Viola chelmea

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Viola chelmea
Viola chelmea subsp. vratnikensis (Syn .: Viola vilaensis Hayek), wild habitat in the sub-Adriatic Dinarides (Orjen)

Viola chelmea subsp. vratnikensis (Syn .: Viola vilaensis Hayek), wild habitat in the sub-Adriatic Dinarides ( Orjen )

Systematics
Rosids
Eurosiden I
Order : Malpighiales (Malpighiales)
Family : Violet family (Violaceae)
Genre : Violets ( viola )
Type : Viola chelmea
Scientific name
Viola chelmea
Boiss. & Hero.

Viola chelmea is a species of the genus violets ( Viola ) fromthe violet family (Violaceae).

It is an endemic - relictic mountain plant of the tree-free zones of the Mediterranean mountains of south-eastern Europe, which is distributed over an area split into individual mountain groups from Greece , Albania , Macedonia , Montenegro , Bosnia and Herzegovina to Croatia . Viola chelmea is adapted in its habitat to dry climates and locations and thus differs from all other European violet species in the Viola section . As one of the morphologically more original species, it is thus placed as the only European representative of the group of the so-called Lignosae Becker . As a special feature within the genus, the individual endemic species of the Lignosae among the Eurasian violets show an overall very low variability. Due to its rarity, its taxonomy as well as its morphological and ecological properties have not been adequately researched.

description

Vegetative characteristics

Viola chelmea is a perennial stemless (lat. Acaulescent from Acaulis) herbaceous plant , which has a short-limbed, woody and thick rhizome that extends vertically into the earth, from which the rosette-like leaves and the flowers in their axils arise; it reaches heights of growth of about 2.5 to 8 centimeters; on this more or less perpendicular basic axis develop laterally a larger number of also short-limbed, strong, upright or ascending branches that stand together like a tuft and, just like the rhizomes, have rosettes of leaves at their ends.

In a basal standing Rosette, long-handled (4-5 cm) true leaves are at ground weak heart-shaped or triangular broad- lanceolate , and are 1 to 3.5 centimeters long and 0.5 to 2 centimeters wide (the ratio length / width: one to 1.5 times); the stalks of the fully grown leaves are preserved for a long time, so that on older plants beneath the new leaves the long, fibrous remnants of petioles of the previous year's leaves can be seen and give the plants a characteristic appearance.

Leaves and flower stalks are hairy protruding; the hairiness is both seasonal and population dependent; Spring leaves are less hairy than summer leaves and in southern Greek populations the individuals in the Peloponnese were more hairy than those in Euboea; The fringes of the narrow, lanceolate stipules are slightly shorter than the wide stipules.

Like all representatives of the Series Eflagellatae, it does not form stolons (above-ground runners) within the Viola subsection .

  • The leaves of Viola chelmea ssp. vratnikensis sit on long petioles.

  • Flower and triangular leaf

Generative characteristics

Viola chelmea forms chasmogamous flowers in spring and blooms from May to early June, depending on the location. Although the plants are predominantly found at altitudes above 1000 m, they bloom as early as the beginning of June, as Nicolaus Bornmüller observed on the Chelmos in 1928 at altitudes between 1900 and 2200 m.

The stemless plants mostly have multi-headed flower stalks that are 3 to 5 centimeters long; the bracts of the flower stalks are below the center; odorless, hermaphrodite flowers are zygomorphic and 10 millimeters in size; the flowers have an upwardly curved, 3 to 5 millimeter long spur, which is darker than the crown. The color of the petals of the flowers varies depending on the breed. Specimens from the Peloponnese are violet, those on Euboea are pale blue-violet to almost white and have a striking pattern. In the subsp. vratnikensis , different white forms have been observed from the sub-Adriatic Orjen in Montenegro, as well as with the variety vilaensis in the Kučka krajina on the Montenegrin border with Albania. For the type locality of the subsp. vratnikensis in Velebit, however, purple flowers with indistinct veins were described as typical. Pen curved gooseneck-like manner on the base, merging at the end with no appendage, and in a short, horizontal or more upwardly directed scars Beak, mostly bald ovary, fruit stalks upright.

During the growing season, kleistogamous flowers are also regularly formed, which open over the year depending on the favorable conditions. The heavy and hairless capsules of the kleistogamous flowers are prostrate and on stems about 2–3 cm long. They usually contain only three to six seeds with a large funiculus . The seeds, which are white when dry, have a distinct oil-containing elaiosome . The length of the seeds is 2.4 to 2.7 mm, the largest width 1.3 to 1.5 mm.

The seeds are spread by ants because they like to eat the nutritious appendix (elaiosome). The ants transport the seeds over long distances and thus help the plant to spread.

The number of chromosomes is 2n = 20.

  • The flower is bearded on the side walls. Potential pollinators are guided to the nectar in the spur through the markings.

  • White-violet form ( Viola chelmea ssp. Vratnikensis )

  • spur

  • Viola chelmea diaspores consist of hard-shell seeds and attached elaiosomes.

etymology

It was named after the Chelmos Mountains in the north of the Peloponnese. Heldreich first found this in 1848 above the Styx waterfall at 7000 feet (approx. 2200 m).

Occurrence and socialization

The Chelmos with the Styx waterfall

The species is irregular, but widespread over a large area. Its south-eastern European area (ssp. Chelmea ) includes parts of the Pindus Mountains, the Peloponnese and, away from the east coast of mainland Greece, more isolated the island of Evia , as well as with the subspecies vratnikensis the more or less coastal Dinaric Mountains between Croatia and northern Albania . To the east, populations in Macedonia ( Jablanica ) have now also been collected.

Viola chelmea grows as a deep-rooted light plant on debris or stony slopes in karst mountains of the Western Balkans. Typically it is Oro-Mediterranean above the pine dry open forest of snake skin pine and black pine , as well as those of the Greek fir .

It colonizes sunny pebbles and dry lime grass between 1400 and 2200 m mostly in the vicinity of snow pits or in the light shade of young pines.

Locations at their northern limit of distribution in Croatia are significantly lower and were given between 500 and 700 m. The nominate form, on the other hand, occurs in Greece between 1300 and 2200 m mostly only more frequently above the tree line. Preference is given to calcareous rendzines that are low in nutrients and rich in bases ( Rendzic Leptosol according to World Reference Base standards ) and rock sites. In the Alto-Mediterranean depth, however, it is also common in the tongue basins of former Pleistocene glaciers. Here the locations are on deeper and more humid lime- brown soils (Chromic Luvic Cambisol), species-rich (Oro) Alti-Mediterranean mats with long snow cover.

Overall, the species is found only occasionally but regularly in the higher elevations of the limestone mountains, but is very rare, especially at the northern limit of distribution in Velebit . With regard to their socialization, the species has a broad ecological amplitude. In addition to pure rock sites on raw humus soils, in which it appears as a pioneer plant in carts , it is associated with Primula elatior and Muscari botryoides as well as Betonica alopecuros on steeper slopes and ridges of the upper tree line . In fat meadows it occurs with Betonica serotina ssp. on.

Evolution and ecology

Herbarium evidence with rhizomes from the Orjen Mountains. In addition to the flower shoot, two underground shoots are formed. This allows the plant to colonize poorly consolidated locations on coarse gravel.

The observed variation of allozymes as well as morphological differences in studies of individual populations of the Greek nominate form (ssp. Chelmea ) showed both morphologically and allozymatically remarkable individuality. In addition, the data of the Chelmos, Kyllini and Euboea (Dirphys) populations examined by Thomas Marcussen (2003) showed that the differences are correlated with geographical isolation. He interpreted the data to the effect that this variation was a consequence of the geographical restriction to mountain habitats. Evolutionary race formation through area fragmentation can therefore be promoted by the development of celestogamous inbreeding, in which losses of the original genetic variation have resulted over a few millennia due to ongoing genetic drift. Marcussen postulates a presumed differentiation of the individual populations for the postglacial epoch after the last glacial peak in the Würm . Due to the lowering of the altitude distribution, mountain habitats were less fragmented in the Würme Ice Age, which means that the areas of mountain plants formed contiguous territories, while the current situation of the distribution of Viola chelmea in relict mountain locations further favors the development of local races.

From an ecological point of view, Viola chelmea is a colonist of rubble and stone corridors of the oromediterranean level and occurs in the northern part of the area also in snake-skin pine as well as sparse fir and spruce forests, in the southern part of the area in Greek fir forests . Adaptations to the Mediterranean climate are given by the lack of runners , small habitus and woody strong rhizome in the form of a deep rhizome . It can also colonize rendzinen and initial or raw floors.

Seed formation does not only take place sexually in the species, the species also regularly forms cleistogamous flowers that appear in later growth periods. Chasmogamous flowers are only formed at the beginning of the growing season. The capsules of the kleistogamous flowers are prostrate and can even remain in the ground. These are non-explosive capsules. For seed dispersal is myrmecochory characteristic. However, there are always no mechanisms of spreading through runners in the group, as shown by many other groups of violets.

With the other types of subsect. Viola, with the exception of the Viola odorata group, Viola chelmea is an element of the meridional - submeridional mountain vegetation. Preservation of the rosette of leaves during the winter of the subsekt. Viola can be understood as a growth characteristic of Mediterranean elements . In general, Viola chelmea is a mesophyllous mountain plant that spreads on locations with little competition, but it has developed striking adaptations to dry periods due to its small stature, the size of the wedge-shaped leaves, a partially woody branched trunk and the lack of stolos. However, there are subterranean branches where roots are not formed. It also settles gravel areas in the mountains. These short ascending shoots below the leaf rosette can develop into flower shoots in subsequent years, which develop next to the mother shoot or, if it dies, replace it. The basal axis sections are also slightly woody.

Systematics

Subspecies and varieties

Distribution map in the sub-Adriatic Orjen (ssp. Vratnikensis )

The taxonomic delimitation of the endemic Viola chelmea belonging to the Efflagelatae Becker section is still uncertain. According to today's view, the species is distributed over the Balkan Peninsula in two subspecies:

  • Viola chelmea Boiss. & Hero. subsp. chelmea
  • Viola chelmea Boiss. & Hero. subsp. vratnikensis Gáyer et Degen

The nominate form subsp. chelmea only in the Pindus in Greece, Euboea and the Peloponnese , the subsp. vratnikensis between the Velebit and the Galičica . Only the length of the fringes (cilia) of the stipules (stipples) serve to differentiate the subspecies. The subsp. vratnikensis longer, finely fringed cilia. The flower color of the nominate form (ssp. Chelmea ) and ssp. vratnikensis is indicated as blue pale violet with only indistinct veins. In the increase made by Trinajstić (1975) the ssp. vratnikensis to the species Viola dinarica , two subspecies - ssp. dinarica , ssp. vilaensis differentiated. For the subspecies ssp. dinarica , purple flowers without venation are indicated as a diagnostic feature. For the subspecies vilaensis from Montenegro (Orjen, Prokletije) pale purple and white-flowered specimens with more pronounced veining.

  • Topotype of the nominate form from Chelmos by Wolfgang Lippert (June 20, 1986)

  • Specimen Viola chelmea ssp. vratnikensis on Orjen by Lujo Adamović (July 1907)

  • White variety (ssp. Vratnikensis ), Bijela gora - Orjen Mountains

  • White-flowered specimens are always darkly veined and come with ssp. vratnikensis Gyer et Degen such as the variety vilaensis Hayek

Locus classicus of the vilaensis variety , on the Vila mountain in the Kučka krajina on the Rikavac lake (Rikavačko jezero)

Viola vilaensis Hayek from the NW Prokletije near the Rikavčko jezeros in the Kučka krajina below the summit of the Vila (2093 m) was described as a high-alpine variety . It is placed in the Flora Europaea with Viola chelmea . In this case, the stipules are slightly hairy at the tip; white-flowered specimens have also been observed more frequently next to the bluish-violet specimens. In addition, according to Hayek, it has no finely papillary, usually more clearly heart-shaped leaves on either side, and also smaller flowers. Overall, however, the few morphological differences are of little taxonomic importance. In the Flora Europaea it is therefore listed as a variant of the nominate form, for which only the slightly hairy stipples at the tip are listed as the only distinguishing feature.

  • Type Viola vilaensis Hayek. Collected by Ignaz Dörfler in 1914 on the border between Montenegro and Albania (Herbarium WU - Vienna Biological University)

  • Document plant Dörflers to Viola vilaensis Hayek ( syn.iola chelmea )

  • Dörfler's information on the location

Another unclear find concerns the viola prenja Beck described by Günther Beck von Mannagetta and Lerchenau in 1886 by Prenj . This was only collected once and can therefore not be clearly assigned to this day (as of 2014) due to a lack of comparisons. While Arpad Degen (1914) and Wilhelm Becker (1925) suspected it probably belonged to Viola chelmea , Hayek (1918) drew a relationship to Viola pyrenaica Ram. in front. Karl Maly (1923) suspected, after he was able to confirm in the gorge of Prača Viola prenja for Bosnia, that Beck's Viola prenja might just be a variety of Viola pyrenaica . In the Flora Europaea, Viola prenja is regarded as probably belonging to Viola pyrenaica . The flowers of Viola prenja were described by Beck as fragrant, which is not the case for Viola chelmea .

Family relationships

According to the classification established by Wilhelm Becker in 1925, the species Viola chelmea was placed in its own series Lignosae within the Nomimium section in the genus Veilchen ( Viola ). With another ten species it is also assigned to a relict group of the series Eflagellatae . These are all montane to high alpine species that are distributed disjointly in Central Europe, North Africa and the Middle East. In the series Eflagellatae , four temperate species ( Viola ambigua , Viola collina , Viola hirta and Viola thomasiana ) contrast with several relic species (including Viola chelmea , Viola libanotica , Viola pyrenaica and Viola sandrasea with four other species from Asia Minor) with discontinuous occurrences.

  • Viola libanotica Boiss., North Lebanon, Cedar Forest

The other species of the Lignosae series belong to the closest relationship of Viola chelmea , all of which are only distributed disjointly in the eastern Mediterranean region. These endemic violets find their center of diversity on the peninsula of Asia Minor, where five species occur, as well as on the Balkan peninsula, where, in addition to Viola chelmea, two forms that are still taxonomically unclear ( Viola vilaensis , Viola prenja ) are included. It is true that they may not be a phylogenetic group, but rather an ecological group that have developed parallel adaptations to drought. The endemic and with the exception of Viola libanotica Boiss. mostly relatively late (1938–2003) newly described species of violets from the series Lignosae Asia Minor are Viola bocquetiana Yıld. , Viola isaurica Contandr. & Quézel , Viola kizildaghensis Dinç & Yıld. , Viola sandrasea Melch. and Viola yildirimlii Dinç &. Bağcı .

Taxonomic classification

Morphological systems

The view of the systematic position of the species has changed several times. Becker introduced Viola chelmea into the newly created group of Scapigerae (= Acaules ) as Lignosae for the first time in 1910 . In 1925 he revised his system and now included viola chelmea , viola prenja , viola libanotica and viola vilaensis in the lignosae . Becker cites the woody rhizome, the upright fruit stalks and the horizontal or more upwardly directed scarab as the main features of the series.

Melchior (1939) and Wilhelm Rössler (1943) came to a different, much more natural classification . Rössler evaluated the occurrence or absence of phloroglucose herbal cells in the leaves of European violets for the group classification. Inclusions were always present in the Caulescentes (= Rostratae ) and Repentes , on the other hand, the "Exkretzellen" were missing in the Acaules (= Uncinatae ) and the Lignosae . Rössler considers the Lignosae to be original relatives of the Acaules . Melchior expressed the same view on the basis of comparative morphological studies. He emphasizes the close phylogenetic relationship of the Lignosae to the Eflagellatae of the Acaules .

According to Melchior, the morphological shoot structure, the lignification of the axes, the rather primitive formation of stylus and stigma, the low variability of the species belonging here and their disjoint distribution are among the characters of the Lignosae . The shape of the style of the Melchior in Viola chelmea Boiss. et Heldr. including ssp. vratnikensis Gay. et Deg. , Viola libanotica Boiss. and Viola vilaensis found Hayek as similar, provided the basis for him to retain the Lignosae , which he however placed in the closest relationship to the Eflagellatae , in which they form their own series. Melchior assumes that closer phylogenetic relationships exist between the Lignosae and the Eflagelatae , in which the Lignosae form an old kinship circle. As a relict group and strict "oreophytes", the lignosae were no longer able to spread after the ice ages and are therefore limited to disjoint occurrences in mountains.

Alexander Schmidt confirmed the view of Melchior and Rössler, for which he also cites the seeds with long elaisomes, which are formed and scattered in lying capsules. The chromosome number 2n = 20 of Viola chelmea investigated by Schmidt is also widespread among the Acaules ; poorly differentiated chromosomes form the rule in the Acaules .

The simplest distinguishing feature from the other Eurasian violets in the Viola section are the small triangular leaves, as well as the overall low growth habit and the small flowers.

Genetic systems

According to comparative genetic studies of allozymes from three populations in Viola chelmea ssp. chelmea in Greece (Chelmos, Dirphys, Kyllini), the species showed no special affinity to other species in the Viola section .

However, weak support for Schmidt's 1964 view of a putative relationship to Viola pyrenaica Ramond exists in a private allele between the two species. However, it also showed a connection in a private allele to Viola hirta L. and Viola ambigua Waldst. & Kit.

However, in addition to the genetic narrowing of the disjoint distribution, Marcussen suspects hybridization processes as a modifying element in the evolution of Viola chelmea , for which he found hybridization processes in the hybrids of Viola chelmea : Viola alba ssp. alba x chelmea and Viola chelmea x odorata lists examples.

type

The type of Viola chelmea ssp. chelmea was collected on July 29, 1848 in the Peloponnese by Theodor von Heldreich (Heldreich no, 2153) at the Chelmos ( rare in regione nivali supra fontem Stygis , 7000). The original protologue of the species was created by Pierre Edmond Boissier and Theodor Heldreich in 1854.

Wilhelm Becker took Viola chelmea 1910 from one collection contained Guenther Beck of Mannagetta and Lerchenau (16 June 1894) for the Velež in the Herzegovina (Montenegro sic). Árpád Degen found Viola chelmea on July 10, 1906, also on the Orjen in Montenegro , but initially considered it to be part of Viola prenja Beck . After F. Dobiasch found one in 1909 at the Vratnik Pass in the Velebit Mountains above Senj , Degen and Julius Gayer revised the violas that had been collected in Croatia and Montenegro (at that time still in Dalmatia) and presented them as a subspecies vratnikensis Viola chelmea . As type evidence of the ssp. vratnikensis the specimen collected by Dobiasch in Croatia was taken. For a collection of Dörfler on the Montenegrin-Albanian border in Prokletije , August von Hayek finally set up the superfluous Viola vilaensis .

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literature

  • Marjan Niketić, Pavle Cikovac, Zoltán Barina, Dániel Pifkó, Ljupčo Melovski, Šemija Duraki, Gordana Tomović 2015: Viola chelmea and Viola jooi ( Violaceae ), new species for the flora of Serbia and their distribution in the Balkan Peninsula and the Carpathians. Bulletin of the Natural History Museum, 2015, 8: 49-74, Belgrade. (PDF)
  • Thomas Marcussen, Liv Borgen, Inge Nordal: Population differentiation and hybridization in the Greek endemism Viola chelmea ssp. chelmea . In: Violets of Subgenus Viola in Europe: variation, evolution and systematics. Faculty of Mathematics and Natural Sciences, University of Oslo 2003.
  • August von Hayek 1918: Viola vilaensis Hayek. Memorandum Akad. Wiss. 94: 154. Vienna.
  • Arpad Degen 1914: About the occurrence of a breed of Viola chelmea Boiss. et Heldr. in Dalmatia and Croatia. Magyar Bot. Lapok 13: 309 (1914), Budapest

Individual evidence

  1. ↑ Muhittin Dinç 2009: Comparative morphological and palynological study on poorly known Viola sandrasea and is closest relative V. kizildaghensis . Biologia, 64/1, 81-87. (Springer: PDF)
  2. ↑ Thomas Marcussen 2003: 'Population Differentiation and Hybridization in the Greek endemism Viola chelmea ssp. chelmea . Thomas Marcussen 2003: Violets of the Subgenus Viola in Europe - Variation, Evolution and systematics. Dissertation, Oslo University.
  3. ↑ Hans Melchior 1939: A new violet from SW Anatolia and the phylogeny of shoot development within the Nomimium section . Feddes Repertorium, 46, 39-42.
  4. ↑ Okamoto, M., Okada, H. & Ueda, K. 1993: Morphology and chromosome number of Viola pilosa and ist systematic position. Taxon, 42, 781-787
  5. ↑ Alexander Schmidt 1964: On the systematic position of Viola chelmea Boiss. et Heldr. ssp. chelmea and V. delphinantha Boiss. Reports of the German Botanical Society, 77, 256-261.
  6. ↑ Kit Tan & Gregoris Iatrou 2001: Endemic Plants of Greece - The Peloponnese . Gads Forlag, Kobenhavn. P. 210
  7. ^ Wilhelm Becker 1925: Viola LA Engler & K. Prantl 1925: The natural plant families . P. 367
  8. ↑ Hans Melchior 1939: pp. 40–41
  9. ↑ a b Hans Melchior 1939: p. 41
  10. ↑ a b c Thomas Marcussen 2003: p. 4
  11. ↑ Alexander Schmidt 1964: On the systematic position of Viola chelmea Boiss. et Hldr. ssp. chelmea and V. delphinantha Boiss. Reports of the German Botanical Society, 77, p. 257
  12. ↑ Muhittin, D. & Yildirimli, S. 2002: A new species of Viola (Violaceae) from Turkey. Botanical Journal of the Linnean Society, 138, 483-487. P. 485
  13. ↑ Alexander Schmidt 1964: p. 258
  14. ↑ Alexander Schmidt 1964: p. 257
  15. ↑ Marcussen, T., Borgen, L. & Nordal, I. 2003: Population differentiation and Hybridisation in the Greek endemism Viola chelmea ssp. chelmea . Thomas Marcussen 2003: Violets of the Subenus Viola in Europe - Variation, Evolution and systematics (Dissertation Oslo University).
  16. ↑ Christian Bräuchler, Pavle Cikovac 2010: Vegetational diversity in a hyper-karstic, hyper-humid oro-Mediterranean environment . 08/2010; In proceeding of: 40th Anniversary Conference of the Ecological Society of Germany, Austria and Switzerland, Volume: 40, Page 405-430, Held at Justus Liebig Universität Gießen, The future of Biodiversity
  17. ↑ Arpad Degen 1914: About the occurrence of a breed of Viola chelmea Boiss. et Heldr. in Dalmatia and Croatia. Magyar Botanikai Lapok, 13, 309-311.
  18. ↑ Arpad Degen 1937: Flora Velebitica - Volume II . Hungarian Academy of Sciences, Budapest. P. 439
  19. ↑ Kit Tan & Gregoris Iatrou 2001: p. 210
  20. ↑ Thomas Marcussen 2003: Population differentiation and Hybridisation in the Greek endemsm Viola chelmea ssp. chelmea - Paper V. Thomas Marcussen 2003: Violets of the Subenus Viola in Europe - Variation, Evolution and systematics. Dissertation University of Oslo. P. 4
  21. ↑ Thomas Marcussen 2003: p. 5
  22. ↑ Kit Tan & Gregorias Iatrou: Endemic plants of Greece - the Peloponnese. Gads Forlag, Copenhagen 2001. p. 210
  23. ↑ Meusel, H., Jäger, E., Rausert, S. & Weinert, E. 1978: Comparative Chorology of Central European Flora - Volume II . Gustav Fischer, Jena. Pp. 28-29
  24. ↑ a b c Thomas Marcussen 2003: p. 7
  25. ^ Raus, T. 1986: Viola LS 608-640 in Strid, A. 1986: Mountain Flora of Greece . Cambridge, Cambridge University Press.
  26. ↑ Alexander Schmidt 1964: On the systematic position of Viola chelmea Boiss. et Heldr. ssp. chelmea and V. delphinantha Boiss. Reports of the German Botanical Society, 77/1, 256–261, October 1964 (Wiley: PDF)
  27. ↑ Arpad Degen 1914: p. 310
  28. ^ DH Valentine, H. Merxmüller, A. Schmidt 1968: Viola L. In: TG Tutin, VH Heywood, NA Burges, DH Valentine, SM Walters, DA Webb (eds.) 1968: Flora Europaea - Volume 2: Rosaceae to Umbelliferae . 1968, Cambridge University Press, Cambridge, 270-282. ISBN 0-521-06662-X . P. 273 (Google Books)
  29. ↑ Ivo Trinajstić 1975: Analitićka Flora Jugoslavije . 3, 6, pp. 47-48
  30. ↑ August von Hayek 1918: Contribution to the knowledge of the flora of the Albanian-Montenegrin border area . Memoranda of mathem.-naturw. Class, 94, p. 54 (country museum: PDF)
  31. ^ Flora Europaea - Google Books
  32. ^ Günther Beck 1886: Flora of South Bosnia and the Bordering Herzegovina - Volume III . P. 81 (99), plate IV, Figs. 1-4. (State Museum: PDG)
  33. ↑ MUHİTTİN DİNÇ, YAVUZ BAĞCI & ŞİNASİ YILDIRIMLI 2003: A new species of Viola L. (Violaceae) from South Anatolia. Botanical Jornal of the Linnean Society, 141, 477-484 A new species of Viola L. (Violaceae) from South Anatolia - 2003 - Botanical Journal of the Linnean Society - Wiley Online Library
  34. ^ Wilhelm Becker 1925: Viola LA 363-376. Engler & K. Prantl 1925: The natural plant families 21, Leipzig. P. 376
  35. ↑ Alexander Schmidt 1964: pp. 258-259
  36. ↑ Hans Melchior 1939: p. 42
  37. ↑ T. Marcussen & L. Borgen 2000: Allozymic Variation and relationships within Viola subsection Viola (Violaceae). Plant systematics and Evolution, 223, 29-57

Web links

Commons : Viola chelmea  - Collection of images, videos and audio files