NOMENCLATURAL AND TAXONOMIC NOTES ON THE FLORA OF SERBIA AND THE BALKAN PENINSULA. I. CARYOPHYLLACEAE

The nomenclatural analysis included 12 autochthonous and mostly endemic species from the genera Atocion Adans., Cerastium L., Heliosperma (Rchb.) Rchb., and Silene L. (Caryophyllaceae). In conformity with ICBN, 12 new infraspecific taxa are described with locus classicus on the territory of Serbia. Also presented are 24 new nomenclatural combinations, including several for endemic taxa not present in Serbia. The nomenclature of certain taxa is supplemented by a short taxonomic-chorological review. Results of these studies will be incorporated in the next volume of “Flora of Serbia”.


INTRODUCTION
The ten volume edition "Flora of Serbia" (J o s i f o v i ć , 1970-1976; 1977; S a r i ć and D i k l i ć , 1986) is a capital-value work of Serbian botany, filling a void that existed for almost more than a century in studies on the flora of this part of Balkan Peninsula, more precisely since the publishing of Pančić's "Flora of the Principality of Serbia" and its "Addendum" (Pančić, 1874;1884)."Flora of Serbia" was finished in an unusually short time for this type of large-scale synthetic work.It united mostly fresh knowledge on taxonomy, distribution and ecological preferences of taxa that were known since Pančić's time, as well as the many new ones recorded within the boundaries of modern Serbia."Flora of Serbia" initiated intensive floristic and chorological studies of the vascular flora in Serbia during the last 30 years.Since 1986, immense quantities of fresh material have been collected on distribution, taxonomy and ecology of plant taxa; new monographs on genera have been published; new regional floras have been issued for the Balkan countries; and new nomenclatural principles have been accepted in European plant taxonomy, elucidating in a new way the validity and taxonomic position of species and genera.Altogether, conditions are now suitable for starting a new edition of Flora of Serbia, and the Committee for Flora and Vegetation of SASA accepted this challenge.The main intent of the new edition is to remove the deficiencies of the first edition (non-uniform taxonomic principles in taxon studies, incomplete and/or false data on taxon distribution, failure to utilize relevant modern taxonomic literature, failure to check herbarium material, etc.).The first volume of the new edition of Flora of Serbia was published in 1992, and the second one is planned, after a long break, for 2008.It will include the orders Ranales (continuing with fam.Berberidaceae), Papaverales, Urticales, Fagales, Betulales, Juglandales, and Caryophyllales.The manuscript of the second volume of the new edition was prepared ten years ago, but unfortunately the publishing procedure was never initiated.More than 15 years have passed since the first edition, and numerous papers have appeared in the meantime in the European botanical literature, including new nomenclatural and taxonomic principles which should be incorporated in the new, expanded manuscript of the second volume.The main goal of this paper was to, through new interpretation of certain taxonomic challenges, facilitate the use of synonyms in the next volume of the second edition of Flora of Serbia.Certain new taxa and nomenclatural combinations are here mentioned for the first time, and in the mentioned publication they will be individually presented within the completely processed genera and species of the family Caryophyllaceae.This paper deals with certain representatives of the genera Atocion Adans., Cerastium L., Heliosperma (Rchb.)Rchb.and Silene L., which were presented in the previous edition of Flora of Serbia by G a j i ć (1970: 156-176) and S l a v n i ć (1970: 204-240).In order to fully present the infraspecific variability of certain species, we have included new combinations for some endemic subspecies that do not appear on the territory of Serbia but are closely related to taxa analyzed in Flora of Serbia.Basic data on distribution are included for newly described taxa., 2007).There are clear definitions of monophyletic lines that are genetically and morphologically distinct from other genera, primarily from the genus Silene (although until recently they were considered an integral part of that genus).This is particularly true of the genera Atocion, Heliosperma, Eudianthe (Rchb.)Rchb.and Viscaria Bernh.On the other hand, certain taxa which were considered as belonging to separate genera are now added to the genus Silene -for example Silene baccifera (L.) Roth, which used to be known as Cucubalus baccifer L. It was therefore necessary to assign new combinations to certain infraspecific taxa within the genera Atocion and Heliosperma, but also within the genus Silene, if they were previously absent.This was not done only for formal reasons, but also due to the need to, as much as possible, elucidate numerous taxonomic and chorological problems.Several new taxa are therefore described on this occasion.
Additionally, activities of describing and interpreting the very complicated infraspecific variability within the genus Cerastium, started by N i k e t i ć (1999), are continued here, with the accent on separating subspecies within the species C. decalvans Schlosser & Vuk. (subsp. glutinosum and subsp. leontopodium).

MATERIAL AND METHODS
The study of 12 investigated species is based on the examination of available herbarium material (BEO, BEOU, LJU, SARA, UPA, SO, SOM).Acronyms follow Index Herbariorum (Holmgren cronyms follow Index Herbariorum (Holmgren et al., 1990; and http://sciweb.nybg.org/science2/IndexHerbariorum.asp).Synonymization and Synonymization and description of new taxa were done in conformity with ICBN (McNeill et al., 2006) and are based on surveyed herbarium material and published data.

Cerastium decalvans
The polyploid complex of the species C. decalvans Schlosser & Vuk. is in the taxonomic sense perhaps the most complicated within the whole genus.So far there are a great many described infraspecific taxa (N i k e t i ć , 1999).Besides the type subspecies, the only other definitively distinct subspecies mentioned in the literature is subsp.orbelicum from the silicate mountains of Southern Bulgaria and Eastern Macedonia (G r e u t e r et al., 1984; J a l a s , 1993; S t r i d 1997; N i k e t i ć , 1999), while the present paper also includes subsp.glutinosum.
It should be noted that all the names of distinct species and subspecies described by G e o r g i e v (1933; 1935) were very soon moved to the synonym list very soon (B u s c h m a n n , 1938).As she had a very conservative approach, these taxa (as well as the taxa described by other authors) were not accorded any recognized status and were treated as synonyms.This includes even current subspecies, for example subsp.orbelicum.The only infraspecific category recognized by this author was var.leontopodium (Stoj.& Stefanov) Buschm., to which were added Correns's (C.histrio) and Bornmüller's (C.lanigerum var.isophyllum and C. balcanicum var.anisophyllum), but not Georgiev's (C.macedonicum, C. cernjavskii) or Baldacci's (C.grandiflorum var.albanicum) taxa.The main records of differential characters and chorology of Buschmann's taxon leontopodium are acceptable even by present-day taxonomy, as they are supported by karyological data.In contrast to the type tetraploid (neodiploid) species, which is probably distributed only in the Dinarides, this taxon represents an extremely polymorphic polyploid complex in the central and southern part of the species range, and it probably originated by reticular evolution.In the morphological sense, it is characterized by more compact structure, narrower leaves, and characteristic axillary fascicles composed of narrow leaflets.For all these features it should be awarded subspecific status.
In regard to proper naming of the subspecies leontopodium, it should be noted that, according to the rules of recognizing a new taxonomic combination (M c N e i l l et al., 2006: Art.11.2) and results of research, the oldest legitimate name of a taxon on the subspecific level must be chosen.For this taxon, there are several appropriate names listed in the same paper by G e o r g i e v (1935) (Greuter and Burdet, 1982: 37).However, due to inclusion of a multitude of names in this paper, in conformity with ICBN (McNeill et al., 2006: Art. 11.5, 11.6) we chose the name subsp.leontopodium as a derived autonym of C. leontopodium subsp.soskianum Georgiev (Georgiev, 1935: 416), although the same paper also explicitly cited C. lanigerum subsp.albanicum (Bald.)Georgiev and C. lanigerum subsp.nikolovii Georgiev.The name leontopodium (and not albanicum or nikolovii) was chosen due to the fact that in most of the range there are individuals that more or less morphologically match the description of C. banaticum var.leontopodium Stoj.& Stefanov from the locus classicus -Mt Slavjanka ["Ali-Botuš"] (excl.f. nikolovii).
In Central Serbia, this subspecies is very common on serpentine substrates.The populations in the eastern part of Western and Southwest Serbia, on Mts.Paštrik and Koritnik, and in the broader area of the Šar Planina Mountains, are probably in the introgression zone with the type subspecies.It is therefore difficult to determine their infraspecific position.The freshest karyological data indicate that representatives from these regions are closely related to subsp.leontopodium, but without always matching the recorded morphological characters.var.cernjavskii (Georgiev)  f. stevanovicii Niketić, f. nova -Typus: Flora macedonica: Mt.Scardus (Ljuboten), in rupestribus calcareis, 2200 m., leg.V. Stevanović, 6-Jul-1979, sub "C. grandiflorum ?" (Holotypus: 1576 BEOU).-Patula, formae scardico valde simile sed inflorescentia eglandulosa.Ser, Ma.
Cerastium malyi (Georgiev)  Molecular study of the genus and species (F r a j m a n and O x e l m a n , 2007) showed that the range of this subspecies probably includes the Tatras, Carpathians, and mountainous areas of the central and southern parts of the Balkan Peninsula (Montenegro, Serbia, Bulgaria, Albania, Macedonia, Greece).In the zone where the range overlaps with subsp.monachorum (the Prokletije, and Šar Planina Mountains, Macedonian and Greek mountains) there was probably some introgression.The phylogenetic connection with a morphologically very similar plant from the Alps, described as Silene pudibunda Hoffmanns, was not proven.G r i s e b a c h (1843: 182) believed that this plant is also present on the Balkan Peninsula, citing it under the name H. pudibunda (Hoffmanns.)Griseb.2: 236 (1970).
This subspecies has a sporadic distribution in mountainous regions of the central part of the Balkan Peninsula (Northeast and Eastern Serbia, Central Bulgaria: middle part of the Stara Planina Mts.).The status of this taxon of Pančić has remained doubtful until the present day (C h a t e r and Wa l t e r s , 1964: 173, G r e u t e r et al., 1984: 270).According to C h a t e r et al. (1993: 211), it is a synonym of Silene chromodonta Boiss.& Reuter, known from the Olympus Mountains in Greece.Besides the great similarities between these two taxa, it is recorded that specimens from the Greek population have longer calyces and relatively long anthophores, which are only 2-3 times shorter than the capsule.According to G r e u t e r (1995: 25) and G r e u t e r et al. (1997: 294), the Greek population belongs to a special, locally endemic taxon, Silene pusilla Waldst.& Kit.subsp.chromodonta (Boiss.& Reuter) Greuter.We also believe that the Greek taxon and the Pančić taxon should be treated separately, but within H. pusillum (Waldst.& Kit.) Hoffmanns.
These two species also greatly differ from the ecological point of view.S. otites mostly inhabits open limestone rocky ground and open thermophilous oak and oriental hornbeam forests from lowlands to altitudes of 1000 m, while S. hungarica prefers lowland steppe and sand plain communities.& Janch. in Österr. Bot. Z. 55: 429 (1905) [basion.].

Silene parnassica
The species S. parnassica Boiss.& Spruner, from the complex S. saxifraga L., according to G r e u t e r (1995: 129) and G r e u t e r et al. (1997: 286-287) grows exclusively on the Balkan Peninsula (the data from the Apennines are actually for other taxa).There are five subspecies, including S. parnassica subsp.serbica (Vierh.et Adamović) Greuter, which was described from limestones of the southern part of Mt.Kopaonik (Treska).The type subspecies, subsp.parnassica, was also recorded for the flora of Serbia, from the foothills of Šar Planina Mountains (S t e v a n o v i ć and N i k e t i ć , 1999: 361, 517).According to G r e u t e r et al. (1997: 286), these taxa are very closely related to S. hayekiana Hand.-Mazz.& Janch., which grows in the eastern Alps and mountains of Northwest Croatia, and which would be best considered another subspecies of S. parnassica.A new combination was therefore proposed.This taxon is morphologically and phytogeographically closest to subsp.serbica.
.A new taxon for the flora of Serbia.Previously known only for central Bulgaria (middle part of the Stara Planina Mountains), it was recently recorded on Mt.Rtanj, in the Djerdap Gorge [Iron Gate] (Mt.Veliki Štrbac), in the Suva Planina Mountains, and in the Sićevačka Gorge.It is interesting to note that all three mentioned broad-leaved taxa have wider distribution than the narrow-leaved type variety (var.moehringiifolium), recorded locally in gorges and on mountains in Northeast Serbia.

BiH, Ser, Mtg, Al, Mac, Gr.
of this taxon in Southern Serbia probably pertained to subsp.moehringiifolium.This taxon grows in the Šar Planina Mountains (near the village of Brod) as well as in the mountains of Western Macedonia.It was long regarded as a subspecies, due to its lanceolate acute coronal scales (G r e u t e r et al., 1984: 271, S t e v a n o v i ć and N i k e t i ć 1999: 360, 517).However, it was later observed that size and acuteness of the coronal scales may vary even within the same population of H. pusillum.var.lanceolatumNiketić & Stevanović, var.nov.Typus: Flora serbica: the Prokletije Mountains (Mt.Žljeb, along the Rožaj-Peć road), 2000 m., leg.V.Both mentioned taxa have broader leaves than the type variety (var.monachorum), which occurs locally in gorges of Serbia and Montenegro.