Seed Morphological and Anatomical Structures as Taxonomy Tool for Turkish Hyacinthella Schur (Asparagaceae) Taxa

Seed Morphological and Anatomical Structures as Taxonomy Tool for Turkish Hyacinthella Schur (Asparagaceae) Taxa | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Seed Morphological and Anatomical Structures as Taxonomy Tool for Turkish Hyacinthella Schur (Asparagaceae) Taxa Hüseyin EROĞLU, Mehmet Cengiz KARAİSMAİLOĞLU, Mehmet FİDAN, Süleyman Mesut PINAR This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-1635226/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract This paper offers the first taxonomic assessment of the morphological and anatomical characters of the seeds of 11 Hyacinthella taxa from Turkey, 10 of them are endemic, with the cluster analysis and principal component analysis. Macro-morphologically, the outcomes display that the species are different from each other as per seed shape and size. The studied seeds are separated into 5 shapes; triangularis, circularis, ellipticus, ellipticus-late and ovatus. Seed sizes range from 1.51 mm to 3.06 mm in length, from 1.06 mm to 2.04 mm in width. Micro-morphologically, seed surface ornamentation is grouped into 7 types: rugose, reticulate-pusticulate, verrucate, tuberculate, ruminate, reticulate-foveate and alveolate. The most common type is verrucate, as rugose (in H. acutiloba ), reticulate-pusticulate (in H. campanulata ), ruminate (in H. hispida ) and reticulate-foveate (in H. lineata ) ornamentation types are taxon specific. Anatomically, testa structures of the studied seeds are usually consisted of 1 layer, the epidermis, formed in the sclerenchymatous tissue. The epidermis layer demonstrates significant differences in cell form, comprising of crushed, flat or polygonal cells, in 1–7 layers, and has surged or straight wall form. Also, the subepidermis layer is found in some of the studied taxa ( H. glabrescens , H. heldreichii and H. siirtensis ). The thickness of testa layers varies between 15.77 µm (in H. micrantha ) and 49.76 µm (in H. campanulata ). In the systematics of the genus Hyacinthella , the seed morphological and anatomical characters are very important characteristics that reveal inter-specific relationships among the studied species. Also, a dichotomous key is provided for the identification of the examined species based on seed characters. Anatomy Diagnostic key Seed Taxonomic study Testa Turkey Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Figure 8 Introduction The family Asparagaceae is of 7 subfamilies, 114 genera and circa 2900 taxa in worldwide (Chase et al 2009 ; Christenhusz and Byng 2016 ). It represents with 19 genera and 182 species in the Flora of Turkey (Güner et al 2012 ). Hyacinthella Schur consists of 17 species, 1 subspecies and 1 variety belonging to 2 sections, and it disperses from Balkan Peninsula to Turkey and along northern Iran (Persson and Wendelbo 1981 ; Speta 1998 ; Govaerts et al 2021 ). According to Mathew ( 1987 ), Turkey is one of the diversity centers of the genus. The genus was placed within the family Liliaceae in its first classification based on limited macromorphological characters. As a result of molecular studies, Hyacinthella was first transferred to the family Hyacinthaceae and then to the family Asparagaceae (APG III 2009). The genus Hyacinthella was reported as 10 taxa with 9 species and 1 hybrid species ( H. micrantha (Boiss.) Chouard × H. heldreichii (Boiss.) Chouard.) to Flora of Turkey (Persson and Wendelbo 1984 ). Subsequently, it was defined that the samples belonging to the declared hybrid species were not hybrid and they were new species ( H. lazulina K. Perss. and Jim. Perss.) (Persson and Persson 1992 ). Afterwards, another new species was published from Turkey (Persson 2000 ). Hyacinthella millingenii (Post) Feinbrun endemic to Cyprus was involved in the latest update flora checklist performed in Turkey; however, no specific specimens or location of the species was included, and it was noted that "the status of the species needs to be clarified" (Ekim 2012 ). Therefore, the number of taxa belonging to the genus Hyacinthella in Turkey has updated as 11 species, including 10 endemic species. The main characters utilized in taxonomy of the genus Hyacinthella are pedicel presence-absence and pedicel length, flower color, lobe-flower ratio, anther-filament ratio, flower shape, leaf shape, leaf edge, secondary leaf to primary leaf ratio (Persson and Wendelbo 1984 ; Persson 2000 ). According to Persson and Wendelbo ( 1984 ), the characters used in the taxonomy of the genus may not be consistent in terms of features such as color, length and number. Accordingly, new distinctive and consistent characters are needed to contribute to the systematics of the genus. The seeds are of the characteristic morphological features such as shape, colour, size and surface ornamentation types, and they can offer valuable data in of the elucidating taxonomic problems and in of the establishing systematic relationships (Barthlott 1981 ; Karaismailoğlu and Erol 2018 ). Besides, the anatomical characters are typically as helpful as morphological characters for plant taxonomy, and they often are valuable in the separation of closely correlated taxa (Karamian et al 2012 ; Karaismailoğlu 2015 ). The importance of the seed in genera within the family Asparagaceae has often been overlooked. The only study showing the importance of comprehensive seed morphology and anatomy within the family was carried out by Eroğlu et al. ( 2021 ) on Turkish Muscari , which is a genus closely related to Hyacinthella . In the mentioned study, it was seen that seed morphological and anatomical characters easily distinguished 36 Muscari taxa from each other. Some morphological, anatomical, cytological and palynological studies on some taxa of the genus Hyacinthella have been made, so far (Feinbrun 1961 ; Persson and Wendelbo 1981 , 1982 ; Puizina et al 2003 ; Selvi et al 2008 ; Yetişen et al 2012 ; Tekin and Meriç 2013 ; Tzonev and Panovska 2017 ; Doğu 2019 ). However, the morphological and anatomical characteristics of seeds have been ignored in the classification of taxa belonging to the genus. The aim of this work is to examine the morphological and anatomical characters of seeds of Turkish Hyacinthella taxa for the first time, and discuss the systematic practice of these characteristics. Material And Methods Plant materials Species were gathered from different phytogeographical areas of Turkey and were stored at VANF (Van Yüzüncü Yıl University Herbarium). The studied species and their locations were offered in Table 1 . Table 1 The collections data of the investigated Hyacinthella taxa and their locations. Species Locality Voucher no. Hyacinthella acutiloba B7 Malatya: Yazıhan, Girmana Canyon, rocky places, 1063 m, 31 March 2021. H.Eroglu 1593 H. campanulata C4 Konya: Meram, Konya-Beyşehir road, 5 km to Altınapa Dam, marly steppe, 1310 m. 26 May 2021. H.Eroglu 1719 H. glabrescens C5 Mersin: Tarsus, south of Cevizliçeşme Village, rocky Pinus yards, 1030 m, 17 March 2020. H.Eroglu 1530 H. heldreichii C3 Antalya: Korkuteli, from the old road from Antalya to Korkuteli, northwest of Güllük Mountain, Pinus yards, 702 m. 19 March 2020. H.Eroglu 1532 H. hispida C5 Mersin: Tarsus, Çamalan Village, southeast of martyrdom, Pinus and Juniperus yards, 776 m, 17 March 2020. H.Eroglu 1529 H. lazulina C4 Mersin: Silifke, between Demircili and İmamlı villages, 1 km to İmamlı Village, maquis yards, 718 m, 18 March 2020. H.Eroglu 1531 H. lineata C2 Denizli: Honaz, south of Gürleyik Village, around of marble quarry, marly steppe, 424 m, 20 March 2020. H.Eroglu 1533 H. micrantha A4 Kastamonu: Kastamonu University Kuzeykent campus, around of football stadium, Pinus yards, 820 m, 18 March 2019. H.Eroglu 1450 H. nervosa C6 Gaziantep: Şahinbey, Gaziantep-Kilis road, 1 km from Küçükkızılhisar, stony-rocky steppe, 859 m, 16 March 2020. H.Eroglu 1527 H. siirtensis C8 Mardin: Artuklu, between Mardin and Diyarbakır, Akresta pass, Hut Farm road, streamside, steppe, 1138 m, 16 March 2020. H.Eroglu 1524 H. venusta C4 Konya; between Taşkent and Başyayla, Bolay highland, roadside, steppe, 1677 m, 26 May 2021. H.Eroglu 1717 Macro And Micro-morphological Analysis Macro-morphological structures of the seeds containing colour, shape and size were studied with 100 seeds of 10 specimens per species using a Leica EZ4 binocular microscope with a HD camera (Fig. 1 , Table 2 ). Micro-morphological structures of the seeds including surface ornamentation, anticlinal and periclinal cell walls, and epidermal cells were analyzed with a Scanning Electron Microscope (SEM) (Fig. 2 , Table 2 ). Firstly, seeds were positioned on the stubb with a carbon tape and covered with gold, then observed with a Zeiss LEO 440 SEM (Karaismailoğlu 2015 ). Table 2 The seed morphological characters of the studied Hyacinthella taxa. Taxa Color Shape Seed sizes L/W Seed surface Anticlinal Periclinal Epidermal L (mm) W (mm) ornamentation cell wall cell wall cell structure H. acutiloba Black Triangularis 2.58 ± 0.10 1.94 ± 0.12 1.32 Rugose Unclear Unclear Unclear H. campanulata Black Circularis 1.51 ± 0.08 1.27 ± 0.06 1.18 Reticulate-Pusticulate Raised Concave Polygonal cells H. glabrescens Black Ellipticus 2.43 ± 0.12 1.85 ± 0.10 1.31 Verrucate Flat Convex Unclear H. heldreichii Black Ovatus 1.95 ± 0.06 1.55 ± 0.12 1.25 Tuberculate Sunken Concave Alveolar cells H. hispida Black Triangularis 1.98 ± 0.04 1.59 ± 0.06 1.24 Ruminate Unclear Unclear Unclear H. lazulina Black Ellipticus-late 2.09 ± 0.04 1.73 ± 0.08 1.20 Verrucate Flat Convex Unclear H. lineata Black Ovatus 1.59 ± 0.06 1.06 ± 0.04 1.50 Reticulate-Foveate Raised Concave Polygonal and alveolar cells H. micrantha Brown Ovatus 2.01 ± 0.04 1.48 ± 0.06 1.35 Alveolate Raised Concave Alveolar cells H. nervosa Black Ovatus 1.82 ± 0.08 1.35 ± 0.04 1.34 Verrucate Flat Convex Unclear H. siirtensis Black Triangularis 3.06 ± 0.12 2.04 ± 0.08 1.50 Alveolate Raised Concave Alveolar cells H. venusta Black Ellipticus-late 1.80 ± 0.06 1.61 ± 0.08 1.11 Tuberculate Sunken Concave Alveolar cells L = length, W = width Anatomical Analysis The examination of the seed anatomical structures was performed with dry herbarium materials. Cross-sections were obtained from the middle of the seed with a fully automatic microtome (Thermo Shonda Met Finesse, Thermo). They were treated with a series of alcohol and xylene, stained with hematoxylin and eosin-Y in a staining device (ASC 720 Medite) and covered with using Entellan (Fig. 3 , Table 3) (Karaismailoğlu 2015 ; Karaismailoğlu and Erol 2018 ). Anatomical structures were surveyed with an Olympus CX31 light microscope and Kameram Imaging Software (KAMERAM12 CCD, Argenit Micro System Ltd., Turkey). The terminology utilized for seed morpho-anatomical features is suitable with Stearn ( 1985 ). Statistical analysis Clustering of taxa was made with utilizing the grouping analysis method (UPGMA) in MultiVariate Statistical Package (MVSP) as per the 42 characteristics in Tables 2 –3 (Fig. 4 ). The characters in statistical analysis were utilized seed colour: black (1), brown (2); seed shape: triangularis (3), circularis (4), ellipticus (5), ovatus (6), ellipticus-late (7); seed size: length (8), width (9), length/width (10); seed ornamentation type: rugose (11), reticulate-pusticulate (12), verrucate (13), tuberculate (14), ruminate (15), reticulate-foveate (16), alveolate (17); anticlinal cell wall: unclear (18), flat (19), raised (20), sunken (21); periclinal cell wall: unclear (22), concave (23), convex (24); epidermal cell structure: unclear (25), polygonal (26), alveolar (27); anatomical structure of epidermis: crushed (28), flat (29), polygonal (30); presence of subepidermis layer (31); anatomical structure of subepidermis: flat (32), polygonal (33); testa thickness (34); cotyledon size: length (35), width (36); embryo shapes: elongated (37), ellipticus (38), ovatus (39), circularis (40); embryo size: length (41), width (42). The dissimilarity matrix of the examined species was formed with MVSP (Kovach 2007 ) (Table 4 ). A dendrogram was formed. Also, the cophenetic correlation coefficient was calculated to explain the relation between the dendrogram and dissimilarity matrix (Table 4 , Fig. 4 ). Table 3 The seed anatomical characters of the examined Hyacinthella taxa Taxa Epidermis layers Cotyledon sizes Embryo Embryo sizes Epidermis structures Subepidermis structures Thickness (μm) L (μm) W (μm) shape L (μm) W (μm) H. acutiloba 3-5 layers, crushed cells - 32.59±1.44 1851.49±4.97 1279.35±4.88 Elongated 859.31±1.35 297.14±1.23 H. campanulata 5-7 layers, crushed cells - 49.76±2.31 1913.63±11.86 1605.52±10.79 Ellipticus 487.92±3.08 339.26±4.88 H. glabrescens 3-4 layers, crushed cells 1 layer, flat cells 39.25±4.99 1652.47±5.70 1327.18±8.64 Elongated 940.25±4.95 290.69±3.29 H. heldreichii 4-5 layers, flat cells 1 layer, flat cells 44.08±1.16 1928.54±2.65 1428.39±3.27 Ovatus 446.08±2.21 152.10±1.08 H. hispida 2-3 layers, flat cells - 25.61±0.88 1975.48±7.29 1425.76±3.15 Ellipticus 245.17±1.26 314.22±3.14 H. lazulina 1-2 layers, flat cells - 20.13±0.53 1869.36±3.89 1196.25±2.76 Elongated 532.15±1.43 180.16±0.89 H. lineata 2-3 layers, crushed cells - 29.89±1.35 1854.82±6.55 1441.38±4.84 Elongated 901.66±2.42 193.51±1.35 H. micrantha 1 layer, flat cells - 15.77±0.92 1679.51±3.64 856.44±5.89 Ovatus 484.19±1.29 179.22±2.11 H. nervosa 2 layers, polygonal cells - 26.14±0.99 1964.69±2.88 1409.22±2.54 Ellipticus 625.11±0.95 368.49±3.07 H. siirtensis 2-3 layers, flat cells 1-2 layers, polygonal cells 47.91±1.27 1848.44±3.71 1382.63±3.66 Elongated 753.04±1.83 195.66±2.48 H. venusta 3-4 layers, flat cells - 36.29±1.56 1792.35±3.24 1231.37±1.09 Circularis 471.66±2.05 332.32±2.59 L=length, W=width Table 4 The dissimilarity matrix of the examined taxa Taxa 1 2 3 4 5 6 7 8 9 10 11 H. acutiloba (1) 0 - - - - - - - - - - H. campanulata (2) 500.259 0 - - - - - - - - - H. glabrescens (3) 220.327 593.938 0 - - - - - - - - H. heldreichii (4) 469.139 261.575 591.528 0 - - - - - - - H. hispida (5) 643.679 310.305 773.267 263.080 0 - - - - - - H. lazulina (6) 357.922 444.546 493.273 257.240 405.344 0 - - - - - H. lineata (7) 197.022 472.466 254.995 463.769 678.517 443.988 0 - - - - H. micrantha (8) 602.783 801.702 665.816 626.211 697.988 392.232 739.987 0 - - - H. nervosa (9) 299.449 247.763 457.996 284.402 384.312 314.003 346.740 665.353 0 - - H. siirtensis (10) 180.315 380.598 292.653 323.504 538.976 291.533 160.999 615.634 246.873 0 - H. venusta (11) 396.656 393.956 500.079 300.937 350.818 185.054 502.22 421.107 293.772 352.165 0 Results Macro-morphologically, this study evaluates the seed structures of the studied species containing colour, shape and size. Seed color is black in all examined taxa except H. micrantha , which is of the brown seeds. The shape and size of seeds differ noticeably. The studied seeds are separated into 5 shapes; triangularis, circularis, ellipticus, ellipticus-late and ovatus. Ovatus is the most common type (noted in 4 species). However, circularis and ellipticus are typical forms for Hyacinthella campanulata and H. glabrescens , respectively. Seed sizes vary between 1.51 mm and 3.06 mm in length, and between 1.06 mm and 2.04 mm in width. As H. siirtensis is of the largest seeds, H. campanulata and H. lineata have the smallest seeds (Table 2 , Fig. 1 ). Micro-morphologically, the surface ornamentation, anticlinal and periclinal cell walls, and epidermal cell forms of the studied seeds have been assessed in this work. Seed surface ornamentation is clustered into 7 forms: rugose, reticulate-pusticulate, verrucate, tuberculate, ruminate, reticulate-foveate and alveolate. The most common type is verrucate, as rugose, reticulate-pusticulate, ruminate and reticulate-foveate ornamentation types are found as taxon-specific (Table 2 , Fig. 2 ). The rugose, reticulate-pusticulate, ruminate and reticulate-foveate ornamentation types are shown by only one taxon. The structures of the anticlinal cell walls of the examined species are unclear, raised, flat and sunken. As raised cell walls are generally observed in the reticulate-pusticulate, reticulate-foveate and alveolate ornamentation types, the tuberculate ornamentation type is found where epidermal cells are surrounded by sunken walls. Rugose and ruminate types are connected with unclear type (Table 2 ). There is no connection between concave or convex periclinal cell walls and surface ornamentation type. In the ruminate and rugose forms are seen only unclear periclinal cells. Also, the shape of epidermal cells on the seed surface has exhibited variety. It is polygonal, alveolar and unclear structures. The most common cell type are unclear and alveolar, as polygonal is fairly rare (Table 2 ). The anatomical characters of the seeds are shown in Fig. 3 and Table 3. Testa structures of the studied seeds are usually consisted of 1 layer, the epidermis, formed in the sclerenchymatous tissue. The epidermis layer demonstrates significant differences in cell form, comprising of crushed, flat or polygonal cells, in 1–7 layers, and has surged or straight wall form. The most common kind are flat and crushed while the fewest one is the polygonal type (Table 3, Fig. 3 ). Also, the subepidermis layer is found in some of the studied taxa ( H. glabrescens , H. heldreichii and H. siirtensis ). The subepidermis layer comprises of flat or polygonal cells in 1–2 layers. The most frequently found type is flat, however; the rarest one is polygonal type. The thickness of testa layers ranges from 15.77 µm (in H. micrantha ) to 49.76 µm (in H. campanulata ). The cotyledon sizes vary between 1652.47 µm (in H. glabrescens ) and 1913.63 µm (in H. campanulata ) in length, between 856.44 µm (in H. micrantha ) and 1605.22 µm (in H. campanulata ) in width. In addition to these findings, embryo sizes of the seeds range from 245.17 µm (in H. hispida ) to 940.25 µm (in H. glabrescens ) in length, from 152.10 µm (in H. heldreichii ) to 368.49 µm (in H. nervosa ) in width, and embryos in seeds are formed elongated ( H. acutiloba , H. glabrescens , H. lazulina , H. lineata and H. siirtensis ), elliptical ( H. campanulata , H. hispida and H. nervosa ), oval ( H. heldreichii and H. micrantha ) and circular ( H. venusta ) in shapes (Fig. 3 ). The numerical assessment of the seed morpho-anatomical characters permits the formation of a dendrogram, which reveals the differences-similarities among the studied species. A dendrogram is built as a consequence of the cluster analysis of the studied taxa of Hyacinthella based on the variation of 42 characteristics in 11 species. The cophenetic correlation coefficient is determined to learn the relative between the dendrogram and dissimilarity matrix (Fig. 4 , Table 4 ). The cophenetic correlation between the dissimilarity matrix and dendrogram has been computed as 0.64, representing a good match. Our cluster analysis has separated the taxa into 2 main groups of A and B: Cluster A comprises H. glabrescens , H. siirtensis , H. lineata and H. acutiloba . Cluster B1 includes H. hispida , H. heldreichii , H. nervosa and H. campanulata . Cluster B2 includes H. venusta and H. lazulina. H. micrantha has produced a clade outside these clusters in the dendrogram (Fig. 4 ). H. siirtensis and H. lineata are the most closely related species (dissimilarity coefficient: 160.999), as H. micrantha and H. campanulata are the most distantly related species (dissimilarity coefficient: 801.702) (Table 4 ). Cluster B has the highest number of species compared to another cluster (Fig. 5 ). Discussion The morphological structures of the seeds present significant data in relation to evolutionary relations of the flowering plants (Corner 1976 ; Karaismailoğlu and Erol 2018 ). However, seed morpho-anatomical characters have so far not been comprehensively applied to clarify inter-species relations within genera of the family Asparagaceae, except the genus Muscari (Eroğlu et al 2021 ). This is the first research to exhibit the morpho-anatomical structures of the seeds of the genus Hyacinthella , and it will be a model for following surveys for various closely related genera. The macro-morphological characteristics of the seeds demonstrate differences among the studied Hyacinthella species (Fig. 1 and Table 2 ). The species studied in our research are not very distinct with regard to seed color. Black dominates in the genus, whereas H. micrantha has brown seeds unlike other species. H. glabrescens (ellipticus)- H. hispida (triangularis), H. campanulata (circularis)- H. heldreichii (ovatus), H. acutiloba (triangularis)- H. lineata (ovatus) and H. siirtensis (triangularis)- H. nervosa (ellipticus-late) taxa are more or less similar in terms of leaf, scape and flower structures, but they can be simply distinguished with applying of seed shape. The researches including surface micro-morphological structures of seeds in various plant families provide systematically useful information (Karaismailoğlu 2015 ; Candan et al 2016 ; Karaismailoğlu and Erol 2018 ; Karaismailoğlu et al. 2018 ; Karaismailoğlu and Güner 2019 ; Eroğlu et al 2021 ; Şirin and Karaismailoğlu 2021 ; Gavrilović and Janaćković 2022 ). Also, the importance and effectiveness of scanning electron microscopy in clarifying of taxonomic difficulties and in characterizing of species have been highlighted by many researchers (Heywood 1971 ; Barthlott 1981 ; Eroğlu et al 2021 ). However, there are few papers on the significance of seed micromorphology in the family Asparagaceae (Yıldırım 2015 ; Eroğlu et al 2021 ). This research on 11 Hyacinthella species reveals that seed micro-structures are helpful characters in discrimination of the taxa within the family, as in the genus Muscari (Eroğlu et al 2021 ). All of the examined species have been analyzed for the first time. We noted seven seed surface ornamentation types in this work. In the genus, the most common seed ornamentation type is verrucate. Unlike this work, reticulate or reticulate-areolate ornamentation forms have been frequently found among species from many angiosperm families (Barthlott 1981 ; Erol et al 2006 ; Bona 2013 ; Karaismailoğlu 2015 ; Özbek et al 2018 ; Karaismailoğlu et al 2018 ; Karaismailoğlu and Erol, 2018 ). The rugose (in H. acutiloba ), reticulate-pusticulate (in H. campanulata ), ruminate (in H. hispida ) and reticulate-foveate (in H. lineata ) ornamentation types are taxon specific. Also, ornamentation types are effective in distinguishing closely related taxa from each other, such as H. glabrescens (verrucate)- H. hispida (ruminate), H. campanulata (reticulate-pusticulate)- H. heldreichii (tuberculate), H. acutiloba (rugose)- H. lineata (reticulate-foveate) and H. siirtensis (alveolate)- H. nervosa (verrucate) taxa (Fig. 2 and Table 2 ). Besides, past seed surface surveys have shown that the views and forms of anticlinal and periclinal cell walls are diagnostic characteristics in the creation of inter-taxa interactions (Barthlott 1981 ; Eroğlu et al 2021 ). The types of anticlinal and periclinal cell walls, and epidermal cell structures of the studies species reveal variations among taxa. The macro and micro-morphological outcomes of this investigation are separated all taxa examined from each other, and they are suitable with the former studies done with exomorphic features of seeds of the family Asparagaceae (Yıldırım 2015 ; Eroğlu et al 2021 ; Tugay et al 2021 ). Testa anatomical studies are effective in answering of the taxonomical problems in many plant families (Vaughan et al 1976 ; Manning and Staden 1987 ; Meyer 1991 ; Karaismailoğlu 2015 ; Karaismailoğlu and Erol 2018 ; Eroğlu et al 2021 ). Koul et al. ( 2000 ) and Eroğlu et al. ( 2021 ) have stated that testa forms can be used as a useful characters in the discrimination of the species and in the explanation of their phylogenetic relations. This study is the first such research for the genus and is the precursor to following works. In this study, it is observed that the testa layers usually contain one layer as the epidermis in the sclerotic form. The epidermis structures display variations among the species. This 1–7 layered epidermis can compose of flat, crushed, or polygonal cells. The most common form is crushed, as the rarest is polygonal type. Some of the studied species, which are H. glabrescens , H. heldreichii and H. nervosa , have the subepidermis layer under epidermis occurring polygonal or flat cells. The presence of the subepidermis layer is sufficient to distinguish some closely related taxa such as H. glabrescens - H. hispida , H. heldreichii - H. campanulata and H. nervosa - H. siirtensis . Also, the differences in testa thicknesses and cotyledon sizes of the seeds of Turkish Hyacinthella have been detected. The most distant species are H. micrantha and H. campanulata . The embryos of Hyacinthella taxa display a broad variation in terms of shapes and sizes. Embryos are elongated form in H. acutiloba , H. glabrescens , H. lazulina , H. lineata and H. siirtensis , elliptical form in H. campanulata , H. hispida and H. nervosa , oval form in H. heldreichii and H. micrantha , circular form in H. venusta . Closely related taxa such as H. glabrescens - H. hispida , H. campanulata - H. heldreichii , H. siirtensis - H. nervosa that are morphologically similar can easily be distinguished from each other according to their embryo shape (Table 3 and Fig. 3 ). In this study, the seed anatomical characters such as the structures of the epidermis and subepidermis, thickness of the testa, cotyledon sizes and embryo shapes are rather efficient and useful in separating almost all of the studied Hyacinthella taxa. This can be explained as follows: the seed anatomical characters are useful additional characteristics in the Hyacinthella , and they are able to assist in the taxonomy of this genus. The obtained outcomes are compatible with parallel earlier works conducted on seed structure of some taxa of the genera Crocus L. and Romulea Maratti in the closely related family Iridaceae, and genus Muscari in the family Asparagaceae (Grilli Caiola et al 2010 ; Karaismailoğlu 2015 ; Karaismailoğlu et al 2018 ; Eroğlu et al 2021 ). A dendrogram has been created to assess the morpho-anatomical characteristics of the seeds of Turkish Hyacinthella taxa with UPGMA cluster analysis (Fig. 4 ). The dendrogram, displaying two main clusters, is partly consistent with the results of Persson and Wendelbo ( 1984 ). According to the descriptions in the Flora of Turkey, the systematic affinity in the closely related H. campanulata - H. heldreichii , and H. acutiloba - H. lineata taxa is partially preserved. However, H. siirtensis - H. nervosa and H. glabrescens - H. hispida taxa are located in different clusters in the dendrogram (Figs. 4 – 5 and Table 4 ). In addition, H. micrantha has been described as an isolated species with a very different description because of its unique leaf and veining properties within the genus in the Flora of Turkey. This species has showed similar characteristics in the dendrogram created according to seed morphological and anatomical features, and formed a clade apart from 2 main clusters. As a result, this work display that morphological and anatomical seed characteristics of Turkish Hyacinthella species presents significant and consistent insights into the taxonomy of species within the genus. Key to studied Hyacinthella taxa, based on seed characters 1.Seed colour is brown………………………...………………………………… H. micrantha 1.Seed colour is black………………………………………………………………………….2 2.Seed shape is circularis, ellipticus or ellipticus-late………………………………………....3 2.Seed shape is triangularis or ovatus………………………………………………………….6 3.Seed shape is circularis……………..………………….………………...….. H. campanulate 3.Seed shape is ellipticus or ellipticus-late..…………………………………………………...4 4.Ellipticus…………………………………………………………………...… H. glabrescens 4.Ellipticus-late………………………………………………………………………………...5 5.Seed surface ornamentation is verrucate…………………………………………. H. lazuline 5.Seed surface ornamentation is tuberculate……………………….……………….. H. venusta 6.Seed shape is triangularis…………………..………………………………………………..7 6.Seed shape is ovatus……………………………………………………….…………………9 7.Seed surface ornamentation is alveolate………………………………………… H. siirtensis 7.Seed surface ornamentation is rugose or ruminate…………………………………………..8 8.Rugose…………………………………………………………………………... H. acutiloba 8.Ruminate……………………………………………………………….…………. H. hispida 9.Seed surface ornamentation is tuberculate or verrucate…………………………………….10 9.Seed surface ornamentation is alveolate or reticulate-foveate……………………………...11 10.Tuberculate………………………………………………………...………… H. heldreichii 10.Verrucate……………………………………………………………...………… H. nervosa 11.Alveolate……………………………………………………………………… H. micrantha 11.Reticulate-foveate………………………………………………...……………… H. lineata Declarations Acknowledgement The authors thank Haydar Öztaş, Barış Bani, and Hüseyin Güldal for their assistance and contributions in the study. Contributions M.C. Karaismailoğlu and H. Eroğlu contributed to the study conception and design. Material preparation, data collection and analysis were performed by M.C. Karaismailoğlu, S.M. Pınar, H. Eroğlu, and M. Fidan. The first draft of the manuscript was written by M.C. Karaismailoğlu, and H. Eroğlu and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript. Funding No financial support was used for this study. Conflict of interest On behalf of all authors, the corresponding author states no conflict of interest. Competing Interest The authors have no competing interests Data Availability Statement The datasets generated during and/or analysed during the current study are available from the corresponding author on reasonable request. References APG (2009) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161(2): 105–121. Barthlott W (1981) Epidermal and seed surface characters of plants: Systematic applicability and some evolutionary aspects. Nordic Journal of Botany 1: 345–355. Bona M (2013) Seed-coat microsculpturing of Turkish Lepidium (Brassicaceae) and its systematic application. Turkish Journal of Botany 37: 662–668. 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Yıldırım H (2015) Muscari atillae (Asparagaceae): a new species from Eastern Anatolia, Turkey. Phytotaxa 213: 291–295. Additional Declarations No competing interests reported. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-1635226","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":105843256,"identity":"bcf7e680-272e-4655-8ec5-27bc56d0ef77","order_by":0,"name":"Hüseyin EROĞLU","email":"","orcid":"","institution":"Yüzüncü Yıl University","correspondingAuthor":false,"prefix":"","firstName":"Hüseyin","middleName":"","lastName":"EROĞLU","suffix":""},{"id":105843257,"identity":"ab9a863b-efa0-47df-b0f3-9760cb8e1ac2","order_by":1,"name":"Mehmet Cengiz KARAİSMAİLOĞLU","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA5klEQVRIie3PMQrCMBSA4VcepEtqV4uDV6hLi6h4lYqgi2fQQqAuoqtO3qKuLQEv0aWTc7qIiINpdHCx7SiYf3oJ+UgCoNP9YC6CkZSDjXhWO8SsJ6CIsyYzgEASrCPwJu6OeopAHfFNTBJx5/I6es2LW9xtyaeKYvGd9BkJ0sOWg4/WqbcPsl6EgM4hrngYpy63NlxaK+7QIDMkIWg1IXK4lGTcjNCbIqQkk1ry+ks4B4cRz9nPsmmEBqv8i2/zNBePAdg2v7TFMBsd1ywVRQVRGRFbhR/L8NvBjx7LBod0Op3ub3sCWBJNr3JzGosAAAAASUVORK5CYII=","orcid":"","institution":"Bartın University","correspondingAuthor":true,"prefix":"","firstName":"Mehmet","middleName":"Cengiz","lastName":"KARAİSMAİLOĞLU","suffix":""},{"id":105843258,"identity":"23bb944e-bff8-4449-9a46-f3abc863048c","order_by":2,"name":"Mehmet FİDAN","email":"","orcid":"","institution":"Siirt University","correspondingAuthor":false,"prefix":"","firstName":"Mehmet","middleName":"","lastName":"FİDAN","suffix":""},{"id":105843259,"identity":"2ae5962e-ddfc-4330-a5c3-03baa936e86e","order_by":3,"name":"Süleyman Mesut PINAR","email":"","orcid":"","institution":"Bartın University","correspondingAuthor":false,"prefix":"","firstName":"Süleyman","middleName":"Mesut","lastName":"PINAR","suffix":""}],"badges":[],"createdAt":"2022-05-08 11:59:08","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-1635226/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-1635226/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":21579662,"identity":"b91ee0d1-cbd3-430f-bddb-ac94c51f337a","added_by":"auto","created_at":"2022-05-17 18:46:12","extension":"jpeg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":421769,"visible":true,"origin":"","legend":"\u003cp\u003eThe seeds of the studied \u003cem\u003eHyacinthella\u003c/em\u003e taxa; \u003cstrong\u003e1:\u003c/strong\u003e \u003cem\u003eH. acutiloba\u003c/em\u003e, \u003cstrong\u003e2:\u003c/strong\u003e \u003cem\u003eH. campanulata\u003c/em\u003e, \u003cstrong\u003e3:\u003c/strong\u003e \u003cem\u003eH. glabrescens\u003c/em\u003e, \u003cstrong\u003e4:\u003c/strong\u003e \u003cem\u003eH. heldreichii\u003c/em\u003e, \u003cstrong\u003e5:\u003c/strong\u003e \u003cem\u003eH. hispida\u003c/em\u003e, \u003cstrong\u003e6:\u003c/strong\u003e \u003cem\u003eH. lazulina\u003c/em\u003e, \u003cstrong\u003e7:\u003c/strong\u003e \u003cem\u003eH. lineata\u003c/em\u003e, \u003cstrong\u003e8:\u003c/strong\u003e \u003cem\u003eH. micrantha\u003c/em\u003e, \u003cstrong\u003e9:\u003c/strong\u003e \u003cem\u003eH. nervosa\u003c/em\u003e, \u003cstrong\u003e10:\u003c/strong\u003e \u003cem\u003eH. siirtensis\u003c/em\u003e and \u003cstrong\u003e11:\u003c/strong\u003e \u003cem\u003eH. venusta\u003c/em\u003e (Scale bars= 1 mm).\u003c/p\u003e\u003cp\u003e\u003cbr\u003e\u003c/p\u003e","description":"","filename":"floatimage1.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-1635226/v1/db34c2e9704170300727f448.jpeg"},{"id":21579041,"identity":"a7a79917-8eae-4ee0-95bd-b3ab1b523fe2","added_by":"auto","created_at":"2022-05-17 18:41:12","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":2871158,"visible":true,"origin":"","legend":"\u003cp\u003eThe SEM pictures of the examined \u003cem\u003eHyacinthella\u003c/em\u003e taxa; \u003cstrong\u003e1:\u003c/strong\u003e \u003cem\u003eH. acutiloba\u003c/em\u003e, \u003cstrong\u003e2:\u003c/strong\u003e \u003cem\u003eH. campanulata\u003c/em\u003e, \u003cstrong\u003e3:\u003c/strong\u003e \u003cem\u003eH. glabrescens\u003c/em\u003e, \u003cstrong\u003e4:\u003c/strong\u003e \u003cem\u003eH. heldreichii\u003c/em\u003e, \u003cstrong\u003e5:\u003c/strong\u003e \u003cem\u003eH. hispida\u003c/em\u003e, \u003cstrong\u003e6:\u003c/strong\u003e \u003cem\u003eH. lazulina\u003c/em\u003e.\u003c/p\u003e\u003cp\u003e\u003cbr\u003e\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-1635226/v1/f5028ff48cd856b7aed13b89.png"},{"id":21579037,"identity":"3d642bd7-d496-422b-b66b-1d073f2a725b","added_by":"auto","created_at":"2022-05-17 18:41:12","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":2419042,"visible":true,"origin":"","legend":"\u003cp\u003eThe SEM pictures of the examined \u003cem\u003eHyacinthella\u003c/em\u003e taxa; \u003cstrong\u003e7:\u003c/strong\u003e \u003cem\u003eH. lineata\u003c/em\u003e, \u003cstrong\u003e8:\u003c/strong\u003e \u003cem\u003eH. micrantha\u003c/em\u003e, \u003cstrong\u003e9:\u003c/strong\u003e \u003cem\u003eH. nervosa\u003c/em\u003e, \u003cstrong\u003e10:\u003c/strong\u003e \u003cem\u003eH. siirtensis\u003c/em\u003e and \u003cstrong\u003e11:\u003c/strong\u003e \u003cem\u003eH. venusta\u003c/em\u003e (black arrow indicates secondary structures on the surface).\u003c/p\u003e\u003cp\u003e\u003cbr\u003e\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-1635226/v1/262aa754231a77d487386a8e.png"},{"id":21579996,"identity":"8c99ff15-e2e2-497a-bc72-73d17e4cd8b4","added_by":"auto","created_at":"2022-05-17 18:51:12","extension":"jpeg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":1164271,"visible":true,"origin":"","legend":"\u003cp\u003eThe seed anatomical structures of the examined \u003cem\u003eHyacinthella\u003c/em\u003e taxa; \u003cstrong\u003e1:\u003c/strong\u003e \u003cem\u003eH. acutiloba\u003c/em\u003e, \u003cstrong\u003e2:\u003c/strong\u003e \u003cem\u003eH. campanulata\u003c/em\u003e, \u003cstrong\u003e3:\u003c/strong\u003e \u003cem\u003eH. glabrescens\u003c/em\u003e, \u003cstrong\u003e4:\u003c/strong\u003e \u003cem\u003eH. heldreichii\u003c/em\u003e (t= testa, co= cotyledon, e= epidermis, se= subepidermis, sm= storage material, em= embryo).\u003c/p\u003e\u003cp\u003e\u003cbr\u003e\u003c/p\u003e","description":"","filename":"floatimage4.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-1635226/v1/c9272e442b0aaae4f1d37432.jpeg"},{"id":21579044,"identity":"298d8042-9eb2-4379-947b-47fdc931a408","added_by":"auto","created_at":"2022-05-17 18:41:12","extension":"jpeg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":1001339,"visible":true,"origin":"","legend":"\u003cp\u003eThe seed anatomical structures of the examined \u003cem\u003eHyacinthella\u003c/em\u003e taxa; \u003cstrong\u003e5:\u003c/strong\u003e \u003cem\u003eH. hispida\u003c/em\u003e, \u003cstrong\u003e6:\u003c/strong\u003e \u003cem\u003eH. lazuline\u003c/em\u003e, \u003cstrong\u003e7:\u003c/strong\u003e \u003cem\u003eH. lineata\u003c/em\u003e, \u003cstrong\u003e8:\u003c/strong\u003e \u003cem\u003eH. micrantha\u003c/em\u003e (t= testa, co= cotyledon, e= epidermis, se= subepidermis, sm= storage material, em= embryo).\u003c/p\u003e\u003cp\u003e\u003cbr\u003e\u003c/p\u003e","description":"","filename":"floatimage5.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-1635226/v1/4a6542f17dcce5fc23ea348d.jpeg"},{"id":21579667,"identity":"af416846-f2a2-47d2-b573-69d408c56757","added_by":"auto","created_at":"2022-05-17 18:46:12","extension":"jpeg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":774487,"visible":true,"origin":"","legend":"\u003cp\u003eThe seed anatomical structures of the examined \u003cem\u003eHyacinthella\u003c/em\u003e taxa; \u003cstrong\u003e9:\u003c/strong\u003e \u003cem\u003eH. nervosa\u003c/em\u003e, \u003cstrong\u003e10:\u003c/strong\u003e \u003cem\u003eH. siirtensis\u003c/em\u003e and \u003cstrong\u003e11:\u003c/strong\u003e \u003cem\u003eH. venusta\u003c/em\u003e (t= testa, co= cotyledon, e= epidermis, se= subepidermis, sm= storage material, em= embryo).\u003c/p\u003e\u003cp\u003e\u003cbr\u003e\u003c/p\u003e","description":"","filename":"floatimage6.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-1635226/v1/13690d391b30215c8fd04b1d.jpeg"},{"id":21579043,"identity":"9dedc0a4-a838-4c89-a3fe-ad78de4bd793","added_by":"auto","created_at":"2022-05-17 18:41:12","extension":"jpeg","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":79972,"visible":true,"origin":"","legend":"\u003cp\u003eThe dendrogram of the examined \u003cem\u003eHyacinthella\u003c/em\u003e taxa.\u003c/p\u003e","description":"","filename":"floatimage7.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-1635226/v1/7c43538ce962320273742df0.jpeg"},{"id":21579038,"identity":"08512863-20da-4fa7-b312-48260814b536","added_by":"auto","created_at":"2022-05-17 18:41:12","extension":"jpeg","order_by":8,"title":"Figure 8","display":"","copyAsset":false,"role":"figure","size":88037,"visible":true,"origin":"","legend":"\u003cp\u003ePrincipal component analysis of the examined \u003cem\u003eHyacinthella\u003c/em\u003e taxa.\u003c/p\u003e\u003cp\u003e\u003cbr\u003e\u003c/p\u003e","description":"","filename":"floatimage8.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-1635226/v1/bdf027cfe2395728bbcf2323.jpeg"},{"id":21822275,"identity":"c8639737-3451-4946-8813-2189ecc856a8","added_by":"auto","created_at":"2022-05-24 13:14:30","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3113919,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-1635226/v1/58e1e9b4-311a-47db-b1ac-f6751765b463.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Seed Morphological and Anatomical Structures as Taxonomy Tool for Turkish Hyacinthella Schur (Asparagaceae) Taxa","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe family Asparagaceae is of 7 subfamilies, 114 genera and circa 2900 taxa in worldwide (Chase et al \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Christenhusz and Byng \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). It represents with 19 genera and 182 species in the Flora of Turkey (G\u0026uuml;ner et al \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). \u003cem\u003eHyacinthella\u003c/em\u003e Schur consists of 17 species, 1 subspecies and 1 variety belonging to 2 sections, and it disperses from Balkan Peninsula to Turkey and along northern Iran (Persson and Wendelbo \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1981\u003c/span\u003e; Speta \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Govaerts et al \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). According to Mathew (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e1987\u003c/span\u003e), Turkey is one of the diversity centers of the genus. The genus was placed within the family Liliaceae in its first classification based on limited macromorphological characters. As a result of molecular studies, \u003cem\u003eHyacinthella\u003c/em\u003e was first transferred to the family Hyacinthaceae and then to the family Asparagaceae (APG III 2009).\u003c/p\u003e \u003cp\u003eThe genus \u003cem\u003eHyacinthella\u003c/em\u003e was reported as 10 taxa with 9 species and 1 hybrid species (\u003cem\u003eH. micrantha\u003c/em\u003e (Boiss.) Chouard \u0026times; \u003cem\u003eH. heldreichii\u003c/em\u003e (Boiss.) Chouard.) to Flora of Turkey (Persson and Wendelbo \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1984\u003c/span\u003e). Subsequently, it was defined that the samples belonging to the declared hybrid species were not hybrid and they were new species (\u003cem\u003eH. lazulina\u003c/em\u003e K. Perss. and Jim. Perss.) (Persson and Persson \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1992\u003c/span\u003e). Afterwards, another new species was published from Turkey (Persson \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). \u003cem\u003eHyacinthella millingenii\u003c/em\u003e (Post) Feinbrun endemic to Cyprus was involved in the latest update flora checklist performed in Turkey; however, no specific specimens or location of the species was included, and it was noted that \"the status of the species needs to be clarified\" (Ekim \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Therefore, the number of taxa belonging to the genus \u003cem\u003eHyacinthella\u003c/em\u003e in Turkey has updated as 11 species, including 10 endemic species.\u003c/p\u003e \u003cp\u003eThe main characters utilized in taxonomy of the genus \u003cem\u003eHyacinthella\u003c/em\u003e are pedicel presence-absence and pedicel length, flower color, lobe-flower ratio, anther-filament ratio, flower shape, leaf shape, leaf edge, secondary leaf to primary leaf ratio (Persson and Wendelbo \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Persson \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). According to Persson and Wendelbo (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1984\u003c/span\u003e), the characters used in the taxonomy of the genus may not be consistent in terms of features such as color, length and number. Accordingly, new distinctive and consistent characters are needed to contribute to the systematics of the genus.\u003c/p\u003e \u003cp\u003eThe seeds are of the characteristic morphological features such as shape, colour, size and surface ornamentation types, and they can offer valuable data in of the elucidating taxonomic problems and in of the establishing systematic relationships (Barthlott \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1981\u003c/span\u003e; Karaismailoğlu and Erol \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Besides, the anatomical characters are typically as helpful as morphological characters for plant taxonomy, and they often are valuable in the separation of closely correlated taxa (Karamian et al \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Karaismailoğlu \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The importance of the seed in genera within the family Asparagaceae has often been overlooked. The only study showing the importance of comprehensive seed morphology and anatomy within the family was carried out by Eroğlu et al. (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) on Turkish \u003cem\u003eMuscari\u003c/em\u003e, which is a genus closely related to \u003cem\u003eHyacinthella\u003c/em\u003e. In the mentioned study, it was seen that seed morphological and anatomical characters easily distinguished 36 \u003cem\u003eMuscari\u003c/em\u003e taxa from each other.\u003c/p\u003e \u003cp\u003eSome morphological, anatomical, cytological and palynological studies on some taxa of the genus \u003cem\u003eHyacinthella\u003c/em\u003e have been made, so far (Feinbrun \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1961\u003c/span\u003e; Persson and Wendelbo \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1981\u003c/span\u003e, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Puizina et al \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Selvi et al \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Yetişen et al \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Tekin and Meri\u0026ccedil; \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Tzonev and Panovska \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Doğu \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). However, the morphological and anatomical characteristics of seeds have been ignored in the classification of taxa belonging to the genus. The aim of this work is to examine the morphological and anatomical characters of seeds of Turkish \u003cem\u003eHyacinthella\u003c/em\u003e taxa for the first time, and discuss the systematic practice of these characteristics.\u003c/p\u003e"},{"header":"Material And Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003ePlant materials\u003c/h2\u003e \u003cp\u003eSpecies were gathered from different phytogeographical areas of Turkey and were stored at VANF (Van Y\u0026uuml;z\u0026uuml;nc\u0026uuml; Yıl University Herbarium). The studied species and their locations were offered in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe collections data of the investigated \u003cem\u003eHyacinthella\u003c/em\u003e taxa and their locations.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"3\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eLocality\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eVoucher no.\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eHyacinthella acutiloba\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eB7 Malatya: Yazıhan, Girmana Canyon, rocky places, 1063 m, 31 March 2021.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1593\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. campanulata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eC4 Konya: Meram, Konya-Beyşehir road, 5 km to Altınapa Dam, marly steppe, 1310 m. 26 May 2021.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1719\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. glabrescens\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eC5 Mersin: Tarsus, south of Cevizli\u0026ccedil;eşme Village, rocky \u003cem\u003ePinus\u003c/em\u003e yards, 1030 m, 17 March 2020.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1530\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. heldreichii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eC3 Antalya: Korkuteli, from the old road from Antalya to Korkuteli, northwest of G\u0026uuml;ll\u0026uuml;k Mountain, \u003cem\u003ePinus\u003c/em\u003e yards, 702 m. 19 March 2020.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1532\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. hispida\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eC5 Mersin: Tarsus, \u0026Ccedil;amalan Village, southeast of martyrdom, \u003cem\u003ePinus\u003c/em\u003e and \u003cem\u003eJuniperus\u003c/em\u003e yards, 776 m, 17 March 2020.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1529\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. lazulina\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eC4 Mersin: Silifke, between Demircili and İmamlı villages, 1 km to İmamlı Village, maquis yards, 718 m, 18 March 2020.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1531\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. lineata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eC2 Denizli: Honaz, south of G\u0026uuml;rleyik Village, around of marble quarry, marly steppe, 424 m, 20 March 2020.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1533\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. micrantha\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eA4 Kastamonu: Kastamonu University Kuzeykent campus, around of football stadium, \u003cem\u003ePinus\u003c/em\u003e yards, 820 m, 18 March 2019.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1450\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. nervosa\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eC6 Gaziantep: Şahinbey, Gaziantep-Kilis road, 1 km from K\u0026uuml;\u0026ccedil;\u0026uuml;kkızılhisar, stony-rocky steppe, 859 m, 16 March 2020.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1527\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. siirtensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eC8 Mardin: Artuklu, between Mardin and Diyarbakır, Akresta pass, Hut Farm road, streamside, steppe, 1138 m, 16 March 2020.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1524\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. venusta\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eC4 Konya; between Taşkent and Başyayla, Bolay highland, roadside, steppe, 1677 m, 26 May 2021.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH.Eroglu 1717\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e\u003ch2\u003eMacro And Micro-morphological Analysis\u003c/h2\u003e\u003cp\u003eMacro-morphological structures of the seeds containing colour, shape and size were studied with 100 seeds of 10 specimens per species using a Leica EZ4 binocular microscope with a HD camera (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Micro-morphological structures of the seeds including surface ornamentation, anticlinal and periclinal cell walls, and epidermal cells were analyzed with a Scanning Electron Microscope (SEM) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003e, Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Firstly, seeds were positioned on the stubb with a carbon tape and covered with gold, then observed with a Zeiss LEO 440 SEM (Karaismailoğlu \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe seed morphological characters of the studied Hyacinthella taxa.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"10\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eTaxa\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eColor\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eShape\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003eSeed sizes\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eL/W\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSeed surface\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eAnticlinal\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003ePericlinal\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eEpidermal\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eL (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eW (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eornamentation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003ecell wall\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003ecell wall\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003ecell structure\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. acutiloba\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTriangularis\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.94\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eRugose\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eUnclear\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eUnclear\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eUnclear\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. campanulata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCircularis\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.51\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.27\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eReticulate-Pusticulate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eRaised\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eConcave\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003ePolygonal cells\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. glabrescens\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEllipticus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.43\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.85\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVerrucate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eFlat\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eConvex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eUnclear\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. heldreichii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eOvatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.95\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eTuberculate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eSunken\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eConcave\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAlveolar cells\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. hispida\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTriangularis\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.98\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.59\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eRuminate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eUnclear\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eUnclear\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eUnclear\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. lazulina\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEllipticus-late\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.73\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVerrucate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eFlat\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eConvex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eUnclear\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. lineata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eOvatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.59\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eReticulate-Foveate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eRaised\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eConcave\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003ePolygonal and alveolar cells\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. micrantha\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBrown\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eOvatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAlveolate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eRaised\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eConcave\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAlveolar cells\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. nervosa\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eOvatus\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVerrucate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eFlat\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eConvex\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eUnclear\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. siirtensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTriangularis\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.06\u0026thinsp;\u0026plusmn;\u0026thinsp;0.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAlveolate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eRaised\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eConcave\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAlveolar cells\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eH. venusta\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eBlack\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEllipticus-late\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eTuberculate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eSunken\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eConcave\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAlveolar cells\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"10\" nameend=\"c10\" namest=\"c1\"\u003e \u003cp\u003eL\u0026thinsp;=\u0026thinsp;length, W\u0026thinsp;=\u0026thinsp;width\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003ch2\u003eAnatomical Analysis\u003c/h2\u003e\u003cp\u003eThe examination of the seed anatomical structures was performed with dry herbarium materials. Cross-sections were obtained from the middle of the seed with a fully automatic microtome (Thermo Shonda Met Finesse, Thermo). They were treated with a series of alcohol and xylene, stained with hematoxylin and eosin-Y in a staining device (ASC 720 Medite) and covered with using Entellan (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e, Table\u0026nbsp;3) (Karaismailoğlu \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Karaismailoğlu and Erol \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Anatomical structures were surveyed with an Olympus CX31 light microscope and Kameram Imaging Software (KAMERAM12 CCD, Argenit Micro System Ltd., Turkey).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe terminology utilized for seed morpho-anatomical features is suitable with Stearn (\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1985\u003c/span\u003e).\u003c/p\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eClustering of taxa was made with utilizing the grouping analysis method (UPGMA) in MultiVariate Statistical Package (MVSP) as per the 42 characteristics in Tables\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e\u0026ndash;3 (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The characters in statistical analysis were utilized seed colour: black (1), brown (2); seed shape: triangularis (3), circularis (4), ellipticus (5), ovatus (6), ellipticus-late (7); seed size: length (8), width (9), length/width (10); seed ornamentation type: rugose (11), reticulate-pusticulate (12), verrucate (13), tuberculate (14), ruminate (15), reticulate-foveate (16), alveolate (17); anticlinal cell wall: unclear (18), flat (19), raised (20), sunken (21); periclinal cell wall: unclear (22), concave (23), convex (24); epidermal cell structure: unclear (25), polygonal (26), alveolar (27); anatomical structure of epidermis: crushed (28), flat (29), polygonal (30); presence of subepidermis layer (31); anatomical structure of subepidermis: flat (32), polygonal (33); testa thickness (34); cotyledon size: length (35), width (36); embryo shapes: elongated (37), ellipticus (38), ovatus (39), circularis (40); embryo size: length (41), width (42). The dissimilarity matrix of the examined species was formed with MVSP (Kovach \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e4\u003c/span\u003e). A dendrogram was formed. Also, the cophenetic correlation coefficient was calculated to explain the relation between the dendrogram and dissimilarity matrix (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e4\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003ctable width=\"0\"\u003e\n\u003ctbody\u003e\n\u003ctr\u003e\n\u003ctd colspan=\"9\" width=\"907\"\u003e\n\u003cp\u003e\u003cstrong\u003eTable 3\u003c/strong\u003e The seed anatomical characters of the examined \u003cem\u003eHyacinthella\u003c/em\u003e taxa\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"0\"\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd rowspan=\"2\" width=\"97\"\u003e\n\u003cp\u003eTaxa\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"256\"\u003e\n\u003cp\u003eEpidermis layers\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\u0026nbsp;\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"187\"\u003e\n\u003cp\u003eCotyledon sizes\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eEmbryo\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"3\" width=\"198\"\u003e\n\u003cp\u003eEmbryo sizes\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003eEpidermis structures\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003eSubepidermis structures\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003eThickness (\u0026mu;m)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003eL (\u0026mu;m)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003eW (\u0026mu;m)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eshape\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003eL (\u0026mu;m)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003eW (\u0026mu;m)\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. acutiloba\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e3-5 layers, crushed cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e32.59\u0026plusmn;1.44\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1851.49\u0026plusmn;4.97\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1279.35\u0026plusmn;4.88\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eElongated\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e859.31\u0026plusmn;1.35\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e297.14\u0026plusmn;1.23\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. campanulata\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e5-7 layers, crushed cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e49.76\u0026plusmn;2.31\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1913.63\u0026plusmn;11.86\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1605.52\u0026plusmn;10.79\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eEllipticus\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e487.92\u0026plusmn;3.08\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e339.26\u0026plusmn;4.88\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. glabrescens\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e3-4 layers, crushed cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e1 layer, flat cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e39.25\u0026plusmn;4.99\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1652.47\u0026plusmn;5.70\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1327.18\u0026plusmn;8.64\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eElongated\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e940.25\u0026plusmn;4.95\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e290.69\u0026plusmn;3.29\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. heldreichii\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e4-5 layers, flat cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e1 layer, flat cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e44.08\u0026plusmn;1.16\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1928.54\u0026plusmn;2.65\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1428.39\u0026plusmn;3.27\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eOvatus\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e446.08\u0026plusmn;2.21\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e152.10\u0026plusmn;1.08\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. hispida\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e2-3 layers, flat cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e25.61\u0026plusmn;0.88\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1975.48\u0026plusmn;7.29\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1425.76\u0026plusmn;3.15\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eEllipticus\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e245.17\u0026plusmn;1.26\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e314.22\u0026plusmn;3.14\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. lazulina\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e1-2 layers, flat cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e20.13\u0026plusmn;0.53\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1869.36\u0026plusmn;3.89\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1196.25\u0026plusmn;2.76\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eElongated\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e532.15\u0026plusmn;1.43\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e180.16\u0026plusmn;0.89\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. lineata\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e2-3 layers, crushed cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e29.89\u0026plusmn;1.35\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1854.82\u0026plusmn;6.55\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1441.38\u0026plusmn;4.84\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eElongated\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e901.66\u0026plusmn;2.42\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e193.51\u0026plusmn;1.35\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. micrantha\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e1 layer, flat cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e15.77\u0026plusmn;0.92\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1679.51\u0026plusmn;3.64\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e856.44\u0026plusmn;5.89\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eOvatus\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e484.19\u0026plusmn;1.29\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e179.22\u0026plusmn;2.11\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. nervosa\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e2 layers, polygonal cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e26.14\u0026plusmn;0.99\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1964.69\u0026plusmn;2.88\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1409.22\u0026plusmn;2.54\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eEllipticus\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e625.11\u0026plusmn;0.95\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e368.49\u0026plusmn;3.07\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. siirtensis\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e2-3 layers, flat cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e1-2 layers, polygonal cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e47.91\u0026plusmn;1.27\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1848.44\u0026plusmn;3.71\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1382.63\u0026plusmn;3.66\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eElongated\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e753.04\u0026plusmn;1.83\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e195.66\u0026plusmn;2.48\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e\u003cem\u003eH. venusta\u003c/em\u003e\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"149\"\u003e\n\u003cp\u003e3-4 layers, flat cells\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"107\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"88\"\u003e\n\u003cp\u003e36.29\u0026plusmn;1.56\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"90\"\u003e\n\u003cp\u003e1792.35\u0026plusmn;3.24\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"97\"\u003e\n\u003cp\u003e1231.37\u0026plusmn;1.09\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"81\"\u003e\n\u003cp\u003eCircularis\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"110\"\u003e\n\u003cp\u003e471.66\u0026plusmn;2.05\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd colspan=\"2\" width=\"88\"\u003e\n\u003cp\u003e332.32\u0026plusmn;2.59\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd colspan=\"9\" width=\"907\"\u003e\n\u003cp\u003eL=length, W=width\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"0\"\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u0026nbsp;\u003c/p\u003e\n\u003ctable width=\"0\"\u003e\n\u003ctbody\u003e\n\u003ctr\u003e\n\u003ctd colspan=\"12\" width=\"842\"\u003e\n\u003cp\u003e\u003cstrong\u003eTable 4 \u003c/strong\u003eThe dissimilarity matrix of the examined taxa\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003eTaxa\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e1\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e2\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e3\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e4\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e5\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e6\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e7\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e8\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e9\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e10\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e11\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. acutiloba\u003c/em\u003e (1)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. campanulata\u003c/em\u003e (2)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e500.259\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. glabrescens \u003c/em\u003e(3)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e220.327\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e593.938\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. heldreichii \u003c/em\u003e(4)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e469.139\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e261.575\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e591.528\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. hispida \u003c/em\u003e(5)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e643.679\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e310.305\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e773.267\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e263.080\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. lazulina \u003c/em\u003e(6)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e357.922\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e444.546\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e493.273\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e257.240\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e405.344\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. lineata\u003c/em\u003e (7)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e197.022\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e472.466\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e254.995\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e463.769\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e678.517\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e443.988\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. micrantha\u003c/em\u003e (8)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e602.783\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e801.702\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e665.816\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e626.211\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e697.988\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e392.232\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e739.987\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. nervosa\u003c/em\u003e (9)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e299.449\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e247.763\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e457.996\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e284.402\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e384.312\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e314.003\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e346.740\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e665.353\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. siirtensis\u003c/em\u003e (10)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e180.315\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e380.598\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e292.653\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e323.504\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e538.976\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e291.533\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e160.999\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e615.634\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e246.873\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e-\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003ctr\u003e\n\u003ctd width=\"154\"\u003e\n\u003cp\u003e\u003cem\u003eH. venusta\u003c/em\u003e (11)\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e396.656\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e393.956\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e500.079\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e300.937\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e350.818\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e185.054\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e502.22\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e421.107\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e293.772\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"66\"\u003e\n\u003cp\u003e352.165\u003c/p\u003e\n\u003c/td\u003e\n\u003ctd width=\"23\"\u003e\n\u003cp\u003e0\u003c/p\u003e\n\u003c/td\u003e\n\u003c/tr\u003e\n\u003c/tbody\u003e\n\u003c/table\u003e "},{"header":"Results","content":"\u003cp\u003eMacro-morphologically, this study evaluates the seed structures of the studied species containing colour, shape and size. Seed color is black in all examined taxa except \u003cem\u003eH. micrantha\u003c/em\u003e, which is of the brown seeds. The shape and size of seeds differ noticeably. The studied seeds are separated into 5 shapes; triangularis, circularis, ellipticus, ellipticus-late and ovatus. Ovatus is the most common type (noted in 4 species). However, circularis and ellipticus are typical forms for \u003cem\u003eHyacinthella campanulata\u003c/em\u003e and \u003cem\u003eH. glabrescens\u003c/em\u003e, respectively. Seed sizes vary between 1.51 mm and 3.06 mm in length, and between 1.06 mm and 2.04 mm in width. As \u003cem\u003eH. siirtensis\u003c/em\u003e is of the largest seeds, \u003cem\u003eH. campanulata\u003c/em\u003e and \u003cem\u003eH. lineata\u003c/em\u003e have the smallest seeds (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMicro-morphologically, the surface ornamentation, anticlinal and periclinal cell walls, and epidermal cell forms of the studied seeds have been assessed in this work. Seed surface ornamentation is clustered into 7 forms: rugose, reticulate-pusticulate, verrucate, tuberculate, ruminate, reticulate-foveate and alveolate. The most common type is verrucate, as rugose, reticulate-pusticulate, ruminate and reticulate-foveate ornamentation types are found as taxon-specific (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The rugose, reticulate-pusticulate, ruminate and reticulate-foveate ornamentation types are shown by only one taxon. The structures of the anticlinal cell walls of the examined species are unclear, raised, flat and sunken. As raised cell walls are generally observed in the reticulate-pusticulate, reticulate-foveate and alveolate ornamentation types, the tuberculate ornamentation type is found where epidermal cells are surrounded by sunken walls. Rugose and ruminate types are connected with unclear type (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). There is no connection between concave or convex periclinal cell walls and surface ornamentation type. In the ruminate and rugose forms are seen only unclear periclinal cells. Also, the shape of epidermal cells on the seed surface has exhibited variety. It is polygonal, alveolar and unclear structures. The most common cell type are unclear and alveolar, as polygonal is fairly rare (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe anatomical characters of the seeds are shown in Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e and Table\u0026nbsp;3. Testa structures of the studied seeds are usually consisted of 1 layer, the epidermis, formed in the sclerenchymatous tissue. The epidermis layer demonstrates significant differences in cell form, comprising of crushed, flat or polygonal cells, in 1\u0026ndash;7 layers, and has surged or straight wall form. The most common kind are flat and crushed while the fewest one is the polygonal type (Table\u0026nbsp;3, Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Also, the subepidermis layer is found in some of the studied taxa (\u003cem\u003eH. glabrescens\u003c/em\u003e, \u003cem\u003eH. heldreichii\u003c/em\u003e and \u003cem\u003eH. siirtensis\u003c/em\u003e). The subepidermis layer comprises of flat or polygonal cells in 1\u0026ndash;2 layers. The most frequently found type is flat, however; the rarest one is polygonal type. The thickness of testa layers ranges from 15.77 \u0026micro;m (in \u003cem\u003eH. micrantha\u003c/em\u003e) to 49.76 \u0026micro;m (in \u003cem\u003eH. campanulata\u003c/em\u003e). The cotyledon sizes vary between 1652.47 \u0026micro;m (in \u003cem\u003eH. glabrescens\u003c/em\u003e) and 1913.63 \u0026micro;m (in \u003cem\u003eH. campanulata\u003c/em\u003e) in length, between 856.44 \u0026micro;m (in \u003cem\u003eH. micrantha\u003c/em\u003e) and 1605.22 \u0026micro;m (in \u003cem\u003eH. campanulata\u003c/em\u003e) in width. In addition to these findings, embryo sizes of the seeds range from 245.17 \u0026micro;m (in \u003cem\u003eH. hispida\u003c/em\u003e) to 940.25 \u0026micro;m (in \u003cem\u003eH. glabrescens\u003c/em\u003e) in length, from 152.10 \u0026micro;m (in \u003cem\u003eH. heldreichii\u003c/em\u003e) to 368.49 \u0026micro;m (in \u003cem\u003eH. nervosa\u003c/em\u003e) in width, and embryos in seeds are formed elongated (\u003cem\u003eH. acutiloba\u003c/em\u003e, \u003cem\u003eH. glabrescens\u003c/em\u003e, \u003cem\u003eH. lazulina\u003c/em\u003e, \u003cem\u003eH. lineata\u003c/em\u003e and \u003cem\u003eH. siirtensis\u003c/em\u003e), elliptical (\u003cem\u003eH. campanulata\u003c/em\u003e, \u003cem\u003eH. hispida\u003c/em\u003e and \u003cem\u003eH. nervosa\u003c/em\u003e), oval (\u003cem\u003eH. heldreichii\u003c/em\u003e and \u003cem\u003eH. micrantha\u003c/em\u003e) and circular (\u003cem\u003eH. venusta\u003c/em\u003e) in shapes (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe numerical assessment of the seed morpho-anatomical characters permits the formation of a dendrogram, which reveals the differences-similarities among the studied species. A dendrogram is built as a consequence of the cluster analysis of the studied taxa of \u003cem\u003eHyacinthella\u003c/em\u003e based on the variation of 42 characteristics in 11 species. The cophenetic correlation coefficient is determined to learn the relative between the dendrogram and dissimilarity matrix (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e4\u003c/span\u003e, Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The cophenetic correlation between the dissimilarity matrix and dendrogram has been computed as 0.64, representing a good match.\u003c/p\u003e \u003cp\u003eOur cluster analysis has separated the taxa into 2 main groups of A and B: Cluster A comprises \u003cem\u003eH. glabrescens\u003c/em\u003e, \u003cem\u003eH. siirtensis\u003c/em\u003e, \u003cem\u003eH. lineata\u003c/em\u003e and \u003cem\u003eH. acutiloba\u003c/em\u003e. Cluster B1 includes \u003cem\u003eH. hispida\u003c/em\u003e, \u003cem\u003eH. heldreichii\u003c/em\u003e, \u003cem\u003eH. nervosa\u003c/em\u003e and \u003cem\u003eH. campanulata\u003c/em\u003e. Cluster B2 includes \u003cem\u003eH. venusta\u003c/em\u003e and \u003cem\u003eH. lazulina. H. micrantha\u003c/em\u003e has produced a clade outside these clusters in the dendrogram (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e4\u003c/span\u003e). \u003cem\u003eH. siirtensis\u003c/em\u003e and \u003cem\u003eH. lineata\u003c/em\u003e are the most closely related species (dissimilarity coefficient: 160.999), as \u003cem\u003eH. micrantha\u003c/em\u003e and \u003cem\u003eH. campanulata\u003c/em\u003e are the most distantly related species (dissimilarity coefficient: 801.702) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Cluster B has the highest number of species compared to another cluster (Fig.\u0026nbsp;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e5\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe morphological structures of the seeds present significant data in relation to evolutionary relations of the flowering plants (Corner \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1976\u003c/span\u003e; Karaismailoğlu and Erol \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). However, seed morpho-anatomical characters have so far not been comprehensively applied to clarify inter-species relations within genera of the family Asparagaceae, except the genus \u003cem\u003eMuscari\u003c/em\u003e (Eroğlu et al \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This is the first research to exhibit the morpho-anatomical structures of the seeds of the genus \u003cem\u003eHyacinthella\u003c/em\u003e, and it will be a model for following surveys for various closely related genera.\u003c/p\u003e \u003cp\u003eThe macro-morphological characteristics of the seeds demonstrate differences among the studied \u003cem\u003eHyacinthella\u003c/em\u003e species (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e and Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The species studied in our research are not very distinct with regard to seed color. Black dominates in the genus, whereas \u003cem\u003eH. micrantha\u003c/em\u003e has brown seeds unlike other species. \u003cem\u003eH. glabrescens\u003c/em\u003e (ellipticus)-\u003cem\u003eH. hispida\u003c/em\u003e (triangularis), \u003cem\u003eH. campanulata\u003c/em\u003e (circularis)-\u003cem\u003eH. heldreichii\u003c/em\u003e (ovatus), \u003cem\u003eH. acutiloba\u003c/em\u003e (triangularis)-\u003cem\u003eH. lineata\u003c/em\u003e (ovatus) and \u003cem\u003eH. siirtensis\u003c/em\u003e (triangularis)-\u003cem\u003eH. nervosa\u003c/em\u003e (ellipticus-late) taxa are more or less similar in terms of leaf, scape and flower structures, but they can be simply distinguished with applying of seed shape.\u003c/p\u003e \u003cp\u003eThe researches including surface micro-morphological structures of seeds in various plant families provide systematically useful information (Karaismailoğlu \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Candan et al \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Karaismailoğlu and Erol \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Karaismailoğlu et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Karaismailoğlu and G\u0026uuml;ner \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Eroğlu et al \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Şirin and Karaismailoğlu \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Gavrilović and Janaćković \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Also, the importance and effectiveness of scanning electron microscopy in clarifying of taxonomic difficulties and in characterizing of species have been highlighted by many researchers (Heywood \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1971\u003c/span\u003e; Barthlott \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1981\u003c/span\u003e; Eroğlu et al \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). However, there are few papers on the significance of seed micromorphology in the family Asparagaceae (Yıldırım \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Eroğlu et al \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This research on 11 \u003cem\u003eHyacinthella\u003c/em\u003e species reveals that seed micro-structures are helpful characters in discrimination of the taxa within the family, as in the genus \u003cem\u003eMuscari\u003c/em\u003e (Eroğlu et al \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). All of the examined species have been analyzed for the first time. We noted seven seed surface ornamentation types in this work. In the genus, the most common seed ornamentation type is verrucate. Unlike this work, reticulate or reticulate-areolate ornamentation forms have been frequently found among species from many angiosperm families (Barthlott \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1981\u003c/span\u003e; Erol et al \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Bona \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Karaismailoğlu \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; \u0026Ouml;zbek et al \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Karaismailoğlu et al \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Karaismailoğlu and Erol, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The rugose (in \u003cem\u003eH. acutiloba\u003c/em\u003e), reticulate-pusticulate (in \u003cem\u003eH. campanulata\u003c/em\u003e), ruminate (in \u003cem\u003eH. hispida\u003c/em\u003e) and reticulate-foveate (in \u003cem\u003eH. lineata\u003c/em\u003e) ornamentation types are taxon specific. Also, ornamentation types are effective in distinguishing closely related taxa from each other, such as \u003cem\u003eH. glabrescens\u003c/em\u003e (verrucate)-\u003cem\u003eH. hispida\u003c/em\u003e (ruminate), \u003cem\u003eH. campanulata\u003c/em\u003e (reticulate-pusticulate)-\u003cem\u003eH. heldreichii\u003c/em\u003e (tuberculate), \u003cem\u003eH. acutiloba\u003c/em\u003e (rugose)-\u003cem\u003eH. lineata\u003c/em\u003e (reticulate-foveate) and \u003cem\u003eH. siirtensis\u003c/em\u003e (alveolate)-\u003cem\u003eH. nervosa\u003c/em\u003e (verrucate) taxa (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003e and Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Besides, past seed surface surveys have shown that the views and forms of anticlinal and periclinal cell walls are diagnostic characteristics in the creation of inter-taxa interactions (Barthlott \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1981\u003c/span\u003e; Eroğlu et al \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The types of anticlinal and periclinal cell walls, and epidermal cell structures of the studies species reveal variations among taxa. The macro and micro-morphological outcomes of this investigation are separated all taxa examined from each other, and they are suitable with the former studies done with exomorphic features of seeds of the family Asparagaceae (Yıldırım \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Eroğlu et al \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Tugay et al \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTesta anatomical studies are effective in answering of the taxonomical problems in many plant families (Vaughan et al \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1976\u003c/span\u003e; Manning and Staden \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Meyer \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Karaismailoğlu \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Karaismailoğlu and Erol \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Eroğlu et al \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Koul et al. (\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2000\u003c/span\u003e) and Eroğlu et al. (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) have stated that testa forms can be used as a useful characters in the discrimination of the species and in the explanation of their phylogenetic relations. This study is the first such research for the genus and is the precursor to following works. In this study, it is observed that the testa layers usually contain one layer as the epidermis in the sclerotic form. The epidermis structures display variations among the species. This 1\u0026ndash;7 layered epidermis can compose of flat, crushed, or polygonal cells. The most common form is crushed, as the rarest is polygonal type. Some of the studied species, which are \u003cem\u003eH. glabrescens\u003c/em\u003e, \u003cem\u003eH. heldreichii\u003c/em\u003e and \u003cem\u003eH. nervosa\u003c/em\u003e, have the subepidermis layer under epidermis occurring polygonal or flat cells. The presence of the subepidermis layer is sufficient to distinguish some closely related taxa such as \u003cem\u003eH. glabrescens\u003c/em\u003e-\u003cem\u003eH. hispida\u003c/em\u003e, \u003cem\u003eH. heldreichii\u003c/em\u003e- \u003cem\u003eH. campanulata\u003c/em\u003e and \u003cem\u003eH. nervosa\u003c/em\u003e- \u003cem\u003eH. siirtensis\u003c/em\u003e. Also, the differences in testa thicknesses and cotyledon sizes of the seeds of Turkish \u003cem\u003eHyacinthella\u003c/em\u003e have been detected. The most distant species are \u003cem\u003eH. micrantha\u003c/em\u003e and \u003cem\u003eH. campanulata\u003c/em\u003e. The embryos of \u003cem\u003eHyacinthella\u003c/em\u003e taxa display a broad variation in terms of shapes and sizes. Embryos are elongated form in \u003cem\u003eH. acutiloba\u003c/em\u003e, \u003cem\u003eH. glabrescens\u003c/em\u003e, \u003cem\u003eH. lazulina\u003c/em\u003e, \u003cem\u003eH. lineata\u003c/em\u003e and \u003cem\u003eH. siirtensis\u003c/em\u003e, elliptical form in \u003cem\u003eH. campanulata\u003c/em\u003e, \u003cem\u003eH. hispida\u003c/em\u003e and \u003cem\u003eH. nervosa\u003c/em\u003e, oval form in \u003cem\u003eH. heldreichii\u003c/em\u003e and \u003cem\u003eH. micrantha\u003c/em\u003e, circular form in \u003cem\u003eH. venusta\u003c/em\u003e. Closely related taxa such as \u003cem\u003eH. glabrescens\u003c/em\u003e-\u003cem\u003eH. hispida\u003c/em\u003e, \u003cem\u003eH. campanulata\u003c/em\u003e-\u003cem\u003eH. heldreichii\u003c/em\u003e, \u003cem\u003eH. siirtensis\u003c/em\u003e- \u003cem\u003eH. nervosa\u003c/em\u003e that are morphologically similar can easily be distinguished from each other according to their embryo shape (Table\u0026nbsp;3 and Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn this study, the seed anatomical characters such as the structures of the epidermis and subepidermis, thickness of the testa, cotyledon sizes and embryo shapes are rather efficient and useful in separating almost all of the studied \u003cem\u003eHyacinthella\u003c/em\u003e taxa. This can be explained as follows: the seed anatomical characters are useful additional characteristics in the \u003cem\u003eHyacinthella\u003c/em\u003e, and they are able to assist in the taxonomy of this genus. The obtained outcomes are compatible with parallel earlier works conducted on seed structure of some taxa of the genera \u003cem\u003eCrocus\u003c/em\u003e L. and \u003cem\u003eRomulea\u003c/em\u003e Maratti in the closely related family Iridaceae, and genus \u003cem\u003eMuscari\u003c/em\u003e in the family Asparagaceae (Grilli Caiola et al \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Karaismailoğlu \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Karaismailoğlu et al \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Eroğlu et al \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eA dendrogram has been created to assess the morpho-anatomical characteristics of the seeds of Turkish \u003cem\u003eHyacinthella\u003c/em\u003e taxa with UPGMA cluster analysis (Fig.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The dendrogram, displaying two main clusters, is partly consistent with the results of Persson and Wendelbo (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1984\u003c/span\u003e). According to the descriptions in the Flora of Turkey, the systematic affinity in the closely related \u003cem\u003eH. campanulata\u003c/em\u003e-\u003cem\u003eH. heldreichii\u003c/em\u003e, and \u003cem\u003eH. acutiloba\u003c/em\u003e-\u003cem\u003eH. lineata\u003c/em\u003e taxa is partially preserved. However, \u003cem\u003eH. siirtensis\u003c/em\u003e-\u003cem\u003eH. nervosa\u003c/em\u003e and \u003cem\u003eH. glabrescens\u003c/em\u003e-\u003cem\u003eH. hispida\u003c/em\u003e taxa are located in different clusters in the dendrogram (Figs.\u0026nbsp;\u003cspan refid=\"Fig7\" class=\"InternalRef\"\u003e4\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig8\" class=\"InternalRef\"\u003e5\u003c/span\u003e and Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e4\u003c/span\u003e). In addition, \u003cem\u003eH. micrantha\u003c/em\u003e has been described as an isolated species with a very different description because of its unique leaf and veining properties within the genus in the Flora of Turkey. This species has showed similar characteristics in the dendrogram created according to seed morphological and anatomical features, and formed a clade apart from 2 main clusters.\u003c/p\u003e \u003cp\u003eAs a result, this work display that morphological and anatomical seed characteristics of Turkish \u003cem\u003eHyacinthella\u003c/em\u003e species presents significant and consistent insights into the taxonomy of species within the genus.\u003c/p\u003e \u003cp\u003e\u003cstrong\u003eKey to studied \u003cem\u003eHyacinthella\u003c/em\u003e taxa, based on seed characters\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e1.Seed colour is brown\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;...\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u003cem\u003e H. micrantha\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e1.Seed colour is black\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;.2\u003c/p\u003e\n\u003cp\u003e2.Seed shape is circularis, ellipticus or ellipticus-late\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;....3\u003c/p\u003e\n\u003cp\u003e2.Seed shape is triangularis or ovatus\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;.6\u003c/p\u003e\n\u003cp\u003e3.Seed shape is circularis\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;..\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;.\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;...\u0026hellip;.. \u003cem\u003eH. campanulate\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e3.Seed shape is ellipticus or ellipticus-late..\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;...4\u003c/p\u003e\n\u003cp\u003e4.Ellipticus\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;...\u0026hellip; \u003cem\u003eH. glabrescens\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e4.Ellipticus-late\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;...5\u003c/p\u003e\n\u003cp\u003e5.Seed surface ornamentation is verrucate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;. \u003cem\u003eH. lazuline\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e5.Seed surface ornamentation is tuberculate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;.\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;.. \u003cem\u003eH. venusta\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e6.Seed shape is triangularis\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;..\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;..7\u003c/p\u003e\n\u003cp\u003e6.Seed shape is ovatus\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;.\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;9\u003c/p\u003e\n\u003cp\u003e7.Seed surface ornamentation is alveolate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip; \u003cem\u003eH. siirtensis\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e7.Seed surface ornamentation is rugose or ruminate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;..8\u003c/p\u003e\n\u003cp\u003e8.Rugose\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;... \u003cem\u003eH. acutiloba\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e8.Ruminate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;.\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;. \u003cem\u003eH. hispida\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e9.Seed surface ornamentation is tuberculate or verrucate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;.10\u003c/p\u003e\n\u003cp\u003e9.Seed surface ornamentation is alveolate or reticulate-foveate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;...11\u003c/p\u003e\n\u003cp\u003e10.Tuberculate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;...\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip; \u003cem\u003eH. heldreichii\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e10.Verrucate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;...\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip; \u003cem\u003eH. nervosa\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e11.Alveolate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip; \u003cem\u003eH. micrantha\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e11.Reticulate-foveate\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;...\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip;\u0026hellip; \u003cem\u003eH. lineata\u003c/em\u003e\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors thank Haydar \u0026Ouml;ztaş, Barış Bani, and H\u0026uuml;seyin G\u0026uuml;ldal for their assistance and contributions in the study.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eContributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eM.C. Karaismailoğlu and H. Eroğlu contributed to the study conception and design. Material preparation, data collection and analysis were performed by M.C. Karaismailoğlu, S.M. Pınar, H. Eroğlu, and M. Fidan. The first draft of the manuscript was written by M.C. Karaismailoğlu, and H. Eroğlu and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNo financial support was used for this study.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eOn behalf of all authors, the corresponding author states no conflict of interest.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting Interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors have no competing interests\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData Availability Statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets generated during and/or analysed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAPG (2009) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161(2): 105\u0026ndash;121.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBarthlott W (1981) Epidermal and seed surface characters of plants: Systematic applicability and some evolutionary aspects. Nordic Journal of Botany 1: 345\u0026ndash;355.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBona M (2013) Seed-coat microsculpturing of Turkish \u003cem\u003eLepidium\u003c/em\u003e (Brassicaceae) and its systematic application. Turkish Journal of Botany 37: 662\u0026ndash;668.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCandan F, Uysal T, Tugay O, Bozkurt M, Ertuğrul K, Demirelma H (2016) The examinations of achene ultrastructural features of section \u003cem\u003eAcrolophus\u003c/em\u003e (\u003cem\u003eCentaurea\u003c/em\u003e, Asteraceae) via scanning electron microscopy. 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London. pp.\u0026nbsp;119\u0026ndash;144.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYetişen K, \u0026Ouml;zdemir C, K\u0026uuml;\u0026ccedil;\u0026uuml;k\u0026ouml;d\u0026uuml;k M, Akyol Y (2012) A morphological and anatomical study of \u003cem\u003eHyacinthella glabrescens\u003c/em\u003e (Liliaceae). Phytologia Balcanica 18(3): 319\u0026ndash;322.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYıldırım H (2015) \u003cem\u003eMuscari atillae\u003c/em\u003e (Asparagaceae): a new species from Eastern Anatolia, Turkey. Phytotaxa 213: 291\u0026ndash;295.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Anatomy, Diagnostic key, Seed, Taxonomic study, Testa, Turkey","lastPublishedDoi":"10.21203/rs.3.rs-1635226/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-1635226/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThis paper offers the first taxonomic assessment of the morphological and anatomical characters of the seeds of 11 \u003cem\u003eHyacinthella\u003c/em\u003e taxa from Turkey, 10 of them are endemic, with the cluster analysis and principal component analysis. Macro-morphologically, the outcomes display that the species are different from each other as per seed shape and size. The studied seeds are separated into 5 shapes; triangularis, circularis, ellipticus, ellipticus-late and ovatus. Seed sizes range from 1.51 mm to 3.06 mm in length, from 1.06 mm to 2.04 mm in width. Micro-morphologically, seed surface ornamentation is grouped into 7 types: rugose, reticulate-pusticulate, verrucate, tuberculate, ruminate, reticulate-foveate and alveolate. The most common type is verrucate, as rugose (in \u003cem\u003eH. acutiloba\u003c/em\u003e), reticulate-pusticulate (in \u003cem\u003eH. campanulata\u003c/em\u003e), ruminate (in \u003cem\u003eH. hispida\u003c/em\u003e) and reticulate-foveate (in \u003cem\u003eH. lineata\u003c/em\u003e) ornamentation types are taxon specific. Anatomically, testa structures of the studied seeds are usually consisted of 1 layer, the epidermis, formed in the sclerenchymatous tissue. The epidermis layer demonstrates significant differences in cell form, comprising of crushed, flat or polygonal cells, in 1\u0026ndash;7 layers, and has surged or straight wall form. Also, the subepidermis layer is found in some of the studied taxa (\u003cem\u003eH. glabrescens\u003c/em\u003e, \u003cem\u003eH. heldreichii\u003c/em\u003e and \u003cem\u003eH. siirtensis\u003c/em\u003e). The thickness of testa layers varies between 15.77 \u0026micro;m (in \u003cem\u003eH. micrantha\u003c/em\u003e) and 49.76 \u0026micro;m (in \u003cem\u003eH. campanulata\u003c/em\u003e). In the systematics of the genus \u003cem\u003eHyacinthella\u003c/em\u003e, the seed morphological and anatomical characters are very important characteristics that reveal inter-specific relationships among the studied species. Also, a dichotomous key is provided for the identification of the examined species based on seed characters.\u003c/p\u003e","manuscriptTitle":"Seed Morphological and Anatomical Structures as Taxonomy Tool for Turkish Hyacinthella Schur (Asparagaceae) Taxa","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2022-05-17 18:41:10","doi":"10.21203/rs.3.rs-1635226/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"e154c57c-b7cb-4652-99dd-a9fcda742fe0","owner":[],"postedDate":"May 17th, 2022","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2022-05-24T13:14:19+00:00","versionOfRecord":[],"versionCreatedAt":"2022-05-17 18:41:10","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-1635226","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-1635226","identity":"rs-1635226","version":["v1"]},"buildId":"_2-kVJe1T_tPrBINL-cwx","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}