Research Article |
Corresponding author: Piotr Kosiński ( cosinski@gmail.com ) Academic editor: Alexander Sennikov
© 2021 Piotr Kosiński, Tomasz Maliński, Marcin Nobis, Magdalena Rojek-Jelonek, Dominik Tomaszewski, Monika Dering, Jerzy Zieliński.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kosiński P, Maliński T, Nobis M, Rojek-Jelonek M, Tomaszewski D, Dering M, Zieliński J (2021) Rubus kaznowskii (Rosaceae), a new bramble species from south-central Poland. PhytoKeys 185: 27-41. https://doi.org/10.3897/phytokeys.185.71193
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Based on field research in south-central Poland, supplemented with a review of herbarium materials, we identified a stable bramble biotype with a range large enough (190 km distance between the outermost stands) to be described as a new regional agamic species, Rubus kaznowskii sp.nov. It belongs to the series Subthyrsoidei(sect. Corylifolii). Although R. kaznowskii has a unique combination of features, it can be potentially mistaken for R. gothicus. It differs from the latter species in many aspects, including: pruinose primocanes, denser indumentum of the abaxial leaf surface, and more curved prickles on the petiole. R. kaznowskii has mainly been observed on rusty soils, in habitats of mixed coniferous and mixed broadleaf forests, usually in sunny places, along forest margins and roads, in clearings and roadside thickets.
distribution, ecology, genome size, morphology, taxonomy
The genus Rubus is one of the taxonomically most complex group of angiosperms. It encompasses 400–500 sexual species and more than twice as many agamic species (
A recent study on Rubus flora of Poland is based on this modern concept of agamic species (
During field studies in the Małopolska Upland (south-central Poland), we came across a remarkable bramble morphotype from the section Corylifolii Lindley, which was different from any other species of the section. The subsequent herbarium survey showed that this bramble had been relatively frequently collected in some areas where detailed floristic investigations were carried out. This allows us to suppose that the so far known area of its occurrence may be still somewhat underestimated.
We classified this bramble to ser. Subthyrsoidei (Focke) Focke (sect. Corylifolii). The series includes species that are supposed to originate by hybridisation of Rubus caesius L. and species of the series Discolores (P.J. Müller) Focke or Rhamnifolii (Bab.) Focke (
Field studies were conducted mainly during 2014 and 2019. The position of each stand (latitude, longitude and elevation) was determined using a handheld GPS unit. The distribution category was assigned following
For the nuclear DNA content estimation in R. kaznowskii, flow cytometry was used. Leaves of R. kaznowskii and the internal standard Solanum lycopersicum L. ‘Stupicke’ (1.96 pg/2C DNA;
Poland. Łódź Province: Łask District, 250 m SE of Gucin village, alongside the road in the pine forest, 186 m alt., 51°32'46"N, 19°14'54"E, 14 Jul 2014, P. Kosiński & J. Zieliński s.n.(holotype: KOR 51366; isotype: KOR 55451).
Rubus kaznowskii can be mistaken for the similar R. gothicus. However, it differs from the latter species in several aspects. Primocanes of R. kaznowskii are pruinose and covered with usually smaller and slightly curved (not straight) prickles, more numerous hairs and stalked glands. Prickles on the petiole are strongly curved and smaller. Similarly, the inflorescence axis in R. kaznowskii is covered by smaller prickles and usually more numerous stalked glands than that in R. gothicus. The serration of leaf blade margins is more regular and finer; the apex of the terminal leaflet is more gradually narrowed and shorter. The abaxial surface of the lamina (yellowish when dry) is covered by a denser indumentum and has distinct whitish, protruding veins. Although indumentum in both species consists predominantly of fasciculate hairs, their proportion to simple hairs in R. kaznowskii is higher than that in R. gothicus. The lack of long hairs in the indumentum, both fasciculate and simple, allows distinguishing R. kaznowskii not only from R. gothicus but also from other Polish representatives of Subthyrsoidei. For detailed differences between R. kaznowskii and R. gothicus, see Table
Some similarities may link R. kaznowskii with two other species from the series Subthyrsoidei, R. holandrei P.J.Müller (= R. grossus H.E.Weber) and R. storhii H.E.Weber & M.Ranft., which has not been found in Poland until now. Both species differ from R. kaznowskii by the absence of stalked glands on the vegetative stems (R. kaznowskii – usually numerous), less hairy leaf undersides, straight or slighty curved prickles on the petioles (R. kaznowskii – hooked), and flower colours: the former has white, and the latter pink petals (R. kaznowskii – white or pinkish-white and pinkish in the bud).
Shrub, usually up to 70 cm tall. First-year stems low arching, 4–6(–8) mm in diameter, angulate, with flat sides (sometimes slightly furrowed or bluntly angled), flushed violet-red or purple on the side exposed to the sun, more or less pruinose, without or with scattered 0.4–0.6-mm long fasciculate hairs (up to 15 per 1 cm length of stem side); stalked glands and acicles up to 0.4 mm long, rather numerous: usually 5–15(–25) per 1 cm length of stem side; prickles somewhat uneven and shorter than stem diameter, (2.5–)3–5(–6) mm long, quite numerous, 10–15(–19) per 5 cm of stem length, usually little curved, rarely straight and declining, abruptly tapering from the 2.5–3.5-mm broad base; intermixed with sparse to quite numerous stout small prickles. Leaves on the first-year stem moderately large (14–)18–20(–25)-cm long, 5-foliolate, indistinctly pedate, with flat or somewhat convex (when alive) leaflets arrangement; leaflets partly imbricate with flat or slightly convex/concave laminas; matt green and subglabrous above (with 0–10 adpressed simple hairs per 1 cm2, especially towards leaf margin and on main veins, and with scattered subsessile glands); light or whitish- or yellowish-green and soft to the touch beneath because of dense indumentum (50–90% of the intercostal area) of fasciculate hairs and less numerous simple hairs, with clearly visible bright and protruding veins. Terminal leaflet lamina (7–)8–10(–11)-cm long, usually ovate and widest below the middle (37–50% of its length); shallowly cordate at the base, gradually narrowed, with an apex 8–15 mm long; petiolule moderately, (17–)20–35(–39) mm, long (25–35% of its lamina length). Basal leaflets mostly sessile or sometimes with short petiolules up to 1–2 mm long; their lamina narrowly ovate to obovate, usually shorter than petioles (about 90% of their length on average). Leaf margin periodically serrate, with incisions 2–3(–5) mm deep, sometimes with 1–2 quite distinct lobes near the middle of the lamina; teeth usually triangular, broader than long, with a thin narrow apex. Petioles sparsely hairy, with scattered stalked glands and usually with 9–13 strongly curved, 1–2-mm long prickles. Stipules filiform, with scattered hairs and stalked glands. Inflorescence up to 20–30 cm long, broadly paniculate, rounded near apex, with erectopatent lateral branches up to 7–10 cm long; usually leafless above, with 3(–5)-foliolate leaves below. Inflorescence axis hairy with adpressed small fasciculate (stellate) hairs and scattered longer both fasciculate and simple hairs; stalked glands 50–80 per 1 cm of the axis length, 0.2–0.5 mm long; prickles usually 5–10 per 5 cm of axis length, declining, slender, subulate, slightly curved, 2–3 mm long (1–2 × as long as the axis thickness). Flower pedicels (5–)10–20(–25) mm long, densely hairy and with 40–80 stalked glands; prickles 4–8(–14), slender, slightly curved and declining, 1–2 mm long. Sepals patent after anthesis, 6–9 mm long, grey-green-felted with numerous stalked glands and several pricklets on the outer surface. Petals white or pinkish-white, wrinkled, broadly ovate or obovate, 12–13 mm broad and 14–16 mm long; often present an additional, incomplete whorl composed of smaller petals. Stamens longer or as long as the yellowish-green styles; anthers glabrous, yellowish-white; filaments whitish. Carpels glabrous or sparsely hairy, receptacle usually with long hairs protruding among carpels. Flowering June-July.
Main morphological differences between Rubus kaznowskii and Rubus gothicus.
Features | R. kaznowskii | R. gothicus |
---|---|---|
Primocane | ||
Wax layer | More or less pruinose | Unpruinose |
Hairs (number per 5 cm of the stem length) | (0–)10–50 fasciculate hairs | 0–10 simple hairs |
Stalked glands (number per 5 cm of the stem length) | 25–75 | 1–10 |
Prickles | Usually slightly curved and shorter than stem diameter | Usually straight, the longest as long as the stem diameter or longer |
Leaf | ||
Apex of the terminal leaflet | Gradually narrowed, 0.8–1.5 cm | Abruptly narrowed, 1.5–2.0 cm |
Blade margin of the terminal leaflet | Usually slightly periodical | Usually distinctly periodical |
Prickles on petiole | Strongly curved, not longer than petiole diameter | Slightly curved or straight and declining, longer than petiole diameter |
Indumentum cover (abaxial side of the lamina) | Dense (above 50%) | Sparse (below 50%) |
Long simple (>0.5 mm), large fasciculate (>0.35 mm), and glandular trichomes (abaxial side of the lamina) | Absent | Numerous |
Short (<0.25 mm) and medium (0.25–0.5 mm) simple hairs (abaxial side of the lamina) | Numerous | Rare |
Inflorescence axis | ||
Prickles length/rachis diameter ratio | 1–2 | >2 |
Prickles form | slightly curved | slightly curved or straight and declining |
Stalked glands (number per 5 cm of the axis length) | 20–40(–50) | 10–25(–50) |
The nuclear DNA content of R. kaznowskii is 2C = 1.43 ± 0.01 pg. Comparison of its PI fluorescence with triploid, tetraploid, and hexaploid Rubus species, of which ploidy was previously established by chromosome counting, revealed that R. kaznowskii is DNA tetraploid. In the European Rubus flora, tetraploids predominate (2n = 28) (
The epithet “kaznowskii” refers to Kazimierz Kaznowski (1876–1943), teacher, naturalist and batologist from the Świętokrzyskie (Holy Cross Mts) region (Poland); the oldest known herbarium specimen of R. kaznowskii was collected by him.
Rubus kaznowskii is a regionally distributed bramble species. The distance between outermost stands reaches more than 190 km. More than two-thirds of all its stands are located in the Kielce Upland, and one-fifth of them in the Central Masovian Upland, between the Warta and Vistula rivers. It was confirmed in nine AFE units: 34UDB4, 34UEB2, 34UDB1, 34UCC4, 34UDB3, 34UEB1, 34UDB2, 34UEB3, 34UEC2 (Fig.
The species occurs mainly on rusty soils (brunic arenosols), in semi-dry to mesic habitats of mixed coniferous and mixed broadleaf forests, preferring open places with favourable light conditions: along forest margins and in clearings and in roadside thickets, among others (Suppl. material
Poland. Łódź Province, Łask District: 2 km N of Kamostek, pine-oak forest, roadside thickets, 181 m a.s.l., 51°31'46"N, 19°04'12"E, 10 Sep 2019, P. Kosiński, T. Maliński & J. Zieliński (KOR 55452); Opoczno District: Wierzchowisko, roadside thickets, 228 m a.s.l., 51°13'04.26"N, 20°13'04.45"E, 23 Aug 2010, A. Trojecka-Brzezińska (KRA); Piotrków District: 0.5 km NW of Poniatów, broadleaved forest margin, 203 m a.s.l., 51°23'55.33"N, 19°45'02.22"E, 14 Jul 2014, P. Kosiński & J. Zieliński (KOR 51359); NW outskirts of Poniatów, edge of the brdoadleaved forest, 203 m a.s.l., 51°23'55"N, 19°45'02"E, 14 Jul 2014, P. Kosiński & J. Zieliński (KOR 51357); SEE of Piotrków Tybunalski, edge of the pine plantation on mixed deciduous forest habitat, 203 m a.s.l., 51°23'46"N, 19°45'29"E, 24 Aug 1999, J. Zieliński (KOR 41323); Mazovia Province, Białobrzegi District: Ksawerów Stary, roadside thickets, 135 m a.s.l., 51°38'58"N, 21°07'42"E, 13 Sep 2019, P. Kosiński, T. Maliński & J. Zieliński (KOR 55454); Ostrowiec District: Łysowody near Ćmielów, roadside thickets, 203 m a.s.l., 50°54'22"N, 21°33'55"E, 8 Aug 2002, R. Piwowarczyk (KRA 0354524); Radom District: W of Młodocin Mniejszy, pine forest, 194 m a.s.l., 51°20'25"N, 21°01'10"E, 3 Aug 2005, M. Nobis (KRA); E outskirts of Waliny, alongside the railway and road with the ditch, 189 m a.s.l., 51°21'07.8"N, 21°00'37"E, 4 Aug 2005, M. Nobis (KRA); 2.8 km SSE of Maliszów, mixed coniferous forest, 194 m a.s.l., 51°16'08"N, 21°07'53"E, 30 Aug 2002, M. Nobis (KRA); Kolonia Dąbrówka Zabłotnia, pine forest margin, 182 m a.s.l., 51°19'06.52"N, 21°03'17.71"E, 24 Jun 2003, M. Nobis (KRA 0318820); Kresy near Wierzbica, railway gorge, 203 m a.s.l., 51°16'22.55"N, 21°01'21.32"E, 23 Jun 2003, M. Nobis (KRA 0318832); 2 km W of Pakosław, pine-oak forest, 220 m a.s.l., 51°12'22.23"N, 21°07'58.29"E, 2 Jul 2004, M. Nobis (KRA 0317627); S of Malczew, sandpit, 179 m a.s.l., 51°21'23.68"N, 21°09'55.12"E, 27 Aug 2005, M. Nobis (KRA 0365787); Skarżysko District: N of Kierz Niedźwiedzi, forest margin, 228 m a.s.l., 51°10'53"N, 20°57'06.6"E, 10 Aug 2003, M. Nobis (KRA 0318815); Szydłowiec District: Szydłowiec, roadside thickets near the artificial lake, 219 m a.s.l., 51°13'07.16"N, 20°51'20.65"E, 22 Jul 2003, M. Nobis (KRA 0320963); 1 km N of Zaława, roadside thickets, 205 m a.s.l., 51°16'04.64"N, 20°45'59.97"E, 22 Jun 2004, M. Nobis (KRA 0317573 & 0317574); between Aleksandrów and Budki I, roadside thickets/forest margin, 242 m a.s.l., 51°12'50.98"N, 20°46'18.60"E, 25 Aug 2003, M. Nobis (KRA 0318819); Wymysłów, roadside thickets, 218 m a.s.l., 51°14'09.39"N, 20°49'08.00"E, 1 Aug 2003, M. Nobis (KRA 0318822); Wola Zagrodnia, roadside thickets, 226 m a.s.l., 51°14'43.59"N, 20°44'49.88"E, 11 Jun 2003, M. Nobis (KRA 0321855); Lipienice, alongside railway and road, 205 m a.s.l., 51°14'45.8"N, 20°58'41.6"E, 18 Aug 2002, M. Nobis (KRA); W part of Gąsawy Rządowe Niwy, forest margin, 240 m a.s.l., 51°11'57.64"N, 20°55'29.66"E, 31 Jul 2003, M. Nobis (KRA 0320129); Gąsawy Rządowe, forest margin, 237 m a.s.l., 51°12'16.47"N, 20°56'46.8"E, 8 Aug 2003, M. Nobis (KRA 0318823); Zwoleń District: 0.5 km NWW of Górki, pine plantation on the mixed deciduous forest habitat, 161 m a.s.l., 51°19'59"N, 21°35'39"E, 13 Sep 2019, P. Kosiński, T. Maliński & J. Zieliński (KOR 55453); Przyłęk, alongside the forest road with ditch, 143 m a.s.l., 51°18'30"N, 21°44'50"E, 1 Aug 2012, ? (KOR 55461); Świętokrzyskie Province, Jędrzejów District: 3 km NW of Zagórze, roadside thickets, 259 m a.s.l., 50°36'41.4"N, 20°10'33.7"E, 8 Aug 2011, B. Piwowarski (KRA); 2 km E of Małogoszcz, 1 km N of Bocheniec, pine-oak forest, 249 m a.s.l., 50°48'37"N, 20°18'27"E, 11 Jun 2011, G. Łazarski (KRA 0472546); Kielce District: 5.8 km NNW of Mniów, 300 m of Gliniany Las, silver fir forest/hornbeam-oak forest, 325 m a.s.l., 51°01'50"N, 20°24'28"E, 23 Aug 2012, M. Podgórska (KOR 55457); 0.5 km NNE of Milechowy, pine-oak forest, 262 m a.s.l., 50°49'38"N, 20°20'10"E, 4 Sep 2014, G. Łazarski (KRA 0472934); between Szewce and Łaziska, forest margin, 306 m a.s.l., 50°51'00.23"N, 20°27'41.69"E, 15 Jul 2014, P. Kosiński & J. Zieliński (KOR 51383); 2 km SW of Łaziska, Piekoszów, thickets at the edge of the wet, mixed forest, 313 m a.s.l., 50°51'29"N, 20°20'10"E, 2 Aug 2012, G. Łazarski (KRA 0472550); W of Niestachów, slopes of Mt Otrocz, pine plantation on the mixed deciduous forest habitat, 321 m a.s.l., 50°50'24"N, 20°42'60"E, 28 Jun 1934, K. Kaznowski (KOR 10598); between Raków and Sadków, edge of the pine plantation on the mixed deciduous forest habitat, 276 m a.s.l., 50°42'40.07"N, 21°03'33.83"E, 17 Jul 2014, P. Kosiński & J. Zieliński (KOR 51365); Masłów, pine plantation on the oak forest habitat, 328 m a.s.l., 50°54'30"N, 20°42'30"E, 12 Aug 1987, J. Zieliński (KOR 31452); Stara Słupia, roadside thickets, 319 m a.s.l., 50°51'37.7"N, 21°04'35.5"E, 16 Sep 1986, R. Kapuściński (SKPN); Słowik, roadside thickets, 244 m a.s.l., 50°50'08"N, 20°32'24"E, 5 Jul 1932, K. Kaznowski (KOR 10609); Końskie District: 2 km NE of Gaworczów, wilderness “Kopaczka”, thermophilous oak forest, 264 m a.s.l., 51°17'34"N, 20°27'34"E, 20 Jul 2012, M. Podgórska (KOR 55458); 1 km W of Cisownik, clearing of pine and hornbeam-oak forests, 290 m a.s.l., 51°05'17"N, 20°25'07"E, 18 Jul 2010, M. Podgórska (KRA); Opatów District: 2.2 km S of Julianów, edge of the pine-oak forest on the mixed deciduous forest habitat, 194 m a.s.l., 50°52'50"N, 21°36'50"E, 13 Sep 2019, P. Kosiński, T. Maliński & J. Zieliński (KOR 55457); Ostrowiec District: Łysowody near Ćmielów, roadside thickets, 215 m a.s.l., 50°54'07"N, 20°26'32"E, 8 Aug 2002, R. Piwowarczyk (KRA 0354532); Łysowody near Ćmielów, roadside thickets, 215 m a.s.l., 50°54'20"N, 21°33'20"E, 8 Aug 2002, R. Piwowarczyk (KRA 0354531); Skarżysko District: 1.5 km S of Grzybowa Góra Mała, roadside thickets/pine forest margin, 237 m a.s.l., 51°7'30.42"N, 20°56'29.80"E, 30 Jun 2003, M. Nobis (KRA 0318817); Skarżysko-Kamienna, Piękna street, roadside thickets, 228 m a.s.l., 51°07'12.37"N, 20°54'24.81"E, 21 Aug 2003, M. Nobis (KRA 0318828 & 0318829); Starachowice District: Jagodne Małe, forest margin, 227 m a.s.l., 51°08'44.49"N, 20°59'41.41"E, 11 Jul 2003, M. Nobis (KRA 0320966 & 0320965); Marcinków, railway embankment, 228 m a.s.l., 51°05'34.88"N, 20°58'54.03"E, 9 Aug 2003, M. Nobis (KRA 0320960); Wąchock, thickets by ruins of the manor house, 214 m a.s.l., 51°4'41.9"N, 21°00'55"E, 29 Jul 2002, M. Nobis (KRA); W of road between Lubienia and Marcule, along the road in the mesic deciduous forest, 242 m a.s.l., 51°04'23.88"N, 21°11'41.77"E, 7 Aug 2002, M. Nobis (KRA 0322795 & 0322796); Szydłowiec District: S of Barak Niwy, roadside thickets/forest margin, 252 m a.s.l., 51°10'49.45"N, 20°51'31.49"E, 26 Jun 2003, M. Nobis (KRA 0318818); Gąsawy Rządowe Niwy, roadside thickets/forest margin, 237 m a.s.l., 51°11'25.89"N, 20°56'41.12"E, 11 Jul 2003, M. Nobis (KRA 0318824).
The study was supported by the Institute of Dendrology, the Polish Academy of Sciences, Kórnik, Poland, and by the Poznań University of Life Sciences, Poznań, Poland.
Map and photos
Data type: species data