Guy L. Nesom
2925 Hartwood Drive
Fort Worth, TX 76109, USA
Study of plants variously identified within the Glandularia bipinnatifida group indicates that
the USA entities are appropriately recognized within six species: G. bipinnatifida (Nutt.) Nutt, G.
wrightii (A Gray) Umber, G. chiricahuensis Umber, Glandularia latilobata (Perry) Nesom, comb,
et stat. nov, Glandularia longidentata (Perry) Nesom, comb, et stat. nov, and Glandularia pubera
(Greene) Nesom, comb. nov. These species are distinguished in a key, described, and mapped.
Summaries of typification are provided. Glandularia gooddingii is sometimes confused with these
species and its distinction is discussed.
KEY WORDS: Glandularia bipinnatifida, Verbenaceae, taxonomy
The taxonomy of Glandularia bipinnatifida (Nutt.) Nutt. and its close relatives has been
studied by Perry (1933), Moldenke (1962-1965), Umber (1979, 1980), Turner (1998, 1999), and
Turner and Powell (2005) but identification of these plants lias remained problematic. In connection
with the development of an overview and taxonomic treatment for the Flora of North America North
of Mexico (FNANM), I have reviewed the nomenclatural types and variation patterns in these plants
and offer yet another taxonomic perspective.
Concepts of species and application of names in the Glandularia bipinnatifida group have
varied both widely and subtly. Details regarding some of the recent usage are discussed under the
pertinent species. In general, Perry's study provided an excellent beginning but typological
identifications blurred species boundaries. Even Umber's treatment of this group seems to have been
marked by typology and over-inclusiveness, as he described segregates that I am not able to affirm
and he overrepresented the morphological and geographical extent of G. bipinnafida. Turner
nomenclaturally linked USA entities to a Mexican taxon, but my study indicates that the more
northern species are discrete in morphology and geography, thus not calling for formal taxonomic
connections with those further south.
To be complete, however, the present study would need to give detailed attention to the
central Mexico representatives of the group, which have been abundantly collected. Umber (1979)
and Peralta (1985) treated all of the Mexican plants of this group broadly as Glandularia (or Verbena)
bipinnatifida, but there are geographical elements among them. Verbena ciliata Benth.  =
Glandularia (Verbena) ciliata (Benth.) Botta is the oldest name among the Mexican plants of this
In the present study, which centers on plants of the FNANM flora area, I have attempted to
delimit taxa and their geographical distributions and then to find the correct names. An advantage of
the protracted nature of this study (begun in 2008), looking and then looking again at various sets of
specimens, is that discrete geographic boundaries have become more evident. Perception of ranges of
variability also underlies, at least in part, differences with earlier studies in interpretation of
Glandularia racemosa (Eggert) Umber is perhaps a closely related member of this group, but
its annual habit and small white corollas make it relatively simple to identify consistently.
Glandularia gooddingii (Briq.) Soibrig produces leaves that vary from unlobed with merely toothed
margins to leaves that are deqily dissected with linear lobes — the latter are sometimes confused with
plants of the species treated here. Glandularia racemosa is included in the key but is not among the
species described and mapped; identification of G. gooddingii is discussed in a separate section
KEY TO THE SPECIES
1. Calyces eglandular to sparsely glandular with sessile to subsessile glands.
2. Floral bracts shorter to longer than the calyces; corolla tubes 9-14 mm, limbs mostly 9-15 mm
in diam.; central USA Glandularia bipinnatifida
2. Floral bracts shorter than the calyces; corolla tubes 7-13, limbs 6-10(-12) mm in diam.; Arizona
and New Mexico.
3. Stems decumbent to ascending or ascending-erect, 12^10 cm; calyces 5-7 mm; corollas
purplish to blue or lavender, tubes 7-11, limbs 6-9 mm in diam.; habitats at 1300-2500 m
3. Stems ascending to erect, 20-80 cm; calyces 8-10 mm; corollas usually pink to purplish pink,
less commonly violet, tubes 10-13 mm, limbs 7-10(-12) mm in diam.; habitats at 2300-2800(-
3000) m Glandularia chiricahensis
1. Calyces densely stipitate-glandular.
4. Plants annual; corollas white, sometimes bluish tinged; corolla tubes 6-7.5 mm, limbs (5.5-)6-9
mm in diam GLANDULARIA RACEMOSA
4. Plants annual or perennial; corollas pink to purplish or blue; corolla tubes 7-15 mm, limbs 7-12
mm in diam.
5. Stems densely hirsute to pilose-hirsute or hirsutulous, at least some of the hairs deflexed;
corollas consistently bright pink to purplish pink and drying similarly, tubes 12-15 mm, limbs
8-12 mm in diam Glandularia pubera
5. Stems stiffly hirsute with hairs spreading at right angles; corollas usually drying purple,
purple to bluish or pinkish when fresh, tubes 7-12 mm, limbs 7-12 mm in diam.
6. Plants perennial; stems decumbent-ascending to ascending-erect or erect; leaves 1-2-
pinnatifid, midstem ovate to lanceolate-ovate in outline, 2— 4(— 5) cm; corolla tubes 8-12 mm;
western Texas to New Mexico and Colorado Glandularia wrightsi
6. Plants annual; stems procumbent to decumbent-ascending,; leaves 2-3-pinnatifid, midstem
broadly ovate to deltate in outline, 1.5-3(-4) cm; corolla tubes 7-9 mm; south Texas (Rio
Grande plains) Glandularia longidentata
GLANDULARIA BIPINNATIFIDA (Nutt.) Nutt, Trans. Amer. Philos. Soc, n.s. 5: 184. 1836.
Verbena bipinnatiflda Nutt., J. Acad. Nat. Sci. Philadelphia 2: 123. 1821. TYPE: USA.
Arkansas. [Red River], T. Nuttall s.n. (probable hoiotype: PH digital image!; isotype: GH).
Umber (1979) indicated that he saw an isotype at NY, but I did not find one there nor is one
included in the NY online type database.
Verbena demareei Moldenke, Amer. Midland Naturalist 24: 752. 1940. TYPE: USA. Arkansas.
Clark Co.: P.O. Okolona, chalk and marl ditches, 350 ft, 30 Jun 1938, D. Demaree 17198
(hoiotype: NY?; isotypes: GH, MO!, NY digital image!). The NY specimen that is pictured
online (NY 00138170) was annotated by Moldenke in 1941 as an isotype; Moldenke (1962,
p. 210), however, noted that the holotype is deposited at NY.
Plants perennial, sometimes annual. Stems prostrate, decumbent, ascending, or ascending-
erect, sometimes adventitiously rooting on the proximal decumbent portions, 5-30(-50) cm,
moderately to densely hirsute, eglandular. Leaves ovate to broadly ovate in outline, membranaceous,
blades 1.2-2 times longer than wide, 1-2-pinnatifid, 3-parted, or deeply incised, ultimate segments
linear to oblanceolate, midstem 1-6 cm. hispid-hirsute to strigose or hirtellous, eglandular; petioles 5-
15(-20) mm. Inflorescence elongating to 5-15(-20) cm in fruit; floral bracts shorter to longer than
the calyces. Calyces 7-10 mm, hispid and strigose. eglandular or sometimes sparsely stipitate-
glandulai. Corollas purple, lavender, or bluish, rarely white, tubes 9-14 mm, limbs mostly 9-15 mm
in diam. Nutlets cylindric, 2.5-3.2 mm, not broadened at the base, black, commissure not reaching
the apex, 1/3-1/2 as wide as the nutlet apex, apical appendage absent. 2n = 30.
Flowering Apr-Jun(-Oct). Prairies, ridges, bluffs, limestone outcrops, gypsum hillsides, low
woods, roadsides, fencerows; 50-600 m; Ala., Ark., Colo., Ga., *Ind., *Iowa. Kans., *Ky., La., Miss.,
Mo., Nebr., Okla., S.Dak., Tex., *Wis., Wyo.; Mexico (Coahuila). Map in Fig. 4. *=cultivated?
I have seen only two collections of Glandularia bipinnatifida sensu stricto from Mexico (as
mapped here on Fig. 4): Coahuila: Mpio. Villa Acuna, limestone hill near Santo Domingo, 3 Jul
1936, Wynd & Mueller 450 (MO); Mpio. Villa Acuna, Rancho El Rincon, on SW margin of Serranias
del Burro (part of Sierra del Carmen), ca. 80 km SE of Big Bend Natl. Park, Texas, 1400-2100 m, 22
Jun 1991, Ruiz 54 (TEX).
The Mssissippi-Alabama plants consistent!) hai e minutd\ stipibte-glandular calyces
(vouchers seen from Scott and Oktibbeha cos. — Miss; Autauga, Dallas, Greene, Hale, Lowndes,
/Marengo, Montgomery, Pickens, Sumter, Wilcox cos. — Ala). 1 have not seen a voucher for the
extreme eastern outlyer in Georgia (Houston Co., see Fig. 4; ranked SI by the Georgia Natural
Heritage Program), but presumably it is similar to those in Alabama and eastern Mssissippi. The
population system east of the Mssissippi River might be recognized at varietal rank, emphasizing its
°to nphi,al distuio'iun Kit the sj hse ^lindubi r* o ~uis poiudt^alK tin u^h ves *m populahons
and such a minor and quantitative distinction hardly seems to justify formal recognition.
Turner (1998, 1999) treated Glandularia bipinnatifida broadly to include several putatively
intergrading geographic varieties. Within his concept of G. bipinnatifida var. ciliata (Benth.) Turner
(= Glandularia ciliata) were three entities treated here as distinct species: G. longidentata, G.
wrightii, and G. pubera. In the concqrt here, Glandularia ciliata sensu stricto is a species of central
Mexico (southern Durango, Nayarit, and Jalisco eastward to San Luis Potosi, Hidalgo, Estado
Mexico, and Puebla) and does not appear to be genetically continuous with the more northern taxa.
Umber's concept of Glandularia bipinnatifida (1979, 1980) was even broader. Within it, he
included G ciliata sensu stricto and plants identified here as G, latilobata, G longidentata, and G.
wrightii. He was mostly unable to find any meaningful geographic trends in morphology, but he did,
however, find "chemical races"(based on flavonoid profiles) that bear some correspondence to taxa
recognized in the present study.
GLANDULARIA WRIGHTII (A. Gay) Umber, Syst. Bot. 4: 92. 1979. Verbena wrightii A Gray,
Synopt. Fl. N. Amer. 2: 337. 1878. TYPE: USA. Texas. [El Paso Co.: Franklin Mountains]
Mountains near Fronteras, 22 Mar 1852, C. Wright 1504 (holotype: GH! — Fig. 1 in the present
report; isotypes: F digital image!, NY! digital image!; probable isotype: MO!).
Verbena ambrosiifolia Rydb. ex Small, Fl. S.E. U.S., 1011, 1327. 1903. TYPE: USA. Colorado.
Otero Co.: Rocky Ford, 8 Tun 1900, G.E. Osterhout s.n. (holotype: NY!, digital image!).
Verbena wrightii forma albiflora Moldenke, Phytologia 11: 497. 1965. TYPE: USA. Texas. Brewster
Co.: Alpine, Sul Ross College campus, 2 Apr 1936, B.H. Warnock 44 (holotype: NY: isotype:
US digital image!).
Plants perennial. Stems decumbent-ascending to ascending-erect or erect, (15-)25-60 cm,
stiffly hirsute with hairs spreading at right angles, eglanduiar or very sparsely stipitate-glandular.
Leaves ovate to lanceolate-ovate in outline, membranaceous, blades 1.2-2(-2.5) times longer than
wide, 1-2-pinnatifid, ultimate segments mostly lanceolate to oblanceolate, midstem mostly 2-4(-5)
cm, hirsute-strigose, eglanduiar: petioles 4-15 mm. Inflorescence elongating to 2-5(-ll) cm, in
fruit; floral bracts shorter than to nearly equalling the calyces. Calyces 6-8 mm, sparsely to
moderately hirsute-pilose, densely minutely stipitate-glandular. Corollas mostly purple-pink, rose-
purple, purple, or lavender-blue, tubes 8-12 mm, limbs 7-12 mm in diam. Nutlets cylindric, 2.5-
3.2(-3.5) mm, slightly broadened at the base, black, commissure not reaching the apex, 1/3-1/2 as
wide as the nutlet apex, apical appendage absent. 2n = 30.
Flowering Apr-Sep. Creosote bush, creosote bush-acacia-mesquite, shrubland, juniper, pine-
juniper, pine-juniper-oak, limestone slopes, grasslands, roadsides, fencerows, rocky hillsides, lava
beds, gravelly banks, arroyos; 1300-2400 m; Colo., N.Mex., Tex.; Mexico (Chihuahua, Coahuila,
Durango, Nuevo Leon, Zacatecas). Glandularia wrightii occurs in Oklahoma — in Cimarron Co., in
the western tip of the panhandle. Map in Fig. 5.
In the interpretation here, the type specimen of Glandularia wrightii does not correspond to
the morphological concept adopted by Umber (1979) and by Turner and Powell (2005). What they
identified as G. wrightii is instead identified here as G. pubera. Measurements from the holotype of
G. wrightii (Fig. 1): calyces 7-8 mm long, lobes linear-filiform; corollas "light purple" (fide A Gray),
tubes 8-11 mm long, limbs ca. 8-10 mm in diam., lobes 2.5-3.5 mm wide; floral bracts and calyces
hirsute, densely stipitate-glandular; stems sparsely glandular, hirsute-hirsutulous, hairs spreading at
light angles, 0.2-0.9 mm long.
The southern boundary of Glandularia wrightii is not determined with certainty in the
present study. I have mapped the species as far south as northern Zacatecas (Fig. 5), where it
apparently becomes sympatic with G. ciliata, but this needs detailed study and documentation.
Turner saw only a single taxon, which he recognized as G. bipinnatifida var. ciliata; if his biological
interpretation is correct. Verbena ciliata Benth. would be the oldest name for the species.
Glandularia wrightii along the eastern edge of its range is parapatric to slightly sympatric
with typical G. bipinnatifida (Figs. 4, 5). The two taxa both a hexaploid and clearly are very similar
in morphology but the differences (especially the vestiture) as outlined in the couplet below are
consistent and sharply defined.
1. Inflorescence elongating to 2-5(-ll) cm, in fruit; calyces sparsely to moderately hirsute-pilose,
densely minutely stipitate-glandular; corolla tubes 8-12 mm, limbs 7-12 mm in diam.
1. Inflorescence elongating to 5-15(-20) cm in fruit; calyces hispid and strigose, eglanduiar or
sometimes sparsely stipixa*e-glandu!ar; corolla tubes 9-14 mm, limbs mostly 9-15 mm in diam.
Turner (1998, p. 5) observed that Glandularia bipinnatifida "is relatively uniform over most
of its distribution, but just west of the Pecos River in trans-Pecos Texas it appears to intergrade over a
relatively short distance in [G. wrightii]. ... Once the Pecos River is reached going eastward (only
ca. 15 mi or so), the populations of [G. wrightii] are replaced by [G. bipinnatifida]." In Pecos
County, where they are sympatric within about a 50-mile radius of Fort Stockton (west of the Pecos
River), intermediates have been noted between the two (Turner 97-33, 97-43, 97-79; all TEX); at
other sites in the same area, they appear to grow together without intemiediacy (e.g., Turner 24-53,
typical G. wrightii, observed to be growing with typical G. bipinnatifida). Even in view of these
hybrids in the narrow zone of sympatry/parapatry, the morphological distinction between the two
appears to be essentially discontinous and in justification of their treatment as separate species. Still,
it is a subjective judgement and their treatment as conspecific varieties also might be justified in view
of the hybridization.
GLANDULARIA LONGIDENTATA (L.M. Perry) Nesom, comb, et stat. nov. Verbena ciliata
Berth, var. longidentata L.M. Perry, Ann" Missouri Bot. Gard. 20: 331. 1933. TYPE:
Mexico. Tamaulipas. Matamoros, Apr 1836, J.L. Berlandier 3020 (holotype: MO!, digital
image!; isotypes: GH!, NY-digital image!).
Plants annual. Stems branching profusely from crown, procumbent to decumbent-
ascending, 1.5-3(^1) cm, pilose to hirsute or hispid-hirsute, eglandular. Leaves broadly ovate to
deltate in outline, membranaceous, blades 1.5-2 times longer than wide, 2-3 -pinna tifid, ultimate
lobes lanceolate to oblanceolate, midstem 1.5-3(^1) cm, hirsute to hirsute-strigose, eglandular, rarely
sparsely stipitate-glandular; petioles 4-10 mm. Inflorescence elongating to 2-5(-7) cm in fruit;
floral bracts shorter than the calyces. Calyces 6-7 mm, hispid-hirsute, densely minutely stipitate-
glandular. Corollas rose-iavender, blue-lavender, or purple to violet, tubes 7-9 mm, limbs 8-1 1 mm
in diam. Nutlets subcylindric, 2.2-3.2 mm, broadened at the base, black, commissure not reaching
the apex, 1/3-1/2 as wide as the nutlet apex, apical appendage absent. In = 30 (as inferred from
reports of counts of "G. bipinnatifida" from Duval and Hidalgo counties; Lewis & Oliver 196 1).
Flowering Dec- Apr. Fields, vacant lots, roadsides, ditch banks, low hills, low plains, thorn
scrub, salirie flats, sand, sandy loam, silty clay, caliche, marl; 2-60(-500) m; Texas; Mexico (Nuevo
Leon, Tamaulipas). Map in Fig. 6.
Glandularia longidentata is essentially parapatric with typical G. bipinnatifida (Figs. 6 and
4) but both species have been collected in Maverick, McMullen, Nueces, and San Patricio counties. I
have not seen an unequivocal intermediate, though occurrence of hybrids would not be unexpected.
The former is distinct in its annual duration, consistently decumbent-ascending stems and radiating
from the base (producing a bowl-shaped habit), shorter, relatively broader, and more dissected leaves,
and shorter corolla tubes.
Perry's concept of Verbena ciliata var. longidentata corresponds to the taxon treated here at
specific rank. Umber (1979) did not recognize it as distinct treating the name as a synonym of his
broadly defined Glandularia bipinnatifida. Turner (1998) mapped the population system of G.
longidentata as part of G. bipinnatifida var. ciliata, showing its distinct range in Texas — similar to
the distribution shown here in Fig. 6 and (also as interpreted here) similar in its essential parapatry
with typical G. bipinnatifida.
Collections examined from Mexico. Nuevo Leon. 2 mi N of Sabinas Hidalgo, sandy loam
bottom, 26 Mir 1944, Barkley et al. 14512 (MO); 16 km W of Sabinas Hidalgo, Villaldama road,
among large rocks, 7 Apr 1962, Dominguez & McCart 8263 (BRIT); 2 mi N of Sabinas Hidalgo,
sandy loam bottom. 26 Mar 1944, Heard et al. 14512 (TEX); Sabinas Hidalgo, 16 Aug 1937.
Kenoyer s.n. (MO); 12 mi W of Monterrey, rock and gravel in xerophytic canyon. 27 Feb 1944,
Painter et al. 14281 (TEX); 16 mi S of Nuevo Laredo on Hwy 85, 19 Mar 1967, Wilson 12167
(TEX). Tamaulipas. 22 mi S of Matamoros, black clay, 21 Mar 1964, Montemayor 31 (BRIT,
TEX); 22 mi S of Matamoros, Natl. Hwy 101, sandy soil, 21 Mar 1964, Serna 34 (BRIT).
GLANDULARIA LATILOBATA (L.M. Perry) Nesom, comb, et stat. nov. Verbena bipinnatifida
Nutt. var. latilobata L.M. Perry, Ann, Missouri Bot. Gard. 20: 325. 1933. Glandularia
bipinnatifida (Nutt.) Nutt. var. 'latilobata (L.M. Perry) B.L. Turner, Lundellia 2: 63. 1999.
TYPE: Mexico. Sonora. Between San Pedro and Fronteras, 20-24 Sep 1890, C.V. Hartman
906 (holotype: GH!).
Plants perennial. Stems decumbent to ascending or ascending-erect, 12-40 cm, hirsute and
hirtellous, eglandular or sparsely stipitate-glandular. Leaves ovate to obovate in outline,
membranaceous, blades 1.3-2 times longer than wide, 1-2-pinnatitid, ultimate lobes linear to
oblanceolate, midstem 2^1 cm, hirsute to strigose-hirsute, eglandular or very sparsely stipitate-
glandular; petiuL> 4-10 mm. Inflorescence elongating to 2-4(-6) cm in fruit; floral bracts shorter
than the calyces. Calyces 5-7 mm, hirsute and strigose, eglandular to sparsely sessile to subsessile
glandular. Corollas purplish to blue or lavender, tubes 7-11, limbs 6-9 mm in diam. Nutlets
subcylindric, 2.5-3.2, broadened at the base, black, commissure not reaching the apex, 1/3-1/2 as
wide as the nutlet apex, apical appendage absent. 2n = unknown.
Flowering (Apr-)Jun-Sep. Pinyon-juniper, pine-oak. oak, alluvial benches, roadsides; 1300-
1800(-2300) m; Ariz., N.Mex.; Mexico (Chihuahua, Durango. Sonora). Map in Fig. 7.
Calyces of Glandularia latilobata characteristically appeal eglandulai but cl< si inspection
may show the presence of minute, translucent, sessile to subsessile glands. Arizona plants with
■.tipim ■>! nds m 1 tentatively identified as G. latilobata should be reexamined for the possibility that
they are instead G. gooddingii with dissected leaves. In Socorro Co., New Mexico, where G
latilobata is sympatic with G wrightii (with densely stipitate-glandular calyces), the contrast in
vestiture usually provides good confidence in their distinction. The flowers of G wrightii also are
larger than those of G latilobata although the differences are overlapping.
Umber (1979) treated this entity among many synonyms of Glandularia bipinnatifida var.
pinnatifida, while Turner (1999) followed the earlier observation of Perry and recognized it as G.
bipinnatifida var. latilobata (although he broadened the concept by including G, chiricahuensis as a
synonym). With Perry and Turner, I agree that "latilobata" is distinct, but its distribution (Fig. 7) is
disjunct from that of typical G bipinnatifida. At the southern end of its distribution in Mexico, G
latilobata apparently is sympatric with the Mexican endemic G. ciliata.
Turner's rationale for treating Glandularia latilobata as conspeeific with G bipinnatifida was
explicit (p. 64): "This taxon is similar to [G. bipinnatifida] var. ciliata but lacks the glandular
pubescence of that taxon. It grades into var. ciliata to the south and southeast." His distribution map,
however, shows only a single intermediate symbol where their ranges contact, and my own
observations suggest that they do not intergrade to the extent that would indicate their appropriate
treatment as conspeeific.
GLANDULARIA CHIRICAHENSIS Umber, Syst. Bot. 4: 92. 1979. TYPE: USA. Arizona.
Cochise Co.: Chiricaliua Mountains, Rustler's Park, 19 Aug 1973, R. Umber 640 (holotype:
GH!; isotypes: ARIZ, MO!, NMC, NY, UC).
Verbena ambrosiifolia Rydb. ex Small forma eglandulosa L.M. Perry, Ann. Missouri Bot. Gard. 20:
331. 1933. TYPE: USA. New Mexico. Grant Co.: vicinity of Silver City, Fort Bayard, Santa
Rita, Fierro, the Mimbres Valley and East Canyon tributary to it, and on the G.O.S. Ranch, in
canyons within 10 mi of the ranch house, Hannover [1ft], 1 Aug 1911, J.M. Holzinger s.n.
wide, usually l-2(-3)-pinnatifid, ultimate segments lanceolate to oblanceolate, midstem 3.5-7 cm,
strigose on both surfaces; petioles 5-15 mm. Inflorescence elongating only slightly in fruit to 2-4(-
5) cm; floral bracts slightly shorter to longer than the calyces. Calyces 8-10 mm, hirsute and
stuguSL eglandniai oi 'e>s -oimnonh ^|Mi^d\ 'iisnuteK stipitate-glandular. Corollas usually pink to
purplish pink, less commonly violet, tubes 10-13 mm, limbs 8-11 mm in diam. Nutlets cylindric,
2.5-3.4 mm, not broadened at the base, black, commissure not reaching the apex, 1/3-1/2 as wide as
the nutlet apex, apical appendage absent. 2n = 20 (Umber 1979).
Flowering Jun-Sq>. Pine, pine-oak, pine-oak-birch, clearings, subaipine meadows, disturbed
areas; 2200-2800(-3000) m; Ariz, N.Mex. Map in Fig. 8,
Glandularia chiricahensis is a distinct species recognized by its relatively tall, erect stems,
short fruiting spikes, and eglandular calyces. The corollas commonly are relatively large and usually
bright pink, similar to the characteristic corolla color in G. pubera (sometimes also seen in G.
gooddingii. The flavonoid profiles of G. chiricahensis and G. pubera are essentially identical (Umber
1980) and they share a tetraploid chromosome number — perhaps evidence of close common
ancestry and distinction as a pair apart from the hexaploids of the group (G bipinnatifida, G wrightii,
G. longidentata, and probably G latilobata).
Glandularia chiricahensis apparently is restricted to relatively high-elevation habitats in
central to southeastern Arizona and adjacent New Mexico (Fig. 8). Additional collections examined
(see type of Verbena ambrosiifolia forma eglandulosa, cited above) from New Mexico: Grant Co.: S
end of the Black Range. Mimbres River, ca. 5500 ft. 20 Aug 1904 Metcalfe 1231 (TEX); Redstone
[ca. 20 km NNE of Silver City], 13 Aug 1895, Mulford 896 (MO); mts. near Pinos Altos [ca. 11 km
NNE of Silver City], 26 Jun 1936, Stewart s.n. (MO); just N of jet Hwy 90 and 61 at San Lorenzo,
juniper zone, 14 Aug 1973, Umber 625 (TEX). Hidalgo Co.: Peloncillo Mts, Clanton Canyon, ca. 2
mi below (east of) summit, Coronado Natl. Forest, 19 Aug 1952, Tucker 2441 (LL); 9 mi NW of jet
US 70 and US 90 on Hwy 90 Lordsburg, 15 Aug 1973, Umber 631 (TEX).
GLANDULARIA PUBERA (Greene) Nesom, comb. nov. Verbena pubera Greene, Pittonia 5: 136.
1903. Verbena ciliata Benth. var. pubera (Greene) L.M. Perry, .Ann. Missouri Bot. Gard. 20:
332. 1933. TYPE: USA. Texas. Jeff Davis Co.: Davis Mountains, 28 Apr 1902, S.M. Tracy
and F.S. Earle 33 7 (holotype: ND-G digital image!, Fig. 2 of the present report). Tracy &
Earle 162 has been cited or notated as the type, but the group of ''162" duplicates instead
constitutes a set of paratypes: Texas. Jeff Davis Co.: Davis Mts., 23 Apr 1902, Tracy &
Earle 162 (F digital image!, MO-2 sheets!, MO digital image!, ND-G digital image! Fig 2 of
present report, NY digital image!, US digital image!). Tracy & Earle 1 62a (Fig. 3 of present
report) also is a paratype.
Glandularia bipinnatifida (Nutt.) Nutt. var. brevispicata Umber, Syst. Bot. 4: 89. 1979. Verbena
bipinnatifida Nutt. var. brevispicata (Umber) Moldenke, Phytologia 45: 470. 1980. TYPE:
USA. New Mexico. Otero Co, : Mountain Park, yard of Trinity Baptist Church, corollas pink-
purple, flowers with an acrid smell, [no date], R. & J. Umber 607 (holotype: GH; isotypes:
ARIZ, MO! MO-digtal urn*, 1 NY-2 ^beet—iiutal i njges't This collection is the voucher
for a chromosome count of In = 30 (Umber 1979). At NY (00137539) and TEX (00375018
and 00375019) are specimens labeled by Umber as his collection 607 and annotated by him
as "isotype," but they have slightly different label information from the protologue and from
specimens with label data corresponding exactly to the protologue: "Abundant on the rocky,
somewhat sandy hillsides in Mountain Park, New Mexico. This collection just NE of Trinity
Baptist Church, not fragrant, viscid. Strong perennial." These variants should be regarded as
Plants strongly perennial. Stems erect to ascending-erect, 15-40 cm, densely hirsute to
pilose-hirsute or hirsutulous, at least some of the hairs prominently deflexed, eglandular. Leaves
ovate to obovate in outline, membranaceous, blades 1-2 times longer than wide, l-2(-3)-pinnatifid,
ultimate segments oblong-lanceolate to lanceolate, midstem 4-5 cm, hirsutulous to strigose-hirsute,
eglandular; petioles 3-7 mm. Inflorescence elongating to (2-)3-6 cm in fruit; floral bracts slightly
shorter than the calyces. Calyces 8-10 mm, softly hirtellous, densely minutely stipitate-glandular.
Corollas magenta to pink, tubes 12-15 mm, limbs 8-12 mm in diam. Nutlets subcylindric, 2.6-3.2
mm, slightly broadened at the base, black, commissure not reaching the apex, 1/2 as wide as the nutlet
apex, apical appendage absent. In = 20 (Turner & Powell 2005; Umber 1979, identified as G.
wrightii; Lewis & Oliver 1961, identified as G. wrightii); 2n = 30 (Umber 1979, identified as G.
bipinnatifida var. brevispicata).
Flowering Feb-Aug. Rocky hillsides, rolling grassland, open oak woodland, pine-juniper,
lava flows, road shouldei md ^ii' iaW^iS (S00-) 1400-2300 m; Ariz., N.Mex., Tex.; Mexico
(Chihuahua, Coahuila). Map in Fig. 9.
Glandularia pubera in the concept of the present study includes plants identified as G.
wrightii by Umber (1979, p. 92), who noted that he was "applying the name Glandularia wrightii to
those tetraploid populations [in trans-Pecos Texas] with an erect habit, compact spikes in flower and
in fruit, a viscid inflorescence, and soft grayish leaves which form an acute angle with the stem."
Turner and Powell's morphological concept of G wrightii (2005) W3h es«enijally the same as
Umber's but as observed here, that name has been incorrectly applied.
The description of Glandularia bipinnatifida var. brevispicata by Limber (1979) and his
concept of it also are a close fit to the trans-Pw <-><* 1 exas pi nils that Ik identified as G. wrightii (here
= G. pubera). Umber observed (pp. 90-91) that var. brevispicata is "Easily recognized by its
abbreviated congested spikes, short calyx teeth, viscid inflorescence, tall, erect or strongly ascending
habit, and pink-purple flowers with tubes that are at least twice as long as the calyx." He also noted
that "The fragrance of the flowers is not at all pleasant, unlike the usually sweet fragrance of the type
variety [here = G. wrightii].. . [but instead] reminiscent of an acrid chemical smell" — the fragrance
is not diagnostic, however, as the holotype was observed to have a. resinous smell while paratypes
from very near the type locality were without any fragrance (as observed by Umber). Umber's key
(p. 86) distinguished var. brevispicata by its corolla tubes 2-2.5 times longer than the calyx (vs. 1.5-2
times longer in G. pubera) and calyx lobes ca. 1 mm long (vs. 1,5-3.5 mm long in G. pubera), but
these differences are not consistent.
Umber noted (1979, p. 91) that Glandularia bipinnatifida var. brevispicata is "primarily
found in mountains and high plateaus of New Mexico, northeastern Arizona, and northern Mexico"
(Coahuila and Nuevo Leon on his Figure 7 of 1979; his Fig. 3 of 1980). Remarkably, he mapped the
popisl.ati.ons of G. wrightii (here identified as G. pubera) as interposed directly between the New
Mexico/Arizona populations and the Mexico populations of var. brevispicata. Umber's var.
brevispicata and his G wrightii are essentially identical in flavonoid profiles (Umber 1980), and the
present study interprets all of these plants as presenting a single evolutionary entity, identified here
as G pubera (Fig. 9).
Chromosome number of Glandularia pubera.
Turner (1998) included Glandularia pubera within his broad concept, of G. bipinnatifida var.
ciliata. Later, he and Powell (Turner & Powell 2005, p. 1660) recognized it as distinct, observing
that in trans-Pecos Texas it "often grows in close proximity to G. bipinnatifida [here = G. wrightii]
without showing signs of hybridization." Turner and Powell found that the two entities in that area
also are distinct in ploidy level, with G. pubera consistently tetraploid (2n = 20, 4 populations
counted from 3 counties), while G. wrightii is consistently hexaploid (2n 1T - 30. 18 populations
counted from 7 counties) — confirmed here from vouchers at TEX.
The voucher for a tetraploid count, 2n = 20, from Apache Co., Arizona (Lewis & Oliver
1961) was identified by Umber as as Glandularia bipinnatifida. It seems likely that this was a plant
of G. pubera in the present sense; I have not seen the voucher. The voucher for a tetraploid count, 2n
= 20, from Apache Co., Arizona (ENE of Springerville on US 60, 4 Aug 2001, Windham 2530, MO!)
is unequivocally a plant of Glandular ia pubera.
Two inconsistencies are noted. (1) The voucher for a hexaploid count (Brewster Co., Texas,
Turner 24-159, TEX; reported by Turner and Powell 2005) is a plant of typical Glandularia pubera.
A duplicate of 24-159 at SRSC, however, is G. wrightii in the present sense (fide Mike Powell, pers.
comm.) and more likely represents the species from which the chromosome sample originated. (2)
Umber reported the chromosome number of the type collection of G. bipinnatifida var. brevispicata
as hexaploid; this was the only count he reported from among the plants he identified as var.
brevispicata. If Umber's count is correct and correctly vouchered, it documents the only record of a
hexaploid within G. pubera, which otherwise apparently is composed of tetraploid populations.
Identification of Glandularia gooddingii.
Glandularia gooddingii in its typical form, with unlobed leaf margins, is easy to identify
correctly. The combination of short floral bracts, densely stipitate-glandular calyces, short calyx
lobes, and broad corolla limbs (10-14 mm in diam.) also is diagnostic. In Arizona, variants of G.
gooddingii with deeply divided leaves and linear ultimate segments often are confused in
identification with species of the G. bipinnatifida group — they are contrasted here, Glandularia
gooddingii is currently documented unequivocally from New Mexico by only one collection from
Hidalgo Co. (Nesom 2010).
1. Calyces egianduiar or nearly so.
2. Calyces 5-7 mm; corolla tubes 7-1 1, limbs 6-9 mm in diam., purplish to blue or lavender
2. Calyces 8-10 mm; corolla tubes 10—13 mm, limbs 8—1 1 mm in diam., usually pink to purplish
pink, less commonly violet Glandularis chiricahensis
1. Calyces prominently stipitate-glandular.
3. Calyces (5-)6-7(-8) mm; corolla tubes 8-10 mm, limbs 10-14 mm in diam.; leaves commonly
with 3 main divisions Glandularia gooddingii
3. Calyces 8-10 mm; corolla tubes 12-15 mm, limbs 8-12 mm in diam.; leaves usually pinnatifid
Glandularia gooddingii (Briq.) Soibrig, Madrono 15: 50. 1959. Verbena gooddingii Briq., Annuaire
Conserv. Jard. Bot. Geneve 10: 103. 1907, Verbena bipinnatifida var. gooddingii (Briq.)
Jepson, Fl. Calif. 3: 382. 1943. TYPE: USA. Nevada. [Lincoln Co.]: Kernan, Meadow
Valley Wash, sandy bottoms, 28 Apr 1902, L.N. Goodding 645 (holotype: G-DEL; isotypes:
F digital image!, GH, MO! digital image!, POM, RM LB digital image!).
Verbena gooddingii var. nepetifolia Tidestrom, Proc. Biol. Soc. Wash. 38: 15. 1925. TYPE: USA.
Nevada. Lincoln Co.: El Dorado Canon, near Colorado River, 600 m, 2 May 1919, /.
Tidestrom 8835 (holotype: US digital image!).
Plants perennial. Stems erect, basally ascending, 10^10 cm, pilose-hirsute, stipitate-
glandular at least on the distal half, uncommonly egianduiar. Leaves deltate to ovate or broadly ovate
in outline, membranaceous, blades 1.5-2 times longer than wide, incised to coarsely toothed with
deltate to triangular teeth, ultimate segments deltate to ovate or lanceolate, to deeply divided with
ultimate segments sublinear, midstem 2-5 cm, hirsute-strigose to strigose adaxially, hirsute abaxially,
eglandular to very sparsely stipitate-glandular; petioles 3-10 mm. Inflorescence mostly remaining
compact, elongating to 2-5(-8, -10) cm in fruit; floral bracts 1/2-3/4 as long as the calyces. Calyces
(5-)6-7(-8) mm, densely minutely stipitate-glandular, pilose-hirsute, lobes 0.5-1 mm, shallowly
deltate to triangular-deltate or triangular, usually apiculate. Corollas pink to purple or lavender, tubes
8-10 mm, limbs 10-14 mm in diam. Nutlets cylindric, 2.5-3(-3.5) mm, not broadened at the base,
black, commissure not reaching the apex, 1/3-1/2 as wide as the nutlet apex, apical appendage absent.
2n = 30.
Flowering (Feb-)Mar-Jun (-Dec, sporadically). Talus, tlats, hillsides, sandy ridges and
bottoms, washes, river banks, streambeds, desert grassland, rocky slopes, roadsides, canyon bottoms,
brittlebrush-tamarisk, oak, thornscrub, pinyon-juniper, ponderosa pine-juniper, oak-juniper-
liackberiy, oak chaparral, baccharis-mesquite-Chilopsis, oak-ash, Larrea-Yucca-Fouquieria-Opuntia,
Washingtonia; 400-1 500(-2000) m; Ariz., Calif., N.Mex., Nev., Utah; Mexico (Baja California,
I am grateful to staff at MO, SMU-BRIT, TEX-LL, and UNM where collections of
Glandularia were studied and annotated on site, to UNM for a loan of specimens (to TEX), to staff at
GH for sending specimens and digital images of critical specimens, to Barbara Hellenthal at ND for
sending digital images and observations on the type of G. pubera, and to Bill Weber for reviewing the
Colorado collections of Glandularia at COLO. I am grateful to Mike Powell for comments on the
manuscript and to Billie Turner for his continuing interest in Glandularia, his comments on the
manuscript, and his always-ready willingness to look from a different pei^p^tne I also tluui Sim
Henrickson for his opinions on Glandularia taxonomy and a copy of his preliminary manuscript and
notes (ca. 2008) dealing with the G, bipinnatifida complex — and look forward to the completion of
his study, which includes an assessment of the Mexican plants. Dr. Henrickson and I agree on
delimitation of some of the entities, but the nomenclature here does not correspond with what he has
proposed and the new names proposed here do not preempt those in his taxonomy. We disagree
variously (fide his preliminary manuscript) with respect to the rank at which taxa should be
recognized, choice of types, and the placement of synonyms. The present study has been supported in
part under contract from the Flora of North America Association, in conjunction with prq>aration of
the taxonomic treatment of Verbenaceae for FNANM.
COLO Specimen Database. 2010. Specimen Database of Colorado Vascular Plants. Univ. of
Colorado Museum of Natural History, Boulder.
<http : //cum useum . Colorado . edu/Research/Botany/Databases/search. php>
Lewis, W.H. and R.C. Oliver. 1961. Cytogeography and phytogeny of the North American species
of Verbena. Amer. J. Bot. 48: 638-643.
Moldenke, H.N. 1961. Verbenaceae. Pp. 13-87, in Flora of Texas, Vol. 3. Texas Research
Foundation, Renner, Texas.
Moldenke, H.N. 1962-1965. Materials toward, a monograph of the genus Verbena. Verbena
ambrosiifolia: (II) Phytologia 8: 149-152. 1962. Verbena bipinnatifida: (III) Phytologia 8:
207-216. 1962 and (IV) 8: 230-244. 1962. Verbena ciliata: (IX) Phytologia 9: 12-29. 1963.
Verbena demareei: (III) 8: 175-216. 1962. Verbena wrightii: (XXIX) Phytologia 11: 486-
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
Nesom, G.L. 2010 (to be published). Glandularia gooddingii (Verbenaceae) in New Mexico.
Peralta, B.L.C. 1985. Verbena L. Pp. 286-290 in J. Rzedowski and G.C. Rzedowski. Flora
Fanerogamica del Valle de Mexico, Vol. II, Dicotyledoneae (Euphorbiaceae-Compositae).
Publ. 15, Instituto de Ecologia, Mexico, D.F.
Perry, L.M. 1933. A revision of the North American species of Verbena. Ann. Missouri Bot. Gard.
Turner, B.L. 1998. Texas species of Glandularia. Lundellia 1: 3-16.
Turner, B.L. 1999. A reevaluation of the Mexican species of Glandularia (Verbenaceae). Lundellia
Turner, B.L. and A.M. Powell. 2005. Chromosome numbers of Glandularia (Verbenaceae) from
central and trans-Pecos Texas. Sida21: 1657-1661.
Umber, R.E. 1979. The genus Glandularia (Verbenaceae) in North America. Syst. Bot. 4: 72-102.
Umber, R.E. 1980. The flavonoid chemistry of Glandularia bipinnatifida (Verbenaceae) and allied
species. Amer. J. Bot. 67: 935-941.
Figure 1. Holotype of Glandularia (Verbena) wrightii, Wright 1504, GH.
Nesom: Glandularia bipinnatifida taxonomy 13
Figure 2. Holotype of Verbena pub era, Tracy & Earle 337, ND-Greene — top two plants. Paratype
of Verbena pubera, Tracy & Earle 162 — bottom plant.
: Glandularis bipinnatifida taxonomy 14
Figure 3. Paratype of Glandularia (Verbena) pubera, Tracy & Earle 162a, ND-Greene (see text).
Figure 4. Distribution of Glandularia bipinnatifida. Open circles are records from literature,
vouchers not seen.
Figure 5. Distribution of Glandularia wrightii. Open circles are from the COLO Specimen Database
(2010), vouchers not seen in the present study.
bipinnatifida taxonomy \ J
Figure 6. Distribution of Glondularia longidentata.
Nesom: Glandularis bipinnatifida taxonomy 18
>_W I ,
Figure 7. Distribution of Glandularia latilobata.
: Glandularis bipinnatifida taxonomy 19
Figure 8. Distribution of Glcindularia chiricahensi
Nesom: Glandularis bipinnatifida taxonomy 20
Figure 9. Distribution of Glandularia puber