Botanical Journal of the Linnean Society, 2004, 146, 97—102. With 2 figures

Table of contents

A new species of Orobanche (Orobanchaceae) from south-eastern Spain ANTONIO J. PUJADAS-SALVA* and MANUEL B. CRESPO 2 'Departamento de Ciencias y Recursos Agrícolas y Forestales, Universidad de Córdoba, Apdo. 3048, E-14080 Córdoba, Spain 2 CIBIO (Instituto Universitario de la Biodiversidad), Universidad de Alicante, Apdo. 99, E-03080 Alicante, Spain Received May 2003; accepted for publication March 2004

A new species, Orobanche portoilicitana A. Pujadas & M. B. Crespo (sect. Trionychon Walk.) is described from the south-eastern coast of the Iberian Peninsula. This species inhabits coastal sand dunes where it is parasitic on Centaurea species. It is characterized by its dense, many-flowered inflorescences, which bear usually erect, pale bluish flowers with long apiculate anthers. Several morphological features relate the new species to O. olbiensis (Coss.) Nyman, a plant known only from southern France, with which it had been misidentified. A complete description and the diagnostic characters allowing differentiation from other taxa in the section are reported. Further data on ecology and conservation status are also presented. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 146, 97-102.

ADDITIONAL KEYWORDS: broomrape – endemism – Iberian Peninsula – parasitic plants – taxonomy.

INTRODUCTION Orobanche L. is a genus that is quite diversified in the Iberian Peninsula, where 32 taxa were reported in Flora Iberica (Foley, 2001). Four of these taxa are endemic to Spain and Portugal, and another three grow exclusively in North Africa and the southern Iberian Peninsula. With regard to section Trionychon Wallr., nine taxa were annotated by Foley (2001) in his revision of the Iberian representatives of the genus. However, he did not include O. olbiensis (Coss.) Nyman, a plant reported by Pujadas-Salva & Crespo (2000) from coastal sand dunes of the southern Alicante province, the first citation for Spain. This taxon was first described in the genus Phelypaea L. by Cosson (1849: 8) from Porqueroles (Hyéres Islands, southern France), parasitizing Helichrysum stoechas (L.) Moench. Identifications of Cosson's taxon by PujadasSalva & Crespo (2000) and then Pujadas-Salva (2001, 2002) were based on comparisons of the Alicantine

*Corresponding author. E-mail: [email protected]

plants to one herbarium sheet at the Institute Botanique de Montpellier (MPU), which was labelled as Phelypaea olbiensis Cosson and certainly appeared to be close to the Spanish plants at first sight. All efforts to localize the type specimen in several European herbaria were unfruitful, even in Cosson's collections at Paris (P), where no vouchers of that taxon seem to exist (Ph. Morat, pers. comm.). Nonetheless, the finding was published quickly in the hope that it would be included in the review of the Iberian Orobanche that was about to be finished at that time; unfortunately, no mention was made of this taxon in Flora Iberica. However, more detailed studies of several specimens of Orobanche olbiensis collected in Porquerolles by Henri Michaud (Consevatoire Botanique National Méditerranéen de Porquerolles) and a new comparison of the Alicantine specimens against the material at MPU, have allowed us to reconsider our previous erroneous identification. In view of the remarkable morphological differences between plants from Alicante and other taxa of sect. Trionychon, namely O. olbiensis and O. mutelii F.W. Schultz (cf. Pujadas-Salva & Crespo, 2000), evidence is here presented to describe a new species from the south-eastern Spain.

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MATERIAL AND METHODS Observations were made on both fresh material collected from the natural populations and herbarium specimens held at the herbaria ABH and COA. Other herbaria (ALME, BC, BCC, 'Colegio La Salle' of Almeria, `Col.legi Oficial de Farmacéutics de les Illes Balears', HJBS, `Herbarium Orell-Casanovas', UIB, MA, MAF, MGC, MUB, MPU, VAB and VAL) were also revised in search of new or misidentified specimens to complete its description and distribution area. Acronyms according to Holmgren, Holmgren & Barnett (1990) and Holmgren & Holmgren (1993). Morphological characterization of the Alicantine plant against O. olbiensis was made on the basis of Cosson's (1849: 8) protologue, and also after a careful study of both the single specimen of that taxon at MPU and plants from its classic locality, parasitic on Helichrysum stoechas, that Dr Michaud kindly collected. Biogeographical and bioclimatic features follow Rivas-Martínez et al. (2001). Authors of taxon names are presented according to Brummitt & Powell (1992), and those of syntaxon names follow the standardization by Izco (in RivasMartínez et al., 2002).

RESULTS OROBANCHE PORTOILICITANA A. PUJADAS

&

M. B. CRESPO, SP. NOV. Orobanche olbiensis sensu A. Pujadas-Salva & M. B. Crespo in Collect. Bot. (Barcelona) 25(2): 231—237. 2000. Diagnosis: Planta (11—) 15—34 cm alta, caulis simplex aut ramosus, ramificatus sub plano soli, in profunditate. Caulis firmus aut tennis, albidus, saepe leviter latior ad medium quam ad basim, (3—) 7—12 mm latus ad basim, vix auctus (3—) 5—13 mm latus ad medium, regione superiore pubescenti-glandulosa, pilis brevissimis 0.1—0.3 mm longis, densis. Folia supera 8— 14 x 2—5 mm, ovata vel lanceolate, densa. Inflorescentia (4)7—15 cm longa, 30—80-flora, densissima. Flores sessiles, erectae vel erecto-patentes. Bracteae 6—9 mm longae, calyce breviores. Calyx 8—11 mm longus, nervis paulo conspicuis. Corolla 16—21 mm longa, cum pilis glandulosis 0.3—0.5 mm longis dorsodensis, marginibus labiorum ciliatis, pilis 0.3—0.4 nun longis, dilute caesia. Stamina 4—6 mm supra basem tubi corollae inserta, glabra. Antherae 1.3—1.7 mm longae, oblongae; apiculus 0.3—0.5 mm longus. Stylus albus, cum pilis glandulosis brevibus dispersis ad apicem densioribus. Instructus supra radices generis Centaureae parasitica. Description: Herbs up to (11—) 15—34 cm high, undivided or ramose, commonly broadly ramified under -

ground (each branch sometimes resembling a single individual). Stem stout or sometimes slender, whitish (3—) 7—12 mm in diameter at the base, usually somewhat swollen in the middle part (3—) 5—13 mm in diameter; subglabrous below, pilose-glandulose upwards, densely covered with very short hairs 0.1— 0.3 mm long. Basal leaves 4—10 x 4—8 mm, triangular to lanceolate, imbricate; upper leaves 8—14 x 2—5 mm, ovate to lancolate, shortly pubescent and densely arranged. Inflorescence (4—) 7—15 x 2—3.2 cm, very dense and 30—80 flowered, cylindrical, usually attenuate in the upper part and rounded at the apex. Flowers sessile, erect to erect-patent. Floral bracts 6— 9 x 1.8—3 mm, broadly lanceolate, shorter than calyx. Bracteoles 6—8 x 0.3—0.8 mm, linear, equalling the bracts. Calyx 8—11 mm long, with the tube equalling lobes in length; calyx lobes white or tinged with pale blue, lanceolate-subulate, covered with short glandulose hairs (c. 0.3 mm), and weakly nerved. Corolla 16— 21 mm long, pale blue and whitish towards the base, densely clothed with short (0.3—0.5 mm) glandular hairs in the outer side and margins of lips ciliate (cilia 0.3—0.4 mm long); during flowering more or less straight, tubulose or slightly infundibuliform, somewhat galeate at the back, 3—6 mm wide at the throat and 1.5—2 (—3) mm wide near the base, not swollen downwards and very slightly contracted in the insertion of the stamen filaments (in fruit, the corolla is strongly swollen at the base and notably contracted in the insertion of stamen filaments); upper lip shortly bilobed and somewhat emarginate, with lobes straightforward or patent; lower lip with subequal lobes, straightforward or deflexed, ovate, subobtuse and shortly apiculate, dentate on margins. Filaments of stamen glabrous, white, inserted 4—6 mm above the base of corolla tube; anthers white, oblong, 1.3— 1.7 mm long (including the apiculum, 0.3—0.5 mm long), with several hairs c. 0.5 mm long at the base, sometimes glabrous. Ovary white, 7—9 mm long, glabrous. Style white, loosely pilose-glandulose, with short glandulose hairs more densely disposed at the apex. Capsule 7—10 x 3.5—5 mm, ovate. Parasite on roots of Centaurea, specially on C. seridis L and C. aspera L. ssp. stenophylla (Dufour) Nyman. It belongs to sect. Trionychon Wallr. Etymology: The epithet `portoilicitana' refers to Portus Ilicitanus, the Roman name of the current Santa Pola (Alicante province), in the surroundings of which most of the known populations of the new species grow. Holotype: SPAIN, Alicante: Elx, La Marina, pr. Hostal Galicia, UTM: 30SYH0724, 3 m, 26.ív 2002, ubi M. B. Crespo, A. Pujadas, E. Triano & C. Burgadella legerunt (COA 31083).

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A NEW OROBANCHE FROM SOUTH-EASTERN SPAIN 99

Isotype: ABH, MA. Iconography: Figure 1; Pujadas-Salva (2000: 234, ut O. olbiensis).

& Crespo

ECOLOGY AND PHYTOSOCIOLOGY Orobanche portoilicitana grows on deep sandy soils, sometimes weakly nitrified by human effect, in coastal sand dunes with a moderate influence of sea spray. It forms part of different psammophilous plant communities of natural ecosystems. On the one hand, it grows in dwarf shrubby vegetation belonging to the association Loto cretici-Crucianelletum maritimae Alcaraz et al., 1989 (All. Crucianellion maritimae Rivas Goday & Rivas Mart. 1963, Ord. Crucianelletalia maritimae Sissingh 1974, Cl. Ammophiletea Braun-Blanq. & Tüxen ex Westhoff et al., 1946), which occupies the leeward areas of well-established secondary dunes with slightly movable substrates. On the other hand, O. portoilicitana also participates of subnitrophilous hemicryptophytic communities belonging to the association Eryngio maritimi-Sporoboletum arenarii (Arenes ex Géhu & Biondi 1994) Rivas Mart. & Cantó in Rivas-Martínez et al., 2002 [All. Sporobolion arenarii ( Géhu & Gehu-Franck ex Géhu & Biondi 1994) Rivas Mart. & Cantó in Rivas-Martínez et al., 2002, Ord. Ammophiletalia Braun-Blanq. 1933, Cl. Ammophiletea Braun-Blanq. & Tüxen ex Westhoff, Dijk & Passchier 1946), which grows usually in flat areas nearby footpaths and tracks leading to the seashore across the dune system and in other sandy areas nitrified and compacted by human activity, mostly during the summer. In all cases, it parasitizes on Centaurea seridis L., C. aspera L. ssp. stenophylla (Dufour) Nyman, and the hybrid between them: C. x subdecurrens Pau nothossp. albuferae (M. J. Costa) M. J. Costa, M. B. Crespo & Mateo.

DISTRIBUTION, BIOCLIMATOLOGY AND BIOGEOGRAPHY Orobanche portoilicitana is known only from the south-eastern areas of the Iberian Peninsula. It should be currently regarded as a stenochorous endemism of the coastal ecosystems of Alicante province, where it grows in a reduced and discontinuous area from Cabo de las Huertas (north of Alicante city) to La Marina (east of Elche) (Fig. 2). Bioclimatically, these territories are located in the Thermomediterranean Semiarid stage of the Alicantine chorological sector (Murciano-Almeriense province; sensu RivasMartínez et al., 2001). The climate is typically Mediterranean maritime, with a mild and rather rainy winter and a very hot and dry summer. The drought period usually extends over six months or even more.

Both host taxa on which O. portoilicitana parasitizes (C. aspera ssp. stenophylla and C. seridis) are mostly distributed along the coastal areas of the eastern and south-eastern Iberian Peninsula and the Balearics, with some unconfirmed localities in northern Africa and even southern France (cf. Boles & Vigo, 1996). Therefore, the area of O. portoilicitana could probably extend into other surrounding Iberolevantine territories (including the Balearic Islands). However, our fieldwork in the neighbouring provinces where both hosts grow together has not yielded new populations to date. DISCUSSION In the context of sect. Trionychon, O. portoilicitana shows a number of characteristics, described in the diagnosis, not present jointly in any other known taxon of that group. Although similarities exist that relate it to O. olbiensis, important morphological divergences between both taxa warrant recognition at the species rank (Table 1). With regard to O. olbiensis, the new species differs mainly by being taller (up to 35 cm high) and more robust. Stems are up to 13 mm wide (slightly broadened around the middle part) and usually ramified, with many underground branches each of them resembling single individuals, and the upper leaves longer (up to 14 mm long). Inflorescences are bigger, very dense and compact, up to 15 cm long, many-flowered (up to 80 flowers). Flowers are erect to erect-patent, and show bigger corollas (16-21 mm) with rather long (0.4-0.5 mm) hairs in the outer side and also longer (0.3-0.4 mm) cilia on the margins of lips. Anthers bear a long apiculum and are comparatively greater (1.3-1.7 mm). There are other differences in host plants and ecological behaviour. Orobanche portoilicitana parasitizes Centaurea species colonizing exclusively the first vegetation belt of sand dune ecosystems, whilst 0. olbiensis was only reported as parasitic on Helichrysum which grew in littoral scrubs. The new species can be also considered close to coastal O. mutelii due to some similarities in length of calyx and corolla. However, O. portoilicitana can be easily differentiated because its stem is somewhat swollen in the middle part, its inflorescence is very dense and many flowered, with flowers erect to erectpatent; the calyx is weakly nerved, with pale blue corolla, somewhat galeate; filaments are glabrous, and anthers long apiculate. CONSERVATION STATUS AND PROPOSALS During the last few years, the known populations of O. portoilicitana have remained almost unaltered,

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Alicante, Elx, La Marina, pr. Hostal Galicia, COA Figure 1. Orobanche portoilicitana A. Pujadas & M. B. Crespo, appearance; B, flower, front view; C, flower, side view; D, bract; E, open calyx and bracteoles; F, open 31083, holotypus: A, corolla and androecium; G, anther; H, gynoecium.

© 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 146, 97-102

A NEW OROBANCHE FROM SOUTH-EASTERN SPAIN 101 with about 250 individuals. However, most of those individuals are concentrated in southern populations (e.g. La Marina) whilst in northern populations (e.g. Cap de les Hortes and Urbanova, near Alicante) less than five specimens were counted. This fact together with the intrinsic fragility of the sand dune ecosystems and the pressure of human activities over the last decades, leads us to suggest labelling this new species as `endangered' – EN Blab(i, ii, iii, iv, v) + 2c(iv); C2a(i); D – according to the categories given in IUCN (2001). For these reasons, conservation measures should be established urgently to preserve the populations and natural habitats of O. portoilicitana. For this purpose, the declaration of new microreserves of flora (cf. Laguna, 1996, 2001) has proved a very effective tool for plant conservation. Fortunately, two of the known populations are already included in such protected

0

500 km

0

40 km

Figure 2. Distribution of Orobanche portoilicitana A. Pujadas & M. B. Crespo: A, general distribution; B, sites in Alicante province.

areas (namely, Platja del Carabassí and Platja de Pinet). However, a new microreserve should be established in the exiguous sand dune areas next to Cap de les Hortes, because other rare taxa such as Orobanche tunetana Beck and O. elatior ssp. icterica (Pau) A. Pujadas are also found there, and this site is dramatically threatened by urbanization.

MATERIAL STUDIED Orobanche portoilicitana A. Pujadas & M.B. Crespo, sp. nov. SPAIN. Alicante: Alicante, Cap de les Hortes, 30SYH2648, sobre Centaurea aspera L. ssp. stenophylla (Dufour) Nyman 26.iv.2002, A. Pujadas, E. Triano, C. Burgadella, COA 31084. Alicante, Urbanova, dunas costeras, 30SYH1639, 5 m, sobre C. seridis L., 28.v.1997, E. Camuñas & M. B. Crespo, ABH 36528. Ibídem, 30SYH1739, 3 m, 13.v.1999, E. Camuñas, ABH 42152. Elx, Arenals del Sol, Platja del Carabassí, 30SYH1735, 5 m, sobre Centaurea x subdecurrens Pau nothossp. albuferae (M.J. Costa) M. J. Costa et al. 18.iv.1997, A. Pujadas & M. B. Crespo, COA 17597. Elx, Arenals del Sol, Platja del Carabassí, 30SYH1735, 5 m, sobre Centaurea x subdecurrens Pau nothossp. albuferae ( M. J. Costa) M. J. Costa et al. 26.iv.2002, M. B. Crespo, A. Pujadas, E. Triano & C. Burgadella, COA 31085. Santa Pola, Playa del Pinet, 30SYH0826, 5 m, sobre C. seridis L., 29.iv.1998, M. B. Crespo, ABH 43121. Santa Pola, Playa del Pinet, 30SYH0826, 22.iv.2000, M. B. Crespo, COA29597. Elx, La Marina, pr. Hostal Galicia, UTM 30SYH0724, 3 m, 26.iv. 2002, M. B. Crespo, A. Pujadas, E. Triano & C. Burgadella, COA 31083, Isotypus. Orobanche olbiensis (Coss.) Nyman, Syll. Fl. Eur. 133 (1854)

Table 1. Main differences between Orobanche portoilicitana and O. olbiensis

Height (cm) Stem diameter in the middle part (mm) Stem branching Upper leaf length (mm) Inflorescence length (cm) Number of flowers Flower position Corolla length (mm) Corolla hairs length (mm) Lip cilia length (mm) Anther length (mm) Apiculum length (mm) Host plant (genus)

O. portoilicitana

O. olbiensis

(11—) 15—35 (3—)5—13 branched or simple 8—14 (4—)7—15 30—80 erect to erect-patent 16—21 0.3—0.5 0.3—0.4 1.3—1.7 0.3—0.5

4—15 (1—) 1.5—2.5 simple 5—6 2.5—6 7—20 erect-patent 12—15 c. 0.1 c. 0.2 0.9—1.1 0.1

Centaurea

Helichrysum

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FRANCE: Hyéres (83), Porquerolles á la plage d'Argent, sur Helichrysum stoechas, 5.vi.2001, H. Michaud, COA 31082. Var, Roquebrune, Guirigui, lieux decouverts dans les bois, vs. 1943, R. -, Herbarium C. Bertrand, MPU-Herbarium Coste (sub Phelypaea olbiensis).

ACKNOWLEDGEMENTS We wish to thank: Elena Camuñas (University of Alicante) for her support and information on natural populations; Dr Henri Michaud for the plant material from Porquerolles; Santos Cabello for the revised Latin description; the curators of the herbaria ABH, ALME, BC, BCC, `Colegio La Salle de Almería', `Col.legi Oficial de Farmacéutics de les Illes Balears', HJBS, `Herbarium Orell-Casanovas', `Herbari Universitat Illes Balears', MA, MAF, MGC, MUB, MPU, NCY, P, VAB, VAL, and W, for the loan of plant material; and Laura Plaza for the editing of the images. Concetta Burgadella and Enrique Triano kindly assisted us during the fieldwork.

Holmgren PK, Holmgren NH. 1993. Additions to Index Herbariorum ( Herbaria), Edition 8 — Second Series. Taxon 42: 489—505. Holmgren PK, Holmgren NH, Barnett LC. 1990. Index Herbariorum. Part 1. The herbaria of the World: Regnum Vegetabile 120, 8th edn. New York: New York Botanical

Garden. IUCN. 2001. IUCN Red List Categories, Version 3.1. Prepared by the IUCN Species Survival Commission. Gland: World Conservation Union. Laguna E. 1996. Conservación in situ mediante microrreservas de flora en la Comunidad Valenciana. Boletín de la Real Sociedad Española de Historia Natural, Tomo Extraordinario del 125 Aniversario: 379—381. Laguna E. 2001. The micro-reserves as a tool for conservation of threatened plants in Europe. Nature and Environment

121. Strasbourg: Council of Europe. Pujadas-Salva AJ. 2001. Aportació al coneixement del genere Orobanche L. als Paisos Catalans. Orsis 16: 71—88. Pujadas-Salva AJ. 2002. Orobanche L. In: López-Sáez JA, Catalán P, Sáez LI, eds. Plantas parásitas de la Península Ibérica e Islas Baleares. Madrid: Mundi-Prensa, 348—

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Bolas O, Vigo J. 1996. Flora deis paisos Catalans 3. Barcelona: Barcino. Brurnmitt RK, Powell CE. 1992. Authors of plants names. Kew: Royal Botanic Gardens. Cosson ESC. 1849. Notes sur quelques plantes critiques. Paris: Librairie de Victor Masson. Foley MJY. 2001. Orobanche L. In: Paiva J, Sales F, Hedge IC, Aedo C, Aldasoro JJ, Castroviejo S, Herrero A, Velayos M, eds. Flora Iberica 14. Madrid: Real Jardín Botánico, CSIC, 32—72.

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© 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 146, 97—102

Introduction Material and methods Results O. portoilicitana Diagnosis Description Etymology Holotype Iconography Ecology and phytosociology Distribution, bioclimatology and biogeography Conservation status and proposals Discussion Material studied Acknoledgements References Figure 1. Icon Figure 2. Distribution Table 1. Main differences...