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Nomenclatural notes on Menthinae (Mentheae — Lamiaceae) native to Brazil

Menthinae, one of the five subtribes of Mentheae (Lamiaceae), with ca. 735 species, is represented in Brazil by a wide range of herbs and shrubs. During an investigation of the taxonomic history of the group for the Flora e Funga do Brasil project, several nomenclatural issues with typifications in the group have been identified. This study attempts to resolve them, making suggestions of new types and re-evaluation of synonyms. Thirteen lectotypifications and one neotypification are being proposed, along with the recognition of Thymus selloi Spreng. ex J.A.Schmidt as a nomen nudum. Most of the suggested changes are due to the loss of many of Sellow’s types at B during WWII, lack of specific herbaria references and collector numbers in the protologues, or to existence of designated syntypes in the referred collections. Our study is a contribution to the nomenclatural and taxonomic knowledge of Menthinae, providing a stable foundation for future study of its taxonomy, biogeography, and evolution.

Mentheae, the largest tribe of Lamiaceae, together with tribe Ocimeae, encompasses most of the diversity of Lamiaceae in the Neotropics (Drew & Sytsma, 2012). Among the five Mentheae subtribes, Menthinae stands out as a group comprising ca. 40 genera and 735 species spread throughout the temperate, subtropical, and tropical regions of all continents, except for Antarctica, and is mostly associated with Europe, Asia, and the Americas. The species of the subtribe are frequently found in high altitude vegetation formations, occupying mostly open phytophysiognomies, such as grasslands, riverine forests, bogs, rocky montane areas, forest glades or margins and, rarely, deserts (Harley et al. 2004;Bräuchler et al. 2010).
Menthinae is represented in Brazil by 42 native species in 8 genera and distributed mostly in open vegetation types, such as the campos de altitude (herbaceous to shrubby vegetation associated with igneous or metamorphic outcrops, composed mainly of granites, gneisses, or nepheline syenites, that had been uplifted during the Tertiary, and starting at altitudes above 1500 m.a.s.l.), the campos rupestres (herbaceous to shrubby vegetation on quartzite, arenite, and ironstone soils and outcrops of the Pre-Cambrian origin, mostly at elevations above 900 m.a.s.l.), both with high levels of endemism, the campos limpos (grasslands at lower altitudes dominated by herbs, mainly by members of Poales, among a diversity of other herbaceous taxa, and slender shrubs), or along forest margins of the Atlantic Forest, the Cerrado and the Pampa domains (Rizzini, 1997;Vasconcelos, 2011;Antar et al. 2022).
The taxonomy of the group had been initially presented by Sprengel (1827), who did not specify herbaria or collectors for the newly described species, followed by that of Bentham (1834Bentham ( , 1848, who on several occasions had referred to more than one herbarium and/or collectoroften without collector's numbersfor a single species, and that of Briquet (1896Briquet ( , 1887. Epling (1936) and Epling & Stewart (1939) made significant contributions to the advancement of taxonomic study of the group, including several lectotypifications (sometimes inadvertentsee Prado et al. 2015), and the proposal of several new taxa. Thereafter, new species or combinations have been proposed, but the taxonomy of most of the group remained largely stable (Brade, 1943;Epling, 1951;Epling & Mathias, 1957;Epling & Játiva, 1963;Irving, 1980).
With the completion of the Menthinae genera project for the online Flora e Funga do Brasil effort (Antar et al. 2022), there remained several unresolved nomenclatural issues for the native Brazilian species of Cunila D.Royen, Glechon Spreng., Hesperozygis Epling, Hoehnea Epling, and Rhabdocaulon (Benth.) Epling. These genera are represented in the Atlantic Forest domain, with Hoehnea restricted to it and the others having it as their main site of occurrence in Brazil (Epling, 1936;Epling & Stewart, 1939;Antar et al. 2022). Cunila, Glechon and mainly Rhabdocaulon are also represented in the Cerrado domain, while the first of those and Hesperozygis are found in the Pampa domain (Epling & Stewart, 1939;Antar et al. 2022).
The objective of our study is to establish correct typifications for the Menthinae species in Brazil, incorporating new nomenclatural decisions and summarizing the ones already made (Epling, 1936(Epling, , 1939Xifreda & Malo, 2004, 2006, thus establishing a stable foundation for future study of their taxonomy, biogeography, and evolution.

Results and discussion
Most of the required lectotypes selections are due to the loss of Friedrich Sellow's specimens housed at B during WWII (BGBM, 2021). They had been initially designated as syntypes by George Bentham (1834Bentham ( , 1848, but later inadvertently lectotypified or proposed as types of new taxa by Carl C. Epling (1936), who provided for them originally absent collection numbers, as can be seen in the photos of the types in the Field Museum Negatives collection. They represent Epling's interpretations of them based on specimens with collection numbers from other herbaria, such as P, and should be treated with caution.
Possible duplicates of the specimens lost at B may be found at several herbaria in Europe and the Americas, sometimes even carrying the original label from B and thus indicating direct links to Sellow's collections analyzed by Bentham. It is worth mentioning that B still holds many type specimens for these taxa, but they represent a later transfer, likely from HAL, of the Rudolf Baschant Herbarium (Hiepko, 1980), and as such cannot be original holotypes or lectotypes. Similar cases of problematic lectotypifications of some of the Brazilian taxa of Menthinae have been referred to by Xifreda & Mallo (2004, 2006. BRITTONIA [VOL Problematic cases with the designated syntypes are also common, as is the case for many species o f R h a b d o c a u l o n , w h e r e s e c o n d s t e p lectotypifications are required to stabilize prioritization among possible lectotypes. Details of remaining, historic, syntypes, disregarded as lectotypes by Epling's decisions (1936), are also provided. We also found rare cases of the complete lack of herbarium citations, as for example for Hesperozygis kleinii Epling & Játiva.
We present a nomenclatural treatment for the following taxa of Menthinae. The conclusions reflect our nomenclatural decisions after numerous analyses of type specimens housed in different herbaria, together with the protologues and other references, leading to the proposition of thirteen lectotypifications, one neotypification, and one nomen nudum: Notes.-Bentham (1834) described Cunila fasciculata based on a specimen Sellow s.n., housed at B. Epling (1936) cited the collection number 1524 for the Sellow specimen at B. However, the specimens at B had not been numbered, as can be seen in the Field Museum Negatives N°2 8974. Nevertheless, most of them had been seen and annotated by Bentham or Epling and confirmed as types. The specimen at B was lost during WWII, but one of Sellow's types that comes from the Rudolf Baschant Herbarium is still housed at B (Hiepko, 1980). Bentham (1834) referred to southern Brazil ("in Brasilia merid.") as the general geographic region for the type. W0027334 is the only specimen matching that general location, that is more precisely restricted to the state of Rio Grande do Sul, part of the Brazilian South (Antar et al. 2022). There cannot be much certainty that all the specimens identified as Sellow s.n. are in fact duplicates. Nevertheless, a combination of Bentham's annotation, the stamp of the Herbarium Benthamianum, and the designation by Epling (1936) as an isotype at K offers enough confidence to select this material ( Fig. 1) as the lectotype.
Schmidt (1858) considered Thymus selloi to be a synonym, reportedly in an annotation on the holotype of Gardoquia capitata. Since no description for that name had been made, and in accordance with Art. 38, Ex.1, Rec. 50B of the Shenzhen Code (Turland et al. 2018), it is here considered a nomen nudum.   Notes.-Bentham (1834) described Cunila galioides based on a specimen Sellow s.n., housed at B and on the material A.F.C.P Saint-Hilaire s.n., housed at P. Epling (1936) cited the specimen housed at B as the type, an action interpreted in Prado et al. (2015) as an inadvertent lectotype selection according to the ICN (Turland et al. 2018). Epling (1936) also cited the collection number 4400 for the Sellow specimen at B. However, the specimens at B had not been originally numbered, as can be seen in the Field Museum Negative N°2 8975. The only numbered collection is that of one specimen at P, but the calligraphy on its label corresponds to that of Bentham, while the original Sellow's collection at B does not have a collection number. The specimens at B, however, were lost during WWII, and are now represented solely by a photograph attached to the UC 61941 material, as confirmed by an annotation left by Epling who defined it as the type. Nevertheless, B still houses one of Sellow's types, the one coming from the Rudolf Baschant Herbarium (Hiepko, 1980). Bentham (1834) cited southern Brazil ("in Brasilia merid.") as the general geographic reference for the type. The only specimens matching that information are K000640692 and W0027331, even though the species is widely distributed throughout the South, Southeast and parts of the Center-West of Brazil (Antar et al. 2022). There is not much certainty that all specimens identified as Sellow s.n. are in fact duplicates, as Sellow could have collected the species more than once. Nevertheless, taking into consideration Bentham's annotations, its origin from the Herbarium Benthamianum, and Epling's annotation stating its status as an isotype, we hereby propose the specimen from K (Fig. 2) as the replacement lectotype. The Saint-Hilaire unnumbered specimens, housed at P and marked as types, are thus recognized as the remaining syntypes.

Cunila galioides
The synonym Cunila galioides var. hispida was proposed by Schmidt (1858) using one of Sellow's types at B, with no references to an herbarium or collection number. Later on, Epling (1936) inadvertently proposed Sellow 4524 as the lectotype. That material was lost during WWII, leaving it without an existing type. A neotype (Fig. 3) is hereby proposed based on the quality of preserved morphology of the specimen matching the description in the protologue. Notes.-Bentham (1834) described Cunila incisa based on a specimen Sellow s.n., housed at B. Epling (1936) inadvertently designated the collection number 3161 as the type, which is not found among examined types except for UC 61953 in its label annotations written at a later date, presumably by the same author. The specimen at B was lost during WWII and is now represented solely by a photograph attached to the UC 61955 material, as confirmed by a note left by Epling defining it as the type. B, within the Rudolf Baschant Herbarium, still houses one of Sellow's types (Hiepko, 1980). Bentham (1834) had southern Brazil ("in Brasilia merid.") as the general geographic reference for the type. The only available specimen that matches that information is W0027327, with a distribution restricted to the states of Santa Catarina and Rio Grande do Sul, parts of the Brazilian South (Antar et al. 2022). There is not much certainty that all specimens identified as Sellow s.n. are in fact duplicates, but a combination of Bentham's annotations, the stamp of the Herbarium Benthamianum, and the designation as an isotype in K by Epling (1936) gives us enough confidence to select the referred material (Fig. 4)  Notes.-Bentham (1834) described Cunila menthoides based on a specimen Sellow s.n., housed at B. Epling (1936) inadvertently designated the number 3990 as the type, which is presently found only in UC 61960 in the annotations written at a later date on its label, presumably by the same author, and in a photograph of the material attached to UC 61961. No other specimen has a collector's number, although most of them are marked as seen by Bentham or Epling. That specimen at B, however, was lost during WWII. No other available duplicates have collectors' numbers. Bentham (1834) cited southern Brazil ("in Brasilia merid.") as the general geographic reference for the type. The only specimen matching that information is HAL0104706, more precisely that of the states of Santa Catarina and Rio Grande do Sul (Antar et al. 2022). The specimens at G and HAL, though representative of the species, are identified as Cunila pulegioides Benth., a name that was never published -highlighted by Dr. Christian Bräuchler in a note on the second specimenand that should not be confused with Cunila pulegioides L., the basionym of Hedeoma pulegioides (L.) Pers.

Cunila incisa
There is no certainty that all specimens identified as Sellow s.n. are in fact duplicates, but a combination of Bentham's annotations, the stamp of the Herbarium Benthamianum, its designation as an isotype in K by Epling (1936), and an Epling's note treating it as an isotype should be enough to confidently select the referred material (Fig. 5)  Notes. -Epling (1936) described Glechon discolor based on the specimen Sellow 4210, housed at B. The specimen at B, however, was lost during WWII. There is no certainty that the specimens at K identified as Sellow s.n. are in fact duplicates but, considering Epling's annotations on them and his decision to treat them as isotypes at K, they are here treated as such. Taking into consideration the herbarium where Epling was working (Stafleu & Cowan, 1976), and the designation of an isotype at LA, later transferred to UC, we treat the material chosen by Epling (Fig.  6) as the most adequate option for the lectotype.
It is noteworthy to mention that even though the province of São Paulo was defined by Epling (1936) in the protologue as the area of occurrence for the type, no collections matching that information have been found. The only geographic reference more specific than the country ("in Brasilia merid.", southern Brazil) was found in a note written by Bentham on the sheet of the K000485444 specimen. This geographic term is fairly general and does include São Paulo, but that state is not part of the known distribution of the species, restricted to the states of Santa Catarina and Rio Grande do Sul (Epling & Stewart, 1939;Antar et al. 2022 Epling & Játiva (1963) did not define where the type materials were housed. Upon investigation, three specimens were uncovered. Among those, we consider the specimen at UC (Fig. 7), previously housed at LA, the most suitable candidate for lectotypification, since it was housed at LA where Epling was mainly working (Stafleu & Cowan 1976) and therefore likely the main specimen on which the description of the new species was based. 7. Hesperozygis nitida (Benth.) Epling, Repert. Spec. Nov. Regni Veg. Beih. 85: 133 (1936 specimen, which led to a lectotypification by Epling (1936), who designated the type specimen as housed in K, and an isotype in LA (later transferred to UC). The appointed collection for the type, however, holds two specimens, which brought the necessity for a second step lectotypification.
Although Epling stated on its label that the specimen K000485460 should be recognized as the lectotype, it presents the contradictory information that it was collected in Minas Gerais state on a printed note, which is not in agreement with the species protologue. Furthermore, the only specimen clearly mentioning Serra dos Órgãos Mountains is UC1943458, but the note concerning that location was written later, as confirmed by its distinct calligraphy. Finally, the specimen K000485461, suggested as an isotype on a note made by Epling, has its location appointed as "Brazil" with a handwritten note stating "Near summit (…)" followed by words which are almost illegible, but were interpreted, as seen in the UC1943458 isotype, as "(…) Organ Mts.", the same locality presented in the protologue. Therefore, we hereby choose the last K specimen (Fig. 8) as the second step lectotype due to the collection where it is housed, the notes left by Bentham, and its available geographical information.   8. Hesperozygis ringens (Benth.) Epling, Repert. Spec. Nov. Regni Veg. Beih. 85: 134 (1936 Notes.-Bentham (1834) described Glechon ringens based on a specimen Sellow s.n., housed at B. Later, Epling (1936) proposed a new combination Hesperozygis ringens for the new genus Hesperozygis, and inadvertently designated the collection number 3305 as its type, which can only be seen in the photograph of the duplicate described by him as the type and attached to UC 70231 and at a later date annotated in Epling's notes attached to the selected lectotype. No other specimen, including the one at B seen in the Field Museum Negative N°28968, displays a collector's number, although most of them are marked as seen by Bentham or Epling. The types at B have been lost during WWII, even though B still houses one of the Sellow's type, that from the Rudolf Baschant Herbarium (Hiepko, 1980). Bentham (1834) designated "in Brasilia merid." as the general geographic reference for the type. No similar information was found among the examined specimens, even though the species is restricted to the state of Rio Grande do Sul in the extreme South of Brazil (Antar et al. 2022). There is no certainty that all specimens identified as Sellow s.n. are in fact duplicates, but a combination of Bentham's annotations, the stamp of the Herbarium Benthamianum, the designation of an isotype at K by Epling (1936) and his note on the material treating it as an isotype should be sufficient to confidently select the referred material ( Fig. 9)  Notes.-Schmidt (1858) described Keithia minima using a specimen Sellow s.n., housed at B. Epling (1936) assigned to it the collection number 4441, what is here understood as an inadvertent lectotypification. Later on, Epling & Stewart (1939) proposed Hoehnea minima, based on Epling's new genus Hoehnea. The lectotype at B was lost during WWII and is only preserved as a negative photograph in the UC 64015 material. The replacement lectotype (Fig. 10), marked as an isotype in Epling's annotation, represents a complete specimen adequate for detailed morphological observation, and therefore most suitable to be treated as such. significant parts of Keithia Spreng. into the new genus Rhabdocaulon, and inadvertently lectotypified the specimens by citing the Sellow s.n. specimen at B as the type. The specimen at B, however, was lost during WWII and can be seen only in a photograph of a specimen in the Field Museum Negative N°28957, without, however, certainty that it represents the specimen selected by Epling.
Though Bentham (1834) described the occurrence of the species in southern Brazil ("in Brasilia merid."), and Epling (1936) cited Minas Gerais as the province of its occurrence, such geographic references have not been found among the analyzed specimens. Other geographic references, found on the label of the HAL0114392, were "Brasil aequin." (Which may be interpreted as a reference to Minas Gerais), and, in a note to the K000485595 isolectotype, St. Paulo, part of southern Brazil. Both latter regions are in fact parts of the actual distribution for the species (Antar et al. 2022).
A combination of Bentham's annotations, the stamp of the Herbarium Benthamianum, the designation of an isotype at K by Epling (1936), and his note treating it as a "cotype"or, rather, an isotypemakes the designated specimen (Fig. 11) the most suitable replacement as the lectotype. It is also worth mentioning that both materials at K are part of the same sheet, though with different barcodes and associated notes.
Upon investigation, we found that there are in fact two specimens with the same collector's number in the collection, which necessitates a second-step lectotypification. Considering prioritizing between the two specimens, we decided on the one (Fig. 12) with the higher quality of preserved morphology and more closely matching the description. A third specimen, from Saint-Hilaire's collection and numbered C2-2791-10, is designated as a syntype. Considering the decisions made, this last specimen should be recognized as a remaining syntype.
Notes.-Bentham (1834) described Cunila stricta using a specimen Sellow s.n., housed at B, and the material A.F.C.P. Saint-Hilaire, housed in the collector's collection. Later on, Epling (1936) proposed the name Rhabdocaulon strictum as a combination in the newly created genus Rhabdocaulon, and made an inadvertent lectotypification by designating the Sellow material as the type. The material at B, however, was lost during WWII. Bentham (1834) used "in Brasilia merid." as a general geographic reference for the type. The only specimen in accordance with such reference is W0013599, even though the species is only found in the states of Santa Catarina and Rio Grande do Sul, parts of the Brazilian South (Antar et al. 2022). A combination of Bentham's annotations, the stamp of the Herbarium Benthamianum, and the designation as an isotype at K by Epling (1936) gives us enough confidence to select the referred material (Fig. 14) as the replacement lectotype. The specimens at P and US and marked as types are part of Saint-Hilaire's collection and should be recognized as remaining syntypes.