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Pedicularis L. Genus: Systematics, Botany, Phytochemistry, Chemotaxonomy, Ethnopharmacology, and Other

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Pedicularis L. Genus: Systematics, Botany, Phytochemistry, Chemotaxonomy, Ethnopharmacology, and Other plants Review Pedicularis L. Genus: Systematics, Botany, Phytochemistry, Chemotaxonomy, Ethnopharmacology, and Other Claudio Frezza 1,* , Alessandro Venditti 2 , Chiara Toniolo 1, Daniela De Vita 1, Ilaria Serafini 2, Alessandro Ciccòla 2, Marco Franceschin 2, Antonio Ventrone 1, Lamberto Tomassini 1 , Sebastiano Foddai 1, Marcella Guiso 2, Marcello Nicoletti 1, Armandodoriano Bianco 2 and Mauro Serafini 1 1 Dipartimento di Biologia Ambientale, Università di Roma “La Sapienza”, Piazzale Aldo Moro 5, 00185 Rome, Italy 2 Dipartimento di Chimica, Università di Roma “La Sapienza”, Piazzale Aldo Moro 5, 00185 Rome, Italy * Correspondence: [email protected]; Tel.: +39-0649912194 Received: 28 June 2019; Accepted: 12 August 2019; Published: 27 August 2019 Abstract: In this review, the relevance of the plant species belonging to the Pedicularis L. genus has been considered from different points of view. Particular emphasis was given to phytochemistry and ethnopharmacology, since several classes of natural compounds have been reported within this genus and many of its species are well known to be employed in the traditional medicines of many Asian countries. Some important conclusions on the chemotaxonomic and chemosystematic aspects of the genus have also been provided for the first time. Actually, this work represents the first total comprehensive review on this genus. Keywords: Pedicularis L. genus; Orobanchaceae family; phytochemistry; chemotaxonomy; ethnopharmacology 1. Systematics Pedicularis L. is a genus of hemiparasitic plants, originally included in the Scrophulariaceae family but now belonging to the Orobanchaceae family [1]. The rest of the systematic classification is the following: order Scrophulariales, subclass Asteridae, class Magnoliopsida, division Magnoliophyta, superdivision Spermatophyta, subkingdom Tracheobionta. The genus comprises 568 accepted species, 335 synonymous species, and 450 unresolved species [2]. 2. Etymology of the Name The etymology of the genus name is Latin, with the term “pediculus” meaning “louse”, which refers to the fact that, according to an ancient English belief, cattle which grazed on these plants were soon found to be infested with lice [3]. 3. Botany Plants of the genus Pedicularis are generally herbaceous and perennial, with a height which can reach up to 50 cm. Annual or biennial species are quite rare but present. From the morphological standpoint, these species are characterized by big and fleshy roots, often taproots, which contain specific organs (haustoria) for their feeding on the lymph of the near plants. The stem is erect and ascendant and may be simple or branched (Figure1). The leaves are basal and cauline. The former ones are disposed to form a rosette and are petiolate. The latter ones are opposite, alternated or verticillated, Plants 2019, 8, 306; doi:10.3390/plants8090306 www.mdpi.com/journal/plants Plants 2019, 8, 306 2 of 62 Plants 2019, 8, x FOR PEER REVIEW 2 of 53 Plants 2019, 8, x FOR PEER REVIEW 2 of 53 and sessile, instead. Both of these have a lanceolate shape and dentate margins which are rarely entire. are rarely entire. Bracts are also present and are similar to the cauline leaves, even if they are smaller Bractsare rarely are also entire. present Bracts and are are also similar present to theand caulineare similar leaves, to the even cauline if they leaves are smaller, even if (Figurethey are1 ).smaller More or (Figure 1). More or less dense terminal spikes generally constitute the inflorescence. The flowers are less(Figure dense 1). terminal More or spikes less dense generally terminal constitute spikes generally the inflorescence. constitute The the inflorescence. flowers are big, The hermaphrodite, flowers are big, hermaphrodite, zigomorphic, and tetrameric or pentameric. They can be sessile or zigomorphic,big, hermaphrodite, and tetrameric zigomorphic, or pentameric. and tetrameric They can beor sessile pentameric. or pedunculated. They can The be floreal sessile formula or pedunculated. The floreal formula is X, K(5), [C (2+3), A 2+2], G (2), (superior), capsule. The calyx is ispedunculated. X, K(5), [C (2+ The3), Afloreal 2+2], formula G (2), (superior),is X, K(5), [C capsule. (2+3), A 2+2], The calyxG (2), is(superior), gamosepalous, capsule. formed The calyx by is five gamosepalous, formed by five lobes that may be dentate or not. The corolla is gamopetalous and gamosepalous, formed by five lobes that may be dentate or not. The corolla is gamopetalous and lobesbilabiate that maywith be a dentatecylindrical or not.shape The slightly corolla compressed is gamopetalous on its andsides. bilabiate Its color with ranges a cylindrical from pink shape to bilabiate with a cylindrical shape slightly compressed on its sides. Its color ranges from pink to slightlywhite, compressed passing through on its sides.red, purple Its color, and ranges yellow. from The pink androecium to white, passing possesses through four red, didinamous purple, and white, passing through red, purple, and yellow. The androecium possesses four didinamous yellow.stamens The with androecium the filaments possesses well included four didinamous into the base stamens of the withcorolla. the The filaments anthers well are includedhidden among into the stamens with the filaments well included into the base of the corolla. The anthers are hidden among basedense of thehairs corolla. and may The be anthers mucronate. are hidden The pollen among maturation dense hairs is contemporaneous and may be mucronate. to the stigma. The pollenThe dense hairs and may be mucronate. The pollen maturation is contemporaneous to the stigma. The maturationovary is superior, is contemporaneous formed by two to carpels the stigma., and bilocular. The ovary The isstylus superior, is inserted formed in the by ovary two carpels, apex and and ovary is superior, formed by two carpels, and bilocular. The stylus is inserted in the ovary apex and bilocular.is filiform. The stylusThe stigma is inserted is simple in the ovaryand protrud apex anded isbeyond filiform. the The corolla stigma hat is simplein order and to protruded avoid is filiform. The stigma is simple and protruded beyond the corolla hat in order to avoid beyondself-pollination the corolla (Figure hat in 1). order The to fruit avoid is an self-pollination acuminated bivalve (Figure capsule1). The with fruit an is oval an acuminated-lanceolate shape bivalve self-pollination (Figure 1). The fruit is an acuminated bivalve capsule with an oval-lanceolate shape capsule(Figure with 1). The an oval-lanceolateseeds are numerous shape or (Figurenot and1 present). The seedsan angular are numerous geometry. orReproduction not and present occurs an (Figure 1). The seeds are numerous or not and present an angular geometry. Reproduction occurs angularthrough geometry. pollination Reproduction by insects or occurs dispersion through [4,5]. pollination by insects or dispersion [4,5]. through pollination by insects or dispersion [4,5]. (a) (b) (c) (d) (a) (b) (c) (d) Figure 1. Examples of the morphological features of Pedicularis species—stem (a), leaves (b), flowers FigureFigure 1. 1.Examples Examples of of the the morphologicalmorphological features of of PedicularisPedicularis speciesspecies—stem—stem (a), (a leaves), leaves (b), (b flowers), flowers (c), fruits (d) [source Google images]. (c),(c fruits), fruits (d ()d [source) [source Google Google images]. images]. 4. Distribution and Habitat 4.4. Distribution Distribution and and Habitat Habitat The species of this genus are distributed in Europe, especially in the mountainous areas of the TheThe species species of of this this genus genus areare distributeddistributed in Europe Europe,, especially especially in in the the mountainous mountainous areas areas of the of the Mediterranean Basin, and in Northern Asia and America (Figure 2). The highest biodiversity is MediterraneanMediterranean Basin, Basin and, and in Northernin Northern Asia Asia and and America America (Figure (Figure2). The 2). highestThe highest biodiversity biodiversity is present is present in Europe, with about 70 species, India, with about 83 species, and China, with about 350 inpresent Europe, in with Europe about, with 70 about species, 70 India,species, with India about, with 83 about species, 83 species and China,, and China with, aboutwith about 350 species, 350 species, 271 of which are endemic [6–8]. In North America, the present species are 36 with two 271species, of which 271 are of endemicwhich are [6 –endemic8]. In North [6–8]. America, In North the America present, the species present are 36species with twoare 36 endemisms with two [9]. endemisms [9]. These species have been reported in Africa and Australia only as imported plants. endemisms [9]. These species have been reported in Africa and Australia only as imported plants. TheseThe speciespreferred have habitat been is reporteda temperate in Africa mountain andous Australia one. The only soil as must imported be quite plants. acidic The and preferred little- The preferred habitat is a temperate mountainous one. The soil must be quite acidic and little- habitatdraining is a. temperateThe typical mountainous areas where one.these The species soil mustcan be be found quite acidicare meadows and little- and draining. lawns with The little typical draining. The typical areas where these species can be found are meadows and lawns with little areasother where vegetation these species[3]. can be found are meadows and lawns with little other vegetation [3]. other vegetation [3]. Figure 2.Worldwide
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