Botanical Journal of the Linnean Society (1999), 129: 155–164. With 3 figures

Article ID: bojl.1998.0210, available online at http://www.idealibrary.com on

A new Biscutella (Brassicaceae) species from thewestern Balearic Islands

JOSEP A. ROSSELLO∗

Botanica, Departamento de Biologıa Vegetal, Facultad de Ciencias, Universidad de Valencia,E-46100 Burjassot, Valencia, Spain

NESTOR TORRES

Apartat de Correus 64, 07800 Eivissa, Balearic Islands, Spain

LLORENC SAEZ

Real Jardın Botanico, CSIC, Plaza de Murillo 2, E-28014 Madrid, Spain

Received January 1998; accepted for publication August 1998

A new diploid species, Biscutella ebusitana is described from coastal cliffs of the westernBalearic islands. It is related to B. sempervirens L. and B. montana Cav. from which it can bedistinguished by a combination of vegetative (shape, size, petiole and hairs of leaves, petiolateleaves) and reproductive features (shape of inflorescence, calyx indument). Earlier Balearicrecords of B. frutescens Coss. should be referred to B. ebusitana.

1999 The Linnean Society of London

ADDITIONAL KEY WORDS:—endemism – insular flora – western Mediterranean –taxonomy.

CONTENTS

Introduction . . . . . . . . . . . . . . . . . . . . . . . 156Material and methods . . . . . . . . . . . . . . . . . . . 156Results . . . . . . . . . . . . . . . . . . . . . . . . 157

Biscutella ebusitana sp. nov. . . . . . . . . . . . . . . . 157Comparative morphology . . . . . . . . . . . . . . . . . 157Chromosome number . . . . . . . . . . . . . . . . . . 160Distribution and ecology . . . . . . . . . . . . . . . . . 160

Discussion . . . . . . . . . . . . . . . . . . . . . . . 161Acknowledgements . . . . . . . . . . . . . . . . . . . . 163References . . . . . . . . . . . . . . . . . . . . . . . 163Appendix . . . . . . . . . . . . . . . . . . . . . . . . 164

∗Corresponding author. E-mail: rossello@uv.es

1550024–4074/99/020155+10 $30.00/0 1999 The Linnean Society of London

J. A. ROSSELLO ET AL.156

INTRODUCTION

Perennial taxa of the genus Biscutella have been traditionally placed in seriesLaevigatae Malin. (sect. Biscutella), in contrast with the annuals which are locatedin series Lyratae (sect. Biscutella) and Jondraba (Medik.) DC. Series Laevigataetherefore comprises woody or subwoody herbs, characterized by petals truncate andauriculate at the base, staminal filaments not winged, all nectaries extrastaminaland fruits with a membranaceous margin. Taxa belonging to this series are mor-phologically highly diversified and several names have been recently proposed fordiscrete entities, mainly from the western Mediterranean basin (Crespo, Guemes &Mateo, 1992; Mateo & Crespo, 1993; Raffaelli & Fiesoli, 1993; Raffaelli & Baldani,1997). Since the initial record of B. frutescens Coss. from the islands of Eivissa (Ibiza)and Formentera (Knoche, 1922), various members of the B. sempervirens L. grouphave been recorded from the western Balearics. Later on, the same populationswere referred to B. montana Cav. by Font Quer (1928) who described them as a newvariety restricted to the Balearic archipelago (B. montana var. densiflora Font Quer).This view was not followed by Guinea & Heywood (1964) and Greuter, Burdet &Long (1986) who reported both B. frutescens and B. sempervirens from the BalearicIslands. Heywood (1964) pointed out differences in leaf features of the insular plantwhen compared to the B. sempervirens from the mainland, although including theBalearic plant under the typical variety. Bolos & Vigo (1974, 1990) agreed with theearlier proposal of Font Quer (1928) concerning the endemic nature of the pytiusicplant and transferred it to a broadly circumscribed B. laevigata [as B. laevigata subsp.montana var. densiflora (Font Quer) O. Bolos & Masclans]. A contrary view was takenby Grau & Klingernberg (1993), who were doubtful about the presence of B.sempervirens in the Balearics. Aguilella et al. (1994) restricted B. montana to the Spanishmainland populations and suggested that the insular plants should be referred toother unspecified taxa closely related to B. sempervirens.

Recent intensive field work devoted to Biscutella populations in Eivissa andFormentera has revealed the presence of a single entity. The combination ofits morphological characters is consistent and does not match earlier taxonomicidentifications of other described taxa. Accordingly, the Balearic populations arehere recognized as an independent species, B. ebusitana.

MATERIAL AND METHODS

Besides the living material examined in the field, herbarium specimens were studiedfrom the following herbaria: BC, BCC, MA, MAF, VAL and VAF (representativespecimens of non-Balearic origin are listed in the Appendix). Type specimens of B.sempervirens (LINN 831.8; cf. Heywood, 1964) and B. montana (MA 475433) were alsoexamined for comparative purposes.

For cytological studies root tips were pretreated in an aqueous solution of 0.2%colchicine for 3–5 hours, rinsed in distilled water and stained for 24 hours in aceticorcein at room temperature. Fresh leaves were fixed in FAA, sectioned in a rotarymicrotome at 10 lm intervals and stained in toluidine blue O. Leaves from herbariumspecimens were rehydrated (12–24 h), decoloured (2–4 h) in a commercial solutionof sodium hypochlorite and observed at a magnification of 400 ×.

A NEW BISCUTELLA FROM THE BALEARIC ISLANDS 157

RESULTS

Biscutella ebusitana Rossello, N. Torres & L. Saez, sp. nov.=B. montana var. densifloraFont Quer, Butll. Inst. Catalana Hist. Nat., 26: 104 (1926); in Cavanillesia 1: 31(1928; B. laevigata L. var. densiflora (Font Quer) O. Bolos & Masclans, Butll. Inst.Catalana Hist. Nat. 38: 76 (1974)

Diagnosis. Biscutella montanae Cav. et Biscutella sempervirenti L. similis sed habitu ro-bustiore, statura majore (usque ad 100 cm); inflorescentia ramosa, paniculata; racemisbrevioribus (usque ad 4 cm), densissimis (usque ad 20 flores per cm); calycibuspubescentibus et siliculis minoribus (3–6× 7–9 mm). Ob foliis latioribus spathulato-ellipticis, lobatis, a Biscutella sempervirenti L. satis diversa videtur.

Holotypus. INSULAE BALEARICAE, EBUSUS: In rupestribus maritimis loco dictoEs Penyal de s’Aguila, 31S62 ubi N. Torres et J.A. Rossello 9.x.1993 legerunt (VAB950061)

Isotypes. BCC, MA, VAL

Material Examined. EIVISSA: Vedra, Gros, 19.vii.1920 (BC 4998); Vedranell, 25 m,Font Quer, 7.v.1920 (BC 4996; BCF 35574); Cala d’Aubarca, Font Quer, 11.v.1920(BC 4997); Cala de les Torretes, Gros, 8.vii.1920 (BC 5000); ibidem, Gros, 29.v.1918(BC 4993); Cala de Lladalt, Gros, 7.vii.1920 (BC 4999; BCF 35575); ibidem, Gros,1918 (BC 4994); ibidem, 29.v.1918, Gros (BC 4993); Punta de la Creu, 9.iv.1919, FontQuer (BC 4995); Ses Balandres, Rossello & Torres, 23.vii.1996. FORMENTERA: LaMola, Gros, 11.v.1918 (BC 4985); ibidem, 24.v.1920 (BC 4984).

Illustration. Figure 1.

Description. Perennial plant, woody at the base, with few stems, tomentose. Basalleaves rosulate, 5–20 × 1.5–3.5 cm, tomentose, light green, spatulate-elliptical,sinuate-dentate at the margins; petiole slightly canaliculate, 1/3–1/5 as long as theblade, 2–4 mm wide. Lower cauline leaves oblong, obtuse, semi-amplexicaule,auriculate at base, dentate to crenate, tapering until the end of the stem. Stem20–100 cm tall, erect, branched in the upper half. Inflorescence paniculate, profuselybranched, with primary branches 15–25 cm long, erect to erect-patent; flowersrestricted to the upper third. Pedicels erect, 5–10 mm long. Calyx 2–2.5 mm long,hairy; petals 4–7 mm long, cuneate, deep yellow. Silicula 3–6 × 7–9 mm, glabrousor covered with sparsely claviform hairs, style persistent, 2–3.5 mm long.

Comparative morphology

Petiole anatomy. The petioles of B. ebusitana are subcircular, with lateral wings, withconvex abaxial and flat adaxial surfaces in transverse sections, with simple, unicellulartrichomes on both sides; hypodermis is present in both surfaces. The vascularstructure consists of a triple central bundle and several smaller ones in the wings;bundles are collateral and U-shaped. These observations contrast with those pre-viously reported in the genus (Olowokudejo, 1987) in several ways. First, the centralvascular structure of the petiole has been described as composed of a single bundlefor all examined species of Biscutella. Second, the hypodermis has been noticed only

J. A. ROSSELLO ET AL.158

Figure 1. Biscutella ebusitana sp. nov. A, sepal; B, petal; C, habit; D, flower; E, silicula. Scalebars: A,B,D,E = 1 mm; C = 1 cm.

A NEW BISCUTELLA FROM THE BALEARIC ISLANDS 159

T 1. Indumentum features of the adaxial blade of Biscutella sempervirens, B. montana and B. ebusitanasp. nov. Measurements are in lm.

B. sempervirens B. montana B. ebusitana sp. nov.

Length 140–1035 180–900 120–670Width at base 20–40 6–30 20–42Width at 1/2 length 12–20 4–12 12–23

Figure 2. Morphological variability in leaves (left) and hairs (right) of Biscutella ebusitana sp. nov.(A), B. montana (B) and B. sempervirens (C). Material examined: B. ebusitana: (BC 4995, BC 4999, BC4997, BC 4985); B. montana (BC 4982; BC 4989, BC 4980, BC 4986); B. sempervirens (BC 4974, BC631382, BC 631368, BC 107796, BC 631384). Scale bars = 1 cm (leaves); 100 lm (hairs).

in B. frutescens but an examination of the figures given by Olowokudejo (1987) suggestthat it may be present in other species like B. intermedia and B. auriculata.Indumentum. Biscutella sempervirens, B. montana and B. ebusitana have pubescent leaveswith an indumentum composed of single unicellular hairs (Table 1). Biscutella montanashows two hair types: cylindrical (flexuous) with a narrow base (6 lm) and flat hairswith a broader base (22–30 lm). Both B. sempervirens and B. ebusitana have onlycylindrical hairs. However, the hairs of the latter are usually flexuous (from themiddle up to the tip), whereas B. sempervirens has rigid and straight hairs (Fig. 2).

J. A. ROSSELLO ET AL.160

Formentera

10 km

MediterraneanSea

Eivissa

SPAIN

BalearicIslands

Figure 3. Distribution of B. ebusitana sp. nov. based on herbarium specimens and field records.

Chromosome number

Plants from the type locality have a chromosome number of 2n=18. Thechromosomes are small, homogeneous in size and shape and without visible primaryconstrictions.

Distribution and ecology

Biscutella ebusitana grows on shadowy coastal limestone cliffs and abrupt slopes,usually on northern, western or eastern exposures. The plants can be found fromnearly the sea level up to 400 m altitude. Its distribution is restricted to the northwestcoast of Eivissa, the northern and eastern coasts of Formentera and the islets Vedraand Vedranell, southeast of Eivissa (Fig. 3). In Eivissa and the small islets B.ebusitana occurs together with many rupicolous taxa of endemic or narrow westernMediterranean distribution such as Helichrysum rupestre DC., Scabiosa cretica L., Lamotteadianiae (Webb) G. Lopez, Limonium ebusitanum (Font Quer) Pignatti, Hippocrepis balearicaJacq. subsp. grosii (Pau) Mus et al., Thymus richardii Pers. subsp. ebusitanus (Font Quer)

A NEW BISCUTELLA FROM THE BALEARIC ISLANDS 161

Jalas, Diplotaxis ibicensis (Pau) Gomez-Campo, Asperula paui Font Quer, Cheirolophusintybaceous (Lam.) Dostlal, Silene hifacensis Rouy ex Willk, and Galium crespianum J.J.Rodr. In some localities several subnitrophilous elements, such as Withania frutescens(L.) Pauquy, Ballota hirsuta Benth. and Salsola oppositifolia Desf. can be found.

DISCUSSION

The general morphology of the western Balearic Biscutella plants falls well withinthe scope of the B. sempervirens complex. The shape of median nectaries, vascularanatomy and petiole section morphology and the silicula without swollen marginsclearly show that the insular plant should not be included within B. frutescens, acommon misidentification in the past (Knoche, 1922; Guinea & Heywood, 1964;Hernandez Bermejo et al., 1986).

Taxonomic recognition within the B. sempervirens group is controversial and noagreement has been achieved regarding the number and rank of the taxa to beincluded in it. Two entities, B. sempervirens and B. montana, have been named toaccount for the opposite poles of variation but their recognition as distinctive taxahas not been followed by many authors. Instead, both species have been usuallymerged as extreme forms of a single taxon with continuous morphological variation(Heywood, 1964; Guinea & Heywood, 1964; Olowokudejo, 1986a; Grau & Klin-genberg, 1993). However, some caution should be applied to this view sinceconclusive evidence is lacking regarding the nature and extent of such variation.Phenotypic polymorphism has been invoked to explain it, although it should benoted that the morphological attributes of the Balearic plant have been retained incultivation for several years, suggesting that they are genetically fixed. Similarconclusions have been made concerning the differing features of B. sempervirens andB. valentina L. populations (Olowokudejo, 1986a, c). These results contrast withearlier reports claiming that some vegetative characters (e.g. leaf-shape) can bemodified by different growth conditions (Heywood, 1964). Moreover, populationsof south-east Spain (from where B. montana was described) were not sampled whenthe morphological variation of B. sempervirens s.l. was surveyed (Olowokudejo, 1986a)or when petiole anatomy, nectaries, cotyledon and seedling features were studiedin the whole genus (Olowokudejo, 1986b, c, d, 1987). Clearly, the taxonomic statusattributed to the Balearic plant should be in accordance with that recognized fortheir diploid relatives, B. sempervirens and B. montana, with which it is more closelyrelated and from which presumably it arose through isolation of the mainland andwestern insular populations. Unfortunately, morphological evidence does not provideclear-cut patterns for recognition of the perennial taxa of Biscutella and cytogeneticgaps are restricted to few taxa, where polyploid cytotypes have been detected(Schonfelder, 1968; Olowokudejo & Heywood, 1984).

In the absence of more conclusive evidence, we think the criteria for speciesdelimitation in the genus necessarily should be based on evaluation of: (i) thediscriminating morphological features, (ii) the relationship between morphologicaland geographical patterns of variation and (iii) the nature and extent of transitionalforms, if any, linking discrete entities. Morphological differentiation between typicalforms of B. sempervirens, B. montana and the Balearic plant is mainly based on theshape and size of the basal leaves and the architecture of the inflorescence (Table 2).

J. A. ROSSELLO ET AL.162

T 2. Morphological differences between Biscutella sempervirens, B. montana and B. ebusitana sp.nov.

B. sempervirens B. montana B. ebusitana sp. nov.

Leaves oblong-lanceolate to spatulate-elliptical, spatulate-elliptical,lanceolate, attenuate petiolatepetiolate

Stem up to 50 cm, up to 60 cm, up to 100 cm,loosely branched loosely branched profusely branched

Raceme length (cm) up to 25 up to 12 up to 4Pedicel length (mm) 5–17 6–16 5–10Calyx length (mm) 2–3 2–3 2–2.5

glabrous glabrous hairySilicula (mm) 4.5–6×7–12, 4.5–7×8–12, 3–6×7–9

hairy or glabrous usually glabrous usually hairyStyle length (mm) 3–5 3–5 2–3.5

The leaves of the island plant differ distinctively from those of both B. montana andB. sempervirens (Fig. 2). Its inflorescence is more branched (with shorter racemes andmore flowers per raceme) than that of its relatives, the sepals are hairy, the stylelength and the silicula size are shorter and the coloration of the petals is deeper,although this is barely visible in the herbarium specimens. The presence of claviformhairs in fruit valves is not exclusive to the new species since some populations of B.sempervirens share this feature. On the other hand, B. montana very rarely has hairson the fruits. These features do not change in culture under homogeneous greenhouseenvironments and provide key characters on which the three taxa can be separated.

This morphological variation is correlated with a geographical pattern of dis-tribution. Plants belonging to the B. montana morphotype are restricted to easternSpain (Valencia and Alicante provinces), B. sempervirens populations extend to thesouth of the lberian peninsula, and the third variant is restricted to the BalearicIslands. Specimens having intermediate morphological features between B. sempervirensand B. montana have been reported from south and south-east Spain (HernandezBermejo et al., 1986; G. Mateo, pers. comm.). Ecological convergence or, perhaps,hybridization and introgression processes should not be ruled out to explain thismorphological intermediacy since sympatry is common in that region. Plants fromeastern Spain resembling B. sempervirens could, in fact, be hybrids between B. montanaand B. valentina.

Taking all the available evidence into account, we think that the three entitiesshould be granted some formal taxonomic recognition as has been done in thepast. We refrain from using the subspecific status to name the most prominentmorphological variants within the B. sempervirens complex in the absence of anybiosystematic work covering the woody perennial taxa of the western Mediterraneanbasin. Merging these entities into a single taxon creates a highly polymorphic species,whose monophyletic nature is for the moment untested. Bearing in mind the syntheticnature of this approach and the few clear discriminating characters present in thegenus, there is no sound reason for lumping together many woody taxa of Biscutellaseries Laevigatae under a single species (B. laevigata) as has been proposed by someauthors (Bolos & Vigo, 1974).

Recognition of the Balearic Biscutella as a separate species from the related B.montana adds a new endemic element for the Pityusic islands and emphasizes the

A NEW BISCUTELLA FROM THE BALEARIC ISLANDS 163

degree of divergence between vicarious taxa inhabiting restricted localities insoutheast Spain and the western Balearic islands (Eivissa and Formentera) in thesame way as the vicariance pattern of eastern vs western islands.

ACKNOWLEDGEMENTS

We thank G. Mateo, G. Nieto Feliner and J. Fuertes Aguilar for their usefulsuggestions and criticisms. This work was supported by DGICYT research grantPB 93–0350.

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APPENDIX

Material examined

Biscutella semperivensALMERIA: Barranco de los Castanos, c. Vacares, 3.vi.1929, Gros (BC 107795); Sierra de Gador,

near Dalias, 600 m, 12.vi.1978, J. Olowokudejo (BC 631368); between Almeria and Enix, 800 m,12.vi.1978, L. Olowokudejo (BC 631382); Barranco del Caballar, 17.vi.1921, Gros (BC 4974); sierrade Enix, rochers, 12.v.1932, Jeronimo (BC 76836); Malaga: Nerja, 28.iv.1919, Gros (BC 4971);Alhama, camino del balneario, 30S VF19, 800 m, 8.iv.1980, Martinez Parras (BCF 35570); GRANADA:sierra de Yedra, 10 km N of Alfacar, 1200 m, 14.vi.1978, J. Olowokudejo (BC 631368); JAEN: Cerrola Vieja, macizo de Magina, 15.vi.1926, Cuatrecasas (BC 5045); La Silleta, vertiente N, 1200 m,10.vi.1925, Cuatrecasas (BC 5047); La serrezuela de Beolmar, 1350 m, 9.vi.1926, Cuatrecasas (BC5042); Sierra de Carboneras, 1300 m, 21.vi.1925, Cuatrecasas (BC 5043); Barranco del Madrono,27.iv.1926, Cuatrecasas (BC 5041); versus Molinos de Cuadros, 20.iv.1926, Cuatrecasas (BC 5040);MALAGA: Siera Tejeda, in cacumine cerro Lucano, 9.v.1926, Gros (BC 197840); Sierra de Mijas,v.1935, M. Laza (BCF 35588); CADIZ: Gibraltar, 100 m, 15.v.1935, Font Quer (BC 86572); Cerrode San Cristobal, pr. Grazalema, ad, 1250 m alt., 14.vi.1922, Gros (BC 87744).

Biscutella montanaALICANTE: In Monte Motngo, prope Dianium, 300 m, 2.v.1923, Gros (BC 4980, BC 4989);

Ibidem, 400 m, 16.vi.1950, O. Bolos (BC 114247); Ibidem, 2.iv.1980, Molero & al. (BCF s.n.); Sagarria,8.v.1923, Gros (BC 4982); Alicante: Cap de Sant Antoni, 16.v.1923, Font Quer (BC 4987); Ibidem,10.vi.1983, Peris & al. (BC 646961); Barranc de Mascarat, pr. Calp, 50 m, 13.v.1923, Font Quer (BC4991); Ifac, pr. Calp, 100 m, 12.v.1923, Font Quer (BC 4990, BC 4998); penon de Calpe (Alicante),23.v.1953, s.r. (BC 600036); Serra de Bemia, 21.vi.1923, Gros, (BC 4978); Serra de Xeresa, 28.v.1923,Font Quer (BC 4979; BCF 35578); Serra d’Altana, 1000 m, 15.vi.1923, Font Quer (BC 107802);Alcoi, la Serreta, 900 m, 11.v.1934, Pau (BCF 35576; BCF 35577); Sierra de Benicadell, 29.vi.1946,Borja (BC 100981); In Regno Valentino: Cova Alta d’Albaida, 10.vi.1923, Gros (BC 4987); VALENCIA:c. cap Cullera, 22.v.1923, Gros (BC 107790).