Systematic Botany (2015), 40(3): pp. 900–913 © Copyright 2015 by the American Society of Plant Taxonomists DOI 10.1600/036364415X689320 Date of publication September 22, 2015 A Taxonomic Revision of the Eurasian/Northwestern African Senecio doria Group (Compositae)

Joel Calvo1,2 and Carlos Aedo1

1Real Jardín Botánico-CSIC, Plaza Murillo 2, 28014 Madrid, Spain. 2Author for correspondence ([email protected])

Communicating Editor: Chuck Bell

Abstract—The taxonomic complexity of the Senecio doria group (Compositae, Senecioneae) is reflected in the heterogeneous treatments that have been proposed and the poor knowledge of some species. This species group consists of perennial herbs distributed in Europe, western and central Asia, and northwestern Africa. The first worldwide revision of this group recognizing seven species is presented here (i.e. S. altissimus, S. bithynicus, S. doria, S. fontanicola, S. legionensis, S. morisii,andS. umbrosus). In this new taxonomic treatment S. legionensis and S. fontanicola are recognized at the specific rank. On the other hand, S. macrophyllus from eastern Europe/western Asia has been synonymized to S. doria. Senecio bithynicus and S. morisii are described as new species. Eleven names are lectotypified. Descriptions, distribution maps, and an identification key are provided for all the species included. Keywords—Asteraceae, lectotypification, new species, Senecio section Doria, taxonomy.

The Senecio doria group (Compositae, Senecioneae) is a uted throughout northwestern Africa (Morocco), Europe taxonomically complex assembly of taxa centered around (lacking in the north), and western Asia (Caucasus, north- S. doria L. This group belongs to S. sect. Doria (Fabr.) Godr., western Turkey, and Aralo-Caspian region), growing from sea which comprises between 14 and 24 taxa according to the level to 2,300 m. Over time, many classifications have been available regional works (Roldugin 1966; Matthews 1975; proposed for this group, although a comprehensive taxo- Wiebe 1997; Greuter 2008; Yilin et al. 2011). Along with S. sect. nomic revision was hitherto still needed. Senecio, S. sect. Crociseris (Rchb.) Boiss., and S. sect. Jacobaea The concept of the S. doria group was indirectly established (Mill.) Gray, section Doria is one of the four main sections of by Chater and Walters (1976), who treated all related taxa to western Eurasian/northern African Senecio species that are S. doria as subspecies (i.e. subsp. doria, subsp. kirghisicus (DC.) classically recognized (Willkomm 1865; Boissier 1875; Chater, subsp. legionensis (Lange) Chater, and subsp. umbrosus Wissjulina 1962; Chater and Walters 1976; Konechnaya 1994; (Waldst. & Kit.) Soó). Before them, Schischkin (1961) and most Menitsky and Konechnaya 2008). Recent phylogenetic studies of the central European authors who worked on this group carried out by Pelser et al. (2007) and Calvo et al. (2013) (Reichenbach 1853; Nyman 1879; Hegi 1928) recognized suggest that species of S. sect. Doria and S. sect. Crociseris most taxa of the group as different species. Concerning the (Rchb.) Boiss. form a clade, and therefore that both sections placement of S. kirghisicus DC. within S. doria as proposed should be synonymized. However, only four sect. Doria spe- by Chater and Walters (1976), recent studies (Calvo et al. cies were included in the analyses. To firmly establish the 2011, 2014a, 2015) suggest that it shares close morphological phylogenetic relationships between both sections a molecular similarities with S. racemosus, and therefore, it is best placed study including all the sect. Doria species is required. To this in S. sect. Crociseris. aim, the limits and the species composition of S. sect. Doria In the geographic frame of the Iberian Peninsula, Pérez have to be further studied. Morales et al. (1989) followed a narrow concept of S. doria The widespread S. nemorensis L. and its allied taxa is another s. s. and they differentiated four taxa: S. doria, S. laderoi Pérez controversial group within section Doria. It is distributed Morales, M. E. García & Penas subsp. laderoi, S. laderoi subsp. from western Europe to central Asia and northeastern Asia cantabricus Pérez Morales, M. E. García & Penas, and (i.e. Mongolia, China, and Japan according to Yilin et al. S. legionensis Lange. Likewise, Grulich and Hodálová (1994) 2011), and it includes highly variable taxa with numerous carried out a taxonomic revision of the group in central and populations displaying intermediate forms (Hodálová 1999; southeastern Europe. It represents the only recent revision of Oberprieler et al. 2010). Senecio sarracenicus L. (= S. fluviatilis the group in that region. The authors recognized S. doria, Wallr.) is another species belonging to section Doria that has S. macrophyllus M. Bieb., S. umbrosus Waldst. & Kit., and they a wide distribution (Europe, eastern Anatolia, central Asia, described S. fontanicola Grulich & Hodálová as a new species. central Siberia) and it should be thoroughly revised along its More recently, Greuter (2008) accepted only four species in whole range. A poorly known species, S. racemulifer Pavlov., the circum-Mediterranean countries: S. doria, S. legionensis, occurs in the Tashkent Region (Uzbekistan). The architecture S. macrophyllus,andS. umbrosus.However,theauthornoticed of the involucres and the number of ligulate florets (8–10) the uncertain taxonomic position of some taxa (S. fontanicola, lead us to tentatively consider it a member of S. sect. Doria. S. doria subsp. laderoi (Pérez Morales, M. E. García & Penas) These are some examples that highlight the need to revise Blanca), and he tentatively included them within S. doria.Chater the taxonomy of the remaining sect. Doria species to clarify and Walters (1976) also underlined the doubtful status of some the relationships between sect. Doria and its allied relatives. plants from Spain with tomentose-lanate indumentum. Like- The S. doria group includes rhizomatous herbs with leaves wise, the circumscription of S. doria and S. macrophyllus is not undivided and decreasing in size up the stem, involucre clear and the geographical delimitation of both species differs with supplementary bracts, and yellow radiate capitula with according to the authors (Grulich and Hodálová 1994; Greuter 5–10 ligulate florets. The achenes are subcylindrical with 2008). All these reasons highlight that S. doria group remains pappus, glabrous or with indumentum. The group is distrib- poorly known and thus it requires a thorough revision.

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The aim of the research presented here is to revise the Taxonomic Treatment S. doria group and to propose a modern and useful taxonomic The main morphological characters revealed as useful for treatment, including updated nomenclature, descriptions, dis- distinguishing between species are the number and shape of tributions maps, drawings, and an identification key. supplementary bracts, length of involucral bracts, number of lig- ulate florets, achene indumentum, and the shape of basal leaves. Senecio doria Group—Rhizomatous perennial herbs. Stem Materials and Methods 0.5–2 m, solid or fistulous, glabrescent to covered with long multicellular trichomes (sometimes slightly arachnoid), green- Around 420 dried specimens were studied from the following ish or with the lower half brick-red coloured. Basal leaves 8– herbaria: BC, CAG, FI, G, GE, HGI, LEB, LOU, MA, MT, and SAV. Only – photographs of specimens or additional information were available from 46 cm long, 1.8 18 cm wide, oblanceolate, linear-lanceolate to other institutions: BEO, BM, BP, CL, COI, KW, LE, LJS, P, PAD, PO, widely lanceolate, elliptic or ovate, obtuse to acute, attenuate PRC, SOM, and TO. to rounded (sometimes decurrent along the petiole), entire A comprehensive synonymy of the S. doria group is compiled. Types to distantly dentate, glabrous to arachnoid-floccose beneath, of all accepted names and most synonyms have been studied, although – we had difficulties while locating the type of the name S. umbrosus var. with a petiole 2 24 cm long slightly widened at the base. Cau- subtuberculatus Borbás ex Formánek. line leaves narrowly lanceolate to ovate or pandurate, sessile A general description of the whole group and a description for each to amplexicaul. Capitula 6–350, 12–24 mm diam; involucre species with the relevant characters are provided. Qualitative characters cylindrical-campanulate or subcylindrical; involucral bracts were studied directly by eye or with the aid of binocular lenses, while (10–)12–13(–16), 3.7–8.2 mm long, 0.7–2.6 mm wide, linear to the quantitative characters were recorded using a Mitutoyo digital caliper, CD-15DC. The most frequent values are given by percentiles and lanceolate, acute, glabrous to weakly arachnoid; supplemen- are shown without parentheses, and the extreme values are included in tary bracts (1–)3–5(–10), 1.2–5.1 mm long, 0.3–0.7 mm wide, parentheses. Habitat, elevation, and flowering period are compiled from linear to linear-triangular or slightly widened at the base, herbarium labels. a quarter to a half as long as involucral bracts, glabrous to The leaf length provided in the descriptions corresponds to the lamina weakly arachnoid (rarely slightly floccose). Ligulate florets length, and the number of supplementary bracts indicated is the number – – – – of bracts inserted at the base of the involucre. Those appearing on the 5 8( 10), 6 18.2 mm long. Achenes 2.9 4.5 mm long, sub- stem near the involucre are considered stem bracts of the synflorescence. cylindrical, glabrous or hairy.

Key to the Species of SENECIO DORIA Group 1. Cauline leaves pandurate, arachnoid-floccose beneath, ± discolorous; supplementary bracts 4–5.1mmlong...... S. bithynicus 1. Cauline leaves narrowly lanceolate to ovate-elliptic, glabrous to arachnoid beneath, concolorous; supplementary bracts 1.2–3.7mmlong...... 2 2. Achenes with intercostal trichomes ...... 3 3. Involucral bracts 3.7–5.3 mm long; supplementary bracts (4–)5–8(–10) ...... S. altissimus 3. Involucral bracts 5.2–5.9 mm long; supplementary bracts (1–)2–5 ...... S. legionensis 2. Achenes glabrous or with some trichomes concentrated at the summit and base ...... 4 4. Involucral bracts 3.9–6.1mmlong...... 5 5. Ligulate florets 5(–6); supplementary bracts 3–5(–6),linear...... S. doria 5. Ligulate florets 6–8; supplementary bracts (4–)5–8, slightly widened at base ...... S. morisii 4. Involucral bracts 5.4–8.2mmlong...... 6 6. Basal leaves ovate to widely lanceolate, 7.5–18 cm wide; ligulate florets 8(–10); stem usually hairy ...... S. umbrosus 6. Basal leaves oblanceolate to narrowly lanceolate, 2–5.5 cm wide; ligulate florets 5–7; stem glabrous ...... S. fontanicola

1. Senecio altissimus Mill., Gard. Dict. ed. 8, n.° 9. 1768. Senecio laderoi subsp. cantabricus PérezMorales,M.E.García& Senecio carnosus Lam., Fl. Franç. 2: 131. 1779, nom. illeg. Penas [“cantabrica”], Stud. Bot. Univ. Salamanca 8: 124. superfl. (McNeill et al. 2012, ICN Art. 52.1).—TYPE: 1989.—TYPE: SPAIN. Palencia: Valcobero, Arroyo del Herb. Miller n.° 319 (lectotype, designated here: BM- Hornillo, 30TUN54 [42°51′N 4°46′W], 18 Sep 1988, 1125235 (digital image!)). F. Gómiz s. n. (holotype: LEB-26987 (digital image!)). Senecio doria var. canescens Porta, Atti Imp. Regia Accad. Senecio doria var. incanescens Lange, nom. nud., in sched. Roveretana ser. 2, 9: 135. 1891.—TYPE: SPAIN. (McNeill et al. 2012, ICN Art. 38.1). Granada: ad rivulos et aquaeductos circa la Sagra, [37°58′N2°32′W], 1890, P. Porta & G. Rigo 562 (lecto- Senecio doria var. iserniana Rivas Goday, nom. nud., in sched. type, designated here: BM-1025977 (digital image!); iso- (McNeill et al. 2012, ICN Art. 38.1) (MA-245334!, MT s. n.!). lectotypes: K-844192 (digital image!), K-844193 (digital Senecio doria var. lanuginosus A. Blanco [“lanuginosa”], nom. image!), G-442487!). nud., in sched. (McNeill et al. 2012, ICN Art. 38.1) Senecio laderoi Pérez Morales, M. E. García & Penas, Stud. (P-3674479 (digital image!)). Bot. Univ. Salamanca 8: 124. 1989. Senecio doria subsp. Senecio doria var. pseudoauricula Rivas Goday, nom. nud., in laderoi (Pérez Morales, M. E. García & Penas) Blanca, sched. (McNeill et al. 2012, ICN Art. 38.1) (MA-200056!). Lagascalia 18: 308. 1996.—TYPE: SPAIN. León: Alto de Casares, 30TTN75 [42°56′N 5°46′W], 18 July 1988, Senecio pseudodoria Schur [“pseudo-doria”], nom. nud., in C. Pérez Morales & E. Puente s. n. (holotype: LEB-40600 sched. (McNeill et al. 2012, ICN Art. 38.1) (K-844190 (digital image!)). (digital image!)). 902 SYSTEMATIC BOTANY [Volume 40

Stem 0.6–1.5 m, solid, glabrescent to covered with scattered 3–5(–6), linear in S. doria] are also distinctive characters, as multicellular trichomes (sometimes slightly arachnoid towards well as the indumentum of the involucre (usually ± arach- the base), greenish. Basal leaves 9–37 cm long, 3–13 cm wide, noid in S. altissimus vs. glabrous or nearly so in S. doria). oblanceolate to widely elliptic, obtuse to acute, cuneate to Senecio altissimus overlaps its range with S. legionensis in attenuate, entire to distantly dentate, glabrescent or covered northern León Province (northwestern Spain). The number with scattered multicellular trichomes (sometimes ± arachnoid, of supplementary bracts [(4–)5–8(–10) in S. altissimus vs. (1–) more densely beneath), with a petiole 2–10 cm long slightly 2–5inS. legionensis], the length of the involucral bracts (3.7– widened at the base. Cauline leaves lanceolate to ovate, semi- 5.3 mm in S. altissimus vs. 5.2–5.9 mm in S. legionensis), and amplexicaul, concolorous. Capitula 24–350, 14–18 mm diam; the width of the basal leaves (3–13 cm in S. altissimus vs. involucre cylindrical-campanulate; involucral bracts 12–13, 1.8–6.1 cm in S. legionensis) are useful characters to discrimi- 3.7–5.3 mm long, 0.7–1.9 mm wide, linear to lanceolate, acute, nate each other. glabrescent to weakly arachnoid near the base; supplementary The provenance of the material that Schur invalidly bracts (4–)5–8(–10), 1.9–2.9 mm long, 0.4–0.6 mm wide, linear, named S. pseudodoria (in sched.) is uncertain. It undoubtedly slightly widened at the base, a quarter as long as involucral corresponds to S. altissimus because of its hairy achenes. bracts, glabrescent to weakly arachnoid. Ligulate florets 5(–6), Representative Specimens Examined — FRANCE. Alpes-de-Haute- ′ ′ 7.9–10.4 mm long. Achenes 2.9–3.9 mm long, subcylindrical, Provence: Basses Alpes, Digne, 44°5 N6°13E, 27 June 1892, F. Norris s. n. (GE). Aude: Narbonne, la Clappe, à la fontaine du Rég, 43°9′N3°7′E, with intercostal trichomes. Figure 1A. 1929, M. Chassagne s. n. (P). Bouches-du-Rhône: Aix-en-Provence, 43°33′N Additional Illustration—Pérez Morales et al. (1989: 5°27′E, 11 July 1964, G. Gavelle s. n. (MA). Hautes-Alpes: Orcières, 44°41′N 125–126, Figs. 3–4). 6°19′E, July 1984, G. Sag 12709 (G, MA). Hérault: bords du Lez, à Montpellier, ′ ′ Chromosome Number—2n = 40 (Löve and Kjellqvist 1974, 43°37 N 3°53 E, 5 June 1894, fr. Sennen 401 (P). Var: Fontaine l'Evêque, 43°43′N 6°11′E, 11 July 1964, G. Gavelle s. n. (MA). sub S. doria). ′ ′ — ITALY. Lazio: Poggi Palanzana, 42°24 N 12°8 E, July 1885, L. Macchiati Distribution and Habitat Senecio altissimus is found in s. n. (FI). Piedmont: vallis del Mastalone, 45°50′N8°12′E, G. Biroli s. n. (TO). France, Italy (including Sicily), Morocco, and Spain. This spe- Sicilia: Madonie, 37°52′N 14°1′E, 1847, A. Franqueville s. n. (P); Caltagirone, cies grows in damp meadows, edges of streams and ditches, 37°14′N14°31′E, June 1893, H. Ross 31 (FI, G, GE, P). ′ ′ reed beds, grassy places, shrubs, and edge of woods of MOROCCO. Fès-Boulemane: Immouzer, ad aquas, 33°27 N 4°17 W, – July 1913, M. Mouret 1471 (P). Meknès-Tafilalet: Moyen Atlas, Timhadit, Quercus, Pinus, Populus, etc., between elevations of 0 2,300 m bords du Guigous, 33°14′N 5°4′W, 10 July 1924, E. Jahandiez 796 (G, P). (Fig. 2). Souss-Massa-Drâa: Amesker de abajo, al N de El Kelâa-des-Mgouna, Phenology—Flowering from May to October. 31°29′N 6°14′W, 4 July 1997, F. Muñoz Garmendia et al. 5323 (MA). Tadla- Notes—The taxonomic and geographic delimitation of Azilal: Takreda, S.-E. de la prov. de Ntifa; S.-E. de la prov. de Demnat, 31°39′N 6°22′W, 30 June 1882, A. Ibrahim 2930 (G, MT, P). S. doria s. s. has been confusing over time, in part because SPAIN. Albacete: à Riopar, 38°30′N 2°25′W, 24 July 1850, E. Bourgeau the distribution indicated by Linnaeus (1763) was completely 725 (G, P). Barcelona: llano de Vich, 41°56′N 2°16′E, July 1884, F. Trèmols vague “Habitat in Oriente, Austriae sylvis, Monspelii ad s. n. (MA). Burgos: Santo Domingo de Silos, pie de peña de Cervera, Ladi ripas”. This conjuncture led the botanists from central/ 41°57′N 3°24′W, 17 July 1979, A. Susanna et al. 361 (B, G). Girona: Alt ′ ′ eastern Europe and from southwestern Europe (France, Italy, Empordà, Riumors, a les Roquetes, 42°13 N3°2E, 3 July 1996, P. Gesti s. n. (HGI). Granada: Sierra Nevada, ad aquas pr. Guejar, 37°9′N 3°25′W, and Spain) to use the name S. doria for their respective 25 July 1879, R. Huter et al. 1045 (G, P). Guadalajara: Molina de Aragón, populations. Our study highlights the morphological differ- salida del pueblo dirección Rillo, pr. el Lavadero, 40°51′N 1°54′W, ences between the mentioned groups and the need to separate 20 Aug 2014, J. Calvo & É. Ross-Nadié 6792 (MA). Jaén: Barranco de ′ ′ them. It is also strongly supported by their distant geographic Valentina, 37°53 N 2°50 W, July 1904, E. Reverchon 1172 (MA, P). León: Barrios de Luna, Vega de Caballeros, orilla río Luna, 42°48′N 5°49′W, areas. Since the populations from central/eastern Europe have 24 July 2009, J. Calvo 3962 (MA). Navarra: prope Yesa in Navarra, in been traditionally considered as S. doria in its narrowest sense, valle fluvii Aragon et alibi in Navarra, 42°36′N 1°11′W, June 1850, we proposed to maintain this name for such entity (Calvo M. Willkomm 264 (G, P). Soria: pr. Abejar, 41°48′N 2°47′W, 29 July 2014, et al. 2014c). Consequently, we retrieve Miller’s name J. Calvo & É. Ross-Nadié 6724 (MA). Teruel: Valacloche, 40°11′N 1°5′W, S. altissimus for the populations from southwestern Europe July 1893, E. Reverchon 847 (G, GE, P). and Morocco, which have been recently treated as S. doria 2. Senecio morisii J. Calvo & Bacch., sp. nov.—TYPE: ITALY. subsp. laderoi in some regional works (Blanca 2002; Blanca Sardinia: Seui, Anulù, 39°51′N9°21′E, 19 June 2013, and Quesada 2009). G. Bacchetta & A. Delage s. n. (holotype: MA!; isotype: CAG!). Although the species described by Miller came from culti- – vated plants, the author explicitly stated in the protologue Stem 0.5 1.5 m, solid, glabrescent (with remnants of that S. altissimus grows naturally in France. We found a spec- arachnoid indumentum on the upper part when young), – – imen in Miller’s herbarium (BM-1125235) that perfectly greenish. Basal leaves 12 34 cm long, 3.2 7cmwide, matches the characters of the plants growing in southwest- oblanceolate, obtuse to acute, attenuate, subentire to distantly – ern Europe. The mentioned sheet is here designated as lecto- dentate, glabrescent, with a petiole 5 12 cm long slightly type of the name S. altissimus. widened at the base. Cauline leaves lanceolate, semi- Senecio altissimus is characterized by its achenes with inter- amplexicaul, concolorous. Capitula 20–55, 13–16 mm diam; costal trichomes. The leaf and stem indumentum is variable involucre cylindrical-campanulate; involucral bracts 12–13, along its distribution area, and therefore, it is not a valuable 4.4–5.7 mm long, 0.9–1.8 mm wide, linear to lanceolate, acute, character to differentiate infraspecific taxa. The presence of glabrescent or with scattered arachnoid trichomes; supple- intercostal trichomes over the achene is a character only mentary bracts (4–)5–8, 2.4–3.7 mm long, 0.3–0.7 mm wide, shared with S. legionensis, although the indumentum in the linear, slightly widened at the base, a third to a half as long as latter species is clearly sparser. This character is useful to dis- involucral bracts, with scattered arachnoid trichomes (some- tinguish S. altissimus from S. doria s. s., which has glabrous times weakly floccose). Ligulate florets 6–8, 6–8.9 mm long. achenes. The number and shape of the supplementary bracts Achenes 3.2–3.8 mm long, subcylindrical, glabrous. Figure 1B. [(4–)5–8(–10), slightly widened at the base in S. altissimus vs. Chromosome Number—unknown. 2015] CALVO AND AEDO: TAXONOMY OF SENECIO DORIA GROUP 903

Fig. 1. A. Senecio altissimus. a1. Capitulum. a2. Supplementary bract. a3. Inner involucral bract. a4. Outer involucral bract. a5. Ligulate floret. a6. Achene with pappus [Based on Segura Zubizarreta s. n. (MA)]. B. Senecio morisii. b1. Capitulum. b2. Supplementary bract. b3. Inner involucral bract. b4. Outer involucral bract. b5. Ligulate floret [Based on Arrigoni & Nardi s. n. (FI)]. b6. Achene with pappus [Based on Moris s. n. (FI)]. C. Senecio doria. c1. Capitulum. c2. Supplementary bract. c3. Inner involucral bract. c4. Outer involucral bract. c5. Ligulate floret. c6. Achene with pappus [Based on Vitek & Walter 13–0417 (MA)]. 904 SYSTEMATIC BOTANY [Volume 40

Fig. 2. Distribution map for Senecio altissimus (open circle), S. legionensis (triangle), and S. morisii (closed circle).

Distribution and Habitat—This species is only known basis of our field investigations, this species is rare and from CE Sardinia (Italy). It grows along the edges of streams scattered; only six populations are currently known. in woods of Ostrya carpinifolia Scop. accompanied by Taxus Representative Specimens Examined — ITALY. Sardegna: Laconi, fra ′ ′ baccata L. and Ilex aquifolium L., on calcareous soils (lime- Corona Sa Guardia e Murtedu, 39°51 N 9°3 E, 27 May 1968, S. Alias s. n. (FI); regione Corongiu Irau (Villanovatulo), 39°48′N 9°11′E, 15 June 1963, stone, travertine, and conglomerate), between elevations of P. V. Arrigoni s. n. (FI); Seui, foresta Montarbu, pendici sopra il Rio 700–1,200 m (Fig. 2). Baccu 'e Pira andando al Fundu de Tonneri, 39°52′N9°23′E, 27 June Phenology—Flowering from late May to July. 1972, P. V. Arrigoni & E. Nardi s. n. (FI); Laconi, Rio Bau Onu, 39°52′N ′ Notes—Until now, this species has been identified as 9°5 E, 1 July 2000, G. Bacchetta s. n. (CAG); Laconi, Bau Crabone, 39°52′N9°5′E, 1 July 2013, G. Bacchetta s. n. (CAG); Laconi, Funtanamela, S. doria (Pignatti 1982; Conti et al. 2005). However, the achene 39°52′N9°5′E, 15 July 2009, G. Bacchetta & C. Dessì s. n. (CAG); Rio del morphology and the number of ligulate florets allow us to Taquisara, 39°51′N 9°28′E, 27 May 1879, A. Biondi s. n. (FI); Sardinia, separate it from S. altissimus and S. doria. Although Moris Ch. Guebhard 292 (P); Sardinia centralis, juxta rivulos Laconi, 39°50′N (1840–1843) also identified the Sardinian populations as 9°3′E, G. G. Moris s. n. (FI); circondario di Aritzo, 39°57′N 9°11′E, 1935, ′ ′ S. doria, he noticed the peculiarity of its glabrous achenes: G. Porru s. n. (FI); Laconi, Su Lau, 39°50 N9°4E, 5 June 1988, A. Scrugli & L. Mura s. n. (CAG). “Akenia glaberrima: caetero cum vulgari stirpe omnino ” congruit [Glabrous achenes: the remaining characters 3. Senecio doria L., Syst. Nat. ed. 10, 2: 1215. 1759, nom. & completely match the common form]. It is noteworthy that typ. cons. prop. (Calvo et al. 2014c). Jacobaea doria (L.) ’ “ ” Moris s concept of common form referred to S. altissimus, G. Gaertn., B. Mey. & Scherb., Oekon. Fl. Wetterau 3(1): which displays hairy achenes. We name this new species 214. 1801. Doria vera (L.) Fourr., Ann. Soc. Linn. Lyon – S. morisii in honor of Giuseppe Giacinto Moris (1796 1869), ser. 2, 16: 404. 1868.—TYPE: AUSTRIA. Lower Austria: Italian botanist who worked as a professor at the University 0.5 km SW of church of village Stopfenreuth, N fringe of – of Cagliari from 1822 1829. the meadow “Brücklwiese”, N of dam, 48°08′N 16°52′E, This species is morphologically similar to S. altissimus and 3 Sep 2013, E. Vitek & J. Walter 13–0417 (Typ. cons. S. doria. Senecio morisii differs from the former in its achene prop.: W!; isotypes: B!, BC!, C!, E!, G!, K!, L!, M!, MA!, indumentum (glabrous vs. hairy in S. altissimus), and in the MO!, NY!, US!, WU!). number of ligulate florets (6–8 vs. 5(–6) in S. altissimus). It differs from S. doria in its numerous supplementary bracts Senecio orientalis Mill., Gard. Dict., ed. 8: Senecio n.° 10. 1768, [(4–)5–8 vs. 3–5(–6)], slightly widener at the base, and in the nom. rej. prop. (Calvo et al. 2014b). Senecio coriaceus indumentum of the base of the involucre (± arachnoid in Aiton, Hort. Kew. 3: 195. 1789, nom. illeg. superfl. S. morisii vs. glabrous or puberulous in S. doria). The number (McNeill et al. 2012, ICN Art. 52.1).—TYPE: ‘Jacobæa of ligulate florets is also higher in S. morisii [6–8 vs. 5(–6)]. orientalis, latifolia, altissima’, Herb. Sherard (lectotype This species is apparently endemic of the Sarcidano and vel neotype, designated by Calvo et al. 2014b: OXF- Ogliastra regions, in central-eastern Sardinia, Italy. On the Sher-5346 (digital image!)). 2015] CALVO AND AEDO: TAXONOMY OF SENECIO DORIA GROUP 905

Senecio macrophyllus M. Bieb., Fl. Taur.-Caucas. 2: 308. 1808, Stem 0.5–1.5 m, solid, glabrescent to covered with ± nom. cons. prop. (Calvo et al. 2014b). Jacobaea macrophylla scattered multicellular trichomes (sometimes slightly arach- (M.Bieb.)C.A.Mey.,Verz.Pfl.Casp.Meer.:81.1831. noid in the insertion point of the leaves), greenish. Basal Senecio doria var. biebersteinii Lindem., Bull. Soc. Imp. leaves 13–36 cm long, 3.2–12.5 cm wide, oblanceolate to Naturalistes Moscou 40(2): 532. 1867. Senecio doria widely lanceolate (rarely ovate), obtuse to acute, attenuate to subsp. macrophyllus (M.Bieb.)Nyman,Consp.Fl.Eur.: rounded, distantly dentate to subentire, glabrescent or cov- 353. 1879. Senecio doria var. macrophyllus (M. Bieb.) Schmalh., ered with scattered multicellular trichomes (rarely slightly Fl. ssredn. jushn. Rossii 2: 88. 1897. Senecio biebersteinii arachnoid), with a petiole 2.5–14 cm long slightly widened at (Lindem.) Grec., Suppl. Consp. Fl. Roman.: 91. 1909. the base. Cauline leaves lanceolate to ovate, semi-amplexicaul, Senecio doria subsp. biebersteinii (Lindem.) Borza, Trif & concolorous. Capitula 35–300, 15–20 mm diam; involucre Ploaţă, Bul. Grad. Bot. Univ. Cluj 20, App. 2: 5. 1940.— cylindrical-campanulate; involucral bracts (11–)13, 3.9–6.1 mm TYPE: Ex Caucaso, M. Bieberstein s. n. (lectotype, desig- long, 0.8–2.5 mm wide, linear to lanceolate, acute, glabrous; nated by Calvo et al. 2014b: LE-1010062 (digital image!)). supplementary bracts 3–5(–6), 1.4–2.9 mm long, 0.3–0.5 mm wide, linear, a third to a half as long as involucral bracts, Senecio doria var. microcephalus Trautv. [“microcephala”], Bull. glabrescent with a tuft of hairs at the summit. Ligulate florets Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 12: 351. 5(–6), 8–13 mm long. Achenes 3.1–4.5 mm long, subcylindrical, 1854, nom. inval. (McNeill et al. 2012, ICN Art. 26.2). glabrous. Figure 1C. Senecio doria var. genuinus Lindem., Bull. Soc. Imp. Naturalistes Additional Illustration—Jacquin (1774: 51, Table 185); Moscou 40(2): 532. 1867, nom. inval. (McNeill et al. 2012, Reichenbach (1853: 42, Fig. 82–1); Schischkin (1936: 372, ICN Art. 24.3). Fig. 691, sub S. schwetzowii); Wissjulina (1962: 384, Fig. 72, sub S. schwetzowii); Nyárády (1964: 581, pl. 113, Fig. 1, Fig. 2, Senecio schwetzowii Korsh. [“schwetzowi”], Tent. Fl. Ross. Orient. sub S. doria var. biebersteinii); Vladimirov (2012: 447, Table 89–1). 519. 1898.—TYPE: RUSSIA. Bashkortostan: in gub. Ufa: Chromosome Number—2n=40 (Murín and Májovský SW, prope Dawlekanowo, [54°13′N 55°05′E], 10 July 1896, 1992; Konechnaya 1994, sub S. schvetzovii; Letz et al. 1999). I. M. Schwetzow s. n. (lectotype, designated here (vide Distribution and Habitat—Senecio doria is the most wide- Konechnaya in sched. 1978): LE-1010071 (digital image!)). spread species in the group. It is found in Austria, Bulgaria, Senecio doria subvar. danubialis Zenari,Lav.Bot.Ist.Bot. Czech Republic, Hungary, Kazakhstan, Moldova, Russia, Univ. Padova: 490. 1947.—TYPE: AUSTRIA. Vienna: in Romania, Serbia, Slovakia, and Ukraine, growing in damp pratis prope Aspern ad Danubium, [48°11′N16°29′E], meadows, grassy places, watersides, shrubs, glades of oak E. Woloszczak 237 (lectotype, designated here: G-443556!; groves, between elevations of 50–800 m (Fig. 3). isolectotype: BC-29026!). Phenology—Flowering from June to September.

Fig. 3. Distribution map for Senecio bithynicus (triangle) and S. doria (closed circle). 906 SYSTEMATIC BOTANY [Volume 40

Notes—Senecio doria and S. macrophyllus have been recently frontière de E de Chèrsone de la Bessarabie sur la Dnièster dans une recognized as separate species by several authors (Grulich vallée, 47°23′N 29°9′E, 3 Aug 1886, L. B. Montrésor s. n. (KW). and Hodálová 1994; Menitsky and Konechnaya 2008; Greuter ROMANIA. Alba: Transsilvania, distr. Târnava micâ, in pratis prope oppidum Blaj, 46°10′N 23°54′E, 13 Aug 1922, I. Pop s. n. (MA, G). Cluj: 2008; Calvo et al. 2014b), although their distribution ranges Kolozsvár, Szénafü, 46°49′N 23°35′E, 26 July 1904, W. Gugler s. n. (BP). remain poorly delimited and the available bibliography dif- Ialomiţa: fluvii Ialomiţa, prope pagum Coşereni, 44°41′N 26°34′E, 24 fers on their limits. According to Grulich and Hodálová Aug 1923, G. P. Grinţescu s. n. (G). (1994), who studied the group in central and southeastern RUSSIA. Chechnya: Chechenia-Ingushetia ASSR, between the rivers ′ ′ Europe, S. macrophyllus occurs in eastern Poland and Ukraine Terek and Sunzha, 1.8 km downstream of Braguny, 43°26 N 46°7 E, 2 Aug 1966, V. B. Kuvaev s. n. (LE). Kabardino-Balkaria: near Aryk and its southern limit is the river Dniester. Otherwise, Greuter village, 43°34′N 44°7′E, 8 Oct 1924, O. E. Knorring 405 (LE). Karachay- (2008) broadened the southern limit to Romania and Bulgaria, Cherkessia: Kuban Oblast, Batalpashinsk [Cherkessk], 44°14′N 42°2′E, and also indicated its presence in southwestern Russia (includ- 1 July 1890, V. I. Lipsky s. n. (LE). Stavropol Krai: Kislovodsk, 43°55′N ′ ing the Caucasus Mountains). 42°43 E, 14 July 1882, I. J. Akinfiev s. n. (LE). SERBIA. Belgrade: ad confines Hungariae meridionalis et Serbiae, ad On the other hand, Lindeman (1867) treated S. macrophyllus Danubium prope oppidum Belgrad, 44°49′N 20°31′E, J. Bornmüller 3440 as a subspecies of S. doria under the name S. doria var. (G, MT). biebersteinii. He separated this variety from the typical one on SLOVAKIA. Bratislava: Bratislava, Petržalka, 48°6′N 17°7′E, F. Nábělek the basis of its glabrous achenes (hairy in the typical variety s. n. (SAV). Trnava: com. Pozsony, prope pag. Sikabony, 47°59′N 17°35′E, invalidly named “genuinus”). Since all the species of the 9 Aug 1939, Á. Pénzes s. n. (BP). UKRAINE. Mykolaiv Oblast: Mykolaiv region, Yelanets'kyi district, S. doria group occurring in Ukraine have glabrous achenes, it vicinity of Kalynivka, 47°38′N 31°54′E, 16 Sep 2003, S. M. Voronov & is feasible to think that Lindemann based his S. doria var. A. F. Shcherbakov s. n. (KW). Sumy Oblast: Lebedynskyi Raion, genuinus on material from southwestern Europe, which corre- Mykhailivska Tsilyna Nature Reserve, north part, 50°44′N 34°10′E, 23 sponds to S. altissimus. July 1959, G. I. Bilyk s. n. (KW). Vinnytsia Oblast: Podolskaya, Olgopol district, vicinity of Verbka, 48°12′N 29°17′E, 26 July 1853, A. S. Rogowicz On the basis of our taxonomic study, S. macrophyllus fall in s. n. (KW). the variability encompassed by S. doria. We did not find any valuable taxonomic character to discriminate between both 4. Senecio legionensis Lange, Pug. Pl. Hispan. 2: 131. 1861. species. In agreement with our criterion, Trautvetter (1854) Senecio doria subsp. legionensis (Lange) Chater, Bot. synonymized S. macrophyllus to S. doria.Hestatedthatthe J. Linn. Soc. 68(4): 276. 1974.—TYPE: SPAIN. León: ad specimens from the region of Kiev (e.g. bei Wassilkow pagum Manzanal, [42°35′N 6°13′W], 10 July 1852, [Vasilkov], Bjelaja-Zerkow [Bila Tserkva], between Taraschtscha J. Lange s. n. (lectotype, designated here (vide Silva- and Swenigorodka [Tarascha, Zvenigorodka]) undoubtedly Pando in sched. 2008): C-10007880 (digital image!); iso- corresponded to S. macrophyllus and that they could not be lectotypes: G-442488!, P-3688094 (digital image!)). distinguished from S. doria. Senecio doria var. subintegrus Merino [“subintegrum”], Fl. In 2014 we realized that the names S. orientalis and Galicia 3: 599. 1909.—TYPE: SPAIN. Ourense: loco dicto S. macrophyllus were synonyms, the former having priority [en pantanos entre Cadones y Bande], [42°01′N 7°58′ over the latter, and we proposed to conserve the name W], without data, B. Merino s. n. (lectotype, designated S. macrophyllus (Calvo et al. 2014b). This nomenclatural by Silva-Pando et al. 2009: LOU-1222/3 (digital image!); action was done on the supposition that S. macrophyllus was isolectotype: MA-130611!). a distinct taxonomic entity. Senecio doria is a variable species mainly regarding the leaf Stem 0.5–1.1 m, usually fistulous, glabrous, greenish. Basal indumentum. Some specimens from Caucasus stand out by leaves 8–26 cm long, 1.8–6.1 cm wide, narrowly lanceolate to displaying leaves with ± scattered multicellular trichomes, linear-lanceolate, obtuse to acute, attenuate, entire to dis- whereas those from central Europe tend to have glabrescent tantly dentate, glabrous, with a petiole 7–21 cm long slightly leaves. The number of supplementary bracts varies between widened at the base. Cauline leaves narrowly lanceolate, 3–5(–6), although any geographic pattern is observed. sessile, concolorous. Capitula 6–65, 16–22 mm diam; invo- We did not have a chance to study material of this taxon lucre cylindrical-campanulate; involucral bracts 10–13(–16), from Kazakhstan, but we include this country in the distri- 5.2–5.9 mm long, 0.9–1.6 mm wide, linear, acute, glabrous; bution according to Fedtschenko and Fedtschenko (1912), supplementary bracts (1–)2–5, 1.5–2.8 mm long, 0.5–0.7 mm who reported this species from Uralsk and Turgai regions wide, linear-triangular, a quarter as long as involucral bracts, under the name S. doria var. macrophyllus. Later, Pavlov glabrous. Ligulate florets 5(–7), 9.7–11.6 mm long. Achenes (1938) and Roldugin (1966) also recorded it under the hetero- 3.4–4 mm long, subcylindrical, with scattered intercostal tri- typic synonym S. schwetzowii. chomes. Figure 4B. — Representative Specimens Examined AUSTRIA. Lower Austria: 4 km Chromosome Number—unknown. NE of church of Hainburg, ESE “Jägerhaussiedlung”, at the narrow point — between “Auglarn” dead river branch and way following Danube river, Distribution and Habitat Senecio legionensis is endemic 48°9′N16°59′E, 3 Sep 2013, J. Walter & E. Vitek 13–0416 (MA). to the northwestern region of the Iberian Peninsula (Portugal BULGARIA. Veliko Tarnovo: Pavlikeni district, NW of Nedan village, and Spain). It grows in wetlands, damp meadows, and near left bank of Lomya river, 43°19′N25°14′E, 17 Aug 2013, S. Stoyanov & the edges of deciduous woods, between elevations of 400– V. Goranova s. n. (SOM). š 1,450 m (Fig. 2). CZECH REPUBLIC. South Moravian Region: Vy kov, inter pagum — Křenovice et Holubice, 49°9′N 16°49′E, 1 Sep 1967, V. S křivánek s. n. (MT). Phenology Flowering from May to July. HUNGARY. Central Transdanubia: opp. Esztergom, 47°47′N18°43′E, Notes—Senecio legionensis is a glabrous species that can be 8 Sep 1907, S. Jávorka s. n. (BP). Central Hungary: insula Danubialis Csepel distinguished by the morphology of its leaves, which are ′ ′ prope pagum Sciget-Ujfalu, 47°14 N18°55E, 29 July 1874, J. A. Tauscher s. n. narrowly lanceolate to linear-lanceolate. The involucre bears (G). Northern Hungary: Abauj-Torna m. Gilbárt, Hernád partján, 48°18′N 21°9′E, 31 July 1937, Á. Kiss s. n. (BP). a low number of short supplementary bracts, and the achenes MOLDOVA. Chişinãu: Kischinewia, 47°2′N 28°54′E, 20 July 1882, have scattered intercostal trichomes. These characters are use- N. M. Zelentzky s. n. (G). Dubãsari: Jahorlik [Iagorlîc], Podolie, sur la ful to easily separate it from S. altissimus, with which it 2015] CALVO AND AEDO: TAXONOMY OF SENECIO DORIA GROUP 907

Fig. 4. A. Senecio umbrosus. a1. Capitulum. a2. Supplementary bract. a3. Inner involucral bract. a4. Outer involucral bract. a5. Ligulate floret [Based on Weber s. n. (MA)]. a6. Achene with pappus [Based on Podpěra & Laus s. n. (MA)]. B. Senecio legionensis. b1. Capitulum. b2. Supplementary bract. b3. Inner involucral bract. b4. Outer involucral bract. b5. Ligulate floret [Based on Fernández Díez s. n. (MA)]. b6. Achene with pappus [Based on Penas et al. s. n. (LEB)]. C. Senecio fontanicola. c1. Capitulum. c2. Supplementary bract. c3. Inner involucral bract. c4. Outer involucral bract. c5. Ligulate floret [Based on Fiori s. n. (FI)]. c6. Achene with pappus [Based on Béguinot 1771 (FI)]. 908 SYSTEMATIC BOTANY [Volume 40 partially overlaps its area of distribution (see also comments quarter as long as involucral bracts, glabrous. Ligulate under S. altissimus). florets 5–7, 12.1–15.6 mm long. Achenes 3.8–4.4 mm long, Senecio legionensis is similar to S. fontanicola from Carinthia subcylindrical, glabrous or with few trichomes at the summit and northeastern Italy, mainly due to their basal leaves and base. Figure 4C. narrower than in the remaining taxa of the group. Both spe- Additional Illustration—Vreš et al. (2012: 2, Fig. 1, cies also bear a low number of supplementary bracts con- as photo). spicuously shorter than the involucral ones. However, the Chromosome Number—2n=40 (Grulich and Hodálová 1994). achene indumentum is a useful character to distinguish Distribution and Habitat—This species occurs in Austria, between the two species (with scattered intercostal trichomes Italy, and Slovenia, growing in marshy spring areas, fens, in S. legionensis vs. glabrous or nearly so in S. fontanicola). and reed beds, between elevations of 20–850 m (Fig. 5). Representative Specimens Examined — PORTUGAL. Alto Minho: Phenology—Flowering from May to early July. ′ ′ Melgaço, Castro Laboreiro, entre Teso e Campelo, 42°2 N 8°8 W, 4 Aug Notes—This is a distinctive species characterized by glabrous 1998, F. B. Caldas & J. J. Honrado s. n. (PO); Melgaço, Castro Laboreiro, Chã de Matança, 4 2°0′N8°11′W, June 1996, F. B. Caldas & H. Nepomuceno basal leaves (narrower than most of the related taxa), involucral Alves s. n. (PO). bracts relatively long, a low number of supplementary bracts, SPAIN. A Coruña: Sobrado dos Monxes, prados de la laguna, 43°2′N and glabrous achenes or nearly so. It is similar to S. legionensis ′ 8°0 W, 14 July 1983, F. Gómez Vigide s. n. (LOU). León: Garrafe de Torío, from northwestern Spain (see comments under it). Fontanos de Torío, 42°46′N 5°33′W, 28 Aug 2007, F. Egido s. n. (LEB); León, P. de Manzanal, 42°35′N 6°13′W, 20 June 1904, M. Gandoger 579 We observed that the populations from southern Carinthia (G, P, WU); valle de Pardomino [Pardaminos], 42°55′N 5°16′W, 12 July (Austria) have glabrous achenes whereas those specimens 1995, F. Gómiz s. n. (LEB); ad pagum Manzanal, prov. legion., 42°35′N from Friuli and Veneto (Italy) usually have some trichomes ′ ′ ′ 6°13 W, 9 July 1852, J. Lange s. n. (C, K); Villagatón, 42°38 N 6°9 W, concentrated at the summit and base, but in no case over the 20 June 1995, Á. Penas et al. s. n. (LEB, MA). Lugo: entre Cuiña y Vilalvite, 42°58′N7°31′W, 5 June 2002, J. Amigo s. n. (GDA, SANT); Castro whole achene. More collections in fruit should be studied to del Rey, Azumara, 43°11′N7°23′W, 18 July 1997, J. Amigo & J. Izco s. n. improve the understanding of the variability of this character. (SANT); Fervedoira, 43°1′N7°31′W, 27 May 1991, F. Gómez Vigide s. n. We include the infraspecific taxa described by Zenari (1947) (LOU); Guitiriz, pr. San Alberto, 43°10′N7°47′W, 20 July 1985, F. Gómez in Friuli within the variability encompassed by S. fontanicola. ′ ′ Vigide s. n. (LOU); Villalba, pr. Parrocha, 43°14 N7°40W, 5 June 1987, F. J. This species does not overlap its distribution area with Silva-Pando 4034 (G, LEB, LOU, MA, SANT); Murás, Viveiró, 43°28′N 7°37′W, 3 Aug 1987, F. J. Silva-Pando 4471 (LOU). Ourense: monte Grande any of the other members of the group. — de Bande, 42°3′N7°56′W, 13 Aug 1981, F. J. Silva-Pando 38 (LOU). Representative Specimens Examined AUSTRIA. Carinthia: Velden am Wörthersee, 46°36′N 14°2′E, July 1903, Müller s. n. (G); Ferlach, 5. Senecio fontanicola Grulich&Hodálová,Phyton(Horn) 46°31′N 14°18′E, June 1905, Müller s. n. (GE). ′ ′ 34(2): 261. 1994.—TYPE: AUSTRIA. Carinthia: Dobratsch- ITALY. Friuli: Porcia-Rio da Pieve, 45°57 N 12°36 E, 18 May 1946, ′ ′ S. Zenari s. n. (PAD); Roraigrande di Pordenone-Lago della Burida- Abstieg, Heil.Geist-Pogöriach, [46°36 N 13°47 E], 11 July insenatura settentrionale, 45°57′N 12°38′E, 28 May 1943, S. Zenari s. n. 1907, K. Ronniger s. n. (holotype: W-23368 (digital image!)). (PAD); Zoppola, rive del fiume Brentella a nord di Ovoledo, 45°58′N 12°46′E, 4 June 1946, S. Zenari s. n. (PAD); Pordenone, lago della Bureda, Senecio doria var. angustifolius Pirona, Fl. Forojul. Syll.: 82. parte settentrionale, 45°57′N 12°38′E, 28 May 1947, S. Zenari s. n. (PAD); 1855, nom. inval. (McNeill et al. 2012, ICN Art. 38.1). Fontanafredda, Rio Laguzza, Ponte Frait, 45°57′N 12°33′E, 18 May 1946, S. Zenari s. n. (PAD); Ai Camoi, prati acquitrinosi Marcile tra Sacile e Senecio doria var. subdecurrens Zenari, Lav. Bot. Ist. Bot. Univ. Fontanaffreda, 45°57′N 12°32′E, 25 May 1943, S. Zenari s. n. (PAD). Padova: 489. 1947.—TYPE: ITALY. Friuli: fra Roraipiccolo Friuli–Venezia Giulia: paludi di Talmassone, 45°55′N13°4′E, 16 May ePorcia,[45°57′N12°37′E], 31 May 1945, S. Zenari s. n. 1875, C. Marchesetti s. n. (FI). Veneto: Venetia, Padova, Loreggia, loco Carpane dicto, 45°35′N 11°56′E, June 1911, A. Béguinot 1771 (FI, GE, P, (lectotype, designated here: PAD s. n. (digital image!)). TO); Padovano ad Abbazia-Pisani, 45°37′N 11°51′E, 23 May 1898, A. Fiori s. n. (FI). Senecio doria subvar. intermedius Zenari,Lav.Bot.Ist.Bot.Univ. č — SLOVENIA. Upper Carniola: Julian Alps, upper Sava valley, Rate e, NR Padova: 489. 1947. TYPE: ITALY. Friuli: Pordenone, lago Zelenci, 46°29′N 13°44′E, 20 June 2011, B. Vreš & I. Dakskobler s. n. (LJS). della Burida, sorgente del boschetto (Casa Zenari), [45°57′ Senecio umbrosus N12°38′E], 18 May 1946, S. Zenari s. n. (lectotype, desig- 6. Waldst. & Kit., Descr. Icon. Pl. Hung.: nated here: PAD s. n. (digital image!)). 232, Table 210. 1805. Senecio doria var. latifolius Thomé, Fl. Deutschl. 4: 299. 1889. Senecio doria [unranked] Senecio doria var. golae Zenari, Lav. Bot. Ist. Bot. Univ. umbrosus (Waldst. & Kit.) Schmalh., Fl. ssredn. jushn. Padova: 489. 1947.—TYPE: ITALY. Friuli: Roraigrande Rossii 2: 89. 1897 (McNeill et al. 2012, ICN Art. 37.3). di Pordenone-Lago della Burida-vallette a boscaglia, Senecio doria var. umbrosus (Waldst. & Kit.) Stoj. & Stef., [45°57′N12°38′E], 14 May 1943, S. Zenari s. n. (lectotype, Fl. Bulg. 2: 1146. 1925. Senecio doria subsp. umbrosus designated here: PAD s. n. (digital image!)). (Waldst. & Kit.) Soó, Erdész. Kisérl. 46: 292. 1946.—TYPE: SLOVAKIA. Without locality or date, P. Kitaibel s. n. (lec- Senecio doria subvar. forojuliensis Zenari, Lav. Bot. Ist. Bot. totype, designated by Kováts 1992: BP Herb. Kit. XXX/ Univ. Padova: 489. 1947, nom. inval. (McNeill et al. 61 (digital image!)). 2012, ICN Art. 26.2). Senecio doria var. macrocephalus Trautv. [“macrocephala”], Stem 0.5–0.95 m, solid, glabrous, greenish. Basal leaves Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint-Pétersbourg 9.5–25 cm long, 2–5.5 cm wide, oblanceolate to narrowly lan- 12: 352. 1854.—TYPE: UKRAINE. “Gal. Podol.” [Galicia, ceolate, acute to obtuse, attenuate, distantly dentate to crenate, W. S. J. G. Besser s. n. glabrous, with a petiole 3–9cmlongslightlywidenedatthe Podolia], (lectotype, designated base. Cauline leaves narrowly lanceolate, semi-amplexicaul to here: KW s. n. (digital image!)). sessile, concolorous. Capitula 12–40, 17–20 mm diam; involucre Senecio umbrosus var. subtuberculatus Borbás ex Formánek, cylindrical-campanulate; involucral bracts 13, 6.2–8.2 mm long, Oesterr. Bot. Z. 40: 85. 1890.—TYPE: BOSNIA AND 0.9–1.8 mm wide, linear, acute, glabrous; supplementary HERZEGOVINA. Liskovica in Bosnia, V. Borbás s. n. bracts 1–3, 1.5–2.2 mm long, 0.3–0.4 mm wide, linear, a (not found at BP). 2015] CALVO AND AEDO: TAXONOMY OF SENECIO DORIA GROUP 909

Fig. 5. Distribution map for Senecio fontanicola (open circle) and S. umbrosus (triangle).

Senecio urumovii Velen. [“Urumovi”], Oesterr. Bot. Z. 52: Additional Illustration—Waldstein and Kitaibel (1805: 52. 1902.—TYPE: BULGARIA. Plovdiv: Tikisky Balkan, 232–233, Table 210); Reichenbach (1853: 42, Fig. 82–2); [42°44′N 24°35′E], 1901, I. K. Urumoff 129 (lectotype, Nyárády (1964: 577, pl. 112 Fig. 1); Vladimirov (2012: 425, designated here: PRC-455075 (digital image!)). Table 84–1). Chromosome Number—2n=40 (Hindáková 1978; Dobeš Senecio umbrosus f. transiens Nyár., Bul. Şti. Acad. Republ. ţ Ş et al. 1996). Populare Romîne, Sec . ti. Biol. Agron. Geol. Geogr. 3(1): Distribution and Habitat—Senecio umbrosus is found in 35. 1951 [in Nyárády and Soó, Kv. Fl. 7: 547. 1943, cum Austria, Bosnia-Herzegovina, Bulgaria, Croatia, Czech Republic, diagn. hung., nom. inval. (McNeill et al. 2012, ICN — Hungary, Poland, Romania, Slovenia, Serbia, Slovakia, and Art. 39.1)]. TYPE: ROMANIA. Cluj: Rezervatiunea din Ukraine. This species grows in damp meadows, woods of Fânatele Clujului, Copârsae din Valea prima, [46°50′N Pinus, Picea, etc., and usually on calcareous soils, between 23°37′E], 25 July 1934, E. I. Nyárády s. n. (lectotype, elevations of 200–1,150 m (Fig. 5). designated here: CL-259929 (digital image!); isolectotype: Phenology—Flowering from July to September. CL-193456 (digital image!)). Notes—This species is easily distinguishable from the Stem 0.8–2 m, solid, covered with long multicellular tri- other species of the group by the high number of ligulate chomes or rarely glabrescent, usually with the lower half florets, which ranges from 8–10, and by its slightly larger brick-red coloured. Basal leaves 23–46 cm long, 7.5–18 cm involucre with short supplementary bracts. The stem is wide, ovate to widely lanceolate, obtuse, rounded to atten- usually covered with a dense indumentum of long multi- uate and usually decurrent along the petiole, distantly den- cellular trichomes, and it often has the lower half brick-red tate (rarely subentire or crenate), glabrescent above, covered coloured. The stem indumentum is a variable character, with scattered multicellular trichomes beneath, sometimes ± as well as the indumentum of the abaxial surface of the dense along the midrib and secondary veins, with a petiole leaves. Some populations have the stem, and the midrib 4–24 cm long slightly widened at the base. Cauline leaves and the secondary veins densely covered with long multi- ovate to elliptic or lanceolate, semi-amplexicaul to amplexicaul, cellular trichomes (e.g. Lúčky in northern Slovakia, Tara concolorous. Capitula 17–80, 19–24 mm diam; involucre Mountain in western Serbia), whereas others are glabrous cylindrical-campanulate; involucral bracts (12–)13(–14), 5.4– or nearly so (e.g. pr. Hristo Danovo in the central Balkan 7.7 mm long, 1.2–2.6 mm wide, linear to lanceolate, acute, Mountains). The existence of intermediate specimens led glabrous; supplementary bracts (3–)4–5(–6), 1.2–3.2 mm us to treat them as part of infraspecific variability of a long, 0.3–0.6 mm wide, linear-triangular, a quarter as long single species. as involucral bracts, with some scattered trichomes or The populations from southeastern Poland and western glabrescent. Ligulate florets 8(–10), 10.4–18.2 mm long. Ukraine have been recently ascribed to S. macrophyllus Achenes 3.7–4.3 mm long, subcylindrical, glabrous. Figure 4A. (Grulich and Hodálová 1994; Czarnecka and Kucharczyk 910 SYSTEMATIC BOTANY [Volume 40

2001; Czarnecka 2009; Czarnecka and Denisow 2014). The as S. umbrosus. Besser (1822) already identified the plants indumentum on the stem, midrib, and secondary veins, occurring in Podolia (Ukraine) as S. umbrosus.Trautvetter the short supplementary bracts, and the number of ligulate (1854), who later studied some of Besser’sspecimensfrom florets (8–10) allow us to clearly identify these populations this region, considered them more similar to S. doria because

Fig. 6. Senecio bithynicus. a. Lower part of the stem. b. Middle cauline leaf. c. Upper part of the stem and synflorescence. d. Detail of the adaxial sur- face of the leaf. e. Detail of the abaxial surface of the leaf. f. Capitulum. g. Capitulum without ligulate florets. h. Supplementary bract. i. Detail ofthe apex of the supplementary bract. j. Involucral bract. k. Ligulate floret. l. Tubular floret [Based on Post s. n., 20 July 1940 (G)]. m. Achene with pappus. n. Achene [Based on Post s. n., 7 Sep 1940 (G)]. 2015] CALVO AND AEDO: TAXONOMY OF SENECIO DORIA GROUP 911 of their scarce arachnoid indumentum. However, he stressed ca. 25 cm long, 13 cm wide, pandurate, decreasing in size up that these specimens could be distinguished by their bigger to the stem and becoming ovate-elliptic, obtuse, auriculate capitula, and named them S. doria var. macrocephalus.Among and semi-amplexicaul, distantly dentate, glabrescent above, the material that we could study from KW, we found a Besser arachnoid-floccose beneath, discolorous. Capitula ca. 65, ca. specimen from Galicia-Podolia that matches the information 12 mm diam; involucre subcylindrical; involucral bracts (11–)13, and provenance indicated by Trauvetter. It is here designated 6.8–7.6 mm long, 0.9–1.3 mm wide, linear, acute, glabrescent as the lectotype of the name S. doria var. macrocephalus. or with some scattered arachnoid trichomes; supplementary Senecio umbrosus occurs through central and southeastern bracts 2–3(–4), 4–5.1 mm long, 0.4–0.7 mm wide, linear, a half Europe, where S. doria also grows. However, we detected as long as involucral bracts, glabrescent or with some scattered that their ranges of distribution strictly overlap only in few arachnoid trichomes. Ligulate florets 5–6, 9.2–11.5 mm long. localities such as nearby Cluj (Romania). The presence of Achenes 3.3–4.1 mm long, subcylindrical, glabrous. Figure 6. S. umbrosus in Bosnia-Herzegovina was reported by Hayek Chromosome Number—unknown. (1931) and Slišković (1983). Unfortunately we did not study Distribution and Habitat—Senecio bithynicus is only known any collection from this country. Godicl (1982), and Wraber from Abant Lake (Bolu Province, NW Turkey), growing in and Skoberne (1989) recorded the presence of S. doria in marshes at elevations of ca. 1,330 m (Fig. 3). Slovenia, but we did not find any herbarium specimen from Phenology—Flowering in July and probably until August. there either. The proximity of some S. umbrosus populations Notes—Until now, plants from Abant Lake (Bolu Province) in Croatia lead us to think that they probably correspond have been identified as S. doria subsp. umbrosus (≡ S. umbrosus) to this species. On this basis we include Slovenia in (Matthews 1975; Uçar Türker and Güner 2003). However, it the distribution. is morphologically clearly different from S. umbrosus and The label of the lectotype of S. umbrosus only shows the from the other species of the group. The relevant characters name of the species in Kitaibel’s handwriting (Kováts 1992). distinguishing this taxon from the others are its pandurate However, the indication provided in the protologue reveals cauline leaves with arachnoid-floccose indumentum beneath that it was collected “ad thermas Lucskienses Comitatus and its linear supplementary bracts a half as long as the Liptoviensis”, which is located in the Žilina region of Slovakia. involucral ones. It is the only member of the group with It has to be noted that there are several duplicates of the lec- discolorous leaves with a constriction in the lower half of the – totype kept at the Kitaibel herbarium (BP). lamina. The length of the supplementary bracts (4 5.1 mm) Representative Specimens Examined — AUSTRIA. Lower Austria: in is also much longer than that of the other taxa. This species valle Grabenweg-Tal prope Pottenstein, 47°57′N 16°4′E, Aug 1907, has the stem fistulous, a character only seen in some speci- ′ ′ K. Ronniger 5095 (G); Austriae inferioris, ad Pottenstein, 47°58 N 16°6 E, mens of S. legionensis. 6 Aug 1896, F. A. Tscherning s. n. (MA, P). BULGARIA. Lovech: Mt Stara Planina, Troyanski Balkan region [close To date, S. bithynicus is only known from the shores of the to the town of Troyan], 42°51′N 24°42′E, S. Grancharov s. n. (SOM). Stara Abant Lake, where it grows in elevations of ca. 1,330 m (Uçar Zagora: Central Stara Planina, E. Mazalat, valley of the river, 42°44′N Türker and Güner 2003). Further material is needed to under- 25°7′E, 29 July 1979, B. A. Kuzmanov s. n. (SOM). stand its variability and to properly describe the basal leaves. ž ′ ′ CROATIA. Bei Warasdiner Tepliz [Vara dinske Toplice], 46°12 N 16°24 E, Since the basal leaves were not present in the studied bloom- Aug 1853, L. Farkaš Vukotinović s. n. (P); in valle fluvii Kolpa prope pagum Kuželj, 45°29′N14°48′E, 2 Aug 1978, E. Mayer & M. Mayer s. n. (LJS). ing specimen, we suppose that they are withering early. CZECH REPUBLIC. South Moravian Region: Moravia Merid,-Orient., The epithet bithynicus refers to the ancient region called Strážnice, secundum rivum Járkovec supra Jiříkovci prope Radějov, Bithynia, where the species occurs, in the northwest of Asia Minor. ′ ′ ě 48°51 N 17°23 E, 11 Aug 1929, J. Podp ra & J. Laus 482 (G, MA, MT); Representative Specimens Examined — TURKEY. Bolu: Abant Gölu, ě ′ ′ Montes Bilé Karpaty, Jarkovec prope Rad jov, 48°51 N 17°20 E, 26 Aug 40°36′N 31°17′E, 7 Sep 1940, B. Post s. n. (G). 1932, L. F. Weber s. n. (G). Zlín Region: Montio Bilé Karpaty, Moravie, 49°3′N 18°0′E, 28 July 1932, L. F. Weber s. n. (MT). HUNGARY. Western Transdanubia: in campestribus elatioribus prope Acknowledgments. We are grateful to the curators of the herbaria pag. Sz. Gothárd [Szentgotthárd] transsilvaniae centralis, 46°57′N 16°17′E, mentioned in the text, as well as to Gianluigi Bacchetta (CAG), Zoltán 20 July 1869, V. Janka s. n. (G). Barina (BP), Iva Hodálová and Tatiana Mihalikova (SAV), Olga POLAND. Lublin: Tomaszów Lubelski County, Central Roztocze Korniyenko (KW), Rossella Marcucci (PAD), Svetlana Bancheva (SOM), Highland (Biała Góra), 50°28′N 23°29′E, as photo. Mihai Puscas (CL), Branko Vreš (LJS), Patrik Mráz (PRC), Mark Spencer ROMANIA. Sud: Prahova, pr. Sinaia, m. Paduchiosul, 45°19′N 25°31′ (BM), Chiara Nepi (FI), Simonetta Peccenini (GE), Laura Guglielmone E, 9 Aug 2009, C. Aedo 16820 (MA); Sinaïa, 45°21′N 25°32′E, July 1892, (TO), Marjan Niketić (BEO), Francisco Javier Silva-Pando (LOU), Elena de D. Grecescu s. n. (P). Paz Canuria (LEB), Joan Font (HGI), María de Lourdes Rico Arce (K), SERBIA. Zlatibor: PN de Tara, loco dicto Verica Batura, 43°54′N Paulo Alves (PO), Roman Ufimov (LE), and Patricia Barberá (MA) for 19°19′E, 13 Sep 2004, A. Charpin 27449 (G); Tara planina, Kruščica, ad providing material or useful information. We also thank Juan Castillo for rivum Beli Rzav, 43°54′N 19°23′E, 7 Oct 1997, M. Niketić s. n. (BEO). the drawings, Roman Ufimov and Ramón Morales for their Cyrillic and SLOVAKIA. Žilina: distr. Turč. Martin, prope pagum Kralovany, 49°9′ German translations respectively, and Élise Ross-Nadié for helpful N 19°8′E, July 1936, J. Dostál s. n. (P); circa thermas Lucsky, 49°8′N suggestions. This work has been funded by the Flora iberica project 19°24′E, Aug 1882, J. Pantocsek 1581 (P). (CGL2011-28613-C03-01). UKRAINE. Lviv Oblast: Rajon Zolochivs'ky, Podolje, Mt. Lysa Gora, 49°47′N 24°44′E, July 1989, I. Hodálová & A. Kagalo s. n. (SAV); Rajon Zolochivs'ky, Podlesje, Mt. Biela Gora, 49°55′N 24°50′E, July 1989, I. Hodálová & A. Kagalo s. n. (SAV). Literature Cited 7. Senecio bithynicus J. Calvo, sp. nov.—TYPE: TURKEY. Besser, W. S. J. G. 1822. Enumeratio plantarum hucusque in Volhynia, Podolia, Bolu: Abant Gölu, [40°36′N 31°17′E], 20 July 1940, Gub. Kiioviensi, Bessarabia Cis-Tyraica et circa Odessam collectarum [...]. Vilnae: Typis Josephi Zawadzki Universitatis Typographi. B. Post s. n. (holotype: G-443529!). Blanca, G. 2002. Senecio L. Pp. 665–669 in Catalogue des plantes vasculaires du Nord du Maroc, incluant des clés d′identification, vol. 2, eds. B. Valdés, Stem ca. 1 m, fistulous, with scattered arachnoid trichomes M. Rejdali, A. Achhal El Kadmiri, J. L. 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Pp. 371–411 in Flora URSR, vol. 11, ed. Lindeman, E. E. 1867. Florula Elisabethgradensis. Additamentum ad Floram E. D. Wissjulina. Kiev: Vydavnyctvo Akademii Nauk Ukrajinskiej RSR. Chersonensem. Bulletin de la Société Impériale des Naturalistes de Moscou Wraber, T. and P. Skoberne. 1989. Rdeči seznam ogroženih praprotnic in 40: 448–544. semenk SR Slovenije. Varstvo Narave (Ljubljana) 14–15: 1–430. 2015] CALVO AND AEDO: TAXONOMY OF SENECIO DORIA GROUP 913

Yilin, C., B. Nordenstam, C. Jeffrey, and L. Vincent. 2011. Senecio L. E. Jahandiez, 796 (1); V. Janka, 20 July 1869 (6); S. Jávorka, 20 Aug 1911 Pp. 508–536 in Flora of China, vol. 20–21 (Asteraceae), eds. Wu, (6), 8 Sep 1907 (3). Z. Y., P. H. Raven, and D. Y. Hong. Beijing: Science Press; St. Louis: Z. Kárpáti, 2 Aug 1946 (3); Á. Kiss, 31 July 1937 (3); I. Klášterský, 9 Aug Missouri Botanical Garden Press. 1946 (6); O. E. Knorring, 405 (3); F. Kovács, 23 Aug 1929 (3); L. Kralik, Zenari, S. 1947. Intorno ad un Senecio delle risorgive friulane. Lavori di 6 June 1847 (1); V. B. Kuvaev, 2 Aug 1966 (3); B. A. Kuzmanov, 29 July botanica, Istituto di Botanica e di Fisiologia Vegetale dell′Università di 1979 (6). Padova (volume pubblicato in occasione del 70° genetliaco del Prof. M. Ladero et al., 4 Oct 1979 (1); M. Laínz, 3 July 1956 (4); J. Lange, 7 July Giuseppe Gola): 487–491. 1860 (4), 8 July 1852 (4), 9 July 1852 (4), 10 July 1852 (4); E. M. Lavrenko, 203 (3), 9 July 1923 (3); G. Lengyel, 30 Aug 1913 (6), June 1910 (3); G. L'Hermite, July 1941 (1); E. E. Lindemann, 20 July 1873 (3); V. I. Lipsky, 1 July 1890 (3), 28 June 1890 (3); A. E. Lomax, 22 July 1891 (1); G. López & Appendix 1. List to Exsiccatae—Specimens are listed alphabetically S. Bondia, 1397 (1). by collector, followed by collection number or date when collector number L. Macchiati, July 1885 (1); R. Maire, 13 July 1921 (1), 18 July 1938 (1), is unavailable. The number in parentheses corresponds to the number of 27 July 1921 (1), 28 June 1938 (1); C. Marchesetti, 16 May 1875 (5); K. the accepted species in the treatment. Marusyak, 255 (3); E. Mayer & M. Mayer, 2 Aug 1978 (6); L. Medina C. Aedo, 16701 (1), 16820 (6); I. J. Akinfiev, 14 July 1882 (3), 4 July 1902 et al., 3615 (1); L. Medina & L. M. Ferrero, 2823 (1), 3702 (1); Yu. L. (3); J. A. Alejandre & P. Urrutia, 1 Aug 1986 (1), 31 July 1986 (1); S. Alias, Menitsky & V. Nikolaev, 18 Sep 1988 (3); B. Merino, 70 (4); L. B. 27 May 1968 (2); F. S. Alioth, 16 July 1853 (1); J. Amigo, 5 June 2002 (4), Montrésor, 3 Aug 1886 (3); M. Mouret, 1471 (1); Müller, June 1905 (5), 8 July 2000 (4); J. Amigo et al., 23 July 1994 (4); J. Amigo & J. Izco, 18 July July 1903 (5); F. Muñoz Garmendia, 04 July 1974 (1); F. Muñoz 1997 (4); V. J. Arán, 5014 (1), 3 Aug 1991 (1); V. J. Arán & M. J. Tohá, 16 Aug Garmendiaetal.,5323(1). 1999 (1); R. Arias, 12 May 2002 (4); P. V. Arrigoni, 15 June 1963 (2); P. V. M. Niketić, 7 Oct 1997 (6); F. Norris, 27 June 1892 (1); I. V. Arrigoni & E. Nardi, 27 June 1972 (2). Novopokrovsky, 1261 (3), 2 July 1901 (3); E. I. Nyárády, 25 July 1934 G. Bacchetta, 1 July 2000 (2), 1 July 2013 (2); G. Bacchetta & A. Delage, (6); M. Nydegger, 37218 (1). 19 June 2013 (2); G. Bacchetta & C. Dessì, 15 July 2009 (2); P. Bariego, 3385 J. Pantocsek, 979 (6), 1581 (6); S. Patino, P. M. Uribe-Echebarría & J. (1); T. Barta, 2002–281 (3), 20 Aug 2006 (3); J. Barth, 16 Aug 1889 (3), 16 Valencia, 10 July 1990 (1); E. Paz Canuria, 27 June 1995 (1), 29 July 2004 Aug 1908 (3), 30 July 1890 (3); Bayern, 20 Aug 1866 (3), 28 July 1866 (3); (1); J. Pedrol & J. Ascaso, 645 (1); abbé Pellat, July 1872 (1), July 1883 (1); A. Béguinot, 1771 (5); F. Bellot, 22 Aug 1965 (1); F. Beltrán, Aug 1914 (1); Á. Penas et al., 13 Aug 1994 (4), 20 June 1995 (4); Á. Pénzes, 9 Aug 1939 G. I. Bilyk, 23 July 1959 (3); A. Biondi, 27 May 1879 (2); A. Blanco, 524 (1); (3); C. Pérez, 28 Aug 1947 (1); C. Pérez Morales, 28 July 1984 (1); C. Pérez O. Bolòs, 19 July 1972 (1); O. Bolòs & V. Codina, 26 June 1955 (1); J. Morales&E.Puente,18July1988(1);J.Podpěra & J. Laus, 482 (6); I. Pop, Bornmüller, 3440 (3); E. Bourgeau, 725 (1), 2514 (1); A. H. Brotherus & V. F. 13 Aug 1922 (3); P. Porta & G. Rigo, 562 (1), 8 July 1895 (1); B. Post, Brotherus, 497 (3). 20 July 1940 (7), 7 Sep 1940 (7). A. Caballero, 21 Aug 1936 (1); F. Cabezas et al., 496 (1); B. Cabezudo & C. Quesada et al., 2 Aug 1984 (1); C. Quesada & G. Blanca, 9 Aug 1984 (1). J. M. Nieto, 16 July 1982 (1); F. B. Caldas & H. Nepomuceno Alves, June E. Reverchon, 59 (1), Aug 1891 (1), Aug 1892 (1), July 1893 (1), July 1894 1996 (4); F. B. Caldas & J. J. Honrado, 4 Aug 1998 (4); J. Calvo, 2653 (1), 2667 (1), July 1900 (1), July 1901 (1), July 1902 (1), July 1904 (1); L. Richter, July (1), 3962 (1), 3971 (1), 4064 (1); J. Calvo & É. Ross-Nadié, 6724 (1), 6733 (1), 1895 (6), Aug 1876 (3); A. Rigual, 27 Aug 1959 (1), 6 July 1975 (1); S. Rivas 6792 (1); M. Campá, July 1868 (1); B. Casaseca, 18 July 1962 (1); A. Charpin, Goday, 28 July 1947 (1), 28 Aug 1947 (1); A. S. Rogowicz, 26 July 1853 (3); T. 19815 (1), 27449 (6); A. Charpin & F. J. Fernández Casas, 10539 (1); S. Romero, 14 July 1982 (1); C. Romero, July 1973 (1); A. Romo, 2 July 1979 (1); Cirujano, 22 July 1992 (1); A. H. C. Coincy, 10 June 1890 (1), 19 June 1892 (1), K. Ronniger, 5095 (6), 11 July 1907 (5); H. Ross, 31 (1). 9 July 1897 (1); H. Correvon, 27 June 1920 (1); A. Cuénod, 23 July 1927 (1). G. Sag, 12709 (1); E. Saint-Lager, 30 June 1908 (1); Z. A. Sarycheva et al., M. Danciu, 2 Aug 1974 (6); J. Dostál, July 1936 (6); J. C. Ducommun, 7 Aug 1958 (3); I. M. Schwetzow, 10 July 1896 (3); A. Scrugli & L. Mura, 5 June May 1857 (1). 1988 (2); A. Segura Zubizarreta, 24 July 1966 (1), 27 July 1966 (1), 2 Sep 1972 F. Egido, 14 July 2009 (4), 28 Aug 2007 (4), 30 June 2009 (4); fr. Elías, (1), 11 Aug 1974 (1), 11 Aug 1990 (1); fr. Sennen, 401 (1); fr. Sennen & fr. Elías, 25 July 1909 (1), 4 July 1906 (1); F. Esteve, 19 Oct 1947 (1). 25 July 1906 (1); P. F. Serdjukow, 30/348 (3); F. J. Silva-Pando, 38 (4), 4034 (4), J. L. Fernández Alonso, 09 Aug 1982 (1); F. J. Fernández Díez, 3 Aug 1979 4471 (4); P. E. E. Sintenis & M. B. G. Sintenis, 387 (3); V. Skřivánek, 1 Sep (1); A. Fiori, 23 May 1898 (5); P. Font i Quer, 11 June 1932 (1), 29 June 1948 1967 (3); A. St.-Yves, 4 July 1899 (1); W. Steinitz, 5 Sep 1880 (3); S. Stoyanov (1); P. Font i Quer & W. Rothmaler, 14673 (1); A. Forestier, July 1849 (1). & V. Goranova, 17 Aug 2013 (3); A. Susanna et al., 361 (1); A. Susanna & A. P. Galán Cela & G. López, 3198 (1); E. F. Galiano, July 1961 (1); M. Pons Sorolla, 641 (1). Gandoger, 579 (4); R. García, 1356 (1); A. García-Villaraco, 19 July 1983 (1); J.A. Tauscher, 29 July 1873 (3), 29 July 1874 (3); Thélesphose, 30 June 1875 P. Geissler, 15 July 1972 (1); D. Gerevanov, 8 July 1914 (3); P. Gesti, 26 May (1); F. Trèmols, July 1884 (1); F. A. Tscherning, 6 Aug 1896 (6). 1998 (1), 3 July 1996 (1); L. Girod, July 1913 (1); F. Gómez Vigide, 14 July 1983 I. K. Urumoff, 129 (6). (4), 20 July 1985 (4), 27 May 1991 (4); F. Gómiz, 12 July 1995 (4), 18 Sep 1988 E. Valdés-Bermejo et al., 11195 (4); S. M. Voronov & A. F. Shcherbakov, (1); D. Grecescu, July 1892 (6); G. P. Grinţescu, 24 Aug 1923 (3); Ch. 16 Sep 2003 (3); B. Vreš & I. Dakskobler, 20 June 2011 (5). Guebhard, 292 (2); W. Gugler, 26 July 1904 (3). J. Walter & E. Vitek, 13–0415 (3), 13–0416 (3), 13–0417 (3); L. F. 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