a taxonomic revision of the genus tamonea …redbiblio.unne.edu.ar/pdf/62_o'leary 2008 a...

A taxonomic revision of the genusTamonea (Verbenaceae)

NATALY O’LEARY*, PAOLA PERALTA and MARÍA. E. MÚLGURA

Instituto de Botánica Darwinion, Labardén 200. CC 22 (B1642HYD), San Isidro,Buenos Aires, Argentina

Received 6 November 2007; accepted for publication 9 November 2007

A taxonomic revision of the genus Tamonea is provided. Tamonea is a small genus of Verbenaceae, placed in thetribe Casselieae. It is represented by six species, distributed in tropical and temperate America, from Mexico andthe Caribbean in the north, to Brazil, Paraguay and Bolivia in the south. Four new combinations are introduced:Tamonea curassavica var. australis, T. curassavica var. cardenasii, T. curassavica var. parvifolia and T. curassavicavar. yucatanensis; a new synonym is proposed for T. curassavica var. curassavica. The species Tamonea juncea islectotypified. Detailed morphological descriptions are given for each taxon, in addition to a key for theiridentification, illustrations, actualized synonymy, distribution and specimens lists, as well as discussion about therelationship between closely related taxa. © 2008 The Linnean Society of London, Botanical Journal of theLinnean Society, 2008, 157, 357–371.

ADDITIONAL KEYWORDS: Casseliae Ghinia – taxonomy – tropical and temperate America.

INTRODUCTION

The name Tamonea Aubl. was founded twice: oncereferring to species of Verbenaceae (Aublet, 1775: 659)and, without realizing it, in the same work (Aublet,1775: 440), also referring to a species of Melastomata-ceae. When Aublet (1775) realized this error, hereplaced the name Tamonea for Melastomataceae byFothergilla Aubl., leaving the name Tamonea for thespecies of Verbenaceae. Tamonea for Melastomata-ceae is not accepted (Rehder et al., 1935: 508;Lanjouw, 1952: 124; Gibson, 1970: 228). Schreber(1789) founded Ghinia to replace Tamonea, consider-ing this last was an invalid name. Several authorsfollowed Schreber in the use of the Ghinia name(Gmelin, 1791; Willdenow, 1797; Swartz, 1800; Stand-ley, 1924; Britton & Wilson, 1925; Moldenke, 1941,1942, León & Liogier, 1957; Moldenke, 1981a, b, c;Correl & Correl, 1982), even though Tamonea is theaccepted and valid name for the Verbenaceae genus.Many other authors, such as Swartz (1788), Jussieu(1789), Persoon (1807), Schlechtendal & Chamisso

(1830, 1831), Walpers (1845), Schauer (1847), Urban(1911), Gibson (1970), Troncoso (1974), Howard(1983), Zaruchi (1984), Martínez & Múlgura (2003)and Atkins (2004), used the correct name Tamonea forVerbenaceae.

The taxonomic position of Tamonea within theVerbenaceae family has been controversial. Schauer(1847) placed Tamonea in the tribe VerbeneaeSchauer, subtribe Casselieae Schauer, along with thegenus Casselia Nees et Mart. Briquet (1895) placedTamonea in the subfamily Verbenoideae Briq., tribeEuverbeneae Briq., along with Verbena L. & Hiero-botana Briq. Junell (1934: 196), on the basis of theovary morphology, agreed with Schauer’s classifica-tion. Troncoso (1974) placed Tamonea in the tribeVerbeneae Schauer, but withdrew subtribe CasselieaeSchauer from it and treated this last as tribe Casse-lieae (Schauer) Tronc., placing only the genus Casse-lia in it. Sanders (2001: 309) placed Tamonea togetherwith genus Casselia in a broadly enlarged tribe Ver-beneae, encompassing all Verbenaceae genus withsuffrutescent to herbaceous habit and spiciform inflo-rescences and, in general, the presence of glandularanther connectives. In the present contribution, we*Corresponding author. E-mail: [email protected]

Botanical Journal of the Linnean Society, 2008, 157, 357–371. With 8 figures

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 357–371 357

mailto:[email protected]

follow the criteria of Martínez & Múlgura (2003), alsoaccepted by Atkins (2004), who suggest that Tamonea,along with Casselia and Parodianthus Tronc., shouldbe grouped together under the tribe Casselieae,because of the ovary with false partitions being fusedto the carpel walls only at the top and bottom andthe inflorescences indeterminate and with leafymain axes, with axillary partial inflorescences in theform of racemes (Martínez, Botta & Múlgura, 1996)(Fig. 1). These three genera differ in fruit characters:in Tamonea, the fruit is entire, subdrupaceous andfour seeded; in Parodianthus it is a drupaceousschizocarp, with two two-seeded mericarps; while inCasselia it is also a drupaceous schizocarp, but withtwo one-seeded mericarps by abortion of the adaxialcarpel.

MATERIAL AND METHODS

This taxonomical revision is based on herbarium col-lections from the following herbaria: G, HAL, K, LIL,MO, NY, P, RB, S, SI, U and UPRPP. Flower mea-surements were taken from material rehydrated byboiling. Fruit measurements were taken from driedspecimens. The descriptive terminology of the inflo-rescences used here is in accordance with Martínezet al. (1996); the morphological terms used followHickey (1974). Vein formations are defined as: pin-natifid, with one major vein and secondary veinsoriginating from the major vein; actinodromous, withthree or more major veins diverging from one point;and hyphodromous, with only one major vein present,secondary veins absent or hidden within the meso-phyll. For the description of pubescence, the termsstrigose, hispid, hirsute, puberulous, scabrous orpilose are used, following Lawrence (1951).

The distribution and habitat of taxa were takenfrom the herbarium specimen labels.

TAXONOMIC TREATMENT

TAMONEA AUBL. Hist. Pl. Guiane 2: 659 (1775).Ghinia Schreb., General Pl. 19 (1789). Type: Tamoneaspicata Aubl.Ischnia A. DC. ex Meisn., Pl. Vasc. General 1: 298,2: 206 (1840). Type: not known.

Perennial HERBS and SUBSHRUBS, stems erect,tetragonal. LEAVES opposite, sessile or petiolated,venation pinnatifid, actinodromous or hyphodromous.SINFLORESCENCES bracteose, arranged in a homoteticpleiobotryum, inflorescences consisting of 1 or 2 pairsof paracladia, the basal paracladia longer than thedistal ones. Flowers arranged in spiciform rachimoseflorescences, with 1–15 briefly pedicelated flowers,alternated along the rachis. FLOWERS five-merous,calyx campanulated, persistent in fruit, with 5 sharp-pointed, equal-length teeth; corolla violet, blue orwhite coloured, funnelform or hypocrateriform,2-labiated, unequally 5-lobed; stamens 4 insertednear middle of the corolla tube, posterior pair withglandular thickening of connective, style deciduous orpersistent in fruit, included, stigma oblique; ovary2-carpellated, 2-locular carpels, 4 ovules. FRUIT

drupaceous, dry at maturity, obovate, 0.4–0.6 ¥ 0.3–0.5 cm, woody exocarp, with 4 spine-like horns orwithout horns.

Distribution and habitat: Tamonea is a genus of sixspecies, distributed in Tropical America, from Mexicoand the Caribbean to southern Brazil, Paraguay andBolivia. Species of Tamonea are not abundant, manyspecies have a very restricted area of distribution and

Figure 1. Diagram of inflorescence of Tamonea spicataAubl. with paracladia in pairs, dark cillindres representflorescences: racemes (redrawn from Martínez et al., 1996).

358 N. O’LEARY ET AL.

© 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157, 357–371

there are many endemisms. That is the reason why ithas been a poorly collected genus.

Notes: In horned fruits there are generally four horns,two of them longer than the other two. Nevertheless,there are specimens [e.g. Duarte 7541 (SI), Tamoneacurassavica] where two of the horns are fusedtogether, looking like a three-horned fruit.

1. TAMONEA BOXIANA (MOLDENKE) R. A. HOWARD

(Fig. 2)

Phytologia 65: 288 (1988). Ghinia boxiana Moldenke,Known Geogr. Distr. Verb. & Avicen. 77 (1942),replaced name: Ghinia spinosa (Sw.) Britton &Wilson, Scient. Survey Puerto Rico 6: 139 (1925),nom. illeg., non G. spinosa (Sw.) Willd., Sp. pl. 1: 114(1797), nom. illeg. Type: Antigua and Barbuda, H. E.Box 1091 (Neotype: here designated, BM, photographSI!; isotype: NY, photograph SI!).

SUBSHRUBS, to 50–60 cm high, puberulous or sca-brous pubescence. LEAVES sessile or briefly petiolated,

hyphodromous venation, dimorphic, the basal leaves0.7–1 ¥ 0.2–0.5 cm, oblong, cuneate base, obtuse apex,lobed margins, the apical leaves 1 ¥ 0.1 cm, linear,acute apex, entire margins. INFLORESCENCES withlong pair of basal paracladia and short solitaire distalparacladia. Florescences 3–6 cm long., with 5–8flowers briefly pedicelated c. 0.5 cm long.; floral bracts0.2 cm long, linear. Calyx 0.25 cm long, teeth 0.5 mmlong; hypocrateriform corolla, violet coloured, tube0.4 cm long. FRUIT 4 horned, longer pair of horns 4 mmlong, shorter pair 2 mm long.

Iconography: Britton, 1932: 5, figure 547; Howard,1989: 241, figure 98.

Common name: ‘cardero’ (Moldenke, 1981b: 417).

Phenology: Flowering and fruiting in spring andsummer.

Distribution and habitat: This species grows inPuerto Rico and the Lesser Antilles. It is found in dryforests.

KEY TO THE SPECIES OF TAMONEA

1a. Fruit without horns...........................................................................................................5. T. spicata1b. Fruit with horns..............................................................................................................................22a. Leaves with actinodromous venation ................................................................................................... 32b. Leaves with pinnatifid or hyphodromous venation ................................................................................. 43a. Fruits with short horns, 1 mm long.; plants with hispid or hirsute pubescence, calyx with glandular trichomes

.......................................................................................................................................4. T. juncea3b. Fruit with long horns, 2–3 mm long.; plants with strigose pubescence, calyx with few, short, adpressed, non-

glandular trichomes..................................................................................................3. T. euphrasiifolia4a. Dimorphic leaves, hyphodromous venation, the basal leaves oblong, 0.2–0.5 cm lat, lobed margins, the apical

leaves linear, 0.1 cm lat., entire margins .............................................................................. 1. T. boxiana4b. Homomorphic leaves, pinnatifid venation, blades ovate, 0.3–2 cm lat., serrate margins.................................55a. Florescences of 1–2 flowers..............................................................................................6. T. subbiflora5b. Florescences of 3–15 flowers .............................................................................................................. 66a. Fruiting calyx reaches the base of the fruit horns; horns brief 0.5–1 mm long; leaves shorter than 1.5 cm long.,

with puberulous pubescence.....................................................................2e. T. curassavica var. parvifolia6b. Fruiting calyx never reaches the base of the fruit horns; horns well developed, 1–4 mm long.; leaves longer than

1.5 cm long., with pilose, hispid, strigose or puberulous pubescence..........................................................77a. Leaves with pilose pubescence, with non-white translucid trichomes, 0.5–0.6 mm long...................................

........................................................................................................... 2b. T. curassavica var. australis7b. Leaves with hispid, strigose or puberulous pubescence, with white opaque trichomes, shorter than 0.5 mm long

.................................................................................................................................................... 88a. Leaves with truncate base, obtuse apex, pubescence puberulous, trichomes 0.1–0.25 mm long. Endemic of Yucatán,

in Mexico ......................................................................................... 2d. T. curassavica var. yucatanensis8b. Leaves with cuneate base, acute apex, pubescence hispid or strigose, with trichomes 0.25–0.5 mm long. Native

from Bahamas, Bolivia, Cuba, Guatemala, Mexico and Paraguay ............................................................. 99a. Floral bracts 0.15–0.2 cm long.; leaves hispid on abaxial surface, glandular trichomes on both surfaces............

..........................................................................................................2c. T. curassavica var. cardenasii9b. Floral bracts 0.2–0.5 cm long.; leaves strigose on main veins of abaxial surface, hispid on secondary veins .......

........................................................................................................2a. T. curassavica var. curassavica

TAXONOMIC REVISION OF TAMONEA 359


Notes: Ghinia spinosa Britton & P. Wilson (1925) isan illegitimate name because it is a homonym tothe previous name by Willdenow (1797). Besides, thislast one is also an illegitimate name, because it isbased on a superfluous name: Tamonea spinosaSw. (see notes under T. curassavica). So Moldenke(1942) replaces this illegitimate name by G. boxiana,without mentioning any type specimen. Later on,Moldenke (1981a: 417), refers to some specimens sentfrom herb. BM by Dr Box ‘Antigua: Box 1091’, butnever explained where the holotype was deposited.Howard (1988: 288) combines Tamonea boxiana(Moldenke) Howard and later (Howard, 1989: 241) hetypifies this taxon with the specimen mentioned byMoldenke (1981a: 417), but does not chose an holo-type. As original material from Swartz could not befound, the Box specimen chosen by Moldenke (1981a:417) and Howard (Howard, 1989: 241) is the suitableneotype.

Specimens examined: ANTIGUA and BARBUDA:Barbuda, August 1891, Nicholls s.n. (K). PUERTORICO: Cabo Rojo: Bo. Boquerón, along ravine on N ofPunta Melones, 5 February 1995, F. Axelrod & A.Axelrod 8735 (UPRPP); Salinas de Cabo Rojo ad litoralmarino, 5 February 1885, Urban 632 (G, K, MO, LIL,P, S); Cerro Mariquita, Sierra Bermeja, on slope, 16August 1992, Liogier 36998 (MO); Ponce: Bo. Canas,along stream bed E of prison, 20 November 1994, FAxelrod & A. Axelrod 8515 (UPRPP); Prope Guanica,in Ponticetis ad la Montalba, 2 February 1886, Urban3554 (G, K, MO, P, S, US); Route 120, km 9.6 inMaricao Forest, 28 Set 1968, Wagner 1667 (U).

2. TAMONEA CURASSAVICA (L.) PERS

Syn. Pl. 2: 139 (1807). Verbena curassavica L., Sp. Pl.19 (1753). Zapania curassavica (L.) Lam., . Encycl.Méth. Bot. [Illustr. General] 1: 59 (1791). Ghinia

Figure 2. Tamonea boxiana. A, leafy branchlet with inflorescence. B, basal leaf, abaxial face. C, apical leaf, adaxial face.D, fruiting calyx. E, fruit. [From Urban 632 (MO).]



curassavica (L.) Oken, Allg. Naturgesch., Bot. 2: 1104(1841). Ghinia curassavica (L.) Millsp., Publ. FieldColumb. Mus. Bot. 2: 174 (1906), nom. illeg. hom.Type: [icon.] ‘Veronicae similis fruticosa Curassavica’in Herman, Parad. Bat. t. opp. p. 240. 1698. (Lecto-type: designated by Méndez Santos & Cafferty, 2001:1139).

Tamonea spinosa Sw., Prodr. 94 (1788), nom. illeg.superfl. Ghinia spinosa (Sw.) Willd., Sp. pl. 1: 114(1797). comb. illeg. Type: Curaçao 1782, s/legit. (Lec-totype: here designated, S, photograph SI!)

Ghinia verbenacea Sw., Fl. Indias Occidentales 2:1089 (1800). nom. illeg. superfl. Tamonea verbenaceaSw., Fl. Indias Occidentales 2: 1089 (1800). nom.inval. Tamonea verbenacea (Sw.) Walpers, Repert.Bot. Syst. 4: 115 (1845), comb. illeg. Tamonea ver-benacea ex Schauer, in DC Prodr. 11: 528 (1847),comb. illeg. Type: unknown.

Tamonea scabra Schltdl. & Cham., Linnaea 5: 99(1830). Type: México: Tierra caliente, Puente del Rey,Hacienda de la Laguna, October 1828/29, F. Deppe &C. J. W. Schiede s.n. (Holotype: HAL 98486, photo-graph SI!)

Ischnia verbenacea A. DC., Prodr. 9: 257 (1845).Type: Mexico 1787–1803, M. Sessé & J. M. Mociño s.n.(Holotype: G).

HERBS or SUBSHRUBS, from 35 to 100 cm high,stems woody at the base, subglabrous or pubescent.LEAVES with a brief petiole 0.3–1 cm long., pinnatifidvenation, blade 0.6–6 ¥ 0.3–2 cm, ovate, cuneate ortruncate base, obtuse or acute apex, serrate margins.INFLORESCENCES with long pair of basal paracladiaand short solitaire distal paracladia. Florescences10–25 cm long, with 3–15 flowers briefly pedicelated0.1–0.3 cm long.; floral bracts 0.15–0.5 cm long.,linear. Calyx 0.3–0.55 cm long, teeth 0.5–1.5 mm long;hypocrateriform corolla, violet or blue coloured, tube0.5–0.7 cm long. FRUIT 4 horned, horns 0.5–4 mmlong., generally a pair of horns longer than the other.

Notes: Varieties were considered when a group oforganisms with characters of gradual variation wasobserved, which would indicate an incomplete segre-gation of the incipient species, sharing the same geo-graphical area. We followed the Suttill & Allen (1992)concept of variety used when the taxon is poorlydifferentiated and the variation is mostly ecotypical,not geographical. In this species, five varieties arerecognized, principally distinguished because of thetype of pubescence, length and morphology of theleaves and floral bracts. Trichomes can be glandularor non-glandular, in the latter group trichomes areusually white opaque, sometimes non-white andtranslucid trichomes are found.

Tamonea spinosa Sw., T. verbenacea Sw. and Ghiniaverbenacea Sw. are all illegitimate superfluous names,

because in all cases the author is referring to Verbenacurassavica L. (a valid name), so following the Viennacode (Art. 52, McNeill et al., 2006) this last one shouldbe the name of the taxon. The combinations byWalpers (1845: 115) and Schauer (1847: 528) areillegitimate because they are based upon illegitimatenames.

2A. TAMONEA CURASSAVICA VAR. CURASSAVICA

Verbena pedicellata Moldenke, Phytologia 25(7): 431(1973). Type: Brazil. Mato Grosso, Campo Gde.,Anhandui, 17 October 1972, G. Hatschbach 30513(Holotype: TEX, photo SI; isotype: MBM!, US, photoSI, SI!). syn. nov.

BRANCHES with short patent glandular and non-glandular with white opaque trichomes. LEAF blade1.5–2.5 ¥ 0.3–1.3 cm, ovate, cuneate base, acute apex,adaxially subglabrous or strigose, abaxially strigoseon the main veins, secondary veins with hispid, withwhite opaque trichomes, 0.25–0.5 mm long. FLORAL

BRACTS 0.2–0.5 cm long., calyx 0.3–0.35 cm long., gla-brous or strigose, fruiting calyx never reaches thebase of the fruit horns. FRUIT 4 horned, horns 1.5–3.5 mm long., generally a pair of horns longer thanthe other.

Iconography: Correl & Correl, 1982: 1229, figure 529(sub Ghinia curassavica); Méndez Santos, 2003: 18,figure 3; Nash & Nee, 1984: 123, figure 14; Rze-dowsky & Calderón de Rzedowsky, 2002: 116.

Common name: ‘flor morada’, ‘spiny-fruted vervain’,‘stachliche Traubennuss’, ‘tamonèe èpineuse’, ‘thorny-fruited tamonea’, ‘zapane de Curaçao’ (Moldenke,1981b: 454).

Phenology: Flowering and fruiting from March toDecember.

Distribution and habitat: Cuba, Guatemala, Mexicoand Paraguay. According to Moldenke (1981a: 454)also in Bahamas (New Providence). It grows in lime-stone hills. Berazaín Iturralde et al. (2005: 48) con-sider this species endangered in Cuba, referring to itsstate as ‘vulnerable’ following the listings of the Inter-national Union for the Conservation of Nature andNatural Resources (IUCN; 2001).

Specimens examined: MEXICO. Chiapas: La Hoya1841–43, Liebman 11307 (SI, U); Ocozocoautla deEspinoza, 1 km N of Ocozocoautla, 830 m, 25 Set1971, Breedlove 19784 (MO). Tamaulipas: Aldama,sierras de Tamaulipas, region of Rancho Las Yucas,c. 40 km, NNW of Aldama, 23°14′N, 98°10′W, 10October 1957, Dressler 2337 (MO); N of Limon, 15



June 1935, Clark 6826 (MO). San Luis Potosi:Masate, Tamazunchale, 7 November 1937, Edwars514 (MO); Puente del Rey, July 1828, Schiede s.n.HAL 98487 (HAL). Veracruz: along Hwy 180 betweenTampico and Pozarica, 12 miles N of Azuluana, 38 kmN of Naranjos, 110 m, 5 June 987, Croat 66099 (MO);8.6 miles S of Tantoyuca off highway 120, 25 June1965, Maxwell 50 (MO); Cordillera, Jun–October1840, Galeotti 767 (P, SI); Córdoba, 5 August 1882,Fournier 28 (K, P, SI) 1882, Kerber 28 (G, K, P, SI);Schiede 91 (K, SI); Cuitlahauc, 17 May 1937, Matuda1457 (MO); Dos Ríos, Cerro Gordo, 600 m, 20 April1970, Ventura 924 (MO); Puente Nacional, Mata deCaña, 100 m, 3 August 1981, Ventura 18912 (MO);Region of San Andrés Tuxtla, along river drainingLaguna de Catemaco, 31 August 1953, Dressler 224(MO); 29 July 1969, Martinez Calderón 1956 (MO);Río de Santa María, November 1906, Purpus 2220(MO); Zacuapan, Abr 1931, Purpus (NY, SI). GUATE-MALA. Huehuetenango: 4 miles E of Teopisca,3100 m, 13 August 1975, Stoerker 123 (MO). CUBA.Camagüey: alrededores del poblado Martín,Guaimaro, Méndez 7175 (SI). La Habana: Lomas deCamoa, limestone hills, 25 km SE of Habana, 7November 1921, Ekman 13468 (G, K, S, SI, U), inruderaris, near hause, 26 June 1922, Ekman 14127(G, S, SI); Ramon de la Sagra, Herb. Richard s.n. (P,SI). PARAGUAY. Boquerón: Parque ‘Valle Natural’,12 km S de Filadelfia, 28 February 1991, Vanni et al.2437 (CTES).

2B. TAMONEA CURASSAVICA VAR. AUSTRALIS

(MOLDENKE) O’LEARY, P. PERALTA & MÚLGURA,COMB. NOV. (Fig. 3)

Ghinia curassavica var. australis Moldenke, Phytolo-gia 24(5): 498 (1972). Type: Brazil. Bahía EspigaoMestre, about 25 km W of Barreuras, in the valley Riodas Ondas, 600 m, 3 March 1971, H. S. Irwin, R. M.Harley & G. L. Smith 31404 (Holotype: TEX, photo-graph SI!; isotype: F, K, MO, US, SI!).

BRANCHES and LEAVES with pilose pubescence,translucid trichomes 0.5–0.6 mm long., translucidand abundant glandular trichomes. LEAF blade 1.5–2 ¥ 0.8–1 cm, ovate, truncate base, acute or obtuseapex. FLORAL BRACTS 0.25–0.3 cm long, hispid, calyx0.5–0.55 cm long., hispid, with some glandular tri-chomes, fruiting calyx never reaches the base of thefruit horns. FRUIT 4 horned, horns 2–4 mm long.,generally a pair of horns longer than the other.

Phenology: Flowering and fruiting in March.

Distribution and habitat: This variety grows inBrazil, in the ‘Cerrado’, at 600 m.

Specimens examined: BRAZIL. Bahía: Riachao dasNeves, G. Hatschbach 42120 (NY, SI). Minas Gerais:Varzea da Palma, Fazenda do Jatobá, 25 November1962, Duarte 7541 (SI).

2C. TAMONEA CURASSAVICA VAR. CARDENASII

(MOLDENKE) O’LEARY, P. PERALTA & MÚLGURA,COMB. NOV.

Ghinia cardenasii Moldenke, Bull. Torrey Bot. Club68: 504 (1941). Tamonea cardenasii (Moldenke)Tronc., Darwiniana 18: 323 (1974). Type: Bolivia.Santa Cruz: Ipias-Chiquitos, in grassy pampas, alt.230 m, October 1934, M. Cárdenas 2946 (Holotype: F755045, photograph SI!; isotype: NY, photograph SI!).

BRANCHES hispid, with glandular thrichomes andnon-glandular short, white opaque thrichomes. LEAF

blade 1.5–6 ¥ 0.8–2 cm, ovate, cuneate base, acuteapex, scarcely pubescent on adaxial surface, hispid onabaxial surface, non-glandular short, white opaquetrichomes, 0.25–0.5 mm long. and glandular tri-chomes on both surfaces. FLORAL BRACTS 0.15–0.2 cmlong., calyx 0.4–0.45 cm long., with glandular andnon-glandular thrichomes, fruiting calyx neverreaches the bases of the fruit horns. FRUIT 4 horned,horns 4 mm long., generally a pair of horns longerthan the other.

Iconography: Troncoso, 1974: 322, figure 5.


Distribution and habitat: Bolivia & Paraguay. Mold-enke (1981a) mentioned specimens from Brazil, in thestates of Bahia and Mina Gerais. It is found in sandysoils and pastures, also in xerophictic forest.

Specimens examined: BOLIVIA. Chuquisaca: LuisCalvo, Ipauso Laguna del Medio, 20°43′S, 63°06′-W,700 ms.m., 9 April 1993, Saravia Toledo et al. 11526(CTES, SI). Santa Cruz:10 km al N Santa Cruz, 20April 1977, Krapovickas et al. 31457 (CTES, K, SI);Carandatay, March 1935, T. Rojas 7358 (LIL). PARA-GUAY. Nueva Asunción: bosque al término N de laautopista militar y alrededores, 20°43′S, 61°56′W, 24March 1986, Brunner 1558 (MO, PY, SI).

2D. TAMONEA CURASSAVICA VAR. PARVIFOLIA

(MOLDENKE) O’LEARY, P. PERALTA & MÚLGURA,COMB. NOV. (Fig. 4)

Ghinia curassavica f. parvifolia Moldenke, Phytologia47(6): 455 (1981). Type: Mexico. San Luis Potosí,



valles, 3 August 1937, G. L. Fisher s.n. (Holotype: NY,photograph SI!).

BRANCHES with adpressed white opaque, non-glandular thrichomes. LEAF blade 1–1.5 ¥ 0.8–1 cm,ovate, cuneate base, acute or obtuse apex, bothsurfaces puberulous with short and white opaque,non-glandular trichomes 0.1–0.25 mm long. FLORAL

BRACTS 0.15–0.20 cm long., calyx 0.3–0.35 cm long.,fruiting calyx reaches the base of the fruit horns.FRUIT 4 horned, horns 0.5–1 mm long., generally apair of horns longer than the other.

Phenology: Flowering from April to October.

Distribution and habitat: México. It is found in lime-stone soils and rocky forests, 10–2050 m.

Specimens examined: MEXICO. San Luis Potosi:Minas de San Rafael, May 1911, Purpus 5291(MO); San Dieguito, 13–16 June 1904, Palmer125 (K, MO, SI); Palmer 133 (K, MO, NY, SI);Rascon, August 1911, Purpus 5290 (MO). Sinaloa:San Antonio, Tanjas, 7 November 1978, Alcorn 1581(NY, SI). Tamaulipas: 1931, von Rozynski 378 (G,SI); 14 October 1932, von Rozynski 521 (P, SI);Nogales, von Rozynski 483 (SI); 27 km SE ofMiquihuana on road to Palmilla, 23°37′N, 99°39′W,

Figure 3. Tamonea curassavica var. australis. A, plant general aspect. B, leaf, abaxial face. C, flower with floral bract.D, fruiting calyx. [From Hatschbach 42120 (NY).]



Figure 4. Tamonea curassavica var. parvifolia. A, plant general aspect. B, leaf, adaxial face. C, fruiting calyx. D, fruit.[From Palmer 133 (K).]



2025 m, 13 August 1941, L.R. Stanford et al. 886(MO).

2E. TAMONEA CURASSAVICA VAR. YUCATANENSIS

(MOLDENKE) O’LEARY, P. PERALTA &M. E. MÚLGURA, COMB. NOV. (Fig. 5)

Ghinia curassavica var. yucatanensis Moldenke, Publ.Carnegie Institute Wash. 522 (17): 152–153 (1940).Type: Mexico. Yucatán 1895, G. F. Gaumer 834b (Holo-type: NY, photograph SI!; isotype: BM!, photographSI!, G, photograph SI!, MO!,US, photograph SI!).

BRANCHES glabrous or scarcely puberulous. LEAF

blade 1.5–2.5 ¥ 0.6–2 cm, ovate, truncate base, obtuseapex, adaxially glabrous or subglabrous, abaxially

surface glabrous or puberulous, with white opaque,non-glandular trichomes 0.1–0.25 mm long. FLORAL

BRACTS 0.2 cm long., calyx 0.3–0.35 cm long. puberu-lous, fruiting calyx never reaches the base of the fruithorns. FRUIT 4 horned, horns 1–3 mm long., generallya pair of horns longer than the other.

Phenology: Flowering and fruiting from June toNovember.

Common name: ‘chancolenexnuc’, ‘chan-ko-xnuk’,‘chanxnuk’ (Moldenke, 1981b: 457).

Distribution and habitat: This variety is endemicfrom Yucatán, in Mexico. It is found in wetlands.

Figure 5. Tamonea curassavica var. yucatanensis. A, branch detail. B, leaf, abaxial face. C, fruit. D, detail of stempubescence [A, B, from Gaumer 834 (K); C, D, from Gaumer 834b (SI isotype)].



Specimens examined: MEXICO. Yucatán: Izamal1917–1921, Gaumer 24097 (G, K, SI); Gaumer 24228(G); Gaumer et al. 23384 (SI); Lundell 8176 (NY, SI);18 Set 1925, Gaumer 834 (G, K, P, SI); San Anselmo,Gaumer 1988 (K, SI); Valladolid, camino rumbo a SanLorenzo, 20°42′53′′N, 88°15′45′′W, 22 m, 4 July 1988,Remmers, G. & H. de Voeyer 16 (MO); a 5 km al S deTelchac Puerto, sobre la carretera a Telchac Pueblo,24 December 1985, E. Cabrera 10312 (MO).

3. TAMONEA EUPHRASIIFOLIA B. L. ROB. (Fig. 6)

Proc. Amer. Acad. Arts 44: 613 (1909). Ghiniaeuphrasiifolia (B. L. Rob.) Standl., Contr. U.S. Natl.Herb. 23(5): 1236 (1924). Type: Mexico. Tamaulipas,Alta Mira, 14–22 May 1898, E. W. Nelson 4415 (Holo-type: GH, photograph SI!; isotype: US, photographSI!, SI!).

Tamonea scabra var. minor Schltdl. & Cham.,Linnaea 6: 373 (1831). Ghinia curassavica var. minor(Schltdl. & Cham.) Moldenke, Phytologia 47(2): 88(1980). Type: Mexico. ad littora maris arenosa propeLaguna salada, reg. calid, March 1828/29, C. J. W.Schiede & F. Deppe s.n. (Holotype: HAL 98481, pho-tograph SI!; isotype: BM, photograph SI!, SI!, W).

SUBSHRUBS, to 30–60 cm high, stems woody at thebase, strigose pubescence, hispid at the ribs. LEAVES

with a brief petiole 0.1–0.2 cm long., actinodromousvenation, blades 0.5–1 ¥ 0.4–1 cm, ovate, truncatebase, obtuse apex, serrate margins, adaxialy subgla-brous, adpressed trichomes on abaxial face. INFLO-RESCENCES with solitaire paracladia. Florescences5–10 cm long., with 3–5 flowers briefly pedicelated0.2 cm long.; floral bracts 0.2 cm long., sublinear.Calyx 0.45–0.6 cm long. with teeth 1–2 mm long.,with adpressed, short, non-glandular trichomes; fun-

Figure 6. Tamonea euphrasiifolia. A, plant general aspect. B, leaf, abaxial face. C, flower with floral bract. D, fruitingcalyx. [From Palmer 538 (K).]



nelform corolla, violet coloured, tube c. 1.5 cm long.FRUIT 4 horned, horns 2–3 mm long, generally a pairof horns longer than the other.

Phenology: Flowering in May.

Distribution and habitat: This species is endemicfrom eastern Mexico. It is found in sandy and limesoils.

Notes: Robinson mentioned the type material ofTamonea euphrasiifolia B. L. Rob., which was depos-ited at GH and US, not mentioning which one was theholotype. Moldenke (1981b: 459) says the specimenfrom GH is the holotype, so the US specimen is anisotype.

Tamonea euphrasiifolia shares with T. junceaSchauer the same kind of actinodromous venation.Nevertheless, they differ because T. juncea has densehispid pubescence, with glandular hairs and the fruithorns are very brief, while T. euphrasiifolia has scarestrigose pubescence, without glandular hairs and thefruit horns are longer, c. 0.3 cm long. Apart from that

T. euphrasiifolia grows in Mexico and T. juncea growsin the state of Bahia, in Brazil.

Specimens examined: MEXICO. Tamaulipas:Tampico, 4 V 1921, Rutten et al. 529 (SI). Veracruz:Near Pueblo Viejo, 9 km S Tampico 1910, Palmer 538(K, NY, SI).

4. TAMONEA JUNCEA SCHAUER (Fig. 7)

Prodr. 11: 529 (1847). Ghinia juncea (Schauer) Mold-enke, Phytologia 1(4): 169 (1935). Type: Brazil. Bahia,no date, J. Blanchet 3397 (Lectotype: here designated,BM, photograph SI!; isotype: FM 17397 (photograph),G, NY, photograph SI!, P, photograph SI!, US, photo-graph SI!, SI!, US, photograph SI!).

SUBSHRUBS, to 50 cm high, rigid, erect, woodybranches; hispid or hirsute glandular pubescence.LEAVES rigid and adpressed to the stem, subsessile orsessile, blades 0.8 ¥ 0.6 cm, actinodromous venation,subtriangular, truncate base, obtuse apex, serratemargins, hispid pubescence, mainly on abaxialsurface veins. INFLORESCENCES with solitaire para-cladia. Florescences 5 cm long., with 1–2 flowers,briefly pedicelated 0.2 cm long.; floral bracts 0.25 cm

Figure 7. Tamonea juncea. A, plant general aspect. B, leaf, abaxial face. C, flower with floral bract. D, fruiting calyx.[From Harley 16667 (K).]



long., sublinear. Calyx 0.4–0.5 cm long. with teeth1–2 mm long., dense hispid or hirsute pubescencewith glandular trichomes; funnelform corolla, violetcoloured, tube 1 cm long. FRUITS 4 horned, horns1 mm long.


Distribution and habitat: Tamonea juncea grows inBrazil, in the state of Bahia. It is found in rocky,sandy lands and in the ‘cerrados’.

Notes: Schauer (1847) in the protologue of this speciesmentions two specimens: Blanchet 2566 and Blanchet3397 (sintypes). The specimen Blanchet 3397 washere chosen as the lectotype because it is a goodspecimen, with many isotypes distributed in severalherbaria; the material from BM was selected as lec-totype because Schauer worked there.

Specimens examined: BRAZIL. Bahia: Serra doCurral Feio, 16 km NW Lagoinha, 4 March 1974,Harley 16667 (K, P, RB, SI); s. loc., Paganucci deQueirolo 850 (K); s. loc., Blanchet 2566 (G).

5. TAMONEA SPICATA AUBL.

Hist. Pl. Guiane 2: 660. t. 268 (1775). Tamoneamutica Sw., Prod. Veg. Ind. Occ. 94 (1788) nom. illeg.superfl. Ghinia mutica (Sw.) Willd., Sp. pl. 1: 114(1797). Ghinia mutica (Sw.) Sw., Fl. Ind. Occid. 2:1090 (1800). nom. illeg. Ghinia spicata (Aubl.) Mold-enke, Phytologia 1(4): 169 (1935). nom. illeg. Type:French Guiana. Aublet s.n. (Holotype: BM, photo-graph SI!).

HERBS or SUBSHRUBS, 30–100 cm high, woodystems, scare pubescence with retrorse hairs. LEAVES

with a brief petiole 0.5–1 cm long., blades 2–3 ¥ 1–2 cm, pinnatifid venation, ovate, truncate base, acuteor subobtuse apex, serrate margins, subglabrous onadaxial surface, hispid or strigose on abaxial surface.INFLORESCENCE with solitaire paracladia or in pairs.Florescences 3–8 cm long., with 3–5 flowers; brieflypedicelated 0.2 cm long.; floral bracts 0.25 cm long.,sublinear. Calyx 0.35–0.4 cm long. teeth 5–15 mmlong., hispid pubescence; funnelform corolla, violetcoloured, tube 0.7 cm long. FRUIT not horned, some-times with 4 small 0.05–0.1 mm long. bulges.

Iconography: Aublet, 1775: 660. figure 268; Gibson,1970: 229, figure 45; Lopez-Palacios, 1977: 315, fig-ure 75; Jansen-Jacobs, 1988: 74, figure 21; Aymard,2005: 439, figure 369.

Phenology: Flowering and fruiting in spring andsummer.

Distribution and habitat: It grows from Mexico in thenorth to tropical Brazil in the south. It grows insandy, rocky grounds.

Notes: Tamonea spicata distinguishes from the rest ofthe species of Tamonea because of the absence ofhorns on the fruits.

Stevens et al. (2001) mentions it is employed as amedicinal plant for menstrual and stomach pain.Aymard (2005) says it is a poisonous for the cattle.

Link (1820) refers to a specimen number ‘389’ fromherb. Willdenow (B), named in herb. as Leptocarpuschamaedrifolius, but that corresponds to Tamoneaspicata, so the Leptocarpus chamaedrifolius name isnot a validly published name (Art. 32, McNeill et al.,2006).

Specimens examined: MEXICO. Chiapas: Ococingo,Cofolasa, 24 km SE crucero Corozal, 25 February1985, Martínez 11257 (MO); Ococingo, 16 km NWBoca Lacantum, 20 January 1986, Martínez 16454(MO); Ococingo, 16 km NW Boca Lacantum, 31October 1985, Martínez 14514 (MO).

BELIZE. s. loc., 1837, Schomburg 74.S (K);Maskall, 5–20 m, 17.52.42 N 088.18.47 W, 10 March1934, Gentle 1181 (NY, MO); Colonel English PineRidge, 16 xii 1957, Gentle 9473 (S); Ridge LagoonPlantation, 18 January 1974, Liesner 1429 (MO);Western Highway, The Place, 5 June 1981, Whitefoord3140 (MO); Western highway, Parrots Wood, 17 Set1980. Whitefoord 2586 (MO); SW Belize city, nearBelize Zoo, 12 August 1992, Worthington 21276 (MO);Western, 2 May 1981, Whitefoord 2681 (MO). Cayo:near Augustine, 26 August 1980, Whitefoord 2220(MO); along road at Rio Ma Cal, 28 May 1973, Croat23513 (MO); 2–5 miles N Blancaneaux lodge, 12 July1973, Dwyer 11641 (MO). Toledo: 12 miles SanAntonio Punta Gorda Road, 10 March 1949, Gentle6687 (G, K, NY, S, SI). Stann Creek: Millins Riverroad, 3 December 1954, Gentle 8482 (G, S). HONDU-RAS. Gracias a Dios: orilla E Laguna Caratasca, 28January 1984, Nelson & Cruz 8620 (MO). NICARA-GUA. Cabo: 13 July 1972, Seymour 5916 (MO). Kiaya:Tuapi, 15 July 1993, Coe et al. 4161 (MO). Zelaya:Tuapi, 15 July 1993, Coe 4160 (MO); 1 km E Lam-blaya, 9 March 1981, Douglas Stevens 19610 (MO);Rio Likas, near Silima Lila, 5 March 1979, Pipoly4068 (MO); Rio Likas, near Silima Lila, 5 March1979, Pipoly 4124 (MO). PANAMA. Panamá: cerroJefe, 29 June 1978, Hammel 3708 (MO).

BRAZIL. Bahia: s. loc., 1909, Bierens de Haan 117(U); s. loc., Blanchet 168 (MO) 1835, Blanchet 1216(G), 24 April 1839, Blanchet s.n. (G), 1853, Blanchet337 (G), 1840. Blanchet s.n. (G); s. loc., Salzmann(MO); 14.18S 038.59 W, 31 March 1974, Harley et al.



17560 (K, MO, RB, U); 5 km SE Maraú, 30 March1974, Harley 17497 (K, MO, U); 2 February 1977,Harley et al. 18491 (K; MO), Coastal Zone, 16 May1980. Harley et al. 22138 (K, U); Feira de Santana,campus da UEFS, Queiroz & Crepaldi 1510 (K);Ilheus, fazenda Porto Seguro, 26 August 1961, Duarte6102 (RB); region Cacaueria da Bahia, area doPousio, 22 April 1982, Cavalho et al. 1288 (K); Serrado Sincorá, 4 February 1974, Harley 15881 (K, MO, P,U); Serra Grande, Itacaré, 26 August 1992, Amorim674 (NY, SI); Theobroma, August 1988, Sobral &Mattos Silva 6047 (ICN, SI); Vittoria, Sellow s.n. (K).FRENCH GUIANA. Cayenne: 1835, Leprieur s.n. (K);Cayenne: Layon forestier, 10 v 1990. Cremer & Hoff11378 (U); km 23, crique Anguille, W of R du Tour del’Ile Savanna, Leeuweuberger 11685 (U); bord de laroute de l’Est, Prévost 1581 (U); Upper Takutu-UpperEssequibo, Karanambo, Savanna near airstrip, 4 Set1988, Maas et al. 7245 (K, NY, U). GUYANA. EbiniExp. Intermediate Savanna, Berbice River, 14 Abr1958, Harrison 858 (K); Waranama ranch, Interme-

diate Savanna, Berbice river, 6 Set 1958, Harrison &Persand 1042 (K). SURINAME. Lobim, savanna nterZanderij, 26 Set 1958, Donselaar 124 (U); Republick,23 April 1960, Hekking 80 (U); Republick, 4 Set 1948,Lanjouw & Lindeman 135 (U); Pará, 12 April 1974,Vreden 14529 (U); Lindeman 27 (NY, SI); Tibitisavanne, 10 January 1949, Lanjouw & Lindeman1764 (NY, U). VENEZUELA. Bolivar: cerro Pichacho,N of Las Nieves, February 1961, Steyermark 89285(G); Páez, La Grulla, 6 km SE Upata, July 1978,Delascio et al. 6811 (MO); 61 km SE Upata, 2 Decem-ber 1973, Davidse 4633 (MO); Reserva ForestalImataca, January 1983, Stergios et al. 4919 (MO).

6. TAMONEA SUBBIFLORA URB. & EKMAN (Fig. 8)

Ark. Bot. 22A(10): 106 (1929). Ghinia subbiflora (Urb.& Ekman) Moldenke, Rev. Sudamer. Bot. 4: 17 (1937).Type: Haití. La Española, ‘Ile la Gonave, La Source’, 7August 1927, E. L. Ekman 8844 (Holotype: S, photo-graph SI!; isotype: NY, photograph SI!, SI!)

Figure 8. Tamonea subbiflora. A, plant general aspect. B, leaf, adaxial face. C, fruiting calyx. [From Ekman 4500 (K).]



HERBS or SUBSHRUBS, 20–60 cm high, stems woodyat the base, puberulous pubescence. LEAVES witha brief petiole 0.25–0.3(1) cm long., puberulous,with or without glandular trichomes, blade 0.6–1.6(2.5) ¥ 0.3–0.6 (1.3) cm, pinnatifid venation, ovate,truncate base, obtuse apex, serrate margin, adaxiallypuberulous or glabrous, abaxially puberulous, overvenation. INFLORESCENCES with solitaire paracladiaor in pairs. Florescences to 2 cm long., with 1 or 2flowers; briefly pedicelated 0.1 cm long.; floral bracts0.1–0.2 (0.3) cm long, linear, with glandular trichomesor not. Calyx 0.2–0.25 cm long., teeth c. 1 mm long.FRUIT 4 horned, horns 3–4 mm long., generally a pairof horns longer than the other.

Phenology: Flowering and fruiting in July andAugust.

Distribution and habitat: Haití and eastern Cuba. Itgrows in wet places and along the routes, at 400 m.

Notes: According to Berazaín Iturralde et al. (2005:48), this species is endangered in Cuba and its stateis ‘critical’ following IUCN (2001). No collected speci-mens from Cuba could be found.

Specimens examined: HAITI. Hispaniola: Civ.,Department Nord-Quest, plateau of Bombarte moistplace on the road near Vallière, 8 July 1925, Ekman4500 (K, S).

ACKNOWLEDGEMENTS

We thank the curators of the herbaria cited in the textfor the loan of specimens and assistance in the searchof type material.

REFERENCES

Atkins S. 2004. Verbenaceae. In: Kubitzki K, ed. The familiesand genera of vascular plants, vol. 7. Berlin: Springer-Verlag, 449–468.

Aublet F. 1775. Histoire des plantes de la Guiane françoiseLondres. Paris: P. F. Didot Jeune, 1–393.

Aymard GA. 2005. Verbenaceae. In: Berry PE, Yatskievych K,Holst BK, eds. Flora of the Venezuelan Guayana, vol. 9. StLouis, MO: Missouri Botanical Garden, 407–445.

Berazaín Iturralde R, Areces Berazaín F, Lazcano LaraJC, González Torres LR. 2005. Verbenaceae. Lista Rojade la Flora Vascular Cubana. Gijón: Jardín Botánico Atlán-tico Gijón, 47–48.

Briquet J. 1895. Verbenaceae. In: Engler A, Prantl K, eds.Die Natürlichen Pflanzenfamilien, vol. 4 (3a). Berlin:Dunker und Humboldt, 132–182.

Britton NL. 1932. Ghinia spinosa. Addisonia 17: 5, pl. 547.

Britton NL, Wilson P. 1925. Scientific survey Porto Rico andVirgin Islands, vol. 6. New York: New York Academy Sci-ences.

Correl DS, Correl HB. 1982. Verbenaceae. Flora of theBahama Archipelago. Vaduz: Cramer, 1214–1252.

Gibson DN. 1970. Verbenaceae. In: Standley PC, WilliamsLO, eds. Flora of Guatemala, Fieldiana Botany 24: 167–236.

Gmelin JF. 1791. Verbenaceae. In: Systema Naturae, vol. 2,13. Lipsiae: Impensis G E Beer, 37.

Hickey LJ. 1974. Clasificación de la arquitectura de las hojasde dicotiledóneas. Bological Sociedad Argentina de Botánica16: 1–24.

Howard RA. 1983. The plates of Aublet’s histoire del plantesde la Guiane Françoise. Journal of Arnold Arboretum 64:255–292.

Howard RA. 1988. Nomenclatural notes on Antillean plants.Phytologia 65: 285–289.

Howard RA. 1989. Verbenaceae. Flora of the Lesser Antilles,Jamaica Plain, vol. 6. Jamaica Plain: Arnold Arboretum,Harvard University, 212–244.

International Union for the Conservation of Nature andNatural Resources (IUCN). 2001. Categorías y criteriosde la Lista Roja. Versión 3.1. Gland, Suiza y Cambridge:Comisión de supervisión de las especies de la IUCN.

Jansen-Jacobs M. 1988. Verbenaceae. In: Görst-van RijnARA, ed. Flora of the Guianas. Series A. Phanerogams,vol. 148. Konigstein: Koeltz Scientific Books, 3–116.

Junell S. 1934. Zur Gynäceummorphologi und Systematikder Verbenaceen und Labiaten. Symbolae Botanicae Upsa-liensis 4: 1–219.

Jussieu AL. 1789. Verbenaceae. In: Genera Plantarum. Paris:Apud Viduam Herissant et Theophilum Barrois, 106–110.

Lanjouw J. 1952. International Code of Botanical Nomen-clature. Regnum Vegetabilum 3: 1–228.

Lawrence GH. 1951. Taxonomy of vascular plants. NewYork: Macmillian.

León H, Liogier H. 1957. Verbenaceae. Contributions Oca-sionales del Museo de Historia Natural del Colegio de LaSalle 4: 279–323.

Link HF. 1820. Der botanische Garten bei Berlin. Jahrbücherder Gewächskunde 3: 51.

Lopez-Palacios S. 1977. Verbenaceae. Flora de Venezuela.Mérida: Talleres Gráficos Universitarios, 1–654.

McNeill J, Barrrie FR, Burdet HM, Demoulin V, Hawk-sworth DL, Marhold K, Nicolson DH, Prado J, SilvaPC, Skog JE, Wiersema JH. 2006. International code ofbotanical nomenclature (Vienna code). Regnum Vegetabile146: 1–568.

Martínez S, Botta S, Múlgura ME. 1996. Morfología de lasinflorescencias en Verbenaceae–Verbenoideae I: Tribu Ver-beneae. Darwiniana 34: 1–17.

Martínez S, Múlgura de Romero ME. 2003. The taxonomicposition of Parodianthus (Verbenaceae): a morphologicalsurvey of the gynoecium and inflorescence. Kew Bulletin 58:929–938.

Méndez Santos IE. 2003. Verbenaceae. In: Greuter W,Rankin Rodríguez R, eds. Flora de la República de Cuba,vol. 7. Königstein: Koeltz Scientific Books, 1–126.



Méndez Santos IE, Cafferty S. 2001. Typification of Lin-naean names of taxa of Verbenaceae s.str. described fromthe Greater Antilles. Taxon 50: 1137–1141.

Moldenke HN. 1941. New species and varieties of Verben-aceae from Central and South America. Bulletin of theTorrey Botanical Club 68: 498–506.

Moldenke HN. 1942. The known geographic distribution ofthe members of the Verbenaceae and Avicenniaceae. NewYork: Privately published.

Moldenke HN. 1981a. Notes on the genus Ghinia (Verben-aceae). Phytologia 47: 404–419.

Moldenke HN. 1981b. Additional notes on the genus Ghinia.Phytologia 47: 447–461.

Moldenke HN. 1981c. Additional notes on the genus Ghinia.Phytologia 48: 111–116.

Nash DL, Nee M. 1984. Verbenaceae. Flora de Veracruz,vol. 41. Xalapa: Instituto Nacional de Investigaciones SobreRecursos Bióticos, 1–154.

Persoon CH. 1807. Verbenaceae. In: Synopsis Plantarum,vol. 2. Paris: C. F. Cramerum, 138–147.

Rehder A, Weatherby CA, Mansfeld R, Green ML. 1935.Conservation of later generic Homonymus. Bulletin of Mis-cellaneous Information 6–9: 341–544.

Rzedowsky J, Calderón de Rzedowsky G. 2002. Flora delBajío y de regiones adyacentes. Familia Verbenaceae 100.Patzcuaro: Instituto de Ecología, 1–145.

Sanders RW. 2001. The genera of Verbenaceae in the south-eastern United States. Harvard Papers in Botany 5: 303–358.

Schauer JC. 1847. Verbenaceae. In: De Candolle AL, eds.Prodromus Systematis Naturalis Regni Vegetabilis, vol. 11.Paris: Sumptibus sociorum Truttel et Würtz, 522–700.

Schlechtendal DFL, Chamisso LKA. 1830. PlantarumMexicanarum. Linnaea 5: 72–174.

Schlechtendal DFL, Chamisso LKA. 1831. PlantarumMexicanarum. Linnaea 6: 352–384.

Schreber JCD. 1789. Ghinia. In: Genera Plantarum, 8thedn, vol. 1. Frankfurt: Varrentrapii et Wanneri, 19.

Standley PC. 1924. Trees and shrubs of Mexico. Contribu-tions from the United States National Herbarium 23:1236.

Stevens WD, Ulloa CA, Pool U, Montiel OM. 2001. Ver-benaceae. Flora de Nicaragua. Monographs in SystematicBotany from the Missouri Botanical Garden 85: 2497–2525.

Suttill TA, Allen GA. 1992. Morphological and chromosomalvariation in Dodecatheon pulchellum (Primulaceae). Cana-dian Journal of Botany 70: 2476–2483.

Swartz O. 1788. Nova genera que species plantarum;seu, Prodromus descriptionum vegetabilium. Stockholm,Sweden: Holmiae, Bibliopoliis Acad. M. Swederi.

Swartz O. 1800. Ghinia. In: Florae Indiae Occidentalis,vol. 2. Erlangae: Jacob Palmi, 1087–1090, tab xxi.

Troncoso NS. 1974. Los géneros de Verbenáceas deSudamérica Extratropical. Darwiniana 18: 295–412.

Urban I. 1911. Verbenaceae. Symbolae Antillanae 4: 529–539.

Walpers GG. 1845. Verbenaceae. In: Repertorium BotanicesSystematicae, vol. 4. Lipsiae: Sumtibus Friderici Hofmeis-ter, 5–134.

Willdenow KL. 1797. Ghinia. In: Species Plantarum, vol. 1.Berlin: Impensis G. C. Nauk, 114.

Zaruchi JL. 1984. The treatment of Aublet’s generic namesby his contemporaries and by present-day taxonomists.Journal of Arnold Arboretum 65: 215–242.



a taxonomic revision of the genus tamonea …redbiblio.unne.edu.ar/pdf/62_o'leary 2008 a...

Documents