A distinctive group of species allied to Taraxacum danubium (T. sect. Erythrosperma, Compositae-Crepidinae): a taxonomic revision

Within Taraxacum section Erythrosperma, several relatively distinct species groups are recognized. One of them, characterized by leaves with numerous, usually patent, very narrow lateral segments, frequently with their most distal part dilated, and outer phyllaries patent or arcuate-recurved, often with tips approaching the involucre base, with narrow whitish borders, includes species similar to the Central European Taraxacum danubium. This group comprises nine species, the majority of them (five species) being confined to the Balkan Peninsula. Another diversity centre of the group is the southernmost Ukraine. The most widespread species is T. persicum, extending from Iran and the lower river Volga to Central Europe. A detailed taxonomic revision of the whole group is presented, and descriptions, illustrations and lists of specimens studied are given. Two Balkanic species are newly described, the name T. danubium is lectotypified, another three names are relegated to the synonymy of the accepted names, and an unclear name, T. pineticola, is discussed. The species concept in Taraxacum is briefly expounded.


Introduction
Taraxacum sect. Erythrosperma (H. Lindb.) Dahlstedt (1921) ranks among the most diverse sections in this complicated genus. More than 215 accepted names (and about 40 synonyms) are reported to belong to this section, and, in almost whole of its European and Near Eastern distribution range, its species diversity varies between ten and twenty species per medium-sized region (Štěpánek and Kirschner 2012).
The diagnostic characters of T. sect. Erythrosperma include low growth, usually a well developed tunic of dry remnants of old petioles, deeply divided leaves, often with narrow lateral segments, involucre with outer phyllaries patent, usually arcuate-recurved or arcuate-reflexed, sometimes subappressed or erect, short, linear-lanceolate, lanceolate to ovate-lanceolate, usually with narrow, inconspicuous borders, sometimes borders more distinct but always narrow, phyllary apex usually minutely corniculate, and achenes usually 3.2-4.0 (-4.5) mm long, of various colourations, most often reddish, brown and combinations of the two colours (but frequently also variously fulvous or light greyish stramineous-brown), body densely spinulose above, most often subabruptly but also subgradually to ± abruptly narrowing into usually cylindrical cone most often 0.6-1.5 mm long.
Taraxacum sect. Erythrosperma is related to T. sect. Erythrocarpa Handel-Mazzetti Kirschner and Štěpánek 2022b), and, on the basis of a relative similarity, also to T. sect. Dissecta van Soest, T. sect. Obliqua Dahlst., and more remotely to T. sect. Suavia Kirschner & Štěpánek (see also a comparison in Kirschner and Štěpánek 2022a). In terms of morphology, the latter section is easily distinguished by the almost orbicular to broadly ovate outer phyllaries and bigger fruits. The section Erythrocarpa, on average, has longer fruits (longer than 4.5 mm, often over 5 mm), more robust growth and longer, often appressed outer phyllaries with broader and / or more distinct paler borders. Taraxacum sect. Dissecta is very similar, but most species have much broader pale border to the outer phyllaries of a broader shape (most often broadly ovate to ovate-lanceolate), achene cones are usually more conical and shorter. The section Obliqua was extended recently to cover both the original couple of species (T. obliquum and T. platyglossum) and plants of the T. pyrenaicum group (all names and their authors follow Kirschner et al. 2008), and their most conspicuous structural difference is the growth in lax cushions (multiple root heads with a leaf rosette each); this group, nevertheless, awaits a revision. As regards the species of T. sect. Erythrosperma occurring in north-central and Northern Europe, there is an extremely helpful handbook with descriptions and a detailed photographic documentation of high quality (Wendt and Øllgaard 2015), covering more than 50 species.
In addition to the similar sections, there are also several species morphologically intermediate between sections, probably of hybridogenous intersectional origin. The group of T. langeanum Dahlst. represents a link between T. sect. Erythrosperma and T. sect. Palustria (Lindb. fil.) Dahlst., and T. lacistophylloides Dahlst. is intermediate between T. sect. Taraxacum and T. sect. Erythrosperma.
To understand the methods of the present taxonomic revision, it is useful to summarize the current state of knowledge of T. sect. Erythrosperma, and the level of exploration of major regions within its distribution range. There are substantial differences among regions in this respect. The most profoundly explored area is Scandinavia and Northwestern Europe, including the Baltic Countries, the United Kingdom, the Netherlands and Belgium, and adjacent territories of Central Europe, mainly Germany, Switzerland, Poland, Slovakia and Czechia. The rest of the geographical range has been examined much less thoroughly, although there are areas covered by several important studies, mainly in the Mediterranean (Greece, Bulgaria, Corsica). Most of the eastern and southeastern regions are quite underexplored in this respect (European Russia, Ukraine, Turkey, Iran).
In general, the taxonomic knowledge of T. sect. Erythrosperma is similar to the situation in other, less complicated plant groups in the middle of the 19th century -names published in local floras are evaluated in revisions, with frequent synonymizations, a picture of interspecific relationships is being drawn, and basic features of species' geographical and ecological ranges are established.

The concept of species in Taraxacum
There are several major phenomena and processes controlling the variation patterns encountered in Taraxacum (primarily, they are coexistence of agamospermy and sexuality, fixed hybridity, and polyploidy associated with asexual reproduction). They operate both generally, at the level of taxonomic entities, and in a spatially specific manner.
The main consequence of varied modes of reproduction and variation patterns (Table 1) is that Taraxacum species are remarkably different when their population heterozygosity and genotype diversity are considered (after Richards 1997;Hughes and Richards 1988: 170): Table 1 Impact of reproduction system on population features in Taraxacum 1 strongly depending on the population and migration histories 2 in Taraxacum, see e.g. Zeisek et al. (2015), Kirschner et al. (2013) 3 in Taraxacum, see e.g. Zeisek et al. (2015), Kirschner et al. (1994), Richards (1988, 1989) 4 in Taraxacum, see e.g. Kirschner et al. (2013Kirschner et al. ( , 2016; in van der Hulst et al. (2003) it is shown that a local mixture of agamospermous species ('population') exhibits a high genotype variation, relationships between diploid and triploid taxa were studied by Šuvada et al. (2012)  The three reproduction systems above roughly correspond to the 'types' of species. Other factors to be considered are (a) history of the population structure, (b) spatial and phylogenetic proximity of sexual and agamospermous entities, and (c) ecological differentiation among species (and sections). On this basis, we recognize allogamous, variable sexual species, such as T. erythrospermum Bess. s. str. or T. serotinum (W. & K.) Fisch., or distinctive autogamous species, such as T. besarabicum (Hornem.) Hand.-Mazz. or T. aristum Markl., or diplosporous agamospermous species (the absolute majority of polyploids in Taraxacum).
In Taraxacum, moreover, a background for the taxonomic evaluation of the above phenomena is formed by a low level of structural differentiation (in terms of morphology) and a relatively high number of entities to be considered, all of which makes the Taraxacum study rather complicated.
As regards the group of T. danubium A. J. Rich., we deal with nine, exclusively agamospermous entities treated as species. It is a noteworthy fact that T. danubium itself frequently coexists with diploid sexual populations of T. erythrospermum s. str. without losing its identity as no hybrids were observed.

Concept of Taraxacum species groups
Agamospermous taxa within a section are not evenly or randomly distributed in the multidimensional network based on their character states. They are apparently clustered in species groups, not always equally distinctive but usually recognizable on the basis of a certain character combination; the most conspicuous groups are recognized, although not formally, as aggregates. These aggregates serve as a practical tool for the taxonomic evaluation of populations studied and for setting up hypotheses of mutual relatedness of agamospermous taxa.
A more profound knowledge of T. sect. Erythrosperma makes it possible to revise whole species groups, a concept used in Štěpánek and Kirschner (2012) for the T. purpureomarginatum group and the T. rubicundum group.
One of the relatively distinct species groups of T. sect. Erythrosperma is the T. danubium group. It comprises species morphologically close to T. danubium A. J. Rich. distributed predominantly in Southeastern and Central Europe. During our studies of T. sect. Erythrosperma we also exploited our material from expeditions to other parts of Southeastern Europe, and results of these studies are summarized in the present taxonomic paper.

Material and methods
The methods of microspecies recognition are outlined in Kirschner et al. (2016Kirschner et al. ( , 2020. The methods of cultivation are described in Kirschner and Štěpánek (1993) and Kirschner et al. (2020). Approaches to the identification of modes of reproduction in dandelions are given in Kirschner et al. (2006Kirschner et al. ( , 2020. Morphological terms follow Ge et al. (2011); achene measurements include the cone. The term 'tunic' is used for a collar of dry remnants of old petioles on root head at plant base. The leaf and capitulum characters are recorded at full anthesis. Only features of ripe fruits are included in the descriptions (unripe achenes may differ in paler colour, less conspicuously developed spinulosity, shorter achene body length and beak length, etc.).
The present study is based upon field studies in Central Europe, Bulgaria and Ukraine. Further material was kindly provided by B. Kuzmanov, C. E. Sonck, I. Uhlemann and R. and E. Willing, partly also cultivated at the Experimental Garden of Institute of Botany, Czech Academy of Sciences. Voucher specimens are deposited at PRA (herbarium codes according to the Index Herbariorum (Thiers 2014+ of agamospermous species is largely an artificial concept, mainly based on mutual similarity. Morphological characters shared by the members of this group can be briefly summarized as follows: Plants small to medium-sized, relatively sparsely hairy, leaves deep to ± dark green, not spotted, rarely with little spots, ± regularly divided into numerous lateral segments, quite frequently with their most distal part dilated, interlobes short, usually not blotched, capitula ± small to medium-sized, stigmas discoloured, pollen absent or present, with grains variable in size, involucre dark green, outer phyllaries patent, arcuate-patent to arcuate-recurved, often with tips approaching the involucre base, relatively short, with a narrow, ± distinct pale or whitish border, corniculate near apex, inner phyllaries ± short, not elongating later, achenes red-brown to dark brown, with a dense spinulosity above, spinules ± erect-patent to suberect, achene body subabruptly to gradually narrowing into ± cylindrical cone 0.6-1.2 (-1.5) mm long. -Agamosperms (only triploids are known).
There are two (similarly artificial) groups that may be comparable with that of T. danubium. First, it is the T. lacistophyllum group of northwestern and Northern Europe; leaves of its members are often ± similar to those of T. danubium, but outer phyllaries are of a broader shape and usually loosely appressed to erect-patent at base and often arcuate distally (Wendt and Øllgaard 2015). Another group resembling that of T. danubium is the T. scanicum group (Vašut 2003;Vašut et al. 2005;Ge et al. 2011). Members of the latter group are characterized by an elongated, conspicuous leaf terminal segment, usually filiform-dentate or even narrowly lobulate at least at the base, and the outer phyllaries are ± irregularly patent-recurved to arcuate-patent. Further groups to be considered are the T. rubicundum group (see Štěpánek and Kirschner 2012) and the group of species similar to T. divinum Sonck and T. gracilens Dahlst., the latter differing primarily in the lower number of suberect to erectpatent outer phyllaries.
Identification key to the members of the T. danubium group A note on the method of identification Only well developed plants, both in flower and with ripe achenes, are suitable for identification. Preferably, several specimens should be evaluated to cover the variation and plasticity ranges of each species. Exsiccates: Taraxaca Exs., numbers 400-404 and 1050 (for introductory notes on the exsiccate series, see Kirschner and Štěpánek 1992).
Etymology Derived from the ancient Roman name of the Danube, Danubius or Danuvius.
Note on the typification The holotype designated by Richards (1970: 108) was reported to come from the Devínska Kobyla, a hill above the Danube in the vicinity of Bratislava, Slovakia. We studied the herbarium material in OXF and SLO (and other collections, including the personal collection of A. J. Richards, now deposited in NMW), and there is not any specimen from that locality under the name of T. danubium. We concluded that the original holotype is not extant. We therefore studied the protologue and the herbarium collections to find elements of the original material of this name, eligible for lectotypification. There is no direct protologue citation of a herbarium specimen other than the original holotype itself. However, several plants of T. danubium are mentioned in the text, from the Kováčovské kopce, from the Soroš Hill near Hlohovec, from Chotin, and from the Devínska Kobyla, all Slovakia (2n = 24 is recorded by Richards from three localities). There are three specimens, elements of the original material, eligible for the typification, all from the Soroš [Hill] near Hlohovec, collected by A. J. Richards and J. Májovský on 2 May 1968, and all with labels written by the latter collector. The specimen deposited in OXF was annotated by A. J. Richards as 'Taraxacum danubium A. J. Richards, holotype'. We therefore select the OXF specimen as the lectotype above, and the remaining two specimens (SLO) become isolectotypes.
Description Plants small to medium-sized, up to 15 (-20) cm tall. Plant base with dense brownish hairs; tunic developed. Leaves patent to erect-patent, sparsely arachnoid to glabrescent, slightly greyish green, leaf blade oblong-elliptical in outline, 3-12 (-20) × 1.5-3 (-4) cm, pinnatisect, with distinct teeth and lobules on interlobes, terminal segment relatively small, ca 1 cm long, in outer leaves helmet-shaped to ± triangular, sometimes distinctly trilobed, apex ± rounded, in inner leaves mucronate, mucro oblong to lingulate, slightly marrowed at the base, distal margin convex, sigmoid to concave, entire, proximal margin concave, entire; lateral segments (3) 4 (6) pairs, small, usually 0.7-2.0 × 0.5-1.0 cm, moderately recurved to patent, triangular or narrowly so, sometimes hamate, obtusely acute to rounded, with an abruptly dilated (drop-shaped) tip, with distal margin distinctly concave, entire, less often with minute teeth, proximal margin concave to straight, entire or with a single conspicuous tooth; interlobes usually short and entire, in inner leaves to 1.5 cm long, often with ± sparse long teeth, bordered dark; midvein pale green, in proximal 1/4 brownish pink; petiole narrow to very narrowly winged, usually pinkish to reddish purple. Scapes arachnoid, later only below capitulum and at base, light brownish pink to purplish pink at base. Capitulum small, to 2.5-3 cm wide, distinctly convex, deep yellow. Outer phyllaries 9-14 (18), narrowly lanceolate to ovate-lanceolate, 6-8 × 2-4 mm, ± regularly arcuate-recurved to subreflexed with tips close to involucre base, adaxially slightly glaucousgreen, rarely reddish, with a very narrow whitish, not very distinct border to 0.1 mm wide, abaxially dark bluish green, minutely corniculate near apex. Outer ligules flat, striped reddish to purplish grey-brown outside. Pollen abundant, irregular in size. Style long, stigmas exserted, discoloured, dark yellow-green to greyish green, with blackish pubescence outside, stigmas blackish when dry. Achenes deep dark red-brown, dark castaneous brown to deep purple-brown, relatively short and thick, (3.0-) 3.3-3.8 × 0.8-1.0 mm, body medium densely to densely spinulose in upper 1/2-1/3, spinules erect-patent, acute, to 0.3 mm long, ± abruptly narrowing into a narrow, cylindrical to subcylindrical cone (0.6-) 0.9-1.0 mm long, sometimes with 1(2) spinules at the base; beak (7-) 8-9.5 mm; pappus white to whitish, (5-) 5.5-6 mm long. Diagnostic notes Within the T. danubium group, this species could be confused with T. persicum only (because it is the only species of this group with the geographical range overlapping that of T. danubium). Central European T. persicum, however, lacks pollen (while pollen in T. danubium is abundantly developed). Moreover, T. persicum has narrower and more distinctly spinulose achenes, and more acute leaf lateral segments. The other members of this group are not sympatric with T. danubium, and the differences are summarized in the key above. In its general habit, T. danubium resembles certain populations of T. erythrospermum from Hungary, but the latter have shorter, less arcuate outer phyllaries, pollen grains of regular size, and a diploid chromosome number. Taraxacum arcuatum is another species that might be confused with T. danubium, but the shorter, erect-patent to suberect-arcuate outer phyllaries, paler stigmas and smaller achenes with a short cone are diagnostic for T. arcuatum. Leaf lateral segments with narrowed, linear middle part and the dilated apical part are also observed in T. lacistophylloides; the latter has conspicuously hairy leaves and light greyish stramineous-brown achenes.
Distribution and habitat Taraxacum danubium, as a morphologically distinctive taxon, was recorded in several important works (Dudáš et al. 2020;Wolanin and Musiał 2018;Trávníček et al. 2010;Vašut 2003) and its overall distribution is relatively well explored, with almost 300 localities known. It is confined to Central Europe, and specimens or reliable records come from northeastern Austria, northern Hungary, most of western and southern Slovakia and southern Poland, and there is a remote locality in Berlin, probably of secondary origin (Uhlemann 2003); the most frequent occurrence of T. danubium is recorded in the Czechia, which not necessarily must be a bias due to the most thorough exploration (Map: Fig. 2).
Taraxacum danubium grows on both calcareous and acid substrates; it is most frequently found in open dry grasslands, along paths and forest margins and clearings in open deciduous woodlands, dry, sandy sites in villages, rocky slopes, often also in old quarries and on castle hills. It is most often found in hilly landscapes, at low or moderate elevations, usually between (130-) 200 and 700 m a. s. l., with a maximum of 920 m recorded in Slovakia (Dudáš et al. 2020). Its IUCN conservation status is estimated as LC, because of the number of localities known, the size of the distribution range, possibility of secondary habitats and the protected status of a number of natural localities.
A selection of representative specimens [A more complete list of specimens studied is given in the Appendix, see also the map (Fig. 2) Exsiccata, no. 1017, PRA, no. det. 33516, etc.).
Etymology The epithet is introduced in honour of our colleague and friend, the late Bogdan Kuzmanov , a specialist in the taxonomy of the Bulgarian flora (Markova 1992).
-Agamosperm (on the basis of pollen analysis).
Diagnostic notes The most conspicuous feature of T. kuzmanovii is its leaf shape. The constricted lateral lobes with a dilated distal appendage of a triangular or rhombic shape is unique among members of the T. danubium group. The closest species is undoubtedly T. danubium itself. Taraxacum kuzmanovii can be distinguished by leaf lateral segments often subhamaterecurved, with the distal appendage, outer phyllaries less conspicuously recurved, more often arcuate-patent, with more distinct, longer horns near their apex, and by achenes with a longer cone and longer beak. Taraxacum danubium is absent from the Balkan Peninsula.
Distribution and habitat Although it is quite distinctive and coming from regions satisfactorily covered by Taraxacum gatherings, it is known from two localities in southern Bulgaria and northern Greece (the distance between the two sites is not more than 125 km). We therefore consider T. kuzmanovii a relatively rare species. It most often grows on dry stony slopes and sunny low-coverage grasslands, open dry woodlands, at relatively low elevations between 150-320 m. Its IUCN conservation status is estimated as VU.
Note For comparison, we mention a species collected at, or near these two sites, T. umbrosum Sonck, Kirschner and Štěpánek in Štěpánek and Kirschner (2015: 161). It was collected at several dozen localities in Bulgaria, Greece and North Macedonia, and the frequency of its occurrence is in a sharp contrast with that of T. kuzmanovii. It also documents the sampling density in Bulgaria and northern Greece, which is also obvious from the distribution map of T. lingulilobum below (Fig. 6).  Type P. Sintenis, It. Transcaspico-persicum 1900/1901, no. 1433(K, no. det. 8807, lectotype, fide van Soest, Catal. Erythrosperma, p. 42, 1966isolectotypes: L 43109, no. det. 19686, BM, no. det. 8446).
Diagnostic notes Taraxacum persicum, usually lacking pollen, is distinct in having triangular or broadly triangular, acute leaf lateral segments, outer phyllaries irregularly arcuate-patent, abaxially dark olivaceous-green to dark bluish green (blackgreen when dry) and pruinose, with a distinct but narrow whitish border. Its achenes are dark purplish brown to dark brown, small and slender, with cone 0.7-1.1 mm long. It is similar to T. egnatiae but leaves of T. persicum are not suffused bronze, interlobes are not blotched, nor bordered, as a rule, and outer phyllaries are distinctly bordered; achenes of T. persicum are darker, often pure dark brown.
Distribution and habitat Taraxacum persicum occupies a large geographical range extending from Central and Eastern Europe to lower Volga River and the Caspian Sea region. However, there are large gaps in our documentation of this distribution, and there is a possibility that the distribution has a really disjunctive nature. In Central Europe, the populations of T. persicum are small and easy to overlook, while in the east, this species may be one of the dominant members of T. sect. Erythrosperma. It grows in a variety of dry, relatively open habitats, including steppe, subsaline sandy sites, calcareous grasslands, etc., mostly at low elevations. Its IUCN conservation status is difficult to estimate because of rather fragmentary data; we consider the LC category as the most appropriate.
Etymology Named after Egnatia Street, Thessaloniki; Egnatia is a township in northern Greece.
Diagnostic notes Taraxacum egnatiae is very similar to T. persicum but differs in a number of features (summarized in Štěpánek and Kirschner 2018: 368), mainly the leaves suffused bronze, with blotched and bordered interlobes, usually ± patent outer phyllaries with a much less distinct and narrower border, floccose-arachnoid scapes, lighter coloured achenes with a longer pappus (6-7 mm long).
Distribution and habitat Taraxacum egnatiae is known from a number of localities in northern Greece, and there is a large disjunction between those sites and a locality on the slopes above Yalta, Crimea, Ukraine. Etymology The epithet 'ziwaschum' is a mangled attempt to derive the name from a local Crimean term Sivash (Sıvaş in Crimean Tatar language, Cивáш in Ukrainian), a wetland and salt lake area in northern and northeastern Crimea, Ukraine. The correct form would be sivasicum, as used in a number of plant names, but the epithet ziwaschum is not correctable, according to the ICN (Turland et al. 2018).
Note Taraxacum ziwaschum is one of the species names in sect. Erythrosperma based on the type gathering only. The 2-3 specimens at LE are of a relatively satisfactory quality and well preserved, and their interpretation is therefore possible. When the type material was compared with our rich material collected in southern Ukraine, it turned out that T. ziwaschum occurs in the famous steppe vegetation reserve, Askania Nova. Our material was used to complete and amend the species' description (the type gathering lacks fully ripened achenes, and only one leaf morphotype is represented). It might be mentioned that Doll (op. c.) considered T. ziwaschum as a member of sect. Scariosa, which is not supported by any morphological characters or other evidence.
Diagnostic notes Taraxacum ziwaschum is characterized by the absence of pollen, relatively short and dark outer phyllaries, and red-brown, long and very slender achenes with a very gradual achene body / cone transition, and a long thin cone. Taraxacum persicum is similar in its triangular, acute leaf lateral segments but differs from T. ziwaschum in achene colour and shape, narrower phyllary border, darker stigmas and a different leaf segment shape. In its general habit, T. ziwaschum approaches T. parnassicum, but the latter has very different achenes, shorter and ± unbordered, and less recurved or ± straight outer phyllaries. T. danubium is distinct in having pollen, a very characteristic leaf shape with dilated most distal parts of lateral segments, a narrower phyllary border, and thicker and shorter achenes. T. kuzmanovii is polliniferous, and has a narrower outer phyllary border.
Distribution and habitat Up to now, T. ziwaschum has been found at Ukrainian localities in the Henichesk and Kherson Districts. It grows in dry, substeppe, steppe or open sites on a lime-rich substrate, in lowland regions only. Its IUCN conservation status is estimated as NT; there is a low number of localities known but they are situated in protected areas.
Etymology The epithet is derived from the tongueshaped distal part of lateral segments of leaves.
Diagnostic notes Diagnostic features of this species include large brown-red achenes, 4.2-4.9 mm long (similar to those of T. sect. Erythrocarpa); it approaches T. kuzmanovii in this respect but can be distinguished by the complicated leaf segmentation pattern with variously long lobules and interlobe teeth. Taraxacum danubium is similar in the leaf shape and outer phyllary characters (with the exception of phyllary colour and less pronounced corniculation) but has achenes up to 3.8 mm long.
Distribution and habitat Taraxacum lingulilobum is a species known from northern Greece and the western part of the southernmost Bulgaria, according to the specimens available (Map: Fig. 6). Its occurrence in the neighbouring countries, e.g. North Macedonia, Albania or Turkey is probable. It grows in dry grasslands, dry slopes, rocky sites, often calcareous, margins of open oak woodlands and along paths, usually between ca 100 m and 2,000 m. Its IUCN conservation status is estimated as LC.
Etymology Derived from Epirus, Epeiros in Greek, a region in the promontories of the Pindus Mts.
-Agamosperm (on the basis of pollen analysis).
Diagnostic notes Taraxacum epirense is characterized by numerous, usually patent, leaf lateral segments, numerous, arcuate-recurved outer phyllaries and a dark brown achene colour. It differs from T. persicum in the achene colour and polliniferous anthers, from T. danubium by thicker achenes of a different colouration, darker stigmas and the leaf shape, and from T. kuzmanovii and T. lingulilobum by the achene size and colour and the leaf shape. Taraxacum epirense is a marginal member of the T. danubium group, probably a link between the latter and species around T. acutiusculum Sonck and T. poliochloroides Doll. Distribution and habitat Taraxacum epirense is relatively widely distributed in the Balkan Peninsula (Bulgaria, Romania, Greece), as pointed out by Štěpánek and Kirschner (2014). This distribution picture is corroborated by further localities from other regions of that area (Albania, Montenegro) and by further numerous localities from Greece. However, van Soest (1966)  Description Plants small to medium-sized, usually 9-13 cm tall. Root head simple, with a well developed dark brown tunic, and with a light greyish-brownish indumentum among petiole bases. Petiole narrow, unwinged (± narrowly winged in the outermost leaves), deep purple. Leaves erect-patent, usually 4-7 × 1-1.5 cm, ± greyish green, not spotted (interlobes ± blotched brown-purple), sparsely arachnoid, later glabrescent, leaf blade lanceolate to ± narrowly elliptic in outline, pinnatisect; terminal segment relatively small, usually 5-9 × 5-11 mm, triangular to broadly triangular, less often helmet-shaped, symmetric to asymmetric, obtusely acute to acute, distal margin subsigmoid or almost straight, entire or with a single asymmetrical incision, proximal margin subsigmoid to ± straight, entire or with a few minute teeth near the lobe base, basal lobules acute, subrecurved; lateral segments in 3-4 (5) pairs, opposite or alternate, relatively broad, usually 5-9 mm long, 4-7 mm wide at base, arcuate-recurved, almost subhamaterecurved, acute to subacute, distal margin convex to sigmoid, entire or seldom with 1-2 minute teeth, proximal margin subconvex, subsigmoid or subconcave, with a single distinct tooth or entire; interlobes short, medium broad, usually 1-3 × 2-3 mm, usually with a single broad tooth or several small teeth, interlobe margin raised, surface distinctly blotched brownpurple; midvein purplish proximally, pale distally. Scapes overtopping leaves, usually 6-10 cm long, suffused purple proximally, otherwise pale greenish, ± densely floccose-arachnoid. Capitulum yellow, relatively small, 1.5-2 cm wide, concave, with dense florets. Involucre ca 7 mm wide and ± rounded at base. Outer phyllaries 17-22, regularly (i.e. not in one row but evenly distributed) arcuate-patent to arcuate-recurved to recurved, lanceolate to narrowly lanceolate, 6-7 × 2-2.5 mm, adaxial surface greyish green, usually suffused purplish, abaxially blackish green (black when dry), border distinct, white, 0.1-0.3 (-0.4) mm wide, margin long ciliate, apex callose to corniculate; inner phyllaries 14-20, usually ca 12 mm long, ± unequally wide, with a darker, callose apex. Outer ligules subcanaliculate, striped purplish grey outside, with apical teeth grey-black, inner ligules subinvolute, short, apical teeth light purplish to grey-purple. Stigmas medium discoloured, yellow-green to greyish green, hairs of pubescence distally dark. Pollen absent (or a few deformed minute grains remaining in the anther tube). Achenes reddish brown to medium brown (castaneous-brown when unripe), 3.2-3.8 × 0.8-0.9 mm, achene body with ± dense erect-patent thin spinules to 0.2 mm long in the upper 1/4, otherwise tuberculate, subabruptly to ± abruptly narrowing in a narrow cylindrical cone 0.8-1.2 mm long (cone sometimes subconical at base and / or slightly thickened distally); beak 8.5-11 mm long, pappus dirty whitish-brownish, 5-6 mm long. Diagnostic notes Taraxacum honestum is characterized by a combination of pollen absence, relatively small, subconcave flower heads, outer phyllaries evenly distributed in an imbricate way, usually arcuate-recurved, with a distinct narrow border, and reddish brown to mid-brown achenes with a relatively short body and a long, cylindrical cone. In its leaf shape it somewhat approaches T. egnatiae but the above character combination is diagnostic.
Distribution and habitat It grows on dry, stony and sandy slopes with low-coverage grasslands. For the time being, it is known from a single macrolocality in the southern Pirin, Bulgaria, and it obviously is a relatively infrequent species in view of the relatively detailed exploration of the neighbouring areas of northern Greece and Bulgaria (on the other hand, it is a tiny plant easy to overlook). Its IUCN conservation status is therefore estimated as VU. Etymology Named after Annette Uhlemann, the wife of I. Uhlemann, an outstanding taraxacologist.
Diagnostic notes Taraxacum annetteae is a species distinct in having patent leaf segments, conspicuously arcuate-reflexed outer phyllaries suffused purple adaxially, dark discoloured stigmas and light brown achenes with long spinules. Within the T. danubium group, it can be distinguished by relatively small light brown achenes, the closest species being T. epirense.
Distribution and habitat Taraxacum annetteae is known from two, rather remote, sites in coastal Croatia (Krk Island, and the Starigrad vicinity). It grows in dry sparse grasslands along paths. Its IUCN conservation status is estimated as VU.
Specimen examined CROATIA.
Comments on Taraxacum pineticolaKlokov, Bot. Mater. Gerb. Bot. Inst. Akad. Nauk SSSR 16: 367. 1954 Finally, we have to mention T. pineticola, a name without any recent interpretation. It is known from the type gathering deposited in LE (see below), a relatively rich set of plants collected in a very late season (June) in the vicinity of Kharkov, Ukraine, too late for a safe identification or comparison. The plants have summer leaves that were preserved under the woodland canopy but do not give any indication as regards the standard leaf shape in the full blossom time (Fig. 10). The characters observed on the plants, particularly the dark stigmas, the lack of pollen, outer phyllaries recurved, ± narrowly bordered, achenes initially red-brown, later dark brown, do not exclude the T. danubium group (nor similar groups, such as the T. scanicum agg.). In conclusion, the name remains to be interpreted, pending the evaluation of erythrospermous plants from the locus classicus region.  Funding Open access publishing supported by the National Technical Library in Prague.