1 PHYTOLOGIA BALCANICA 14 (1): 45 49, Sofia, Typification and taxonomic status of Dianthus balbisii subsp. knappii comb. et stat. nov. (Caryophyllaceae) Lorenzo Peruzzi 1,2 & Dimitar Uzunov 2 1 Dipartimento di Biologia, Orto Botanico & Museo Botanico, Unità di Botanica Generale e Botanica Sistematica, Università di Pisa, via Luca Ghini 5, Pisa, Italy, 2 Museo di Storia Naturale della Calabria ed Orto Botanico, Università della Calabria, Arcavacata di Rende (CS), Italy. Received: July 31, 2007 Accepted: September 27, 2007 Abstract. Morphological observations of Dianthus knappii (here typified) from Herzegovina are presented and critically compared with those of the closely related D. balbisii Ser. s. l.. Dianthus balbisii subsp. knappii comb. et stat. nov. is proposed. Key words: Balkans, Dianthus, Italy, taxonomy, typification Introduction Dianthus knappii (Pant.) Borbás (Caryophyllaceae) is a yellow-flowered carnation species, endemic to Herzegovina and SW Montenegro (Jalas & Suominen 1986; Tutin & Walters 1993; Šilič 2002), included in the Red Data Book of the Flora of Bosnia and Herzegovina, and assigned to the category Vulnerable (Šilič 1996). When the authors visited Bosnia-Herzegovina in 2003, they were surprised by the resemblance between D. knappii and the S Italian endemic D. guliae Janka (for the nomenclature of the latter species, see Peruzzi & Gargano 2006), a critical taxon included in the D. balbisii Ser. aggr., including also D. liburnicus Bartl. ex Bartl. & Wendl. and D. liburnicus var. lucanus Lacaita widespread from SE France to peninsular Italy and the Adriatic part (Liburnia, in classical times) of the Balkan Peninsula (Greuter & al. 1984; Tutin & Walters 1993). Morphological studies aiming to clarify the taxonomic relationships among these taxa were carried out and the name D. knappi was typified. Material and methods Herbarium specimens of D. guliae were used preserved in Florence (FI); the Botanic Garden of Calabria University (CLU); Cluj-Napoca, Romania (CL); Budapest, Hungary (BP), and the British Museum, UK (BM), along with specimens of D. balbisii s.l. conserved in CLU and BM, and specimens of D. knappii (Pant.) Borbàs conserved in Sarajevo, Bosnia and Herzegovina (SARA) and Budapest (BP). Specimina visa: Dianthus balbisii s.l.: Croatia, Quarnero Istria. In herbidis prope Castuam et Voloscam ad sinum flanaticum; solo calc m, s.d., Pichler (BM, topotype of D. liburnicus); Italy, Liguria Sestri Levante (Riviera di Genova), in rup. marit. siliceis, flower deli cate pink, , Lacaita (BM, topotype of D. balbisii); Basilicata Foresta di Gallipoli-Cognato (Basilicata), in collibus herbosis, ca. 600 m, petala rubina, , Lacaita 2422 (BM, type material of D. liburnicus var. lucanus Lacaita); ibidem, forma luxurians, Lacaita (BM, type material of D. liburnicus var. lucanus Lacaita);
2 46 Peruzzi, L. & Uzunov, D. Dianthus balbisii subsp. knappii comb. et stat. nova Calabria Rossano, in dumetis montanis, solo siliceo, ca. 700 m, , Lacaita (BM); Arcavacata di rende (Cosenza) nei pressi dello svincolo per l Università, 250 m, V/2002, Gargano (CLU, no. 2017); primo svincolo Università (Arcavacata di Rende, Cosenza), UTM XD 05.56, 220 m, , Bernardo (CLU, no. 2018); strada per Caloveto (Cosenza), ca. 500 m, 10/ VI/1993, Bernardo & Gangale (CLU, no. 2019); Valle del Fiume Lese nei pressi del ponte sul Lese lungo la strada per Verzino (Crotone), 200 m, , Bernardo, Gangale et Naccarato (CLU, no. 2020); Acque Termali Bruciarello presso lo svincolo sulla S.S. 107 per Cotronei (prov. Crotone, Calabria), 130 m, 29/V/2000, Bernardo (CLU, no. 2588); Cerenzia, nei pressi dello svincolo della SS 107 (Prov. di Crotone, Calabria), 587 m, , Bernardo (CLU, no. 2589); Caloveto, presso la cava di Peppe (Prov. Cosenza, Calabria), affioramento rosso ammonitico, 550 m, UTM 33S XD 51.73, 22/V/1999, Bernardo (CLU, no. 7454). Dianthus guliae: Italy, Tuscany Cetona sopra Sferracavalli, , Corti (FI); Cetona, a Belvedere, , Negri et Corradi (FI); Campania In campestribus dumosis inter Eboli et fl. Sele non procul a Neapoli, , Janka (BP); Lucania prov. Salerno: secus viam inter praga Sacco et Teggiano, in pascuis montanis loco dicto Timpa Muletta, 1050 m, solo schistoso, , Lacaita (FI); Timpa Muletta tra Sacco e Teggiano (Salerno) in pascuis montanis ca m, , Lacaita (BM, FI); In pascuis La Pecora, ca m, inter Sacco et Teggiano (Lucania) ubi copiosissima, flores supra lutei, subtus rubiginosi, , Lacaita (BM); Monte Alburno, s.d., De Philippis (FI); Campania, Vallo di Diano, , Bernardo (CLU, no. 2053); Sassano, Parco del Cilento provincia di Salerno Campania, , Musacchio (CLU, no. 2549, 2550, 2551); Calabria Calabria, Settore orientale del Massiccio del Pollino, sul versante settentrionale del Monte Sellaro, presso il Bifurto (Cerchiara di Calabria, CS), UTM XE 16.13, flysch argillitico calcareo a contatto con roccia calcarea ca. 950 m esp. NE, , Bernardo (FI; CLU, no. 2050, 2051, 2052); Dianthus knappii: Hercegovina Gatačko Pole, Faz lagića/kula, 970 m, , Maly (SARA, no ); Pagum Bijele Rudine prope Bileće, , Živković (SARA, no ); Supra pagum Korita prope Koritska Jama, , Djuran (SARA, no ); Steppen na Gacko, , Maly (SARA, no ); Staza prope pagum Gacko, , Šilič (SARA, no ); Trebeni in dumetis montis Gliva copiose, , Vandas (SARA, no ); SW Montenegro Auf trockenen steppen bei Miljkovac, 1869, Knapp (BP). For SEM studies, the material was directly observed and photographed at 20 kv. The analysis was carried out on very small leaf samples taken from the following herbarium specimens: CLU, no (D. balbisii); CLU, no (D. guliae); SARA, no (D. knappii). Quantitative and qualitative features of D. guliae, D. balbisii and D. knappii derive from observations carried out on herbarium specimens. Results Examining the morphological characters reported in Table 1, there have been wide overlaps among D. guliae, D. balbisii and D. knappii in respect to all quantitatively compared features. The bract is distinctly longer than calyx in some individuals of D. guliae, while in D. balbisii and D. knappii the bract seldom overtops the calyx. D. knappii is always scabrid on the surface of leaves and bracts, on the average has a grea ter number of flowers, and is with a smaller calyx width than in D. guliae and D. balbisii. Scabridity of the leaves in D. knappii is due to the presence of bi-celled, somewhat pyramidal tubercles, ca. 20 μm long, which occur grossly arranged in lines. Those tubercles are absent in D. balbisii and D. guliae (Fig. 1). On the other hand, the size of stomata in D. balbisii and D. knappii completely overlaps, while D. guliae tends to be distinct, thanks to longer and larger stomata (Fig. 2). Table 1. Morphological characters of D. balbisii, D. guliae and D. knappii. Numerical data are expressed in percentiles, with the extreme values in brackets. D. guliae D. balbisii D. knappii foliar surface glabrous glabrous scabrid bract almost equalling to longer than flowers a little shorter to slightly a little shorter to slightly longer than flowers longer than flowers width of leaves (mm) (4) (1.5) (5.5) (2) (4) length of calyx (mm) (16) (21) (16) (21) (17) width of calyx (mm) (4) (6) (3) (6) (5) length of petal limbs (mm) (4) (7.5) (5) (11) 5 7 number of flowers (1) 2 6 (8) (1) (15) number of inflorescences 1 1(2) 1 (2) corolla colour citrine to orange yellow pink to red purplish citrine
3 Phytol. Balcan. 14(1) Sofia Fig. 1. SEM microphotographs of leaf surface of: Dianthus balbisii: A, general view, B, particular; D. guliae: C, general view, D, particular; D. knappii: E, general view, F, particular.
4 48 Peruzzi, L. & Uzunov, D. Dianthus balbisii subsp. knappii comb. et stat. nova 15 Width Stomata dimentions A Width 10 Length balbisii B guliae knappii Length balbisii C guliae balbisii guliae knappii knappii Fig. 2. Comparison of stomata dimensions between D. balbisii, D. guliae and D. knappii: A, scatter plot of length over width; B, boxplots illustrating the var iability of stomata width; C, boxplots illustrating the variability of stomata lenght. In the figure, the outlined central box depicts the middle 50 % of the data extending from upper to lower quartile; the horizontal bar is on the median. The ends of the vertical lines (or "whiskers") indicate the minimum and maximum data values, unless outliers are present, in which case the whis kers extend maximum 1.5 times to the interquartile range. Superimposed grey areas indicate confidence interval bounds around its median (median ± 1.58 times to the interquartile range). Circles indicate outliers, unless extreme outliers are present, in which case the circles extend maximum three times to the interquartile range and the extreme outliers are indicated with asterisks. All measurements are in μm. Discussion The three studied units appear closely related to each other, and it is very difficult to define clearly distinctive characters among them, other than the corolla colour (D. balbisii vs. D. guliae/d. knappii), stomata dimensions (D. guliae vs. D. balbisii/d. knappii) and scabridity (D. knappii vs. D. balbisii/d. guliae). As far as karyology is concerned, D. knappii was repeatedly reported with 2n = 30 (Andersson- Kottö & Gairdner 1931; Rohweder 1934; Gentscheff 1937; Carolin 1957). D. balbisii s.s. also has 2n = 30 chromosomes (Loon & Jong 1978, sub D. ferrugineus Mill.) counted on plants coming from Alassio (Liguria, N Italy) and the same number was found in the S Italian D. guliae from Calabria by Peruzzi (2003). Indeed, according to Fedorov (1969), the chromosome number 2n = 30 is the lowest and the most widespread within the genus Dianthus L., which therefore should be characterized by a basic number x = 15. Conclusions Thanks to the morphological and karyological affinities, it seems appropriate to consider D. knappii as a subspecies of D. balbisii, because of the tendency of the two taxa to occur in distinct areas, even if with few overlaps. Irrespective that D. guliae is not directly object of this study, it probably deserves a taxonomical status as subsp. of D. balbisii and this will be a matter of further investigation. Dianthus balbisii Ser. in DC. [Prodr. 1: 356 (1824)] subsp. knappii (Pant.) Peruzzi & Uzunov comb. et stat. nov. D. liburnicus var. knappii Pant., Österr. Bot. Z. 23: 4 (1873) D. knappii (Pant.) Borbás, Math. Természettud Közlem 13: 196 (1877) Lectotypus (here designated): Auf trockenen steppen bei Miljkovac, 1869, Knapp (BP!, Fig. 3).
5 Phytol. Balcan. 14(1) Sofia The specimen traced at BP is eligible as a lectotype, since it is part of the material originally collected by Knapp used four years later by the author for variety description. The selected type is clearly referable to the systematic unit to which the concerned name currently applies. Acknowledgements. The authors are very grateful to the curators of the herbaria in Florence (FI) for their help; to Dr Abadzic Sabaheta, Sarajevo (SARA), Dr Lajos Somlyay, Budapest, Hungary (BP), and Dr Ana-Maria Csergo, Cluj-Napoca, Romania (CL) for the information and images of D. guliae and D. liburnicus var. knappii; to Dr Nicodemo Giuseppe Passalaqua (University of Calabria) for the images of Dianthus specimens conserved at BM; and to Prof. Franco Russo (University of Calabria) for the SEM study. References Andersson-Kottö, I. & Gairdner, A.E Interspecific crosses in the genus Dianthus. Genetica, 13(1-2): Carolin, R.C Cytological and hybridization studies into the genus Dianthus. New Phytol., 56(1): Fedorov, A.A. (ed.) Chromosome Numbers of Flowering Plants. Nauka, Leningrad (in Russian). Gentscheff, G Experimental and caryological investigation of the relationship among the species of the genus Dianthus L. DSc Thesis. Agric. & Forest. Fac., Sofia Univ., Sofia (in Bulgarian, unpubl.). Greuter, W., Burdet, H.M. & Long, G. (eds) Med-Checklist. A Critical Inventory of Vascular Plants of the Circum-Mediterranean Countries. Vol. 1 (Pteridophyta, Gymnospremae, Dicotyledones: Acanthaceae Cneoraceae). Conservatoire et Jardin Botanique, Med-Checklist trust of OPTIMA, Genève & Berlin. Jalas, J. & Suominen, J Atlas Flora Europaeae. Distribution of Vascular Plants in Europe. Vol. 7. Caryophyllaceae (Silenoideae). The Committee for Mapping the Flora of Europe & Societas Biologica Fennica Vanamo, Helsinki. Loon van, J.C. & Jong de, H Reports. In: Löve, A., IOPB chromosome number reports LIX. Taxon, 27(1): Peruzzi, L Biosistematica e fitogeografia di specie critiche della flora calabra. Ph.D. Thesis. Univ. Calabria (in Italian, unpubl.). Fig. 3. Lectotype of the name D. liburnicus var. knappii Pant., conserved in BP. Peruzzi, L. & Gargano, D Dianthus ferrugineus Mill. vs. D. guliae Janka: nomenclatural considerations on the Italian yellow carnation. Taxon, 55(3): Rohweder, H Beiträge zur Systematik und Phylogenie des Genus Dianthus. Bot. Jahrb., 66(3): Šilič, Č List of the plant species (Pteridophyta and Spermatophyta) for the Red Book of Bosnia & Herzegovina. Glasn. Zemaljsk. Muz. Saraevu, 31: (in Serbo-Croation). Šilič, Č Endemic and rare plants in Blidinje Natural Park. Matica Hrvatska, Ogranak čitluk, Sarajevo (in Serbo- Croation). Tutin, T.G. & Walters, S.M Dianthus L. In: Tutin, T.G. & al. (eds), Flora Europaea. Ed. 2. Vol. 1, pp Cambridge Univ. Press, Cambridge. 7 Phytol. Balcan. 14(1) 2008
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