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In the present work a revision of tribe Verbeneae for Peru is presented, descriptions and illustrations for each taxa are presented. In this country this tribe is represented by 16 taxa: four Glandularia species, five Junellia species, one Mulguraea species and four Verbena species and two varieties; six of these taxa are endemic to Peru. Three new combinations are presented: Glandularia cuneifolia, Junellia clavata and Verbena glabrata var. hayekii. Nine new synonyms are proposed, lectotypes are designated for Verbena cuneifolia, Verbena clavata and Verbena villifolia, and an epitype is designated for Verbena pogostoma.
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Accepted by Federico Luebert: 1 Mar. 2014; published: 26 Mar. 2014 121
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Copyright © 2014 Magnolia Press
Phytotaxa 163 (3): 121–148
www.mapress.com/phytotaxa/Article
http://dx.doi.org/10.11646/phytotaxa.163.3.1
Synopsis of tribe Verbeneae Dumortier (Verbenaceae) in Peru
NATALY O’LEARY* & MARIA EMA MÚLGURA
Instituto de Botánica Darwinion, Labardén 200, CC 22, B1642HYD San Isidro, Buenos Aires, Argentina.
*Email: noleary@darwin.edu.ar
Abstract
In the present work a revision of tribe Verbeneae for Peru is presented, descriptions and illustrations for each taxa are
presented. In this country this tribe is represented by 16 taxa: four Glandularia species, five Junellia species, one
Mulguraea species and four Ve r b e n a species and two varieties; six of these taxa are endemic to Peru. Three new
combinations are presented: Glandularia cuneifolia, Junellia clavata and Verbena glabrata var. hayekii. Nine new
synonyms are proposed, lectotypes are designated for Verbena cuneifolia, Verbena clavata and Verbena villifolia, and an
epitype is designated for Verbena pogostoma.
Key words: Glandularia, Junellia, Mulguraea, Peru, Verbena, Verbeneae, Verbenaceae
Resumen
En este trabajo se presenta una revisión de la tribu Verbeneae en Perú, con descripciones e ilustraciones para todos los
taxones presentes. En este país la tribu está representada por 16 taxones: cuatro especies de Glandularia, cinco especies
de Junellia, una especie de Mulguraea y cuatro especies de Verb e n a y dos variedades; seis de estos taxones son
endémicos de Perú. Se proponen tres nuevas combinaciones: Glandularia cuneifolia, Junellia clavata y Verbena
glabrata var. hayekii. Se presentan nueve nuevos sinónimos, lectotipos son designados para Verbena cuneifolia, Verbena
clavata y Verbena villifolia, y un epitipo para Verbena pogostoma.
Palabras clave: Glandularia, unellia, Mulguraea, Perú, Verbena, Verbeneae, Verbenaceae
Introduction
Peru’s flora has been studied widely by Macbride in his contributions between years 1936–1964, Verbenaceae
being treated in 1960 (Macbride 1960). The author mentions 36 species from the tribe Verbeneae Dumortier (1829:
22), 35 of them under genus Verbena L. (1753: 18), and monotypic Hierobotana Briquet (1895: 148). This last
genus was recently demonstrated to be an endemic genus from Ecuador, not present in Peru (O’Leary & Moroni,
unpubl. mscr.).
Brako & Zarucchi (1993) in the Catalogue of Peru mention ca. 27 species of Verbena, 7 Glandularia J. F.
Gmel. (1791[1792]: 886, 920), and 6 Junellia Moldenke (1940c: 392). Recently, Binder (2002) presented an
unpublished floristic revision of several genera of Verbenaceae in Peru, as a dissertation, in which the author
recognized 10 Verbena, 5 Glandularia and 3 Junellia species in Peru. However, the taxonomy of Verbenaceae has
been recently updated by several significant studies (Marx et al. 2010, Yuan et al. 2010). Molecular phylogenetic
analyses focused on tribe Verbeneae (O’Leary et al. 2009, Yuan & Olmstead 2008a,b) have resulted in a new
circumscription of some genera, and a robust phylogenetic backbone for Verbeneae has been estimated. The
approximately 175 species belonging to tribe Verbeneae have been often combined under Ve r be n a in early
treatments (Schauer 1847, Briquet 1895, Perry 1933, Troncoso 1974). Currently, tribe Verbeneae is integrated by 5
genera, Glandularia, Hierobotana, Junellia, Mulguraea N. O’Leary & P. Peralta in O’Leary et al. (2009: 782) and
O’LEARY & MÚLGURA122 Phytotaxa 163 (3) © 2014 Magnolia Press
Ver b e na , all of them represented in Peru except Hierobotana. All members of tribe Verbeneae can be distinguished
from the rest of the Verbenaceae taxa by the fruit divided into 4 cluses, being a fruit drupaceous, fleshy or if dry,
divided into 2 cluses, in the rest of the Verbenaceae species.
Most species from Verbeneae are found in temperate North or South America. This work presents a complete
synopsis of tribe Verbeneae in Peru.
Material and Methods
This synopsis is based on herbarium collections from the following herbaria: BA, BM, CONC, F, G, GH, K, LIL,
MO, NY, SI, TEX, US, USM (Thiers 2013). Flower measurements were taken from material rehydrated by boiling.
Fruit measurements were taken from dried specimens. The descriptive terminology of the inflorescences used here
is in accordance with Martínez et al. (1996), the morphological terms employed follow Hickey (1974). For the
description of pubescence the terms: strigose, hispid, villous, puberulous, or sericeous are used, following
Lawrence (1951). The term cluse is here employed to describe a one-seeded unit. In tribe Verbeneae the 4 cluses
are derived from a bicarpellate ovary that separates along both the medial and transverse plane of the ovary
(O’Leary et al.. 2012). Illustrations have been drawn by V. Dudás and F. Rojas, from SI. The distribution and
habitat of taxa were taken from the herbarium specimen labels.
Results
Sixteen taxa (species and varieties) of tribe Verbeneae are recognized for Peru: 4 Glandularia, 5 Junellia, one
Mulguraea and 6 Verbena. Six of these taxa are endemic to Peru, the presence of endemisms being probably linked
to the existence of the Andes as a barrier (León et al. 2006). The departments with the highest number of endemic
taxa are found among those with Andean slopes, and wide altitudinal and ecological ranges, similar to Peru's
overall floristic trends.
The analysis of the cluse base morphology has helped sustain the new combination here proposed from
Verbena clavata to Junellia clavata. The most important difference between genera Junellia and Mulguraea from
Ve r b e na and Glandularia lies on the cluse morphology, being narrowed at the base in the first two genera and
enlarged, with basal and transversal commissural fold, in the last two genera (O’Leary et al. 2012).
The length of the style in relation to the length of the ovary has been a diagnostic trait traditionally used to
distinguish Verbena from Glandularia (Schnack & Covas 1944, 1946, Schnack 1964, Troncoso 1974, Botta 1993).
The presence of a long style has helped sustain the combination of Glandularia cuneifolia (formerly Verb e na
cuneifolia). Finally, the identity of a Peruvian narrow endemic has been resolved in the combination at varietal
level of Verbena glabrata var. hayekii.
Taxonomic treatment
Key to the genera of tribe Verbeneae from Peru
1. Subshrubs, suffruticose or plate forming plants, narrowed cluse base, inflorescences generally unbranched (sometimes
with simple branching), base chromosome number x = 9 or x = 10 ..................................................................................... 2
- Herbs or suffruticose plants, enlarged cluse base, branched inflorescences (at times condensed and appearing simple), base
chromosome number x = 5 or x = 7 ...................................................................................................................................... 3
2. Anther connective tissue not surpassing the thecae................................................................................................... Junellia
- Anther connective tissue surpassing the thecae .................................................................................................... .Mulguraea
3. Style long, more than 3 times longer than ovary, mature calyx usually longer than fruit and with teeth generally contorted,
chromosome number x = 5 ................................................................................................................................. Glandularia
- Style brief, less than 3 times longer than ovary, mature calyx shorter than fruit and with teeth not contorted, chromosome
number x = 7............................................................................................................................................................... Verbe na
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SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
Key to the species of tribe Verbeneae, genera Ver b e n a , Glandularia, Junellia and Mulguraea from Peru.
1. Subshrubs, suffruticose or plate forming plants, narrowed cluse base .................................................................................. 2
- Herbs or suffruticose plants, enlarged cluse base................................................................................................................... 7
2. Anther connective tissue not surpassing thecae; if enlarged then with glandular tissue, sessile or pedicelated.................... 3
- Anther connective tissue surpassing thecae; never enlarged .......................................................................... M. arequipense
3. Pricking plants..................................................................................................................................................... J. juniperina
- Not pricking (unarmed) plants ............................................................................................................................................... 4
4. Plate forming plants, entire leaf blades, paucifloral florescences............................................................................ J. minima
- Subshrubs or suffruticose plants, non entire leaf blades, plurifloral florescences ................................................................. 5
5. Leaf blades trisected from the base.......................................................................................................................... J. clavata
- Leaf blades 3–5 sected or 3–5 parted, never from the base, occasionally bipinnatisected or bipinnatiparted ..................... 6
6. Floral bracts 7–9 mm long, notably longer than the calyx length ............................................................................ J. occulta
- Floral bracts 3–6.5 mm long, generally smaller than the calyx length, occasionally slightly longer ................ J. fasciculata
7. Style long, more than 3 times ovary length............................................................................................................................ 8
- Style short, less than 3 times ovary length ............................................................................................................................11
8. Stumpy suffruticose plants, less than 10 cm high .................................................................................................................. 9
- Erect suffruticose plants, more than 10 cm high................................................................................................................. .10
9. Trisect leaf blades, densely hispid, with glandular hairs..................................................................................... G. gynobasis
- Triparted leaf blades, densely strigose, no glandular hairs .............................................................................. G. microphylla
10. Leaf blade generally 3-lobed or 3-parted, calyx teeth 3 mm long ...................................................................... G. cuneifolia
- Leaf blade 3 or 5-parted, lobated or parted segments, calyx teeth less than 1 mm long ..................................... G. laciniata
11. Plants with dense hispid hairs.................................................................................................................................. V. hispida
- Subglabrous plants or with strigose or villous pubescence, never hispid hairs ................................................................... 12
12. Erect herbs or suffruticose plants ........................................................................................................................................ 13
- Prostrate suffruticose plants ................................................................................................................................................ 15
13. Subconical appearance of florescences.................................................................................................................. V. g la br a ta
- Cylindrical appearance of florescences............................................................................................................................... .14
14. Cuneate based leaves............................................................................................................................................... V. litoralis
- Subauriculated based leaves......................................................................................................... V. litoralis var. subglabrata
15. Leaf blades entire with incised dentate margin, sometimes shallowly divided blade, floral bract shorter than the calyx
length ................................................................................................................................................... V. glabrata var. hayekii
- Leaf blades deeply dentate margin, 3- parted blades, floral bract longer than or equal to the calyx length.......... V. villifolia
1. Glandularia J. F. Gmelin (1791: 886, 920)
Glandularia is an exclusively American genus, with nearly 84 species and 8 varieties growing from southern
United States of América to Guatemala, in the northern hemisphere, and from Brazil and Peru to southern
Argentina in the southern hemisphere. In Peru there are 4 species, one of these endemic.
1.1 Glandularia cuneifolia (Ruiz & Pav.) N. O’Leary & Múlgura, comb. nov. Fig. 1.
Basionym: Verbena cuneifolia Ruiz & Pavón (1798: 22) (not Verbena cuneifolia Rafinesque 1808: 360). Lectotype (here
designated):—PERU. Sine loc., 1778–79, H. Ruiz & J. Pavón s.n. (MA barcode 814974!, isotypes MA barcode 814975!,
O’LEARY & MÚLGURA124 Phytotaxa 163 (3) © 2014 Magnolia Press
MA barcode 814976!, B†, photo F neg. 17411!, possible isotypes J. Dombey, P barcode 650811!, P barcode 650810!, P
barcode 650809!, P barcode 752584!).
FIGURE 1. Glandularia cuneifolia (Ruiz & Pav.) N. O’Leary & Múlgura (from Von Rentzell 67, SI). A. Branch general aspect. B.
Detail of stem pubescence. C. Leaf, abaxial surface. D. Detail of leaf abaxial surface pubescence. E. Flower with floral bract. F.
Extended calyx, outer surface. G. Extended corolla, inner surface with androecia. H. Stamen. I. Gynoecia.
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SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
Erect or suffruticose plant, 50–1.5 m, stems densely hispid with dark headed glandular hairs; internodes 3–5 cm
long. Sessile leaves, blade 3.5–6.5 x 1–2 cm, generally 3-lobed to 3-parted, acute apex, cuneiform base, dentate
margin, strigose adaxial surface, abaxial surface densely hispid over venation, strigose with short dark headed
glandular hairs on the rest of the surface. Inflorescences in monobotrya or pleiobotrya with lateral paracladia
surpassing the principal florescence; rachis enlarged in fructification; basal internode 3–4 cm long. Floral bracts 6–
6.5 mm long, linear to narrow ovate, densely hispid with dark headed glandular hairs. Calyx 8.5–9 mm long,
narrow acute teeth 3 mm long, densely hispid glandular pubescence. Corolla purple colored, tube 10–15 mm long,
densely puberulous external surface and puberulous fauce. Superior pair of stamens unappendaged, style long, 7–8
mm long. Cluses 2.8–3.5 mm long, enlarged base with basal and transversal commissural fold, obtuse apex,
reticulate dorsal surface, verrucose ventral surface.
Common names:—“cara huanzuy”, “china-verbena”.
Distribution and habitat:—Endemic to Peru, found in the departments of La Libertad, Lambayeque and
Junín. It grows in open forests, slopes of cliffs and small mountains, 3200–3600 m.
Remarks:—Marticorena & Quezada (1985) mention this taxon to occur in Chile. However no specimen has
been found in local herbaria of CONC and SGO. Therefore G. cuneifolia is here considered to be restricted to Peru.
Glandularia cuneifolia is clearly recognized by its dark headed glandular hairs found on stems, leaves and floral
bracts, this distinguishes this species from the rest of the Glandularia taxa from Peru.
Specimens examined:—PERU. Junín: Entre Chamiseria y Acpalca, sine data, Soukup 3537 (CONC),
Huancayo, 8 July 1966, Von Rentzell 67 (SI). La Libertad: Otuzco, Camino La Piedra Chunga-Millvachaqui, 3
June 1990, Leiva 69 (SI), Otuzco, Shitahuara, al norte de Salpo, 11 June 1992, Leiva 588 (SI), Otuzco, cerro de los
Enamorados, 14 May 1991, Leiva 252 (SI). Lambayeque: Ferreñafe, 22 June 1986, Llata Quiroz 1953 (SI).
1.2 Glandularia gynobasis (Wedd.) N. O’Leary & P. Peralta (2007: 224). Fig. 2.
Verbena gynobasis Weddell (1857: 156). Lectotype (designated by O’Leary et al., Darwiniana 2013: 263):—PERU. Cordillera
de Tacora, camino de Tacna a La Paz, 4000 m, 1839–40, H. A. Weddell s.n. (P barcode 752582!, isolectotypes F barcode
74515F!, SI 76923!).
Stumpy suffruticose plant, up to 10 cm high, stems densely hispid with short glandular hairs; internodes 0.5–1 cm
long. Sessile verticilated leaves, blade 0.5–0.8 x 0.6–0.7 cm, trisected, acute apex, cuneate base, densely hispid
with glandular hairs on both surfaces. Inflorescences in monobotrya or pleiobotrya with lateral paracladia
surpassing or not the principal florescence, this 15–20 mm long; rachis enlarged in fructification; basal internode
4–5 cm long. Floral bracts 4.5–5 mm long, linear to filiform, densely hispid with glandular hairs. Calyx 6–7 mm
long, acute teeth 1–1.5 mm long, densely hispid with glandular pubescence. Corolla purple or pink colored, tube 15
mm long, densely puberulous external surface and puberulous fauce. Superior pair of stamens unappendaged, style
long, 8–12 mm long. Cluses 2–2.5 mm long, enlarged base with basal and transversal commissural fold, obtuse
apex, reticulate dorsal surface, verrucose ventral surface.
Complete synonymy and exsiccate see O’Leary & Peralta (2007: 224) and O’Leary et al. (2013: 263).
Distribution and habitat:—Glandularia gynobasis is found in the north of Chile (Region of Arica and
Parinacota), Bolivia and southern Peru. It grows between 1900 and 4000 m, on loose soils. This species is known
from Peru only from the type specimen.
Remarks:—This species is distinguished by its stumpy suffruticose habit, its small trisected leaf blades and its
densely hispid glandular pubescence.
1.3 Glandularia laciniata (L.) Schnack & Covas (1944: 475). Fig. 3.
Erinus laciniatus Linnaeus (1753: 630). Verbena laciniata (L.) Briquet (1904: 296). Lectoype (designated by O’Leary et al.,
Darwiniana 2013: 263):—Feuillée, Journ. Obs. Phys. Cotes Orient. 3: 35–36; iconography tab 25. 1725.
O’LEARY & MÚLGURA126 Phytotaxa 163 (3) © 2014 Magnolia Press
FIGURE 2. Glandularia gynobasis (Wedd.) N. O’Leary & P. Peralta (from Smith 12, Chile, Región Arica y Parinacota, Cerro
Llaillane, SI). A. Branch general aspect. B. Leaf, adaxial surface. C. Detail of leaf abaxial surface pubescence. D. Flower with floral
bract. E. Extended calyx, inner surface. F. Extended corolla, inner surface with androecia and gynoecia. G, H. Stamens. I. Cluse, dorsal
view. J. Cluse, ventral view.
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SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
FIGURE 3. Glandularia laciniata (L.) Schnack & Covas (from Quezada 164, Chile, Región Biobío, cerro Teta sur, CONC). A. Plant.
B. Detail of stem pubescence. C, Leaf, adaxial surface. D. Detail of leaf abaxial surface pubescence. E. Flower with floral bract. F.
Extended corolla, inner surface with androecia and gynoecia. G. Stamen. H. Cluse, lateral view.
Erect or suffruticose plant, 15–50 cm, stems densely hispid, internodes 30–50 mm long. Leaves petiolated, blade
1.5–3.5 x 2.5–3.5 cm, generally 3 or 5-parted, segments lobated or parted, acute apex, attenuate base, strigose
adaxial surface, abaxial surface strigose and hispid over venation. Inflorescences in monobotrya or pleiobotrya
with lateral paracladia surpassing the principal florescence; rachis enlarged in fructification; basal internode 1–5
O’LEARY & MÚLGURA128 Phytotaxa 163 (3) © 2014 Magnolia Press
cm long. Floral bracts 3–4 mm long, ovate, densely hispid or strigose with ciliate margin. Calyx 5–8 mm long,
acute teeth 0.7–1 mm long, hispid to hirsute, with glandular pubescence. Corolla pink, lilac to purple colored, tube
10–16 mm long, densely puberulous external surface and puberulous fauce. Superior pair of stamens with
glandular anther connective appendix, sessile or surpassing thecae, exceptionally protruding from the corolla limb,
style long 7–8 mm long. Cluses 2.5–3 mm long, enlarged base with basal and transversal commissural fold, obtuse
apex, reticulate dorsal surface, verrucose ventral surface.
Complete synonymy and additional exsiccate see O’Leary et al. (2013: 263–265).
Distribution and habitat:—Frequent in Chile and southern Peru.
Remarks:—This species is distinguished from the rest of the Glandularia taxa from Peru by its divided leaf
blades, 3 or 5–parted with segments lobated or parted.
Specimens examined:—PERU. Cuzco: Calca, Urco, January 1938, Vargas 959 (SI); Sicuaní, 6 February
1903, Hicken s.n. (SI 10518). Lima: Huarochirí, Chicla, 5 June 1949, Velarde Nuñez 1683 (SI); Rio Blanco, 17
May 1959, Lothar Diers 962 (SI).
1.4. Glandularia microphylla (Kunth) Cabrera (1957: 332). Fig. 4.
Verbena microphylla Kunth (1818: 133). Type:—ECUADOR. “Crescit in frigidis Regni Quitensis juxta urbem Nova
Riobamba, alt. 1482 hex.” A. Humboltd & A. Bonpland s.n. (holotype P barcode 650850!, isotypes F barcode 976559!, P
barcode 650851!).
Stumpy suffruticose plant, up to 10 cm high, stems densely hispid, woody based, internodes 0.5–1 cm long. Leaves
petiolated, blade 0.4–1.6 x 0.4–0.8 cm, ovate, triparted, entire or 2–3-lobate segments, obtuse apex, cuneiform
base, densely strigose adaxial and abaxial surfaces. Inflorescences in monobotrya, rachis enlarged in fructification,
basal internode 0.4–1 cm long. Floral bracts 2.5–3 mm long, ovate, strigose, ciliate margin. Calyx 5–7 mm long,
brief triangular acute teeth, strigose-hispid pubescence. Corolla white or lilac colored, tube 11–16 mm long,
scarcely puberulous external surface and puberulous fauce. Superior pair of stamens with glandular anther
connective appendix, surpassing thecae, generally slightly protruding from the corolla limb, style long 8–9 mm
long. Cluses 2.5–3 mm long, enlarged base with basal and transversal commissural fold, obtuse apex, reticulate
dorsal surface, verrucose ventral surface.
Complete synonymy and additional exsiccate see Múlgura et al. (2012: 59).
Distribution and habitat:—G. microphylla is found in northwestern Argentina, from the province of Jujuy to
Mendoza. It also grows in Bolivia, Ecuador and Peru. It is found on rocky and dry soils, between 2000 and 4600 m.
Remarks:—This species is distinguished by its stumpy suffruticose habit and its conspicuous glandular anther
appendixes, protruding from the corolla limb.
Specimens examined:—PERU. Ayacucho: Sucre, El Belén, 16 March 1950, Rea Clavijo 46 (SI). Cuzco:
Choccó, January 1937, sine legit, Universidad Cuzco 241 (SI).
2. Junellia Moldenke (1940c: 392), nom. cons.
Junellia is a South American genus represented by 37 species and 6 varieties, growing from Peru and
Bolivia to Argentina and Chile. In Peru there are 5 species, one of them endemic.
2.1. Junellia clavata (Ruiz & Pav.) N. O’Leary & Múlgura, comb. nov. Fig. 5.
Basionym: Glandularia clavata (Ruiz & Pavón 1798: 21) Botta in Botta et al. (1995: 128). Verbena clavata Ruiz & Pav.
Lectotype (designated here):—PERU. Sine loc., sine data, H. Ruiz s.n. (MA barcode 814980!, isolectotypes F barcode
74507F!, MA barcode 814979!).
= Verbena calcicola Walpers (1843: 378) syn. nov. Type:—PERU. “Peruvia in Pampa Grande de Arequipa”, Mar 1831, F. J . F.
Meyen s.n. (holotype B†, photo F neg. 17409!).
= Verbena fissa Hayek (1908: 165). Type:—PERU. Ancash: Cajatambo, Ocros, 3000 m sm, A. Weberbauer 2768 (holotype B†,
photo F neg. 17416!). Syn fide Brako & Zarucchi (1993: 1177).
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SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
= Verbena clavata f. albiflora Moldenke (1950: 278) syn. nov. Type:—PERU. Arequipa: between Nazca & Chala, 300–350 m
s.m., 7 Nov 1947, R. A. Ferreyra 2507 (holotype NY barcode 0138255!, isotypes F barcode 0074508F!, MO barcode
694794!).
= Verbena clavata var. casmensis Moldenke (1952: 58) syn. nov. Type:—PERU. Ancash: Santa, Lomas de Casma, 250–300 m,
9 Set 1950, R. A. Ferreyra 8031 (holotype US barcode 118682!).
Prostrate suffruticose plant, hispid stems, internodes 7–20 mm long. Sessile leaves, fasciculate, blade 8–15 x 15–20
mm, trisected from the base, occasionally segments divided, sericeous-strigose pubescence, somewhat fleshy
texture, acute apex, cuneate base, entire margin. Inflorescences in pleiobotrya, frondose paracladia, dense
plurifloral florescences, rachis enlarged in fructification, basal internode 0.5–1 cm long. Floral bracts 6–6.5 mm
long, linear, filiform, and densely strigose. Calyx 6–7 mm long, acute teeth 1–1.5 mm long, densely sericeous-
strigose pubescence. Corolla white to pink colored, tube 8–8.5 mm long, densely puberulous external surface and
puberulous fauce. Superior pair of stamens generally with exert glandular anther connective appendix, surpassing
thecae and protruding from the corolla limb, style 5.5–6 mm long. Cluses 2–2.5 mm long., narrowed base, obtuse
apex, reticulate dorsal surface.
FIGURE 4. Glandularia microphylla (Kunth) Cabrera (from Fabris 3601, Argentina, Jujuy, Valle Grande, LP). A. Plant. B. Leaves.
C. Flower with floral bract. D. Bract, external surface. E. Extended calyx, outer surface. F. Superior stamen with glandular appendix.
G. Cluse, ventral view and longitudinal cross section.
O’LEARY & MÚLGURA130 Phytotaxa 163 (3) © 2014 Magnolia Press
FIGURE 5. Junellia clavata (Ruiz & Pav.) N. O’Leary & Múlgura (from Dillon 4826, SI). A. Branch general aspect. B. Leaf, adaxial
surface. C. Flower with floral bract. D. Extended corolla, inner surface with androecia and gynoecia. E. Cluse, lateral view.
Distribution and habitat:—Endemic to southwestern Peru, from the departments of Ica and Arequipa,
between 350–3300 m, growing on sandy, rocky or clay soils.
Remarks:—Junellia clavata is distinguished by its 3-sected leaf blades and the presence of glandular
connective appendix, being together with the Argentine Patagonian species J. micrantha (Phil.) Moldenke, the only
two species of Junellia that have glandular appendix. This species is combined under Junellia as a result of the
presence of a narrowed cluse base, which is a typical trait of this genus.
In herb. MA there are two specimens collected by Ruiz, and both are type material of Verbena clavata. The
specimen with the code bar number 814980 is a better preserved material, and is here chosen as lectotype.
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SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
Specimens examined:—PERU. Arequipa: Islay, Lomas de Mollendo, 400 m, 15 October 1957, Angulo 2600
(SI), idem, August 1940, Vargas 2023 (SI), idem, 17 November 1986, Dillon 4826 (SI), idem, August-September
1932, Stafford 10 (K, SI), idem, September–October 1933, Stafford 300 (K, SI) Caravelé, Lomas de Pongo, Acaví,
350 m, 3 August 1947, Velarde Nuñez 526 (SI), idem, 22 August 1948, Velarde Nuñez 1481 (SI), Lomas de Jahuay,
entre Nazca y Chala, 25 October 1976, Ferreyra 18692 (USM), 11 October 1955, Ferreyra 11492 (USM), Entre
Nazca y Chala, 9 November 1952, Ferreyra 8792 (MO), Caravelé, Lomas de Jahuay, 25 October 1976, Ferreyra
18692 (MO). Ica: Nazca, Lomas de San Nicolás, 22 September 1958, Ferreyra 13394 (USM), Lomas de
Llangarina, 22 September 1958, Ferreyra 13403 (USM), Nazca, Lomas de Llangarina, 22 September 1958,
Ferreyra 13413 (MO, USM).
2.2. Junellia fasciculata (Benth.) N. O’Leary & P. Peralta in O’Leary et al. (2009: 783). Fig. 6.
Verbena fasciculata Bentham (1844: 153). Glandularia fasciculata (Benth.) P. M. Jørgensen (2005: 463)Type:—PERU. Lima:
“Huamantango”, G. W. Barclay s.n. (holotype K barcode 193646!).
= Verbena pogostoma Klotzsch ex Walpers (1845: 31) syn. nov.—Type:—“Verbena pogostoma Klotzsch, Hort. Berol., Oct.
1839” (holotype B, photo F neg. nr. 17440!). Epitype (here designated): PERU. Lima: Mounts near Chosica, 1900 m,
April 1910, A. Weberbawer 5333 (MO!, isoepitype F!).
= Verbena mathewsii Briquet (1907: 104) syn. nov. Type:—PERU. Lima: sine loc., A. Mathews 495 (holotype G barcode
00366378!, isotypes F barcode 0074523F!, K barcode 0470512!, K barcode 0470511!).
= Verbena occidentalis Moldenke (1940b: 479). syn. nov. Type:—PERU, sine loc., 1834, A. Mathews 498 (holotype NY
barcode 138300!).
= Verbena variabilis Moldenke (1944 : 164). Type:—PERU. Huancavelica, Castrovirreina, near Córdoba, 27 March 1942, R.
D. Metcalf 30255 (holotype US barcode 118744!; isotypes GH barcode 96121!, LIL barcode 1388!, MO barcode 357490!,
SI!, UC barcode 694968!).
= Verbena ferreyrae Moldenke (1950: 279) syn. nov. Type:—PERU. Ayacucho: provincia de Lucanas, Puquio, 3200–3300
msm, 19 March 1949, R. A. Ferreira 5491 (holotype NY barcode 138263!, isotypes F barcode 74513F!, SI 168342!, US
barcode 48318!).
= Verbena lucanensis Moldenke (1950: 279). Glandularia lucanensis (Moldenke) Botta in Botta et al. (1995: 128). Junellia
lucanensis (Moldenke) N. O’Leary & P. Peralta in O’Leary et al. (2009: 783). syn. nov. Type:—PERU. Ayacucho:
Lucanas, between Nazca and Puquio, 1500–2000 m s.m., 19 March 1949, R. Ferreyra 5493 (holotype NY barcode
138290!, isotypes F barcode 074522F!, MO!, SI 168344!, US barcode 048317!).
Suffruticose plant, 50–70 cm high, stems densely hirsute with glandular hairs, internodes 15–40 mm long. Sessile
leaves, fasciculate, blade 10–25 x 10 mm, 3–5 sected, never from the base, occasionally bipinnatisected, strigose
pubescence on both surfaces, acute or subobtuse apex, cuneate base, entire margin. Inflorescences in pleiobotrya,
long frondose paracladia, dense plurifloral florescences, rachis enlarged in fructification, basal internode 0.1–1.5
cm long. Floral bracts 3–6.5 mm long, generally smaller than the calyx length, occasionally slightly longer, narrow
ovate, densely hispid-strigose. Calyx 4–7 mm long, acute teeth 1 mm long, densely hispid-strigose pubescence.
Corolla lilac colored, tube 5–8 mm long, slightly puberulous external surface and puberulous fauce. Superior pair
of stamens unappendaged, style 2.5–3 mm long. Cluses 2–2.5 mm long., narrowed base, obtuse apex, reticulate
dorsal surface.
Additional exsiccate see O’Leary et al. (2011: 56).
Distribution:—This species is endemic to western Peru.
Remarks:—Junellia fasciculata is distinguished by its leaf blades 3–5 sected or 3–5 parted, never from the
base, occasionally bipinnatisected or bipinnatiparted and the inflorescences with pleiobotrya with proximal and
basal paracladia long and frondose.
Walpers (1845) described Verbena pogostoma based on a plant cultivated at herb. B, from seeds originating
from “Bonariensibus”. After the Second World War the only reference to this specimen is the photograph of
MacBride F neg. nr. 17440, where it can be read: “Verbena pogostoma Klotzsch, Hort. Berol., Oct. 1839”. This
photograph is here considered as part of the original material. No further material could be found related to this
specimen. Therefore, an epitype is here designated. The specimen from Weberbauer 5333 is here chosen as a
suitable epitype given its congruence with the description in the protologue.
O’LEARY & MÚLGURA132 Phytotaxa 163 (3) © 2014 Magnolia Press
Five new synonyms are here proposed, V. lucanensis and V. ferreyrae are both based on a type material
collected by Ferreira the same day, in the same place, almost at the same time: one Ferreira 5491 and the other
Ferreira 5493. Moldenke describes both taxa in 1950, but mentions nothing to differentiate them.
FIGURE 6. Junellia fasciculata (Benth.) N. O’Leary & P. Peralta (from Ferreyra 5493 (isotipo MO). A. Branch general aspect. B.
Leaf, adaxial surface. C. Detail of leaf abaxial surface pubescence. D. Flower with floral bract. E. Cluse, ventral view.
Phytotaxa 163 (3) © 2014 Magnolia Press 133
SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
V. occidentalis and V. mathewsii are both based on a Mathews specimen, collected both in Lima province,
numbers 495 and 498. Both are probably also part of a same population.
Specimens examined:—PERU. Ancash: Recuay, Bosque de Noqno, sine data, Cano 1717 (MO). Bolognesi,
31 October 1984, Sagástegui 12294 (SI). Ayacucho: Parinacochas, rio de Lomas, May 1911, Weberbauer 5749
(MO). Cajamarca: Chota, Pampa Grande, arriba de Chuyubamba, 6 August 1988, Sagástegui 14051 (SI). Ica:
Nazca, 52 km de Ruquio, 15 March 1971, Ellenberg 4915 (TEX). Lima: Huarachiri, Langa, July 1838, Barclay
2363 (BM, SI); Huarochiri, San José, 20 April 1962, Saunders 798 (K, SI).
2.3. Junellia juniperina (Lag.) Moldenke (1940c: 396). Fig. 7.
Verbena juniperina Lagasca y Segura (1816: 19). Neotype (designated by Peralta et al., Ann. Missouri Bot. Gard. 95: 361.
2008):ARGENTINA. Mendoza. San Carlos, camino a Laguna Diamante, 2000 m, 2 February 1950, A. Soriano 4057 (SI
barcode 03714!).
Suffruticose pricking plant, 30–70 cm high, internodes 5–10 mm long. Leaves dimorphic, macroblast leaves 3-
sected, segments linear, rigid, shooting apex; braquiblast leaves entire or 2–3 lobed to 2–3-sected, smaller.
Inflorescences in monobotrya, florescences 2–3 cm long. Floral bracts 5–7.5 x 1.5–2 mm, ovate, shooting apex,
strigose pubescence. Calyx 6.5–8 mm long, acute shooting apex teeth 1 mm long, strigose pubescence. Corolla
lilac colored, tube10–12.5 mm long, slightly puberulous external surface and glabrous fauce. Superior pair of
stamens unappendaged, style 6.5–8 mm long. Cluses 3.5–4 mm long., narrowed base, obtuse apex, reticulate dorsal
surface.
Complete synonymy and additional exsiccate see Múlgura et al. (2012: 95).
Distribution:—Junellia juniperina is found in southern Peru, Bolivia, north and central Chile and
northwestern Argentina.
Remarks:—This species is distinguished by its shooting apex leaves, which turns it into a pricking plant.
Specimens examined:—PERU. Arequipa: Pampa de los arrieros, sine data, Stafford 723 (K). Ayacucho:
Lucanas, entre Nazca y Pampa Galera, 9 April 1970, Tovar 6779 (SI), Coracora, Parinacochas, April 1970, Cerrate
4859 (SI). Huancavelica: Castrovirreina, near Córdova, sine data, Goodspeed 30268 (LIL). Moquegua: Mariscal
Nieto, between Torata and Carumas, 14 Feb. 1983, Dillon 3336 (F). Tacn a : Tarata, camino a Caro, sine data, La
Torre 1837 (USM).
2.4. Junellia minima (Meyen) Moldenke (1940a: 47). Fig. 8.
Verbena minima Meyen (1834: 451). Neotype (designated by Peralta et al., Ann. Missouri Bot. Gard. 95: 351. 2008):—“Pérou,
sur le gran plateau”, Weddell, H. A. 1871 Chlor. And. 2: 154–155, pl 62 B. Fig. 1E.
Plate forming plant, 1–5 cm high, woody base. Densely imbricate sessile leaves, blade 4–5 x 1–1.5 mm, linear to
narrow ovate, entire, connate base, mucronate apex, slightly strigose pubescence on both surfaces. Inflorescences
in monobotrya, paucifloral florescences, 2–3 flowered. Floral bracts 2.5–3.5 x 1.5–2.5 mm, ovate, acute to obtuse
apex, strigose pubescence. Calyx 2.5–3 mm long, triangular teeth 0.5 mm long, puberulous on the upper half,
glabrous on the lower half. Corolla white or lilac colored, tube 3.5–4.5(–5.5) mm long, glabrous external surface
and puberulous fauce. Superior pair of stamens unappendaged, style 2–2.5 mm long. Cluses 2–2.5 mm long.,
narrowed base, obtuse apex, reticulate dorsal surface.
Complete synonymy and additional exsiccate see Múlgura et al. (2012: 97).
Distribution:—Junellia minima is found in southern Peru, Bolivia, Chile and Argentina.
Remarks:—This species is distinguished by its plate forming habit and its paucifloral florescences.
Specimens examined:—PERU. Arequipa: Cailloma. Sobre Chivay, sine data, Weberbauer 6895 (F). sine
loc., March 1943, Sanderman 3855 (SI). Moquegua: Cordillera above Torata, Weberbauer 7472 (F). Puno:
Juliaca, Stafford 1136 (F). sine loc., 2 March 1971, Ellenberg 4716 (SI). Tacna: Tacna, Cordillera Volcán Tacara,
Ancara, April 1926, Werdermann 1120 (SI).
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FIGURE 7. Junellia juniperina (Lag.) Moldenke (B, from Jörgensen 1027, Argentina, Catamarca, SI; rest from Cabrera 30801,
Argentina, Tucumán, SI). A. Plant. B. Florescence. C. Floral bract. D. Extended calyx, outer surface. E. Extended corolla, inner
surface with androecia and gynoecia. F. Gynoecia. G. Stamen. H. Cluse, dorsal view. I. Cluse, ventral view.
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SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
FIGURE 8. Junellia minima (Meyen) Moldenke (from s.leg. s.n., Argentina, Jujuy, Humahuaca, SI 1496). A. Plant. B. Florescence.
C. Leaves. D. Floral bracts. E. Calyx. F. Corolla. G. Gynoecia. H. Cluse, lateral view. I. Stamen.
2.5. Junellia occulta (Moldenke) N. O’Leary & P. Peralta in O’Leary et al. (2009: 784). Fig. 9.
Verbena occulta Moldenke (1950: 280). Glandularia occulta (Moldenke) P. M. Jørgensen (2005: 463). Type:—PERU. La
Libertad: Prov. Bolívar, cerca al Nevado Cajamarquilla, 12 September 1946, R. Ferreyra 1298 (holotype NY barcode
138301!, isotype SI 168345!).
Suffruticose plant, 30–60 cm high, stems densely hirsute, internodes 5–30 mm long. Sessile leaves, fasciculate,
blade 18–25 x 18–20 mm, 3 sected, never from the base, segments 3–5 mm lat., occasionally bipinnatiparted,
strigose pubescence on adaxial surfaces, hispid on abaxial surface, with scare glandular hairs, acute apex, cuneate
base, entire margin. Inflorescences in monobotrya or pleiobotrya, frondose or bracteose paracladia, dense
plurifloral florescences, rachis enlarged in fructification, basal internode 1–3 cm long. Floral bracts 7–9 mm long,
notably longer than the calyx length, narrow ovate, densely strigose with some glandular hairs. Calyx 5–6 mm
long, acute teeth 0.6–0.8 mm long, strigose pubescence. Corolla lilac colored, tube 9–11 mm long, slightly
puberulous external surface and puberulous fauce. Superior pair of stamens unappendaged, style 4mm long. Cluses
2–2.5 mm long, narrowed base, obtuse apex, reticulate dorsal surface.
Complete synonymy and additional exsiccate see O’Leary et al. (2011: 63).
Distribution and habitat:—Endemic to northwestern Peru, it is found growing at 2625–4000 m.
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FIGURE 9. Junellia occulta (Moldenke) N. O’Leary & P. Peralta (from Sanchez Vega 2702, SI). A. Branch general aspect. B. Leaf,
adaxial surface. C. Detail of leaf abaxial surface pubescence. D. Flower with floral bract. E. Cluse, ventral view.
Remarks:—Junellia occulta is differentiated from the rest of the Junellia taxa from Peru because of its long
floral bracts, exceeding the length of the calyx.
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SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
Specimens examined:—PERU. Ancash: Along road to Pastobueno from Huamachuco-Trujillo road ca. 36
km from Huamachuco and 6 km from Pastobueno, sine data, Duncan 2640 (MO). Cajamarca: Caserío Huacataz,
20 km NE de Cajamarca, 21 November 1981, Sanchez Vega 2702 (MO, SI); Celendín, entre Cajamarca y Celendín,
monte bajo, sine data, Ferreyra 15020 (MO); Cumbemayo, August 1947, Velarde Nuñez 1592 (SI); camino de
Cajamarca a Celendín, October 1948, Scolnik 1304 (SI); entre Cajamarca y la Encañada, 17 August 1973, Sanchez
Vega 1226 (SI); Pampa de la Culebra, cerca de la Encañada, 17 June 1975, Sagástegui 8078a (SI). La Libertad:
sine loc., Goodspeed 10032 (G, SI); Otuzco, Cerro Sango, Montil-Shorey, 28 March 1991, Sagástegui 14423 (SI);
La Libertad. Near Nevado Cajamarquilla, province Bolivar, alt. 3200–3500 m., 12 September 1946, Ferreyra 1298
(NY). Pasco: Between San Rafael and Cerro de Pasco, alt. 3800–3900 m., 31 January 1950, Ferreyra 6586 (NY).
3. Mulguraea N. O’Leary & P. Peralta (2009: 782)
Mulguraea is a genus of 11 species and 2 varieties inhabiting arid zones in southern South America. In Peru there
is one species.
3.1. Mulguraea arequipense (Botta) N. O’Leary & P. Peralta in O’Leary et al. (2009: 782). Fig. 10.
Verbena arequipense Botta (1988: 237). Junellia arequipense (Botta) Botta (1989: 392). Dipyrena arequipensis (Botta)
Ravenna (2008: 43). Type:—PERU. Arequipa: encima de baños de Jesús, 23 April 1961, 2600–2700 m, A. Ferreyra
14261 (holotype SI barcode 3674!, isotype SGO barcode 04245!).
Subshrub, 120–180 cm high, stems hispid. Leaves dimorphic, macroblast leaves 8–20 x 3–7 mm, elliptic, sessile,
acute apex, cuneate base, adaxial surface strigose, abaxial surface hispid; braquiblast leaves similar to macroblast
leaves, smaller, sometimes obovate, obuse apex. Inflorescences in monobotrya, dense plurifloral florescences 10–
16 cm; rachis enlarged in fructification; basal internode 1–2 cm long. Floral bracts 2.5–4 x 0.5 mm, narrow ovate,
densely strigose. Calyx 4–5 mm long, triangular teeth 0.4–0.5 mm long, strigose pubescence. Corolla redish to
yellow colored, tube 13–15 mm long, slightly puberulous external upper surface and puberulous fauce. Superior
pair of stamens with anther connective tissue surpassing thecae, unappendaged, style 11 mm long. Cluses 2.5–3
mm long., narrowed base, obtuse apex, slightly developed lateral wings, reticulate dorsal surface.
Complete synonymy and additional exsiccate see Peralta et al. (2008: 374), under Junellia arequipense and Botta
(1988: 238), under Verbena arequipense.
Distribution and habitat:—Mulguraea arequipense is found in Peru and Chile, on rocky slopes, between
2200 and 4300 m.
Remarks:—This species is distinguished by its yellow to reddish colored corollas, with long tubes.
Specimens examined:—PERU. Arequipa: Arequipa: Estanquillo Ahogado, Chachani, 31 March 1973,
Arenas 84 (SI); 14 km S Arequipa, 14 September 1938, Goodspeed 22121 (SI). Cajamarca: Contumazá, Cruz
Gde., 3000 msm, Sagástegui et al. 6463 (SI). Lima: Cajatambo, cerca de Churin, 25 March 1976, Ferreyra 18706
(SI, USM). Tacna: Tarata, Tisaco, 31 March 1998, Cano 8312 (SI, USM).
4. Verbena Linnaeus (1753: 18).
Ve r b e na comprises 42 species and 10 varieties growing in temperate regions of America, from southern North
America to southern South America, some species have been naturalized in other parts of the world as Europe,
Asia, northern Africa and Australia. In Peru there are 6 taxa, 4 species and two varieties, two of these taxa are
endemic.
O’LEARY & MÚLGURA138 Phytotaxa 163 (3) © 2014 Magnolia Press
FIGURE 10. Mulguraea arequipense (Botta) N. O’Leary & P. Peralta (from Ferreyra 14261, type SI). A. Branch, general aspect. B.
Detail of part of rachis with flower. C. Extended calyx, schematic. D. Extended corolla, inner surface with androecia and gynoecia. E.
Stamens. F. Cluse, transverse cut, schematic.
4.1a. Verbena glabrata Kunth (1818: 276). Fig. 11.
Type:—[COLOMBIA.] “crescit in Andibus Novo-Granatensium prope Almaguer et Pasto, altura 1160–1350m”, A. von
Humboldt & A. Bonpland 2140 (holotype P barcode 00289949!).
Erect herb or suffruticose plant, 0.3–1 m, stems subglabrous to scarcely strigose, internodes 3–5 cm long. Sessile
leaves, blade 2–3 (–7.5) x 1–2.5 cm, entire, ovate-elliptic to obovate, acute apex, amplexicaul base, serrate margin,
principally towards the apex strigose adaxial surface, abaxial surface densely strigose over venation. Inflorescences
in bracteose pleiobotrya with lateral paracladia not surpassing the principal florescence, arranged in trimerous
paracladia; florescence of sub conical appearance, up to 8 cm long in fructification; rachis enlarged in
fructification; basal internode 3–4 cm long. Floral bracts 2–3.3 mm long, ovate, sub glabrous with some few
strigose hairs. Calyx 2–3 mm long, brief triangular teeth, scare strigose pubescence, sometimes few glandular hairs.
Corolla white, lilac colored, tube 3.5–5 mm long, densely puberulous external surface and puberulous fauce.
Superior pair of stamens unappendaged, style brief 1.5 mm long. Cluses 1.8–2 mm long, enlarged base with basal
and transversal commissural fold, obtuse apex, reticulate dorsal surface, verrucose ventral surface.
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SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
FIGURE 11. Verbena glabrata Kunth (from Sagástegui et al. 6463, SI) A. Plant. B. Detail of stem node with two leaves. C. Flower
with floral bract. D. Extended corolla, inner surface with androecia and gynoecia. E. Fruiting calyx. F. Cluse, lateral view.
Complete synonymy and additional exsiccate see O’Leary et al. (2007: 585).
Distribution and habitat:—Verbena glabrata is found from México to Peru, but it is scarcely collected. It
grows on clay, humid soils; it is found in moorlands of Ecuador and Peru, at 3450 m.
Remarks:—This species is easily recognized by the sub conical appearance of its florescences.
Specimens examined:—PERU. Arequipa: Baños de Jesus, 2600 m.s.m., 16 November 1957, Sagástegui &
Aguado 2603 (SI); Moquegua, Ilo, Lomas de Tacahuay, 400–700 m.s.m., FLSP 2191 (US). Cajamarca: Prov.
Cajamarca, hacienda Polloquito, 5 March 1967, Sanchez Vega 272 (SI), Valle de Cajamarca, Shultínm 5 December
1971, Sanchez Vega 853 (SI), Contumazá, Cruz Gde., 3000 m.s.m., 19 April 1967, Sagástegui et al. 6463 (SI).
Junín: Tarma, Palea, 2600–2800 m.s.m., 19 October 1947, Ve l ar d e 670 (SI). La Libertad: Prov. Otuzco, Los Tres
Shulgones, 4 June 1990, Leiva 76 (SI). Lima: Río Blanco, 3500–3600 m.s.m., 19 May 1959, Löthar Diers 982
(SI).
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4.1b. Verbena glabrata var. hayekii (Moldenke) N. O’Leary & Múlgura, comb. nov. Fig. 12.
Basionym: Verbena hayekii Moldenke (1946: 148). nov. nom. for Verbena procumbens Hayek (1908: 163) nom. illeg. homon.
Verbena procumbens Forsskål (1775: 10). Lectotype (designated by Macbride 1960: 622):—PERU. Junín: La Oroya, no
date, 3700–3800 m, A. Weberbauer 2573 (G barcode 366704!, isolectotype SI 160567!).
= Verbena weberbaueri Hayek (1908: 163). syn. nov. Type:—PERU. no date, A. Weberbauer 440 (holotype B† photo F neg.
17459!)
FIGURE 12. Verbena glabrata var. hayekii (Moldenke) N. O’Leary & Múlgura (from Hutchinson 4209, SI) A. Postrate branch,
general aspect. B. Leaf, abaxial surface.
Suffruticose prostrate plant, stems adpressed to the ground, rooting at nodes, internodes brief, 0.5–1 cm long;
strigose pubescence. Sessile leaves, blade 1–2.5 x 0.3–1 cm, entire, acute apex, cuneiform base, incised-dentate
margin, sometimes shallowly divided, strigose on both surfaces. Inflorescences in monobotrya or frondose
pleiobotrya; rachis brief; basal internode 0.5–2 cm long. Floral bracts 2–2.5mm long, linear to narrow ovate,
strigose pubescence. Calyx 2.5–3 mm long, brief acute teeth, strigose pubescence. Corolla lilac, or bluish-white
colored, tube 3–4 mm long, densely puberulous external surface and puberulous fauce. Superior pair of stamens
unappendaged, style brief, 1.5 mm long. Cluses 2 mm long, enlarged base with basal and transversal commissural
fold, obtuse apex, reticulate dorsal surface, verrucose ventral surface.
Distribution and habitat:—This variety is endemic to central Peru, found at high altitudes.
Remarks:—It differs from the typical variety by its prostrate habit, with short internodes 0.5–2 cm long,
rooting nodes and smaller leaves 1–2.5 cm long with leaf blades shallowly divided.
The study of the type material of this taxon, along with the photograph of the destroyed holotype of V.
weberbaueri, allow us to conclude both are the same taxon. Macbride (1960) mentions V. hayekii Moldenke to
occur in the Flora of Peru. However he also mentions Ve r b e n a aretioides Hayek as a synonym of this species,
which is not correct since the latter name is a synonym of Junellia aretioides.
Specimens examined:—PERU. Cuzco: Cuzco, dry bare slopes, 11000 ft., September 1933–May 1934,
Stafford s.n. (K, SI). Junín: Tarma, 15 km Oroya-Tarma rd, 25 December 1961, Saunders 696 (K, SI), Tarma, Rio
Mantaro, entre Oroya y Huancayo, 19 February 1964, Hutchinson 4209 (MO, SI).
4.2. Verbena hispida Ruiz & Pavón (1798: 22). Fig. 13.
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SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
Lectotype (designated by Munir, 2002: 47):—PERU. Sine loc., 1778–88, H. Ruiz & J. A. Pavón 1/34 (MA barcode 814981!,
isolectotypes, BM barcode 900954!, G, MA barcode 814982!, MA 814983!, P barcode 650840!, SI barcode 3924!).
FIGURE 13. Verbena hispida Ruiz & Pavón (from Burkart 30601, Argentina, Jujuy, SI) A. Plant, basal and apical part. B.
Florescence. C. Floral bract, outer surface. D. Flower. E. Gynoecia. F. Fruiting calyx. G. Cluse, lateral view.
O’LEARY & MÚLGURA142 Phytotaxa 163 (3) © 2014 Magnolia Press
Herb or suffruticose plant, 0.3–0.8 m, stems decumbent at the base, erect at the apex, densely hispid. Sessile leaves,
blade 3–6(–10) x 1.5–2.5 cm, entire, ovate-elliptic, sometimes slightly trifid at the base, acute or obtuse apex,
amplexicaul to cuneate base, irregularly serrate margin principally towards the apex, strigose adaxial surface,
abaxial surface densely hispid, with glandular hairs on both surfaces. Inflorescences in bracteose pleiobotrya with
lateral paracladia surpassing the principal florescence, florescence up to 15 cm long in fructification, subtended by
two foliaceous bracts at the base, rachis enlarged in fructification. Floral bracts 3–4.5 mm long, elliptic to ovate,
hispid with glandular hairs. Calyx 2–3.5 mm long, brief triangular teeth, hispid pubescence with glandular hairs.
Corolla white, purple or lilac colored, tube 4–5 mm long, densely puberulous external surface and puberulous
fauce. Superior pair of stamens unappendaged, style brief, 1.5 mm long. Cluses 1.5–2 mm long, enlarged base with
basal and transversal commissural fold, obtuse apex, reticulate dorsal surface, verrucose ventral surface.
Complete synonymy and additional exsiccate see O’Leary et al. (2007: 587) and Botta & Cabrera (1993: 92,
fig. 41).
Distribution and habitat:—Verbena hispida grows in northern Argentina, Paraguay, Chile, Bolivia, and Peru.
It is found in marginal forests, bushes and stubbles, along routes, preferably in lowlands with swampy and clay
soils.
Remarks:—This species is distinguished by its densely hispid glandular pubescence.
Specimens examined:—PERU. Cuzco: Sicuari, 3550 m.s.m., 6 February 1903, Hicken s.n. (SI 10519);
Yucay, Soukup 259 (BA). Cajamarca: Contumazá, Teterillas-Marín, Guzmango, Sagástegui 10377 (NY). Lima:
Lima, Sta. Eulalia, 17 July 1949, Velarde 1664 (SI). Puno: Chunto, September 1965, Anibal Vera s.n. (SI). Tacna:
Tarata, Ticaco, 3600–4000 m.s.m., Cano 8335 (US).
4.3a. Verbena litoralis Kunth (1818: 276). Fig. 14.
Verbena bonariensis var. litoralis (Kunth) Gillies & Hooker ex Hooker (1829: 166). Lectotype (designated by Macbride, 1960:
624):—[PERU.] Trujillo, A. Bonpland s.n. (P barcode 500760!, isolectotype SI 161612!).
Herb or suffruticose plant, 0.8–3 m, erect stems, subglabrous to slightly strigose. Sessile to shortly petiolated
leaves, blade 5–12 x 1–3 cm, entire, ovate-elliptic or obovate, acute or obtuse apex, cuneate base, irregularly
serrate margin to sometimes almost entire margin, subglabrous to strigose on both surfaces. Inflorescences in
bracteose pleiobotrya with lateral paracladia surpassing the principal florescence, florescence cylindrical, up to 18
cm long in fructification, rachis enlarged in fructification. Floral bracts 1.5–4 mm long, ovate, subglabrous. Calyx
2–3.5 mm long, brief triangular teeth, strigose. Corolla white, purple or lilac colored, tube 3–5 mm long, densely
puberulous external surface and puberulous fauce. Superior pair of stamens unappendaged, style brief, 1.5 mm
long. Cluses 1.5–2 mm long, enlarged base with basal and transversal commissural fold, obtuse apex, reticulate
dorsal surface, verrucose ventral surface.
Complete synonymy and additional exsiccate see O’Leary et al. (2007: 593) and Múlgura et al. (2012: 196).
Distribution and habitat:—Verbena litoralis is the most frequent Ve r be n a species. It is distributed all along
America, in Argentina, Chile, eastern Paraguay, Uruguay, southern Brazil, Bolivia, Peru, Colombia, Venezuela,
Honduras and Mexico. It grows near the water, in swampy lowlands, calcareous, sandy soils, along bushes, scrubs,
grasslands, meadows or open areas and disturbed forests, edges of forests, waysides and rocky cliffs.
Remarks:—This species is distinguished by its erect habit, reaching up to 3 m height.
Specimens examined:—PERU. La Libertad: Prov. Trujillo El Cortijo, 3 April 1969, Sagastegui 7192 (SI).
4.3b. Verbena litoralis var. subglabrata (Moldenke) N. O’Leary & Múlgura in Múlgura et al. (2012: 198). Fig. 15.
Verbena brasiliensis Vellozo (1825 [1829]: 17) var. subglabrata Moldenke (1950: 278). Type:—CHILE. San Fernando,
Colchagua, Feb. 1930, E. Barros 8050 (holotype NY barcode 138247!, isotype SI 160594!).
Herb or suffruticose plant, up to 3 m, erect stems, subglabrous to slightly strigose. Sessile leaves, blade 5–10 x 1–3
cm, entire, elliptic or obovate, acute or obtuse apex, subauriculated base, irregularly serrate margin, subglabrous to
strigose on both surfaces. Inflorescences in bracteose pleiobotrya with lateral paracladia surpassing the principal
florescence, florescence up to 10 cm long in fructification, cylindrical, rachis enlarged in fructification. Floral
Phytotaxa 163 (3) © 2014 Magnolia Press 143
SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
bracts 2–3 mm long, ovate, subglabrous. Calyx 2–3.5 mm long, brief triangular teeth, strigose. Corolla white,
purple or lilac colored, tube 3–4 mm long, densely puberulous external surface and puberulous fauce. Superior pair
of stamens unappendaged, style brief, 1.5 mm long. Cluses 1.5–2 mm long, enlarged base with basal and
transversal commissural fold, obtuse apex, reticulate dorsal surface, verrucose ventral surface.
FIGURE 14. Verbena litoralis Kunth var. litoralis (from Burkart 8306, Argentina, Entre Ríos, SI). A. Plant, basal and apical part. B.
Floral bracts, outer and lateral view. C. Extended calyx, internal view from apical part. D. Flower with floral bract. E. Gynoecia. F.
Fruiting calyx. G. Fruit. H. Cluses, ventral and dorsal view.
O’LEARY & MÚLGURA144 Phytotaxa 163 (3) © 2014 Magnolia Press
FIGURE 15. Verbena litoralis var. subglabrata (Moldenke) N. O’Leary & Múlgura (from Rojas 1877, Paraguay, Col. Nva. Germania,
SI). A. Plant, basal and apical part. B. Flower with floral bract.
Phytotaxa 163 (3) © 2014 Magnolia Press 145
SYNOPSIS OF TRIBE VERBENEAE (VERBENACEAE) IN PERU
Complete synonymy and additional exsiccate see Múlgura et al. (2012: 198).
Distribution and habitat:—This variety grows in Argentina, Chile, Bolivia, Paraguay, Peru and southern
Brazil. It has been naturalized in United States of America, being found in the states of Texas, South Carolina and
Massachusetts. It grows on dry soils, stony, salty or too sandy, disturbed areas,
Specimens examined:—PERU. Cuzco: 3350 m.s.m., May, Herrera 17 (SI).
4.4. Verbena villifolia Hayek (1908: 164). Fig. 16.
Type:—PERU. Dep. Junin., prov. Farma, La Oroya , 3700–3800 msm, no date, A. Weberbauer 2582 (holotype B, lectotype
[here designated] G!).
FIGURE 16. Verbena villifolia Hayek (A, from Macbride 942, GH; B, from Pearce s.n., SI; C–F, from Smith 7484, SI). A. Branch. B.
Leaf, abaxial surface. C. Flower with floral bract. D. Extended corolla, inner surface with androecia and gynoecia. E. Fructiferous
calyx with floral bract. F. Cluse, lateral view.
O’LEARY & MÚLGURA146 Phytotaxa 163 (3) © 2014 Magnolia Press
Suffruticose prostrate plant, stems densely villous; rooting at nodes, internodes brief, 0.5–1 cm long. Sessile leaves,
blade 0.5–1.5 x 0.3–1 cm, 3-parted, the middle lobe 3-lobed at the apex, lateral lobes 2–3 lobed, acute apex,
cuneiform base, deeply dentate margin, villous adaxial surface, abaxial surface densely hispid over venation,
villous on the rest of the surface. Inflorescences in monobotrya or frondose pleiobotrya; rachis brief; basal
internode 0.5–1 cm long. Floral bracts 4–6 mm long, linear to narrow ovate, densely villous-strigose. Calyx 3.5–4
mm long, brief acute teeth, densely villous-strigose pubescence. Corolla rose, violet or white colored, tube 5–6 mm
long, strigose external surface and puberulous fauce. Superior pair of stamens unappendaged, style brief, 3 mm
long. Cluses 2.5 mm long, enlarged base with basal and transversal commissural fold, obtuse apex, reticulate dorsal
surface, verrucose ventral surface.
Distribution and habitat:—It is endemic to Peru.
Remarks:—This species is distinguished by its prostrate habit, its densely villous pubescence and its long
floral bracts.
The holotype of V. villifolia was housed at B and is now destroyed, but there exists a photograph of it (F neg.
nr. 17458). A lectotype is here designated from an isotype found at . G.
Specimens examined:—PERU. Ayacucho: Andes de Huanta, 13–14 February 1867, Pearce s.n. (K, SI).
Cajamarca: Jose C. Mariátegui, 4 June 1984, Smith & Sanchez Vega 7484 (SI). Junín: La Oroya, 1922, Macbride
& Featherstone 942 (GH).
Acknowledgments
This work has received financial support from CONICET PIP 112-200801-00177 to the authors. A special thank is
given to curators of herbaria that helped with the search of specimens, especially to Tom Wendt, from TEX, whose
work was of great help, as well as to Francisco Rojas for the illustrations.
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... Most of the studies cited above did not include any of the several Andean species of Verbeneae ( O'Leary et al., 2007 ;Peralta et al., 2008 ;O'Leary, 2014 ;O'Leary and Múlgura, 2014 ), except for the inclusion of the Ecuadorian monotypic Hierobotana in the analysis of Marx et al. (2010) . Th at study, based on plastid DNA, uncovered a tantalizing placement of this northern Andean endemic as sister to the clade of North American Verbena , but included fewer species of tribe Verbeneae than the earlier studies. ...
... In contrast to Glandularia , Junellia , and Mulguraea , Verbena is distributed throughout the Andes, albeit with diminishing diversity to the north ( O'Leary et al., 2007 ;Peralta et al., 2008 ;O'Leary and Múlgura, 2014 ). Th ough never very abundant or diverse in any particular region, Verbena is distributed mostly in the more xeric habitats in the high puna of the southern Andes, arid Andean foothills, and dry inter-Andean valleys. ...
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... Until now, common verbena has been analyzed in the botanical literature mostly from a taxonomic point of view (Martínez et al. 1996;O'Leary and Múlgura 2014) and as a medicinal plant (Cao et.al 2012;Deepak and Handa 2000a, b;Lai et al. 2006;Liu et al. 2012;Makino et al. 2009,). The present report reveals the additional virtues of the species. ...
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... Peru and Ecuador are known centers of endemism for Verbenaceae (O'Leary and Múlgura 2014;O'Leary and Moroni 2014). Andean geography has likely resulted in niche differentiation and speciation, functioning as a migration corridor for North American Verbenaceae derived from South American species (Frost et al. 2017). ...
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A new species of Citharexylum from the foothills of inter-Andean dry valleys in Peru, Citharexylum peruvianum, is described and illustrated herein. It is distinguished from other closely related species with caulinar spines (C. andinum, C. flexuosum, C. herrerae, C. montevidense, and C. weberbaueri), by inflorescence morphology, leaf size, and pubescence. The newly defined taxon has 15-40 flowered racemes, rachis 7-12 cm long, and coriaceous leaves, with the leaf blade being 4-6(8) × 2-3 cm and densely hirsute on the abaxial surface. A brief discussion on closely related species is provided. Citharexylum peruvianum is known exclusively from northwestern Peru, from the departments of Cajamarca, Lambayeque, and La Libertad. The species grows in the Andean foothills at mid-elevations between 1400 and 3000 m a. s. l., in rocky soils and on steep slopes with thorny scrubland. It is occasionally found in moist areas. This species has a restricted distribution and limited numbers of individuals, which will likely result in threatened status following formal review.
... We present the full list, following an ordinal arrangement. (Wood, 2008;O'Leary & Múlgura, 2014; even if they were not included in the treatments of their respective families for Peruvian endemics cf. León, 2007 andGranda, 2007), which may be eligible for conservation and management efforts. ...
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A full catalogue of all vascular plants found in the Gayalopo archaeological site, in Arequipa province, Arequipa department, southern Peru, is provided. Some context regarding the Gayalopo petroglyphs and its current state of conservation is provided. Further remarks on en-demism and threat levels for some of the found species is given. It is concluded that this and further studies will promote the conservation of the place.
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The catalog describes 1362 species of ferns and flowering plants for the department of Arequipa, more than 200 species more as reported in the last systematic inventory by Quipuscoa, Dillon, & Ortíz in 2006. Ninety-five species are mentioned for the first time for Arequipa and 26 species mentioned in the literature for the department were excluded. In addition to a brief description of the species, the catalog includes information on their ecology, distribution, and human use. In addition to information on systematics and phylogeny, the most important synonyms are listed in the appendix.
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Background and aims-The last comprehensive study that estimated the number of Verbenaceae genera and species was published in 2004, and included 34 genera and around 1200 species. Since then, several publications based on morphology and/or molecular data have proposed important changes within the family. Due to the lack of updated literature to cite when referring to the number of Verbenaceae taxa, a review of these estimates is necessary. Key results and conclusion-We present a detailed list of genera currently accepted in Verbenaceae with the number of species contained in each and compare our numbers with the previous estimate. In addition, we indicate the geographic distribution and the most recent important taxonomic or phylogenetic works for each genus. Our compilation shows that Verbenaceae have 32 genera and 800 species currently accepted. This work provides up-to-date numbers and brings a holistic view of the family.
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Verbenaceae is represented in Brazil by 15 genera and ca. 290 species, with most of its richness in the Cerrado and Atlantic Forest domains. The state of Espírito Santo has an area of 46,184.1 km 2 , wholly embedded in the Atlantic Forest. Nonetheless, due to the intense environmental degradation in the state, only 10.5% of its original vegetation remains. The present study is part of the "Flora do estado do Espírito Santo" project and aims to provide a taxonomic treatment to Verbenaceae in the state, as well as to analyze its distribution patterns, together with preliminary conservation assessments for each species. It is based on morphological analysis of herbaria collections, field expeditions, and literature compilation. Distribution maps were made for each species, while the preliminary conservation assessments followed the criteria proposed by the IUCN. Verbenaceae is represented in the state of Espírito Santo by 31 species arranged in 11 genera: Aloysia (2 spp.), Bouchea (1 sp.). Casselia (1 sp.), Citharexylum (1 sp.), Glandularia (1 sp.), Lantana (10 spp.), Lippia (4 spp.), Petrea (1 sp.), Priva (2 spp.), Stachytarpheta (6 spp.) and Verbena (2 spp.). Eight species occurring in Espírito Santo are threatened with extinction. Three new records are verified, including species of the genus Lantana. The dense rainforest, which covers most of the state's territory, presents the most significant number of species (20 spp.), followed by pioneer vegetations (12 spp.), seasonal semideciduous forests (11 spp.), inselbergs (8 spp.), and ecological refuges (2 spp.). The main richness centers for Verbenaceae in the state of Espírito Santo are the municipalities of Linhares, Santa Teresa, Vitória, Vila Velha, and Guarapari in this order. In contrast, the family is noticeably little represented in the northern region of the state, as well as near its borders, where the forest remnants are highly fragmented and scattered. The current results contribute to the taxonomic and biogeographic knowledge of Verbenaceae, to the creation of conservation strategies for threatened species in the State of Espírito Santo, and reinforce the need of fieldwork in several areas of the state.
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Asinopsis of genus Junellia is here presented, following the recent re-circumscription of the genus based on morphologically founded molecular phylogenetic studies. A key to the 37 Junellia species recognized by the latest delimitation of the genus is here proposed, as well as an actualized description of the genus and its differentiation from related genera from tribe Verbeneae. Nine species and one variety of Junellia, never taxonomically treated before, are here described and/or illustrated, or their description is here amended. Two new combinations: Junellia hookeriana var. catamarcensis and Junellia trifida, and eight new synonyms are here proposed.
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Verbena and Glandularia are two genera of Verbenaceae closely related by their morphology. In this work, six species of Verbena from southern Brazil, Uruguay and Chile are transferred to Glandularia: G. catharinae, G. corymbosa, G. dusenii, G. gynobasis, G. hatschbachii and G. jordanensis, based on several typical Glandularia features, such as those related to habit, style length, inflorescence types and position of teeth on the fructiferous calyx, among others. Descriptions, illustrations and a distribution map are also provided.