Taxonomic study of Marsypianthes Mart. ex Benth. (Hyptidinae, Lamiaceae) in Brazil

Marsypianthes is a Neotropical plant genus distributed from southern Mexico to northeastern Argentina. It is composed of six species — M. arenosa, M. burchellii, M. chamaedrys, M. foliolosa, M. hassleri , M. montana — with all occurring in Brazil except for M. arenosa , which occurs in Mexico . Marsypianthes burchellii, M. foliolosa and M. montana are endemic to Brazil, while M. hassleri also occurs in Argentina and Paraguay and M. chamaedrys is widely distributed from southern Mexico to Argentina. Species of Marsypianthes are found in all the biomes of Brazil — Amazonia, Caatinga, Cerrado, Atlantic Forest, Pampa and Pantanal. Goiás is the Brazilian state with the greatest species richness of Marsypianthes , with four of the five species that occur in the country. According to IUCN criteria, three of the species are classified as Least Concern, one as Data Deficient and one as Vulnerable. The present study provides descriptions, an identification key, illustrations, geographic distribution data and information on the conservation status for all the taxa of Marsypianthes in Brazil.


Introduction
Marsypianthes belongs to the order Lamiales, family Lamiaceae, subfamily Nepetoideae, tribe Ocimeae and subtribe Hyptidinae.Hyptidinae is almost exclusively Neotropical, occurring throughout tropical and subtropical America (Harley at al. 2004).The flowers of species of this subtribe are arranged in a diverse array of modified bracteolate cymes, with a cupped inferior corolla lobe enclosing stamens, which provides an explosive pollination mechanism (Harley 1971) for nutlets, which have an expanded areole (Paton & Ryding 1998;O'Leary 2015).
Flowers of species of Marsypianthes are arranged in lax, few-flowered cymes, or in many-flowered cymes, and then in spherical capituliform heads.The calyx lobes are triangular, often reflexing in fruit.The gynoecium possesses a persistent, short, conic stylopodium and is attached to nutlets until maturity; the nutlets are cymbiform with an involute, laciniate margin and concave inner surface (Harley & Pastore 2012).Rudall (1981) mentioned that members of the subtribe Hyptidinae, other than Marsypianthes, possess ovoid nutlets.The gynoecial structure of Marsypianthes is unique within the family, making nutlet morphology a key character for identifying the genus (O' Leary 2015).
Recent phylogenetic studies have verified that Marsypianthes is a monophyletic group (Pastore et al 2011;Harley & Pastore 2012), characterized by nutlets with a concave inner surface and a fimbriated margin.
This work aimed to contribute to the morphological and taxonomical knowledge of Marsypianthes in Brazil by providing full descriptions, botanical illustrations, an identification key, distribution maps and ecological notes.

Materials and methods
This study was carried out based on the examination of specimens from the collections housed at BHCB, CEN, CGMS, HB, HEPH, HJ, HUEFS, HUTO, IBGE, ICN, INPA, JPB, MBM, R, RB, SP, SPF, UB, UFG and UFMT (acronyms according to Thiers); by consulting material deposited in virtual herbaria from INCT, K, KGF, NY, WU; and through the study of specimens collected during fieldwork, which were deposited in the herbarium of UFG.
Collection trips were carried out between October 2011 and May 2013, in the state of Goiás.Specimens were collected from different areas of the Cerrado biomecerrado sensu stricto, campo cerrado (cerrado fields), cerrado rupestre (rupestrian cerrado), campo úmido (humid fields), cerrado denso (dense cerrado) and riparian and gallery forest edges.Material was prepared following standard herborization techniques (Mori et al. 1989).
Collected material was identified using the descriptions and keys in Bentham (1833Bentham ( , 1848)), Schmidt (1858), Briquet (1907), Epling (1936) and Harley & Pastore (2012), and by comparison with type collections.Morphological descriptions follow Hickey (1973) for venation pattern terminology and Radford et al. (1974) for terminology related to pubescense and leaf, flower, fruit and seed morphology.Illustrations were made using a LEICA EZ4 stereoscopic microscope.Common names were acquired, when possible, from the labels of herbarium specimen and specialized literature.
The distribution map was produced in ArcGis 9.0 (Esri 2013) in the Laboratory of Image Processing and Geoprocessing (LAPIG) at UFG. Geographical coordinates were obtained from herbarium specimen labels when available, otherwise they were estimated using the coordinates of the headquarters of the municipality, available in Google Earth version 7.0.38542 (2013).

(HUEFS).
Marsypianthes burchelli is endemic to Chapada dos Veadeiros in Goiás State, growing near waterfalls, between rocks on hill slopes, edges of gallery forests, campo rupestre, campo cerrado, campo úmido, or sandy, loamy or sandyclay soils, from 823 to 1700 m in elevation.Flowering and fruiting throughout the year.The species is classified here as Vulnerable (VU), according to the criteria of IUCN (2017).
It occurs in all the biomes of Brazil in firm and swampy terrains, dunes and sandbank environments.Commonly grows on river or stream banks, under the shade of trees or shrubs, in vacant lots, on roads sides, in crops and pastures, on clay soils, latosols, sandy and rocky outcrops, from sea level to 1950 m elevation.It is considered a weed species (Lorenzi & Matos 2002).Flowering and fruiting throughout the year.Considering its wide occurrence, this species was assessed as Least Concern (LC), according to the criteria of IUCN (2017).
Marsypianthes chamaedrys differs from the other species of the genus by being the only one that lacks an underground system and has peduncle branches (5 mm) and externally glabrous corolla tubes.
Marsypianthes chamaedrys is the best-known species of the genus,and has antirheumatic medicinal properties (Heinrich 1992).Leaf extract is used against snake bite, as well as for diarrhea treatment, by the Tikunas indians in the Amazon region (Heinrich 1992).Pharmacological studies confirm the presence of triterpenoids in the species and reveal that its aqueous extract has analgesic, molluscicide, antibacterial and antifungal functions (Menezes et al. 1999) and anti-inflammatory and edema inhibitory action against the bite of Bothrops jararaca (pit viper) (Menezes et al. 1998).
Brazilian material selected: BRAZIL.Bahia: Formosa do Rio Preto, 11º 06' 33''S, 45º 33' 1''W, 5  Marsypianthes foliolosa is endemic to Brazil where it occurs in the North (TO), Northeast (BA, PI), Central-West (GO, MT) and Southeast (MG) regions.It can be found in the Cerrado biome in campos rupestres and associated to sandy-stony soils with rock outcrops, close to rivers and streams, from 173 to 1600 m in elevation.Flowering and fruiting throghout the year, with a peak in February.The species is classified as Least Concern (LC), according to the criteria of IUCN (2017).
Marsypianthes foliolosa is easily recognized by its elliptic or narrow-elliptic leaves.It is distinguished from the other species of the genus by its clearly smaller leaves (0.3-1.7 cm), shorter peduncle (0.5-7 mm) and few flowered cymes (one-two, rarely three or five), while the rest of the species have larger leaves, peduncles longer than 7 mm and many flowered cymes (three-22).The species can be also recognized by fewer bracteoles (two per flower), onefive flowers per inflorescence, and narrow-ovoid nutlets.Bentham (1848) mentioned that this species has usually one flower or rarely two to three flowers per cyme.The present research confirms Bentham's assertion, however, cymes with up to five flowers were also found.Briq., Bull. Herb. Boiss. Sér. II. 7: 620. 1907.
The species is found in sandy and closed fields, commonly growing in the shade of trees, in sandy loam and moist soils between from 115 to 1060 m in elevation.Flowering and fruiting probably throughout the year.Due to insufficient data to evaluate the conservation status of this species, it is here classified as Data Deficient (DD) according to the criteria of IUCN (2017).
This species can be easily recognized by the stigmas surpassing the anthers.This species morphologically resembles M. montana, by having elliptic and obovate leaf blades and cuneate and decurrent leaf base; however, it differs by having a membranaceous blade (vs.chartaceous in M. montana), a glabrous style (vs.sparsely hirsute in M. montana), smooth light-brown colored nutlets (vs.rough and dark-brown colored nutlets in M. montana), and peduncle measuring 1-1.5 cm long (vs.2.5-8 mm long in M. montana).Benth. Prodr. Syst. Nat. Regni Veg. 17 in elevation.Flowering and fruiting throughout the year.The species is classified as Least Concern (LC), according to the criteria of IUCN (2017).
Marsypianthes montana is easily recognized from the other species of the genus by being a caespitose growing herb and having chartaceous blades with irregularly serrated margins and purple filaments, all of which differ in the other species.