Adansonia33(2)-ENVOI2.indb 323ADANSONIA, sér. 3 • 2011 • 33 (2) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.adansonia.com MOTS CLÉS Asteraceae, Anthemideae, Santolina ageratifolia, nomenclature, taxonomie. KEY WORDS Asteraceae, Anthemideae, Santolina ageratifolia, nomenclature, taxonomy. Rivero-Guerra A. O. 2011. — Typifi cation and synonymy of Santolina ageratifolia Barnades ex Asso (Asteraceae, Anthemideae), an endemic species of the Iberian Peninsula. Adansonia, sér. 3, 33 (2): 323-330. DOI: 10.5252/a2011n2a15. ABSTRACT Patterns of speciation in the angiosperm genus Santolina (Asteraceae, An- themideae) have recently been intensively studied, but progress is hampered by nomenclatural confusion. In this paper, the species S. ageratifolia Barnades ex Asso, S. pectinata Lag. var. paniculata Loscos & J.Pardo and S. longidentata Pau are lectotypifi ed. An invalid name has been found: S. rosmarinifolia L. subvar. virens macrocephala Pau. Th e nomenclature of S. ageratifolia is provided. RÉSUMÉ Typifi cation et synonymie de Santolina ageratifolia Barnades ex Asso (Asteraceae, Anthemideae), une espèce endémique de la péninsule Ibérique. Les patrons de spéciation dans le genre d’angiospermes Santolina (Asteraceae, Anthemideae) ont été récemment étudiés intensivement, mais la progression des connaissances est entravée par la confusion nomenclaturale. Dans cet arti- cle, les espèces S. ageratifolia Barnades ex Asso, S. pectinata Lag. var. paniculata Loscos & J.Pardo et S. longidentata Pau sont lectotypifi ées. Un nom invalide a été découvert : S. rosmarinifolia L. subvar. virens macrocephala Pau. La nomen- clature de S. ageratifolia est détaillée. Aixa O. RIVERO-GUERRA Universidad de Sevilla, Facultad de Biología, Departamento de Biología Vegetal y Ecología, Avda. Reina Mercedes, 41012 Sevilla (Spain) and Centro de Investigação en Biodiversidade e Recursos Genéticos, Campus Agrario de Vairão, Rua Padre Armando Quintas-Crasto, 4485-661 Vairão (Portugal) rivero-guerra@hotmail.com Typifi cation and synonymy of Santolina ageratifolia Barnades ex Asso (Asteraceae, Anthemideae), an endemic species of the Iberian Peninsula 324 ADANSONIA, sér. 3 • 2011 • 33 (2) Rivero-Guerra A. O. INTRODUCTION Th e genus Santolina belongs to the subtribe San- tolininae Willk., tribe Anthemideae Cass., family Compositae Giseke, alternatively named Asteraceae Dumort. Recent investigations have focused on reconstructing phylogeny and interpreting bio- geography of the tribe Anthemideae (Francisco- Ortega et al. 1997; Oberprieler & Vogt 2000; Watson et al. 2000, 2002; Oberprieler 2002, 2004a, b, 2005; Oberprieler et al. 2009). In the course of these evolutionary studies, it has also been judged helpful to revise certain taxa of this tribe in need of taxonomic attention. Such is the case with Santolina, a genus endemic to the Medi- terranean Region. Santolina ageratifolia Barnades ex Asso (2n = 6x = 54), a member of the S. rosmarinifolia L. aggregate (Rivero-Guerra 2011) with the highest ploidy level, is an endemic species of Teruel Province, Spain, of restricted geographical distribution: Ródenas, Cerro San Ginés, and Cerro del Ardal (Almohaja) (Rivero-Guerra 2008). Th is species lives on sand- stone and red limolite (Ródenas population), and quartzite (Cerro San Ginés). Pau (1907a) characterised S. ageratifolia and cited S. rosmarinifolia Asso and S. longidentata Pau as its synonyms. Th e same author in 1895, changed the concept and cited “– S. rosmarinifolia b) heterophylla subv.) virens macrocephala Pau, not. bot., fasc. 2, pag. 28. – S. pectinata var. paniculata? Losc. et Pard.?” as synonyms of S. longidentata. Mateo Sanz (1990) suggested that S. ageratifolia and S. longidentata are synonyms of S. rosmarini- folia L.; however, López Udías et al. (1997) cited S. rosmarinifolia subvar. virens macrocephala Pau, S. longidentata Pau and S. rosmarinifolia auct., non L. as synonyms of S. ageratifolia. Th is study was undertaken to improve our knowl- edge of the taxonomy of the genus Santolina and to improve its nomenclature. A search for suitable lectotypes of S. ageratifolia, S. pectinata var. pan- iculata Loscos & J.Pardo, and of S. longidentata Pau, as well as the historical background and the nomenclatural discussion of these names, has been conducted in the literature and in relevant herbaria, and these results are presented below. HISTORICAL BACKGROUND Asso y del Río (1779) in Synopsis Stirpium Indige- narum Aragoniae, p. 116, published Santolina ros- marinifolia auct., non L. (1753) with the following notation: “805. Santolina rosmarinifolia // Abrotanum femina, virens, vermiculato, serrato folio Barrel. Ic. 464. // Provenit propè Rodenas // Planta suff ruticosa, pedalis. Caules virgati, unifl ori, supernènudi. Folia linearia, acuta, carnosa, duplici denticulorum serie utrinquè instructa. Flores duplò majores quàm in praecedenti. Paleae concavae”. Th is name is the homonym of the validly published S. rosmarinifolia L., and is there- fore illegitimate and unavailable for use under the Article 53.1 of the International Code of Botanical Nomenclature (McNeill et al. 2006). Asso y del Río adopted the Linnaeus’ name “S. rosmarinifolia” for a taxon of the same rank, with explicit exclusion of the original type. Th e same author (1779) men- tioned the fi gure 464 of Barrelier in the protologue of S. rosmarinifolia sensu Asso. Pau (1907b) explained the origin of the name S. ageratifolia. Asso (1784) mentioned the Synopsis Stirpium Indigenarum Aragoniae in the protologue of S. ageratifolia, and changed the number “805” into “850”: “Santolina rosmarinifolia Syn. n 850 // Species à nobis circa pagum Rodenas observata S. rosmarini- folia varietas videbatur. Eandem in herbario Cl. Bar- nades nomine Santolina ageratifolia postea invenimus, unde novam speciem esse perspicue apparuit. Caules in nostra, ut in Exemplari Barnadiano, simplices; folia linearia, carnosa, duplici denticulorum serie in utroque margine”. He attributed the name S. ageratifolia to Barnades, apparently because this name was used by Barnades in the label of the voucher specimen of his herbarium. However, there is no indication that Barnades validly published the name. Th e no- men novum and its diagnosis are validly ascribed to Asso y del Río (1784), under the Article 46.4 of the ICBN (McNeill et al. 2006). Th e same author, in Historia de la Economía política de Aragón (1798) emphasized that S. ageratifolia is a novel taxon “y en Ródenas una especie nueva de Santolina, que el difunto D. Miguel Barnades puso en su herbario con el nombre trivial de ageratifolia”. Th e present study is based on collections from the herbaria reported by Stafl eu & Mennega (1992), 325 Typifi cation and synonymy of Santolina ageratifolia Barnades ex Asso (Asteraceae, Anthemideae) ADANSONIA, sér. 3 • 2011 • 33 (2) where the material used by Asso y del Río (some material in P) and Barnades (BC, BM, C, LINN, MA) are deposited, and on all Spanish herbaria that contain collections of Santolina. Specimens of S. ager- atifolia gathered by Pau (BC 30129, BM 909683, UCLA, JE, W) and by Vicioso (MA 126817) were found. Ibáñez Cortina (2006) showed three speci- mens of the genus Santolina (which classifi ed as S. chamaecyparissus) in the collection of Barnades at the BC herbarium. Th e study of Barnades’s col- lection indicates that these specimens do not fully match the diagnostic characteristics of the genus Santolina, and none of them are from the locus classicus of S. ageratifolia. Th e same is true of the S. chamaecyparissus specimen fi gured by Barnades showed in his book “Herbarium pictum hispanicum” (six volumes). Th ese specimens should be excluded from Santolina. Loscos y Bernal & Pardo y Sastrón (1866-1867) classifi ed as “S. rosmarinifolia L. ex Asso” the speci- mens from “en el soto del Cañar”, gathered by Echeandia and indicated in brackets “acaso S. pec- tinata?”. De Jaime Lorén (2005) explained that Loscos y Bernal probably studied the Asso y del Río’s material in the “Sociedad Económica Aragonesa”. I could not fi nd the specimens from the locus clas- sicus of S. ageratifolia in the Echeandia, Schultz, Schule, Vallier and Loscos y Bernal’s collections of the private herbarium of the “Sociedad Económica Aragonesa de Amigos del País, IberCaja, Zaragoza”, and in Loscos y Bernal and Zapater’s collections of the herbarium of the “Instituto de Educación Secundaria “Vega del Turia”, Teruel”. Th e main objects of this study are to: 1) desig- nate a lectotype of S. ageratifolia, S. pectinata var. paniculata Loscos & J.Pardo, and of S. longidentata Pau, and 2) to determine all the names available as synonyms of S. ageratifolia. TYPIFICATION 1. Santolina ageratifolia Barnades ex Asso Introductio in Oryctographiam, et Zoologiam Aragoniae accedit. Enumeratio stirpium in eadem noviter detectarum, n. 805: 179 (1784). TYPUS (as given in the protologue). — “circa pagum Rodenas”. LECTOTYPUS (designated here). — fi g. 464 (Abrotanum femina, virens, vermiculato, serrato folio) of Plantae per Galliam, Hispaniam et Italiam of Barrelier (1714: p. 117) / tab. Viii.; fi g. 3 of Osservazione quarantissima prima – L’origine, e prima impressione di alcune piante marine, imperfette, come fuchi, corralline zeophite, fungi terrestri, e simili” of Schrockius (1697). EPITYPUS (designated here). — Teruel, Ródenas, 40°38’87’’N, 1°31’12’’W, 1400 m, conglomerates, sand- stone and red limolite, 15.VII.1998, A. O. Rivero-Guerra s.n. (SEV 277730, Fig. 1). REMARKS Th e fi gure 464 (Abrotanum femina, virens, vermicula- to, serrato folio) of Plantae per Galliam, Hispaniam et Italiam of Barrelier (1714: p. 117; http://bibdigital. rjb.csic.es/spa/Libro.php?Libro=1394) shows similar morphological characteristics of S. ageratifolia. Bar- relier (1714: p. 91) wrote: “1019. Santolina foliis Rosmarini, major Inst. R. Herb. 461. Abrotanum foemina, virens, vermiculato, serrate folio Barr. Icon. 464 & Bocc. Muss. Part. 1 tab. Viii. Fig. 3. Abrotanum foemina, foliis Rosmarini, majus C.B. Pin. 137”. Th e author’s quotation “Bocc. Muss. Part. 1 tab. Viii. Fig. 3” refers to Schrockius (1697), Osservazione quarantissima prima – L’origine, e prima impressio- ne di alcune piante marine, imperfette, come fuchi, corralline zeophite, fungi terrestri, e simili, p. 262- 273, pl. V-VIII in Boccone P. Museo di fi sica e di esperienze variato, e decorato di osservazioni naturali, table VIII (before p. 271: http://bibdigital.rjb.csic. es/spa/Libro.php?Libro=4631). Th e fi gure 3 of this plate is the counterpart of Barrelier’s fi gure 464. Probably the material used to illustrated this fi gure is from the south of Italy. However, S. ageratifolia is endemic of the Iberian Peninsula. Unfortunately, Asso y del Río’s and Barrelier’s herbaria have ap- parently been lost or destroyed, and duplicates of the original material were not distributed to major herbaria; as a consequence, no known duplicate material exists. Furthermore, there is no herbarium known for Schrockius. Th e lectotype selection is based on Barrelier/Schrockius-Boccone fi gure, due to the fact that the ICBN does not allow to exclude original material, when no other original material is known. 326 ADANSONIA, sér. 3 • 2011 • 33 (2) Rivero-Guerra A. O. 2. Santolina pectinata Lag. var. paniculata Loscos & J.Pardo Series inconfecta plantarum indigenarum Aragoniae praecipue meridionales: 54 (1863). TYPUS (as given in the protologue). — “Prope Aranda del Conde hanc formam memorabilem legit cl Calavia”. LECTOTYPUS (designated here). — Without date, loc., and collector (private herbarium of “Sociedad Económica Aragonesa de Amigos del País, IberCaja, Zaragoza”). ISOLECTOTYPUS (designated here). — “Prope Aranda del Conde in Aragonia, Calavia, without date and collector (COI-WILLK 00035957). REMARKS The lectotype is the sole original material for this name of the private herbarium of “Sociedad Económica Aragonesa de Amigos del País, IberCaja, Zaragoza”. It has several depauperate fl owering stems. Th e lectotype has a label with the following handwritten notation by Loscos y Bernal in black ink: “Santolina / pectinata var. paniculata”. Th e volume I of the book Francisco Loscos y Ber- nal (1823-1886). Un Botánico Aragonés (Muñoz Garmendia & González Bueno 2001) gathers the correspondence between Loscos y Bernal and other botanists, especially with Willlkomm. Willkomm (letter date October 20, 1862) and Boissier (letter date September 12, 1863) requested some Spanish plants from Loscos y Bernal, including S. pectinata and S. pectinata var. paniculata respectively. None specimen of S. pectinata var. paniculata is conserved in G, but one specimen is preserved in COI-WILLK herbarium. Th e isolectotype (COI-WILLK 00035957) has a sample with several fl owering stems without leaves. Th e sheet has three labels with the following notations in black ink. Label 1 (in the lower left- hand corner): “Herbarium Willkommi [printed] // Santolina pectinata Bth. [underlined] / var.? paniculata [underlined] Losc. Pard. // Prope Aranda del Conde in Aragonia [Willkomm’s handwritten notation] // Legit [printed] Calavia [Willkomm’s handwritten notation]. Label 2 (in the lower left- hand corner): “Herb. Univ. Coimbra // *COI 00035957*”. Label 3 (red) [to right of the speci- men]: “TYPUS”. Th e two specimens display all the characteristics listed in the protologue: “diff er a specie caulibus paniculato-ramosis”. Furthermore, Willkomm in Willkomm & Lange (1870) with regards of the diag- nostic characteristics of this name indicated: “Rami fl oriferi paniculato-ramosi polycephali. Pedunculi eximie angulati et incrassati. An species propria?” Loscos y Bernal & Pardo y Sastrón (1866-1867: 205-206) wrote: “1041 S. pectinata Benth., Lag. forma canescens; cerca de Jaca, Forest. Jul. // b. pan- iculata Losc. Pard. Ser, incof., Wk. Lge. Prodr. (an sp. propria?) Abunda alrededor de Aranda, Calavia”. Willkomm in Willkomm & Lange (1870) cited S. pectinata Lag. var. paniculata Loscos & J. Pardo as “S. pectinata Bth. β. paniculata Losc. Pard.”. However, S. pectinata Benth. (Bentham 1826) is the homonym of the validly published S. pectinata Lag. (Lagasca 1816), and is therefore illegitimate and unavailable for use under the Article 53.1 of the ICBN (McNeill et al. 2006). Th e lectotype and isolectype of S. pectinata var. paniculata indicate that this name is a synonym of S. chamaecyparissus L. sensu stricto. Th is suggests that Loscos y Bernal and Pardo y Sastrón used the name S. chamaecyparissus L. for a more inclusive taxon than recent studies. 3. Santolina rosmarinifolia L. subvar. virens macrocephala Pau, Notas botánicas de la Flora española 2: 28 (1888) (nom. inval. cf. Art. 23.1 of ICBN). LECTOTYPUS. — Not cited. REMARKS Pau (1888), in the protologue of S. rosmarinifolia subvar. virens macrocephala, wrote: “Junto á Albar- racín y en Griegos (Zapater). // No conozco esta forma, y si S. canescens Lag. de Sierra Nevada, de la que se aparta, lo mismo que de S. rosmarini- folia L.”. Pau published S. rosmarinifolia subvar. virens macrocephala on the basis of the specimen collected by Zapater (MA 126816). Th is name is not a binary combination of the name of the genus followed by a single specifi c epithet, and the two words of the epithet are not hyphenated (Article 327 Typifi cation and synonymy of Santolina ageratifolia Barnades ex Asso (Asteraceae, Anthemideae) ADANSONIA, sér. 3 • 2011 • 33 (2) FIG 1. — Epitype of Santolina ageratifolia Barnades ex Asso. 328 ADANSONIA, sér. 3 • 2011 • 33 (2) Rivero-Guerra A. O. 23.1 of the ICBN [McNeill et al. 2006]). Th e two subvarietal terms (virens and macrocephala) are adjectives in the nominative case; such unhy- phenated terms, if in adjectives, are not allowed to be hyphenated. Santolina rosmarinifolia subvar. virens macrocephala Pau has no status under ICBN, however, it has been fully cited as a synonyms of S. ageratifolia without alterations from the origi- nally published form. 4. Santolina longidentata Pau Notas botánicas de la Flora española 6: 60 (1896). TYPUS (as given in the protologue). — “Sierra de Albar- racín, en Griegos (Zap.)”. LECTOTYPUS (designated here). — Albarracín, Griegos, Zapater s.n. (MA 126816). REMARKS Th e lectotype (MA 126816) has seven fragments of the fl owering stems, most of them without leaves. Th e sheet has three labels in the lower right-hand corner with the following notations in black ink. Label 1: “Caroli Pau Herbarium hispanicum [printed and underlined] // Santolina longidentata Pau / (notas bot. fas. 6) p. 28, n. 103 / (1888) // Albarracín (Griegos) // Zapater legit [Pau’s handwritten notation]. Label 2: Santolina rosmarinifolia b) heterophylla subv. virens mac- rocephala Pau, Not. Bot. Fasc. 2: 28 (1888) = / S. longidentata Pau, Not. Bot. Fl. Esp. 6: 60 (1895) // Me parece que esta planta, herborizada por Reverchon en Griegos, Hervier se lo comunicó a Rouy, que la dio como especie nueva y le dio la localidad en el Pirineo, al confundir Griegos con Gedre // C. Pau” [Pau’s handwritten notation]. Label 3: Label of determination of E. Guinea: JARDÍN BOTÁNICO DE MADRID [printed] // Santolina pectinata [underlined] Lagasca // 3 IX 1966 [E. Guinea’s handwritten notation in rose ink]. Th is specimen displays all the characteristics listed in the protologue: “Cabezuelas grandes, semiesféricas, no umbilicadas, solitarias, hojas del tallo parecido á la Achillea microphylla W.”. DISCUSSION Asso y del Río (1779, 1784) considered that plant habit, leaf shape, leaf apex, leaf incision and lobe insertion were good diagnostic characteristics, but in 1784 he included plant colour as a new diagnostic character. In contrast, López Udías et al. (1997) used the presence/absence of plant indument, leaf incision, and insertion of the appendage of the outer bracts to diff erentiate S. ageratifolia from the remaining taxa of the S. rosmarinifolia aggregate. Rivero-Guerra (2011) demonstrated that leaf shape, leaf incision, and lobe insertion are variable between individuals of a given population; therefore, they are not good taxonomic characters. Th e shape of the leaf apex has no taxonomic value except to diff erentiate S. impressa, S. orocarpetana (Rivero- Guerra in press) and S. × oblongifolia nothosp. from the remaining taxa of S. rosmarinifolia aggregate. Plant habit and colour diff erentiate S. ageratifolia from the remaining taxa of the S. rosmarinifolia ag- gregate. Th e receptacle is conical in S. rosmarinifolia subsp. arrabidensis, S. ageratifolia, and S. elegans exclusively. Th e appendage of the involucral bract is dark copperish and fragile in S. ageratifolia, whereas in the remaining taxa, it is hyaline and robust. Th e plants of S. rosmarinifolia subsp. rosmarinifolia, S. ageratifolia, S. rosmarinifolia subsp. arrabidensis (69.09% of the individuals), and S. × oblongifolia (26.81% of the individuals) are glabrous, whereas the remaining taxa are tomentose, glabrescent, tomentose to glabrescent or sericeous. Finally, S. agerati folia does not have viscose glands, except on the fl owers and interseminal bracts, whereas the plants of the remaining taxa have viscose glands. Study of the morphological variation of the S. ros- marinifolia aggregate (Rivero-Guerra 2011) suggests strong relationships between the subspecies of S. pec- tinata (pectinata and montiberica) and S. ageratifolia, but the morphological diff erences between those three taxa are greater than those within S. pectinata. Th e study also suggests that S. ageratifolia probably derived from S. pectinata. Th is work demonstrated that S. pectinata var. paniculata Loscos & J.Pardo is not a synonym of S. longidentata, as Pau (1895) suggested. It is a synonym of S. chamaecyparissus L. sensu stricto. Th e 329 Typifi cation and synonymy of Santolina ageratifolia Barnades ex Asso (Asteraceae, Anthemideae) ADANSONIA, sér. 3 • 2011 • 33 (2) results indicated that S. longidentata Pau is synonym of S. ageratifolia as Pau (1907a) and López Udías et al. (1997) suggested. Th e invalid name S. ros- marinifolia subvar. virens macrocephala Pau has no status under the ICBN; however, it has been cited as synonym of S. ageratifolia. Th ese clarifi cations can help future studies on speciation patterns and process in Santolina and, by extension, in angiosperms in general. Th e fol- lowing full synonymy is accepted. Santolina ageratifolia Barnades ex Asso Introductio in Oryctographiam, et Zoologiam Aragoniae accedit. Enumeratio stirpium in eadem noviter detectarum n. 805: 179 (1784). TYPUS (as given in the protologue). — “circa pagum Rodenas”. LECTOTYPUS (designated here). — fi g. 464 (Abrotanum femina, virens, vermiculato, serrato folio) of Plantae per Galliam, Hispaniam et Italiam of Barrelier (1714: p. 117) / tab. Viii. Fig. 3 of Osservazione quarantissima prima – L’origine, e prima impressione di alcune piante marine, imperfette, come fuchi, corralline zeophite, fungi terrestri, e simili” of Schrockius (1697). EPITYPUS (designated here). — Teruel, Ródenas, 40°38’87’’N, 1°31’12’’W, 1400 m, conglomerates, sand- stone and red limolite, 15.VII.1998, A. O. Rivero-Guerra s.n. (SEV 277730; Fig. 1). Santolina rosmarinifolia L. subv. virens macrocephala Pau, Notas botánicas de la Flora española 2: 28. (1888) (nom. inval. cf. Art. 23.1 of ICBN). — Lectotype not cited. Santolina longidentata Pau, Notas botánicas de la Flora española 6: 60 (1896). — Type (as given in the proto- logue): “Sierra de Albarracín, en Griegos (Zap.)”. — Lectotype (designated here): Albarracín, Griegos, Zapater s.n. (MA 126816). Santolina rosmarinifolia sensu Asso, Synopsis Stirpium Indigenarum Aragoniae n. 805, 116 (1779), non L., Species Plantarum II: 842 (1753). Acknowledgements Th e study of the Santolina collection of the Natu- ral History Museum of Vienna was supported by Synthesys Project: AT-TAF-3669. I am most grateful to herbaria: BC, BM, UCLA, JE, W, and MA for supplying vouchers via loan, to the staff of Sociedad Económica Aragonesa de Amigos del País, IberCaja, Zaragoza, to Josefi na Cabarga for their cooperation, to Dr. M. Laurin for comments and help with the English version of the manuscript, to Dr. C. Romero Zarco for his helpful comments and suggestions in revising a pre-submission draft of the manuscript, to Dr. J. 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