FRAGA-ARGUIMBAU, P. & J. A. ROSSELLÓ (2011). Euphorbia nurae P. Fraga & Rosselló (Euphorbiaceae), a new species from Minorca (Balearic Islands). Candollea 66: 181–190. In English, English and French abstracts.
Euphorbia nurae P. Fraga & Rosselló (Euphorbiaceae sect. Cymatospermum (Prokh.) Prokh.) is described as a new species from coastal sites of Minorca (Balearic Islands). Morphological features suggest that Euphorbia exigua L., Euphorbia dracunculoides Lam. and Euphorbia sulcata Loisel. are the most closely related taxa, but the new species can be easily discriminated by several morphological characters.
Introduction
Euphorbia L. is a world wide distributed genus comprising more than 2000 species showing an astonishing diversity in life forms (Webster, 1994). Native European species mostly belong to subgen. Esula Pers. (Benedí & al., 1997). Within this subgenus sect. Cymatospermum (Prokh.) Prokh. comprises exclusively annual species. Overall morphology between taxa is similar and discriminating characters used to identify species are mainly based on seed features, including seed size and episperm ornamentation (Molero & al., 1996).
The Balearic Islands harbour 22 species of genus Euphorbia including narrow endemic taxa like E. fontqueriana Greuter, E. maresii Knoche and E. margalidiana Kühbier & Lewej. (Benedí & al., 1997). Most of the non-endemic Euphorbia diversity found in the archipelago is from Minorca, where 15 species have been recorded (Fraga & al., 2004), of which four entities (E. exigua L., E. falcata L., E. medicaginea Boiss., and E. peplus L.) belong to sect. Cymatospermum. Only Euphorbia exigua and E. peplus are relatively common in the island whereas E. falcata and E. medicaginea have been recorded from a few localities (Fraga & al., 2004). In Minorca and elsewhere in the distribution area, E. exigua shows, a remarkable polymorphism regarding vegetative features (e.g., branching pattern and leaf shape) (Pignatti, 1982; Meikle , 1985; Bolòs & Vigo, 1990; Benedí & al., 1997). Some extreme morphotypes have been recognized (e.g., var. acuta L., var. retusa L., var. truncataW. D. J. Koch; see Willkomm & Lange, 1880; Fiori, 1925; Vindt, 1953; Bonafè, 1979; Meikle, 1985), but there is not evidence that these variants correlate with the variation detected in seminal features (Molero & al., 1996).
During field work aimed to increasing the floristic knowledge of Minorca we have observed several coastal populations of annual plants that were related to Euphorbia exigua but showed conspicuous differences in plant height, branching pattern, leaf arrangement, female inflorescence, seed size and testa ornamentation. Comparison of these forms with herbarium specimens of E. exigua from non-Balearic sites corroborated their unique nature. They are presented in this paper as a new species, E. nurae.
Material and methods
Morphological observations were made from living plants (in the field and from cultivated plants) and from herbarium specimens (MA, BC, and the personal herbarium of Fraga-Arguimbau). Further, information from taxonomic and floristic sources covering sect. Cymatospermum (Molero & al., 1996), or dealing with the genus Euphorbia in western Mediterranean areas (Moris , 1859; Battandier & Trabut, 1888; Coste, 1903; Fiori, 1925; Vindt, 1953; Quezel & Santa, 1963; Smith & Tutin, 1968; Pignatti , 1982; Meikle 1985; Bol òs & Vigo, 1990; Benedí & al., 1997; Jeanmonod & Gamisans, 2007) was compiled for comparative purposes. The shape and ornamentation of the seeds were observed using stereoscope and scanning electron microscopes (SEM). For SEM they were mounted on stubs with double-sided carbon adhesive tape, gold coated and observed under Jeol Scanning Electron Microscope at 10 kV accelerating voltage. The method of Molero & al. (1996 ) was used to assess the presence of mucilage compounds on the seed episperm.
Results
Euphorbia nurae P. Fraga & Rosselló, spec. nova (Fig. 1A, 1B, 1C–5)
Typus: BALEARIC ISLANDS. Minorca: Es Berrecks de Santa Anna, Ciutadella de Menorca (31SEE807212), ephemeral pastures on thin, limestone soils, 30 m, 31.III. 1996, P. Fraga s.n. (holo-: VAB [202781]) (Fig. 1A,1B, 1C, 2).
Planta annua, minuta (ad 0.5–2.5 cm alta), at Euphorbia exigua, E. dracunculoides et E. sulcata similis, sed prostrata, folia imbricata, pauci carnosa, cyathiis parvulis (usque ad 1 mm); stylus circa 0.2 mm longus, adpresus; semina 0.3–0.6 × 0.6–0.8 mm, parce tuberculata, sine elaiosoma differt.
Annual plant, 0.5–1.5(-2.5) cm high, glabrous, light green to glaucous up to anthesis, turning reddish or orange at fruiting. Usually with a single stem or multi-stemmed, with a few radial branches starting at the base; stems prostrate (except at fruiting), 0.5–5 cm long, 0.2–0.5 mm thick, usually sinuate. Caulinar leaves 0.4–0.8 × 0.8–2(-3) mm, densely imbricate, sessile, slightly carnosulate, linear to linear-spatulate, widest towards the apex; leaf blade carinate at the apex, apex rounded to obtuse, margin entire. Pleiochasial bracts 0.6–0.9 × 2–3.5 mm, linear-lanceolate, attenuate at the apex, apex obtuse to subacute, not rounded, limb plane, margin entire. Dichasial bracts free, 0.9–1.5 × 1.3–3.8 mm, narrowly triangular to triangular-lanceolate, apex obtuse to rounded, margin entire. Dichasial cymes 1–3, up to 5(-10) mm long, not bifurcating. Cyathia glands small (up to 1mm long), glabrous, each cyathium comprising two oblong and two semicircular yellowish to red nectaries, appendiculate; nectaries with parallel to slightly divergent, short appendages (up to 0.2 mm long), obtuse and rounded at the apex. Carpels with short stigmas (up to 0.2 mm long), bifurcate, usually recurved and appressed. Capsule pedicelated (pedicel up to 0.5 mm long, recurvate), 3-angular, ca. 1.1–1.4 × 1.2–1.5 mm, subovoid, sulcate, glabrous; cocci smooth, slightly granulose near the keels. Seeds whitish-grey, not mucilaginous, 0.6–0.8 × 0.3–0.6 mm, ovoid or subglobose, with a very marked longitudinal furrow, papillate, with poorly differentiated tubercles; caruncle completely absent, being only visible the insertion point.
Etymology. — From Nura, an ancient name of Menorca.
Phenology. — Euphorbia nurae flowers between February and April (exceptionally up to mid May) (Fig. 3A). Fruiting individuals have been seen in April and May (Fig. 3B).
Habitat and distribution. — Currently, E. nurae is known from the coastal areas of south Minorca and a single population is present in quaternary sandy soils from the north (Fig. 4). It grows in sandy or clayish calcareous soils formed by the dissolution of Miocenic rocks. The low nutrient concentration and scarce hydric content of these soils favour the establishment of ephemeral communities dominated by therophytes and geophytes of small size. There, E. nurae shows a patchy distribution (isolated individuals or up to 30 plants per square meter) together with Valantia muralis L., Trifolium scabrum L., Plantago weldenii Rchb., Senecio rodriguezii J. J. Rodr., Medicago littoralis Loisel., Medicago minima (L.) L., Desmazeria marina (L.) Druce, Lagurus ovatus L., Hypochoeris achyrophorus L., Sagina maritima G. Don, Bupleurum semicompositum L., Romulea assumptionis Garcias Font, Sedum rubens L., Filago pyramidata L., Valerianella microcarpa Loisel., Linum strictum L., Sherardia arvensis L., Plantago bellardii All., Allium chamaemoly L., Merendera filifolia Camb., Hyoseris scabra L., Asteriscus aquaticus (L.) Less., Galium murale (L.) All., and Campanula erinus L. Concerning its conservation status, most of the known populations are located within protected areas, mainly included in the Natura 2000 network, and only a few of them are at risk due to landscape transformation by construction of touristic buildings. Nevertheless, its narrow distribution range and small populations suggest a classification as Vulnerable (VU), under criteria D2, according to the criteria of the IUCN Red List (IUCN, 2001).
Specimina visa. — MINORCA: De Son Xoriguer a Son Saura, 3.V.1999, P. Fraga s.n. (MA [625575]); Son Olivaret, 3.V.1999, P. Fraga s.n. (MA [624577]); Mallaui, Sa Marjal Nova, 30.III.2004, P. Fraga s.n. (Herbarium Fraga-Arguimbau); Binicalaf Nou, 1.IV.2004, P. Fraga s.n. (Herbarium Fraga-Arguimbau); Marina de s'Arena, Capifort, 2.IV.2004, P. Fraga s.n. (Herbarium Fraga-Arguimbau); Son Saura de Ciutadella, 3.IV.1999, P. Fraga s.n. (Herbarium Fraga-Arguimbau).
Notes. — Morphological features (see Fig. 1, 4) suggest that Euphorbia nurae is closely related to E. exigua L., E. dracunculoides Lam., and E. sulcata Loisel. from which it can be easily differentiated by a combination of vegetative and reproductive features (Table 1, Appendix 1). The superficial similarity shown between the new species and these widespread Mediterranean species could be related to convergence, or to the retention of plesiomorphic characters. Furthermore, the significant reduction of vegetative features in these annual plants makes their recognition difficult unless key diagnostic characters, like seeds, are present.
Despite the acknowledged polymorphism concerning the shape of caulinar leaves in species of sect. Cymatospermum (Molero & al., 1996) we have noted a contrasting leaf shape that is constant in all known populations of Euphorbia nurae so far analyzed: they are linear-lanceolate to linear spatulate, slightly carnulose and carinate. By contrast, herbarium specimens from related taxa (Appendix 1) show leaves with different shapes, usually longer and wider, and not fleshy at all, even in the case of the highly variable E. exigua (Table 1). Moreover, reproductive features from inflorescence architecture, nectaries, flowers, fruits and, mainly, seeds also discriminate the new species from related taxa. Thus, E. nurae always shows simpler and smaller inflorescences, shorter nectaries appendages and stigmas, but relatively larger capsules (excepting E. exigua) than their relatives. In this way, all related taxa examined (Appendix 1) show a lower number of longer dichasial rays and longer individual flowers (Table 1). Seed size and episperm ornamentation have been reported to be good taxonomic features in the whole subgen. Esula Pers. (Baiges & al., 1991) and in sect. Cymatospermum (Molero & al., 1996; Benedí & al., 1997). Euphorbia nurae has seeds conspicuously smaller than its relatives, showing a different shape (ovoid to suborbicular, instead of elliptical) and episperm ornamentation (testa surface is poorly tuberculate with small and obtuse protu berances; Fig. 5). All related taxa examined (Appendix 1) have seed surfaces with conspicuous tubercles or are sulcate (Table 1). Moreover, the most reliable character discriminating the new species is the lack of caruncle, which is uniformly present in all species of sect. Cymatospermum (Molero & al., 1996).
The reduction or complete disappearance of caruncle seems to have evolved independently several times in Euphorbia and it has been reported in other unrelated species from subgenus Esula such as E. pterococca Brot., E. aleppica L. or E. paralias L. (Vindt, 1953; Benedí & al., 1997; Baiges & al., 1991). In Euphorbia the lack or reduction of seed caruncle has been related to dispersal strategies. The presence of an elaiosome for myrmecochorous dispersal may alter the seed aerodynamics, reducing the distance of explosive dispersal (Beattie & Lyons, 1975). Species showing reduced or lacking caruncles or those whose caruncles are readily shed from the seeds would increase the explosive dispersal distance (Narbona & al., 2005).
At the same time the seeds lose their attractiveness, reducing the likelihood of secondary dispersal by ants (Stamp & Lucas, 1990; Narbona & al., 2005). It has been postulated that secondary dispersal by ants is more effective than explosive mechanisms in dispersing the seeds from the parent plant (Stamp & Lucas, 1990) and species that use explosive dispersal maximize either explosive distances or secondary myrmecochorous dispersal (Beattie & Lyons, 1975; Stamp & Lucas, 1990). However, ornithochory and anemochory have been also reported to be involved in the secondary dispersion of Euphorbia species lacking caruncules (Baiges & al., 1991), like the Mediterranean psammophyte E. paralias where dispersal distances up to 100 m have been recorded (Baiges & al., 1991).
Thus, the lack of a conspicuous elaiosome in E. nurae could be linked to a loss of secondary seed dispersal capacity favoring the colonization of suitable neighborhood environments. However, the presence of E. nurae in disjunct populations in Minorca (Fig. 4) might suggest that eventual episodes of long range dispersal might occur. Overall, plants of E. nurae are smaller than those related species, showing at anthesis prostrate rather than erect stems (Fig. 3A) and only at the fruiting stage the stems of E. nurae are upright (Fig. 3B). This could be related to reproductive strategies aimed to be more efficient concerning seed dispersal (Swaine & al., 1979; Stamp & Lucas, 1990; Garrison & al., 2000) to counteract caruncle loss.
Significant reduction of dispersal ability may develop relatively quickly in small, isolated natural populations (Cody & Overton, 1996) and has been documented for species living on both oceanic (Carlquist, 1974) and continental islands (Cody & Overton, 1996; Fresnillo & Ehlers, 2008). Changes in morphological features that are well suited for the colonization of Mediterranean environments with severe summer drought have been reported in Minorcan species of Bellium L. growing in the same habitats as Euphorbia nurae (Fraga & al., 2007). It would be interesting to know which species is sister to E. nurae in order to check the hypothesis that changes in reproductive features could have promoted intraspecific divergence, population isolation and, ultimately, speciation.
Fig. 1.
Euphorbia nurae P. Fraga & Rosselló. A. Habit ; B. Inflorescences ; C. Seeds; D. Representative seed of E. exigua L.
[A–C: Fraga s.n., VAB ; D: Fraga s.n., MA] [Drawings: J. Quintana]
Fig. 3.
Field specimens of Euphorbia nurae P. Fraga & Rosselló at the type locality. A. At anthesis, showing a prostrate habit ; B. At the fruiting stage showing the upright stems.
[Fraga s.n., Herbarium Fraga-Arguimbau]
Fig. 4.
Distribution map of Euphorbia nurae P. Fraga & Rosselló based on field records and herbarium specimens (UTM grid of 1 × 1 km).
Acknowledgements
We thank our colleague Andrés Bermejo for sharing information about the distribution of the new species and for suggesting the specific name and Dr Duncan Ackery for linguistic advice, and Josep Quintana for the drawings.
References
Appendices
Appendix 1.
Additional herbarium material examined.
Euphorbia dracunculoides subsp. glebulosa Coss. & Durieu
Morocco. Gafsa: Canyon of Oued Seldja, near Metlaoui, 25.V.1992, C. Benedí, C. Blanché, J. Molero & J. Vicens s.n. (MA [579274]).
Euphorbia dracunculoides subsp. inconspicua (Ball) Maire
Morocco. Oujda: between Oujda and Jerada, 8 km NW of Guenfouda, 21.V.1991, J. Molero & J. Vicens s.n. (MA [579275]); Safi : Road from Marrakech to Essaouira, cross-road to Korimat and Mouarid, 26.V.1991, J. Molero & J. Vicens s.n. (MA [579276]); Désert Occidental: Tatta, Règion du Djebel Bani, 8.IV.1934, R. Maire & E. Wilczek s.n. (MA [75611]); Moyen Atlas: El Hammam, 10.V.1975, E. Jahandiez s.n. (MA [75612]).
Spain. Almeria: Pilar de Jaravías, 24.III.1986, M.A. Carrasco & M. Velayos s.n. (MA [593988]); Granada: Cerca del Túnel de Izbor, 9.IV.1974, J. Fernández Casas s.n. (MA [395718]).
Euphorbia exigua L. subsp. exigua
Austria. Niederösterreich N. von Wien, Hagenbrun, 24.VII.1999, Ernst Vitek s.n. (MA [642213]).
Belgium. Tilff, Sur la Mont, 3.X.1980, V. Demoulin s.n. (MA [346747]).
Czech Republic.Moravia: 10.VIII.1927, R. Dvo ak s.n. (BC [88099]).
Denmark. Vallensbaek, SW of Copenhagen, 26.VIII.1989, A. Hansen s.n. (MA [497588]).
France. Pegomas, 4.V.1963, G. Gavelle s.n. (MA [178887]); Ablon, Seine-et-Oise, 18.VIII.1912, H. Debaise s.n. (BC [57499]).
Germany. Oberbayern, lehmiger Acker in München-Riem, 15.VII.1963, W. Lippert s.n. (MA [192574]); Munitz, VIII.1933, Zaus s.n. (MA [75567]).
Israel. Judaean Mts., Jerusalem, Givat Ram, 18.V.1956, H. Parmet s.n. (MA [172097]).
Italy. Mont. Catalfano, commune de Bagheria, province de Palerme, Sicilie, 38º 06′N 13º 31′E, 22.IV.1999, A. Certa & A. Carratello s.n. (MA [693144]); Venetia, 1897, Rigo s.n. (MA [75570]).
Morocco. Env. de Martimprey du Kiss, 22.IV.1933, A. Faure s.n. (MA [75604]); Tarquist, 21.V.1927, P. Font Quer s.n. (MA [75565]); Sidi-Musa, Beni-Sicar, 14.III.1935, Hno. Mauricio s.n. (MA [161496]); Tetuan, V.1975, M. G. s.n. (MA [75574]); Yebel Malmusi, 8.V.1927 Font Quer s.n. (MA [75577]); Oran, 10.IV.1908, A. Faure s.n. (BC [57501]).
Netherlands. Rhijnauwen, vic. of Utrecht (prov. Utrecht), 1.IX.1959, W. Punt s.n. (MA [173615]).
Portugal. Madeira, Cabo Girão, 29.V.1954, M. Beliz s.n. (MA [250853]); Caillabé, Coimbra, 25.IV.1949, Y. Cebãlos s.n. (MA [250832]).
Spain. Álava: San Román de Kampezo, Bernedo, Vitoria, 14.IX.1983, J. A. Alejandre s.n. (MA [408763]); Lapuebla de Labarca, Vitoria, 12.V.1983 Goyo Morante & J. A. Alejandre s.n. (MA [408734]); Zambrana, Vitoria, 8.X.1983, J. A. Alejandre s.n. (MA [408762]); Laserna, Laguardia, Vitoria, 21.V.1983, J. A. Alejandre s.n. (MA [408733]). Alicante: Villafranquesa, 23.III.196, A. Rigual s.n. (MA [370816]); Barranco de las Ovejas, 09.V.1959, A. Rigual s.n. (MA [370814]); Barranco de Chirles, 10.VII.1933, M. Martínez s.n. (MA [75526]); Muchamel, 9.V.1962, A. Rigual s.n. (MA [370797]); Moraira la Torre, 13.IV.1996, J. X. Soler & B. Banyulus s.n. (MA [587493]); Murla. S. del Peñon, 1.IV.1983 J. X. Soler & al. s.n. (MA [572184]); Jesús Pobre, 29.V.1994, J. X. Soler s.n. (MA [546387]); Montgó, Denia, 30.V.1988, Pilar Donat s.n. (MA [502801]); Chirles, 9.VII.1933, M. Martínez s.n. (MA [75562]). Barcelona: Castelldefels, 20.V.1917, E. Gros s.n. (MA [75534]); Collsacabra, VII.1885, F. Trémols s.n. (MA [75533]); Gualba, Montseny, 15.V.1915, Font Quer s.n. (BC [57490]); Font d'en Fargas, Horta, IV.1933, Dra. Gallardo s.n. (BC [125051]); Sant Llorenç del Munt, 1.VI.1941, A. de Bolòs s.n. (BC [99624]); Santa Coloma de Gramenet, 28.V.1939, A. de Bolòs s.n. (BC [99268]). Cádiz: Algeciras, 21.V.1962, B. Casaseca s.n. (MA [178881]); Chiclana, 28.V.1957, A. de Bolòs s.n. (BC [116711]). Cantabria: Unquera, Santander, 21.V.1960, F. Bellot & B. Casaseca s.n. (MA [178880]). Castellón: Benicarló, 8.IV.1993, V. J. Arán & M. J. Tohá s.n. (MA [523434]); Embalse de Ulldecona, 7.VI.1999, C. Navarro & al. s.n. (MA [626643]); Fredes, hacia la peña Blanca, 8.VI.1999, J. Güemes & al. s.n. (MA [628568]). Ciudad Real: Sierra de Alhambra, 30.IV.1933, J. González Albo s.n. (BC [84115]). Gerona: Torroella de Montgrí, 25.IV.1948, Font Quer s.n. (BC [106205]); Palafrugell, 16.IV.1922, Font Quer s.n. (BC [99342]); Cadaqués, 12.V.1917, Gros s.n. (BC [99276]). Granada: Guadix, 16.V.1929, Gros s.n. (BC [99329]). Huelva: Huelva, 24.IV.1931, Gros s.n. (BC [99343]). Islas Baleares: Cabrera, 19.IV.1948, P. Ferrer s.n. (MA [75528]); Isla Dragonera, 17.VI.1980, E. Valdés-Bermejo s.n. (MA [405901]); Matorrales debajo de Sa Vicaria, Cabrera, 29.V.1947, P. Ferrer s.n. (MA [75530]); Riera de Palma, 9.VI.1946, P. Ferrer s.n. (MA [75531]); Ibiza, IV.1899, C. Pau s.n. (MA [75542]); Camino de San Juan a Cala Portinatx, 15.VI.1949, P. Ferrer s.n. (MA [75527]); Port des Torrents, Ibiza, 1.V.1980, Rivas Martínez, M. Costa & A. M. Regueiro s.n. (MA [422598]); Puigpunyent, Mallorca, 2.VI.1982, R. Morales & al. s.n. (MA [618132]); Fornalutx, Mallorca, 6.VI.1998, M. Velayos & al. s.n. (MA [618910]); Son Àngel, Menorca, 27.III.1999, P. Fraga s.n. (MA [624572]); Portopí, Mallorca, 1.IV.1956, O. Bolòs & R. Molinier s.n. (BC [136083]); Porto Pi, Mallorca, 26.III.1948, Palau Ferrer s.n. (BC [112256]); Porto Pi, Mallorca, 24.III.1948 Palau Ferrer s.n. (BC [106779]); Cala Llonga, Eivissa, IV.1918, Gros s.n. (BC [99335]); Sant Antoni, Eivissa, 25.III.1918, Gros & Font Quer s.n. (BC [99301]); Torrent d'en Costa, Sant Rafel, Eivissa, 31.V.1918, Gros s.n. (BC [99333]); Pont d'Inca, 6.V.1917, Bianor s.n. (BC [574 80]); Alcaufar, Sant Lluís, Menorca, III.1912, Font Quer s.n. (BC [57477]); Santa Ponsa, Menorca, 10.V.1913, Font Quer s.n. (BC [57478]); Cala en Turqueta, Ciutadella de Menorca, Menorca, 19.V.1959, A. de Bolòs & O. de Bolòs s.n. (BC [121 535]); Espardell, Eivissa, 19.V.1920, Gros s.n. (BC [99328]); Santa Eulàlia, Eivissa, 27.III.1918, Gros & Font Quer s.n. (BC [142376]). La Rioja: Canales de la Sierra, 22.VII.1975, B. Casaseca y Fernández Díez s.n. (MA [204 505]); San Millán de Yécora, 28.V.1988, J. Arizaleta, F. Muñoz Garmendia, J. Pedrol y R. Rodríguez s.n. (MA [438801]). Málaga: Estepona, 16.V.1919, Gros s.n. (BC [57504]). Melilla: Melilla, IV.1912, A. Caballero s.n. (MA [75578]). Murcia: Sierra de la Puerta, Cehegín, 1.V.1982, Caridad Selme s.n. (MA [541799]); Algameca Chica, Cartagena, 20.VII.1902, Ibáñez, Jiménez & Pau s.n. (MA [75 564]); Cabo Cope, Aguilas, 24.III.1978, Fernández Díez s.n. (MA [250400]); Javali N., 11.IV.1986, R. Garcia s.n. (MA [456780]); Castillo de la Azohía, 29.III.1998, A. Carrillo, E. Coy, J. Güemes, A. Hernández, F. Muñoz-Garm. & C. Navarro s.n. (MA [612425]); pr. Portman, 1.III.2004, C. Aedo s.n., (MA [713875]). Navarra: Mendavia, 30TWM 6898, 28.V.1985, J. A. Alejandre s.n. (MA [337872]). Santander: Monegro (Reinosa), VII.1924, L. Atendo s.n. (MA [145087]); San Vicente de la Barquera, VII-1920, J. de la Espada (MA [75539]). Sevilla: Moron, 25.IV.1933, C. Vicioso s.n. (MA [75525]). Tarragona: Litoral prope l'Hospitalet del Infante, 01.VII.1969, L. Carreras & E. Valdés Bermejo s.n. (MA [397257]); Muntanyes de Poblet, Montblanc, 30.IV.1950, F. Masclans s.n. (BC [127549]); La Bisbal de Falset, 31.V. 1974, J. Molero & A. Boldú s.n. (BC [631681]). Valencia: La arboleda, Llíria, 24.VIII.1993, Fco. Morán s.n. (MA [565 502]); El Paller, 29.IV.1982, M. Palasà s.n. (MA [331740]); Serra Perengisa, Torrent, 25.IV.1984, Rivas-Martínez s.n. (MA [337632]); Bétera, 27.III.1984, J.A. Alcober s.n. (MA [383189]); Los Corrales, 21.V.1984, R. Figuerola & G. Mateo s.n. (MA [426932]); Coll d'Eslida, Serra d'Espadà, 24.V.1947, Font Quer s.n. (BC [106115]). Vizcaya: Bilbao, 1.V.1947, E. Guinea s.n. (MA [461173]). Bilbao. zona camino de Iturri-Gorri, V.1947, E. Guinea s.n. (MA [1650 60]). Zaragoza: Calatayud, 21.VII.1897, Iñiguez s.n. (MA [145086]).
Tunisia . Gabés (Menzel), II.1907, C. J. Pitard s.n. (MA [75605]).
Euphorbia exigua subsp. merinoi M. Laínz
Italy. Sardinia: Nouro, Genargentu, 4.VI.2003, C. Navarro & al. s.n. (MA [708348]).
Spain. Cáceres: El Pozuelo, Guadalupe, 20.V.1949, A. Caballero s.n. (MA [75544]); Cercanías del río Guadalupe, 25.V.1949, A. Caballero s.n. (MA [75545]); Las Hurdes, 21.V.1947, A. Caballero s.n. (MA [75547]); Guadalupe, 16.VI.1948, A. Caballero s.n. (MA [75546]). Cuenca: Término de Cuenca, barranco de la Madera, 7.VII.2001, V. J. Arán & al. s.n. (MA [691928]). Orense: Rubió, supra Pardellán, 2.V.1987, F. J. Silva-Pando & al. s.n. (MA [406650]); Barco de Valdeorras, Xaguarras, 14.V.1989, Amigo & al. s.n. (MA [478113]).
Euphorbia sulcata Loisel.
France. Arnas (Rhône), 16.IX.1874, M. Gandoger (MA [145085]); Aix (B. du R.) au Prégnon, IV.1887, Bruyas s.n. (MA [75670]).
Italy: Abruzzo, L'Aquila, Sª Eufemia, 7.VII.2002, J. Aldasoro s.n. (MA [698660]).
Spain. Burgos: Tejada: pie del Pico Valdosa, 3.VII.1979, Fdez. Casas, Lara, Pons-Sorolla & Susanna s.n. (MA [251162]). Cuenca: Laguna del Hito, Montalvo, 19.VI.1975, S. Cirujano s.n. (MA [408772]); Lleida: La Noguera, entre Balaguer y Castelló de Farfanya, 3.V.1986, C. & J. Pedrol s.n. (MA [304602]); Entre Balaguer y la Sentiu de Sió, 6.V.1985, J. Pedrol s.n. (MA [313986]); Logroño: Dehesa de Ausejo, 29.V.1985, J. Pedrol & al. s.n. (MA [438411]); Madrid: Cerca de Aranjuez, 6.IV.1946, Rivas Goday s.n. (MA [276056]); Cerro Negro, 1.VI.2002, J. Isern s.n. (MA [720252]); Murcia: Moratalla, proximidades de El Sabinar, 25.IV.1997, I. Álvarez s.n. (MA [591040]); Salamanca: La Orbada, 25.V.1985, E. Rico s.n. (MA [390778]); Segovia: Arroyo de la Hoz, Cedillo de la Torre, 30.IV.1984, M. J. Illueca s.n. (MA [507332]); Valladolid: Tiedra, 8.VI.1981, Fernández Diez (MA [250608]); Cabezón de Pisuerga, 31.V.1980, A. R. Burgaz s.n. (MA [308989]); Fuenteoco, 15.V.1983, J. L. Fernández Alonso s.n. (MA [406320]).
