Corydalis wendelboi subsp. fragilis, Olcay & Kültür & Dogru-Koca, 2025

Olcay, Bülent, Kültür, ükran & Dogru-Koca, Aslı, 2025, A new subspecies of Corydalis wendelboi Lidén (Papaveraceae) from NW Türkiye, evidence from morphological and molecular analysis, Adansonia (3) 47 (21), pp. 341-351 : 343-349

Corydalis wendelboi subsp. fragilis

Olcay, Doğru-Koca & Kültür, subsp. nov.

( Figs 2; 3)

TYPE MATERIAL. — Turkey • A3 Ankara, Kızılcahamam, Şahinler Nature Park , woodlands; 40°38’10.2”N, 32°26’10.3”E; 1480 m a.s.l.; 7.V.2021; B. Olcay s.n.; holotype: ISTE [ ISTE118601 ]!; isotype: HUB ! • A3 Bolu, Near Aladağ Lake ; 1391 m a.s.l.; 7.V.2021; B. Olcay s.n.; paratype: ISTE [ ISTE118598 ] !).

DIAGNOSIS. — Corydalis wendelboi subsp. fragilis subsp. nov. is closely related to C. wendelboi subsp. wendelboi , C. wendelboi subsp. congesta and C. tauricola , because of its divided bracts; a ratio of pedicel to bract length and obtuse lower petals. However, it differs from C. tauricola by its short pedicels, 1-2 mm (not 7-15 mm) and its slightly incised ± equal lobes of bracts (not central lobe of bract largest). It differs from C. wendelboi taxa by its narrowly lobed leaves, 2-3 mm broad (not 3.5- 5 mm); by its smaller bracts, 5-6 mm long (not 9-10 mm), which are divided into very thin lobes (not divided into thick lobes as in subsp. wendelboi or into moderately thin lobes as in subsp. congesta ) and by its lax racemes (not dense). C. wendelboi subsp. fragilis subsp. nov. also differs from subsp. wendelboi because of its smaller seeds, 1-1.2 mm in diameter ( c. 1.5 mm in diameter in subsp. wendelboi ) (Table 2; Figs 3; 4).

Corydalis wendelboi subsp. fragilis subsp. nov., PP849377, Türkiye ( holotype) 0.99/73 Corydalis wendelboi subsp. fragilis subsp. nov., PQ656721, Türkiye 0.86/63 Corydalis wendelboi subsp. fragilis subsp. nov., PP849378, Türkiye 0.88/63 Corydalis tauricola, PP 849383, Türkiye Corydalis wendelboi subsp. wendelboi, PP 849381, Türkiye 1/93 CLADE A Corydalis wendelboi subsp. wendelboi, PQ 656720, Türkiye 1/99 Corydalis wendelboi subsp. congesta, PP 849382, Türkiye 1/71 Corydalis henrikii , X85476 - X85487 , Türkiye Corydalis “ caucasica alba ”, X85468 - X85469 CLADE B 0.97/73 Corydalis intermedia, MN 662998, Norway 1/85 Corydalis pumila, MN 662999, Norway 0.76/ Corydalis solida subsp. solida, PP 849379, Türkiye 74 Corydalis solida subsp. incisa, PP 849380, Türkiye CLADE C Corydalis solida subsp. solida , X85470 - X85472 0.83/76 Corydalis solida subsp. incisa , X85464 - X85471 , Greece 1/100 Corydalis decumbens, MH 349117, China 1/-- Corydalis decumbens, MH 349119, China Corydalis tauricola , X85478 - X85480 Corydalis repens var. repens, MH 349155 China 1/87 Corydalis repens var. repens, MH 349154 China Corydalis repens var. repens, MH 349153 China 1/100 Corydalis repens var. watanabei, MH 349157 China Corydalis repens var. watanabei, MH 349156 China 1/91 Corydalis ambigua, FJ 469598, China 0.73/-- 1/99 Corydalis linjiangensis, MH 349138, China 1/70 Corydalis linjiangensis, MH 349137, China 1/100 Corydalis turtschaninovii, KY 305172, China Corydalis turtschaninovii, MH 349205, China 0.77/32 0.97/90 Corydalis yanhusuo, MH 349215, China Corydalis yanhusuo, MH 349210, China 0.9/68 Corydalis dajingensis, MH 393121, China Corydalis benecincta, MH 393114, China 1/35 Corydalis glaucescens, MT 924005, China Corydalis nariniana , X85460 - X85463 , Türkiye Corydalis bungeana, MH 349108, China Out Group

ETYMOLOGY. — The epithet ‘fragilis’ alludes to the delicate and fragile nature of the plant.

PHENOLOGY. — Flowering commences at the beginning of April and continues until the end of May.Mature fruits have been collected in May. DISTRIBUTION AND ECOLOGY. — C. wendelboi subsp. fragilis subsp. nov. is endemic to Türkiye and is known from only two localities near the border between the provinces of Bolu and Ankara ( Figs 5; 6). It occurs in the subalpine zone, at altitudes of 1400-1600 m a.s.l, in rocky habitats in sparse Pinus nigra forests and areas sheltered by Juniperus communis subsp. nana .

IUCN STATUS. — C. wendelboi subsp. fragilis subsp. nov. is known from only two locations in Bolu and Ankara provinces (Criterion B2a), with an estimated area of occupancy that is less than 10 km 2 (Criterion B2b). A small population of between 30 and 40 individuals has been observed at both locations. Furthermore, no subpopulation was found to contain more than 50 mature individuals, with the expected continuing decline observed in all cases (Criterion C2). The number of mature individuals is less than 50 (Criterion D). Furthermore, the species’ habitat has been affected by factors such as road construction, overgrazing and human pressure. Therefore, it should be classified as “Critically Endangered (CR)” according to the IUCN Red List criteria ( IUCN 2022). Category: CR. B2ab(ii, iv, v); C2a(i); D.

LOCAL NAME. — The Turkish name “Narin tarlakuşu” is proposed for this subspecies, according to the guidelines of Menemen et al. (2016). ADDITIONAL SPECIMENS EXAMINED. — C. wendelboi subsp. fragilis subsp.nov.: Turkey • A3 Ankara, Kızılcahamam, Şahinler Nature Park , woodlands; 40°38’10.2”N, 32°26’10.3”E; 1450 m a.s.l.; 22.IV.2024; B. Olcay & F. Nur Olcay s.n.; ISTE [ ISTE118781 ] !).

C. wendelboi subsp. wendelboi : Turkey • C2 Denizli, Mount Honaz, Tuzluca Plateau ; 1320 m a.s.l.; 8.IV.1973; E.TuzlacI s.n.; ISTE [ ISTE24024 ] ! • C2 Muğla, Muğla-Göktepe TV transmitting station; 1450 m a.s.l.; 20.IV.1981; M. A. Önür 483; ISTE [ ISTE47753 ] ! • C4 Konya, Ermenek ; 1.IV.1993; A. Byfield 151; ISTE [ ISTE65756 ] ! • C3 Antalya, Elmalı, Çam Kuyusu district ; 1580 m a.s.l.; 16.IV.1993; N. Özhatay & A. Byfield s.n.; ISTE [ ISTE65037 ] • B3 Afyon, Sultandağları, above Dereçine ; 1450 m a.s.l.; 19 IV.1976; A. Baytop s.n.; ISTE [ ISTE34558 ] ! • B1 İzmir, Mount Nif ; 990 m a.s.l.; 26.III.1981; A.&T. Baytop s.n.; ISTE [ ISTE46151 ] ! • C4 Konya, Ermenek, Mount Yeşilgöl ; 1500 m a.s.l.; Remark & Wendelbo s.n.; ISTE [ ISTE101596 ] ! • C3 Konya, Beyşehir, Kurucova, Suludere district ; 1500 m a.s.l.; 8.VI.1975; H. Peşmen & A. Güner 2183; HUB [ HUB01517 ] !• C4 Konya, Seydişehir, Şahinkayası district ; 1500 m a.s.l.; 14.IV.1982; H. Ocakverdi s.n.; ANK [ ANK1358 ] !• C2 Muğla, Köyceğiz, Sultaniye district ; 17.IV.1992; A. Güner H. Duman & A. Dönmez 10450; GAZI [ GAZI 210919 ] ! • A3 Ankara, Ayaşbeli Çal Hill ; 1380 m a.s.l.; 13.IV.1986; M.Vural s.n.; GAZI [ GAZI4030 ] ! • C3 Konya, Beyşehir-Akseki Plateau, Baharan district ; 1400 m a.s.l.; 21.IV.1983, M. Koyuncu 5934; AEF [ AEF13415 ] ! • A3 Bolu, Göynük, Sünnetgölü district ; 965 m a.s.l.; 7.IV.2009; A. Güner, M. Johnson & N. İkinci 15387; NGBB [ NGBB001980 ] ! • B4 Ankara, Beypazarı, Sığıryatağı Hill ; 1400 m a.s.l.; 24.IV.2013; A.Ö. Çimen s.n.; NGBB [ NGBB009180 ] !• B2 İzmir, Ödemiş,above Bozdağ

TABLE 2. — Morphological features comparisons of Corydalis wendelboi subsp. fragilis subsp. nov., C. wendelboi subsp. wendelboi Lidén , C. wendelboi subsp. congesta Lidén & Zetterl. , and C. tauricola (Cullen & P.H.Davis) Lidén.

village; 1300 m a.s.l.; 3.IV.1972; E. Leblebici s.n.; EGE [ EGE15109 ]!. Corydalis wendelboi subsp. congesta : Turkey • A3 Bolu, around Lake Abant ; 1350 m a.s.l.; 28.IV.2019; M.Koçyiğit s.n.; ISTE [ ISTE116560 ] !• A2 (A) Bursa, Uludağ, Kirazlıyayla district ; 24.IV.1964; A & T. Baytop s.n.; ISTE [ ISTE7587 ] !• B3 Eskişehir, between the junction of Kütahya and Eskişehir roads and Bozüyük ; 17.IV.1965; A & T. Baytop s.n.; ISTE [ ISTE7878 ] ! • A5 Amasya, Doğantepe Kayacık village ; 950 m a.s.l.; 11.IV.1982; A & T. Baytop s.n.; ISTE [ ISTE48459 ] ! • A2 Kocaeli, Maşukiye- Kartepe ( Keltepe ); 30.V.1987; N. Özhatay, E. Özhatay & M. Johnson s.n.; ISTE [ ISTE57794 ] ! • A4 Kastamonu, Ilgaz passage, 40 km to Kastamonu; 1770 m a.s.l.; 18.V.1972; A. & T. Baytop s.n.; ISTE [ ISTE21751 ] ! • A3 Bolu, Abant Lake side, under Pinus forest; 11.IV.1973; T. Baytop s.n.; ISTE [ ISTE24198 ] ! • A5 Amasya, Akdağ, above Ormanözü village, Eğricepınar district , wetlands; 1200 m a.s.l.; 29.III.1977; T.Baytop s.n.; ISTE [ ISTE36611 ] !• A1 (A) Balıkesir, Marmara Island , hills around the radar facility; 700 m a.s.l.; 16.III.1977; E. TuzlacI s.n.; ISTE [ ISTE36511 ] !• B2 Kütahya, Mount Murat, Kesik Söğüt district ; 1600 m a.s.l.; 18.V.1978; A. Baytop s.n.; ISTE [ ISTE39400 ] ! • A4 Ankara, between Kızılcahamam and Gerede , near the passage; 1512 m a.s.l.; 30.III.2011; E. Kaya 1807; ISTE [ ISTE99393 ] ! • A3 Bolu, near Abant Lake ; 1320 m a.s.l.; 28.III.2011; E. Kaya 1782; ISTE [ ISTE95481 ] ! • A3 Bolu, Sarıalan-Yazıören Plateau , rocky area; 1677 m a.s.l.; 4.V.2014; H. Arslan 1024; DUOF [ DUOF0002247 ] ! • A3 Düzce, Sinekli Plateau ; 1399 m a.s.l.; 2.IV.2010; N. Koçer 1082; DUOF [ DUOF0003500 ] !• A3 Bolu, Yedigöller National Park , yellow pine forest clearings; 1730 m a.s.l.; 15.IV.2018; A. Ayteğin & Y. Zorlu 2339; DUOF [ DUOF0007674 ] ! • B4 Ankara, Kızılcahamam, Mount Işık ; 1500 m a.s.l.; F.Demircioğlu & M.Koyuncu s.n.; AEF [ AEF7359 ] !• Kastamonu, Mount Ilgaz ; 1800 m a.s.l.; 11.V.1982; Y. Akman & E. Yurdakulol s.n.; ANK [ ANK12457 ] !

C. tauricola : Turkey • C6 Adana, Seyhan, Düldül Dağı, Haruniye ; 1300 m a.s.l.; P.H.Davis s.n.; ANK [ ANK26094 ] ! • C5 Adana, Seyhan, Pozantı, Bürücek ; 1300 m a.s.l.; 3.IV.1957; Davis & Hedge s.n.; ANK [ ANK26350 ] ! • C5 İçel, Namrun ; 1000-1800 m a.s.l.; Balls s.n.; ANK [ ANK176 ] ! • C6 Adana, Saimbeyli, Bozoğlan Mt, Obruk Plateau ; 1400 m a.s.l.; 13.IV.1957; Davis & Hedge s.n.; ANK [ ANK26683 ] ! • C6 Kahramanmaraş, Akkar Mt ; 1524 m a.s.l.; 1.V.1934; Balls s.n.; ANK [ ANK942 ] ! • C4 İçel, Mut to Ermenek ; 1800 m a.s.l.; 14.V.1965; M.J. E. Coode & B.M.G. Jones s.n.; E [ E00410736 ] ! • C4 Konya, Ermenek, Damlaçal, Sedir ormanı; 1750 m a.s.l.; 4.IV.1979; M. Vural s.n.; ANK [ ANK1221 ]!, E [ E00410737 ] • C6 Hatay, Amanus Mts, Karlik tepe from sogukoluk; 900 m a.s.l.; 26.III.1967; M.J. Cheese s.n.; E [ E00410739 ] • C6 Kahramanmaraş, Engirek Mt, Biçmeoluk district ; 1500 m a.s.l.; 21.IV.1987; H. Duman s.n.; GAZI [ GAZI2485 ] ! • B6 Kahramanmaraş, Göksun, Acalmak, Cennet river ; 1800 m a.s.l.; 25.IV.1987; B.YIldIz s.n.; HUB [ HUB01530 ] ! • C4 Mersin, between Silifke and Gülnar , rocky places; 750 m a.s.l.; 7.III.1973; M. Liden s.n.; ISTE [ ISTE23911 ] ! • C5 Niğde, Çamardı district ; 1740 m a.s.l.; M. Liden s.n.; ISTE [ ISTE101598 ] ! • C6 Hatay, Antakya, Harbiye- Yayladağı, near Şenköy village ; 800 m a.s.l.; 20.III.1955; H. Demiriz s.n.; ISTF [ ISTF13986 ] ! • C6 Kahramanmaraş, Hartlap, between Fatih and Hacılar district , Bozarmut passage; 1580 m a.s.l.; 30.IV.2007; A. Güner s.n.; NGBB [ NGBB00240 ] ! • C6 Adana, Saimbeyli, between Karakuyu village and Sarıçiçek plateau; 1530 m a.s.l.; 22.III.2014; A. Güner s.n.; NGBB [ NGBB006377 ] !

DESCRIPTION

Perennial, herbs, tuber simple ± spherical, 10-15(-20) mm in diameter. Stem very slender, erect 10-20(-25) cm long with one yellowish scale leaf at the base. Cauline leaves 2, green, alternate, 3(-4) × ternate, long-stalked with deeply divided leaflets. The segments very narrow, divided into obovate to obtuse laciniae, c. 3 mm wide at the mid-point. Racemes lax, 5-15 flowered. Bracts 5(-6) mm long, obovate, semiamplexicaul, divided deeply into numerous unequal acute to obtuse thin lobes. Bracts always longer than pedicels. Pedicels very short, 1-2(3) mm, patent to recurved in fruit. Corolla pale pink, pale lilac to reddish purple, with inner petals internally

IDENTIFICATION KEY TO SPECIES MORPHOLOGICALLY RELATED TO CORYDALIS WENDELBOI SUBSP. FRAGILIS SUBSP. NOV. IN TÜRKIYE

1. Pedicels 7-15(20) mm, longer than bracts ................................................................................................... 2

— Pedicels 2-7 mm, shorter than bracts ........................................................................................................... 3

2. Bracts divided into lax to dense ± equal lobes ................................................................. C. solida (L.) Clairv. — Bracts divided into lax unequal lobes, central lobe of bracts usually largest .....................................................

........................................................................................................ C. tauricola (Cullen & P.H.Davis) Lidén

3. Racemes very dense 10-25 flowered, lower petal 4-6 mm broad ....... C. wendelboi subsp. congesta Lidén & Zetterl. — Racemes moderately dense to lax 5-15 flowered, lower petal 2-4 mm broad ................................................ 4

4. Racemes moderately dense 10-15 flowered, bracts c. 10 mm long, leaf lobes more than 5 mm wide, seeds more than 1.5 mm in diameter ....................................................................... C. wendelboi subsp. wendelboi Lidén

— Racemes lax 5-15 fowered, bracts c. 5 mm long, leaf lobes less than 3 mm wide, seeds less than 1.5 mm in diameter ................................................. C. wendelboi subsp. fragilis Olcay, Doğru-Koca & Kültür , subsp. nov.

tipped dark purple at the apex. Spur 7-12 mm long, straight to slightly curved; outer petals subspathulate, obtuse; lower petal 2-4 mm broad. Stigma flattened with simple lateral papillae. Nectary 4-6 mm long, c. 1/3-2/5 as long as spur. Capsule lanceolate, 10-17 seeded, (12-)13-18 × 2.5-4 mm. Seeds c. 1.2 mm in diameter, with spirally twisted elaiosome ( Figs 2; 3).

MOLECULAR PHYLOGENY

The samples of the new subspecies Corydalis wendelboi subsp. fragilis subsp. nov. occurred in the clade A. The phylogenetic tree displayed that clade A was strongly supported by posterior probability and high bootstrap value (PP = 1, BS = 99) ( Fig. 1). This clade was subdivided into subclades: three samples from two different populations of C. wendelboi subsp. fragilis subsp. nov. formed a monophyletic subclade with strong posterior probability and moderate bootstrap value (PP = 0.99 BS = 73). C. tauricola was a sister group of the C. wendelboi subsp. fragilis subsp. nov. lineage, with branch support values (PP = 0.86, weak BS = 63). In the second subclade, two samples from different populations of C. wendelboi subsp. wendelboi were on the same lineage with moderate posterior probability and weak bootstrap value (PP = 0.88, weak BS = 63). C. wendelboi subsp. congesta was a sister species of the C. wendelboi subsp. wendelboi lineage group was on the same lineage with strong posterior probability and high bootstrap value (PP = 1, BS = 93). C. henrikii was sister to C. wendelboi group. Regarding clade B, C. intermedia Mérat , and C. ‘ caucasica alba ’, which is a garden-origin specimen and whose identification is uncertain ( Lidén et al. 1995) conducted a clade with a high posterior probability and medium bootstrap value (PP = 0.97, BS = 73). In clade C, C. pumila and the intra-specific taxa of C. solida generate a monophyletic clade supported by a moderate posterior probability and bootstrap value (PP = 0.76, BS = 74). Sequences obtained from the GenBank and in this study demonstrate consistency.