New taxa of Onosma L. (Boraginaceae, Lithospermeae) from Iran

2025  47  16 adansonia Adansonia est une revue en flux continu publiée par les Publications scientifiques du Muséum, Paris Adansonia is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Geodiversitas, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-8571/ ISSN (électronique / electronic) : 1639-4798 Directeur De la publication / Publication director: Gilles Bloch Président du Muséum national d’Histoire naturelle réDacteur en chef / editor-in-chief : Thierry Deroin réDacteurs / editors : Porter P. 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Tohmé (Conseil national de la Recherche scientifique Liban, Beyrouth) J. G. West (Australian National Herbarium, Canberra) J. R. Wood (Oxford) couverture / cover : Réalisée à partir des Figures de l’article/Made from the Figures of the article. Adansonia est indexé dans / Adansonia is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Agriculture, Biology, and Environmental Sciences® – Scopus® Adansonia est distribué en version électronique par / Adansonia is distributed electronically by: – BioOne® (http://www.bioone.org) 299 ADANSONIA, sér. 3 • 2025 • 47 (16) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.adansonia.com KEY WORDS Asterotricha, endemic species, morphology, Flora of Iran, new species. Attar F., Mirtadzadini M., Binzet R. & Sotoodeh A. 2025. — New taxa of Onosma L. (Boraginaceae, Lithospermeae) from Iran. Adansonia, sér. 3, 47 (16): 299-310. https://doi.org/10.5252/adansonia2025v47a16. http://adansonia.com/47/16 ABSTRACT This study describes three new species within the genus Onosma L. (Boraginaceae), from sect. Onosma, subsect. Asterotricha (Boiss.) Gürke, in Iran. These species, determined as O. ligniclada Attar& Mirtadzadini, sp. nov., O. elymaitica Attar& Mirtadz., sp. nov., and O. farsistanica Attar& Mirtadz., sp. nov., were distinguished based on detailed morphological and phylogenetic evidence. Onosma ligniclada Attar& Mirtadzadini, sp. nov. and O. elymaitica Attar& Mirtadz., sp. nov., originating from Ilam province, display distinct morphological features that differ from their closest known species, O. albo-rosea Fisch.& C.A.Mey., O. iransharii Ghahr.& Attar and O. bilabiata Boiss.& Buhse respectively. This differentiation is marked by variations in habit, corolla color and size, and leaf morphology on sterile shoots and indumentum of leaves. In contrast, O. farsistanica Attar& Mirtadz., sp.nov. identified in the Quercus L. forests of Fars province (Quercus brantii var. Persica (Jaub.& Spach) Zohary), exhibits notable distinctions from their closely related species, O. bilabiata, particularly in terms of habitat, growth form, tubercle characteristics, calyx and corolla dimensions. Additionally, the manuscript provides photographic documentation, a distribution map, IUCN conservation assessments, and a diagnostic key for these newly identified species. Farideh ATTAR Central Herbarium of Tehran University, School of Biology, College of Science, University of Tehran (Iran) [email protected] (corresponding author) Mansour MIRTADZADINI Department of Biology, Faculty of Sciences, Shahid Bahonar University of Kerman, Kerman (Iran) [email protected] RızaBINZET Department of Biology, Faculty of Arts and Science, Mersin University, 33343, Mersin (Turkey) [email protected] Arash SOTOODEH Université Bourgogne Europe, Institut Agro Dijon, INRAE, Agroécologie, Dijon (France) [email protected] (corresponding author) Submitted on 15 February 2025 | accepted on 8 March 2025 | published on 20 October 2025 New taxa of Onosma L. (Boraginaceae, Lithospermeae) from Iran 300 ADANSONIA, sér. 3 • 2025 • 47 (16) Attar F. et al. INTRODUCTION Onosma L. (Linnaeus 1762) (Lithospermae, Boraginaceae) is a diverse genus comprising approximately 250 taxa worldwide (Chacón etal. 2016; Naqinezhad& Attar 2016; Attar etal. 2023; Advay etal. 2023). Its distribution spans from northwest Africa to Europe and Asia, with a notable prevalence in Turkey and Iran (Kadereit etal. 2016; Chacón etal. 2016; Cecchi etal. 2016; Binzet& Eren 2018; Attar etal. 2020; Firat& Binzet 2021; Noroozi etal. 2022, 2024). The morphology of Onosma species is varied, yet they commonly exhibit characteristics such as being herbaceous perennials or biennials, with some species manifesting as annuals. Typically, these plants have erect stems ranging from 10 to 100cm in height, depending on the species. The leaves are alternately arranged along the stem, often possessing a rough texture due to their setose nature. A defining feature of Onosma species is their funnel-shaped, cylindrical or clavate flowers, comprising five fused petals that form a tube with five triangular or rounded lobes. The flower color varies, commonly presenting in shades of blue, purple, pink, or yellow. These flowers are arranged in cymes, either terminal or axillary, with five epipetalous stamens. The filament length varies among species; bases can be either glabrous or hairy (sect. Protonosma Popov). The nectariferous annulus may also be glabrous or hairy. Nutlets in these species are generally smooth, rugose, or mamillate. The Flora Iranica mentions ten species belonging to subsect. Asterotricha from Iran, including O. albo-rosea Fisch.& C.A.Mey. (with four varieties), O. armena DC., O.bilabiata Boiss.& Buhse, O. caerulescens Boiss., O. cardiostegia Bornm., O. dasytricha Boiss., O. hebebulba DC., O. kurdica Teppner, O.latifolia Boiss.& Hausskn. ex Boiss., and O.rascheyana Boiss. Teppner (1980) also described O. kurdica from Mt Hamzeharab in Kurdistan province. Subsequently, two new species, O.iranshahrii Ghahr.& Attar (Ghahreman& Attar 1996) and O. bisotunensis Attar& Hamzeh’ee (Attar& Hamzeh’ee 2007), and also O.safaie-fari Advay, Attar& S.A.Ahmad (Advay etal. 2023), O.shehbazii Advay, Attar& S.A.Ahmad (Advay etal. 2023) were described from the west of Iran, along with Onosma mozaffarianii Mehrabian (Mehrabian et al. 2013) from Kermanshah, O. targevarensis Mozaff.& Mehrabian (Mehrabian & Mozaffarian 2018) from Azerbaijan province and O. wendelboi Mehrabian& Mozaff. (Mehrabian & Mozaffarian 2018) from Lorestan Province, O. madjnoniani Mehrabian& Naghizadeh (Mehrabian et al. 2022), O. pseudohebebulba Mozaff.& Mehrabian (Mehrabian et al. 2022), O. atrii Mehrabian& Khajoei Nasab (Mehrabian etal. 2022), O.persica Mehrabian& Naghizadeh (Mehrabian et al. 2022), O.bozghushica Mozaff.& Mehrabian (Mehrabian etal. 2022). All species in subsect. Asterotricha have white, yellow, pink, and red corolla that become dark blue when dry in most species. This is an important characteristic for distinguishing between species. According to Riedl (1967, 1979) and Khatamsaz (2002), the center of diversity for subsect. Asterotricha is in the west of Iran and Turkey. With the recent additions of new taxa, the number of species belonging to subsect. Asterotricha in the Flora of Iran has increased to 30. During several field works and herbarium studies conducted by the authors of this study for a taxonomic treatment of Onosma, they discovered some specimens that were previously unknown to science. MATERIAL AND METHODS New species Three specimens were collected from different populations in Ilam and Fars provinces in Iran (Fig. 1 and Table 1), which were identified as new species. To determine the taxonomic status of the collected specimens, relevant literature including Parsa (1949), Riedl (1967, 1979), Popov (1974), and MOTS CLÉS Asterotricha, espèces endémiques, morphologie, flore d’Iran, espèces nouvelles. RÉSUMÉ Nouveaux taxons d’Onosma L. (Boraginaceae, Lithospermeae) d’Iran. Cette étude décrit trois nouvelles espèces du genre Onosma L. (Boraginaceae), appartenant à la section Onosma L., sous-section Asterotricha (Boiss.) Gürke, en Iran. Ces espèces, identifiées comme O. ligniclada Attar& Mirtadzadini, sp. nov., O. elymaitica Attar& Mirtadzadini, sp. nov. et O. farsistanica Attar& Mirtadzadini, sp. nov., ont été distinguées sur la base de preuves morphologiques et phylogénétiques détaillées. Onosma ligniclada Attar& Mirtadzadini, sp. nov. et O. elymaitica Attar& Mirtadz., sp. nov., originaires de la province d’Ilam, présentent des caractéristiques morphologiques distinctes de leurs espèces les plus proches connues, à savoir O. albo-rosea Fisch.& C.A.Mey., O. iransharii Ghahr.& Attar et O. bilabiata Boiss.& Buhse respectivement. Cette différenciation repose sur des variations concernant l’habitus, la couleur et la taille de la corolle, ainsi que la morphologie foliaire des pousses stériles et l’indument des feuilles. En revanche, O. farsistanica Attar& Mirtadz., sp. nov., identifié dans les forêts de Quercus L. de la province de Fars (Quercus brantii var. persica (Jaub.& Spach) Zohary), se distingue nettement de son espèce la plus apparentée, O. bilabiata, notamment en termes d’habitat, de port, de caractéristiques des tubercules, ainsi que de dimensions du calice et de la corolle. En outre, cet article fournit une documentation photographique, une carte de distribution, des évaluations de conservation selon les critères de l’UICN et une clé diagnostique pour ces nouvelles espèces. 301 Three new Onosma L.L. from the flora of Iran ADANSONIA, sér. 3 • 2025 • 47 (16) Khatamsaz (2002), as well as type photos (K, TARI and W Herbaria) of closely related species, were consulted. Field photographs were taken of all collected species. The indumentum and nutlet were photographed using a Dino-Lite digital microscope AM413T. The holotypes, isotypes, and paratypes were deposited in IRAN, MIR, and TUH (Attar Collection) herbaria (Thiers 2017). Moreover, to examine specific morphological features, meticulous observations were conducted utilizing a scanning electron microscope (SEM). RESULTS AND DISCUSSION Morphology Our investigations of several Onosma specimens from herbaria and field collections have revealed three previously undescribed species within the genus Onosma, specifically in the section Onosma and the subsection Asterotricha. These species are identified as O. ligniclada sp. nov., O. elymaitica sp. nov., and O. farsistanica sp. nov. Detailed morphological comparisons showed that these new species exhibit distinct differences from their closest relatives. Onosma ligniclada sp. nov. and O. elymaitica sp. nov., both originating from Ilam province, display unique morphological features that distinguish them from their closest known species, O. albo-rosea, O. iransharii, and O. bilabiata. These differences include variations in habit, corolla color and size, and leaf morphology on sterile shoots, as well as the indumentum of leaves. In contrast, O. farsistanica sp. nov., identified in the Quercus forests of Fars province (Quercus brantii var. persica), shows notable distinctions from the closely related O. bilabiata. These distinctions are evident in habitat preference, growth form, tubercle characteristics, and calyx and corolla dimensions. Family BoragiNaceae Juss. Tribe Lithospermae (DC.) Gürke Onosma sect. Onosma subsect. Asterotricha (Boiss.) Gürke DiagNosis. — The basal leaves of these species have stellate hairy setae tubercles, consisting of numerous oblong or rounded cells arranged in one, two, or three series. The number and length of hairs on the tubercles vary widely in length and density and may exhibit regular or irregular patterns. These characters, along with other vegetative and reproductive traits, are highly important in species delimitation. SAUDI ARABIA IRAQ SYRIA TURKEY 40°E 40°S 30° 25° 35° 45° 50° 55° 60° 65° 70° IRAN AFGHANISTAN PAKISTAN TURKMENISTAN Caspian Sea Persian Gulf O. farsistanica sp. nov. O. elymaitica sp. nov. O. bilabiata Boiss. & Buhse O. latifolia Boiss. & Hausskn. ex Boiss. O. ligniclada sp. nov. fig. 1. — Distribution map of three new species, O. ligniclada Attar & Mirtadzadini, sp. nov., O. elymaitica Attar & Mirtadzadini, sp. nov. et O. farsistanica Attar & Mirtadz., sp. nov., and their related species. table 1. — Localities of identified Onosma L. species. Species Locality O. ligniclada sp. nov. Iran, W, Kermanshah, c. 11 km to the crossroad of Gilan-e Gharb and Sumar, alt. 1050 m, 34°16’30”N, 46°32’12”E O. elymaitica sp. nov. Iran, W, Ilam, c. 15 km to Badreh from Ilam, alt. 844 m, 33°36’58”N, 46°09’6.83”E O. farsistanica sp. nov. Iran SW, Fars, between Shiraz and Kazerun, Dasht-e Arjan, 29°39’41.33”N, 51°58’59.36”E 302 ADANSONIA, sér. 3 • 2025 • 47 (16) Attar F. et al. Onosma ligniclada Attar& Mirtadzadini, sp. nov. (Figs 2; 3) Type MaTerial. — Iran • W, Kermanshah, c.11km to the crossroad of Gilan-e Gharb and Sumar; 1050m alt.; 34°16’30”N, 46°32’12”E; 28.V.2016; Attar, Mirtadzadini& Raie Niaki 46689; holotype: TUH!; isotype: MIR! • same location; 2.V.2019; Attar& Habibi 48901; paratype: TUH!. DiagNosis. — Differt ab Onosma albo-rosea Fisch.& C. A. Mey.: Caules usque 50cm (nec 10-20cm); folia basalia late elliptica, 20-45× 7-17mm (nec obovato-spathulato lanceolata, 20-60× 6-12[-15]mm); pedicellus 2-4mm longus (nec 1-3mm longus); corolla alba (nec alba, rosea ); filament ±2.5mm longa (nec 8-9mm longa); nuculae 8mm longae (nec 6-7 [-8] mm longae). recogNiTioN. — Onosma ligniclada sp. nov. is related to Onosma albo-rosea, which belongs to the sect. Onosma subsect. Asterotricha (Riedl 1967; Khatamsaz 2002). Onosma albo-rosea is distributed in Iran, Turkey, Iraq and Syria. These two species have some morphological differences such as the height of the plant, the size and color of the corolla, and size of leaves of sterile shoots. Further differences are outlined in Table 2. DisTriBuTioN, haBiTaT aND pheNology. — Onosma ligniclada sp. nov. grows on the dry hillside steppes, at elevation of 1050m. It is an endemic species of Iran. The species that are associated with this species in the area include: Avena sp., Chardinia orientalis (L.) Kuntze, Echinops kermanshahensis Mozaff., Hirschfeldia incana (L.) Lagrèze, Oliveria decumbens Vent., Onopordum acanthium L., Zoegea sp. Flowering in April to June, fruiting in May to July. fig. 2. — Holotype specimen of Onosma ligniclada Attar & Mirtadz., sp. nov. 303 Three new Onosma L.L. from the flora of Iran ADANSONIA, sér. 3 • 2025 • 47 (16) e TyMology . — The specific epithet is derived from the woody base of the plant, referring to the lignified structure present either at the base of the stem or at the lower parts of the branches. coNservaTioN sTaTus. — Onosma ligniclada sp. nov. is currently known from a single locality with a small population. Considering its very restricted area of occupancy (AOO), occurrence at one location, and an inferred continuing decline in habitat quality, we assess it as Endangered (EN) under criterion B2ab(iii) (IUCN 2012). DescripTioN Perennial, woody at base, up to 50cm high. Stem branched, white or sometimes brownish with sterile shoots at the base, erect, covered with white and ±patent setae. Leaves of sterile shoots broad ovate-elliptic, acute, 10-20× 3-8mm (including a c.5mm long petiole); basal leaves of flowering shoots ovate-elliptic, acute, 20-45× 7-17mm; middle and upper leaves similar to the lower ones, subsessile; indumentum consists of white tubercle-based adpressed dense setae on the upper surface and lax on the lower surface; tubercles stellate hairy, rays long and numerous, regular; midrib prominent, adpressed-setose on the abaxial leaf surface. Petiole 2-4cm long. Cymes few-flowered, 5-8cm long. Bracts linear-lanceolate, 20× 3-4mm. Pedicels 5mm long in flower, to 7mm long in fruit, with spreading setae. Calyx ±20mm AB CD fig. 3. — Onosma ligniclada Attar & Mirtadz., sp. nov.: A, stems and inflorescence; B, nutlet; C, D, upper and lower surface of leaves respectively. Scale bars: B, 2 mm; C, 0.5 mm; D, 0.2 mm. table 2. — Comparing the morphological characters between Onosma ligniclada Attar & Mirtadzadini, sp. nov. and its related O. albo-rosea. Characters O. ligniclada sp. nov. O. albo-rosea Height of plant c. 50 cm 10-20 cm Habit Erect Prostrate Shape of basal leaves of flowering shoot Broad elliptic Obovate, spathulate-lanceolate Pedicel length (mm) 2-4 1-3 Calyx length in flowering and fruiting (mm) ± 20, 30-32 14-16, 25 Corolla color White, becoming blue White, pink, becoming blue Corolla length (mm) 26-29 18-30 Anther, filament length c. 8, ± c. 3 mm 7, 8-9 Filament connection point Middle of corolla Below middle Nutlet length and width (mm) 8 6-7(-8) Nutlet shape and color Trigonal, shortly apiculate, ±smooth, shining white Smooth or indistinctly rugose 304 ADANSONIA, sér. 3 • 2025 • 47 (16) Attar F. et al. long in flower, 3-5mm wide, accrescent, 30-32mm long in fruit, 7-8mm wide in the fruiting state; calyx lobes 5, divided to near the base, united for 5mm, outside covered with sparse white patent setae, tubercles stellate hairy, pubescent between them, inside similar outside but fewer. Corolla cream at first, turning bluish towards upper, 26-29mm long, campanulatecylindrical; lobes triangular, obtuse, 2.5mm wide and 2mm long, pubescent outside, glabrous inside. Annulus 0.5mm wide, glabrous, lobulate, crescent. Anthers sagittate, included, c.8mm long. Filaments c.3mm long, arising from approximately the middle of the corolla tube. Nutlets 8× 7mm, often solitary, cream, smooth, shiny, ovate-three-faced, with dorsal keel, with a middle vein on the adaxial surface, gradually beaked (Figs 2; 8). Onosma elymaitica Attar& Mirtadz., sp. nov. (Figs 4; 5) Type MaTerial.— Iran • W Ilam, c.15km to Badreh from Ilam; 844m alt.; 33°36’58”N, 46°09’6.83”E; 15.V.2015; Attar& Zamani 46138; holotype: TUH! DiagNosis. — Differt ab O. bilabiata Boiss.: folia basalia ellipticoorbicularia, excluso petiolo 15-20× 13-18mm (nec oblanceolatoobovata, petiolo excluso 25-100× 7-25mm); corolla ±15mm longa (nec 25-30mm longa); corolla flava, in sicco coerulea (nec flavoalba et in sicco coerulea); filament ±2.5mm longa (nec 2-3mm longa); nuculae 7mm longae, distincte rugosae (nec 6mm longae, foveolato-punctatae). fig. 4. — Holotype specimen of Onosma elymaitica Attar & Mirtadz., sp. nov. 305 Three new Onosma L.L. from the flora of Iran ADANSONIA, sér. 3 • 2025 • 47 (16) recogNiTioN.— Onosma elymaitica sp. nov. is closely related to O. bilabiata which belongs to the sect. Onosma subsect. Asterotricha (Riedl 1967; Khatamsaz 2002). The new species is distinguished from O. bilabiata by the shape and size of basal leaves, size, and color of the corolla in a dry state and nutlet characters (Table 3). DisTriBuTioN, ecology aND pheNology. — This species is collected from a hillside small area near Ilam. Vegetation is limited to annual grasses and scattered shrubs and bushes. Due to low elevation, summer is too warm and severe for perennial plants in this region. The following species are presented in their habitat: Amygdalus haussknechtii (C.K.Schneider.) Bornm. var. pubescens Bornm., Anchusa strigosa Labill., Bromus tectorum L., B. danthoniae Trin. in C.A.Mey.var. danthoniae, Cerastium inflatum Link ex Desf., C. intricata Boiss. subsp. kermanshahensis Wagenitz, Cephalaria dichaetophora Boiss., Cousinia jaccobsii Rech.f., C. calocephala Jaub.& Spach, Daphne mucronata Royle, Ferulago angulata Schlecht.& Boiss. subsp. angulata, Heteranthelium piliferum (Banks& Soland.) Hochst., Onopordon carduchorum Bornm.& Beauv., Onosma alborosea Fisch.& C.A.Mey. var. albo-rosea, O. albo-rosea Fisch.& C.A.Mey. var. macrocarpa Bornm., Stachys benthamiana Boiss., Quercus brantii Lindl., var. persica (Jaub.& Spach) Zohary, Stipa sp. Flowering April to May, fruiting May to June. eTyMology. — The specific epithet refers to the type locality “Elyma or Elymais” the ancient name of today Ilam [and most parts of Khuzestan] province in the west of Iran. coNservaTioN sTaTus. — Onosma elymaitica sp. nov. is restricted to a very small area near Ilam and is known from a single, small population. Based on its limited AOO, occurrence at one location, and an inferred continuing decline in habitat quality, we assess it as Endangered (EN) under criterion B2ab(iii) (IUCN 2012). DescripTioN Perennial, woody at base, up to 25cm high, with horizontal stolon, with several sterile rosettes at the base. Stem erectascending, covered with white appressed setae. Leaves of sterile shoots broad elliptic, shortly mucronate, blade 15-20× 13-18mm, petiole 6-15cm long; basal leaves of flowering shoots elliptic-orbicular, acute, 15-18mm long, 7-12mm wide, shortly petiolate; middle and upper leaves obovate, remote, subsessile; indumentum yellowish-greyish; upper surface consists of indistinctly tubercle-based appressed very dense setae; lower surface covered by lax tubercled setae; tubercles stellate hairy, rays long and numerous; between setae shortly pubescent. Cymes scorpioid, many-flowered, 3.5-5cm long in fruiting time. Bracts lanceolate, 10× 4mm. Pedicels setose, to 5mm long in fruit. Calyx 10× 3mm, accrescent at fruiting time, 15-17mm long, lobes broad lanceolate, divided to near the base, outside densely setose, tubercles A DE BC fig. 5. — Onosma elymaitica Attar & Mirtadz., sp. nov.: A, B, stems and inflorescence; C, nutlet; D, E, upper and lower surface of leaf respectively. Scale bars: C, 2 mm; D, 0.5 mm; E, 0.2 mm.