Webbia Journal of and Geography

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The genus Paeonia L. in : taxonomic survey and revision

N. G. Passalacqua & L. Bernardo

To cite this article: N. G. Passalacqua & L. Bernardo (2004) The genus Paeonia L. in Italy: taxonomic survey and revision, Webbia, 59:2, 215-268, DOI: 10.1080/00837792.2004.10670771

To link to this article: http://dx.doi.org/10.1080/00837792.2004.10670771

Published online: 14 Apr 2013.

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Download by: [Università di Pisa] Date: 05 November 2015, At: 02:39 Webbia 59(2): 215-268.2004

The genus Paeonia L. in Italy: taxonomic survey and revision

N. G. PASSALACQUA, L. BERNARDO Museo eli Storia Naturale della Calabria ed Orto Botanico, Universita della Calabria 1-87030 Arcavacata eli Rende (CS)

Ricevuto il4 Agosto 2003 Accettato il30 Settembre 2004 Il genere Paeonia L. in Italia: indagine tassonomica e revisione- Le peonie ita­ liane finora sono state distinte essenzialmente in base alia morfologia delle foglie, tuttavia la forma, le dimensioni ed il numero di segmenti fogliari possono variare anche nella medesima popolazione; cio ha generato in passato diverse incertezze ed errori. Lo stesso LINNEo (1753) nel descrivere le due varieta corrispondenti, attual­ mente, a Paeonia officina/is e Paeonia mascula non chiarisce del tutto le differenze fra le due. TI nostro studio, basato sull' osservazione sia di exsiccata in diversi erbari che di popolazioni vive nella penisola Italiana, in Sicilia, in Sardegna ed in Canton Ticino (Svizzera), accompagnato da misure morfometriche, da indagini cariologiche ed ecologiche, ci ha permesso di individuare otto entita, suddivise in due gruppi: Paeo­ nia mascula gr. e Paeonia officina/is gr. Fra i caratteri osservati, assumono particola­ re rilevanza la morfologia dell'apparato ipogeo, dei fiori, dei frutti e soprattutto del­ le foglie, nonche la qualita e la quantita della peluria. In particolare le foglie sono composte, hi- o triternate, con segmenti ultimi che possono essere interi oppure partiti e/o lobati. Si inten dono partiti quei segmenti che sono divisi per oltre la meta della loro lunghezza, lobati nel caso in cui sono divisi per non oltre la meta. Gruppo Paeonia officina/is L., presente con due entita ben distinte fra loro: Paeonia officina/is e . La primae specie temperata sia di bosco che di radura, caratterizzata da apparato sotterraneo tuberiforme e da segmenti fogliari per lo piu partiti e/o lobati, di numero variabile, mediamente fra 15 e 30. In base ai nostri studi sono individuabili quattro sottospecie lungo l'arco alpino e l'Appenni­ no settentrionale e centrale. • Paeonia officina/is L. subsp. o/ficinalis, presente sulla Catena Alpina dal Pie­ monte al Friuli Venezia Giulia (ad esdusione del Carso), sulle Alpi Apuane e sul- 1' Appennino settentrionale dalla Toscana all'Emilia Romagna; L. Downloaded by [Università di Pisa] at 02:39 05 November 2015 • Paeonia of/icinalis subsp. huthii Soldano, in poche localita della Liguria; • Paeonia o/ficinalis L. subsp. italica subsp. nov., endemica dell'Appennino centrale dalle Marche al Lazio e all' Abruzzo; • Paeonia officina/is L. subsp. banatica (Rochel) So6, attualmente rinvenuta so­ lo sui Carso Triestino. Per quanto riguarda I' altra specie del gruppo: Paeonia peregrina Mill. e specie orientale con disgiunzione d' areale in Italia me­ ridionale, con poche popolazioni ai confini fra Calabria e Basilicata. Essa e morfo­ logicamente affine a P. officinalis, se ne distingue essenzialmente per i petali forte­ mente concavi, di colore rosso cupo invece che violacei e per i segmenti terminali ultimi, caratteristicamente tridentati all' apice. Gruppo Paeonia mascula (L.) Mill.: caratterizzato da apparato sotterraneo cilin­ drico-affusolato e segmenti fogliari interi, da ovali ad ellittici, di numero inferiore a 18. In Italia e presente con tre entita di sottobosco. • Paeonia mascula subsp. mascula, rara dal Lazio alia Basilicata; 216 N.G. PASSALACQUA,L. BERNARDO

• Paeonia mascula subsp. mascula var. russoi (Biv.) comb. nov., in numerose lo­ calita della Sicilia, rara in Calabria. • Paeonia morisii Cesca, Bernardo & N.G. Passal., descritta recentemente per la Sardegna. Tutte le popolazioni indagate presentano numero cromosomico 2n=20 ad esclusione di quelle attribuite a Paeonia morisii 2n=10. Key words: Paeoniaceae, Paeonia, taxonomy, chorology, Italy, Sicily, Sardinia. The genus Paeonia is almost homogeneous in the Italian territory, showing all herbaceous individuals that die at the end of summer and grow up at the end of winter; they are all perennial with a tuberous stem and fleshy roots that provide the nourishment of the plant during the period of quiescence. The flower, gen­ erally one, spirocyclic, dialisepal and dialipetal, is hermaphrodite, with a high number of and several carpels, completely free, laying on a fleshy disk. Growth form, phenology and usually floral characters apart, most of the morphological characters of have a large range of variability, which has led to the description of numerous units not well defined and difficult to dif­ ferentiate. The numerous investigations in the field have allowed us to verify how the individuals with extreme morphological characters within the same population can look quite different, even if they are connected by a continuum of variability from an extreme to the other.

Historical taxonomic treatment The genus was established by LINNAEUS in 1753 with one , Paeonia officina/is L., and two varieties, afoeminea and 13 mascula, split on the base of different leaflet morphology. It is evident that Linnaeus went back to earlier botanists who recognized two different species, but he did not (" limites inter species non reperi") and lumped then both under the same species. The famous gardener and botanist Miller, observing cultivated , raised the var. mas­ cula to species level (MILLER, 1768). Both Linnaeus and Miller indicate the Downloaded by [Università di Pisa] at 02:39 05 November 2015 mountains of as its locus classicus. At the same time, Miller de­ scribed a coming from the Levant (i.e. ) with deep red and rounded petals as P. peregrina. ANDERSON (1818) published the first Monograph on the genus, based mainly on leaf morphology and hairiness, but in some case misleading botanists as to the identification of plants, especially as he gave no clear distribution of taxa. Earlier records of peonies in Italy come from TENORE ( 1811-15) who recorded P. o//icinalis for the Abruzzi and P. corallina (= P. mascula (L.) Mill.) for Calabria. At the same time, two new taxa close toP. mascula (L.) Mill. were described for Sicily: P. Russi Bivona (1816), in the mountains around Palermo, and P.fla­ vescens C. Pres! (1822). TilE GENUS PAEONIA L. IN ITALY 217 The first organic taxonomic treatment of Italian peonies comes from BER­ TOLONI (1844) who recorded three species and gave a good account of syno­ nyms (tab. 1) and localities. He indicated: P. officina/is L., in central northern Italy from Tuscany to Piedmont and Friuli; P. corallina Retz. (=P. mascula (L.) Mill.), in Tuscany (Florence and Volterra) and Veneto (Bassano), and P. Russi Biv., Sicily (Mountains of Palermo and Pizzuta) and Sardinia. BERTOLONI dis­ tinguished P. corallina toP. Russi by leaf morphology, but also carpel morphol­ ogy, recently used as diagnostic character for a new taxon from Sardinia (i.e. P. morisii, see below).

TABLE 1-Historical view of main reports of the genus Paeonia L. for Italy

Author taxon localities Bertoloni, 1844 P. officina/is L. P. coral/ina sensu Bert. Marches, Tuscany (Apuan P. paradoxa sensu Ten. Alps), Emilia Romagna, Piedmont, Lombardy, Trentino Veneto Friuli. P. coral/ina Retz. P. officina/is L. .B mascula Veneto, Tuscany (Florence, Volterra) P. Russi Biv. P. flavescens Presl Sicily, Sardinia Huth, 1891 P. coral/ina Retz. P. officina/is L. .B mascula a tiQica Latium, Sicily ~ fl_avescens Presl Sicily (Madonie, Nebrodi) y Russi Bivona Sicily (M. te of Palermo, M.te S. Venere Nebrodi) P. peregrina Miller P. officina/is a foeminea L. P. f!.ubens Rchb. a officina/is Retz. Trentino, Veneto, Friuli, Is tria ~ villosa P. p_aradoxa DC. Abruzzi (Maiella) Fiori & Paoletti, P. offjcinalis L. 1898 subsQ. euoffjcinalis a [oeminea L. !stria, Italian Peninsula ~ peregrina Miller with the typical var. and Downloaded by [Università di Pisa] at 02:39 05 November 2015 Qrob. more freguent y villosa Des£. P. paradoxa Anders. Maritimes Alpes, Central P. pubens Sims. Apennines and perhaps elsewhere subsQ. corollina () mascula L. P. coral/ina Retz. !stria, Veneto, Lombardy, Tuscany, Latium, Campania, Puglia, Basilicata, Calabria, Sicily, Sardinia e triternata Biv. P. coral/ina var. pubescens Calabria, Sicily, Sardinia Moris P. officina/is e P. Russi Fiori (next page) 218 N.G. PASSALACQUA, L. BERNARDO (continuer/) Author taxon synonym localities Stern, 1946 P. officina/is L. P. pubens Rchb. North Italy from Piedmont to Friuli P. humilis Retz. var. villosa P. paradoxa Anderson Abruzzi P. arietina Anderson Friuli Emilia P. mascula (L.) Miller P. coral/ina Retz. !stria, Friuli, Tuscany P. corallina var. flavescens (Florence) (Presl) Guss. P. Russi Bivona P. corallina var. pubescens Sicily (M.te of Palermo), Moris Sardinia Pignatti, 1982 P. officina/is L. subsp. officina/is Piedmont, Lombardy, Trentino Veneto Friuli subsp. villosa (Huth) P. paradoxa Anderson Maritimes Alpes, Cullen et Heywood Northern-Central Apennines P. peregrina Miller P. pubens Sims Abruzzi (Maiella) P. mascula (L.) Miller subsp. mascula P. coral/ina Retz. Lombardy, Veneto, Friuli, Tuscany, Latium, Campania, Puglia, Basilicata subsp. russii (Biv.) Calabria, Sicily, Sardinia Cullen et Heywood P. coriacea Boiss. P. coral/ina Retz. var. Sardinia leiocarpa Cosson

Several later records, even if influenced by ANDERSON's Monograph, can someway be taken back to BERTOLONI taxonomic setting, until at the end of the century when HuTH (1891) published a new monograph with a different taxo­ nomic arrangement that influenced the following Italian records for Paeonia: P. peregrina Mill. a officinalis Retz., from Trentino to !stria; P. peregrina Mill. p villosa, Maiella (Abruzzi); P. corallina Retz. a tipica, in Lazio and Sicily; P. cor­ allina Retz. Pflavescens C. Presl, in Sicily (Madonie and Nebrodi); P. corallina Retz. y Russi Bivona, in Sicily too (mountains of Palermo, Nebrodi and M. Downloaded by [Università di Pisa] at 02:39 05 November 2015 Santa Venere). Except for nomenclature and distribution, the only new taxon is P. peregrina Mill. Pvillosa, described from , South and Italy. FIORI & PAOLETTI (1898) recognised again a single species P. o/ficinalis L., and included all Italian taxa as subspecies and varieties of this: subsp. euoffici­ nalis a /oeminea (L.) Fiori & Paoletti for !stria and Italian peninsula; p pereg­ rina (Mill.) Fiori & Paoletti, with the type, perhaps more frequent; y villosa (Desf.) Fiori & Paoletti for Liguria (Marittimes Alps) and Central Apennines; subsp. corallina () mascula (L.) Fiori & Paoletti, almost everywhere (from !stria to Lombardy, from Tuscany to Calabria, Sicily and Sardinia) and e triternata (Pallas) Fiori & Paoletti (= P. Russi. Biv.) for Calabria, Sicily and Sardinia. Again, the main characters are leaflet morphology and hairiness. THE GENUS PAEONIA L. IN ITALY 219 A new global Monograph, with the ingression of a new taxon into Italian territory, was published by STERN in 1946: he recorded P. officinalis L., from Piedmont to !stria; P. mascula (L.) Mill. for Tuscany and !stria; P. russiBivona, for Sicily and Sardinia, and the var. reverchoni LeGrand, that: "closely resem­ bles P. Russi except that its carpels are glabrous", for Sardinia but on the base of a specimen quoted for P. Russi too; P. humilis Retz. var. villosa (Huth) Stem for Abruzzi, but alsoP. arietina G. Anderson for Friuli (Karst) and Emilia Ro­ magna (Apennines); this new taxon will be later recognised to be restricted to Anatolian peninsula. STERN took into account the work of STAPF (1918) who gave a full history of the name P. peregrina Mill. limiting it to the Balkan plant. The first edition of Flora Europaea (CuLLEN & HEYWOOD, 1964a) was not clear with the distribution of taxa, recording for Italy P. peregrina Mill. (Cen­ tral Italy); P. officinalis L. subsp. officinalis (from France to and Al­ bany) and subsp. villosa (Huth) Cullen & Heywood (from South France to Central Italy); P. mascula (L.) Mill. subsp. mascula (supposed) and subsp. russii (Biv.) Cullen & Heywood (Islands of W. Mediterranean) and P. coriacea Boiss. (possibly Sardinia); CULLEN & HEYWOOD mainly again used as diagnostic char­ acters leaflet morphology and hairiness. Flora d'Italia (PIGNA'ITI, 1982) maintains the same taxonomic setting but with a better definition of distributions (tab. 1) and, following on, the new edi­ tion of Flora Europaea (AKEROYD, 1993) and Med-Checklt'st (GREUTER et al., 1989) maintained the same taxonomic arrangement of the genus in Italy. ScHMITT (1997, 1998, 1999) published a series of contributions that con­ tain a great deal on the knowledges of taxonomy of the whole genus Paeonia, even if he did not modify very much what was known for Italy, except that he accepts observations from SoLDANO (1993) on L. subsp. vil­ losa (Huth) Cullen & Heywood, proposing the new name P. officinalis subsp. huthii Soldano. Recently, PoLDINI et al. (2001, 2002) reported Paeonia officinalis L. subsp.

Downloaded by [Università di Pisa] at 02:39 05 November 2015 banatica (Rachel) So6 for Friuli (North-East Italy), which should be the West­ em limit of distribution for the taxon, and CESCA et al. (2001) described a new peony from Sardinia, P. morisii Cesca, Bernardo & N.G. Passal. formerly in­ cluded in P. mascula subsp. russoi. The present arrangement of Paeonia in Italy is as follows: Paeonia L. Paeonia officinalis L. subsp. officinalis subsp. huthii Soldano subsp. banatica (Rochel) So6 Paeonia peregrina Mill. 220 N.G. PASSALACQUA, L. BERNARDO Paeonia mascula (L.) Mill. subsp. mascula subsp. russoi (Biv.) Cullen & Heywood P. coriacea Boiss. P. morisii Cesca, Bernardo & N.G. Passal.

MATERIALS AND METHODS

The study is based on specimens from herbaria, living populations and cultivated plants. We believed important to study samples and populations also from other countries (especially Swit­ zerland, and ), to resolve some of the problems inherent to the definition of taxa. The herbaria consulted were: AQUI, BM, BOLO, CAG, CAT, CLU, FI, PAD, PAL, PESA, PI, RO, ROV, SO, TO, TSB, UPA, VER, and they have been important to point out taxonomic problems; other herbaria have been contacted (LINN, BE, DR, OXF, SBT, PH, MW, KIEL, GOET, P) to have information about type and/or original material. The populations investigated have been selected after herbaria and bibliography consultation indicated the ones which showed the best variability and/or appeared crucial to solve taxonomic questions.

Living populations: Italy Sardinia: Bosco lungo la strada nel vallone Rio Aratu (Fonni, Nuoro, Sardegna), Bosco di leccio e roverella, 1100 m; Serra Edele-Bosco di Ghivine (Cala Gonone, Nuoro, Sardegna), lecceta aperta, in radure, 540 m, su calcare; Monte Bruncu Spina (Gennargentu, Nuoro, Sardegna), piccolo bosco ripario ad Alnus glutinosa, 1600 m, su calcare; Monte Albo (Lula, Nuoro, Sar­ degna), bosco di leccio, 1000 m ca., su calcare; Monte D' lscudu (Gennargentu, Nuoro, Sar­ degna), gariga a Genista corsica e Rosa sera/inii,1300 m, su calcare. Sicily: Ficuzza, presso 1' Alpe di Cucco, 980 m; Nebrodi, Bosco del Flascio, C.da Serra di Cara­ cozzo, bosco a Quercus congesta, 970 m; Etna, Bosco Maletto aile falde del monte, bosco a leccio e Quercus congesta, 1100 m; Nebrodi, Bosco di Grappida, bosco a cerro, 1300 m; Iblei, valle Cava Grande, bosco a leccio, 550 m; Madonie, Piano Battaglia, bosco di faggio in radure, 1500 m. Latium: Alta Vaile Rio Fuggio, Leonessa, Rl, 1240 m.; Bosco Riguardata-Molara al C. dei Guardi­ ani (Rocca di Papa, Lazio, ltalia), bosco di castagno, 500 m s.l.m. Calabria: Colle Marcione, Civita, 900 s.l.m.; Monte Camara, San Paolo Albanese (CS); Piano di Marco, ai piedi della Mula, S.Donato di Ninea (CS) 1220 m; Valle Olivella, Casiglia, sotto Downloaded by [Università di Pisa] at 02:39 05 November 2015 Monte La Mula (S.Sosti, CS); Monte Sellaro, presso il Bifurto, Cerchiara di Calabria, CS, 950 m; Monte Basilica, Bocchigliero, Sila Greca, CS, 900 m. Apulia: Gargano, nella foresta Umbra lungo la strada per Vieste, bosco di cerro. Basilicata: Potenza, Bosco Pallareta, vicino monte Grosso, bosco di cerro, 1150 m Greece: Makrolivado (prov Phthiotis, Grecia) 900s.l.m.; Karia (M. Stavroras, lsole Lefkas, Gre­ cia) 900 s.l.m., gariga sassosa. Abruzzi: *Prati di Tivo, Gran Sasso d'ltalia, 1450 m s.l.m.;Valle dell'Orfento, presso Sfischia, Maiella; Valle del Vasto, Gran Sasso d'ltalia. Tuscany: *Base del Monte Pisanino vicino Orto di Donna, Alpi Apuane Liguria: *Vers. orientale di Monte Toraggio, Bordighera, IM, 1750 m Piedmont: *Val di Lanzo, lungo la strada Balme e Pian di Mussa,TO, 1500 m Lombardy: Monte Maddalena sopra Botticino, BS; Monte Barro, Galbiate, CO, 920 m Trentino: Terra Rossa, vers. merid. del Monte Finonchio sopra Scottini-Pomal, Trento; Versante Sud di Coma Piana, Monte Baldo, Trento, 1400 m, faggeta e rupi; *Monte Coma Piana, Monte Baldo, Trento, 1600-1700 m, pascoli d'altitudine. THE GENUS PAEONIA L. IN ITALY 221 Friuli: *Sagrado di Sgonico, strada forestale verso Monte Lanaro, Monrupino, TS; Gropada, Carso Triestino, TS. Switzerland: *Monte Generoso in localita Pianche (Canton Ticino); 1130 m s.l.m. The field investigation of seven populations of P. officina/is gr. (marked with*) was accompa­ nied by morphometric measurements (characters in tab. 2) on twenty individuals for each popu­ lation, that was submitted to statistical and multivariate analysis with Data Desk 6.1 software. Some plants from each population investigated were cultivated in the Botanical Garden of the University of Calabria. On these plants the following were carried out: - micromorphological analysis, by observation of the lower leaf surface; - caryological analysis on mitotic metaphase plates of meristematic cells taken from root tips; - morphological and phenological observations on different stages of development.

P. officinalis and P. mascula: general morphology and other evidence

Morphology Rootstock To discriminate between the two main groups of Italian peonies is by the different morphologies of the rootstocks which, although known in the past and shown in the drawings of MATTIOLI (1565, fig. 1) and others (LoBEL, 1581, Downloaded by [Università di Pisa] at 02:39 05 November 2015

Fig. 1-Woodcut of (a) P./oemina (=P. officina/is L.) and (b) P. mas (=P. mascula (L.) Mill.) from MAT­ nou (1565). 222 N.G. PASSALACQUA,L. BERNARDO

fig. 2; FucHs, 1542), were ignored by many later botanists (STEARN & DAVIS, 1984). LINNAEUS (1753), who examined leaf morphology only, expressed doubt on the discrimination of the two groups in Paeonia, now called P. mas­ cula and P. officina/is. The subterranean persistent part consists of a stem and roots merged in a rootstock, elongated and tapering as in the Paeonia mascula gr. (fig. 3b; including P. morisii), or fusiform to subglobose, as in the Paeonia officina/is gr. (fig. 3a; including P. peregrina), where slender attachments like strings bringing short and thick roots and hanging fusiform tubers grow from the rootstock.

Stem From buds on the rootstock develop one or more aerial annual stems, which are herbaceus, 30-90 em tall, simple and erect-flexuosus, fleshy, cylin- Downloaded by [Università di Pisa] at 02:39 05 November 2015

Fig. 2-Woodcut of (a) P. foemina (=P. officina/is L.) and (a) P. mas L. (=P. mascula (L.) Mill.) from Lo­ BEL (1581). THE GENUS PAEONIA L. IN ITALY 223 Downloaded by [Università di Pisa] at 02:39 05 November 2015

Fig. 3-Different rootstock morphology in P. officina/is gr. (a) and P. mascula gr. (b). (from STEARN & DAVIS, 1984) 224 N.G. PASSALACQUA, L. BERNARDO dric with subangled grooves; they can be glabrous, subglabrous or hirsute be­ low the flower. The height and the dimensions of the whole plant varies even in the same population, but generally in the P. mascula group the samples are on the average more reduced than in P. officinalis group. The colour changes from pale green to purplish red: generally the P. offici­ nalis group, exhibits only slight red veins on the green stem, while in the P. mascula group the stem is very often reddish. However, in the Italian peonies, at the beginning of the growth, the stem, the petiole and the rachis of leaves are purplish-red, but when mature the plant have lost this colour, with exception of plants from Sardinian that keep stems and petioles purplish-red, while in Si­ cilian peonies the stem is reddish coloured only at the base at maturity. In any case, the strength of reddening on the plants could be due to the light intensity of the place where the plants grow up. At the base of the stem there are some sheaths, alternate, obtuse and ob­ long, from green with a reddish edge to completely reddish.

Leaf

In the past the shape of leaves has been of a great importance in the tax­ onomy of the genus Paeonia, even though this is very variable, even in the same population. The leaves are alternate, without stipules and range in number from 3 to 8. The P. mascula group had generally few leaves (less then 5) while in the P. officinalis group there may be up to 10. The leaves are stalked and always compound, biternate as seen in fig 4. From the common petiole 3 primary divisions arise with petiolules, then every division is divided into 3 primary segments or leaflets. The central pri­ mary segment is always petiolulate, the lateral ones may be sessile or with Downloaded by [Università di Pisa] at 02:39 05 November 2015 petiolules. According to the taxon, the primary segments may be entire or di­ vided in secondary segments, once again entire or divided into leaflets or eventually two- or three-lobed. Thus the number of ultimate segments and lobes may range from few (7-9) to many (30-40). Also the shape of the ulti­ mate leaflets is very variable: it may be orbicular and broadly obovate or nar­ rowly lanceolate, acuminate. In general, the P. officinalis group has mainly partite segments, linear lanceolate to elliptic, compared with the P. mascula group where the segments are mainly entire and orbicular to elliptic. An­ other useful character for separating the taxa is the hair-covering, when present, by a minute, often scattered pubescence, which can be spread only on the lower leaf surface. THE GENUS PAEONIA L. IN ITALY 225

Fig. 4-Leaf segmentation diagram in P. officina/is gr. (continuous line) and P. mascula gr. (dotted line). (from STEARN & DAVIS, 1984, modified).

Flower

Although the flower in Paeonia is very remarkable, it has no great impor­ tance for the discrimination of the Italian taxa. With the exception of Paeonia

Downloaded by [Università di Pisa] at 02:39 05 November 2015 peregrina Mill., showing a cup-shaped flower with petals strongly concave and red coloured, nearly all the Italian variants have flowers opened wide with slightly concave petals, the colour varying from purplish-red (P. officina/is group) to pinkish-mauve (P. mascula group). Furthermore, in Sicilian peonies the flower may be completely white or white striped, but, in our opinion, it is not a significant taxonomic character. The calyx is enveloping (imbricate), covering the petals in flower, reflected or falling at fructification. It shows a change in shape from the outside inwards: lower sepals (1-3) are elongated leaflet -like, the inner ones (3-5) are broadly el­ liptical, entire, concave and rounded at the base; green with membranous and purplish edge, glabrous or tomentose on the outer surface. The stamens are 226 N.G. PASSALACQUA, L. BERNARDO very numerous, with white or pink filaments and yellow anthers dehiscing mar­ ginally. The pollen grains are yellow, spheroid prolate, 3-colporate, with the colpi narrow and extending to the poles, the exine is finely reticulate. The carpels range from 1 to 8, they are erect, free and surrounded at the base by a depressed fleshy disc, tomentose and with a very short style. The stigma is broadly recurving, glabrous and red coloured.

Fruit

After fertilisation the carpels develop into follicles, always tomentose in the Italian peony taxa. They are oblong, mosdy abrupdy narrowed to an obtuse apex or tapering and gradually attenuate in their upper part, like in peonies from Sardinia. When mature the follicles may be suberect or outspread or re­ flexed and fully curved backwards, opening along a ventral suture and showing several anatropous seeds in two rows. Seeds are broadly ellipsoid, 7-9 mm long, smooth, blackish or red, probably sterile in the latter.

Caryology and molecular evidences

Italian peonies show a basic chromosome number x=5, which is expressed both with diploid populations, 2n=10, and tetraploid ones, 2n=20. All popula­ tions studied of the P officina/is group are tetraploid, 2n=20, (Peruzzi, pers. com.). Also reported in the literature is 2n=20 for the taxa of this group (BAR­ BER, 1941; Gregory, 1941; STERN, 1944; CuLLEN & HEYWOOD, 1964a; FE­ DOROV, 1969; TZANOUDAKIS, 1983; SCHWARZACHER-ROBINSON, 1986; ELENA RosSELLO et al., 1987; BERNARDO et al., 1995; KOEVA & SARCOVA, 1997), with Downloaded by [Università di Pisa] at 02:39 05 November 2015 the exception of SoPOVA (1971) who found the diploid level for P. peregrina Mill. from Macedonia. As already indicated by CESCA et al. (2001) for P. mascula group many data in the literature are difficult to interpretate because they often appear in con­ tradiction, giving both 2n=10 (SOPOVA, 1971; TZANOUDAKIS, 1983; KOEVA & SARCOVA, 1997) and 2n=20 (BARBER, 1941; STERN, 1944; CULLEN & HEY­ wooD, 1964a; TzANOUDAKIS, 1983 ). The results of the most recent caryological investigation on the P. mascula group in Italy always show the Sardinian popu­ lations to be diploid (CESCA et al., 2001), those from the Italian Peninsula tetra­ ploid (BERNARDO et al., 1995, Peruzzi, pers. com.) and the tetraploid level also in all Sicilian peonies, as in the literature (RAIMoNDO et al., 1983). THE GENUS PAEONIA L. IN ITALY 227 Phylogenetic studies, based on ITS sequences (SANG et al., 1995) provide evidence on the separate evolution between P. mascula s.l. and P. officinalis s.l., but the former were supposed to be of hybrid origin. Later (SANG et al., 1997), comparing ITS and matk sequences, showed the two groups keep their separa­ tion, even if additional hybridization events are hypothesized.

Distribution and ecology

It is difficult to outline a general distribution of the two groups of Paeonia, mainly because there are numerous taxa that are associated with them forming aggregates and recent Floras use leaf morphology as the main diagnostic char­ acter. P. officinalis L. is a South European species (CuLLEN & HEYWOOD, 1964b), with four subspecies that form a geographic series: subsp. microcarpa Nyman (= subsp. humilis (Retz.) Cullen & Heywood) from to Southern France; subsp. huthii Soldano (= subsp. villosa (Huth) Cullen & Heywood) from Southern France to Central Italy; subsp. officinalis from South East France to and subsp. banatica (Rochel) So6 in Hungary, and the former Jugoslavia (in the Banat region). P. peregrina Mill. is a Balkan-Tura­ nian species distributed from Balkan peninsula to Moldavia and Turkey and with one station in Italy (STEARN & DAVIS, 1984; AKEROYD, 1993; BERNARDO et al., 1995). P. mascula subsp. mascula is a Southern European- Western Asiatic element, spreading from to the Caucasus and Northern and Iran (STEARN & DAVIS, 1984); subsp. russoi (Biv.) Cullen & Heywood a Mediterranean element formerly indicated for Ionian islands of Greece, Sicily, Sardinia and Corsica (STEARN & DAVIS, 1984) but now restricted by the de­ scription of P. morisii Cesca, Bernardo & N.G. Passal. that includes its popu­

Downloaded by [Università di Pisa] at 02:39 05 November 2015 lations from Sardinia and Corsica. We have found that the two groups have distributions in Italy which are substantially separate, the P. mascula group in Sardinia, Sicily and Southern­ central part of Italian Peninsula, while the P. officinalis group is in the Alps, Northern-central Apennines and, very localized, in Northern Calabria. This distribution is slightly different from that given by PIGNATTI (1982), mainly be­ cause from our observations (fig. 5) P. mascula does not appear to be present in Emilia, Veneto, Tuscany and Friuli, and moreover, we have found populations of the P. o/ficinalis group in Southern Italy. The two groups show a different ecology, the P. mascula group is almost ex­ clusively in submediterranean and mediterranean woods on deep soil, whereas the P. o/ficinalis group is mainly on mountains in open consolidated scree 228 N.G. PASSALACQUA, L. BERNARDO Downloaded by [Università di Pisa] at 02:39 05 November 2015

Fig. 5-Italian distribution of P. officina/is gr. (dull gray) and P. mascula gr. (light gray). THE GENUS PAEONIA L. IN ITALY 229 slopes, with the exception of Southern Italian and Trieste Karst populations, that live in submediterranean deciduous oak woods. There is a clear phytogeographical distinction between the two groups, the P. mascula group with a Mediterranean character and the P. officinalis group with a European one, in the sense that the first group lives in environments climatically and chorologically characterized by a Mediterranean element, while the second, with certain exceptions, lives in vegetation characterized by European elements.

TAXONOMIC ANALYSIS AND RESULTS Following HALDA (1998), all Italian peonies can be referred to subgenus Paeonia (= sect. Paeon Stern, 1946), section Paeonia characterised by: stems herbaceous, annual, dying down at the end of each growing season. Roots suc­ culent not woody, tapering, fusiform to subglobose. Leaves thin or coriaceous with smooth entire margin, with less then 40 segments, each more then 5 mm wide in the lower leaves. Flowers more or less erect not yellow or cremy, petals thin, much larger than sepals, staminoid disc imperceptible, not lobed at the base of carpels. Seeds cylindric or ovate. HALDA identifies two subsections: subsect. Paeonia with fusiform to subglobose roots, carpels not narrowed and leaf segments dentate, corresponding to what was previusly called the P. offici­ nalis group; and subsect. Foliolatae Stern 1946 (= Masculae (Stern ex Us­ pensk.) Halda 1998, name established with the same type species and then su­ perfluous), with tapering roots, corresponding to the P. mascula group.

Subsection Paeonia The peonies of this group are characterized by pyriform, fusiform or sub­ globose roots, carpels not narrowed, leaf segments dentate, and in Italy they generally also show:

Downloaded by [Università di Pisa] at 02:39 05 November 2015 - a green stem, which is generally more elevated then in P. mascula group; - leaves in greater numbers, bi-triternate and with a greater number of seg- ments (13-55), that are linear-lanceolate to elliptic; - a smaller number of follicles (2-4) that lengthen at maturity and bend downwards; - a European group that lives on consolidated mountain scres, with the ex­ ception of the southern populations and those from Trieste Karst, that live in the woods of deciduous oaks. - · The most recent Floras give P. officinalis L. subsp. of/icinalis for Northern Italy (from the Piedmont to the Friuli Venezia Giulia), subsp. huthii Soldano (Maritime Alps and the Central-Northern Apennines), subsp. banatica (Ro­ che!) So6 (Friuli) and P. peregrina Mill. (Abruzzi, Maiella Massif). 230 N.G. PASSALACQUA, L. BERNARDO Our observations have demonstrated that peonies from Southern Italy are clearly separate from the rest, above all by their corolla that is dark red and in a closed cup, while in the other populations it varies from pink to red and to vio­ let and it is always in an open cup at maturity. Comparing this material with fresh and dried samples from Eastern Europe (Greece and Bulgaria), we have been able to verify that it is P. peregrina Mill., as already intuitively proposed by STEARN & DAVIS (1984) and BERNARDO et al. (1995). Added to this, the differ­ ent corolla, P. peregrina Mill. is also well differentiated by the typical leaf seg­ mentation, showing the last central segments with two or three apical lobes that appear as big teeth. All other populations, included those from Abruzzi, fall in P. officina/is L. s.l. that exhibits a series of morphotypes with separate distributions, intuitively recognizable in the field, but that does not show clear diagnostic characters. The chromosome counts have not given any evidence to separate the differ­ ent populations of the P. officina/is group, since they are always tetraploid, and as already noted, this agrees with the greatest part of the known data in the lit­ erature. Further studies are planned for a more detailed karyotype analysis. A biometric investigation was undertaken (tab. 2) on the following popula­ tions to point out possible groups of homogeneity (see also "MATERIALS AND METHODs"): Gran Sasso (Abruzzi), for the morphotype of the Central Apen­ nines; M. Pisanino (Tuscany), for the Apuane Alps and Northern Apennines; M. T oraggio (Liguria), for the Ligurian and Maritime Alps; Valleys of Lanzo (Piedmont), for the Piedmont Alps and pre-Alps; M. Generoso (Canton Ti-

TABLE 2 - Morphological characters used for biometric investigation. -characters 2.5 and 3.5 (hair covering) are classed in five categories of coverage.

1 Habitus 1. stem hight, 2. stem number, 3.leaf number 2 Lower leaf l.lenght Downloaded by [Università di Pisa] at 02:39 05 November 2015 2. segments number (2.1 entire, 2.2 partite, 2.3lobed) 3. biggest segment (3 .1lenght, 3 .2 width, 3 .3 form, 3 .4 area) 4. smaller segment (4.1lenght, 4.2 width, 4.3 form, 4.4 area) 5. hairs cover on lower surface (5.1lamina, 5.2 rib, 5.3 petiolus) 3 Upper leaf l.lenght 2. segments number (2.1 entire, 2.2 partite, 2.3 lobed) 3. biggest segment (3.1lenght, 3.2 width, 3.3 form, 3.4 area) 4. smaller segment (4.1lenght, 4.2 width, 4.3 form, 4.4 area) 5. hairs cover on lower surface (5.1lamina, 5.2 rib, 5.3 petiolus) 4 Bract l.lenght 2. width 3. segments number (2.1 entire, 2.2 partite, 2.3 lobed) 5 Sepal 1. number 6 Fruit 1. number THE GENUS PAEONIA L. IN ITALY 231 cino, Switzerland), for Switzerland and the Lombardy Alps; M. Baldo (Tren- tina), for the Trentino, Venetian and Friulian Alps; Sagrado di Sgonico (Carst) for the Karst of Trieste. To have an overview of similarity among all populations, we computed a Principal Components Analysis of 30 characters (tab. 3, fig. 6) that showed 57,5 % of variance on the first three axes.

TABLE 3-Report of Principal Component Analisys in P. officinalis s.l. group (see also tab. 2)

Principal Component Analysis Based on Covariance 140 total cases of which 8 are missing EigenValues 95% Interval Variance Values Lower UQQer ProQortion e1 15922,32 12818,06 21010,65 35,5 e2 5778,05 4651,54 7624,55 12,9 e3 4065,98 3273 27 5365 36 91 Eigen Vectors Unrotated Factor Matrix Character V1 V2 V3 F1 F2 F3 1.1 0,146 0,206 0,181 0,450 0,384 0,283 1.3 -0,089 0,135 0,051 -0,298 0,272 0,086 2.1 0,184 0,144 0,101 0,583 0,275 0,163 2.2 -0,254 -0,046 0,012 -0,800 -0,087 0,020 2.2.1 0,086 0,117 -0,002 0,281 0,231 -0,003 2.2.2 -0,242 -0,046 0,013 -0,755 -0,087 0,020 2.2.3 -0,093 -0,089 -0,061 -0,315 -0,182 -0,104 2.3.1 0,206 0,230 0,162 0,655 0,440 0,261 2.3.2 0,288 0,056 -0,105 0,908 0,105 -0,167 2.3.3 -0,198 0,106 0,375 -0,623 0,202 0,598 2.4.1 0,242 0,174 0,061 0,766 0,332 0,098 2.4.2 0,277 0,068 -0,113 0,872 0,129 -0,180 2.4.3 -0,116 0,174 0,338 -0,367 0,330 0,540 2.5.1 -0,211 0,263 -0,171 -0,710 0,534 -0,290 2.5.2 -0,215 0,271 -0,136 -0,721 0,546 -0,230 2.5.3 -0,058 0,251 -0,275 -0,194 0,506 -0,464 Downloaded by [Università di Pisa] at 02:39 05 November 2015 3.1 0,162 0,011 0,192 0,504 0,020 0,302 3.2 -0,113 -0,220 0,074 -0,376 -0,439 0,123 3.2.1 -0,007 0,206 0,029 -0,023 0,413 0,049 3.2.2 -0,076 -0,271 0,084 -0,252 -0,542 0,140 3.2.3 -0,048 -0,054 -0,092 -0,266 -0,178 -0,256 3.3.1 0,182 0,133 0,216 0,583 0,257 0,350 3.3.2 0,251 0,060 -0,141 0,788 0,114 -0,224 3.3.3 -0,159 0,061 0,416 -0,497 0,115 0,657 3.4.1 0,206 0,234 0,043 0,668 0,457 0,071 3.4.2 0,235 0,073 -0,151 0,752 0,141 -0,244 3.4.3 -0,068 0,245 0,325 -0,219 0,475 0,528 3.5.1 -0,217 0,269 -0,133 -0,727 0,543 -0,225 3.5.2 -0,214 0,279 -0,134 -0,717 0,564 -0,226 3.5.3 -0,163 0,302 -0,234 -0,539 0,601 -0,390 232 N.G. PASSALACQUA, L. BERNARDO Downloaded by [Università di Pisa] at 02:39 05 November 2015

Fig. 6-Principal Component Analysis based on 30 morphometric characters (see also tab. 3) in P. offi­ cina/is s.l.: - (a) first (Ul) and second (U2) axes of regression - (b) first (Ul) and second (U3) axes of regression THE GENUS PAEONIA L. IN ITALY 233 On the first two axes of regression (fig. 6a), representing 35,3 and 11,9% of variance, three groups of individuals are coarsely separate: group A composed by populations from Tuscany and Piedmont and individuals from M. Gen­ eroso and Trentino; group B composed by population from Friuli and again in­ dividuals from M. Generoso and Trentino; group C composed by populations from Liguria and the Abruzzi. On the third axis of regression (fig. 6b), representing 9,1 % of variance, the groups A and B widely merge in a cloud of individuals, while group C split in two, separating the Ligurian plants (Cl) from those of the Abruzzi (C2) into two clearly identified sub groups. PCA indicates that plants from Liguria, Abruzzi, Friuli and Piedmont-Tus­ cany are identified as homogeneous morphotype in their populations, while populations from M. Generoso and Trentino have a wider variability and indi­ viduals scatter to all directions overlapping and connecting to the variability of other populations. In some cases, populations which seem to be morphologically separate enough in the field are very difficult to separate when considering the wide populations, because they are connected by individuals that fall in the overall range of variability. There is a continuum of morphological variability that goes from one extreme to the other taking in single populations. The PCA points out as principal diagnostic characters the hairiness, the el­ evated leaf segmentation and the dimensions of the segments, on the first axe of regression, while the second and third ones point out the type and the form of the segments; other indicative characters are the length of the stem and leaves and the number ofleaves. The hairiness on the lower surface of the leaves has been used in the past to distinguish P. o/ficinalis L. subsp. huthii Soldano(= subsp. villosa (Huth) Cul­ len & Heywood), where different authors have at time included the popula­ tions from the Northern Apennines and sometime also those from Central Ap­ Downloaded by [Università di Pisa] at 02:39 05 November 2015 ennines. In effect, this is a character that seems to be particularly meaningful: peonies from the Abruzzi and Liguria have long hair, widened to the base, in great quantity on the lower leaf surface, on the ribs and on the stem (fig. 7a-b), while the other peonies have short hairs, cylindrical, varying in quantity (the populations from Friuli are subglabrous, while those from Tuscany are hairy), but never dense as above (fig. 7c-f). Correlated with the hairiness, another meaningful diagnostic character is the segmentation of the leaves. The peonies from Liguria and the Abruzzi are characterized by a general greater splitting up of the lamina, with a more el­ evated number of ultimate leaflets (tab. 4), whereas the peonies from Liguria tend to have a higher number of lobed segments;. The range of variability of 234 N.G. PASSALACQUA, L. BERNARDO Downloaded by [Università di Pisa] at 02:39 05 November 2015

Fig. 7 - Different kind of hairs of the lower leaf surface in different populations of P. officina/is s.l.: (a) Liguria; (b) Abruzzi; (c) Tuscany; (d) C. Ticino; (e) Trentino; (f) Trieste Karst

this character range from Liguria to Abruzzi and then to Trentino and Canton Ticino, slightly overlapping in the lower values. The dimension of the larger segment of the lower leaf (fig. 8, x-axes) does not show differentiation between the populations, except for the population from Liguria that tends to have very small segments, compared to those from Piedmont and Tuscany that show individuals with the largest segments. It is THE GENUS PAEONIA L. IN ITALY 235

TABLE 4 - Number of segments of lower leaf in P. officina lis s.l. f.t: mean; M: maximum value; P90: percetile 90; PlO: percentile 10; m: minimum value.

Total Entire Partite Lobed

f.L M P90 P10 mf.L M P90 PlO m f.L M P90 PlO mf.L M P90 PlO m Abruzzi 38,1 66,0 50,3 26,8 22,0 1,3 8,0 4,2 0,0 0,0 34,9 58,0 45,2 22,9 21,0 1,9 8,0 7,0 0,0 0,0 Liguria 56,2 95,0 84,8 37,7 33,0 1,9 14,0 4,9 0,0 0,0 40,9 65,0 57,6 25,8 21,0 13,3 41,0 26,4 0,0 0,0 Trentino 26,5 55,0 33,3 17,9 16,0 4,9 13,0 10,2 1,0 0,0 15,8 35,0 22,5 6,6 3,0 5,8 19,0 12,0 0,0 0,0 C. Ticino 24,1 40,0 32,7 14,9 11,0 2,0 7,0 6,1 0,0 0,0 19,2 36,0 26,3 7,9 4,0 2,9 23,0 6,3 0,0 0,0 Piedmont 20,0 35,0 27,5 14,0 12,0 4,7 9,0 7,2 0,9 0,0 13,2 27,0 20,6 7,9 6,0 2,2 11,0 8,1 0,0 0,0 Tuscany 19,3 33,0 27,2 13,9 12,0 3,4 7,0 6,0 1,8 0,0 13,3 22,0 18,1 8,0 6,0 2,6 18,0 8,7 0,0 0,0 Carst 22,9 36,0 31,0 14,9 7,0 0,9 4,0 3,1 0,0 0,0 19,8 31,0 29,1 12,5 5,0 2,2 8,0 6,0 0,0 0,0 Downloaded by [Università di Pisa] at 02:39 05 November 2015

Fig. 8-Form (= lenght /widht; x-axes) and size (= lenght x widht; y-axes) variability of the biggest seg­ ment in the lower leaf in P. officinalis s.l. 236 N.G. PASSALACQUA, L. BERNARDO possible to see how populations from Tuscany cover nearly all the range of variability of this character. The form of the segments (fig. 8, y-axes) shows how the plants from the Abruzzi have segments from lanceolate to lineare-lanceolate (length/width 4-8.5). They are fairly distinguished from all other populations in that they have segments from elliptic to lanceolate (length /width (1-) 2-4), in a fairly restricted range that overlaps the range of the Abruzzi only for extreme indi­ viduals. The segment regression analysis (tab. 5, fig. 9-10) shows, with few excep­ tions, that the various populations have the same scheme of development both in the lower leaf (fig. 9) and, also along the stem, from the lower to the upper leaf (fig. 10). The individuals broadly overlap excepting some plants from Piedmont, that have the tendency for the dimension of the segments notably reduce from the lower leaf to the upper one (fig. lOa), and in part in those from the Abruzzi, that show a greater regression in the transition from the largest segment to the smallest in the lower leaf (fig. 9). For the plants from Piedmont, we can hypothesize that the most greatest reduction is due to the larger size of the plants and, therefore, to the fact that there is a greater distance between the lower and the upper leaves (fig. lOb). For the plants from the Abruzzi we be­ lieve that there is a real difference in the plan of the development of the leaf.

TABLE 5-Segment regression values in P. officina/is s.l. (l: mean; P90: percetile 90; PlO: percentile 10. - values are calculated as follow: l.lower leaf regression: biggest segment lenght the lower leaf (2.3.1) smallest segment lenght the lower leaf (2.4.1) 2.lower to upper leaf regression: biggest segment lenght in the lower leaf (2.3.1) biggest segment lenght in the upper leaf (3.3.1) 3.lower to upper weighted: lower to UQQer leaf regression x 100 stem hight Downloaded by [Università di Pisa] at 02:39 05 November 2015 Lower to upper Lower leaf regression Lower to upper weighted leaf regression fl PlO P90 fl PlO P90 fl PlO P90 Abruzzi 3,92 2,55 6 1,14 0,78 1,53 2,48 1,70 3,19 Liguria 2,55 1,89 3,09 1,05 0,68 1,45 2,71 1,61 3,99 Trentino 1,99 1,29 2,71 1,24 0,96 1,53 2,83 2,03 3,86 C. Ticino 2,04 1,5 2,65 1,25 0,92 1,71 2,36 1,87 3,05 Piedmont 2,12 1,51 2,73 1,5 1,13 1,9 2,09 1,48 2,83 Tuscany 1,95 1,49 2,45 1,17 0,94 1,41 2,66 1,69 3,41 Carst 2,31 1,47 3 1,1 0,76 1,46 2,22 1,24 3,18 THE GENUS PAEONIA L. IN ITALY 237

Fig. 9-Segment regression variability of lower leaf in P. officina/is s.l. (explanations under tab. 5)

DISCUSSION

On our observations, the populations of this group, with the exclusion of those from Southern Italy already assigned toP. peregrina Mill., can be only as­ signed to a single species: P. o/ficinalis L. It is interesting to note that this tax­ onomy is in agreement with BERTOLONI's opinion way back in 1840 (tab. 1). The variability found among the different populations corresponds to best subspecific rank. The type subspecies of/icinalis includes the populations from the Alps, from Piedmont to Friuli (except Trieste Karst), and it includes the plants from the

Downloaded by [Università di Pisa] at 02:39 05 November 2015 Northern Apennines, all characterized by the lower leaf having on average a smaller number of segments (13-30), of which (35-) 25-8 (-3) are partite and (0-) 2-7 (-13) entire; the segments are greater (the largest is 140-85 x 65-18 mm) and generally lanceolate to elliptic, with short and cylindrical hairs varying in quantity from subglabrous to hispid on the lower surface. Subspecies o/ficinalis lives preferentially on consolidated mountain screes. The name P. officina lis L. has been recently investigated by ScHMITT (2003) who designated a lectotype (n° 692.1, LINN). He pointed out that this name has been attributed to the plant growing on M. Generoso (Switzerland), where he also stated that the epitype of var.foeminea must be a taxonomic synonym of the type variety, var. of/icinalis. We have compared the lectotype with differ- 238 N.G. PASSALACQUA, L. BERNARDO Downloaded by [Università di Pisa] at 02:39 05 November 2015

Fig. 10-Segment regression variability in P. officina/is s.l. (explications under tab. 5): (a) lower to upper leaf regression; (b) lower to upper leaf regression weighted with stem height; the outlined central box depicts the middle half of the data extending from the low hinge (roughly 25% of points) to the high hinge (roughly 75% of points); the horizontal bar is at the medium; the whiskers depict the extent of the main body of the data and extend from the box to the highest data value not above: high hinge+ 1.5(high hinge-low hinge) and from the box to the lower data value not below: low hinge- 1.5(high hinge-low hinge); extreme data value, beyond these limits, are plotted with a circle. THE GENUS PAEONIA L. IN ITALY 239 ent populations of P. officina/is and confirm this view that the lectotype corre­ sponds well with the plant from M. Generoso. Considering that a lot of past Floras and revisions are founded mainly on the leaf characters for the recognition of peony taxa, we think that very prob­ ably the report of P. mascula (L.) Mill. for Northern Apennines has been a misi­ dentification of individuals with few, large, elliptic segments. The plants from Liguria fall into the variability of P. o//icinalis L. subsp. huthii Soldano, characterized by a villose hairiness on the lower surface of the leaves, composed by long pungent hairs widened to the base and flattened, and by a great number of leaf segments (22-59) of small dimensions (80-36 x 25-10 mm). This subspecies lives on consolidated mountain screes. This taxon was described by HuTH (1891) from Southern France asP. per­ egrina var. villosa and later transferred in the new combination P. officina/is subsp. villosa by CuLLEN & HEYWOOD (1964b), who noted that even if the same combination was proposed earlier by CIFERRI & GIACOMINI (1954), as it was without reference to the place of publication of the basionym it was thereby invalid under the International Code of Botanical Nomenclature (GREUTER et al., 2000). SoLDANO (1993) noted that this combination was in fact published by FIORI (1924), but in reference to a different taxon, P. villosa Desf., which is an illegitimate name (superfluous) and not a nomen nudum (so that it was not validly published, as proposed by CuLLEN & HEYWOOD, op. cit.), and so is valid as reference for a new combination and thus Soldano proposed the new name P. officina/is subsp. huthiz; which is the correct name for this taxon. Starting from HuTH (1891), the plants from the Central Apennines have been included in the preceding subspecies by many botanists. This is because they both possess the same type of hairiness and an elevated number of leaf segments (25-55). However, we prefer to distinguish them from subsp. huthii because of their different form and size of the segments, lineare-lanceolate and

Downloaded by [Università di Pisa] at 02:39 05 November 2015 110-60 x 23-12 mm. They also have a slightly different ecology, being charac­ teristic of mountain grassland. This peony has a long taxonomic history. The first report was by TENORE (1811) who recorded P. officina/is, but later changed his opinion TENORE (1830) reporting P. peregrina. However, after a year he called the same plant P. paradoxa (TENORE, 1831) and, lastly, P. pubens Rchb. (TENORE, 1835). HuTH (1891) identified it in his new taxon P. peregrina var. villosa, followed by FIORI & PAOLETTI (1898) in their new combination P. officina/is subsp. euofficinalis var. villosa (Desf.) Fiori & Paoletti, and by Stern (1946) in the combination P. humilis var. villosa (Huth) Stern. Finally, PIGNATTI (1982) reported for the Central Apennines both P. officina/is subsp. villosa and P. peregrina, but limit- 240 N.G. PASSALACQUA,L. BERNARDO ing the latter to the Maiella Massif (Abruzzi). In our opinion, there is only one variant that deserves to be considered as a different subspecies and we propose P. officina/is L. subsp. italica, subsp. nov. The recent reports of P. officina/is L. subsp. banatica (Rachel) So6 (PoLDINI et al., 2001; 2002) fit well with our own observations, and also the comparison of samples from the Karst show a good similarity to herbaria specimens of P. banatica. P. banatica is treated by CuLLEN & HEYWOOD (1964b) as a subspecies of P. officina/is L., distributed in Romania, Hungary and Jugoslavia ( and ), living on sandy steppes and deciduous forest. So6 (1960) made a thorough investigation on the taxon, based on the variability of leaf morphol­ ogy and hairiness of specimens from many places. He found that the "apical leaflet consist namely almost invariably of 2-3 lobes or 2-3 segments, and are very seldom entire, while the lateral leaves are as a rule entire." and that " Some terms of the original description - e. g. foliolis ... subtus arachnoideo-incanis­ are not always valid ... The hairs of the leaves are variable, below with very little, short hairs or becoming glabrous.". Moreover he stated that it "differs both in its habit and the partition of its leaves from the typical P. officina/is, whose lat­ eralleaves, too, are generally divided into 2-4 segments, on the specimens from South-Tyro! examined by myself (cf. Herb. Norm. 3501, FEAH. 90, ... , chiefly from Monte Baldo and Val di Ledro), leaves 2,0, 2,5,-3 em wide and 6-11 em long.". He concluded that "P. banatica is varying but remains within the above indicated limits of variation a uniform population which, although it is partly suggestive with its entire and broader leaves to a certain extent of P. mascula, it belongs, however, to the Formenkreis of P. officina/is ... ", referring the possible confusion to P. mascula particularly for plants from Banat-Bazias, growing "near the Monastery and in the forest" (i.e. in the same habitat of population from Karst), that NYARADI (1953) described asP. coral/ina var. triternati/ormis, but stating that "as an extreme variety may remain in the Formenkreis of P. ba­ natica". Downloaded by [Università di Pisa] at 02:39 05 November 2015 As we have seen above, the population from the Triestino Karst has been identified in the PCA (fig. 6a), but does not seem to show any real differentia­ tion in comparison to the plants from Alps and Tuscany. These Trentino plants have segments very similar to the typical subspecies for dimensions (124-60 x 53-27 mm) and number (8-30), but with almost complete dominance of partite segments (9-28) and almost absence of entire (0-2). The individuals observed are always subglabrous with the same kind of hair of those of the typical sub­ species. This morphotype lives in deciduous oaks woods of the hilly plain in deep soil (while all other populations of P. o//icinalis live on mountains in con­ solidated screes). These morphological, ecological and geographical characters separate this variant when compared to the other peonies, so that we confirm THE GENUS PAEONIA L. IN ITALY 241 that plants from the Karst can be attributed to P. officina/is L. subsp banatica (Rochel) So6. As for plants from Bazias, very probably the reports as P. mascula (L.) Mill. can be attributed to the misidentification of this taxon.

Subsection Foliolatae Stern 1946 [subsection Masculae (Stern ex Uspensk.) Halda 1998] The peonies of this group are characterized by cylindrical or fusiform root­ stocks, and in Italy they generally also show: - a more or less purplish stem, which is generally shorter than in subsec­ tion Paeonia; - leaf segments from oval to elliptic, mainly entire, varying in number from 9- 13 (-17); - a higher number of follicles {3-8) that lengthen at maturity bending downwards; - a Mediterranean distribution that lives in deep soil of deciduous woods, but also in evergreen oak woods and in beech woods if within its southern limits. According to the most recent Floras, in Italy there are four taxa in this group: P. mascula (L.) Mill. subsp. mascula, present in continental Italy from Calabria to Tuscany and in Friuli; P. mascula subsp. russoi (Biv.) Cullen & Heywood, present in Sicily and Calabria; P. morisii Cesca, Bernardo & N.G. Passal., present in Sardinia, and finally P. coriacea Boiss., present in Sardinia. However, the record of this last variant seems to be a past error in the sam­ pling and setup of herbarium material (CuLLEN & HEYWOOD, 1964b) or it could be an error linked to STERN' s citation of P. russi var. reverchoni for sar­ dinia (see "Historical taxonomic treatment") wich is a peony with glabrous carpels like P. coriacea Boiss. In Sicily another taxon was recorded: P. flaves­

Downloaded by [Università di Pisa] at 02:39 05 November 2015 cens C. Presl (1822; subsequently named P. corallina var.flavescens (C. Presl.) Guss., 1843) a combination put in synonymy with P. mascula subsp. hellenica Tzanoudakis (STEARN & DAVIS, 1984), a name accepted by Flora Europea (AKEROYD, 1993) that however does not allude to the presence of the subspe­ cies in Sicily.

RESULTS

The observation of living populations from the Italian Peninsula, Sicily and Sardinia, has point out a lot of affinities and divergences (tab. 6). 242 N.G. PASSALACQUA, L. BERNARDO

TABLE 6-Variable morphological and ecological characters among main populations of P. mascula s.l. Peninsula Sicily and Calabria Sardinia (exd. Calabria) gen. completely red Stem green green to slightly red to petioles Leaflets no (9-) 13-18 (9-) 13-18 (7-) 9 (-12) mm 90-180x38-145 90-180x38-145 60-135x34-85 shape ovate-elliptic (ovate) elliptic (ob- )ovate-elliptic texture thin thin thick and coriaceus upperface green opaque green opaque green bright lowerface (sub-) glabrous villous to subglabrous wolly to subglabrous Corolla red purplish red purplish to white mauve rose Fruit no (1-) 3 (-5) (1-) 3 (-5) (2-) 3-5 (-8) tapering, gradually shape abruptely reduced abruptely reduced - attenuate Supramediterranean Supramediterranean Also meadowlands Habitat deciduous woods deciduous woods and stony ground Altitude (m) 500-1000 500-1500 40-1700 Soil Fresh and deep Fresh and deep Also poor soil

In all populations stem, leaf stalk and main nerve are purplish at the begin­ ning of the development, while at maturity there are some differences among the populations of Sardinia, that have the tendency to maintain the purple col­ oration, those of Sicily, that become green at least at the base of the stem and those of the Peninsula that become completely green. Meaningful variations are observed in the morphology of the leaves; the populations of Sardinia show more constant segmentation with 9 segments in the lower leaf, while the populations from Sicily and the Italian Penisula show Downloaded by [Università di Pisa] at 02:39 05 November 2015 13-17. The segments are ovate to obovate and generally smaller in the Sardinian populations, while the populations of Sicily and the Peninsula have leaf seg­ ments that can be elliptic and of greater dimensions. In the peonies of Sardinia the leaf segments are coriaceous and the upper surface is shiny, while in the peonies of the Peninsula and Sicily the consistence is not coriaceous and the upper surface is matt. A very particular value is done by hair, that can be found, besides on the carpels, also on the lower page of the leaf segments, along the principal ribs and on the stem, varying a lot for quantity and quality. From the qualitative point of view we have two typologies (fig. 11): wooly hairiness with pluricellu- THE GENUS PAEONIA L. IN ITALY 243

Fig. 11 - Different kind of hairs of the lower leaf surface in different populations of P. mascula group (from CESCA et al., 2001): Sardinia (a, b) and Sicily (c, d)

lar crisp hairs, in the Sardinian populations and glabrous to hispid or villous plants, with the hairiness that decreases from South to North, with straight hair, uni- or pluricellular, in Sicily and Calabria. In the rest of the Peninsula the leaves are glabrous, or rarely with a few hairs along the main ribs. The petals vary from wine-red to deep rose in the Peninsular populations to pink in those of Sardinia, while in Sicily individuals flowers striped with white to completely white can be found. The fruits are of two types: in the Sardinian populations the carpels are more numerous (4-5) and become thinner gradually beginning from the me­

Downloaded by [Università di Pisa] at 02:39 05 November 2015 dian part toward the base and the apex, while in the populations of the Penin­ sula and Sicily there is a smaller number of carpels (2-3) which are abruptly truncated at the base and apex. It should be said that in Sicily there is another population, isolated in com­ parison to those considered for making tab. 6. This population, found on the Iblei mountains in a closed Quercus ilex community, consist of only about thirty individuals and is not in the best of health. Mter four years of checking the population at the most opportune time, we have always only found imma­ ture individuals: the number and the dimensions of the leaves is always very modest and the hairiness reduced; moreover no plant has been seen in flower or fruit. For such reasons, this population has not been considered in tab. 6, 244 N.G. PASSALACQUA,L. BERNARDO that refers to observations on mature individuals, in flower or in fruit. Few plants from Iblei have been cultivated: fertile plants have~come up. Surpris­ ingly, one of them was very similar to that from Sardinia for type of hairiness and morphology of the stem and leaves. We believe, therefore, that the popu­ lation from Iblei needs further investigation, even if we attribute it to the same taxon as those from the rest of Sicily. As we have seen, the populations from Sardinia are diploid (tab. 7), while those from the Peninsula and Sicily are tetraploid showing a clear caryological discontinuity; the population from Iblei has both diploid and tetraploid indi­ viduals, partly confirming the morphological affinities to both ploidy level ob­ served in the cultivated plants. A molecular analysis (MusACCHIO et al., 2000) has already shown notable affinities between the Peninsular populations and the Sicilians ones (tab. 7), on the one hand, and between those from Sardinia and partly with those from Iblei, on the other. The comparison of the ITS1 sequences shown by the ten populations examined has allowed to distribute the samples in three groups, each characterized by an elevated degree of homology. Particularly, the first group (100% homology) includes all the Sardinian samples and individuals from Iblei; a second group (100%) includes some populations from Calabria, Etna and Madonie and the last group includes those from the Nebrodi, Colli Albani and Gargano (99,6%). Anyway, these two last groups have a degree of homology among them which is greater (99%) than each has to the Sardinian ones. The comparison with the ITS-1 sequences of SANG et al. (1995) shows

0 TABLE 7-Chromosome number ( ) and variable nucleotide sites in ITSl segments(") in dif- ferent populations of P. mascula group. 0 ( ) Peruzzi L., personal communication (")from MusACCHIO et al., 2000 (modified)

Locality 2n= Site number identity

Downloaded by [Università di Pisa] at 02:39 05 November 2015 1 2 3 4 5 6 7 M.Albo 10 T A T A G T T 100% M. d'Ischudu 10 * * * * * * * Bruncu Seina 10 * * * * * * *

Pollino 20 A G c T c c * Madonie 20 A G c T c c * 100% Etna 20 A G c T c c *

Nebrodi 20 * G c * s * * Gargano 20 w G c w s * * 96% Colli Albani 20 * G c w s * * W=A&T; S=C&G THE GENUS PAEONIA L. IN ITALY 245 that the group with our Sardinian samples fall into the molecular variability of the unit that Sang attributes to P. russoi (Sang's analysis has probably been made on Sardinian plants), the samples from Calabria, the Madonie and Etna are P. mascula subsp. hellenica and the group from Nebrodi, Colli Albani and Gargano, P. mascula subsp. mascula.

DISCUSSION

Morphological analysis, chromosome counts and molecular data lead us to the opinion that the peonies from Sardinia (and possibly some from Iblei), should be attributed to a single taxon, those from the Peninsula to a second and those from Sicily and Calabria to a third one. The nomenclature of three units corresponds to taxa of different hierarchical ranks. The peonies from the Peninsula (with exclusion of Calabria) can be attrib­ uted to P. mascula subsp. mascula, the typical subspecies. Following STEARN & DAVIS (1984) this taxon is to be referred to LINNAEUS (1753) because, even if MILLER ( 17 68) did not cites Linnaeus in the 8th edition of his Gardeners Dic­ tionary, he cite the same pre-Linnean polynomials occurring in Linnaeus's Spe­ cies Plantarum, both authors taking their diagnostic characters from HALLER (1742). The Linnean name, P. officina/is var. mascula, has been typified by ScH­ MITT (2003) who excluded the provenance of this taxon from Switzerland and confirmed that the selected specimen ("Paeonia mas altera, quae tardier J.B. 3. 492", no 211.1. A, Clifford Herbarium, BM) is of doubtful provenance and so designated an epitype from Central France for a better definition of the taxon. Plants from the Italian Peninsula correspond well to this type and to the plant from Central France. The peony from Sardinia (with possibly also some from Iblei) must be ref­ ered toP. morisii Cesca, Bernardo & N.G. Passal., a recently described taxon Downloaded by [Università di Pisa] at 02:39 05 November 2015 (CEscA et al., 2001). The specific rank attributed to this is justified by the fact that it is well separated morphologically, caryologically and by the molecular analysis of its DNA. The peonies from Sicily and Calabria, generally with hairs on the lower sur­ face of the leaf segments and on the stern and with the corolla varying from white to red, can be attributed to another taxon for which we propose a new combination: P. mascula subsp. mascula var. russoi. In our opinion this taxon does not deserve subspecific rank, since the differential characters on which it is found are extremely variable, even within the same population. Growing to­ gether with hairy peonies, there are several individuals nearly glabrous, with wine-red flowers easily attributable toP. mascula subsp. mascula. STERN (1946) 246 N.G. PASSALACQUA, L. BERNARDO raised the difficult of problem the distinction between P. russoi and P. mascula plants from Sicily. The binomial P. russi (P. mascula subsp russoi (Biv.) Cullen & Heywood) was originally introduced by BIVONA (1816) and has created more than a few problems, because the protologue has too few details, lacks a type and so leaves doubt concerning the real constitution of this taxon. His description could be attributed completely to P. corallina Retz. (a binomial used by BIVONA for the real P. mascula subsp. mascula), with the exception of the hairs on the lower surface of leaf segments, that are variable. Besides, the comments that he adds to the protologue leaves some perplexity around his real knowledge of the ge­ nus Paeonia, for he maintained that the taxon he described: " Panormi cum P. corallina et P. o/ficinalis"; this last species has never been known in Sicily. Our research, even with the assistance of local botanists, has not enabled us to relo­ cate the locus classicus, which is given somewhat vague, any different individual from those coming from the Madonie, Etna and the Nebrodi. For that, we be­ lieve that Bivona has attributed this binomial to the pubescent form of P. mas­ cula (L.) Mill. very diffusely distributed throughout the whole of Sicily. Because of the hairiness on the lower surface of leaf segments, the authors after BIVONA have attributed to his taxon, most of peonies from Sicily, but also all peonies from Sardinia, including those that recently have been reported to a different species. We believe that the identification of the Sardinian peony with the taxon described by BIVONA was the result of the fact that many of the dif­ ferential characters of P. morisii Cesca, Bernardo & N.G.Passal., well evident in living plants (leathery of leaf segments, superior leaf surface shiny, stem and principal ribs markedly reddened) disappear in the dry specimens and are, therefore, not evident on the herbarium specimens on which the comparison were usually done. Moreover, also the different types of hairiness can be diffi­

Downloaded by [Università di Pisa] at 02:39 05 November 2015 cult to see if magnification is not available. The cultivation of individuals coming from different Sardinian and Sicilian places, besides the observation of living populations, have allowed us to note well the differences among two taxa. Among the others, the peonies from Sar­ dinia show in cultivation a precocious phenology, often flowering in February, in comparison to those from Sicily and Peninsula, flowering in April. The white corolla of some individuals from Sicily has resulted in the de­ scription of P. flavescens C. Presl, reported by Flora Europaea 2nd ed. as syno­ nym of P. mascula subsp. hellenica Tzanoudakis. However we think this does not have any taxonomic value, since in the Italian island this character is never accompanied by the others reported in the protologue of the Greek taxon (greater dimensions of the corolla and of the whole plant). THE GENUS PAEONIA L. IN ITALY 247

TAXONOMIC CONSPECTUS

subsect. Paeonia 1. P. officina/is L., Sp. Pl. 530 (1753) 1.1 subsp. officina/is (-) P. arietina sensu Stem, A Study of the genus Paeonia: 81 (J946), p.p. (=) P. officina/is subsp. euofficinalis var./oeminea (L.) Fiori & Paol., Fl. Anal. Ita!. 1:527 (1898) p.p. (-) P. peregrina sensu Caruel, Pr. Fl. Tosc.: 19-20 (1860) (=) P. peregrina var. officina/is (L.) Huth, Bot. ]ahrb. Syst. 14:270 (1891). Protologue: PAEONIA. Hort. cliff 212. Hort. ups. 149. Mat. med. 267. Sauv. monsp. 307. feminea a. Paeonia follis difformiter lobatis. Hall. helv. 311. Paeonia communis f. femina. Bauh. pin. 323. Paeonia femina.Fuchs hist. 202. Lob. ic. 602. Habitat in nemoribus montium Idae, Hel­ vetiae. Type: Lectotype: 692.1 (LINN). Epitype: in pascuis M. Generoso, vol. 30 n°122 (Haller Herbarium, P). illustrations: Fuchs (1542, pag. 202); Mattioli (1565, pag. 915); Lobel (1581, pag. 682). Description: Plant 35-90 em, green, hairy to glabrous. Lower leaves bitemate, with 9-15 leaflets deeply cut into (12) 14-34 (45) elliptic or oblong acute partite segments of 70-135 x 20-57 mm, which can be lobed; upper face green and glabrous; lower face glaucous, more or less covered with long stright hair, from villose to glabrous. Flowers red to pinkish. Carpels 2-3 tomentose by white or whitish thick, short hairs. Follicles sub-erect, 2-3 em. Chromosome number: 2n=20 Ecology: open consolidated block screes (rarely in wood) of the montane belt, in rich but stony and thin soil. General distrubution: France, Italy (Val d'Aosta, Piedmont, Lombardy, Venetia, Trentino, Friuli, Emilia and Tuscany, fig. 12), Switzerland (Canton Ticino), Slovenia, , Bosnia, Mon­ tenegro and Albania.

Selected specimens seen: Italy- Val d'Aosta: Vallone eli Machaby, Vaccari, 6.VI.1900 (FI); Val D'Aosta, Col­ line eli "Donnes" presso Gran Barme, Valli Lonzo, Sauro Michele, 19.V.1899 (TO); Arnaud (Valle d'Aosta)-Testa Colois, radure eli lariceto, 1700-1800, M. Bovio, VII.1980 (TO); Piedmont: Serra eli San Michele, Val eli Susa, Fe"ari, 19.V.1899 (FI); Sacra eli S. Michele, boschi sopra Ia borgata S. Pietro ad est della cascina Giocosa (Val eli Susa), Mattirolo e Negri, 12.V.1920 (TO); Pendii lungo il sentiero dalla Sacra eli S. Michele aS. Francesco sotto l'Ancocia (Val eli Susa), Vallino, 7.V.1911 (TO); Boschi presso Ia sagra eli S. Michele della Chiesa (Alpi Cozie), Ferrari e Bmino, 17.V.1891 (TO); Trana, rupi est del M. Pi­ etraborga, P. Fontana, 26.VI.1926 (TO);-Valle della Stura eli Lanzo, riva destra del torrente su Balme verso

Downloaded by [Università di Pisa] at 02:39 05 November 2015 il piano della Mussa, Vallino, 1888 (TO); Balme: nel bosco eli !arid salendo al piano della Mussa, Nigra e Suillot, 5.VII.1920 (TO); Balme: in mezzo ai pini salendo al piano della Mussa, Fe"ari, F. Santi, 21.VIII.1915 (TO); Mattia, presso Ia torre (Val Susa), Montacchini e Aiello, 22.1V.1970 (TO); St. Ambro­ gio; boschi alia Sacra eli S. Michele, P. Fontana, 12.V.1907 (PAD); Val eli Lanzo, lungo Ia strada Balme e Pian eli Mussa,TO, 1500 m, L. Bernardo, N.G. Passalacqua, 21.VI.1995 (CLU); Val eli Lanzo, lungo Ia strada Balme e Pian eli Mussa,TO, 1500 m, L. Bernardo, N.G. Passalacqua, 19.VI.2001 (CLU). Lombardy: Comi eli Canzo, presso Lecco, F. Levier, 29.VI.1871 (FI); Tra BrioneS. Giovanni (BS) quae Ia nella zona boschiva, G. Abba, 25.IV.1968 (TO); M. Dragoncello (BS), 400 m, in Omo-Ostryetum, E. Banfi, 6.VI.1976 (MI); Lombarelia, BG, Val Brembana: San Giovanni in Bianco, A. Piazzolli, 8.VIII.1985 (MI); Corni eli Canzo, Raniero, s.d. (BOLO); Lago eli Garda, M. Pizzoccolo, 1200 m, S. Zenari, 31.V.1925 (PAD); Nelle selve eli Albino e Nembo, Bergamasco, Bertoloni, 1839 (BOLO); Corni eli Canzo, Artaria, Vl.1887 (PI); M. Barro, in saxosis declivi merielionalis, Cesati, 18.VI.1894 (RO); Lago d'Iseo: Lovere Pi­ anico Sellero P. Gaiano eli Budino Vello (lungo strada) zone Marone, Anzalone, 3:V.1980 (RO); Monte Maddalena sopra Botticino, BS, L. Bernardo, N.G. Passalacqua, 24.VI.1995 (CLU); Monte Barro, Galbi­ ate, CO, 920 m, L. Bernardo, N. G. Passalacqua, 25.VI.1995 (CLU); Veneto: Collanti e Bollari (Venezia). P. Bolzon, (FI, Ml); Mons. Pastello prov. Verona, 3000', Brachis, III.1844 (FI); In colle eli Avesa prope Ve­ rona, Manganetti, 2.III.1845 (FI); M. Grappa, VII.1889 (FI); M. Costa-S. Fila, Vaccari, V.1896 (FI); Passo Downloaded by [Università di Pisa] at 02:39 05 November 2015

N -1>. 00

z p ~ Cll Cll ~ ~ r t:lj I

a.,IL- .ll_l!f "--'*---••--~

Fig. 12 - Italian distribution and picture (from M. Generoso, Switzerland) of P. o/ficinalis L. subsp. officina/is THE GENUS PAEONIA L. IN ITALY 249

di S. Ubaldo nelle prealpi bellunesi, presso Vittorio, luoghi soleggiati fra i cespugli, Pampanini, 6.VI.l904 (FI); Sulla montagna di San Lorenzo (Colli di Vittorio), 609 m, R. Pampanini, 20.V.1895 (FI); Bassano aS. Michele, L. Vaccari, 12.IV.1897 (FI); S. Fila, Zarpo, 10.V.1986 (FI); Venezia: in pascuis subalpinis Baldi Montis, alt. 1000, solo calcareo, Riga, VI.1904 (FI); M. Baldo presso la fontana di Nemba, Martelli, l.IX.1890 (FI); M. Baldo, A. Kerner, VII.1871 (FI); Veneto-Schievenin (M. Grappa), L. Vaccari, VIII.1819 (FI); Veneto-Puise (dintorni), L. Vaccari, 19.VI.1919 (FI); M. delle Ungherine presso Verona, Micheletti, (TO); Monte Ball: presso Serravalle (Vittorio), Bianchi, 26.IV.l899 (TO); Venezia, prope Ve­ rona: in pascuis subalpinis Baldi Montis, alt 1000 m, solo calcareo, Riga, 1904 (TO, PAD); M. Pastello prov Verona, A. Goiran, (TO); Mti Lessini, vetta del Pastello (1031 m), A. Forti, 6.VI.l906 (PAD); Vice­ nza, Colli Berici, Raniero, 1834 (BOLO); M. Tondo, Colli Berici, S. Pignatti, 29.IV.1967 (TSB); M. Pas­ tello (Valpolicella) (VR) 1128 m, Bianchini e DiCarlo, 13.VI.1984 (VER); Cavalo, Valpolicella (VR) 600 m, Pisani, 2.VI.1984 (VER); Verona sulle Torricelle verso le case vecchie 300m, Bianchini e DiCarlo, 8.V.1979 (VER); Sopra San Fidenzio (VR), 200m, Bianchini e DiCarlo, 16.V.1979 (VER); Avesa nel Vajo Gallina (VR), 100m, Bianchini e DiCarlo, 12.IX.1979 (VER); Castel Cerino, sopra Soave (VR), 500 m, Bi­ anchini, 22.V.1956 (VER); Baize calcaree esposte, a! M. Valdaldo m 400 Comedo (VI), A. Lucato, 15 .IV.1969 (VER); Sottobosco, in Prado m 1000, S. Zeno di Montagna (VR), A. Lucato, 28.V .1970 (VER); Prati sommitali, selletta a nord della cima del Pastel, m 1000, Fumane (VR), A. Lucato, 26.V.1979 (VER); In Vico Tremmiaco, loco dicto bosco di Ferrari, M. Precastio, A. Massalongo, 1850 (VER); In collibus su­ pra vicum Quinto, A Goiran, s.d. (VER); Passo di S. Ubaldo nelle Prealpi Bellunesi, presso Vittorio, R. Pampanini, 15.VI.1903 (PAD); Trail Colletto ed il Summano, m 900, pascoli e boschi di abeti, Frigo, 8.VI.1942 (PAD); Dintorni prossimi alia chiesa del Summano presso la croce. Versante sud, m 1290, Frigo, 23 .VIII.1942 (PAD); Colle Montorio (Verona), Berengere, 1840 (PAD); Frese sorgentive di Miane a Pollina, Berengere, VI.1843 (PAD); Summano, 7 comuni, Spranzi, s.d. (PAD); Verona in collibus di Avesa, Raniero, 1838 (BOLO); Le selve nei monti di Angarano, Montini, s.d. (BOLO); Verona colles, Brachi, 1840 (PI); In collibus di Aveda prope Veronam, Reiner, V.1864 (RO); Monte Ball, presso Serravalle (Vit­ torio), D. G. Bianchini, 26.IV.1899 (RO); In colle delle Ungherine prope Veronam, A. Menganotti, s.d. (RO); Colli veronesi, Goiran, V.1870 (RO); Trentino: Valsugana in M. Cofre, Ombrosi, s.d. (FI); Tirolia australis, Val di Ledro, in pascuis montanis sol. calcar., 1900-2000, Porta, VI.1893 (FI); Venezia Triden­ tina-Brentonico, arbusteti suM. Altissimo, M. Tamanini, 11.VI.1984 (FI); M. Altissimo (M. Baldo), B. Peyronel L. Rocco, 28.VI.1958 (TO); Ex Capo di Tenna a! prato diS. Antonio, Marzialetti, 1837 (BOLO); Frequente nei boschi e nei prati tra Andalo e Fai (Trento), Tonzig, 24.V.1938 (PAD); Mte Finonchio Bei Rovereto. Kalk, 1200 m, W. Pfaff, 14.V.1894 (PAD); Valsugana in sylvis monti Efre, Ambrogi, 1850 (BOLO); Nei monti di Rovereto de' distretti di Arco e Riva e ne' confinanti veronesi e bresciani, Bertoloni, s.d. (BOLO); Mte Baldo, A. Barbiero, 1827 (BOLO); Mte Baldo: Wiesen bei S. Valentino, 1300 m, Be­ hrendsen, 24.VI.1903 (PI); Versante Sud di Corna Piana, Monte Baldo, Trento, 1400 m, faggeta e rupi., L. Bernardo, N.C. Passalacqua, 18.VI.1996 (CLU); Terra Rossa, vers. merid. del Monte Finonchio sopra Scottini-Pornal, Trento, L. Bernardo, N.C. Passalacqua, 18.VI.l996 (CLU); Monte Corna Piana, Monte Baldo, Trento, 1600-1700 m, Pascoli d'altitudine, L. Bernardo, N.C. Passalacqua, 18.VI.1996 (CLU); Friuli: Flora Italiae, Prov. di Pordenone prealpi Carniche: M. i Cameroni (estrernita est del gruppo del Cavallo) 950 m s.l.m, L. Poldini, 26.IV.1973 (TSB); Flora Italiae, Prov. di Pordenone prealpi Carniche: M. Ciaurlec, 1000 m, L. Poldini, 29.IV; 1972 (TSB); Emilia: Faggeta alia cima del M. Pareto sopra Solignano (m 900) Parma, P. Bolzon, 26.5.1907 (FI); Baiso M.te Lusino (App. Modenese), Mori, 19.V.l956 (FI); Nella faggeta sopra Calestano (m 600-700) subappennino parmense. Emilia, P. Bolzon, 24.IV.1903 (FI); Colle del Gesso a Scandiano, prov. di Reggio Emilia, A. Fiori, 17 .IV.1984 (FI); Dirupi sopra Casale presso

Downloaded by [Università di Pisa] at 02:39 05 November 2015 Montegibbio, A. Fiori, 10.V.1883 (FI); Fra Vignola e Guiglia (prov. di Modena), Mattei, IV.1880 (FI); Loc. Emilia-prov. di Bologna: in dumetis submontanis locis Rio Maggiore etJano prope pagum Sasso, ab­ bundans, alt. 300m ca. solo siliceo, A. Fiori, 29.IV.l904 (FI, TO, PAD, PI, RO); Colline presso Vignola prov. di Modena, A. Mori, IV.1877 (FI); Colli di Varano prov. di Modena, Mori, V-VI.l885 (TO); Monte Vignola de Conti prope Marzabo in districto Bononiensis, Bertoloni, VII.l827 (BOLO); Colle del Sasso presso Scandiano, prov. di Reggio, A. Fiori, 1l.IV.1885 (RO); Strada per Assella (Valmozzola), L. Mon­ tanari, 2.V.l981 (GE); In monti Parmensi a Cassio, Passerini, VI.1866 (FI); M.te Prinzera nei boschi di quercia, Parlatore, 3.VIII.1838 (FI); Tuscany: Pisanino (Apuane), S. Sommier, l.VI.1871 (FI); M. Sagro (Carrara), Bolzon, VI.1896 (FI); Cima della Penna di Lucchio (Toscana), P. Fantozzi, 5.V.1887 (FI); App. Lucchese, V Martellli, VI.l884 (FI); Prato Fiorito, Sommier, 5.V.1869 (FI); Sul Pisanino, M. Grilli, 21.VII.l857 (FI); Villa Colle Mandina (Garfagnana) Pania di Corfino, roccette esp S-W, alt. m 1400-1500, Poggi, Nepi, Urbani, 3.VI.1987 (FI); Valle di Gremolaggio (Alpi Apuane) in pratis montanis, Sommier, 14.VI.1876 (FI); Appen. Lucchese a! Prato Fiorito, Beccari, 14.VI.1863 (FI); Toscana-Alpi Apuane (M. Altissimo) Passo del Vestito-Galleria Cipollaio, m 1151-900 ca., Corradi, Bavazzano, Contardo, 22.V.1953 (FI); Penna di Lucchio: vetta e cime vicine m 1150-1176, Maggi-Nardi, 20.VI.1967 (FI); Habui ex Tam­ bura alpium Apuanorum, Guidoni, s.d. (BOLO); Prato Fiorito, Bertoloni, 1842 (BOLO); Solchi dell' Apen. Lucchese sui Prato Fiorito, s.l., 1846 (PI); Base del Monte Pisanino vicino Otto di Donna, Alpi 250 N.G. PASSALACQUA, L. BERNARDO

Apuane, L. Bernardo, N.G. Passalacqua, 18.VI.1995 (CLU), Base del Monte Pisanino vicino Orto di Donna, Alpi Apuane, L. Bernardo, N.G. Passalacqua, 17.VI.2001 (CLU). Switzerland: M. Generoso, Ber­ toloni, 1829 (BOLO); M. Generoso, Negri, 21.VIll.1891 (TO); M. Generoso, E. Burnat, 7.VI.1845 (FI); M. Generoso, Gussone, s.d. (NAP).

12 subsp. huthiiSoldano,AttiSoc. Ita!. Sci. Nat. Mus. Civ. Stor. Nat. Milano, 133 (10): 114 (1993) (=) P. humilis var. villosa sensu Stem (1946) p.p. (=) P. officina/is subsp. villosa (Huth) Cullen & Heywood, Feddes Repert. Spec. Nov. Regni Veg. 69: 34 (1964) (=) P. officina/is subsp. villosa (Desf.) Cif. & Giacom., Nomenclator Fl. Ita!. Pars altera Dicotyle­ dones, Fasc. I: 308 (1954) (=) P. officina/is subsp. euofficinalis var. villosa (Desf.) Fiori & Paol., Fl. Anal. Ita!. 1: 527 (1898) p.p. (-) P. peregrina sensu Bicknell, Fl. Bordighera 8: 10 (1896) & sensu Burnat, Fl. Alpes Marit. 1:54 (1892) (=) P. peregrina var. villosa Huth, Bot. Jahrb. Syst. 14: 270 (1891) bas. Protologue: fol.laciniis ovalibus vel oblongis, subtus albido-glaucis villosis supemo sparse pilosis, carp. tomentosis. P. paradoxa DC.! (Anders.?) P. tatarica Mill. t. 19. Type: described on material from Spain, Southern France and Italy. Description: Plant 30-60 em, green, hairy. Lower leaves hi- to triternate, with 15-25leaflets deeply cut into (21) 30-55 (65) elliptic to ovate partite segments of 40-70 x 13-21 mm, often with nu­ merous lobes; upper face green and glabrous; lower face glaucous, strongly covered with long and flat hairs, villose. Flower red to pinkish. Carpels 2-3, tomentouse with white to whitish thick, short hairs. Follicles sub-erect, 2-3 em. Chromosome number: 2n=20 Ecology: open consolidated block screes of the montane belt, on rich but stony and thin soil. General distrubution: Italy (Liguria, fig. 13), F ranee.

Selected specimens seen: Liguria: M. Longo, sopra Zuccol, su Pigna, m 1000, S. Zenari, VI.1922 (PAD); Monte Toraggio sopra Pigna, Alpi Mar. Liguria, C. Bicknell, (PI); Me Toraggio (a 1971 m) nel Circ. diS. Remo Alpi Marittime, G. Mari, 4.VIll.1886 (RO ); Vers. orientale di Monte Toraggio, Bordighera, IM, 1750 m, L. Bernardo, N.G. Passalacqua, 19.VI.1995 (CLU). France: Col de la Sine, Andon, pentes roccailles, alt. 1100 m, 10.V.1968, Gavelle (FI); Montagne du Cheiron, au Col de Negay (Alpes Maritimes), 15.V.1863, E. Bur­ nat (FI); Pierre lmpie pres Sisteron, Basse Alpes, l.VI.1868, A. Burle (FI); Plaine de Caussole, sopra Grasse. Alp. Mar. Francia, C. Bicknell, 5 .V.1889 (GE); Montagne de Bergeande, L. Bertrand, V.1919 (GE).

1.3 subsp. italica subsp. nov. (-) P. humilis var. villosa sensu Stem (1946) p.p. Downloaded by [Università di Pisa] at 02:39 05 November 2015 (-) P. officina/is sensu Tenore, Fl. Napo/.1: 337 (1811-1815) (-) P. officinalz's subsp. euofficinalis var. villosa (Desf.) Fiori & Paol., Fl. Anal. Ita!. 1:527 (1898) p.p. (=) P. paradoxa sensu Tenore, Syll. Fl. Neap. 261 (1831) (-) P. peregrina sensu Pignatti, Flora d'Italia (1982) (=) P. peregrina sensu Tenore Succinta relazione del viaggio /atto in Abruzzo: 71 (1830) (-) P. peregrina var. villosa Huth, Bot. Jahrb. Syst. 14: 270 (1891) p.p. (=) P. pubens sensu Tenore, Syll. Fl. Neap., Appendix IV, 21 (1835) Protologue: P. officina/is L. subsp. huthii Soldano similis sed segmentis longioribus, linare-lanceo­ latis et apicibus acutis differt. Type: Prati di Tivo, Gran Sasso d'ltalia, Abruzzo, 1450 m s.l.m., L. Bernardo, N.G. Passalacqua, 13. Vll. 2001 (CLU). Description: Plant 40-55 em, green, hairy. Lower leaves hi- to tritemate, with 15-20 leaflets deeply cut into (21) 24-45 (55) linear-lanceolate partite segments of 60-103 x 15-20 mm, with few or THE GENUS PAEONIA L. IN ITALY 251

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no lobes; upper face green and glabrous; lower face glaucous, strongly covered with long and flat hairs, villose. Flowers vinous red. Carpels 2-3, tomentouse with white thick and short hairs. Follicles sub-erect, 2-3 em. Chromosome number: 2n=20 Ecology: open ground of the montane belt, in rich and deep soil. General distrubution: Italy (Marches, Umbria, Latium, Abruzzi, fig. 14).

Selected specimens seen: Marches: Appennino di Cingoli, L. Narducci, IV.1877 (FI); App. Piceno, Arquata del Tronto, Forca di Presto suolo calcari 1500 m, A. Fiori, 24.VI.1929 (FI); Legi in appennino Piceno aM. Bimo (Marche), Gennari, VI1.1847 (FI); Pedate sui Pizzo di Sevo (Ascoli Piceno), C. Castelli, 15.VI.1886 (TO, RO); Ex Mte Priore a! Piceno, Marzialetti, s.d. (BOLO); Alia sommita Appennini Semanensis, Ber­ toloni, 1836 (BOLO); Nell'Appennino di Semano, ed a! Mte diS. Vicino, s.l., s.d. (PI); Falde dei monti dei Sibillini nella foce e comunissima in tutto il gruppo del Vettore, Cicioni, 23.VII.l886 (RO); Montes Sibilla, altezza media, E. Marini, 28.VI.1887 (RO); Monti Sibillini: pascoli freschi nel basso versante S del M. Castel Manardo attomo all'Ara del Re, 1550-1575 m s.l.m., suolo calcarea o pill o meno decalcificato, humus piu o meno abbondante, Brilli-Cattarim; L. Gubbellini, 11. VI1.1986 (PESA); Gruppo del Pennino: pascolo nel versante N del Mte Pennino, 1500-1525 m, suolo calcarea o pill o meno decalcificato, humus pill o meno abbondante, Brilli-Cattarini, L. Gubbellini, 28.VI.1985 (PESA); Monti Sibillini: pascoli as­ ciutti sui versante N del M. dell'Amandola lungo !'alto Fosso di Codarda, 1500-1550 m, suolo calcarea, humus da scarso a pill o meno abbondante, Brilli-Cattarini, L. Gubbellini, 24.VI.1983 (PESA); Monti Sibillini: fruticeti e radure erbose lungo il torrente Fiastrone nella Vaile del Farnio, 1300-1325 m, suolo calcarea, humus pill o meno abbondante, Brilli-Cattarini, L. Gubbellini, 22.VI1.1983 (PESA); Monti Sibil­ lini: pascoli freschi sui versante W del M. diVa! di Fibbia a! Capriole, 1500 m, suolo prevalentemente cal­ carea, humus pill o meno abbondante, Brilli-Cattarinz; L. Gubbellini, 9.VII.1981 (PESA); Monti Sibillini: pascoli nel basso versante SE del Pizzo di Meta, 1500 m, suolo calcarea o pill o meno decalcificato, humus abbondante, Brilli-Cattarini, 15. VII.1975 (PESA); Gruppo del M. San Vicino: luoghi semirupestri nel ver­ sante SE del M. San Vicino aile baize della Grotta, 1350-1425 m, suolo calcarea, humus generalmente scarso, Brilli-Cattarini, T. Hegly, R. Widmer, 2l.VI.1967 (PESA); Gruppo del M.S. Vicino: prati e fruti­ ceti nel versante E del M. S. Vicino, ai prati diS. Vicino, circa 1180 m, suolo decalcificato (+humus), Brilli-Cattarini, 2l.VI.1963 (PESA). Umbria: Dintomi di Castelluccio, A. Batelli, VII.l886 (FI, RO); MonteS. Vicino nei prati a 1100 m. Umbria, Bucci, 4.VII.l867 (FI); Alia sommita del Mte S. Vicino. Tro­ vata fruttificata in agosto, Narducci, s.d. (BOLO); Dintomi di Castelluccio di Norcia, A. Batelli, s.d. (PI); Mte Serra Santa sulla destra di Val Sorda, Gualdo-Tadino, s.l., 9.VI.1912 (PI); Mte Vettore (Umbria), Corsini, s.d. (RO); In prati montanis San Vicino, Prolli, s.d. (RO); Gruppo della Serra Santa: fruticeti radi nella bassa V a! Sorda presso le T rasce, 100-1025 m, suolo calcarea pill o meno decalcificato, humus pill o meno abbondante, Brilli-Cattarini, 16.VI.1980 (PESA); Gruppo della Serra Santa: pascoli e radure erbi­ ose nel versante NE del M. Nero aile Rotelle, 1150- 1175 m, suolo prevalentemente calcarea, humus pill o meno abbondante, Brilli-Cattarini, Gubbellini, l.VI.l979 (PESA); Gruppo della Serra Santa: faggete deg­ radate sulla cresta N del M. della Serra Santa, 1350 m, suolo calcarea pill o meno decalcificato, humus ab­ bondante, Brilli-Cattarini, f. Coaz, R. Sialm, 17 .VII.1972 (PESA); Gruppo della Serra Santa: fruticeti era­ dure erbose nel versante E del M. della Serra Santa alia Sella della Chiaravalle, 1250 m, suolo prevalente­ mente calcarea, humus abbondante, Brilli-Cattarini, f. Coaz, R. Sialm, 1o.VI1.1972 (PESA). Latium: M.

Downloaded by [Università di Pisa] at 02:39 05 November 2015 Terminillo: Vallone Rio Fuggio (m 1000-1200) verso Leonessa, Anzalone, 30.VI-2.VII.1993 (RO); Vallepi­ etra (Monti Simbruini), Campo dellaPietra (presso SS Trinita), m 1300-1400,Anzalone, 30.VI.1990 (RO); M. Simbruini, Valico sopra Filettino, Fra Domenico, VI.1985 (RO); Alta Valle Rio Fuggio, Leonessa, RI, 1240 m, L. Bernardo, l.VI1.1993 (CLU). Abruzzi: M. Morrone supra Solie, E. Levier, V.1875 (FI); In Monte Morrone, in silvis montanis ultra 1000 m, S. Sommier, 4.VII.1872 (FI); Supra Caramanico in saxe­ sis regionis fagi Monte Morrone, E. Levier, 4.VII.l877 (FI); In subalpinis montes Majella (V alii d' Orfento a! Piano del Mulino .... ), E. Levier, 6.VIII.l874 (FI); Rocca di Cambio (Abruzzo) base seggiovia per Campo Felice (m 1500), B. Anzalone, l.VI.1971 (FI, RO); In nemorosis: San Pietro d'Isola di Gran Sasso 1600 m, Zodda, VI.1950 (FI); Nei prati di Trapietro, gruppo del Gran Sasso prov. di Teramo, A. Fiori, 14.VII.1907 (FI); Abruzzo, Gran Sasso, Pietracamela, calcare, 800-1000, Siarra, 13.VIII.1927 (FI); M.S. Franco (Aquila) fra valle dell'Infemo e Casa Cappelli 1600-1700, L. Vaccari, 24.VI1.1906 (FI); Gran Sasso d'Italia sopra Assergi, A. Fiori, 15.VII.1886 (FI); Sui monti boscosi. Nella valle dell'Inferno del Monte Grande, nella Costa, A. Cecchetti, 1830 (BOLO); Morrone, Guadagno, s.d. (PI); Valle dell'Orfenta (Ma­ jella), Pedicino, 6.VI1.1872 (PI); Rocca di Cambio (Sirente), presso Ia funivia, margine della faggeta, a 1400-1500 m, Montelucci, 28.VI.1971 (RO); Orfenta, N.A. Pedicino, s.d. (RO); Prati: Pietracamela ai prati di Tivo, 1400-1600 m, Zodda, VI.1956 (RO); M. dei Fiori, E. Fiorini-Mazzanti, s.d. (RO); In pratis monte­ sis M. Como, A. Orsini, s.d. (PI); M. Como alia Schiera, A. Orsini, 1826 (BOLO); Nel fondo di Valle Or­ fenta alia Piana dei Mulini con Lilium martagon, Guadagno, s.d. (PI); M. Morrone (Abruzzo), Pedicino et THE GENUS PAEONIA L. IN ITALY 253

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Mori, 24.VII.1879 (RO); Gran Sasso, Vena del Vannoni, N. A. Pedicino, 26.VII.1877 (RO); M. S. Franco (Gran Sasso), 1300·1500 m, verso Lago Campotosto, Anzalone, 27.VI.1974 (RO); Prato Tondo, Gran Sasso, Furnari, 4.1X.1971 (CT); Monti dell Laga: luoghi erboso-sassosi lungo il Rio Castellano alla diga ENEL della Piana Cavalieri, 1350 m, suolo arenaceo, humus scarso o subnullo, Brilli-Cattarin~ L. Gubbel­ lini, 13.V.1983-6.VI.1983 (PESA); Monti della Laga: praterie e pascoli nel versnate SE del Mte Comuni­ tore presso il Passo di Chino, 1575-1600 m, suolo arenaceo, humus da scarso a pili o meno abbondante, Brilli-Cattarin~ L. Gubbellini, 13.VII.1983 (PESA); Boschi di Chiarino presso !'Aquila, Gussone, IX.1823 (NAP); Piano del Mulino nella Valle dell'Orfento, Gussone, s.d. (NAP); Maiella nella Valle dell'Orfento, Gussone, 20.VII.1834 (NAP); Valle dell'Orfento, presso Sfischia, Maiella, L. Bernardo, N.G. Passalacqua, l.VI.1994 (CLU); Valle del Vasto, Gran Sasso d'ltalia, L. Bernardo, N.G. Passalacqua, 31.V.1994 (CLU).

1.4 subsp. banatica (Rochd) So6, Noveny/oldrajz 146 (1945). (-) P. arietina sensu Stem (1946), p.p. (=) P. banatica Rochd, Pl. Banat. Rar.: 48. t. 11 (1828) bas. (-) P. coral/ina sensu Marchesetti, Flora di Trieste e de' suoi dintorni: 17 (1896-1897). (-) P. mascula subsp. arietina (Anderson) Cullen & Heywood, Feddes Repert. Spec. Nov. Regni Veg. 69: 35 (1964) p.p. (-) P. mascula sensu Stem, A Study of the genus Paeonia (1946) p.p. (-) P. officina/is subsp. coral/ina var. mascula (L.) Fiori & Paol., Fl. Anal. !tal. 1, 527 (1898) p.p. (-) P. officina/is subsp. euofficinalis var./oeminea (L.) Fiori & Paol., Fl. Anal. !tal. 1, 527 (1898) p.p. (-) P. peregrina sensu Marchesetti, Flora di Trieste e de' suoi dintorni, 16 (1896-1897). Protologue: P. foliolis tri-quinquepartitis, laciniis lanceolatis decurrentibus, subtus pallidioribus venosis leviter arachnoideo - incanis, gerrninibus arcuato - patentissimis tomentosis, stigmati­ bus recurvatis. Radix perennis oblique - descendens ramosa nodosa lignosa, radiculis laterali­ bus descendentibus, multicaulis. Caules, basi vaginis membranaceis tecti, uni-bipedales erecti teretiusculi canaliculati glabri simplices foliosi. Folia ima altema petiolata temata: foliolis tri - quinquepartitis, laciniis lanceolatis acutis in petiolum canaliculatum decurrentibus, plicato - undatis caeterum integerrimis; summa simplicia florem stipantia; omnia supra obscure - viridia glabra, subtus leviter arachnoideo - incana, costis venisque rubescentibus picta. Flos speciosus in apice caulis solitarius, foliolis simplicibus, rarissime tripartitis, basi dilatatis am­ plexus. Calyx quinquesepalus, sepalis subrotundis, flore explicato deciduis. Corolla speciosa P. officinalis minor penta - hexapetala sanguinea, petalis subrotundis basi angustatis. Stamina innumera, filamentis subulatis basi dilatatis albis, antheris oblongis flavis. Pistilla rarissime duo-quatuor potissimum tria sub anthesi erecto-patula. Germina villosa; styli subnulli; stig­ mata compressa recurva purpurea. Fructus numero pistillorum ovato - oblongi ventricosi ar­ cuato - recurvi depressi tomento ferrugineo undique vestiti. Semina subrotundo-angulosa lae­ via nitida atropurpurea. Rara in I-ae et II-ae regionis Bido-Berdo clivis arenosis, frequentior Downloaded by [Università di Pisa] at 02:39 05 November 2015 in graminosis versus pagum Ulma occurrens. Flor. Mart. April. Expl. icon, - a. planta integra. - b. fructus maturi. A. Paeonia peregrina. Mill. cui accedit, saris differt: foliolis tri-quinquepartitis, laciniis lanceo­ latis decurrentibus subtus arachnoideo-incanis, fructibus tomentosis arcuato-patentissimis. Hanc olim sub P. corallina. Willd. dedi. Type: described on plants from Bido-Berdo region (Serbia). Illustrations: Rochd A. (1828, pag. 46 tav. 11); Reichenbach (1838-1840, tav. 125). Chromosome number: 2n=20 Ecology: hilly woods of deciduous oaks in deep soil. General distrubution: Italy (Friuli, fig. 15), Serbia, Hungary and Rumania.

Selected specimens seen: Italia- Friuli: M. Vremiza, Marchetti, Vl.1896 (FI); In fruticosis (Seslerio-Ostryetum) Montis M. dei Pini supra Gropada s. calc. 660 m, L. Poldini et T. Wraber, 29.N.1966 (TSB); Prov. di Tri­ este Mte Medved (Mte Orsario) Repentabor, L. Poldini, 30.N.1966 (TSB); Opicina (Trieste) nel Bosco, A. Marcello, 2.X.1951 (PAD); Frequente nei monti del Carso presso Trieste, M. Tommasini, s.d. (PI); Tri­ este: Aurisina, Percedol Universitii, Cividale: Bosco Romagno, Anzalone, 5.X.1980 (RO); Sagrado di Downloaded by [Università di Pisa] at 02:39 05 November 2015

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Sgonico, strada forestale verso Monte Lanaro, Monrupino, TS, L. Bernardo, N. G. Passalacqua, 25.VI.1995 (CLU); Gropada, Carso Triestino, TS, L. Bernardo, N.C. Passalacqua, 26.VI.1995 (CLU). Slovenia: In fru­ ticetis lapidosis prope sacellum Sv. Hieronio in tractu montius Nanos supra visum Razdrto. Solo calc. 100 m. s.m., 24. V. 1969, Drufcovic (FI). Bosnia: Auf den rubinica bei Rusanovic, Bets Rogatica ca. 960 m, V. 1897, Kalk/iala (FI). Hungaria: In elatioribus montium apricis ericetosis ad Bazias, IV-V. 1831, Limkovics (FI); In montibus pr. Bazias, IV. 1888, Degen (FI).

2. Paeonia peregrina Mill., Card. Diet. ed. VIII, n. 3 (1768)

Protologue: PAEONIA (Peregrina) foliis difforrniter lobatis, lobis incisis, petalis florum rotundi­ oribus. Peony with deformed lobated leaves which are cut, and rounded petals to the /lower. Paeonia peregrina, flore saturate rubente. C.B.P. 324. Foreign Peony with a deep red /lower. The third sort grows naturally in the Levant; the roots of this are composed of roundish knobs like those of the second sort [P. officina/is senso stricto, P. /eminea], as are also the leaves, but are of a thicker substance: the stalks do not rise so high, and the flowers have a greater number of petals. This flowers a little after the other. Type: Neotype: Iter trojanum id. dumetiv supra Kuzkoei 12.V 1883, N 334, P. Sintenis (LE), (in M.S. Uspenskaja, L.W. Solowjewa, PAEONIA PEREGRINA MILL. v SSSR i ego tipi/ikazija, Bulletin Mosk. O-va Ispytatelej Prirody otd. Bioi. 1991 t.96 vyd.3 p.131-133.) Description: Stem glabrous, 30-50 em, green. Lower leaves biternate, firm, with some leaflets di­ vided to base, in all with 15-17 principal divisions, some of these deeply cut into 2-3 segments; segments and divisions coarsely toothed at apex; principal divisions 5-12 em long, glossy green above, glaucous, glabrous or sparsely pilose beneath. Flowers deep red, strongly con­ cave. Carpels 1-4, mostly 2, long tomentose. illustrations: Stapf 0. (1918; pag. 144, tav. 8742); Goulandris, Goulimis et Stearn (1968; tav. 22); Jordanov D. (1970; tav. 42); Goulandris N. (in Stearn W. T., Davis P. H., 1984; pag. 69). Chromosome number: 2n=20, 10 Ecology: supramediterranean decidous woods (oak), from 800 to 1200 m, in deep soil. General distrubution: Italy (Calabria, Basilicata; fig. 16), Albania, Greece, Serbia, Bulgaria, Ru­ mania and Turkey.

Selected specimens seen: Italy - Basilicata: Monte Carnara (San Paolo Alabanese, Potenza), 1000 m s.l.m., 26.V.1995, L. Bernardo (CLU); Calabria: Piano di Marco, ai piedi della Mula, S.Donato di Ninea, CS, 1220 m, L. Bernardo, G. Cesca, 21.V.1993 (CLU); Piano di Marco, ai piedi della Mula, S.Donato di Ninea, CS, 1220 m, L. Bernardo, N.G. Passalacqua, l.VI.1993 (CLU); Monte Sellaro, presso il Bifurto, Cercbiara di Calabria, CS, 950 m, L. Bernardo, 15.VI.1994 (CLU); San Donato di Ninea (Cal. Citra) al Piano delle Vacche, 1200 m, Lacaita, 23.VIII.l912 (BM); Un po' sotto Ia regione del faggio prima di arrivare alia

Downloaded by [Università di Pisa] at 02:39 05 November 2015 Grotta di Affrido (San Donato di Ninea), B. Longo, 22.VII.1903 (RO); Greece: Karia, Monte Stavrotas, Isola Lefkas, 900 m, Gariga sassosa a Q. coccifera., L. Bernardo, N. G. Passalacqua, 19 .V .1996 (CLU); Pala­ mas, Lamia, prov. Phtios, 540 m, Bosco di cerro e farnetto., L. Bernardo, N.G. Passalacqua, 16.V.1995 (CLU); Lefkas-Ionian island-Mt Stavrotas NW the village Ajios Ilias alt. 900-1150 m. W of Chapel Profitis Ilias ruins. Fresh places between rocks, E. Stamatiadou, 28.V.l971 (BM); Mte Athos, pr. Chilandari, Hayek, s.d. (BM); Serbia: In collibus circa oppidum Nisch., S. Petrovic, IV.1884 (BM); Albania: Melesine, above Lescovik, 400ft, limestone slope near summit, A. H. G. Alston, N.Y. Sandwith, 19.VI.1933 (BM); Romania: Altenia, distr. Calafat. In silvis, inter pagos Plenita et Verbicioara, alt cca. 120m s.m., Al Buia, M. Pliun, D. Cirtu, G. Pulga, M. Olaru, 20.V.1963 (BM). Bulgaria (translated from cirillic): Mt. Sredna Gora, near Srednogorec, in in open wood on limestones, 28.V.1964, I. Gancev, V. Velcev (154442 SOM); T racika plane- hill, Svilengrad district, near by vill. Mezek, 29. V.1980, M. Marcova (142879 SOM); Sliven district, Joe. Barmuk Bair, in open shrubs and in rocky grasslands on limestones, 5.VI.l964, P. Panov (131733 SOM); Malko Tarnovo district, in open shrubs near crossroad to vill. Stoianovo, 21.VII.1977, D. Peev (137259 SOM); Hascovo district, Uzundjovsca Koria loc., Sevdjata place, near trees and shrubs, 20.V.l974, P. R.ahov (129925 SOM); (in latin) In pascuis dumosis Deli ormani supra urben Varna, 10.V.1899, B. Davido/f(26349 SOM). Downloaded by [Università di Pisa] at 02:39 05 November 2015

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3. Paeonia mascula Mill., Gard. Diet. ed. VIII, n. 1 (1768)

3 .1 subsp mascula (=) P. coral/ina Retz., Obs. iii. 34. (1783) (=) P. officina/is [beta] mascula L., Sp. Pl. 530. (1753) (=) P. officina/is subsp. coral/ina var. mascula (L.) Fiori & Paol., Fl. Anal. Ita!. 1, 527 (1898) (-) P. officina/is subsp. euofficinalis var.foeminea (L.) Fiori & Paol., Fl. Anal. Ita!. 1, 527 (1898) p.p. Protologue: Paeonia (Mascula) foliis lobatis ex ovatolanceolatis. Haller, Hdv. 311. Peony with lo­ bate leaves which are oval and spear-shaped. Paeonia folio nigricante splendido, quae mas. C.B.P. 323. Peony with dark shining leaves, otherwise male Peony. The first sort here enu­ merated, is the common male Peony, which grows naturally on the Hdvetian mountains. Type: Lectotype: "Paeoniamas altera, quaetardiorJ.B. 3. 492", no 211.1. A. (Clifford Herbarium, BM). Epitype: "Paeonia coral/ina Retz. Savigny les Beaune (Cote d'Or) bois, taillis pierreux calcaire, vers la partie superieure de laCombe Vautdoy alt. 400 m env. 24 mai 1911legi Ldevergnes" (P). Description: Stem 30-80 em, glabrous, purplish when young, completdy green when mature (rardy slightly purplish in the lower parts). Lower leaves with 13-18leaflets (rardy 9), from ovate to dliptic, 390-180 x 38-145, entire or bifurcate, pale green and glabrous above, gla­ brous or with few stright hair bdow. Flower fuchsia coloured, falling precociously. Follicles (1) 3 (5), covered with white tomentum, rounded at apex, sub-erect, curved backward when mature. illustrations: Mattioli P.A. (1565; tav. 914); Lobelius (1581; pag. 683-684); Sowerby et Smith (1806; tav. 1513). Chromosome number: 2n=20 Ecology: supramediterranean decidous woods (oak, chestnut), from 500 to 1000 m a.s.l., in deep soil. General distrubution: Italy (Latium, Apulia and Basilicata, fig. 17) and Greece; other localities in Southern Europe and Asia Minor are to be verified.

Selected specimens seen: Tuscany: Bosco della Certosa a Firenze, A. Tongiorgi-Tozzetti, 1833 (BOLO); In luogo affatto sdvaggio in settentrione Volterra, Amidei, 1841 (BOLO); In una sdvetta di lecci ed altro, Volterra, Amidei, (PI); Etrurio in umbrosis junta coenobium Carthusianorum, E. Levier, IV.1886 (PI); Firenze, bosco della Certosa, Caruel, 19.IV.1859 (PI)- Nel bosco dei frati della Certosa di Firenze, Arcan­ Downloaded by [Università di Pisa] at 02:39 05 November 2015 geli, 8.IV.1971 (RO); Latium; In silvaticis Albano, Rolli, 13.IV.1863 (RO); Villa della Rufenella, 0. Grampini, 10.V.1896 (RO); S. Cesareo, nd bosco di castagni, frequente, A. Cacciato, (RO); Sdva del Tus­ colo, Panebianco, 6.V.1880 (RO).; Abbondante nel castagneto sopra Rocca Priora, 550-600 m.s.m., Anzal­ one, 15.V.1995 (RO); Castelli Romani (Monti Albani), Bosco della Malora, Anzalone, IX.1956 (RO); RR nei castagneti presso Camaldoli (Frascati) a 550 m s.m., G. Lusina, 3.VII.1939 (RO) San Polo, Sanguinetti, s.d. (RO); Colli Albani, presso Bosco Riguardata-Molara, bosco di castagno, L. Bernardo et N.G. Passalac­ qua, 15.VI.1996 (CLU); Apulia: Gargano, nella foresta Umbra lungo !a strada per Vieste, bosco di cerro, L. Bernardo et N.G. Passalacqua, 29.VI.1995 (CLU); Bosca Caputi alle Murge di Ruvo di Puglia, Palanza, 14.IV.1897 (FI); Bosco di .... presso !a ValleS. Antonio (Gargano), V. Martelli, 21.V.1895 (FI); Bosco Um­ bro presso a! Cutino d'Otra, V. Martell~ 21.V.1893 (Fl); Murgia di Belluomo between Martina Franca, Vitantonio, VI.1883 (FI); Basilicata: Potenza in silva Pallarita, loco dicto valle delle Pile, alt. 1200 m, 0. Gavioli, 9.V.1927 (Fl).; Potenza in silva Aria Silvana, suolo argilloso calcareo, alt. 1200 m, 0. Gavioli, VI.1922 (Fl); Pignola in silva Cugni dicta, solo argilloso, alt. 1000-1200 m, 0. Gavioli, 7.V.1927 (FI); Po­ tenza, Bosco Pallareta, vicino monte Grosso, bosco di cerro, 1150 m, L. Bernardo et S. Oliveti, 5. VII.1996 (CLU); Lucignano (selva di Venusio) Bosco a Quercus macedonica, Matera, R. Corti, E. Francini, G. Negri, 22.V.1951 (FI); Pignola Bosco dell'Aria Silvana, Gavioli, V.1962 (FI); Potenza, Bosco della Pal­ larita, A. Fiori, VI.1910 (FI)- Lucania-Brindisi di Montagna in Silva Grancita alt. 1150, 0. Gavioli, THE GENUS PAEONIA L. IN ITALY 259

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10.VI.1928 (Fl); In nemore Acqua di Maio, solo argilloso, alt. 1000, 0. Gavioli, 7 .V.1927 (FI); Tra blocchi di arenaria presso una fonte Masseria del Monte, turrito Colombano, Basilicata, G. Bruno, s.d. (RO); Pomarico in Basilicata, P. Giordano, IX.1860 (RO).

3.1.1 P. mascula Mill. subsp. mascula var. russoi (Biv.), comb. nov. (=) Paeonia coral/ina var. /lavescens (C. Presl) Gussone, Fl. Sicul. Syn. 2:26 (1843) (=) P. /lavescens C. Presl, Velie. Prag. 5 (1822) ( =) P. mascula (L.) Mill. subsp. russoi (Biv.) Cullen & Heywood, Feddes Repert. Spec. Nov. Regni Veg. 69: 35 (1964) p.p. (=) P. officina/is subsp. corallina var. Russi (Biv.) Fiori & Paol., Fl. Anal. Ita!. 1, 527 (1898) (=) P. russiBiv.,Man. Fl. Sic. iv. 12. (1816) bas. Protologue: Peonia foliis biternatis, foliolis ellipticis, integris capsulis recurvatis pilosis, radice fusiformi. Radix fusiformis, simplex vd ramosa, ramis 1-2, pariter fusiformibus, patentibus. Caulis erectus, pedalis et altior, teres, subangulatus glaber, simplex, foliosus, purpurascens uti petioli. Folia inferiora biternata, summa saepe ternata, potiolo tereti, superne sulcato, foliolis obscure viridibus, ellipticis, integris, supra glabris, subtus vix pubescentibus. Calycis foliola quinque, quorum 2-4 parva, oblonga, concava, reliqua majora, elliptica, integra, raro biloba. Petala 5-6, obovata, chermisina. Capsulae duo, recurvatae, pilosae, pills flavis, latere interiori dehiscentes. Panormi in montibus cum Peon. officinalis et corallina. Perennis Floret Majo. Observ. Paeoniae humili quam corallinae proximior, at differt ab ilia forma radicis, integritate foliolorum et directione capsularum. Dixi in honorem solertissimi patriae Botanices cultoris D. Joachimi Russo Cassinensis. Type: Neotypus (designated here): Pizzuta, presso Piana degli Albanesi (Palermo, Sicilia), bosco aperto di querce decidue, 700 m s.l.m. ca, L. Bernardo & N.G. Passalacqua, 28.V.1996 (CLU; Neosyntype in Fl). Description: Plant 30-60 em, green, purplish only in the lower parts (rarely completely green or totally purplish). Lower leaves with (9) 13-18 leaflets, from elliptic to ovate (rardy bifurcate), 390-180 x 38-145, pale green and glabrous above, glaucous and villous to subglabrous below. Flower red, red purplish, white-striped or white. Follicles (1) 3 (-6), covered with white to­ mentum, rounded at apex, sub-erect, curved backward when mature. Chromosome number: 2n=20 Ecology: supramediterranean decidous woods (oak, chestnut, beech), from 500 to 1500 m a.s.l., in deep soil. General distrubution: Italy (Calabria and Sicily, fig. 18); reports from Greece are to be verified.

Specimina visa: Sicily: Bosco della Ficuzza, Minissale, 22.IV.1990 (CT); Piana degli Albanesi, Brullo e Spampi­ nato, 25.IV.1991 (CT); Bosco sotto M.S. Angelo, Brullo e Signorello, 6.VI.1982 (CT); Lecceta-Piano Zuc­ chi (Madonie), S. Brullo, 6.XI.l981 (CT); Bosco di Issullo, Bartolo, Brullo, Pulvirenti, Scelsi e spampinato, Downloaded by [Università di Pisa] at 02:39 05 November 2015 l.VIII.1990 (CT); Monte Cuccio presso Palermo, Gussone, IV (NAP); Palermo a! monte vicino San Mar­ tino, Gussone, IV.1839 (NAP); Ficuzza, Gussone, V (NAP); Madonie, Gussone, V (NAP); Boschi di Man­ danici, Gussone, 14.VI (NAP); Sagana presso Partinico, Gussone, V (NAP); Cauni, Gussone, V (NAP); Altavilla, Gussone, VII (NAP); Palermo alia Pizzuta, Gussone, IV.1839 (NAP); Pizzuta sopra Ia Piana, Gussone, V (NAP); Palermo alia Portella di S. Anna, Gussone, V (NAP); Selva dell'Etna, Cosentini, 1834 (NAP); San Michele, Gussone, V (NAP); Etna a! MonteS. Leo, Tornabene, 1847 (NAP); Pizzuta, A. To­ daro, V.1828 (PAL); Militello nei luoghi ombrosi ed incolti, A. Todaro, 1820 (PAL); Sopra A vola, A. To­ daro, IV.1840 (PAL); San Martino, s.l., V (PAL); Monte Cuccio, s.l., V.1850(PAL); Madonie all'ortaggio, s.l., V.1866 (PAL); Madonie, Parlatore, VIII.1871 (Fl); Palermo a Mte Cuccio, Parlatore, III.l842 (FI); In montis Panormitanis, Parlatore, VIII.1842 (FI); Panormo e Pizzuta, Parlatore, VIII.1847 (FI); In pasquis elatis montis Pizzuta (1500-2000 m.s.m.), e. etA. Huet du Pavillon, 23.V.1855 (Fl); In pascuis montosis et in silvaticis. Palermo, H. Ross, IV.1896 (FI); Ficuzza in silva, Sommier, 1l.V.1895 (FI); Boschi della Ficuzza-Sicilia- a 900 m, Biondi, 1l.V.1895 (FI); Ponticelli, Minii, VII.1846 (FI); Sicilia, Mistretta, bosco della Giumenta (Sambughetti), V. Martelli, 11-19.VII.1906 (Fl); Palermo, San Martino, D. Lanza, VI.1990 (RO); Sopra l'Etna, Polgano, s.d. (RO); Madoni, Gasparrini, s.d. (RO); Boschi di Collebasso, ter­ ritorio di Castiglione siculo, Cesati, (RO); Nebrodi, dopo il passo della Portella della Femminamorta, 1000 m, Bianchini, DiCarlo, 11.V.1996 (VER); Costone calcareo ad arbusti, pendici M. Renna o Matassaro, ol- THE GENUS PAEONIA L. IN ITALY 261

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tre Giacalone, m 780, Pioppo (Palermo), A. Lucato, 12.VII.1978 (VER); Monti di Palermo, Gussone, 1824 (BOLO); Castel Termini, bosco Buonanotte, V. Martelli, s.d. (PI); In sylvaticis montis alia Pizzuta, To­ daro, 1843 (BOLO, PI); Palermo in silvaticis montosis alia Pizzuta, D. Lanza, s.d. (PAD, RO, FI); Bosco della Ficuzza: lungo Ia strada fra l'Abbazia e Rocca Busambra, 900 m ca, L. Bernardo, N.G. Passalacqua, 28.V.1996 (CLU); Ficuzza, presso I' Alpe di Cucco, 980 m, L. Bernardo et N.G. Passalacqua, 28.V.1996 (CLU); Nebrodi, Bosco del Flascio, C.da Serra di Caracozzo, bosco a Quercus congesta, 970 m, L. Ber­ nardo, N.G. Passalacqua et G. Spampinato, 25.V.1996 (CLU); Etna, Bosco Maletto alle falde del monte, bosco a leccio e Quercus congesta, 1100 m, L. Bernardo, N.G. Passalacqua et G. Spampinato, 25.V.1996 (CLU); Nebrodi, Bosco di Petrosino, bosco a Quercus congesta, 900 m, L. Bernardo, N.G. Passalacqua et G. Spampinato, 25.V.1996 (CLU); Nebrodi, Bosco di Grappida, bosco a cerro, 1300 m, L. Bernardo, N.G. Passalacqua et G. Spampinato, 25.V.1996 (CLU); Madonie, Piano Battaglia, bosco di faggio in radure, 1500 m, L. Bernardo et N.G. Passalacqua, 27.V.1996 (CLU); Calabria: S. Demetrio Corone, CS, Giuliani, IV.1927 (NAP); Valle Olivella, Casiglia, sotto Monte La Mula (S.Sosti, CS), L. Bernardo, N.G. Passalac­ qua, l.VI.1994 (CLU); Valle Olivella, Casiglia, sotto Monte La Mula (S.Sosti, CS), N.G. Passalacqua, 3.VI.1996 (CLU); Vallata sotto Colle Marcione, Civita,CS, 800 m, L. Bernardo, 30.V.1995 (CLU); S. Demetrio Corone, CS, D. Puntillo, 26.V.1994 (CLU); Piano di Marco, ai piedi della Mula, S.Donato di Ninea, CS, 1220 m, L. Bernardo, G. Cesca, 21.V.1993 (CLU); Monte Sellaro, presso il Bifurto, Cerchiara di Calabria, CS, 950 m, L. Bernardo, l.VII.1995 (CLU); Monte Basilica, Bocchigliero, Sila Greca, CS, 900 m, L. Bernardo, G. Cesca, 18.V.1996 (CLU); La Mula, S.Donato di Ninea, CS, 1935 m, D. Puntillo, 9.V.1983 (CLU); Monte Camara, S.Paolo Albanese,PZ, L. Bernardo, 26.V.1995 (CLU); San Giovanni in Fiore reg. Pirillo, 1000 m, G. Lopez, 10.V.1910 (Fl); Bosco di Rosamo, Pasquale, IV.1850 (Fl); Bosco di Rosamo, N. A. Pedicino, s.d. (RO).

4. Paeonia morisii Cesca, Bernardo et N.G.Passal., Webbia 56(2): 229 (2001). (=) P. corallina var. pubescens Moris, Fl. Sardoa (1837). (=) P. corallina var. pubescens f. hypoleuca Briq., Prodr. Fl. Corse, 1910. (-) P. mascula subsp. russoi auct. fl ita!. p.p. (-) P. russi auct. fl. sarda Protologue: Planta perennis, erbacea, simplex, 25 - 60 em alta. Caulis erectus, teres, subangulatus, glaber, paullo vd omnino purpurascens. Folia 4- 7, alterna, inferiora 18- 30 x 16- 28 em, bit­ ernata, segmentis (8-) 9 (-10) (raro 7- 20), summa gradatim reducta dein segmentis 3 (-1); fo­ liola coriacea, dliptico-ovata, 3 - 12 x 2 - 9 em, apiculata, basi inequilatera, breviter angustata, supra Iucida, glabra, obscure viridia, subtus glauca, sparse pilosa usque lanosa, pills longis crispatis; petiolis supra vd ambo latere purpurascentibus, superne saepe reductis. Flos plerumque unicus (raro 2 - 3), terminalis; calix zygomorphus, in fructu reflexus, sepalis 3, viridibus saepe purpurascentibus, extra pubescentibus; petala (5-) 6- 8 (-9), obovata, purpu­ rea, rosea vd albido- variegata (raro). Folliculi 3 -5 (raro 2- 7), 2 - 3 x 0,7 - 1,5 em, irregulares, e medio ad apicem angustati, in stigmatem stylo tenui suffultum desinentes, suberecti, matu­ ritate leviter reclinati, virides vd purpurascentes, tomento lutescente tecti. DIAGNOSIS - a P. mascula (L.) Mill. et P. russoiBiv. pills crispatis, foliis coriaceis et follicolis maturis minori­ bus superne angustioribus differt; a P. coriacea Boiss.et P. cambessedesii (Willk.) Willk. fol­ Downloaded by [Università di Pisa] at 02:39 05 November 2015 liculis glabris differt. ETYMOLOGIA- Species Magistro G. G. Moris (1796- 1869), florae sardae perspicaci investigatori, grata memoria dicata. Type: Holotype: Monte D' Iscudu (Gennargentu, Nuoro, Sardegna), gariga a Genista corsica e Rosa sera/ini,l300 m, su calcare, L. Bernardo et N.G. Passalacqua, 22.VI.1996 (CLU). Isotypes: Fl, PAL, SS. Description: Plant 25-60 em (rardy less), the colour may vary from green and becoming purplish only in the lower parts (rardy completdy green) to totally purplish. Lower leaves biternate, with 9leaflets (rardy 7-20); upper leaves progressivdy reduced till3 (1); leaflets elliptic to ovate, 3-12 x 2-9long, acuminate, usually asymmetrical and decurrent at the base; upper face deep green, glabrous and bright; lower face blue-green, more or less covered with long rip­ pled hair. Flower pinkish mauve or white-striped, falling precociously. Carpds (2-) 4-6, to­ mentose with white to whitish thick, short hairs. Follicles, irregularly shaped, tapering in the upper half and ending with the stigma carried by a thin style, sub-erect, curved backward when mature, from green to red and covered with yellowish tomentum. THE GENUS PAEONIA L. IN ITALY 263

illustrations: Moris G.G., 1837: Flora Sardoa, 1: 64, t. 4; Anne Maury, sub. P. mascula (L.) Mill. subsp. russoi (Biv.) Cullen & Heywood, in: Camarda I., Corrias B., Diana S., Valsecchi F., 1992: Piante di Sardegna, Ed. Chiarella, Sassari;Jordan A, Fourreau}. (1903; tav. 322). Chromosome number: 2n=10 Ecology: it grows in different kind of woods (more or less clear) but even in meadowlands, in stony ground and among bushes above 400 m. It can be found on every type of geological substrate (on calcareous or siliceous rocks, sedimentary and magmatic, metamorphic or not), and it grows also on a very poor soil, even if it prefers deep, rich and wet soils. General distrubution: Sardinia (fig. 19) and Corsica; the Sicilian population (M. Iblei) needs fur­ ther investigation.

Selected specimens seen: Sardinia: Gennargentu verso Fonni, 800 m, B. DC., 15.V.1984 (VER); Sardinia pro­ vincia di Sassari: in silva domanialis di Bultei, loco fiorentino dicto, copiosa, alt. 600 m, solo granitico, A. Fiori et S. Tiana, 24.V.1912 (FI, PAD, RO, PI); In silvis montanis ex Sardinia, Moris, 1826 (BOLO); Mon­ tagna di Ala dei Sardi, Martelli, 13.14.VI.1898 (PI); Insula Sardinia, Reg centr. or. in Monte Oliena, C. Forsyth, 12.V.1884 (PI); In pratis montanis editis prope Pulam Sardinia, U. I. Miiller, IV (FI); Gennar­ gentu a Girgini, Martelli, 2.VI.1896 (FI); In insula Tavolara rupestribus calcareis rocce della Madonna, 25.V.1885 (FI); Italia, Sardaigne, prov. Nuoro, Gennargentu, au dessu du Rifugio Baraum Spina, forme de magnifiques touffes, 1570-1700 m, A. Chaipin, M. Dittrichi, 27.V.1983 (FI); Arrondissement de Tern­ pia. Monte Limbardo, maquis decouverts sur granit, E. Reverchon, 16.V et 14 VII.1882 (FI); Mti d'Aritzo, VII.1859 (CAG); Monti Urtigu (CAG); Gennargentu Arcu D'Iscudu, Martinoli, 2l.IV. 1948 (CAG); Cala Gonone, G.Miliaet I. Camarda,22.V.1975 (CAG); Desulo, SuAu, GoiA., IV.1983 (CAG), Saramini (Vil­ lagrande Strisaili), Floreddu, 18.V.1983 (CAG); Mte Linas, Chiappini et Angiolino, 1984 (CAG); Acquaf­ rida, M. G. Biagini, 20.IV.1986 (CAG); Mte Tonneri-Semi, M. Ballero, VI.1987 (CAG); Punta Sebbera, Domus De Maria (Cagliari), A. R. Collu, IV.1988 (CAG); Talasaggia, Tonara (Nuoro), C. Fogu, V.1990 (CAG); Bosco lungo Ia strada nel vallone Rio Aratu (Fonni, Nuoro, Sardegna), Bosco di leccio e roverella, 1100 m, L. Bernardo et N. G. Passalacqua, 22 .VI.1996 (CLU); Serra Edele-Bosco di Ghivine (Cala Gonone, Nuoro, Sardegna), lecceta aperta, in radure, 540 m, su calcare, L. Bernardo et N.G. Passalacqua, 23.VI.1996 (CLU); Monte Bruncu Spina (Gennargentu, Nuoro, Sardegna), piccolo bosco ripario ad Al­ nus glutinosa, 1600 m, su calcare, L. Bernardo et N.G. Passalacqua, 22.VI.1996 (CLU); Monte Albo (Lula, Nuoro, Sardegna), bosco di leccio, 1000 m ca., su calcare, L. Bernardo et N.G. Passalacqua, 21.VI.1996 (CLU).; Monte D' lscudu (Gennargentu, Nuoro, Sardegna), gariga a Genista corsica e Rosa sera/ini,1300 m, su calcare, L. Bernardo et N. G. Passalacqua, 22.VI.1996 (CLU). Sicily: Iblei, valle Cava Grande, bosco a leccio, 550 m, L. Bernardo et N.G. Passalacqua, 26.V.1996 (CLU).

TABLE 8-Taxonomic setting of the genus Paeonia L. in Italy

Genus: Paeonia L. Downloaded by [Università di Pisa] at 02:39 05 November 2015 Sub gen. Paeonia Section: Paeonia Subsection: Paeonia Paeonia peregrina Mill. Paeonia officina/is L. subsp. officina/is subsp. huthii Soldano subsp. italica subsp. nov. subsp. banatica (Rochel) Soo Subsection: Foliolatae Stem 1946 Paeonia morisii Cesca, Bernardo et N.G. Passal. Paeonia mascula (L.) Mill. subsp. mascula subsp. mascula var. russoi (Biv.) comb. nov 264 N.G. PASSALACQUA, L. BERNARDO Downloaded by [Università di Pisa] at 02:39 05 November 2015 THE GENUS PAEONIA L. IN ITALY 265

K£y FOR IDENTIFICATION OF THE ITALIAN TAXA

la Underground rootstock cylindrical or fusiform; leaf segments entire, ovate to elliptic, varying from (7) 9 to 17 (19) ...... 2 lb Underground rootstock pyriform, fusiform or subglobose; leaf segments mainly partite, vary- ing from (7) 13 to 41 (60) ...... 4 2a Leaves coriaceous with (7-) 9 (-12) segments; upper face of leaves bright green, lower face from slightly hairy to villose-floccose with long curled hair; follicle (2-) 3-5 (-8), becoming thinner gradually beginning from the median part toward the base and the apex a~ matu~it1...... Paeorua mortsu 2b Leaves never coriaceous with (9-) 13-17 segments; upper face ofleaves deep green, lower face glabrous or slightly hairy with long straight hairs; fruits (1-) 3 (-5), oblongs, abruptly truncates at apex and base ...... 3 3a Leaf stalk and lower surface of lamina glabrous or rarely with some hair scattered on rib; co­ rolla violet ...... Paeonia mascula subsp. mascula 3b Leaf stalk and lower surface of lamina hairy to villose; corolla from violet to white or rose stripped ...... Paeonia mascula subsp. mascula var. russoi 4a Corolla in a closed cup, deep red; leaf segments often with (2) 3 coarse teeth .at the a~ex ...... Paeonta peregnna 4b Corolla soon an open cup, violet to red and pink; leaf segments partite and lobed, but rarely with teeth at the apex...... 5 5a Lower surface of leaves and stalk villose-floccose with hairs opaque flattened...... 6 5b Lower surface of leaves and stalk hairy but never villose-floccose; hairs bright cylindrical .. 7 6a Lower leaf with elliptic segments, the largest 36-80xl0-15 mm, generally with many lobes ...... Paeonia officinalis subsp. huthii 6b Lower leaf with linear-lanceolate segments, the biggest of 60-110 x 12-23 mm, generally with few lobes...... Paeonia officinalis subsp. italica 7a Last segments of lower leaf nearly exclusively partite; lower surface subglabrous; plants living in hilly deciduous oak woods...... Paeonia officinalis subsp. banatica 7b Lower leaf generally with 3-11 entire last segments; lower surface more or less hairy; plants liv- ing on mountain rocks and screes ...... Paeonia officinalis subsp. officinalis

AcKNOWLEDGEMENTS We wish to thank Directors and Keepers of mentioned Herbaria who have kindly provided us with the exiccata of studied species; particularly, we wish to thank Dr. Charlie Jarvis (BM), for all information and suggestion, and Dr. Annalisa Santangelo (NA) and all staff of Herbarium Cent­ rale ltalicum (FI), for the assistance.

Downloaded by [Università di Pisa] at 02:39 05 November 2015 Moreover, we are grateful to Dr. Sandro Bonacquisti, Dr. Fabio Conti, Dr. Walter Good, Dr. Michele Codogno, Prof. Giovanni Spampinato, for the useful help and for information on field investigation, and Dr. Lorenzo Peruzzi, for the caryological data. We are also grateful to Dr. Carsten Burkhart for the useful information found on the Web site http://www.paeon.de. The financial support by MIUR (Contribution ex 60 %) is gratefully acknowledged.

BmuOGRAPHY

AKERoYDJ. R.,1993.- Paeonia L. -In: TuTIN T.G. et al. (2nd ed.), Flora Europea 1:292-294. Cambridge, Uni- versity Press. ANDERSON G.,1818. -A Monograph of the Genus Paeonia. Trans. Linn. Soc. London 12:248-290. BARBER H. N.,1941.- Evolution of the genus Paeonia. Nature 148:227-228. BERNARDO L., BRUNO F., CESCA G., PASSALACQUA N. G., 1995. - Specie critiche della flora calabra: problemi tassonomici e nuove segnalazioni. Boll. Soc. Sard. Sci. Nat. 30:435-445. 266 N.G. PASSALACQUA, L. BERNARDO

BERTOLONI A.,1844.- Flora Italica. 5: 392-395. Bonomia. BICKNELL C., 1896. - Paeonia L. Fl. Bordighera 8: 10 BIVONA-BERNARDI A.,1816.- Paeonia russi. Stirp. Rar. Sicilia 4: 12. BURNAT E., 1892.- Paeonia L. Fl. Alpes Maritim. 1: 54 CAESALPINO A., 1583.- De Plantis Libri. 588. Firenze. CARUEL T., 1860.- Paeonia L. Pr. Fl. Tosc.: 19-20. CESCA G., BERNARDO L. & PASSALACQUA N.G., 2001.- Paeonia morisii sp. nov. (Paeoniaceae), a new species /rom Sardinia. Webbia 56(2): 229-240. CIFERRI R. & GIACOMINI V., 1954.- Nomenclator Fl. Ital. Pars altera Dicotyledones. Fasc. 1: 308-309, Ticini ex Typis C. Busca. CuLLEN J. & HEYWOOD V. H., 1964a.- Paeonia L.- In: TUTIN T.G. eta!. (Eds.), Flora Europea 1: 242-244. Cambridge, University Press. CULLEN}. & HEYWOOD V. H., 1964b.- Notes on the European species o/Paeonia. Feddes Repert. Spec. Nov. Regni Veg. 69: 32-35. ELENA· RossELLO J .A., GoNzALEs-ZAPATERO A. & NAvARRO-ANDRES F., 1987. - Numeros cromosomaticos de plantas occidentales: 412-413. AnnalesJardin Botanico de Madrid 43:417-420. FEDOROV A., 1969. -Chromosome numbers offlowering plants. Reprint by Koeltz. Sci. Pub!. Koenigstein. 479- 480. FioRI A., 1924.- Nuova Flora Analitica d'Italia. 1: 689-690. Firenze. FIORI A. & PAOLETTI G., 1898.- Paeonia. In- Flora Analitica d'Italia. 1:526-527. Padova. FucHs L., 1542.- De Historia stirpium Commentarii Insignes. 202. Basel. GESNER K., 1561.- Horti germaniae: 270. Strasbourg. GoULANDRIS N., GoULIMIS C.N. & STEARN W. T., 1968.- Wildflowers a/Greece. The Goulandris Bot. Mu­ seum, Kifissia. GREGORY W.C., 1941.- Phylogenetic and cytological studies in the R.anunculaceae fuss. Trans. Amer. Phil. Soc. 31(5): 443-520 GREUTER W., BURDET H.M. & LoNG G., (Eds.) 1989.- Med-Checklist 4:266-269. Geneve. GREUTER W et a!. (Eds.) 2000. - International Code o/ Botanical Nomenclature (Saint Lous Code) Regnum Vegetable. GussoNE G., 1843.- Paeonia coral/ina var. flavescens (C. Pres/). Fl. Sicul. Syn. 2: 26. HALDA, J .]., 1997. -Systematic treatment of the genus Paeonia L. with some nomenclatoric changes. Acta Mus. Richnov. Sect. Natur, Sect. Nat. 4(2): 25-32. HALDA, J.J., 1998.- Notes on the observations upon the structure of the Paeonia seeds, fruit and roots. Acta Mus. Richnov. Sect. Natur, Sect. Nat. 5(1): 4. HALLER A. VoN. 1742.- Enumeratio Methodica Stirpium Helvetiae indigenarum. 1: 311. Gottingae. Hum E., 1891.- Monographie der Gattung Paeonia. Bot. Jahrb. Syst. 14: 258-276. JoRDAN C.T.A. & FoURREAu J.P., 1903.- Icon. Fl. Bur. 2:37-38, tt. 318-323. ]oRDANOV D., 1970.- Paeonia L.- ln:]ORDANOV D. & KozUHAROV S., Fl. Reipubl. Popularis Bulg. 4:216-223. Bulg. Acad. Sci., Sofia. KoEvA]. & SARCOVA S., 1997.- Karyological study of three species o/Paeonia (Paeoniaceae) in Bulgaria. Boc- conea 5: 553-556. LINNAEUS C., 1737.- Paeonia. Hortus Cli/fortianus.: 211-212. Amstelaedami. LINNAEUS C., 1748.- Paeonia. Hortus Upsaliensis: 149. Stockholm. LINNAEUS C., 1753.- Paeonia. Species Plantarum. 1:530. Holmiae.

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RETZIUs A.]., 1783.- Observ. Bot. 3: 34. Siegfried Lebrecht, Leipzig. RacHEL A., 1828.- Plantae Banatus rariores, iconibus et descriptionibus illustratae: 48, tt. 11-12. SANG T., CRAWFORD D.]. & STUESSY T. F.,1995.- Documentation of reticulate evolution in peonies (Paeonia) using internal transcribed spacer sequences of nuclear ribosomal DNA: implications /or biogeography and concerted evolution. Proc. Nat. Acad. Sci. USA 92: 6813-6817. SANG T., CRAWFORD D. J. & STUESSY T. F., 1997. - Chloroplast DNA Phylogeny, reticulate evolution, and bio­ geography of Paeonia (Paeoniaceae). Amer.]. Bot. 84(8): 1120-1136. ScHMITI E., 1997.- Les Pivoines. Etude systematique du genre Paeonia L. Pl. Mont. 181: 176-186. ScHMITT E., 1998.- Les Pivoines. Etude systematique du genre Paeonia L. Pl. Mont. 186: 372-382; 188: 466- 473. ScHMITI E., 1999.- Les Pivoines. Etude systematique du genre Paeonia L. Pl. Mont. 189:501-508. ScHMITT E., 2003.- Typification of the Linnean names of the genus Paeonia L. Candollea 58: 183-188. ScHWARZACHER-ROBINSON T.1986. -Meiosis, SC-/ormation and karyotype structure in diploid Paeonia tenui/o- lia and tetraploid P.officinalis. PI. Syst. Evol. 154: 259-274. SoLDANO A., 1993.- Nuovi dati nomenclaturali su piante della/lora italiana e medite"anea. Atti Soc. Ital. Sci. Nat. Mus. Civ. Stor. Nat. Milano 133(10): 114. So6 R., 1945.- Noveny/Oldrajz: 146. So6 R., 1960.- What is Paeonia banatica Roche!? Acta Bot. Acad. Sci. Hung. 6(1-2): 139-141. SorovA M., 1971. -Cytological study oftwo Paeonia species /rom Macedonia. Fragm. Bale. Mus. Macedon. Sci. Nat. 8(16):137-142. SoWERBY & SMITII, 1806.- Paeonia. Engl. Bot. 22, T. 1513. STAPF 0. 1918.- Paeonia peregrina. Curtis's Botanical Magazine 144: t. 8742. STEARN W. T. & DAVIS P.H., 1984.- Peonies of Greece. The Goulandris Natural History Museum, Kifissia. STERN F.C, 1944.- Geographical distribution of the genus Paeonia. Proc. Linn. Soc. London 155:76-79. STERN F.C.,1946.- A study of the Genus Paeonia. The Royal Horticultural Society, London. TENORE M., 1811-1815.- Fl. Napol. 1: 300-301, 337-338. TENORE M., 1830. - Succinta relazione del viaggio /atto in Abruzzo ed in alcune parti dello Stato Ponti/icio dal Cavaliere Tenore nell'esta de/1929: 71. Stamperia Societa Filomatica, Napoli. TENORE M., 1831. - Syll. Fl. Neap.: 261. TENORE M., 1835.- Syll. Fl. Neap. App. IV: 21. TzANOUDAKIS D.,1983. - Karyotypes offour wild Paeonia species /rom Greece. Nord. J. Bot. 3: 307-318. UsPENSKAJA M.S. & SOLOWJEWA L. W., 1991. - Paeonia peregrina Mill. v SSSR i ego tipi/ikazija. Bulletin Mosk. O-va Ispytatelej Prirody otd. Bioi. 3: 131-13 3.

Summary

The taxa of the genus Paeonia in Italy need a thorough rearrangement of their taxonomy and geographical distributions. The genus Paeonia has always shown bearing many controversies since it was establishment by Linnaeus (1753), who perceived the close relationships between the two units he described. Downloaded by [Università di Pisa] at 02:39 05 November 2015 Several authors have successively amended the genus. At the present, the Italian peonies are rec­ ognized essentially by the morphological features of their leaves; but usually the shape, the size and the number of leaf-segments is variable in the same population. On account of this, there were, in the past, a lot of uncertainties and some mistakes in their classification. The present study is based on both living populations and the dried collections of several her­ baria; field investigation through continental Italy, Sardinia and Sicily was carried out by bio­ metrical analyses and was supported by laboratory analysis and bibliographical investigations. As a result, we have identified: Paeonia peregrina Mill. s.s., a Balcanian- Turanian element with a separate area in Southern Italy, a few populations in Basilicata and Calabria. Paeonia officina/is s.l., a temperate element mainly in open habitat, with four taxa across the Alpine Chain and on the North and Middle Apennines: P. officina/is L. subsp. officina/is, from Piedmont to Friuli Venezia Giulia (except Trieste Karst), Apuan Alps and Northern Apennines; P. officina/is L. subsp. huthii Soldano, with few populations in Liguria; P. officina/is L. subsp. italica subsp. nov., on the Middle Appennines from Marche to Latium and Abruzzo; P. officina/is L. subsp. banatica (Rachel) So6, on the Trieste Karst. 268 N.G. PASSALACQUA, L. BERNARDO Paeonia mascula s.l., a eurimediterranean element of woody places, with two infraspecific taxa occurring in continental Italy, from Lazio to Calabria, in Sardinia and in Sicily: P. mascula (L.) Mill. subsp. mascula var. mascula, from Latium to Basilicata; Paeonia mascula (L.) Mill. subsp. mascula var. russoi (Biv.) comb. nov., occurring in Sicily and Calabria. Paeonia morisii Cesca, Bernardo & N.G. Passal., in Sardinia. Downloaded by [Università di Pisa] at 02:39 05 November 2015