Knautia goecmenii Yıldırım, 2022

Yildirim, Hasan, Özdöl, Tuğkan, Can, Ilgin Deniz, Erdem, Seval, Oğur, Erdinç & Özüdoğru, Bariş, 2022, Knautia goecmenii (Caprifoliaceae): an unexpected caespitose suffrutescent new species from the Western Anatolia, Turkey, Phytotaxa 531 (2), pp. 97-110 : 99-106

Knautia goecmenii Yıldırım sp. nov. ( Figs. 1‒3 )

Type: — TURKEY. B1 İzmir, Kemalpaşa, Nif Mountain , slopes below the summit, 1400 m elevation, 11 September 2018, H.Yıldırım 7731 & E.Oğur (holotype: EGE 43195!; isotypes: EGE 43196!, HUB!, NGBB!, ANK!); paratype :— TURKEY. İzmir, Kemalpaşa, Nif Mountain , slopes below the summit, 1290 m elevation, 10 July 2019, H.Yıldırım 8094 (EGE 43198!) .

Diagnosis: — Knautia goecmenii has no known closely related species. It is easily distinguished by its distinct suffrutescent and caespitose habitus, with 22–152 stems at base; stems lacking any leaf rosette at base.

Description: —Suffrutescent, caespitose perennial herbs, 22–152 stemmed at base, with thick cylindrical taproot. Stems 20–60 × 0.1–0.4 cm, branched or unbranched, densely hairy; hairs intermixed with pubescent, hirsute, crisped and glandular hairs; glandular hairs much denser at upper stem. Leaves, densely intermixed with tomentose-pubescent and short hirsute hairs; leaf rosettes absent; lower cauline leaves 3–11 × 0.5–3.5 cm (including petiole), lyrate-pinnatisect, rarely oblanceolate-spathulate or linear to oblanceolate (if oblanceolate-spathulate or linear to oblanceolate then 2–3 deeply lobed or entire); upper cauline leaves 0.5–6.5 × 0.1–11 cm, sessile, linear to linear-oblanceolate or linearlanceolate, sometimes spathulate, rarely with one lateral segment, usually entire. Peduncles 4–18 cm long, densely glandular and pubescent hairy. Inflorescence solitary on each branch. Capitula 1–1.8 × 0.7–1.2 cm, 12–30 flowered, radiant. Involucral bracts 8–16, in 2 rows, 2–5 mm long, outer ones slightly wider than inner ones, ovate to lanceolate, sometimes linear-lanceolate, pilose. Receptacle pilose hairy. Receptacular bracts subherbaceous, 4–8 mm × 0.2–0.5 mm, outer bracts longer than inner bracts, linear. Involucel 1–2 mm long in lowering time and 3-5 mm in fruiting time, densely villose-pilose and minute hairy, with several tiny teeth at apex. Calyx cupuliform, 0.3–0.7 × 0.8–1.6 mm, pubescent-tomentose, with 8 setae; setae 0.7–2.5 mm long, short hairy. Corolla 6–12 mm long, very pale whitish-lilac, shortly villose-pilose hairy on tube, short pubescent-pilose hairy at throat; lobes 1–2 × 1–1.5 mm long, villose hairy at dorsally, glabrescent at apex. Stamens 6–8 mm long; anther ca. 2 mm long; filaments 4–6 mm long; pollen grains yellow. Style 0.5–1 cm long. Fruit 5–8 × 1–2 mm, dorsally compressed, linear, oblanceolate, mostly 2 horned, rarely 4 horned, if with 4 horn then two lateral ones conspicuously longer, straw coloured. Seed 3.5–5.5 mm long, shortly silky hairy, shiny green. ( Figs. 1–3 ).

Eponymy: —This new species is named in memory of Prof. Bayram Göçmen, who was an expert of Reptiles and produced outstanding contribution in the field of Protozoology. Unfortunately, he died tragically in cancer in March 2019. The Turkish name of this species is proposed as “efeçenberi”, according to the guidelines of Menemen et al. (2013).

Phenology: — Flowering time from July to September; fruiting time from August to September.

Ecology, distribution and conservational status: — Knautia goecmenii is a local endemic to the Nif Mountain, a part of Bozdağ Mountains series in İzmir, Western Anatolia, which belongs to the Mediterranean floristic region ( Fig. 5 ). It colonizes mountain slopes on calcareous substrates in Kemalpaşa district of İzmir province, preferably with a western and north-west orientation, at elevations between 1420 to 1505 m. It has only a single population. The new species has a very restricted distribution, with an area of occupancy (AOO) of 0.07 km 2, in which about 1700 individuals were observed. That territory is under pressure of overgrazing by sheep and goats along with anthropogenic effects due to road construction. Given all these threats and the limited distribution of the species, K. goecmenii is here evaluated as “Critically Endangered”, CR [B1+B2ab (ii,iii,v)] according to the IUCN (2016) criteria.

Pollen morphology: — Pollen dispersal unit was monad. The pollen grains are isopolar, radial symmetric, and triporate, spheroidal or oblate-spheroidal; outline in equatorial view is circular or oval to oblate, in polar view circulartriangular; large-sized. The long equatorial diameter (EL) was 97.15 ± 1.13 µm, short equatorial diameter (ES) was 92.27 ± 1.22 µm Sculpture exine is heteroechinate, spinulate-spinulose ( Fig. 6 ).

Phylogenetic trees: —The aligned ITS dataset included 43 sequences belonging to 42 species and was 754 bp long, of which 146 were potentially parsimony informative and 123 were variable but not informative. The phylogenetic backbone of the ITS tree coincides with the previous phylogenetic study dealing with the genus Knautia ( Rešetnik et al. 2014) . The results of phylogenetic analyses, as shown in Figure 6 , support the monophyly of the genus with maximum posterior probabilities (PP) and bootstrap support (BS) (PP=1.00 and BS=100). Concordantly with the results of Resetnik et al. (2014), K. orientalis Linnaeus (1753: 101) , from sect. Knautia , was sister to the remaining two sections, and all studied members of sect. Trichera formed a clade with 1.00 PP and 0.74 BS. The new species K. goecmenii fell into the highly supported monophyletic clade (PP=1, BS=100) with the members of sect. Tricheroides plus K. degenii Borbás ex Formánek (1895: 135) (sect. Knautia ).

Taxonomic relationships:— Among the 11 Knautia species distributed in Turkey, only K. involucrata Sommier & Levier (1893: 46) and K. drymeja Heuffel (1856: 53) are perennial. Knautia goecmenii differs from both latter species mainly by its many-stemmed and suffrutescent and caespitose habitus, as well as by many features such as leaf shape, indumentum and flower colour. Although K. goecmenii nests phylogenetically close to the annual Knautia byzantia, K. degenii , and K. integrifolia , it can easily be distinguished from them by several noticeable morphological characters (see details in Table 1 ).

Contrary to its outstanding morphology, the phylogenetic placement of K. goecmenii is largely concordant with geography and as explained above, all the species distributed Anatolia and the Balkan Peninsula fell into the same clade together with K. goecmenii . The relationship between Anatolian and Balkan biotas has long been known, however, divergence/connections of Anatolian and the Balkans has been shown mostly for animals with the exception of a plant genus Bornmuellera Haussknecht (1897: 70) [ Brassicaceae Burnett (1835: 854) ]. Both the divergence between Anatolian and Balkan clades of this genus in Plio-Pleistocene transition and non-monophyletic relationships of its Anatolian clades has been shown recently ( Özüdoğru & Mummenhoff 2020). It is clear that more Anatolian representatives and more markers are needed to reconstruct a precise biogeographical history for K. goecmenii .

Knautia goecmenii is a unique species within the genus due to its several morphological features especially its suffrutescent and caespitose habitus with very rich stemmed base (up to 152 stems). These features are unique for K. goecmenii have never been observed in any Knautia species to date. In accordance with these characters, unlike most of the perennial or biennial Knautia taxa with leaf rosettes, K. goecmenii lacks rosulate leaves.

Among about 40 perennial Knautia species , K. dinarica Borbás (1894: 399) (subcaespitose), K. velutina Briquet (1902: 94) (rather caespitose), K. carinthiaca Ehrendorfer (1962: 335) (rather caespitose), K. albanica Briquet (1902: 125) (rather caespitose), K. velebitica Szabó (1910: 50) (rather caespitose), K. arvensis ( Linnaeus 1753: 99) Coulter (1823: 29) (laxly caespitose); K. kitaibelii ( Schultes 1809: 18) Borbás (1904: 60) (laxly caespitose), and K. rupicola ( Willkomm 1893: 72) Font Quer (1920: 228) (densely caespitose) have caespitose habitus. But, none of them have a suffrutescent base and such a large number of stems as in K. goecmenii ( Fig. 2 ). The suffrutescent habitus of K. goecmenii has been observed for the first time in the genus Knautia and therefore, it has no known close relatives.

The genus Knautia was traditionally divided into three sections, namely Knautia, Trichera, and Tricheroides. Among them sect. Trichera was mostly characterized by perennial and polyploid species ( Rešetnik et al. 2014). Contrary to this species-rich section, sect. Knautia and sect. Tricheroides include annual, diploid species and is restricted to the Balkan Peninsula and Northwest Anatolia, with the exception of the relatively widely distributed K. integrifolia ( Linnaeus 1753: 99) Bertoloni (1835: 32) (see Rešetnik et al. 2014).

Although K. goecmenii has a perennial life-span, like members of K. sect. Trichera, it is easily separated from the other members of that group by clear morphological and molecular evidence. As said before, the distinct suffrutescent and caespitose habit with many stems at base are unique features in the genus. According to our phylogenetic analyses, one unexpected finding of the present study was the placement of K. goecmenii along with members of sect. Knautia and sect. Tricheroides in a well-supported clade ( Fig. 6 ), which might perhaps indicate that K. goecmenii evolved from annual members of the genus. This result is somewhat surprising given the cited morphological distinctiveness of K. goecmenii . However, further studies with the inclusion of new molecular markers (mostly plastid ones) are needed to test this phylogenetic hypothesis, and to shed light on the tribal affiliation of K. goecmenii , which could not be determined in this study with the available data.

Figures

FIGURE 1. Knautia goecmenii: A. Lower stem; B. Inflorescence; C. Capitula; D. Flower; E. Corolla inside; F. Upper cauline leaves; G. Lower cauline leaves; H. Achene.

FIGURE 2. Knautia goecmenii: A−E. Habitus in habitat.

FIGURE 3. Knautia goecmenii: A. Capitula in flowering and fruiting periods; B. Capitula in early flowering period; C. Capitula in late flowering period; D. Capitula with immature fruits; E. Capitula with mature fruits.

FIGURE 5. Known distribution of Knautia goecmenii.

FIGURE 6. Phylogenetic placement of Knautia goecmenii based on Internal Transcribed Spacers (ITS) sequences. Bayesian posterior probabilities (> 0.5) are shown above branches, whereas ML bootstrap values (> 50 %) are shown below branches only for the main nodes of Knautia. Members of Knautia sect. Knautia, sect. Tricheroides, sect. Trichera were highlighted by different colours.