Jump to content

Cirsium greimleri

From Wikipedia, the free encyclopedia

Cirsium greimleri
1895 illustration by Johann Gottfried Hallier

Endangered  (IUCN 3.1)[1]: 122 [2]: 111 
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Asterales
Family: Asteraceae
Genus: Cirsium
Species:
C. greimleri
Binomial name
Cirsium greimleri
Bureš
Cirsium greimleri is located in Dinaric and Eastern Alps
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Cirsium greimleri
Range of C. greimleri (precise locations only).[3][1][4]
Synonyms[1]
Synonymy
  • Cirsium pauciflorum Spreng.
  • Cirsium waldsteinii Rouy ante 2018

Cirsium greimleri, German: Greimler-Kratzdistel lit.'Greimler's thistle',[5] is a species of flowering plant belonging to the family Asteraceae.[6]

Its native range is in Eastern Europe.[6] It was only described as a separate species in 2018, and belongs to the minority species discovered through karyological analysis.[1]

Etymology[edit]

Named after botanist Josef Greimler, whose chromosome count spurred the study that separated it from C. waldsteinii.[1]: 123 

Distribution[edit]

It is found in the Eastern[7] and Dinaric[8] Alps. A partial but detailed map is designated Obr. 32b in Vavrinec 2020, which also provides distribution maps for its hybrids.[2] A complete map was published in 2018, which includes flora-based ranges in the Dinarides, from which few precise coordinates exist yet.[9][1]: 111 

The more precise ranges it has been found on include the Ennstal Alps, Rottenmann and Wölz Tauern, Seckau Tauern, Seetal Alps,[10][11] Saualpe, Koralpe, Karawanks, Kamnik–Savinja Alps, Pohorje,[12] Julian Alps, Snežnik,[13] Srnetica, Klekovača,[3]: 36  Cincar, Vranica, Bjelašnica, Visočica, Treskavica, Zelengora, Jahorina,[14] Komovi, Golija, and Kopaonik.[15][1]: 122 

On its own, its distribution is similar to that of Cardamine waldsteinii,[16] and somewhat similar to those of Achillea clusiana,[17]: 84  Moehringia ciliata,[18]: 36  and Vicia oroboides.[19]: 171 

When taken together with C. waldsteinii, its circum-Pannonian distribution is similar to that of Crocus vernus agg.,[16] including a polyploidy factor.[20] Or to the acidophile species Hieracium transylvanicum.[19] With less overlap, to Centaurea uniflora agg. or Pilosella alpicola. Including the Pontic and Caucasus, to Koeleria eriostachya. Including part of the Apennines, to Aposeris foetida.[19]: 168 

Habitat[edit]

It is a subalpine[19][1] and high montane[1][2] species,[21][19] found from 800 to 2000 m above sea level.[1] It grows on the moist leeward 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.[1][22]

In the Ellenberg system of indicator values, C. greimleri scores 9 for light availability,[a] 5 for temperature,[b] 9 for climatic continentality,[c] 5 for soil moisture,[d] 5 for soil pH,[e] 8 for soil fertility,[f] 0 for soil salinity.[g][2] As calculated by these values, the coterminous Cirsium species with greatest ecological similarity was C. arvense, followed by C. heterophyllum and C. spinosissimum.[2] The species is calcifuge, but does sometimes grow on calcareous substrates.[1]

Description[edit]

1853 illustration by Carl Schnorr.
Representative specimen.

Root is an oblique, cylindrical rhizome.[1]

Plants grow to a height of 110-180[h] cm,[1] with the exception of a form that grows to 30 cm identified in 1902 and designated C. greimleri f. depressum by Eugen Johan Khek,[23] and sterile basal rosettes.[1] Growth is erect but nodding below the capitula.[1] Stems are shallowly ribbed.[1] Stems are simple or have few branches.[24] The form with branches was designated C. greimleri var. ramosum by Khek after he identified it in 1906.[25] Their colour is green or reddish green but pilose or subglabrous at the base, whitish green and sparsely arachnoid in the middle, and white and densely arachnoid below the capitula.[1] Leaves grow all along the stem.[24]

Basal leaves are 11-24[i] cm wide and 1-2[j] cm longer than wide in mature specimens.[1] Their morphology ranges from ovate, to orbicular,[24] to deltate especially in younger leaves. Peduncular leaves amplexicaul, often auriculate.[26] Their shallow pinnate lobes are doubly serrate to pinnatipartite, with weak yellowish to brownish-purple spinules up to 2 mm situated at the margins,[24] subglabrous to scattered pubescent above and densely arachnoid-lanate below.[24] All leaves are soft and herbaceous.[1][27]

Flowers are capitula, each with 1-8[k] flowers, solitary or corymbosely terminally clustered, rarely solitary on 1-5[l] lateral pedicels.[1] Involucre[m] dimensions are 13-21[n] mm long at flowering.[24] Phyllaries in 6-7[o] rows.[1] Individual bracts bracts flare out from the bud, with distinct vittae, the outer and inner bracts being distinguished by the presence and absence, respectively, of a visible spine.[24] Involucres are purplish brown to purplish black.[1]

Corolla length is 17-21[p] mm in hermaphrodites, 15-19[q] mm in females.[1] At full 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 (hermaphrodite) or ochre synantherium. The style is always whitish except for the ruby shortly bi-lobed stigma, which fades to deep crimson or brownish-purple.[r][1][29]

Strongly anemochorous, its fruits contain oblong, compressed, asymmetric greyish ochre achenes, 16-19[s] mm in hermaphrodites, 14-18[t] mm in females. They are attached to 4-5[u] mm pappi.[24] The pappus is plumose, whitish, or stramineous.[1][30]

Lookalikes[edit]

  • C. waldsteinii Rouy 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). This 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.[1]
  • C. hypoleucum DC.. Its nodding flowers are sometimes ruby red like those of C. greimleri, which is rare for the genus.[1] The leaves of C. hypoleucum, sometimes resembling C. waldsteinii more than C. greimleri,[1] 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,[21] and the involucres themselves are narrower. The idumentum of subcapitular stem is sparser.[1]
  • C. oblongifolium K.Koch. It has longer, more oblong leaves. Even the undersides of the leaves are glabrous.[21]
  • C. pseudopersonata Boiss. & Balansa ex Boiss.. It is distinguished by its flower colour, more regular for the genus than that of C. greimleri, visible green through sparse idumentum of subcapitular stem,[1] and glabrous leaf undersides.[21]
  • C. sychnosanthum Petr.. Even the undersides of the leaves are glabrous.[21]
  • C. uliginosum (M.Bieb.) Fisch.
  • C. carniolicum Scop.. Very similar to C. greimleri in vegetative stage, but with yellow to white flowers[v] with tougher phyllaries and spiny sepals, visible green through pilosity of subcapitular stem, and more distinctive spines on upper cauline leaves with subglabrous undersides.[1]
  • C. alpis-lunae Brilli-Catt. & L.Gubellini. A C. carniolum like species with spiny phyllaries and longer, stiffer leaf spines.[1]
  • C. latifolium Lowe has similarly wide, whole leaves, but little resemblance otherwise.[1]
  • A number of species have a similar flower colour but little resemblance otherwise: C. rivulare var. 'Atropurpureum' (Jacq.) All., C. borealinipponense Kitam., C. hachimantaiense Kadota, C. hidakamontanum Kadota, C. shimae Kadota, C. chokaiense Kitam., C. douglasii DC., C. occidentale (Nutt.) Jepson.[1]
  • 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).[1]: 124  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.[1]: 128–129 

Micromorphology[edit]

Stomatal guard cells are about 17[w] μm long and 9[x] μm wide.[1]

Life cycle[edit]

Perennial.[22][24][1]

Reproduction[edit]

Flowering is from late June to late July, to early August at sites of higher shade or altitude.[1]

Plants are gynodioecious,[31] 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 7-9[y]  mm, compared to female synantheria, at 4-6[z] mm. The same applies to styles, 20.0-25[aa] mm in hermaphrodites, and 18-22[ab] mm in females. Although both hermaphrodite and female stigmas can be straight, only female stigmas can be twisted, and usually are.[1]

The smaller size of female achenes relative to hermaphrodite achenes in this species is unusual.[1] 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.[32][33] Suggested explanations for the aberrance in C. greimleri and C. waldsteinii include larger achene count, higher germanation rates, and lower infestation with achene predators like the Tephritidae and Curculionidae.[1]

Hybridisation[edit]

C. greimleri is a diploid species with sporophytic chromosome number 2n=2x=34,[34] without variation.[1] Its somatic nuclear DNA size is about 1930[ac] Mbp.[1]

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.[2] A similar situation exists for C. alsophilum, C. bertolonii, and C. carniolicum.[35] 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.[1]: 122 

The following hybrids have been reported:[2]

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.[2] 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.[1]

Despite high range overlap, C. greimleri does not hybridise with C. arvense,[2] which may be due to intersectional incompatibility.[21]

Taxonomy[edit]

Internally, C. greimleri populations are more genetically distinct from one another than C. waldsteinii populations, whose intrapopulation diversity however is higher than C. greimleri.[1]: 110  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.[1]: 128 

Externally, a close relationship between C. greimleri, C. waldsteinii, and C. hypoleucum is suspected on morphological grounds.[1]: 124  The relationship between C. greimleri and C. waldsteinii has already been confirmed genetically, although an allopolyploid origin of C. waldsteinii is strongly suggested.[1]: 128  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.[35]

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.[36]

Taxonomic history[edit]

Precovery specimen collected by Michael Hölzl in 1837.

Its holotype[ad] was collected by Petr Bureš at 1739 m from 47°16′19″N 14°24′23″E / 47.271972°N 14.406250°E / 47.271972; 14.406250 in the Rottenmann and Wölz Tauern on 15 September 2015.

C. greimleri was considered part of C. waldsteinii until 2018.[1]

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, at Bürgersee near Judenburg, under the name Cnicus pauciflorus Kit..[37] 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.[38]

First published under its present genus as Cirsium pauciflorum Spreng. by Anton Eleutherius Sauter following its discovery in Rottenmann Tauern.[39]

In 1905, C. pauciflorum was recognised to be a younger homonym of C. pauciflorum Lam., which is now designated Carduus defloratus. Because of this, they assigned it a new name, C. waldsteinii.[40][1]

In 2018, following the discovery that Alpine-Dinaric populations of "C. waldsteinii" were invariably diploid, whereas Carpathian populations were invariably tetraploid, together with physical and genetic evidence of separation, the two populations were split into two species.[1]

Ecology[edit]

It grows in forests dominated by Acer pseudoplatanus, Alnus incana, Betula pendula, Corylus avellana, Fagus sylvatica, Larix decidua, Picea abies, Pinus cembra, and Sorbus aucuparia.[41][1]

It is often found in association with other subalpine tall forbs, such as Adenostyles alliariae, Athyrium felix-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 agg., Senecio ovatus, Struthiopteris spicant, Telekia speciosa, Vaccinium myrtillus, Valeriana tripteris, Veratrum lobelianum, and Veronica urticifolia.[42][1]

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.[42][43][19]

It is forms part of the Pinion mugo type of Krummholz vegetation in the Dinarics.[44]: 157 

It is sometimes an element of the Scabioso hladnikianae-Grafietum golakae Čarni association.[45]: 246 

It is a rare element of the Calamagrostion arundinaceae association,[46]: 56  the Huperzio-Alnetum viridis Mulgedio-Aconitetea Hadac & Klika ex Klika (1948) subassociation,[47]: 136 [46] the Polysticho-lonchitis-fagetum rhododendrotosum hirsuti Boštjan & Rakaj subassociation.[13]

It is also found in the associations EU 6170 Adenostylion,[48]: 245  Cirsio cani-Filipenduletum ulmariae, Caricion davallianae, and Veronico-Calitrichetum.[3]

Agriculture[edit]

The average TKW[ae] is 0.4 g, as calculated following the collection of seeds[af] for the Millennium Seed Bank in 2014.[49]: 12 

See also[edit]

Notes[edit]

  1. ^ Value L
  2. ^ Value T
  3. ^ Value K
  4. ^ Value F
  5. ^ Value R
  6. ^ Value N
  7. ^ Value S
  8. ^ [70-]110-180[-210]
  9. ^ [8.2-]11.5-23.3[-26.8]
  10. ^ [1.0-]1.2-2.1[-2.4]
  11. ^ 1-8[-12]
  12. ^ 1-5[-10]
  13. ^ In sensu Asteraceae.
  14. ^ [11.6-]12.9-21.0[-21.7]
  15. ^ [5-]6-7[-9]
  16. ^ [14.8-]17.3-21.0[-23.0]
  17. ^ [13.9-]15.1-18.6[-19.2]
  18. ^ Colour values are given in the Methuen system.[28]
  19. ^ [15.0-]16.0-19.0[-20.0]
  20. ^ [13.5-]14.5-18.0[-18.5]
  21. ^ [3.7-]3.9-5.0[-5.6]
  22. ^ Pinkish in the Pyrenean subspecies C. carniolicum subsp. rufescens.
  23. ^ [12.3-]16.5[-23.4]
  24. ^ [6.1-]9.2[-18.4]
  25. ^ [6.3-]6.7-9.3[-9.4]
  26. ^ [4.2-]4.3-5.7[-6.5]
  27. ^ [18.3-]20.0-25.0[-25.7]
  28. ^ [16.5-]18.0-22.4[-23.3]
  29. ^ [1827-]1929[-2125]
  30. ^ BRNU 658042
  31. ^ Thousand Kernel Weight, a popular value giving the dry weight of 1000 seeds.
  32. ^ Catalogued as AT-0-GZU-14 141.

References[edit]

  1. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc bd Bureš, Petr; Knoll, Aleš; Michálková, Ester; Šmarda, Petr; Šmerda, Jakub; Vavrinec, Martin (2018-05-09). "Cirsium greimleri: a new species of thistle endemic to the Eastern Alps and Dinarides". Preslia. 90 (2): 105–134. doi:10.23855/preslia.2018.105. eISSN 2570-950X.
  2. ^ a b c d e f g h i Vavrinec, Martin (2020-02-13). Geografická analýza mezidruhové hybridizace rodu Cirsium ve střední Evropě (Diplomová práce) (Thesis).
  3. ^ a b c Milanović, Đorđije; Brujić, Jugoslav; Stupar, Vladimir; Bucalo, Vojin; Travar, Jovan; Cvjetićanin, Rade (2015). "Vaskularna flora planine Klekovače u Bosni i Hercegovini". Glasnik Šumarskog fakulteta Univerziteta u Banjoj Luci. 23: 15–83. eISSN 2303-694X.
  4. ^ GBIF (2024). "Cirsium greimleri Bures Occurrence Download". Global Biodiversity Information Facility. doi:10.15468/dl.nac5jc.
  5. ^ Schratt-Ehrendorfer, L. (2022). "Cirsium". Rote Liste der Farn- und Blütenpflanzen Österreichs (3rd ed.). Retrieved 2024-07-04.
  6. ^ a b "Cirsium greimleri Bureš | Plants of the World Online | Kew Science". Plants of the World Online. Retrieved 2024-06-08.
  7. ^ Nyman, Carl Fredrik (1879). "C. pauciflorum". Conspectus florae Europaeae, seu enumeratio methodica plantarum phanerogamarum europae indigenarum, indicatio distributionis geographicae singularum. Vol. 2. p. 408. 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.
  8. ^
  9. ^ Greuter, Werner (2006–2024). "Cirsium greimleri". Euro+Med PlantBase.
  10. ^ Ritter von Eichenfeld, Michael (1889) [presented 1889-05-17]. "Botanischer Discussionsabend am 17. Mai 1889". 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.
  11. ^ Zernig, Kurt; Berg, Christian (2018). "Durch das Winterleitenkar: Floristische Kleinode der Seetaler Alpen". Tuexenia (11): 171–188. ISSN 0722-494X.
  12. ^ Frajman, Božo; Turjak, Martin (2005). "Poročilo o delu botanične skupine". Raziskovalni tabor študentov biologije, Lovrenc na Pohorju 2005 (PDF). Društvo študentov biologije. pp. 11–26. ISBN 978-961-91041-8-7.
  13. ^ a b Surina, Boštjan; Rakaj, Marash (2007). "Subalpine Beech Forest with Hairy Alpenrose (Polysticho-lonchitis-fagetum rhododendrotosum hirsuti subass. nova) on Mt. Snežnik (Liburnian Karst, Dinaric Mts)". Hacquetia. 6 (2) (published 2008-01-10): 195–208. doi:10.2478/v10028-007-0011-y. eISSN 1854-9829.
  14. ^ Petronić, Slađana; Bratić, Nataša (2016). "Asteraceae in the flora of the Jahorina Mountain (Bosnia and Herzegovina)". VII International Scientific Agriculture Symposium, "Agrosym 2016", 6-9 October 2016, Jahorina, Bosnia and Herzegovina: Proceedings. pp. 1895–1904. ISBN 978-99976-632-7-6.
  15. ^ Lakušić, Dmitar (1996). "Pregled flore Kopaonika (J.Z. Srbija, Jugoslavija)" [Survey of the flora of Mt. Kopaonik (S.W. Serbia, Yugoslavia)]. Ekologija. 31 (2): 1–35. ISSN 0531-9110.
  16. ^ a b Drescher, Anton (2018). "Kolline und montane Waldgesellschaften in der illyrisch geprägten Südwest-Steiermar". om Buchen-Urwald zum Kastanienhain – Waldlandschaften im illyrischen Entfaltungszentrum der mitteleuropäischen Buchenwälder [Virgin forests with beech and chestnut groves – forested landscapes in the illyric developmental centre of Middle European beech forests] (PDF). pp. 38–43.
  17. ^ Stevanović, Vladimir (1996). "Analysis of the Central European and Mediterranean orophytic element on the mountains of the W. and CentraI Balkan Peninsula, with special reference to endemics" (PDF). Bocconea. 5 (1): 77–97. ISSN 1120-4060.
  18. ^ Stevanović, Vladimir; Niketić, Marjan; Lakušić, Dmitar (1990–1991). "Distribution of the Vascular Plants in Yugoslavia (Serbia, Montenegro) and Macedonia I" (PDF). Glasnik Instituta za botaniku i Botaničke bašte Univerziteta u Beogradu (24–25) (published 1993): 33–54. ISSN 0351-1588.
  19. ^ a b c d e f Zupančič, Mitja; Vreš, Branko (2018). "Fitogeografska oznaka Slovenije" [Phythogeographic analysis of Slovenia]. Folia Biologica et Geologica. 59 (2) (published 2019-01-24): 159–211. doi:10.3986/fbg0048. eISSN 2335-2914.
  20. ^ Mosolygó-Lukács, Ágnes; Sramkó, Gábor; Barabás, Sándor; Czeglédi, Levente (2016-04-28). "Molecular genetic evidence for allotetraploid hybrid speciation in the genus Crocus L. (Iridaceae)". Phytotaxa. 258 (2): 121–136. doi:10.11646/phytotaxa.258.2.2. ISSN 1179-3155.
  21. ^ a b c d e f Лукьяновна Харадзе, Анна (2001) [originally published 1963 in Russian]. "Cirsium". Flora Unionis Rerumpublicarum Socialisticarum Sovieticarum. Vol. 28. Translated by Dhote, A. K. Science publishers. pp. 51–213. ISBN 9781886106451.
  22. ^ a b Reichenbach, Heinrich Gustav (1853). "Cirsium". 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. Vol. 15. pp. 67–101. Mächst auf grasigen Wiesen, in Wäldern. Sommer.♃
  23. ^ Khek, Eugen Johan (1905). "Floristisches aus Steiermark" (PDF). Allgemeine botanische Zeitschrift für Systematik, Floristik, Pflanzengeographie. 11 (3): 41–42. Cirsium pauciflorum Spreng. f. depressum. Durch 3 Jahre beobachtet. Eine 30 cm hohe, auffallend niedrige Form in mehreren Exemplaren bei Trieben.
  24. ^ a b c d e f g h i Werner, Klaus (1976). "Cirsium Miller". Flora Europaea. Vol. 4. pp. 232–242. ISBN 0-521-08717-1.
  25. ^ Khek, Eugen Johan (1908). "Seltene Cirsienbastarde aus Steiermark". Allgemeine botanische Zeitschrift für Systematik, Floristik, Pflanzengeographie. 14 (3): 33–46. 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).
  26. ^ Kitaibel, Pál; von Waldstein, Franz de Paula Adam Norbert Wenzel Ludwig Valentin (1803). "Cnicus pauciflorus". Descriptiones et icones plantarum rariorum Hungariae. Vol. 2. pp. 175–176.
  27. ^ Reichenbach, Heinrich Gustav (1853). "Cirsium". 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. Vol. 15. pp. 67–101. 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.♃)
  28. ^ Kornerup, Ann-Mari; Wanscher, J. H. (1989). Methuen Handbook of Colour. E. Methuen. ISBN 0413334007.
  29. ^ Fritsch, Karl (1905). "Blütenbiologische Untersuchungen verschiedener Pflanzen der Flora von Steiermark" (PDF). Mitteilungen des Naturwissenschaftlichen Vereines für Steiermark. 42: 267–282. ISSN 0369-1136. Cirsium pauciflorum Spr. ¶ Anfangs Juli 1905 übersendete mir Herr K. Pilhatsch aus den Judenburger Alpen blühende Exemplare von Cirsium pauciflorum Spr., sowie von den Hybriden dieser Art mit Cirsium Erisithales (L.) Scop., Cirsium palustre (L.) Scop. und Cirsium heterophyllum All. Da alle diese Formen in Bezug auf ihre Blüteneinrichtungen noch nicht näher untersucht sind, so nahm ich ihre Untersuchung vor. Möglich ist, daß infolge der Übersendung mit der Post und der nachfolgenden Einfrischung im Wasser die bei den einzelnen Formen im folgenden angegebenen Maße und Farben etwas von jenen abweichen, welche direkt am Standorte der Pflanzen festgestellt würden. ... ¶ 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. Im Tubus der Blumenkrone ist viel Honig enthalten. 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.
  30. ^ Reichenbach, Heinrich Gustav (1853). "Cirsium". 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. Vol. 15. pp. 67–101. 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.
  31. ^ Николаевич Годин, Владимир (2020-01-05). "Распространение гинодиэции у цветковых растений" [Distribution of Ginodioecy in Flowering Plants]. Ботанический журнал. 105 (3): 236–252. doi:10.31857/S0006813620030023. eISSN 2658-6339.
  32. ^ Shykoff, Jacqui A.; Kolokotronis, Sergios-Orestis; Collin, Carine L.; López-Villavicencio, Manuela (2003-02-11). "Effects of male sterility on reproductive traits in gynodioecious plants: a meta-analysis". Oecologia. 135 (1): 1–9. Bibcode:2003Oecol.135....1S. doi:10.1007/s00442-002-1133-z. ISSN 0029-8549. PMID 12647098.
  33. ^ Billard, Emmanuelle; Dufay, Mathilde (2011-03-31). "How much better are females? The occurrence of female advantage, its proximal causes and its variation within and among gynodioecious species". Annals of Botany. 109 (3): 505–519. doi:10.1093/aob/mcr062. eISSN 1095-8290. PMC 3278283. PMID 21459860.
  34. ^ Dobeš, Christoph; Kiehn, Michael; Vitek, Ernst (1996). "Beiträge zur Gefässpflanzen-Flora von Österreich: Chromosomenzählungen III". Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien. 133: 301–318. ISSN 1025-4749.
  35. ^ a b Michálková, Ester; Šmerda, Jakub; Plačková, Klára; Knoll, Aleš; Bureš, Petr (2023-05-24). "Hybridization may endanger the rare North Apennine endemic Cirsium bertolonii". Plant Systematics and Evolution. 309 (20). Bibcode:2023PSyEv.309...20M. doi:10.1007/s00606-023-01854-2. eISSN 1615-6110.
  36. ^ Bureš, Petr; et al. (2023-02-27). "Evolution of genome size and GC content in the tribe Carduinae (Asteraceae): rare descending dysploidy and polyploidy, limited environmental control and strong phylogenetic signal". Preslia. 95 (1): 185–213. doi:10.23855/preslia.2023.185. eISSN 2570-950X.
  37. ^ Host, Nicolaus Thomas (1831). "Cnicus pauciflorus". Flora austriaca. Vol. 2. pp. 445–446. 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.
  38. ^ Nyman, Carl Fredrik (October 1879). "C. pauciflorum". Conspectus florae Europaeae, seu enumeratio methodica plantarum phanerogamarum europae indigenarum, indicatio distributionis geographicae singularum. Vol. 2. p. 408. 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.
  39. ^ Reichenbach, Heinrich Gustav (1853). "Cirsium". 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. Vol. 15. pp. 67–101. 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!)
  40. ^ Rouy, Georges; Foucaud, Julien; Camus, Edmond Gustave (1905). Flore de France. Vol. 9. p. 84. 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.
  41. ^ Benz, Robert (1922). Vorarbeiten zu einer pflanzengeographischen Karte Österreichs: Die Vegetationsverhältnisse der Lavanttaler Alpen (PDF). Abhandlungen der zoologisch-botanischen Gsellschaft in Wien. Vol. 13, 2.
  42. ^ a b Murr, Josef (1899). "Einiges Neue aus Steiermarlc, Tirol und Oberösterreich" (PDF). Allgemeine botanische Zeitschrift für Systematik, Floristik, Pflanzengeographie etc. 5 (published 1900): 23–24. 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).
  43. ^ Beck von Mannagetta und Lerchenau, Günther (1913). "Vegetationsstudien in den Ostalpen III". Sitzungsberichte der Akademie der Wissenschaften Mathematisch-Naturwissenschaftliche Klasse. 122. ISSN 0371-4810.
  44. ^ Ilić, Tijana; Kuzmanović, Nevena; Vukojičić, Snežana; Lakušić, Dmitar (2023). "The alpine scrubs and dwarf heaths of the Balkan Peninsula - an exceptional center of floristic richness and endemism". Botanica Serbia. 47 (1): 145–161. doi:10.2298/BOTSERB2301145I. eISSN 1821-2638.
  45. ^ Čarni, Andraž (2007). "Vegetation of forest fringes on shallow dolomite bedrock in Central Slovenia". Зборник на трудови посветени на академик Кирил Мицевски: по повод 80-годишнината од раѓањето [Collection of Papers Devoted to Academician Kiril Micevski]. Makedonska akademija na naukite i umetnostite. pp. 237–248. ISBN 9789989101809.
  46. ^ a b de Foucault, Bruno; Corriol, Gilles (2013). "Contribution au prodrome des végétations de France: les Mulgedio alpini – Aconitetea variegati Hadac et Klika ex Klika 1948". Le Journal de Botanique. 61: 49–87. doi:10.3406/jobot.2013.1173. eISSN 2741-4884.
  47. ^ Daskobler, Igor; Rozman, Andrej; Seliškar, Andrej (2013). "Forest and scrub communities with green alder (Alnus viridis) in Slovenia". Hacquetia. 12 (2): 95–185. doi:10.2478/hacq-2013-0012. eISSN 1854-9829.
  48. ^ Dobravec, Jurij; Wraber, Tone (2005). "Alpinska in subalpinska travišča na karbonatnih tleh (EU_6170)" (PDF). Opredelitev območij evropsko pomembnih negozdnih habitatnih tipov s pomočjo razširjenosti značilnih rastlinskih vrst. pp. 200–286.
  49. ^ "Accessions for the Millenium Seed Bank, Royal Botanic Gardens Kew, 2014". Samentauschverzeichnis des Botanischen Gartens der Karl-Franzens-Universität Graz: Ernte 2014 [Seed list of the Botanical Gardens of the University of Graz: Harvest 2014] (PDF). pp. 14–20.

Literature[edit]