Andre A Naranjo

Rahul R Pathirickal

Kanchi N Gandhi

Piero G Delprete

Riccardo M Baldini

Arnoldo Santos

Kanchi N Gan- Dhi

Arnoldo Santos-Guerra

Láza- Ro Sánchez-Pinto

Javier Francisco

ORCID Fred W Stauffer

Javier Francisco-Ortega

Philip Barker

Honoring Sabin Berthelot: Nomenclature and botanical history of Berthelotia DC. (Asteraceae, Inuleae)

Keywords: Asteraceae, Berthelotia, Canary Islands, India, Sonora Desert, Mexico, USA, Nomenclature, Phylogeny, Sabin Berthelot

The genus Berthelotia DC. (presently a synonym of the accepted name Pluchea Cass.), Asteraceae, Inuleae has been regarded as comprising two species, B. lanceolata DC. and B. sericea (Nutt.) Rydb. It honors the French naturalist Sabin Berthelot (1794-1880) for his contribution to the studies of the flora and fauna of the Canary Islands. Under the leadership of Philip B. Webb (1793-1854), Berthelot co-authored the most important work pertinent to the natural history of this archipelago: Histoire Naturelle des Îles Canaries. The nomenclature of Berthelotia and its associated names are revisited here. The holotype of Polypappus sericeus Nutt. (K000974653) was located, and lectotypes were assigned to B. lanceolata (G00450458) and B. lanceolata var. senegalensis DC. (G00452584). Phylogenetic analyses based on nucleotide sequences of the nrDNA ITS were conducted focusing on members of what is known as the Clade J of the Inuleae. A well-supported clade (100% P.P.

) that we have labeled as "Clade I" was uncovered, and it is composed of three species that belong to the Pluchea indica Clade plus P. dioscoridis and P. ovalis; but this lineage did not support a previous hypothesis to include P. lanceolata. The two species of Berthelotia did not form a monophyletic group and seem to be distantly related; furthermore, none of these two taxa is sister to Andre A. Naranjo et al.

Tessaria, a South American genus that has been suggested to be a close relative to Berthelotia. Ethnobotanical uses of B. lanceolata and B. sericea are reviewed.

INTRODUCTION

The botanical history of the Canary Islands has been highly influenced by the British botanist Philip B. Webb (1793Webb ( -1854) ) and the French naturalist Sabin Berthelot (1794-1880; Figure 1). They were the editors and main authors of the Histoire Naturelle des Îles Canaries (thereafter "Histoire Naturelle"), a multivolume work published between 1835 and 1850, that includes the Phytographia Canariensis (thereafter "Phytographia"), the most important single work ever published on plant taxonomic diversity of the Canaries (Webb andBerthelot 1835-1850).

The journal Webbia honors Webb in recognition for establishing in his will that his extensive herbarium, botanical documents, and personal house located in Paris were to be bequeathed to the Grand Duke of Tuscany Leopold II of Lorraine (1797Lorraine ( -1870)). After the demise of Webb, his house in Paris was sold to establish an endowment for the curation of the archives and herbarium collection that he donated to Tuscany. Currently, Webb's specimens are a central part of the FI herbarium of the Sezione Botanica "F. Parlatore" of the Natural History Museum of the University of Florence, constituting the Herbarium Webbianum. His entire library and archives are kept in the Biblioteca di Scienze -Fondi di Botanica and Fondi Archivistici, also at the University of Florence.

In a recent work published in the journal Webbia, a botanical history and nomenclature study was presented, including a discussion of the description of the genus Webbia by three different botanists who published this name independently, referring to three taxa belonging to different families (Asteraceae, Hypericaceae) or tribes within Asteraceae (Astereae and Vernonieae). [START_REF] Francisco-Ortega | Honoring Philip Barker Webb: The three intriguing stories of Webbia as a genus name[END_REF] recognized the legacy of this outstanding British botanist, and the research that we are presenting in this issue of Webbia is in many aspects a follow up of the work that we previously published on the genus Webbia, as it provides nomenclature and historical insights into Berthelotia DC. (Asteraceae, Inuleae), a genus that the Swiss botanist Augustin Pyramus de Candolle (1778-1841, Figure 1) dedicated to Sabin Berthelot encompassing two species, B. lanceolata DC. and B. sericea (Nutt.) Rydb.

There have been a few extensive studies pertinent to the life of Sabin Berthelot (e.g., Zerolo 1881;[START_REF] Drouin | Sabin Berthelot et la Géographie Botanique dans l'Histoire Naturelle des Îles Canaries (1836-1850[END_REF]Instituto de Estudios Canarios 1980;[START_REF] Belin | Los frutos de una amistad[END_REF][START_REF] Brun | Un Francés entre Guanches[END_REF][START_REF] Brun | El legado de Sabin Berthelot: Una primera aproximación a su biblioteca y sus colecci-ones (1878-1894). Vegueta[END_REF], and he also wrote autobiographical accounts (1838)(1839)(1840)1980) that help to reconstruct the most relevant aspects of his life. Therefore, a full biog- raphy regarding his achievements and challenges are outside the scope of this contribution. However, information pertaining to his botanical contributions should be reviewed in order to fully understand why one of the most important plant taxonomists of the 19 th century, Augustin P. de Candolle, described the genus Berthelotia to honor him. An overview of Berthelot's biography and botanical work is therefore presented below.

Sabin Berthelot: A biographical sketch

Unlike Webb, Berthelot did not have any formal academic training in biological disciplines. Born in Marseille (April 4, 1794), between 1804 and 1809 he only undertook secondary education studies in the Liceum of Marseille (also known as Lycée Impérial). Afterwards, he served in the French Navy from 1809 to 1814. Berthelot was uninterested in formal academic studies, and this could explain why he never pursued higher education (Le Brun 2016: 64-65). After this military service, he started to work as a civil sailor in 1816, and little is known about his professional activities between this year and 1819 (Le Brun 2016: 70-71). He arrived in the Canary Islands (Tenerife) in 1820. Berthelot (1838Berthelot ( -1840: 11-14) : 11-14) indicated that he reached the Canaries by coincidence, as a trip he took to Senegal had several misfortunes that caused the vessel in which he travelled to change route to Tenerife. Once in the Canaries, in May 1824 he opened a high school in the city of La Orotava under the name of Liceo de la Orotava, but this initiative was halted in April 1825 by the Catholic clergy as they opposed the establishment of liberal education centers in Tenerife (Le Brun 2016: 120). During this time he was also connected with the Botanic Garden of La Orotava through its founder Alonso de Nava y Grimón (VI Marqués de Villanueva del Prado). During this time, in his publications, Berthelot (1827aBerthelot ( , 1827bBerthelot ( , 1828aBerthelot ( , 1828b) cited his job title as the "Director" of the garden; however, he never had any official appointment for this or any position in this garden (Le Brun 2016: 121). Founded in 1788, it is the oldest botanic institution of Macaronesia and the second oldest of Spain. It was part of a network of Jardines de Aclimatación (acclimation gardens) that the Spanish Crown had in the Canaries and mainland Spain (Puerto Sarmiento 1988: 201;2002: 34;Cioranescu 2010: 150-151). Berthelot (1844) had an interest in plant acclimation protocols to cultivate plants into temperate regions, and this could explain Alonso de Nava's interest in having him associated with this botanic garden.

In 1828, eight years after his arrival to the archipelago, he met Webb in one of the gorges located near the main harbor of the island in Santa Cruz de Tenerife.

It seems that they met by chance when they were botanizing in this area (Santos Guerra 2007); importantly, this casual meeting led to a massive joint project that resulted in the afore-mentioned work entitled Histoire Naturelle des Îles Canaries. Webb's original plan was to continue his travel to Brazil to study its natural history; however, his plans were cancelled, and for two years Webb and Berthelot travelled through the archipelago, with Berthelot employed by Webb as his field assistant. Together they collected natural history specimens and gathered information pertinent to the Canarian flora, fauna, geography, and geology; with the exception of La Gomera and El Hierro, they visited all islands (including La Graciosa) (Relancio Menéndez and Breen 2006: 38, 56, 61). In 1830, Webb and Berthelot traveled back to Europe aiming to produce an extensive account on the natural history of the Canaries. After this long trip across the continent, they arrived in Paris in 1833.

Webb and Berthelot had several disagreements regarding the structure and focus of this work that was exacerbated by 1841 when their partnership collapsed (Suárez Martín 2018: 169-170). Webb's vision was to follow the approach done in other well-known natural history or flora projects that were published by naturalists such as [START_REF] Ruiz | Flora Peruvianae, et Chilensis Prodromus[END_REF] in their Florae Peruvianae, et Chilensis or Augustin de Saint-Hilaire (1829-1833) in his Flora Brasiliae Meridionalis. These were based mostly on solid taxonomic treatments and were richly illustrated with color plates. In contrast, Berthelot preferred that an important component of the work to be written in French, mostly including aspects pertinent to the ethnography, society traditions, and history of the Canaries. These topics were rarely covered in other books that followed a scientific approach to the natural history of a particular area during this time. The volume-parts of the Histoire Naturelle that had Berthelot as the single author were written in French and did not have any formal plant taxonomic components [START_REF] Berthelot | Observations sur les forêts Canariennes, leurs changements et leurs alternances dans les différentes îles[END_REF](Berthelot -1842[START_REF] Berthelot | Coup d'Oeil sur les Forêts Canariennes, sur Leurs Changements et Leurs Alternances[END_REF](Berthelot -1839(Berthelot , 1838[START_REF] Berthelot | Ethnographie. Annales de la Conquête[END_REF][START_REF] Berthelot | Ethnographie. Annales de la Conquête[END_REF](Berthelot -1842)). Regarding Berthelot's botanical contributions to this project, he was the sole author of the Geographie Botanique, totaling 181 pages and represents the first part of Volume 3 of this work [START_REF] Berthelot | Observations sur les forêts Canariennes, leurs changements et leurs alternances dans les différentes îles[END_REF](Berthelot -1842)).

All taxa described in the Phytographia that involved Berthelot had him as the second author and Webb as the first author. Authorship of this extensive floristic and taxonomic treatment and of the Géographie Botanique was one of the reasons behind the intense arguments that they had in 1841 and 1842 (Suárez Martín 2018: 171, 213, 219), and it is likely that they also pertained to the author order of the new taxa included in the Phytographia. As far as we are aware, prior to his involve-ment in the Histoire Naturelle, Berthelot (1827b) validly published only a single new species by himself, the Tenerife endemic Viola teydea Berthel. (alternative name V. canariensis Berthel.); however, this is a heterotypic synonym of the previously validly published species V. cheiranthifolia Bonpl. in 18071 . Unlike Berthelot's background, Webb had a solid botanical training acquired in Oxford, with expertise in classical Greek and Latin languages, and importantly a good understanding of taxonomic methods and protocols. After their split in 1842, Berthelot followed other endeavors that included traveling across the Western Mediterranean to perform fisheries studies for the French government (Le Brun 2016: 285-291). Eventually, he returned to Tenerife in 1847 to start working in the French Consulate of the island, where in due course he was appointed First Class Consul in 1874 (Le Brun 2016: 360).

After 1842, several parts of the Phytographia were published until the project ended in 1850. Between 1842 and Webb's death (31 August 1854) they exchanged nine letters (Suárez Martín 2018: 38-39, 216-239, pers. comm.). However, only one of the letters (October 11, 1844) makes reference to galley proofs revised by Berthelot and discussions regarding the meaning of common plant names from the Canaries, suggesting that in spite of the intense arguments they had in 1842, Berthelot was still somehow associated with the Histoire Naturelle project.

Based on his life-time publication record, it is evident that the vast majority of Berthelot's single-author publications were devoted to the general public and had a wide range of topics. Appendix 1 lists the 29 botanical publications authored by Berthelot, 17 of them focused on the Canary Islands and the rest covered introductory works to Caribbean plants, ivies, larches, Orchilla lichens, palms, violets, and the importance of plant acclimation. These popular publications were important outreach initiatives aimed at generating interest in the general public, particularly those published in Spanish for Canary Islands magazines such as Revista de Canarias, Eco de Comercio, and El Amigo del País contributed to increase public awareness for the study and conservation of the botanical history heritage of these islands.

The importance of Berthelot's contributions to the botanical research history of the Canary Islands cannot be understated as he assisted Webb to make his floristic and taxonomic assessments. As Webb's project employee, Berthelot was actively involved in organizing the herbarium collections that were used for the extensive floristic component of this work, and there is no doubt that they had discussions pertinent to the identification of the plant material that was examined for the Phytographia. Indeed, when Candolle (1836: 375) described the sunflower genus Berthelotia as a homage to Berthelot's, he states "Fruticulum elegantem dicavi cl. [clarissimus] Berthelot florae faunaeque Canariensis strenuo ilustratori" [I have dedicated this elegant small shrub to honorable Berthelot, an active interpreter of the flora and fauna of Canary Islands], acknowledging the importance of Berthelot's input in the Histoire Naturelle to the study of the unique flora and fauna of these islands.

BOTANICAL HISTORY AND SYSTEMATICS

When Webb and Berthelot worked on the publication of the Histoire Naturelle des Îles Canaries, they were able to interact with some of the most prominent botanists from Europe. A total of 15 of the family treatments found in this work were written by some of these distinguished plant taxonomists (Relancio Menéndez and Breen 2006: 176;[START_REF]Los estudios botánico-zoológicos de A. Moquin Tandon y su relación con los archipiélagos macaronésicos[END_REF]. Among them was the Swiss botanist Alphonse Pyramus de Candolle (1806Candolle ( -1893)), who prepared the revision of Myrsinaceae (currently merged within the Primulaceae). He was the son of Augustin Pyramus de Candolle who was the founder of the Candolle's botanical dynasty, a family based in Geneva which provided consequential botanical research contributions during four consecutive generations, with the last member of this lineage being Richard Émile Augustin de Candolle (1868Candolle ( -1920)).

Augustin P. de Candolle did not write any text for the Histoire Naturelle, but he and Webb had extensive correspondence that included 38 letters housed in the Conservatoire et Jardin botaniques de Genève-Archives (22 letters) and in the University of Florence-Science Library (16 letters). Candolle exchanged with Berthelot only four letters, which are currently housed in the Archives of the Botanic Garden of Geneva. The content of these numerous letters includes many botanical inquires pertinent to the seminal work of Webb and Berthelot on the Canary Island flora, and it is unclear why this prominent Swiss botanist was not involved in preparing any of the plant taxonomy accounts of the Histoire Naturelle. Plausible reasons were that he already had a large commitment with the publication of his Prodromus, and that he suffered poor health during his last years of life and eventually passed away in 1841, when the Phytographia was still in progress. This section of the Histoire Naturelle was by far the largest (1403 pages and 287 plates) of this multivolume enterprise, and it was published in 106 independent livraisons (booklets) ranging from 1835 to 1850 [START_REF] Stearn | On the dates of publication of Webb and Berthelot's "Histoire Naturelle des Îles Canaries[END_REF].

Augustin de Candolle's autobiography has a few details regarding his interactions with Webb and Berthelot that confirm that they sought his help as they were preparing the Phytographia (Candolle 2004: 459). As already mentioned, in 1830 they moved from the Canaries to continental Europe to prepare the Histoire Naturelle. However, before reaching their final destination in Paris in 1833, they travelled extensively for three years in North Africa, the French Mediterranean region, Italy, and Switzerland. According to Candolle (2004: 459), during this journey Webb and Berthelot spent a few months in Geneva where they shared herbarium specimens with, and studied Canarian collections that were made by botanists who visited these islands, particularly Pierre M. Auguste Broussonet (1761-1807) from France, Antoine Courant from Switzerland, and Christen Smith (1785-1816) from Norway. Evidently, Berthelot had high respect for Candolle, and one year after he and Webb settled in Paris, he wrote a biographical account on him [START_REF] Berthelot | Anonymous author[END_REF]) that was one of the few published biographies produced before Candolle passed away in Geneva in 1841 (Candolle 1862: 588). During his life Berthelot (1839aBerthelot ( , 1839b) ) wrote only two other botanical biographies; they were for the French botanists Joseph P. Tournefort (1656-1708) and Valmont de Bomare .

It is worth mentioning that Augustin de Candolle's descriptions of Berthelotia and Webbia were published in volume 5 of his monumental Prodromus (Candolle 1836: 72, 375-376). Although, both genera belong to Asteraceae, Berthelotia is a member of the tribe Inuleae, whereas Webbia is placed within the tribe Vernonieae. The contents of two of the letters that Candolle exchanged with Webb and Berthelot reveal that he had other sunflower plants in mind to pay homage to his French and British friends. One of these letters is located in the University of Florence-Science Library (dated 13 April 1833, from Candolle to Webb, document #73.9.1-73.9.6), the second one is housed in the Archives of the library of Botanic Garden of Geneva (dated 12 July 1833, from Berthelot and Webb to Candolle, unnumbered document). The second of these letters has a more extensive account regarding Candolle's plans to describe Berthelotia than the other one. Therefore, an annotated translation of the latter is presented in Appendix 2, since it also deals with the taxonomic identity of Asteraceae specimens that Webb and Berthelot examined for their Canary Island flora work. The information found in these two letters is also relevant to the botanical history of Berthelotia. From these letters it appears that, at least initially, Candolle wanted to publish the new genus Berthelotia to accommodate both the Canarian-Salvagen endemic Chrysocoma sericea L.f., and a new species, restricted to Gran Canaria, that was going to be named "B. glaberrima." It seems that when Candolle exchanged these letters, he was unaware of Schizogyne Cass. (Asteraceae), which was published earlier by Cassini [START_REF] Cuvier | Dictionnaire des Sciences Naturelles dans Lequel On Traite Méthodiquement Des Differents Êtres de la Nature[END_REF]. Schizogyne, endemic to the Canary and Savage islands, is the taxonomic choice to transfer Chrysocoma sericea to a different genus and to accommodate the new taxon to which Candolle assigned the species epithet of "glaberrima." Later, Candolle (1836: 75) was aware of Schizogyne as the taxonomic option for these rearrangements, because in the same work in which he described Berthelotia (Candolle 1836: 375-376), he made the new combination S. sericea (L.f.) DC. and validly published S. glaberrima DC. as a new species (Candolle 1836: 473). Candolle's plans to dedicate a genus name to Berthelot did not change, and he still described Berthelotia, but it was not based on Canary Islands plants but on a species that he named B. lanceolata DC. (Candolle 1836: 376, Figures 2-3), which has a relatively widespread distribution in India, Pakistan, Afghanistan, and Tropical Africa (Figure 4; King-Jones 2001: 82-84).

Following the well-known treatment of [START_REF] Bentham | Compositae[END_REF] on the classification of Compositae plant genera, taxonomists have not recognized Berthelotia as a distinct genus, and currently Berthelotia is treated as part of Pluchea Cass. (Compositae Working Group -CWG 2022). [START_REF] Bentham | Compositae[END_REF] generic concepts for the Asteraceae have been greatly influential in the taxonomic history of this family [START_REF] Cronquist | The Compositae revisited[END_REF]; however, from their work it is not readily evident whether Bentham and Hooker (1873: 291) recognized Berthelotia (printed in small cap font) as part of Pluchea but without any taxonomic rank, or they (1873: 550) proposed Berthelotia [printed in italics and regular font] to be a section of Pluchea. Nevertheless, it was not Bentham and Hooker, but Clarke (1876: 94) who first made the legitimate transfer of Berthelotia to Pluchea when he published the combination P. lanceolata (DC.) C.B.Clarke. Berthelotia can be distinguished from other species of Pluchea s. str. by their lack of resin ducts, flattened filament collar cells, and divided styles (Anderberg 1991b).

Pluchea lanceolata has ample ethnobotanical use in India (Shanker and Srivastava 2017; Figure 2; see section on ethnobotany below), and it can also be a noxious weed [START_REF] Inderjit | Pluchea lanceolata: A noxious perennial weed[END_REF]. Because of its medicinal properties and widespread distribution, this species has been the subject of phytochemical (e.g., Srivastava and Shanker 2012; Pandey 2018), in vitro culture (e.g., [START_REF] Kumar | In vitro propagation of Pluchea lanceolata, a medicinal plant, and effect of heavy metals and different aminopurines on quercetin content[END_REF]), and ecological (e.g., Inderjit 1998[START_REF] Inderjit | Allelopathic effect of Pluchea lanceolata on growth and yield components of mustard (Brassica juncea) and its influence on selected soil properties[END_REF] studies.

Pluchea sericea (Nutt.) Coville is a species restricted to arid parts of northeastern Mexico and western United States (Figures 567). This species is isolated among North American Pluchea, with its woody perennial habit and densely arranged waxy eglandular leaves, and [START_REF] Nesom | Pluchea[END_REF] stated that "Pluchea sericea is more similar to the segregates than to the herbaceous American groups [of the genus]." In 1848, it was originally described as Polypappus sericeus Nutt., and in 1906 this species was accommodated in Berthelotia (as B. sericea (Nutt.) Rydb.) by the Swedish-American botanist Per Axel Rydberg (1860-1931, Figure 1) who was the first curator of the New York Botanical Garden herbarium [START_REF] Nybg | Per Axel Rydberg Records (RG4). Archives & Manuscripts[END_REF]. The species is the type of Aven Nelson's genus Eremohylema A.Nelson.

Pluchea sericea has also ethnobotanical applications, and among Native Americans it was used both as material for building mud houses and to make arrows [START_REF] Moerman | Native American Ethnobotany Database[END_REF]. Its roots have therapeutic properties to treat diarrhea, and indigestion as well as eye and dermatological inflammations. However, this species does not have any documented wide ethnobotanical use as compared to P. lanceolata in India.

Both Pluchea sericea and P. lanceolata are shrubs/ subshrubs with white sericeous branches and sessile lanceolate leaves. Their floral parts are similar as well, with pink to purplish phyllaries, campanulate involucres, and pink to purplish corollas. Th ese two species mainly differ in their growth habit, with P. sericea reaching up to 5 m in height (Barkley et al. 1993), while P. lanceolata usually reaches 1.5 m in height (King-Jones 2001). Th ey also diff er slightly in their leaves, with P. sericea characterized by narrower leaves (1-5 x 0.2-1 cm) than P. lanceolata (1-7.5 x 0.4-1.5 cm).

Molecular phylogenetics

Pluchea has been circumscribed to comprise taxa lacking the characteristic features of other smaller genera in the tribe Inuleae. Using fl oral micro-characters, Anderberg (1991b) found that Pluchea is not monophyletic. Th e genus as a whole lacks any diagnostic apomorphic characters; instead, it is defi ned by absence of characters found in related groups.

King-Jones's ( 2001) monograph for the Old World species of Pluchea included a phylogenetic analysis based on morphological data. Th is previous study suggested that P. lanceolata belongs to a monophyletic group with seven species that she informally named the "Pluchea indica Clade" and that was weakly supported with a low jackknife value of 63%. Additional taxonomic relationships pertinent to Berthelotia have been suggested by [START_REF] Nesom | New species, new sections, and a taxonomic overview of American Pluchea (Compositae: Inuleae)[END_REF][START_REF] Nesom | Pluchea[END_REF] to "Clade J" as recovered in Nylinder and Anderberg's (2015) molecular phylogeny; however, this study did not include P. lanceolata. As a result, we aimed to analyze relationships across New and Old World members of Clade J of Pluchea to better understand relationships between the two Berthelotia taxa with P. indica and monotypic Tessaria as putative relatives of them.

MATERIAL AND METHODS

We chose ITS nucleotide sequences of 19 species representing two main clades of the Inuleae uncovered in recent literature, as part of Clade J in Nylinder and Anderberg (2015). They belong to the genera Allopter-igeron, Coleocoma, Epaltes, Pluchea, Streptoglossa, Tessaria, and Thespidium, as our ingroups (including the two species described as Berthelotia as well as P. indica). Two species of Epaltes, which were found to be sister to Clade J (Nylinder and Anderberg 2015), are used as outgroup. For each species selected, ITS sequences were downloaded from GenBank (accessions in Table 1); all of them were previously published by Nylinder and Anderberg (2015) except that of P. lanceolata the unpublished one which was obtained by Rahul Pathirickal et al. in 2017. ETS data were not available for Pluchea lanceolata; therefore, we did not include this marker in our study.

Alignments were performed in MAFFT v.7 (Katoh and Standley 2013) for ITS. Matrices were visually inspected in order to check for false homologies. Phylogenetic analyses were performed across maximum likelihood and Bayesian methods. For the maximum likelihood analysis, the ITS gene tree was inferred for each of the individual gene alignments using RAxML v.8 (Stamatakis 2014), with 100 bootstraps (BS) using the GTRGAMMA model. For the Bayesian analysis, MrBayes was run for 1 million generations under the GTRGAMMA model, with 20% burn-in removed.

RESULTS AND DISCUSSION

The aligned length of the concatenated 21-sample, ITS matrix was 844 bp. For our Bayesian analysis, we recovered strong phylogenetic signal in the ancestral nodes of the ingroup, with low support for the clade including the members of Tessaria and Pluchea from both the New and Old World (68% posterior probability, P.P.). An additional clade with strong posterior probability support, identifi ed here as "Clade I", was recovered. Clade I is composed of three species of the Pluchea indi-ca Clade (i.e., P. indica, P. bojeri, and P. lyciodes) plus P. dioscoridis and P. ovalis (Figure 8). Th e latter two species were placed inside the "Pluchea discoridis clade" by [START_REF]Revision of Pluchea Cass. (Compositae, Plucheeae) in the Old World[END_REF]. Allopterigeron fi lifolius, P. caroliniensis and P. sericea, formed a polytomy with the clade that has the rest of the species included in our analyses (Figure 8). Clade I exhibited equally high support (100% P.P., Figure 8) and the lineage composed of P. sagittalis and Coleocoma centaurea was also strongly supported (90% B.S.). Th erefore, our analyses did not support the Pluchea indica Clade as a monophyletic group as defi ned by King-Jones's (2001) treatment. Maximum likelihood analysis of our ITS dataset showed similar results to our Bayesian analyses, with Clade I members resolved with high bootstrap support (99 B.S.). Almost all of the other shallower nodes in the ingroup had very low support.

In comparison to previous phylogenetic studies of the Inuleae and Pluchea (Anderberg 1991b; King-Jones 2001; Nylinder and Anderberg 2015), we found similarly low resolution at shallow nodes, however, our inclusion of P. lanceolata here reveals that: (1) the two species of Berthelotia are perhaps more distantly related than pre- As indicated above, the monotypic genus Tessaria is placed in a weakly supported clade (68% P.P.) that has all the sampled species of Pluchea, except P. carolinensis (Jacq.) G. Don and P. sericea (Figure 8). If future studies confirm these relationships with greater phylogenetic resolution, this will have a profound impact on the taxonomy of the group, as the name Tessaria (described in 1794) holds precedent over the younger name Pluchea, which was only described in 1817. Tessaria also has priority over Berthelotia (described in 1836) and the other genera of Clade J (Allopterigeron, described in 1981; Colecoma, described in 1857; Epaltes, described in 1818, Thespidium, described in 1862; and Streptoglossa, described in 1863). Even if a large number of Pluchea species were renamed within a larger Tessaria, the latter genus would still likely be polyphyletic under current taxonomic definitions, and because of the phylogenetic placement of other genera that compose the Inuleae; therefore, this would require extensive nomenclatural modifications in order to have a species-rich monophyletic Tessaria (Nylinder and Anderberg 2015). Within the J Clade several ways of dealing with these generic mismatches can be proposed once nodes of these clade are better resolved; one of them would involve reinstating genera such as Berthelotia as described by [START_REF] Candolle | Prodromus Systematis Naturalis Regni Vegetabilis[END_REF] and creating new monophyletic genera (likely most of them with few species). Such a solution could lead to consider B. sericea as a monotypic genus, in that case it will need to be treated as Eremohylema sericea. With Tessaria currently restricted to T. integrifolia, another solution would be for Tessaria to be expanded more broadly to include the two species listed within Berthelotia (P. sericea and P. lanceolata) and the other members of the J Clade.

ETHNOBOTANY AND MEDICINAL USE OF PLUCHEA LANCEOLATA

The ethnobotanical use of Pluchea lanceolata in India has its roots in the ancient Ayurveda, a natural system of medicine still widespread in the country (Figure 2), which originated more than 3,000 years ago and was part of the Vedic sacred literature of India [START_REF] Padma | [END_REF]. The age of Ayurveda has been traced back Table 1. Species included in the ITS phylogenetic analyses, along with their GenBank accession numbers, and their voucher specimen numbers (with herbaria acronyms). All sequences published by Nylinder and Anderberg (2015) between 4500 and 600 BC [START_REF] Kapoor | CRC Handbook of Ayurvedic Medicinal Plants[END_REF]). The Ayurvedic plant naming structure does not have a binomial system of nomenclature as in Western botany, but it is based on polynomials, where one plant can be defined by a group of synonyms (one to fifty), each relating to its characters; such as morphology, habit, habitat, qualities, biological actions, therapeutic use, etc. Sometimes a synonym can be given to more than one plant resulting in a therapeutic "taxonomy" that helps an Ayurvedic physician in selecting a plant for its medicinal use [START_REF] Viswanathan | A brief introduction to Ayurvedic system of medicine and some of its problems[END_REF]. This has also led to confusions, as technically precise descriptions of plants are not given. Therefore, this system of having numerous names for a single medicinal plant does not provide any morphological features sufficient for establishing its botanical identity. India is a place of diversity in all its forms including use of various languages/dialects in different regions, many tribal practices and folklore medicines contributing to use more than one name for the same plant, and sometimes different plants are having same names in different states of the country.

Only five species of Pluchea have widespread use as medicinal plants (Shanker and Srivastava 2017: 35-39). Three of them are from the New World (i.e., P. carolinensis (Jacq.) G.Don, and P. odorata (L.) Cass., P. sagittalis (Lam.) Cabrera), and at least one of them has been the subject of ethnobotanical field studies [START_REF] Hodges | The ethnobotany of Pluchea carolinensis (Jacq.) G.Don (Asteraceae) in the Botánicas of Miami, Florida[END_REF]. The aforementioned P. indica also has medicinal value in India and Southeast Asia (Shanker and Srivastava 2017: 35). Eleven additional species are also reported to have ethnobotanical interest, among them is Pluchea sericea (Shanker and Srivastava 2017: 35, 39).

Though plants of Pluchea lanceolata are commonly known as "rasna" in northern parts of India, they are also known by many other names in different regions of the country [i.e., rayasana, phaar (Hindi), rasnapat (Assam), chithramoolaka, dumme-rasna (Kannada), rasna, rashna, rayasana (Marathi), sanna rashtramu (Telugu), reshae, raasana, reshamee-sunnai, ra-sunna (Punjabi)]. The word "rasna" means "the process of tasting" or "perceptible by tongue". The species has been described in Sanskrit using many synonyms such as elaparni, mukta, rasna, rasya, sreyasi, sugandha, surabhi, surasa, suvaha, yukta. Consequently, the Ayurvedic text (Sanskrit) for rasna has been interpreted in different ways. The term "elaparni" could be interpreted for plants that have their leaf shape similar to the shape of Cardamom fruits (Elettaria cardamomum (L.) Maton), as considered by the northern Indian Ayurvedic physicians (Vaidyas). However, it can also be interpreted as plants bearing leaves whose shape is similar to the leaf shape of Cardamom, as it usually occurs in southern parts of India. The words "sugandha" and "suvaha" can refer to tubers having fragrance, which can be found in different species of plants. Thus, while the leaves of P. lanceolata are mostly used in northern India, the rhizomes of Alpinia galanga (L.) Willd. (Zingiberaceae) are widely utilized in the Southern parts of India. They both share the Ayurvedic name of rasna and have similar therapeutic properties. Notwithstanding the controversy regarding the identity of the species, the Ayurvedic Pharmacopeia of India (API 2001) has recognized P. lanceolata as the plant "rasna" whose leaves (Figure 2) and roots are recommended for preparation of various herbal formulations, and A. galanga as the approved substitute in the Ayurvedic Formulary of India (AFI 2003).

In practice, the leaves of Pluchea lanceolata are usually traded as rasna, rasnapatti, vaaya-surai, or baisurai, while its roots are mostly traded as rasnamoola. However, at least 17 other plant species have their parts traded under the name rasna in various parts of the country [START_REF] Viswanathan | A brief introduction to Ayurvedic system of medicine and some of its problems[END_REF][START_REF] Khare | Indian Medicinal Plants[END_REF]FRLHT Database 2016). A few examples of them include plants whose rhizomes or roots have medicinal use [e.g., Alpinia galanga, A. calcarata (Andrews) Roscoe, Leucoblepharis subsessilis Arn. (Asteraceae); and Vanda tessellata Roxb. (Orchidaceae)]; and others whose leaves exhibit similar therapeutic properties (e.g., Dodonaea viscosa (L.) Jacq., Sapindaceae). This medicinal plant classification system has made identification of the right material for Ayurvedic medicine preparation a daunting task. Some of the important Ayurvedic formulations containing rasna (P. lanceolata) are dasamularishta, devadarvarishta, karpasasthyadi taila, rasnadi kvatha churna, rasnairandadi kvatha churna. Some of these formulations are for ingestion (arishta/churna) and some are for external applications (taila/churna) [START_REF] Shanker | Monograph of Rasana (Pluchea lanceolata): An Important Medicinal Plant. Standarization and Validation of Key Ingredients of Ayurvedic medicines[END_REF]). However, drug formulations containing rasna (P. lanceolata) are all indicated for treatment of inflammation, rheumatism, asthma, fever, dyspepsia, skin diseases, rheumatoid arthritis, gout, etc. Other of the widespread ethno-botanical uses associated with P. lanceolata in India include its action as a laxative, nervine tonic, uterine relaxant, and pain relief from scorpion stings. In the former Sind region of British India, the species was known as koura-sana or koura-sunna. Its leaves were used as aperient medicine, and the roots were regarded as an excellent purgative [START_REF] Murray | The Plants and Drugs of Sind[END_REF][START_REF] Dymock | Pharmacographia Indica[END_REF][START_REF] Khare | Indian Medicinal Plants[END_REF]).

In the traditional Ayurvedic treatments and literature there are references for the use of Pluchea lanceolata in poly-herbal formulations as it was never recommended as a single herb. Nevertheless, there has been a steadily increasing trend for naturopathic herbal compa-nies to promote the use of single-herbs (such as rasna) that come from traditional Ayurvedic medicine. These companies are also making new combinations of Ayurvedic drug formulations, resulting in Ayurvedic medicinal patents that are associated with property rights (Patwardhan 2016). Products such as rasna churna (churna translates as powder), rasna capsules, and rasna tablets, which are not prescribed in the Ayurvedic classical medicine texts, fall in this category. They are marketed as herbal dietary supplements and advertised as having similar beneficial attributes as rasna formulations that have been used in India for millennia.

Typification notes

Berthelotia DC. is typified by B. lanceolata DC.; Nelson (1924: 55) erroneously treated Berthelotia as a later homonym (non Bertholletia Bonpl. 1807) and included Berthelotia as a synonym of Eremohylema A.Nelson, and thus the latter genus name was superfluous and illegitimate when published. Nevertheless, Nelson cited E. sericea (Nutt.) A.Nelson as the type of Eremohylema. Therefore, Eremohylema was published as the name of a new genus, has its own type, and is not typified by the type of Berthelotia (see Shenzhen Code Art. 7.5). Candolle (1836: 375-376) treated Berthelotia as a monospecific genus, and he included two varieties, viz., B. lanceolata DC. var. indica DC., and var. senegalensis DC. He did not include B. lanceolata DC. var. lanceolata or var. typica, or any such autonym. He did not cite any type collection for the species name B. lanceolata, but he did cite collections for vars. indica and senegalensis. His specific description for B. lanceolata was limited to a short text stating: ♄ frutic. pedalis. An species forsan duae admittendae?, which indicates Candolle's belief that the species was composed of two morphs (var. indica and var. senegalensis). Nevertheless, Candolle inadvertently created the autonym B. lanceolata DC. var. lanceolata (vide Art. 26.3).

In the above situation, one may question whether either var. indica or var. senegalensis is homotypic with var. lanceolata, and if so, whether such a varietal name is valid. In this regard, Art. 26.1 of the Code states that "The name of any infraspecific taxon that includes the type of the adopted, legitimate name of the species to which it is assigned is to repeat the specific epithet unaltered as its final epithet […]". It is ascertained here that since the species name B. lanceolata remained untypified at the time of its publication, neither var. indica nor var. senegalensis included the type of B. lanceolata, that Art. 26.1 does not apply here, and that the two varietal names were validly published. We add that theoretically, even if the names B. lanceolata and var. indica were typified with the same specimen and made homotypic, the validity of the varietal name indica would not be affected (cf.,Art. 26.2 Ex. 4).

Prior to our study, King-Jones (2001: 82) provided nomenclatural assessments for these varietal names; however, we do not concur with two of her conclusions, as addressed below. King-Jones's treatment did not include any discussion on the nomenclature of Berthelotia; therefore, her reasoning behind a few of her nomenclatural conclusions is not clear.

Regarding the typification of the name Berthelotia lanceolata var. senegalensis, Candolle referred to a gathering collected in Senegal (in Lampsar and Gandiole) by George S. Perrottet (1793-1870), a Swiss-French botanist who performed field work in this country from 1824 to 1829 (Stafleu & Cowan 1983: 175). One specimen housed in G (G00452584) fits the protologue quite well and is designated here as the lectotype for the variety. The protologue of this variety includes a reference to the name "Conyza proteiformis Perr. in litt.!" (sic). The published epithet "proteiformis" may imply resemblance to the habit of the genus Protea. The designated lectotype specimen sheet, however, shows Perrottet's annotation as "Conyza proteifolia nob.", which suggests that he intended to allude resemblance to foliage of Protea. It is evident that there was a typographical error of the epithet spelling, which was overlooked by Candolle in the publication. In our assessment, we treat "Conyza proteiformis" as a typographical error.

Furthermore, King-Jones' (2001: 82) conclusion pertaining to the typification of Berthelotia lanceolata needs to be revisited. She mentioned that Clarke (1876: 94) typified this name ("-Lectotype (designated by Clarke, Compos. Ind.: 94. 1876 No. 3029 longe discrepat, videl …" (sic). Clarke did not indicate the type element by direct citation of the term "type" (typus) or an equivalent and thus did not meet the mandatory requirement needed for a typification of a previously validly published name. (see Art. 7.11).

Although [START_REF]Revision of Pluchea Cass. (Compositae, Plucheeae) in the Old World[END_REF] cited a lectotype for the name Pluchea lanceolata, she did not inadvertently lectotypify the name because she did not use the phrase "designated here" (hic designatus) or an equivalent, which is required for typifications made from 1 January 2001 (Art. 7.11). Therefore, neither Clarke nor King-Jones typified the name, and we herewith designate the lectotype (G00450438; isolectotypes: K000974690, E00098190) of this taxon.

The protologue of Berthelotia lanceolata var. indica refers to three different specimens collected by the Danish physician and botanist Nathaniel Wallich or the Scottish physician and botanist Francis Buchanan-Hamilton (1762-1829). The former lived in India during 1814-1846 (Stafleu and Cowan 1988: 37), and most of his career was associated with the Royal Botanic Garden, Calcutta. Buchanan-Hamilton also resided in India (between 1795 and 1815), and worked for the Botanic Garden of Calcutta, but for a shorter period (1814-1815;Stafleu and Cowan 1979: 35). According to the protologue, these three specimens were annotated as "Conyza rubra Ham." (G00452579), "Conyza lanceolata Wall." (G00450438, G00450458), and "Saussurea mucronata Spreng." (P01816057). Of the preceding three names, S. mucronata was proposed as a pro synonym of B. lanceolata var. indica by Candolle (1836: 376) and is invalid. The two other names are found in Wallich Catalogue (http://wallich.rbge.info/node/13421) without a description; they were treated by Candolle (1836: 376) as synonyms of B. lanceolata var. indica.

Since Candolle used the specific epithet lanceolata to coin for his new species name B. lanceolata, and since he did not cite any collection for his new species, it is construed here that his inclusion of the name "Conyza lanceolata" as a pro synonym of var. indica implies that he intended to treat this variety as the typical variety of the species. Thus, among the three specimens suitable for lectotypification we have designated G00450458 as the lectotype of B. lanceolata and of B. lanceolata var. indica (isolectotypes: G00450438, K000974690, E00098190).

Polypappus sericeus

Typification notes

Polypappus sericeus was described by the British botanist and ornithologist Thomas Nuttall (1786-1859, Figure 1), who lived in USA during 1808-1841, and 1847-48 [START_REF] Beidleman | Some biographical sidelights on Thomas Nuttall, 1786[END_REF][START_REF] Graustein | Thomas Nuttall Naturalist. Explorations in America 1808-1841[END_REF], based on material collected in "Upper California, towards the Rocky Mountains" by botanist William Gambel (1823-1849), who actively collected especially in California and New Mexico (Graustein 1967: 342-344;Fischer, 2001: 26-27); Gambel also collected plant materials for Nuttall (Graustein 1967: 350).

Regarding Nuttall's herbarium and type specimens, Stafleu and Cowan (1981: 781) stated that "The Nuttall herbarium (5.750 species, acquired in 1860) is at BM. However, it would appear that up to the publication of the 'Genera [of North American plants]...' in 1818 he kept few or no specimens for himself, presenting a complete series of his plants to the Academy of Natural Sciences (PH) so that there repose his early types" (Pennell 1936). Further Nuttall specimens are found in CGE, DUKE, DWC, E, F, FI, G, GH, K, LIV, MANCH, MO, NY, OXF." Since Nuttall published the name Polypappus sericeus in 1848, it is reasonable to look for the Gambel's collection in the preceding herbaria. In our internet searches, we located only a single specimen, which is housed at the herbarium of the Royal Botanic Gardens, Kew (K000974653). Furthermore, in a personal communication, John Pruski (MO) informed us that he checked the PH microfiche for the name P. sericeus and did not find any specimen.

The Kew specimen, which we have identified as the original material, was examined and annotated by Nuttall. The specimen label shows the remark "I have only one bad[?] specimen" indicating that this was the only one specimen available for Nuttall to describe this species; the preceding remark also makes it clear that other herbaria do not have Gambel's collection for this species. Therefore, we recognize the K specimen as the holotype of P. sericeus Nutt. Two of the names (Tessaria borealis Torr. & A. Gray and Pluchea borealis A. Gray) that are relevant for our research were superfluous for Polypappus sericeus and illegitimate when published. Finally, "Berthelotia borealis Wooton", one of the names associated with the accepted Pluchea sericea, is not valid; [START_REF] Wooton | Trees and shrubs on New Mexico[END_REF] provided a description for the genus Berthelotia but not for the listed species, B. borealis.

APPENDIX 2.

Translation of the letter that Sabin Berthelot sent to Augustin Pyramus de Candolle on July 12, 1833. Notice that the last paragraph of this letter was written by Philip Barker Webb. Original letter is in French. Footnote bibliographies are listed within the Reference section of this contribution. Courtesy of Conservatoire et Jardin botaniques de Genève-Archives. Translation by Piero G. Delprete.

[Page 1] Dear Sir, In the hope of finding you in Paris, we are answering the interesting letter that you wrote at the eve of your departure from Geneva. When it arrived here, we learned with sadness that you left a few days before. We are sorry about this, which we are remembering as much unfortunate for us, and we are hasting to write to reply to your several questions and observations related to our Canarian Compositae.

The species about which you are asking some explanations and is with nº 3 bis in the collection of our radiate, is not cited in our catalogue and we have reasons to believe that this specimen cannot be other than a duplicate of nº 3, which following your announcement hereafter will be Berthelotia glaberrima 4 . We encountered this species only in Gran Canaria, in the volcanic peninsula called La Isleta 5 [and it] can be considered as the principal population, although in this locality is quite rare. This plant grows in this locality with Berth. sericea 6 , which is found in abundance. We never observed intermediary stages between the two species; the first one was always with glabrous leaves, green, almost lucid and a little fleshy, characters which make it remarkable even from a distance; whereas the other is constantly conserved on hers [its leaves] with silky hairs that distinguish it and a grayish-green. Unfortunately, this brilliant color that gives to Berth. glaberrima a physiognomy so characteristic is lost during desiccation [when plants are dried] and despite all our cares we were unable to prevent our specimens from blackening. Before passing to another subject, being the most inter- esting part, I should not finish this one without proper thanks. I accept with gratitude the dedication that you made for me for the new genus Berthelotia; it is unfortunate that the botanical laws oblige you in this case to maintain the old specific name sericea 7 , because villosa, or even pilosa would be more in agreement, as both these adjectives would have alluded (more divi Linnée) to the more evident external character of my individual [plant] and that we could have been referred to that [the leaf indumentum character] of the species type. But I know too well that your decisions are final judgements, therefore leave to me these reflection on the silky hairs of my species and make the case that it deserves. Mr. Webb will also be very charmed of seeing his name dedicated to a genus from Greece 8 ; you say that you have one [name] in reserve near Berthelotia; that one will establish a sort of confraternity between the two friends, they will be delighted of this friendship and they will feel glorified of being registered on the golden book under the auspices of such celebrated patron 9 . I can count on the gratitude of this new godson 10 . Regarding the other genus Webbia, which we have discussed in Geneva, it is true that Pavón 11 had the intention of publishing it and he also traced the characters in a manuscript; but certainly, the Spanish botanist published nothing on it 12 . My nº 6 is Erigeron inuloides Willd. 13 and the Chrysoco-7 It refers to Chrysocoma sericea L.f. (Asteraceae, Inuleae), which is the basionym of Schizogyne sericea (L.f.) DC. See note 6. 8 The genus Webbia DC. (Asteraceae, Vernonieae; nom. illeg., non Webbia Spach, Hist. Nat. Veg. Phan. 5: 408 June 1836) does not occur in Greece and it is restricted to Africa. Therefore, Augustin Pyramus de Candolle did not refer to an Eastern Mediterranean species when he described Webbia. See further details on the botanical history of Webbia in [START_REF] Francisco-Ortega | Honoring Philip Barker Webb: The three intriguing stories of Webbia as a genus name[END_REF], as under this name three different genera were described by three plant taxonomists independently. 9 It seems that in this sentence Berthelot is referring to a close association between Berthelotia (suggested in the letter as a Canary Island genus) and Webbia (suggested in the letter as a Greek genus) as both genera were described by Augustin Pyramus de Candolle. See additional details in note 8. 10 Berthelot seems to metaphorically "treat" the genus Berthelotia as a close family relative of him. . This is primarily a continental African species (Morocco, Cameroon, eastern Africa, South Africa) that is also found in the Canaries (La Gomera, Gran Canaria, Tenerife, and La Palma). See https://africanplantdatabase.ch/en/nomen/97596. 41 These two names refer to Conyza sericea (L.f.) W.T. Aiton and "Berthelotia sericea DC. " (both with accepted name Schizogyne sericea (L.f.) DC., Asteraceae, Inuleae). See note 6 as the name "B. sericea" was never published by Candolle. 42 The names "Conyza glaberrima" and "Berthelotia glaberrima" were never published; however, they both clearly refer to Schizogyne glaberrima DC. (Asteraceae, Inuleae). See note 4. 43 It refers to Phagnalon scariosum DC. ex Lowe (Asteraceae, Gnaphalieae), species published by the British botanist Richard Thomas Lowe (1802-1874) using the name and brief description provided by Augustin Pyramus de Candolle. This is a poorly known species that seems to be morphologically similar to Phagnalon rupestre (L.) DC. (Lowe 1868: 443) The latter is a native (no-endemic) species in the Canary Islands. 44 These two names refer to Conyza sordida L. (accepted name Phagnalon sordidum (L.) Rchb., Asteraceae, Gnaphaliea) and P. umbel-Gonospermum fruticosum 45 [Gonospernum] multiflorum 46

[Page 3] and finally, the Carlowizia xeranthemoides 47 . I still did not have decided anything on my method of publication. I have great need of your advice on this, and your departure from Paris at the moment when I arrived to profit from your advice is for me a fatality. Is it better to deal with the same publisher for the different parts that I wish to publish on the natural history of the Canaries so that the parts are published successively with the same format that will be the same of our drawings? And which is, according to you, the publisher that I should prefer? As I presume that you will not answer from Paris immediately, as you will be without doubt on the road towards Lugano at the reception of my letter, I am giving you the address of Mr. Webb in England. Farewell, my dear Mister, please convey my compliments to all your family and rest assured of my sincere attachment.

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Figure 1 .

 1 Figure 1. Portraits of relevant botanists involved in the botanical history of Berthelotia DC. (Asteraceae, Inuleae). (A) French naturalist Sabin Berthelot to whom Berthelotia is dedicated. (B) Swiss botanist Augustin Pyramus de Candolle who described Berthelotia and B. lanceolata (accepted name Pluchea lanceolata (DC.) C.B. Clarke). (C) British botanist Thomas Nuttall who described Polypappus sericeus (accepted name Pluchea sericea (Nutt.) Coville). (D) Swedish-American botanist Per A. Rydberg author of the combination Berthelotia sericea (Nutt.) Rydb. (A) Courtesy of the Museo Municipal de Bellas Artes, Santa Cruz de Tenerife, image kindly provided by Carlos Gaviño de Franchy. (B) Courtesy of the Royal Botanic Gardens, Kew. (C) Courtesy of Harvard Art Museums. (D) Courtesy of Hunt Institute for Botanical Documentation.

Figure 2 .

 2 Figure 2. Selection of photos pertinent to Berthelotia lanceolata DC. (accepted name Pluchea lanceolata (DC.) C.B.Clarke). (A) Shop where Ayurveda medicinal plants are commonly sold, Kozhikode, Kerala, India. (B) Flowering branch. (C) Leaf fragments as sold in medicinal plant shop. (D) View of capitula in full flowering through stereomicroscope. Photo credits: Rahul R. Pathirickal (A, C, D), and D.S. Rawat (B).

Figure 3 .

 3 Figure 3. Illustration of Berthelotia lanceolata DC. (accepted name Pluchea lanceolata (DC.) C.B.Clarke) published by Delessert and Candolle (1840). Drawn by Jean-Christophe Heyland (1791-1866) and engraved by Eugenia Taillant (?-?). Courtesy of the Linnean Society of London.

  , who indicated that the placement of Tessaria is relevant to understand the actual arrangement of Berthelotia in the Inuleae. Th e polyphyletic nature of the genus was further reiterated by Nylinder and Anderberg's (2015) molecular phylogenetic studies based on nucleotide sequences of ITS and ETS (nuclear ribosomal DNA) and three chloroplast genome regions. In this work, Pluchea species are found in several diff erent places in the recovered phylogeny, and many of the smaller genera of the tribe, such as the South American monotypic genus Tessaria Ruiz & Pav. (Anderberg 1994; Anderberg and Eldenäs 2007) became ingroups related to different Pluchea groups. Th erefore, Pluchea as presently circumscribed is a diverse polyphyletic grade which is poorly understood and needs major taxonomic revisions. Tessaria (one species; Tessaria integrifolia Ruiz & Pav.), Pluchea indica (L.) Less., and P. sericea belong

Figure 4 .

 4 Figure 4. Distribution map of Berthelotia lanceolata DC. (accepted name Pluchea lanceolata (DC.) C.B. Clarke). Map is based on King-Jones (2001) and was created using QGIS 3.26 in conjunction with base maps from MapTiler.

Figure 5 .

 5 Figure 5. Berthelotia sericea (Nutt.) Rydb. (accepted name Pluchea sericea (Nutt.) Coville) in USA. (A) Plants in habitat in Dead Valley National Park, California. (B) Informative panel highlighting the ethnobotanical use of the species among Native Americans and its importance as a landmark species for the landscapes of Dead Valley National Park. (C) Young stems showing capitula in early developmental stages. (D) Capitula at different developmental stages. Photo credits: Mike Duran (A, B), Ron Vanderhoff (C, D).

Figure 6 .

 6 Figure 6. Illustration of Berthelotia sericea (Nutt.) Rydb. (accepted name Pluchea sericea (Nutt.) Coville). Courtesy Flora of North America Association, Volume 19; Barbara Alongi, illustrator; published by Nesom (2006).

Figure 7 .

 7 Figure 7. Distribution map of Berthelotia sericea (Nutt.) Rydb. (accepted name Pluchea sericea (Nutt.) Coville). Map was created using QGIS 3.26, in conjunction with base maps from MapTiler

Figure 8 .

 8 Figure 8. Bayesian topology for Pluchea species and related genera belonging to Clade J sensu Nylinder & Anderberg (2015) with Epaltes divaricata and E. gariepina as the outgroup. Species underlined in red were previously described as belonging to the "Pluchea indica Clade" (King-Jones, 2001). Node labels refer to Posterior Probability scores.

  (=) Berthelotia lanceolata var. senegalensis DC., Prodr. 5: 376. 1836. (=) Conyza proteifolia Perr. ex DC., Prodr. 5: 376. 1836 ["proteiformis"], pro syn., inval. Type: [Senegal] "In arenosis circa Lamsar [Lampsar] et Gandiole mense junio legit cl. Perrottet. Conyza proteifolia ("proteiformis") Perr.! in litt. Folia infer. ex Perr. in litt. magis serrata. (v.s. comm. a cl. inv.)" (sic). Lectotype designated here: 15-June-1825, G.S. Perrottett 2 /Conyza proteifolia nob.: G [G00452584 (photo!)].

  Nutt., Proc. Acad. Nat. Sci. Philadelphia 4: 22. 1848. (≡) Tessaria borealis Torr. & A.Gray, Mem. Amer. Acad. Arts ser. 2, 4(1): 75, adnot. 1849, nom. superfl. & illegit. for Polypappus sericeus Nutt. (≡) Pluchea borealis A. Gray, Proc. Amer. Acad. Arts and Sci. 17: 212. 1882, nom. superfl. & illegit. for Polypappus sericeus Nutt.(≡) Pluchea sericea (Nutt.) Coville, Contr. U.S. Natl. Herb. 4: 128. 1893. (≡) Berthelotia sericea (Nutt.) Rydb., Bull. Torrey Bot. Club 33: 154. 1906. (≡) Eremohylema sericea (Nutt.) A.Nelson, Univ. Wyoming Publ. Sci., Bot. 1: 54. 1924. (≡) Tessaria sericea (Nutt.) Shinners, Sida 3: 122. 1967. Type: "In Upper California, towards the Rocky Mountains". W. Gambel Polypappus sericeus [/] Eastern California [/] I have only one bad [?] specimen [holotype, K: K000974653 (photo!)].

  except that of Pluchea lanceolata (MG273761) that is published by us in this contribution.

	Species	GenBank	Vouchers
	Allopterigeron filifolius (F.Muell.) Dunlop	LN607417	Short & Dunlop 4758 (DNA)
	Coleocoma centaurea F.Muell.	LN607432	Albrecht 10563 (S)
	Epaltes australis Less.	LN607444	Anderberg & Anderberg 7938 (S)
	Epaltes divaricata (L.) Cass.	LN607446	Bremer et al. 43 (S)
	Epaltes gariepina (DC.) Steetz	LN607447	Wanntorp & Wanntorp 769 (S)
	Pluchea bojeri (DC.) Humbert	LN607514	Malcomber & Leeuwenberg 1137 (S)
	Pluchea dentex Benth.	LN607516	Crawford 1076 (CANB)
	Pluchea dioscoridis (L.) DC.	LN607516	Ryding & Ermias 1279 (UPS)
	Pluchea dunlopii Hunger	LN607519	Hunger & Kilian 3948 (B)
	Pluchea fastigiata Griseb.	LN607520	Beck & Lieberman 9613 (S)
	Pluchea indica (L.) Less.	LN607524	Kilian et al. NK 4601 (S)
	Pluchea lanceolata (DC.) C.B.Clarke	MG273761	S.P. Geetha 8769 (CMPR)
	Pluchea lycioides (Hiern) Merxm.	LN607528	Smook 765 (WIND)
	Pluchea ovalis (Pers.) DC.	LN607534	Gilbert & Thulin 328 (UPS)
	Pluchea sagittalis (Lam.) Cabrera	LN607539	Chung & Anderberg 1171 (S)
	Pluchea sericea (Nutt.) Coville	LN607542	Davis & Lightowlers 66328 (B)
	Pluchea yucatanensisi G.L. Nesom	LN607546	Jones & Jones 12656 (TEX)
	Tessaria integrifolia Ruiz & Pav.	LN607592	Daly et al. 6392 (S)
	Thespidium basiflorum (F.Muell.) F.Muell.	LN607593	Cowie & Dunlop 3923 (S)

  Pluchea lanceolata (DC.) Oliv. & Hiern in Oliv.,Fl. Trop. Afr. 3: 329. 1877, isonym. (≡) Conyza lanceolata Wall.,Numer. List n. 2991. 1831, nom. nud., inval.

	(≡) Type: "Conyza lanceolata Wall.! Cat. et herb. n. 2991.
	comp. n. 101". N. Wallich 101, Conyza lanceolata Wall. in
	herb. / 1830 [Lectotype designated here, G [G00450458
	(photo!)]; isolectotypes, G [G00450438 (photo!)], K
	[K000974690 (photo!)], E [E00098190].
	(=) Conyza rubra Buch.-Ham. in Wall., Numer. List n.
	2991B. 1831, nom. nud., inval.
		TAXONOMY & NOMENCLATURE
	(=) Saussurea mucronata Spreng. ex DC., Prodr. 5: 376.
	1836, pro syn., inval.	Berthelotia DC. 2 (Asteraceae, Inuleae).
		Berthelotia DC., Prodr. 5: 375. 1836.
		(≡) Pluchea Cass. subgen. Berthelotia (DC.) A. Gray, Syn.
		Fl. N. Amer 1(2): 225. 1884.
		Type: Berthelotia lanceolata DC., Prodr. 5: 375-376.
		1836.
		(=) Eremohylema A.Nelson, Univ. Wyoming Publ. Sci.,
		Bot. 1(3-4): 54. 1924, nom. superfl. & illegit. for Berth-
		elotia DC.
		Type: Eremohylema sericea (Nutt.) A.Nelson, Univ. Wyo-
		ming Publ. Sci., Bot. 1(3-4): 54. 1924 [Basionyn: Polypap-
		pus sericeus Nutt., Proc. Acad. Nat. Sci. Philadelphia 4:
		22. 1848].
		(=) Pluchea Cass., Bull. Sci. Soc. Philom. Paris 1817: 31.
		1817.
		Type: Pluchea marilandica (Michx.) Cass. in F.Cuvier,
		Dict. Sci. Nat., ed. 2 42: 2. 1826.
		['marylandica'] [Basionym: Conyza marilandica Michx.
		Fl. Bor.-Amer. 2: 126. 1803].
		Berthelotia lanceolata DC., Prodr. 5: 376. 1836.
		(≡) Berthelotia lanceolata var. indica DC., Prodr. 5: 376.
		1836.
		(≡) Pluchea lanceolata (DC.) C.B.Clarke, Compos. Ind.:
		94. 1876.

  ):[India], Cawnpure [= Kanpur 26?28'N, 80?21'E], 1830, Wallich 2991/101 (E!, G!"). However, Clarke's treatment of this species shows "Pluchea lanceolata i.e., Berthelotia lanceolata DC. V. p. 376 Conyza lanceolata Wall. Cat. 2991 … Hab. Ad ripam Gangis prope Cawnpore in 1820 coll. Wallich. In omni Oude, India boreali-occidentali, Punjab et Afghanistan communis. Cl. Bth. in Gen. Pars. II. p. 291 Berthelotiam lanceolatam DC. cum Pluchea wallichiana DC. conjunxit. Berthelotia lanceolata est vera Pluchea sed Conyza lanceolata Wall. Cat. No. 2991 a Conyza sessilifolia Wall. Cat.

4

  It refers to Schizogyne glaberrima DC. (Asteraceae, Inuleae). The species was described by Augustin Pyramus de Candolle in the same work in which he described the genera Berthelotia and Webbia. Schizogyne Cass. (two species) is endemic to the Canary and Salvage Islands and was described by the French botanist Alexandre de Cassini in 1828. Candolle never described "B. glaberrima." Schizogyne glaberrima is endemic to Gran Canaria.5 It refers to the small peninsula of La Isleta located in the main city of the island, Las Palmas de Gran Canaria.6 It refers to Schizogyne sericea (L.f.) DC. (Asteraceae, Inuleae). The species is found in the Savage Islands and in the Canaries (all islands except Lanzarote; https://www.biodiversidadcanarias.es/biota/). Candolle did not ever describe "Berthelotia sericea". See additional details in note 4.

  11 It refers to the Spanish botanist José Antonio Pavón Jiménez (1754-1840) who together with Hipólito Ruiz López led one of the most important botanical endeavors of the Spanish Crown: The Expedición Botánica al Virreinato del Perú. Some of the specimens of this expedition reached Webb's herbarium[START_REF] Steinberg | The collectors and collections in the Herbarium Webb[END_REF].12 See[START_REF] Francisco-Ortega | Honoring Philip Barker Webb: The three intriguing stories of Webbia as a genus name[END_REF] regarding Pavón's plans to dedicate the genus Webbia to Webb.13 We did not find the name "Erigeron inuloides Willd. " (Asteraceae, Inuleae) in the consulted taxonomic literature, and we initially interpreted that Berthelot referred to Erigeron inuloides Poir. (accepted name Pulicaria inuloides (Poir.) DC.). The protologue of E. uniloides Poir. is for a perennial herbaceous species that was collected by Pierre M. A. Broussonet [see additional details on Broussonet in note 16] in the Canaries(Poiret 1817: 464). Unfortunately, no original material collected by Broussonet has been ma dichotoma 14 , Canaries, according to our research in the herbarium of Mr. Bouchet 15 . The label was written by him and dictated by Broussonet 16 .[Page 2] In the same herbarium we found only Carlina sulphurea 17 with the habitat of Tanger and with no species of this genus with the name of simplex. We also did not see any Carthamus rigidus 18 from Gibraltar cited by Broussonet 19 or by Durand 20 . The Carlina xeranthemoides 21 in the Brouss.[Broussonet] herbarium is only a variety with large leaves of Carlowizia salicifolia 22 . This plant is found in the collection of Mr. Bouchet with two labels written by him and found that allow us to properly interpret this name. Pulicaria arabica (L.) Cass. ssp. hispanica (Boiss.) Murb. is the only Canary Island native species of the genus to which this taxon could refer to. However, the text of this line of the letter can also be interpreted as Broussonet treating "Erigeron inuloides" as a synonym of Chrysocoma dichotoma L.f. We cannot rule this second possibility, because the account of C. dichotoma (as Allagopappus dichotomus (L.f.French botanist and director of the Botanical Garden of Montpellier between 1803 and 1807. He lived in the Canary Islands between 1799 and 1803 where he was appointed French Commissioner for Trade. During this period, he actively botanized in this archipelago. Between 1794 and 1795, he exiled to the Iberian Peninsula where he was supported by botanists and friends from Spain, Britain, and Portugal[START_REF] Motte | Broussonet (or Broussonnet or Broussounet)[END_REF][START_REF] Caillé | Un Savant Montpelliérain: Le Professeur Auguste Broussonet[END_REF]). 17 Carlina sulphurea Desf. (accepted name C. racemosa L., Asteraceae, Cardueae) is restricted to the Western Mediterranean (including Morocco) and does not reach the Canary Islands (López Martínez and Devesa 2014; http://www.floraiberica.es/PHP/generos_lista.php). by Broussonet. Mr. Bouchet confirmed that all the specimens of Carlina xeranthemoides 23 sent to other botanists by Broussonet, passed by his hands and they are all duplicates of the specimen that belongs to him. Therefore, our Carlina xeranth. [xeranthemoides] is not present in the Broussonet herbarium. In this case the herbaria of Linnaeus and Banks 24 could be useful in clarifying the doubts that we have about this species. The Centaurea calcitrapa 25 is indigenous of to the Canaries; we can find it very frequently in the field surrounding the cities and in almost whole of the maritime region. We can say the same for Galactites tomentosa 26 [tomentosus] and Silybum marianum 27 . Regarding Carthamus tinctorius 28 , this species is cultivated only in the Canaries [Canary Islands] and the Carthamus teneriffae Steud. 29 , which we do not know, is probably just a variety of C. tinctorius 30 caused by the climate and that some good German could have considered as an indigenous plant. During my visit at Montpellier, I made a drawing of your Gonospermum elegans 31 that probably I have in my herbarium as a variety, this drawing is the copy of that in the collection of the garden. If you wish to have it, I can send it to you, otherwise I will publish it with mine.Here is, more or less, all the information that I can supply on the several species that you mentioned in your letter; if you wish that I do other searches on this subject or on other plants, you only have to indicate this to me, and with pleasure I will try to satisfy your desire. I will remain here another fifteen days and then I will leave for London where I will stay for about one month to check the conditions of my collections and give a glance at the Banks 3223 See note 21.24 It refers to the herbarium of Sir Joseph Banks (1743-1820) who was the unofficial director of the Royal Botanic Gardens, Kew and one the most prominent figures in the history of science of his era. His herbarium is part of the collections of Natural History Museum of London (BM). for comparing the Canarian species collected by Masson 33 . I will then return to Paris to start working on my publications. I already finished the editing of several families and I hope that at my return to England Hayland34 will have almost finished the drawings of the plants that I gave him in Lyon, as well as of our more important Compositae that you would give him for this purpose. Here is the list: Centaurea canariensis Brouss. Willd 35 Carduus clavulatus 36 Centaurea cynaroides 37 Conyza canariensis Willd = Jasonia dichotoma DC. in Litt. 38 Buphthalmum laevigatum Brouss. = Jasonia laevigata DC. in litt. 39 Conyza gouanii Willd. 40 Conyza sericea = Berthelotia sericea DC. in litt. 41 Conyza glaberrima = B. glaber. DC. in litt. 42 Phagnalon scariosum 43 Conyza sordida n°1 bis in herb. Willd. B = Phagnalon umbelliforme DC. in litt. 44 33 It refers to the Scottish botanist Francis Masson (1741-1805), first official collector of the Royal Botanic Gardens, Kew who performed field work in the Azores, Canaries and Madeira between 1776 and 1779 (Francisco-Ortega et al. 2008). 34 It refers to the German artist Jean-Christophe Heyland (1791-1866). 35 It refers to Centaurea canariensis Brouss. ex Willd. (accepted name Cheirolophus canariensis (Brouss. ex Willd.) Holub, Asteraceae, Cardueae). This is a species endemic to Tenerife. 36 Carduus clavulatus Link (Asteraceae, Cardueae) is a species endemic to the Canary Islands. 37 It refers to Centaurea cynaroides Buch (accepted name Rhaponticum canariense DC., Asteraceae, Cardueae), a Tenerife endemic. 38 These two names refer to Conyza canariensis Willd. and Jasonia dichotoma (L.f.) DC. (accepted name for both of them: Allagopappus canariensis (Willd.) Greuter, Asteraceae, Inuleae, see note 14). 39 These two names refer to Buphthalmum laevigatum Brouss. ex Willd. and Jasonia laevigata (Brouss. ex Willd.) DC. (accepted name for both of them: Vieraea laevigata (Brouss. ex Willd.) Sch.Bip., Asteraceae, Inuleae). The genus Vieraea is monotypic and endemic to Tenerife. 40 It refers to Conyza gouanii (L.) Willd. (accepted name Erigeron gouanii L., Asteraceae, Inuleae)

		25 Centaurea calcitrapa L. (Asteraceae, Cardueae) is a Mediterranean
		species that reaches the Canaries where it is found in all islands except
		La Palma (Devesa 2014a).
		26 Galactites tomentosus Moench (Asteraceae, Cardueae) is a Mac-
		aronesian-Mediterranean species, it is found in all Canaries except
	18 Carthamus rigidus Willd., nom. illeg. (Asteraceae, Cardueae) refers to	Fuerteventura and Lanzarote (Devesa 2014b).
	Carduncellus arborescens (L.) Sweet a species restricted to the Western	27 Silybum marianum (L.) Gaertn. (Asteraceae, Cardueae) is a species with
	Mediterranean that does not reach the Canary Islands (López González	a widespread distribution that reaches the Canary Islands (Devesa 2014c).
	2014a; http://www.floraiberica.es/PHP/generos_lista.php).	28 Carthamus tinctorius L. (Asteraceae, Cardueae) has an unknown bio-
	19 See note 16.	geographical origin and it is cultivated in the Canaries (López González
	20 It refers to the French priest and botanist Philippe Durand (1795-	2014b).
	1815) who collected in southern Spain and Morocco. He worked in the
	Botanical Garden of Montpellier as Conservateur des Collections de
	Botanique (Veldkamp 2001; Michaud 2018).

) Cass.) presented by Schultz Bipontinus

(1844-1850: 224)

in

Webb and Berthelot'

s Histoire Naturelle des Iles Canaries lists Conyza inuloides Aiton as a synonym of A. dichotomus. There is a chance that Berthelot had an oversight and wrote "Erigeron inuloides Willd" instead of "Conyza inuloides Willd;" interestingly,

Willdenow (1803Willdenow ( : 1938) )

did list C. inuloides Aiton for the island of Tenerife. 14 It refers to Chrysocoma dichotoma L.f. (accepted name Allagopappus canariensis (Willd.) Greuter, Asteraceae, Inuleae). Allagopappus Cass. (two species) is endemic to the Canaries, with A. canariensis occurring in Gran Canaria, La Gomera, and Tenerife. See note 13 for additional details on the interpretation of this line of the letter.

15

It refers to the French botanist Dominique Bouchet (1770-1845) whose collections are housed at MPU (Herbier de l'Université de Montpellier II) and includes the herbarium of Pierre-Auguste-Marie Broussonett

(Stafleu and Cowan 1976: 290;[START_REF] Brun | Philip Baker & Sabino Berthelot: Una amistad a tres bandas[END_REF]

. See additional details on Broussonet in note 16.

16

Pierre-Auguste-Marie Broussonett (1761-1807) was a 21 Carlina xeranthemoides L.f. (Asteraceae, Cardueae) is a species endemic to Tenerife.

22

Carlowizia salicifolia (L.f.) Moench (accepted name Carlina salicifolia (L.f.) Cav., Asteraceae, Cardueae) is a species endemic to the Canary Islands and Madeira. dictated 29 This is a nomen nudum

(Steudel 1821: 164)

and we are not certain of the taxonomic identity of this species. It could refer to Carthamus lanatus L. (Asteraceae, Cardueae) or C. tinctorius L. 30 See notes 28 and 29. 31 Gonospermum elegans DC. (accepted name Gonospermum canariense Less, Asteraceae, Anthemideae) is endemic to La Palma and El Hierro. The genus is restricted to the Canary Islands and it has seven species. 32 See note 24.

herbarium

  48 I join my kinds regards to those of Berthelot, & should you have anything to make out in the herbaria of Linnaeus, Banks 49 or Lambert 50 beg leave to offer my humble services [illeg.] in London. My address is at my Brother's Milford House near Godalming, Surrey. I wish you a pleasant excursion to Italian Switzerland. Parolini 51 writes me that he shall send you hither our Greek Composite. P. B.W. liforme DC., respectively. The latter is a Canary Island endemic that occurs in all islands except Fuerteventura and Lanzarote. According to the protologue of P. umbelliforme, this specie is closely related to P. sordidum(Candolle 1836: 396), and this could explain why Berthelot listed Conyza sordida and Phagnalum umbeliforme together in this entry.45 It refers to Gonospermum fruticosum (Buch) Less. (Asteraceae, Anthemideae). This species is restricted to the islands of La Gomera, El Hierro, and Tenerife. See note 31 regarding biogeography of Gonospermum.46 It refers to Gonospermum multiflorum DC. (accepted name G. fruticosum (Buch) Less., Asteraceae, Anthemideae). See note 31 regarding biogeography of Gonospermum and note 45 on the distribution of this species in the Canary Islands.47 We have not found "Carlowizia xeranthemoides" in the taxonomic literature; however, this entry clearly refers to Carlina xeranthemoides L.f. (Asteraceae, Cardueae). See note 21.48 This part of the letter was written by Philip Barker Webb, and it is in English.49 See note 24.50 It refers to the British naturalist Aylmer Bourker Lambert (1761-1842) who was of the first fellows of the Linnean Society of London and owner of the largest private herbarium in Europe at that time[START_REF] Miller | The herbarium of Aylmer Bourke Lambert: Notes on its acquisition, dispersal, and present whereabouts[END_REF].51 It refers to the Italian botanist Alberto Parolini (1788-1867), a close friend of Webb to whom he dedicated the Canary Island endemic genus Parolinia (Brassicaceae), see further details in[START_REF] Francisco-Ortega | Honoring Philip Barker Webb: The three intriguing stories of Webbia as a genus name[END_REF].

Alexander von Humboldt (1769[START_REF] Berthelot | De l'acclimatation en Algérie des principales essences forestières des Îles Canaries[END_REF] and Aimé Bonpland (1773-1858) have been usually considered as the coauthors of the species name Viola cheiranthifolia; however, Bonpland alone is credited for the text of the two volumes of Plantae aequinoctiales (title page: "in Ordinem Digessit: Amatus Bonpland");Stafleu and Cowan (1979: 367, #3141) also mentioned that Bonpland is the author of the text. Therefore, the correct authorship for this species name is: Viola cheiranthifolia Bonpl. as indicated in the websites www.ipni.org and plantsoftheworldonline.org/

Honoring Sabin Berthelot: Nomenclature and botanical history of Berthelotia DC. (Asteraceae, Inuleae)

Accepted names indicated in bold font.

ACKNOWLEDGEMENTS

We dedicate this publication to Carlos Gaviño de Franchy in recognition for his contributions as publisher of books centered in the humanities of the Canary Islands. Nomenclature conclusions were part of the workshop and graduate course "Advanced Latin American and Caribbean Island Workshop on Plant Taxonomy and Nomenclature" (25-29 April 2022) jointly sponsored by Montgomery Botanical Center and the Kimberly Green Latin American and Caribbean Center of Florida International University (FIU), Miami and delivered by GC (Griffith et al. 2022). Javier Francisco-Ortega is grateful for the support received by the Montgomery Botanical Center to conduct botanical research projects in summer 2022. Our gratitude to Tammy Charron (Missouri Botanical Garden), Andrea Deneau (Linnean Society of London), Sylvie Rivet (Société de Géographie, France), Stephen Sinon (Hunt Institute for Botanical Documentation), and Jeff Steward (Harvard Art Museum) for their help in locating relevant bibliography and/ or historical images. Andrés Delgado (Asociación de Amigos del Museo de Naturaleza y Arqueología, Tenerife, Spain) helped with the layout of historical photos. Jean Philippe Chassot (G) kindly located for us relevant letters housed in the Conservatoire et Jardin botaniques de Genève-Archives. John Pruski (MO) checked PH microfiche for Gambel's collection of Polypappus sericeus Nutt., and confirmed the absence of original material at the PH herbarium. We thank Arne Anderberg (S) for his correspondence and advice on Inuleae systematics. Our gratitude to Guy L. Nesom (TEX) and Fred Stauffer (G) for critically reading the paper.