ARTÍCULO

Four new species of Onosma (Boraginaceae) from Iran as an important diversity center of the genus in the world

AHMAD REZA MEHRABIAN¹, VALIOLLAH MOZAFFARIAN² & SIAVASH NAGHIZADEH¹

¹Department of Plant Sciences and Biotechnology, Faculty of Life sciences and Biotechnology, Shahid Beheshti University, IR-1983969411 Tehran, Iran
²Department of Plant Botany, Research Institute of Forests and Rangelands, IR-13185116 Tehran, Iran

ORCID iD. A. R. MEHRABIAN: https://orcid.org/0000-0001-6633-3092, V. MOZAFFARIAN: https://orcid.org/0000-00002-7630-9069, S. NAGHIZADEH: https://orcid.org/0000-0003-2284-9796

Author for correspondence: A. R. Mehrabian (a_mehrabian@sbu.ac.ir)

Editor: A. Susanna

ABSTRACT
Four new species of Onosma (Boraginaceae) from Iran as an important diversity center of the genus in the world.— In this study, four new species of Onosma subsect. Asterotricha from Northwest, West and North of Iran are described and illustrated. An extensive review of literature and a comparative study with specimens from several herbaria lead to determine these new species. Our argument for mentioned innovations is the significant divergence in the most important morphological characters. A diagnostic key to the new species as well Asterotrichous members of the genus known from Flora Iranica region is provided. In addition, detailed description and distribution as well as comparison tables with closely related species, are included.
KEY WORDS: Boraginaceae; ecology; geobotany; Irano-Turanian; Lithospermeae; SW Asia.

RESUMEN
Cuatro nuevas especies de Onosma (Boraginaceae) de Irán un importante centro de diversidad del género en el mundo.— En este estudio, se describen e ilustran cuatro nuevas especies de Onosma subsect. Asterotricha del noroeste, oeste y norte de Irán. Estas especies se han descrito tras una revisión exhaustiva de la literatura y un estudio comparativo de especímenes de diversos herbarios. Esta propuesta se basa en la divergencia significativa de los caracteres morfológicos más importantes. Se incluye una clave diagnostica para las nuevas especies y para los miembros de la sect Asterotrichous reconocidos en Flora Iranica. Se incluye también una descripción detallada y la distribución de las especies, así como una tabla comparativa con las especies más estrechamente relacionadas.
PALABRAS CLAVE: Boraginaceae; ecología; geobotánica; Lithospermeae; región Irano-Turaniana; SW Asia.

Received 27 March 2021; accepted 10 February 2022; published on line 13 May 2022

Cómo citar este artículo / Citation

Mehrabian, A. R., Mozaffarian, V. & Naghizadeh, S. 2022. Four new species of Onosma (Boraginaceae) from Iran as an important diversity centre of the genus in the world. Collectanea Botanica 41: e003. https://doi.org/10.3989/collectbot.2022.v41.003

CONTENTS

ABSTRACT

RESUMEN

INTRODUCTION

MATERIALS AND METHODS

RESULTS AND DISCUSSION

ACKNOWLEDGEMENTS

REFERENCES

APPENDIX

INTRODUCTIONTop

The current study aimed to evaluate the most important diagnostic characteristics of unidentified species compared to their related species, justify them as new species, and provide taxonomic as well as ecological descriptions of the new taxa.

MATERIALS AND METHODSTop

During field studies in various regions of the country, some unknown samples were collected. They were also collected by the authors during their scientific excursions in the Azerbaijan, Ilam and Alborz provinces. The new taxa were compared with fundamental diagnostic keys, including Boissier (1875Boissier, P. E. 1875. Flora orientalis, sive enumeratio plantarum in Oriente a Graecia et Aegypto ad Indiae fines hucusque observatarum 4. H. Georg, Geneva, Basilea & Lugdunum.), Flora of USSR (Popov, 1953Popov, M. G. 1953. Boraginaceae. In: Shishkin, B. K. (Ed.), Flora of USSR 19. Keter Publishing House, Jerusalem.), Johnston (1954Johnston, I. M. 1954. Studies in the Boraginaceae XXVI. Further evaluations of the genera of the Lithospermeae. Journal of Arnold Arboretum. 35: 1–8. https://doi.org/10.5962/bhl.part.8314), Post (1966), Riedl (1967Riedl, H. 1967. Boraginaceae In: Rechinger, K. H. (Ed.), Flora Iranica 48. Akademische Druck-u. Verlagsanstalt, Graz: 170–212. ), Polunin (1969Polunin, O. 1969. Flowers of Europe. Oxford University Press, Oxford.), Ball (1972Ball, P. W. 1972. Onosma L. In: Tutin, T. G., Heywood, V. H., Burges, N. A., Moore, D. M., Valentine, D. H., Walters, S. M. & Webb, D. A. (Eds.), Flora Europaea 3. Cambridge University Press, Cambridge: 89–95.), Riedl (1979Riedl, H. 1979. Onosma L. In: Davis, P. H. (Ed.), Flora of Turkey and the east aegean islands 6. Edinburgh University Press, Edinburgh: 326–376.) Nasir & Ali (1979Nasir, E. & Ali, S. I. 1979. Flora of West Pakistan, an annotated catalogue of vascular plants of West Pakistan and Kashmir by RR Stewart. Fakhri Press, Karachi. ), and Khatamsaz (2002Khatamsaz, N. 2002. Boraginaceae L. In: Assadi, M., Massoumi, A. A., Mozaffarian, V. & Jamzad, Z. (Eds.), Flora of Iran. Research Institute of Forests and Rangelands, Tehran.). Additionally, the results of several studies on the newly introduced species of Onosma (Mehrabian & Mozaffarian, 2018Mehrabian, A. R. & Mozaffarian, V. 2018. Seven new species of Onosma L. (Boraginaceae) with emphasis on their habitats in Iran. Taiwania 4: 366–388., Mehrabian et al., 2022Mehrabian, A. R., Naghizadeh, S., Khajoei Nasab, F., Moradi Zeinab, H. & Mozaffarin, V. 2022. Two new species and a new record for Onosma L. (Boraginaceae) from northwestern Iran. Feddes Repertorium 133: 1–21. https://doi.org/10.1002/fedr.202000036) were assessed and compared with the new taxa. More than 2800 herbarium accessions of Onosma, were examined, including type specimens and image types, in several European herbaria (W!, WU!, LE, BG!, JE!, C!, B!, G!, and K) and Iranian herbaria (HSBU, TARI, IRAN) (acronyms follow Thiers, 2019Thiers, B. 2019 [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Retrieved January 15, 2019, from http://sweetgum.nybg.org/science/ih/ ) to reinforce the taxonomic reasons for introducing the mentioned new species (Appendix). Furthermore, morphological characters including trichome type (cauline and basal leaves and nectary ring), shape and size of the calyx, shape and size of the corolla, nutlet morphology, anther length and connection as well as their exertion and the type and size of pollen grains were studied and compared with closely related species.

Holotypes and isotypes were deposited and preserved in the Herbarium of Shahid Beheshti University (HSBU) as well Herbarium of Research Institute of Forests and Rangelands of Iran (TARI) with the collector and acronyms of herbariums in parentheses, according to Index Herbariorum (Thiers, 2019Thiers, B. 2019 [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Retrieved January 15, 2019, from http://sweetgum.nybg.org/science/ih/ ). In addition, the author’s names are included in accordance with IPNI.

The upper and lower surfaces of basal and cauline dried leaves as well as pollen grains were mounted on stubs using double-sided adhesive tape and coated with gold by a DC magnetron sputtering desktop machine. Afterward, the samples were photographed by a scanning electron microscope (Cam Scan Hitachi SU3500). In addition, 15–30 pollen grains were assessed through analysis, and mature dried nutlets were photographed by a dinolite microscope. The morphological terminology follows that of Riedl (1967Riedl, H. 1967. Boraginaceae In: Rechinger, K. H. (Ed.), Flora Iranica 48. Akademische Druck-u. Verlagsanstalt, Graz: 170–212. ), Harris & Harris (2001Harris, J. G. & Harris, M. W. 2001. Plant identification terminology, an illustrated glossary (2nd ed.). Lake Publishing, Spring Lake.), Simpson (2006Simpson, M. 2006. Plant Systematics. Elsevier Academic Press, Burlington, San Diego & London.), Punt et al. (2007Punt, W., Hoen, P. P., Blackmore, S., Nilssont, S. & Thomas, L. 2007. Glossary of pollen and spores terminology. Review of Paleobotany and Palynology 143:1–81. https://doi.org/10.1016/j.revpalbo.2006.06.008), Halbritter et al. (2008Halbritter, H., Weber, M., Zetter, R., Frosch-Radivo, A., Buchner, R. & Hesse, M. 2008. PalDat- illustrated handbook on pollen terminology. Society of the Promotion of Palynological Research in Austria, Vienna.), and Hesse et al. (2009Hesse, M., Halbritter, H., Zette, R., Weber, M., Buchner, R., Frosch-Radivo, R. A. & Ulrich, S. 2009. Pollen Terminology. Springer Verlag, Wien.). Lastly, morphological and micro-morphological evidence was provided to compare new introduced taxa with other asterotrichous and heterotrichous species of Onosma (Tables 1 and 2). A synoptic key to nearest taxa to new taxa was provided for decision making (Appendix).

Table 1. Main characters distinguishing Onosma madjnoniani and Onosma riahii from its morphologically similar species. MXc: Mediterranean xeric-continental, MPc: Mediterranean pluviseasonal continental, Ig: Igneous Rocks, Me: Metamorphic, Vo-Se: Volcano-Sedimentary, Se; Sedimentary, QD: Quaternary Deposits, LS: Lithosols from igneous rocks, brown soils, and Sierozome Soils CL: Calcareous Lithosols-Brown Soils and Chestnut Soils, BS: Brown Soils, Op: Ophiolithic.

Characters	*Onosma madjnoniani*	*Onosma riahii*	*Onosma rascheyana*	*Onosma armena*	*Onosma kurdica*	*Onosma moussavi*
Plant height (cm)	20–25	30–35(40)	12–25	15–30	30–35	22–30
Rosette leaves size (mm)	90–140(150) × 4–9	30–40 × 2–3	20–55 × 3-10	15–35 × 1–3.5	15–22 × 3–4	20–44(45)
Rosette leaves shape	lanceolate to lanceolate-linear	linear to linear-lanceolate	spathulate to linear-spathulate	linear-spathulate	linear, linear-lanceolate	spatulate-lanceolate
Basal leaf size (mm)	25–45 × 4–7	30–40 × 2–3	20–40 × 3–8	20–40 × 2–3.5	100–200 × 3–5	15–25
Basal leaf shape	lanceolate to lanceolate-linear	linear-lnceolate	spathulate to linear-spathulate	linear-spathulate	narrowly obovate, lanceolate	spatulate-lanceolate
Cauline leaf size(mm)	25–90 × 5–9(10)	50–90 × 4–4.5(5.5)	20–40 × 3–8	20–35 × 2–3.5	100–200 × 3–5	30–50 × 2–6
Cauline leaf shape	lanceolate to lanceolate-linear	linear-lnceolate	oblong	spathulate	linear-lanceolate	linear-lanceolate
Indumentum	sub-dense, sub-adpressed	dense, sub-adpressed	sub-dense, sub-adpressed	dense sub-adpressed	dense, adpressed	sub-dense, adpressed
Tubercle rays	short, semi-dense	long, semi-dense	long and dense	long, dense	long, dense	long, dense
Pilies	sub-dense	sub-dense	sparsely	sub-dense	sparsely	sparsely
Inflorescence	up to 8(9) flowers	up to 4 flowers	up to 7–9 flowers	up to 8–10 flowers	up to 7–9 (10) flowers	up to 9 flowers
Corolla color	yellow, violet at the apex	yellow	yellow turning to top	white, pale yellow	reddish yellow	yellow
Corolla shape	clavate	tubular to clavate	clavate	tubular, tubular-sub clavate	clavate	clavate to clavate-campanulate
Corolla length (mm)	up to 25	23–25	20–25	16–20	25–26	17–18
Pedicel length (mm)	ca. 1.5	2	ca. 2	ca. 1.5	ca. 10	ca. 7
Calyx lenghth (mm)	16–18(19)	15–17	19–20	9–10	20–25	12–16
Bract shape	lanceolate	linear-lanceolate	lanceolate	linear	linear-lanceolate	linear-lanceolate
Bract length (mm)	15–20(22)	18–20(25) ×2-3	7–8	9–10	20–25	ca. 10
Anther length (mm)	ca. 8	ca. 8	7–8	5–6	5–6	ca. 4
Annulus	glabrous	glabrous	glabrous	glabrous	glabrous	glabrous
Nutlet (mm)	(2.8)3 × 3.8(4)	-	3–3.2 × 2-3	3.5–4 × 2-2.5	4.5–6.5 × 3-3.5	3.8–4×2.5–2.8
Elevation range (a.s.l., in m)	2100–2200	2100–2280	2100–3200	1700–2500	2200–2600	2800–3235
Phytochorion	Atropatene	Atropatene	Atropatene, Kurdistan-Zagros	Atropatene	Kurdistan-Zagros	Atropatene
Geomorphologic Unit	Alborz	Alborz	northwest	northwest	west	northwest
Geologic formation	Ig	Ig	Ig, Me, Se, Vo-Se, QD	Ig, Me, Se, Vo-Se, QD	Se	Op
Bioclimatic Unit	MXc	MXc	MXc, MPc	MXc, MPc	MPc	MXc
Soil	LS	LS	Cl-BS	Cl-BS	BS	Cl-BS

Table 2. Main characters distinguishing Onosma pseudohebebulba and Onosma bozghushica from its morphologically similar species. MXc: Mediterranean xeric-continental, MPc: Mediterranean pluviseasonal continental, Ig: Igneous Rocks, Me: Metamorphic, Vo-Se: Volcano-Sedimentary, Se; Sedimentary, QD: Quaternary Deposits, LS: Lithosols from igneous rocks, brown soils and Sierozome Soils CL: Calcareous Lithosols-Brown Soils and Chestnut Soils, BS: Brown Soils.

Characters	*Onosma pseudohebebulba*	*Onosma bozghushica*	*Onosma iranshahri*	*Onosma bilabiata*	*Onosma hebebulba*
Plant height (cm)	25–30	22–25(30)	30–35	15–25	20–35
Rosette leaves size (mm)	30–80(85) × 8–10(12)	40–75(85) × 7–10(12)	25–50 × 10–25	30–110 × 7–25	70–100 × 25–30
Rosette leaves shape	spathulae	spathulae	spathulae-oblong	oblanceolate, lanceolate, oblanceolate-spathulate, obovate-lanceolate	obovate-spathulate
Basal leaf size (mm)	12–25 × 4–5	15–32 × 2–4	20–35 × 10–20	10–15 × 3–5	15–5 × 10–20
Basal leaf shape	spathulate	spathulate	pathulae-oblong	spathulate	obovate-spathulate
Cauline leaf size (mm)	18–35(40) × 7–9(10)	30–35 × 4–7	25–50 × 10–20	30–80 × 10–18	20–50 × 5–12
Cauline leaf shape	spathulate	spathulate	spathulae-lanceolate	lanceolate-oblong	lanceolate to lanceolate-oblong
Indumentum	sub-dense, semi-adpressed	dense, adpressed	sub-dense, sub-adpressed	sub-dense, sub-adpressed	sub-dense, sub-adpressed
Tubercle rays	short, sparse	short, sub-dense	long, severely dense	short, sub-dense	short, sub-dense
Pilies	dense	sparse	sparse	dense	dense
Inflorescence	dense	dense	sub-dense	dense	sub-dense
Corolla color	yellow	yellow	yellow	white to yellow	yellow
Corolla shape	clavate, tubular to clavate	tubular to campanulate	clavate-campanulate	tubular-campanulate	tubular-tubular-campanulate
Corolla length(mm)	17–19 (20)	20–22	18–20	25–30	18–20
Pedicel length (mm)	ca. 7	ca. 7	ca. 5–9	ca. 3	ca. 4
Calyx lenghth (mm)	18–20(22)	16–18(20)	18–20	18–27	15–17
Bract shape	lanceolate to lanceolate-linear	lanceolate	lanceolate	lanceolate	lanceolate-linear
Bract length (mm)	12–25	18–24 × 5–6(7)	9–10	10–18	10–12
Anther length (mm)	ca. 7	ca. 10	ca. 8.5	9–10	6–8
Annulus	glabrous	glabrous	sparsely pilose	glabrous	glabrous
Nutlet (mm)	3.5–4 × 2.5–3	-	-	5–6.5 × 3–3.5	-
Elevation range (a.s.l., in m)	1374	2560	1800–1900	2000–2600	1900–2200
Phytochorion	Kurdistan-Zagros	Atropatene	Kurdistan-Zagros	Atropatene	Atropatene- Kurdistn-Zagros
Geomorphologic Unit	west	Alborz	west	west	northwest, west
Geologic formation	Ig, Se	Ig	QD, Ig	Ig, Vo-Se, QD	Se, QD
Bioclimatic Unit	MPc	MPc	MPc	MXc	MXc, MPc
Soil	CL-BS	LS	CL-BS	BS-LS	CL, LB

RESULTS AND DISCUSSIONTop

Iran and particularly a segment of the prominent natural plateau of Iran featuring several mountainous barriers (Zagros, Alborz, Kopet Dagh, and Makran mountain ranges) and scattered internal mountains is considered one of the main centers of diversity and endemism for angiosperms in the world (Hedge & Wendelbo, 1978Hedge, I. C. & Wendelbo, P. 1978. Patterns of distribution and endemism in Iran. Notes from the Royal Botanic Garden, Edinburgh 36: 441–464.; Jalili & Jamzad, 1999Jalili, A. & Jamzad, Z. 1999. The Red Data Book of Iran. Research Institute of Forests and Rangelands, Tehran. Iran.) because of the area’s variability in paleo-biogeographical events (Takhtajan, 1986Takhtajan, A. 1986. Floristic regions of the world. University of California Press, Berkeley. ) and climatic and geomorphologic heterogeneity (Khalili, 2005Khalili, A. 2005. Climate of Iran. In: Banaei et al. (Eds.), The soils of Iran. New achievements in perception. Management and use Soil and Water Research Institute of Iran, Tehran: 24–71 [Persian].; Stöklin, 1968Stöklin, J. 1968. Structural history and tectonics of Iran: a review. AAPG Bulletin 52: 1229–1258. https://doi.org/10.1306/5D25C4A5-16C1-11D7-8645000102C1865D).

The Zagros as well as the northwest geomorphologic unit of Iran are reported to be the main centers of diversity and endemism for Onosma in Iran (Mehrabian, 2015Mehrabian, A. R. 2015. Distribution patterns and diversity of Onosma in Iran: with emphasis on endemism conservation and distribution pattern in SW Asia. Rostaniha 16: 36–60 (in Persian).; Moradi, 2017Moradi Zeinab, H. 2017. Geobotany evaluation of endemic species of Onosma L. (Boraginaceae) in order to protect and manage habitats in Ardabil and East Azerbaijan provinces. Master Thesis, Shahid Beheshti University, Tehran.; Naghizadeh, 2017Naghizadeh, S. 2017. Ecological evaluation of endemic species of Onosma L. (Boraginaceae) in order to protect and manage habitats in West and East Azerbaijan provinces. Master Thesis, Shahid Beheshti University, Tehran.; Khajoei Nasab et al., 2020aKhajoei Nasab, F., Mehrabian, A. R. & Namati Porshokouh, A. 2020a. Distribution patterns and endemism of the genus Onosma L. (Boraginaceae) in central Alborz. Iranian Journal of Applied Ecology 9: 1–18. https://doi.org/10.47176/ijae.9.2.12551). Accordingly, the mentioned zones were evaluated by our team through several field studies, which led to the discovery and identification of numerous new species of Onosma (Mehrabian et al., 2013Mehrabian, A. R., Sheidai, M. & Mozaffarian, V. 2013. Three new species of Onosma L. (Boraginaceae) from Iran. Feddes Repertorium. 124: 69–79. https://doi.org/10.1002/fedr.201300007; Mehrabian & Amini Rad, 2018Mehrabian, A. R. & Amini Rad, M. 2018. Onosma moussavi sp. nov. (Boraginaceae) from Zagros Mountain (s), Iran. Feddes Repertorium 129: 1–8. https://doi.org/10.1002/fedr.201700020; Mehrabian & Mozaffarian, 2018Mehrabian, A. R. & Mozaffarian, V. 2018. Seven new species of Onosma L. (Boraginaceae) with emphasis on their habitats in Iran. Taiwania 4: 366–388., Mehrabian et al., 2022Mehrabian, A. R., Naghizadeh, S., Khajoei Nasab, F., Moradi Zeinab, H. & Mozaffarin, V. 2022. Two new species and a new record for Onosma L. (Boraginaceae) from northwestern Iran. Feddes Repertorium 133: 1–21. https://doi.org/10.1002/fedr.202000036). Moreover, this area is one of the main targets for scientific field work in the region, leading to the collection and determination of unknown species. Accordingly, a comprehensive literature review and the evaluation of numerous Iranian and European herbaria led to the identification of these novelties. Our argument for these novelties is based on the prominent divergence in the most important morphological characteristics (e.g. trichome type, shape and size of the calyx, shape and size of cauline and basal leaves, annulus hairs, shape and size of the corolla, anther length, and connection as well as their exertion, nutlet morphology, and pollen type and size) compared with their closest allied species that were highlighted by literature (Boissier, 1875Boissier, P. E. 1875. Flora orientalis, sive enumeratio plantarum in Oriente a Graecia et Aegypto ad Indiae fines hucusque observatarum 4. H. Georg, Geneva, Basilea & Lugdunum.; Popov, 1953Popov, M. G. 1953. Boraginaceae. In: Shishkin, B. K. (Ed.), Flora of USSR 19. Keter Publishing House, Jerusalem.; Johnston, 1954Johnston, I. M. 1954. Studies in the Boraginaceae XXVI. Further evaluations of the genera of the Lithospermeae. Journal of Arnold Arboretum. 35: 1–8. https://doi.org/10.5962/bhl.part.8314; Riedl, 1967Riedl, H. 1967. Boraginaceae In: Rechinger, K. H. (Ed.), Flora Iranica 48. Akademische Druck-u. Verlagsanstalt, Graz: 170–212. ; Polunin, 1969Polunin, O. 1969. Flowers of Europe. Oxford University Press, Oxford.; Ball, 1972Ball, P. W. 1972. Onosma L. In: Tutin, T. G., Heywood, V. H., Burges, N. A., Moore, D. M., Valentine, D. H., Walters, S. M. & Webb, D. A. (Eds.), Flora Europaea 3. Cambridge University Press, Cambridge: 89–95.; Nasir & Ali, 1979Nasir, E. & Ali, S. I. 1979. Flora of West Pakistan, an annotated catalogue of vascular plants of West Pakistan and Kashmir by RR Stewart. Fakhri Press, Karachi. ; Post, 1986Post, G. E. 1986. Flora Syria, Palestine and Sinai. The American Press, Beirut.; Khatamsaz, 2002Khatamsaz, N. 2002. Boraginaceae L. In: Assadi, M., Massoumi, A. A., Mozaffarian, V. & Jamzad, Z. (Eds.), Flora of Iran. Research Institute of Forests and Rangelands, Tehran.; Mehrabian et al., 2013Mehrabian, A. R., Sheidai, M. & Mozaffarian, V. 2013. Three new species of Onosma L. (Boraginaceae) from Iran. Feddes Repertorium. 124: 69–79. https://doi.org/10.1002/fedr.201300007; Sayadi et al., 2017Sayadi, S. & Mehrabian, A. R. 2017. Distribution patterns of Convolvulaceae in Iran: priorities for conservation. Rostaniha 18: 181–197.; Mehrabian & Amini Rad, 2018Mehrabian, A. R. & Amini Rad, M. 2018. Onosma moussavi sp. nov. (Boraginaceae) from Zagros Mountain (s), Iran. Feddes Repertorium 129: 1–8. https://doi.org/10.1002/fedr.201700020; Mehrabian & Mozaffarian, 2018Mehrabian, A. R. & Mozaffarian, V. 2018. Seven new species of Onosma L. (Boraginaceae) with emphasis on their habitats in Iran. Taiwania 4: 366–388.). In addition, these new species have been confirmed by the noticeable geobotanical divergence emphasized in recent studies (Cecchi et al., 2011Cecchi, L., Coppi, A. & Selvi, F. 2011. Evolutionary dynamics of serpentine adaptation in Onosma (Boraginaceae) as revealed by ITS sequence data. Plant Systematics and Evolution 297: 185–199. https://doi.org/10.1007/s00606-011-0506-3; Mehrabian, 2015Mehrabian, A. R. 2015. Distribution patterns and diversity of Onosma in Iran: with emphasis on endemism conservation and distribution pattern in SW Asia. Rostaniha 16: 36–60 (in Persian).; Moradi, 2017Moradi Zeinab, H. 2017. Geobotany evaluation of endemic species of Onosma L. (Boraginaceae) in order to protect and manage habitats in Ardabil and East Azerbaijan provinces. Master Thesis, Shahid Beheshti University, Tehran.; Nikjouyan, 2018Nikjouyan, M. J. 2018. Geobotany evaluation of species of Onosma L. (Boraginaceae) in order to protect and manage habitats in Central Zagros. Master Thesis, Shahid Beheshti University, Tehran. ; Nemati, 2019Nemati Porshokooh, A. 2019. Geobotanical assessment of habitats of Onosma L. (Boraginaceae) in order to conservation and habitat management in central Alborz. Master Thesis, Shahid Beheshti University, Tehran.; Naghizadeh et al., 2020Naghizadeh, S., Mehrabian, A. R., Moradi Zeinab, H., Mostafavi, H. & Khajoei Nasab, F. 2020. Distribution patterns, diversity and conservation Priorities of Onosma L. (Boraginaceae Juss.) in part of Armeno-Iranian Province of Iran. Journal of Plant Research 34: 451–463 [in Persian].). Thus, the new species represent a significant differentiation compared with the species last introduced from the area under study as well as neighboring regions and improve our arguments that these unknown specimens can be considered as new species.

Onosma madjnoniani Mehrabian & Naghizadeh, sp. nov. (Fig. 1)

Figure 1. Holotype of Onosma madjnoniani Mehrabian & Naghizadeh sp. nov.

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Type: Iran. Eastern Azerbaijan province: Oscoo, Kord Abad to Gonbarf Village, 2200 m a.s.l., 37º 43’ 13” N, 46º 12’ 40” E, 28 May 2017, Naghizadeh 2018658 (Holotype: HSBU).

Diagnosis: Onosma madjnoniani is closest to O. armena DC. Nevertheless, it is distinguished by its rosette leaf size, lanceolate to lanceolate-linear leaves, short and sparse dense rays, and shorter calyx and bracts. In addition, it is similar to O. rascheyana Boiss., but O. madjnoniani is differentiated by its lanceolate to lanceolate-linear leaves, long rosettes, long bracts, short and sparse trichomes, and sparse hairs. Additionally, it is similar to O. kurdica Teppner, a restricted endemic species found in western Iran; however, it differs from O. kurdica in having lanceolate to lanceolate-linear cauline leaves, long basal leaves, short pedicels, nearly dense tuberculate rays, and sub-dense hairs. Moreover, it is apparently close to O. moussavi Mehrabian & Amini Rad, a restricted endemic species found in western Iran, but distinguished by long rosettes, lanceolate to lanceolate-linear rosette and cauline leaves, short and sub-dense tubercles of setae, nearly dense hairs and longer corolla, bracts, and calyx (Table 1).

Description: Perennial, with woody rootstock. Stems branched at the base, partly diffuse, flowering stems up to 20(25) cm tall. Indumentum asterotrichous (Fig. 2 A–L). Rosette leaves lanceolate to lanceolate-linear, 90–140(150) × 4–9 mm, attenuate at the base into a short petiole, obtuse to acute at apex, involute, covered by nearly dense monomorphic hairs, each consisting of an antrorsely appressed central seta, 1.2–1.8 mm long inserted on a multicellular tubercle with 8–10 asterosetulae, ca. 5–6 times shorter than the central seta. Basal leaves similar to rosette leaves, 25–45 × 4–7 mm. Cauline leaves sessile, lanceolate to lanceolate-linear, 25–90 × 5–9(10) mm, tapering into a short petiole, obtuse to acute at apex involute, setae similar to rosette leaves. Inflorescence simple to bifurcate, with 2–3 apical cymes, each cyme up to 8(9) flowers, pedicel ca. 1.5 mm long; bracts lanceolate, 15–20(22) mm. Flowering calyx 16–18(19) mm long, divided to base in linear to oblanceolate lobes, covered with short hairs mixed to yellowish setae, sparse near the margins and along the central vein, dense at the base and on the pedicel. Corolla clavate, up to 25 mm long and 1 mm wide, puberulent outside. Stamens included, inserted at the upper 3/4 of the corolla tube, anthers connate at the base, ca. 8 mm, apically bifid. Style protruding outside the corolla limb, up to 29 mm. Stigma bilobed. Nutlet not seen.

Figure 2. Scanning electron micrographs of Onosma madjnoniani: (A–C), indumentum of cauline leaves in the ventral surface; (D–F), indumentum of of cauline leaves in the dorsal surface; (G–I), indumentum of basal leaves in the ventral surface; (J–L), indumentum of basal leaves in the dorsal surface; (M–P), pollen in polar and equatorial view

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Pollen: Heteropolar, tricolporate, prolate, polar axis 13.8–16.8 µm, mean: 14.3µm, equatorial axis 11.1–12.8 µm, mean: 11.8µm, exine granulate 0.15–0.28 µm wide (Fig. 2 M–P).

Phenology: These plants flower from May to June and fruit from June to July.

Etymology: This plant was named in honor of Professor Henric Madjnonian, a pioneering, outstanding as well famous environmental conservationist of Iran, who has made extensive efforts and several scientific references to protecting Iran’s nature as well Protected areas.

Distribution and habitat: Onosma madjnoniani is a sub-alpine species confined to the NW geomorphologic unit of Iran in volcano-sedimentary rocks and brown soils (lithosols) as well as the Mediterranean xeric-continental (Mxc) global bioclimatic unit. Additionally, it is distributed in the steppes as cushion-shaped plant formations on the western slopes of the Sahand Mountains in the Irano-Armenian province of the Irano-Turanian region (Fig. 3).

Figure 3. Geographic location of the type locality of Onosma madjnoniani, O. riahii, O. pseudohebebulba and O. bozghushica in Iran.

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Conservation status: The estimated area of occupancy is less than 5000 km². The number of mature individuals in the population is less than 20. The status of critically endangered (CR) is suggested based upon the IUCN criteria (Bland et al., 2017Bland, L. M., Keith, D. A., Miller, R. M, Murray, N. J. & Rodríguez, J. P. (Eds.) 2017. Guidelines for the application of IUCN Red List of Ecosystems Categories and
Criteria (Version 1.1). IUCN, Gland.) because of its restricted distribution range and existing threatening issues in its habitats (e.g. over-grazing, land-use change, etc.).

Onosma riahii Mehrabian, sp. nov. (Fig. 4)

Type: Iran. Alborz, Taleghan, North of Taleghan Dam, 2140–2250 m a.s.l., 36º 11’ 23” N, 50º 39’ 12” E, 13 June 2018, Mehrabian 2018981 (Holotype: HSBU).

Figure 4. Holotype of Onosma riahii Mehrabian sp. nov.

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Diagnosis: Onosma riahii is similar to O. rascheyana but, it is characterized by linear to linear-lanceolate, cauline leaf size, dense hairs, longer calyx, and denser inflorescences. It is similar to O. armena, but, it is distinguished by its shorter rosette leaves, linear to linear-lanceolate leaves, oblong cauline leaves, sparse hairs, and longer calyx. Also similar to O. kurdica, it is nonetheless differentiated from it by its lanceolate-linear basal leaves, shorter cauline leaves, sparse hairs and pedicels, and long calyx. It is apparently also similar to O. moussavi, a restricted endemic species in northwestern Iran, but distinct in its rosette leaf size, linear to linear-lanceolate leaves, nearly dense hairs, and long corolla and pedicels (Table 1).

Description: Perennial, with woody rootstock and long narrow rhizomes. Stems partly erect, the flowering ones up to 35(40) cm, covered at the base with scaly remains of the pervious years shoots. Indumentum asteritericous (Fig. 5 A–L). Rosette leaves linear to linear-lanceolate, 30–40 × 2–3 mm, acute to acute-acuminate at apex, involute, tapering into a long petiole, hispid, white to yellowish white, upper surface coverd by dense as well monomorphic setae, each consisting of an antrorsely appressed, central setae 1.2–1.5 mm long, inserted on a multicellular tubercle with 10–12 asterosetulae, ca. 4–5 times shorter than the central seta. Basal leaves, sessile, other characters similar to rosette leaves. Cauline leaves linear-lnceolate, sessile, 50–90 × 4–4.5(5.5) mm. Inflorescence, bifurcate, 2–3 apical cymes, each cymes with 3–4 flowers, Pedicel ca. 2 mm long in fruit. Bracts linear-lanceolate, pubescent, 18–20(25) × 2–3 mm. Flowering and fruiting calyx divided to base, 15–17 mm long, linear lobes, covered with short yellow hairs very dense at the base and on the pedicel. Corolla tubular to clavate, yellow, with very reflexed lobes short, up to 24 mm long yellow, glabrous to puberulent outside. Nectary ring glabrous. Stamens excluded, inserted, connate at base, inserted upper 4/5 corolla tube, anthers up to 8 mm, apically bifid, Style protruding outside the corolla limb, up to 26 mm long. Stigma bilobed. Nutlets unknown.

Figure 5. Scanning electron micrographs of Onosma riahii: (A–C), indumentum in the ventral surface of basal leaves; (D–F), indumentum in the dorsal surface of basal leaves; (G–I), indumentum in the ventral surface of cauline leaves; (J–L), indumentum in the dorsal surface of cauline leaves; (M–P), pollen in polar and equatorial view

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Pollen: Heteropolar, tricolporate, prolate, polar axis 12.9–15.2 µm, mean: 13.8 µm, equatorial axis 8.5–10.8 µm, mean: 9.8 µm, exine granulate 0.14–0.26 µm wide (Fig. 5 M–P).

Phenology: This species flowers from May to June and fruits from June to July.

Etymology: The specific epithet is based on in honor of the eminent Phycologist-Mycologist Prof. Dr. Hossein Riahi in Iran, who has played a prominent role in improving our knowledge in this field in Iran.

Distribution and habitat: Onosma riahii is known only from the restricted zone in igneous formations in the lithosol (from igneous rocks), brown soils, and seirozome zone of Taleghan (Alborz). It is distributed in the Mediterranean xeric-continental (Mxc) global bioclimatic units of Iran (Fig. 3).

Conservation status: Onosma riahii is known only from the restricted zone on the southern slopes of the Western Alborz Mountains with an estimated area of occupancy less than 5000 km². Critically endangered status (CR) is proposed based on IUCN criteria (Bland et al., 2017Bland, L. M., Keith, D. A., Miller, R. M, Murray, N. J. & Rodríguez, J. P. (Eds.) 2017. Guidelines for the application of IUCN Red List of Ecosystems Categories and
Criteria (Version 1.1). IUCN, Gland.).

Onosma pseudohebebulba Mozaffarian & Mehrabian, sp. nov. (Fig. 6)

Figure 6. Holotype of Onosma pseudohebebulba Mozaffarian & Mehrabian sp. nov.

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Type: Iran. Charvar to Ivan, Tange Dalab 1374 m a.s.l., 33º 42’ 24” N, 46º 28’ 28” E, 5 May 2005, Mozaffarian 92841 (Holotype: TARI).

Diagnosis: Onosma pseudohebebulba is similar to O. hebebulba DC., found in western and northwestern Iran, though it is differentiated by its spathulate and shorter rosette leaves, shorter cauline leaves, and shorter pedicels and bracts. Onosma pseudohebebulba is similar to O. iranshahri Ghahr. & Attar, a restricted species endemic to western Iran. Regardless of its similarities, it differs in having shorter cauline leaves, sub-dense and sub-adpressed setae, shorter pedicels, and dense hairs. In addition, it is apparently close to O. bilabiata Boiss., a species endemic to western and northwestern Iran, yet distinguished by shorter rosette and cauline leaves, spathulate rosette leaves, thinner cauline leaves, sparse tubercle rays, shorter corolla, longer pedicels, and shorter anthers (Table 2).

Description: Caespitose perennial, with woody rootstock, creeping rhizome. Stems partly diffuse, branched at the base, covered at base with scaly remains of the previous year’s shoots and bearing sterile leaf rosettes, the flowering stems up to 25 cm tall. Indumentum asterotrichous (Fig. 7 A–L). Rosette leaves spathulate, involute, obtuse to acute at apex, hispid, white to yellowish, 30–80(85) × 8–10(12) mm, upper surface covered by very dense and monomorphic seta and hairs, nearly dense setose, each consisting of an antrorsely appressed, central seta, 1.5–2 mm long, inserted on a multicellular tubercle with 14–16 asterosetulae, ca. 4–5 times shorter than the central seta. Basal leaves spathulate, 12–25 × 4–5 mm, obtuse to acute at apex, seta similar to rosette leaves. Cauline leaves sessile, spathulate, 18–35(40) × 7–9(10) mm acute to obtuse at apex. Inflorescence capaitate, with 5–6 apical cymes, each cyme with 8–12 flowers, fruiting pedicel ca. 7 mm long. Bracts lanceolate to lanceolate-linear, 14–25 × 1–1.2 mm. Flowering Calyx divided near to base in oblanceolate lobes, 14–16(18) mm long, ca. 1 mm wide, Fruiting calyx 18–20(22) mm long, ca. 2.5 mm wide, covered by very dense seta and hairs. Corolla tubular, clavate to tubular-clavate, yellow, 17–19(20) mm long, yellow, densely puberulent outside., lobes ca. 1.5 mm long, ca. 1.5 mm wide; annulus glabrous. Stamens included, inserted at the upper two-thirds corolla tube anthers connate at base, 6–7 mm, apically bifid. Style little out of corolla, stigma bilobed. Nutlet ovoid, 3.5–4 × 2.5–3 mm long, beak to 1 mm long, navicular, green.

Figure 7. Scanning electron micrographs of Onosma pseudohebebulba: (A–C), indumentum in the ventral surface of basal leaves; (D–F), indumentum in the dorsal surface of basal leaves; (G–I), indumentum in the ventral surface of cauline leaves; (J–L), indumentum in the dorsal surface of cauline leaves.

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Phenology: This plant flowers from May to June and fruits from June to July.

Etymology: The name of this species relates to its similarity to O. hebebulba.

Distribution and habitat: Onosma pseudohebebulba is distributed in open oak forests with well fertilized soils (Fig. 3).

Conservation status: Onosma pseudohebebulba is known only from Chavar (the restricted zone of Ilam province) to Ilam city and Tang Dalab on the western slopes of the Southern Zagros Mountains. It has an estimated area of occupancy less than 5000 km². The status of critically endangered (CR) is proposed based on IUCN criteria (Bland et al., 2017Bland, L. M., Keith, D. A., Miller, R. M, Murray, N. J. & Rodríguez, J. P. (Eds.) 2017. Guidelines for the application of IUCN Red List of Ecosystems Categories and
Criteria (Version 1.1). IUCN, Gland.).

Onosma bozghushica Mozaffarian & Mehrabian, sp. nov. (Fig. 8)

Figure 8. Holotype of Onosma bozghushica Mozaffarian & Mehrabian sp. nov.

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Type: Iran. Eastern Azerbaijan, Bozghoosh Mountains, Neshagh top Mountains 2560 m a.s.l. 37º 45’ 54” N, 47º 41’ 43” E, 24 June 2010, Mozaffarian 94598 (Holotype: TARI).

Diagnosis: O. bozghushica is similar to O. iranshahri, but differing in its longer rosette leaves, shorter cauline leaves, dense and adpressed setae, dense hairs, and long corolla and bracts. It is also apparently similar to O. bilabiata. Nevertheless, it differs by having shorter and spathulate rosette leaves, sub-dense tubercle rays, longer pedicels, and shorter bracts and calyx. Moreover, it is similar to O. hebebulba, although differing by having longer rosette leaves and spathulate leaves (Table 2).

Description: Caespitose perennial, with woody rootstock as well long rhizome. Stems diffuse, the flowering stems 22–25 (30) cm tall. Indumentum asterotrichous (Fig. 9 A–L). Rosette leaves spathulate, 40–75 (85) × 7–10 (12) mm, acute at apex, involute, attenuate at the base to a long petiole, hispid, upper surface covered by very dense, monomorphic seta and hairs, each consisting of an antrosely appresse, central seta 1.5–2 mm long, inserted on a multicellular tubercle with 16–20 asterosetulae, 5–6 times shorter than the central seta. Basal leaves similar to rosette leaves, 15–32 × 2–4 mm. Cauline leaves sessile, 30–35 × 4–7 mm, spathulate, acute to obtuse at apex, involute Inflorescence 2–3 apical cymes, each cymes with 12–15 (17) flowers, pubescent. Bracts widely lanceolate, leaf like, pubescent, 18–24 × 5–6 (7) mm. Pedicel ca. 7 mm long. Calyx divided to base in oblanceolate lobes, 16–18 (20) mm long, covered with short and dense hairs mainly at the base. Corolla tubular to campanulate, yellow, ca. 22 mm long, glabrous to puberulent outside, lobes ca. 1mm long, ca. 1 mm wide; annulus glabrous. Stamens a little out of corolla, inserted at the upper 3/4 of corolla tube, connate at the base, ca. 10 mm, apically bifid. Style little out of corolla Limb. Stigma bilobed. Nutlet not found.

Figure 9. Scanning electron micrographs of Onosam bozghushica: (A–C), indumentum in the ventral surface of basal leaves; (D–F), indumentum in the dorsal surface of basal leaves; (G–I), indumentum in the ventral surface of cauline leaves; (J–L), indumentum in the dorsal surface of cauline leaves; (M–O), Pollen in polar and equatorial view.

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Pollen: Heteropolar, tricolporate, prolate, polar axis 11.7–12.9 µm, mean: 12.1 µm, equatorial axis 10.7–11.2 µm, mean: 10.9 µm, exine granulate 0.09–0.29 µm wide (Fig. 9 M–O).

Phenology: This plant flowers from May to June and fruits from June to July.

Etymology: The specific epithet is based on the geographical origin of the species (Bozghoosh Mountain) a prominent mountain range in northwestern Iran.

Distribution and habitat: Onosma bozghushica is known only from restricted zones in igneous formations in the lithosol (from igneous rocks), brown soils, and seirozome zone of Azerbaijan, on the southern slopes of the Bozghoosh Mountains. It is distributed in Mediterranean xeric-continental (Mxc) global bioclimatic units (Fig. 3).

Conservation status: Onosma bozghushica is known only from the restricted zone of Bozghoosh Mountains, Neshagh top mountains (Eastern Azerbaijan province) with an estimated area of occupancy less than 5000 km². Due to the narrow distribution range of the species, a status of critically endangered (CR) is proposed based on the IUCN criteria (Bland et al., 2017Bland, L. M., Keith, D. A., Miller, R. M, Murray, N. J. & Rodríguez, J. P. (Eds.) 2017. Guidelines for the application of IUCN Red List of Ecosystems Categories and
Criteria (Version 1.1). IUCN, Gland.).

ACKNOWLEDGMENTSTop

We thank Mr. Javadi in Central Laboratory of Shahid Beheshti University for his valuable helps in taking SEM images.

REFERENCESTop


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APPENDIX. Additional specimens examined, synoptic key to Onosma L. species in Flora Iranica region, and some additional notes.Top

Additional specimens examined

Onosma armena: Armenia, Aucher (Holotypus G 2358), Iran, West Azerbaijan, Makoo to Khoy, 2400–2650 m, Assadi & Mozaffarian (TARI 30353); Khoy to Ghotor, Sharif (IRAN 2693); Urmia, Ghooshchi, 1400–2100 m, Mousavi & Tehrani (IRAN 2692); Silvana, 1700 m, Runemark & Forooghi (TARI 19595); Zanjan, Mahneshan, Alam kandi, Ghalan Mts, 1900–2500 m, Mousavi et al. (IRAN 2689).

Onosma rascheyana: Iran, Between Rasch and Damascus, Boissier (G, Holotypus), West Azerbaijan, Bazargan, 2100–2250 m, Rechinger (W 06099); Azerbaijan, Kani Ziarat, Habasi bala, 2300–3000 m, Rechinger & Renz (W 05734); Chali Kuh be Rajan, 2600–3200 m, Rechinger & Renz (W 05728); Zanjan, Mahneshan to Takab, Belqeys Mts., 2700 m, Mehrabian (HSBU 281); Mahneshan, Angoran, 2700–2900 m (TARI 64841); 10 km E. Sanandaj, Furse (W 12625).

Onosma kurdica: Iran, Bijar, Kūh-e Ḩamzeh ‘Arab, 2200–2600 m, Lamond & Termeh (Holotype W 42261, isotype W 06112).

Onosma moussavi: Iran, West Azerbaijan, Salmas, Kozeh-Rash, Haravil Mountain, 2800–3235 m, 18 July 2016, Amini Rad & Bahrami Shad (Holotype IRAN 71503!, isotype HSBU 2016876).

Onosma hebebulba: Iran, Kurdistan, 45 km Kerend, Dalahoo Mts, 2000–2200 m, Assadi (TARI 60835); Kermanshah, Eslam Abad to Illam, Sharif (IRAN 5329); Illam, N.W. Ziam, Malegavan, 2200 m, Mozaffarian (TARI 95495).

Onosma iranshahri: Iran, Kurdistan, Marivan to Paveh, Gardan-e Tate, between Dezli and Hanigarmaleh, 1800–2600 m, Mozaffarian (isotype TARI 75701).

Onosma bilabiata: Iran, East Azerbaijan, Northern slopes of Sahand, 2300–2600 m, Assadi & Mozaffarian (TARI 30764); Mianeh, Termeh (W 579); Ardabil, Hashjin towards Barandigh, 1500m, Moradi (HSBU 2018229).

Synoptic key to Onosma L. species in Flora Iranica region

1. Haplotrichous

O. angustiloba Riedl, O. asperrima Bornm., O. assadi Mozaff. & Mehrabian, O. azarbaidjanensis Mehrabian, O. bakhteganensis Mozaff. & Mehrabian, O. barsczewskyi Lipsky, O. bodeana Boiss., O. bulbotricha DC., O. chitralica I.M. Jhonst., O. chrysochaeta Bornm., O. cornuta Riedl, O. demawendica Riedl, O. dichroantha Boiss., O. gaubae Bornm., O. griffithi Vatke, O. haussknechtii Bornm., O. hispida Steven, O. khorassanica Attar & Joharchi, O. khyberiana I. M. Johnst., O. kilouyensis Boiss. & Hausskn., O. kotschyi Boiss., O. limitanea I. M. Johnst., O. liwanovii Popov, O. longiloba Bunge, O. macrorhiza Popov, O. marivanensis Mozaff. & Mehrabian, O. microcarpa DC., O. oreodoxa Boiss. & Heldr., O. pachypoda Boiss., O. platyphylla Riedl, O. rechingeri Riedl, O. rostellata Lehm., O. sabalanica Ponert, O. sanandajensis Mehrabian & Mozaff., O. sarvestanica Mozaff. & Mehrabian, O. sericea Willd., O. sheidaii Mehrabian, O. stenosiphon Boiss., O. striata Riedl, O. subsericea Freyn, O. sulaimaniaca Riedl, O. trachytricha Boiss., O. yamagatae Kitamura, O. zerizamina Lipsky

-. Asterotrichous or Heterotrichous........................................................................................2

2. Cauline leaves very long (up to 150 mm), with prominent veins.............................................O. macrophylla Bornm., O. nervosa Riedl, O. mozaffariani Mehrabian

-. Cauline leaves short (up to 80 mm), lack of prominent veins..................................................3

3. Basal leaves obovate, widely spathulate.............................................O. dasytricha Boiss., Onosma albo-rosea Fisch. & C. A. Mey., O. ovalifolia Kotschy & Boiss., O. qandilica Rech. f. & Riedl, O. bisotunensis Attar & Hamzehee, O. hawramanensis Ahmad, O. hebebulba A. DC.

-. Basal leaves linear to linear-lanceolate, lanceolate, Oblong, oblanceolate, spathulathe...............4

4. Nectary ring pilos.............................................O. straussi (Riedl) Khatamsaz, O. lanceolata Boiss. & Hausskn., O. iranshahrii Ghahr. & Attar

-. Nectary ring glabrous........................................................................................................5

5. Basal leaves up to 200 mm.............................................O. chlorotricha Boiss. & Noë, O. kurdica Teppner

-. Basal leaves up to 110 mm...............................................................................................6

6. Bract up to 30 (32) mm. Rosette leaves oblong-spathulate, linear-spathulate. Cauline leaves ovate-lanceolate.............................................O. cardiostegia Bornm.

-. Bract up to 25 mm ..........................................................................................................7

7. Basal leaves up to 150 mm, linear-lanceolate. Corolla violet at the apex, up to 25 mm.............................................O. madjnoniani

-. Basal leaves up to 110 mm...............................................................................................8

8. Bract shorter than 19 (20) mm.........................................................................................9

-. Bract longer than to 25 mm.............................................................................................10

9. Setae mixed or heterogeneous (haplotricha and asterotricha).............................................O. elwendica Wettst, O. olivieri Boiss.

-. Setae heterotricha..........................................................................................................11

10. Corolla up to 27 mm. Rosette leaves linear-spathulate. Bracts linear-lanceolate, up to 25 mm .............................................O. targevarensis Mozaffarian & Mehrabian

-. Corolla up to 22mm. Bracts widely lanceolate, up to 24 mm. Rosette leaves spathulate.............................................O. bozgushica

11. Calyx up to 27 mm. Cauline leaves linear-oblanceolate, Oblanceolate-spathulate.............................................O. bilabiata Boiss. & Buhse

-. Calyx up to 22 mm ........................................................................................................12

12. Bract up to 10 mm.......................................................................................................13

-. Bract up to 20 mm..........................................................................................................14

13. Pedicel up to 2 mm. Corolla up to 25 mm.............................................O. rascheyana Boissier

-. Pedicel up to 7 mm. Corolla up to 18 mm.............................................O. moussavi Mehrabian & Amini Rad

14. Rosette leaves linear-Lanceolate, Corolla up to 23 mm..............................................O. riahii

-. Rosette leaves linear- spathulate. Corolla campanulate-clavate. Bract up to 10 mm..............................................O. armena A. DC.

15. Cauline leaves up to 60 mm. Bract up to 17 mm .............................................O. wendelboi Mehrabian & Mozaffarian (2018: 380), O. caerulescens Boiss.

-. Cauline leaves up to 35 mm. Bract up to 25 mm.............................................O. pseudohebebulba

Some additional notes

Onosma microcarpa DC.

O. microcarpa is a widespread species distributed in southwest Asia with morphological diversity as well high polymorphism, which has caused some populations to be described as separate species.

Syn. O. microcarpa DC.var. spathulata (Wettst.) Bornm.

Syn. O. microcarpa DC.var. stapfii (Wettst.) Bornm. & Riedl.

Syn. O. paboti Riedl.

Syn. O. lorestanica Attar & Sotoodeh

Onosma elwendica Wettst.

O. elwendica introduced from the Zagros region, which also spreads to Alborz with a very high variation. The density as well ratio of haplo and hetero setae showing high variation in different populations.

Syn. O. demawendica Riedl.

Syn. O. trachytricha Riedl.

Onosma gaubae Bornm.

The observed morphological differences (e.g. height and size of the corolla, anther, pedicel, basal leaves, and style protruding) are severely affected by environmental variation, especially climate factors. Therefore, no significant differentiation was observed in Tar Lake populations (that named as O. ghahremanii Attar & Naqinezhad) compared with other populations of O. gaubae (Khajoei Nasab et al., 2020bKhajoei Nasab, F., Mehrabian, A. R. & Namati Porshkouh, A. 2020b. Taxonomic revision of the Onosma gaubae sensu lato (Boraginaceae) based on morphological and ecological analysis. Plant Biosystems 155: 1019–1031. https://doi.org/10.1080/11263504.2020.1813827).

Syn. O. borragoidinum Ponert.

Syn. O. ghahremanii Attar & Naqinezhad

Onosma straussii (Riedl) Khatamsaz

Syn. O. sultanabadensis Ranjbar & Almasi

Onosma nervosa Riedl

Syn. O. maculata Ranjbar & Almasi

Onosma armena DC.

Syn. O. kerendica Attar & Mirtadz

Onosma bilabiata Boiss & Buhse

Syn. O. khoramabadensis Attar, Mirtadz. & Sotoodeh