Medusantha multiflora (Pohl ex Benth.) Harley & J.F.B.Pastore, 2012

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 : 29-36

Medusantha multiflora (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov.

≡ Hyptis multiflora Pohl ex Bentham (1833: 96) ≡ Mesosphaerum multiflorum (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: inter Rio Jequitinhonha et Columbis, Pohl 3163 (lectotype W!, designated by Epling 1936b; isolectotype K!).

Medusantha plumosa (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis plumosa Bentham (1848: 94) ≡ Mesosphaerum plumosum (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: Formigas, July 1840, Gardner 5086 (holotype K!; isotype OXF!).

Medusantha simulans (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis simulans Epling (1936b: 282) . Type:— BRAZIL. Ceará: Serra do Araripe in “caatinga”, 17 April 1910, Löfgren 566 (holotype S!; isotype UC!, fragment only).

Mesosphaerum Browne (1756: 257) . Type:— Mesosphaerum suaveolens (L.) Kuntze [= Ballota suaveolens L., lectotype designated by Kuntze 1891].

= Hyptis sect. Mesosphaeria Benth. (1833: 122) , pro parte. Type:— Hyptis suaveolens (L.) Poit., lectotype designated by Epling 1936b [= Mesosphaerum suaveolens (L.) Kuntze].

= Brotera Sprengel (1802: 151 , t. 12), nom. illeg. [non Cavanilles 1799]. Type:— Brotera persica Spreng. [= Mesosphaerum pectinatum (L.) Kuntze].

= Schaueria Hasskarl (1842: 25) , nom. illeg. [non Nees 1838]. Type:— Schaueria graveolens (Blume) Hassk. [= Mesosphaerum suaveolens (L.) Kuntze].

= Gnoteris Rafinesque (1838: 76) . Type:— Gnoteris cordata Raf. , lectotype designated here [= Mesosphaerum suaveolens (L.) Kuntze].

= Hyptis sect. Subumbellaria Epling (1933: 79) , syn. nov. Type:— Hyptis asperifolia Standley [= Mesosphaerum asperifolium (Standl.) Harley & J.F.B.Pastore ].

= Hyptis sect. Mesosphaeria subsect. Fruticosae Epling (1936b: 246) , syn. nov. Type:— Hyptis melissoides Kunth [= Mesosphaerum melissoides (Kunth) Kuntze ].

= Hyptis sect. Mesosphaeria subsect. Pectinaria (Benth.) Epling (1936b: 241) ≡ Hyptis sect. Pectinaria Bentham (1833: 127) . Type:— Hyptis pectinata (L.) Poit. [= Mesosphaerum pectinatum (L.) Kuntze].

Bentham (1833) included only one species, H. pectinata , in his Hyptis section Pectinaria , which was reduced to a subsection of sect. Mesosphaeria by Epling (1936b), with seven other species and later expanded to 14 ( Epling 1949).

= Hyptis sect. Mesosphaeria subsect. Eriocephalae Epling (1936b: 246) . Type:— Hyptis eriocephala Benth. [= Mesosphaerum eriocephalum (Benth.) Harley & J.F.B.Pastore ].

Mesosphaerum , as delimited in this paper, is characterized by an inflorescence with several to many flowers usually in lax or more congested cincinnate, pedunculate, sometimes weakly capitate cymes, flowers shortly pedicellate, subtended by small bracteoles not forming an involucre, and with a subactinomorphic calyx, often with white hairs in the throat, and the gynoecium without a stylopodium. There are about 25 species, with a primarily Andean distribution, extending also into the mountains of Central America and Mexico. Mesosphaerum sidifolium (including Hyptis umbrosa Salzm. ex Benth. ) has a wider distribution, extending also into eastern Brazil, while M. irwinii is endemic to the mountains of Bahia, Northeast Brazil. Two species, M. pectinatum and M. suaveolens , have become common weeds, widespread in the tropics and extending into the Old World. See Fig. 2F .

Epling (1949) originally recognized 32 species in Hyptis sect. Mesosphaeria , divided into five subsections: Eriocephalae Epling (14 species) largely Andean, Spicaria (Benth.) Epling (two species), Pectinaria (Benth.) Epling (c. 14 species, including two pantropical weeds. H. pectinata (L.) Poit. and H. suaveolens (L.) Poit.), and two monotypic subsections Ocimoideae Epling and Plectranthodon Epling. The molecular evidence ( Pastore et al. 2011) demonstrates that sect. Mesosphaeria is polyphyletic, comprising at least two different lineages. One of these, composed of species of Hyptis sect. Mesosphaeria subsect. Pectinaria pro parte, subsect. Spicaria and subsect. Plectranthodon has now been included here in the newly recognized genus Cantinoa , along with former members of Hyptis sect. Polydesmia . Cantinoa is shown from molecular studies to be sister to Hyptis sensu stricto. The other lineage, which comprises Hyptis sect. Mesosphaeria subsect. Pectinaria pro parte , including H. pectinata and H. suaveolens , and Hyptis sect Mesosphaeria subsect. Eriocephalae , is here recognized as the genus: Mesosphaerum P. Browne. No attempt at this stage, has been made to recognize sections within Mesosphaerum , especially as there is still doubt how these will eventually be disposed. Indeed the monotypic Hyptis subsect. Ocimoideae remains to be analysed. Wider sampling of the constituent species is needed.

In the list below, we have included all species considered definitely to belong to the genus Mesosphaerum . The necessary combination for twelve of the species had already been made by Kuntze (1891).

Mesosphaerum argutifolium (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis argutifolia Epling (1936b: 245) . Type:— ECUADOR. Loja: Yangana, 18 December 1876, André 4592 (holotype K!).

Mesosphaerum asperifolium (Standley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis asperifolia Standley (1930: 40) . Type:— HONDURAS. Dept. Comayagua: near Siguatepeque, 14−27 February 1928, Standley 56231 (holotype US!; isotypes F!, G!, K!).

Mesosphaerum chacapoyense (Briq.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis chacapoyensis Briquet (1898: 204) . Type :— PERU. Amazonas : Chachapoyas, Mathews 3151 (holotype G!; isotypes K!, OXF!).

= Hyptis polyantha var. longiflora Bentham (1848: 124) . Type :— PERU. Amazonas : Chachapoyas, Mathews 3151 (holotype K!; isotypes G!, OXF!).

The variety described by Bentham is based on the same type collection as Briquet’s species, although the former was based on a specimen at Kew and the latter based on a specimen in Geneva.

Mesosphaerum collinum (Brandegee) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis collina Brandegee (1893: 164) . Type:— MEXICO. Baja California: San José del Cabo, 16 September 1891, Brandegee 468 (holotype UC!; isotypes G!, NY!).

Mesosphaerum diffusum (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis diffusa Epling (1936b: 242) . Type:— COLOMBIA. Magdalena: Santa Marta. Near Onaea, 4 December 1898, Smith 1373 (holotype US!; isotypes BR!, COL!, G!, K!, NY!, P!, S!).

Mesosphaerum diversifolium (Benth.) Kuntze (1891: 526) ≡ Hyptis diversifolia Bentham (1844b: 144) . Type:— PERU. Loja: “Catamayo, prope Loxa”, Hartweg 803 (holotype K!; isotypes G!, NY!, OXF!, P!, W!).

Mesosphaerum eriocephalum (Benth.) Kuntze (1891: 526) ≡ Hyptis eriocephala Bentham (1848: 124) . Type:— PERU. Amazonas: Chachapoyas, 1835, Mathews 1530 (holotype K!; isotype OXF!).

= Hyptis kuntzeanum Briq. ( Mesosphaerum kuntzeanum ) (1896: 787), syn. nov. Type:— BOLIVIA. Cochabamba: Cochabamba, April 1892, Kuntze s.n. (holotype G!; isotype NY!).

For the above, and a number of other taxa, Briquet published the name of the new species under Hyptis , followed by Mesosphaerum in parenthesis. This latter name is here treated as invalid.

Mesosphaerum gymnocaulon (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis gymnocaulos Epling (1936b: 246) . Type:— ECUADOR. Galapagos Islands: Albemarle, Cowley Bay , 10 August 1905, Stewart 3326 (holotype GH; isotypes K!, NY!, US!) .

Mesosphaerum irwinii (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis irwinii Harley (1974: 130) . Type:— BRAZIL. Bahia: Valley of the Rio das Ondas, ca. 24 km N of Seabra , 28 February 1971, Irwin et al. 31277 (holotype K!; isotype NY!, UB!).

Mesosphaerum lachnosphaerium (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis lachnosphaeria Epling (1936b: 247) . Type:— ECUADOR. Chimborazo, Huigra, Hitchcock 20365 (holotype US!; isotypes K!, NY!).

Mesosphaerum marrubiifolium (Epling & Mathias) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis marrubiifolia Epling & Mathias (1957: 229) . Type:— PERU. Amazonas : 10 km SE of Balzas, Evinger 508 (holotype US!) .

Mesosphaerum melissoides (Kunth) Kuntze (1891: 526) ≡ Hyptis melissoides Kunth in Humboldt, Bonpland & Kunth (1818: 320). Type:— COLOMBIA. “Prope pagum El Tablón et ripam fluminis Juanambu, Regno Novo Granatensi”, Bonpland 2129 (holotype: P!).

Epling (1949) erroneously placed the type collection in Ecuador. This and all subsequent collections are from Colombia ( Fernández-Alonso 1995).

Mesosphaerum oblongifolium (Benth.) Kuntze (1891: 526) ≡ Hyptis oblongifolia Bentham (1848: 125) . Type:— MEXICO. Oaxaca: Juquila, Jurgensen 71 (holotype K!; isotype G!, OXF!).

= Hyptis viejensis Oersted in Bentham & Oersted (1854: 34) ≡ Mesosphaerum viejensis (Oerst.) Kuntze (1891: 527) . Type:— NICARAGUA. Nueva Segovia: Volcan Viejo, Oersted s.n. (holotype C!; isotype K!).

The specimen at K is numbered 44, but appears to be the same collection. Epling (1933, 1949) gives Volcan Viejo as from Costa Rica, but this is erroneous, as Oersted refers to Nicaragua in the protologue. There are apparently no other records of Hyptis oblongifolia from Costa Rica.

= Hyptis nicaraguensis Oersted in Bentham & Oersted (1854: 34), syn. nov. Type:— NICARAGUA. Granada: prope Granada, Oersted 15761 (holotype C!).

= Hyptis vulcanica Seemann (1854: 188) ≡ Mesosphaerum vulcanicum (Seem.) Kuntze (1891: 526) . Type:— PANAMA. Chiriqui: Mt. Chiriqui, Seemann 1603 (holotype BM!; isotype K!).

= Hyptis chapalensis Briquet ( Mesosphaerum chapalense Briq. ) (1898: 203). Type:— MEXICO. Jalisco: rich mountains near Lake Chapala, 9 December 1889, Pringle 2440 (holotype G!; isotypes BM!, F!, K!, LE!, M!, MEL!, MO photo!, P!).

= Hyptis arborescens Epling (1933: 100) , syn. nov. Type:— NICARAGUA. Nueva Segovia: Volcán El Viejo, 18–22 January 1903, Baker 18 (holotype MO!; isotype NY!).

Mesosphaerum obtusatum (Benth.) Kuntze (1891: 526) ≡ Hyptis obtusata Bentham (1846: 241) . Type:— ECUADOR. Pichincha: “Ad pontem Guapulo prope Quito”, Hartweg 1322 (holotype K!; isotype OXF!).

The specimen at OXF erroneously gives the collection locality as Colombia.

Mesosphaerum pectinatum (L.) Kuntze (1891: 525) ≡ Nepeta pectinata Linnaeus (1759: 1097) ≡ Bystropogon pectinatus (L.) L’Héritier (1789: 19) ≡ Hyptis pectinata (L.) Poiteau (1806: 474). Type:— JAMAICA. Without locality, Browne s.n., Herb. Linneanum 726.31 (LINN!, lectotype designated by Howard 1989).

= Brotera persica Sprengel (1802: 151 , t. 12) ≡ Hyptis persica (Spreng.) Poiteau (1806: 471) . Type:—Cultivated material from Halle Botanic Garden, from seed collected in Iran (Persia) by Oliver & Bruguière, not found, although two specimens in the Willdenow herbarium (B-W 10839-010! [Image ID 361203], B-W 10839-020! [Image ID 361211) might be relevant, but lack collection data .

= Hyptis nepetoides Fisch. ex Schrank (1822: 52) . Type:—A cultivated plant, probably from Brazil, not located. Neotype selected here: BRAZIL. Bahia: Feira de Santana, 16 September 2003, Ribeiro 68 (HUEFS). There is a specimen of Mesosphaerum pectinatum at M, possibly collected by Martius, which might represent the living plant from which H. nepetoides was described, which was said to have come originally from Brazil. There is, however, insufficient information available on the sheet. Schrank often did not preserve dried material of the cultivated plants to which he gave names, and therefore many of these have not been unequivocally identified. It is therefore necessary to choose a neotype.

Mesosphaerum perbullatum (Fern.Alonso) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis perbullata Fernández-Alonso (1995: 475−476) . Type:— COLOMBIA. Boyaca: W of Tibarrosa , between Duitarama and Sogamoza, Wood 3762 (holotype COL!; isotype K!).

Mesosphaerum pilosum (Benth.) Kuntze (1891: 526) ≡ Hyptis pilosa Bentham (1833: 124) . Type:— PERU. Without locality, 1827, Pavon s.n. (holotype OXF!; isotypes G!, MA!, P!).

Mesosphaerum pseudoglaucum (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis pseudoglauca Epling (1936b: 247) . Type:— ECUADOR. Province unknown: San Pedro, Townsend A-107 (holotype US!) .

Mesosphaerum purdiaei (Benth.) Kuntze (1891: 527) ≡ Hyptis purdiaei Bentham (1848: 125) . Type:— COLOMBIA. Magdalena: Santa Marta, July 1844, Purdie s.n. (holotype K!; isotype UC!).

Mesosphaerum septentrionale (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis septentrionalis Epling (1833: 98) . Type:— MEXICO. Durango: San Ramón, 21 April–18 May 1906, Palmer 111 (holotype US!; isotypes K!, G!, NY!, MO!).

Mesosphaerum sidifolium (L'Hérit.) Harley & J.F.B.Pastore , comb. nov. ≡ Bystropogon sidifolius L'Héritier (1787: 19) ≡ Hyptis sidifolia (L'Hérit.) Briquet (1898: 204) . Type:— PERU. Without locality, Dombey s.n. (holotype P!).

= Hyptis polyantha Poiteau (1806: 470) ≡ Mesosphaerum polyanthum (Poit.) Kuntze (1891: 526) . Type:— ECUADOR. Loja: Gonzanama, Bonpland s.n. (holotype P!; B photo!).

= Hyptis silvestris Epling (1936b: 249) . Type:— COLOMBIA. Tolima: La Virginia, Libano, 22 December 1917, Pennell 3274 (holotype NY!; isotype US!) .

= Hyptis umbrosa Salzm. ex Bentham (1833: 125) . Type:— BRAZIL. Bahia: “in umbrosis”, Salzmann s.n. (lectotype K!, designated by Epling 1936b; isolectotypes E!, G-DC!, P!).

= Hyptis graveolens Salzm. ex Bentham (1833: 125) , nom. illeg. (non Hyptis graveolens Schrank 1822: 52 ). Type:— BRAZIL. Bahia: “in maritimis“, Salzmann s.n. (holotype K!; isotypes E!, G-DC!).

This species is extremely variable especially in leaf morphology and indumentum. Fernández-Alonso (1995) has placed Hyptis silvestris in synonymy of “ Hyptis sidifolia ”, as it differs primarily in leaf indumentum and peduncle length, and after careful study we have accepted this, as these characters occur individually in material from other parts of the range of H. sidifolia . For fuller details on synonymy of this species see Harley (2006) .

Mesosphaerum suaveolens (L.) Kuntze (1891: 525) ≡ Ballota suaveolens Linnaeus (1759: 1100) ≡ Hyptis suaveolens (L.) Poiteau (1806: 474) Type:— JAMAICA. Browne s.n. (LINN 737.6!, lectotype designated by Epling 1949).

= Hyptis ebracteata R.Brown in Aiton (1811: 391), nom. superfl.

= Hyptis plumieri Poiteau (1806: 473) . Type:— VENEZUELA. Distrito Federal: Caracas, Humboldt & Bonpland s.n. (holotype?P).

= Hyptis graveolens Schrank (1828: 56) . Type:— A cultivated specimen, not traced with certainty, from seed collected by Martius in Brazil .

The type specimen of H. graveolens has not been definitely located, although there is a specimen of Mesosphaerum suaveolens , annotated by Martius as Hyptis graveolens at M! (see Harley 1985a ).

= Hyptis congesta Leonard (1927: 70) . Type:— HAITI. S. Michel de l’Attalaye , Leonard 7594 (holotype US!) .

Mesosphaerum urticoides (Kunth) Kuntze (1891: 527) [as “ urticodes ”] ≡ Hyptis urticoides Kunth in Humboldt, Bonpland & Kunth (1818: 320). Type:— MEXICO. Tabasco: prope Jalapa, Bonpland s.n. (holotype P!).

= Hyptis lilacina Chamisso & Schlechtendal (1830: 101) . Type:— MEXICO. Tabasco: Jalapa, Schiede & Deppe 152 (lectotype B†, designated by Epling 1936b, replacement lectotype HAL!, designated here; isolectotype BM?).

The material at BM lacks a collection number, so must remain doubtful.

Oocephalus (Benth.) Harley & J.F.B.Pastore , comb. et stat. nov. ≡ Hyptis sect. Oocephalus Bentham (1833: 84) ≡ Hyptis sect. Polydesmia subsect. Oocephalus (Benth.) Epling (1936b: 250) . Type:— Oocephalus lacunosus (Pohl. ex Benth.) Harley & J.F.B.Pastore [= Hyptis lacunosa Pohl ex Benth. , lectotype designated by Epling 1936b].

= Hyptis sect. Polydesmia subsect. Glomeratae Bentham (1848: 119) , pro parte, syn. nov. Type:— Oocephalus oppositiflorus (Schrank) Harley & J.F.B.Pastore [= Hyptis glomerata Mart. ex Schrank , lectotype designated by Epling 1936b].

Shrubs or subshrubs, rarely herbs with usually small or medium-sized leaves, sessile or shortly petiolate, distinguished by the possession of an inflorescence of congested, pedunculate or sessile cymes, few- to manyflowered, not forming a globose or semi-globose capitulum, but ± ovoid in form, enveloped by an involucre of usually broad, ovate or lanceolate bracteoles, especially in bud, rarely narrower and not enveloping the cymes. Flowers sessile or subsessile, corollas with an elongate tube, lobes usually short. Gynoecium without stylopodium. 14 species are recognized here, typically in campo rupestre of the Serra do Espinhaço of Minas Gerais and Bahia and in similar habitats in Goiás. One species, O. oppositiflorus , is spreading through Brazil into disturbed habitats and has a wider distribution, extending into eastern Bolivia. Further research is needed to define more clearly the characters of the genus. See Figs. 1A , 4H .

Oocephalus argyrophyllus (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis argyrophylla Harley (1985b: 614) . Type:— BRAZIL. Bahia: Barra da Estiva on Ibicoara Road by the Rio Preto , 27 January 1974, Harley et al. 15515 (holotype CEPEC!; isotypes AAU!, K!, NY!, SPF!, U!, UEC!, US!) .

var. argyrophyllus

var. pedunculatus (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis argyrophylla var. pedunculata Harley (1985b: 615) . Type:— BRAZIL. Bahia: Serra do Sincorá, 8 km SW of Mucugê, on road from Cascavel near Fazenda Paraguaçu, 6 February 1974, Harley et al. 16076 (holotype CEPEC!; isotype K!).

Oocephalus crassifolius (Mart. ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis crassifolia Mart. ex Bentham (1833: 94) ≡ Mesosphaerum crassifolium (Mart. ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Bahia: “in petrosis ad Vila do Rio de Contas”, Martius Obs. 1976 (holotype M!).

Oocephalus foliosus (A.St.-Hil. ex Benth.) Harley & J.F.B.Pastore, comb. nov. ≡ Hyptis foliosa A.St. -Hil. ex Bentham (1833: 117) ≡ Mesosphaerum foliosum (A.St.-Hil. ex Benth.) Kuntze (1891: 526). Type:— BRAZIL. Goiás: Saint-Hilaire C3 709 (holotype P!; isotype UC!).

Oocephalus hagei (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis hagei Harley (1986a: 145) . Type:— BRAZIL. Bahia: Lençóis, Serra da Larguinha, ca. 2 km NE of Caeté Açu (Capão Grande), 25 May 1980, Harley et al. 22533 (holotype CEPEC!; isotype K!).

The bracteoles of this species are atypical for the genus, in being very slender and not enveloping the flowers. However Rudall (pers. com. in Harley 1986a ) has demonstrated the very characteristic leaf anatomy which places this species unequivocally with the genus Oocephalus , confirmed by the molecular results.

Oocephalus halimifolius (Mart. ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis halimifolia Mart. ex Bentham (1833: 94) ≡ Mesosphaerum halimifolium (Mart. ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Bahia: “campis altis petrosis, Vila do Rio de Contas”, October 1818, Martius s.n. (holotype M!; isotype M!).

Oocephalus lacunosus (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis lacunosa Pohl ex Bentham (1833: 94) ≡ Mesosphaerum lacunosum (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: Santa Ingrazia, Pohl 3194 (lectotype W!, designated by Epling 1936b; isolectotype K!).

= Hyptis cordifolia Glaziou (1911: 551) , nom. nud. Reference specimen: BRAZIL. Minas Gerais: “Biribiry a Mocotó”, 28 March 1892, Glaziou 19703 (K!, P!).

Oocephalus lythroides (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis lythroides Pohl ex Bentham (1833: 118) ≡ Mesosphaerum lythroides (Pohl ex Benth.) Kuntze (1891: 526) [as “ lythrodes ”]. Type:— BRAZIL. Goiás: Fazenda Água Fria, Pohl 1483 (holotype W!; isotype K!).

Oocephalus niveus (Epling) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis nivea Epling (1936b: 252) . Type:— BRAZIL. Goiás (?): entre Rio Tocantins et Os Porcos , 3 January 1895, Glaziou 21934 (holotype K!; isotype P!).

The type locality has not been traced. The greater part of the the Rio Tocantins is in Tocantins State (formerly a part of Goiás) though a part of the upper reaches of the river extend into Goiás State. The only extant population known is in the Chapada dos Veadeiros, Goiás .

Oocephalus nubicola (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis nubicola Harley (1992: 578) . Type:— BRAZIL. Bahia: Rio de Contas, Pico das Almas, 18 March 1977, Harley et al. 19614 (holotype CEPEC!; isotype K!).

Oocephalus oppositiflorus ( Schrank) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis oppositiflora Schrank (1822: 52) . Type :— Probably a cultivated plant, not localized. Neotype designated here: BRAZIL. Distrito Federal : Samambaia, 4 May 1996, Rezende 459 (CEN).

= Hyptis glomerata Mart. ex Schrank (1828: 55) . Type:— BRAZIL. Minas Gerais: “in humidis campis ad Burityzais deserti”, Martius s.n. (holotype M!).

= Hyptis pauciflora Pohl ex Bentham (1833: 117) ≡ Mesosphaerum pauciflorum (Pohl) Kuntze (1891: 526) . Type:— BRAZIL. Goiás: Megaponte [Meiaponte, now Pirenópolis], Pohl 2795 (holotype W!; isotype K!).

= Hyptis glomerata var. villosa Bentham (1848: 120) . Type:— BRAZIL. Minas Gerais: without locality, Claussen s.n. (holotype K!; isotype K!).

Unfortunately, the earliest name for this species is that of Schrank, which supersedes the well-known epithet of “ glomerata ”. The cultivated plant on which the current name is based was grown from seed collected in Brazil by Martius, but no specimen appears to exist.

Oocephalus pauciflorus (Harley) Harley & J.F.B.Pastore , comb. & stat. nov. ≡ Hyptis halimifolia Mart. ex Benth. var. pauciflora Harley (1985b: 612) . Type:— BRAZIL. Bahia: Serra das Almas, ca. 25 km WNW of town of Rio de Contas , 19 March 1976, Harley et al. 19676 (holotype CEPEC!; isotypes AAU!, IPA!, K!, NY!, SPF!, U!, UEC!, US!) .

Field observations and study of the substantial number of collections of both this taxon and of Oocephalus halimifolius indicate that they are morphologically quite distinct and have differing altitudinal ranges and ecological requirements and are therefore best treated as distinct species.

Oocephalus petraeus (A.St.-Hil. ex Benth.) Harley & J.F.B.Pastore, comb. nov. ≡ Hyptis petraea A.St. -Hil. ex Bentham (1833: 117) ≡ Mesosphaerum petraeum (A.St.-Hil. ex Benth.) Kuntze (1891: 526). Type:— BRAZIL. Minas Gerais: "un endroit pierreux près Taioba", Saint-Hilaire B1 1740 (holotype P!; isotype UC!).

Oocephalus piranii (Harley) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis piranii Harley (1992: 572) . Type:— BRAZIL. Minas Gerais: Grão Mogol, descida para Jambeiro , 15 July 1990, Pirani et al. CFCR 13069 (holotype SPF!; isotype K!).

Oocephalus silvinae (Harley) Harley & J.F.B.Pastore comb. nov. ≡ Hyptis silvinae Harley (1985b: 624) . Type:— BRAZIL. Bahia: Pico das Almas, 19 March 1977, Harley et al. 19701 (holotype CEPEC!; isotypes AAU!, E!, IPA!, K!, NY!, U!, UEC!, US!, SPF!, K!, NY!).

Physominthe Harley & J.F.B.Pastore , gen. nov. Hypeniae habitu virgato cum internodiis superis inflatis, glabris et pruinosis conveniens sed inflorescentia cymis pedunculatis paucifloris usque 3−6-floribus instructis et basi calycis bracteolis binatis destituto differt. Type:— Physominthe vitifolia (Pohl ex Benth.) Harley & J.F.B.Pastore [= Hyptis vitifolia Pohl ex Benth ].

Shrubs or subshrubs, often aromatic; stems virgate, erect, upper internodes glabrous, pruinose, usually strongly fistulose and inflated, nodes and lower stems often with long, rigid setose hairs; leaves lobed, sometimes weakly so; inflorescence thyrsoid, lax, formed of pedunculate, subumbellate, few-flowered cymes (with intercalary axes reduced), 3- to 6-flowered, flowers shortly pedicellate from the axils of minute, subulate bracts; flowers small with calyx ± actinomorphic, turbinate at anthesis, becoming ± campanulate in fruit, bracteoles reduced, when present borne near base of cyme, never paired at base of calyx, style deciduous from below nutlets, stylopodium absent.

Similar, in its virgate habit with upper internodes inflated, glabrous and pruinose, to Hypenia , in which it was formerly included, and some species of Eriope , and differing from all species of both these genera in lacking a pair of bracteoles at base of calyx. Chromosome number 2 n = 28, differing from numbers found in Eriope (2 n = 20) and Hypenia (2 n = 20, 12). See Harley & Heywood (1992). The genus is restricted to upland cerrado areas of Bahia, Minas Gerais, Goiás and São Paulo. One species so far described with another awaiting publication. See Figs. 1B , 3E .

Physominthe vitifolia (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis vitifolia Pohl ex Bentham (1833: 138) ≡ Mesosphaerum vitifolium (Pohl ex Benth.) Kuntze (1891: 527) ≡ Hypenia vitifolia (Pohl ex

Benth.) Harley (1988: 91). Type:— BRAZIL. Goiás: “ad Santa Cruz”, Pohl 6059 (holotype W!) .

= Hyptis calophylla A.St. -Hil. ex Bentham (1833: 138) ≡ Mesosphaerum calophyllum (A.St.-Hil. ex Benth.) Kuntze (1891: 526). Type:— BRAZIL. Minas Gerais: Morro do Indaia, Minas Novas, St.-Hilaire B1 1155 (holotype P!; isotype P!).

= Hyptis glaziovii Briquet (1894: 716) . Type:— BRAZIL. São Paulo: Campos do Bocaina , prés S. José dos Barreiros, 6 April 1892, Glaziou 13047 (holotype G!; isotype P!) .

Rhaphiodon Schauer (1844: 345) . Type:— Rhaphiodon echinus (Nees & Mart.) Schauer [= Zappania echinus Nees & Mart. ].

= Hyptis sect. Xanthiophaea Bentham (1833: 70 . Type:— Hyptis sideritis Mart. ex Benth. , lectotype designated by Epling (1936a: 186).

Epling excluded two of the three species which Bentham recognized in Hyptis Sect. Xanthiophaea. However he misspelt the genus as Raphiodon , a name frequently encountered in herbaria.

Rhaphiodon is a monotypic genus, with R. echinus the only species. It is a prostrate, aromatic herb, with flowers in long-pedunculate spherical heads, with spinose involucral bracteoles and the calyx with lobes composed of up to ca 11 spines, corolla long-tubular, bright purple, the spinose fruiting head, with only one nutlet per flower, falling as a unit. It occurs in seasonally damp, disturbed areas, especially along roadsides in the caatingas of Northeast Brazil extending southwards to N Minas Gerais State. See Figs. 1G , 3G .

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.