Journal of Ethnopharmacology 118 (2008) 343353

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Journal of Ethnopharmacology
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Review

Ethnobotany, biochemistry and pharmacology of Minthostachys (Lamiaceae)

A.N. Schmidt-Lebuhn
Abteilung Panzenkologie, Institut fr Biologie/Bereich Geobotanik, Martin-Luther-Universitt Halle-Wittenberg, Am Kirchtor 1, 06108 Halle, Germany

a r t i c l e

i n f o

a b s t r a c t
Aims of the study: The South American mint genus Minthostachys is of great importance in the Andes as a medicinal, aromatic, culinary and commercial essential oil plant. After decades of taxonomic confusion and virtual indeterminability of specimens, new systematic and taxonomic work has been conducted in recent years. The present paper attempts to summarize the state of knowledge about Minthostachys with a focus on ethnobotany, analyses of essential oil content and pharmacology, to identify the currently accepted species names for the plants examined in these previous studies, and to assess where additional research is needed. Materials and methods: All available studies on Minthostachys were obtained and evaluated. Herbaria were contacted to identify voucher specimens cited in the respective publications. Results and conclusions: The great majority of published studies was conducted on a single species, Argentinean Minthostachys verticillata. In contrast, the most widely distributed and well-known species (Minthostachys mollis) as well as several locally important and intensively used species (e.g., Minthostachys acutifolia) have received disproportionately little attention, and virtually nothing is known about the local endemics among the 17 species currently recognized. In many cases, however, it is difcult to relate the results to taxonomic entities due to the lack of voucher specimens. Future research efforts should especially be directed at studying the chemistry and potential for use of several common but so far neglected species of the central and northern Andes, at disentangling environmental and genetic inuences on essential oil composition, at prerequisites for cultivation, and at the pharmacological basis of the most important traditional uses. Because of the morphological complexity of the genus, future researchers are urged to deposit voucher specimens of the plants used in their studies to facilitate species identication and to make the results more comparable and reproducible. 2008 Elsevier Ireland Ltd. All rights reserved.

Article history: Received 4 March 2008 Received in revised form 22 May 2008 Accepted 28 May 2008 Available online 4 June 2008 Keywords: Labiatae Lamiaceae Minthostachys Muna Peperina

Contents 1. 2. 3. 4. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Morphology, anatomy, systematics and cytology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Essential oils and other chemical components . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Harvesting, cultivation, and the reasons underlying the variability in oil composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1. Harvesting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2. Cultivation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3. Variability in oil composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ethnobotany and practical uses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.1. Common names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.2. Uses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3. Protection of stored tubers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pharmacology and antiparasitic potential . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . State of knowledge about the individual species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 344 345 345 346 346 346 347 347 347 348 349 349 350 351 351 351

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6. 7. 8.

Tel.: +49 345 5526261; fax: +49 345 5527228. E-mail address: schmidtleb@yahoo.de. 0378-8741/$ see front matter 2008 Elsevier Ireland Ltd. All rights reserved. doi:10.1016/j.jep.2008.05.030

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1. Introduction Minthostachys (Benth.) Spach (Lamiaceae, Nepetoideae, Mentheae) is a genus of 17 species of scandent aromatic shrubs restricted to Andean South America (Schmidt-Lebuhn, 2008b). It is of great ethnobotanical, pharmacological and commercial interest because of the essential oils found in the plants. It does not only nd use, as a condiment or tea, in the traditional cuisine of the Andes, but is one of the most important plants in the folk medicine of the area. On the basis of this observation, it receives growing attention from modern pharmacology and medicine, as plant decoctions and extracted essential oils are tested for pharmacological effects. In Argentina and Peru, essential oil is extracted on a commercial scale. Unfortunately, the genus is taxonomically complicated and exhibits a high morphological variability that has, in the past, severely hampered the understanding of its diversity. The most comprehensive taxonomic treatment of the past recognized 12 species (Epling, 1936), but the same researcher later changed his opinion and suggested treating Minthostachys as monotypic (Epling and Jtiva, 1963). Consequently, the same plants were sometimes, with varying degrees of success, determined to species according to the rst approach, sometimes universally labeled as Minthostachys mollis (HBK) Griseb., and sometimes dismissed as undeterminable, leading to much confusion and lack of comparability in the growing number of ethnobotanical, biochemical, and pharmacological studies. In contrast to Epling and Jtiva (1963) assumptions, several morphologically dened and ecologically distinct species of Minthostachys formed clusters in analyses based on Amplied Frag-

ment Polymorphism data, indicating that the recognition of various species is feasible (Schmidt-Lebuhn, 2007). On the basis of these results, extensive eld work, and the morphological examination of ca. 1000 herbarium specimens, 17 species are now recognized (Schmidt-Lebuhn, 2008b; Table 1), following a similar species concept as that of Epling (1936). Of these, seven are usually morphologically very distinct local endemics restricted to only one Andean valley or a similarly small area. The best known and most notable species are morphologically variable Minthostachys mollis (Fig. 1b) with three varieties occurring from Venezuela to Bolivia (and including the formerly distinct species Minthostachys mandoniana (Briq.) Epling and Minthostachys tomentosa (Benth.) Epling), southern Peruvian Minthostachys acris Schmidt-Leb. (Fig. 1c), and Minthostachys verticillata (Griseb.) Epling of Argentina (Fig. 1a). The present paper is intended to summarize ethnobotanical, biochemical, pharmacological, and other literature on the genus as a whole, and thus to provide a concise source of information to researchers of Minthostachys, especially those working on a local scale. In addition, an attempt is made to relate the results of the individual studies to taxonomic entities as currently understood. Reliable identication of the study species of these publications should facilitate comparison of their results, and allow to get a clearer picture about which species and what aspects of research may have previously been neglected. In many cases, all species of Minthostachys are subsumed under the catch-all name Minthostachys mollis regardless of provenance. On the other hand, Minthostachys acris is usually treated as separate in Peruvian publications, but then erroneously called Minthostachys glabrescens (Benth.) Epling due to a mistake dating back to Epling (1936) (see Schmidt-Lebuhn, 2008b, for details).

Table 1 List of species and subspecies accepted in Minthostachys in the recent taxonomic revision (Schmidt-Lebuhn, 2008b), with synonyms and distribution Accepted name Minthostachys andina (Britt. ex Rusby) Epling Minthostachys dimorpha Schmidt-Leb. Minthostachys fusca Schmidt-Leb. Minthostachys verticillata (Griseb.) Epling Minthostachys elongata Schmidt-Leb. Minthostachys glabrescens (Benth.) Epling Minthostachys salicifolia Epling Minthostachys latifolia Schmidt-Leb. Minthostachys septentrionalis Schmidt-Leb. Minthostachys setosa (Briq.) Epling Synonyms Bystropogon andinus Britt. ex Rusby Distribution Bolivia: Sorata Peru: Machu Picchu Bolivia: Tipuani valley Northwestern and central Argentina Bolivia: Tarija Ecuador: Loja Southern Peru Andean slopes of southern Peru and western Bolivia Venezuela, Colombia Central Bolivia

Xenopoma verticillatum Griseb.; Bystropogon kuntzeanus Briq.; Minthostachys verticillata var. eupatorioides Epling Bystropogon glabrescens Benth.

Bystropogon setosus Briq.; Bystropogon setosus var. menthiodorus Kuntze; Bystropogon setosus var. citronella Kuntze Bystropogon spicatus Benth. Minthostachys glabrescens auct., non (Benth.) Epling quoad typum Bystropogon ovatus Briq.; Bystropogon andinus var. hypoleucus Briq. Bystropogon mollis HBK; Mentha mollis (HBK) Benth.; Bystropogon canus Benth.; Bystropogon pavonianus Benth.; Bystropogon tomentosus Benth.; Minthostachys tomentosa (Benth.) Epling Bystropogon mollis HBK; Mentha mollis (HBK) Benth.; Bystropogon canus Benth.; Bystropogon pavonianus Benth.; Bystropogon tomentosus Benth.; Minthostachys tomentosa (Benth.) Epling Bystropogon mandonianus Briq.; Minthostachys mandoniana (Briq.) Epling

Minthostachys diffusa Epling Minthostachys spicata (Benth.) Epling Minthostachys acris Schmidt-Leb. Minthostachys acutifolia Epling Minthostachys ovata (Briq.) Epling Minthostachys rubra Schmidt-Leb. Minthostachys mollis (HBK) Griseb.

Bolivia: Yungas Coastal cordillera of Peru and southernmost Ecuador Peru: southern highlands Bolivia: La Paz Central Bolivia Ecuador: Carchi Venezuela, Colombia, Ecuador, Peru, Bolivia

Minthostachys mollis var. mollis

Venezuela, Colombia, Ecuador, Peru

Minthostachys mollis var. hybrida Schmidt-Leb. Minthostachys mollis var. mandoniana (Briq.) Schmidt-Leb. Sequence of species and varieties follows that of the revision.

Venezuela, Colombia Southern Peru, western Bolivia

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Fig. 1. Important representatives of the genus Minthostachys. (A) Minthostachys verticillata is, to current knowledge, the only Argentinean species and endemic to that country. Despite this, the peperina is the most thoroughly studied species. (B) Conspiciously hairy Minthostachys mollis has the widest distribution of all species. (C) Minthostachys acris (previously sometimes called Minthostachys glabrescens) is the most abundant species in the southern Peruvian highlands and is intensively used by the local population. (D) Very little, if anything, is known about the essential oils and potential uses of Minthostachys acutifolia, the predominant species near La Paz, Bolivia.

2. Morphology, anatomy, systematics and cytology Apart from what has been touched in the taxonomic literature (Epling, 1936; Schmidt-Lebuhn, 2008b), only very little work has been done on morphology and anatomy. Bonzani and Ariza Espinar (1993) presented an examination of leaf and stem anatomy of Minthostachys verticillata. Arevalo Gonzales (1995) examined morphological variability and ecological preferences of three populations of Minthostachys in Colombia. It appears that the differences between populations found in her study with regard to leaf shape, indument and ecological niche correspond to Minthostachys mollis and newly described Minthostachys septentrionalis Schmidt-Leb. (Schmidt-Lebuhn, 2008b). While Minthostachys was traditionally held to be closely related to Macaronesian Bystropogon LHr. and North American Pycnanthemum Michx. (e.g., Epling, 1936), recent studies based on pollen morphology (Wagstaff, 1992) and molecular data (SchmidtLebuhn, 2007, 2008a; C. Bruchler, pers. comm.) point to a close relationship with South American Mentheae, particularly the former genera Xenopoma Willd. and Gardoquia HBK, both currently treated as part of a broadly circumscribed Clinopodium L. (Harley and Granda, 2000). To explain its seemingly chaotic variability, Epling and Jtiva (1963) hypothesized that Minthostachys could represent a polyploid

complex. At present, only a very limited number of chromosome counts is available for Minthostachys. Harley and Heywood (1992) presented four counts, all with 2n = 46. Vouchers at K for three of the four counts could be identied as all belonging to Minthostachys mollis var. mollis from Ecuador. A recent survey of nuclear DNA content in 10 of the 17 species of the genus (Schmidt-Lebuhn et al., 2008) did not reveal differences of a degree that would indicate polyploidy, suggesting that this phenomenon did at least not play a major role in the diversication of Minthostachys. On the other hand, Ordonez et al. (2002) rather surprisingly reported 2n = 24 for one population and 2n = 42 for three populations of Minthostachys verticillata. Considering the sharp contrast between these results and the cytological uniformity observed in the other studies cited here as well as the fact that presumed close relatives of Minthostachys also show chromosome numbers of 2n = 46 (48) (Harley and Heywood, 1992), it would be rewarding to study more accessions to see whether their observations can be reproduced. 3. Essential oils and other chemical components As in many Lamiaceae, the essential oils are the chemical components of Minthostachys plants that are of commercial and pharmaceutical interest. It is therefore not surprising that numerous analyses of the oil composition have been conducted. Their

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great majority, however, examined a single species: Argentinean Minthostachys verticillata. Scientic interest in this species, colloquially called peperina, was sparked as early as 1913 when the exploitation and use of the species were described and protocols for the distillation of the oil were published (Doering, 1913). One of the rst analyses of the essential oil lists menthol as a main component (Preioni, 1932), the same substance found copiously in the oil of the commercial mints of Mentha L. subgenus Mentha. Later researchers (Fester et al., 1947; Fester and Martinuzzi, 1949, 1952; Fester, 1958) soon corrected this error and, like most subsequent studies, showed menthone and pulegone to be the main components of peperina oil, with menthone in some cases accounting for ca. 5060% of the mixture. Improved methods of analysis subsequently made possible the identication of minor components, notably isomenthone, limonene (Fester and Martinuzzi, 1950; Fester et al., 1950, 1956), menthol (Fester et al., 1948), and -pinene (Fester et al., 1960). More recent studies of the same species have drawn a more complicated picture. On the one hand, ever more substances have been identied to be present in small amounts, among them carvone, pinene, piperitenone, sabinene, myrcene, and (E)- -ocimene (de Feo et al., 1998). On the other hand, oil composition appears to be subject to high intraspecic variability (see Section 4.3 for more detail), with some samples very much pulegone-dominated (Manfredini and Montes, 1962; Rufnengo et al., 2005; Primo et al., 2001; Gonzlez Pereyra et al., 2005), and others even having carvone, thymol or carvacrol as the principal component (Lizzi and Retamar, 1975; Zyglado et al., 1996). Arguing that menthol was economically more interesting than pulegone and menthone, protocols have been proposed to convert the oil of Minthostachys verticillata into menthol using hydrogenation methods (Retamar and Mazzola, 1963; Retamar et al., 1996). Much less information is available for the other 16 species of the genus Minthostachys, if at all. Minthostachys mollis var. mandoniana (Briq.) Schmidt-Leb. from Bolivia was examined once (Fournet et al., 1996), and it also contained high amounts of pulegone (25.5%) and menthone (24.9%). Essential oil of a Minthostachys from Venezuela, either Minthostachys mollis var. mollis or Minthostachys septentrionalis (a voucher was deposited in MERF but not seen by myself), was very much dominated by pulegone (>75%) (Rojas and Usubillaga, 1995). On the other hand, the predominant compounds of the essential oil of a specimen of Minthostachys mollis var. mollis from Ecuador were, in one case, neomenthol (29.3%), menthol (20.6%), menthone (24.0%) and piperitone (9.0%) (Alkire et al., 1994). The oil of another Ecuadorian Minthostachys, tentatively identied as probably an aberrant individual of Minthostachys mollis, contained menthone (16%), carvacryl acetate (10%), pulegone (10%) and carvacrol (9%) (Malagn et al., 2003). Other studies cannot as reliably be related to species names due to the lack of voucher specimens. Nevertheless, pulegone and menthone were the principal components of the oil in unidentied plants from various areas outside Argentina, e.g. Jauja in Peru (33.1 and 48.2%, respectively; Aliaga and Feldheim, 1985), Lima in central Peru (47.4% and 25.3%; Minthostachys mollis or Minthostachys spicata (Benth.) Epling; Senatore, 1998), southern Peru (45 and 18%; probably Minthostachys acris; Svendsen et al., 1987) and Bolivia (ca. 5861% and 3%; Munoz et al., 1990; ca. 3665% and 1932%; Munoz-Collazaros et al., 1993). Exceptionally, Senatore (1995) and Senatore and de Feo (1995) analyzed the essential oil of what were most probably specimens of Minthostachys acris (inferred from provenance; vouchers were cited, but attempts to nd them or contact the respective herbarium failed) and found a very complicated composition without one or two dominant components. The most abundant components were

menthone (14.5%), 2-hydroxy-p-menth-1-en-3-one (=diosphenol, 12.5%), 3,4,5-trimethoxytoluene (12.4%), and pulegone (9.7%). Apart from the essential oils, little is known about the chemical components of Minthostachys plants, but Minthostachys mollis var. mollis from Colombia has been screened for mucilage and lectin, a sugar binding protein (Fernandez-Alonso et al., 2003). Its seeds show no presence of mucilage, and lectin activity is low unless induced by treatment with the Pectinex enzyme. 4. Harvesting, cultivation, and the reasons underlying the variability in oil composition For an efcient and sustainable exploitation of Minthostachys as a source of essential oils, it is necessary to understand how and to what degree it can be harvested in the wild, how it can be cultivated, and how a maximum yield of the desired chemical components can be obtained. Again, the majority of studies about these aspects has been conducted on Argentinean Minthostachys verticillata. 4.1. Harvesting Bustos and Bonino (2005) examined the socio-economic importance of the exploitation of Minthostachys verticillata in the Crdoba province of Argentina. They found that the collection of Minthostachys was, on average, the most important source of income for the collectors, and claimed that the demand is covered exclusively by collection from the wild. Exploitation in the Crdoba area was described as being effected by independent collectors without any degree of training or organization. Plants were collected on private farmland, sometimes on those of others against a percentage of the proceeds or illegally without the knowledge of the landowner. Plant material was air-dried and then sold to intermediaries who further sold it to companies using it as a raw material for their products. The whole process of exploitation was described as operating outside of scal and ecological statutes set by the government. An overwhelming majority of polled collectors agreed that the plant was becoming less abundant in the previous years, while its price was increasing. It appears that collection was generally destructive (i.e., collection of juvenile plants, of plants with roots, and cutting of plants before the owering period). Similarly, Ojeda et al. (2004b) expressed concern about increasing trade in aromatic species in the Argentinean Crdoba province being based mainly on harvesting of wild plants, thus endangering natural populations of Minthostachys verticillata. 4.2. Cultivation While even an Argentinean online source for would be Minthostachys farmers is available (http://www.herbotecnia. com.ar/aut-peperina.html), relatively little research has been published about the possibility of cultivation. The most interesting study is that of Ojeda et al. (2004a) who tested the best form of cultivation by subjecting eight different Minthostachys verticillata populations to ve different cropping regimens. They found signicant differences in plant mortality, productivity and essential oil yield between the regimens, but also that some populations were better adapted to cropping than others. They recommend a single-cropping regimen with cropping in April and a careful selection of populations to be brought into cultivation. Maquera Lupaca et al. (2007) tested plants of what they called Minthostachys mollis for their productivity in cultivation under three different spacing regimens. Differences in growth height and moist and dry mass gain were found to be negligible, but plants from the middle group planted 1.25 m apart were slightly more productive. Results may have suffered from limiting growth globally

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through the cultivation of the plants under suboptimal conditions, as the authors mention drought stress occurring. No vouchers were deposited, but judging from geographic provenance (Hunuco, Peru) it is likely that the study plants really were Minthostachys mollis. Minthostachys verticillata has been successfully cultivated in vitro, and the regenerated plants showed negligible differences in essential oil composition in comparison with the paternal plants (Chebel et al., 1998). The regenerative ability of parts of plants from the high altitude valleys (28003500 m above sea level) of Cochabamba, Bolivia has also been examined under in vitro culture conditions (Castillo and Jordan, 1997). On the basis of geographic and elevational provenance, the plants most likely belong to Minthostachys ovata (Briq.) Epling, but no voucher appears to have been deposited. Minthostachys seeds germinate in 514 days and generally show a high viability (>90%; Pacheco Farfan, 1984; A.N. Schmidt-Lebuhn, pers. obs.). So far, germination does not seem to have been observed under different microclimatic conditions, nor appears anything to be known about seed viability after varying storage times and conditions. 4.3. Variability in oil composition Highly diverse factors have been shown to inuence essential oil composition in the same plant, notably stress, elevation, and season. In addition, variability between plants of the same species appears to be relatively high. Dramatic increases in the concentration of the two most abundant monoterpenes, menthone and pulegone, occurred in mechanically wounded leaves of Minthostachys verticillata after 1 day (Banchio et al., 2005a). Values returned to normal after 2 days. Banchio et al. (2005b) then tested the effect of herbivory by four different insects on essential oil composition in the same species. They found the concentration of menthone decreasing and the concentration of pulegone increasing after 24 and 48 h after most types of leaf damage. Only for leaf scraping thrips, the plants showed a different reaction, with menthone concentration remaining constant and pulegone concentration stopping its increase after 24 h. In contrast to mechanical wounding (Banchio et al., 2005a), the damage by insects also induced a systemic response in that adjacent undamaged leaves showed the same changes in essential oil composition. Valladares et al. (2002), contrarily found a decrease of pulegone content in Minthostachys verticillata following herbivory in the wild by a leaf-miner and a gall insect, and an increase in menthone content in the rst case. A singular study of seasonal variation of the essential oil of Minthostachys verticillata found a distinctive drop in aromatic oil content during the winter rest, with highest yield obtained from plants in full bloom (Bandoni et al., 2002). In addition, the relation of the two most abundant components of the oil was also subject to seasonal change, ranging from 58.3% pulegone and 24.1% menthone at the beginning of the growth period to ca. 9% pulegone and ca. 73% menthone in winter. Plants of a Venezuelan Minthostachys (either Minthostachys mollis or Minthostachys septentrionalis, voucher not seen) transplanted from an elevation of 3600 to 1600 m showed a slightly lower oil content in the new habitat (Rojas and Usubillaga, 1995). The results of a study of the essential oil composition of several populations of Minthostachys collected in the Cochabamba department of Bolivia (Munoz-Collazaros et al., 1993) may indicate an economically relevant positive correlation between elevation and the ratio of pulegone to menthone. Unfortunately, the species examined in the study cannot be identied due to the lack of voucher specimens, so that the observed gradient could also have a genetic instead of

an ecological basis, as at least three species of the genus occur in the study area. A high degree of intraspecic biochemical variability is obvious from the aromatic scent of wild plants alone. Individuals of Minthostachys verticillata from the Taf del Valle area, for example, variously have odors similar to spearmint or oregano; in a single population of Minthostachys mollis in Ecuador, adjacent plants can smell of mint, of lemon and unpleasantly musty, respectively (A.N. Schmidt-Lebuhn, pers. obs. during eld work). Analyses of aromatic oil composition in Minthostachys verticillata indicate a correspondingly high diversity, but a higher variation between populations than within (Ojeda et al., 2004b) and a higher variation between regions of Argentina than within (Zyglado et al., 1996). In a single study of this one species (Zyglado et al., 1996), menthone content varied between 3.8 and 29.6%, limonene between 1.4 and 25.0%, pulegone between 1.2 and 47.0%; thymol and carvacrol were only found in populations from one region (Catamarca), but there they were of major importance (10.531.8 and 25.334.0%, respectively). Retamar et al. (1995) found their samples from Tucumn dominated by carvone and those from Crdoba dominated by pulegone and menthone. Until more detailed studies have been conducted, it must remain unclear how much of the observed variation derives from ecological factors and how much from genetic diversity. Nevertheless, a genetic component seems to be indicated by the observed variability in single populations sampled under the same environmental conditions, and this suggests that prospective cultivars for sustainable economic exploitation would have to be carefully selected and presumably propagated vegetatively to guarantee a desired composition of oil in the harvest.

5. Ethnobotany and practical uses 5.1. Common names The genus Minthostachys is known to the local population under regionally differing names (Schmidt-Lebuhn, 2008b). The best known of these are muna in the area from central Peru to Bolivia and peperina in Argentina, where muna is more frequently used for species of Clinopodium [Xenopoma]. Others include poleo and tipo in Ecuador, chancua in northern Peru, and sometimes more generic names for aromatic Lamiaceae like oregano(te) (especially in Colombia), menta or toronjil. Ethnobotanical notes on herbarium specimens indicate that individual species of the genus rarely seem to be reliably differentiated by common names, and that supercially similar other Lamiaceae are sometimes also called muna (Schmidt-Lebuhn, 2008b). The only impression of a more discerning approach gives a report on popular taxonomy for aromatic Lamiaceae in the southern Peruvian highlands by Ugarte Ochoa et al. (1984). The surveyed locals knew four munas, the rst two of them yuraq muna/misti muna/muna blanca (white mint) and yana muna/muna negra (black mint) as subshrubs growing between 2000 and 3500 m and distinguished by their stem color, presumably related to different indument densities. Judging from these descriptions and geographic provenance, they probably represent Minthostachys mollis var. mandoniana and Minthostachys acris, respectively. Both together were called hatun muna or muna grande (large mint). The third, khunu muna/muna menuda (small mint), grew above 3500 m and was described as a lower and smaller subshrub with a very strong aromatic scent. This may also be Minthostachys acris, and the differences to muna negra may be due to ecological factors, but it is also possible that the name could refer to Clinopodium bolivianum. The fourth, pacha muna/pampa muna, was the small creeping herb scientically known as Hedeoma mandonianum Wedd., another Lamiaceae with minty scent.

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Table 2 List of uses documented for species of Minthostachys in scientic literature (roman) and as collectors notes on herbarium specimens (italics) Drink and food Liquor Guarapos (a herbal alcoholic beverage) Yerba compuesta (variant of Argentinean yerba mate) Tea, mate Candy Seasoning

Minthostachys verticillata: Bonzani and Ariza Espinar (1993), Ojeda et al. (2004a,b), Parodi in the preface to Epling (1938, p. 7) Minthostachys mollis var. hybrida: Trujillo et al. 18373 (MO) Minthostachys verticillata: Bustos and Bonino (2005) Minthostachys mollis: Macbride & Featherstone 1516 (F, MO); Minthostachys verticillata: Bocco et al. (1997) Minthostachys verticillata: Ojeda et al. (2004a,b) Minthostachys acris: Davis et al. 1476 (F, MO, USM), Hermann 465 (NY); Minthostachys dimorpha: Minthostachys Kessler (pers. comm.); Minthostachys mollis var. mollis: Andr 776 (NY), Cern 7116 (MO, QCNE), Cerrate 365 (US), Ferreyra 7272 (US), 7667 (MO), Leiva et al. 286 (F, GOET), Llatas Quiroz 1543, 2416 (both F), Macbride & Featherstone 399 (F, MO), Snchez Vega 3548 (F), Shonle 6 (MO), Smith & Buddensiek 12447 (F, HUT, MO); Minthostachys spicata: Fabris & Sgastegui 7553 (F, HUT), Leiva et al. 1154 (GOET), Sagstegui & Fabris 285 (NY), Sagstegui 12717 (HUT, MO, NY); unidentied: Ugarte Ochoa et al. (1984), Ulloa (2006) Minthostachys acris: Duran 7 (LPB); Minthostachys dimorpha: M. Kessler (pers. comm.); presumably Minthostachys mollis or Minthostachys spicata: de Feo (1992); Minthostachys mollis var. mandoniana: Carter s.n. (F); Minthostachys mollis var. mollis: Arguello 87, 133 (both QCA), Cern 1490 (AAU, MO, QCNE), Ellemann 91663 (QCA), Smith & Buddensiek 12447 (F, HUT, MO); Minthostachys ovata: Carter s.n. (F), Coca & Padilla 8 (LPB), Erquicia 34 (LPB); Minthostachys verticillata: Bandoni et al. (1972), Bonzani and Ariza Espinar (1993) Minthostachys verticillata: Martnez and Planchuelo (2003); Bonzani and Ariza Espinar (1993) Minthostachys mollis var. mollis: Milanowski & Shonle 49 (MO) presumably Minthostachys mollis or Minthostachys spicata: de Feo (1992) Minthostachys mollis: White (1982) Minthostachys acris: Infantes 6191 (B) Minthostachys mollis: White (1982) Minthostachys mollis var. mollis: Cern 7116 (MO, QCNE) Minthostachys mollis var. mollis: Cern 6892, 7150 (both MO, QCNE), Smith 12447 Minthostachys mollis var. mollis: Cern 1490 (AAU, MO, QCNE), Ellemann 66621 (QCA) Minthostachys septentrionalis: Uribe Uribe 4587 (NY) Minthostachys mollis: White (1982); Minthostachys mollis var. mollis: Cern 7116 (MO, QCNE) Minthostachys mollis var. mollis: Arguello 133 (QCA) Minthostachys verticillata: Martnez and Planchuelo (2003) Minthostachys mollis var. mollis: Milanowski & Shonle 49 (MO) Minthostachys mollis var. mollis: Camp E-2312 (NY) presumably Minthostachys mollis or Minthostachys spicata: de Feo, 1992 Minthostachys mollis var. mollis: Camp E-2312 (NY); unidentied (probably Minthostachys acris): Ormachea (1980) Minthostachys acris: Hermann 465 (NY), Infantes 6191 (B); unidentied (probably Minthostachys acris): Ormachea (1980) unidentied (probably Minthostachys acris): Ormachea (1980)

Medicinal Diarrhea, colics, atulence, and stomach pain in general

Digestive Nausea Sedative Headache Bruises Bronchitis, asthma Cold Cough Flu Misc. respiratory ailments Induction of menstruation Preparation of birth Aphrodisiac Impotence Rheumatism Biopesticide or repellent Antimycotic, antiparasitic Flea infestations in the house (by sweeping the oor with branches) Protection of stored tubers (by alternating tuber and Minthostachys leaf layers) Plant pests in the eld (by applying decoctions to the pests) Other Commercial production of aromatic oil Mummication (historical)

unidentied: Ulloa (2006) Minthostachys acris: Infantes 6191 (B)

In all cases, above-ground parts of the plants are used, preferably leaves and young stems collected before owering, sometimes whole inorescences. Herbarium acronyms follow Holmgren et al. (1990).

5.2. Uses The use of plants called muna has been recorded as early as the 16th century (Heinrich, 1992, and citations therein). Even though their taxonomic afliation must remain ambiguous (see Section 5.1), it can be assumed that the aromatic properties of the genus Minthostachys must have been readily discovered by the people of the Andes, given its commonness in areas favorable to human settlement. Ethnobotanists have documented a surprisingly wide array of traditional uses for Minthostachys plants (see Table 2). These include use as carminative, digestive, antispasmodic, and for the production of liqueur (Minthostachys verticillata; Bonzani and Ariza Espinar, 1993), for yerba compuesta, a variant of the iconic Argentinean yerba mate with Ilex paraguariensis L. as the main ingredient (Minthostachys verticillata; Bustos and Bonino, 2005), for antimycotic and antiparasitic purposes, as a vermifuge or sedative, and against diarrhea, colics, and respiratory illnesses (presumably Minthostachys mollis or Minthostachys spicata; de Feo, 1992), against ea infestations and rheumatism (Minthostachys mollis; Joyal, 1987), against bronchitis, asthma, and headache, to induce

menstruation, and as a condiment (Minthostachys mollis; White, 1982). Acha (2001) names Minthostachys as a potential biopesticide. Ugarte Ochoa et al. (1984) described the use of Minthostachys from southern Peru for seasoning, tea, medicine against colds and stomach ache, and especially for the preservation of stored potatoes. Ulloa (2006) lists uses of Minthostachys like condiment for soups, stew, and even modern Pizza, medicinal infusions, and commercial production of aromatic oil. She also presents two culinary receipts containing Minthostachys (for chupe verde and lagua de maz). Franquemont et al. (1990) mention Minthostachys acris as a condiment for soups and other dishes prepared by the Chinchero people of Peru. Ormachea (1980) recorded traditional uses of Minthostachys in the vicinity of Cuzco and Puno: the protection of stored potato and oca tubers from pests, the application of water boiled while containing Minthostachys leaves or even of fresh plant parts against aphids on crop plants, and use in the house against ea infestations. The species in question was most likely Minthostachys acris, as it is by far the most frequent in the Cuzco area and especially at the elevations mentioned by the author.

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An ethnobotanical study in Argentina characterized Minthostachys verticillata as a hot species in the dualistic view of traditional native American medicine, most probably because of the pungent aromatic odor (Martnez and Planchuelo, 2003). It was said to be used as an aphrodisiac and a digestive. In a survey, the same species surfaced as one of the nine most important of 66 medicinal plants native to the Argentinean Ro Cuarto Department (Bocco et al., 1997). Even among these nine, it was said to be the most collected. Here, the plant was said to be predominantly used for tea and mate. All available sources indicate that the most frequently documented and observed traditional uses of Minthostachys plants are related to treatments of the digestive system, followed by the treatment of various illnesses of the respiratory system (Table 2). This is not surprising considering that other plants containing the dominant essential oil components found in the genus, like menthone, pulegone, carvacrol, carvone, etc., are likewise chiey employed as digestives, carminatives, antispasmodics, and antitussives (Teuscher et al., 2004). 5.3. Protection of stored tubers Of the various uses described above and in Table 2, especially the preservation of potato tubers has been tested systematically for practicability on a larger scale. Munares et al. (1978) tested leaves, emulsion and extracts of Minthostachys for their ability to inhibit sprouting of stored potatoes and the volatile components of the plants for their antibacterial and antifungal activities. While low temperatures and high humidity in storage were more effective in reducing weight loss of the tubers, the application of Minthostachys also resulted in signicantly less bud growth. Minthostachys plants originated from Comas between Jauja and Satipo, but no vouchers were deposited, so that they cannot be determined to species. Aliaga and Feldheim (1985) obtained 1000 kg of Minthostachys plant material of unknown taxonomic afliation (no voucher) from Jauja in Peru to extract essential oil, analyzed it, and tested the practicability of using it to repress shoot formation and thus preserve tuber weight and quality in the storage of potatoes. They found that potatoes stored for 48 months in an atmosphere saturated with Minthostachys oil lost distinctively less weight than the control and did not develop shoots. One day after removal from the storage compound, the potatoes had lost the mint smell and could be consumed as if they were freshly harvested. Main ingredients of the oil were pulegone (33.1%) and menthone (48.2%), so that additional tests were conducted storing potatoes under the inuence of pure pulegone, menthone, and a pulegonementhone mixture. The results were similar, but interestingly, utilization of either pulegone or menthone alone resulted in the loss of tuber viability. The authors suggested large-scale testing of this form of potato storage as a possible alternative to what they saw as often unhealthy synthetic growth repressors and energy-consuming cooling. Hurtado et al. (1987) conducted a similar experiment with emulsions prepared from whole Minthostachys leaves and in an atmosphere saturated with the essential oils alone. The latter preparation produced the best results. Munares et al. (1978) successfully tested leaves, emulsions and oil extracts of what was probably Minthostachys mollis from Peru for their capacity to inhibit the growth of potato tubers. Raman et al. (1987) showed that dried leaves of unidentied Peruvian Minthostachys either shredded or ground to dust signicantly reduced potato tuber moth and rotting damage in potato storage. Similarly, dried plant material of Minthostachys as well as its essential oils signicantly deterred the potato tuber moth from laying eggs on stored potatoes (Guerra et al., 2007). While the study plants, all from the same locality near Cuzco, were said to

belong to Minthostachys spicata, Minthostachys glabrescens and Minthostachys mollis, they most likely represent Minthostachys acris or possibly Minthostachys mollis var. mandoniana, but no voucher specimen is available. 6. Pharmacology and antiparasitic potential In addition to the traditional uses described above, often based on direct consumption or application of fresh or dried plant material, pharmacological and pesticidal applications of the extracted oil of Minthostachys have repeatedly been explored. The simplest approach is that of dousing a parasite or pest with the oil in different concentrations, or subjecting it to an oil-saturated atmosphere. In this manner, Minthostachys oil has been tested successfully against pig lice (Caballero Osorio, 1984a), alpaca scabies (Caballero Osorio, 1984b), potato tuber worms (Chvez, 1984), and corn leaf rollers (Molleda Meneses, 1984), in the rst two cases as a viable alternative to traditionally popular but unhealthy petroleum. More sophisticated studies have explored the effect of the essential oil against diverse parasites or cell cultures under laboratory conditions, usually together with an analysis of the oil components. Most of them were conducted on Argentinean Minthostachys verticillata. The oil of this species was tested for inhibitory activity of the oil against eight bacterial strains, seven of which were affected by the oil: Bacillus subtilis Cohn, Staphylococcus aureus Rosenbach, Streptococcus faecalis Orla-Jensen, Bacillus cereus Frankland & Frankland var. micoides, Proteus mirabilis Hauser, Escherichia coli Casellani & Chalmers, and Salmonella typhi (de Feo et al., 1998). It also exhibited immunomodulating and anti-allergic properties in human cell lines (Gonzlez Pereyra et al., 2005) and antiviral activity in vitro against herpes and pseudorabies viruses and antibacterial activity especially against Gram-positive bacteria (Primo et al., 2001). For higher organisms, Minthostachys verticillata oil showed repellent properties against the mite Varroa destructor Anderson & Trueman (Rufnengo et al., 2005), though its toxicity was lower than that of other essential oils tested in the same study. Sutil et al. (2006) tested the cytotoxicity of the oil on larvae of the crustacean Artemia salina and on Vero and HEp-2 cell cultures. They found that pulegone was the most toxic component examined (LD50 0.30 mg/ml on Artemia salina), while menthone, limonene and also the essential oil as a whole were less toxic (LD50 1.12, 5.25, 2.10 mg/ml, respectively). The results obtained from cell lines and the crustacean were comparable. Oil samples in all these studies were dominated by pulegone and menthone. Cariddi et al. (2007) found the oil of Minthostachys verticillata to suppress allergic reactions of basophiles and lymphocytes from 30 allergic patients, but higher concentrations were toxic. Without examining the oil composition, other studies demonstrated decoctions of this species to have immunogenic and mitogenic activities in rabbits (Cariddi et al., 2005) and antibacterial properties against Staphylococcus aureus as well as immunomodulating properties (Maldonado et al., 2001), and the extract of the plant material to show antiviral activity in assays against the pseudorabies virus (Zanon et al., 1999). Decoctions of Minthostachys verticillata inhibit onion root growth but do not show signicant genotoxic activity (Casasnovas et al., 2006). As with the oil compositions, much less work has been done on the other species of the genus, and in several cases it remains unclear which species has been examined. Fournet et al. (1996) tested insecticidal properties of Minthostachys mollis var. mandoniana oil, also rich in pulegone and menthone, against the insects transferring Chagas disease and found toxicity against both insect species. Hurtado et al. (1987) demonstrated fungicidal and insecticidal effects of Minthostachys oil in laboratory experiments. The plants were obtained from Peru, from Comas between Jauja and

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Table 3 The publications discussed in this paper broken down according to the species they have (in some cases, most likely) examined and the type of studies that were conducted Genus-level study Systematics, taxonomy, anatomy or cytology Germination Essential oil analysis Biochemistry (other than oil) Variability of oil Exploitation and protection Ethnobotany Tuber preservation Pest control Pharmacology and antiparasitic potential Cultivation Sum 7 verticillata 2 12 8 2 2 mollis var. mollis 2 2 1 2 1 3 2 1 2 3 4 3 1 1 19 mollis var. mandoniana acris ovata unidentied Sum 11 1 19 1 10 3 7 4 4 14 3

1 1 1 1 8 1

10 1 37

Note: outside of genus-level studies, only four species of the genus are known with some certainty to have been studied at all.

Satipo at the end of the Valle de Mantaro, but apparently no vouchers were deposited. Geographically, they could belong to Minthostachys mollis or Minthostachys spicata. Ciccia et al. (2000) tested extracts of Minthostachys for insecticidal activity against larvae of Aedes aegypti L., a vector of yellow fever, and described the results as promising. The sample included in the study under the name of Minthostachys setosa (Briq.) Epling was collected either in Argentina or Peru, so that it is certainly misidentied. A voucher was cited from the Museo de Farmacobotnica, Facultad de Farmacia y Bioqumica, Universidad de Buenos Aires, but it does not seem to have actually been deposited there (A. Gurni, pers. comm.). Carvajal and Thilly (1988) studied possible mutagenic effects in human AHH1 lymphoblast cells exposed for three days to different concentrations of Minthostachys oil. They found decreasing survival rates and increasing mutagenic activity with higher oil concentrations. The plants were collected in the Peruvian Puno area, so that they most likely represented Minthostachys mollis var. mandoniana, but perhaps Minthostachys salicifolia Epling, Minthostachys latifolia Schmidt-Leb. or even Minthostachys acris. Unfortunately, no voucher is mentioned in the publication. Beltran and Cedillo (1994) demonstrated antihistaminic effect of the essential oil of either Minthostachys mollis or Minthostachys septentrionalis (no voucher available) in preparations of guinea pig intestines comparable to that of chloropheniramine but slightly weaker. 7. State of knowledge about the individual species The review of the available literature shows that much more research effort has been directed at Minthostachys verticillata than at any other species (Table 3). In addition, for some species of the genus, virtually nothing is known about oil composition, uses or the possibility of cultivation. Next to Minthostachys verticillata, the best studied species appear to be Minthostachys mollis and Minthostachys acrisnot surprisingly, as the rst is the most widely distributed and very abundant at least from Ecuador to central Peru, and the second is ubiquitous in the Cuzco area and traditionally much used. On the other hand, the two very frequent species of the Bolivian highlands, Minthostachys acutifolia Epling of La Paz (Fig. 1d) and Minthostachys ovata (Briq.) Epling of Cochabamba, Sucre and Potos, have been very much neglected, and it cannot with certainty be said if they have been studied at all. This leads to the second problem concerning our knowledge about the species. While the occurrence of other species in the region bordering Bolivia cannot be ruled out, the abundance of material from the Tucumn and Crdoba regions available for taxonomic and oristic study allows us to conclude with much certainty that Minthostachys verticillata is the only species of its genus to

be found there. Virtually all Argentinean studies as discussed here have taken place in these areas, and so we can be quite optimistic that they do, in fact, deal with Minthostachys verticillata, even if no voucher material is available for identication (Table 4). Unfortunately, the same is not true for other areas, where between two (Colombia and Venezuela) and nine (Bolivia) morphologically and ecologically divergent species can be found in the same country. Munoz et al. (1990), for example, studied the essential oils of two species they called Minthostachys andina and Hedeoma mandonianum. Both were most certainly misidentied: Minthostachys andina (Britt. ex Rusby) Epling occurs only at Sorata, so chances are great that the samples collected from seven different places (!) really represent Minthostachys acutifolia or Minthostachys ovata or perhaps both, but the true identity of the plants will remain a mystery, as no vouchers are mentioned in the publication. Hedeoma mandonianum, in reality a tiny creeping herb, is described as a shrub of 1.5 m height, so that the second examined species was most probably Clinopodium bolivianum. The study by Munoz-Collazaros et al. (1993) dealing with a possible correlation between elevation and the ratio of pulegone to menthone in the essential oils has also already been discussed critically above. It is another good example, because verication of the taxonomic afliation of the samples used in the study is once more impossible due to the lack of vouchers. The dry highlands of Cochabamba are inhabited by Minthostachys ovata, but another species, Minthostachys setosa, is found on moister Andean slopes of the same geographic area, so that the observed gradient could also have a genetic instead of an ecological basis. In addition, it cannot even be ruled out that the rarer Minthostachys mollis var. mandoniana was also sampled.

Table 4 The studies discussed in this paper broken down according to the criteria available for species identication Criteria available Voucher seen Voucher exists, could not be seen, but species identity could be inferred from provenance No voucher exists but species identity could be inferred from provenance Educated guess from other data (elevation, ecology, descriptions) Voucher exists but could not be seen, species name thus unveried No voucher exists, species name thus unveried Studies 12 (16%) 11 (14%) 31 (40%) 4 (5%) 3 (4%) 16 (21%)

Note: the number of publications with unveried identity of the plant material would be much higher if not for the fact that only one Minthostachys occurs in Argentina.

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These two cases illustrate the general problem. Without voucher specimens available for identication in publicly accessible herbaria, it is in some cases a matter of guesswork, in others utterly impossible to nd out which species has been covered by an individual study. This severely limits the usefulness and comparability of their results. 8. Conclusions One of the most striking results of this literature survey is the realization that the vast majority of scientic studies, particularly those with a pharmacological or economic background, have been conducted on only one species, Argentinean Minthostachys verticillata. It is to be hoped that the future will see more research effort directed towards the other 16 species of Minthostachys, several of which have not even been examined for their oil composition, let alone their medicinal and economic potential and requirements for cultivation. It is possible that there are notable differences in oil composition between some species that remain undetected, as indicated by the observation that certain species have characteristic aromatic odors (e.g., often pleasantly fresh in Minthostachys spicata, sharply minty in Minthostachys acris; A.N. Schmidt-Lebuhn, pers. obs.), and this could be of some pharmacological or commercial interest. For Minthostachys verticillata, a wealth of information is already available, including a good overview of the variability of aromatic oil content as well as possible underlying reasons for this variability. In the face of possible overexploitation, some researchers have begun to address issues related to the cultivation of Minthostachys verticillata. It remains to be seen to what degree their ndings can be adapted for other species, e.g. Peruvian Minthostachys acris, which also appears to be intensively used. Even for the best examined species, however, several intriguing aspects have been neglected so far. More studies are needed to identify to what degree ecological and seasonal parameters inuence aromatic oil composition, and to what degree it is genetically determined; both aspects have, so far, only been examined separately. Comparatively little has also been published about optimal growth conditions (soil, elevation, light, nutrients, humidity, etc.), a question that is of as much economical as ecological interest. Germination experiments under varying conditions and after varying seeds storage times would be equally rewarding. It is also apparent that the pharmacological basis for the most commonly cited traditional uses of Minthostachys, against ailments of the respiratory and digestive systems, has received much less scientic attention than the effects of its essential oil on selected pests and parasites. The latter observation should be especially relevant from an ethnopharmacological perspective, hopefully prompting further research into the use of Minthostachys in folk medicine and possible pharmacological insights that can be derived from it. Future progress in this area will require a closer connection between ethnobotanical and pharmacological approaches than achieved previously in most studies of the group. Like several other economically interesting Lamiaceae (e.g., Mentha, Thymus L.) the genus Minthostachys exhibits high morphological variability, and identication of species is difcult. To ensure that the results of future pharmacological and ethnobotanical studies are comparable, it is desirable to increase awareness for the necessity to deposit voucher specimens in publicly accessible herbaria and that they should also be properly cited. In several cases presented in this review, data on oil composition or uses will never with any certainty be related to accepted taxonomic names simply because of the lack of voucher specimens or failure to actually locate them in the herbaria mentioned in publication, rendering it much less valuable than it could have been.

Acknowledgements The author would like to thank Ingeborg Hempel, Irene Lett, Marybel Morales, Marta Susana Ojeda, Daniel Renison and Doris Seidel for help with obtaining elusive articles, the staff of the herbaria BAF, BCN, CUZ, K and USM for providing information on or images of voucher specimens, Michael Kessler, Irene Lett and Thekla Pleines for helpful discussions, and Birgit Drger and two anonymous reviewers for helpful comments on a previous version of the manuscript. The authors research is currently funded by the Deutsche Forschungsgemeinschaft (DFG grant SCHM 2449/1-1).

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