish,
almost glabrous and shiny or very sparsely puberulous. Leaves usually glabrous adaxially,
puberulous on the veins abaxially, lobes obtuse or rounded at the apex." Munir also noted that var.
erecta is known to occur in several countries of Eastern Europe, North Africa, and the Mddle East.
In context of Munir's account as summarized here, it appears that var. erecta would be appropriately
treated at specific rank. Without commenting on patterns of variability, O'Leary et al. (2010) noted
only that "El analisis del ejemplar tipo permitio determinar que este taxon es un sinonimo de V.

VERBENA OFFICINALIS L., Sp. PI. I: 20. 1753. TYPE: "Habitat in Europae Mediterraneae
ruderatis," probably from central Europe (see Mchael 1997). LECTOTYPE (Verdcourt in
Jarvis et al., Regnum Veg. 127: 98. 1993): Herb. Clifford: 11, Verbena 6, sheet 6 (BM).

Verbena officinalis L. var. prostrata Gren. & Godr., Fl. France 2: 718. 1853. TYPE: France. Sables
des environs de Bayonne, bords des chemins et decombres, Jun-Oct, no collector or date
specified. Described in the protologue simply as "Tige etalee-couchee."

Verbena spuria L., Sp. PL 1: 20. 1753. Verbena officinalis L. var. spuria (L.) Hook, Companion Bot.
Mag. 1: 176. 1836. TYPE: O'Leary et al. (2010) designated (from photos) "Muhlenberg 68,"
"PH 3073 no visto" as the neotype. Typifieation of V spuria is under further study by Nesom
(in manuscript).

Verbena riparia Raf ex Small & Heller, Mem. Torrey Bot. Club 3: 12. 1892. Verbena urticifolia var.
riparia (Raf. ex Small & Heller) Britton, Mem. Torrey Bot. Club 5: 276. 1894. LECTOTYPE:
(Moldenke 1964, p. 101): USA. North Carolina. Caldwell Co: Along the John's River near
Globe, 3 Jul 1891, J.K. Small s.n. with AA. Heller (NY-digital nragj isolectotype: MO!).
O'Leary et al. (2010) designated a ieetotype for this, not realizing that it had been done
previously.

Verbena domingensis Urb., Symb. Antill. 5: 484. 1908. LECTOTYPE (O'Leary et al. 2010):
Hispaniola. Dominican Republic. Ad Angostura del Rio Yaqui, "in rupibus calcareis, ad
ripam fluminis," 210 m, 8 May 1887, H.F.A. von Eggers 1828 (NY-digital image!). Placed
by O'Leary et al. (2010) as a synonym of V. menthifolia.

Verbena russellii Moldenke, Phytologia 2: 55. 1941. TYPE: Mexico. Sinaloa, Vicinity of Culiacan,
moist field, 21 Apr 1910, J.N. Rose 14850 (holotype: NY digital image!; isotypes: US digital

1). O'Leary et al. (2010) placed V russellii as a synonym of V.

Verbena domingemis Urb. var. cubensis Moldenke, Plrytologia 50: 309, 1982. TYPE: Cuba. Campo
Florido, 13 Mar 1905, A.H. Curtiss 677 (holotype: NY-digital image!; isotype: F-digital
image!). Placed by O'Leary et al. (2010) as a synonym of V. menthifolia.
Verbena domingemis Urb. forma foliosa Moldenke, Phytologia 34: 19. 1976. TYPE: Hispaniola.
Dominican Republic. [Prov.] Pedernales. Near Canote, ca. 5 mi W of Aceitillar, Baoruco
Mts., in thickets at the bottom of the gorge, ravine on limestone, 1400 m, 9 Nov 1969, A.H.
Liogier 16846 (holotype: NY-digital image!). Placed by O'Leary et al. (2010) as a synonym
of V menthifolia.

Plants annual or weakly perennial herbs, taprooted. Stems mostly 1 from the base, erect, 30-
140 cm, glabrous to scabrous to sparsely liispid-strigose or hirsute-strigose along the angles,
eglandular, becoming sparsely stipitate-glandular distally into the inflorescence. Leaves relatively
evenly distributed on stems, narrowly obovate to oblanceolate or lanceolate in outline, midstem
blades 2-7 cm x 8-^10 mm, even-sized or sometimes largest at midstem, hirsute-strigose abaxially and
adaxially, eglandular at maturity but stipitate-glandular abaxially when young, margins coarsely and
unevenly crenate to incised-crenate or crenate-serrate, basal often 3-lobed or rarely pinnately lobed,
not revolute or only slightly so; petioles 5-15(-20) mm, narrowly or broadly winged. Fruiting
spikes (3-)5-15 in panicles, elongate and slender, 3-15 cm, rachis sparsely stipitate-glandular; floral
bracts ovate-acuminate to triangular, 1.5-2 mm, shorter than the calyces. Calyces 2-2.2 mm, hirsute
to hispid-strigose, minutely stipitate-glandular, lobes shallow!;' deltate, not connivent. Corollas blue
to purplish, rarely pink, tubes 2.5-3 mm, 0.5-1 mm longer than the calyces, limbs 2-3 mm in diam.
Nutlets separating at maturity, 1.4-2 mm, commissural faces reaching the nutlet tips, acicular-
papillate, rarely bare. 2n = 14.

Flowering Jun-Aug. Ri\u lnn ! s dunes l sulside* 'mils t 5-700 m; introduced; Alabama,
California, Colorado, Connecticut. Delaware, District of Columbia, Florida, Georgia, Kentucky,
Louisiana, Maryland, Massachusetts, Michigan, Mssissippi, New Jersey, New Mexico, New York,
North Carolina, Oregon, Pennsylvania, Rhode Island, South Carolina, Tennessee, Virginia,
Washington, West Virginia, Wisconsin; Europe, Asia; introduced also in West Indies, Bermuda,
South America (Chile). Asia (Bhutan, China, India, Japan, Pakistan, Thailand), Africa (Canary
Islands, Natal, South Africa), Atlantic Islands (Azores), Pacific Islands (New Zealand), Australia.
Records from Colorado, Mssissippi, New Mexico, Oregon, and Wisconsin (as reported by PLANTS
Database) have not been confirmed in the present study.

Some plants in New York state are identified by the PLANTS Database as Verbena officinalis
var. prostrata, perhaps based on the record in House (1924, p. 589), but House identified the New
York plants as V. officinalis, citing "K spuria L.; Torr. Fl. N.Y. 2: 52. 1843" only as a synonym.

Verbena officinalis is not common in the USA outside of cultivation. At least some
collections perhaps reflec pljnts that are pe.vi te.it Mom cultivation and not clearly naturalized.
Plants and populations along river banks in montane North Carolina, Tennessee, Virginia, and West
Virginia, however, apparently have long been established, apparently naturalized, and often identified
as V riparia. It seems unusual that they would already have been well-established in these habitats
by the time that Rafinesque observed them (in the 1820s or 1830s), but they appear to be similar to V.
officinalis in all features. On the other hand, a number of collections identified as V. officinalis were
made in the same regions in the 1840s through the 1890s — from Alabama, Delaware, Georgia, New
Jersey, New York, North Carolina, Pennsylvania, Tennessee, Virginia, and others — suggesting that
this species perhaps was cultivated at that time for its medicinal properties and commonly escaped to
become more abundant then than it is now.

Additional collections identified, as "Verbena riparia" (if not with exclamation, then as cited
by Small and Vail 1893; Perry 1933; Moldenke 1964c). North Carolina. Stanly Co.: near falls of
Yadkin, 18 Aug, 1891, Small and Heller s.n. (F). Virginia. [Isle of Wight County]: Ragged Island,
[date], Fernald and Long 12453 (NY). Smyth Co.: banks of Staley Creek at Marion, valley of Middle
Fork of Holston River, 2200 ft, 16-? May 1892, Small and Vail s.n. (NY); banks of Cedar Creek,
valley of Mddle Fork of Holston River, 1900 ft, 16-? May 1892. Small and Vail s.n. (NY); bluffs of
Mddle Fork Holston River, near Marion, 2100 ft, 22 Jun 1892, Small s.n. (NY, OKL, TEX!, UC);
Mddle Fork, Holston River, 2100 ft, 1 Jul 1892, Small s.n. (BH-3 sheets, CAS, CINC, CM DUKE,
FLAS, NCU, NY, P, PENN, POM US, WVA); Marion, 2100 ft, 6 Jul 1892, Small s.n. (ARIZ, GH,
MO); Middle Fork Holston Rive!', 6 Jul 1892, Small s.n. (CAN. CS, ISC, MO, NY-2 sheets, NYS,
OC, US); about Marion, 2100 ft, 20 Jul 1892, Small s.n. (MO); Marion, Mddle Fork Holston River, 6
Jul 1922, 2100 ft, Small s.n. (MO, SMU).

These collections, also from riverbanks and clearly the same species as those cited above as

"Verbena riparia," were identified as V. officinalis: Tennessee. Carter Co.: banks of Doe River, 16-
17 Jul 1891, Small and Heller 84 (SMU), Small and Heller 484 (MO); Knox Co.: Knoxville,
roadside, Jul 1893, Ruth s.n. (MO). Virginia. Marion, 2100 ft, 29 Jun 1892, Small s.n. (MO)

In the protologue of Verbena domingensis, Urban (1908) noted that V officinalis differs from
V domingensis in its more brandling stems, smaller, mostly toothed (vs. pinnatifid or pinnately
parted) leaves, and smaller flowers. Perry (1933) treated V domingensis as a synonym of V
officinalis, but it was maintained as distinct by Moldenke (1964) and by Liogier (1957, 1994). The
type collections of V domingensis var. cubensis and V domingensis forma foliosa appear to be of
plants typical of V. officinalis in leaf morphology, suggesting th it the smaller-lea\ >_d p'jnf. pi * il 1\
are populational variants of naturalized V. officinalis.

Early reports of Verbena officinalis from the Valley of Mexico confused its identification
with V. menthifolia (fide Rzedowski & Rzedowski 1985). Moldenke (1964) cited as V. officinalis
only a single collection from Mexico: Nuevo Leon, Pennell 16880, PH, MEXU, US); I have not seen
this collection but the plants are probably V madrensis or V menthifolia. Most other collections from
western stales (e.g., Chihuahua, Durango, Sonora) originally determined as V. officinalis have proved
to be V. menthifolia.

Verbena russellii Moldenke (cited above) from Sinaloa is unequivocally is V. officinalis sensu
stricto. The only other representative of V. officinalis I have seen from Mexico was collected from
the same immediate area as the type of V. russellii: Sinaloa. Maraton, 12 mi W of Culican, heavy clay
of coastal plain, thorn forest, 100 ft, common annual, 14 Mar 1944, Gentry 701 1 (ARIZ).

A collection from Guatemala was cited as Verbena menthifolia by Moldenke (1964a) and
Gibson (1970): Huehuetenango, between Nenton and Las Palmas, Steyermark 51655 (US; cited by
O'Leary et al. 2010 as V menthifolia). I have not seen the Guatemala specimen, but distinctive plants
(CAS-DS, TEX-LL, MO) from adjacent southeastern Chiapas, e.g., Amatenango del Valle, Comitan
de Dominguez, Frontera Comalapa, Venustiano Carranza, near Trinitaria, near Cuahtemoc, and
others, probably represent the same entity, definitely not V menthifolia. These plants have relatively
few spikes in a panicle, the fruits becoming remote proximally. and glandular rachises, but the leaves
are uncharacteristically elongate (oblong-lanceolate) and the teeth and lobes are sharp-pointed. They
are remarkably similar to the Australian native V officinalis var. gaudichaudii Brio., recognized by
Mchael (1997) at specific rank If not that same entity, they will be justifiably treated as a distinct
species, previously undescribed.

Michael (1997) segregated at specific rank three native Australian taxa often identified as
infraspecific taxa of Verbena officinalis: V. macrostachya F. Mueii. (= V. officinalis var.
macrostachya (F. Mueii.) Benth.), V. gaudichaudii (Briq.) P.W. Michael (= V. officinalis var.
gaudichaudii Briq.), and V. africana (Fernand. &. Verde.) P.W. Michael (= V officinalis subsp.
africana Fernand. & Verde). These taxa differ in vestiture, leaf shape, and corolla size, their
treatment as species correlative with Michael's observation only that they are "sufficiently distinct"
from V. officinalis sensu stricto, which is naturalized in Australia. Munir (2002), however, treated
these three at varietal rani: within V. officinalis, adding V. officinalis var. monticola Munir (native), V.
officinalis var. eremicola Munir (native), and V. officinalis var halei (introduced) as inhabitants of
Australia. Munir's explicit rationale also was minimal — he noted (p. 80) that "the present author
believes that the above-named infraspecific taxa do not merit the status of species because they differ
from the typical form chiefly by the size of their spikes and flower parts and dentation of leaf blades.
These characters are very variable and there are intermediates between these taxa."

In review of the Australian taxonomic pattern, and to use Munir's ranks, vars. gaudichaudii
and africana occur over broad ranges and are mostly sympatric with each other; each of vars.
eremicola, monticola, and macrostachya has a narrowly restricted range and is allopatric with the
other two, but all three occur within the broader ranges of vars, gaudichaudii and africana; var.
officinalis and var. halei are each known from scattered collections over a broad area, essentially
sympatric with vars. gaudichaudii and africana. Verdcourt (1992) observed that two entities — V.
officinalis sensu stricto and V. africana sensu Michael— with "quite marked" differences occur
together in South Africa. If morphology supports distinct recognition of the five Australian endemics,
then each of them probably is appropriately treated at specific rank. Verbena halei clearly should be
treated so.

In addition to those discussed above, a group of South American species also is hypothesized,
to be closely related to Verbena officinalis: V. caniuensis Moldenke, V. demissa Moldenke, V.
filicaulis Schauer, V. gracilescens (Chamisso) Herter, V. grisea Rob. &. Greenm., V. swiftiana
Moldenke, and V townsendii Svenson. These species are discussed in relation to the formal
delimitation of series Verbena (Nesom 2010).

VERBENA HALEI Small, Bull. Torrey Bot. Club 25: 617. 1898. Verbena officinalis L. subsp.
halei (Small) S. C. Barber, Syst. Bot. 7: 454. 1982. Verbena officinalis L. var. halei (Small)
Munir. J. Adelaide Bot. Card. 20: 93. 2002. LECTOTYPE (Moldenke 1963, p. 162): USA.
Louisiana. No other collection information, Dr. J. Hale 245 (NY-digital image!). Small cited
"Hale 245," but the last digit of the collection number- on the type label is somewhat
ambiguous (either "5" or "3").
Verbena leucanthemifolia Greene, Pittonia 5: 135. 1903. TYPE. USA, Texas. Taylor Co.: Abilene, 19
May 1902, S.M. Tracy 7996 (holotype: ND-G 43312; isotjpes: GH, MO!, MSC, NY-digital
image!, TEX! digital image!, US-digital image!). The ND-G sheet has the annotation of
"type" in Greene's hand, fide Barbara Hellenthal.

Plants short-lived perennial to annual, taprooted. Stems mostly 1-3 from the base, erect,
(25-)40-8Q cm, glabrate to scabrous, or sparsely hirsute-strigose with upturned hairs on the ridges,
eglandular. Leaves (basal and lower cauline) spatulate to ovate or oblong-obovate, margins coarsely
toothed to incised, revolute, cauline reduced in size and narrower, 1-2-pinnatifid, upper cauline
commonly bracteate, linear and entire, veins impressed adaxially, basal and proximal blades 2-6 cm x
7-25(-30) mm, strigose to strigillose adaxially. hirsute to hispid-hirsute abaxially, eglandular; petioles
10-60 mm. Fruiting spikes (3-)5-15 in panicles, elongate and slender, 8-25 cm, rachis eglandular;
floral bracts ovate-lanceolate, 1.6-2.5 mm, shorter than the calyces. Calyces 2.2-3.2 mm, strigillose,
eglandular, lobes deltate to triangular, not connivent. Corollas bluish to lavender-blue or purple,

rarely white, tubes 3-4 mm, 0.8-1 mm longer than the calyx, limbs 4-5(-7) mm in diam. Nutlets

separating at maturity, 1.6-2 mm, commissural faces ending below the nutlet tips, with huliate to
papillate plates. 2n = 14.

Flow^ering Mar-Jun(-Oct). Rocky hillsides, prairies, pastures, fields, beaches, dunes, shell
banks, post oak, pine, cedar glades, oak-mesquite, mesquite. roadsides, disturbed sites; 5-200(-800)
m; Alabama, Arkansas, Florida, Georgia, Louisiana. Missouri, Mississippi, North Carolina,
Oklahoma. South Carolina, Texas; Mexico (Coahuila, Nuevo Leon. Tamaulipas, Sonora, Veracruz);
introduced in Australia.

Verbena halei is a distinctive North American species. With V. officinalis it stores closely
strigillose calyces, small corollas, and mostly glabrate to sparsely hirsute-strigose stems, but the two
taxa are continentally disjunct in native range and discontinuous in morphology. Verbena halei is
completely eglandular, it produces more numerous spikes often in congested panicles, and the leaves
are primarily basal and proximal and thicker with revolute margins and veins impressed adaxially.

Barber's submergence of Verbena halei into V. officinalis (1982) was based on her encounter
of specimens of the former misidentified as the latter. "These mis identifications were often justified
because the plants would really fit neither taxon because of some intermediate character exhibited by
the specimen. The problem of deciding which species a specimen belongs to would be solved with
the merger ..." (p. 452). In contrast to Barber's assessment, and in agreement with Perry (as noted
above). V. officinalis is as similar or more to V menthifolia than to V. halei (though probably not most
closely related to each other (see Nesom 2010). And because of the rarity of V. officinalis
(naturalized or otherwise) within the geographic range of V. halei, the two rarely if ever have the
opportunity even to hybridize, much less to intergrade. They can be separated by these contrasts.

1. Basal and lower cauline leaves persistent, relatively thick, large and spatulate, margins revolute,
cauline leaves quickly reduced in size distally and becoming linear-entire; rachis and calyces
eglandular Verbena halei

1. Basal leaves usually deciduous, relatively thin, margins not revolute, cauline leaves relatively even-
sized upwards or largest near midstem; rachis and calyces stipitate-glandular Verbena offidsialis

Moldenke (1963) cited collections of Verbena halei from Durango (Waterfall & Wallis
13371, TEX, = V. menthifolia]), Queretaro (Arsene 10242, MO! = V. menthifolia':), Sinaloa (Gentry
7011, ARIZ, = V. officinalis\, see comments above), and Chihuahua (Pringle s.n., 11 Apr 1887, not
seen). All other collections that I have seen from Chihuahua, Durango, Sinaloa, and Sonora identified
as Verbena halei by Barbel" and by Moldenke have instead been V. menthifolia. Collections identified
as V. halei from Arizona (SEINET 2009; Kearney & Peebles 1960; Moldenke 1963) either have
proved to be or apparently are V. menthifolia. I have not seen evidence that V. halei occurs further
west in the USA than Texas. The single collection upon which attribution of V. halei to the New
Mexico flora has been based (Otero Co., Jones 26229) is identified here as V. menthifolia (see
comments above). A collection from southeastern New Mexico previously identified as V.
menthifolia (Chaves Co., as cited by Peterson and David 1998, see comments above under V.
menthifolia) instead probably is V. halei, but I have not seen a voucher.

ACKNOWLEDGEMENTS

This study is based primarily on study of collections from ARIZ, BRIT-SMU, MO, and TEX-
LL. I'm grateful to TEX staff for facilitating a critical loan of specimens from ARIZ (to TEX). Deb
Trock and Rebecca Peters (CAS-DS), Jane Mygatt (UNM), Ken Heil (SJNM), and Rich Spellenberg
(NMC) provided information on various collections and Barbara Hellenthal (ND-G) provided
information on the holotype of Verbena leucanthemifolia. Alan Weakley (NCU) posed questions
about V. riparia that stimulated stronger documentation and more pointed comments. I am grateful to

Nataly O'Leary for providing the pre-print copy of the 2010 Verbena revision This research was
done in conjunction with preparation of the FNA treatment of Verbena and supported by the Flora of
North America Association.

LITERATURE CITED

Alfred, K.W. 2009. Flora Neomexicana I: The Vascular Plants of New Mexico. An annotated

checklist to the names of vascular plants, with synonymy and bibliography. Published by

Lulu.com.
Barber, S. 1982. Taxonomic studies in the Verbena striata complex (Verbenaceae). Syst. Bot. 7:

433-456.
House, H.D. 1924. Annotated list of the ferns and flowering plants of New York State. New York

State Mus. Bull. No. 254. University of the State of New York, Albany.
Gibson D.N. 1970. Verbenaceae. In P.C. Standley and L.O. Williams (eds.), Flora of Guatemala.

Fieldiana Bot. 24(9): 167-236.
Kartesz, J.T. 2010. Floristic synthesis of North America. A Synonymized Checklist and Atlas with

Biological Attributes for the Vascular Flora of the United States. Canada, and Greenland.

Ver. 1.0.3679.36568 (January 2010).
Kearney, T.H. and R.H. Peebles [and collaborators]. 1960. Arizona flora (Second Edition and

Supplement by J.T. Howell, E. McClintock, and collaborators). Univ. of California Press,

Berkeley.
Liogier, E.E. [Hno. Alain]. 1957. Flora de Cuba. Vol. IV. P. Fernandez y Cia, La Habana.
Liogier, AH. 1994. La Flora de la Espanola. VI. San Pedro de Macoris, R.D. Santo Domingo,

Republica Dominicana.
Martin, W.C. and C.R. Hutchins. 1981. A Flora of New Mexico, Vol. 2. J. Cramer, Vaduz.
Michael, P.W. 1997. Notes on Verbena officinalis s.s. and V. macrostachya (Verbenaceae) with new

combinations in two closely related taxa. Telopea 7: 293-297.
Moldenke, H.N. 1980. A sixth summary of the Verbenaceae, Avicenniaceae, Stilbaceae,

Chloanthaceae, Symphoremaceae, Nyctanthaceae, and Eriocaulaceae of the world as to valid

taxa. geographic distribution and synonymy. Phytologia Mem., Vol. 2. Plainfield, New

Moldenke, H.N. 1963 and 1964. Materials toward a monograph of the genus Verbena. Phytologia 9:

160-175. 1963 (V halei); 10: 138-145. 1964a (V menthifolia); 10: 194-236. 1964b (V

officinalis); 11: 100-103. 1964c (V riparia); 11: 251-261. 1965 (V supina).
Munir, A.A 2002. A taxonomic revision of the genus Verbena L. (Verbenaceae) in Australia. J.

Adelaide Bot. Gard. 20: 21-103.
Nesom, G.L. 2010. Infrageneric classification of Verbena (Verbenaceae). Phytoneuron 2010-11: 1-

15.
O'Leary, N. 2006. Typifications in Verbena (Verbenaceae). Darwiniana 44: 493-499.
O'Leary, N., M.E. Mulgura, & O. Morrone. 2010. Revision taxonomic de las especies del genero

Verbena (Verbenaceae). II, Serie Verbena. Ann. Missouri Bot. Garden, in press.
Perry. L.M. 1933. A revision of the north American species of Verbena. Ann. Mssouri Bot. Gard.

20: 239-363.
Peterson, R.S. and I.L. David. 1998. Plants of Bitter Lake National Wildlife Refuge, Chaves County,

New Mexico. New Mexico Naturalist's Notes 1: 1-23.
Rzedowski, J. y G. Calderon de Rzedowski (eds.). 1985. Flora Fanerogamica del Valle de Mexico.

Volumen II: Dicotyledoneae (Euphorbiaceae-Compositae). Instituto de Ecologia, Publ. 15.

Diseno Editorial: Myriam Cerda, Mexico, D.F.
SEINET. 2009. Southwest Environmental Information Network. Arizona State Univ. and partners.

<http://swbiodiversity.org/seinet/index.php>
Shreve, F. and I.L. Wiggins. 1964. Vegetation and flora of the Sonoran desert, Vol. II. Stanford

Univ. Press, Stanford, Calif.

Small, J.K. and M.A. Vail. 1893. Report of the botanical exploration of southwestern Virginia

during the season of 1 892. Mem. Torrey Bot. Club 5 : 93-1 71 .
USDA, NRCS. 2010. The PLANTS Database. National Plant Data Center, Baton Rouge, Louisiana.

<http://plants.usda.gov> Accessed April 2010.
Verdcourt, B. 1992. Verbenaceae. In R.M. Polhill, Fl. Trop. E. Africa 8: 1-155. A.A Balkema,

Rotter dam/Brookfield.
Yuan, Y-W. and R.G. Olmstead. 2008. Evolution and phylogenetic utility of the PHOT gene

duplicates in the Verbena complex (Verbenaceae): Dramatic intron size variation and

footprint of ancestral recombination. Amer. J. Bot. 95: 1166-1176.

Figure 1 . Holotype of Verbena madrensis Nesom.

Figure 2. Geographic distribution of Verbena madrensis Nesom.