Orobanche Apuana (Orobanchaceae) a New Species Endemic to Italy

Phytotaxa 207 (1): 163–171 ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ PHYTOTAXA Copyright © 2015 Magnolia Press Article ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.207.2.2

Orobanche apuana (Orobanchaceae) a new species endemic to Italy

GIANNIANTONIO DOMINA1* & ADRIANO SOLDANO2 1 Dipartimento di Scienze Agrarie e Forestali, via Archirafi 38, 90123 Palermo, Italy; e-mail: [email protected] 2 Largo Brigata Cagliari 6, 13100 Vercelli, Italy *author for correspondence

Abstract

Orobanche apuana, a new species belonging to Orobanche sect. Orobanche, is described and illustrated from the Apuan Alps, Central Italy. Its relationships with the other species of the group of O. caryophyllacea (O. grex Galeatae) and with other Orobanche that parasitize Santolina species are examined. The names Boulardia latisquama, Orobanche lutea and O. teucrii are here lectotypified.

Key words: broomrape, Santolina pinnata, Mediterranean flora

Introduction

Orobanche Linnaeus (1752: 632), as unanimously accepted by all modern studies (Carlón et al. 2008, Crespo & Pujadas 2006, Domina 2009, Schneeweiss et al. 2004a, 2004b), in Europe and the Mediterranean area includes at least two well distinct groups from the morphological and karyological points of view. Despite this, researchers are divided with regard to the nomenclature to be used by grouping this taxon under a single genus with different sections (Crespo & Pujadas 2006, Domina 2009) or under different genera (Schneeweiss et al. 2004a, 2004b, Carlón et al. 2008). The taxonomic research on Orobanche s.l. in Europe is in full progress. In the last years, relationships between several taxa have been studied from the taxonomic and biological points of view (Jeanmonod 2007, Carlón et al. 2008, Piwowarczyk et al. 2014). These studies led to the discovery of new localities and to the description of new taxa (i.e. Pujadas-Salvà & Arguimbau 2008, Nobis et al. 2014a, 2014b). In Europe and the Mediterranean area 124 species are known: 43 belonging to Orobanche sect. Trionychon Wallroth (1822: 314) and 81 to Orobanche L. sect. Orobanche (Domina & Raab-Straube 2010 onwards). In Italy 40 (9+31) species occurs (Domina 2015). Four of them are endemic to very restricted areas: Orobanche australis Bertoloni (1846: 440) and O. denudata Moris (1828: [1]) in Sardinia; O. ebuli Huter & Rigo in Huter (1907: 354) in Central Italy, O. chironii Lojacono-Pojero (1878: 12) and O. thapsoides Lojacono-Pojero (1882: 60) in Sicily. During the revision of this genus in the Mediterranean, from the nomenclatural and taxonomic points of view (Domina et al. 2005, Domina & Arrigoni 2007, Domina & Mazzola 2007, 2009, Domina & Raimondo 2009, Domina & Stepanek 2009, Domina & Mazzola 2011, Domina et al. 2011, 2013, Domina & Danin 2014), we studied plants collected in the Apuan Alps in Central Italy. After meticulous field research and herbarium studies, we realized that this species belongs to the group of Orobanche caryophyllacea Smith (1798: 169) (O. grex Galeatae Beck 1930: 222) but we have not found in herbaria or in literature material referable to this taxon. So, we decided to describe it as a new species to science and to name it Orobanche apuana.

Material and Methods

This plant was first collected during a field trip to the Apuan Alps by Adriano Soldano in 1996. Although the place where this broomrape was found, along the Via Vandelli, on the western slope of Mt. Tambura, is one of the tracks that are mostly used by hikers and naturalists in the Apuan Alps, no records of broomrape referable to this species can be

Accepted by Lorenzo Peruzzi: 14 Apr. 2015; published: 8 May 2015 163 found in the local floras (Baroni 1897–1908, Pellegrini 1941, Ferrarini 1967, Barbero & Bono 1973, Ferrarini et al. 1997, Ansaldi & Maccioni 2006), in the Floras and checklists of Italy (Bertoloni 1846, Caruel 1884, Béguinot 1902, Pignatti 1982, Conti et al. 2005) or in the main floras and checklists of Europe (Chater & Webb 1972, Greuter et al. 1989, Domina & Raab-Straube 2010 onwards) and monographs (Beck 1890, 1930). We verified in the BOLO, FI, PAL, PI, RO and TO herbaria (acronyms follow Thiers 20015) that host the largest collections from this area and Italy; but no material referable to this taxon was found. Further excursions were done in the same place during the last years to observe its ecology and distribution. The new species is here compared with the type material of the species related by morphology or sharing the same host. The measures and character states of the compared taxa are taken directly from the studied specimens. This to preserve the taxa delimitation in their author’s sense. For O. apuana further observations and measures were done in the field on about 30 living individuals. In the typifications when a part of the handwriting in the labels was illegible it is reported here as […]. The handwritings of Baumgarten, Durieu and Holandre were identified by direct comparison with those in the herbarium labels of specimens of reliable source housed in P.

Description of the new species

Orobanche apuana Domina & Soldano, spec. nov. (Figs. 1, 2)

Type:—ITALY. Apuan Alps, Via Vandelli (Massa), 44°05’48”N, 10°13’09”E, 920 m a.s.l., dry pastures on carbonate lithosol, on Santolina pinnata Viv., 13 July 2014, A. Soldano 16532 (holotype PAL!, isotypes FI!, Herb. Soldano!).

Plant 15–30 cm tall with stem slender and erect, reddish-brown or yellow-brown, densely glandular-pubescent, laxly scaled. Lower scales triangular to oval, sparsely glandular pubescent; upper scales linear-lanceolate and sparsely glandular-pubescent, brown, erect. Inflorescence short and lax with 10–25 flowers. Bracteoles absent. Floral bract reddish-brown or yellow-brown, glandular pubescent, acute, about as long as or slightly shorter than the corolla, deflexed in the distal part. Calyx segments entire or bifid, 8–12 mm long, with teeth shorter than the tube, the upper tooth longer than the lower one or subequal. Flowers erect-spreading at first, bent forward later, with few purple veins. Corolla 17–24 mm long, inflated above the insertion of the stamens. Dorsal line of the corolla slightly curved in the lower part, almost straight in the middle and strongly flexed forwards near the upper lip. Upper lip of the corolla almost entire or slightly emarginated in two crenate lobes, with glandular hairs. Lower lip of the corolla with three, equal, crenate, deflexed lobes. Stamen inserted 3–5 mm above the base of the corolla, with filaments pubescent at the base. Anthers pubescent in the line of fusion. Style glandular-pubescent; stigma yellow, bilobate. Host:—Orobanche apuana is exclusively parasitic on the Apuan Alps endemic Santolina pinnata Viviani (1802: 31) (Asteraceae). The latter name, although dismissed by Garbari & Bechi (1992) and Peruzzi et al. (2014, 2015), is correct, because its alleged earlier homonym S. pinnata Donn (1800: 107) is not validly published (see, in particular, Art. 38.2 Ex. 3 of the ICN, McNeill et al. 2012). Flowering time:—Flowering and fruiting from late June, at lower altitudes, to early August in the higher ones. Ecology and Distribution:—Orobanche apuana was collected on the Apuan Alps, on the western slopes of Monte Tambura from 600 to 1200 m a.s.l. on sunny carbonate rocks and dry pastures on lithosol. A photo of this plant from Campocecina (Massa-Carrara), at 1250 m, the 11.07.2010, by Melania Marchi was seen also on the forum of Acta Plantarum (http://www.actaplantarum.org/floraitaliae/viewtopic.php?f=40&t=19583&p=133289&hilit=orobanche+c ampocecina#p133289). It is it plausible that O. apuana grows also in other localities of the Apuan Alps where its host, S. pinnata occurs locally abundant from 500 to 1500 m on pioneer thermo-xerophilous formations of cliffs with open woods. This vegetation hosts a rich component of taxa endemic to Central and Northern Italy (Ansaldi & Maccioni 2006), as Moltkia suffruticosa (Linnaeus 1763: 1667) Brand (1902: 1999) subsp. bigazziana Peruzzi & Soldano (2010: 385), Buphthalmum salicifolium Linnaeus (1753: 904) subsp. flexile (Bertoloni 1854: 413) Garbari (1971: 191), Leontodon anomalus Ball (1850: 9), Silene pichiana Ferrarini & Cecchi (2001: 246), Rhamnus glaucophylla Sommier (1894: 19). Further specimens seen (paratype):—ITALY. Via Vandelli (Massa), 44°05’48”N, 10°13’09”E, 920 m a.s.l., on Santolina pinnata Viv., July 1996, A. Soldano (Herb. Soldano!).

164 • Phytotaxa 207 (2) © 2015 Magnolia Press DOMINA & SOLDANO FIGURE 1. Orobanche apuana: a) habit; b) floral bracts; c) calyx segments; d) frontal view of the corolla; e) flower; f) corolla in section; g) pistil; h) seeds (Drawings by G. Domina on the original material).

Taxonomic relationships:—By the curvature of the dorsal line of the corolla, bent, straight and humped near the apex, this species can be ascribed to the group of O. caryophyllacea Sm. (O. grex Galeatae G.Beck). Schneeweiss et al. (2004a) consider the character of the shape of the dorsal line of the corolla unstable and, on the basis of nuclear ITS sequences, demonstrated that this group is polyphyletic. In fact, in Italy, it includes O. caryophyllacea Smith (1798: 169), O. lutea Baumgarten (1816: 215) and O. teucrii Holandre (1829: 322) that are morphologically close, but in the West part of the Mediterranean region also O. latisquama (Schultz 1848: 104) Battandier (1890: 659) occurs, that is

Orobanche apuana a new species endemic to Italy Phytotaxa 207 (2) © 2015 Magnolia Press • 165 a quite different taxon (Carlón et al. 2008). Orobanche apuana can be distinguished from the other species similar to O. caryophyllacea by the shape and the reduced size of the corolla, the shape of the calyx, the colour of the plant when dry and the shape of the floral bract (Fig. 3). The diagnostic characters are summarized in Table 1. All these species are faithfully recurring on the same host plant or family: O. apuana only on Santolina pinnata (Asteraceae), O. caryophyllacea on Rubiaceae, O. lutea on Fabaceae, O latisquama and O. teucrii on Lamiaceae (Rosmarinus and Teucrium respectively). In the western Mediterranean region, in the Iberian Peninsula, Balearic Islands and in the southwest of France O. santolinae Loscos & Pardo (1863: 79) occurs, parasitizing different species of Santolina at altitudes from 20 to 1900 m and flowering from May to August (Pujadas-Salvà 2010). In Corse O. cyrnea Jeanmonod et al. (2005: 257) was recently described. These broomrapes share only the host genus (Santolina) with O. apuana but there is no close taxonomic relationship between them. In fact O. apuana has larger flowers with differentiated curvature of the dorsal line of the corolla and belongs to the group of O. caryophyllacea (O. grex Galeatae Beck 1930: 222), O. santolinae and O. cyrnea have smaller flowers with constant curvature of the dorsal line of the corolla, thus belonging to the group of O. minor Smith (1797: 422) (O. grex Minores Beck 1930: 167). Jeanmonod et al. (2005), on the basis of morphology and rbcL sequences analysis, consider difficult the assignment of O. cyrnea to one of the actually recognized group. On the forum Acta Plantarum (see above), the photos of O. apuana are identified as Orobanche elatior Sutton (1798: 178). However, we are not able to find significant similarities with this species that, unlike O. apuana, has long, dense spike, dorsal line of the corolla with constant curvature, calyx segments bifid, rarely entire, lips of the corolla deeply emarginated the upper and deeply lobed the lower one.

FIGURE 2. Orobanche apuana in the field on Santolina pinnata (Photo by M. Soldano).

FIGURE 3. Comparison of flowers: a) O. apuana, b) O. caryophyllacea, c) O. teucrii, d) O. lutea, e) O. latisquama, f) O. cyrnea, g) O. santolinae. (Drawings by G. Domina on the type materials).

166 • Phytotaxa 207 (2) © 2015 Magnolia Press DOMINA & SOLDANO (Asteraceae) 2–4 dense 14–18 absent shorter narrow light brown bent forward O. santolinae mostly bidentate linear-lanceolate purple (yellow in whitish to pinkish equalling or shorter apochromatic plants*) Santolina equalling the lateral ones (Asteraceae) 2–2.5 dense 13–16 absent shorter narrow O. cyrnea lanceolate dark brown bent forward mostly bidentate whitish to purple yellow or whitish equalling or shorter Santolina equalling the lateral ones (Lamiaceae) 4–6 dense 17–35 yellow shorter narrow dark brown mostly entire inconspicuous O. latisquama elliptic-oblong erect spreading pink to deep purple equalling or little longer equalling the lateral ones Rosmarinum 3–6 dense broad 20–32 longer yellow shorter O. lutea Fabaceae light brown bent forward mostly bidentate linear-lanceolate yellow to pinkish highly accentuated equalling the lateral ones (Lamiaceae) lax 3–5 20–35 longer narrow O. teucrii light brown the later ones inconspicuous erect spreading mostly bidentate linear-lanceolate purple (yellow in equalling or longer apochromatic plants) apochromatic plants*) Teucrium Teucrium equalling or longer than pink to purple (yellow in lax 1–4 shorter lanceolate Rubiaceae very broad dark brown (17–)20–35 inconspicuous erect spreading mostly bidentate purple (yellow in O. caryophyllacea shorter or equalling apochromatic plants*) in apochromatic plants) equalling the lateral ones pink to deep purple (yellow (Asteraceae) lax 3–5 broad 17–24 longer yellow O. apuana light brown bent forward mostly entire inconspicuous linear-lanceolate yellow to pinkish equalling or shorter Santolina equalling the lateral ones . Diagnostic characters of the taxa considered. The measures and character states are directly taken from the type material and on studied specimens. The measures and character states are . Diagnostic characters of the taxa considered. Character/Species when dry Stem colour Spike flowers Mature Floral bract shape Floral bract in comparison with the corolla Calyx Calyx teeth in tube the with comparison colour Corolla length (mm) Corolla near Hump of the corolla the apex lobes of the Upper corolla Central lobe of the lower lip of the corolla Stigma colour Insertion of the stamens the base of from (mm) corolla Host *Character state observed in living plants. TABLE 1 TABLE

Orobanche caryophyllacea Smith (1798: 169). Type (lectotype designated by Foley 2001: 230):—ITALY. Valcimara, 29 April 1787, s.c. s.n. (LINN-HS 1087.23!).

Orobanche cyrnea Jeanmonod et al. (2005: 257). Type:—FRANCE. Secteur Rotondu, au NW de Santa-Lucia-di-Mercurio sur la D41 [42°20’N 9°13’E], fruticée basse rocailleuse, assez abondant, sur Santolina corsica, 810 m, 29 May 2000, D. Jeanmonod, A. Schlüssel et C. Habashi J6439 (holotype G 68949/1, left individual!)

Orobanche teucrii Holandre (1829: 322). Type (lectotype here designated):—FRANCE. Orobanche teucrii n., […….], para- site sur les racines de la germandrée petit chène elle croit sur la cote st. Quentin près Metz, [.….], Coll. Marchal 1828 [manu Holandre] (MPU023457!). Note:—A single specimen was found in the herbarium of the Universite´ Montpellier 2 (MPU) collected by E. Marchal and with a long description by Jean Joseph Jacques Holandre. This specimen is collected from one of the locality reported in the protologue, it includes two well-developed individuals togheter with the host plant clearly coming from the same collection and fits the original description by the author; thus we here selected it as lectotype.

Orobanche lutea Baumgarten (1816: 215). Type (lectotype here designated):—ROMANIA. Herbarium Baumgartenianum, Orobanche lutea Baumg., aliis O. major, in herbosis arenosis supra Riuta am Siechenberg, 5, June 1816, [manu Baumgarten] (CL10947, the most complete individual in the centre of the sheet!). Note:—Two specimens were found in the Herbarium of the A. Borza Botanical Garden, Babes-Bolyai University (CL) where the Herbarium of Johann Christian Gottlob Baumgar- ten is kept. They are CL10946! from Kohalom (the Hungarian variant name for the village of Rupea, Brasov) and CL10947 between Riuta the old name of the village Roandola (Sibiu) and Siechenberg, most probably a hill in the area of the village. Both specimens refer to the same taxon. We have chosen the most complete one that better fits to the original description of the species.

Orobanche latisquama (F.W.Schultz) Batt. in Battandier & Trabut (1890: 659) ≡ Boulardia latisquama Schultz (1848: 104). Type (lectotype here designated):—ALGERIA. N. 14 Notes prises sur la plante vivante [….] Boulardia latisquama / N. 14 Sur le Micromeria inodora Benth. et plus rarement sur le Rosmarinus officinalis L. jamais sur d’autres plantes. Mostaganem. Coteaux maritimes, 20 April 1844 [manu Durieu] (P02977809, the individual on the left formed by two parts: the base and the spike!). Note:—Two specimens were found in the Herbarium of the Muséum National d’Histoire Naturelle in Paris (P). In this herbarium are housed the duplicates of the herbarium of Maire which, is kept in MPU and, in turn, includes the Bat- tandier and Pomel herbaria (Domina et al. 2013). These specimens are P02977809 and P06775839! and both are coming from Mostaganem (Algeria). Both specimens refer to the same taxon. We have chosen the most complete one that better fits to the original description of the species and includes a long handwritten description.

Orobanche santolinae Loscos & Pardo (1863: 79). Type (lectotype Designated by Pujadas 2010: 960):—SPAIN. in Aragonia australi [North Aragona], on Santolina Chamaecyparisus, 1860, Loscos s.n. (COI00056915!).

Acknowledgements

This study was supported by the International Foundation pro Herbario Mediterraneum. We thank the keepers of the cited herbaria for technical support. In particular Mihai Puscas (Cluj-Napoca, Romania) and Constantin Dragulescu (Sibiu, Romania) for digital loan and help in reading the historical labels. A special thank to Maurizio Soldano (Pietrasanta, Italy) for the photos of the plant and to Melania Marchi (Marina di Carrara, Italy) for the information on her photos.

References

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