Condea sect. Condea

Harley, R. M. & Pastore, J. F. B., 2012, A generic revision and new combinations in the Hyptidinae (Lamiaceae), based on molecular and morphological evidence, Phytotaxa 58 (1), pp. 1-55 : 13-29

Condea sect. Condea

= Hyptis sect. Minthidium Bentham (1833: 128) , syn. nov. Type:— Hyptis verticillata Jacq. , lectotype designated by Epling 1936b [= Condea verticillata (Jacq.) Harley & J.F.B.Pastore ].

= Hyptis sect. Minthidium subsect. Campanulatae Briquet (1897b: 337) , syn. nov. Type:— Hyptis fasciculata Benth. , lectotype designated here [= Condea undulata (Schrank) Harley & J.F.B.Pastore ].

= Hyptis sect. Hypenia subsect. Pubescentes Briquet (1897b: 335) , syn. nov. Type:— Hyptis floribunda Briq. , lectotype designated by Epling 1936b [= Condea floribunda (Briq.) Harley & J.F.B.Pastore ].

Condea section Condea is characterized by an indumentum of simple trichomes and with flowers solitary or in fascicles, in the axils of reduced or leaf-like bracts. In some species, the nutlets bear a swollen, corky swelling, which may act as a flotation device to aid dispersal. There are 16 species, centred in the Caribbean and in South America, with one species, Condea verticillata , more widespread, occurring in the Caribbean, Central America and western South America. See Fig. 1K .

Condea americana (Poir.) Harley & J.F.B.Pastore , comb. nov. ≡ Satureja americana Poiret in Lamarck (1805: 571) [as Satureia americana ] ≡ Hyptis americana (Poir.) Briquet (1897b: 338) [non (Aubl.) Urban 1918: 322] ≡ Mesosphaerum americanum (Poir.) Kuntze (1891: 525) . Type:— “ America” [ DOMINICAN REPUBLIC?], " Condea frutescens Satureiae foliis , flore albo, Mss. Descript. Plant. Amer.", Pouppé-Desportes s.n. (holotype P-LA!).

= Hyptis scoparia Poiteau (1806: 475 , t.31, f.2). Type :— DOMINICAN REPUBLIC. Cap Français, environs S. Martin , Poiteau s.n. (holotype P!) .

= Hyptis escobilla Urban (1919: 143) , nom. illeg., p.p. excl. type. See note under Condea urbanii Harley & J.F.B.Pastore.

Condea chyliantha (Urb. & Ekman) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis chyliantha Urb. & Ekman in Urban (1929: 48). Type:— HAITI. Massif du Nord, St. Michel de Atalaye , 21 December 1927, Ekman 9430 (holotype S!; isotypes B †, BM!, C!, G!, K!, NY!, US!) .

Condea cubensis (Urb.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis cubensis Urban (1912: 367) . Type:— CUBA. Partido de Consolación , Wright 3150 (holotype B †; lectotype designated here: K!; isolectotypes BM!, G!, K!, MA!, P!, US!) .

Condea domingensis (Urb.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis domingensis Urban (1912: 368) . Type :— DOMINICAN REPUBLIC. Prope Constanza, February 1910, Türckheim 2897 (holotype B†; lectotype designated here: NY!; isolectotypes: BR!, G!, M!).

Condea elegans (Briq.) Harley & J.F.B.Pastore , comb. nov. ≡ Eriope elegans Briquet (1889: 114) ≡ Hyptis elegans (Briq.) Briquet (1897a: 19 , t. 58). Type:— PARAGUAY. Paraguari: Paraguari, 25 March 1875, Balansa 979 (holotype G!; isotypes BM!, BR!, K!).

Condea fastigiata (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis fastigiata Bentham (1833: 130) ≡ Mesosphaerum fastigiatum (Benth.) Kuntze (1891: 526) ≡ Hyptis fasciculata var. fastigiata (Benth.) Schmidt (1858: 142) ≡ Hyptis fasciculata subsp. fastigiata (Benth.) Harley (1985a: 14) . Type:— BRAZIL. "Brasilia meridionalis" Sellow 2110 (holotype B†; lectotype K!, designated here).

= Hyptis diaphora Briquet (1896: 786) ≡ Mesosphaerum diaphorum (Briq.) Kuntze (1898: 260) . Type:— BRAZIL. Minas Gerais: Contendas, Kuntze s.n. (holotype G!).

Recent fieldwork ( Harley unpubl.) clearly indicates that C. undulata and C. fastigiata should be treated as two separate though closely related species, as Bentham (1833) originally proposed. For further details of diagnostic characters, see Harley (1985) .

Condea floribunda (Briq.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis floribunda Briquet (1897a: 18) . Type:— PARAGUAY. Cordillera: Arroyos y Esteros , July 1875, Balansa 980 (holotype G!; isotype K!, P!).

Condea mixta (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis mixta Epling (1944: 495) . Type:— MEXICO. Oaxaca: Tuxtepec, Chiltepec and vicinity, July 1940, Martinez-Calderón 251 (holotype US!; isotype UC!).

Condea rivularis (Britton) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis rivularis Britton (1920: 100) . Type:— CUBA. “Santa Clara cerca Trinidad a lo largo Río Toyaba ”, Britton & Wilson 5567 (holotype NY!).

Condea scandens (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis scandens Epling (1940: 239) ≡ Hyptis ascendens (sphalm.) Epling (1949: 246). Type:— GUATEMALA. Petén: El Paso, April 1932, Lundell 4421 (holotype MICH!; isotype F!).

Condea scoparioides (Urb.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis scoparioides Urban (1912: 366) . Type:— DOMINICAN REPUBLIC. La Vega: “prope Constanza in pineto 1200 m ”, Türckheim 3543 (holotype B†; lectotype K!, designated here; isolectotypes BM!, BR!, M!, NY!).

Condea thyrsiflora (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis thyrsiflora Epling (1936b: 237) . Type:— BRAZIL. Minas Gerais: “ Fazenda do Dias, Rio dos Peixes”, Pohl 2720 (holotype K!; isotypes BR!, OXF!, UC!, W!).

Condea trichopes (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Eriope trichopes Epling (1944: 495) ≡ Hyptis trichopes (Epling) Harley (1973: 24) . Type:— CUBA. Isle of Pines, vicinity of San Pedro, pine lands, 15−17 February 1916, Britton et al. 14469 (holotype NY!; isotypes K!, US!).

Condea undulata (Schrank) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis undulata Schrank (1822: 51−52) . Type:— BRAZIL. A plant formerly cultivated in the Munich Botanical Garden, now apparently lost. Neotype designated here: BRAZIL. São Paulo: Parque do Estado, Instituto de Botânica, 26 March 2004, Cordeiro & Harley 2805 (HUEFS!).

Schrank cultivated Hyptis undulata from seeds collected by Martius in Brazil, but apparently failed to keep a herbarium specimen. The name was included by Bentham, in the synonymy of the protologue of Hyptis fasciculata , with a question mark, and the short description fairly clearly indicates its identity.Unfortunately this means a change from the well-known name of this species.

= Hypothronia undata Schrank (1824: 85) . Type:—NOT LOCALIZED. Neotype designated here: BRAZIL. Santa Catarina: Garuva , 2 May 2008, Harley 55879 (HUEFS!).

The generic name Hypothronia was treated as doubtful by Bentham (who used the spelling Hippothronia ) and by Epling, both of whom placed it in synonymy. The generic name is based on Hypothronia undata , which was published simultaneously. (It is possible that this is a misprint for Hyptis undulata Schrank , published a few years earlier). As is the case with many Schrank names, H. undata was probably based on a cultivated specimen, and no type material has ever been located. The description given by Schrank is difficult to relate to any particular taxon, although both Bentham (1833: 130) and Epling (1949: 156) place this tentatively in the synonymy of Hyptis fasciculata [= Condea undulata ]. Following their opinion, we accept that Hypothronia undata Schrank , may well be this, and have chosen a neotype to that effect.

= Clinopodium verticillatum Vellozo (1829: 242) [nec Condea verticillata (Jacq.) Harley & J.F.B.Pastore ]. Type:— BRAZIL. Rio de Janeiro: Fl. Flumin. Icon. 6: t. 4. (1831), lectotype designated here [the original plate on parchment of "Flora Fluminensis" is in the Manuscript Section of the Biblioteca Nacional of Rio de Janeiro]. Epitype designated here: BRAZIL. Santa Catarina: Garuva, Sol Nascente , 9 May 1981, Hatschbach 43874 (MBM; isoepitypes C!, K!, MU).

The epitype is to establish the identity of Vellozo’s name as Condea undulata . It is impossible to determine which species was intended from the illustration, which could equally well have been either C. undulata or C. fastigiata .

= Hyptis fasciculata Bentham (1833: 130) , syn. nov. ≡ Mesosphaerum fasciculatum (Benth.) Kuntze (1891: 516) . Type:— BRAZIL. "Brasilia meridionalis", Sellow 2030 (holotype B†; lectotype K!, designated here).

Bentham published this name in the protologue as: “174. H. FASCICULATA (Benth: in Herb. Mus. Reg. Berol. MSS.)” Bentham travelled widely in Europe and visited many herbaria, annotating specimens. The holotype, now destroyed, was therefore in Berlin.

= Hyptis fasciculata Benth. var. tomentella Bentham (1848: 129) ≡ Hyptis fasciculata var. tomentosa Schmidt (1858: 142) . Type:— BRAZIL. Minas Gerais: “circa João Gomes”, Pohl 3768 (holotype W!).

Bentham apparently indicated a specimen from Minas Gerais for this variety by putting the words: “specimina magis tomentosa” after it. The same reference can be found also in his earlier work ( Bentham 1833: 130), in which he cites the full locality, but without providing a name.

= Hyptis eriocalyx A.St. -Hil. ex Bentham (1833: 131) ≡ Mesosphaerum eriocalyx (A.St.-Hil. ex Benth.) Kuntze (1891: 526). Type:— BRAZIL. Minas Gerais: “ruisseau de Simão Pereira”, Saint-Hilaire B1 36 (holotype P!).

= Lepechinia anomala Epling (1960: 146) . Type:— BRAZIL. Santa Catarina: “in capoeira ad Pilões, Palhoça”, Reitz & Klein 3227 (holotype UC!; isotypes HBR!, MO!).

Condea urbanii Harley & J.F.B.Pastore , nom. nov. ≡ Hyptis escobilla Urban 1919: 143 , nom. illeg. (excl. syn. Satureja americana Poir. ). Type:— DOMINICAN REPUBLIC. Santo Domingo: Puerto Plata, 16 April 1887, Eggers 1545 (lectotype K!, designated here; isolectotypes BM!, G!, M!, P!).

Although Hyptis escobilla Urb. is an illegitimate and superfluous name because the protologue included Satureja americana Poir. in synonymy, it is not typified by the type of S. americana (≡ Condea americana ) because Urban designated a different type (Art. 7.5 of the Vienna Code, McNeill et al. 2006). Moreover Urban’s description as well as the type appears to belong to a previously undescried taxon, which now requires a new name. The specimen of Hyptis escobilla in Urban’s own herbarium at B was obviosuly destroyed, and so we have selected a lectotype from K.

In spite of Urban’s description, the apparent lack of ripe fruit, both in the type material and other collections cited by Epling, raises the possibility that this might be a hybrid with Condea americana as one of the parents.

Condea verticillata (Jacq.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis verticillata Jacquin (1787b: 101) ≡ Mesosphaerum verticillatum (Jacq.) Kuntze (1891: 525) . Type:— DOMINICAN REPUBLIC. Without locality, but cultivated probably in Vienna: Jacquin, Icon. Pl. Rar.: t. 113, opp. p. 11 (1787), lectotype designated here. An epitype is also designated here: DOMINICAN REPUBLIC. Peravia, 18 November 1981, Mejia & Pimental 18188 (K!).

Dates of publication of many of the plates are uncertain, see Stafleu & Cowan (1979: 411) and Schubert (1945). Jacquin’s cultivated specimen, from seed originating in the Dominican Republic and perhaps formerly located in W, has never been found.

= Mentha hyptiformis Poir. in Lamarck (1797: 110). Type :— MARTINIQUE. Anonymous 342 (holotype P!).

= Hyptis parviflora Martens & Galeotti (1844: 186) . Type:— MEXICO. Veracruz: Cordillera, June−October 1840, Galeotti 677 (holotype BR!).

= Hyptis pringlei Fernald (1900: 565) . Type:— MEXICO. San Luis Potosí: Tamasopo canyon, 5 August 1890, Pringle 3223 (lectotype GH!, designated by Epling 1933; isolectotypes BM!, JE!, K!, M!, NY!, P!, UC!).

= Hyptis axillaris Fernald (1900: 565) . Type:— MEXICO. Puebla: Metlaltoyuca, Goldman 48 (lectotype US!, designated by Epling 1933; isolectotypes GH).

Condea sect. Laniflorae (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis sect. Laniflorae Epling (1933: 89) . Type:— Condea laniflora (Benth.) Harley & J.F.B.Pastore [= Hyptis laniflora Benth. , lectotype designated by Epling 1933].

= Hyptis sect. Umbellaria Benth. subsect. Eriocalycinae Briquet (1897b: 337) , syn. nov. Type:— Hyptis laniflora Benth. , lectotype designated here [= Condea laniflora (Benth.) Harley & J.F.B.Pastore ].

= Hyptis sect. Buddleioides subsect. Umbellatae Epling (1936b: 221) , syn. nov. ≡ Hyptis sect. Umbellatae Epling (1949: 197) . Type:— Hyptis tafallae Benth. [= Condea tafallae (Benth.) Harley & J.F.B.Pastore ].

Shrubs with usually dendroid trichomes and flowers in pedunculate or sessile, subumbellate cymes, arranged in elongate spiciform or paniculate inflorescences; flowers without a stylopodium. There are nine species from the deserts of SW United States, Mexico, especially Baja California, and south to Guatemala, and two species in western South America. See Fig. 4F .

Condea albida (Kunth) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis albida Kunth (in Humboldt, Bonpland & Kunth 1818: 319) ≡ Mesosphaerum albidum (Kunth) Kuntze (1891: 527) . Type:— MEXICO. Michoacan: Lago de Cuitzeo, Humboldt & Bonpland s.n. (holotype P!).

Condea anitae (Epling & Játiva) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis anitae Epling & Játiva (1968: 298) . Type:— MEXICO. Baja California: Sierra de la Giganta, near Portezuelo. 3 October 1965, Carter 5104 (holotype UC!; isotype BM!).

Condea decipiens (M.E.Jones) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis decipiens Jones (1933: 53) . Type:— MEXICO. Baja California: Triunfo, 6 October 1930, Jones 27299 (holotype RSA; isotypes BM!, HUH, MEXU, MO!, UC!, US!) .

Condea emoryi (Torr.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis emoryi Torrey (1861: 20) ≡ Mesosphaerum emoryi (Torr.) Kuntze (1891: 526) . Type:— USA. Arizona: The Upper Colorado (River), 14 January 1857, Newberry s.n. (holotype NY!: isotype K!).

= Hyptis lanata Torrey (1858: 129) , nom. nud. [non Pohl ex Bentham 1833].

= Hyptis palmeri Watson (1889: 68) ≡ Mesosphaerum palmeri (S.Watson) Goldman (1916: 363) . Type:— MEXICO. Guaymas: without locality, 1885, Palmer 278 (holotype GH; isotypes BM!, C!, K!, US!).

Epling (1949) cited as type of Hyptis emoryi an early Emory collection, now in the NY herbarium, from the same general locality (near Yuma), but this does not accord with the protologue. In an earlier paper, Torrey (1858) had erroneously named the Emory material Hyptis lanata , citing Bentham (1844a: 42). This must be an error for Hyptis laniflora Benth. , which is the only Hyptis species from Baja California mentioned in that publication. Although the Emory collection is actually H. emoryi , the specimen cited in the protologue is the Newberry specimen, as given above. Hyptis lanata Pohl ex Bentham (1833) is an earlier legitimate name for a Brazilian species.

Condea iodantha (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis iodantha Epling (1939: 16) . Type:— MEXICO. Sonora: Zapo, 18 November 1936, Hinton 9844 (holotype UC!; isotypes BM!, F!, G!, K!, US!) .

Condea jacobi (Fern.Alonso) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis jacobi Fernández-Alonso (2010: 127) . Type:— COLOMBIA. Santander: Vía Málaga-Bucaramanga, Vereda Buenavista, 29 June 2009, Fernández-Alonso 28193 (holotype COL!; isotypes COL!, G, HUA, K, M!, MA, MO, UIS, US) .

This recently described taxon is well illustrated in the original paper, and clearly represents a distinct new species, related to Condea tafallae , but with dendroid trichomes as in the Central American species.

Condea laniflora (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis laniflora Bentham (1844: 42, t. 20) ≡ Mesosphaerum laniflorum (Benth.) Kuntze (1891: 526) . Type:— MEXICO. Baja California: Cabo San Lucas, 1841, Hinds s.n. (holotype K!; isotype K!).

= Mesosphaerum insulare Standley & Goldman (1911: 375) ≡ Hyptis insularis (Standl. & Goldm.) Standley (1924: 1276) ≡ Hyptis laniflora var. insularis (Standl. & Goldm.) Johnston (1922: 1150) . Type:— MEXICO. Baja California: Isla Espiritu Santo, 7 February 1906, Nelson & Goldman 7503 (holotype US!; isotype UC!).

Condea subtilis (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis subtilis Epling (1933: 79) . Type :— MEXICO. State unknown: Sadovi, 1842, Liebmann 15251 (holotype C!; isotype F!).

= Hyptis perpulchra (as perpulcher) Epling (1939: 15). Type:— MEXICO. Temascaltepec: Pungarancho, 18 October 1935, Hinton et al. 8574 (holotype UC!; isotypes BM!, F!, K!, MO!, NY!, US!) .

Condea tafallae (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis tafallae Bentham (1833: 132) ≡ Mesosphaerum tafallae (Benth.) Kuntze (1891: 527) . Type:— PERU. Without locality, Ruiz s.n. [ Tafalla s.n.] (holotype OXF Lambert Herbarium†; lectotype MA!, designated here; isolectotypes B†, P!).

The specimen in Madrid appears, according to the label, to have originally been in the Lambert Herbarium (OXF).

= Hyptis tafalloides Mansfeld (1925: 288) . Type:— PERU. Ayacucho: between Tambo and Rio Apurimac, Weberbauer 5608 (lectotype B†, designated by Epling 1936b; replacement lectotype F!, designated here).

= Mesosphaerum grandiflorum Rusby (1912: 116) . Type:— BOLIVIA. La Paz: Apolo, 2 July 1902, Williams 1512 (holotype NY!; isotype K!).

The species lacks dendroid trichomes, found in other members of the section.

Condea tephrodes (A.Gray) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis tephrodes Gray (1862: 164) ≡ Mesosphaerum tephrodes (A.Gray) Kuntze (1891: 527) . Type:— MEXICO. Baja California: Cape San Lucas, August 1859 − January 1860, Xantus 72 (holotype GH; isotypes K!, NY!, US).

Condea tomentosa (Poit.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis tomentosa Poiteau (1806: 469) ≡ Mesosphaerum tomentosum (Poit.) Kuntze (1891: 527) ≡ Hyptis incana Willdenow ex Steudel (1841: 794) , nom. nud. Type:— MEXICO. Guerrero: Acapulco, Bonpland & Humboldt s.n. (holotype P!; isotypes B-W 10844-010! [Image ID 291330], P!).

Cyanocephalus (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. et stat. nov. ≡ Hyptis sect. Cyanocephalus Pohl ex Bentham (1833: 84) . Type:— Cyanocephalus lippioides (Pohl. ex Benth.) Harley & J.F.B.Pastore [= Hyptis lippioides Pohl ex Benth. , lectotype designated by Epling 1936b].

= Hyptis sect. Cyrta subsect. Rigidae Bentham (1848: 91) , nom. illeg., homonym (see Harley 1985c: 627 ) of Hyptis sect. Polydesmia Benth. subsect. Rigidae Bentham (1848: 116) .

= Hyptis sect. Cyrta subsect. Cordifoliae Bentham (1848: 94) ≡ Hyptis sect. Cyanocephalus subsect. Virgatae Epling (1936b: 275) , nom. superfl. Type:— Hyptis cardiophylla Pohl ex Benth. , lectotype designated by Epling 1936b [= Cyanocephalus cardiophyllus (Pohl. ex Benth.) Harley & J.F.B.Pastore ].

= Hyptis sect. Cyrta subsect. Lobatae Briquet (1897b: 346) . Type:— Hyptis tripartita Briq. [= Cyanocephalus tripartitus (Briq.) Harley & J.F.B.Pastore ].

= Hyptis sect. Cyrta subsect. Argenteae Briquet (1897b: 346) ≡ Hyptis sect. Cyanocephalus subsect. Argenteae (Briq.) Epling (1936b: 270) . Type:— Hyptis incana Briq. , lectotype designated by Epling 1936b [= Cyanocephalus incanus (Briq.) Harley & J.F.B.Pastore ].

= Hyptis sect. Cyanocephalus subsect. Rugosae Epling (1933: 86) . Type:— Hyptis pedalipes Griseb. [= Cyanocephalus pedalipes (Griseb.) Harley & J.F.B.Pastore ].

Epling clearly intended that Hyptis rugosa Benth. should be the type of this subsection ( Epling 1936b). However the subsectional name was published three years earlier ( Epling 1933) with only one species cited: the Cuban Hyptis pedalipes , which therefore automatically becomes the type.

= Hyptis sect. Cyanocephalus subsect. Longifoliae Epling (1936b: 270) . Type:— Hyptis tenuifolia Epling [= Cyanocephalus tenuifolius (Epling) Harley & J.F.B.Pastore ].

= Hyptis sect. Cyanocephalus subsect. Lippiastra Epling (1936b: 272) . Type:— Hyptis lippioides Pohl ex Benth. [= Cyanocephalus lippioides (Pohl ex Benth.) Harley & J.F.B.Pastore ].

Cyanocephalus is composed of herbs and subshrubs, often strongly aromatic, with usually small leaves, entire or toothed to deeply pinnatifid. It is distinguished by its pedunculate capitula, globose even when immature, with narrowly linear, soft bracteoles (see note), which form an involucre, inconspicuous at anthesis, the calyx with linear, usually clavate lobes, with an oblique throat and tube usually down-curved to deflexed, above the middle. Gynoecium without stylopodium, style capitate, with stigmatic branches usually reduced. The sectional classification presented by Epling (1949) has proved unusable ( Harley, 2006 ). There are 25 species currently recognized, mainly found in the cerrados of central Brazil, extending to eastern Paraguay and Bolivia. The type species is endemic to Cuba. See Fig. 1D, F .

Note:—The bracteolar nature of the involucres of the Hyptidinae capitulum can be demonstrated by a study of the structure of congested inflorescences in a number of species with less compact inflorescence units, where the flowers and bracteoles subtending them become reoriented, the flowers moving to an upper position related to the inflorescence axis, and the bracteole to a lower position, with foreshortening of the axes separating them (cincinnus or drepanium).

Cyanocephalus adpressus (A.St.-Hil. ex Benth.) Harley & J.F.B.Pastore, comb. nov. ≡ Hyptis adpressa A.St. - Hil. ex Bentham (1833: 84) ≡ Mesosphaerum adpressum (A.St.-Hil. ex Benth.) Kuntze (1891: 526). Type:— BRAZIL. Goiás: in campis prope Forquilla, St.-Hilaire C1 853 (holotype P!; isotypes F!, US!).

Cyanocephalus apertiflorus (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis apertiflora Epling (1936b: 273) . Type :— BRAZIL. Locality unknown, Sellow s.n. (holotype B†; lectotype UC!, designated here; isolectotype ZT!).

Cyanocephalus bombycinus (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis bombycina Epling (1936b: 276) . Type:— BRAZIL. Mato Grosso: Coxim, Hoehne 2852 (holotype UC!).

Cyanocephalus caprariifolius (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis caprariifolia (as “ caprariaefolia ”) Pohl ex Bentham (1833: 83) ≡ Mesosphaerum caprariifolium (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: Cristalina “Serra dos Chrystais”, Pohl 1020 (lectotype W!, designated by Epling 1936b; isolectotypes K!, F!, UC!).

Cyanocephalus cardiophyllus (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis cardiophylla Pohl ex Bentham (1833: 84) ≡ Mesosphaerum cardiophyllum (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: “Rio Urubu, Ouro Fino, Santa Luzia, Cap. Goyaz et ad Paracatu Cap. Min. G.” Pohl 958 (lectotype W!, designated by Epling 1936b; possible isolectotype K!).

The label data on the type specimen in W indicates a range of specimens, from both Goiás and Minas Gerais. Epling (1949) cites the type as from Minas Gerais state, although there is no way to determine where the specimen came from .

Cyanocephalus coriaceus (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis coriacea Bentham (1848: 95) ≡ Mesosphaerum coriaceum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: Diamantina, July 1840, Gardner 5090 (holotype K!; isotype G!, OXF!, W!, B† photo!).

Cyanocephalus cretatus (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis cretata Epling (1936b: 276) . Type:— BRAZIL. Minas Gerais: Uberaba, Riedel & Lund 2430 (holotype UC!; isotypes K!, LE!, NY!, US!) .

Cyanocephalus cuneatus (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis cuneata Pohl ex Bentham (1833: 86) ≡ Mesosphaerum cuneatum (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: Rio Parahybuna, Pohl 624 (holotype W!; isotype K!).

In the protologue Bentham does not mention a specimen at K, as he normally did. In this case he merely states: “Hab. in Brasilia: in campis desertorum ad Rio Parahybuna provinciae Minas Gerais, Pohl! (v.s.sp. in herb. Mus. Caes. Reg. Bras. Vind.)”. Bentham travelled widely in Europe, visiting many herbaria, whose specimens he annotated, and sometimes received duplicates of material at a later date.

= Hyptis clavellifera Bentham (1848: 91) ≡ Mesosphaerum clavelliferum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: between Arraias and São Domingos, May 1840, Gardner 4307 (holotype K!; isotypes OXF!, B† photo!).

Cyanocephalus delicatulus (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis delicatula Harley (1985c: 630) . Type:— BRAZIL. Bahia: Serra do Sincorá, S of Mucugê, by Rio Paraguaçu, 7 February 1979, Harley et al. 16103 (holotype CEPEC!; isotypes AAU!, E!, IPA!, K!, NY!, P!, U!, US!, SPF!).

Cyanocephalus desertorum (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis desertorum Pohl ex Bentham (1833: 83) . Type:— BRAZIL. Minas Gerais: Rio Parahybuna, Pohl 542 (holotype W!).

= Hyptis lasiocalyx Pilger (1901: 190) . Type:— BRAZIL. Mato Grosso: Cuiabá, 22 April 1849, Meyer 512 (holotype B†).

Cyanocephalus digitatus (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis digitata Harley (1985c: 632) . Type:— BRAZIL. Goiás: Alto Paraíso de Goiás, Chapada dos Veadeiros, 28 September 1975, Hatschbach & Kummrow 37241 (holotype MBM!; isotypes K!, MO!).

Cyanocephalus incanus (Briq.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis incana Briquet (1889: 111) [non Hyptis incana Willdenow ex Steudel (1841: 794) , nom.nud.] ≡ Hyptis albicoma Epling (1936b: 276) , nom. superfl. Type:— PARAGUAY. Guaira: entre Santa Bárbara y Borja, Balansa 1001 (holotype G!; isotypes K!, P!).

Epling erected a new name for Hyptis incana Briq. , believing incorrectly that the earlier Willdenow name was valid.

Cyanocephalus lanatus (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis lanata Pohl ex Bentham (1833: 84) ≡ Mesosphaerum lanatum (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: Água Quente et Trahiras, Pohl 1879 (holotype W!; isotypes K!, W!).

Cyanocephalus lippioides (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis lippioides Pohl ex Bentham (1833: 86) ≡ Mesosphaerum lippioides (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: Chapada São Marcos ad Cabesseira Ribeirão Batalha, Pohl 2866 (holotype W!; isotype K!).

= Hyptis fragilifolia A.St. -Hil. ex Bentham (1833: 85) ≡ Mesosphaerum fragilifolium (A.St.-Hil. ex Benth.) Kuntze (1891: 526). Type:— BRAZIL. Minas Gerais: Penha, St.-Hilaire 1164 (holotype P!).

= Hyptis nervosa Pohl. ex Bentham (1833: 85) ≡ Mesosphaerum nervosum (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: “inter Estiva et Rio Preto”, Pohl s.n. (holotype W!; isotype K!).

There are two specimens on the type sheet of H. nervosa in W, of which the right hand specimen is labelled as type. Below it is a label listing three different collection numbers, at the top right hand side and the locality data: “Inter Estiva et Rio da Prata , inter Calumbis et Barreros, Cap. Minas Geraes”. The left hand specimen is unlabelled, but could well be part of the same collection or a different one from the other .

= Hyptis candida Pohl ex Bentham (1833: 85) . Type:— BRAZIL. Minas Gerais: “inter Rio Jequitinhonha et Barreros”, Pohl 6170 (holotype W!; isotype K!).

= Hyptis rigida Pohl. ex Bentham (1833: 85) ≡ Mesosphaerum rigidum (Pohl ex Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Minas Gerais: “inter Calumbis et Barreros”, Pohl 6171 (holotype W!).

= Hyptis subnuda Briquet (1898: 231) ≡ Mesosphaerum subnudum Briquet (1898: 231) . Type:— BRAZIL. São Paulo: Bocaina, Glaziou 11308 (holotype G!; isotype LE!, P!).

Cyanocephalus nitidulus (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis nitidula Bentham (1848: 91) ≡ Mesosphaerum nitidulum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: Chapada da Mangabeira, September 1839, Gardner 3392 (holotype K!; isotypes BR!, K!, OXF!).

Cyanocephalus pedalipes (Griseb.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis pedalipes Grisebach (1866: 213) ≡ Mesosphaerum pedalipes (Griseb.) Kuntze (1891: 526) . Type:— CUBA. “Occidentalis”, unlocalized, Wright 3152 (holotype GOET; isotypes NY!, P!, G!, K!).

Cyanocephalus peduncularis (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis peduncularis Bentham (1833: 87) ≡ Mesosphaerum pedunculare (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: Volta de Serra, October 1818, Sellow (lectotype B†, designated by Epling 1936b, replacement lectotype K!, designated by Harley 2006 ).) For comment on type locality and date of collection see Harley (2006) .

= Hyptis camporum Bentham (1848: 92) ≡ Mesosphaerum camporum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: Diamantina, July 1840, Gardner 5091 (holotype K!; isotypes W!, OXF!).

Cyanocephalus poliodes (Briq.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis poliodes Briquet (1897: 35) . Type:— PARAGUAY. Cordillera: Valenzuela, 18 January 1884, Balansa 4557 (holotype P!).

Epling (1949) wrongly gives the name as Hyptis polioides Briq.

Cyanocephalus rugosus (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis rugosa Bentham (1833: 86) ≡ Mesosphaerum rugosum (Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Minas Gerais: Saquinho, Sellow 1478 (lectotype B†, designated by Epling 1936b, replacement lectotype K!, designated here).

= Hyptis albipes A.St. -Hil. ex Bentham (1833: 88) ≡ Mesosphaerum albipes (A.St.-Hil. ex Benth.) Kuntze (1891: 526). Type:— BRAZIL. Minas Gerais: Rio Salgado, St.-Hilaire 1856 (holotype P!; isotype F! fragment).

= Hyptis arguta Pohl ex Bentham (1833: 86) ≡ Mesosphaerum argutum (Pohl ex Benth.) Kuntze (1891: 526) ≡ Hyptis rugosa var. villosissima Schmidt (1858: 94) . Type:— BRAZIL. Minas Gerais: Rio Parahybuna, Pohl 539 (holotype W!).

= Hyptis bisdentata Pohl ex Bentham (1833: 87) . Type:— BRAZIL. Minas Gerais: Fazenda Tallaio, inter Piedade et Fanado , Pohl s.n. (holotype W!; isotype K!). The label on the type sheet has three numbers: 2344, 3044, 540.

= Hyptis brunnescens Pohl ex Bentham (1833: 83) ≡ Mesosphaerum brunnescens (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: Engenho dos Bois, Pohl 1361 (holotype W!; isotype K!).

= Hyptis incisa A.St. -Hil. ex Bentham (1833: 87) ≡ Mesosphaerum incisum (A.St.-Hil. ex Benth.) Kuntze (1891: 526) ≡ Hyptis rugosa var. incisa (A.St.-Hil. ex Benth.) Epling (1936b: 274). Type:— BRAZIL. Minas Gerais: Salgado, St.-Hilaire 1857 (lectotype P!, designated by Epling 1936b).

= Hyptis longipes A.St. -Hil. ex Bentham (1833: 88) ≡ Mesosphaerum longipes (A.St.-Hil. ex Benth.) Kuntze (1891: 526). Type:— BRAZIL. Minas Gerais: Minas Novas, “in sylvis siccis prope pagum Sucuriu”, St.- Hilaire 1392 (holotype P!; isotype K!).

= Hyptis reflexa A.St. -Hil. ex Bentham (1833: 83) ≡ Mesosphaerum reflexum (A.St.-Hil. ex Benth.) Kuntze (1891: 527). Type:— BRAZIL. Minas Gerais: Araxá, St.-Hilaire 461 (holotype P!; isotype F!).

= Hyptis araripensis Bentham (1848: 93) ≡ Mesosphaerum araripense (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Ceará: Serra do Araripe, October 1838, Gardner 1805 (holotype K!: isotypes OXF!, P!, W!, MANCH!).

= Hyptis subsessilis Bentham (1848: 92) . Type:— BRAZIL. Piaui: Serra da Batalha, May 1840, Gardner 2927 (lectotype K!, designated by Epling 1936b; isolectotype OXF!).

= Hyptis viscidula Bentham (1848: 92) ≡ Mesosphaerum viscidulum (Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Minas Gerais: between Rio Claro and São Romão, June 1840, Gardner 5088 (lectotype K!, designated by Epling 1936b; isolectotype G!, OXF!).

As indicated by the extensive synonymy and the number of varieties which have been recognized, the species is morphologically extremely diverse. Epling (1949) recognized three varieties, but subsequent collections make these of doubtful value. A full study is required to determine the status of these forms.

Cyanocephalus selaginifolius (Mart. ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis selaginifolia Mart. ex Bentham (1833: 87) ≡ Mesosphaerum selaginifolium (Mart. ex Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Minas Gerais: “in campis, Serra de Santo Antonio”, Martius s.n. (holotype M!).

Cyanocephalus tacianae (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis tacianae Harley (2006: 95) . Type :— BRAZIL. Distrito Federal : Samambaia , Parque Boca da Mata, 28 May 1998, Rezende 410 (holotype CEN!; isotypes K!, HUEFS!, UFG!).

Cyanocephalus tagetifolius (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis tagetifolia Harley (1974: 134) . Type:— BRAZIL. Goiás: ca. 20 km N of Alto Paraíso de Goiás, 25 March 1971, Irwin et al. 33180 (holotype K!; isotypes NY!, UB!).

Cyanocephalus tenuifolius (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis tenuifolia Epling (1936b: 270) . Type:— BRAZIL. São Paulo: “inter São Simão et Casa Branca”, 1885, Regnell III 923 (holotype S!).

Cyanocephalus tripartitus (Briq.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis tripartita Briquet (1889: 111) . Type:— PARAGUAY. Caaguazú: Caaguazú, Balansa 999 (holotype G!; isotypes BM!, K!, P!, S!).

Cyanocephalus viaticus (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis viatica Harley (1985c: 628) . Type :— BRAZIL. Minas Gerais: BR 4 , km 878−879, 30 January 1965, Pereira 9807 (holotype HB!; isotype K!) .

Eplingiella Harley & J.F.B.Pastore , gen. nov. Hyptidendro similis sed gynoecio sine stylopodio, et corollis forma propria lobis patentibus azureo-violaceis coloratis, et ab omnibus generibus Hyptidinarum combinatione foliiorum parvorum xeromorphorum et floribus brevemente pedicillatis in cymulis plusminusve paucifloribus bracteolatis, bracteolis parveolis inconspicuis, calycibus fructificantibus aut actinomorphis lobis curtis fauce barbato trichomatibus albis aut zygomorphis tubo ad medium deflexo, fauce non barbato, lobis subulatis haud clavatis differt. Type:— Eplingiella fruticosa (Salzm. ex Benth.) Harley & J.F.B.Pastore [= Hyptis fruticosa Salzm. ex Benth ].

This genus is characterized by its shrubby habit, with small xeromorphic leaves, the flowers in 2−18- flowered pedunculate cymes, subtended by bracts similar to leaves, of shortly pedicellate flowers with small, inconspicuous, narrowly linear bracteoles. Corolla characteristic, with spreading lobes, blue to violet-blue. Fruiting calyx variable, that of E. cuniloides actinomorphic with short lobes and with dense white trichomes in throat and E. fruticosa zygomorphic, with mid-tube strongly curved and lobes subulate, not clavate and throat not bearded. Gynoecium without a stylopodium, style capitate or with stigmatic lobes very short. Nutlets narrowly ellipsoid, dark brown, strongly mucilaginous when wet.

There are two species at present recognized, from semi-arid, sandy areas in the upland interior of Northeast Brazil, the type species descending to coastal sands and the other endemic to Morro do Chapéu, Chapada Diamantina, Bahia. See Fig. 3C .

Both species of Eplingiella were originally included by Epling (1936b) in Hyptis sect. Umbellaria , together with a group of other species, later removed to Hyptidendron sect. Umbellaria ( Harley 1988) , and characterized by the possession of a well developed stylopodium. Lacking this feature, the two species mentioned above remained unranked for a number of years, as they did not appear to fit in any of the sections of Hyptis recognized at that time. The molecular analysis, presented by Pastore et al. (2011), has shown that they form a distinct clade and justifies their recognition at generic level. The genus is named in honour of Carl Epling, whose contribution to the study of the New World Lamiaceae is immense.

Eplingiella cuniloides (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis cuniloides Epling (1947: 517) . Type:— BRAZIL. Bahia: Morro do Chapéu, April 1944, Schery 587 (holotype MO!; isotype UC!).

Eplingiella fruticosa (Salzmann. ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis fruticosa Salzm. ex Bentham (1833: 123) ≡ Mesosphaerum fruticosum (Salzm. ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Bahia: “in collibus aridis circa Bahiam” (Salvador), 1830, Salzmann s.n. (lectotype K!, designated by Epling 1936b; isolectotypes BR!, E!, G!, HAL!, P!).

Eriope Humb. & Bonpl. ex Bentham (1833: 142) . Type:— Eriope nudiflora Humb. & Bonpl. ex Benth. [= Eriope crassipes Benth. ].

= Hyptis sect. Siagonarrhen Mart. ex Benth. subsect. Nudiflorae Bentham (1848: 134) , syn. nov. Type:— Hyptis latifolia Mart. ex Benth. , lectotype designated by Epling 1936b [= Eriope latifolia (Mart. ex. Benth.) Harley ].

= Hyptis sect. Mixtae (Epling) Epling (1949: 215) , syn. nov. Type:— Hyptis salviifolia Pohl ex. Benth. [ Eriope salviifolia (Pohl ex. Benth.) Harley ].

Trees, shrubs or subshrubs, often aromatic; stems often geoxylic, sometimes virgate, often with conspicuous white, spreading setose hairs and a few species as in Hypenia with pruinose, fistulose internodes below inflorescence. Inflorescence elongate, raceme-like, simple or branched, with 1-flowered pedunculate cymes from the axils of caducous bracts, the peduncle appearing as a pedicel (pseudo-pedicel), but with a pair of usually inconspicuous bracteoles at its apex, at base of calyx. Calyx turbinate to campanulate, 5-lobed, the posterior lobes often becoming connate and indistinct in fruit, when throat closed by dense white hairs in many species; corolla strongly 2-lipped, violet or pink and often yellowish in bud, tube widening above and sometimes dorsally gibbous, usually constricted near base, posterior lip internally with dark striae and a whitish area at base. Gynoecium with conspicuous persistent stylopodium overtopping ovary. Nutlets slightly flattened to ovoid, rarely narrowly winged.

There are over 30 species occurring in savannas, especially cerrados of Central Brazil and in campo rupestre in mountain areas of Eastern Brazil, and in campinas in Amazonia, with a few species extending into neighbouring countries. See Fig. 2E .

Harley (1976) revised the genus, raising the number of species recognized by Epling (1936) from 18 to 20, by various taxonomic changes, reducing several to synonymy, describing four new species and transferring three species from Hyptis . Later ( Harley 1992 ) further changes and additions were made.

Eriopidion Harley (1976: 103) ≡ Eriope sect. Tubiflorae Epling (1936a: 190) . Type:— Eriopidion strictum (Benth.) Harley [= Eriope stricta Bentham (1848: 142) ].

The only species is a low perennial herb, sometimes slightly woody at base, strongly aromatic. It shares many of the characters found in species of Eriope , including the slender raceme-like inflorescence composed of 1-flowered cymes, but differs in the persistent bracts, the narrowly campanulate calyx with a broad hygroscopic posterior lobe, which folds when dry to close mouth of calyx, a gynoecium with stylopodium absent and with ±triquetrous nutlets.

A monotypic genus ( Harley 1976 ), with Eriopidion strictum a rare plant of dry, sandy areas within the caatinga zone of NE Brazil, reappearing in semi-arid vegetation by the lower Orinoco River, in Venezuela. See Fig. 1J .

Gymneia (Benth.) Harley & J.F.B.Pastore , comb. et stat. nov. ≡ Hyptis Sect. Gymneia Bentham (1833: 77−78) . Type:— Gymneia platanifolia (Mart. ex Benth.) Harley & J.F.B.Pastore [= Hyptis platanifolia Mart. ex Benth. , lectotype designated by Epling 1936b].

= Hyptis sect. Spiciformes Schmidt (1858: 83) , syn. nov. Type:— Hyptis platanifolia Mart. ex Benth. , lectotype designated here [= Gymneia platanifolia (Mart. ex Benth.) Harley & J.F.B.Pastore ].

Gymneia is composed of herbs or subshrubs, characterized by terminal inflorescences of elongate, congested or interrupted spikes, with flowers arranged in ± globose verticillasters in the axils of reduced bracts, the verticillasters formed of strongly congested cincinni with filiform bracteoles. Flowers with corollas small, fruiting calyx with strongly curved tube and oblique mouth, gynoecium without stylopodium, style ± capitates, stigma lobes much reduced. A very homogeneous genus of at least six species, occurring in the cerrados of central Brazil, with one species occurring in waste places in the caatingas of NE Brazil and disjunctly in eastern Bolivia, where it may have been introduced. See Figs. 2A , 3A .

Gymneia ampelophylla (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis ampelophylla Epling (1936b: 279) . Type:— BRAZIL. Tocantins: Porto Nacional [“ Goiás, Porto Real”], 21 February 1829, Burchell 8671-2 (holotype K!; isotypes K!).

Gymneia interrupta (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis interrupta Pohl ex Bentham (1833: 77) ≡ Mesosphaerum interruptum (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: Santa Cruz, Pohl 2758 (lectotype W!, designated by Epling 1936b; isolectotypes K!, W!).

= Hyptis ovalifolia Bentham (1848: 87) ≡ Mesosphaerum ovalifolium (Benth.) Kuntze (1891: 526) , syn. nov. Type:— BRAZIL. Goiás: Serra Dourada, Pohl 1497 (holotype W!; isotype K!).

The extensive collections now available show a range of intermediates between Hyptis ovalifolia and G. interrupta , especially in the interrupted or congested spikes of vertillasters and in the variable leaf shape, previously considered diagnostic, which requires that the two species be united.

Gymneia malacophylla (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis malacophylla Bentham (1848: 86) ≡ Mesosphaerum malacophyllum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Tocantins: between Natividade & Arraias, February1841, Gardner 3930 (holotype K!; isotype G!, OXF!).

Gymneia platanifolia (Mart. ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis platanifolia Mart. ex Bentham (1833: 77) ≡ Mesosphaerum platanifolium (Mart. ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Bahia: “in campis ad Joazeiro”, Martius Obs. 2361 (lectotype M!, designated by Epling 1936b; isolectotype M!).

Gymneia virgata (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis virgata Bentham (1833: 77) ≡ Mesosphaerum virgatum (Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Minas Gerais: Sellow 1497 (lectotype B†, designated by Epling 1936b, replacement lectotype K!, designated here).

Hypenia (Mart. ex Benth.) Harley (1988: 91) ≡ Hyptis sect. Hypenia Mart. ex Bentham (1833: 136) . Type:— Hypenia reticulata (Mart. ex Benth.) Benth. [= Hyptis reticulata Mart. ex Benth. , lectotype designated by Epling 1936b].

The genus is composed of shrubs or subshrubs of virgate habit, usually slightly aromatic, with erect, virgate stems and with at least the upper internodes pruinose, fistulose and sometimes slightly inflated, lower stems often with conspicuous white, spreading setose hairs. The inflorescence lax or congested and composed of 1-flowered cymes, rarely 3-flowered, the peduncle appearing as a pedicel, but with a pair of narrow and inconspicuous or broad, persistent bracteoles at base of calyx. In many species the peduncles are elongate and very slender, while others have short peduncles and congested inflorescences. Calyx subactinomorphic, campanulate to tubular and distinctly 5-lobed with subequal lobes; corollas pale blue, lilac, red or yellow, small with short tube, to large, and then with elongate tube, tube usually contracted near base and with relatively short lobes; gynoecium with a very short stylopodium, much shorter than the ovary; nutlets ovoid. The species with a lax inflorescence of large red corollas, many of which have resupinate flowers (Atkinson, unpublished thesis), appear to be adapted to bird pollination, and are often visited by humming-birds.

The genus, with ca. 23 species, is typical of the upland lateritic savannas or cerrados and the shallow sandy soils of the campo rupestres in Goiás and the Serra do Espinhaço range of Eastern Brazil, extending to eastern Paraguay and Bolivia, with one species widespread in the caatingas of NE Brazil and with records from the Guayana Highlands of Venezuela. See Figs. 4C, E .

Harley (1988) erroneously published the combination Hypenia durifolia (Epling) Harley. The correct name should be based on the earlier Hyptis sclerophylla Epling , which is illegitimate under that genus.

Hypenia sclerophylla (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis sclerophylla Epling (1936b: 234) ≡ Hyptis durifolia Epling (1949: 235) ≡ Hypenia durifolia (Epling) Harley (1988: 92) . Type:— BRAZIL. Goiás: between Meiaponte (Pirenópolis) and Caisara, 23 October 1827, Burchell 6325 (holotype K!).

This is to correct the illegitimate combination made by Harley (1988) .

Hypenia simplex (A.St.-Hil. ex Benth.) Harley & J.F.B.Pastore, comb. nov. ≡ Hyptis simplex A.St. -Hil. ex Bentham (1833: 138) ≡ Mesosphaerum simplex (A.St.-Hil. ex Benth.) Kuntze (1891: 527) ≡ Eriope simplex (A.St.-Hil. ex Benth.) Harley (1988: 93) . Type:— BRAZIL. Goiás: “inter saxa in Serra Dourada”, St.-Hilaire 775 (holotype P!)

= Hyptis campanulata Bentham in (1848: 137) ≡ Mesosphaerum campanulatum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Tocantins: Serra da Natividade , January 1840, Gardner 3923 (holotype K!).

The molecular data ( Pastore et al. 2011) place this species as sister to the rest of Hypenia . This position is confirmed by an examination of the gynoecium structure, where the stylopodium is very small or absent, as in Hypenia and unlike that in Eriope where it overtops the nutlets. The species was originally removed from Hyptis to Eriope ( Harley 1988) , due to the similarity of its corolla, which bears a remarkable likeness to those of the latter genus.

Hyptidendron Harley (1988: 90) ≡ Hyptis Sect. Buddleioides Bentham (1833: 132) . Type:— Hyptis membranacea Benth. , lectotype designated by Epling 1936b [= Hyptidendron asperrimum (Spreng.) Harley ]. = Hyptis sect. Siagonarrhen Mart. ex Bentham (1833: 133) ≡ Hyptis sect. Siagonarrhen subsect. Cymosae Bentham (1848: 133) . Type:— Hyptis scabra Benth. , lectotype designated by Epling 1936b [= Hyptidendron canum (Benth.) Harley ].

= Hyptis sect. Latiflorae Epling (1936a: 224) , syn. nov. Type:— Hyptis eximia Epling.

The genus is composed of trees, shrubs or subshrubs, of varied habit, with an indumentum on leaves and stems of simple to in some species dendroid hairs, and with small to often large, coriaceous, sometimes aromatic leaves. The inflorescence is composed of thyrsoid, often paniculate flowers arranged in dichasial or monochasial cymes in the axils of usually foliar bracts, these sometimes conspicuous with white or purplish hairs. The pedicellate flowers have inconspicuous bracteoles, and an actinomorphic to weakly zygomorphic calyx with tube cylindrical to infundibuliform, straight, 5-lobed with subequal lobes; corollas bluish purple to lilac or rarely white, the tube cylindrical to infundibuliform; gynoecium with stylopodium overtopping ovary; nutlets ovoid to slightly flattened or narrowly winged.

The genus contains about 18 species, all occurring in S America from the Guayana Highlands and the Andes to Central Brazil, Bolivia and Paraguay. Several species are small trees or shrubs in the cerrados of Central Brazil ( Harley 1988 ), and H. arboreum Benth. ) Harley can grow up to 20 m high in relictual montane forests in Northern and Western S America. See Figs. 1C, E .

Molecular studies ( Pastore et al. 2011) indicate that Hyptis eximia Epling , placed by Epling in a monotypic section Latiflorae , falls within this genus, making the following new combination necessary. The molecular data shows some conflict with the subsections recognized by Harley (1988) . Until fuller data are available, it is proposed not to recognize them here.

Hyptidendron eximium (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis eximia Epling (1936a: 223) . Type:— BRAZIL. Mato Grosso: Barão de Melgaço , June 1918, Kuhlmann 2279 (holotype UC!).

Hyptis Jacquin (1787a: 101) , nom. cons. ( Briquet 1906). Type:— Hyptis capitata Jacq. , typ. cons. (lectotype designated by Green 1929: 107).

Hyptis Jacq. was conserved against Condea Adans. and Mesosphaerum P.Browne. However the types of these two genera no longer fall within our concept of Hyptis , so the names again become available.

A genus of annual or perennial herbs, subshrubs, or shrubs, of variable habit, with flowers in pedunculate to sessile, cymose capitula with an involucre of bracteoles, and borne singly in the axils of foliaceous or reduced bracts and sometimes forming complex synflorescences. Flowers usually small, with narrow, cylindrical calyx, often accrescent in fruit, usually with subequal lobes; corolla small often white or occasionally lilac and with spotted posterior lip, tube cylindrical; Nutlets ovoid or narrowly ellipsoid.

A genus of around 144 species, distributed in tropical and subtropical zones from North America to the Caribbean and southward to Argentina and Peru, often occurring in humid savannas. A few species extend to the Old World tropics, mainly as weeds. See Figs. 1H , 2C , 3B, 3D , 4G .

The molecular studies ( Pastore et al. 2011) on which the taxonomic changes provided in this paper are largely based, require a much reduced genus Hyptis . Of the 26 sections previously recognized by Epling (1949), only ten remain, with a few, such as Hyptis sections Induratae and Pachyphyllae now shown to fall within H. sect. Eriosphaeria Benth. Hyptis sect. Polydesmia , partially dismembered, and a sister group to the reduced Hyptis clade, is now treated at generic level as Cantinoa and includes several species formerly in H. sect. Mesosphaeria . The genus Peltodon is shown to be grouped within Hyptis . There are still questions to be answered with respect to inter-relationships within this remodelled Hyptis , but it is hoped that future studies, with much wider sampling, will enable a reevaluation of its infrageneric classification. The ten sections of Hyptis remaining, which need further investigation, are: Hyptis sect Hyptis , H. sect. Pusillae Epling, H. sect. Hilaria , H. sect. Cyrta , H. sect. Plagiotis Benth., H. sect. Myriocephala Benth., H. sect. Muellerohyptis Briq., H. sect. Xylodontes , H. sect. Apodotes Benth. and H. sect. Eriosphaeria . The only changes we propose at this time are as follows:

Hyptis section Peltodon (Pohl) Harley & J.F.B.Pastore , comb. & stat. nov. ≡ Peltodon Pohl (1827: 66) . Type:— Peltodon radicans Pohl , lectotype designated by Epling 1936b [= Hyptis radicans (Pohl) Harley & J.F.B.Pastore ].

Epling (1936b) wrote: “A genus scarcely separate from Hyptis and coordinate with its sections”—a comment confirmed by the molecular data ( Pastore et al. 2011).

Hyptis sect. Peltodon is easily recognized by the small, spreading, foliaceous appendage at the apex of each calyx lobe. The flowers are arranged in robust, subglobose capitula, with a conspicuous involucre of broad, often coloured bracteoles.

Five species are known, mainly occurring in cerrado or other similar savanna formations or in margins of Atlantic forest, in eastern and southern Brazil extending into eastern Paraguay and Argentina. See Fig. 1H .

Hyptis campestris Harley & J.F.B.Pastore , nom. nov. ≡ Peltodon tomentosus Pohl (1827: 69) [non Hyptis tomentosa Poiteau 1806 ]. Type:— BRAZIL. Minas Gerais: “Fazenda de Almas, in via de Paracatu do Príncipe ad Rio S. Antônio”, Pohl 3352 (holotype W!; isotype K!).

Hyptis comaroides (Briq.) Harley & J.F.B.Pastore , comb. nov. ≡ Peltodon comaroides Briquet. (1889: 110) . Type:— PARAGUAY. Guairá: “Itapé dans les prairies”, Balansa 1008 (holotype G!; isotype K!).

= Peltodon longipes A.St. -Hil. ex Bentham (1833: 63) [non Hyptis longipes A.St. -Hil. ex Bentham 1833: 88]. Type:— BRAZIL. Rio Grande do Sul: “ad rivulos prope Santa (?São) Borja in Missionibus Uruguayensibus”, St.-Hilaire 2656 (holotype P!).

Hyptis meridionalis Harley & J.F.B.Pastore , nom. nov. ≡ Peltodon rugosus Tolmatchew (1923: 62) [non Hyptis rugosa Bentham 1833: 86 ]. Type:— BRAZIL. Paraná: in campis montosis, prope Castro, March 1826, Riedel 361 (holotype LE!).

Epling (1936b) gives Dusén s.n. from Paraná ( S) as holotype, but this is not in accord with the protologue !

Hyptis pusilla (Pohl) Harley & J.F.B.Pastore , comb. nov. ≡ Peltodon pusillus Pohl (1827: 67) . Type:— BRAZIL. Tocantins (as “Goyaz”): “circa Trahiras et Natividade”, Pohl 2407 (holotype W!; isotype K!).

Hyptis radicans (Pohl) Harley & J.F.B.Pastore , comb. nov. ≡ Peltodon radicans Pohl (1827: 68) . Type:— BRAZIL. Minas Gerais: Ouro Preto [“Villa Ricca”], December 1820 – January 1821, Pohl 3640 (holotype W!; isotype K!).

= Clinopodium repens Vellozo (1829: 242) [non Clinopodium repens Roxburgh (1814: 44 , nom.nud.) 1832: 13. ≡ Peltodon repens (Vell.) Kuntze (1898: 260) . Type:— BRAZIL. Rio de Janeiro: Fl. Flumin. Icon. 6: t. 7 (1831).

Epitype designated here: BRAZIL. São Paulo: Ubatuba, Picinguaba , 6 February 1988, Ribeiro & Cunha 177 ( HUEFS!) .

Leptohyptis Harley & J.F.B.Pastore , nom. et stat. nov. ≡ Hyptis sect. Minthidium Benth. subsect. Tubulosae Briquet (1897b: 377) ≡ Hyptis sect. Polydesmia Benth. subsect. Tubulosae (Briq.) Harley (1985b: 615) . Type:— Leptohyptis macrostachys (Benth.) Harley & J.F.B.Pastore [= Hyptis macrostachys Benth. , lectotype designated by Harley 1985b ].

= Hyptis sect. Leptostachys Epling (1936b: 262) ≡ Hyptis sect. Polydesmia Benth. subsect. Leptostachys Epling (1949: 298) [non Leptostachys G.Mey. (1818): Poaceae ] Type:— Hyptis macrostachys Benth. , lectotype designated by Epling 1936b [= Leptohyptis macrostachys (Benth.) Harley & J.F.B.Pastore ].

An easily recognized genus, of slender shrubs bearing elongate, slender spikes of sessile cymes, distinguished especially by the delicate triangular scale in the sinus between each lobe of the calyx, and by the slender, tubular corollas with short lobes.

There are five species, mostly restricted to drier montane areas of NE Brazil, in Bahia, Pernambuco and Minas Gerais. See Figs. 1L , 4D .

Leptohyptis calida (Mart. ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis calida Mart. ex Bentham (1833: 131) ≡ Mesosphaerum calidum (Mart. ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: “in Serra Santo Antonio in deserto Serro Frio”, Martius s.n. (holotype M!; isotypes UC!, K!).

Leptohyptis leptostachys (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis leptostachys Epling (1936b: 263) . Type:— BRAZIL. Minas Gerais: Ouro Preto, 1882, Glaziou 14194 (holotype K!; isotype P!).

subsp. leptostachys

subsp. caatingae (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis leptostachys Epling subsp. caatingae Harley (1985b: 616−617) . Type:— BRAZIL. Bahia: 10 km from town, on road to Marcolino Moura, roadside through caatinga, 25 March 1977, Harley et al. 20012 (holotype CEPEC!; isotypes AAU!, IPA!, K!, NY!, SPF!, U!, UEC!, US!). Leptohyptis macrostachys (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis macrostachys Bentham (1848: 130) ≡ Mesosphaerum macrostachyum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Bahia: Jaguarari, Serra Jacobina, Blanchet 2582 (lectotype K!, designated by Epling 1936b; isolectotypes BM!, E!, G!, K!, P!, US!).

Leptohyptis pinheiroi (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis pinheiroi Harley (1985b: 623−624) . Type:— BRAZIL. Bahia: Umburanas, Delfino, Serra do Curral Feio, 16 km NW of Lagoinha on the side of road to Minas do Mimoso, 950−1000m, 4 March 1974, Harley et al. 16689 (holotype CEPEC!; isotypes K!, NY!).

Leptohyptis siphonantha (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis siphonantha Harley (1974: 132) . Type:— BRAZIL. Bahia: Seabra, Serra da Água de Rega , 28 km N of Seabra , 23 February 1971, Irwin et al. 30757 (holotype K!, MO photo!; isotypes M!, NY!, UB!).

Marsypianthes Mart. ex Bentham (1833: 64) . Type — Marsypianthes hyptoides Mart. ex Bentham (1833: 64) , nom. illeg. [= Marsypianthes chamaedrys (Vahl) Kuntze ].

Subshrubs or perennial herbs, glandular-viscid, sometimes geoxylic with membranous leaves and axillary, few- to many-flowered cymes from the axils of foliar bracts, cymes either pedunculate with a lax sub-globose cluster of many flowers subtended by elliptic-lanceolate to linear bracteoles or sessile and 1- to few-flowered. Flowers pedicellate, calyx actinomorphic, broadly infundibuliform, with equal ±deltate to lanceolate lobes, connivent to erect at first, often becoming spreading to strongly reflexed in fruit; corolla bluish lilac or pale yellow, with tube cylindrical and with anterior lobe much shorter than the others; gynoecium with a persistent, thickened and angled stylopodium which is fused throughout its length to the inner face of the four ovary lobes; on ripening, the four mericarps (nutlets) break free and have a domed outer surface and a concave inner surface with a thin laciniate involute margin. The gynoecial structure is unique within the family, the shed nutlets strongly resembling minute tortoises!

Marsypianthes contains about five species, primarily in cerrado, in Central , West and Northeast Brazil, extending to Paraguay and Argentina. M. chamaedrys (Vahl) Kuntze is a very polymorphic herb or subshrub, mainly of disturbed ground extending from Mexico and the Caribbean , southwards to Peru, Bolivia and N Argentina. Species limits in the genus still remain to be clearly defined. See Fig. 3F .

Martianthus Harley & J.F.B.Pastore , nom.et stat. nov. ≡ Hyptis sect. Leucocephala Bentham (1848: 89) [non Leucocephala Roxburgh (1832: 612): Eriocaulaceae ]. Type:— Martianthus leucocephalus (Mart. ex Benth.) Harley & J.F.B.Pastore [= Hyptis leucocephala Mart. ex Benth. ].

The genus Martianthus bears flowers in compact, globose, pedunculate capitula with an involucre of narrowly linear, membranous bracteoles, obscured when the capitulum matures. The fruiting calyx tube is usually curved downward, from the middle, and the gynoecium has no stylopodium. These same morphological characters are shared with Cyanocephalus . Martianthus differs from the latter genus, in its non-clavate calyx lobes, the corolla lobes are usually very dark vinaceous or purple, sometimes dark pink, especially at the apex, with a paler tube and the stigma is usually distinctly lobed, rather than capitate. There are four species, three restricted to the caatingas or dry montane semi-arid areas of Northeast Brazil, with an outlying species in similar semi-arid conditions in Huarochi, coastal Peru. (On the other hand, the genus Cyanocephalus is a characteristic component of cerrado, a seasonal vegetation type centred in Central Brazil). See Fig. 4B .

Martianthus elongatus (Benth) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis elongata Bentham (1833: 88) ≡ Mesosphaerum elongatum (Benth.) Kuntze (1891: 526) . Type:— PERU. Province unknown: Tonga, Ruiz & Pavon s.n. (lectotype OXF!, designated by Epling 1936b; isolectotypes G!, MA!, P!).

Martianthus leucocephalus (Mart. ex Benth.) J.F.B.Pastore , comb. nov. ≡ Hyptis leucocephala Mart. ex Bentham (1833: 89) ≡ Mesosphaerum leucocephalum (Mart. ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Bahia: prope Joazeiro, fl. S. Francisco, Martius s.n. (holotype M!).

Martianthus sancti-gabrielii (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis sancti-gabrielii Harley (2001: 686) . Type:— BRAZIL. Bahia: São Gabriel, Alto da Lagoa Nova, 28 May 2000, Harley & Giulietti 53920 (holotype HUEFS!; isotype MO).

Martianthus stachydifolius (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis stachydifolia Epling (1936b: 264) . Type:— BRAZIL. Bahia: Serra de São Ignácio, February 1907, Ule 7550 (holotype K!; isotype G!, HBG!).

Medusantha Harley & J.F.B.Pastore , nom. et stat. nov. ≡ Hyptis sect. Trichosphaeria Bentham (1833: 95) [non Trichosphaeria Fuckel (1870) : Fungi]. Type:— Medusantha eriophylla (Pohl ex Benth.) Harley & J.F.B.Pastore [= Hyptis eriophylla Pohl ex Benth. , lectotype designated by Epling 1936b].

= Hyptis sect. Trichosphaeria subsect. Plumosae Epling (1936b: 280) , syn. nov. Type:— Hyptis plumosa Benth. [= Medusantha plumosa (Benth.) Harley & J.F.B.Pastore ].

= Hyptis sect. Trichosphaeria subsect. Crinitae Epling (1936b: 281) , syn. nov. Type:— Hyptis crinita Benth. [= Medusantha crinita (Benth.) Harley & J.F.B.Pastore ].

Medusantha is a small, rather homogeneous group, easily recognized by the flowers in globose capitula, with an involucre of soft, filiform bracteoles and the flowers with calyx tube straight, and long filiform calyx lobes (hence “medusoid”— Epling 1936b), a very slender corolla tube and a gynoecium without a stylopodium, style capitate with stigmatic branches reduced. Butterflies are strongly attracted to at least some of the species. There are eight species, mostly occurring in the cerrados of central Brazil. Medusantha martiusii is a characteristic shrub of the caatinga vegetation in semi-arid areas of the Brazilian North-east. See Fig. 4A .

Medusantha carvalhoi (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis carvalhoi Harley (1986a: 141 , 143). Type:— BRAZIL. Bahia: Serra do Sincorá, 4 km S of Ibicoara road, 25 March 1980, Harley et al. 20934 (holotype CEPEC!; isotype K!, NY!).

Medusantha crinita (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis crinita Bentham (1833: 95) ≡ Mesosphaerum crinitum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: Serra do Galheiro, Sellow s.n. (lectotype B†, designated by Epling 1936b, replacement lectotype K!, designated here; isolectotype LE!).

= Hyptis crinita Benth. var. polycephala Bentham (1833: 95) . Type :— BRAZIL. Without locality, Sellow s.n. (holotype K!).

= Hyptis spiraeifolia Mart. ex Bentham (1833: 95) ≡ Mesosphaerum spiraeifolium (Mart. ex Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Minas Gerais: in campis desertis Serro Frio ad flumen Jequitinhonha, Martius s.n. (holotype M!).

There are two Sellow specimens of Medusantha crinita in K. Epling erroneously annotated the wrong specimen as isotype of Hyptis crinita , as this sheet clearly says “ Hyptis crinita ss Benth.” in Bentham’s hand. This refers to Hyptis crinita var. ss polycephala Benth., foliis minoribus magis tomentosis, capitulis numerosis. The specimen clearly matches this description and is here taken as holotype of this variety. It is the other Sellow specimen in K, which is here designated as a replacement lectotype of the typical variety.

Medusantha eriophylla (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis eriophylla Pohl ex Bentham (1833: 96) ≡ Mesosphaerum eriophyllum (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: ad Megaponte [Meiaponte], Santa Lucia, Pohl 1072 (lectotype W!, designated by Epling 1936b; isolectotype K!).

var. eriophylla

var. coriifolia (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis eriophylla var. coriifolia Bentham (1833: 96) . Type:— BRAZIL. Minas Gerais: Corrego Boa Vista, Serra Bom Jardim, Pohl 2880 (lectotype W!, designated by Epling 1936b; isolectotype K!).

Medusantha martiusii (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis martiusii Bentham (1833: 95−96) ≡ Mesosphaerum martiusii (Benth.) Kuntze (1891: 526) (as “ Martinsii ”). Type:— BRAZIL. Bahia: inter Santa Anna et Santo Antonio das Queimadas, Martius s.n. (lectotype M!, designated by Epling 1936b).

= Hyptis brachyphylla Mart. ex Bentham (1833: 96) ≡ Mesosphaerum brachyphyllum (Mart. ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: in campis, Serra Frio, Martius s.n. (holotype M!).

Medusantha mollissima (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis mollissima Bentham (1833: 85) ≡ Mesosphaerum mollissimum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: Paranna, Sellow 1479 (lectotype B†, designated by Epling 1936b, replacement lectotype K!, designated here).

Figures

FIGURE 1. A. Oocephalus silvinae; B. Physominthe vitifolia; C. Hyptidendron caudatum; D. Cyanocephalus rugosus; E. Hyptidendron canum; F. Cyanocephalus lanatus; G. Rhaphiodon echinus (fr.); H. Hyptis radicans; J. Eriopidion strictum; K. Condea undulata; L. Leptohyptis macrostachys. Photos A, E, G, H, J−L by R. Harley; B by A. Paton; C, D, F by H. Moreira.

FIGURE 2. A. Gymneia malacophylla; B. Asterohyptis stellulata; C.. Hyptis imbricatiformis; D. Cantinoa carpinifolia; E. Eriope crassipes; F. Mesosphaerum suaveolens. Photos A−C, E, F by R. Harley; D by D. Cardoso.

FIGURE 3. A. Gymneia virgata; B. Hyptis recurvata; C. Eplingiella cuniloides; D. Hyptis ramosa; E. Physominthe vitifolia; F. Marsypianthes burchellii; G. Rhaphiodon echinus (fl.). Photos A, B, F by H. Moreira; C, G by R. Harley; D, E by A. Paton.

FIGURE 4. A. Medusantha eriophylla; B. Martianthus stachydifolius; C. Hypenia marifolia; D. Leptohyptis macrostachys; E. Hypenia sp.; F. Condea albida; G. Hyptis crenata; H. Oocephalus oppositiflorus. Photos A−C, F, H by R. Harley; D by J.F. Pastore; E by H. Moreira; G by D. Cardoso.