TAXON 58 (1) • February 2009: 271–276 Šmarda & al. • On Festuca pannonica, F. valesiaca and F. pseudodalmatica Taxonomic and nomenclatural notes on Festuca pannonica, F. valesiaca and F. pseudodalmatica (Poaceae) Petr Šmarda1, Jiří Danihelka1,2 & Bruno Foggi3 1 2 3 Department of Botany and Zoology, Masaryk University, Kotlářská 2, 611 37 Brno, Czech Republic. smardap@sci.muni.cz (author for correspondence) Institute of Botany, Department of Ecology Brno, Academy of Sciences of the Czech Republic, Poříčí 3b, 603 00 Brno, Czech Republic Department of Botany, University of Florence, via La Pira, 4, 50121 Firenze, Italy The taxonomic identity of Festuca pannonica Wulfen (1809) is analysed, and lectotypes are designated for F. valesiaca Schleich. ex Gaudin (1811) and F. pseudodalmatica Krajina ex Domin (1929). The nomenclatural and taxonomic histories of these taxa are briefly described, and the lectotypes of their names are analysed morphologically. As the lectotype of F. pannonica is unambiguously referable to F. valesiaca, rejection of the name F. pannonica is desirable and is proposed separately. KEYWORDS: Central Europe, fescue, lectotypification, taxonomy INTRODUCTION Narrow-leaved taxa of Festuca L. subg. Festuca represent the most species-rich and taxonomically difficult group of the genus Festuca. Numerous taxa classified within the subgenus differ in a few qualitative morphological and anatomical characters and sometimes even only in quantitative characters with considerable overlaps. The modern taxonomy of fescues goes back to Hackel (1882), who introduced anatomical characters, such as sclerenchyma arrangement in leaf cross sections, and standardised measurements of spikelet length. With chromosome counts and measurements of the relative DNA content in numerous population samples becoming a standard part of taxonomic studies, the taxonomy of narrow-leaved fescues has witnessed substantial progress (e.g., Auquier & Kerguélen, 1977; Pils, 1981, 1984; Wallossek, 1999; Šmarda & Kočí, 2003; Šmarda & al., 2005, 2007). To tackle nomenclatural confusion and stabilise nomenclature within the group, extant names have to be typified and interpreted. Here we would like to discuss the taxonomic and nomenclatural histories of F. pannonica Wulfen, F. valesiaca Schleich. ex Gaudin and F. pseudodalmatica Krajina ex Domin, including typification of the latter two. FESTUCA PANNONICA The name Festuca pannonica was proposed by Wulfen (in Host, 1809) for some narrow-leaved fescues of the F. ovina group sensu latissimo from eastern Austria and Hungary (“In collibus, locis asperis Austriae, Pannoniae”), and remained neglected for a long time. It was resurrected by Hackel (1882), who reduced it to varietal rank as F. ovina var. pannonica, as he did with several other taxa recognised nowadays at species level. Hackel (1882: 98) presented a detailed description, applicable to both diploid and tetraploid plants of F. pallens s.l., i.e., F. pallens Host and F. csikhegyensis Simonk., respectively, as classified recently (Šmarda & al., 2007). This description substantially differs from the original diagnosis and description (Host, 1809: 36; see below) at least in leaf characters: “laminae crasse junceae (1 mm diam. et ultra) rigidae, […] laeves, v. dorso scaberulae”. Hackel included in this variety a specimen from the herbarium Host and plants collected by A. Kerner in the western periphery of the city of Budapest (“Budae ad Auwinkel [ = Zugliget]”; WU-Kerner!). While the specimen W-Host 2227 is now very fragmentary (Foggi & al., 2004: 608) and cannot by identified with certainty (see below), Kerner’s specimens represent robust plants of F. pallens s.l. Hackel himself (1882: 98) was sometimes not sure about the delimitation of F. pannonica (as F. ovina var. pannonica) from F. pallens (as F. ovina var. glauca). As Hackel’s monograph remained a standard reference work for the taxonomy of fescues for many decades, the name F. pannonica became firmly connected with F. pallens s.l. in all later treatments. Soó (1973: 285), Dostál (1989: 1332) and Farkas (1999: 345) used the name F. pannonica when referring to the robust plants of F. pallens s.l. that occur on limestone and andesite rocks in northern Hungary and southern Slovakia. Whereas Hungarian authors treated F. pannonica as a subspecies of F. pallens, Markgraf-Dannenberg (1980: 146) and Dostál (1989: 1332), influenced by the former, granted F. pannonica species status. In fact, morphological descriptions of this taxon (cf. Hackel, 1882; Markgraf271 Šmarda & al. • On Festuca pannonica, F. valesiaca and F. pseudodalmatica Dannenberg, 1980) do not substantially differ from those of F. pallens (s.str.), and in some recent accounts (e.g., Englmaier, 1994, 2008; Simon, 2001), F. pannonica is reduced to the synonymy of F. pallens (s.str.). The protologue of Festuca pannonica (Host, 1809: 36 & Plate 62) consists of three elements: a diagnosis, a detailed description, and an illustration, Plate 62. The description and to a major part also the diagnosis include two important character groups used in the contemporary taxonomy of narrow-leaved Festuca species. These are the morphology of tiller leaves (“folia compresso-setacea, dorso scabra”) and of lemmas (“valvula exterior ciliata, apice brevi arista instructa, dorso convexa, laevis”). The species in question are F. csikhegyensis, F. pallens s.str., F. psammophila (Hack. ex Čelak.) Fritsch, and F. vaginata Waldst. & Kit. ex Willd. of the F. pallens group, and F. pseudodalmatica, F. pseudovina Hack. ex Wiesb., F. rupicola Heuff., and F. valesiaca s.str. of the F. valesiaca group. The leaf characters apply best to the F. valesiaca group and exclude the F. pallens group due to the rigid and mostly smooth tiller leaves in that group. The short awns, however, certainly exclude F. pseudodalmatica and probably also F. rupicola and F. valesiaca, which all usually have longer awns. By contrast to the diagnosis, the plant depicted in Plate 62 has the general appearance of F. pallens s.l. However, its rather short awns (ca. 1/4 of lemma length) do not make it possible to assign it with certainty to any particular taxon within F. pallens s.l. Within the F. valesiaca group only F. pseudovina has awns usually as short as about 1/4 of lemma length. Further, the plate is clearly discordant with the description, stating “folia compresso setacea” (Host, 1809: 36 & Plate 62). An analysis of the Festuca treatment by Host (1802, 1809) has shown that he clearly recognised morphology, habitat preferences and geographic distribution of main taxonomic groups of lowland and upland narrow-leaved fescues occurring in eastern Austria and Hungary. For the saxicolous populations (“In Austriae, Pannoniae collibus, rupestribus montanis alpinis”; Host 1802: 63), now classified as F. pallens and F. csikhegyensis (Šmarda & al., 2007), Host introduced the name F. pallens. The populations of non-rocky sites, such as rather dry meadows and sandy places (“In collibus, pratis siccioribus, locis arenosis”; Host, 1802: 61), now recognised as F. rupicola Heuff., were given the superfluous illegitimate name F. hirsuta Mygind in Host. Festuca vaginata from sandy hills in the western outskirts of Vienna (“In agro vindobonensi in collibus prope Weinhaus”; Host, 1802: 63; see also Kerner, 1882: 102–103) was erroneously attributed to F. amethystina L., while F. pseudovina, another species occurring in the surroundings of Vienna, was treated as F. ovina L. (Host 1802: Tab. 84; Hackel, 1882: 103; Hackel in Kerner, 1882: 103). Only the remaining fine-leaved taxa of dry non-rocky sites, i.e., F. valesiaca s.l. (including 272 TAXON 58 (1) • February 2009: 271–276 F. pseudodalmatica and F. valesiaca), remained neglected in the second volume of the grass monograph and, in our opinion, were later described as F. pannonica (“In collibus, locis asperis Austriae, Pannoniae”; Host, 1809: 36). From this point of view, the attribution of F. pannonica to F. valesiaca s.l. is less contradictory than any other solution; it is also very unlikely that Host would have distinguished two very similar saxicolous taxa, i.e., F. pallens and F. pannonica, while neglecting rather widespread steppic plants common in the outskirts of Vienna. A question may arise as to what constitutes original material of F. pannonica. The protologue consists of a diagnosis, ascribed to F.X. Wulfen (1728–1805), and of a description, written by Host. Either can be taken as validating the name. However, the ascription of the name according to Art. 46.3 (McNeill & al., 2006) is less clear, and there are two alternative ways of interpreting the typographic arrangement used by Host (1809; see http://www .botanicus.org/item/31753002806617): (1) The name and the diagnosis are a single statement, interrupted only by a reference to Plate 62, in which case both the name and the validating diagnosis are ascribed to Wulfen. (2) Only the diagnosis is ascribed to Wulfen but not the name. If the first option is accepted, original material should be searched for in the herbarium Wulfen, while if the second option is accepted, the name can be considered to be validly published on the basis of two different sets of descriptive material, the analysis by Wulfen and the description by Host. In that case, both the material that can be assumed to have been available to Wulfen while preparing the diagnosis and the material that Host had while preparing the description represent the original material for F. pannonica, and a lectotype may be selected from either. An analysis of Host’s publications (Host, 1802, 1809) has shown that it was not Host’s practice to ascribe a name directly to its original author, but instead to ascribe the diagnosis to him (i.e., the name and the diagnosis represent one entity, interrupted for typographical reasons by the reference to the corresponding plate), as can be documented for instance by the treaments of F. calamaria Sm., F. uniglumis Sm., F. ciliata [sensu] Pers., F. elatior L., and F. amethystina L. This suggests that both the name F. pannonica and the validating diagnosis are being ascribed by Host to Wulfen (a similar situation is described in Art. 46 Ex. 14; McNeill & al., 2006), and we prefer this option in this paper. Our opinion can indirectly be supported by the circumstance that Wulfen referred to the plants considered here as F. pannonica already in his herbarium (see Table 1). The original material of F. pannonica is deposited in the Herbarium of the Natural History Museum in Vienna (W). The name was typified along with other Festuca names published by N.T. Host by Foggi & al. (2004), unfortunately without a detailed analysis of the specimens considered, which is provided here. TAXON 58 (1) • February 2009: 271–276 Šmarda & al. • On Festuca pannonica, F. valesiaca and F. pseudodalmatica The material from herbarium Wulfen consists of four sheets (Table 1, sheets 1–4) with 1–9 culms, most of them with mature panicles; on three sheets, some fragments of tiller leaves are present. Cross sections of tiller leaves from sheets 1 and 3 are 0.41 and 0.42 mm high, respectively. They have five vascular bundles and three thick sclerenchyma strands at the midrib and edges, two thin strands in between and the valesiaca-like epidermis described below. These characters suggest that both sheets bear plants that may represent F. pseudovina, F. rupicola, F. pseudodalmatica, or F. valesiaca. However, the large panicles and linear lanceolate lemmas never occur in F. pseudovina, and a comparison of quantitative characters (Tables 1 and 2) with reliable literature (Markgraf-Dannenberg, 1980: 152– 153, Englmaier, 2008: 1161–1162) and our personal data (Šmarda, unpub.) shows that the specimens from Wulfen’s herbarium are taxonomically homogenous and referable to F. valesiaca s.str. as currently understood. It can be assumed from the texts on the labels of specimens 2–4 that they were collected in 1797 in the Erlangen botanical garden by Johann Ch.D. Schreber (1739–1810), who served as its director from 1773. He had received the seeds from an unknown person from Hungary and later sent at least three herbarium sheets of F. pannonica to F.X. Wulfen in Klagenfurt. The correspondence between Schreber and Wulfen is documented by Wulfen’s letter to J.A. Frölich (1766–1841), where also the taxonomy of Festuca is discussed (Leute, 1979). Schreber was also the first to use the name F. pannonica. The origin of specimen 1 (lectotype) with a label written only by Wulfen is uncertain: it may represent Wulfen’s own gathering from the wild or, more likely, it was also communicated by Schreber. There is also a sheet of F. pannonica in Host’s herbarium (W-Host 2227; Table 1, sheet 5; see Foggi & al. 2004: 598). It bears a single culm with two stem leaves and a well developed panicle in flower. Hence, the cross section of the tiller leaves could not be observed. The stem leaves are scabrid and the lower one has six vascular bundles, three main sclerenchyma strands (at midrib and edges) and another two thin strands in between. Also its epidermis with longitudinal ribs, observed as tubercles in the leaf cross section, is typical of F. pseudovina, Table 1. Analysis of Festuca pannonica specimens from herbaria W-Wulfen and W-Host. Nr. Inventory number and label text Description of the specimens Inflorescence characters Identification 1 W-Wulfen s.n. Festuca pannonica [m. Wulfen] 9 culms, of those 7 with mature panicles and one with small fragment of a tiller leaf PL: 7.7 cm (7) SL: 5.8 mm (9) LL: 4.0 mm (9) AL: 1.1 mm (9) Lemma ciliate Festuca valesiaca 2 W-Wulfen s.n. Festuca pannonica dum floreret lecta 1797 in h. Erl[angensi]. [m. Schreber?] 1 culm with mature panicle PL: 8.5 cm (1) SL: 6.0 mm (4) LL: 4.0 mm (4) AL: 0.85 (4) Lemma ciliate Festuca valesiaca 3 W-Wulfen s.n. Festuca pro ovina data, a qua tamen differt. Anne peculiaris species? [m. Schreber?] Schrebero pro Festuca ovina data, a qua calycibus usque 10-floris differt. Festuca pannonica [m. Wulfen] 4 culms, of those 2 with mature panicles, and fragments of tiller leaves PL: 7.0 cm (2) SL: 5.5 mm (6) LL: 3.8 mm (6) AL: 1.1 mm (6) Lemma ciliate Festuca valesiaca 4 W-Wulfen s.n. Exemplaria juniora h. a. colligam et mittam. Hujus Festucae semina ex Hungaria accepi sub nomine Festucae ovinae. An distincta species? [m. Schreber?] Ex Hungaria a Schrebero. Festuca pannonica [m. Wulfen] 5 culms, of those 3 with mature panicles, and some fragments of tiller leaves PL: 7.0 cm (3) SL: 5.65 mm (6) LL: 3.55 mm (6) AL: 1.0 mm (6) Lemma ciliate Festuca valesiaca 5 W-Host 2227 Festuca pannonica [m. Host] 1 culm with panicle and 2 stem leaves PL: 9.0 cm (1) SL: 6.95 mm (4) LL: 4.85 mm (4) AL: 1.15 mm (4) Lemma sparsely ciliate Festuca sp. AL = median length of 2nd lemma awn, LL = median lemma length, PL = maximum panicle length, SL = median spikelet length; numbers in parentheses indicate the number of measurements. 273 Šmarda & al. • On Festuca pannonica, F. valesiaca and F. pseudodalmatica F. rupicola, F. valesiaca s.l., and F. stricta Host. Considering the length and form of the panicle, lemmas, and awns, only F. pseudovina can be excluded with certainty but more precise identification is impossible. As described above, the name F. pannonica is referable to the taxon widely known as F. valesiaca. Its introduction into the botanical literature would have undesirable effect on the stability of nomenclature, so a rejection proposal under Art. 56 of the ICBN (McNeill & al., 2006) is presented elsewhere in this issue (Danihelka & al., 2009). Festuca pannonica Wulfen in Host, Icon. Descr. Gram. Austriac. 4: 36, t. 62. 1809, nom. utique rej. prop. (Danihelka & al. in Taxon 58: 295. 2009). Lectotypus (vide Foggi & al. in Ann. Naturhist. Mus. Wien, ser. B, 105: 608, Fig. 2. 2004): “Festuca pannonica [s.loc., s.coll., s.d.]” (W-Wulfen sine no.!). TAXON 58 (1) • February 2009: 271–276 FESTUCA VALESIACA The name F. valesiaca Schleich. ex Gaudin is applied to diploid narrow-leaved fescues growing mainly in the extensive steppe regions of southern Siberia, Central Asia, Asia Minor and southeastern Europe (Seidel & Jäger in Conert, 1998: 603), extending west to the inner-Alpine valleys in Switzerland, where it was first recognised and named after the canton of Valais, and to the Massif Central in France (Portal, 1999: 285). The name first appeared (as a nomen nudum) in exsiccata lists of Schleicher (1807) and Thomas (1806–1809; for bibliographic information see Moret, 1999) and was first validly published by Gaudin (1811). The only specimen of F. valesiaca found in Gaudin’s herbarium (LAU) has two original labels, written by Gaudin (Moret, pers. comm.; Figs. 1, 2B, C). The smaller label contains the locality information (De Table 2. Some quantitative morphological characters of the lectotype specimens of Festuca valesiaca, F. pannonica and F. pseudodalmatica ; numbers in parentheses indicate the number of measurements. Character F. valesiaca (LAU-Gaudin, plant B) F. pannonica (W-Host) F. pseudodalmatica (PRC) Maximum stem length (cm) 20 (3) 36 (1) 42 (14) Maximum panicle length (cm) 2.8 (3) 7.7 (7) 8.3 (14) Spikelet length (mm): median/maximum 5.6/5.6 (3) 5.8/6.0 (9) 7.0/7.0 (6) 2nd lemma length (mm): median/maximum 3.8/4.0 (3) 4.0/4.1 (9) 4.9/5.0 (6) 1.2/1.3 (3) 1.1/1.3 (9) 2.35/2.7 (6) 0.35/0.40 (6) 0.41/– (1) 0.45/0.55 (10) 2nd lemma awn length (mm): median/maximum Diameter of tiller leaves (mm): median/maximum Fig. 1. Lectotype of Festuca valesiaca (plant B; LAU-Gaudin), and cross sections of tiller leaves of plants C and D (the cross sections of A and B have the same arrangement of sclerenchyma strands as D). 274 TAXON 58 (1) • February 2009: 271–276 Šmarda & al. • On Festuca pannonica, F. valesiaca and F. pseudodalmatica Fig. 2. Labels of the lectotype specimen of Festuca valesiaca (LAU-Gaudin). For the transliterations and information about authors see text. Branson) identical with that given in the protologue (Fig. 2B), the larger label a note about origin and identification of the specimens (Fig. 2C): “Cette espèce croît à Branson en Valais. M. Villars qui l’a vue l’a appelée F. glauca maritima avec interrogation; elle me paroît [ = parâit] bien prononcée.” Ingeborg Markgraf-Dannenberg considered it the holotype, as noted on a separate label (Fig. 2A). The herbarium sheet bears four plants, three of them fully corresponding to the protologue of F. valesiaca and the current use of the name (Fig. 1, plants A, B, D). However, the third plant from the left (Fig. 1, plant C) belongs to F. trichophylla (Gaudin) K. Richt., so plant B is proposed here as lectotype (for measurements see Table 2). Although a similar cross section is found also in the alpine F. halleri, three of Gaudin’s plants clearly differ from this species in having entirely open tiller leaf sheaths, much shorter awns, smaller spikelets and thinner scabrid tiller leaves (see Markgraf-Danneberg, 1980: 143; Portal, 1999: 172–175; Englmaier, 2008: 1158–1159). Festuca valesiaca Schleich. ex Gaudin, Agrost. Helv. 1: 242. 1811. Lectotypus (hic designatus): “Festuca glauca marit ? Vill. De Branson [s.coll., s.d.]”, planta secunda a latere sinistro (LAU-herb. Gaudin!; Fig. 1, plant B). FESTUCA PSEUDODALMATICA Festuca pseudodalmatica Krajina ex Domin is very similar and closely related to F. valesiaca. In contrast to the diploid F. valesiaca, it has been shown to be tetraploid (Šmarda & al., 2005; Šmarda, unpub. data). It differs from the latter in quantitative morphological characters of panicles, spikelets, florets, and tiller leaves. Although considerable overlaps exist in all characters, statistically significant correlations have been found between several characters and ploidy level (Šmarda & al., unpubl. data), and most plants can be attributed to one or other of the two taxa at least if populations samples are considered. Festuca pseudodalmatica was first recognised during a joint expedition of K. Domin, V. Krajina and M. Deyl to the Kováčov Hills in southern Slovakia on 20 May 1929 when the first two collected it (together with other species) for the exsiccate series Flora Čechoslovenica exsiccata. Vladimír Krajina intended to publish the description of this newly recognised species in his monographic treatment of Czechoslovak fescues, published on labels to the second centuria of this exsiccate series and also separately (Krajina, 1930). For this purpose, he returned to the site one month later and collected many additional specimens, now deposited in PRC. He used these gatherings when compiling the description, as can be seen from numerous notes and drawings attached to herbarium sheets. The values measured on these specimens are given in the description of F. pseudodalmatica in the monograph (Krajina, 1930). However, the name has to be ascribed to K. Domin, who published a diagnosis of F. pseudodalmatica on the label of Cerastium brachypetalum, collected in the type locality of F. pseudodalmatica on the same day and issued already in the first centuria (Domin, 1929). Consequently, the lectotype has to be selected from the specimens collected for and later issued in the exsiccate series in May 1929 and not among those collected by V. Krajina a month later. As the specimens were gathered rather early in the season, some plants have not fully developed spikelets and might also be referable to the diploid F. valesiaca, rarely cooccurring with tetraploid plants of F. pseudodalmatica in the type locality. For this reason, we propose as lectotype one of well developed specimens of this exsiccate collection now deposited at PRC, the home institution of K. Domin and V. Krajina. Isolectotypes are found in many other herbaria but the identity of some plants may be uncertain. Our lectotype matches well both the diagnosis by Domin (1929) and the later description by Krajina (1930), and corresponds to the current use of the name. Festuca pseudodalmatica Krajina ex Domin in Acta Bot. Bohem. 8: 61. 1929. Lectotypus (hic designatus): “Festuca pseudodalmatica Krajina apud Domin […] Slovakia australis: in montibus andesiticis Kováčovské hory supra flumen Dunaj inter ostia rivorum Hron et Ipel in Quercetis mixtis, altitudine circa 150–380 m s. m. 20. V. 1929.” Flora Čechoslovenica exsiccata, no. 126, K. Domin & V. Krajina s.n. (PRC 451022!). 275 Šmarda & al. • On Festuca pannonica, F. valesiaca and F. pseudodalmatica ACKNOWLEDGEMENTS We are much obliged to Jean-Louis Moret (LAU) for his kind assistance during the inspection of type specimens in Lausanne and for deciphering the French label texts, as well as to Jan Štěpánek (PRC) and Ernst Vitek (W), curators of the herbaria. Thanks are also due to Manfred Fischer (Vienna) for making some rare literature available to us. We also would like to thank two anonymous reviewers for their advice concerning the taxonomy and nomenclature of Festuca, as well as to John McNeill and Jan Kirschner for their nomenclature expertise. The research was supported by the Ministry of Education of the Czech Republic through research grants MSM0021622416 and LC06073, and by the Czech Academy of Sciences (through the long-term research plan AV0Z60050516 of the Institute of Botany, Czech Academy of Sciences). LITERATURE CITED Auquier, P. & Kerguélen, M. 1977. Un groupe embrouillé de Festuca (Poaceae): les taxons désignés par l’épithète «glauca» en Europe occidentale et dans les régions voisines. Lejeunia, s.n., 89: 1–82. Conert, H.J. 1998. Festuca. Pp. 530–633 in: Conert, H.J. (ed.), Gustav Hegi, Illustrierte Flora von Mitteleuropa, ed. 3, vol. 1/3. Parey, Berlin. Danihelka, J., Šmarda, P. & Foggi, B. 2009. (1864) Proposal to reject the name Festuca pannonica (Poaceae). Taxon 58: 295–296. Domin, K. 1929. Schedae ad floram Čechoslovenicam exsiccatam. Centuria 1. Acta Bot. Bohem. 8: 44–79. Dostál, J. 1989. Nová květena ČSSR, vol. 2. Academia, Praha. Englmaier, P. 1994. Festuca. Pp. 993–1005 in: Fischer, M. 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