Hyptidinae

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 : 8-13

Conspectus of Hyptidinae with new classification including new combinations, synonyms and typifications

Asterohyptis Epling (1932: 17) . Type:— Asterohyptis stellulata (Benth.) Epling (1932: 17) Ξ Hyptis stellulata Bentham (1833: 129) .

This genus, which is the only one of the tribe which does not occur in South America, has the flowers arranged in elongate, spike-like inflorescences, composed of few-flowered verticillasters, in axils of reduced bracts. The calyx lobes are subulate or filamentous, often rigid, spreading, corollas small, white, weakly 2- lipped with 5 subequal lobes and lacking the explosive pollination mechanism, with the thickened hinge at base of anterior corolla lip. The three or possibly four species ( Turner 2011) extend from Mexico to Central America, in dry, open habitats. See Fig. 2B .

Cantinoa Harley & J.F.B.Pastore , nom. et stat. nov. Ξ Hyptis sect. Polydesmia Bentham (1833: 114) [non Polydesmia Boudier (1885) : Fungi]. Type:— Cantinoa carpinifolia (Pohl ex Benth.) Harley & J.F.B.Pastore [= Hyptis carpinifolia Pohl ex Benth. , lectotype designated by Epling 1936b].

= Hyptis section Spicaria Bentham (1833: 78) Ξ Hyptis sect. Mesosphaeria subsect. Spicaria (Bentham) Epling (1933: 96) , syn. nov. Type:— Hyptis spicigera Benth. , lectotype designated by Epling 1936b [= Cantinoa americana (Aubl.) Harley & J.F.B.Pastore ].

= Hyptis sect. Polydesmia subsect. Rigidae Bentham (1848: 116) , syn. nov. Type:— Hyptis carpinifolia Benth. , lectotype designated by Epling 1936b [= Cantinoa carpinifolia (Benth.) Harley & J.F.B.Pastore ].

= Hyptis sect. Polydesmia subsect. Vulgares Bentham (1848: 120) , syn. nov. Type:— Hyptis mutabilis (Rich.) Briq. , lectotype designated by Epling 1936b [= Cantinoa mutabilis (Rich.) Harley & J.F.B.Pastore ].

= Hyptis sect. Polydesmia subsect. Mutabiles Epling (1933: 102) , syn. nov. Type:— Hyptis mutabilis (Rich.) Briq. [= Cantinoa mutabilis (Rich.) Harley & J.F.B.Pastore ].

This subsectional name is illegitimate, being a superfluous name for Hyptis subsection Vulgares . Epling (1933) mentions it only in his account of N American Hyptis , subsequently ignoring the name.

= Hyptis sect. Mesosphaeria subsect. Plectranthodon Epling (1936b: 237) , syn. nov. Type:— Hyptis plectranthoides Benth. [= Cantinoa plectranthoides (Benth.) Harley & J.F.B.Pastore ].

= Hyptis sect. Polydesmia subsect. Malvastra Epling (1936b: 254) , syn. nov. Type:— Hyptis vestita Pohl ex Benth. , lectotype designated by Epling 1936b [= Cantinoa althaeifolia (Pohl ex Benth.) Harley & J.F.B.Pastore ].

Cantinoa is a genus composed of herbs, subshrubs or shrubs with often aromatic, mesomorphic leaves, sessile to long-petiolate. The flowers are borne in congested 12−26-flowered, ± ovoid cymes (cymules), not forming hemispherical or spherical capitula, the flowers subsessile and surrounded by rather slender to ovate, sometimes scarious bracteoles apparently equal in number to the flowers and sometimes investing them to form an involucre. The calyx has a straight tube with five usually subequal ± subulate lobes, these rarely absent or reduced, the corolla with well-developed, spreading limb. Gynoecium without stylopodium. The cymules often form branched synflorescences or may be congested to form ± oblong, terminal spikes. There are 23 species currently recognized, extending from SE United States to the Caribbean and S America as far as Argentina, and occurring in a variety of mostly mesophytic habitats. Two species are widespread and have been introduced into the Old World: Cantinoa americana (Aubl.) Harley & J.F.B.Pastore is frequent in the tropics as a weed of cultivation, and C. mutabilis (Rich.) Harley & J.F.B.Pastore has been recorded from S Africa. See Fig. 2D .

Epling (1949) recognized five subsections under Hyptis sect. Polydesmia . Of these, Hyptis subsects. Oocephalus and Glomeratae have here been placed in another genus, Oocephalus (Benth.) Harley & J.F.B.Pastore. H y ptis subsect. Malvastra, was erected by Epling (1936b) solely on the basis of the presence of dendroid hairs, but it can no longer be upheld, as these have now been found to occur in other species, e.g. C. carpinifolia (Benth.) Harley & J.F.B.Pastore (of subsect. Rigidae Benth.). Hybrids have now been recognized involving C. mutabilis (H. subsect. Vulgares ) with both C. carpinifolia (H. subsect. Rigidae) and C. althaeifolia (Pohl ex Benth.) Harley & J.F.B.Pastore (H. subsect. Malvastra). As a result there now seems little justification in maintaining these subsections at any rank.

The genus is named in honour of Dr Philip Cantino, Ohio, whose researches on Lamiaceae , especially the delimitation of its suprageneric taxa and his studies on the distinguishing characters between this family and the Verbenaceae , have greatly advanced our understanding of the group.

Cantinoa americana (Aubl.) Harley & J.F.B.Pastore , comb. nov. ≡ Nepeta americana Aublet (1775: 623) ≡ Hyptis americana (Aubl.) Urban (1918: 322) , nom. illeg. [non Hyptis americana (Poiret in Lamarck 1805: 571) Briquet 1897b: 338 = Condea americana (Poir.) Harley & J.F.B.Pastore ] ≡ “ Nepeta foliis serratis , ovatoacutis, spicis imbricatis, acuminatis ” Burmann in Plumier (1758: 155). Type:—ANTILLES: without locality, Plumier (1758: t. 162, f. 2), lectotype designated here.

The original Aublet specimen, presumably from French Guiana , has not been found. However he cites the above polynomial and plate from the earlier Plumier volume, which enables the choice of Plumier’s plate as lectotype, although this was an illustration of a specimen from the Antilles , without further indication of provenance. To assist identification of the name we also propose an epitype, designated here: MEXICO. Morelos: Cuernavaca , 23 September 1896, Pringle 6559 ( K!) .

= Hyptis spicigera Lamarck (1789: 185) ≡ Mesosphaerum spicigerum (Lam.) Kuntze (1891: 527) . Type:— SIERRA LEONE (?). Smeathman s.n. (holotype P!, a specimen given to Lamarck by de Beauvois).

= Hyptis lophantha Martius ex Bentham (1833: 78) ≡ Mesosphaerum lophanthum (Mart. ex Benth.) Kuntze (1891: 526) Type:— BRAZIL. Minas Gerais: “in herbidis pascuis Serro Frio”, Martius s.n. (holotype M!).

= Hyptis madagascariensis Bojer (1837: 251) , nom. nud.

= Hyptis subverticillata Andersson (1855: 197) ≡ Mesosphaerum subverticillatum (Andersson) Kuntze (1891: 527) . Type:— ECUADOR. Galapagos Islands: Albemarle, Andersson 207 (holotype S!; isotype P!).

= Hyptis gonocephala Wright ex Grisebach (1866: 212) ≡ Mesosphaerum gonocephalum (Wright ex Griseb.) Kuntze (1891: 526) . Type:— CUBA. La Punta de La Junta, Wright 3154 (holotype GOET; isotypes BM!, K!, NY!, S!, US!).

This species occurs as a weed in both the New and Old World tropics. Its native distribution is uncertain, though clearly in the Americas.

Cantinoa althaeifolia (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis althaeifolia (as althaeaefolia) Pohl ex Bentham (1833: 115) Ξ Mesosphaerum althaeefolium (sic!) (Pohl ex Benth.) Kuntze (1891: 526). Type:— BRAZIL. Minas Gerais: Fazenda Vieira, Pohl 2540 (lectotype W!, designated by Epling 1936b; isolectotype F!, K!).

= Hyptis vestita Bentham (1833: 114) Ξ Mesosphaerum vestitum (Benth.) Kuntze (1891: 527) . Type:— BRAZIL. “In Brasilia meridionali”, Sellow s.n. (lectotype B†, designated by Epling 1936b, replacement lectotype K!, designated here; isolectotypes A!, E!, F!, HAL!, LE!, UC!).

Cantinoa carpinifolia (Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis carpinifolia Bentham (1833: 115) Ξ Mesosphaerum carpinifolium (Benth.) Kuntze (1891: 526) . Type:— BRAZIL. “In Brasilia meridionali”, Sellow s.n. (lectotype B†, designated by Epling 1936b; replacement lectotype K!, designated here).

= Hyptis aquatica Pohl ex Bentham (1833: 116) Ξ Hyptis carpinifolia var. aquatica (Pohl ex Benth.) Schmidt (1858: 130) . Type:— BRAZIL. Minas Gerais: Rio da Prata, Pohl 2924 (holotype W!; isotype K!).

Cantinoa colombiana (Epling) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis colombiana Epling (1936b: 257) . Type:— COLOMBIA. Magdalena: Santa Marta, Smith 1490 (holotype US!; isotypes BR!, COL!, G!, K!, NY!, P!).

Cantinoa dubia (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis dubia Pohl ex Bentham (1833: 122) Ξ Mesosphaerum dubium (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Minas Gerais: Rio Abaite, Pohl 3276 (holotype W!; isotype K!).

= Hyptis expansa Pohl ex Bentham (1833: 122) Ξ Mesosphaerum expansum (Pohl ex Benth.) Kuntze (1891: 526) . Type:— BRAZIL. Without locality, Pohl s.n. (holotype W!).

Cantinoa duplicatodentata (Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis duplicatodentata Bentham (1833: 114) Ξ Hyptis vestita Benth. var. duplicatodentata (Pohl ex Benth.) Schmidt (1858: 128) Ξ Mesosphaerum duplicatodentatum (Pohl ex Benth.) Kuntze (1898: 260) . Type:— BRAZIL. Minas Gerais: Rio Jequitinhonha, Pohl 1912 (lectotype W!, designated by Epling 1936b; isolectotypes F!, K!).

= Hyptis duplicatodentata Pohl ex Benth. var. virescens Pohl ex Bentham (1833: 114) . Type:— BRAZIL. Minas Gerais: Manoel Pereira, Pohl 2968 (holotype W!; isotypes A!, K!).

Cantinoa erythrostachys (Epling) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis erythrostachys Epling (1936b: 258) . Type:— BRAZIL. Minas Gerais: Congonhas do Campo, Stephan 55 (holotype BR; isotype fragment UC!).

Cantinoa heterodon (Epling) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis heterodon Epling (1936b: 243) . Type:— BRAZIL. São Paulo: Apiaí, Puiggari 3234 (holotype P!; isotype UC!).

Cantinoa impar (Epling) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis impar Epling (1936b: 257) . Type:— BRAZIL. Mato Grosso: Triunfo, Rios Cuiabá e São Lourenço , Hoehne 4563 (holotype UC!).

Cantinoa indivisa (Pilg.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis indivisa Pilger (1901: 190) . Type:— BRAZIL. Mato Grosso: Cuiabá, 2 April 1899, Pilger 399 (holotype B†, photo!).

Cantinoa macrotera (Briq.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis macrotera Briquet (1898: 210) . Type:— BRAZIL. Minas Gerais: entre “Barbacena et Sitio”, 23 June 1879, Glaziou 11314 (holotype G!; isotypes BR!, C!, K!, P!, UC!).

Cantinoa multiseta (Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis multiseta Bentham (1848: 122) . Type:— BRAZIL. Minas Gerais: Serra das Araras, June 1840, Gardner 5104 (holotype K!; isotypes BM!, E!, OXF!, P!, US!, W!).

Epling (1949) treats this species in Hyptis sect. Mesosphaeria , and in the same publication also places it in synonymy under H. mutabilis in Hyptis sect. Polydesmia .

Cantinoa muricata (Schott ex Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis muricata Schott ex Bentham (1833: 119) . Type:— BRAZIL. Rio de Janeiro: Schott 6167 (holotype W!).

Cantinoa mutabilis (Rich.) Harley & J.F.B.Pastore , comb. nov. Ξ Nepeta mutabilis Richard (1792: 110) Ξ Mesosphaerum mutabile (Rich.) Kuntze (1891: 525) Ξ Hyptis mutabilis (Rich.) Briquet (1896: 788) . Type:— FRENCH GUIANA. Cayenne, Le Blond s.n. (holotype P!; isotype G!).

= Hyptis spicata Poiteau (1806: 474 , t. 28, f. 2). Type:— DOMINICAN REPUBLIC. Santo Domingo, Richard s.n. (holotype P!).

= Hyptis canescens Kunth in Humboldt, Bonpland & Kunth (1818: 321). Type:— VENEZUELA. Distrito Federal: Caracas, Humboldt & Bonpland s.n. (holotype P!).

= Hyptis polystachya Kunth in Humboldt, Bonpland & Kunth (1818: 321). Type:— MEXICO. Michoacán: inter Pátzcuaro & Ario Mexicanorum, Humbold & Bonpland s.n. (holotype P!).

= Hyptis micrantha Pohl ex Bentham (1833: 120) . Type:— BRAZIL. Goiás: ad Villa Boa, Pohl 1619 (holotype W!; isotype BR!, K!).

= Hyptis tenuiflora Bentham (1833: 121) . Type :— BRAZIL. Without locality, Sacramento s.n. (holotype P!).

= Hyptis rostrata Salzm. ex Bentham (1833: 121) . Type:— BRAZIL. Bahia: near Salvador, Salzmann s.n. (lectotype K!, designated by Epling 1936b; isolectotypes G!, P!, W!).

= Hyptis arvensis Poeppig ex Bentham (1835: 712) . Type:— PERU. Huánuco: Cuchero, Poeppig 1097 (holotype BM!; isotype G-DC!).

= Hyptis aspera Martins & Galeotti (1844: 189) . Type:— MEXICO. Veracruz: Mirador, Galeotti 620 (holotype BR!; isotype K!).

= Mesosphaerum yungasense Britton ex Rusby (1895: 246) . Type:— BOLIVIA. La Paz: Yungas, Bang 622 (holotype NY!; isotype K!, US!) .

= Hyptis trichocalyx Briquet (1897a: 21) . Type:— PARAGUAY. Caaguazu: Cosme, Balansa 996 (holotype P!).

= Hyptis singularis Glaziou (1911: 554) , nom. nud. Reference specimen:— BRAZIL. Minas Gerais: Serra de Mantiqueira, João Aires , Glaziou 1130 (P!).

= Hyptis kerberi Gandoger (1918: 66) . Type:— MEXICO. Jalisco: Atoyac, Kerber 139 (holotype LY; isotype P!).

= Hyptis canaminensis Rusby (1927: 342) . Type:— BOLIVIA. La Paz: Cañamina, Rusby 61 (holotype NY!; isotype UC!, fragment).

Cantinoa × obvallata (Sprengel ex Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis obvallata Sprengel ex Bentham (1833: 115) , pro spec. Type:— BRAZIL. Minas Gerais: Carandahy, Sellow 1489 (lectotype B†, designated by Epling 1936b, replacement lectotype K!, designated here; isolectotypes A!, G-DC!, HAL!, P!, W!).

For further details see Harley (1999) .

Cantinoa pinetorum (Epling) Harley & J.F.B.Pastore , comb.nov. Ξ Hyptis pinetorum Epling (1933: 103) . Type:— MEXICO. Jalisco: San Sebastián, Mexia 1396 (holotype UC!; isotypes BM!, G!).

Cantinoa plectranthoides (Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis plectranthoides Bentham (1833: 122) Ξ Mesosphaerum plectranthoides (Benth.) Kuntze (1891: 525) . Type:— BRAZIL. “Brasilia meridionali” [Minas Gerais: “Matheus Lemes”, fide Epling], Sellow 1942 (lectotype B†, designated by Epling 1936b, replacement lectotype K!, designated here; isolectotypes G!, HAL!).

= Hyptis communis A.St. -Hil. ex Bentham (1833: 123). Type:— BRAZIL. Minas Gerais: without locality, St.- Hilaire 277 (lectotype P! designated by Epling 1936b; isolectotype F! fragment, K!).

The sheet in P originally annotated as type by Epling is different from the above and was selected in error. The specimen which Bentham cited in the protologue came from Minas Gerais, which was not the case of the annotated specimen, but Epling obviously realized his error before publication .

= Hyptis pumila Pohl ex Bentham (1833: 122) Ξ Mesosphaerum pumilum (Pohl ex Benth.) Kuntze (1891: 525) . Type:— BRAZIL. Goiás: “ad Serra de Chrystais”, Pohl 6060 (holotype W!).

Cantinoa propinqua (Epling) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis propinqua Epling (1936b: 243) . Type:— BRAZIL. Rio de Janeiro: Serra dos Orgãos, May 1837, Gardner 574 (holotype K!; isotypes BM!, G!, OXF!, UC!, W!).

Cantinoa racemulosa (Mart. ex Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis racemulosa Mart. ex Bentham (1833: 126) Ξ Mesosphaerum racemulosum (Mart. ex Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Minas Gerais: Villa Rica, Martius Obs. 780 (lectotype M!, designated here, but see note below).

In the protologue, after the species epithet, Bentham (1833) added “Martius MSS” in parenthesis, and he clearly referred to a specimen he saw in Munich (only). Epling (1936b) lectotypified Hyptis racemulosa with the Martius duplicate at Kew ( Epling 1936b: 243), saying that he could find no specimen either at M nor in the Martius herbarium (in BR). However a specimen exists in M, Martius Obs. 780, which is annotated by Bentham as Hyptis racemulosa , and also by Epling as “ Type ”.

Cantinoa rubicunda (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis rubicunda Pohl ex Bentham (1833: 118) Ξ Mesosphaerum rubicundum (Pohl ex Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Minas Gerais: Barra do Rio das Velhas, Pohl 1769 (lectotype W!, designated by Epling 1936b; isolectotype K!).

= Hyptis rubicunda var. grandifolia Bentham (1848: 119) . Type:— BRAZIL. Goiás: Serra da Santa Brigida, April 1840, Gardner 3925 (holotype K!).

Cantinoa similis (Epling) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis similis Epling (1936b: 258) . Type:— BRAZIL. Mato Grosso: Cuiabá, Caxipó do Ponte , Hoehne 4548 (holotype UC!).

Cantinoa stricta (Benth.) Harley & J.F.B.Pastore , comb. nov. ≡ Hyptis stricta Bentham (1833: 79) ≡ Mesosphaerum strictum (Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Rio Grande do Sul: Cerro Agudo, 23 March 1823, Sellow 4580 (lectotype B†, designated by Epling 1936b, replacement lectotype K!, designated here).

Bentham published this name with the protologue given as: “6.? H. STRICTA (Benth: in Herb. Mus. Reg. Berol. MSS.)”. See remarks under Condea undulata (Schrank) Harley & J.F.B.Pastore.

Cantinoa subrotunda (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis subrotunda Pohl ex Bentham (1833: 118) ≡ Mesosphaerum subrotundum (Pohl ex Benth.) Kuntze (1891: 527) . Type:— BRAZIL. Goiás: Ponte Feito, Pohl 2678 (holotype W!; isotype K!).

= Hyptis parvifolia Pohl ex Bentham (1833: 118) Ξ Hyptis subrotunda var. angustifolia Schmidt (1858: 131) . Type:— BRAZIL. Goiás: Megaponte [Meiaponte, now Pirenópolis], Pohl 2789 (holotype W!).

Although Pohl material from this locality is cited as Megaponte by Epling, it appears that Pohl wrote “Meyaponte” on the label. There has been much confusion over this name.

Cantinoa × sylvularum (A.St.-Hil. ex Benth.) Harley & J.F.B.Pastore, comb. nov. Ξ Hyptis sylvularum A.St. - Hil. ex Bentham (1833: 119), pro spec. Ξ Mesosphaerum sylvularum (A.St.-Hil. ex Benth.) Kuntze (1891: 527). Type:— BRAZIL. Minas Gerais: capoeiras, Saint-Hilaire 555 (holotype P!; isotype P!).

For further details, see Harley (1999) .

Cantinoa villicaulis (Epling) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis villicaulis Epling (1941: 554) . Type:— BRAZIL. Mato Grosso: Marimbondo, Rio São Lourenço , Hoehne 2866 (holotype UC!).

Cantinoa violacea (Pohl ex Benth.) Harley & J.F.B.Pastore , comb. nov. Ξ Hyptis violacea Pohl ex Bentham (1833: 116) . Type:— BRAZIL. Goiás: Trahiras, Pohl 1880 (holotype W!; isotype K!).

= Hyptis arida A.St. -Hil. ex Bentham (1833: 116) Ξ Mesosphaerum aridum (A.St.-Hil. ex Benth.) Kuntze (1891: 525). Type:— BRAZIL. Goiás: “in desertis meridionalibus” Saint-Hilaire 816 (holotype P!; isotype P!).

Condea Adanson (1763: 504) . Type:— Condea americana (Poir.) Harley & J.F.B.Pastore [= Satureja americana Poiret in Lamarck (1805: 571), lectotype designated by Kuntze 1891].

= Hypothronia Schrank (1824: 85) . Type:— Hypothronia undata Schrank (1824: 85) [= Condea undulata (Schrank) Harley & J.F.B.Pastore ]. See note under this species.

The genus Condea is characterized by an often spiciform or raceme-like thyrse of pedunculate or sessile cymes in which the intercalary cyme-axes are contracted. When cymes pedunculate, the flowers, which often bear long pedicels, appear subumbellate, or appear fasciculate when peduncle absent. Flowers usually small, gynoecium without stylopodium. There are 26 species, ranging from North America (One species in the western United States) and Central America to the Caribbean and South America.

The genus is divided into two sections here, which correspond to three sections recognized by Epling under Hyptis (1949) . Condea sect. Condea is exactly equivalent to H. sect. Minthidium , while H. sect. Umbellatae Epling is placed in synonymy under C. sect. Laniflorae (Epling) Harley & J.F.B.Pastore.

Figures

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.