Abstract
Orobanche ritro, which is frequently considered a synonym of O. elatior, is lectotypified here from Grenier’s original material in the herbarium of the Muséum National d’Histoire Naturelle, Paris (P). A morphological description focusing on the particular characters which distinguish it from related species is provided. The conspecificity of O. echinopis with O. ritro is pointed out. O. ritro occurs in central and southern areas of Europe, from north-east Spain to south-west European Russia and warrants recognition at species level.
Résumé
Orobanche ritro, souvent considéré comme un synonyme d’O. elatior, est lectotypifié ici à partir du matériel original de l’herbier du Muséum national d’histoire naturelle, Paris (P). Nous fournissons sa description morphologique mettant l’accent sur les caractères particuliers qui le distinguent des autres espèces proches. La conspécificité d’O. echinopis avec O. ritro est démontrée. O. ritro habite dans les zones centrales et méridionales de l’Europe, du nord-est de l’Espagne au sud-ouest de la Russie européenne, et mérite d’être reconnu comme espèce.
Introduction
Orobanche ritro was first described by Grenier and Godron in Flore de France 2: 635 (1853) on material collected by Grenier in Gap and by Boullu in Guillestre (France), parasitizing on roots of Echinops ritro L. (Asteraceae). Reichenbach (Citation1862: 97) confirmed its presence in France and Loscos (1878–1880: 125) deemed it a non-rare species in Castelserás (Teruel, Spain). Also, Saint-Lager (Citation1883: 610) initially reported it as O. rhytrosepiphyta in southern France and subsequently (Saint-Lager, Citation1889: 649) designated it as O echinopophya, both names being superfluous.
Pančić (Citation1868: 80) reported its presence in Romania by referring to the same host (“…in radice Echinopis Ritronis (?)”[sic.]) in describing O. echinopis Panc. Nyman (Citation1881: 560) deemed O. ritro dubious (?) in southern France and recognized O. echinopis, albeit as a subspecies subordinated to O. salviae F.W. Schultz.
For no apparent reason, Beck (Citation1890: 171) considered it a form subordinate to O. major L., a taxonomic rank he maintained (Beck, Citation1930: 251) in reporting its presence in France (Dép. Hautes Alpes, Gap, le Devès bei Rabou, Guillestre) and Italy [Im Ceviere Tale der Cottichen Alpen (Rostan)]. However, Beck (Citation1893: Citation1083; 1930: 250) referred to O. echinopis as a mere synonym for O. major.
Willkomm (Citation1893: 189) used the previous reference by Loscos to grant this taxon the status of variety [O. major β ritro (Gren. & Godr.) Willk.]. Lázaro-Ibiza (Citation1896: 810) also dealt with it as a variety. Rouy (Citation1909: 181) initially considered it a mere race and subsequently (Rouy, Citation1927: 204) a subspecies. Douin (Citation1926: 97) also deemed it a subspecies, subordinate to O. major.
However, some prominent botanists have refused to consider O. ritro an autonomous taxon and dealt with it as a synonym for O. major (Jackson, Citation1895: 374; Fiori, Citation1902: 480; Coste, Citation1937: 70; Borza, Citation1949: 253, sub O. echinopis; Fournier, Citation1977: 199) or O. elatior Sutton (Chater & Webb, Citation1972: 359; Pignatti, Citation1982: 614; Greuter et al., Citation1989: 260; Zázvorka, Citation1997: 513–514; Foley, Citation2001: 71; Domina & Raab-Straube, Citation2009). Also, other authors have included the genus Echinops L. among the hosts for O. major (Novopokrosky & Tzvelev, Citation1958: 132; Mądalski, Citation1967: 51) or O. elatior (Gilli, Citation1974: 496; Tzvelev, Citation1981: 334; Uhlich et al., Citation1995: 135) without referring to O. ritro, not even as a synonym.
Recently, Carlón et al. (Citation2003: 31) reinstated its varietal rank and Pujadas-Salvà (Citation2005) its species rank.
Finally, Zázvorka (Citation2010: 82) includes it as a synonym for O. kochii FW. Schultz, without ever having studied any original material from Grenier. The images of the lectotypes of O. kochii and of O. echinopis (= O. ritro) chosen by Zázvorka (Citation2010: 102, Fig. 12; 103, Fig. 13) have a dissimilar appearance and confirm that these two taxa are different rather than conspecific. To the naked eye, O. kochii has a lax inflorescence and O. echinopis a dense inflorescence. In addition, closer inspection of O. ritro reveals other important morphological differences from O. kochii regarding calyx shape, corolla shape and color, and number of flowers on the inflorescence, among others. Moreover, neither Schultz (Citation1847), nor Maly (Citation1842, Citation1851, Citation1857) cited Echinops as a possible host of O. kochii.
This paper revises the taxonomic status of O. ritro and examines its morphology in contrast with O. salviae and with O. elatior (= O. major L., p.p. [nom. rejic.]), to which it has previously been subordinated, and also with O. icterica Pau, with which it can co-exist and may also be confused. O. ritro is here lectotypified; its distribution, habitat and phenology are provided.
Material and methods
Orobanche ritro specimens from the COI, JACA, MA, P, PRC, W and WU herbaria, and the private herbarium of J.V. Ferrández (JVF), were studied (see Appendix 1). These specimens were also examined in search for original material from Grenier and Godron.
Pančić’s, O. echinopis type material from the BEOU and PAD herbaria, was also examined in order to confirm whether this taxon is in fact conspecific with O. ritro.
Reported data from Koch (Citation1833: 458-459), Reichenbach (Citation1862: 96; t. 179, MDCCC), Beck (Citation1930: 267), Gilli (Citation1974: 501-502) and Chater & Webb (Citation1972: 293) were used to distinguish and separate O. ritro from O. salviae.
The O. ritro specimens were compared with O. elatior and O. icterica material from the ABH, BC, COA, JACA, K, MA, MUB, VAB, VAL herbaria (cf. Pujadas-Salvà et al., Citation2005). Reported data from Grenier & Godron (Citation1853) and Pančić (Citation1868) were also used in the analysis.
Results
Lectotypification
Orobanche ritro is lectotypified here on the basis of material reported as collected in the places named in the protologue, namely: “collines sèches des environ de Gap, en allant à Rabou et à la Grangette (Gren.); Guillestre (Boullu)”. Only two sheets with material from such places could be found in the P and W herbaria, namely: “Gap, 1857, Grenier, W” and “Rabou près Gap, sur l’Echinops ritro, 14-VIII-1848, Grenier, P”. Both sheets contained material collected by Grenier in Gap (a place in southern France named in the protologue), but only the latter material was collected before the description of the species in 1853.
Material conserved as a «topo» type in the P herbarium [viz. Orobanche ritro Gren. / Rabou Lagarde juillet / style garni de quelques poils glanduleux / au dehors du stigmate qui est jaune / Blanc 1854] is not used for lectotypification here, however, as this locality was not mentioned in the protologue.
Because no additional testimonies of the material studied by Grenier could be found, we chose to use the sheet voucher kept as type material in the P herbarium and selected as lectotype here.
Orobanche ritro Gren. & Godr., Fl. France 2: 635. 1853 ≡ Orobanche rhytrosepiphyta St. Lager, Cat. Flore du Bassin du Rhône: 610. 1883 [nom. illeg.] ≡ Orobanche echinopophya St. Lager, in Cariot, Étude des fleurs, éd. 8, II: 649. 1889 [nom. illeg.] ≡ Orobanche major f. ritro (Gren. & Godr.) Beck, Biblioth. Bot. 19: 171. 1890 ≡ Orobanche major var. ritro (Gren. & Godr.) Willk., Suppl. Prodr. Fl. Hispan.: 188. 1893 ≡ Orobanche major race ritro (Gren. & Godr.) Rouy, Fl. France 11: 181. 1909 ≡ Orobanche major subsp. ritro (Gren. & Godr.) Douin, Fl. Compl. France, Suisse & Belg. 8: 97. 1926 – Ind. loc.: «Hab. Sur les racines de l’Echinops ritro; collines sèches des environs de Gap, en allant à Rabou et à la Grangette (Gren.); Guillestre? (Boullu) Juillet.»
-Lectotype (designated here): Orobanche ritro - nob. / couleur jaune pâle. / Rabou près Gap. Sur l’Echinops ritro / 14 août 1848 / Grenier / (handwritten by Grenier)
In addition, the sheet contains the following handwritten label by Reuter:
Belle espèce que je ne connais pas! / probablement nouvelle. / (Reuter) 1851 / (P!)
= Orobanche echinopis Pančić in Österr. Bot. Zeit.: 80 (1868) [“Habitat copiosissima in arena mobili ad Vakarec et Fontina fetje agris Romanorum in Banatu parasitica in radice Echinopis Ritronis (?). Floret Julio”]
= Orobanche salviae subsp. echinopis (Pančić) Nyman, Consp. Fl. Eur.: 560 (1881)
| - | Orobanche major auct. p.p. (eg. Fiori, Fl. Italia 2: 480. 1902; Coste, Fl. Descr. France 3: 70. 1937) non L. | ||||
| - | Orobanche elatior auct. p.p. (eg. Chater & Webb, Fl. Eur. 3: 359.Citation1972; Greuter et al., Med-Checklist 4: 260. Citation1989) non Sutton | ||||
| - | Orobanche kochii auct. p.p. (eg. Zázvorka, Acta Mus. Morav., Sci. Biol. 95(2), 77-119. 2010) non F.W. Schultz | ||||
[Iconography. Fig. (Photography); Reichenbach (Citation1862: t. 170, MDCCXCI); Pujadas-Salvà (Citation2005) (Photography).
Fig. 1 Orobanche ritro. France, Rabou près Gap, sur l’Echinops ritro, 14 VIII 1848, Grenier (P). Lectotype.
Morphological description
The description of Orobanche ritro (Grenier and Godron Citation1853: 635) is quite comprehensive and includes several distinct features such as its having bracts smaller than its corolla; ovate, bifid, several-nerved sepals; tubular, slightly pubescent to glandular, sulphur-yellow coloured corolla with a near-straight back; villous filaments in the lower half and sparse glandular hairs in the apex. However, studying the material examined here allowed us to discover various additional morphological features which may further facilitate its identification:
Plant 20–56 cm. Stem 4–10 mm in diameter at half height, robust, scarcely swollen at base [8–10(15) mm in diameter], shortly glandular hairy with rusty glandular short hairs 0.1–0.5(0.6) mm in size, creamy yellow stem (cream-like with a red ting when dry). Basal leaves 8–12(15) × 5–8 mm, triangular to deltoid in shape, sub-imbricate to sparse; upper leaves 12–20(25) × 3.5–6(9) mm, widely lanceolate to ovate, erect–patent to patent, rusty glandular hairy, with short hairs (0.2–0.3 mm, up to 0.7 mm at base). Inflorescence 7–22 × 2.9–3.5 cm, cylindrical, rounded, occasionally sub-comose apex, dense, with many flowers. Bracts (12)13–20(24) × (3.5)4–6.5 mm, smaller than the flowers, but equalling the corolla lower lip, widely lanceolate, deflected, with rusty glandular hairs 0.1–0.6 mm, the longest lying near bract bases. Calyx segments, 10–16(19) × (3.5)4–5.5 mm, several-nerved with prominent veins, segments ± ovate, broad-based, free, separated or contiguous, rarely connate abaxially (some inflorescences contain both free and connate calyx segments), two short teeth, unequal to sub-equal, shortly bipartite to bifid, occasionally emarginate, teeth smaller or as long as one-half the segment length [1/2–1/3(1/5) segment length], triangular teeth with 2–3 veins running lengthwise, calyx scarcely hairy to glabrescent at base, with glandular hairs up to 0.2 mm, hairy teeth, with glandular hairs c. 0.2 mm long and up to 0.6 mm at the margin, creamy coloured calyx, occasionally red-tinged. Corolla 17–23 mm, erect to erect–patent, rarely patent, tubular, generally straight, straight or occasionally slightly curved back, glabrescent at the lower half, with short glandular hairs (< 0.2 mm), hairy in the upper half with glandular hairs 0.1–0.3 mm, more abundant especially on the back, colour pale yellow, sulphur-yellow or creamy, limb veins rarely tinged reddish-brown (straw-coloured corolla with red-tinged limb when dry). Corolla margin erose, glabrous or glabrescent, with scant, sparse hairs c. 0.1 mm long; widely bilobate, scarcely emarginated upper lip with porrect, occasionally patent lobes, and deflected, trilobate lower lip with sub-equal lobes, the central lobe slightly larger than the lateral ones, widely obovate. Obliquely inserted filaments, adaxial filaments inserted (3)4.5–6 mm above corolla base, abaxial filaments inserted at (1.5)2–4 mm, slightly geniculate, densely villous in the basal ½– ¾, with long hairs 0.3–0.7 mm, and glandular hairs, c. 0.1 mm in size, below anthers, occasionally sub-glabrous. Anthers 1.5–1.9 mm (apiculum 0.2–0.4 mm), oblong, papillate hairs on suture, yellowish or creamy coloured. Ovary glabrescent with scant hairs (< 0.2 mm) spread at apex, occasionally glabrous. Style with glandular hairs (< 0.3 mm), more abundant below stigma, occasionally sub-glabrous. Stigma bilobate, included, bright yellow.
Habitat, distribution and phenology
Orobanche ritro grows in open spaces of xerophytic bush areas and also, occasionally, in degraded scrubland, meadows, dry pasture, sub-ruderalized zones under a generally dry, sunny climate, occasionally on sandy or gypsiferous soils. The taxon is known to parasitize roots of Echinops ritro (Asteraceae), and has also been encountered on E. sphaerocephalus L. (cf. Saint-Lager Citation1889: 650). In addition, Beck (Citation1890: 36) named E. bovei Boiss. (apud Debeaux) and E. ruthenicus M. Bieb. (apud Borbás) as hosts of O. major L, when in fact they are highly likely to be O. ritro hosts. This species occurs at heights from 300 to 1300 m and flowers from May to August. It can be found in areas of central and southern Europe from the north-east of the Iberian Peninsula (Spain) to the south-west of Russia, in the Volgograd Region (European Russia).
Relationship to other species
Orobanche ritro exhibits a number of morphological characteristics not present jointly in any other taxon known to date.
O. ritro can be easily distinguished from O. salviae mainly by its calyx segments shortly bipartite to bifid, with triangular teeth (vs. entire or unequally bidentate, with acuminate often sub-filiform teeth in O. salviae); corolla straight or occasionally slightly curved, tubulose (vs. corolla leaning forward conspicuously curvate, tubulose slightly enlarged above the insertion of the filaments); corolla margin glabrous or glabrescent (vs. corolla margin hairy); ovary glabrescent (vs. ovary very hairy). Consequently, we consider that these are two different taxa and that the subordination by Nyman (Citation1881: 560) of O. ritro to O. salviae (O. salviae subsp. echinopis) is not consistent. In addition no other author has considered them as related species.
Compared to O. elatior, O. ritro has a thinner, only slightly swollen stem at base (8–10 mm); sub-imbricate to sparse basal leaves; widely lanceolate to ovate upper leaves; free, rarely connate calyx segments that are shortly bipartite to bifid, occasionally emarginate, and have a scarcely hairy to glabrescent base with glandular hairs up to 0.2 mm (Fig. ); corolla with a straight to slightly curved dorsal line and short glandular hairs up to 0.3 mm; staminal filaments with hairs 0.3–0.7 mm at base; and usually shorter anthers (1.5–1.9 mm) (Table ).
Fig. 2 Calyx segments. Orobanche ritro: a) France, Rabou (P) (lectotype), b) Spain, Huesca, Ortilla, (JACA 344296); c) Romania, Ad vallum Romanorum legionum Serbio-banaticae, (WU 771). O. elatior: d) England, Hants, Bury Hill, Andover (K); e) Hungary, National Park of Hortobágy (COA 16491); O. icterica: f) Spain, Alicante, Cap de les Hortes (COA 25532); g) Spain, Almería, Vélez Blanco (COA 37658).
Table I. Main differences between O. ritro Gren. & Godr., O. elatior Sutton and O. icterica Pau.
Orobanche ritro can also be considered close to O. icterica on the grounds of some similarities in appearance, spike density, leaf size, corolla length, stigma colour and, occasionally, overall plant colour. However, O. ritro, can be easily distinguished by its having sub-imbricate to sparse basal leaves; a usually longer [10-16(19) mm] calyx, shortly bipartite to bifid, occasionally emarginate calyx segments, and scarcely hairy to glabrescent base with glandular hairs up to 0.2 mm (Fig. ); tubular corolla with a straight to slightly curved back with short (< 0.3 mm) glandular hairs; densely pubescent staminal filaments with hairs 0.3–0.7 mm at base, adaxial filaments being inserted at slightly greater heights [(3)4.5-6 mm above corolla base] and non-auriculate abaxial filaments with a straight margin; and usually shorter (1.5–1.9 mm) anthers (Table ).
There are also some differences between host plants. Thus, O. ritro parasitizes Echinops species, whilst O. elatior and O. icterica tend to parasitize various Centaurea and Rhaponticoides (Asteraceae) species.
Although it shares some characteristics with O. elatior and O. icterica, major morphological differences between these taxa warrant recognizing of O. ritro as a separate species.
Acknowledgements
The author would like to thank the keepers and staff of the ABH, BC, BEOU, COI, JACA, JVF, K, MA, MUB, P, PAD, PRC, VAB, VAL, W and WU herbaria for providing the specimens used in this study, and Enrique Triano for his helpful comments on the manuscript.
Related Research Data
References
- Beck , G. 1890 . Monographie der Gattung Orobanche . Biblioth. Bot , 19 : 1 – 275 .
- Beck , G. 1893 . Flora von Nieder-Österreich . Carl Gerold’s Sohn, Wien , 2 ( 2 ) : 895 – 1396 .
- Beck, G., 1930. Orobanchaceae. In Das Pflanzenreich, A. Engler (ed.), Engelmann, Leipzig, 96 (IV.261), 348 p.
- Borza , A. 1949 . Conspectus Florae Romaniae . Cartea Romaneasca, Cluj , 2 : 161 – 360 .
- Carlón , L. , Gómez-Casares , G. , Laínz , M. , Moreno-Moral , G. and Sánchez-Pedraja , Ó. 2003 . Más a propósito de algunas Orobanche (Orobanchaceae) del norte y este de la Península Ibérica . Documentos Jard. Bot. Atlántico (Gijón) , 2 : 1 – 45 .
- Chater , A.O. and Webb , D.A. 1972 . “ Orobanche ” . In Flora Europaea , Edited by: Tutin , T.G. Vol. 3 , 286 – 293 . Cambridge : Cambridge University Press .
- Coste, H., 1937. Flore descriptive et illustrée de la France, de la Corse et des contrés limitrophes. Librairie Scientifique et Technique, Albert Blanchard, Paris, 3, 729 p.
- Domina, G. and E. von Raab-Straube, 2009. Orobanchaceae. In Euro+Med Plantbase - the information resource for Euro - Mediterranean plant diversity. Retrieved 19 March 2010 from http://ww2.bgbm.org/mcl/home.asp.
- Douin, R., 1926. Orobanche. In Flore complète illustrée en couleurs de France, Suisse et Belgique. G. Bonnier and R. Douin (eds.), Librairie Générale de l’Enseignement, E. Orlhac, Paris, 8 (78), 91–100.
- Fiori, A., 1902. Flora analitica d’Italia. Tipografia del Seminario, Padova, 2, 492 p.
- Foley, M.J.Y., 2001. Orobanche. In Flora iberica, J. Paiva et al. (eds.), Real Jardín Botánico, Madrid, 14, 32–72.
- Fournier, P., 1977. Les quatre flores de la France, Corse comprise, 2e éd., Editions Lechevalier, Paris, 1, 1106 p.
- Gilli, A., 1974. Orobanchaceae. In Gustav Heg, Illustrierte Flora von Mitteleuropa, 2nd ed. D. Hartl and G. Wagenitz (eds.), Paul Parey, Berlin and Hamburg, 6, 1, 470–505.
- Grenier , J.C.M. and Godron , D.A. 1853 . Flore de France . J.B. Baillière. Paris , 2 : 393 – 760 .
- Greuter, W.R., H.M. Burdet and G. Long, 1989. Med-checklist, (Lauraceae-Rhamnaceae). Conservatoire et Jardin Botaniques de la Ville de Genève, Genève, 4, 458, CXXIX p.
- Jackson, B.D., 1895. Index Kewensis, Clarendon Press, Oxford, 2, 1299 p.
- Koch, W.D.J., 1833. Orobanche. In Deutschlands Flora, ed. 3. J.C. Röhling and W.D.J. Koch (eds.), Friedrich Wilmans, Frankfurt am - Main, 4, 427–469.
- Lázaro-Ibiza, B., 1896. Compendio de la Flora Española, Librería de Hernando y Compañía, Madrid, 2, 1038 p.
- Loscos, F., 1878–1880. Tratado de plantas de Aragón, Parte segunda. Establecimiento Tipográfico del Hospicio, Madrid, 249 p.
- Mądalski, J., 1967. Orobanchaceae. In Flora Polska – Rosliny Naczyniowe I Ziem Osciennych. B. Pawlowski (ed.), Panstwowe Wydawnictwo Naukowe, Warszawa-Krakow, 11, 25–52.
- Maly, J., 1842. Die Orobanchen der Gratzer Gegend. Flora (Regensb.) 25(1), Beibl. (8), 124–127.
- Maly , J. 1851 . Ueber Orobanche Kochii Schultz . Oesterr. Bot. Wochenbl , 1 ( 42 ) : 337 – 338 .
- Maly , J. 1857 . Ueber Orobanche Kochii Schultz . Oesterr. Bot. Wochenbl , 7 ( 18 ) : 143
- Novopokrosky, I.V. and N.N. Tzvelev, 1958. Orobanchaceae. In Flora URSS, English edition, 2000. V.L. Komarov (ed.), Bishen Singh Mahendra Pal Singh and Koeltz Scientific Books, Mosqua and Leningrad, 23, 19-117.
- Nyman , C.F. 1881 . Conspectus Florae Europaea . Typis officinae Bohlinianae, Orebro , 3 : 493 – 677 .
- Pančić , J. 1868 . Zur Flora des Banates . Österr. Bot. Zeit , 3 : 78 – 84 .
- Pignatti, S. 1982. Flora d’Italia. Edagricole, Bologna, 2, 732 p.
- Pujadas-Salvà, A.J., 2005. Orobanche. In: Atlas de la Flora de Aragón. D. Gómez (ed.), Retrieved 18 January 2010 from http://www.ipe.csic.es/floragon.
- Pujadas-Salvà, A.J., L. Plaza-Arregui, E. Triano-Muñoz and J.A. Algarra-Ávila, 2005. Orobanche icterica (Orobanchaceae) revisited. Bot. J. Linn. Soc. 148, 117–124.
- Reichenbach, H.G., 1862.- Icones florae Germanicae et Helveticae. Typogr. Guilielmi Baensch, Lipsiae, 20, 125 p.
- Rouy, G., 1909. Flore de France. Les Fils d’Émile Deyrolle, Paris, 11, 429 p.
- Rouy, G., 1927. Conspectus de la flore de France. Paul Lechevalier, Paris, 319 p.
- Saint-Lager, J.B., 1883. Catalogue des plantes vasculaires de la flore du bassin du Rhône. Librairie H. Georg, Lyon, 886 p.
- Saint-Lager, J.B., 1889. Orobanche. In Étude des fleurs. Botanique élémentaire, descriptive et usuelle, 8ª ed. A. Cariot and J.B. Saint-Lager (eds.), Vitte and Perrussel, Lyon, 2, 647–652.
- Schultz, F.W., 1847. Orobanche kochii, eine neue deutsche Art. Flora (Regensb.) 30(5), 65–67, tab. 2.
- Tzvelev, N.N., 1981. Orobanchaceae. In Flora parties europeae URSS. A. Federov (ed.), Nauka, Leningrad, 5, 317–336.
- Uhlich, H., J. Pusch and K.J. Barthel, 1995. Die Sommerwurzarten Europas. Westarp Wissenschaften, Magdeburg, 235 p.
- Willkomm, H.M., 1893. Supplementum prodromi florae hispanicae. E. Schweizerbart (E. Koch), Stuttgart, 370 p.
- Zázvorka, J., 1997. Orobanche. In Flóra Slovenska. K. Goliašová (ed.), VEDA, Bratislava, 5(2), 478–529.
- Zázvorka , J. 2010 . Orobanche kochii and O. elatior (Orobanchaceae) in central Europe . Acta Mus. Morav., Sci. Biol. , 95 ( 2 ) : 77 – 119 .
Appendix 1
Herbarium specimens of Orobanche ritro Gren. and Godr. studied FRANCE: Rabou près Gap, sur l’Echinops ritro, 14-VIII-1848, Grenier, s.n., P; Rabou Lagarde, VII-1854, Blanc, s.n., P; Gap, 1857, Grenier, s.n., W 14149; Gap, Coréo, chemin de Rabou, 21-VII-1861, Burle, s.n., W; HUNGARY: Inter Lepsény et Siófok ad Pelsionem lacum in radicibus Echinopis, s/f, Staub, s.n., COI; PRC; WU 667 (sub O. major); Liofok [Siófok] (comté de Wesprem), prés sablonneux sur les bords du Balaton, parasite sur les Echinops, 16-VII-1884, Hermann, s.n., MA 170768; WU 1628 (sub O. echinops); Sivkof, Veszprim, 16-VII-1884, Hermann, s.n., PRC; WU 1108 (sub O. echinopis); Erin Cotta Albenoi e vineis e radice Ulmus campestris, 27-VII-1869, J. Tauscher, s.n., WU (sub O. elatior Sutton by Beck); ROMANIA: Agri Romanorum in Banatu loco Fontina fetje dicto supra Echinops, VII-1857, J. Pančić, s.n., BEOU (sub O. echinopis, corrected by Beck as O. major) [digital image]; Agri Romanorum loco Vakarec dicto in Banatu, VII-1867, J. Pančić, s.n., BEOU (sub O. echinopis corrected by Beck as O. major) [digital image]; In arenosis ad Vakarec, Banatu, VII, J. Pančić, s.n., PAD 7791 (sub O. echinopis) [digital image]; Ad vallum Romanorum legionum Serbio-banaticae, VI-1869, Sonklar, s.n., WU 771 [sub O. ritro G. G. / Echinopis Pančić, corrected by Beck as gr. O. major L.]; RUSSIA: Fl. Sareptana, 1853, Becker, s.n., W 42435 (sub O. elatior); SERBIA: Prope oppidum Nisch, VII-1883, S. Petrovic, s.n., PRC (sub O. echinopis); SPAIN: Huesca, Ortilla, cerca del corral Purrudán, 30TXM9969, 555 m, 8-VI-1996, J.V. Ferrández, s.n., JACA 344296 (sub O. lutea Baumg); Huesca, Barbastro, Pr. Torre Joaquina, yesos, sobre Echinops ritro, 31TBG6452, 360 m, 22-V-2003, J.V Ferrández, s.n., JVF 13303 (sub O. cf. elatior); Huesca, Almunia de San Juan, al E de Valmayor, yesos, claro de carrascal y sarda, sobre Echinops ritro, 31TBG7445, 400 m, 21-VI-2004, J.V. Ferrández, s.n., JVF 10404; Zaragoza, Calatayud, Marivella, 25-V-1908, Vicioso, s.n., MA 436911.