Botanical Journal of the Linnean Society, 2003, 143, 331—335. With 2 figures

A new species of

Valantia

(Rubiaceae) from Spain

JUAN A. DEVESA* and ANA ORTEGA-OLIVENCIA Departamento de Biología y Producción Vegetal (Botánica), Facultad de Ciencias, Universidad de Extremadura, E-06071 Badajoz, Spain Received March 2003; accepted for publication July 2003

A new species of Valantia, V. lainzii Devesa & Ortega-Olivencia, endemic to the coastal zone of Granada (southern Spain) is described. The species recalls V. muralis L. in its general aspect, and V. deltoidea Brullo in the type of fructiferous body. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143, 331– 335.

ADDITIONAL KEYWORDS: Flora Iberica – Iberian Peninsula – Rubieae – Rubioideae – taxonomy.

INTRODUCTION The genus Valantia L. is one of the 550 genera of the Rubiaceae, a monophyletic family (Bremer et al., 1999) for which around 9000 species have been described (Judd et al., 1999), or according to Mabberley (1997) up to 630 genera and 10 200 species. Its main diagnostic character is having a fructiferous body originating from the fusion of the accrescent peduncle of the inflorescence and the flower pedicels (Figs 1E, 2) that houses and protects the fruit. The genus, which has major affinities with Cruciata Mill. and the representatives of Galium sect. Platygalium (DC.) W.D.J. Koch (e.g. Galium rotundifolium L. and G. scabrum L.), is regarded as included in the subfamily Rubioideae and the tribe Rubieae, a monophyletic group (Ehrendorfer, Manen & Natali, 1994; Manen, Natali & Ehrendorfer, 1994; Natali, Manen & Ehrendorfer, 1995) especially well represented in temperate zones. After its delimitation by Linnaeus (1753), who described Valantia muralis L. and V. hispida L., the genus has increased in taxonomic diversity to the present seven species (Brullo, 1979, 1980, 1991; Aiello, Brullo & Piccione, 1981). The two Linnaean species, until now the only ones present in the flora of the Iberian Peninsula, are widely represented in the Mediterranean region ( V. hispida also reaches the Macaronesian and Irano-Turanian regions), whereas *Corresponding author. E-mail: [email protected]

the others have a more restricted distribution. Thus, V. columella (Ehrenb. ex Boiss.) Bald. is present in north and east Africa; V aprica (Sibth. & Sm.) Boiss. & Heldr. in Crete and the southern Balkans; V. eburnea Brullo is endemic to Cyprus; and V calva Brullo and V. deltoidea Brullo are restricted to Sicily, in particular to the Isle of Linosa and the Rock of Busambra, respectively. All the genus's species are annual except for V aprica, which is also considered to be the most primitive member and is the only one with allogamy as its reproductive system (Ehrendorfer, 1971, 1988). The rest are autogamous, and form diaspores that, with the exception of V. columella ( which is anemochorous), are more or less strongly adapted to dissemination by epizoochory. Evolutionary processes within the genus have also involved major chromosome rearrangements (Ehrendorfer, 1988). From the basic primitive number, x= 11 (2n = 22, in V aprica), there would have arisen by dysploidy the basic secondary numbers 9 (2n = 18, V calva, V. muralis, and V. hispida) and 10, with polyploidy also playing an important role in speciation, 2n = 40 (x = 10, in V. columella) and 2n = 36 (x = 9, in V. deltoidea; Brullo, 1980). Its taxonomic diversity in the Iberian Peninsula was thought to be known following its treatment by Lange (1868) and more recently by Ehrendorfer (1976). However, the revision that the authors are currently carrying out for the Flora Iberica (Castroviejo et al. eds.) has revealed individuals from southern Spain that,

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Figure 1. A. Habit of Valantia lainzii Devesa & Ortega-Olivencia (MA 435243). B. Leaf. C. Male flower. D. Hermaphroditic flower. E. Fructiferous body. F. Mericarp. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143, 331—335

A NEW SPECIES OF VALANTIA 333

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Figure 2. A. Fructiferous body of Valantia hispida L. (SEV 9234). B. Fructiferous body of V. muralis L. (MA 509150). C. Fructiferous body of V. deltoidea Brullo (UNEX 30820).

because of their singular characteristics, are here described as a new species. This description is based on six individuals from the same collection. MATERIAL AND METHODS

All the morphological observations are based on stereoscopic examination of the material kept in the herbaria BC, GDA, GDAC, JAEN, MA, MGC, SEV, and UNEX (abbreviations according to Holmgren, Holmgren & Barnett, 1990). DESCRIPTION VALANTIA LAINZII DEVESA & ORTEGA-OLIVENCIA SP. NOV.

Herba annua, glabra. Caules ut plurimum 6 cm, ramificati. Folia (0.5)1—5 x 0.5–1.7(2) mm, 4-verticillata, elliptica, ovata vel obovata, obtusa, crassiuscula, margine leviter non-numquam revoluta. Cymae triflorae in verticillos plus minusve approximatos quaternatim dispositae, in pedunculos brevissimos impositae atque ii tandem accreti, recurvati et coaliti cum pedicellis ita ut mericarpia coniunctim plus minusve cooperiant. Flores proterandri, actinomorphi, laterales masculi atque trimeri, medius hermaphroditus atque tetramerus. Calyx nullus. Corolla 0.4–0.6 mm longa, 0.7– 1.4 mm diametro, cupulato-rotata, flavido-viridula. Stamina 3–4. Ovarium biloculare; styli 2; stigmata capitata. Corpus fructiferum 2.5–3 mm, deltoideum, albidum ant albido-luteolum, dorso laeve ant aculeis apicalibus (1)3(5) tantum praeditum, rigidis, conicis vel compressis (triangularibus), rectis ant leviter curvis, hyalinis; margine inferiore laevi aut pilis conicis non 0.1 mm attingentibus praedito. Mericarpia 1– 1.2 x 0.6–0.8 mm, plerumque 1 in unoquoque flore, reniformia, laevia, nigra, fasciculis linearibus rhaphidium albidorum c. hilum praedita.

Annual. STEMS of up to 6 cm, ramified from the base, ascendent or decumbent, tetragonal, at times with the angles slightly thickened, glabrous. LEAVES (0.5)1— 5 x 0.5–1.7(2) mm, in whorls of 4, elliptic, ovate, or obovate, obtuse, attenuate or shortly petiolate, somewhat fleshy, with the main vein more conspicuous, especially on the abaxial surface, and the lateral veins hardly visible, leaf margins occasionally slightly revolute, glabrous, finally reflexed, with ± abundant linear oblique bundles of raphides, whitish, generally dry or absent in flowering. Partial inflorescences cymose, axillary, with 3 flowers, arranged in whorls, 4 in each, ± approximate along the stems, on very short peduncles, accrescent at maturity, recurved and coalescent with the flower pedicels to form a fructiferous body that covers the mericarps. BRACTS in whorls of 4, similar to the leaves, decreasing in size towards the apex of the flowering branches. FLOWERS hermaphroditic or unisexual (plants polygamous), protandrous, actinomorphic, the lateral flowers of each cyme trimerons, male, and the central tetramerous, hermaphroditic. CALYX absent. COROLLA 0.4—0.6 mm in length and 0.7— 1.4 mm in diameter, sympetalous, cup-shaped to rotate, glabrous, yellowy-greenish; tube c. 0.1– 0.2 mm; lobules c. 0.3–0.5 mm, ovate, obtuse. ANDROECIUM with 3–4 stamens, alternipetalous, included, with smooth filaments, adnate to the corolla tube; ANTHERS 0.05–0.1 mm, oblong–ovoid, dorsifixed. OVARY with two locules, each with 1 ovule. STYLES 2, included; stigmas capitate. FRUCTIFEROUS BODY 2.5— 3 mm, deltoid, whitish or pale-yellowish, with dorsal side smooth or with only (1)3(5) stiff apical bristles, conical or compressed (triangular), straight or slightly curved, hyaline, and lower edge smooth or with conical hairs of less than 0.1 mm. MERICARPS 1–1.2 x 0.6– 0.8 mm, in general only one per flower, reniform, smooth, black, with linear bundles of whitish raphides in the region of the hilum.

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Typus: GRANADA, Calahonda. Playa de Calahonda, en arenal marítimo. 8.iii.1980. F. Pérez Raya (Holotype: MA 435243; designated here). Etymology: Dedicated to Dr Manuel Laínz, friend and untiring student of the Spanish flora.

leaves, which are glabrous rather than sparsely hirsute and ciliate. This is a microspecies that requires chorological and cytotaxonomic studies to delimit its true area of distribution in the Iberian Peninsula and its possible origin and affinities.

Iconography: Figure 1. Chorology, flowering time and ecology: Valantia lainzii is known only on the maritime sands of Calahonda beach in the province of Granada (southern Spain) where it was collected in early March 1980, cohabiting with V. hispida L. Although its area of distribution may extend to the neighbouring provinces of Málaga and Almería under similar ecological conditions, this possibility requires future prospecting.

MORPHOLOGICAL RELATIONSHIPS TO OTHER VALANTIA SPECIES Identification of the species of Valantia is especially straightforward when the examples are fruiting, because then the flower pedicels and peduncles fuse to form a mericarp dispersal unit (fructiferous body) with great taxonomic value. The morphology of this fructiferous body has been studied in all the known species (Aiello et al., 1981; Brullo, 1991). V. columella and V. calva are those that have the most clearly differentiated fructiferous body, covered with long hairs that are flattened in the former and smooth in the latter. In the other species, the fructiferous body is more or less aculeolate dorsally (e.g. V. hispida, Fig. 2A), with V. muralis, V. deltoidea, and V. lainzii being the only species to present a dorsal protuberance, deltoid or cylindrical, aculeolate at the tip (Figs 1E, 2B,C). V. lainzii has great morphological similarity with V. muralis. In fact, some coastal populations of V. muralis have the same habit and are somewhat succulent, condensed, and glabrescent, and there are populations occurring geographically next to Calahonda (e.g. Almuñecar, southern Granada), the only known population of V. lainzii. However, this new species differs from V. muralis most noticeably in the morphology of the fructiferous body. Thus, whereas V. muralis presents a very conspicuous dorsal protuberance, more or less cylindrical, culminating in 6–15 stiff bristles that are conical or compressed, in V. lainzii the fructiferous body is deltoid and culminates in (1)3(5) bristles. This structure is similar to that described in V. deltoidea (Brullo, 1980), but the internal edges of the crests are either smooth or possess conical hairs of less than 0.1 mm, rather than conspicuously fimbriate and with hairs 0.2–0.3 mm long as is common in V. deltoidea (and in V. muralis). It differs from V. deltoidea also in its very approximate flower whorls almost completely hiding the internodes, and in its

ACKNOWLEDGEMENTS We thank Professor S. Brullo for sending the V. deltoidea material, Dr M. Laínz for transcribing the Latin diagnosis, the curators of the herbaria referred to in the text and an anonymous reviewer for valuable comments on the manuscript. The work was financed by Spain's Ministry of Science and Technology through the projects Flora Iberica V and VI (PB96-0447 and REN2002-04634-0O5-04, respectively).

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© 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 143, 331–335