Cirsium greimleri explained

Cirsium greimleri, German: Greimler-Kratzdistel,[1] is a perennial species of flowering plant in the family Asteraceae. It was described as a separate species from Cirsium waldsteinii in 2018, from which it can be distinguished visually by its shallower lobes and deeper flower colour. Additionally, its leaves are unusually broad for the genus. It is one of a minority of species discovered through karyological analysis. It is a tall herbaceous plant with nodding purple flowers growing in high montane to subalpine habitats on exposed acidic slopes. It is native to Eastern Europe, but with an unusual geographic distribution. It is found only in the Eastern Alps and Dinaric Alps. It hybridises readily, to the extent that there is concern about genetic erosion for most populations.

Description

Stems

Cirsium greimleri is a perennial species. Plants grow from oblique, cylindrical rhizomes to heights between 110 and 180 centimetres, with the exception of a rare form that only grows to 30 cm,[2] and occasional sterile basal rosettes. Growth is erect but nodding below the capitula. Stems are shallowly ribbed. They are simple or have few branches. Their colour is green or reddish green but finely haired (pilose), or almost smooth (subglabrous) at the base, whitish green and sparsely webbed (arachnoid) in the middle, and white and densely arachnoid below the capitula.[3]

Leaves

Leaves grow all along the stem. Ground level (basal) leaves are 11.5 to 23.3 cm wide and 1.2 to 2.1 cm longer than wide in mature specimens. Their morphology ranges from egg-shaped (ovate) to circular (orbicular) to triangular (deltate), especially in younger leaves. Stalk (peduncular) leaves are lobed (amplexicaul), often eared (auriculate).[4] Their shallow feathered (pinnate) lobes are doubly toothed (serrate) to pinnatipartite, with weak yellowish to brownish-purple spinules up to 2 millimetres situated at the margins, subglabrous, to scattered hairy (pubescent), above and densely webbed-woolly (arachnoid-lanate) below.[5] All leaves are soft and herbaceous.[6]

Micromorphology

Stomatal guard cells are about 16.5 micrometres long and 9.2 μm wide.

Flowers

The flowers of C. greimleri are grouped in flower heads (capitula),[7] with each head containing between 1 and 8 individual flowers. These flower heads can be solitary or arranged in clusters (corymbosely) at the top of the stem, sometimes appearing on 1 to 5 side branches (lateral pedicels). The protective casing around the flowers (the involucre) measures 12.9 to 21.0 mm in length during flowering. Phyllaries are arranged in 6 to 7 rows. bracts flare out from the bud, with distinct vittae. The outer bracts having a visible spine, while the inner ones do not. Involucres are purplish brown to purplish black. The flower heads abound with nectar.[8] The pollen is coarsely spined.[9]

Petals are is 17.3 to 21.0 mm long in hermaphrodites, and 15.1 to 18 mm in females. At full flowering (anthesis), corollae average deep violet, but can be greyish violet or even ruby. During fading, corollae average dark purple, but can be lighter, dark magenta, or even dark ruby or dark violet. They ultimately fade to whitish, as does the brownish-purple (in hermaphrodites) or ochre synantherium. The style is always whitish except for the ruby shortly bi-lobed stigma, which fades to deep crimson or brownish-purple.[10]

Fruit

Strongly wind-dispersed (anemochorous), its fruits contain oblong, compressed, asymmetric greyish ochre achenes, 16 to 19 mm in hermaphrodites, 14.5 to 18.0 mm in females. They are attached to either 4 or 5 mm pappi. The pappus is whitish and feathery or strawlike.[11] The average dry weight of 1000 seeds is 0.4 g.[12]

Chromosomes

C. greimleri is a diploid species with sporophytic chromosome number 2n=2x=34,[13] without variation. Its somatic nuclear DNA size is about 1929 Mega base pairs.

Similar species

C. waldsteinii is genetically close and almost identical in appearance. In the field, the main distinguishing traits are lobe depth and flower colour. Although their basal leaves are roughly equal in width, the cauline leaves of C. waldsteinii are significantly wider, though the significantly deeper lobes of the latter may create the appearance of narrowness. The apparent broadness of C. greimleri leaves is distinctive within the genus. The corollae of C. waldsteinii are lighter in colour at any given stage, with little overlap in hue after opening (though both can be greyish violet at full bloom). Any comparison must be made between flowers of roughly equal stage. The green colour of its stems below the capitula are slightly visible, unlike the completely covered C. greimleri.

C. hypoleucum . Its nodding flowers are sometimes ruby red like those of C. greimleri, which is rare for the genus. The leaves of C. hypoleucum, sometimes resembling C. waldsteinii more than C. greimleri, are white-tomentose beneath, in contrast to the greyish-arachnoid leaves of C. greimleri. The involucral bracts of C. greimleri gradually become longer inward. The involucres of C. hypoleucum differ markedly in colour from those of C. greimleri, and the involucres themselves are narrower. The idumentum of subcapitular stem is sparser.

C. carniolicum . Very similar to C. greimleri in vegetative stage, but with yellow to white flowers with tougher phyllaries and spiny sepals, visible green colour through the pilosity of the subcapitular stem, and more distinctive spines on upper cauline leaves with subglabrous undersides. C. alpis-lunae is a C. carniolum like species with spiny phyllaries and longer, stiffer leaf spines but no overlap with C. greimleri.

A number of species of the Caucasus can be difficult to distinguish from the species: C. oblongifolium (longer, oblong leaves, glabrous leaf undersides), C. pseudopersonata (lighter flower colour, thinner subcapitular idumentum, glabrous leaf undersides), C. sychnosanthum (glabrous leaf undersides), C. uliginosum .

A number of species have a similar flower colour but little resemblance otherwise: C. rivulare var. 'Atropurpureum', C. borealinipponense, C. hachimantaiense, C. hidakamontanum, C. shimae, C. chokaiense, C. douglasii, C. occidentale . Such a flower colour can result from hybrids between yellow-flowered C. eristhales and purple-flowered species (such as C. alsophilum, C. palustre, C. pannonicum, C. rivulare). Hybridisation between the ancestor of C. greimleri and a yellow-flowered species like C. eristhales or C. carniolicum may be the reason the former has a different flower colour from C. waldsteinii.

The unusually wide leaves of C. greimleri are even rarer in the genus. Apart from C. carniolicum, their width is approached only by Madeira endemic C. latifolium, with which there is little risk of confusion otherwise.

Taxonomy

Etymology

The species is named after botanist Josef Greimler, whose chromosome count spurred the study that separated it from C. waldsteinii.

Phylogeny

Internally, C. greimleri populations are more genetically distinct from one another than C. waldsteinii populations, whose intrapopulation diversity, however, is higher than C. greimleri. The high interpopulation diversity could have more to do with interspecific hybridisation, but it could also be solely the result of two effects of tetraploidy: one which slows genetic drift, and another which homogenises migrating populations more effectively, such as during recolonisation following the Last Glacial Period. The low intrapopulation diversity of C. greimleri could be thanks to differences in the severity of glaciation between the Alps and the Carpathians.

Externally, a close relationship between C. greimleri, C. waldsteinii, and C. hypoleucum is suspected on morphological grounds. The relationship between C. greimleri and C. waldsteinii has already been confirmed genetically, although an allopolyploid origin of C. waldsteinii is strongly suggested. The relationship between C. greimleri and C. waldsteinii is so close that the intraspecific differences between females and hermaphrodites are higher than the interspecific differences.[14]

C. waldsteinii was grouped with C. eristhales over C. greimleri in a 2023 paper, but the phylogeny was based on genome size, GC-content, achene length, and guard cell length, rather than genetics.[15]

Taxonomic history

Its holotype BRNU 658042 was collected by Petr Bureš at 1739 m from 47.272°N 14.4063°W in the Rottenmann and Wölz Tauern on 15 September 2015. C. greimleri had been considered part of C. waldsteinii until 2018. In 2018, following the discovery that Alpine-Dinaric populations of "C. waldsteinii" were invariably diploid, whereas Carpathian populations were invariably tetraploid, the Alpine-Dinaric and Carpathian populations were split into two species. This decision was corroborated by physical and genetic evidence of separation.

All records before 2018 were published under other names, but the earliest published report of proleptic C. greimleri can be considered that of Host 1831, collected at near Judenburg and classified as Cnicus pauciflorus .[16] Unpublished, the first record of what is now considered C. greimleri may have been a Tyrolian record in an 1807 catalogue by Johann Christoph Schleicher, but this is dubious.[17]

It was first published within its present genus as Cirsium pauciflorum by Anton Eleutherius Sauter following its discovery in Rottenmanner Tauern.[18] In 1905, C. pauciflorum was recognised to be a younger homonym of C. pauciflorum, which is now designated Carduus defloratus. Because of this, they assigned it a new name, C. waldsteinii.[19]

The form that only grows to 30 cm was identified in 1902 and designated C. greimleri f. depressum by Eugen Johan Khek, The form with branches was designated C. greimleri var. ramosum by Khek after he identified it in 1906.[20]

Distribution and habitat

Distribution

It is found in the Eastern[21] and Dinaric Alps. A partial but detailed map is designated Obr. 32b in Vavrinec 2020, which also provides distribution maps for its hybrids. A complete map was published in 2018, which includes flora-based ranges in the Dinarides, from which few precise coordinates exist yet.[22] The precise ranges it has been found on include the Ennstal Alps, Rottenmann and Wölz Tauern, Seckau Tauern, Seetal Alps,[23] [24] Saualpe, Koralpe, Karawanks, Kamnik–Savinja Alps, Pohorje,[25] Julian Alps, Snežnik,[26] Srnetica, Klekovača,[27] Cincar, Vranica, Bjelašnica, Visočica, Treskavica, Zelengora, Jahorina,[28] Komovi, Golija, and Kopaonik.[29]

On its own, its distribution is similar to that of Cardamine waldsteinii,[30] and somewhat similar to those of Achillea clusiana,[31] Moehringia ciliata,[32] and Vicia oroboides. When taken together with C. waldsteinii, its circum-Pannonian distribution is similar to that of Crocus vernus, which also exhibits geographic differences in ploidy.[33] Or to the acidophile species Hieracium transylvanicum. With less overlap, to Centaurea uniflora or Pilosella alpicola. Including the Pontic and Caucasus, to Koeleria eriostachya. Including part of the Apennines, to Aposeris foetida.

Habitat

It is a subalpine and high montane species,[34] [35] found from 800 to 2000 m. It grows on moist downwind slopes and on roadsides in valleys with running water in the more montane part of its range, but also in more open scree forests in the more subalpine part of its range.

C. greimleri has an extremely low shade tolerance, 5 for temperature, requiring a continental climate with moderate soil moisture, It has no halotolerance. The overlapping, or coterminous Cirsium species with the greatest ecological similarity is C. arvense, followed by C. heterophyllum and C. spinosissimum. The species is calcifuge, requiring moderately acidic soil pH, but does sometimes grow on calcareous substrates, so long as it is very richly fertile soil.

Ecology

Reproduction

Flowering is from late June to late July, to early August at sites of higher shade or altitude.

Plants are gynodioecious,[36] some hermaphrodite and others female (rudimental synantheria, without developed pollen), in addition to sterile individuals that do not progress beyond rosette stage. Hermaphrodite flowers can be distinguished visually by the protrusion of their synantherium from the corolla, by colour of their synantherium, by the pollen pushed out by an elongating style at full anthesis (female synantheria lack pollen). Hermaphrodite synantheria are longer, at 6.7 to 9.3 mm, compared to female synantheria, at 4.3 to 5.7 mm. The same applies to styles, 20 to 25 mm in hermaphrodites, and 18.0 to 22.4 mm in females. Although both hermaphrodite and female stigmas can be straight, only female stigmas can be twisted, and usually are.

The smaller size of female achenes relative to hermaphrodite achenes in this species is unusual. Usually the reverse is the case, which has been explained as a compensation for the genetic disadvantage of females relative to hermaphrodites, and as a result of the absence of inbreeding depression.[37] [38] Suggested explanations for the aberrance of C. greimleri and C. waldsteinii include larger achene count, higher germanation rates, and lower infestation with achene predators like the Tephritidae and Curculionidae.

Hybridisation

The following hybrids have been reported:

Of the Cirsium species native to the northern part of its range, the greatest geographic overlap is with C. palustre, C. arvense, C. vulgare, and C. oleraceum. There is high overlap with C. eristhales and C. heterophyllum. There is moderate overlap with C. rivulare. There is low overlap with C. spinosissimum and C. carniolicum. There is very little overlap with, C. pannonicum, C. acaule, and no overlap with C. alsophilum, C. brachycephalum, C. canum. Though the lack of overlap with C. alsophilum, also a mountainous species with a presence in the Alps and Dinarides, is not certain at least for the Dinaric portion of its range.

Despite high range overlap, C. greimleri does not hybridise with C. arvense, which may be due to intersectional incompatibility.

Associations

C. greimleri grows in forests dominated by Acer pseudoplatanus, Alnus incana, Betula pendula, Corylus avellana, Fagus sylvatica, Larix decidua, Picea abies, Pinus cembra, and Sorbus aucuparia.[39]

It is often found in association with other subalpine tall forbs, such as Adenostyles alliariae, Athyrium filix-femina, Calluna vulgaris, Cerastium lanatum, Chaerophyllum hirsutum, Chamaenerion angustifolium, Cicerbita alpina, Doronicum austriacum, Dryopteris filix-mas, Erigeron glabratus, Gentiana asclepiadea, Hieracium laevigatum, Hieracium lachenalii, Hieracium sylvaticum, Lactuca muralis, Peucedanum ostruthium, Phyteuma spicatum, Pleurospermum austriacum, Polypodium vulgare, Prenanthes purpurea, Senecio nemorensis, Senecio ovatus, Struthiopteris spicant, Telekia speciosa, Vaccinium myrtillus, Valeriana tripteris, Veratrum lobelianum, and Veronica urticifolia.

Among the shorter species it associates with are Aremonia agrimonoides, Artemisia umbelliformis, Cardamine enneaphyllos, Cardamine trifolia, Cerastium lanatum, Lamium orvala, Omphalodes verna, Sanicula epipactis, and Vicia oroboides.[40] [41]

It forms part of the Pinion mugo type of Krummholz vegetation in the Dinarics.[42] It is sometimes an element of the Scabioso hladnikianae-Grafietum golakae association.[43] It is a rare element of the Calamagrostion arundinaceae association, the Huperzio-Alnetum viridis Mulgedio-Aconitetea (1948) subassociation,[44] [45] and the Polysticho-lonchitis-fagetum rhododendrotosum hirsuti subassociation. It is also found in the associations EU 6170 Adenostylion,[46] Cirsio cani-Filipenduletum ulmariae, Caricion davallianae, and Veronico-Calitrichetum.

Conservation

It readily forms hybrids, with the highest degree of promiscuity in its genus among the Cirsium species of the Pannonian Basin, leaving it vulnerable to genetic erosion through local imbalances in pollen production, leading to unidirectional geneflow,[47] for which reason it is regarded as an endangered species. Most populations consist of only a few to several hundreds of individuals, and only the Koralpe and Seetaler Alpen have known populations with over a thousand.[48] Within its genus, an erosion risk is not unique to C. greimleri, applying also to C. alsophilum, C. bertolonii, and C. carniolicum.

A collection of seeds for the Millennium Seed Bank was made in 2014.

See also

Further reading

Notes and References

  1. Web site: 2022 . Luise . Schratt-Ehrendorfer . Cirsium . Rote Liste der Farn- und Blütenpflanzen Österreichs . 3rd . 2024-07-04 . de.
  2. 1905 . Eugen Johan . Khek . J.J. Reiff . Karlsruhe . Floristisches aus Steiermark . 41–42 . 2024-10-18 . Allgemeine botanische Zeitschrift für Systematik, Floristik, Pflanzengeographie . 11 . 3 . Cirsium pauciflorum Spreng. f. depressum. Durch 3 Jahre beobachtet. Eine 30 cm hohe, auffallend niedrige Form in mehreren Exemplaren bei Trieben. . de . Floristics from Styria.
  3. Book: Reichenbach, Heinrich Gustav . 1853 . Heinrich Gustav Reichenbach . Ambrosius Abel . Leipzig . Cirsium . 67–101 . https://books.google.com/books?id=dyUCzGbZ_5AC&pg=PA67 . Deutschlands Flora, mit höchst naturgetreuen, charakteristischen Abbildungen aller ihrer Pflanzenarten in natürlicher Größe und mit Analysen auf Kupfertafeln, als Beleg für die Flora Germanica Excursoria und zur Aufnahme und Verbreitung der neuesten Entdeckungen innerhalb Deutschlands und der angrenzenden Länder: Belgien und Holland, Dänemark, Preussen, Galizien, Polen, Siebenbürgen, Ungarn, Kroatien, Dalmatien, Istrien, Venedig, der Lombardei, Piemont, der Schweiz, dem Elsaß . 15 . Mächst auf grasigen Wiesen, in Wäldern. Sommer.♃ . de . The Flora of Germany, with Highly Lifelike, Representative Illustrations of all its Plant Species in Natural Size and with Analyses on Copper Plates, as Evidence for the Flora Germanica Excursoria and for the Recording and Dissemination of the Latest Discoveries within Germany and the neighboring countries: Belgium and Holland, Denmark, Prussia, Galicia, Poland, Transylvania, Hungary, Croatia, Dalmatia, Istria, Venice, Lombardy, Piedmont, Switzerland, the Alsace.
  4. Book: 1803 . Pál . Kitaibel . Pál Kitaibel . Franz de Paula Adam Norbert Wenzel Ludwig Valentin . von Waldstein . Franz de Paula Adam von Waldstein . Matthias Andreas Schmidt . Vienna . Cnicus pauciflorus . 175–176 . https://bibdigital.rjb.csic.es/viewer/11189/?offset=#page=105 . Descriptiones et icones plantarum rariorum Hungariae . 2 . la . Descriptions and Depictions of Rare Plants of Hungary.
  5. Book: Werner, Klaus . 1976 . University Press . Cambridge . Cirsium Miller . 232–242 . https://archive.org/details/floraeuropaea0004tuti/page/232 . Flora Europaea . 4 . 0-521-08717-1.
  6. Reichenbach 1853. "Cirsium pauciflorum Sprengel: einfach, flockigspinnwebig, Stängel wenig-blüthig, oben nackt, unterste Blätter breit länglichdreieckig, grob buchtig gezähnt, am Rande stachlich, ober sitzend, am Grunde beiderseits geöhrt, schmäler, unterseits spinnwebig ... Blätter oberseits mit kleinen Haaren. Mächst auf grasigen Wiesen, in Wäldern. Sommer.♃" (in German) [Cirsium pauciflorum Sprengel: simple, flaky, spider-webbed, stems with few flowers, naked above, lowest leaves broadly elongated-triangular, coarsely sinuously toothed, spiny on the edge, sessile above, auriculate on both sides at the base, narrower, spider-webbed below ... leaves with small hairs on the upper side. Grows on grassy meadows, in forests. Summer. Perennial]
  7. 1905 . Karl . Fritsch . Karl Fritsch . Naturwissenschaftlicher Verein für Steiermark . Graz . Blütenbiologische Untersuchungen verschiedener Pflanzen der Flora von Steiermark . Flower Biology Research on Various Plants of the Flora of Styria . 267–282 . 2024-10-18 . Mitteilungen des Naturwissenschaftlichen Vereines für Steiermark . 0369-1136 . 42 . de.
  8. Fritsch 1905. "Im Tubus der Blumenkrone ist viel Honig enthalten." (in German) [The tube of the corolla contains a lot of honey.]
  9. Fritsch 1905. "Der Griffel ist während des weiblichen Stadiums der Anthese 22 mm lang, wovon nur 3 mm auf die dicht aneinandergepreßten Griffeläste entfallen. Die langen Fegehaare bilden einen auffallenden Kranz. Der Pollen ist grobstachelig." (in German) [During the female stage of anthesis, the style is 22 mm long, of which only 3 mm are the tightly pressed style branches. The long hairs form a conspicuous wreath. The pollen is coarsely spined.]
  10. Fritsch 1906. "Die großen Köpfchen des Cirsium pauciflorum Spr. zeichnen sich durch sehr dunkle, trübpurpurne Färbung aus. Zu dieser dunklen Gesamtfärbung tragen hauptsächlich die fast schwarzpurpurnen Griffeläste bei, welche viel dunkler sind als die Blumenkrone. Am Grunde (nämlich in dem nicht dem Lichte ausgesetzten Teule) sind alle Blütenteile, namentlich die Röhre der Blumenkrone und der Griffel, weißlich, gegen die Spitze zu wird die Färbung immer lebhafter purpurn. Die Blumenkrone ist 16–17 mm lang; hievon entfallen 5–6 mm auf den stielförmigen Teil der Röhre. Die Zipfel sind stets von ungleicher Länge, der kürzeste ist 3 mm, der längste 5 mm lang." (in German) [The large heads of Cirsium pauciflorum Spr. are characterized by a very dark, dull purple color. The almost black-purple style branches, which are much darker than the corolla, contribute to this dark overall color. At the base (namely in the part not exposed to light), all parts of the flower, especially the corolla tube and the style, are whitish; towards the tip the color becomes increasingly more vibrant purple. The corolla is 16–17 mm long; 5–6 mm of this is the stalk-shaped part of the tube. The tips are always of unequal length; the shortest is 3 mm, the longest 5 mm.]
  11. Reichenbach 1853. "Köpfe wenige, 2–4, Hüllschuppen lanzettlich, am Rand und auch gegen Rücken bis auf Mittellinie fitzig, undeutlich gekielt, an der Spitze umgeschlagen. ... Blüthchen dunkelpurpurn, Gaum bis über Mitte gespalten, Spitzen der Abschnitte verdickt. Pappusfedern an der Spitze spatelig erweitert. Pollen gut entwickelt. Achenien länglich zusammengedrückt, am Grunde schmäler, dunkelbraun." (in German) [Heads few, 2–4, involucral scales lanceolate, striate at the margin and also towards the back up to the midline, indistinctly keeled, folded over at the tip. ... Florets dark purple, palate split up to the middle, tips of the sections thickened. Pappus feathers widened spatulately at the tip. Pollen well developed. Achenia elongated and compressed, narrower at the base, dark brown.]
  12. Book: Universität . Graz . Accessions for the Millenium Seed Bank, Royal Botanic Gardens Kew, 2014 . 14–20 . Samentauschverzeichnis des Botanischen Gartens der Karl-Franzens-Universität Graz: Ernte 2014 . 2024-10-18 . de . Seed Exchange List of the Botanical Garden of Karl-Franzens-University in Graz: Harvest 2014.
  13. 1996 . Christoph . Dobeš . Michael . Kiehn . Ernst . Vitek . Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft . Vienna . Beiträge zur Gefässpflanzen-Flora von Österreich: Chromosomenzählungen III . 301–318 . 2024-10-18 . Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien . 1025-4749 . 133 . de . Contributions to the Vascular Plant Flora of Austria.
  14. 2023-05-24 . Ester . Michálková . Jakub . Šmerda . Klára . Plačková . Aleš . Knoll . Petr . Bureš . Hybridization may endanger the rare North Apennine endemic Cirsium bertolonii . Plant Systematics and Evolution . 1615-6110 . 309 . 20. 10.1007/s00606-023-01854-2 . 2023PSyEv.309...20M . free .
  15. 2023-02-27 . Petr . Bureš . etal . Evolution of genome size and GC content in the tribe Carduinae (Asteraceae): rare descending dysploidy and polyploidy, limited environmental control and strong phylogenetic signal . 185–213 . 2024-10-18 . Preslia . 2570-950X . 95 . 1. 10.23855/preslia.2023.185 . free .
  16. Book: Host, Nicolaus Thomas . 1831 . Beck . Vienna . Cnicus pauciflorus . 445–446 . Flora austriaca . 2024-10-18 . 2 . C. foliis serratis spinoso-ciliatis dorso villosis, inferioribus petiolatis subovatis sinuatis, superioribus oblongo-ovatis amplexicaulibus lanciniatis, anthodii squamis lineari-subulatis patulis. ... In Styria superiore in alpibus judenburgensibus inter saxa ad Bürger-See. In Hungaria in sylvis Comitatus Marmarosiensis sub alpe Petrosa. Fl. Julio, Augusto. Per. Radix crassa, fibris validis aucta. Caulis erectus, altissimus, striatus, setis inspersus et lanugine tectus. Folia ampla, serrata, spinulis ciliata, facie subglabra, dorso villo arachnoideo tecta; radicalia, et caulina inferiora petiolo auriculato affixa, subovata, sinuata; superiora oblonga, in lacinias inaequales divisa; his proxima sensim minora, minus divisa; summa minima. Flores cernui. Pedunculi languinosi. Anthodium constat squamis supra medium patentibus. Corollulae purpurascunt. Filamenta pubescentia. Semen striatum, nitidum. . la.
  17. Nyman 1879. "C. pauciflorum Spr. – Carn. Croat. Hung. Banat. Transs. Galic. Bucov. Serb. Bosn. Herceg.(r.). Etiam Cephal. m. Oenos sec. Rchb. f. Sec. cl. Naegeli hybr. est (C. Eristhali heterophyllum). – C. helenioides Bmg. * C. ambiguum K. (non All.) Cnicus ambigus Schleich. cat. 1807. Tyrol. etc." (in Latin) [C. pauciflorum Spr. – Carn. Croatian Hung. Banat Trans. Gaelic Bukov. Serbian Bosnia Herceg. (r.). Even Cephal. m. Oenos sec. Rchb. f. Sec. cl. Naegeli hybr. is (C. Eristhali heterophyllum). – C. helenioides Bmg. * C. ambiguum K. (not All.) Cnicus ambigus Schleich. cat. 1807. Tyrol. etc.]
  18. Reichenbach 1853. "In Radelwäldern der Marmaros an dem Felsen Petrosa bei Borsa in Ungarn Kitaibel. Rottenmannertaurn in Steiermark Sauter! Um Triest soll es von Herrn v. Tommasini gessamelt woren sein. (Am M. Oena auf Cephalonian Mazziari!)" (in German) [In the Radelwalden of the Marmaros on the Petrosa rock near Borsa in Hungary Kitaibel. Rottenmannertaurn in Styria Sauter! It is said to have been collected around Trieste by Mr. v. Tommasini. (On M. Oena on Cephalonian Mazziari!)]
  19. Book: 1905 . Georges . Rouy . Georges Rouy . Julien . Foucaud . Julien Foucaud . Edmond Gustave . Camus . Edmond Gustave Camus . Geneva Botanical Garden . Asnières-sur-Seine . Flore de France: ou, Description des plantes qui croissent spontanément en France, en Corse et en Alsace-Lorraine . 9 . 84 . 2024-10-18 . Le Cirsium pauciflorum Spreng. (1826), Cnicus pauciflorus Waldst. et Kit. (1805), bien postérieur au C. pauciflorum Lamk.)1778), devra prendre le nom de Cirsium Waldstinii Ruoy. . fr . Flora of France: or, a Description of the Plants that Grow Spontaneously in France, Corsica and Alsace-Lorraine.
  20. 1908 . Eugen Johan . Khek . J. J. Reiff . Karlsruhe . Seltene Cirsienbastarde aus Steiermark . 33–46 . 2024-10-18 . Allgemeine botanische Zeitschrift für Systematik, Floristik, Pflanzengeographie . 14 . 3 . Cirsium pauciflorum Sprengel β. ramosum E. Khek.: C. pauciflorum Scop. ist bei Trieben sehr häufig, immer aber zeigt dasselbe ungeteilte Stengel. Im vergangenen Jahre traf ich zum erstenmale Exemplare, eren Stengel verzweigt waren, oder bei welchen aus den Blattachseln lange Seitnäste entsprangen, welche wieder gehäufte Köpfchen trugen. Es handelte sich dabei keineswegs um putierte Formen (1. Juli 1907). . de . Rare Cirsium Hybrids from Styria.
  21. Book: Nyman, Carl Fredrik . 1879 . Carl Fredrik Nyman . Officina Bohliniana . Örebro . C. pauciflorum . 408 . Conspectus florae Europaeae, seu enumeratio methodica plantarum phanerogamarum europae indigenarum, indicatio distributionis geographicae singularum . 2024-10-18 . 2 . C. pauciflorum Spr. – Carn. Croat. Hung. Banat. Transs. Galic. Bucov. Serb. Bosn. Herceg.(r.). Etiam Cephal. m. Oenos sec. Rchb. f. Sec. cl. Naegeli hybr. est (C. Eristhali heterophyllum). – C. helenioides Bmg. * C. ambiguum K. (non All.) Cnicus ambigus Schleich. cat. 1807. Tyrol. etc. . la . Survey of Flora of Europe, or a Methodical Enumeration of Native Phanerogamous Plants of Europe, Indicating Geographical distributions of Individuals.
  22. Web site: 2006–2024 . Werner . Greuter . Werner Greuter . Cirsium greimleri . 2024-10-18 . Euro+Med PlantBase.
  23. 1889 . presented 1889-05-17 . Michael . Ritter von Eichenfeld . Kaiserlich-Königliche Zoologisch-Botanische Gesellschaft . Vienna . Botanischer Discussionsabend am 17. Mai 1889 . 2024-10-18 . Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien . 39 . 68–70 . Herr Dr. M. R. v. Eichenfeld besprach und demonstrierte nachfolgende Pflanzen, welche sämmtlich auf einer zwei Stunde von Judenburg in Steiermark liegenden Voralpenwiese der Seethaler Alpen von ihm und dem Judenburger Floristen Herrn B. Przybylski, welcher diesen Standort entdeckt hat, gesammelt worden waren. ... Cirsium pauciflorum Spr., welches sich auf der erwähnten Wiese in grösserer Anzahl findet; diese Pflanze wurde auch, etwa drei Stunden von diesem Standorte entfernt, in der Nähe der ungefähr 1700–1800 m hoch gelegenden Winterleiten-Seen von Herrn Przybylski in grösserer Anzahl aufgefunden. . de . Botanical Discussions on the Morning of 17. May 1889.
  24. 2018 . Kurt . Zernig . Christian . Berg . Durch das Winterleitenkar: Floristische Kleinode der Seetaler Alpen . 171–188 . 2024-10-18 . Tuexenia . 0722-494X . 11 . de . Through the Winterleitenkar: Floristic Gems of the Seetal Alps.
  25. Book: 2005 . Božo . Frajman . Martin . Turjak . Društvo študentov biologije . Ljubljana . Poročilo o delu botanične skupine . 11–26 . Raziskovalni tabor študentov biologije, Lovrenc na Pohorju 2005 . 978-961-91041-8-7 . 2024-10-18 . sl . Exploration Camp of Biology Students, Lovrenc on Pohorje 2005.
  26. 2008-01-10 . 2007 . Boštjan . Surina . Marash . Rakaj . Založba ZRC . Ljubljana . Subalpine Beech Forest with Hairy Alpenrose (Polysticho-lonchitis-fagetum rhododendrotosum hirsuti subass. nova) on Mt. Snežnik (Liburnian Karst, Dinaric Mts) . 195–208 . 2024-10-18 . Hacquetia . 1854-9829 . 6 . 2. 10.2478/v10028-007-0011-y . free .
  27. 2015 . Đorđije . Milanović . Jugoslav . Brujić . Vladimir . Stupar . Vojin . Bucalo . Jovan . Travar . Rade . Cvjetićanin . Vaskularna flora planine Klekovače u Bosni i Hercegovini . 15–83 . 2024-10-18 . Glasnik Šumarskog fakulteta Univerziteta u Banjoj Luci . 2303-694X . 23 . sr . Vascular Flora of the Klekovača Mountain in Bosnia and Herzegovina.
  28. Book: 2016 . Slađana . Petronić . Nataša . Bratić . University of Life Sciences-SGGW . Warsaw . Asteraceae in the flora of the Jahorina Mountain (Bosnia and Herzegovina) . 1895–1904 . https://www.cabidigitallibrary.org/doi/pdf/10.5555/20173053905 . VII International Scientific Agriculture Symposium, "Agrosym 2016", 6–9 October 2016, Jahorina, Bosnia and Herzegovina: Proceedings . 978-99976-632-7-6.
  29. 1996 . Dmitar . Lakušić . Unija bioloških naučnih društava Jugoslavije . Zemun . Pregled flore Kopaonika (J.Z. Srbija, Jugoslavija) . 1–35 . 2024-10-18 . Ekologija . 0531-9110 . 31 . 2 . Survey of the flora of Mt. Kopaonik (S.W. Serbia, Yugoslavia) . sr.
  30. Book: Drescher, Anton . 2018 . Kolline und montane Waldgesellschaften in der illyrisch geprägten Südwest-Steiermar . 38–43 . Vom Om Buchen-Urwald zum Kastanienhain – Waldlandschaften im illyrischen Entfaltungszentrum der mitteleuropäischen Buchenwälder . 2024-10-18 . Virgin forests with beech and chestnut groves – forested landscapes in the illyric developmental centre of Middle European beech forests . de.
  31. 1996 . Vladimir . Stevanović . Analysis of the Central European and Mediterranean orophytic element on the mountains of the W. and CentraI Balkan Peninsula, with special reference to endemics . 77–97 . 2024-10-18 . Bocconea . 1120-4060 . 5 . 1.
  32. 1993 . 1990–1991 . Vladimir . Stevanović . Marjan . Niketić . Dmitar . Lakušić . Institut za botaniku i Botaničke bašte Univerziteta . Belgrade . Distribution of the Vascular Plants in Yugoslavia (Serbia, Montenegro) and Macedonia I . 33–54 . 2024-10-18 . Glasnik Instituta za botaniku i Botaničke bašte Univerziteta u Beogradu . 0351-1588 . 24–25.
  33. 2016-04-28 . Ágnes . Mosolygó-Lukács . Gábor . Sramkó . Sándor . Barabás . Levente . Czeglédi . Molecular genetic evidence for allotetraploid hybrid speciation in the genus Crocus L. (Iridaceae) . 121–136 . 2024-10-18 . Phytotaxa . 1179-3155 . 258 . 2. 10.11646/phytotaxa.258.2.2 .
  34. Book: Лукьяновна Харадзе, Анна . 2001 . originally published 1963 in Russian . A. K. . Dhote . Anna Kharadze . Cirsium . 51–213 . https://www.biodiversitylibrary.org/page/29988913 . Flora Unionis Rerumpublicarum Socialisticarum Sovieticarum . 28 . Science publishers . . 9781886106451.
  35. 2019-01-24 . 2018 . Mitja . Zupančič . Branko . Vreš . Fitogeografska oznaka Slovenije . 159–211 . 2024-10-18 . Folia Biologica et Geologica . 2335-2914 . 59 . 2 . 10.3986/fbg0048 . free . sl . Phytogeographic Characteristics of Slovenia.
  36. 2020-01-05 . Владимир . Николаевич Годин . Распространение гинодиэции у цветковых растений . 236–252 . 2024-10-18 . Ботанический журнал . 2658-6339 . 105 . 3 . 10.31857/S0006813620030023 . Distribution of Ginodioecy in Flowering Plants . ru.
  37. 2003-02-11 . Jacqui A. . Shykoff . Sergios-Orestis . Kolokotronis . Carine L. . Collin . Manuela . López-Villavicencio . Effects of male sterility on reproductive traits in gynodioecious plants: a meta-analysis . 1–9 . 10.1007/s00442-002-1133-z . Oecologia . 0029-8549 . 135. 1 . 12647098 . 2003Oecol.135....1S .
  38. 2011-03-31 . Emmanuelle . Billard . Mathilde . Dufay . How much better are females? The occurrence of female advantage, its proximal causes and its variation within and among gynodioecious species . 505–519 . 2024-10-18 . Annals of Botany . 1095-8290 . 109 . 3. 10.1093/aob/mcr062 . 21459860 . 3278283 .
  39. Book: Benz, Robert . 1922 . Geographische Zeitschrift . Wiesbaden . Vorarbeiten zu einer pflanzengeographischen Karte Österreichs: Die Vegetationsverhältnisse der Lavanttaler Alpen . 2024-10-18 . Abhandlungen der zoologisch-botanischen Gsellschaft in Wien . 13,2 . de . Preliminary Work on a Phytogeographical Map of Austria: The Vegetation Conditions of the Lavanttal Alps.
  40. 1900 . 1899 . Josef . Murr . Josef Murr . J.J. Reiff . Karlsruhe . Einiges Neue aus Steiermarlc, Tirol und Oberösterreich . 23–24 . Allgemeine botanische Zeitschrift für Systematik, Floristik, Pflanzengeographie etc. . 5 . 2024-10-18 . Ergigeron ... glabratus Hoppe et Hornsch. ... Standort: Gottstahlgraben bei Mautern an einer einzigen Stelle auf Felsen und in Felsritzen bei ca. 1300 m in Gesellschaft von Cerastium lanatum Lam., Pleurospermum Austriacum Hoffm., Artemisia laxa (Lam.) Fritsch, Cirsium pauciflorum Spreng. und C. Scopolianum Schultz Bip. (=C. Eristithales × pauciflorum)..
  41. 1913 . Günther . Beck von Mannagetta und Lerchenau . Günther Beck von Mannagetta und Lerchenau . Akademie der Wissenschaften . Vienna . Vegetationsstudien in den Ostalpen III . 2024-10-18 . Sitzungsberichte der Akademie der Wissenschaften Mathematisch-Naturwissenschaftliche Klasse . 0371-4810 . 122 . de . Vegetation Studies in the East Alps III.
  42. 2023 . Tijana . Ilić . Nevena . Kuzmanović . Snežana . Vukojičić . Dmitar . Lakušić . The alpine scrubs and dwarf heaths of the Balkan Peninsula - an exceptional center of floristic richness and endemism . 145–161 . 2024-10-18 . Botanica Serbia . 1821-2638 . 47 . 1. 10.2298/BOTSERB2301145I . free .
  43. Book: Čarni, Andraž . 2007 . Vegetation of forest fringes on shallow dolomite bedrock in Central Slovenia . 237–248 . https://www.researchgate.net/publication/372958241 . Зборник на трудови посветени на академик Кирил Мицевски: по повод 80-годишнината од раѓањето . Makedonska akademija na naukite i umetnostite . Skopje . 9789989101809 . Collection of Papers Devoted to Academician Kiril Micevski.
  44. 2013 . Igor . Daskobler . Andrej . Rozman . Andrej . Seliškar . Forest and scrub communities with green alder (Alnus viridis) in Slovenia . 95–185 . 2024-10-18 . Hacquetia . 1854-9829 . 12 . 2. 10.2478/hacq-2013-0012 . free .
  45. 2013 . Bruno . de Foucault . Gilles . Corriol . Contribution au prodrome des végétations de France: les Mulgedio alpini – Aconitetea variegati Hadac et Klika ex Klika 1948 . 49–87 . 2024-10-18 . Le Journal de Botanique . 2741-4884 . 61. 10.3406/jobot.2013.1173 . fr . Contribution to the Prodrome of the Vegetation of France: the Mulgedio alpini – Aconitetea variegati Hadac et Klika ex Klika 1948.
  46. Book: 2005 . Jurij . Dobravec . Tone . Wraber . MOPE . Alpinska in subalpinska travišča na karbonatnih tleh (EU_6170) . 200–286 . https://www.ckff.si/dokumenti/n2k/200_286.pdf . Opredelitev območij evropsko pomembnih negozdnih habitatnih tipov s pomočjo razširjenosti značilnih rastlinskih vrst . sl . Ljubljan . Definition of Ranges of Important European Unforested Habitat Types by the Distribution of Type Plant Species.
  47. 2020-02-13 . Martin . Vavrinec . Geografická analýza mezidruhové hybridizace rodu Cirsium ve střední Evropě (Diplomová práce) . 2024-10-18 . cs . Geographical Analysis of Interspecies Hybridization of the Genus Cirsium in Central Europe (Practice for Diploma).
  48. 2018-05-09 . Petr . Bureš . Aleš . Knoll . Ester . Michálková . Petr . Šmarda . Jakub . Šmerda . Martin . Vavrinec . Cirsium greimleri: a new species of thistle endemic to the Eastern Alps and Dinarides . 105–134 . 2024-10-18 . Preslia . 2570-950X . 90 . 2. 10.23855/preslia.2018.105 . free .

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