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. 2026 Apr 11;15(8):1180. doi: 10.3390/plants15081180

Taurus, an Important Diversification Center for the Genus Aethionema (Brassicaceae): Four New Species from the Central Taurus Mountains in Türkiye

Kuddisi Ertuğrul 1,*, Tuna Uysal 1, Meryem Bozkurt 1, Emrah Şirin 1, Hakkı Demirelma 1, Burcu Yılmaz Çıtak 1
PMCID: PMC13119092  PMID: 42075382

Abstract

This paper explores the role of the Taurus Mountains in shaping species differentiation and biogeographical regions, including distinct ecological zones at various altitudes. The genus Aethionema is a monophyletic group in tribe Aethionemeae, subfamily Aethionemoideae (Brassicaceae), a sister group to the rest of the family. Aethionema has significant taxonomic complexity, particularly in Türkiye, where the genus has the highest species diversity. In this study, four new species (Aethionema kadriyeae, A. uysalii, A. beysehirense, and A. ermenekense) are described based on morphological, palynological and phylogenetic analyses. The diagnoses, detailed descriptions, distribution maps, and illustrations of the new species are provided. Pollen and seed morphology, including detailed measurements of size, ornamentation, and shape, is given. Phylogenetic analyses using DNA sequences from nuclear region (ITS) and chloroplast region (rpl32-trnLUAG) were conducted to determine the evolutionary relationships within the genus. Overall, this research provides new insights into the biodiversity and evolutionary history of Aethionema in Türkiye, highlighting the significance of the Taurus Mountains in supporting rich ecological diversity.

Keywords: Anatolia, Cruciferae, kayagülü, phylogeny, stonecress

1. Introduction

Türkiye is notable for its remarkable species diversity, hosting three biodiversity hotspots within the Palearctic region: the Mediterranean, the Caucasus, and the Irano-Anatolian regions. Positioned at the crossroads of Asia, Europe, and Africa, Anatolia benefits from a unique geographic location that has contributed to its rich biological diversity. Over the course of glacial and interglacial periods, various floral and faunal elements (e.g., Siberian, African, Mediterranean, Boreal, and Eremial) have migrated into Anatolia, enriching its ecological landscape. These periods also facilitated the formation of numerous refugia, further enhancing the region’s biodiversity. Among Anatolia’s key ecological features, the Taurus Mountains play a particularly significant role in supporting and preserving its biological richness [1].

The species diversity observed along the Mediterranean phytogeographic region and the Anatolian Diagonal can be associated with past tectonic events. The belt was proposed as one of the most significant topographic barriers between the eastern and western parts of Anatolia through delimiting distributional ranges of many species. Similar to the formation of the Alborz, Zagros and Kopeh-Dag Mountain ranges in Iran, the formation of the Taurus and Pontic Mountain ranges in Türkiye is part of the Alpine orogeny caused by the collision of the African and Arabian plates with the Eurasian plate [2,3,4]. In particular, the collision of the Arabian and Eurasian plates during the Miocene accelerated the uplift of the Anatolian plateaus and allowed the formation of many different habitat types between the Mediterranean and the formed plateaus [5]. The formation and uplift of these areas caused the aridification of the Irano-Turanian region, turning the surrounding regions into a melting pot of climatic characteristics [5,6]. This has made the Irano-Turanian and Mediterranean regions in Türkiye a home where species boundaries intersect, coexist, and hybridize. In addition, orobiomes and different altitudinal boundaries in different parts of the Taurus Mountains have allowed differentiation of different taxa [7].

Mountain ranges exceeding 1000 m in elevation within the same climatic zone often lead to vertical ecological zonation. This stratification typically supports three distinct orobiomes. For example, in the Taurus Mountains, the lower zone, ranging from the coast to approximately 1000–1200 m, serves as the primary habitat for typical Mediterranean vegetation, including Calabrian pine (Pinus brutia Ten.) and maquis communities. The middle zone, situated between 1200 and 2000 m, is characterized by forests dominated by cedar (Cedrus libani A.Rich.), Taurus fir (Abies cilicica (Antoine & Kotschy) Carrière) and Anatolian black pine (Pinus nigra J.F.Arnold). Pure stands of Calabrian pine forest grow only under Mediterranean climate conditions characterized by mild rainy winters and hot and dry summers, while cedar, fir and Anatolian black pine grow under cold and snowy winters and cool and slightly rainy summers. Therefore, the boundary extending between the two different forest belts is considered the main orobiome boundary of the Taurus Mountains. Above this, the upper zone corresponds to a subalpine region, where alpine and steppe plant species are found [7,8]. The mountain ranges of Anatolia, particularly the Taurus Mountains, play a pivotal role in speciation and in delineating biogeographical subregions and provinces. These ranges, characterized by numerous valleys, especially in the southern regions, significantly shape the area’s biodiversity. The Cilician and western Taurus largely define the East Mediterranean province, while the southeastern Taurus serves as a boundary between the Irano-Anatolian and Mesopotamian provinces. Additionally, the Pontic Taurus facilitates the exchange of Irano-Anatolian elements with Euxinic elements, whereas the eastern Taurus creates pronounced distinctions between central and eastern Anatolia [9]. All four new Aethionema species grow in the middle section of the Taurus Mountains along the “middle zone” line of the Mediterranean vegetation.

As the sister group to the rest of Brassicaceae, Aethionema W.T. Aiton is the sole genus in tribe Aethionemeae and subfamily Aethionemoideae [10]. The complexity of Aethionema arises from the limited number of macro-morphological traits, such as those related to fruits and leaves, that can be used to distinguish species. Aethionema is a taxonomically difficult genus representing 68 species worldwide [11], and its center of highest diversity is Türkiye, with fewer species in neighboring Iran, Caucasus, and Greece. The distribution of individual species extends eastward to Kazakhstan and westward to Spain and Morocco [12,13,14]. The genus Aethionema was represented by 30 species in the first volume of the Flora of Turkey [15]. Ertuğrul [16] recognized 40 species of Aethionema in Türkiye, but subsequent discoveries increased it to 57 [17,18,19,20,21,22,23]. Following recent taxonomic revisions to the genus, the number of species in Türkiye was reduced to 47 [24].

This study aims to investigate the influence of the Taurus Mountains on species differentiation and biogeographical structuring by describing four new species of Aethionema from Türkiye. Utilizing an integrative approach that combines morphological, palynological, and phylogenetic analyses, this research seeks to resolve the taxonomic complexity within the genus, clarify evolutionary relationships, and contribute new insights into the biodiversity and evolutionary history of Aethionema in one of its primary centers of diversity. As a result of the completed systematic and phylogenetic studies, numerous samples belonging to various species were collected during recent extensive fieldwork. Following comprehensive herbarium studies, including all type collections, it was concluded that these samples represent four new species, hereafter recognized as Aethionema. kadriyeae Ertuğrul, A. uysalii Ertuğrul & Demir., A. beysehirense Ertuğrul & Şirin, and A. ermenekense Ertuğrul & Uysal.

2. Results

2.1. Palynomorphological Characteristics

All studied species possess radially symmetrical, isopolar pollen grains, with tricolpate as the predominant aperture type; however, Aethionema armenum Boiss., A. spicatum Post, and A. yildirimlii Kılıç additionally exhibit tetracolpate pollen, whereas A. kadriyeae displays syncolpate grains. In all taxa, the pollen outline is triangular or orbicular in polar view and elliptic in equatorial view, and the colpus is long and sunken and possesses distinct, regular margins with an ovate end (Figure 1). Pollen shape and size vary considerably among species: A. kadriyeae exhibits prolate-spheroidal pollen (P: 20.01 ± 0.90 μm, E: 12.42 ± 0.42 μm) with a microreticulate exine and narrow muri, whereas A. turcicum H.Duman & Aytaç has prolate pollen (P: 20.04 ± 2.31 μm, E: 13.21 ± 1.79 μm) and a reticulate exine with wider muri. Among A. uysalii, A. coridifolium DC., and A. spicatum, pollen shape is consistently prolate-spheroidal, yet aperture morphology differs: A. uysalii and A. coridifolium are tricolpate, while A. spicatum exhibits both tricolpate and syncolpate grains, with pollen dimensions of P: 17.64 ± 1.13 μm, E: 15.24 ± 1.85 μm; P: 16.50 ± 1.16 μm, E: 15.75 ± 0.82 μm; and P: 20.44 ± 1.00 μm, E: 20.22 ± 0.46 μm, respectively, and all three possess reticulate pollen but differ in muri size (Figure 1 and Table 1). Aethionema beysehirense exhibits subprolate pollen (P: 21.14 ± 2.41 μm, E: 16.91 ± 3.45 μm) with a reticulate exine, whereas A. armenum (P: 18.12 ± 0.78 μm, E: 16.23 ± 0.50 μm) and A. schistosum (P: 16.41 ± 1.04 μm, E: 13.66 ± 1.03 μm) are prolate-spheroidal and microreticulate, with all three varying in muri size (Table 1). Finally, A. ermenekense, A. spicatum, and A. yildirimlii share prolate-spheroidal pollen, with dimensions of P: 17.64 ± 1.13 μm, E: 15.24 ± 1.85 μm; P: 20.44 ± 1.00 μm, E: 20.22 ± 0.46 μm; and P: 15.98 ± 0.99 μm, E: 15.56 ± 0.87 μm, respectively, and exine ornamentation varies from microreticulate in A. ermenekense to reticulate in A. spicatum and A. yildirimlii, with corresponding differences in muri size (Figure 1 and Table 1).

Figure 1.

Figure 1

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The comparative pollen morphological characters of Aethionema species by scanning electron microscope (SEM). (A1,A2) A. kadriyeae., (B1,B2) A. turcicum, (C1,C2) A. uysalii, (D1,D2) A. spicatum, (E1,E2) A. coridifolium, (F1,F2) A. beysehirense, (G1,G2) A. schistosum, (H1,H2) A. armenum, (I1,I2) A. ermenekense, (J1,J2) A. yildirimlii.

Table 1.

The comparative pollen features of Aethionema species (values in μm, n = 30).

Taxa/Pollen Characters Polar Axes Equatorial Axes P/E Ratio Pollen Shape Colpus Length (Clg) Colpus Width (Clt) Exine Intine Apocolpium Aperture Sculpture Lumina
A. kadriyeae 20.01 ± 0.90 12.42 ± 0.42 1.15 Prolate-spheroidal 16.98 ± 1.06 3.10 ± 0.46 1.44 ± 0.07 0.62 ± 0.03 3.44 ± 0.47 Tricolpate Microreticulate 0.87 ± 0.10
A. turcicum 20.04 ± 2.31 13.21 ± 1.79 1.51 Prolate 15.77 ± 1.41 3.02 ± 1.48 1.15 ± 0.04 0.59 ± 0.05 3.07 ± 1.06 Tricolpate Reticulate 1.03 ± 0.17
A. uysalii 17.64 ± 1.13 15.24 ± 1.85 1.15 Prolate-spheroidal 15.21 ± 0.62 4.22 ± 0.42 1.34 ± 0.17 0.66 ± 0.06 3.18 ± 1.67 Tricolpate Reticulate 1.19 ± 0.30
A. spicatum 20.44 ± 1.00 20.22 ± 0.46 1.01 Prolate-spheroidal 22.42 ± 0.48 2.12 ± 1.07 1.43 ± 0.18 0.59 ± 0.08 3.34 ± 1.15 Tricolpate-Tetracolpate Reticulate 1.01 ± 0.26
A. coridifolium 16.50 ± 1.16 15.75 ± 0.82 1.04 Prolate-spheroidal 14.64 ± 0.3 5.06 ± 0.46 1.26 ± 0.20 0.62 ± 0.05 2.52 ± 0.34 Tricolpate Reticulate 1.26 ± 0.25
A. beysehirense 21.14 ± 2.41 16.91 ± 3.45 1.25 Subprolate 16.09 ± 4.63 3.61 ± 0.92 1.22 ± 0.19 0.58 ± 0.05 4.34 ± 0.55 Tricolpate-Syncolpate Microeticulate 1.59 ± 0.25
A. schistosum 16.41 ± 1.04 13.66 ± 1.03 1.20 Subprolate 14.49 ± 0.93 3.12 ± 0.19 0.94 ± 0.21 0.48 ± 0.04 4.19 ± 0.11 Tricolpate Microeticulate 1.19 ± 0.29
A. armenum 18.12 ± 0.78 16.23 ± 0.50 1.11 Prolate-spheroidal 14.66 ± 0.71 5.71 ± 1.10 1.15 ± 0.22 0.54 ± 0.06 2.20 ± 0.59 Tricolpate-Syncolpate-Tetracolpate Microreticulate 0.83 ± 0.15
A. ermenekense 17.64 ± 1.13 15.24 ± 1.85 1.15 Prolate-spheroidal 15.21 ± 0.62 4.22 ± 0.42 1.34 ± 0.17 0.66 ± 0.06 3.18 ± 1.67 Tricolpate Reticulate 1.19 ± 0.30
A. yildirimlii 15.98 ± 0.99 15.56 ± 0.87 1.02 Prolate-spheroidal 14.52 ± 1.37 5.67 ± 0.38 1.36 ± 0.17 0.57 ± 0.11 4.71 ± 0.74 Tricolpate-Tetracolpate Microreticulate 0.76 ± 0.09
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2.2. Seed Morphology

The investigated Aethionema taxa have ovate to broadly ovate seeds with obtuse base and rounded apex and vary in color from light-brown to brown, though A. ermenekense differs from the rest by having brown-grey seeds. The common seed sculpture is reticulate-verrucate. The seed size of A. kadriyeae is 2.17 × 1.26 mm and of A. turcicum is 1.65 × 0.86 mm (Figure 2, Table 2), and both have ovate seeds with a striate pattern. Seed sizes are 2.11 × 1.42 mm in A. uysalii, 2.29 × 1.25 mm in A. spicatum, 2.87 × 1.73 mm in A. coridifolium, 2.33 × 1.25 mm in A. beysehirense, 2.11 × 1.00 mm in A. armenum, 2.02 × 1.18 mm in A. schistosum, 2.79 × 1.49 mm in A. ermenekense, and 2.68 × 1.47 mm in A. yildirimlii (Table 2). The largest seeds are found in A. ermenekense, and the smallest are found in A. turcicum. The cells of seed-surface testa are orbicular with irregular epidermal cells (Figure 2 and Figure 3). A. kadriyeae, A. coridifolium, A. schistosum, and A. yildirimlii have broadly ovate seeds that vary in epidermal cell size. Both A. kadriyeae and A. turcicum have regular cells, though the anticlinal walls are clearer in the former. The verrucae of epidermal cells are convex in all the species studied except A. yildirimlii (Figure 2 and Table 2).

Figure 2.

Figure 2

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The comparative seed morphological characters of Aethionema species by scanning electron microscope (SEM). (A1–A3) A. kadriyeae, (B1–B3) A. turcicum, (C1–C3) A. uysalii, (D1–D3) A. spicatum, (E1–E3) A. coridifolium.

Table 2.

The comparative seed characteristics of Aethionema species (n = 20).

Taxa/Seed Characters Length (mm) Width (mm) L/W Ratio Seed Shape Seed Color
Mean Mean
A. kadriyeae 2.17 1.26 1.76 Broadly ovate Light-brown and brown
A. turcicum 1.65 0.86 1.84 Ovate Light-brown and brown
A. uysalii 2.11 1.42 1.86 Ovate Light-brown
A. spicatum 2.29 1.25 1.75 Ovate Light-brown and brown
A. coridifolium 2.87 1.73 1.66 Broadly ovate Light-brown
A. beysehirense 2.33 1.25 2.03 Ovate Light-brown
A. schistosum 2.02 1.18 1.70 Broadly ovate Brown
A. armenum 2.11 1.00 2.03 Ovate Light-brown
A. ermenekense 2.79 1.49 1.86 Ovate Brown-grey
A. yildirimlii 2.68 1.47 1.75 Broadly ovate Brown
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Figure 3.

Figure 3

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The comparative seed morphological characters of Aethionema species by scanning electron microscope (SEM). (A1–A3) A. beysehirense, (B1–B3) A. schistosum, (C1–C3) A. armenum, (D1–D3) A. ermenekense, (E1–E3) A. yildirimlii.

2.3. Phylogenetic Studies

The ITS and rpl32-trnL dataset includes sequences of 14 Aethionema taxa. The length of the aligned datasets is 617 (ITS) and 636 (rpl32-trnLUAG) base pairs. PAUP and MrBayes programs were used to construct phylogenetic trees, and tree topologies in NJ, MP, ML and BI were similar. Due to such similarities, the following discussion will focus only on NJ (Figure 4). Based on the ITS sequence data, the Neighbor-Joining (NJ) tree resolved the Aethionema taxa into two major clades. The first main clade, comprising A. turcicum, A. kadriyeae, A. dumanii, and A. aytachii, is strongly supported (PP = 1.00; BS = 94% for MP/95% for ML). The remaining Aethionema taxa are grouped within the second main clade. Similarly, the NJ tree constructed from the rpl32-trnL sequence data also divided the Aethionema taxa into two main clades. The first main clade includes A. kadriyeae, A. dumanii, and A. aytachii. The second main clade is further subdivided into two subclades: the first subclade consists solely of A. turcicum (PP = 0.82; BS = 68 for MP/67 for ML), while the second subclade contains the remaining Aethionema taxa and is strongly supported (PP = 0.93; BS = 93 for MP/87 for ML) (Figure 4). In the phylogenetic trees, certain taxa (such as A. ermenekense, A. yildirimli, and A. spicatum for ITS and A. yildirimli and A. coridifolium for rpl32-trnL) remain unresolved, forming partial polytomies. These patterns suggest the presence of complex evolutionary relationships that cannot be fully captured by tree-based approaches. These complex evolutionary relationships observed in the trees were resolved using network analyses.

Figure 4.

Figure 4

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Neighbor-Joining trees showing phylogenetic relationships based on nuclear (A) and plastid (B) sequences and network analyses (C) based on plastid sequences. The numbers on the branch are given as posterior probability (PP) and bootstrap (BS; MP and ML), respectively (ITS; CI:0.853, RI:0.857, HI:0.147 and rpl32; CI:0.884, RI:0.889, HI:0.116). The letters A and B, indicated in blue on the phylogenetic tree, represent the major clades.

2.4. Taxonomy

2.4.1. Aethionema kadriyeae Ertuğrul, sp. Nov. (Figure 5 and Figure 6)

Type: Türkiye. Karaman: Kayaönü village, south slopes of Karadere, marly steppe, 1620 m, 19 May 2021, K. Ertuğrul 6339 and B. Yılmaz Çıtak (holo KNYA; isotypes GAZI, ANK).

Figure 5.

Figure 5

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The fruit light microscope microphotographs of examined Aethionema species. (A) A. kadriyeae, (B) A. turcicum, (C) A. uysalii, (D) A. spicatum, (E) A. coridifolium, (F) A. beysehirense, (G) A. schistosum, (H) A. armenum, (I) A. ermenekense, (J) A. yildirimlii.

Figure 6.

Figure 6

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Habit, flowers and infructescence of Aethionema kadriyeae.

Diagnosis: A. kadriyeae is similar to A. turcicum, but it differs by its elliptic ovary (vs. oblong), edentate inner filaments (vs. dentate), 6–8 × 5.5–7 mm fruits (vs. 8.5–11 × 8–9), cordate base (vs. sagittate) at base, 1.5–2.5 mm length wing (vs. 2.5–3 mm) and open (vs. closed) sinus (Table 3).

Table 3.

Comparison of morphological characters of A. kadriyeae, A. uysalii and related taxa.

Characters A. kadriyeae A. turcicum A. uysalii A. spicatum A. coridifolium
Stem leaves elliptic, attenuate at base ovate, rounded at base oblanceolate, attenuate at base obovate, attenuate at base linear–oblanceolate, attenuate at base
Raceme cylindrical cylindrical capitate capitate cylindrical
Infructescence lax lax compact compact lax
Sepals (mm) oblong, 2.5–3.5 × 1.5–1.6 oblong, 2.7–3.1 × 1.3–1.6 oblong, 3–3.5 × 0.8–1.2 oblong, 2–4 × 1–1.5 (–2) oblong, 2.7–3.5 × 1.1–1.8
Petals (mm) obovate, 6.5–7.5 × 2.5–4 obovate, 5–6.5 × 2.5–4 spatulate–obovate, 7.5–8 × 2.7–5 obovate, 4–6 × 1.5–2.1 (–3) obovate, 5.5–6 × 3.2–4
Inner filaments (mm) not toothed, 3–3.5 toothed, 2–3 not toothed, 1.8–2.8 not toothed, 1.8–2.5 not toothed, 2.5–3
Pedicel in fruit (mm) erect, 5–5.5 horizontal–recurved, 4–7 adpressed, 4.5–5 ascending, 3.5–6 erect–recurved, 3.5–6.5
Ovary elliptic oblong elliptic obovate ovate-orbicular
Fruits (mm) oblong–obovate, cordate at base, 6–8 × 5.5–7 oblong, sagittate at base, 8.5–11 × 8–9 obovate, cordate at base, 7.5–9.5 × 8–11 obovate–orbicular, cordate at base, 6–11 × 5–11 ovate–orbicular, cordate at base, 6–8.5 × 5.5–6.5
Wing (mm) 1.5–2.5 2.5–3 4.5–5.5 2–3.5 2–3.5
Cell (mm) 3.5–5.5 × 2–2.5 (–3.5) 5.5–7 × 2–3 3–4 × 2.5–3.5 3–3.5 × 2–3.5 2.5–4 × 2–3.5
Septum (mm) 4–5.5 × 1–2 5–7 × 1.5–2.1 3.5–4 × 1–1.3 3–5 × 0.9–1 3.5–4.5 × 1.4–2.5
Sinus (mm) open, 0.5–0.8 closed, 0.7–1.1 open, (3.5–) 4–5 closed, 2–4 closed, 1–2.5
Style (mm) 1–1.7 1–1.5 0.5 0.5–1 0.2–0.3
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Perennial, woody at base, many stemmed; stems 7–21 cm long, erect–ascending, sometimes branched towards the apex, glabrous, glaucous. Upper stem leaves sub-fleshy, glaucous, subsessile, alternate, elliptic, attenuate at base, obtuse and mucronulate at apex, 5.5–9 × 3–5.5 mm, lower stem leaves sub-fleshy, glaucous, subsessile, alternate rarely opposite, ovate, attenuate–rounded base, obtuse at, 4.5–5.5 × 2–3 mm, glomerule leaves more prominent during fruiting period, sub-fleshy, glaucous, subsessile, rosulate–alternate, oblong–elliptic, attenuate, acute at apex, 2–5 × 0.7–1.5 mm. Racemes 1–3 cm long, cylindrical, elongated in fruit. Pedicels 4–5 mm long in flowers, 5–5.5 mm in fruit, erect. Sepals oblong, violet margined with green band, 2.5–3.5 × 1.5–1.6 mm. Petals obovate, truncate, 6.5–7.5 × 2.5–4 mm, purple, claw and blade not distinct. Filaments free; inners not toothed, dilated at base, 3–3.5 mm long; outers 1.8–2.1 mm long; anthers yellow, apiculate, 0.5–0.8 mm long; stigma capitate; ovary elliptic. Infructescence lax, 3–9 cm long; fruit weakly cymbiform, dehiscent, oblong–obovate, 6–8 × 5.5–7 mm, bilocular, 2-seeded per locule, cordate at base, cell 3.5–5.5 × 2–2.5 (–3.5) mm, septum 4–5.5 × 1–2 mm, undulate at margin, wing 1.5–2.5 mm long, sinus open and 0.5–0.8 mm deep, style 1–1.7 mm long. Seed light-brown or brown, 2.148–2.205 × 1.24–1.28 mm, broadly ovate, verrucate.

Etymology: This new species is named in honor of the first author’s wife Kadriye Ertuğrul for her contribution to most fieldwork. The vernacular name of A. kadriyeae was proposed as “Sultan kayagülü” according to guidelines published by Menemen et al. [25].

Distribution, habitat and ecology: A. kadriyeae grows on marly steppes around Kayaönü in the Ayrancı district of Karaman Province at altitudes of 1260–1680 m (Figure 7).

Figure 7.

Figure 7

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Distribution map of A. ermenekense (●), A. uysalii (♦), A. kadriyeae (■) and A. beysehirense (▲).

Additional examined specimens: A. kadriyeae: Türkiye. Karaman: Ayrancı, Kayaönü, south slopes of Karadere, marly steppe, 1620 m, 4 July 2021, K. Ertuğrul 6616 (KNYA); 20 May 2022, K. Ertuğrul 6815 (KNYA). Türkiye. Karaman: Between Küçükkoraş-Kayaönü villages, white marly slopes, 1600 m, 20 June 2012, K. Ertuğrul 4587 & T. Uysal (KNYA). Ayrancı, Kayaönü village, Quercus openings, limestone soils, 19 June 2013, T. Uysal 3055 & al. (KNYA).

A. turcicum: Ankara Ayaş road, Aysantı pass, marly slopes, 1180 m, 25 May 2021, K. Ertuğrul 6379 & E. Şirin (KNYA).

The infructescence and fruit shape of A. kadriyeae resemble those of A. turcicum but differ by its basally attenuate (vs. rounded) upper leaves, erect (vs. horizontal and recurved) fruiting pedicels, edentate inner filaments 3–3.5 mm long (vs. dentate and 2–3 mm long), outer filaments 1.8–2.1 mm long (vs. 1.3–1.8 mm), elliptic (vs. oblong) ovary, fruits 6–8 × 5.5–7 mm (vs. 8.5–11 × 8–9 mm), with cordate (vs. sagittate) fruit base, locule 3.5–5.5 mm (vs. 5.5–6.5 mm) long, septum 4–5.5 mm (vs. 5–7 mm) long, wing 1.5–2.5 mm (vs. 2.5–3 mm) long, and open (vs. closed) sinus.

2.4.2. Aethionema uysalii Ertuğrul & Demir., sp. Nov. (Figure 5 and Figure 8)

Type: Türkiye. Antalya: Akseki, between Salamut-Morca plateaus, stony slopes, 1987 m, 8 July 2020, K. Ertuğrul 6124 & E. Şirin (holo KNYA; isotypes GAZI, ANK).

Figure 8.

Figure 8

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Habit, flowers and infructescence of Aethionema uysalii.

Diagnosis: A. uysalii is similar to A. spicatum and A. coridifolium but differs from A. spicatum by its oblanceolate (vs. obovate) stem leaves, 7.5–8 × 2.7–5 mm (vs. 4–6 × 1.5–2.1 (–3) mm) petals, and elliptic (vs. obovate) ovary. It differs from A. coridifolium by its capitate (vs. cylindrical) raceme, compact (vs. lax) infructescence, 8–11 mm (vs. 5.5–6.5 mm) fruit width, and 0.5 mm (vs. 0.2–0.3 mm) long styles (Table 3).

Perennial, with woody caudex, multi-stemmed at the base; sterile shoots, 10.5–36 cm long, erect, unbranched, glabrous, and glaucous. Stem leaves not fleshy, glaucous, subsessile, alternate, oblanceolate, attenuate at base, obtuse and sometimes mucronulate at apex, 12–25 × 2–4.5 mm; leaves of sterile stems glaucous, petiolate, opposite rarely alternate, oblanceolate, attenuate at base, obtuse at apex (2.5-) 4–10 × 1.5–3 mm. Racemes capitate, 0.7–2 cm long, elongated in fruit. Pedicels 3–4 mm long in flowers, 4.5–5 mm in fruit, adpressed. Sepals oblong, white margined with violet band, 3–3.5 × 0.8–1.2 mm. Petals spatulate–obovate, 7.5–8 × 2.7–5 mm, pink, claw and blade not distinct. Filaments free; inners not toothed, dilated at base, 1.8–2.8 mm long; outers 1–1.5 mm long; anthers yellow, truncate, 0.5–0.9 mm long; stigma capitate, papillose; ovary elliptic. Infructescence compact, imbricated, 1.5–5 cm long; fruits cymbiform, dehiscent, obovate, 7.5–9.5 × 8–11 mm, bilocular, 1-seeded per locule, cordate at base, cell 3–4 × 2.5–3.5 mm, septum 3.5–4 × 1–1.3 mm, undulate at margin, wing dilated towards apex and 4.5–5.5 mm long, sinus open and (3.5-) 4–5 mm deep, style 0.5 mm long. Seed light-brown, ovate, 2.77–2.82 × 1.48–1.51 mm, verrucate.

Etymology: This new species is named in honor of Süleyman Uysal, an amateur photographer interested in wild flowering plants in Antalya. The vernacular name of A. uysalii was proposed as “Akseki kayagülü” according to guidelines published by Menemen et al. [25].

Distribution, habitat and ecology: A. uysalii grows on rock crevices and stony slopes around Salamut-Morca plateaus in the Akseki district of Antalya province at altitudes of 1450–1987 m (Figure 7).

Additional examined specimens: A. uysalii: Türkiye. Antalya: Akseki, environs of Morca plateau, rocky areas, 1965 m, 8 July 2020, K. Ertuğrul 6140 & E. Şirin (KNYA); 6 August 2020, H. Demirelma 3444 & E. Şirin (KNYA). Türkiye. Antalya: Akseki, Fersin plateau, Evlek pass, rock crevices, 1710 m, 6 July 2020, H. Demirelma 3441 & E. Şirin (KNYA). Evlek pass, rocky areas, 1718 m, 8 July 2020, K. Ertuğrul 6123 & E. Şirin (KNYA). Akseki, Murtiçi, transmitter road of Gülen Mountain, rocky areas, 1450 m, 27 May 2022, K. Ertuğrul 6866 (KNYA). Akseki, Güzelsu, Gidefi Mountain, Gülek location, cliffs, 1733 m, 27 May 2022, K. Ertuğrul 6869 (KNYA).

A. coridifolium: Türkiye. Kahramanmaraş: Göksun, above Değirmendere, around Şarlak, stony slopes, 5 June 2022, K. Ertuğrul 6940 & E. Şirin (KNYA), Mersin, Ayvagediği, above Çandır Castle, around pond, clearings of Pinus forest, 1110 m, 27 June 2019, K. Ertuğrul 2940 (KNYA).

A. spicatum: Türkiye. Adana, Pozantı, Hamidiye to Çamlıbel, clearings of Pinus forest, 1290 m, 7 June 2020, K. Ertuğrul 6017 & H. Demirelma (KNYA), Mersin, Kaynakkeşli village, around Kızılasma bridge, clearings of Pinus forest, 760 m, 17 May 2019, K. Ertuğrul 5693 & H. Demirelma (KNYA).

A. uysalii resembles A. spicatum in having capitate racemes and compact infructescence and A. coridifolium in the linear–oblanceolate stem leaves. The novelty differs from A. spicatum by its non-fleshy, oblanceolate stem leaves and up to 25 mm long (vs. fleshy, obovate, up to 14 mm long) that are petiolate, oblanceolate on sterile stems (vs. subsessile, obovate), adpressed (vs. ascending) fruiting pedicels, petals 7.5–8 × 2.7–5 mm (vs. 4–6 × 1.5–2.1 (–3) mm), and elliptic (vs. obovate) ovary. It differs from A. coridifolium by having outer filaments 1–1.5 mm long (vs. 1.5–2.1 mm), elliptic (vs. circular) ovary, compact (vs. lax) infructescence, weakly cymbiform (vs. strongly cymbiform) fruits 8–11 mm (vs. 5.5–6.5 mm) wide, septum width 1–1.3 mm (vs. 1.4–2.5 mm), wing 4.5–5.5 mm long (vs. 2–3.5 mm), sinus 4–5 mm deep (vs. 1–2.5 mm) and style 0.5 mm long (vs. 0.2–0.3 mm).

2.4.3. Aethionema beysehirense Ertuğrul & Şirin, sp. Nov. (Figure 5 and Figure 9)

Type: Türkiye. Konya: Konya-Beyşehir road, Gündoğdu-Gönen junction area, calcareous slopes, 1163 m, 12 July 2022, K. Ertuğrul 7063 (holo KNYA, isotypes GAZI, ANK).

Diagnosis: A. beysehirense is similar to A. schistosum and A. armenum, but it differs from A. schistosum by having obovate (vs. orbicular) fruits 5–7.5 mm (vs. 9–12 mm) wide and styles more than 0.5 mm (vs. less than 0.5 mm). It differs from A. armenum by its spatulate (vs. oblanceolate-obovate) petals and fruits more than 5 mm (vs. less than 5 mm) wide, and wings with closed sinus and more than 2 mm long (vs. open sinus and less than 2 mm long) (Table 4).

Table 4.

Comparison of morphological characters of A. beysehirense, A. ermenekense and related taxa.

Characters A. beysehirense A. schistosum A. armenum A. ermenekense A. yildirimlii
Stem leaves linear, rarely involute and falcate, attenuate at base linear, attenuate at base linear-oblanceolate, attenuate at base oblong, attenuate at base oblanceolate, attenuate at base
Raceme cylindrical capitate capitate capitate capitate
Infructescence lax compact lax compact compact
Sepals (mm) oblong rarely obovate, 2.5–3.5 × 1–1.5 oblong–obovate, 2.5–3 × 0.8–1.2 oblong, 2–3.5 × 1–1.7 oblong, 2.4–2.5 × 1–1.2 oblong, 3–3.5 × 1–1.5
Petals (mm) spatulate, 5–7 × 2–4 spatulate, 4.5–7 × 2.5–4 oblanceolate or obovate, 4–6 × 1.8–3 spatulate, 4.5–6 × 2.1–2.5 spatulate-obovate, 6–7 × 2.4–3
Inner filaments (mm) not toothed, 1.7–2.7 not toothed, 1.8–2.6 not toothed, 2.2–3.5 not toothed, 1.5–1.6 not toothed, 2.5–2.8
Pedicel in fruit (mm) adpressed–recurved, 4–6 adpressed–recurved, 4.5–5 adpressed–ascending, 3–5 adpressed and recurved, 3.5–4 adpressed—recurved, 4.5–5
Ovary obovate obovate–oblong oblong elliptic ovate
Fruits (mm) obovate, cordate at base, 6.5–7.5 × 5–7.5 orbicular, cordate at base, 7–10.5 × 9–12 obovate rarely oblong, cordate at base, 4–6.5 × 3–5 obovate-orbicular, cordate at base, 7–9 × 7–10 obovate-orbicular, cordate at base, 7–8.5 × 7–9.5
Wing (mm) 2–4 4–5 1–2 2.5–3.5 3.5–4.5
Cell (mm) 3–4.5 × 2–3 3.5–4.2 × 2–3.5 2–4 ×1–2.2 3.5–4.5 × 2.5–3 3–3.5 × 2.2–3
Septum (mm) 3.5–5 × 1–1.6 3–4 × 0.8–1.2 3–4 × 0.7–1.5 3.5–4.5 × 1.1–2 3.5–4.5 × 0.9–1.1
Sinus (mm) closed, 0.8–1 closed, 3–3.5 open, 0.5–1 closed, 1.5–3 closed, 2–3
Style (mm) 0.5–1 0.2–0.5 0.5–1 0.1–0.2 0.3–0.5
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Perennial, with woody branches at the base; stems 5–22 cm long, erect–ascending, glabrous. Leaves heteromorphic: upper ones glaucous subsessile, alternate, linear, rarely involute and falcate, attenuate at base, usually acute at apex, 5.5–13 (–16) × 1–1.5 mm; glomerule leaves more prominent during fruiting period, glaucous, subsessile, rosulate-opposite rarely alternate, linear–oblong rarely oblanceolate, attenuate at base, acute, rarely obtuse at apex, 2–5 (–7) × 0.5–1 mm. Racemes cylindrical, 0.5–2.5 cm long, elongated in fruit. Pedicels 3–4.5 mm long in flowers, 4–6 mm in fruit, adpressed and recurved. Sepals oblong or rarely obovate, white margined with green band, rarely violet margined, 2.5–3.5 × 1–1.5 mm. Petals spatulate, 5–7 × 2–4 mm, violet, claw and blade almost distinct. Filaments free; inners not toothed, dilated at base, 1.7–2.7 mm long; outhers 1–1.6 mm long; anthers yellow, apiculate, 0.5–0.8 mm long; stigma capitate; ovary obovate. Infructescence lax, 1–7.5 cm long; fruit weakly cymbiform, dehiscent, obovate, 6.5–7.5 × 5–7.5 mm, bilocular, 1-seeded per locule, cordate at base, cell 3–4.5 × 2–3 mm, septum 3.5–5 × 1–1.6 mm, irregular dentate at margin, wing 2–4 mm long, sinus closed and 0.8–1 mm deep, style 0.5–1 mm long. Seed light-brown, ovate, 2.02–2.03 × 0.97–0.99 mm, verrucate.

Etymology: The specific epithet was derived from the name of Beyşehir district from where the first specimens of the new species were collected. The vernacular name of A. beysehirense was proposed as “Beyşehir kayagülü” according to guidelines published by Menemen et al. [25].

Distribution, habitat and ecology: A. beysehirense grows on gypsum, marly and limestone slopes on the roadside of the Konya to Beyşehir, Gündoğdu-Gönen junction area, on the roadside of Hüyük to Konya, Konya, Kızılören Mountain foothills, around Halkapınar village of Ereğli in Konya, west of Büyükkarapınar village of Başyayla in Karaman, and south of Ulukışla in Niğde, at altitudes of 1163–1683 m (Figure 7).

Additional examined specimens: A. beysehirense: Türkiye. Konya: Beyşehir road, Gündoğdu junction area, calcareous slopes, 1163 m, 3 June 2022, K. Ertuğrul 6905 (KNYA); Konya-Beyşehir road, Gündoğdu-Gönen junction, slopes, 1163 m, 22 May 2022, K. Ertuğrul 6853 (KNYA). Türkiye. Konya: Beyşehir, Hüyük-Konya road, after Selki, limestone slopes, 1232 m, 29 May 2022, K. Ertuğrul 6896 (KNYA); 12 July 2022, K. Ertuğrul 7062 (KNYA); Konya, Kızılören, above Kent Forest, Kızılören Mountain foothills, openings of Juniperus, 25 June 2021, K. Ertuğrul 6608 (KNYA); Ereğli, Halkapınar, Kayasaray village, Düğünlük stream, Kocaoluk area, limestone slopes 1650 m, 15 May 2022, H. Demirelma 3453 & E. Şirin (KNYA). Entrance of Düğünlük stream, Quercus openings, 1630 m, 25 May 2019 K. Ertuğrul 5747 & E. Şirin (KNYA); 12 July 2019, K. Ertuğrul 5910 & al. (KNYA); Ereğli, 5–10 km southwest of Büyükdede village, clearings of Juniperus forest, 1810 m, 20 June 2017, K. Ertuğrul 5386 & H. Dural (KNYA). Ereğli, Halkapınar, Kayasaray village, Karaman: Ermenek, Tekeçatı to Balkusan, limestone slopes, Juniperus and Pinus openings, 1620 m, 20 May 2022, K. Ertuğrul 6819 (KNYA); Ermenek, west of Büyükkarapınar village, limestone slopes, 1594 m, 21 May 2022, K. Ertuğrul 6824 (KNYA); 7 July 2022, K. Ertuğrul 7053 (KNYA), 1683 m, 2 July 2022, H. Demirelma 3471 & E. Şirin (KNYA); 1610 m, 15 July 2019, K. Ertuğrul 5918 (KNYA);1685 m, 2 July 2022, H. Demirelma 3469A & E. Şirin (KNYA); Niğde: South of Ulukışla, around Evrenhasan Tepe, eroded clayey slopes, 1600 m, 8 May 2022, K. Ertuğrul 6801 (KNYA); 1560 m, 15 June 2017, K. Ertuğrul 5337 & H. Dural (KNYA); Cedrus plantation area, 1550 m, 12 July 2019, K. Ertuğrul 5883 & al. (KNYA); gypsum and limestone slopes, 1670 m, 12 July 2019, K. Ertuğrul 5888, 5888A & al. (KNYA); 1700 m, 12 July 2019, K. Ertuğrul 5894 & al. (KNYA).

A. schistosum: Türkiye: Konya, Seydişehir, west of Madenli willage, slopes, 1710 m, 4 July 2019, K. Ertuğrul 5863 (KNYA), Adana: Tufanbeyli to Tomarza, between Ayvat village and Tomarza, stony slopes, 2003 m, 14 June 2022, K. Ertuğrul 7036 & E. Şirin (KNYA), Adana: Pozantı to Çamardı, between Fındıklı and Kamışlı villages, Pinus openings, 1200 m., 8 May 2022, K. Ertuğrul 6797 (KNYA).

A. armenum: Kayseri, Alidağı, west foothills, stony slopes, 1378 m, 12 June 2021, K. Ertuğrul 6513 & E. Şirin (KNYA); Kayseri, Pınarbaşı, between Aşağıbeyçayırı and Yukarıbeyçayırı villages, serpentine slopes, 12 June 2021, K. Ertuğrul 6524 & E. Şirin (KNYA).

A. beysehirense resembles A. schistosum in having linear stem leaves and spatulate petals and A. armenum in the shape of stem leaves and lax infructescence. It differs from A. schistosum by having obovate (vs. orbicular) fruits 5–7.5 mm (vs. 9–12 mm) long, with fruit wing less than 4 mm (vs. more than 4 mm) long, and style more than 0.5 mm (vs. less than 0.5 mm) long. It differs from A. armenum by having spatulate (vs. oblanceolate–obovate) petals differentiated (vs. undifferentiated) into claw and blade, and fruits more than 5 mm (vs. less than 5 mm) wide with wing more than 2 mm (vs. less than 2 mm) long and closed (vs. open) sinus.

Figure 9.

Figure 9

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Habit, flowers and infructescence of Aethionema beysehirense.

2.4.4. Aethionema ermenekense Ertuğrul & Uysal, sp. Nov. (Figure 5 and Figure 10)

Type: Karaman: Sarıeliler, Göktepe (Karalar) plateau, high mountain steppe, 1830 m, 19 June 2017, K. Ertuğrul 5367 & H. Dural (KNYA).

Diagnosis: A. ermenekense is similar to A. spicatum and A. yildirimlii, but it differs from A. spicatum by violet margined with green band sepals (vs. white margined with violet band), spatulate (vs. obovate) petals, elliptic (vs. obovate) ovary, and weakly cymbiform (vs. cymbiform) fruits with style, 0.1–0.2 mm (vs. 0.5–1 mm) long. It differs from A. yildirimlii by its unbranched (vs. basally branched) stems, 2.4–2.5 mm long (vs. 3–3.5 mm) sepals, elliptic (vs. ovate) ovary, 2.5–3.5 mm long (vs. 3.5–4.5 mm) wing, and style 0.1–0.2 mm long (vs. 0.3–0.5 mm) (Table 4).

Perennial, with woody caudex, multi-stemmed at the base; stems (2.5–) 4–12 cm long, erect–ascending, unbranched, glabrous, and glaucous. Leaves heteromorphic; stem leaves sub-fleshy, glaucous, subsessile or rarely sessile, alternate, oblong near to inflorescence, attenuate at base, obtuse at apex, (5–) 6–10 (–11) × 1–2.5 mm; glomerule leaves dense towards the base, fleshy, glaucous, subsessile, alternate–opposite, obovate, attenuate at base, obtuse at apex, 2.5–5 × 1.2–2.5 mm. Racemes capitate, corymbose, 0.5–1 cm long, slightly elongated in fruit. Pedicels 2–3 mm long in flowers, 3.5–4 mm in fruit, adpressed and recurved. Sepals oblong, violet margined with green band, 2.4–2.5 × 1–1.2 mm. Petals spatulate, 4.5–6 × 2.1–2.5 mm, pink, differentiated into claw and blade. Filaments free, inner not toothed, dilated at base, 1.5–1.6 mm long; outer 1–1.2 mm long; anthers yellow, apiculate, 0.5–0.8 mm long; stigma capitate, papillous; ovarium elliptic. Infructescence compact, imbricated, 1–2 cm long; fruits weakly cymbiform, dehiscent, obovate-orbicular, 7–9 × 7–10 mm, bilocular, 1-seeded per locule, cordate at base, cell 3.5–4.5 × 2.5–3 mm, septum 3.5–4.5 × 1.1–2 mm, entire or rarely undulate at margin, wing 2.5–3.5 mm long, sinus closed and 1.5–3 mm deep, style 0.1–0.2 mm long. Seed brown-gray ovate, 2.77–2.82 × 1.48–1.51, verrucate.

Etymology: The specific epithet was derived from the name of Ermenek district from where the first specimens of the new species were collected. The vernacular name of A. ermenekense was proposed as “Göktepe kayagülü” according to guidelines published by Menemen et al. [25].

Distribution, habitat and ecology: A. ermenekense grows on steppe, stony slopes in the Ermenek-Göktepe district of Karaman province at altitudes of 1830 m. This region is part of the Mediterranean floristic region (Figure 7).

Additional examined specimens: A. ermenekense: Türkiye. Karaman: Sarıveliler, west of Göktepe (Karalar) plateau, high mountain steppe, 1830 m, 23 April 2018, K. Ertuğrul 5547 (KNYA). Ermenek, Sarıveliler, Göktepe (Karalar) plateau, 1830 m, stony slopes, 15 July 2019, K. Ertuğrul 5912 (KNYA); Sarıveliler, Göktepe (Karalar) plateau, high mountain steppe, 1830 m, 21 May 2022, K. Ertuğrul 6832 (KNYA); Ermenek-Sarıvelliler, west of Göktepe (Karalar) plateau, 1830 m, 2 July 2022, H. Demirelma 3464 and E. Şirin (KNYA). A. spicatum: Türkiye. Adana, Pozantı, Hamidiye to Çamlıbel, clearings of Pinus forest, 1290 m, 7 June 2020, K. Ertuğrul 6017 & H.Demirelma (KNYA), Mersin, Kaynakkeşli village, around Kızılasma bridge, clearings of Pinus forest, 760 m, 17 May 2019, K. Ertuğrul 5693 & H.Demirelma (KNYA). A. yildirimlii: Türkiye: Konya, Derebucak, Çamlık village, around Kızıldağ, forest clearings, serpentine slopes, 1534 m., 22 May 2022, K. Ertuğrul 6839 (KNYA), Kızıldağ, forest clearings, serpentine slopes, 1580 m, 15 June 2021, T. Uysal 4358 (KNYA).

A. ermenekense resembles A. spicatum and A. yildirimlii in its wider stem leaves and compact infructescences. It differs from A. spicatum by its oblong (vs. oblanceolate–obovate) stem leaves, 1.2–2.5 mm (vs 2.5–4.5 mm) wide, glomerulate leaves 1–2.5 mm (vs. 2–4.5 mm) wide, flowering pedicels 2–3 mm (vs. 3.5–4.5 mm) long, violet margined sepals with green band (vs. white margined with violet band), spatulate petals 2.1–2.5 mm wide (vs. obovate and 1.5–2.1 mm wide), inner filaments 1.5–1.6 mm (vs. 1.8–2.5 mm) long, and elliptic (vs. obovate) ovary. It differs from A. yildrimlii by the unbranched stems 4–12 cm long (vs. basally branched and 12–20 cm long), stem leaves 6–11 mm (vs. 4–6 mm) long, pedicels 2–3 mm long in flower and 3.5–4 mm in fruit (vs. 3–4.5 mm in flowers and 4.5–5 mm in fruit), sepals 2.4–2.5 mm (vs. 3–3.5 mm) long, petal differentiated into claw and blade and 4.5–6 mm long (vs. undifferentiated and 6–7 mm long), inner filaments 1.5–1.6 mm (vs. 2.5–2.8 mm) long, elliptic (vs. ovate) ovary, fruit wing 2.5–3.5 mm (vs. 3.5–4.5 mm) long, and style 0.1–0.2 mm (vs. 0.3–0.5) mm long.

Figure 10.

Figure 10

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Habit, flowers and infructescence of Aethionema ermenekense.

3. Discussion and Conclusions

In Mediterranean Türkiye, different vegetations are formed under changing climatic conditions and topography [26]. Geographic, ecological, and climatic factors as well as interspecific hybridization (introgression) and polyploidy play important roles in speciation and diversification of Aethionema [24,27].

Mohammadin et al. [28] concluded that the center of origin of Aethionema is most probably along the Anatolian Diagonal, from which it spread to the Irano-Turanian region after the uplift of Iranian and Anatolian plateaus and the formation of mountain ranges in Iran and Türkiye. This concept is confirmed by the present study as well as by Ertuğrul et al. [24], revealing that two local endemics occupy an ancestral position compared to those of the Anatolian Diagonal, where half of the 55 taxa in Türkiye are distributed. It is clear, however, that the Taurus Mountains play an important role in the evolution and differentiation of the genus but not as much as the Anatolian Diagonal.

As a result of the uprise of the Zagros Mountains, different habitats appeared due to the orogeny in Iran. A similar orogenic formation occurred for Central Taurus, where numerous new habitats have contributed to the evolution of many new Aethionema species. The Mediterranean vegetation in Türkiye is divided into two Mediterranean biome and orobiome ecoregions, with the former extending along the Mediterranean coastline and the latter covering mountainous areas above 1000 m [26]. Ertuğrul et al. [24] indicated that ten local endemics (A. lycium I.A. Andersson et al., A. demirizii Davis & Hedge, A. subulatum (Boiss. & Heldr.) Boiss., A. alanyae H. Duman, A. karamanicum Ertuğrul & Beyazoğlu, A. eunomioides (Boiss.) Bornm., A. compactum (Hartvig & Strid) Yıld., A. yıldırımlii Kılıç, A. thesiifolium Boiss. & Heldr., and A. balansae Boiss.) are localized especially in the orobiome ecoregion (on calcareous slopes and rock crevices of rocky slopes). Serpentine rocks or their fragmented surfaces rarely provide a special habitat for some endemics, such as A. compactum and A. yildirimlii. Apart from this, Mesozoic limestone, karst landforms and reddish Mediterranean soils are spectacular habitats for some Aethionema taxa. For example, dolines are very special geological formations and host many relict and endemic species. They are generally found in the upper parts of the Taurus Mountains. For example, A. ermenekense is distributed in red soil meadows surrounded by sparsely distributed and degraded cedar forests on sloping limestone rocky areas. Several other narrow endemics (e.g., Muscari vuralii Bagci & Dogu, Centaurea ermenekensis Şirin & Uysal, Sedum ermenekensis Yıld. & Dinç, Gladiolus izzet-baysalii Eker & Sagiroglu and Haplophyllum ermenekense Ulukuş & Tugay) were discovered from the same region. Aethionema uysalii is a subalpine that grows on limestone rocky crevices east of Antalya. Recently discovered novelties such as Bilacunaria aksekiensis A. Duran & B. Doğan, Polygonum uysalii S. Makbul, Coskunç. & S. Kundakçi, Chaerophyllum aksekiense A. Duran & H. Duman, and Linaria dumanii A. Duran & Y. Menemen occupy different altitudes and habitats within two different Mediterranean ecobiomes. Thus, one can conclude that these special habitats on the Taurus Mountains accommodate many endemics, of which dozens of novelties were discovered in the last decade.

Interestingly, both A. uysalii and A. ermenekense are morphologically related to A. spicatum, but they differ in their phylogenetic relationships. A. spicatum is widely distributed in the Eastern Mediterranean, where it grows on limestone, serpentine rocks, or slopes in forest clearings. In ITS-based phylogenetic analyses, A. spicatum has been shown to exhibit a relatively weak phylogenetic affinity with A. beysehirense and A. armenum, which are distributed between the Anatolian Plateau and the Mediterranean phytogeographical region. In contrast, plastid genes (rpl32–trnL intron) indicate that the species displays a closer phylogenetic relationship with A. ermenekense compared to nuclear genomic data. Accordingly, phylogenetic relationships inferred from chloroplast DNA sequences are congruent with morphological characters. In contrast, ITS-based phylogenetic inferences are incongruent with morphology. This discordance is most plausibly explained by introgressive hybridization resulting from gene flow mediated by interspecific pollination with a sympatrically distributed species. Furthermore, network analyses provide strong evidence that A. spicatum and A. ermenekense are closely related phylogenetically, indicating a high degree of genetic affinity between the two taxa. Similarly, A. uysalii exhibits incongruence between maternal (chloroplast) and nuclear phylogenetic signals. Specifically, chloroplast-based inference indicates a closer evolutionary affinity to A. schistosum than to A. coridifolium, in contrast to relationships inferred from nuclear data. A. beysehirense, another new subalpine species, is distributed in limestone slope clearings of semi-shrub forest that display a transition from the Mediterranean orobiome to the Anatolian plateaus. Although the species is morphologically similar to A. schistosum, it is phylogenetically related to A. armenum. This species appears to be adapted to transition areas among the Mediterrenean and the Anatolian plateaus rather than the dolines. By contrast, A. kadriyeae, another transition zone species, is very locally distributed on limestone slopes of Kayaönü, an area with many endemics (e.g., Linum ciliatum Hayek, Aethionema karamanicum, Polygala inexpectata Peşmen & Erik, etc.). The phylogenetic position of all four new Aethionema indicates differentiation in the ITS and chloroplast sequence data, though an inconsistency occurs. While A. kadriyeae shares a common position with A. dumanii and A. aytachii from the gypsum areas of the Anatolian plateau in the plastid inheritance, it shows a closer relationship with A. turcicum according to the nuclear ITS sequences (BS and PP values above the branches of the trees). Similar incongruencies have been reported in much earlier studies, e.g., [28,29,30,31]. The ITS phylogeny is congruent with the findings of Mohammadin et al. [28] and Lenser et al. [29], supporting the delineation of the two principal clades (A and B). The rpl32-trnL-based phylogeny likewise corroborates the presence of these major clades, with the exception of A. turcicum. Furthermore, network analysis inferred from rpl32-trnL sequences provides additional support for the separation of these major clades (Figure 4).

Seed morphology was previously shown to have some taxonomic value in Brassicaceae, e.g., [32,33,34,35]. Bona [36] found that seed morphology in the 14 Lepidium L. taxa from Türkiye is tuberculate, ruminate, reticulate, reticulate-areolate, reticulate-tuberculate, or reticulate-fovate type. By contrast, Kaya et al. [37] demonstrated that the seed morphology is also useful in Malcolmia W.T. Aiton, Strigosella Boissier and Zuvanda Askerova (now Plagioloba Reichenbach) at the species level. By contrast, seed morphology was shown by Pınar et al. [38] to support the division of Hesperis L. into sections Hesperis, Mediterreraneae, Pachycarpos, Delicate, Diaplictos and Contorta.

In their study of 17 taxa of Aethionema, Pınar et al. [38] showed that seed shape, color and ornamentation are taxonomically useful in distinguishing species. Four types of seed ornamentation (reticulate, reticulate-clavate, reticulate-verrucate and verrucate) are found in Aethionema, and Demirpolat [39] showed that the seeds of A. sancakense are broadly ovoid, brown, and reticulate-verrucate. Our present results overlap significantly with the above-mentioned studies and support the value of overall seed morphology (including SEM data) in the diagnoses and recognition of the four new species. For example, A. kadriyeae differs from the related A. turcicum by having broadly ovate (vs. ovate) seeds; A. uysalii resembles A. spicatum in having ovate seeds that differ from A. coridifolium, which has broadly ovate seeds; and A. beysehirense has ovate seeds similar to A. armenum but differs from A. schistosum with broadly ovate seeds. Ovate seeds are observed in A. ermenekense and A. spicatum, while A. yildirimlii has broadly ovate ones. The seed color is not found useful for delimitation taxa because close species have the same seed color. All new species and their close relatives have reticulate-verrucate seed sculpture. Finally, the smallest seeds were observed in A. turcicum, while the largest ones are in A. ermenekense.

Palynological data, especially shape and sculpture, is also useful in the separation of four new taxa from their close relatives. For example, A. kadriyeae has prolate-spheroidal-shaped pollen grains, while the related A. turcicum has prolate ones. However, A. uysalii and close relatives A. spicatum and A. coridifolium cannot be distinguished according to pollen shape and sculpture of exine. Therefore, the aperture type can separate A. spicatum, but it is not useful between A. uysalii and A. cordifolium. Pollen shape is subprolate in A. beysehirense, as it is in A. schistosum, but differs from A. armenum, which has prolate-spheroidal. Both tetracolpate and tricolpate aperture morphology was seen in A. beysehirense, while A. schistosum had only tricolpate. Similarly, A. ermenekense has only tricolpate aperture that differs from its close relatives A. yildirimlii and A. spicatum, which have tricolpate-tetracolpate. Çeter et al. [40] reported that the pollen shape, size, surface ornamentation, and muri and lumina shape and size are important in distinguishing Aethionema species, and their results are congruent with ours.

The infructescence morphology, especially compact (vs. lose), is one of the main characters in the separation of taxa in Aethionema. Furthermore, fruit morphology (especially shape and base, sinus length and opening, septum and style length, cell structure and shape, wing length) is very useful in determining species boundaries, and these are discussed under each of the four new species.

4. Materials and Methods

Field trips were mainly conducted in 2020–2024. Approximately 1500 Aethionema samples were collected from different habitats throughout Türkiye. Morphological observations and measurements were carried out in the field as well as in the laboratory. Distribution maps of all the species were prepared based on records from both our fieldwork and the localities from examined herbarium specimens.

Pollen of various species was taken from herbarium materials and prepared following Wodehouse [41]. The non-acetolysis pollen grains were separated by anthers and were stained with glycerin-jelly and covered with coverslips. Pollen was observed using a Leica DM 1000 light microscope with a Canon 450 D camera (Ota City, Tokyo, Japan) and measured using Kameram 21 software (Argenit, İstanbul, Türkiye). At least 30 or more pollen were investigated per sample.

At least 20 mature fruits or seeds per sample were investigated for micromorphological analyses. For scanning electron microscope (SEM) studies, pollen and seeds were directly transferred to small aluminum stubs, and they were coated with gold aid of a sputter coater. Photographs of pollen and seeds were taken with a Zeiss Evo LS 10 microscope (Carl Zeiss NTS GmbH, Oberkochen, Germany). Refs. [38,40,41,42] were followed for palynological and seed micromorphological terminology.

For molecular phylogenetic studies, samples of various species of Aethionema and Noccaea (as outgroup) were collected from different regions of Türkiye (Table 5). Total DNA isolation was performed using the 2X CTAB method described by Doyle and Doyle [43] and modified by Soltis et al. [44] and Cullings [45]. For PCR amplification, primers were used for ITS1 and ITS4 regions [46] and rpl32-trnLUAG [47]. Amplification conditions and procedures were carried out in accordance with previously established methodologies reported in [48] and [47], respectively. The raw sequences were edited using the Chromas Lite program. All sequences were aligned in the Bioedit program (v.7.0.5.3 version; [49]). Neighbor Joining (NJ), Parsimony and Maximum Likelihood (ML); the SYM + G and GTR + G evolutionary model was used for ITS and rpl32-trnL, respectively; the default option in PAUP analyses was performed in the PAUP program (v.4.0b10 version; [50]). Bootstrap analyses were performed with 1000 repetitions for the reliability of branches. Both the retention index (RI) and consistency index (CI) were given for the strict consensus trees, with the exclusion of uninformative characters. Bayesian inference (BI) of the two datasets was calculated using MrBayes v3.2.6 [51]. The best available model of molecular evolution required for Bayesian estimations of phylogeny was selected using Akaike information criteria (AIC) for both datasets as implemented in the software MrModelTest v2.2 [52]. The best fitting models for the ITS and rpl32-trnL datasets were TIM2ef + G and K81uf + G, respectively. Bayesian analyses for ITS and rpl32-trnL were conducted using random starting trees, which were then run for 6.1 × 104 and 10 × 104 generations, for 2 independent 4 Metropolis-coupled chain runs. For every 1000 generations that were run, only one was recorded. The run output was examined for convergence by considering the standard deviation of the split frequencies that were near 0.001. The first 1000 samples (20%) were considered to be burn-in and were removed after they were visually examined with regards to the likelihood score plots. The stationary of runs as well as the convergence between the runs were examined using Tracer software (v.1.7.0 version) [53]. Phylogenetic network analyses were performed with the Network program [54]. Bayesian (BI), Maximum Likelihood (ML), and Maximum parsimony (MP) trees based on nuclear (ITS) and plastid (rpl32-trnL) sequences are provided in the Supplementary Materials.

Table 5.

Voucher specimens for the ITS and rpl32-trnL study.

Taxa Collection No ITS Genebank rpl32-trnL Genebank
A. turcicum K.Ertuğrul 5754 MW791189 (Uysal et al. In: Ertuğrul et al. [22]) PV166694 (In this study)
A. kadriyeae K.Ertuğrul 6339 PV156495 (In this study) PV166695 (In this study)
A. dumanii K.Ertuğrul 5755 MW791190 (Uysal et al. In: Ertuğrul et al. [22]) PV166696 (In this study)
A. aytachii K.Ertuğrul 5757 MW791188 (Uysal et al. In: Ertuğrul et al. [22]) PV166697 (In this study)
A. ermenekense K.Ertuğrul 5912 PV156496 (In this study) PV166698 (In this study)
A. yildirimlii H.Demirelma 2917(ITS), K.Ertuğrul 5849 (rpl32) PV156497 (In this study) PV166699 (In this study)
A. spicatum K.Ertuğrul 6016 PV156498 (In this study) PV166700 (In this study)
A. uysalii K.Ertuğrul 6124(ITS), K.Ertuğrul 6123 (rpl32) PV156499 (In this study) PV166701 (In this study)
A. coridifolium K.Ertuğrul 6557 PV156500 (In this study) PV166702 (In this study)
A. beysehirense K.Ertuğrul 6905 PV156501 (In this study) PV166703 (In this study)
A. armenum K.Ertuğrul 6521 PV156502 (In this study) PV166704 (In this study)
A. schistosum K.Ertuğrul 6851 PV156503 (In this study) PV166705 (In this study)
Noccaea oppositifolia K.Ertuğrul 6836 PV156504 (In this study) PV166706 (In this study)
N. iberidea K.Ertuğrul 6818 PV156505 (In this study) PV166707 (In this study)
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Acknowledgments

The authors thank TUBITAK and S.U-BAP for their support. We thank Ihsan A. Al-Shehbaz for reviewing and editing the manuscript.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/plants15081180/s1. S1: Bayesian (BI), Maximum Likelihood (ML), and Maximum parsimony (MP) trees showing phylogenetic relationships based on nuclear (ITS) and plastid (rpl32-trnL) sequences.

plants-15-01180-s001.zip (5.8MB, zip)

Author Contributions

K.E., T.U., H.D. and E.Ş. were responsible for the collection of plants and carried out the species identification and nomenclature work. K.E., H.D. and E.Ş. conducted morphological analysis. T.U. and M.B. prepared the DNA samples, conducted the sequencing and performed molecular phylogenetic analyses. B.Y.Ç. was responsible for palynology and seed micromorphology analyses. K.E., T.U., H.D., E.Ş. and B.Y.Ç. wrote the original manuscript. All authors have read and agreed to the published version of the manuscript.

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding authors.

Conflicts of Interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as potential conflicts of interest.

Funding Statement

This study was supported by the Scientific and Technological Research Council of Türkiye (TUBITAK) under Grant Number 118Z995 and by Selcuk University, Scientific Research Coordination Office (S:U. BAP) 20401025. 50% of the APC fee was funded by the Selcuk University Scientific Research Coordination Office.

Footnotes

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Associated Data

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Supplementary Materials

plants-15-01180-s001.zip (5.8MB, zip)

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding authors.

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