An apomictic Limonium with a single pollen-stigma combination (B type), Limonium interjectum, is proposed as a new species to replace the invalid name Statice dianiae Pau. The new species has a restricted distribution on the sea costs of eastern Spain (Alicante Province). It shows a striking morphological similarity to L. virgatum (Willd.) Fourr. and L. girardianum (Guss.) Girard; on this basis it is suggested that L. interjectum is a hybrid derived from the two species.Se describe Limonium interjectum como especie nueva del Levante peninsular español, el cual había sido previamente denominado con el nombre ilegítimo de Statice dianiae Pau. L. interjectum es una especie apomíctica que presenta una sola combinación polínico estigmática (tipo B). Sus afinidades morfológicas son próximas a L. virgatum (Willd.) Fourr. Y L. girardianum (Guss.) Girard, y se sugiere que L. interjectum es un taxon hibridógeno que pudo haberse originado a partir de ambas especies

Rosello, Josep A.

Soler, Jaume X.

English

Anales del Jardín Botánico de Madrid (Real Jardín Botánico - CSIC)

ON THE STATUS OF STATICE DIANIAE PAU (PLUMBAGINACEAE)* by JAUME X. SOLER & JOSEP A. ROSSELLÓ** Resumen SOLER, J.X. & J.A. ROSSELLÓ (1997). Consideraciones sobre Statice dianiae Pau (Plumbaginaceae). Anales Jará. Bot. Madrid 55(1): 9-15 (en inglés). Se describe Limonium interjectum como especie nueva del Levante peninsular español, el cual había sido previamente denominado con el nombre ilegítimo de Statice dianiae Pau. L. interjectum es una especie apomíctica que presenta una sola combinación polínico-estigmática (tipo B). Sus afinidades morfológicas son próximas a L. virgatum (Willd.) Fourr. y L. girardianum (Guss.) Girard, y se sugiere que L. interjectum es un taxon hibridógeno que pudo haberse originado a partir de ambas especies. Palabras clave: Plumbaginaceae, Limonium, taxonomía, táxones apomícticos, hibridación, flora mediterránea. Abstract SOLER, J.X. & J.A. ROSSELLÓ (1997). On the status of Statice dianiae Pau (Plumbaginaceae). Anales Jard. Bot. Madrid 55(1): 9-15. An apomictic Limonium with a single pollen-stigma combination (B type), Limonium interjectum, is proposed as a new species to replace the invalid ñame Statice dianiae Pau. The new species has a restricted distribution on the sea costs of eastern Spain (Alicante Province). It shows a srriking morphological similarity to L. virgatum (Willd.) Fourr. and L. girardianum (Guss.) Girard; on this basis it is suggested that L. interjectum is a hybrid derived ftom the two species. Key words: Plumbaginaceae, Limonium, taxonomy, apomictic taxa, hybridization, Mediterranean flora. INTRODUCTION efforts, some taxonomic and nomenclatura! issues remain largely unresolved, mainly in The western Mediterranean Basin is one of the Iberian Peninsula and the Balearic Islands. the most important regions where the genus One of the ñames awaiting clarification is that Limonium, notably section Limonium, has of Statice dianiae Pau. This taxon was diversified. Comprehensive treatements of obscurely published by PAU (1898) on the PIGNATTI (1972, 1982), ERBEN (1993) and basis of plants gathered at one locality in ARRIGONI & DIANA (1993) have greatly southeast Spain (Alicante Province). He improved the taxonomy, nomenclature and tentatively related the new plant to distribution of most taxa. Despite these L. virgatum (Willd.) Fourr. and L. girar-* Part three of the series "A taxonomic and biosystematic revisión of the genus Limonium (Plumbaginaceae)". For part two see Anales Jard. Bot. Madrid 54:285-289 (1996). ** Botánica, Departamento de Biología Vegetal, Facultad de Ciencias, Universidad de Valencia. E-46100 Burjassot (Valencia). 10 ANALES JARDÍN BOTÁNICO DE MADRID, 55(1) 1997 Fig. 1 .-Original specimens of Statice dianiae Pau (MA). J.X. SOLER & J.A. ROSSELLÓ: STATICE DIANIAE 11 dianum (Guss.) Girard (sub Statice). But the only information given by the author is the very short diagnosis of S. dianiae which refers to its narrow leaves. Since then, the taxonomic status of 5. dianiae has not been reappraised in any of the papers dealing with the European species of Limonium (cf. GREUTER & al., 1989). In fact, the ñame is not included in the recent comprehensive treatment of the Iberian taxa (ERBEN, 1993). The examination of the original material of S. dianiae (MA; fig. 1) revealed that the plant is very similar to L. virgatum, but constant and conspicuous morphological differences do exist between them. More herbarium specimens from the surroundings of the type locality which resembled S. dianiae became available (ABH, MA) when the collections of L. virgatum were examined. Such new material was labelled either as L. virgatum or as a hybrid between this species and L. rigualii Crespo & Erben, a restricted triploid species growing in the same área (ERBEN, 1991,1993). Living plants collected from the type locality permitted detailed study, revealing that S. dianae is an apomictic species sharing several diagnostic features with L. virgatum and L. girardianum. These data suggest that S. dianae could be of hybrid origin, arising from a cross between the former two species. Unfortunately, the ñame Statice dianiae was not validy published, since PAU (1898) stated that it was a provisional ñame. The use of the word "provisionally" precludes valid publication under the rules of the Botanical Nomenclatural Code and therefore it could not be used as basionym for its transfer to Limonium. No other available ñames exist. For these reasons we propose the southeast Spanish plant as a new species, L. interjectum. RESULTS Limonium interjectum Soler & Rosselló, sp. nov. (figs. 2,3) = Statice virgata var. macroclada Pau, Actas Soc. Esp. Hist. Nat. 27: 199 (1898) = Statice dianiae Pau, Actas Soc. Esp. Hist. Nat. 27: 199 (1898), nom. inval. A Limonium girardianum et L. virgatum valde similis. A L. virgatum affinis sed rami omnia fértiles, folia glauca, leviter, florendi tempore persistentia; spicae dense dispositis, minoribus, atque bractea inferior minoribus differt. A L. girardianum similis sed inflorescentia ramosa, folia minoribus, anguste obovata, spiculae subarcuate differt. Derivatio nominis: from the latin, interjectum, i = intermediate. Typus. Hispania, Lucentina, locus dictus Montanyar (31SBC59) prope oppidulum Xábia, in rupestribus maritimus ubi J.X. Soler legi 5-VII-1992 (holotypus: VAB 933865; isotypus: ABH, G, MA, M). Perennial with many stems, glabrous. Caudicles 1.5-15 cm long, spirally leafly in upper third. Basal leaves erect, usually green at anthesis, 15-80 x 4-9(12) mm, glaucous. Blade lanceolate, tip acute, with a short terminal apiculum; flattened at the margins, smooth on both faces, usually 3-nerved. Petiole canaliculate, 3/4 as long as the blade, 1-3.5 mm wide. Stem 15-45 cm long, erect, branched in the lower half (rarely in the lower third); usually without sterile branches (rarely with 1-2). Inflorescence paniculate, loosely branched, triangular. Branches up to 16 cm long, subcurved at the ends, obliquely inserted (45-60°), flowering in the upper third; non-flowering branches absent. Spikes 10-30 mm long, straight to curved, obliquely inserted, with 5-8 spikelets per cm. Spikelets 6-7.2 mm long, 2-5(8) flowered. Outer bract 2.9-3.4 x 2-2.3 mm, triangular-ovate, acute, margin membranous highly variable; central partsubfleshy, 2-2.5 x 1.8-2 mm, triangular to oblong; tip 0.9-1 mm, acute, the acumen reaching the margin. Middle bract 2.4-3 x 1.7-2 mm, broadly oblong-obovate, blunt to subemarginate, membraneous. Inner bract 5.2-6 x 3-3.6 mm, oblong-elliptical, blunt to acute, with a thin membranous margin; central part subfleshy, 4-5 x 2-3 mm, oblong, acuminate, acumen 0.9-1.2 mm long, not reaching the margin. Calyx 5.5-6.5 mm long, overtopping by 1-2 mm the inner bract, tube hairy, with short eglandular hairs; tooth 12 ANALES JARDÍN BOTÁNICO DE MADRID, 55(1) 1997 Fig. I.-Limonium interjectum: Habit. I.X. SOLER & J.A. ROSSELLÓ: STATICE DIANIAE 13 ca. 0.8-1 mm, semielliptic to triangular-ovate; midrib usually reaching the calyx lobes. Corolla funnel-shaped. Petáis 9 x 3 mm, cunéate, violet. Flowering: July-September Material examined SPAIN. AUCANTE: Denia, Palmar del Pi, VI-1897, Pau, MA 92085. Xábia, 17-VII-1973, Segura, MA 310850. Xábia, Montanyar, l-VII-1990, Soler, VAB; ibidem, 2-VIÜ-1993, Soler, VAB. Xábia, Montanyar, 2-VII-1990, Soler, VAB; ibidem, 2-VIII-1993, Soler, VAB. Xábia, Montanyar de Baix, 15-VÜ-1992, Barber, ABH 1972, 1973, 1974, 1975, 1976, 1977. Xábia, Arenal, VII-1990, Soler, VAB; ibidem, 2-VIIM993, Soler, VAB. Xábia, Cala Blanca., 2-Vm-1991,¿agu«a & al., ABH 301. Pollen-stigma combination andfertility All individuáis from the three sampled populations of L. interjectum had a reduced fertility. Pollen fertility from 32 individuáis ranged from 0 to 5.1% (mean = 0.6%; n = 3.206 pollen grains) as determined by the staining method used. Shape and length irregularities were very frequent and also involved variation in the number of colpi. The few stained pollen grains which could be found in some individuáis have a greater length and are presumed to be of diploid or polyploid nature. The only pollen/stigma combination found in L. interjectum is the B type. The surrounding populations of L. virgatum so far studied also present pollen abnormalities and have the same pollen-stigma combination, which agrees with data from other distant localities (LLORENS, 1986; Rosselló, unpublished data). On the other hand, Limonium girardianum has a pollen/stigma combination of the A type (Rosselló, unpublished data). Ecology and distribution The new species grows on calcarenites of quaternary fossil sand beaches which are located near to the sea. Associate species includes Limonium delicatulum (Girard) Kuntze, L. virgatum (Willd.) Fourr., Crithmum maritimum L., Halimione portulacoides (L.) Aellen, Juncus mariti-mus L., Inula crithmoides L. and Salicornia ramosissima J. Woods. Limonium inter-jectum is restricted to very few populations scattered through the Xábia Bay (North of Alicante Province). There is herbarium evidence that this species occurred at one locality (Palmar del Pi) situated 15 km apart from the extant populations, but an intensive search for these plants at that site was not successful. In the last decades, urbanistic development affecting the eastern coastal environments has dramatically changed most of the main saltshore communities, so it is possible that the distribution of L. interjectum was broader in the past. As far as we are aware, over 400 individuáis of L. interjectum are currently known. This, together with the fact that some of the populations are located at sites where urbanistic alterations are planned, render L. interjectum an endangered species. DISCUSSION Limonium interjectum has striking similarities with L. virgatum, with which it has been confused in the past. Despite this, the lack of non-fiowering branches, the smoothness and glaucous leaves, the smaller spikes and outer bracts, easily sepárate both taxa (table 1). However, both species have similar sized inner bracts and calyx tubes, as well as similarly shaped leaves. The characters of L. interjectum not present in L. virgatum are reminiscent of those of a L. girardianum-like plant. Limonium virgatum and L. interjectum occur together and L. girardianum also grows in the vicinity (Soler, unpublished data) so it is conceivable to interpret L. interjectum as having arisen by a cross between the other two taxa. Although the morphological and chorological data do not conflict, this evidence alone is by no means conclusive and molecular data need to be obtained before the origin of L. interjectum can be firmly established. Limonium virgatum and L. girardianum belong to different subsections of section Limonium as currently circumscribed (BOISSIER, 1848). This taxonomic divergence does not threaten the proposed origin of L. interjectum, since the available pattern of reticulation in the 14 ANALES JARDÍN BOTÁNICO DE MADRID, 55(1) 1997 Fig. S.-Limonium interjectum: a, inner bract; b, middle bract; c, teeth of calyx; d, outer bract; e, calyx; f, spikelet; g, leaves. J.X. SOLER & J.A. ROSSELLÓ: STATICE DIANIAE 15 TABLEI MORPHOLOGICALFEATURESONL/MON/Í/AÍ INTERJECTUM, L. VIRGATUM ANDL. GIRARDIANUM [Data of L. virgatum and L. girardianum are taken from ERBEN (1993)] Leaves: blade apex roughness colour Sterile branches Spikes Outer bract Inner bract Calyx L. virgatum lanceolate variable verrucate deep green many 20-80(120) mm 1.9-2.8 x 1.9-2.5 mm 5.1-6.5 x 2.9-4 mm 5.3-6.1 mm L. interjectum narrowly lanceolate to oblanceolate acute smooth green-glaucous absent (rarely 1-2) 10-30 mm 2.9-3.4 x 2-2.3 mm 5.2-6 x 3-3.6 mm 5.5-6.5 mm L. girardianum oblanceolate-spatulate acute smooth green-glaucous absent 8-16 1.8-2.3 x 2-2.6 mm 3.8-4.6 x 3.8-4.4 mm 4.1-4.8 mm genus, at least in the western Mediterranean Basin, is not only intrasectional but also intersectional (ERBEN, 1993). On the other hand, since the proposed parents of L. interjectum are apomictic and have severe pollen irregularities it must be shown that such biological barriers could be effectively overeóme to allow gene flow between them. Records of gene flow between apomictic or between sexual and apomictic Limonium taxa are not rare (ERBEN, 1993; Rosselló & Sáez, unpublished data), but unfortunately only morphological data support this reproductive avenue. Molecular work is now in progress to assess i) the proposed parentage of L. interjectum, ii) the ovule and pollen donor of the cross, and iii) whether múltiple events are involved in the origin of L. interjectum. ACKNOWLEDGMENTS The authors are much indebted to R. Silva and L. Sáez for the illustrations and to W. Greuter and F. Muñoz for nomenclatura! advice. This work was partially supported by a research grant PB 93-0350 of the spanish DGIC YT. REFERENCES ARRIGONI, P.V. & S. DIANA (1993). Contribution á la connaissance du genre Limonium en Corsé. Candolka 48:631-677. BOISSIER, E. (1848). Plumbaginaceae. In: A.P. de Candolle, Prodomus systematis naturalis regni vegetabais. Vol. 12:617-696. Paris. ERBEN, M. (1991). Bermerkungen zur Taxonomie der Gattung Limonium VI. Mitt. Bot. Staatssamml. München 30:459-478. ERBEN, M. (1993). Limonium. In: S. Castroviejo & al. (eds.), Flora iberica. Vol. 3:2-143. Madrid. GREUTER, W., H.M. BURDET & G. LONG (1989). Med-Check List 4: Dicotylledones (Lauraceae-Rhamnaceae). Ed. Conserv. Jard. Bot. Genéve. LLORENS, L. (1986). Revisión sistemático-taximétrica del género Limonium Miller en la isla de Mallorca (I). Lazaroa 8:11-68. PAU, C. (1898). Noticia de algunas plantas curiosas o nuevas. Actas Soc. Esp. Hist. Nat. 27:196-200. PIGNATTI, S. (1972). Limonium. In: T.G. Tutin & al. (eds.), Flora Europaea. Vol. 3: 38-50. Cambridge University Press. PIGNATTI, S. (1982). Flora d'Italia. Vol. 2. Ed. Edagricole. Bologna. Aceptado para publicación: 20-H-1996 

Consideraciones sobre Statice dianiae Pau (Plumbaginaceae)

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Consideraciones sobre Statice dianiae Pau (Plumbaginaceae)

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