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Phytotaxa 58: 1–55 (2012) 

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Copyright © 2012 Magnolia Press 

ISSN 1179-3155 (print edition) 

PHYTOTAXA 

ISSN 1179-3163 (online edition) 

A generic revision and new combinations in the Hyptidinae (Lamiaceae), based 

on molecular and morphological evidence 

R.M. HARLEY¹ & J.F.B. PASTORE² 

1 

Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK and Post-graduate Programme in Botany, Depto de Ciências 

Biológicas, Universidade Estadual de Feira de Santana, Km 03, BR116, Campus, Feira de Santana, 44031-460 Bahia, Brazil. 

E-mail: rharley05@aol.com 

2 

Post-graduate Programme in Botany, Depto de Ciências Biológicas, Universidade Estadual de Feira de Santana, Km 03, BR116, 

Campus, Feira de Santana, 44031-460 Bahia, Brazil. 

Table of contents 

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 

Taxonomic History of Hyptis and relations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 

Infrageneric classification of Hyptis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 

Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 

Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 

Key to Genera of Hyptidinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 

Conspectus of Hyptidinae with new classification including new combinations, synonyms and typifications . . . . . . . . . . . . . . . . . . . 8 

Sections unplaced to genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 

Appendix 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 

Abstract 

An earlier molecular study demonstrated the monophyly of the Hyptidinae and most of the genera within it. However, 

the largest genus, Hyptis, is paraphyletic and all other genera seem to be derived from a Hyptis ancestor. Most of the 

different lineages which comprise Hyptis are already established sections, some of which are now raised to generic rank, 

augmenting the subtribe to 19 genera and with a greatly reduced but monophyletic Hyptis, in which the genus Peltodon is 

included as a section. The sections Mesosphaeria and Polydesmia are also shown not to be monophyletic, making it 

necessary to reassign some species from the former to the latter, and then raising the two sections to generic rank as 

Mesosphaerum and Cantinoa respectively, the latter a new genus name. A new genus Oocephalus is also created from 

two former subsections of Hyptis sect. Polydesmia. The genus Condea, formed from three former sections of Hyptis, is 

itself divided into two sections. A key to all recognized genera is included, together with a generic conspectus, with brief 

distributional data, differentiating the new genera and listing their component species. The necessary nomenclatural 

changes comprise 142 new combinations, 30 new or replacement lectotypifications, four neotypes, 23 new synonymies, 

eight stat. nov., six newly coined generic names, five new epitypes and four new names. 

Key words: generic key, Hyptis, molecular data, morphology, new genera, taxonomy 

Introduction 

The Lamiaceae have recently undergone major changes of both delimitation and reorganization of constituent 

taxa, as a result of morphological and molecular phylogenetic studies (e.g., Cantino 1992a, 1992b, Wagstaff et 

al. 1995, 1998), with many genera, formerly placed in Verbenaceae, being incorporated (Harley et al. 2004). 

Accepted by Hajo Esser: 18 Jun. 2012; published online in PDF: 27 Jun. 2012 1

Currently, the family comprises about 7200 species, the largest within the order Lamiales, with 240 

genera, divided among seven subfamilies (Harley et al. 2004), of which subfam. Nepetoideae contains over 

50% of all species. Molecular studies on various groups of Nepetoideae have been published, in particular on 

Mentheae by Bräuchler et al. (2010), on Ocimeae by Paton et al. (2004) and more recently by Zhong et al. 

(2010) on the phylogeny of Isodon (Schrad. ex Benth.) Spach (Ocimeae) and allies, suggesting a close link 

between Hyptidinae and Hanceolinae. All these studies are causing a re-evaluation of morphological and 

other non-molecular characters, as well as advancing our understanding of probable evolutionary lines and 

past geographical dispersion. The Ocimeae, distributed through tropical regions, are characterized by having 

stamens declinate above the anterior corolla lip. Although with a largely Old World distribution, one subtribe, 

the Hyptidinae, which forms the subject of this paper, is almost exclusively neotropical, occurring throughout 

tropical and subtropical America, with perhaps two species extending their natural range to Africa, as well as 

several weedy species which are introduced into the Palaeotropics (Harley et al. 2004). Most species are to be 

found in a wide range of primarily savanna habitats, often in upland areas. The Hyptidinae are characterized 

by flowers arranged in variously modified bracteolate cymes, with corollas bearing stamens decurrent and 

held within the hinged anterior corolla lobe, which provides an explosive pollination mechanism (Harley 

1971), and by the possession of nutlets with an expanded areole (Paton & Ryding 1988). 

Taxonomic History of Hyptis and relations 

Bentham (1833), who published the first detailed account of the Labiatae, recognized four genera in the 

group: Peltodon Pohl, Marsypianthes Mart. ex Benth., Hyptis Jacq. and Eriope Humb. & Bonpl. ex Benth., 

considering it a natural group, although the name Hyptidinae did not receive formal recognition until it was 

proposed by Endlicher (1838). The largest of the genera is Hyptis, currently with about 280 species, of which 

Bentham wrote: “it would be more convenient to divide it into a number of genera”, but he was unable to find 

a satisfactory means of separating the wide range in inflorescence form that he encountered. Instead he treated 

the various groups he recognized as sections, later amplifying these in De Candolle’s Prodromus (Bentham 

1848). Also at this time, Schauer (1844) separated a fifth genus, the monotypic Rhaphiodon Schauer, treated 

by Bentham as a species of Hyptis. Kuntze (1891), in the Revisio Genera Plantarum, having discovered an 

earlier name for Hyptis, published a very large number of new combinations under, the generic name of 

Mesosphaerum P.Browne (1756). These however were not generally accepted, leading to Hyptis Jacq. being 

made a conserved name (see McNeill et al. 2006) and considerably adding to the synonymy. No further 

generic additions were made, until Epling (1932) further dismembered Hyptis by publishing the Central 

American genus Asterohyptis Epling, with three species, characterized by the much reduced corollas and 

lacking the explosive pollination mechanism. This was followed by the recognition of Eriopidion Harley 

(1976), a monotypic genus, removed from Eriope, and the subsequent creation of Hyptidendron Harley and 

Hypenia (Mart. ex Benth.) Harley (Harley 1988), removing three sections from Hyptis to raise the number of 

generic taxa in the Hyptidinae to nine. Eight of these were accepted in the most recent account (Harley et al. 

2004), although Atkinson (1999, unpublished thesis) suggested placing all species of Hypenia within Eriope, 

a change not justified according to Pastore et al. (2011) and not followed in the present paper. 

Up to the mid-20 th century, undoubtedly the most important contributions to the taxonomy of New World 

Labiatae were made by Carl Epling, those dealing with the Hyptidinae being particularly relevant here. Two 

publications, A Synopsis of South American Labiatae (Epling 1935, 1936a, 1936b, 1937) and the Revisión 

del género Hyptis (Labiatae) (Epling 1949), remain standard works, and established the infrageneric 

classification in Hyptis (see below). The most recent overview of the group (Harley et al. 2004) recognizes 

eight genera: Hyptidendron, Eriope, Hypenia, Marsypianthes, Hyptis, Peltodon, Rhaphiodon and 

Asterohyptis. Eriopidion was here placed in the synonymy of Eriope. 

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HARLEY & PASTORE

Infrageneric classification of Hyptis 

Bentham (1833) recognized 19 sections within Hyptis (Table 1), further dividing sect. Cephalohyptis Benth. 

into a number of subsections. Subsequently, he modified this system and recognized 20 sections (Bentham 

1848), further dividing six of these into a series of subsections. Many of these taxa are still in current use. A 

number of minor changes were made by Schmidt (1858) in an account of Labiatae for the Flora Brasiliensis. 

He also introduced an entirely new infrageneric classification for Hyptis, based on inflorescence structure, 

which has been ignored by later workers. John Briquet, from Geneva, published an extensive series of papers 

on Labiatae, mostly in the latter half of the 19 th century, which included many new taxa of Hyptidinae, 

especially from Paraguay. He also extensively revised infrageneric classifications, especially of Hyptis (Table 

1), in Engler & Prantl’s Pflanzenfamilien (Briquet 1897b). Epling (1933, 1935, 1936a, 1936b, 1937) further 

modified the infrageneric classification of Hyptis, basing his system mainly on Bentham, but with some 

changes proposed by Briquet. Later, in his revision of the genus (Epling 1949), he recognized a total of 26 

sections, of which 15 had originally been proposed by Bentham and one by Briquet (1897), although 

subsequent changes had resulted in very different sectional delimitations (Table 1). Changes included Hyptis 

sects. Spicaria Benth. and Pectinaria Benth. being treated as subsections under H. sect. Mesosphaeria Benth.; 

Hyptis sect. Oocephalus Benth. became a subsection under H. sect. Polydesmia Benth.; Hyptis sect. 

Xanthiophoea Benth. was removed, with the three constituent species placed elsewhere: one becoming the 

type of the genus Rhaphiodon, another one, Hyptis lagenaria A.St.-Hil. ex Benth., being placed in H. sect. 

Cyrta Benth., and the third, Hyptis lobata A.St.-Hil. ex Benth., becoming the only representative of a new 

section, H. sect. Hilaria Epling; Hyptis sect. Turbinaria Benth. was placed in synonymy of Hyptis sect. 

Umbellaria Benth. [= Hyptidendron sect. Umbellaria (Benth.) Harley], H. sect. Siagonarrhen Mart. ex Benth. 

subsect. Cymosae Benth. in synonymy of sect. Buddleioides Benth., and H. sect. Siagonarrhen subsect. 

Nudiflorae Benth. into synonymy of Hyptis sect. Hypenia subsect. Densiflorae Benth. Epling (1933, 1936, 

1937) also created 10 new sections (Table 1). He also raised H. sect. Cephalohyptis Benth. subsect. 

Xylodontes Benth. to sectional rank as H. sect. Xylodontes (Benth) Epling. 

In the introduction to his monograph of the genus Hyptis, Epling (1949) followed Bentham in suggesting 

that a case could perhaps be made for the further division of Hyptis into a greater number of genera. While he 

felt at that time that this revolutionary step was not justified, increased knowledge, based on vastly increased 

collections, has now indicated that the idea was not without foundation. 

Harley (1976) also made a number of changes, including the change of the name of the section which 

contains the generic type, H. sect. Cephalohyptis, to H. sect. Hyptis to conform to ICBN rules (see McNeill et 

al. 2006). Five sections accepted by Epling were placed in synonymy or received a change of status. Harley 

transferred the only species of Hyptis sect. Mixtae Epling to Eriope (Harley 1976). In the same publication he 

also removed the species of Hyptis sect. Siagonarrhen subsect. Nudiflorae, including its type, Hyptis latifolia 

Mart. ex Benth., into the synonymy of Eriope. Later, Hyptis sect. Pachyphyllae (Epling) Harley, based on H. 

sect. Eriosphaeria subsect. Pachyphyllae Epling, was created (Harley 1986b). He also transferred H. sect. 

Umbellaria and H. sect. Buddleoides to the genus Hyptidendron, the former as a subsection and the latter in 

synonymy, and, in the same paper, raised H. sect. Hypenia to generic rank (Harley 1988). For a summary of 

these changes see Table 1. As a result of all these taxonomic changes, the number of accepted sections 

currently stands at 24. Most of these taxa had been defined by characters of the inflorescence and floral 

structure, and many form well-defined groups. 

Concurrently with the taxonomy, a series of other studies has substantially contributed to our 

understanding of the interrelationships with the Hyptidinae. Among these are a series of anatomical studies by 

Rudall (1979, 1980, 1981b, 1981c), a brief palynological survey (Rudall 1981a), which now suggests a more 

detailed study would provide significant data to support the new taxonomy, and a survey of chromosome 

numbers (Harley & Heywood 1992) in which some possible relationships were discussed. More recently, 

Pastore et al. (2011) published a molecular phylogenetic analysis of the Hyptidinae, using nuclear sequences 

from the ITS and ETS regions, complemented by sampling four plastid regions (trnL-F, matK, trnS-G, trnD- 

A GENERIC REVISION OF HYPTIDINAE (LAMIACEAE) 

Phytotaxa 58 © 2012 Magnolia Press 3

T). The sample included all genera of Hyptidinae, 22 of the 24 sections of Hyptis and totalled up to 180 

species. Most of the large number of subsections recognized by Epling were also sampled, only a very few of 

these being unobtainable. 

TABLE 1. Changes in the classification of Hyptidinae. 

Hyptis sections are in normal type and genera are in bold-face. The arrows indicate the changes made. 

The analyses demonstrate the monophyly of the Hyptidinae and of its constituent genera except for 

Hyptis, provided that Hypenia vitifolia (Pohl ex Benth.) Harley is excluded from Hypenia, Eriope simplex 

(A.St.-Hil. ex Benth.) Harley is transferred to Hypenia and Hyptis eximia Epling is transferred to 

Hyptidendron. These taxonomic changes are supported by the morphology. Indeed, for some time, the 

removal of Hyptis vitifolia from Hypenia had been considered by the senior author on purely morphological 

grounds. Hyptis, meanwhile, is shown to be highly polyphyletic, and represented by nine different lineages, 

often composed of single former sections of Hyptis. Two sections, H. sect. Mesosphaeria and H. sect. 

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HARLEY & PASTORE

Polydesmia, require some relocation of species, while H. sect. Minthidium Benth., H. sect. Polydesmia 

subsect. Oocephalus and H. sect. Laniflorae Epling are grouped to form a clade. The recognition of these at 

the generic level would leave Hyptis monophyletic but still very diverse. At the same time, the former genus 

Peltodon could now only be recognized at sectional level within Hyptis. There is however a need for much 

wider sampling within Hyptis sensu stricto, to resolve its internal sectional structure. 

Materials and Methods 

Taxonomic studies of a large taxon such as the subtribe Hyptidinae, requires a synthesis of data obtained from 

a wide range of disciplines, summarized above, especially detailed studies of herbarium material 

representative of the taxa studied. Over many years, the senior author has been studying this group, both in the 

field and in the herbaria listed here: AAU, ALCB, B, BM, BR, C, CEN, CEPEC, COL, CTES, E, F, FLOR, G, 

HAL, HBR, HUEFS, INPA, IPA, IS, JBSD, JE, K, LE, LINN, M, MA, MANCH, MBM, MG, MO, MS, NY, 

OXF, P, R, RB, S, SP, SPF, TEPB, UB, UC, UEC, UFG, US, W, Z, ZT, with a view to preparing accounts for 

Flora Neotropica. Detailed morphological studies have provided an insight into their relationships, which with 

the new data from molecular studies enables us to set out a new classification. For the purposes of this paper, 

the original bibliography has been examined wherever possible, to check dates of publication, protologues 

and typification, and the volumes of Taxonomic Literature by Stafleu & Cowan published between 1976 and 

1988 have been frequently consulted. 

Conclusions 

The results from the molecular studies (Pastore et al. 2011) clearly require major changes to be made to the 

current classification of the Hyptidinae. One solution would be to reduce all other genera, including 

Marsypianthes and Eriope, to synonymy under Hyptis. This would leave a genus of over 400 species with a 

range of morphology, both vegetative and reproductive which would transcend that found in possibly any 

other genus of Angiosperms. The other alternative, chosen here, is to reorganize groupings along cladistic 

lines, redefining groupings where indicated, and recognizing a much larger number of genera, which would be 

morphologically more homogeneous, more narrowly defined and justified also on morphological, and in 

many cases by chromosomal and anatomical evidence also. A diagram displaying the phylogenetic 

relationships between the genera recognized here is displayed in Fig. 5. Nevertheless, these changes leave an 

open field for more detailed studies on a range of characters, often not previously considered significant, 

which will hopefully provide further support for the taxonomic changes now proposed. 

The decision to erect a substantial number of new genera has resulted in a careful assessment of names 

already available, and names which need to be created. Hyptis was conserved against both Condea Adans. and 

Mesosphaerum P.Browne, which are earlier names (McNeill et al. 2006). Both of these now become 

available, as their types no longer fall within Hyptis as defined in this paper. Only two of the appropriate 

sectional names in Hyptis can be raised to generic rank: Cyanocephalus Benth. and Gymneia Benth., the 

others being already occupied. Apart from the large number of changes proposed for many New World 

species, there are also a number of weedy species introduced into the Old World tropics which are affected: 

Hyptis suaveolens (L.) Poit. and Hyptis pectinata (L.) Poit. become Mesosphaerum suaveolens (L.) Kuntze 

and M. pectinatum (L.) Kuntze respectively, and Hyptis mutabilis (Rich.) Briq. becomes Cantinoa mutabilis 

(Rich.) Harley & J.F.B.Pastore. Hyptis spicigera Lam. takes up the earlier Aublet epithet, which could not be 

used under Hyptis, to become Cantinoa americana (Aubl.) Harley & J.F.B.Pastore. Other species, such as 

Hyptis brevipes Poit., H. lanceolata Poir. and H. capitata Jacq., remain unchanged. 

The following text provides a provisional key to genera, a conspectus of Hyptidinae, with descriptions 

and new combinations, where needed. The establishment of the new classification and nomenclature is a 



priority and opens up the need for a more detailed search for characters which may prove to be diagnostic of 

the newly recognized genera, beyond those already used. Also, a number of species, which have been difficult 

to obtain, still require analysis to confirm their taxonomic position, although their absence does not affect the 

overall taxonomy and these are listed in the conspectus as “unplaced”. Within Hyptis (sensu stricto) much 

wider sampling is still required to evaluate infrageneric groupings and this is a task for the future. A list of 

Hyptidinae, with original names, the pre-1950 taxa as recognized by Epling, with their new taxonomic 

position, is provided (Appendix 1). Bibliographic references indicate where accepted names were published, 

if these occurred prior to this paper, and names which contain a new epithet are also listed. Epithet changes 

which only involve a change of gender are not provided. 

Key to Genera of Hyptidinae 

1. Anterior lip of corolla thickened at base and reflexing at anthesis to release the stamens explosively. Flowers 

arranged variously. Plants of tropical and subtropical America ............................................................................. 2 

1*. Anterior lip of corolla not thickened at base, nor reflexing at anthesis to release stamens explosively. Flowers in 

slender, elongate spikes, arranged in sessile or subsessile verticillasters, often 2 per leaf-axil, 6−12-flowered. 

Plants of Mexico and Central America ................................................................................................. Asterohyptis 

2. Cymes usually 1-flowered (rarely up to 3−6-flowered), flowers with often inconspicuous paired bracteoles at base 

of calyx, or if not as described, then flowering stems waxy with often inflated internodes ................................... 3 

2*. Cymes usually many-flowered, flowers sometimes congested or modified into bracteate heads or capitula†, or in 

axillary fascicles. Paired bracteoles not present at base of calyx. Flowering stems never waxy or inflated .......... 6 

3. Cymes up to 3−6-flowered, pedunculate, flowers shortly pedicellate, calyx lacking inconspicuous paired 

bracteoles at base ................................................................................................................................ Physominthe 

3*. Cymes usually uniflorous and forming a raceme-like, often branched inflorescence, calyx with inconspicuous 

paired bracteoles at base, above a long or short peduncle (pseudo-pedicel), rarely some cymes 3-flowered and 

then pedicels long, slender ...................................................................................................................................... 4 

4. Calyx in fruit zygomorphic, with lobes unequal, posterior lip rounded or with posterior lobes partly connate. 

Corolla at anthesis, with tube abruptly contracted near base, usually lilac, pink or violet, sometimes yellowish in 

bud ........................................................................................................................................................................... 5 

4*. Calyx in fruit actinomorphic, or almost so, with subequal lobes. Corolla at anthesis, with tube not abruptly 

contracted at base, lilac or pale blue, cream, yellow or red ........................................................................ Hypenia 

5. Calyx throat open, though sometimes closed by dense white hairs. Corolla tube often broadly campanulate or 

funnel-shaped. Base of style persistent, with stylopodium overtopping nutlets. Nutlets broad, slightly flattened or 

rarely winged .................................................................................................................................................. Eriope 

5*. Calyx hygroscopic, throat closed by upper lobes when dry, lobes with a row of rigid hairs within. Corolla tube 

shortly and narrowly cylindrical. Stylopodium absent. Nutlets elongate, ±triquetrous .......................... Eriopidion 

6. Flowers in lax, few-flowered cymes, or cymes many-flowered and then in ± spherical capituliform heads. Calyx 

lobes triangular, often reflexing in fruit. Gynoecium with persistent, short, quadrangular stylopodium equalling 

nutlets and attached to them until maturity. Nutlets cymbiform with an involute, laciniate margin and concave 

inner face ........................................................................................................................................... Marsypianthes 

6*. Flowers arranged variously. Calyx lobes not reflexing in fruit. Stylopodium if present not as above. Nutlets ovoid 

or flattened, never concave nor laciniate................................................................................................................... 7 

7. Flowers in a spherical capitulum, dropping as a unit in fruit. Corolla tubular, deep purple. Calyx with 5--10 

unequal spines ....................................................................................................................................... Rhaphiodon 

7*. Flowers not as above, if in a capitulum or a capituliform head not dropping as a unit in fruit. Calyx lobes 5, not 

spinose, but sometimes subulate and rigid at apex ................................................................................................. 8 

8. Flowers sessile to subsessile, in an ovoid, hemispherical or globose capitulum or capituliform head, surrounded 

by a distinct involucre of filamentous, ligulate to ovate bracteoles; when filamentous these sometimes obscured 

when capitula globose at anthesis. Capitula often pedunculate, forming panicles or corymbs or sometimes axillary 

from reduced or leaflike bracts, rarely sessile and forming elongate spiciform inflorescences ............................. 9 

8*. Flowers subsessile to long-pedicellate, variously arranged in a pedunculate or subsessile, congested or lax cyme 

but not in a capitulum, sometimes forming a long, spiciform inflorescence, or flowers in compact, cincinnate 

verticillasters or in fascicles, rarely solitary and then usually long-pedicellate, or in subumbellate, pedunculate or 

weakly globose cymes, bracteoles not forming an involucre, or if involucre present, bracteoles usually slender and 

enclosing a 10–15-flowered cymule or obscured in a dense, elongate broadly spiciform inflorescence ............. 13 


HARLEY & PASTORE

9. Flowers in ovoid heads, sessile or pedunculate and often enclosed by broad concave bracteoles when immature. 

Corollas long tubular, with short lobes, not spotted or marked on upper lip ........................................ Oocephalus 

9*. Flowers in hemispherical to globose capitula, with an involucre of ovate to ligulate, subulate or filamentous 

bracteoles. Corollas various, often spotted on upper lip ....................................................................................... 10 

10. Calyx lobes 3–4 times longer than the slender tube and terminating in a long filamentous apex. Capitula globose 

with long filamentous involucral bracteoles ........................................................................................ Medusantha 

10*. Calyx lobes shorter, filamentous to ovate, capitula hemispherical or globose ..................................................... 11 

11. Capitula globose, > 10 mm diam., with filamentous or narrowly linear bracteoles which are often obscured by 

reflexing flowers at anthesis. Calyx tube usually strongly deflexed in mid-tube. Peduncles usually longer than 

adjacent internode .................................................................................................................................................. 12 

11*. Capitula hemispherical, with involucre of subulate, ligulate or lanceolate to ovate bracteoles, these usually not 

reflexing at maturity, or if globose with calyx tube deflexed, then less than 10 mm diam. and peduncles shorter 

than adjacent internode ................................................................................................................................... Hyptis 

12. Calyx lobes clavate, widening slightly near apex, stigma ± capitate. plants typical of cerrado and similar 

formations . ....................................................................................................................................... Cyanocephalus 

12*. Calyx lobes subulate, never clavate, stigmas bilobed, plants of sandy areas in semi-arid regions ..... Martianthus 

13. Stylopodium present, flowers in usually lax cymes. Trees or shrubs, rarely herbs ........................... Hyptidendron 

13*. Stylopodium absent, flowers disposed variously. Shrubs, subshrubs or herbs, rarely trees ................................. 14 

14. Flowers sessile or subsessile, in few-flowered sessile cymes, with slender bracteoles, in the axils of reduced, 

inconspicuous bracts, forming slender, elongate, spiciform inflorescences. Calyx with ± scarious, deltoid flanges 

in sinus between calyx lobes, corolla tubular .........................................................................................Leptohyptis 

14*. Flowers arranged in fascicles in the axils of leaf-like bracts, or in sub-umbellate or congested, pedunculate cymes, 

or in globose verticillasters or cincinnate or few-flowered cymes, if inflorescence spiciform, usually not slender. 

Calyx lobes without flanges in sinus ..................................................................................................................... 15 

15. Flowers in fascicles, rarely solitary or flowers few on long pedicels, in the axils of often leaf-like bracts, or in 

shortly pedunculate sub-umbels, often forming elongate, raceme-like inflorescences, or rarely panicles. Corolla 

never blue. In varied habitats, sometimes subject to inundation ................................................................. Condea 

15*. Flowers not in fascicles or pedunculate sub-umbels.............................................................................................. 16 

16. Flowers in a dense, head-like cyme, on a short peduncle from the axils of leaf-like bracts. Leaves small. Corolla 

blue or violet-blue. Shrubs of sandy, semi-arid areas in NE Brazil ........................................................ Eplingiella 

16*. Flowers not as above. Leaves and corolla various................................................................................................. 17 

17. Flowers in dense subglobose or globose verticillasters, formed from congested cincinni, in the axils of reduced 

bracts, and forming an elongate, often interrupted or congested terminal spike, with leaves often developed 

toward base of flowering stem. Bracteoles setaceous, rigid and almost spine-tipped, calyx strongly deflexed in 

mid-tube ...................................................................................................................................................... Gymneia 

17*. Flowers in cincinnate or shortly dichotomous cymes, but never forming subglobose verticillasters. Bracteoles 

various, but not as above. Calyx tube straight . ..................................................................................................... 18 

18. Inflorescence usually an elongate spiciform or racemose thyrse. Flowers in pedunculate or sessile cymes, not 

forming cincinni, bracteoles ovate to lanceolate, often red-tinged and sometimes paleaceous, and often forming a 

small involucre around cymules, or bracts narrower, calyx lobes subequal or with posterior lobe broader ............. 

.................................................................................................................................................................... Cantinoa 

18*. Inflorescence an elongate, spiciform thyrse or often a diffuse leafy panicle of pedunculate, often cincinnate cymes 

or shortly pedunculate, few-flowered cymes in axils of foliose bracts, or compact, long-pedunculate, shortly 

cincinnate cymes forming a globose head. Bracteoles inconspicuous, never forming an involucre Mesosphaerum 

† Note on the capitulum: In Hyptidinae, the flowers are arranged in a diverse array of forms. Especially in Hyptis sensu 

stricto, the flowers form a spherical or hemispherical capitulum. The flowers are sessile or almost so (pedicel less than 

0.5 mm) and borne on a slightly swollen receptacle and surrounded by an involucre of linear to ovate bracteoles, which 

are not associated with the individual flowers. Some other groups, such as in the genus Cantinoa, the flowers are often in 

congested cymes or capituliform heads, usually fewer-flowered and with a numerical relationship between flowers and 

bracteoles, but the heads are not spherical or hemispherical in form. See other note on capitulum structure, under 

Cyanocephalus. 



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

typifications 

Asterohyptis Epling (1932: 17). Type:—Asterohyptis stellulata (Benth.) Epling (1932: 17) ≡ 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 2lipped 

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. ≡ 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) ≡ 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. Hyptis 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. 


HARLEY & PASTORE

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. ≡ Hyptis althaeifolia (as 

althaeaefolia) Pohl ex Bentham (1833: 115) ≡ 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) ≡ 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. ≡ Hyptis carpinifolia Bentham (1833: 

115) ≡ 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) ≡ 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. ≡ 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. ≡ Hyptis dubia Pohl ex Bentham (1833: 

122) ≡ 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) ≡ 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. ≡ Hyptis duplicatodentata Bentham 

(1833: 114) ≡ Hyptis vestita Benth. var. duplicatodentata (Pohl ex Benth.) Schmidt (1858: 128) ≡ 

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. ≡ 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. ≡ 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. ≡ 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. ≡ 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. ≡ 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. ≡ 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. ≡ 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. ≡ Nepeta mutabilis Richard (1792: 110) ≡ 

Mesosphaerum mutabile (Rich.) Kuntze (1891: 525) ≡ 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!). 


HARLEY & PASTORE

= 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. ≡ 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. ≡ 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. ≡ Hyptis plectranthoides Bentham 

(1833: 122) ≡ 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) ≡ 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. ≡ 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. ≡ Hyptis racemulosa Mart. ex 

Bentham (1833: 126) ≡ 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. ≡ Hyptis rubicunda Pohl ex 

Bentham (1833: 118) ≡ 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. ≡ 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. ≡ 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) ≡ 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. ≡ Hyptis sylvularum A.St.- 

Hil. ex Bentham (1833: 119), pro spec. ≡ 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. ≡ 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. ≡ 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) ≡ 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 


HARLEY & PASTORE

ear 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. 

Key to sections of Condea 

Flowers solitary or in fascicles, in the axils of reduced or leafy bracts, trichomes simple.............................. Sect. Condea 

Flowers in subumbellate, pedunculate cymes, or if cymes ±sessile, then trichomes dendroid Sect. Laniflorae 

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 


HARLEY & PASTORE

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!). 


HARLEY & PASTORE

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. 


HARLEY & PASTORE

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!). 


HARLEY & PASTORE

= 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 2−18flowered 

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. 


HARLEY & PASTORE

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. 


HARLEY & PASTORE

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!). 


HARLEY & PASTORE

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!). 


HARLEY & PASTORE

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). 

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!). 


HARLEY & PASTORE

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. 


= 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!). 


HARLEY & PASTORE

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. 


= 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!). 


HARLEY & PASTORE

= 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 2n = 28, differing from numbers found in 

Eriope (2n = 20) and Hypenia (2n = 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. 

Sections unplaced to genus 

Hyptis sect. Rhytidea Epling (1933: 80), with two species, H. rhytidea Bentham (1839: 21) and H. 

pseudolantana Epling (1941: 553), may possibly belong in Condea Adans. Hyptis rhytidea was shown to be 

sister to the Condea clade (Pastore et al. 2011). Further studies are needed to explore this relationship. 

Hyptis sect. Hilaria Epling (1936b: 280). Type:—Hyptis lobata A.St.-Hil. ex Bentham (1833: 97). A 

monotypic section, which is only known from the type: BRAZIL. São Paulo: São José dos Campos? [São 

Jozé], Saint-Hilaire 664 (holotype P!; isotype F!). This species may well be related to Hyptis lagenaria A.St.- 

Hil. ex Bentham (1833: 98), in Hyptis sect. Cyrta, but further material is needed to establish its correct 

position. 

Acknowledgements 

The authors would like to thank the Programa de Biodiversidade do Semi-árido (PPBIO), Instituto do Milênio 

do Semi-árido (IMSEAR) of the Brazilian Ministério de Ciência e Tecnologia, and the Conselho Nacional de 

Desenvolvimento Científico e Tecnológico, for financial support for fieldwork, the Brazilian government 

(CAPES, PDEE) for financial support for the second author’s Doctoral programme and also the staff of the 

Departamento de Ciências Biológicas of the Universidade Estadual de Feira de Santana (UEFS), Brazil and 

the Keeper and staff of the Herbarium and Library of the Royal Botanic Gardens, Kew, UK for providing 

facilities and much assistance. We would also like to thank the staff of the many herbaria, cited in the text 

above, and visited mostly by the senior author over many years, for permission to use their facilities, their 

welcome and assistance. Thanks are especially due, in RBG Kew, to Alan Paton, Gemma Bramley, Lesley 

Walsingham for constant help or advice, and to the staff of the IPNI office, especially Irina Belyaeva, and also 

to Dick Brummitt, for guidance over nomenclatural matters. We particularly wish to acknowledge the great 

help given to us by John McNeill over Condea urbanii, which caused us quite a few headaches. We also wish 

to thank Alan Paton, Henrique Moreira, Domingos Cardoso and Sergio Bordignon for permission to use some 


HARLEY & PASTORE

of their images of Hyptidinae. Those not acknowledged were taken by the authors. The study visit by the 

senior author to the University Herbarium of Copenhagen helped resolve a number of problems. Support was 

made available by the European Community–Research Infrastructure Action under the FP6 Structuring the 

European Research Area Programme (grant DK-TAF-1865). We would like to thank Hajo Esser, of the 

Botanische Staatssammlung, Munich, for editorial guidance and valuable advice, on Roxburgh’s Clinopodium 

repens and many other matters. Also we should like to thank the two reviewers who, with their constructive 

criticism, ensured a much better final product, and finally we thank especially Ana Maria Giulietti for 

constant advice, encouragement and support at all stages of the preparation of this paper. 

References 

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HARLEY & PASTORE

APPENDIX 1. Register of earlier names of Hyptidinae with changes accepted in this paper. 

This is a list of species names recognized by Epling (1949) with the addition of names of more recent taxa published subsequently, 

followed by the genus to which they are assigned in this paper and in the last column a reference to the work in which this change was 

published, if not in the present work. 

FORMER 

GENUS SPECIES AUTHOR ACCEPTED GENUS 

Eriope alpestris Mart. ex Benth. Eriope 

Eriope anamariae Harley Eriope 

Eriope angustifolia Epling Eriope 

Eriope arenaria Harley Eriope 

Eriope complicata Mart. ex Benth. Eriope 

Eriope confusa Harley Eriope 

Eriope crassifolia Mart. ex Benth. Eriope 

Eriope crassipes Benth. Eriope 

Eriope exaltata Harley Eriope 

Eriope filifolia Benth. Eriope 

Eriope foetida Pohl ex Benth. Eriope 

Eriope glandulosa (Harley) Harley Eriope 

Eriope hypenioides Mart ex Benth. Eriope 

Eriope luetzelburgii Harley Eriope 

Eriope macrostachya Mart ex Benth. Eriope 


NOTES OF 

CHANGE 

Eriope micrantha Benth. in DC. Hypenia Harley 1988 

Eriope monticola Harley Eriope 

Eriope obovata Epling Eriope 

Eriope parvifolia Mart. ex Benth. Eriope 

Eriope polyphylla Mart. ex Benth. Eriope 

Eriope simplex (A.St.-Hil. ex Benth.) Harley Hypenia see Hyptis simplex 

Eriope sincorana Harley Eriope 

Eriope stricta Benth. Eriopidion Harley 1976 

Eriope tumidicaulis Harley Eriope 

Eriope velutina Epling Eriope 

Eriope xavantium Harley Eriope 

Hyptis actinocephala Griseb. Hyptis 

Hyptis adamantium A.St.-Hil. ex Benth. Hyptis 

Hyptis adpressa A.St.-Hil. ex Benth. Cyanocephalus 

Hyptis alata (Raf.) Shinners Hyptis 

Hyptis albicoma Epling Cyanocephalus 

Hyptis albida Kunth Condea sect. Laniflorae 

Hyptis alpestris A. St.-Hil. ex Benth. Hyptis 

Hyptis althaeifolia Pohl ex Benth. Cantinoa 

Hyptis alutacea Pohl ex Benth. Hyptis 

Hyptis amaurocaulos Briq. Hyptis 

Hyptis americana (Poir.) Briq. Condea sect. Condea 

Hyptis americana (Aubl.) Urb. Cantinoa 

= Cyanocephalus 

incanus 

...... continued 


APPENDIX 1 (continued) 

FORMER 

NOTES OF 

GENUS SPECIES AUTHOR ACCEPTED GENUS CHANGE 

Hyptis amethystoides Benth. Hyptidendron Harley 1988 

Hyptis ammotropha Wright ex Griseb. Hyptis 

Hyptis ampelophylla Epling Gymneia 

Hyptis angulosa Schott ex Benth. Hyptis 

Hyptis angustifolia Pohl ex Benth. Hyptis 

Hyptis anitae Epling & Játiva Condea sect. Laniflorae 

Hyptis apertiflora Epling Cyanocephalus 

Hyptis arborea Benth. Hyptidendron Harley 1988 

Hyptis arbuscula Epling Hyptidendron Harley 1988 

Hyptis arenaria Benth. in DC. Hyptis 

Hyptis argentea Epling & Mathias Hyptis 

Hyptis argutifolia Epling Mesosphaerum 

Hyptis argyrophylla Harley Oocephalus 

Hyptis aristulata Epling Hypenia Harley 1988 

Hyptis armillata Epling Hyptis 

Hyptis asperifolia Standley Mesosphaerum 

Hyptis asperrima (Spreng.) Epling Hyptidendron Harley 1988 

Hyptis asteroides A.St.-Hil. ex Benth. Hyptis 

Hyptis atrorubens Poit. Hyptis 

Hyptis australis Epling Hyptis 

Hyptis bahiensis Harley Hyptis 

Hyptis balansae Briq. Hyptis 

Hyptis blanchetii Benth. Eriope Harley 1988 

Hyptis bombycina Epling Cyanocephalus 

Hyptis brachiata Briq. Hyptis 

Hyptis brachypoda Epling Hyptis 

Hyptis brachystachys Pohl ex Benth. Hypenia Harley 1988 

Hyptis brevipes Poit. Hyptis 

Hyptis caduca Epling Hyptis 

Hyptis caespitosa A.St.-Hil. ex Benth. Hyptis 

Hyptis calida Mart. ex Benth. Leptohyptis 

Hyptis calycina Pohl ex Benth. Hypenia Harley 1988 

Hyptis cana Pohl ex Benth. Hyptidendron Harley 1988 

Hyptis capitata Jacq. Hyptis 

Hyptis caprariifolia Pohl ex Benth. Cyanocephalus 

Hyptis cardiophylla Pohl ex Benth. Cyanocephalus 

Hyptis carpinifolia Benth. Cantinoa 

Hyptis carvalhoi Harley Medusantha 

Hyptis caudata Epling & Játiva Hyptidendron Harley 1988 

Hyptis chacapoyensis Briq. Mesosphaerum 

Hyptis chyliantha Urb. & Ekman Condea sect. Condea 

Hyptis claussenii Benth. Hyptidendron Harley 1988 



HARLEY & PASTORE


FORMER 

NOTES OF 


Hyptis colligata Epling & Jativa Hyptis 

Hyptis collina Brandegee Mesosphaerum 

Hyptis colombiana Epling Cantinoa 

Hyptis colubrimontis Epling & Játiva Hyptis 

Hyptis complicata A.St.-Hil. ex Benth. Hyptis 

Hyptis concinna Benth. in DC. Hypenia Harley 1988 

Hyptis conferta Pohl ex Benth. Hyptis 

Hyptis conspersa Benth. Hyptidendron Harley 1988 

Hyptis coriacea Benth. in DC. Cyanocephalus 

Hyptis corymbosa Benth. in DC. Hyptis 

Hyptis crassifolia Mart. ex Benth. Oocephalus 

Hyptis crassipes Epling Hyptis 

Hyptis crenata Pohl ex Benth. Hyptis 

Hyptis cretata Epling Cyanocephalus 

Hyptis crinita Benth. Medusantha 

Hyptis crispata Pohl ex Benth. Hypenia Harley 1988 

Hyptis cruciformis Epling Hyptis 

Hyptis cubensis Urb. Condea sect. Condea 

Hyptis cuneata Pohl ex Benth. Cyanocephalus 

Hyptis cuniloides Epling Eplingiella 

Hyptis cymulosa Benth. in DC. Unplaced ? 

Hyptis decipiens M.E. Jones Condea sect. Laniflorae 

Hyptis delicatula Harley Cyanocephalus 

Hyptis densiflora Pohl ex Benth. Hypenia Harley 1988 

Hyptis desertorum Pohl ex Benth. Cyanocephalus 

Hyptis dictiocalyx Benth. Hyptidendron Harley 1988 

Hyptis dictyodea Pohl ex Benth. Hyptis 

Hyptis diffusa Epling Mesosphaerum 

Hyptis digitata Harley Cyanocephalus 

Hyptis dilatata Benth. in DC. Hyptis 

Hyptis ditassoides Mart. ex Benth. Hyptis 

Hyptis divaricata Pohl ex Benth. Hyptis 

Hyptis diversifolia Benth. Mesosphaerum Kuntze 1891 

Hyptis domingensis Urb. Condea sect. Condea 

Hyptis dubia Pohl ex Benth. Cantinoa 

Hyptis dumetorum Morong Hyptis 

Hyptis duplicato-dentata Pohl ex Benth. Cantinoa 

= Hypenia 

Hyptis durifolia Epling Hypenia 

sclerophylla 

Hyptis effusa S.Moore Hypenia = Hypenia micrantha 

Hyptis elegans (Briq.) Briq. Condea sect. Condea 

Hyptis elongata Benth. Martianthus 





FORMER 

NOTES OF 


Hyptis emoryi Torrey Condea sect. Laniflorae 

Hyptis eriocauloides Rich. Hyptis 

Hyptis eriocephala Benth. in DC. Mesosphaerum Kuntze 1891 

Hyptis eriophylla Pohl ex Benth. Medusantha 

Hyptis erythrostachys Epling Cantinoa 

see Condea 

Hyptis escobilla Urb., nom. illeg. Condea sect. Condea americana 

Hyptis eximia Epling Hyptidendron 

Hyptis fallax Harley Hyptis 

Hyptis fasciculata Benth. Condea sect. Condea See Condea undulata 

Hyptis fastigiata Benth. Condea sect. Condea 

Hyptis ferruginosa Pohl ex Benth. Hyptis 

Hyptis floribunda Briq. Condea sect. Condea 

Hyptis florida Benth. Hyptis 

Hyptis foliosa A.St.-Hil. ex Benth. Oocephalus 

Hyptis frondosa S. Moore Hyptis 

Hyptis fruticosa Salzm. ex Benth. Eplingiella 

Hyptis fulva Epling Hyptis 

Hyptis gardneri Briq. Hyptis 

see Oocephalus 

Hyptis glomerata Mart. ex Schrank Oocephalus 

oppositiflorus 

Hyptis glutinosa Benth. Hyptidendron Harley 1988 

Hyptis goyavensis A.St.-Hil. ex Benth. Hyptis 

Hyptis guanchezii Harley Hyptis 

Hyptis gymnocaulos Epling Mesosphaerum 

Hyptis hagei Harley Oocephalus 

Hyptis halimifolia Mart. ex Benth. Oocephalus 

Hyptis halimifolia var. pauciflora Harley Oocephalus 

= Oocephalus 

pauciflorus 

Hyptis hamatidens Epling & Jativa Hyptis 

Hyptis hassleri Briq. Hyptis 

Hyptis havanensis Britton ex Epling Hyptis 

Hyptis heterodon Epling Cantinoa 

Hyptis heterophylla Benth. in DC. Hyptis 

Hyptis hilarii Benth. Hyptis 

Hyptis hirsuta Kunth Hyptis 

Hyptis hispida Benth. in DC. Hyptis 

Hyptis homalophylla Pohl ex Benth. Hyptis 

Hyptis huberi Harley Hyptis 

Hyptis humilis Benth. in DC. Hyptis 

Hyptis hygrobia Briq. Hyptis 

Hyptis hypoleuca Benth. Eriope Harley 1976 

Hyptis imbricata Pohl ex Benth. Hyptis 



HARLEY & PASTORE


FORMER 

NOTES OF 


Hyptis imbricatiformis Harley Hyptis 

Hyptis impar Epling Cantinoa 

Hyptis incana Briq. Cyanocephalus 

Hyptis indivisa Pilg. Cantinoa 

Hyptis inelegans Epling Hypenia Harley 1988 

Hyptis inodora Schrank Hyptis 

Hyptis intermedia Epling Hyptis 

Hyptis interrupta Pohl ex Benth. Gymneia 

Hyptis involucrata Benth. Hyptis 

Hyptis iodantha Epling Condea sect. Laniflorae 

Hyptis irwinii Harley Mesosphaerum 

Hyptis jacobi Fernándo-Alonso Condea sect. Laniflorae 

Hyptis kramerioides Harley & J.F.B.Pastore Hyptis 

Hyptis lachnosphaeria Epling Mesosphaerum 

Hyptis laciniata Benth. Hyptis 

Hyptis lacunosa Pohl ex Benth. Oocephalus 

Hyptis lacustris A.St.-Hil. ex Benth. Hyptis 

Hyptis lagenaria A.St.-Hil. ex Benth. Hyptis 

Hyptis lanata Pohl ex Benth. Cyanocephalus 

Hyptis lanceolata Poir. Hyptis 

Hyptis laniflora Benth. Condea sect. Laniflorae 

Hyptis lantanifolia Poit. Hyptis 

Hyptis lanuginosa Glaz. ex Epling Hyptis 

Hyptis lappacea Benth. Hyptis 

Hyptis lappulacea Mart. ex Benth. Hyptis 

Hyptis latifolia Mart. ex Benth. Eriope Harley 1976 

Hyptis lavandulacea Pohl ex Benth. Hyptis 

Hyptis leptoclada Benth. in DC. Hyptis 

Hyptis leptostachys Epling Leptohyptis 

Hyptis leucocephala Mart. ex Benth. Martianthus 

Hyptis leucophylla Pohl ex Benth. Hyptidendron Harley 1988 

Hyptis linarioides Pohl ex Benth. Hyptis 

Hyptis lippioides Pohl ex Benth. Cyanocephalus 

Hyptis lobata A.St.Hil. ex Benth. Unplaced ? 

Hyptis longifolia Pohl ex Benth. Hyptis 

Hyptis longipes A.St.-Hil. ex Benth. Cyanocephalus 

Hyptis lorentziana O.Hoffm. Hyptis 

Hyptis loseneriana Pilg. Hyptis? 

Hyptis lucida Pohl ex Benth. Hyptis 

Hyptis lutescens Pohl ex Benth. Hyptis 

Hyptis luticola Epling Hyptis 


= Cyanocephalus 

rugosus 




FORMER 

NOTES OF 


Hyptis lythroides Pohl ex Benth. Oocephalus 

Hyptis machrisae Epling Eriope Harley 1988 

Hyptis macrantha A.St.-Hil. ex Benth. Hypenia Harley 1988 

Hyptis macrocephala M.Martens & Galeotti Hyptis 

Hyptis macrosiphon Briq. Hypenia Harley 1988 

Hyptis macrostachys Benth. in DC. Leptohyptis 

Hyptis macrotera Briq. Cantinoa 

Hyptis malacophylla Benth. in DC. Gymneia 

Hyptis mariannarum Briq. Hyptis 

Hyptis marifolia Benth. Hypenia Harley 1988 

Hyptis marrubifolia Epling & Mathias Mesosphaerum 

Hyptis marrubioides Epling Hyptis 

Hyptis martiusii Benth. Medusantha 

Hyptis maya Harley Hyptis 

Hyptis melissoides Kunth Mesosphaerum Kuntze, 1891 

Hyptis microphylla Pohl ex Benth. Hyptis 

Hyptis microsphaera Epling Hyptis 

Hyptis minutifolia Griseb. Hyptis 

Hyptis mixta Epling Condea sect. Condea 

Hyptis mociniana Benth. Asterohyptis Epling 1932 

Hyptis mollis Pohl ex Benth. Hyptis 

Hyptis mollissima Benth. Medusantha 

Hyptis monticola Mart. ex Benth. Hyptis 

Hyptis muelleri Briq. Hyptis 

Hyptis multibracteata Benth. Hyptis ? 

Hyptis multiflora Pohl ex Benth. Medusantha 

Hyptis multiseta Benth. in DC. Cantinoa 

Hyptis muricata Schott ex Benth. Cantinoa 

Hyptis mutabilis (Rich.) Briq. Cantinoa 

Hyptis nigrescens Pohl ex Benth. Hyptis 

Hyptis nitidula Benth. in DC. Cyanocephalus 

Hyptis nivea Epling Oocephalus 

Hyptis nubicola Harley Oocephalus 

Hyptis nudicaulis Benth. Hyptis 

Hyptis oblongifolia Benth. in DC. Mesosphaerum Kuntze 1891 

Hyptis obtecta Benth. in DC. Hyptis 

Hyptis obtusata Benth. Mesosphaerum Kuntze,1891 

Hyptis obtusiflora Presl ex Benth. Hyptis 

Hyptis odorata Benth. Hyptis 

Hyptis orbiculata Pohl ex Benth. Hyptis 

Hyptis origanoides Pohl ex Benth. Hyptis 

Hyptis ovalifolia Benth. Gymneia = Gymneia interrupta 



HARLEY & PASTORE


FORMER 

NOTES OF 


Hyptis ovata Pohl ex Benth. Hyptis 

Hyptis pachyarthra Briq. Hyptis 

Hyptis pachycephala Epling Hyptis 

Hyptis pachyphylla Epling Hyptis 

Hyptis paludosa A.St.-Hil. ex Benth. Hyptis 

Hyptis paniculata Benth. Hypenia Harley 1988 

Hyptis paradisi Harley Hypenia Harley 1988 

Hyptis parkeri Benth. Hyptis 

Hyptis passerina Mart. ex Benth. Hyptis 

Hyptis paupercula Epling Hyptis 

Hyptis pectinata (L.) Poit. Mesosphaerum Kuntze 1891 

Hyptis pedalipes Griseb. Cyanocephalus 

Hyptis peduncularis Benth. Cyanocephalus 

Hyptis penaeoides Taub. Hyptis 

Hyptis perbullata Fernándo-Alonso Mesosphaerum 

Hyptis personata Epling Hyptis 

Hyptis petiolaris Pohl ex Benth. Hyptis 

Hyptis petraea A.St.-Hil. ex Benth. Oocephalus 

Hyptis pilosa Benth. Mesosphaerum 

Hyptis pinetorum Epling Cantinoa 

Hyptis pinheiroi Harley Leptohyptis 

Hyptis piranii Harley Oocephalus 

Hyptis platanifolia Mart. ex Benth. Gymneia 

Hyptis plectranthoides Benth. Cantinoa 

Hyptis plumosa Benth. in DC. Medusantha 

Hyptis poliodes Briq. Cyanocephalus 

Hyptis propinqua Epling Cantinoa 

Hyptis proteoides A.St.-Hil. ex Benth. Hyptis 

Hyptis pruinosa Pohl ex Benth. Hypenia Harley 1988 

Hyptis pseudoglauca Epling Mesosphaerum 

Hyptis pseudolantana Epling Condea ? Unplaced 

Hyptis pseudosinuata Epling Hyptis 

Hyptis pulchella Briq. Hyptis 

Hyptis pulegioides Pohl ex Benth. Hyptis 

Hyptis purdiaei Benth. in DC. Mesosphaerum Kuntze 1891 

Hyptis pycnocephala Benth. in DC. Hyptis 

Hyptis pyriformis Epling & Játiva Hyptis 

Hyptis racemulosa Mart. ex Benth. Cantinoa 

Hyptis ramosa Pohl ex Benth. Hyptis 

Hyptis recurvata Poit. Hyptis 

Hyptis remota Pohl ex Benth. Hyptis 

Hyptis reticulata Mart. ex Benth. Hypenia Harley 1988 





FORMER 

NOTES OF 


Hyptis rhabdocalyx Mart. ex Benth. Hyptidendron Harley 1989 

Hyptis rhomboidea M Martens & Galeotti Hyptis 

Hyptis rhypidiophylla Briq. Hyptis 

Hyptis rhytidea Benth. Condea ? Unplaced 

Hyptis riparia Harley Hyptis 

Hyptis rivularis Britton Condea sect. Condea 

Hyptis rondonica Harley Hyptidendron Harley 1988 

Hyptis rondonii Epling Hyptis 

Hyptis rotundifolia Benth. Hyptis 

Hyptis rubicunda Pohl ex Benth. Cantinoa 

Hyptis rubiginosa Benth. Hyptis 

Hyptis rugosa Benth. Cyanocephalus 

Hyptis salicina J.A.Schmidt Hyptis 

Hyptis salviifolia Pohl ex Benth. Eriope Harley 1976 

Hyptis salzmannii Benth. Hypenia Harley 1988 

Hyptis sancti-gabrielii Harley Martianthus 

Hyptis savannarum Briq. Hyptis 

Hyptis saxatilis A.St.-Hil. ex Benth. Hyptis 

Hyptis scandens Epling Condea sect. Condea 

Hyptis sclerophylla Epling Hypenia 

Hyptis scoparioides Urb. Condea sect. Condea 

Hyptis seemannii (Gray) Epling Asterohyptis Epling 1932 

Hyptis selaginifolia Mart. ex Benth. Cyanocephalus 

Hyptis septentrionalis Epling Mesosphaerum 

Hyptis sericea Benth. Hyptis? 

Hyptis shaferi Britton Hyptis 

Hyptis sidifolia (L’Hérit.) Briq. Mesosphaerum 

Hyptis silvestris Epling Mesosphaerum 

Hyptis silvinae Harley Oocephalus 

Hyptis similis Epling Cantinoa 

see also Eriope 

Hyptis simplex A.St.-Hil. ex Benth. Hypenia 

simplex 

Hyptis simulans Epling Medusantha 

Hyptis sinuata Pohl ex Benth. Hyptis 

Hyptis siphonantha Harley Leptohyptis 

= Cantinoa 

Hyptis spicigera Lam. Cantinoa 

americana 

Hyptis stachydifolia Epling Martianthus 

Hyptis stellulata Benth. Asterohyptis Epling 1932 

Hyptis stricta Benth. Cantinoa 

Hyptis suaveolens (L.) Poit. Mesosphaerum Kuntze 1891 

Hyptis subrosea Harley Hypenia Harley 1988 



HARLEY & PASTORE


FORMER 

NOTES OF 


Hyptis subrotunda Pohl ex Benth. Cantinoa 

Hyptis subtilis Epling Condea sect. Laniflorae 

Hyptis subviolacea Briq. Hyptis 

Hyptis tacianae Harley Cyanocephalus 

Hyptis tafallae Benth. Condea sect. Laniflorae 

Hyptis tagetifolia Harley Cyanocephalus 

Hyptis tenuifolia Epling Cyanocephalus 

Hyptis tephrodes A. Gray Condea sect. Laniflorae 

Hyptis tetracephala Bordignon Hyptis 

Hyptis tetragona Pohl ex Benth. Hyptis? 

Hyptis thyrsiflora Epling Condea sect. Condea 

Hyptis tomentosa Poit. Condea sect. Laniflorae 

Hyptis tricephala A.St.-Hil. ex Benth. Hyptis 

Hyptis trichopes (Epling) Harley Condea sect. Condea 

Hyptis tripartita Briq. Cyanocephalus 

Hyptis tumidicalyx Epling & Játiva Hyptis 

Hyptis turnerifolia Mart. ex Benth. Hyptis 

Hyptis uliginosa A.St.-Hil. ex Benth. Hyptis 

Hyptis uncinata Benth. Hyptis 

Hyptis unilateralis Epling Hyptidendron Harley 1988 

Hyptis urticoides Kunth Mesosphaerum Kuntze 1891 

Hyptis vauthieri Briq. Hyptidendron Harley 1988 

Hyptis velutina Pohl ex Benth. Hyptis 

Hyptis vepretorum Mart. ex Benth. Hyptidendron Harley 1988 

Hyptis verticillata Jacq. Condea sect. Condea 

Hyptis viatica Harley Cyanocephalus 

Hyptis vilis Kunth & Bouche Hyptis 

Hyptis villicaulis Epling Cantinoa 

Hyptis villosa Pohl ex Benth. Hyptis 

Hyptis viminea Epling Hyptis 

Hyptis violacea Pohl ex Benth. Cantinoa 

Hyptis virgata Benth. Gymneia 

Hyptis vitifolia Pohl ex Benth. Physominthe see Hypenia vitifolia 

Hyptis xanthiocephala Mart. ex Benth. Hyptis 

= Hypenia 

Hypenia durifolia (Epling) Harley Hypenia 

sclerophylla 

Hypenia vitifolia (Pohl ex Benth) Harley Physominthe see Hyptis vitifolia 

Marsypianthes burchellii Epling Marsypianthes 

Marsypianthes chamaedrys (Vahl) Kuntze Marsypianthes 

Marsypianthes foliolosa Benth. in DC. Marsypianthes 

Marsypianthes hassleri Briq. Marsypianthes 

Marsypianthes montana Benth. Marsypianthes 





FORMER 

NOTES OF 


Peltodon comaroides Briq. Hyptis sect. Peltodon 

Peltodon longipes A.St.-Hil. ex Benth. Hyptis sect. Peltodon = Hyptis comaroides 

Peltodon pusillus Pohl Hyptis sect. Peltodon 

Peltodon radicans Pohl Hyptis sect. Peltodon 

Peltodon rugosus Tolmatchew Hyptis sect. Peltodon = Hyptis meridionalis 

Peltodon tomentosus Pohl Hyptis sect. Peltodon = Hyptis campestris 

Rhaphiodon echinus (Nees & Mart.) Schauer Rhaphiodon 


HARLEY & PASTORE

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. 


HARLEY & PASTORE

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. 


HARLEY & PASTORE

FIGURE 5. Phylogenetic relationships of genera of Hyptidinae 


Phytotaxa 58 © 2012 Magnolia Press • 55

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