Veronica austriaca Linnaeus (1759: 849)

Rojas-Andrés, Blanca M. & Martínez-Ortega, Montserrat, 2016, Taxonomic revision of Veronica subsection Pentasepalae (Veronica, Plantaginaceae sensu APG III), Phytotaxa 285 (1), pp. 448-450 : 448-450

2. Veronica austriaca Linnaeus (1759: 849) . ( Figs. 2 , 3 )

Stems (4) 10–50 (70) cm long, slightly ascending to erect, covered by eglandular hairs 0.2–1.2 (1.9) mm long, incurvate, appressed and antrorse, sometimes arranged in 2 opposite lines along the stem; apical shoot bearing 1–9 (13) pairs of leaves. Leaves (8) 10–45 (63) × (0.8) 3.5–20 (30) mm, variable in outline, rounded or truncate to cuneate at the base, serrate, dentate to deeply dentate, pinnatifid to pinnatisect or bipinnatifid to bipinnatisect with linear-lanceolate to narrowly elliptic segments, variable in width, rarely subentire, revolute to subrevolute, subglabrous or pilose, covered by hairs (0.07) 0.09–1.2 (1.5) mm long, sessile to shortly petiolate; apical shoot leaves opposite, linear, lanceolate, narrowly elliptic, entire, dentate-serrate or pinnatifid, revolute to subrevolute. Racemes axillary, opposite, bearing (8) 10–60 (70) flowers, loosely to densely arranged; peduncles 1.7–10.5 cm long, covered by the same type of indument as that of the stem, although a bit shorter; bracts (1.7) 2–10.5 mm long, variable in outline and division, pilose, glabrous or subglabrous; pedicels (0.7) 2.0–6.5 (10) mm long. Calyx (2) 3– 6 (6.5) mm long, (4) 5 sepals, linear-lanceolate, usually shorter than the capsule, glabrous, subglabrous or pilose, covered by eglandular hairs. Corolla (6) 7.5–14 (18) mm in diameter, light blue, intense blue or violet. Capsule (2.3) 2.5–6.5 (7) × 2–6 (6.5) mm, usually longer than wide, suborbicular or widely elliptic to obovate or obcordate, rounded or ± truncate at the base, emarginate or rounded at apex, sinus up to 1 (1.5) mm depth, usually puberulent, with eglandular hairs, or glabrous. Style (2.5) 3–7 mm long. Seeds (0.8) 0.9–2.3 × (0.6) 0.7–1.8 mm, ca. 1–15 per capsule.

Chromosome Number: —2 n = 16, 32, 48, (64), (80).

Habitat: —Grassy to dry and stony meadows, steppes, forest glades and edges, shrublands, alpine and subalpine meadows, rocky slopes; preferentially on calcareous substrate, occasionally on granites, basalt, gneiss, andesites, sandy soils, schistose soils, serpentines, dolomites, marble and quartzites; (0) 60–2,000 (2,250) m.

Distribution: —C, E and SE Europe, Turkey, and according to Borissova (1955) extending to Moldavia, Belarus, European Russia and the Caucasus region ( Fig. 4 ).

Phenology: —Flowering from (April) May to July (August).

Notes: —One of the most complicated cases regarding species delimitation within V. subsect. Pentasepalae is the V. austriaca complex. Both the taxonomic status and the number of biological entities recognized within it have varied a lot throughout history. Whereas some authors have accepted them at the specific rank (e.g., Fischer , 1991; Martínez-Ortega, 1999), others have treated these taxa as subspecies or varieties (e.g., Watzl, 1910; Fischer , 2011). In addition, the existence of populations whose members show particular morphological characters in states considered intermediate between those considered typical for the recognized taxonomic entities has been interpreted in different ways. Watzl (1910) considered that they represent transitional forms between different subspecies of V. austriaca , whereas Lehmann (1937) and Scheerer (1937, 1949) considered them as taxonomic entities of putative hybrid origin.

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The taxonomic entities accepted here as members of the V. austriaca complex (i.e., V. austriaca subsp. austriaca , V. austriaca subsp. dentata , V. austriaca subsp. jacquinii ) show a gradient of morphological variation regarding leaf incision. Veronica austriaca subsp. dentata is distributed in central Europe and has serrate or dentate medium leaves; V. austriaca subsp. austriaca is distributed eastwards and southwards from the area occupied by subspecies dentata , and shows deeply dentate, pinnatifid to pinnatipartite medium leaves; V. austriaca subsp. jacquinii is widely distributed in the Balkan Peninsula and shows pinnatipartite or bipinnatipartite to pinnatisect or bipinnatisect medium leaves. In addition, these taxa are mostly hexaploid, except for a few diploid populations of V. austriaca subsp. jacquinii distributed in a narrow range in the western Balkans. We consider that the populations composed of individuals that show character states intermediate between two accepted subspecies are transitional forms distributed along a gradient of ecological conditions (i.e., phenotypic plasticity of one large and variable species). Therefore, we propose to resurrect the classical treatment of V. dentata and V. jacquinii ( Watzl 1910) at the subspecific level and subordinated to V. austriaca .

2.1. Veronica austriaca subsp. austriaca ( Fig. 2 )

Stems (10) 15–30 (58.5) cm long, erect, covered by hairs 0.2–1 (1.3) mm long; apical shoot bearing 3–9 (13) pair of leaves. Leaves (11) 17–45 (48) × (1) 4.5–12 (23) mm, the basal and medium ones similar, lanceolate, narrowly elliptic, ovate, rarely elliptic, deeply dentate, pinnatifid to pinnatipartite, ± subrevolute, subglabrous or pilose, sparsely covered by hairs (0.07) 0.09–0.8 (1.3) mm long, usually the shortest ones along the margin, sometimes arranged on the primary vein and the margin (on the adaxial and abaxial face); apical shoot leaves linear, lanceolate or narrowly elliptic, entire to dentate-serrate, sometimes subrevolute. Racemes bearing (8) 15–60 (70) flowers; peduncles 3.7–10.5 cm long; bracts 2–10.3 mm long, linear or narrowly elliptic, entire, pilose, glabrous or subglabrous; pedicels (0.7) 4–5 (8.9) mm long. Calyx (2) 3–5.5 (5.8) mm long, pilose, glabrous or subglabrous. Corolla (6) 7.5–12 (14) mm in diameter, intense blue or violet. Capsule (2.3) 2.5–4.5 (5.1) × (2) 2.5–4 (5.1) mm, obovate-cordate, ± emarginate, sinus up 0.7 mm depth. Style (2.5) 3–6 (6.5) mm long. Seeds 0.8–1.7 × 0.6–1.5 mm, ca. 1–15 per capsule.

Chromosome Number: —2 n = 48.

Habitat: —Dry meadows, grassy meadows, forest glades, shrublands; preferentially on calcareous substrate, occasionally on andesites and tertiary sands; 100– 760 m.

Distribution: —E Europe; Poland, Slovakia, Romania, Croatia ( Fig. 4 ). According to Watzl (1910), present in Bosnia and Herzegovina.

Phenology: —Flowering from May to June.

Notes: —Given that it was not possible to directly revise the specimens used for the chromosome counts reported in the literature, we have only considered ploidy level estimations done by flow cytometry using our own collected material.

The epithet “macrodonta ” has been used by some authors (at the species level or combined as a subspecies or variety under V. austriaca ) to name a few populations occurring in Romania ( Rojas-Andrés et al. 2016). After examining this material, we consider that there are not enough morphological differences to recognize these populations as a separate taxon. Pending further molecular analyses, we include this material under the variation of V. austriaca subsp. austriaca .

Specimens Examined: — CROATIA. Zadar: Gračac, Crnopac, prados calizos secos, 761 m, 44º15'02.2''N, 15º48'35.5''E, 8 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR94bis (SALA149374); Gračac, Crnopac, prados calizos secos, 761 m, 44º15'02.2''N, 15º48'35.5''E, 8 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR94 (SALA149373).

POLAND. Kuyavia-Pomerania: Czyżkówker (illegible word) bei Bromberg, 15 June 69, C . KÖhler s.n. ( B100325806 , B100325808 ) ; Lublin: Z Flory Królestwa Polskiego, Gubernii lubelskiej, Okolic Lublina, Hügel im Dorf, 20 June 1882, Ferdynand Karo s.n. ( B100325825 ); Lublin, (illegible group of words), 20 June 1882, Karo s.n. ( K); Lubelskie, Lysaków prope Lublinum, in decliviis apricis angustisrum, in territorio loessaceo, 31 May 1951, Fijalkowski s.n. ( K) .

ROMANIA. Alba: Transsilvania, in collibus Scholten [= Cenade], 200 m, 1 June 1888, J . Barth s.n. ( PR); Transsilvania , in collibus graminosis Scholten [= Cenade], 10 June 1888, J . Barth s.n. ( B, K, PR, PRC) ; Transsilvania, in collibus Scholten , 13 June 1888, J . Barth s.n. ( B, E, K); Transsilvania, in collibus Scholten [=

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Phytotaxa 285 (1) © 2016 Magnolia Press • 13 Cenade], 1 June 1889, Barth s.n. ( B, K); Transsilvania, pr. Scholten [= Cenade], in collibus graminosis, 200 m, 4 June 1893, Barth s.n. ( B); in collibus Scholten [= Cenade], 100 m, 1 June 1896, J . Barth s.n. ( PR); Transsilvania, in collibus graminosis Langenthal [Valea Lungă], 100 m, 26 May 1897, Barth s.n. ( PRC); Monora [= Mănărade], in collibus, 4 June 1880, Barth s.n. ( K); Monora, in collibus, 100 m, 10 June 1901, Barth s.n. ( B, PR); Transylvania, Langenthal, Valea Lunga, E-seite del Tales, Gebüsche, Trockenwiesen, tertiäre Sande, ca. 300–500 m, 1 June 1981, E. & F . Krendl s.n. ( W); Valea Lungă, hacia el monasterio del Tauri, prados herbosos en la cara este, 316 m, 46º08'26.1''N, 24º03'26.2''E, 20 July 2014, X. Giráldez , M. M . Martínez Ortega, B. Rojas Andrés & N. López González MO 6106 ( SALA157051 ); Sibiu: in pratis: Hermannstadt [= Sibiu], 400 m, 16 June 1906, J . Barth s.n. ( W); Hermannstadt [= Sibiu] (illegible word), 29 June 1906, J . Barth s.n. ( WU); unknown province: Collinis pilisiensis, in saxis calcareis supra opp. Kestelcs [= Kesztölc], 6 May 1972, Weber s.n. ( PR); Schossen, in collibus, 300 m, 26 May 1889, J . Barth s.n. ( MA112521 ); Transsilvanien, 10 June 85, Basch. s.n. ( B) .

SLOVAKIA. Banská Bystrica: Murán, Muranska Planina, Poludnika, repisas herbosas sobre calizas, 48.76667ºN, 20.05000ºE, 2 July 2002, M. Martínez Ortega, X. Giráldez & Muñoz Centeno MO1550 (SALA124611); Muráň, Muránska Planina, prado en sotobosque de Larix , Betula y Pinus , 649 m, 48.75112ºN, 20.04861ºE, 16 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González MO 6098 (SALA157057); Košice: Comit. Szepes, in herbidis saxosis, calcareis montis “Ihrik” secus fluvium Hernád infra pag. Káposztafalv [= Hrabušice], ca. 620 m, 7 June 1915, Filarszky & Timkó s.n. (BM, C, K, PR); Letanovce, Slovensky raj, tomando la senda que sale de Trsteny potola y sube al monte Ihrik, plataforma arbustiva y rocosa (sobre calizas) con Spiraea , 619 m, 48.95525ºN, 20.43775ºE, 17 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González BR 265 (SALA153002); Vinné, en la subida a Viniansky Hrad (castillo), claros de bosque en las caídas del monte y prados sobre calizas en la subida al castillo, 164 m, 48.81709ºN, 21.94442ºE, 18 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González NLG 128 (SALA155836); Černochov, alrededores del pueblo en el monte de la antena, espinares de Prunus y Rosa sobre andesitas, 245 m, 48.43022ºN, 21.70586ºE, 18 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González NLG 129 (SALA157058).

UKRAINE. Zakarpattia: Uzhorod, in decliviis fruticosis supra Chlumec , ca. 200 m, 24 May 1930, J . Bucek s.n. ( C) .

DUBIOUS OR UNKNOWN LOCATION: Preussen: Göritz, 10 May 1954, unknown collector (B100325864).

2.2. Veronica austriaca subsp. dentata ( Schmidt 1793: 20) Watzl (1910: 53) . ( Fig. 2 )

Stems (4) 10–30 (43) cm long, erect, covered by hairs 0.2–1.0 (1.9) mm long, rarely subglabrous; apical shoot bearing (2) 4–9 (11) pair of leaves. Leaves (8) 10–45 (63) × (0.8) 3.5–12 (13.5) mm, the basal and medium ones similar, linear, lanceolate, ovate or narrowly elliptic, sometimes elliptic, rarely oblanceolate, serrate or dentate, rarely subentire and subrevolute, subglabrous or pilose, sparsely covered by hairs (0.08) 0.1–0.8 (1.1) mm long, usually the shortest ones along the margin, sometimes arranged on the primary vein and the margin (on the adaxial and abaxial face); apical shoot leaves linear or linear-lanceolate, sometimes very narrowly elliptic, entire or subentire, sometimes subrevolute. Racemes bearing 10–50 (63) flowers; peduncles 1.7–9.5 cm long; bracts (1.7) 2.5–7.0 (10.5) mm long, linear or narrowly elliptic, entire, pilose, glabrous or subglabrous; pedicels (0.7) 2.0–6.5 (6.8) mm long. Calyx (2.9) 4–6 (6.4) mm long, pilose, glabrous or subglabrous. Corolla (8) 9–14 (15.5) mm in diameter, intense blue or violet. Capsule 3–5.5 (6.5) × 2–4 (5) mm, obovate-cordate, emarginate, sinus up to 0.6 mm depth. Style 3.5–5.5 mm long. Seeds 1.2–2.2 × 1.0– 1.8 mm, 5–15 per capsule ( Kulpa 1968).

Chromosome Number: —2 n = 48.

Habitat: —Stony meadows, dry meadows, grassy meadows, forest glades, rarely on steppes; usually on limestones, rarely on basalt, gneiss or granite; 200– 560 m.

Distribution: —C and E Europe; Austria, Czech Republic, Germany, Hungary, Poland, Romania, Slovakia ( Fig. 4 ). According to Borissova (1955), it occurs in European Russia, Belarus, Ukraine, Moldavia and S Caucasus.

Phenology: —Flowering from May to June (July).

Notes: —We have examined herbarium material from the Republic of Macedonia labelled as V. austriaca subsp. dentata . These specimens have a particular morphology and from our point of view it is not clear whether they should be ascribed to V. austriaca subsp. dentata . Whereas Watzl (1910) mentioned a form of this taxon ( V. austriaca subsp. dentata f. praeterita ) from the southern part of this country, he doubted about the presence of the typical V. austriaca subsp. dentata in the northern part.

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The revised old herbarium specimens from western Germany have been provisionally assigned to this taxon, but it was not possible to confirm the presence of the plant in this area.

This taxon has also been cited from S Italy ( Fischer 1982). However, from our point of view this taxon does not reach the southern part of the Italian Peninsula. No herbarium material from this region has been found during this taxonomic revision (see notes under V. orsiniana ).

Specimens Examined: — ALBANIA. Flora albanica, ¿ Ljubelrui?, 30 June 1898, J . Bornmüller s.n. ( B100325887 ) .

AUSTRIA. Burgenland: Zitsmanndorfer Wiesen bei Podersdorf , 19 May 1973, M. A . Fischer s.n. (WU);

Lower Austria: Wiener Becken, Fischa-Wiesen bei Gramatneusiedl, trockener Riegel, 30 May 1968, M. A. Fischer s.n. ( WU) ; in graminosis ad Laxenburg in agro Vindobonensi , solo calcareo, 200 m, Braun s.n. ( B, BM, K, WU) ; Perchstoldorf , in graminosis, solo calcareo, 300 m, July 1902, Krebs s.n. ( B, BM) ; Vindobonae , in apricis circa Perchtoldsdorf, Schott s.n. ( B) ; wiesen am Fischau , June 1916, illegible collector ( PR) ; wiesen am (illegible word) der Hohen Wand, June 1906, illegible collector ( PR) ; auf der Fischerwiese am Teufelstein bei Kaltenleutgeben, 24 May 1893, Keller s.n. ( PR) ; Niederösterreichische Voralpen, Pölla bei Grillenberg nächst Berndorf , am Rand eines P. nigra - Waldes, kalk, 6 May 1972, Polatschek s.n. ( W) ; Wachau , zwischen Weissenkirchen und Dürnstein, an den SE-Hängen der Wenaheide, Trockenhänge, gneisfelsen, ca. 250–300 m, 19 May 1978, Krendl s.n. ( W) ; Thermalpen, Fluxberg , SW Gumpoldskirchen, kalk, 2 June 1957, Krendl s.n. ( W) ; Kalksburg bei Wien, Parapluieberg , 1 June and 10 June 1874, Wiesbaur s.n. ( K) ; Kalkburg , (illegible word) bei Wien, 24 May 72, ???baur?? s.n. ( K) ; Kalke Weide a. Bierhäuselberg b. Rodaun (südrand von Wien), 13 June 1908, B. Watzl s.n. ( WU) ; Kalke Weide a. Bierhäuselberg b. Rodaun (südrand von Wien), 30 May 1908, B. Watzl s.n. ( WU) ; Wien, bei Münchendorf , June 1870, Dr. W. v. Renfs s.n. ( B100325835 ) ; Geissberg bei Perchtoldsdorf nächst Wien, 26 May 1867, ¿ J B? s.n. ( WU) ; auf dem Nordlang des Braunsberges bei Hainburg o. d. Donau, May 1909, Wettstein s.n. ( WU) ; unter Gebüsch am Calvarienberg bei Baden, 18 May 1876, Skrenbergen s.n. ( WU) ; Unter Gebüsch am Calvarienberg bei Baden , Nudbestr., 18 May 1876, Skrenbergen s.n. ( MA112519 ) ; hainbueger Berge, W-Sporn des Hundscheimes Berges : waldsteppe, ca. 350 m, 3 June 1984, M. A. Fischer s.n. ( WU) ; Wien,

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Phytotaxa 285 (1) © 2016 Magnolia Press • 15 Kalksburg, Neuberg, ‘Himmelswiese’, ¿Paunon?, trockenrasen, ca. 330–340 m, 29 May 1964, M. A. Fischer s.n. (WU); Wien-Kalksburg, S-Teil der Himmelswiwiese (Neuburg), 340 m, 29 May 1964, M. A. Fischer s.n. (WU); Hundsheimer Berg, (bei Hainburg/Donau), N-Ecke, kleine Waldlichtung, 3 May 1968, M. A. Fischer s.n. (WU); Auf bergwiesen bei Gumpolds Kirchen-Kalk, 560 m, 15 May 1894, L. Keller s.n. (WU); ober dem Jägerhaus bei B ade n i m l ic ht en ¿F öre rw al d?, k al k, 400 m, 17 M ay 190 3, H. F. v. Han del - M azz et ti s.n. (W U); Niederösterreichische Kalkvoralpen: Hohe Wand: unterhalb Gh. Almfrieden, kalk, July 1991, Solinger s.n. (W); Kalksburg, 18 June 1886, illegible collector (PR); Braunsberg bei Hainburg, kalk, 200–300 m, 13 May 1906, H. F. v. Handel-Mazzetti s.n. (WU); Perchtoldsdorf: in graminosis, solo calc., 300 m. s.m., July 1902, Otto Krebs s.n. (B100325839); (illegible word) bei Rodaun, Wien, June 1854, Keller s.n. (LD1361145); Braunsberg b. Hainburg, 23 May 67, J. Wiesbauer s.n. (B100325829); Auf (illegible word) a, Brauns berge bei Hainburg an der Donau in N.Ö., May 1886, C. Aust s.n. (B100325697); NW Alpenostrand ("Thermenlinie"), ca. 1 Km. W Gumpoldskirchen (S Mödling), bosque aclarado de Quercus pubescens , ca. 350 m, 15 May 2001, M. Martínez Ortega MO1099 (SALA124613); Hainburg, Braunsberg, prados sobre calizas, 48º09'13''N, 16º57'77''E, 3 June 2001, M. Martínez Ortega MO1303 (SALA121497); Berndorf, Grillenberg, Steinbruch, 28 June 2001, M. Martínez Ortega & G. Fischer MO1450 (SALA121496); Krems, entre Weissenkirchen y Dürnstein, prados secos sobre gneis, 259 m, 48º24'18.1''N, 15º31'04.4''E, 22 July 2011, X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR 178 (SALA149043); Styria: Gösting bei Graz, August 1894, Molisch s.n. (W); bei Oberkurzheim nächst Pöls ob Judenburg, im trockerasen eines Hügels aus Kalk, 21 July 1972, Melzer s.n. (W); Laguar?a Abfängen?nt?r der (illegible word) Gösting bei Graz, 450 m, 19 May 1887, Preissmann s.n. (B); (illegible group of words) Gösting bei Graz, 450 m, 19 May 1887, E. Preissmann s.n. (B100325843); Flora Styriaca. (illegible group of words) unter der Ruine Gösting bei Graz, 450 m, 19 May 1887, E. Preissmann s.n. (B100325840); Vienna: Wien, in graminosis montanis, Kováts s.n. (B); Wien, ¿Mädling Fennyberg?, June 1894, Krebs s.n. (B); Kalksburg. En la subida desde la calle "In der Klausen", 338 m, 48º08'28.0''N, 16º14'50.2''E, 24 July 2011, X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés MO6041 (SALA149383); Perchtoldsdorf, Kalksburg, 28 June 2001, M. Martínez Ortega & G. Fischer MO1452 (SALA121495).

CULTIVATED. Ex. h. Bot. Berol. sub hoc nomine D. Delhardingius, unknown collector (B100325877).

CZECH REPUBLIC. Central Bohemia: locis stepposis collis Doutnác prope pagum Srbsko , solo calacareo, May 1930, P. Sillinger s.n. ( BM, LD1360425 , MA411803 ) ; Ad arcem Karlstein in Bohemia, unknown collector ( K) ; Vellika hora, bei karlstein in Böhmen, 1871, Pala ??? s.n. ( WU) ; Vom Berge Welikahora bei Karlstein , unknown collector ( WU) ; in rupestribus calcareis ad Karlstein , 06 June 1888, illegible collector ( B100325804 ) ; in fruticetis in colle Velká Hora prope vicum Karluv Týn , 19 May 1939, Milos Deyl s.n. ( C) ; Moravia-Silesia: Auf dem Berge Kotouc bei Stramberg , 4 June 1906, Janchen s.n. ( WU) ; Olomouc: Nebotein b. Olmütz [= Hnevotín, Olomouc], May 1907, Laus & Olmitz s.n. ( MA112515 ) ; Gryrgov prope Olomone , collis, 25 May 1932, L. F. Weber s.n. ( MA 112516 ) ; Olmütz, (illegible word), ¿steppenhügel?, June 1930, Laus s.n. ( K) ; Olmütz, [illegible group of words], June 1939, K. Laus s.n. ( MA112520 ) ; Olmütz: Steppenrelikte auf Kalk b. Grügau , June 1931, H. Laus s.n. ( B100325805 ) ; Prague: In Bohemia circa Pragam , 1821, Mann s.n. ( B100327716 ) ; Bei Prag, Wagner s.n. ( B, BM, K) ; South Moravia: Galgenberg bei Nikolsburg, Südmähren , 22 May 1913, Ronniger s.n. ( W) ; Mähren, auf grasigen Boden am Galgenberg bei Nikolsburg , 22 May 1913, Korb s.n. ( W) ; Ústí nad Labem: Montes Doupovské hory: in declivi occidentali montis Úhost' , in saxosis basalticus ad cacumen, ca. 550 m, 22 July 1951, Dr. Ivan Klásterský s.n. ( C) ; Montes Doupovské hory, in declivi occcidentali montis Úhost , in saxosis ad cacumen, ca. 550 m, 22 July 1955, Klásterský s.n. ( W) ; unknown province: E. Bohemia (illegible group of words), unknown collector ( K) .

GERMANY. Baden-Württemberg: Würtemberg, Hegelmeiner s.n. ( K); Freiburg (illegible word), im Rädel, 26 May 1896, illegible collector ( B) ; Bavaria: Marienhöhe bei Nördlingen, 3 June 1933, H . Paul s.n. ( MA 185334 ); Nórdlingen, granitboden, 460 m, 21 May & 21 June 1904, Gugler s.n. ( PR); prope Monaium, Garbmayr s.n. ( K); in ericetis prope Garching , on the?oar N ? of München, 10 June 1883, J. E . Weiss s.n. ( K); München, in ericetis prope Garching , 16 June 1883, Weiss s.n. ( BM); München: Garchinger Heide, 18 June 1920, Prof. R . Hay s.n. ( B100325866 ); prope Monach, unknown collector ( B100325831 ); unknown state: Flora von Deutschland, Laborice, Waldwiese unterhalb der Gipfels des Lobesch, 20 June 65, W . Hempel s.n. ( B100325872 ) .

HUNGARY. Budapest: Budae-Pestini, in montibus ¿ Budae?, June 1885, Vinc. de Borbás s.n. ( B100325802 ); In fruticetis montis Rupphegy , ad opp. Budapest , ca. 200 m, 20 May 1929, A . Bartha s.n. ( B100325821 ); In monte Swabenberg prope Budam, Láng s.n. ( K); Nagyszénás-B?ye c. Budapest, 03 June 37 ,

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Baschant s.n. (B100325809); Komárom-Esztergom: In decl. stepposis inter pagos Labatlant, et Bajot, 6 May 1972, Weber s.n. ( PR) ; Pest: Pilis Szanto supra oppidum Szentkereszt , 7 May 1972, Weber s.n. ( PR) ; Heves: Gyöngyös, Gyöngyösi Sárhegy , prados sobre calizas, 482m, 47º48'36.1''N, 20º00'02.7''E, 25 July 2014, X. Giráldez , M. M . Martínez Ortega, B. Rojas Andrés & N. López González BR 275 ( SALA155883 ) .

POLAND. Lesser Poland: Poloniae meridionalis districtus Miechów, ad pagum Kalina Mala , ad declive cretaceum siccum, 17 May 1968, Sychowa & Jasiewicz s.n. ( BM, C) ; Polonia merid., distr. Miechów, Zbocze stepowe, in graminosis substeposis, 8 June 1952, Blaszczyk s.n. ( BM) ; pr. Tunel distr. Miechów, Polonia merid., in graminosis substepposis, 08 June 1952, H. Blaszczyk s.n. ( C) ; Tunel prope Miechów, in graminosis substepposis prope Biara Góra , 8 June 1952, Blaszczyk s.n. ( K) ; Polonia merid., Jaksice prope Miechów , 9 June 1959, Brzyski s.n. ( BM) .

ROMANIA. Alba: Mănărade, predos secos en laderas cercanas a la carretera y a la vía de tren, 270m, 46º08'43.9''N, 23º58'41.3''E, 20 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González MO 6105 (SALA157044); Cluj: Cluj Napoca, Fanatele Clujului area, prados sobre calizas, 503m, 46º50'15.7''N, 23º37'14.3''E, 20 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González MO 6102 (SALA155862); Cluj Napoca, Cojocna, en las afueras del pueblo hacia Ceanu Mare, prados, 323m, 46º44'32.8''N, 23º50'18.5''E, 19 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González MO 6100 (SALA155863).

SLOVAKIA. Nitra: Parkan in collibus Kovacovské kopce, in decliviis aridis meridion., in rupibus supra, fl. Dunaj, ca. 250 m, 2 May 1934, Krist & Unzeitig s.n. (K); Trenčín: In rupibus calcareis m. Velikij Manin (illegible word) hungaraie, June 1878, Holuby s.n. (K); Bratislava: Thebener Kogel, prados sobre calizas, 3 June 2001, M. Martínez Ortega MO1315 (SALA124612); Košice: Čingov. P.N. Slovensky raj (Spisšká Nová Ves ), cruce con el hotel "Grand Hotel Spisš", prado herboso sobre calizas, 515m, 48º56'58.8''N, 20º29'20.4''E, 17 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González BR 266 (SALA155861).

DUBIOUS OR UNKNOWN LOCATION: In collibus, pratis Hungaria, vicinaeque Austria, unknown collector (W); In apricis ¿Kohl?, unknown collector (K); In graminosis Matrae, June–August, unknown collector (K); Neustadt, Eberswalde in Marchionata, 1824, D. Haenefeld s.n. (B100327802).

Material provisionally assigned to V. austriaca subsp. dentata : — REPUBLIC OF MACEDONIA ( FYROM). Rosoman: In faucibus "Dol. Klisura" fluvii "Rajek Reka" pr. Drenovo, c. 300 m, 11 May 1918, J. Bornmüller s.n. ( B); Saraj: Üsküb [= Skopje]: in faucibus fl. Treska, in buxetis; 4–700 m; 4–10 May 1917, J . Bornmüller s.n. ( B); Zeden-Gebirge bei Skopje, Hänge zwischen Grasje und Radusa, 21 May 1968, M . Bässler, I . Quasdorf s.n. ( B) .

SERBIA. Kosovo: Prope Kacanik (ad fl. Lepensac) prope Skoplje , 22 May 1906, Lakatos s.n. ( PRC455007 ) .

2.3. Veronica austriaca subsp. jacquinii ( Baumgarten 1816: 26) Watzl (1910: 63) . ( Fig. 3 )

Stems (12) 25–50 (70) cm long, slightly ascending to erect, covered by hairs (0.6) 0.8–1.2 (1.5) mm long; apical shoot bearing 1–5 pairs of leaves. Leaves (12) 20–30 (50) × (2) 10–20 (30) mm; the basal ones suborbicular or widely ovate, deeply dentate, pinnatifid or bipinnatifid; the medium ones pinnatipartite or bipinnatipartite, pinnatisect or bipinnatisect, with linear-lanceolate to narrowly elliptic segments, variable in width; revolute to subrevolute, pilose, covered by incurvate or crooked hairs (0.3) 0.4–1.2 (1.5) mm long; apical shoot leaves pinnatifid (very rarely only deeply dentate), revolute to subrevolute. Racemes bearing 10–50 flowers; peduncles 2– 10 cm long; bracts (2) 5–8 (9) mm long, the lower ones entire to pinnatisect, the upper ones linear and entire, glabrous, subglabrous or covered by hairs similar to those covering the leaves; pedicels (1) 3–6 (10) mm long. Calyx (2) 3–4 (6.5) mm long, pilose (covered by crooked hairs) or subglabrous. Corolla 10–14 (18) mm in diameter, light blue or dark blue. Capsule (4) 5–6.5 (7) × (3.7) 4.5–6 (6.5) mm, suborbicular or widely elliptic to obovate or obcordate, emarginate or rounded at apex, sinus up to 1 (1.5) mm depth. Style 4–7 mm long. Seeds 1.4– 2.3 × 1.1–1.8 mm, ca. 1–13 per capsule ( Kulpa 1968).

Chromosome Number: —2 n = 16, 32, 48, (64), (80).

Habitat: —Dry and stony meadows, grassy meadows, steppe-forest, steppes, forest glades and forest edges, shrublands, alpine and subalpine meadows, rocky slopes; usually on calcareous soils, occasionally on andesites, sandy soils, schistose soils, serpentines, dolomites, gneiss, marble, granites and quartzites; (0) 60–2,000 (2,250) m.

Distribution: —CE and SE Europe and Caucasus; Albania, Austria, Bosnia and Herzegovina, Bulgaria, Croatia, Greece, Georgia, Hungary, NE and SE Italy, Montenegro, Republic of Macedonia, Romania, Serbia,

TAXONOMIC REVISION OF VERONICA SUBSECTION PENTASEPALAE

Phytotaxa 285 (1) © 2016 Magnolia Press • 17 Slovakia, Slovenia, Turkey and Ukraine ( Fig. 4 ). According to Borissova (1955) it also occurs in European Russia, Belarus, Ukraine and Moldavia.

Phenology: —Flowering from (April) May to July (August).

Notes: —There is one sheet of V. austriaca subsp. jacquinii lodged at herbarium K and collected in the south of Italy (Basilicata region). We have not found populations of V. austriaca subsp. jacquinii during our field work in that area, although appropriate habitat exists. Additional searches would be necessary to confirm the presence of this taxon in S Italy, which would represent a further example of past contacts in the amphi-Adriatic area.

Specimens Examined: — ALBANIA. Lezhë: Lezhe, cerca de Fishte, pastos secos en flysch, 56 m, 41.89112ºN, 19.67781ºE, 17 June 2015, M . Martínez Ortega, X. Giráldez, N. Padilla & N. López NPG 48 ( SALA157035 ) ; Kukës: NNW von Peshkopia , W Exp. Felswand bei Slavica, an der Strasse Peshkopia-Kukës , ca. 25 km NNW von Peshkopia , 550 m, 2 August 1982, M . Baltisberger & A . Lenherr 82/1381 ( WU) ; Shkodër: Humusreiche Stellen zw. Felsen auf dem Maranaj nördl. v. Skutari, 1230 m, 2 June 1916, DÖrfler s.n. ( WU) ;

Tirana: Dajti , meadows in limestone hills, 4000 ft., 26 May 34, Pennington s.n. ( K) ; Limestone slope leading to peak of Dajti , 3500 ft., 27 May 34, Pennington s.n. ( K) ; Vlorë: distr. Kuci , in pratis ad (illegible word), 24 June 1900, Baldacci s.n. ( K) ; South Albania, mountains above Borsh , broken limestone, 2500 ft., 5 June 1933, Alston & Sandwith s.n. ( K) ; Austria Bileze, Südostga ?zien, 1884, illegible collector ( K) ; unknown province: Cla ??? Vallbone , 5000 ft., 16 June 30, G?iseff s.n. ( K) .

AUSTRIA. Upper Austria: Worf. Enns (Steyr) im Rasan a. v. Hal ???, June 1853, ¿Büschank? s.n. ( MA 163389 ) ; Styria: 1842, Alexander s.n. ( K) ; gebirgewiesen an der (illegible word), Kockbek s.n. ( WU) ; in monte Wotsch, solo dolomitico, May, Eques a Pittoni s.n. ( B) .

BOSNIA AND HERZEGOVINA. Central Bosnia Canton: Vlasic, 1931, Thompson s.n. (K); entre Travnik y Babanovac, Vlašić, prados rocosos calizos, 1464 m, 44.27533ºN, 17.60089ºE, 14 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NPG 45 (SALA157032); Travnik, Vlašić, prados sobre calizas, 1482 m, 44º16'29.4''N, 17º35'59.9''E, 12 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés SA382 (SALA149389); Herzeg-Bosnia Canton: an Acker-rainen bei Grkovci, Kalk, 780 m, 7 July 1907, Janchen & Watzl s.n. (WU); auf dem Kamme der Ilica Planina, mittlerer Teil, Kalk, 1450–1600 m, 10 July 1907, Janchen & Watzl s.n. (WU); auf der Karsthochfläge zwischen Glamoc und Hrasticevo, am wege nördlich der Rudic kosa, 1440 m, 15 July 1904, Handel Macetti & al. s.n. (WU); Bosnia, inter Suica et Livno, May 1910, Vandas s.n. (PR); in der Senkung zwischen Male poljanice un dem Troglavkessel, Kalk, 1500–1600 m, 4 July 1907, Janchen & Watzl s.n. (WU); In graminosis m. Slavic pr. ¿Glamoc?, July 1905, Vandas s.n. (PR); Paklina Planina am übergange von ¿Zupanian? nach Raono, Kalk, 1100 m, 19 July 1907, Stadlmann & al. s.n. (WU); Herzegovina-Neretva Canton: Hercegovina centralis, Porim Pl. , June 1895, illegible collector (PR); Hercegovina, montis Malý Prenj supra vicum Glogosnica, solo calcareo, 1650–1900 m, July 1933, Sillinger & Deyl s.n. (PR); Municipality of Foča: Sutjeska, Tjentiste, claros de bosque, 637 m, 43.36919ºN, 18.71119ºE, 10 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López MO 6124 (SALA157052); Municipality of Nevesinje: Hercegovina, Nevesinje, Velez , in graminosis, 15–1600 m, 18 June 1911, Sagorski s.n. (PR); Municipality of Pale: Trebevic bei sarajevo (illegible group of words), 1500 m, 11 July 1929, Ronniger s.n. (W); in declivibus graminosis montis Trevevic, ca. 1000 m, 12 June 1905, K. Maly s.n. (K); in declivibus septen. m. Trebevic prope Sarajevo, 860 m, 17 June 1906, Maly s.n. (BM); Sarajevo, Trebević, prados rocosos secos, 1224 m, 43º49'42.9''N, 18º26'06.5''E, 12 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés SA379 (SALA149388); Sarajevo, Trebevik, prados sobre calizas, 1175 m, 43.83104ºN, 18.43563ºE, 12 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NLG 142 (SALA157056); Municipality of Trebinje: Four miles on road from Gacko to Foca along Vrba valley, grassy open parts on limestone slope with few Pinus nigra , Ornithogalum, Anthyllis , flowers sky blue, 1150 m, 22 June 1965, Wrigley 65/1404 s.n. (K); entre Brgat y Trevinje, claros de bosque de Carpinus y Paliurus , 415 m, 42.65472ºN, 18.21275ºE, 10 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López MO 6118 (SALA157020); entre Gacko y Tjentiste, prados calizos subalpinos, 1076 m, 43.1847ºN, 18.56578ºE, 10 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López MO 6123 (SALA157024); entre Trebinje y Dubrovnik, sobre rocas calizas en zonas aclaradas, 282 m, 42º41'02.1''N, 18º17'49.2''E, 14 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR108 (SALA149284); entre Gacko y Tjentiste, prados calizos subalpinos, 1076 m, 43.1847ºN, 18.56578ºE, 10 June 2015, M. Martínez Ortega, X. Giráldez, N. Padilla & N. López MO 6120 (SALA157021); entre Gacko y Tjentiste, prados calizos subalpinos, 1076 m, 43.1847ºN, 18.56578ºE, 10 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López MO 6121 (SALA157022);

18 • Phytotaxa 285 (1) © 2016 Magnolia Press

ROJAS-ANDRÉS & MARTÍNEZ-ORTEGA entre Gacko y Tjentiste, prados calizos subalpinos, 1076 m, 43.1847ºN, 18.56578ºE, 10 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López MO 6122 (SALA157023); entre Brgat y Trevinje, claros de bosque de Carpinus y Paliurus , 283 m, 42.68289ºN, 18.28949ºE, 10 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López MO 6119 (SALA157047); entre Gacko y Tjentiste, prados calizos subalpinos, 1076 m, 43.1847ºN, 18.56578ºE, 10 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López MO 6123bis (SALA157025); Sarajevo Canton: ager Sarajevensis, ???one Bakije, ca. 700 m, 30 August 1906, K. Maly s.n. (K); along railway embankment, Miljacka valley, May 1931, Gilliat Smith s.n. (K); An Felsen der Miljackatals bei Sarajevo, 14 May 1906, Janchen s.n. (WU); Bosna, in collibus prope oppidum Sarajevo, 1 July 1933, Sillinger & Deyl s.n. (PR); unknown canton/municipality: 1848, Sendtner s.n. (K); Hercegovina centralis, Slitar Pl. , 16 June 1890, Begovic s.n. (PR); in declivibus graminosis montis Trevenic, ca. 1200 m, 12 August 1906, K. Maly s.n. (K).

BULGARIA. Blagoevgrad: Belasica mt., grassy places by village Kolarovo, 10 June 1980, Kuzmanov s.n. (B); in herbidis inter dumetis, mt. Alibotus, supra pag. Paril, ca. 1500 m, 25 May 1934, Achtaroff s.n. (PR); Montes Pirin (centralis), ad oppidum Bansko Vallis “Banderica”, July 1936, Klásterský & Deýl s.n. (PR); Kyustendil: Bulg. occid., Rila Planina in declivibus graminosis “Kara Bunar” dictis supra monastirum, 2250 m, Novák s.n. ( PRC); Lovech: In graminosis ad Lovec, 1894, Urumoff s.n. (WU); Pazardzhik: Batak Swamp, moist ground, 7 June 1933, unknown collector (BM); entre Batak y Beglika, borde de bosque de Pinus y Picea , 1549 m, 41º50'43.7''N, 24º09'29.4''E, 17 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente BR20 (SALA149367); entre Batak y Beglika, pastizal perenne, 1522 m, 41º50'27.1''N, 24º08'47.9''E, 17 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente BR21 (SALA149368); Pernik: Lulin Pl., 6 August 22, Turrill s.n. (K); M. Ljulin, in graminosis cacumine Strazarnic, solo andesitico, ad ca. 1100 m, 3 June 1951, B. Zelezova s.n. (K, MA182294, PR, W); cerca de Tran, entre Vukan y Konsturintsi, prados secos en bosque de Quercus pubescens , 786 m, 42º46'20.5''N, 22º35'27.8''E, 14 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente MO4576 (SALA149375); entre Staychovtsi y Dolna Melna, prados herbosos, 950 m, 42º41'03.9''N, 22º31'56.8''E, 14 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente MO4577 (SALA149376); Plovdiv: In montibus Balkan Stara Planina, in rupibus et abruptis fruticosis vallis “Sucurum” dicti ad oppidum Karlovo, 8 June 1936, Klásterský & Deyl s.n. (PR); Montes Rhodope centrales ad vicum Backovo, June 1936, Klásterský & Deyl s.n. (PR); Central Rhodopes, Asenovgrad, near Biospehere Reserve, calcareous meadow, 1130 m, 41º53'42''N, 24º55'46''E, 12 June 2001, D. Albach DA485 (SALA124579); entre Oreshets y Dobrostan, junto a la carretera, pastos secos sobre pedragales calizos, 1109 m, 41º53'44.2''N, 24º55'58.0''E, 18 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente MS1221 (SALA149384); Shumen: Madara, arriba en la fortaleza de Madara, prado subnitrófilo en la cumbre, 400 m, 43º16'38.9''N, 27º07'14.6''E, 21 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente MS1242 (SALA149386); Silistra: in graminosis prope Silistra, 1803, Urumoff s.n. (WU); Silistra, 189?, Urumoff s.n. (PR); Sliven: Sinite Kamani, prados secos sobre sustrato ácido, 689 m, 42º43'21.3''N, 26º17'39.8''E, 20 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente BR32 (SALA149370); Sofia: N of Dragoman, dry limestone hills, 1000 m, 2 August 22, Turrill s.n. (K); Sofia-city: Montes Vitosa ad urbem Sofia, 600–1900 m, 6 June 1936, Klástersky & Deyl s.n. (PR); Mount Vitosha, Sofia, multistemmed perennial in meadow, 1700 m, 23 July 1990, Bowen s.n. (RNG); Sofia, en la subida a Vitosha, prados junto a la carretera que se acerca al teleférico, prados, 1578 m, 42º35'23.7''N, 23º18'44.8''E, 15 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente BR18 (SALA149366); Vitosa Mountain, near point where road comes closest to gondola, meadow, 1580 m, 42º35'19''N, 23º18'43''E, 17 June 2001, D. Albach DA525 (SALA124581); Vitosa Mountain, mid-station of gondola, forest clearing, 1350 m, 17 June 2001, D. Albach DA522 (SALA124576); Vitosa Mountain, Ljulimin, meadow beneath ski slope, meadow, 20 June 2001, D. Albach DA527b (SALA124582); Vitosa Mountain, Ljulimin, meadow beneath ski slope, 20 June 2001, D. Albach DA527a (SALA124583); Stara Zagora: Nova Mahala , cerca de Nikolaev, junto a la carretera y cultivos de lavandín, prados secos sobre sustrato ácido en borde de bosque de Ostrya , 353 m, 42º38'02.2"N, 25º45'18.3"E, 19 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente MO4595 (SALA149377); Targovishte: Popovo, 1902, Urumoff s.n. (PR); Popovo, 1902, Urumoff s.n. (PR); Varbitza: entre Veliki Preslav y Varbitsa, pastos perennes sobre calizas junto a la carretera, 159 m, 43º06'01.2''N, 26º47'16.8''E, 21 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente MS1235 (SALA149385); Varna: hills south of Varna, 10 May 24, Gilliat-Smith s.n. (K); hills south of Varna, 19 May 1923, Gilliat-Smith s.n. (K); hills south of Varna, 22 July 24, Gilliat-Smith s.n. (K); hills south of Varna, 22 May 24, Gilliat-Smith s.n. (K);

TAXONOMIC REVISION OF VERONICA SUBSECTION PENTASEPALAE

Phytotaxa 285 (1) © 2016 Magnolia Press • 19 Prope Varna, in collinis ad Fracka , 2 June 1907, Scheneider s.n. ( K) ; prope Varna, in graminosis siccis ad ¿ Emirler ?, 18 May 1907, Scheneider s.n. ( BM, K) ; unknown province: Ad Tronovo , 1809, Urumoff s.n. ( PR) ; an wüsten orten am Celinska stena bei Loivtscha, unknown date, Urumoff s.n. ( WU) ; Bulgaria, Rhodopy Orechoro (illegible group of words), 25 September 1980, Sádlo s.n. (PRC); In m. Preslovski Balkan prope Pantelejna , 12 May 1928, Stojanoff s.n. ( K) ; prope?icum Trojacia Macedonica , 15 August 1918, Nikoloff s.n. ( K) ; Struma valley, grassy places on the hill Malkija Kozuh , 9 June 1980, Kuzmanov s.n. ( B) ; Östl. Stara Planina, Emine Balkan, Wegrand , Nähe von Slancev brjag, 450 m, 11 June 1973, Dambodt s.n. ( B) ; Macedonia Jugoslavica, Sipka Montis Stoletovo , June 1963, Weber s.n. ( PR) ; Eastern Stara Planina, Gindzi, 2 km south of Gindzi , roadside, 20 June 2001, D. Albach DA536 ( SALA124584 ) .

CROATIA. Dubrovnik-Neretva: Dalmatia, Ragusa [= Dubrovnik], 1843, unknown collector (K); Dalmatia, Dubrovnik, April 1937, Hillcoat s.n. (BM); Dalmatia, E of Dubrovnik, mountainsides near Čajcovici and Šumet, 29 May 1960, Stud. Biol. Traj. no. 60-940 d.d. s.n. (MA522017); Dalmatien, Ragusa [= Dubrovnik], am (two illegible words) Brgat, 7 April 1926, Ronniger s.n. (W); Dalmatia, in loc. saxosis ad Fort Imperial supr.?agu?am, May 1872, Pichler s.n. (K); Ob Mlini bei Dubrovkik auf Kalkblöcken, 80–100 m, 16 July 1964, Krendl s.n. (W); Dubrovnik, Gromača, prados secos sobre calizas, 320 m, 42º43'28.0''N, 18º01'04.0''E, 14 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR112 (SALA149039); Dubrovnik, entre Sumet y Gornji Brgat, prados sobre calizas, 212 m, 42º38'38.7''N, 18º08'47.2''E, 15 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés SA384 (SALA149286); Brgat, en la frontera entre Bosnia Herzegovina y Croacia, junto a la carretera, 352 m, 42.65892ºN, 18.16279ºE, 10 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López MO 6117 (SALA157019); Istria: Pisino [= Pazin], Istrien, May 1926, Meebold s.n. (K); Karlovac: Josipdol, entre Oštarije y Ribarići, prados sobre calizas, 333 m, 45º13'31.2''N, 15º14'45.7''E, 07 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés MO5528 (SALA149042); entre Josipdol y Plitvička Jezera, Lička Jesenica, prados sobre calizas, 513 m, 45º00'24.6''N, 15º25'40.8''E, 07 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés MO5529 (SALA149380); Lika-Senj: Auf dem Velnac bei Karlopago, 800 m, 28 May 1908, Janchen s.n. (WU); Croatia, ¿Plitvice?, 2000 ft., 18 May 30, Wyatt s.n. (K); Kroatien, zwischen Vrelo und Babin Potok (W Plitvicka Jezera), Kalk, am Rand einer Weidewiese, 10–24 June 1978, Polatschek s.n. (W, WU); entre Bjelopolje y Pecane, prados calizos, 44º40'26''N, 15º44'16''E, 21 June 2001, M. M. Martínez Ortega MO1439 (SALA124567); Plitvička Jezera, entre Prijeboj y Ličko, prados sobre calizas, 670 m, 44º50'44.5''N, 15º41'02.6''E, 7 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés MO5532 (SALA149339); Primorje-Gorski: Inter pagos Fuzine et Vrata in monte Mackovica dicto, in pratis, 11 June 1911, Hermann Bessel Hagen s.n. (MA112524); Istrien, Gebiet des Monte Maggiore [= Učka], beim (illegible word), ca. 950 m, 13 June 1908, Ginzberger s.n. (WU); Istrien, trockene Wiesen auf Monte Maggiore [= Učka], June 31, Baschant s.n. (B); Kroatien, Libursnicher karst, Rupa, ca. 15. km nördlich von Opatija, Karstheiden, illyr. Flaumeichenwald, kalk, ca. 420 m, 5 June 1965, Ehrendorfer s.n. (WU); Prope Fiume [= Rijeka], 28 June 1869, A. M. Smith natae Glennie s.n. (K); Prope Veprinatz, Istria, in pratis montanis, prope Fiume [= Rijeka], 30 May 1872, Smith nata Glennie s.n. (K); Schutzhaus am Mt. Maggiore [= Učka], fecha ilegible, Wihau s.n. (PR); Velika Kapela, Gorska Kotar, P. N. Risnjak, entre Platak y Stab, prados calizos, ca. 900 m, 16 June 2001, M. Martínez Ortega & A. Tribsch MO1390 (SALA124607); Opatija, ca. 15 Km al N en la carretera a Ljubljana, entre Rupa y Lipa, prados calizos, ca. 500 m, 17 June 2001, M. Martínez Ortega MO1394 (SALA124603); Velika Kapela, paso Gornje Jelenje, en el cruce hacia Crikvanica, prados calizos, 880 m, 16 June 2001, M. Martínez Ortega & A. Tribsch MO1391 (SALA124604); Entre Mrkopalj y Begovo Razdolje, prados calizos, 45º18'34''N, 14º52'54''E, 16 June 2001, M. Martínez Ortega & A. Tribsch MO1392 (SALA124605); Split-Dalmatia: Dalmatian coast, Croatia, steep south-west facing hillside at the fork to Imotski, ca. 45 km south Split on road from Makarska to Omis, 22 June 1970, R. Alava & al s.n. (SALA29163); Zadar: Kroatien, Velebiet Gebirge, strasse zw. Obrovac und dem Mali, Alanpass, kalkblockhalden mit Flecken von Pionier-Buchenwald, 1000 m, 10 June 1965, Ehrendorfer s.n. (WU); Gračac, Crnopac, Prados calizos secos, 761 m, 44º15'02.2''N, 15º48'35.5''E, 08 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR92 (SALA149372); Velebit, NP Paklenica, Sattel 0.2 km NE Strazbenica-Tadina staza- Buljma mit Umgebung, hochwüchsige Rasen an der Waldgrenze über Kalk, 1130 – 1410 m, 44.363532N, 15.453496E, 21 June 2001, G. Schneeweiss, P. SchÖnswetter & A. Tribsch PS29 (SALA); Velebit, paso Mali Alan, entre Obrovac y Sveti Rok, en el hayedo de la cara N, prados calizos, ca. 1020 m, 18 June 2001, M. Martínez Ortega MO1406 (SALA124606); Alan Pass, hacia Podprag, paredones calizos secos, 758 m, 44º15'34.1''N, 15º38'15.4''E, 08 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR89

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ROJAS-ANDRÉS & MARTÍNEZ-ORTEGA

(SALA149371); unknown county: Licausko Polje, prados calizos, 740 m, 17 June 2001, M . Martínez Ortega & A. Tribsch MO1404 ( SALA124609 ) .

CULTIVATED. Culta in R . Horto Botanico Patavino e seminibus patriae ignotae, May–June 1912 –1913, Béguinot s.n. ( BM, K); Cultiviert im Botanischen Garten in Prag, unknown collector ( PRC); Jardín Botánico de la Universidad Babes-Bolyai, cultivada, 29 June 2000, X. Giráldez , M . Martínez Ortega & J. A . Sánchez Agudo MO900 ( SALA121509 ) .

GEORGIA. Kakheti: Shirakis vake, en la frontera W de Rajons Dedoplistsqaro , entre Kvemo Bodve y Gamardzhveba, cerca del km 9 de la carretera, prados, 780 m, 41º32'07''N, 45º54'18''E, 18 May 2001, M . Martínez Ortega MO1136 ( SALA124610 ) .

GREECE. Chalkidiki: Cholomon-Gebirge, Strasse E ob Vasilika, Quercus coccifera Gebüsch mit krautiggrasigen Lichtstellen, Kristallin, 600 m, 17 August 1960, Ehrendorfer s.n. (WU); Chalkidiki: Mt. Cholomontas (también llamado Ypsizon), cerca de Paleochora, subiendo por pista forestal, claros de bosque de coscoja ( Q. coccifera ) sobre cuarcitas, 503 m, 40º29'42.7"N, 23º27'12.4"E, 24 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente MO4606 (SALA149379); Corfu: Nomos Kerkyras, ins Kerkyra, mons Pantocrator ad cacumen, in saxosis calc., 850–910 m, 27 July 1982, Georgiou s.n. (UPA); Drama: in latere boreoorientali montis Meniki (Boz Dag Serron) supra pagum Mikropolis, in pratis petrosis, pascuis, caeduis regionis superioris sylvae fagi, solo calcareo, 1400–1500 m, 41º12’N, 23º56’E, 20 July 1978, Greuter s.n. (B); mons Orvilos, in crista meridionali verticis Trisla supra refugium veterum Preseki, in clivis graminosis solo calcareo, 1450 m, 21 August 1978, Greuter s.n. (B, UPA); Nom. Serron/Dramas, Mt. Menikion (Boz Dagh of Serre), ca. 1 Km NNE of the summit, alpine meadows, limestone mixed with small patches of schist, 1750–1850 m, July 1981, Strid al. 18661 s.n. (B); Rhodopengebiet N von Drama, Umgebung der Fortstation Zagradenia, 47 km EN der Fortstation Kentriko Ergotaxion Elatias, Ruderalflur uns Strasse, 13 June 1987, Ch. Oberprieler s.n. (B); Evros: ca. 20 Km of alexandropoulis near Avra, 200–300 m, 18–20 May 1961, Ball & Wagstaffe s.n. (RNG); distr. Alexandhroupolis, between Alexandhropolis Esimi SW of the village Ava, by a corn-field, area with shrubs of Olea , 80–100 m, 22 April 1972, Stamatiadou s.n. (BM); eparhia Alexandhroupolis, ad pontem a pago Avas 3 Km septemtriones versus, in rupestribus calcareis meridiem vel occidentem spectantibus, 100 m, 40º57’N, 25º55’E, 14 July 1978, Greuter s.n. (B); Evrytania: Timphristos zwischen Karpenision und der H. A. C.-Hütte, Kalk, Schulchtsteilhang, ca. 1600 m, 26–27 June 1981, Polatschek s.n. (WU); Grevena: Palaiokastro, Mts. Vourinos, 1465 m, 40º11'56.0''N, 21º38'48.0''E, 1 August 2007, F. Cabezas & al. FJC793 (SALA149233); Imathia: in monte Vermion prope Naussa, reg. fag., sustr. calc., 1200 m, 30 May and 1 June 1936, K. H. & F. Rechinger s.n. (BM); Vermion Gebirge, 10 Km W Naouss gegen Tria-Pende-Pigadia, 1400 m, 8 June 1989, Malicky s.n. (W); Mt. Vermion, prados subalpinos con enebro, 1459 m, 40º33'06.6''N, 22º01'24.8''E, 25 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente MS1251 (SALA149387); Kavala: ad fines prov. Kavala et Serres, in montis Pangeo ditio Deve Karan, in clivis praerruptis boreo-orientem spectantibus, in graminosis subalpinis, solo marmoreo, 1800 m, 17 July 78, Greuter s.n. (B, BM); M. Pangaeon prope Kavalla, in saxosis marmor, regionis superioris, 1700–1900 m, 15 July 1970, Rechinger s.n. (B, W); Kozani/Grevena: Mons Vourinos, 1500 m, 20 June 1970, unknown collector (W); Vourinos-Massiv, östlich von Exarhos, feinerdereicher, felsdurchragter Hang, südesponiert, Serpentingestein, beweidet, 950 m, 30 May 1985, Schwerdtfeger 17674 (B); Vourinos-Massiv, östlich von Exarhos, feinerdereicher, felsdurchragter Hang, südesponiert, Serpentingestein, beweidet, 950 m, 15 August 1983, Hagemann, Ketelhut & Wolf s.n. (B); Mount Athos: Athos bei der Kirche Panajia, 1553 m, 2 July 1938, Behr s.n. (W); Mte. Athos, (illegible word) zum Panagia, in dem Gipfel, 23 June 1891, Sintenis & Bormüller s.n. (WU); Mte. Athos, peninsula Hagion Oros, 23 June 1891, Sintenis & Bormüller s.n. (PR, PRC); Pella: Berg Tzena, aufstieg Pinovon, Grobschotter-Hang, Kalk, 1050–1150 m, 41º6.5’N, 22º6.0’E, 23 August 1990, Pirker, Royl & Fleischer s.n. (B); Phthiotis: Sterea Ellas, distr. Fthiotis Mt. Othris, SE side of Jerakovouni, 1500 m up to the top 1725 m, 13 June 1971, Stamatiadou s.n. (RNG); Rhodope: SW of the village Pandhrosos, on both sides of the river, woodland with Quercus , Ostrya etc, 100–120 m, 17 May 1972, Stamatiadou s.n. (BM); Al N de Komotini, entre Pandrossos y Symvola, prados secos sobre calizas, en claros de bosque con Juniperus , Quercus, Osrya , Rhamnus , 302 m, 41º11'07.5"N, 25º25'35.0"E, 24 June 2009, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & M. Santos Vicente MO4600 (SALA149378); Serres: in monte Boz Dagh prope Serrai (Seres), ad monast. Pradramos, sol. calc., 300 m, 15 July 1936, K. H. & F. Rechinger s.n. (BM); montes Boz Dagh prope Serrai (Seres), sol. calc., 1600 m, 15 July 1936, K. H. Rechinger s.n. (BM); montes Boz Dagh prope Drama, infra granitis, 1 June 1934, K. H. & F. Rechinger s.n. (BM); montes Boz Dagh prope Drama, in saxosis inter ¿Volaka? et Pyrgos, ca.

TAXONOMIC REVISION OF VERONICA SUBSECTION PENTASEPALAE

Phytotaxa 285 (1) © 2016 Magnolia Press • 21 1000 m, 7–8 July 1936, K. H. & F. Rechinger s.n. (BM); Nom. Serron, Ep. Sintikis, Mt. Belles (Kerkini), Neon Triethnes, between the filakion and the summit, alpine meadows and rocky outcrops, gneiss and schist, 1700–1850 m, 4 July 1981, Strid & al. 18567 s.n. (B); Thessaloniki: Hag. Ilias pr. Thessaloniki, 730 m, 1929, Dibowski s.n. (W); unknown regional unit: Hillside above Papignon, 12 June 74, Lancaster s.n. (BM); in stepposis ad Halkali- Tki???, 27 May 32, ¿Scheibe? s.n. (B); Krystallopegae, limestone slopes behind the village, rare, 4000 ft., 19 June 1932, Alston & Sandwith s.n. (BM).

HUNGARY. Bács-Kiskun: Comit Pest, in pratis ad Felsöpeszér , 100 m, 13 May 1934, Boros s.n. ( PR) ; Budapest: Budapest, in silvis ad ¿ Monov?, 30 May 1988, Szépligeti s.n. ( WU) ; Hajdú-Bihar: Hungaria centralis, in collibus ad pagum Nadudvar penes Kalocsa , solo arenoso, 200 m, Haynald s.n. ( B, BM, K, PR, WU) ; Pest: Comit. Pest. in arenosis prope pag. Alsódabas , 120 m, 13 May 1934, Zarpati s.n. ( B) ; unknown county: In declivibus ad Colocam , 12 May 1882, illegible collector ( B) ; inter fruticas prope Szivinicza, 21 May 1910, Petrak s.n. ( PR) .

ITALY. Matera: ¿Matera? 19 June 37, unknown collector (K); Trieste: (illegible word), am M. Spaccato, 16 May 1871, Strobl s.n. (K); auf Wiesen von (illegible word) bei Triest, 18 May 1934, Engelhardt s.n. (B); In pratis montis Karst prope Nabresina, litorale, July 1883, Breindl s.n. (PR, PRC, W); Istrien, trockene Hänge zwischen Rizmanje [= San Giuseppe della Chiusa] und Basovizza ober Triest, 16 May 1902, Handel Mazzetti s.n. (WU); Italie, près Trieste, M. Spaccato, 4 June 84, Cheieviére s.n. (K); Lippiza ¿evom? near Triest, 7 June 39, unknown collector (K); monte Spaccato, 22 May 1895, Engelhardt s.n. (B); SW d’Aurisina, boisements de type illyrien en transition vers la fôret de chênes verts (Quercetum ilicis) sur calcaire maquia rochers et e’boulis, 150 m, 30 May 1976, P.Penret s.n. (MA303802); Tergesti [= Trieste], May 1826, Fleischer s.n. (K); Tergesto [= Trieste], unknown collector (K); Trieste in monte Spaccato, 28 May 1888, K. Richter s.n. (MA112525); Trieste, Carso Triestino, M. Cocusso, in locis graminosis, solo calcareo, cum Bromus erectus , Festuca rupicola , Scorzonera austriaca , Scorzonera villosa , Crocus reticulatus , Leontodon crispus , Narcissus poeticus ssp. radiiflorus , Carex humilis , Centaurea rupestris , 400 m, 28 May 1980, Feoli Chiapella s.n. (RNG, SALAF8756, SEV112514); Cercanías de Aurisina, hacia Trieste, prados herbosos abiertos en claros de robledal con Colymus, 177m, 45.74253ºN, 13.68331ºE, 17 June 2014, X. Giráldez , M. M. Martínez Ortega, N. Padilla García & N. López González NPG9 (SALA155844); Trieste, entre Monrupino y Fernetti, borde de bosque de Quercus junto a un camino, sobre calizas, 329 m, 45.71117ºN, 13.81217ºE, 18 June 2014, X. Giráldez , M. M. Martínez Ortega, N. Padilla García & N. López González MO6086 (SALA155838); Trieste, entre Monrupino y Fernetti, borde de bosque de Quercus junto a un camino, sobre calizas, 329 m, 45.71117ºN, 13.81217ºE, 18 June 2014, X. Giráldez , M. M. Martínez Ortega, N. Padilla García & N. López González MO6086* (SALA155880); Udine: Friuli, Udine, M. Cuarnan ad est di Gemons, 1100–1370 m, 2 July 1949, Contardo s.n. (W); Unknown province: Geröll bei ¿Natresina?, 5 June 1890, Verouvé s.n. (B).

MONTENEGRO. Andrijevica: Montenegro, in m. Balj pr. Andrijevica, ca. 1400 m, August 1906, Rohlena s.n. (BM, PR); Andrijevica, a 1 km en dirección Kolasin, pista que sale a la derecha, prados sobre calizas, 884 m, 42.74523ºN, 19.77552ºE, 8 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NPG 32 (SALA157016); Andrijevica, a 2 km en dirección Kolasin, claros de robledal, calizas, 989 m, 42.73946ºN, 19.76141ºE, 8 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NPG 31 (SALA157015); Andrijevica, a 1 km en dirección Kolasin, pista que sale a la derecha, prados sobre calizas, 884 m, 42.74523ºN, 19.77552ºE, 8 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NPG 32* (SALA157041); Bar: entre Sutorman y Karuci, Rumija Planina, claros de bosque sobre calizas junto a la carretera, 738 m, 42.16105N, 19.09708E, 9 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NLG 136 (SALA157017); entre Sutorman y Karuči, Rumija Planina, prados sobre calizas, 753 m, 42º09'43.9''N, 19º05'52.6''E, 16 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés MO5556 (SALA149285); Cetinje: Between Cattaro [= Kotor] and ¿Cettunga?, Montenegro, 25 May 1905, unknown collector (E); In gramin. ad Njegusi, loco Velji Kraj, June 1936, Pejovic s.n. (PR); Mount Lovcen, 5000 ft., 5 June 67, Huter s.n. (K); Lovcen, claros sobre calizas, 904 m, 42º25'04.9''N, 18º47'38.8''E, 9 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NLG 137 (SALA157018); Niksic: M. ¿Vardar? distr. Piva, July 1905, Rohlena s.n. ( PRC); Plav: Bertiscus (Alpes boreales balcanicae), in monte Greben prope pagun Gusinje, sustr. calc., ca. 1700 m, 22 to 26 July 1933, Rechinger fil. & Scheffer s.n. (K); Montenegro, abhänge des Visitor gegen den See von Plav, Kalk, Fettwiesen mit Pancicia, 900–1500 m, Pachernegg s.n. (WU); Pljevlja: Montenegro, S of Kosanica, in grassland, 5 June 1894, S. J. van Ooststroom & Hennipman s.n. (B); Montenegro, S of Kosanica in

22 • Phytotaxa 285 (1) © 2016 Magnolia Press

ROJAS-ANDRÉS & MARTÍNEZ-ORTEGA grassland, 30 June 1965, S. J. van Ooststroom s.n. (BM); Ljubišnja Planina, Bobovo, Prados secos sobre calizas, 1475 m, 43º16'15.5''N, 19º08'59.4''E, 18 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés SA396 (SALA149390); Plužine: Montenegro, Bozur Planina, June 1905, Rohlena s.n. (PR); Montenegro, Bozur Planina, distr. Piva, July 1905, Rohlena s.n. (PR); Mts. Treskavac, entre Borkovici y Boricje, prados subalpinos sobre calizas, 1383 m, 43º06'24.4''N, 18º54'28.4''E, 17 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR123-4 (SALA149360); Mts. Treskavac, entre Borkovici y Boricje, Prados subalpinos sobre calizas, 1383 m, 43º06'24.4''N, 18º54'28.4''E, 17 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR123-3 (SALA149359); Mts. Treskavac, entre Borkovici y Boricje, prados subalpinos sobre calizas, 1383 m, 43º06'24.4''N, 18º54'28.4''E, 17 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR123-2 (SALA149358); Mts. Treskavac, entre Borkovici y Boricje, prados subalpinos sobre calizas, 1383 m, 43º06'24.4''N, 18º54'28.4''E, 17 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR123-5 (SALA149361); Mts. Treskavac, entre Borkovici y Boricje, prados subalpinos sobre calizas, 1383 m, 43º06'24.4''N, 18º54'28.4''E, 17 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR123-7 (SALA149362); cercanías de Boricje y Plužine, en el comienzo de la pista hacia Boricje, prados sobre calizas, 1489 m, 43.14030ºN, 18.91295ºE, 11 June 2015, M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NPG 39 (SALA157055); hacia el P. N. Durmitor, desde Plužine, entre Plužine y Trsa, prados calizos junto a la carretera, 1102 m, 43.17808ºN, 18.8818ºE, 11 June 2015, M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NPG 37 (SALA157027); Mts. Treskavac, entre Borkovici y Boricje, prados subalpinos sobre calizas, 1383 m, 43º06'24.4''N, 18º54'28.4''E, 17 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR123-1 (SALA149357); Mts. Treskavac, entre Borkovici y Boricje, prados subalpinos sobre calizas, 1383 m, 43º06'24.4''N, 18º54'28.4''E, 17 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR121 (SALA149369); Žabljak: Meždo, prados cortos con enebros, calizas, 1390 m, 43.16384ºN, 19.14908ºE, 12 June 2015, M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NLG 139 (SALA157030); Žabljak, cercanías del pueblo, prados secos sobre calizas con Juniperus , 1392 m, 43º09'49.6''N, 19º09'00.3''E, 18 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés SA392 (SALA149287); unknown municipality: Montenegro, in pratis montanis prope “Krivodo” ad confines Hercegovinae, 1400 m, 21 July 1933, Sillinger & Deyl s.n. (PR).

REPUBLIC OF MACEDONIA (FYROM): Čaška: Macedonia Jugoslavica, Bogumilo, Mt. Jakupica, in saxis supra vic. Dol. Begova, 4 July 1967, Weber s.n. (PR); Jegunovce: Macedonia serbica, in collibus siccis (solo p. p. calcareo p. p. serpentinico) ad secturas Rud, Orasje (etiam Radusa) (Allatini Mines Ltd.) dictas in vicinitate oppidi General Hanriev, haud procul ab op. Skoplje (Vardarska banovina), 600–900 m, May–July 1938, A. Mach s.n. (PR806916); Karpoš: Macedonia, Yugoslavia, Skoplje, in herbosis et fruticosis montis Vodno supra opp., 17 May 1938, Weber s.n. (PR); Makedonien, Nordhang des Vodno bei Skopje, 2 May 1968, Bässler & Quasdorf s.n. (B); Kavadarci: in dumetis montis Radobilj supra pag. Kesendre, Maced. centr., 8–900 m, 12 May 1918, Bornmüller s.n. (B); Kozjak Planina, Majden, prados sobre calizas, 751 m, 41º09'32.4''N, 21º56'38.4''E, 24 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés SA416 (SALA149391); Kruševo: Macedonia jugoslavica, Prilep in fruticetis ad opp. Krusevo, 1 June 1966, Weber s.n. (PR); Mavrovo and Rostuša: Jugoslavien, Makedonien, ca. 3 Km, N Mavrovo selo, (Tetovo), Kalk, buschiger Hang an der Strasse, 10–24 June 1978, Polatschek s.n. (W, WU); South Serbia, supra pag. Senel prope urb. Gostivar, Bistra Mts., in pratis subalpinis, 1700 m, 8 July 1936, Grebenchi?off s.n. (K); Mavrovo, Bistra planina, Prados sobre cuarcitas, 1308 m, 41º43'12.3''N, 20º46'17.8''E, 23 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR129 (SALA149363); Prilep: Sivec, cerca de la cantera de mármol, prados sobre sustrato ácido, 815m, 41º24'25.7''N, 21º36'13.9''E, 10 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González NLG 124 (SALA155839); Prilep, montañas cercanas a la ciudad, sobre mármol, 950 m, 41.32127ºN, 21.63918ºE, 19 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NPG 49 (SALA157037); Prilep, en la pista hacia Toplica, prados secos en claros de robledal con Juniperus oxycedrus , 593 m, 41.39429ºN, 21.75491ºE, 18 June 2015, M. M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NLG 146 (SALA157036); Rosoman: ad drenovo, in saxosis fa??? ¿”Dol. Klisura”? fluvii Rajek-reka, 3–400 m, 13 May 1918, Bornmüller s.n. (B); Saraj: Skoplje, in fauce fluminis Treska ad vicum Glumovo, 20 May 1938, Weber s.n. (PR); Sopište: Macedonia, in decl. dumosis inter Drzilovo et Sahepjeka, ca. 1000 m, 24 August 1922, Vandas s.n. (PR); Skopje, Sopište, St. Trifun, prados secos sobre calizas con Juniperus , 526 m, 41.92604ºN, 21.36663ºE, 22 June 2015, M. Martínez Ortega, X. Giráldez, N. Padilla & N. López NLG 150 (SALA157039); Studeničani: Jugoslavia, Mt. Jakupica Macedonia, 6000–6500 m, 14 June 1955, Sladen s.n. (BM); Macedonia, in decl. siccis Elovo pr. Drzilovo,

TAXONOMIC REVISION OF VERONICA SUBSECTION PENTASEPALAE

Phytotaxa 285 (1) © 2016 Magnolia Press • 23 ca. 900 m, 24 August 1922, Vandas s.n. (PR); Macedonia, Kitka Planina, pr. Skolpje, ca. 1400 m, 29 September 1922, unknown collector (PR); Dračevo, entre Crvena Voda y Paligrad, prados herbosos y cunetas sobre sustrato ácido, 663m, 41.86184ºN, 21.45830ºE, 12 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González BR 262 (SALA155840); Golesnica Planina, entre Cverno Voda y Kitka, Claros de robledal, 828 m, 41º50'42.2''N, 21º28'17.3''E, 22 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés MO5565 (SALA149382); Tetovo: entre Stenče y Gurgurnica, prados cortos sobre calizas, 1291m, 41.87981ºN, 21.04070ºE, 6 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González BR 254 (SALA155841); Valandovo: gebiet von Valandovo, Rabrovo, 1918, Biesalski s.n. (B); Zelenikovo: Skopje, Golesnica Planina, entre Zelenikovo y Tisovica, claros de bosque de Quercus , Carpinus y Juniperus sobre esquistos (ácido), 288 m, 41º52'06.3''N, 21º36'37.4''E, 22 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés MO5562 (SALA149381).

ROMANIA. Caraș-Severin: Domogled bei Herkuleesbad, 5 June 1909, Watzl s.n. (WU); in declivibus occid. montis Domogled vicum Baile Herculanae, 1160 m, 26 June 1970, Soják s.n. (PR); Baile Herculane, cerca del pueblo junto a la carretera 67D, prados sobre calizas y borde de cultivo, 223m, 44.87198ºN, 22.41441ºE, 22 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González NLG 130 (SALA155842); Hunedoara: in collibus apricis vallis Hatzeg, 440 m, unknown date, Barth s.n. (BM, K); in collibus: Deva, 1 June 1883, J. Barth s.n. (BM, MA112526, PRC, W, WU); Iași: Iasi Valea lui David, distr. Iasi, fînat, 14 May 67, Toma s.n. (BM); Sibiu: Hammersdorf [=Gușterița], 1 June 1875, Barth s.n. (K); Hammersdorf [=Gușterița], 1 June 1974, Barth s.n. (PR); Hermannstadt, EN ob Hammersdorf (Gusterita), gegen das Denkmal, Waldränder, Weidewiesen, tertiäre sande, ca. 450–590 m, 30 May 1981, Kredl s.n. (W); in pratis at collibus ad Gross-Scheuern [= Sura Mare], 250 m, July 1889, Barth s.n. (PR); in pratis et collibus ad Gross-Scheuern [= Sura Mare], 250 m, 14 July 1889, Barth s.n. (K); in pratis Hammersdorf [=Gușterița], 420 m, 1 June 1906, Barth s.n. (BM); In pratis Hammersdorf [=Gușterița], 440 m, 10 June 1906, Barth s.n. (W); Sura Mare, alrededores de Hamba, prados herbosos, 494 m, 45º52'16.8''N, 24º12'14.8''E, 21 July 2014, X. Giráldez , M. M. Martínez Ortega, B. Rojas Andrés & N. López González BR 270 (SALA155843); unknown county: (illegible word) dorf, in collibus apricis, 1 June 1874, Barth s.n. (K); In collibus herbidis Banatus, May–June, illegible collector (K); in declivibus graminosis prope pagum Pecenecica, 60–80 m, 22 May 1922, Bujorean s.n. (BM, E, K, PR, W).

SERBIA. Prizren: Jugoslavia, Kosmet i Metohia, Osljak, in graminosis lapidosis solo calcareo, 1850 m, 10 July 1964, Mayer s.n. (B, UPA); Bor: zwischen Zlot und Brestovacka Banja, Kalk , buschiger Hang an der Strasse, 10–24 June 1978, Polatschek s.n. (W, WU); South Bačka: Fruska Gora, 12 June 1907, Prodan s.n. (PR); Deliblát, Kincstári homokpuszta, June 1910, János s.n. (PR); Deliblát, Kincstári homokpuszta, June 1908, W. János s.n. (PR); Deliblatsko sands, ca. 2, 5 mls. E of Vladimirovak station, dry sandy places among 2 ft. Vegetation, 160 m, 10 June 1965, Wrigley 65/1277 s.n. (K); South Banat: Devojački Bunar, Vladimirovac, prados sobre arenas, 155 m, 45º00'10.8''N, 20º56'21.1''E, 27 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés SA429 (SALA149392); Kosovo: Sar Planina, zwischen der Passhöhe (S Doganovic) und Globocnica, Kalk, Serpentin, Wiese im Buchenberei, 10–24 June 1978, Polatschek s.n. (WU); Pec: Bertiscus (Alpes boreales balcanicae), in monte Peklen [=Paklena] prope oppidum Pec (Ipek), in regiam Fagi, sustr. calc., ca. 1200 m, 5 July 1933, Rechinger fil. & Scheffer s.n. (K); Montenegro, in saxosis serpent. prope Pec, August 1914, Vandas s.n. (PR); Montenegro, Paklena (Kukari?a) Pl., pr. Pec, July and September 1914, Vandas s.n. (PR); Yugoslavia, Kosovo Metohija, Rusolija Massif, 10 Km NNW of Pec, soil pockets in limestone slopes, 5500 ft., 21 July 59, Cook, Grubb, Newman & Whitmore s.n. (K); Pirot: Mt. Basara, ca. 10 Km E Pirot, Wiesen W des Gipfels, tiefgründige Mähwiesen, Buchenwaldränder über kalk , ca. 1200 m, 24 July 1960, Ehrendorfer s.n. (WU); Pomoravlje: Serbien Park von Topola, Flaumeichen-Steppebuschwald, Sandstein, 300–400 m, 17 May and 5 June 1962, Ehrendorfer s.n. (WU); Šumadija: Kragujevac, May 1894, Dimitrijevic s.n. (WU); Entre Krepoljin y Despotovac, prados sobre calizas; bordes de bosque de Quercus y Carpinus , 286 m, 44º07'33.1''N, 21º30'22.4''E, 26 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR138 (SALA149365); Zaječar: M. Atanj pr. Aleksinac, in pratis subalpinis, 11 June 1939, Lindtner s.n. (K); Sokobanja, Rtanj. Base del monte Rtanj, prados herbosos sobre calizas, 498 m, 43º47'08.3''N, 21º55'05.6''E, 26 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés BR136 (SALA149364).

SLOVAKIA. Košice: Vinne, Vihorlat Mts. , Viniansky hradny vrch, laderas S y SE , en la subida al castillo, prados secos sobre calizas, ca. 200 m, 48º48'N, 20º57'E, 4 July 2002, M . Martínez Ortega, X. Giráldez & Muñoz Centeno MO1552 ( SALA124575 ); Cernochov, Vychodoslovenska nizina, sobre los viñedos, prados secos sobre andesitas, ca. 200 m, 4 July 2002, M . Martínez Ortega, X. Giráldez & Muñoz Centeno MO1553 ( SALA124608 ) .

24 • Phytotaxa 285 (1) © 2016 Magnolia Press

ROJAS-ANDRÉS & MARTÍNEZ-ORTEGA

SLOVENIA. Central Sava: In Lichten grasigen Wäldern am westlichen Bergabhang oberhal Trifail [= Trbovlje], 1 June 1906, Janchen s.n. ( WU) ; Central Slovenia: Auf trockenen Wiesen b. Laibach [= Ljubljana], unknown date, Graf s.n. ( K, PRC) ; Notranjska, Polhograjsko hribovje. E flank of Polhograjska grmada, Thermophilic meadow, 700 m, 46º05'05.0''N, 14º20'09.0''E, 28 May 2009, B. Frajman 12600 ( SALA149234 ) ; Coastal – Karst: Jugoslavien, Karstwiesen bei Divacca, 31 May 1913, Ronniger s.n. ( W) ; Wald von Lippizza bei Triest , 18 May 1884, illegible collector ( PR) ; Wegrand, Slavnik, 900–1028 m, 30 May 1966, Rainer Petermann s.n. ( MA 303803 ) ; Gorizia: Slovenija, Primorsko, in pratis lapidosis montis Mala Gora (Caven) supra oppidum Ajdovscina, solo calcareo, ca. 1100 m, 14 July 1980, Palma & Trpin s.n. ( B) ; Ternovaner Wald, Caven und Höhle Ledenica Dol, 28 June 1981, Malicky s.n. ( W, WU) ; Littoral – Inner Carniola: Slovenija, Postojna, S slope of Mt. Sovic Alt, rich forest, dry open patch, 600–650 m, 12 June 1971, Uotila s.n. ( RNG) ; Savinja: ¿ Tüffen?, May 18??, Graf s.n. ( BC654157 ) ; in declivibus rupestribus montis Hum prope Tüffer [= Lasko], solo dolomitico, 230–400 m, May 1907, Cetina s.n. ( BM, E) ; Krain, Zidani Most, steile S Hänge, kalk, 200–300 m, 9 June 1956, Ehrendorfer s.n. ( WU) ; Slowenien im Tal zwischen Radece und Dole Kalk, buschiger Hang am Waldrand , 10–24 June 1978, Polatschek s.n. ( W, WU) ; Lasko, monte Hum, en el bosque hacia la cumbre, 14 June 2001, M. Martínez Ortega, A. Tribsch & K. Tremetsberger MO1338 ( SALA124566 ) .

TURKEY. Gümüshane: carretera de Tirebolu a Kürtün (nº 887), ca. 2 Km Kürtün, grietas y rellanos de roquedos entre restos de bosque de Quercus sp. con cultivos de avellanos, granitos y serpentinas, 415 m, 40º42'N, 39º03'E, 24 June 2001, A . Herrero & al. AH1232 ( MA688155 , SALA107967 ) .

UKRAINE. Lviv: Bilcze [= Bilche Zolote] (Podole galic.) na suchych nagiah (two illegible words), 1884, illegible collector (PRC); unknown oblast: Rozyn, distr. Lipowiec, Na suchych trawiastych miejscach, 28 May 1898, Markieiczówna s.n. (K).

DUBIOUS OR UNKNOWN LOCATION: ¿Colni?, auf Stendel zu V. prostrata (illegible group of words), unknown collector (W); Diva ???ia, 600 m, 29 May 1925, Berg s.n. ( B) ; Halb-Saule, i. ¿Schrilgarten?, June 33, Baschant s.n. (B); in alpinis mte. ¿ Orieu?, May 1906, Adamovic s.n. ( K) ; In locis herbidis apricis, June 1867, Golde s.n. ( K) ; In pratis fertilibus, June 1867, Golde s.n. ( K) ; in pratis prope Rudolfswer ???, May 1898, illegible collector ( PRC); Mont ???, nei prati montosi, unknown collector ( K) ; in collibus Ha?tzag, SW Siebenbürgen near ¿ Eiscuthor? Pass, 17 July 1883, Barth s.n. ( K) ; Siebenburgen bei ¿Totfaend?, coletor ilegible (WU); in herbidis et saxosis supra Bijde Skala distr. ¿ Primovije?, 8 June 1898, Antonio Baldacci s.n. ( BM, W, WU) .

Material provisionally assigned to V. austriaca subsp. jacquinii : — BOSNIA AND HERZEGOVINA. Trebinje: Entre Tjentište y Gacko, Prados sobre calizas, 1085 m, 43º11'07.7''N, 18º33'57.7''E, 13 July 2010, S. Andrés , X. Giráldez , M. M. Martínez Ortega & B. Rojas Andrés MO 5552 ( SALA 149274).

Figures

FIGURE 2. Veronica austriaca subsp. austriaca (a–j. Slovakia, Spisšká Nová Ves, Mt. Ihrik, B. Rojas-Andrés et al. BR265, SALA153002): a. Habit; b. Stem indument; c, d. Basal leaves (adaxial face); e, f. Medium leaves (adaxial face); g. Detail of the medium leaf indument (adaxial face); h. Apical shoot leaf (abaxial face); i. Detail of the apical shoot leaf indument (abaxial face); j. Part of a fruiting raceme, with a capsule; Veronica austriaca subsp. dentata (k–o. Slovakia, Bratislava, Thebener Kogel, M. Martínez- Ortega et al. MO1315, SALA124612): k. Habit; l. Medium leaf (adaxial face); m. Detail of the medium leaf indument (adaxial face); n. Apical shoot leaf (abaxial face); o. Detail of the apical shoot leaf indument (abaxial face). Illustration by Juan Castillo.

FIGURE 3. Veronica austriaca subsp. jacquinii [a–e. Georgia, Kakheti, Shirakis vake, Dedophlistkaro, between Kvemo Bodve and Gamardjveba, M. Martínez-Ortega MO1136, SALA124610; f–i. Bosnia and Herzegovina, Trebinje, near Trebinje, B. Rojas-Andrés et al. BR108, SALA149284; j–m. Bosnia and Herzegovina, Central Bosnia, Travnik, Vlašić, S. Andrés-Sánchez et al. SA382, SALA149389]: a. Habit. b. Stem indument. c. Medium leaf (adaxial face). d. Detail of the medium leaf indument (abaxial face). e. Corolla, stamens and style. f. Habit. g. Medium leaf (adaxial face). h. Bract of a distal flower. i. Capsule and calyx. j. Habit. k. Medium leaf (abaxial face). l. Bract of a proximal flower. m. Capsule and calyx. Illustration by Juan Castillo.

FIGURE 4. Distribution map of V. austriaca based on herbarium specimens (black symbols) and collected material (open symbols); V. austriaca subsp. austriaca (circles); V. austriaca subsp. dentata (squares); V. austriaca subsp. jacquinii (triangles).