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Abstract 


In this paper a critical annotated checklist of 256 endemic and near endemic species belonging to 152 genera and 50 families of flowering plants known from Hyrcanian relict forests is presented. Distribution maps of taxa, elevational range, number of known records, chorotypes, life forms, IUCN threat categories and habitat types are also provided. The chorotypes are categorized into eight main patterns: 1) the Omni-Hyrcanian pattern (OH), 2) West Hyrcanian pattern (WH), 3) Manjil-Rudbar pattern (MR), 4) Central Hyrcanian pattern (CH), 5) Central and East Hyrcanian pattern (CEH), 6) East Hyrcanian pattern (EH), 7) Alborz-Hyrcanian pattern (AH), and 8) Euxino-Hyrcanian pattern (XH). The richness and distribution maps were generated based on 5408 records gained from herbarium specimens and literature records. The life form spectra show that the majority of taxa (54.7%) belong to hemicryptophytes, followed by the tuberous, bulbous and parasitic geophytes with 45 species (17.6%) and phanerophytes with 28 taxa (10.9%). The conservation status of species according to IUCN criteria indicates that 30 taxa are Critically Endangered, 52 taxa Endangered, 30 taxa Vulnerable, 25 taxa Near Threatened and 81 taxa are of Least Concern. Our present data were not sufficient to evaluate 38 taxa that are categorized here as Data Deficient. The new combination of Leutea translucens (=Peucedanum translucens) is validated with inclusion of Peucedanum hyrcanicum as its synonym. The disjunct occurrence of the Caucasian species Gentiana grossheimii is reported from the eastern parts of the Hyrcanian forests in Iran for the first time. We conclude that (i) the Hyrcanian forests and associated habitats in the northern slopes of the Alborz Mountains harbour tremendous floristic diversity of high conservation priority, and (ii) the Hyrcanian forest zone is an important and unique center of endemism within the Euro-Siberian region that should be considered a floristic province with a large number of relict species.

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Plant Divers. 2022 Jan; 44(1): 39–69.
Published online 2021 Jul 27. https://doi.org/10.1016/j.pld.2021.07.005
PMCID: PMC8897184
PMID: 35281126

Plant diversity of Hyrcanian relict forests: An annotated checklist, chorology and threat categories of endemic and near endemic vascular plant species

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Supplementary Materials

Abstract

In this paper a critical annotated checklist of 256 endemic and near endemic species belonging to 152 genera and 50 families of flowering plants known from Hyrcanian relict forests is presented. Distribution maps of taxa, elevational range, number of known records, chorotypes, life forms, IUCN threat categories and habitat types are also provided. The chorotypes are categorized into eight main patterns: 1) the Omni-Hyrcanian pattern (OH), 2) West Hyrcanian pattern (WH), 3) Manjil-Rudbar pattern (MR), 4) Central Hyrcanian pattern (CH), 5) Central and East Hyrcanian pattern (CEH), 6) East Hyrcanian pattern (EH), 7) Alborz-Hyrcanian pattern (AH), and 8) Euxino-Hyrcanian pattern (XH). The richness and distribution maps were generated based on 5408 records gained from herbarium specimens and literature records. The life form spectra show that the majority of taxa (54.7%) belong to hemicryptophytes, followed by the tuberous, bulbous and parasitic geophytes with 45 species (17.6%) and phanerophytes with 28 taxa (10.9%). The conservation status of species according to IUCN criteria indicates that 30 taxa are Critically Endangered, 52 taxa Endangered, 30 taxa Vulnerable, 25 taxa Near Threatened and 81 taxa are of Least Concern. Our present data were not sufficient to evaluate 38 taxa that are categorized here as Data Deficient. The new combination of Leutea translucens (=Peucedanum translucens) is validated with inclusion of Peucedanum hyrcanicum as its synonym. The disjunct occurrence of the Caucasian species Gentianagrossheimii is reported from the eastern parts of the Hyrcanian forests in Iran for the first time. We conclude that (i) the Hyrcanian forests and associated habitats in the northern slopes of the Alborz Mountains harbour tremendous floristic diversity of high conservation priority, and (ii) the Hyrcanian forest zone is an important and unique center of endemism within the Euro-Siberian region that should be considered a floristic province with a large number of relict species.

Keywords: Biodiversity, Distribution pattern, Endemic plant, Hyrcanian forests, IUCN category

1. Introduction

The Hyrcanian forests stretch as a green continuous arc from the Talish in Azerbaijan Republic, cover the northern slopes of the Alborz Mountains along three Iranian Provinces (Gilan, Mazandaran and Golestan) and end in Golestan National Park (GNP) (Fig. 1). There are some isolated outliers of the Hyrcanian forests in North Khorassan Province (Jowzak area) and East Azarbaijan Province in the Arasbaran Protected Area (APA) (Akhani et al., 2010 and references therein). The area is climatically and topographically very heterogeneous, with a unique flora and vegetation (Frey and Probst, 1986; Akhani et al., 2010). These forests have a surface area of about 1.9 mio ha in Iran and 0.1 mio ha in Azerbaijan (Anonymous, 2010). Although large parts of the mesic areas are covered by lowland and montane forests and wetlands, there are also transitional habitats in nearby areas produced by the presence of arid and semi-arid Irano-Turanian flora and the high elevations of the Alborz Mountains. The diversity of transition zones between forests and adjacent Irano-Turanian vegetation is extremely high. For example, a total of 1362 species has been documented in GNP, which is almost 19% of the entire Iranian flora (Akhani, 1998, 2005).

Fig. 1

Map of the study area (green). GNP = Golestan National Park located in Golestan and North Khorassan provinces; APA = Arasbaran Protected Area located in East Azarbaijan Province, Iran. The Alborz Mountains harbour the Hyrcanian forests on the northern slopes, and the Irano-Turanian montane steppe and Juniper woodland on the southern slopes and/or in the subalpine to alpine zones of northern and southern slopes. The inset picture represents the boundaries of two main phytogeographic regions (Euro-Siberian and Irano-Turanian) disccused in this paper (Rudov et al., 2020). The vegetation types and boundaries are shown in Akhani et al. (2010): Fig. 2.

The Hyrcanian forests contain Arcto-Tertiary relict elements, which are of high phytogeographical interest. However, the intensification of human activity in this threatened forest zone has made conservation a major priority. Accordingly, in 2019, the Hyrcanian forests were designated a UNESCO World Heritage property (https://whc.unesco.org/en/list/1584).

In our first review paper we provided an overview of the flora, climate, major vegetation units, phytogeography, palaeoecology and conservation status of the Hyrcanian forests (Akhani et al., 2010). In this paper, an annotated critical checklist of endemic and near endemic species is provided, with plant chorology, conservation status and habitats. To evaluate phytogeographical status and threat categories, distribution maps are provided for all taxa. Further, we try to answer the following questions: (i) Is the Hyrcanian forest zone an autochthonous floristic province under the Euro-Siberian (boreal) region? (ii) Which species are endangered and require specific protection measures, and which taxa are Data Deficient based on IUCN threat categories?

2. Material and methods

2.1. Sources for taxonomic evaluation, literature and herbarium data

We searched for any floristic records from the northern provinces of Iran (Gilan, Mazandaran and Golestan), GNP, and the APA. The main sources of our data include 181 volumes of Flora Iranica (Rechinger, 1963–2015), 143 volumes of Flora of Iran (Research Institute of Forests and Rangeland: Assadi et al., 1992–2016), floristic records of the APA (Assadi, 1987, 1988; Hamzeh'ee et al., 2010) and GNP (Akhani, 1998), and our own collections and field studies over the entire Hyrcanian forests over the last three decades.

We constructed a second database consisting of only endemic and near endemic species. Complete information, including location data, elevational range and any ecological information, have been added to the data base. The chorology data were expanded from reliably identified herbarium specimens held in the following herbaria: 1) Herbarium of Halophytes and C4 Plant Research Laboratory (Hb. Akhani, School of Biology, University of Tehran); 2) Herbarium of Iranian Research Institute of Plant Protection (IRAN); 3) Herbarium of Research Institute of Forests and Rangelands (TARI) and 4) Herbarium of Botanical Garden and Botanical Museum of Berlin (B). Access to the Central Herbarium of University of Tehran (TUH) was not possible except for one specimen. For some critical taxa we checked either type specimens in mentioned herbaria or received photos from the herbarium of the Conservatoire botanique de la Ville de Genève (G), and through JSTOR, as well as some photos of specimens from Komarov Botanical Institute (LE).

All literature and herbarium data were georeferenced and converted into coordinates. Our access to reliable records outside of Iran, in particular the Caucasus, was limited. Therefore, we used point maps available in the monumental work, Flora Caucasus (Flora Kavkaza) (Grossheim, 1939, 1940, 1950, 1952, 1962, 1967), converting the points into coordinates. With the same method, we have extracted the point data of some arboreal species from Browicz's chorology of trees and shrubs in SW Asia (Browicz, 1982, 1983, 1984, 1986, 1988, 1991, 1992, 1994, 1996; Browicz and Zieliński, 1984). In total 5408 records (1272 herbarium data and 4136 literature records) have been used to generate distribution data and calculate the frequency of species. Records in the literature and specimens that we have been able to see in the herbarium are considered here as “seen plant”.

Nomenclature of plant families follows APG IV (Byng et al., 2016) and the nomenclature of taxa is mainly based on Flora Iranica (Rechinger, 1963-2015). However, in many outdated volumes we tried to use the latest taxonomic updates. In some cases, we have checked the identity of species using type specimens in herbaria or from their photos. In recent years several new species have been described from the study area mostly by our Iranian colleagues, which may require verification. Despite our efforts to see the types of these taxa, we were unable to check and ensure some of them because of restrictions we confronted with curators/staff. Therefore, these species are in the list of doubtful or excluded species. Voucher specimens were given for many species we studied. Specimens have been deposited in herbaria as indicated in Appendix 1; otherwise, they have been stored in the herbarium of Halophytes and C4 Plants Laboratory, School of Biology, University of Tehran (Herbarium H. Akhani).

2.2. Criteria to include species as endemic or near endemic

Endemic taxa are attributed to those species that exclusively occur in mesic parts of the Hyrcanian forests. Near endemic taxa are species found in the Hyrcanian area as well as in the transitional zones between the Hyrcanian area and the Irano-Turanian region, and/or species distributed through the Hyrcanian area and that have expanded further in the Caucasus and Euxinian areas. Phytogeographic assessment of species occurring in transitional habitats with the neighboring Irano-Turanian flora is problematic. The South Caspian forests on the northern slopes of the Alborz Mountains connect with the Irano-Turanian vegetation on the southern slopes of these mountains, above the timberline and many rain shadow valleys in the northern slopes (Fig. 1). It is controversial to say whether a species growing in transition zone is a dry-adapted Hyrcanian species or a mesic-adapted Irano-Turanian species. In such cases we apply the following measures:

  • 1)

    Those species growing in cliffs surrounded by forests and scrubs or occurring in coastal area with a distribution pattern within the Hyrcanian zone are accepted in our list as Hyrcanian species.

  • 2)

    The endemic species of the Sefidrud Massif (Fig. 1), known in our paper as Manjil-Rudbar pattern, are included in the list of Hyrcanian plants, despite the fact that these species are xerophytes growing in a rain shadow Mediterranean semi-arid climate.

  • 3)

    Species growing in moist transitional habitats such as meadows are considered Hyrcanian/Alborz transition species.

2.3. Distribution maps

All geographical coordinates were imported into DMAP (Morton, 2001) and distribution maps were generated for almost all taxa. The richness map of all species has been calculated using coincidence option for 30’ x 30’ grids. According to the inspection of 254 distribution maps and our own knowledge of the area, eight distribution patterns were defined for the Hyrcanian endemic and near endemic species. These patterns were determined based on the distribution range of each species along the study area. To identify the borders of each pattern, we generated multispecies maps of species that show similar distribution ranges.

2.4. Conservation status

The conservation status of all species has been assessed using IUCN criteria (IUCN, 2019). In some cases, we have only one old type collection. We preferred to classify such taxa as “Data Deficient” (DD) rather than considering them extinct.

We have presented two assessments for each species. In the first step we applied the assessment obtained by GeoCAT (a browser-based tool), which used the extent of occurrence (EOO) and the area of occupancy (AOO) according to criterion B of the IUCN Red List categories (http://geocat.kew.org; Moat, 2007; Bachman et al., 2011; Memariani et al., 2016; Pahlevani et al., 2020). The obtained categories have either been accepted or improved further, based on own field knowledge and field observations that support other IUCN criteria, i.e., A, C and D criteria (IUCN, 2019). The number of records gained from herbaria and literature provides useful information for evaluation of threat categories.

2.5. Habitat data

The Caspian forests and its transitional zones display high habitat heterogeneity (Akhani et al., 2010). The habitat types used in this paper are mostly based on our own field studies and available data in the literature. Our own extensive studies in GNP provide habitat information for a large number of species (Akhani, 1998).

3. Results and discussion

The number of genera and species (including subspecific taxa) of all plant families recorded from three Iranian northern provinces, Talish of Azerbaijan Republic, and two protected areas (GNP and APA) (Fig. 1) is provided in Table S1. A list of all endemic and near endemic taxa of the Hyrcanian area is provided in Appendix 1, with data on distribution, elevation range, number of records, chorotype, life form, threat category, habitat type, references, and one selected voucher specimen. Additional notes on the taxonomy of specific taxa are given in Appendix 2. A photographic selection of 37 representative endemic Hyrcanian species is depicted in Fig. 2, Fig. 3, Fig. 4. Distribution patterns of the Hyrcanian endemics and near endemics are shown in Fig. 5 and the percentage of these taxa within each distribution pattern is given in Fig. 6. The percentage of life forms of the Hyrcanian endemic and near endemic taxa is illustrated in Fig. 7. The numbers and percentages of these endemic and near endemic taxa in each threat category are summarized in Table 1. A richness map of all Hyrcanian endemics and near endemics is provided in Fig. 8. The distribution map of 254 Hyrcanian endemic and near endemic species is presented in Supplemental materials: Maps 1–254.

Fig. 2

Selected photos of Hyrcanian endemics and near endemics. A. Aristolochia hyrcana; B. Ruscus hyrcanus; C. Crocus caspius; D. Lilium ledebourii; E. Cephalanthera caucasica; F. Poa golestanensis; G. Leutea translucens; H. Ilex spinigera; I. Hedera pastuchovii; J. Centaurea hyrcanica; K. Epimedium pinnatum; L. Echium amoenum; M. Hesperis hyrcana; N. Buxus hyrcana. Photos: A, B, E-H, J-N by H. Akhani; C, I by A. Ghorbanalizadeh; D by A.R. Noormohammadi.

Fig. 3

Selected photos of Hyrcanian endemics and near endemics. A. Campanula lourica; B. Lonicera iberica; C. Saponaria bodeana; D. Evonymus velutina; E. Albizzia julibrissin; F. Parrotia persica; G. Quercus castaneifolia; H. Gentiana grossheimii; I. Scutellaria tournefortii; J. Tilia platyphyllos subsp. caucasica; K. Rhynchocorys maxima; L. Paeonia tomentosa; M. Papaver chelidoniifolium; N. Digitalis nervosa. Photos: A-D, F-N by H. Akhani; E by A. Ghorbanalizadeh.

Fig. 4

Selected photos of Hyrcanian endemics and near endemics. A. Primula heterochroma; B. Anemone caucasica; C, D. Pyrus boissieriana; E. Phuopsis stylosa; F. Populus caspica; G. Acer velutinum; H. Saxifraga wendelboi; I. Scrophularia vernalis subsp. clausii; J. Solanum kieseritzkii. Photos: A, B by A. Ghorbanalizadeh; C-J by H. Akhani.

Fig. 5

Distribution patterns of the Hyrcanian endemic and near endemic species. A. Omni-Hyrcanian pattern; B. West Hyrcanian pattern; C. Manjil-Rudbar pattern; D. Central Hyrcanian pattern; E. Central and East Hyrcanian pattern; F. East Hyrcanian pattern; G. Alborz-Hyrcanian pattern; H. Euxino-Hyrcanian pattern.

Fig. 6

Percentage of the Hyrcanian endemic and near endemic species within each distribution pattern.

Fig. 7

Life form spectra of endemic and near endemic species in the Hyrcanian area.

Table 1

Number and percentage of the Hyrcanian endemic and near endemic species in each Red List category.

Threat categoriesNo. of taxaPercent (%)
CR3011.7
EN5220.3
VU3011.7
NT259.8
LC8131.6
No category/DD3814.8
Total25699.9
Fig. 8

Richness map of all Hyrcanian endemic and near endemic taxa.

3.1. Alborz Mountains: Diversity hot spot in SW Asia

The total number of species recorded from three provinces (Gilan, Mazandaran and Golestan) and adjacent areas, including the APA and Talish, shows that 3855 vascular plant taxa (species and subspecies) belonging to 135 families and 893 genera are known from the area (Supplemental file 1: Table S1). There are only 55 species of Pteridophytes (ferns and fern allies) belonging to 15 families and 25 genera, and 10 species of Gymnosperms belonging to three families and five genera. In addition to 91 families, 681 genera and 3125 Dicot taxa and 23 families, 178 genera and 659 species of Monocots, these areas also contain basal Angiosperms, e.g., two species and two genera of Nymphaeales (Nymphaeaceae), and two species and one genus of Magnoliids (Aristolochiaceae); two species of Ceratophyllaceae, which is a sister family of Eudicots (Supplemental file 1: Table S1). The presence of at least 3855 species, which corresponds 52.8% of the total Iranian flora, in only three provinces of the northern Iran covering just 3.5% of the country's surface area is of great conservation importance. These provinces include the whole northern slopes of the Alborz Mountains and fragments of the southern slopes and parts of the Khorassan-Kopet Dagh Mountains (Fig. 1). This figure would increase if the whole Alborz area, i.e., the montane zones of Ardabil, Zanjan, Qazvin, Alborz, Tehran and Semnan provinces, is included. The presence of at least 1623 species in the area of Tehran and its vicinity to central Alborz is evidence of a hot spot center in the Irano-Turanian area. This is related to the complex lithological and elevational diversity, as well as the palaeoecological history of the area (Akhani et al., 2010, 2013) (Figs. 5G and and88).

Our floristic data support the hypothesis that the Alborz Mountains are an important diversity center in the Holarctic. The Irano-Turanian flora predominate the southern slopes and above timberline in the northern slopes and the Euro-Siberian flora dominate the northern slopes with higher precipitation. Parts of the deep valleys, such as Manjil-Rudbar Massif, are characterized by a Mediterranean climate and vegetation. The climatic diversity, presence of many high peaks and rich mountains ridges result in a tremendous heterogeneity for the growth of a wide spectrum of mesophytes, chasmophytes, oreophytes, psammophytes, halophytes, xerophytes, alpine and nival plants (Klein, 1994; Noroozi et al., 2008; Akhani et al., 2010, 2013; Noroozi and Akhani, 2013).

3.2. Hyrcanian forests: A regional center of diversity

In total, 256 endemic and near endemic taxa belonging to 50 families and 152 genera of flowering plants have been identified in the Caspian forests (Appendix 1; Fig. 2, Fig. 3, Fig. 4). This is less than the ca. 280 species which we have previously noted in our previous paper (Akhani et al., 2010). The reason for this is that we have excluded several taxa with doubtful taxonomic status and endemic taxa that require re-evaluation. As 11 species of our list are indeed Euxino-Hyrcanian species, we can accept 245 species belonging to endemic and near endemic species of the Hyrcanian area and nearby Alborz Mountains. By excluding an additional 62 species of the Alborz-Hyrcanian pattern, only 183 species remain as exclusive endemic species of the Hyrcanian forests.

The largest families of the Hyrcanian endemic and near endemic species are Asteraceae (13.3%), Apiaceae (7.8%), Rosaceae (7%) and Lamiaceae (5.8%), comprising one third of these taxa. The most species-rich genera among the Hyrcanian endemics and near endemics are Alchemilla L. (with nine species), Cousinia Cass., Dianthus L., and Veronica L. (each with seven species), and Astragalus L. and Leutea Pimenov (each with six species). The life form spectra (Fig. 7) show noticeable number of phanerophytes within the Hyrcanian endemic taxa. However, hemicryptophytes, with 140 species (54.7%), and geophytes, with 45 species (17.6%), are the most species-rich life forms in the area. The chamaephytes include only 12 species (4.7%), albeit, 10 further species (3.9%) could not well be classified as chamaephyte or hemicryptophyte.

There is consensus between most authors considering Hyrcanian forests as an autochthonous phytogeographical entity with a particular palaeoecological history, and particular flora and vegetation. However, its chorological position has fluctuated between province, subprovince and district. Zohary (1973) considered the area a district of the Pontic province belonging to the Euro-Siberian region. Takhtajan (1986) classified the area as the Hyrcanian province in the Irano-Turanian region. Frey et al. (1999) categorized the area as the Hyrcanian subprovince of the Euxino-Caucasian-Hyrcanian province, and Hedge and Wendelbo (1978) justified its position as the Hyrcanian subprovince of the Euro-Siberian region, supporting their view by the absence of Abies Mill., Picea A. Dietr., Pinus L. and Rhododendron L. in the area. The data in our paper confirms a regional center of diversity equivalent to a floristic province by the presence of 256 endemics and near endemics. The area is not only rich in isolated relicts but also harbors several vicariants with Euxinian and or northern and western Euro-Siberian areas, which probably evolved through allopatric speciation. All three important, widely distributed arboreal Hyrcanian species in the genera Ilex L., Buxus L. and Hedera L. have sister or related species in the Euxinian forests: Ilex spinigera (Loes.) Loes. (its Euxinian sister is Ilex colchica Pojark.) (Manen et al., 2010), Buxus hyrcana Pojark. (its Euxinian sister is B. colchica Pojark.) (Van Laere et al., 2011), Hedera pastuchovii Woron. ex Grossh. (its close affinity is H. colchica (K.Koch) K.Koch) (Vargas et al., 1999; Green et al., 2011).

The presence of 256 endemics and near endemics in a small area of forest zone, the presence of many relict species and a dozen of vicariant species with ecologically sister areas are justification for a floristic province. The Khorassan-Kopet Dagh floristic province located at the easternmost part of the Hyrcanian area is known with 356 endemics (Memariani et al., 2016).

There is some evidence that the Hyrcanian forests played a role in donating species to the Irano-Turanian region. The genus Leutea (Apiaceae) is an exclusively endemic genus in the Hyrcanian and Irano-Turanian region. Species of the arid parts of the Irano-Turanian are characterized by their cylindrical leaves (Pimenov, 1987). Discovery of a broad-leaved species (Leutea laseroides Akhani = Laser rechingeri Akhani) (Supplemental file 2: Map 53) in the moist cliffs of the eastern parts of the Hyrcanian forests, which is phylogenetically sister to the cylindrical-leaved species, is evidence of its ancestral state (Akhani, 1996, 1998; Valiejo-Roman et al., 2006). Both Rubiaceous species Phuopsis stylosa (Trin.) Hook. f (an endemic Hyrcanian genus, Fig. 4E, Supplemental file 2: Map 236). and Crucianella platyphylla Ehrend. & Schönb.-Tem (Supplemental file 2: Map 230). show plesiomorphic features (Schönbeck-Temesy and Ehrendorfer, 1989; Ehrendorfer and Schönbeck-Temesy, 2005). Alyssopsis Boiss. is also an isolated non-arboreal Hyrcanian endemic genus that phylogenetic studies show has an affinity with the Irano-Turanian genera Dielsiocharis O.E.Schulz and Calymmatium O.E.Schulz (Warwick et al., 2010; Toro-Nunez et al., 2015).

3.3. Distribution patterns and endemism

According to the distribution maps of 254 taxa (Supplemental file 2: Maps 1-254), eight geographical distribution patterns have been identified for the Hyrcanian endemic and near endemic species (Fig. 5).

  • 1)

    Omni-Hyrcanian pattern (OH, Fig. 5A)

This is the second largest group of species, including 61 species belonging to 34 families and 51 genera. The core range of most species lies in a continuous belt on the South Caspian forests. However, some more widespread species occur in proper surrounding areas such as isolated forests in the APA, East Caucasus up to South Dagestan, the Jowzak forest (in North Khorassan Province in the eastern part of the Hyrcanian area) and even mesic valleys in Turkmenistan. Some prominent arboreal examples of this group are the Tertiary relict elements such as Parrotia persica (DC.) C.A. Mey. (Fig. 3F; Supplemental file 2: Map 150), Quercus castaneifolia C.A. Mey. (Fig. 3G; Supplemental file 2: Map 147), Alnus subcordata C.A. Mey. (Supplemental file 2: Map 98), Acer velutinum Boiss. (Fig. 4G; Supplemental file 2: Map 239), Pyrus boissieriana Buhse (Fig. 4C and D; Supplemental file 2: Map 216), Frangula grandifolia (Fisch. & C.A. Mey.) Grubov (Supplemental file 2: Map 204) and Ilex spinigera (Fig. 2H; Supplemental file 2: Map 60). Many herbaceous species are associated with forest zones and their distribution ranges perfectly overlap with arboreal species. Examples are Digitalis nervosa Steud. & Hochst. ex Benth. (Fig. 3N; Supplemental file 2: Map 180), Primula heterochroma Stapf (Fig. 4A; Supplemental file 2: Map 196), Scutellaria tournefortii Benth. (Fig. 3I; Supplemental file 2: Map 163), Alyssopsis mollis (Jacq.) O.E. Schulz (Supplemental file 2: Map 108), Echium amoenum Fisch. & C.A. Mey. (Fig. 2L; Supplemental file 2: Map 100), and Centaurea hyrcanica Bornm. (Fig. 2J; Supplemental file 2: Map 68).

  • 2)

    West Hyrcanian pattern (WH, Fig. 5B)

This group includes 39 species belonging to 21 families and 31 genera. They usually occur from the central Mazandaran westwards to the Talish forests on the border of Iran and Azerbaijan. This area matches with the “temperate oceanic” bioclimate (Djamali et al., 2011) and is the most humid part of the Hyrcanian zone, where the annual precipitation may reach or even exceed 2000 mm in some parts. Epimedium pinnatum Fisch. (Fig. 2K; Supplemental file 2: Map 97), Pyrus grossheimii Fedor. (Supplemental file 2: Map 217), P. stylosa (Fig. 4E; Supplemental file 2: Map 236), Scrophularia rostrata Boiss. & Buhse (Supplemental file 2: Map 246) and Potentilla petraea Willd. ex Schlecht (Supplemental file 2: Map 215). are the most prominent species representing the West Hyrcanian pattern.

  • 3)

    Manjil-Rudbar pattern (MR, Fig. 5C)

Sefidrud Massif is a special area in the Hyrcanian lowlands, in which the climate is very dry and forms an island-like sub-desert in the moist zone of Iran. The low elevation and the strong winds create a rain shadow in this valley, forming sand dunes in parts of the area. The bioclimate of this zone is Mediterranean Xeric Oceanic (MXO) (Djamali et al., 2011). A total of 17 species belonging to 11 families and 16 genera are known to be endemic in this small area, including Cousinia erinacea Jaub. & Spach (Supplemental file 2: Map 76), Cousinia hypochionea Bornm. (Supplemental file 2: Map 78), Salvia oligophylla Auch. ex Benth. (Supplemental file 2: Map 158), Atraphaxis aucheri Jaub. & Spach (Supplemental file 2: Map 189) and Asperula sherardioides (Boiss.) Jaub. & Spach (Supplemental file 2: Map 226). A new undescribed species of Heliotropium is also known from this area (Supplemental file 2: Map 102). Most species of this pattern are either very rare, known only from type locality, e.g., Scorzonera persica Boiss. & Buhse (Supplemental file 2: Map 91), or are endangered species such as Aristolochia hyrcana Davis & M.S. Khan (Fig. 2A; Supplemental file 2: Map 1).

  • 4)

    Central Hyrcanian pattern (CH, Fig. 5D)

This pattern includes 15 species belonging to 11 families and 15 genera, with a center of distribution in forest zone of Mazandaran Province. Most species occur in scrubs and cliff habitats. Satureja isophylla Rech. f. (Supplemental file 2: Map 161) and Galium wendelboi Ehrend. & Schönb.-Tem. (Supplemental file 2: Map 235) are found in cliff habitats. Ophrys kojurensis Gölz (Supplemental file 2: Map 29), Hypericum fursei N. Robson (Supplemental file 2: Map 151), Pyrus mazanderanica Schönbeck-Temesy (Supplemental file 2: Map 219), Spiraea sheikhii Zare (Supplemental file 2: Map 222), and Astragalus lacus-valashtii Maassoumi, Podlech & Jalili (Supplemental file 2: Map 137) are examples of scrub habitats and Acer iranicum M. Mohtashamian & A. Rastegar/Acer mazandaranicum Amini, Zare & Assadi (Supplemental file 2: Map 238) a tree species of the forest zone.

  • 5)

    Central and East Hyrcanian pattern (CEH, Fig. 5E)

Species in this group occur from the central Hyrcanian forests and extend their range into the eastern parts up to Khorassan-Kopet Dagh area in some cases. This pattern includes 18 species belonging to 15 families and 18 genera. Except a few species such as Epipactis rechingeri Renz (Supplemental file 2: Map 26) and Heracleum gorganicum Rech. f. (Supplemental file 2: Map 48), which grow in shady forests, most other species, e.g., Berberis orthobotrys Bienert ex C.K. Schneider (Supplemental file 2: Map 96), Campanula lourica Boiss. (Fig. 3A; Supplemental file 2: Map 114), Alcea gorganica (Rech. f., Aell. & Esfand.) Zohary (Supplemental file 2: Map 169), Verbascum sublobatum Murb. (Supplemental file 2: Map 251) and Colutea buhsei (Boiss.) Shap. (Supplemental file 2: Map 142) are elements of cliffs, meadows and scrubs.

  • 6)

    East Hyrcanian pattern (EH, Fig. 5F)

The range of the 33 species (belonging to 14 families and 29 genera) in this group is located from 52°E longitude to the easternmost extensions of the Hyrcanian forests. They usually occupy open scrubs and rocky outcrops, and grasslands adjacent to the forests. Some species of this group are taxonomically and phylogenetically very isolated, such as Leutea laseroides (Supplemental file 2: Map 53), Johrenia golestanica Rech. f. (Supplemental file 2: Map 50), Centaurea golestanica Akhani & Wagenitz (Supplemental file 2: Map 67), Crucianella platyphylla (Supplemental file 2: Map 230) and Corydalis chionophila Czernjak. (Supplemental file 2: Map 177). Sometimes it is very difficult to distinguish these species from the surrounding phytochoria, in particular that of the Khorassan-Kopet Dagh Province of the Irano-Turanian region. According to Memariani et al. (2016) some species of this pattern have been categorized in the checklist of Khorassan-Kopet Dagh Province. GNP and Jahan Nama Protected Area are two important protected areas of Iran, in which many East Hyrcanian endemics occur (Akhani, 1998; Jafari and Akhani, 2008).

  • 7)

    Alborz-Hyrcanian pattern (AH, Fig. 5G)

Several mesophytic Irano-Turanian elements co-occur with the Hyrcanian species in the northern slopes of the Alborz Mountains, usually in deforested areas and rocky outcrops, or at the marginal and transitional parts in the timberline and nearby grasslands. One may consider these species, which constitute the largest group of endemics, as “xerophytic Hyrcanian” species or “mesophytic Irano-Turanian” elements, with 62 species, belonging to 21 families and 43 genera. In some cases, the range of some species extends to Talish in the west and Khorassan-Kopet Dagh Mountains in the east. Some prominent examples of the Alborz-Hyrcanian pattern are Phleum iranicum Bornm. & Gauba (Supplemental file 2: Map 35), Seseli elbursense Pimenov & Kljuykov (Supplemental file 2: Map 58), Dolichorrhiza persica (Boiss.) B. Nord. (Supplemental file 2: Map 84), Eritrichium gracillimum Rech. f. (Supplemental file 2: Map 101), Nepeta pogonosperma Jamzad & Assadi (Supplemental file 2: Map 156), Veronica paederotae Boiss. (Supplemental file 2: Map 186), Dionysia aretioides (Lehm.) Boiss. (Supplemental file 2: Map 195), Paraquilegia caespitosa (Boiss. & Hohen.) Drumm. & Hutch. (Supplemental file 2: Map 201), and Galium aucheri Boiss. (Supplemental file 2: Map 231).

  • 8)

    Euxino-Hyrcanian pattern (XH, Fig. 5H)

We have not included all Euxino-Hyrcanian species in this paper. The 11 selected species belong to 11 families and 11 genera. These species show an inconsistent pattern, ranging from the Euxinian species with some localities in the Hyrcanian forests [Erythronium caucasicum Woron. (Supplemental file 2: Map 20) and Polygonatum glaberrimum C. Koch (Supplemental file 2: Map 14)] or the Hyrcanian species with some localities in the Euxinian or Caucasus area [Froriepia subpinnata (Ledeb.) Baill. (Supplemental file 2: Map 47)], or widespread species occurring in both areas [Anemone caucasica Willd. ex Rupr. (Fig. 4B; Supplemental file 2: Map 198), Orchis adenocheila Czerniak. (Supplemental file 2: Map 31), Hesperis hyrcana Bornm. & Gauba (Fig. 2M; Supplemental file 2: Map 109), Teucrium hyrcanicum L. (Supplemental file 2: Map 166) and Lonicera iberica M. Bieb. (Fig. 3B; Supplemental file 2: Map 116)].

Comparison of the species richness of different distribution patterns (Fig. 6) indicates that the Alborz-Hyrcanian (Fig. 5G) and Omni-Hyrcanian (Fig. 5A) patterns are the richest in the area.

In the Caspian forests the richness of endemics in lowlands and sub-montane zones is low whereas it increases remarkably in montane forests and above timberline towards the limits of the area (Hedge and Wendelbo, 1978) (Fig. 9). This is a known phenomenon in Iran, explained by the geographical isolation in higher elevations (Noroozi et al., 2016). According to the results, the elevational distribution of Hyrcanian endemics and near endemics ranges between sea level in the Caspian lowlands and 4200 m in the alpine areas (Appendix 1). The largest numbers of Hyrcanian endemics and near endemics are found at elevations between 500 and 3000 m. Notably, many endemic and near endemic species, such as Poa masenderana Freyn & Sint. (75–2600 m), Centaurea hyrcanica (50–2810 m), C. zuvandica (Sosn.) Sosn. (0–3000 m), Crepis willemetioides Boiss. (50–2900 m), Echium amoenum (60–3000 m), Rhynchocorys maxima C. Richter (-26–2750 m), Polygala platyptera Bornm. & Gauba (0–2500 m) and Scrophularia rostrata (50–3010 m), have wide elevational distribution ranges in this area. The main reason for this extensive range is likely owed to favorable ecological conditions, including convenient temperature and humidity, from low to high elevations. Annual species become quite rare at high elevations (Mahdavi et al., 2013). These species are not common as endemics or near endemics in the Hyrcanian forests and mostly grow in the lowlands.

Fig. 9

Elevational distribution of endemic and near endemic species in the Hyrcanian area.

3.4. Hyrcanian forests: Irano-Turanian or Euro-Siberian?

Although the majority of phytogeographers have classified the Hyrcanian forests under the circumboreal or Euro-Siberian region (Zohary, 1973; Browicz, 1989; White and Léonard, 1991; Frey et al., 1999), a few, however, prefer to consider the area a mesophytic variant of the Irano-Turanian region (Takhtajan, 1986; Manafzadeh et al., 2017) or Sub-Mediterranean subregion (Meusel et al., 1965). The arguments in favour of the Euro-Siberian enclaves mostly refer to large number of arboreal species which are either circumboreal species such as Carpinus betulus L., Sorbus torminalis (L.) Crantz, Fraxinus excelsior L., Acer campestre L. and Berberis vulgaris L. (Browicz, 1989) or species having close affinity with some widely distributed boreal taxa belonging to Carpinus L., Quercus L., Fagus L., Castanea Mill., Acer L., Evonymus L., Zelkova Spach, and many more. The majority of Orchids and Pteridophytes growing in Iran are restricted to the Hyrcanian area as boreal or Euro-Siberian elements (Renz, 1978; Khoshravesh et al., 2009). Many Tertiary relict Hyrcanian endemic trees such as Parrotia persica (Fig. 3F; Supplemental file 2: Map 150), Quercus castaneifolia (Fig. 3G; Supplemental file 2: Map 147) and Acer velutinum (Fig. 4G; Supplemental file 2: Map 239) survived glaciations as refugees. The fossils of several species or their close affinities have been identified in Europe [(Bernasso in south France (Leroy and Roiron, 1996), the Mesovian Interglacial of Poland (Binka et al., 2003), and Willershausen in Germany (Ferguson and Knobloch, 1998)]. Twenty-nine of the species in Willershausen near Göttingen grow in the Hyrcanian forest zone and 16 further species are almost identical (Probst, 1981).

Parrotia C.A. Mey., which was long considered a monotypic genus, has a sibling Chinese species (P. subaequalis (Hung T. Chang) R.M. Hao & H.T. Wei) known from a few populations in Eastern China in Jiangsu, Anhui and Zhejiang provinces (Li and Del Tredici, 2008). Despite very long distances (6500 km), the two regions share many woody plant genera, including Acer, Albizzia Durazz., Buxus, Castanea, Carpinus, Diospyros L., Fagus, Pterocarya Kunth, Quercus, Sorbus L., Taxus L., Zelkova and Hedera (Valcárcel et al., 2017).

From vegetation point of view, none of the syntaxa of the Hyrcanian forest correspond with Irano-Turanian region. All four classes recently known from the area (Alnetea glutinosae Br.-Bl. et Tx. ex Westhoff et al., 1946, Alno glutinosae-Populetea albae P. Fukarek et Fabijanić 1968, Carpino-Fagetea sylvaticae Jakucs ex Passarge 1968, and Quercetea pubescentis Doing-Kraft ex Scamoni et Passarge 1959) are indeed Euro-Siberian higher syntaxa (Mucina et al., 2016; Gholizadeh et al., 2020).

The chorological analysis of the flora of Golestan National Park, the easternmost end of the Hyrcanian forests with the mixture of deciduous forests in the west and Irano-Turanian Juniper woodlands, Artemisia L. and montane steppes in southern, eastern and northern parts of the area shows that 15% of the species are Euro-Siberian plants (Akhani, 1998). The distribution of the Euro-Siberian species shows that 40% of the species are Omni-Euro-Siberian, 30% are Euxino-Hyrcanian sharing species and 30% are Hyrcanian endemics. Despite its easternmost location, many Euro-Siberian species dominate the mesic parts of the area and some interesting cases have been found as new records from that area, such as Carex pseudocyperus L., Hordelymus europaeus (L.) Harz and Alopecurus aequalis Sobol. (Akhani and Scholz, 1998; Akhani, 1999).

The Irano-Turanian region is characterized by its drought-adapted flora, which evolved under a particular climate differentiated from the Euro-Siberian region by continentality, winter temperature, and precipitation seasonality (Djamali et al., 2012). The climate of the Hyrcanian area has a characteristic temperate feature in the distribution of precipitation over the year, a very short dry period, and high air humidity (Akhani et al., 2010). However, the area deviates from the northern temperate forests by having an average minimum of temperatures in the coldest month above the freezing point. This provides not only support for growing thermophilous trees [such as deciduous species belonging to Parrotia, Pterocarya and Zelkova, and evergreen Buxus, Hedera and Ilex (Iversen, 1944)] but also the development of C4 grasslands in open areas (Akhani and Ziegler, 2002). The main argument that supports excluding the Hyrcanian forests from the Irano-Turanian region is the age difference of their endemics. The Irano-Turanian region harbors many neoendemics, with large numbers of recently radiated lineages adapted to the arid conditions, mostly diversified after orogenic activities since the Oligocene (Manafzadeh et al., 2014). But the endemics of the Hyrcanian area are either paleoendemics that remained as refugees in the area or evolved from common ancestors of widely distributed temperate elements due to geographical vicariance (Tralau, 1963).

The Irano-Turanian elements in the Hyrcanian zones only occur in the marginal habitats of the forests, rain shadow areas, rocky outcrops, above the timberline and sometimes also in the deforested parts. None of the typical Irano-Turanian species and genera are associated with the forests. Therefore, considering Hyrcanian forests as Irano-Turanian is not justified.

3.5. Hyrcanian area with highly threatened species

The endemic and near endemic species of the Hyrcanian forests area have been evaluated using the IUCN Red List categories and criteria. We did not accept preliminary evaluations for several species suggested by Jamzad and Jalili (1999) or Naqinezhad et al. (2015), because of the poor methodological approach of these studies. Our checklist includes 30 taxa (11.7%) as Critically Endangered (CR), 52 taxa (20.3%) as Endangered (EN), 30 taxa (11.7%) as Vulnerable (VU), 25 taxa (9.8%) as Near Threatened (NT), 81 taxa (31.6%) as Least Concern (LC) and 38 taxa (14.8%) are Data Deficient (DD), lacking reliable data to be evaluated (Table 1). Assessment of 44% of the Hyrcanian endemic and near endemic species as threatened is alarming for national and international community. The distribution of different threatened categories among eight known distribution patterns clearly shows that except for the “Omni-Hyrcanian”, “Central and East Hyrcanian” and “Euxino-Hyrcanian” patterns, all other patterns harbor a considerable number of Critically Endangered, Endangered and Vulnerable species (Fig. 10). The Caspian forests are among the most highly threatened ecosystems in Iran. Some of the main factors responsible for threatening populations of indigenous species include urbanization and industrialization, clearing land areas for agricultural purposes, livestock overgrazing, road and dam construction, firing, intensive mining, disposal of waste, and pollution (Akhani et al., 2010).

Fig. 10

Number of threatened endemic and near endemic species within distribution patterns of the Hyrcanian area.

Several threatened species of the Hyrcanian area, including Aristolochia hyrcana (Fig. 2A; Supplemental file 2: Map 1), Hyacinthella persica (Boiss. & Buhse) Chouard (Supplemental file 2: Map 10), Heliotropium sp. nov. (Supplemental file 2: Map 102), Nonea longiflora Wettst. ex Stapf (Supplemental file 2: Map 106), Scabiosa schimperiana Boiss. & Buhse (Supplemental file 2: Map 117) and Atraphaxis aucheri (Supplemental file 2: Map 189), grow in special habitats around Sefidrud valley (MR pattern, Fig. 5C). The area has been heavily damaged by a variety of land use activities such as road and dam construction along with development of agricultural activities (olive plantations). Other species, such as Crocus almehensis Brickell & Mathew (Supplemental file 2: Map 17), Ophrys kojurensis (Supplemental file 2: Map 29), Eriocycla ghafooriana Akhani (Supplemental file 2: Map 46), Leutea laseroides (Supplemental file 2: Map 53), Centaurea golestanica (Supplemental file 2: Map 67), Plantago podlechii Akhani (Supplemental file 2: Map 181) and Saxifraga ramsarica Jamzad (Supplemental file 2: Map 242), are very local endemics with distributions restricted to small areas in the Hyrcanian zone. Additional endemics have suffered from habitat loss in coastal areas, as well as lowland and montane forests, e.g., Daucus littoralis Smith subsp. hyrcanicus Rech. f. (Supplemental file 2: Map 44), Typha caspica Pobed. (Supplemental file 2: Map 39), Dianthus hyrcanicus Rech. f. (Supplemental file 2: Map 120), Polygala platyptera (Supplemental file 2: Map 188), Scrophularia megalantha Rech. f (Supplemental file 2: Map 245). and Veronica francispetae M.A. Fischer (Supplemental file 2: Map 183).

The Data Deficient category should be applied only in cases of unresolved taxonomy or uncertain locality information (Callmander et al., 2005). Many Hyrcanian endemics and near endemics designated as DD in the checklist (Appendix 1), including Tulipa harazensis Rech. f., Bunium scabrellum Korov., Crepis alfredi Bornm., Campanula ghilanensis Pall., Hypericum fursei, Phlomis ghilanensis C. Koch and Delphinium syncarpum Freyn, have incomplete locality information and other taxa require updated taxonomic assessments.

4. Conclusion

The Hyrcanian forests, a UNESCO World Heritage region, harbor highly valuable relicts of Tertiary elements. The area is suffering from many degrading factors threatening the unique ecosystems that contains many rare and endangered species both in the mesic parts of the forests and surrounding highlands. Our studies support the hypothesis that the Hyrcanian forests are a regional center of endemism within the boreal (Euro-Siberian) floristic region. Implementation of protection measures to stop reduction of forest area and reduce destructive factors should be combined with documentation of the status of individual species, identification of unknown populations, census of rare species and in situ and ex situ programs to ensure the survival of rare, endemic and threatened species.

Author contribution

H.A. conceived of the research idea, A.Gh. collected and provided the data, tables and maps. The tables were critically reviewed by H.A., who wrote the manuscript jointly with A.Gh.

Declaration of competing interest

The authors declare no conflict of interests.

Acknowledgments

The herbarium curators of the Plant Pests and Diseases Research Institute (IRAN), the Research Institute of Forests and Rangeland (TARI) and Botanical Garden and Botanical Museum of Berlin (B) are appreciated for allowing us to study available material in those herbaria. We are grateful to Targol Chatrenoor for preparing some photos of herbarium specimens from Komarov Botanical Institute (LE). We appreciate Alireza Noormohammadi for providing the photograph of Lilium ledebourii and Mahmood Shakiba for finding Gentiana grossheimii in Jahan Nama Protected Area.

Footnotes

Peer review under responsibility of Editorial Office of Plant Diversity.

Appendix ASupplementary data to this article can be found online at https://doi.org/10.1016/j.pld.2021.07.005.

Appendix 1. 

Annotated checklist of endemic and near endemic species of the Hyrcanian forests and marginal habitats including distribution along the Caspian forests, elevational range, number of records in surveyed references and herbarium data, chorotypes, life forms, threat categories, habitat types and references (with selected voucher). Threat categories are based on IUCN cateogories and criteria (IUCN, 2019). Superscript numbers are discussed in Appendix 2. Symbols and abbreviations. 1. Provinces and areas: Aras. = Arasbaran, Cauc. = Caucasia, E Azar. = East Azarbaijan, Gil. = Gilan, Gol. = Golestan, Maz. = Mazandaran, N Khor. = North Khorasan, Qazv. = Qazvin, R. Khor. = Razavi Khorassan, Sem. = Semnan, Tehr. = Tehran, Turk. = Turkmenistan, W Azar. = West Azarbaijan; 2. Chorotypes: AH = Alborz-Hyrcanian pattern, CEH= Central and East Hyrcanian pattern, CH= Central Hyrcanian pattern, EH = East Hyrcanian pattern, OH= Omni-Hyrcanian pattern, MR = Manjil-Rudbar pattern, WH= West Hyrcanian pattern, XH = Euxino-Hyrcanian pattern; 3. Life forms: Cham. = Chamaephyte, Geo. = Geophyte, Hem. = Hemicryptophyte, Phan. = Phanerophyte, Ther. = Therophyte; 4. Table heading and threat categories: Elev. range = Elevation range, Rec. = Records, EOO = Extend of occurrence, AOO = Area of occupancy, max. dist. = Maximum distance, Imp. ass. = Improved assessment, CR= Critically Endangered, EN = Endangered, VU= Vulnerable, NT= Near Threatened, LC = Least Concern, DD = Data Deficient; 5. Habitat types: AZ = Alpine zone, CSD= Coastal sand dune, DH = Disturbed habitat [around road, farm and garden], DV = Dry valley, FM= Forest margin, FO= Forest opening (clearing), GR = Grassland [SGR= Subalpine grassland], LF = Lowland forest [ALF = Alluvial lowland forest, CLF= Closed lowland forest, OLF= Open lowland forest], MF = Montane forest [CMF= Closed montane forest, OMF= Open montane forest], MH = Moist and Marshy habitat, MW = Meadow [AMW = Alpine meadow, LMW = Lowland meadow, MMW = Montane meadow, SMW= Subalpine meadow], MX = Mediterranean xeric enclave [PMX= Porphyry soil, SMX= Sandy soil, SEMX= Serpentine hill, SWMX= Shady woodland, STMX= Steppe, SAMX= Stony area], RH = Riverside (riparian) habitat, RO = Rocky outcrop (crevices of rock) [CRO= Calcareous rock, MRO = Marl rock, SRO= Schistous rock, VRO= Volcanic rock], SC= Scrub and shrubland [GSC = Grassy scrub, LSC = Lowland scrub, MSC = Montane scrub, OSC= Open scrub, SASC= Subalpine scrub, SMSC= Submontane scrub], SF= Submontane forest [CSF= Closed submontane forest, OSF= Open submontane forest], SP= Stony and pebble bed, ST= Steppe [LST = Lowland steppe, MST = Montane steppe, SAST= Subalpine steppe, SMST= Submontane steppe], SZ= Subalpine zone, VC= Vertical cliff [LVC = Limestone vertical cliff], WO= Woodland [GOWO = Grassy open woodland, JWO = Juniper woodland, LWO = Lowland woodland, MWO = Montane woodland, SWO= Subalpine woodland]; 6. F. = Flora, Herb. = It means the number of those records that seen in herbarium, Inc. = It means that the distribution map of this species is incomplete in outside Iran (Caucasus); 7. Names: A. = Akhani, A.&Gh. = Akhani & Ghasemkhani, As.&M. = Assadi & Mozaffarian, Fo. = Foroughi, G.&M. = Ghahreman & Mozaffarian, Gho. = Ghorbanalizadeh, J. = Jafari, M. = Mozaffarian, Ma.&Sa. = Maasoumi & Safavi, Ma.&Ja. = Maasoumi & Jalili, Man. = Manuchehri, Mo. = Moosavi Movahedi, N. = Noroozi, R.&Ma. = Runemark & Maassoumi, Re.&Ir. = Renz & Iranshahr, Sa.&Ga. = Sabeti & Gauba, Sh. = Shahsavari, T.&D. = Terme & Daryadel, T. = Terme, Z. = Zargani.

No.Family and species (Figure number)Distribution (Map number)Elev. range (m)Rec. (Herb.)ChorotypeLife formEOO (km2)AOO (km2)10% max. dist. (km)Threat category
Habitat typeReferences and (selected voucher)
GeoCatImp. ass.
Magnoliids and Monocots
Aristolochiaceae
1Aristolochia hyrcana Davis & M. S. Khan, (Fig. 2A)Gil., (Map 1)200–11007 (4)MRGeo.4061504.8EN B1+B2b (i,ii,iii)ENSWMXF. Iranica 26: 1–2; F. Iran 29: 4–6; (A. & al. 20,943)
Amaryllidaceae
2Allium grande Lipsky (= A. chelotum Wendelbo), (Akhani, 2005: Figs. 171–172)Gol., Maz., Sem./Gol. (Map 2)1000–310019 (8)CEHGeo.26,40917,67246.9NTVU C2bMSC, SMSCAkhani (1998); F. Iranica 76: 87–88; Fritsch and Abbasi (2013); (A. 11,305)
3Allium lenkoranicum Miscz., (Akhani, 2005: Fig. 180)Gol., Maz., Talish, Turk., (Map 3)600–160020 (5)OHGeo.127,73217,50080.4LCLCCSF, LMWAkhani (1998); F. Iranica 76: 61; Grossheim (1940); (A. 17,172)
4Allium subvineale Wendelbo, (Akhani, 2005: Fig. 186)Gil., Gol., Maz., (Map 4)1900–20604 (2)AHGeo.26,938750054.8NTEN C2bSMWAkhani (1998); F. Iranica 76:53; (A. 11,311)
5Allium talyschense Miscz.Talish, (Map 5)1800–24004WHGeo.1817384431.1EN B1ab (i,ii)ENROF. Iranica 76: 56–57; Grossheim (1940)
Asparagaceae
6Danaë racemosa (L.) Moench, (Akhani, 2005: Figs. 211–212)Gil., Gol., Maz., Talish, (Map 6)50–192369 (15)OH (disjunctly in East Mediterranean)Phan.59,96460,000179.3LCNTCLF, CSF, RO, VCAkhani (1998); Browicz (1988); F. Iranica 165: 177; (A. 13,387)
7Fessia gorganica (Speta) Speta (=Scilla gorganica Speta), (Akhani, 2005: Fig. 225)Gol., Maz., (Map 7)100–247214 (5)EHGeo.965713,69036.6VU B1b(i,ii)LCLF, MF, SC, SFAkhani (1998); F. Iranica 165: 114–115; Speta (1998); (A. 13,393)
8Fessia greilhuberi (Speta) Speta (=Scilla greilhuberi Speta)Gil., Maz., (Map 8)50–115013 (3)WHGeo.36,86926,41148.7NTNTLFF. Iranica 165: 113; Speta (1998); (A. 14,566)
9Fessia hohenackeri (Fisch. & C.A.Mey.) Speta (=Scilla hohenackeri Fisch. & C. A. Mey.)Talish, (Map 9)9WHGeo.3996308721.3EN B1b(i,ii)ENLF (Possibly)F. Iranica 165: 112–113; Grossheim (1940); Speta (1998)
10Hyacinthella persica (Boiss. & Buhse) ChouardGil., Gil./Qazv., (Map 10)390–7004MRGeo.4.173122.2CR B1b(i,ii,iii)CRSMXF. Iranica 165: 137–138
11Ornithogalum boissieri Bidarlord & F. Ghahrem.Ardabil/Gil., (Map 11)24001AHGeo.DDMMWBidarlord and Ghahremaninejad (2016)
12Ornithogalum bungei Boiss., (Akhani, 2005: Figs. 221–223)Gol., Maz., Sem./Gol., Talish, (Map 12)150–250026 (5)OHGeo.95,80827,50072.1LCLCCMF, FMAkhani (1998); F. Iranica 165: 124–125; Grossheim (1940); (A. 14,570)
13Ornithogalum sintenisii Freyn, (Akhani, 2005: Fig. 224)Gil., Gol., Maz., Talish, (Map 13)50–245256 (8)OHGeo.294,97660,00097.2LCLCLF, MWO, ROAkhani (1998); F. Iranica 165: 129; Grossheim (1940); (A. 13,943)
14Polygonatum glaberrimum C. KochGol., Maz., (Map 14)800–220062XHGeo.511,431127,500171.8LCLCLF, MF, SFF. Iranica 165: 179; Grossheim (1940)
15Pseudomuscari chalusicum (DC. Stuart) Garbari (= Muscari pseudomuscari (Boiss. & Buhse) Wendelbo)Gil. (without point in map), Gol., Maz., Sem./Gol., (Map 15)1100–18006 (1)CEHGeo.2953346834.5EN B1b(i,ii)ENGSC, ROF. Iranica 165: 148; Jafari and Maassoumi (2011); (A. 17,505)
16Ruscus hyrcanus Woron., (Fig. 2B)Gil., Gol., Maz., Talish, (Map 16)0–120070 (25)OHCham.90,25650,00058.4LCLCLF, SFF. Iranica 165: 178; (A. 14,139)
Iridaceae
17Crocus almehensis Brickell & Mathew, (Akhani, 2005: Fig. 226)Gol. (Golestan National Park), (Map 17)1470–20005 (2)EHGeo.79.12731.5CR B1+B2ab (i,ii)CRMMW, SASTAkhani (1998); F. Iranica 112: 5; F. Iran 31: 6–7; (A. & Sh. 6107)
18Crocus caspius Fisch. & C. A. Mey. (Fig. 2C)Gil., Gol., Maz., Talish, (Map 18)−20–130050 (21)OHGeo.69,07845,00053.6LCLCLFF. Iranica 112: 8 & 75; F. Iran 31: 9–10; (A. 13,774)
19Crocus gilanicus MathewGil., Maz., (Map 19)1500–24004WHGeo.4063235227.9EN B1ab (i,ii)ENSZF. Iranica 112: 11 & 75; F. Iran 31: 16
Liliaceae
20Erythronium caucasicum Woron.Gil., Maz., (Map 20)600–140017XHGeo.220,63640,000146.5LCNTLF, SFF. Iranica 165: 103; Grossheim (1940); Mathew (1992)
21Fritillaria kotschyana Herbert [incl. subsp. kotschyana & subsp. grandiflora (Grossh.) Rix], (Akhani, 2005: Figs. 279–280)E Azar.?, Gil., Gol., Maz., Talish, Tehr., (Map 21)1000–370041 (19)AHGeo.129,52725,00077.8LCLCAMW, MFAkhani (1998); F. Iranica 165: 72; (A. 13,386)
22Lilium ledebourii (Baker) Boiss., (Fig. 2D)Ardabil, Gil., Maz., Talish, (Map 22)1350–22509 (4)WHGeo.10,238490034.6VU B1ab (i,ii,iii)EN C1FM, SGRF. Iranica 165: 58–59; (T. & D. 30,061 (IRAN))
23Tulipa harazensis Rech. f.Maz., (Map 23)7001AHGeo.DDSTChristenhusz et al. (2013); F. Iranica 165: 91–92
24Tulipa ulophylla WendelboGol./Sem., Maz., (Map 24)600–25003AHGeo.125025CR B1b(i,ii,iii)DDSCF. Iranica 165: 95–96
Orchidaceae
25Cephalanthera caucasica Kränzl., (Fig. 2E)Aras., Gil., Gol., Maz., Talish, (Map 25)50–200348 (7)OHGeo.280,76155,00092.6LCLCCLF, CMFAkhani (1998); F. Iranica 126: 29–31; F. Iran 57: 15–16; Grossheim (1940); Hamzeh'ee et al., 2010; (A. 11,253)
26Epipactis rechingeri Renz, (Akhani, 2005: Figs. 253–254)Gol., Maz., (Map 26)950–220015 (4)CEHGeo.27,70314,17545.5NTVU C1MF, SFAkhani (1998); F. Iranica 126: 36–37; F. Iran 57: 19–20; (A. 11,982)
27Himantoglossum formosum (Stev.) C. KochTalish, (Map 27)1509WHGeo.22,076777635.6NTEN C1LF, MFF. Iranica 126: 90–91; Grossheim (1940)
28Ophrys × aghemani Renz (Ophrys scolopax × transhyrcana)Gol., (Map 28)12501EHGeo.CR
D
MWF. Iranica 126: 86
29Ophrys kojurensis GölzMaz., (Map 29)650–10207CHGeo.505.4CR B1b(i,ii,iii)CRGSCGölz et al. (2006); Gölz et al. (2007)
30Ophrys sintenisii Fleischamn. & Bornm.Gol., Maz., (Map 30)70–6506EHGeo.310.94690015.5EN B1b(i,ii,iii)ENLMWGölz et al. (2007)
31Orchis adenocheila Czerniak. (= O. stevenii auct Fl. Iranica non Rchb. f.), (Akhani, 2005: Fig. 262)Aras., Gol., Maz., Talish, Turk., (Map 31)150–261976 (12)XHGeo.359,59367,500146.1LCLCFM, LF, MF, SCAkhani (1998); F. Iranica 126: 115–116; F. Iran 57: 57–58; Grossheim (1940); (A. 13,511)
32Orchis caspia Trautv.Gol., Talish, (Map 32)18OHGeo.157,68535,000109.6LCEN
C1
MW, SCF. Iranica 126: 105–106; Grossheim (1940)
Poaceae
33Agropyron bulbosum Boiss.Maz., (Map 33)23002AHHem.DDGRF. Iranica 70: 156–157
34Festuca akhanii TzvelevGol., Maz., (Map 34)1300–17502 (2)CEHHem.405044.8CR B1b(i,ii)NTMSTAkhani (1998); Tzvelev (1997); (A. 10,806)
35Phleum iranicum Bornm. & GaubaMaz., Maz./Qazv., Maz./Sem., Maz./Tehr., (Map 35)2130–28005AHHem.1368172824.4EN B1ab (i,ii)ENMMWF. Iranica 70: 289; Nath and Nielsen (1963)
36Poa golestanensis H. Scholz & Akhani, (Fig. 2F)Gol., (Map 36)1250–238010 (10)EHHem.2099144418.7EN B1b(i,ii)ENMF, MMW, MSCAkhani (1998); (A. 11,329)
37Poa masenderana Freyn & Sint., (Akhani, 2005: Fig. 438)Gil., Gol., Iraq, Maz., Sem., Talish, Tehr., Turkey, (Map 37)75–260033 (8)OH (disjunctly it occurs in Zagros)Hem.524,96947,500115.9LCLCCLF, CMF, RHAkhani (1998); Edmondson (1985); F. Iranica 70: 38; Grossheim (1939); (A. 11,912)
38Trisetum bungei Boiss.Gol., Sem., (Map 38)26003AHHem.653.83645014.6EN B1+B2ab (i,ii)DDGRF. Iranica 70: 318
Typhaceae
39Typha caspica Pobed.Gil., Talish, (Map 39)2WHHem.183.4CR B1b(i,ii)DDMHF. Iranica 71: 3; F. Iran 42: 9
Eudicots
Apiaceae
40Bunium scabrellum Korov.Talish, (Map 40)1WHGeo.DDUnknownF. Iranica 162: 251–252
41Bupleurum flexile Bornm. & GaubaGil., Gol., Maz., (Map 41)1200–2700m15 (7)OHHem.52,69922,50056.6LCLCMF, ROF. Iranica 162: 290; F. Iran 54: 295; (A. 13,709)
42Bupleurum ghahremanii Mozaff.Maz., (Map 42)1850–29503 (3)CHHem.21.2CR B1+B2b(i,ii)CRMF, SPF. Iran 54: 295–296; (G.&M. 8436 (TUH))
43Cervaria cervariifolia (C. A. Mey.) M. Pimen.E Azar., Gil., Gol., Maz., Qazv., Talish, Tehr., Turk., (Map 43)500–350082 (8)OHHem.223,06360,00095LCLCJWO, MF, SCAkhani (1998); F. Iranica 162: 452–453; F. Iran 54: 449–451; Joharchi et al. (2007); (A. 11,157)
44Daucus littoralis Smith subsp. hyrcanicus Rech. f.Gil., Gil./Ardabil, Maz., (Map 44)−20–137016 (1)WHHem.20,24513,44840.9NTVU A1cCSDF. Iranica 162: 139–140; F. Iran 54: 181–183; (A. 15,455)
45Dichoropetalum ramosissimum (Mozaff.) Pimenov &
Kljuykov (=Johrenia ramosissima Mozaff.)
Maz., (Map 45)200–17005 (4)CHHem.228.368362.9EN B1+B2ab (i,ii)ENLF, MF, SFF. Iran 54: 367–368; (M. 72,066 (TARI))
46Eriocycla ghafooriana AkhaniGol., (Map 46)550–10154 (3)EHCham.39213.8CR B1b(i,ii)CRLVCAkhani (1998); F. Iran 54: 343–344; (A. 12,322)
47Froriepia subpinnata (Ledeb.) Baill.Gil., Gol., Maz., Talish, (Map 47)0–165041 (11)XHHem.354,02460,000153.3LCLCFO, SCAkhani (1998); F. Iranica 162: 297–298; F. Iran 54: 296–297; (A. 12,394)
48Heracleum gorganicum Rech. f.Gol., Maz., (Map 48)1220–26008 (4)CEHHem.10,53311,09443.1VU B1ab (i,ii,iii)LCCMFAkhani (1998); F. Iranica 162: 497; F. Iran 54: 483; (A. 10,591)
49Heracleum rechingeri MandenGil., Maz., Tehr., (Map 49)1250–29506WHHem.3760270029.7EN B1b(i,ii)VU A4MFF. Iranica 162: 496; F. Iran 54: 482–483
50Johrenia golestanica Rech. f.Gol., (Map 50)220–200021 (18)EHHem.680.0625046.3EN B1b(i,ii)ENFO, RO, SCAkhani (1998); F. Iranica 162: 376–377; F. Iran 54: 365–366; (A.12,302)
51Leutea glaucopruinosa (Rech.f.) Akhani & Salimian (=Peucedanum pimenovii Mozaff.)2Gol., Maz., (Map 51)1460–18906 (2)CEHHem.447.061204831.6EN B1b(i,ii)ENMF, ROF. Iranica 162: 443–444; F. Iran 54: 437–438; (A. 15,015)
52Leutea gracillima M. PimenovGol. (Golestan National Park), (Map 52)750–20803 (2)EHHem.71.052121.9CR B1b(i,ii)EN A4aCRO, JWOAkhani (1998); F. Iranica 162: 448–449; (A. 12,060)
53Leutea laseroides AkhaniGol. (Golestan National Park), (Map 53)700–20106 (6)EHHem.33.795201.6CR B1b(i,ii)EN A4aLVCAkhani (1998); (A. 12,361)
54Leutea nematoloba (Rech. f.) M. Pimen.Maz., Sem., (Map 54)1000–24007 (1)AHHem.461.596176420.6EN B1b(i,ii)ENCROF. Iranica 162: 447; F. Iran 54: 445; (A.&Gh. 15,473)
55Leutea polyscias (Boiss.) M. Pimen.Gil., Qazv., (Map 55)200–165016 (4)MRHem.140970010.2EN B1b(i,ii)ENVCF. Iranica 162: 446–447; F. Iran 54: 441–442
56Leutea translucens (Rech.f.) Akhani & Salimian (= Peucedanum translucens Rech. f.)3, (Fig. 2G)Maz., Gol., (Map 56)1750–27504 (3)EHHem.687.36276815.8EN B1ab (i,ii)CR C1JWOF. Iranica 162: 444; (A. 15,359)
57Polylophium involucratum (Pall.) Boiss.Maz., (Map 57)2000–30006AHHem.699.3172457.1EN B1+B2b(i,ii)VU A4AZ (MW, SMW)F. Iranica 162: 525; F. Iran 54: 507
58Seseli elbursense Pimenov & Kljuykov (=Trachydium eriocarpum Bornm. & Gauba)Maz., (Map 58)3200–34002AHHem.82.4CR B1+B2b(i,ii)EN A4AZ, CROF. Iranica 162: 181–182
59Seseli kiabii (Akhani) AkhaniGol., Maz., (Map 59)300–200010 (8)EHHem.7664360030.1VU B1ab (i,ii)VUOSC, ROAkhani (1998); Akhani et al. (2016); F. Iran 54: 349–350; (A. 12,370)
Aquifoliaceae
60Ilex spinigera (Loes.) Loes., (Fig. 2H)Gil., Gol., Maz., Talish, (Map 60)30–201868 (31)OHPhan.117,52947,50068.6LCLCLF, MF, SFAkhani (1998); Browicz (1984); F. Iranica 25: 1; F. Iran 130: 3–5; (A. 14,134)
Araliaceae
61Hedera pastuchovii Woron. ex Grossh., (Fig. 2I)Gil., Gol., Maz., Talish, (Map 61)50–1800m76 (16)OHPhan. (Liana)247,81067,500102.4LCLCLF, MF, SFBrowicz (1986); F. Iranica 102: 2–3; (A. 13,661)
Asteraceae
62Amblyocarpum inuloides Fisch. & C. A. Mey.Gil., Gol., Maz., Talish, (Map 62-Inc.)Sea level–680m9 (2)OHTher.53,03812,50057.6LCVU A4cMH, RHF. Iranica 145: 126; (Z. 7162 (IRAN))
63Anthemis mazandaranica IranshahrGil., Maz., (Map 63)900–15205WHTher.4325388836EN B1ab (i,ii)ENUnknownF. Iranica 158: 21; F. Iran 59: 26
64Anthemis talyschensis A. Fedor.Gil., Gil./Ardabil, Maz., Talish, (Map 64)1250–480010WHHem.16,117800040.1VU B1ab(i)VUAZ, MSCF. Iranica 158: 26; F. Iran 59: 34
65Anthemis triumfettii (L.) All. subsp. khorasanica (Rech. f.) IranshahrGil., Gol., Gol./N Khor., Gol./Semn, Maz., Turk., (Map 65)400–300029 (4)CEHHem.83,12316,00064.2LCLCLSC, LWO, MSC, MWOAkhani (1998); F. Iranica 158: 20; F. Iran 59: 22–26; (A. 10,684)
66Carduus transcaspicus Gandog. subsp. transcaspicusGol., Turk., (Map 66)780–17007 (4)EHTher.9961420528.9VU B1ab (i,ii)LCDH, FM, ROAkhani (1998); F. Iranica 139a: 223; (A. 11,476)
67Centaurea golestanica Akhani & WagenitzGol. (Golestan National Park), (Map 67)1200–15504 (4)EHHem.4.0941.0800.6CR B1+B2b(i,ii,iii)CRGOWOAkhani (1998); (A. 11,704)
68Centaurea hyrcanica Bornm., (Fig. 2J)Aras., Gil., Gil./Ardabil, Gol., Maz., Talish, (Map 68)50–2810101 (57)OHHem.130,02665,00081.5LCLCLF, MF, RO, SCAkhani (1998); F. Iranica 139b: 345–346; (A. & al. 21,281)
69Centaurea kandavanensis WagenitzAlborz, Maz., (Map 69)270–250015 (3)AHHem.3562129612.3EN B1b(i,ii)ENRO, SCF. Iranica 139b: 392; (T. & al. 8182 (IRAN))
70Centaurea meyerana Tzvel.Talish, (Map 70)12002WHHem.DDUnknownF. Iranica 139b: 404–405
71Centaurea zuvandica (Sosn.) Sosn.Ardabil, Aras., Gil., Gol., Maz., Sem., Talish, Tehr., (Map 71)0–300090 (49)OHHem.143,49450,00081.1LCLCCRO, VCAkhani (1998); F. Iranica 139b: 403–404; (A. & al. 21,044)
72Cephalorrhynchus gorganicus (Rech. f. & Esfand.) TuislGol., Maz., Talish, (Map 72)27003OHHem.58,033750062.5LCDDSWOF. Iranica 122: 212–213
73Cirsium gadukense PetrakMaz., (Map 73)21501AHHem.DDMST, OSCF. Iranica 139a: 257
74Cousinia alfredii Bornm. & GaubaAlborz, Maz., (Map 74)1300–15002AHHem.986.8CR B1b(i,ii)CRSF (Xeric slopes)F. Iranica 90: 92–93; F. Iranica 139a: 117
75Cousinia decipiens Boiss. & BuhseGol., Gol./N Khor., Gol./Sem., Maz., (Map 75)1000–281018 (8)EHHem.5440510327.4VU B1b(i)LCAMW, JWO, MST, SCAkhani (1998); F. Iranica 90: 158–159; F. Iranica 139a: 126; (A. 16,844)
76Cousinia erinacea Jaub. & SpachGil., (Map 76)300–10006MRHem.716.5161254.6EN B1+B2b(i,ii,iii)ENSTMXF. Iranica 90: 307–308; F. Iranica 139a: 150
77Cousinia gmelini C. Winkl.Gil., Maz., (Map 77)2600–33004AHHem.171543212.4EN B1+B2ab (i,ii)ENAMW, SMWF. Iranica 90: 121–122; F. Iranica 139a: 121
78Cousinia hypochionea Bornm.Gil., (Map 78)400–4501MRHem.DDSEMXF. Iranica 90: 316
79Cousinia rechingerae Bornm.Gol., N Khor., (Map 79)740–130011 (3)EHHem.21664488.5EN B1+B2b(i,ii)VU A4aOSCF. Iranica 90: 93–94; F. Iranica 139a: 117; (A. 14,111)
80Cousinia wendelboi Rech. f.Maz., (Map 80)20001AHHem.DDMSCF. Iranica 90: 167
81Crepis alfredi Bornm.Gil., (Map 81)3001MRHem.DDUnknownF. Iranica 122: 338
82Crepis papposissima BabcockGol., (Map 82)1EHTher.DDUnknownF. Iranica 122: 319
83Crepis willemetioides Boiss.Gol., Maz., Turk., (Map 83)50–290020 (10)CEHGeo.43,88615,00054.5NTLCLF, MSC, MWOAkhani (1998); F. Iranica 122: 302–303; (A. 16,667)
84Dolichorrhiza persica (Boiss.) B. Nord.Gil., Maz., (Map 84)2600–34004AHHem.64817.7CR B1b(i,ii)CRAZF. Iranica 164: 52–53
85Doronicum wendelboi EdmondsonGol., Maz., Sem., (Map 85)2000–26004EHHem.2499129618.1EN B1ab (i,ii)ENMFF. Iranica 164: 46
86Echinops koelzii Rech. f.Ardabil/Gil., Gol., Kho., Maz., Maz./Tehr., Sem., W Azar.?, (Map 86)500–300019 (3)AHHem.153,76627,50074.8LCLCCMF, CSF, OMF, OSF, SCAkhani (1998); F. Iranica 139a: 81; Joharchi et al. (2007); (A. 9827)
87Hieracium hoppeanum SchultesAras., Gil., (Map 87)1200–24005WHHem.6363648035.7VU B1ab(i)VUMMWF. Iranica 122: 172–173
88Hieracium kandawanicum (Rech. f. & Zahn) Rech. f.Maz./Alborz, (Map 88)2100–27003AHHem.1.280.8CR B1+B2b(i,ii)CRSZF. Iranica 122: 173–174
89Klasea quinquefolia (Willd.) Greuter & Wagenitz (= Serratula quinquefolia M. B. ex Willd.)Aras., Ardabil/Gil., Gil., Gol., Maz., Talish, W Azar., (Map 89)385–220036 (10)XHHem.647,24657,500172.2LCLCLF, MF, SFAkhani (1998); F. Iranica 139b: 298–299; Martins (2006); (A. 13,726)
90Leontodon stenocalathius Rech. f.Gol., Gol./Sem., (Map 90)2400–26002EHHem.505.3CR B1b(i,ii)CRCROF. Iranica 122: 128
91Scorzonera persica Boiss. & BuhseGil., (Map 91)1MRHem.DDSTMXF. Iranica 122: 69; (Buhse 984(Photo-G))
92Serratula gracillima Rech. f.Gol., (Map 92)1EHHem.DDUnknownF. Iranica 139b: 291–292
93Tanacetum archibaldii Podl.Maz./Alborz, (Map 93)21901AHHem.DDCROF. Iranica 158: 105
94Tragopogon gongylorrhizus Rech. f.Gol., Gol./N Khor., Maz., (Map 94)450–182015 (5)EHHem.7155821436.7VU B1b(i)VUFO, GR, SCAkhani (1998); F. Iranica 122: 98; (A. 16,225)
95Willemetia tuberosa DC. (= Calycocorsus tuberosus (DC.) RauschertGil., Gol., Maz., Talish, (Map 95)20–200048 (20)OHGeo.195,74945,00077.4LCLCLF, MHAkhani (1998); F. Iranica 122: 289–290; (A. 21,075)
Berberidaceae
96Berberis orthobotrys Bienert ex C. K. SchneiderGil., Gol., Maz., Turk., (Map 96)700–280043 (2)CEHPhan.93,38337,50059.3LCLCSCAkhani (1998); F. Iranica 111: 13; F. Iran 64: 9–12; (A. 11,625)
97Epimedium pinnatum Fisch., (Fig. 2K)Gil., Maz., Talish, (Map 97)100–170047 (9)WHGeo.29,34222,27545NTNTLF, MF, SFF. Iranica 101: 3; F. Iran 56: 4–7; (A. & al. 21,061)
Betulaceae
98Alnus subcordata C. A. Mey. [incl. var. subcordata & var. villosa (Regel) H. Winkl.]Gil., Gol., Maz., Talish, (Map 98)100–190090 (19)OHPhan.103,88645,00068LCLCLF, RH, SFAkhani (1998); (Browicz, 1982); F. Iranica 96: 7–8; (A. 11,995)
Boraginaceae
99Cynoglossum kandavanensis (Bornm. & Gauba) AkhaniAlborz, Aras., Gil., Gol., Maz., (Map 99)Sea level–236016 (8)OHHem.107,25512,50082.1LCLCFO, MMWAkhani (1998); F. Iranica 48: 141–142; F. Iran 39: 440–442; Hamzeh'ee et al., 2010; (A. 13,521)
100Echium amoenum Fisch. & C. A. Mey., (Fig. 2L)Alborz, Ardabil, E Azar.?, Gil., Gol., Maz., Talish, Tehr., (Map 100)60–3000110 (42)OHHem.172,43957,50071.6LCLCFO, MMWAkhani (1998); F. Iranica 48: 215; F. Iran 39: 184–185; Grossheim (1967); (A. & al. 20,932)
101Eritrichium gracillimum Rech. f.Gol., (Map 101)30001AHHem.DDAZF. Iranica 48: 65–66
102“Heliotropium sp. nov.”6Gil., Qazvin, (Map 102)150–8506 (6)MRTher.59.498804.2CR B1b(i,ii,iii)EN C2bSMXUnpublished
103Lepechiniella persica (Boiss.) H. RiedlGol./Sem., Maz., (Map 103)2400–39004AHHem.10,958456338.9VU B1ab (i,ii)DDAZF. Iranica 48: 81–82; F. Iran 39: 335–336
104Lepechiniella wendelboi H. RiedlMaz., Sem., Tehr., (Map 104)1000–41008AHHem.12,330576630.7VU B1ab (i,ii)VUAZ, SZF. Iranica 48: 82; F. Iran 39: 336–338
105Myosotis anomala H. RiedlAras., Ardabil ?, Gil., Maz., N Khor., (Map 105)250–245011OHHem.115,94417,50086.6LCLCLF, MF, MHF. Iranica 48: 266; F. Iran 39: 266–267; Joharchi et al. (2007)
106Nonea longiflora Wettst. ex StapfGil., (Map 106)200–5005MRTher.30.299121.9CR B1b(i,ii,iii)CRSC, STF. Iranica 48: 244; Nejhad-Falatoury et al. (2011)
107Rindera regia (Gmelin) Kusn.Gil., Maz., (Map 107)2500–27502AHHem.0.180.3CR B1+B2b(i,ii)CRAZF. Iranica 48: 127; F. Iran 39: 428
Brassicaceae
108Alyssopsis mollis (Jacq.) O. E. SchulzArdabil, Gil., Gol., Maz., N Khor., Sem., Talish, Tehr., (Map 108)400–320087 (35)OHHem.132,14252,50080LCLCLF, MF, RO, SC, VCAkhani (1998); F. Iranica 57: 200; F. Iran 143: 492–495; (A. 13,707)
109Hesperis hyrcana Bornm. & Gauba, (Fig. 2M)Aras., Gil., Gol., Maz., Cauc., (Map 109)Sea level–2750126 (33)XHHem.497,196147,500169.9LCLCCLF, FM, MF, MH, SFAkhani (1998); Dvořák (1964); F. Iranica 57: 268; F. Iran 143: 648–651; (A. 9349)
110Isatis gaubae Bornm.Gol., N Khor., Maz., Sem., Tehr., (Map 110)570–300039CEHTher.57,43237,50068.9LCLCMSCF. Iranica 57: 85; F. Iran 143: 541-543
111Noccaea umbellata (Steven ex DC.) Al-ShehbazGil., Gol., Maz., Talish, (Map 111)Sea level–193043 (2)OHTher.158,75650,00080LCLCLF, MF, SFF. Iranica 57: 113–114; F. Iran 143: 143; Grossheim (1950); (J. 1866)
112Noccaea hastulata (Steven ex DC.) KhosraviGil., Gol., Maz., Talish, (Map 112)150–250054 (4)OHTher.89,08345,00062.6LCLCFM, MF, MWF. Iranica 57: 115–116; F. Iran 143: 148–151; Grossheim (1950); (A. 13,746)
Buxaceae
113Buxus hyrcana Pojark., (Fig. 2N)Gil., Gol., Maz., Talish, (Map 113)−20–125048 (15)OHPhan.62,51740,00054.2LCNTALFF. Iranica 27: 4; F. Iran 131, 3–5; (A. 21,266)
Campanulaceae
114Campanula ghilanensis Pall.Gil., (unknown point, without Map)1WHHem.DDUnknownF. Iranica 13: 37
115Campanula lourica Boiss., (Fig. 3A)Alborz, Gol., Maz., N Khor., Sem., Tehr., (Map 114)700–340041 (8)CEHHem.34,71635,00052.7NTLCCROAkhani (1998); F. Iranica 13: 24; F. Iran 66: 50–51; (A. 11,396)
Caprifoliaceae
116Lonicera floribunda Boiss. & BuhseGil., Gol., Maz., Turk., Cauc., (Map 115)400–246072 (22)OHPhan.444,40665,000129.7LCLCSCAkhani (1998); (Browicz, 1991); F. Iranica 10: 13; F. Iran 13: 20–23; (A. 16,209)
117Lonicera iberica M. Bieb., (Fig. 3B)Ardabil, Cauc., E Azar., Gil., Gol., Maz., Qazv., Sem., Talish, Tehr., Turkey, W Azar., (Map 116)500–3000136 (5)XHPhan.535,599155,000135.1LCNTMF, JWO, RO, SASCAkhani (1998); Browicz (1988); F. Iranica 10: 9–10; F. Iran 13: 15–16; (A.11,350)
118Scabiosa schimperiana Boiss. & BuhseGil., (Map 117)300–13006MRHem.148.546755.4EN B1+B2b(i,ii,iii)ENPMXF. Iranica 168: 48–49; F. Iran 8: 91–94
Caryophyllaceae
119Cerastium microspermum C. A. Mey.Ardabil, Gil., Talish, (Map 118)500–24007WHTher.2420270426.2EN B1b(i,ii)ENMWF. Iranica 163: 94
120Dianthus agrostolepis Rech. f.Ardabil/Gil., Gil., (Map 119)420–21004WHHem.4232160020EN B1ab (i,ii)ENMFAssadi (1985); F. Iranica 163: 155
121Dianthus hyrcanicus Rech. f.Gil., Maz., (Map 120)0–22504WHHem.179.15867515.4EN B1ab (i,ii)ENLF, MF, ROF. Iranica 163: 144
122Dianthus mazanderanicus Rech. f.Maz., (Map 121)1950–27002AHHem.504.6CR B1b(i,ii)EN
C1
MSTF. Iranica 163: 154–155
123Dianthus orientalis Adams subsp. gilanicus Rech. f.Ardabil/E Azar. ?, Gil., Gil./Qazv., Maz., Sem., (Map 122)200–232010AHCham.19,20312,50053.2VU B1ab(i)LCLSC, MSC, STF. Iranica 163: 158–159
124Dianthus orientalis Adams subsp. gorganicus Rech. f., (Akhani, 2005: Figs. 560–561)Gol., Maz., Sem./Gol., (Map 123)350–260038 (15)EHCham.14,29810,89033.4VU B1b(i)LCRO, SCAkhani (1998); F. Iranica 163: 162; (A. 13,700)
125Dianthus rudbaricus AssadiGil., (Map 124)2002MRHem.0.080.2CR B1+B2b(i,ii,iii)DDSAMXF. Iranica 163: 188
126Dianthus talyschensis Boiss. & BuhseTalish, (Map 125)9001WHHem.DDLFF. Iranica 163: 143–144; (Buhse 802 a (Photo-G))
127Gypsophila modesta Bornm.Gil., (Map 126)400–5002MRTher.323.9CR B1b(i,ii,iii)CRSMXF. Iranica 163: 246
128Saponaria bodeana Boiss., (Fig. 3C)Gol., Maz., (Map 127)400–260020 (12)EHHem.11,74113,68939.3VU B1b(i)VUMFAkhani (1998); F. Iranica 163: 198; Joharchi et al. (2007); (A. 11,092)
129Silene lineata Boiss. & BuhseGil., (Map 128)1MRHem.DDUnknownF. Iranica 163: 386; (Buhse 984 (Photo-G))
130Silene schafta Gmel. Jun. ex Hohen.Gil., Gol., Maz., Talish, (Map 129)820–281033 (12)OHHem.68,79032,50059.2LCLCMSC, MST, ROF. Iranica 163: 466–467; (A. 13,660)
131Silene sojakii Melzh.Gol. (Golestan National Park)?, Sem., (Map 130)2100–28004EHCham.2180307231.8EN B1ab (i,ii)ENCROAkhani (1998); F. Iranica 163: 462; Gholipour and Sheidai (2009)
Celastraceae
132Evonymus velutina (C. A. Mey.) Fisch. & C. A. Mey., (Fig. 3D)Gol., Maz., N Khor., Talish, Turk., (Map 131)300–200048 (18)OHPhan.201,89637,50090.2LCNTOLF, OMF, OSC, ROAkhani (1998); F. Iranica 64: 2; Browicz (1986); Joharchi et al. (2007); (A. 14,121)
Crassulaceae
133Sedum lenkoranicum Grossh.Alborz, Aras., Ardabil/Gil., Gil., Maz., Sem., Talish, (Map 132)1300–250018AHHem.96,99030,00075.7LCLCMF, MSC, ROF. Iranica 72: 12; F. Iran 32: 24–26; Grossheim (1950); (A. 12,047)
134Sempervivum iranicum Bornm. & Gauba, (Akhani, 2005: Figs. 784–785)Alborz, Gil., Gol., Maz., Sem., (Map 133)1650–295025 (8)AHHem.49,69132,50057LCVU A4cSAST, SP, SWOAkhani (1998); F. Iranica 72: 4; F. Iran 32: 11–13; (A. 17,001)
Euphorbiaceae
135Euphorbia mazandaranica PahlevaniMaz., (Map 134)Sea level–14509CHHem.4289.5437525EN B1b(i,ii)VU A4LF, SFGrossheim (1962); Pahlevani and Riina (2014); Pahlevani et al. (2020)
Fabaceae
136Albizzia julibrissin Durazz., (Fig. 3E)Gil., Gol., Maz., Talish, (Map 135)Sea level–62044 (5)OH (disjunctly occurs in SE Asia)Phan.57,57530,00053.6LCNTLFBrowicz (1982); F. Iranica 161: 9; F. Iran 18: 30; (Gho. 4535)
137Astragalus acutifolius Bunge (= A. ammodendroides Podlech & Zarre)Gil., (Map 136)200–3607 (2)MRCham.0.1411.4700.7CR B1+B2b(i,ii,iii)CRSMXF. Iranica 178: 379; (Ma.&Sa. 85,976 (TARI))
138Astragalus lacus-valashti Maassoumi, Podlech & JaliliMaz., (Map 137)530–22685 (1)CHHem.94.112804CR B1b(i,ii)EN C1MSC, SMSC, WOF. Iranica 178: 343; (Ma.&Ja. 82,422 (TARI))
139Astragalus nurensis Boiss. & BuhseAlborz, Gol., Maz., Sem., (Map 138)150–300037 (11)AHHem.25,95326,46042NTNTMSC, MST, ROF. Iranica 178: 44–45; (Ma. 75,471 (TARI))
140Astragalus rosellus Širj. & Rech. f.Alborz, Gil., Gol., Maz., Sem., (Map 139)1500–275011 (2)AHCham.17,82510,08641.2VU B1ab(i)LCSASCF. Iranica 177: 103–104; (Fo. 453 (TARI))
141Astragalus vereskensis Maassoumi & PodlechGil., Gol., Maz., Sem., (Map 140)200–300033 (1)AHHem.31,52732,50053.6NTNTLMW, LSC, LST, MMW, MSC, MSTF. Iranica 174: 236–237; F. Iran 43: 185–186; (Klet 44,087 (IRAN))
142Astragalus yushensis T. Sabaii, Zarre & Podlech. (=A. submitis Boiss. & Hohen. subsp. maassoumii Tietz & Zarre)Maz., (Map 141)500–25006AHCham.62.895803.9CR B1b(i,ii)EN
C1
MSC, MST, SMSC, SMSTF. Iranica 179: 120-121
143Colutea buhsei (Boiss.) Shap.Alborz, Ardabil, Gol., Maz., N Khor., Qazv., R. Khor., Sem., Tehr., Turk., (Map 142)320–3000109 (9)CEH (also in Khorassan-Kopet Dagh)Phan.201,47565,00097.5LCLCDV, FM, OSC, ROAkhani (1998); Browicz (1986); F. Iranica 157: 72–73; (A. 16,616)
144Gleditsia caspica Desf.Gil., Gol., Maz., Talish, (Map 143)−25–39046 (8)OHPhan.63,18932,50053.6LCLCLFBrowicz (1982); F. Iranica 160: 10–11; F. Iran 45: 20–21; (A. & al. 21,030)
145Onobrychis heterophylla C. A. Mey.Talish, (Map 144)900–12007WHHem.3488450029.8EN B1b(i,ii)ENFO, SCF. Iranica 157: 437; Grossheim (1952)
146Onobrychis mazanderanica Rech. f.Alborz, Gol., Maz., (Map 145)500–240013 (1)AHHem.24,92416,20044.8NTNTFO, SCF. Iranica 157: 457; (A. 14,577)
147Trifolium mazanderanicum Rech. f.Maz., (Map 146)4001EHHem.DDFOF. Iranica 157: 313
Fagaceae
148Quercus castaneifolia C. A. Mey., (Fig. 3G)Gil., Gol., Maz., Talish, (Map 147)Sea level–2400119 (36)OHPhan.225,25067,50077.5LCLCCLF, OLF, CSF, OSF, CMF, OMF, SCAkhani (1998); Browicz (1982); F. Iranica 77: 10–11; (A. 14,580)
Gentianaceae
149Gentiana grossheimii Doluch.7, (Fig. 3H)Daghestan, Gol., (Map 148)800–230013 (2)OHHem.9253 (for Daghesten records)3249 (for Daghesten records)19 (for Daghesten records)VU B1b(i,ii) (Daghestan)CR D (Iran)LVC, CRO, OSCDolukhanof (1948); Grossheim (1967); (A. 23,291)
Geraniaceae
150Erodium dimorphum WendelboMaz., Tehr./Sem., (Map 149)2600–33002AHHem.183.5CR B1b(i,ii,iii)CRRO, AZF. Iranica 69: 50; F. Iran 62: 89–91
Hamamelidaceae
151Parrotia persica (DC.) C. A. Mey., (Fig. 3F)Gil., Gol., Maz., Talish, (Map 150)Sea level–1500102 (36)OHPhan.174,91855,00078.6LCLCALF, LF, RHAkhani (1998); Browicz (1982); F. Iranica 53: 1–2; F. Iran 136: 3–5; (A. 14,579)
Hypericaceae
152Hypericum fursei N. RobsonMaz., (Map 151)12001CHHem.CR DSCF. Iranica 49: 16
Lamiaceae
153Ballota platyloma Rech. f.Gil., Maz., Sem. Tehr., (Map 152)400–250017 (5)AHHem.9776768831VU B1b(i)VULSC, LWO, MSC, MWO, RHF. Iranica 150: 352–353; F. Iran 76: 411; (A. 13,749)
154Betonica nivea Stev. subsp. mazandarana (Bornm.) Rech. f.Maz., Tehr., (Map 153)2000–400013 (6)AHHem.5576320020.1VU B1b(i,ii)LCAZ, ROF. Iranica 150: 399–400; (Mo. 14.7.2006, s.n.)
155Eremostachys lanata JamzadMaz., (Map 154)1900–23502 (2)AHHem.CR C2bMSTJamzad (1987); (As.& M. 33,028 (holotype- TARI))
156Lagochilus quadridentatus JamzadMaz., (Map 155)1900–23002 (1)AHHem.DDMSTJamzad (1988); (As.& M. 33,030 (holotype- TARI))
157Nepeta pogonosperma Jamzad & AssadiMaz., Zanjan ?, (Map 156)2900–36006 (1)AHHem.4780450030.3EN B1b(i,ii)ENAZ, ROJamzad and Assadi (1984); Jamzad (1991); (R.&Ma. 21,689 (holotype- TARI))
158Phlomis ghilanensis C. KochGil., (Map 157)1AHHem.DDAZF. Iranica 150: 301–302
159Salvia oligophylla Auch. ex Benth.Gil., Qazv., Zanjan, (Map 158)450–147012MRHem.1629137214.4EN B1b(i,ii,iii)ENMXF. Iranica 150: 466; F. Iran 76: 915
160Salvia shahkuhmahalei AkhaniGol./N Khor., (Map 159)1500–19356EHHem.30.01461.3CR B1+B2b(i,ii)CRSCAkhani et al. (2016)
161Satureja intermedia C. A. Mey.Gil., Talish, (Map 160)1900–23507 (1)WHHem.426.77543212.3EN B1+B2b(i,ii)ENROF. Iranica 150: 498; F. Iran 76: 675
162Satureja isophylla Rech. f.Maz., (Map 161)1600–330013 (1)CHHem.12135768EN B1b(i,ii)ENMF, MSC, MST,F. Iranica 150: 498; F. Iran 76: 687
163Satureja mutica Fisch. & C. A. Mey.Gil., Gol., Maz., N Khor., Talish, Turk., (Map 162)550–208021 (8)OHHem.174,54127,50089LCLCRO, VCAkhani (1998); F. Iranica 150: 499; F. Iran 76: 694; (A. 12,263)
164Scutellaria tournefortii Benth., (Fig. 3I)Gil., Gol., Maz., Talish, (Map 163)20–213086 (11)OHHem.124,96270,00068LCLCCLF, CMFAkhani (1998); F. Iranica 150: 52–53; F. Iran 76: 92; Grossheim (1967); (A. 9278)
165Stachys laxa Boiss. & BuhseGol., Maz., Sem., Tehr., (Map 164)500–281043 (8)CEHHem.36,95629,58442.7NTLCROAkhani (1998); F. Iranica 150: 389–390; Salmaki et al. (2012); (A. 12,326)
166Stachys talyschensis KapellerTalish, (Map 165)1WHHem.DDSPF. Iranica 150: 379; Grossheim (1967)
167Teucrium hyrcanicum L.Gil., Gol., Maz., Talish, (Map 166)−20–200097 (30)XHHem.495,070115,000149.1LCLCLF, MF, OSC, WOAkhani (1998); F. Iranica 150: 36–37; Grossheim (1967); (A. 11,960)
Linaceae
168Linum bungei Boiss.Gol., Maz., Sem., (Map 167)2100–320014 (3)AHHem.27,04415,12941.1NTNTMMW, MSC, MSTF. Iranica 106: 12; F. Iran 34: 29; (Furse 2769 (IRAN))
Lythraceae
169Peplis hyrcanica Sosn.Talish, (Map 168)1WHTher.DDMHF. Iranica 51: 6–7
Malvaceae
170Alcea gorganica (Rech. f., Aell. & Esfand.) Zohary [incl. A. popovii Iljin & A. sycophylla Iljin & Nikitin, A. mazandaranica Pakravan & Ghahreman]Gol., Maz., N Khor., W Azar.?, Turk., (Map 169)100–205035 (12)CEHHem.123,61421,16046.2LCLCFM, FO, OSC, WOAkhani (1998); F. Iranica 120: 72, 76–77, 83; F. Iran 58: 77–80; (A. 11,197)
171Alcea hyrcana (Grossh.) Grossh.Gil., Maz., Talish, (Map 170)100–21005WHHem.20,362544533NTNTLF, MFF. Iranica 120: 57; F. Iran 58: 108–110
172Tilia platyphyllos Scop. subsp. caucasica (Rupr.) Loria9, (Fig. 3J)Gil., Gol., Maz., Talish, (Map 171)100–2400141 (4)XHPhan.1,452,474217,500252.9LCNTCLF, CMFAkhani (1998); F. Iranica 148: 6–7
Orobanchaceae
173Orobanche eriophora Bornm. & Gauba10Maz., (Map 172)23001AHGeo.DDMSC?F. Iranica 5: 10
174Orobanche schwingenschussii GilliMaz./Alborz, (Map 173)28602AHGeo.0.080.2CR B1+B2b(i,ii)DDMSC?F. Iranica 5: 10; F. Iran 86: 50-51
175Rhynchocorys maxima C. Richter, (Fig. 3K)Alborz, Gil., Gol., Maz., Talish, Tehr., (Map 174)−26–275070 (35)OHHem.185,77162,50088.8LCLCDH, FM, FO, LF, MF,Akhani (1998); Burbidge and Richardson (1970); F. Iranica 147: 209; (A. & al. 20,929)
Paeoniaceae
176Paeonia tomentosa (Lomak.) N. Busch11, (Fig. 3L)Gil., Gol., Maz., Talish, (Map 175)1000–300056 (6)OHGeo.119,30840,00068.8LCLCCMF, FM, FO, SCAkhani (1998); F. Iranica 60: 4; F. Iran 137: 4–6; Hong and Zhou (2003); Rukšãns and Zetterlund (2014); (A. 10,310)
177Paeonia wendelboi Rukšāns & ZetterlundArdabil-Gil., (Map 176)2000–23005WHGeo.9.72731CR B1+B2b(i,ii,iii)CRSMW, RORukšāns and Zetterlund (2014)
Papaveraceae
178Corydalis chionophila Czernjak.Gol., N Khor., R. Khor., Turk., (Map 177)1650–234012 (11)EH KHORASSAN-KOPET DAGH (Memariani)Geo.32,75712,94343.3NTNTMMW, WOAkhani (1998); F. Iranica 110: 20; (A. 10,203)
179Corydalis hyrcana WendelboArdabil ?, Gol., Maz., (Map 178)1000–24505 (3)OHGeo.32,704750060NTVU A4cMMW, WOF. Iranica 110: 14–15; (J. 1590)
180Papaver chelidoniifolium Boiss & Buhse, (Fig. 3M)Gil., Maz., Talish, (Map 179)−22-180049 (2)OHTher.57,91939,76247.2LCLCFM, LF, SC, SFF. Iranica 34: 19; F. Iran 127: 99–102; (A. 5335)
Plantaginaceae
181Digitalis nervosa Steud. & Hochst. ex Benth., (Fig. 3N)Aras.,Ardabil/Gil., Gil., Gol., Maz., Sem., Talish, (Map 180)200–320088 (32)OHHem.262,97475,00093.6LCLCFM, FO, LF, MF, RO, SCAkhani (1998); F. Iranica 147: 169–170; F. Iran 68: 152; Grossheim (1967); (A. 13,704)
182Plantago podlechii AkhaniGol. (Golestan National Park), (Map 181)1600–17002 (2)EHHem.0.980.7CR B1+B2b(i,ii)CRMST, OSCAkhani (1998); (A. 17,084)
183Veronica bungei Boiss.Maz./Sem., (Map 182)Ca. 23001EHTher.DDMFF. Iranica 147: 97–98; (Bunge 15 (holotype- Photo-G-Boiss.))
184Veronica euphrasiifolia LinkGil. (unknown point, without map)1AHHem.DDAZF. Iranica 147: 124; (Willdenow 203 (type-B-Willd.))
185Veronica francispetae M. A. FischerGil., Gol., Maz., (Map 183)20–12009OHTher.15,78910,64739.1VU B1ab(i)VULF, SFF. Iranica 147: 104–105; Saeidi-Mehrvarz (1999)
186Veronica gaubae Bornm.Gil., Gol., Maz., Khor. Sem., Tehr., Turk., (Map 184)2000–320035 (1)AHTher.103,50237,50084.5LCLCMF, MST, ROAkhani (1998); F. Iranica 147: 112–113; F. Iran 68: 115; (A. 12,036)
187Veronica mazanderanae WendelboArdabil, Gil., Gol., Maz., (Map 185)520–261913 (4)AHTher.89,21217,50063.9LCLCLST, MSTF. Iranica 147: 78–79; Saeidi-Mehrvarz (1999); (J. 1779)
188Veronica paederotae Boiss.Maz., Tehr., (Map 186)2700–41007 (1)AHHem.10084869.1EN B1+B2 b (i,ii)ENAZ, MST, SPF. Iranica 147: 144–145; Saeidi-Mehrvarz (1999); (N. 942)
189Veronica siaretensis LehmannGol., Maz., Talish, (Map 187)400–150017 (3)OHTher.84,72215,00062.6LCLCLF, MFAkhani (1998); F. Iranica 147: 98–99; Saeidi-Mehrvarz (1999); (A. 10,403)
Polygalaceae
190Polygala platyptera Bornm. & GaubaMaz., (Map 188)Sea level–250042 (4)CHHem.19,83311,76028.4VU B1b(i)LCLF, MF, SF,F. Iranica 124: 7–8; F. Iran 49: 15–17; (Abai 12,645 (IRAN))
Polygonaceae
191Atraphaxis aucheri Jaub. & SpachGil., (Map 189)300–6004MRCham.55.222273.2CR B1b(i,ii,iii)CRSTMXF. Iranica 56: 35
192Atraphaxis radkanensis S.Tavakkoli, Kaz.Osaloo & Mozaff.Gol., (Map 190)1400–17623EHCham.11.35931.1CR B1+B2b(i,ii)CRCRO, MROTavakkoli et al. (2013); (A. 23,283)
193Polygonum hyrcanicum Rech. f., (Akhani, 2005: Figs. 747–748)Ardabil, Gil., Gol., Maz., N Khor., Qazv., Sem., (Map 191)10–175033 (11)OHHem.179,48340,00090.3LCLCDH, LMW, MMWAkhani (1998); F. Iranica 56: 64–65; (A. 12,307)
194Rumex kandavanicus (Rech. f.) Rech. f.Maz./Alborz, (Map 192)28001AHHem.DDMH, SZF. Iranica 56: 9–10
Primulaceae
195Cortusa matthioli L. subsp. iranica Iranshahr & WendelboMaz., (Map 193)2500–30002 (1)AHHem.CR C2bMF?F. Iran 25: 59; Iranshahr and Wendelbo (1976); (Re.&Ir. 16,804 (type-IRAN))
196Cyclamen elegans Boiss. & Buhse (= C. coum auct. Flora Iranica 9: 30, 1965 non Miller)Gil., Gol., Maz., Talish, (Map 194)50–175033 (7)OHGeo.62,66335,00055.3LCLCCLF, CSFClennett (2002); F. Iranica 9: 30–31; F. Iran 25: 67–69; Grossheim (1967); (A. 13,795)
197Dionysia aretioides (Lehm.) Boiss.Gil., Maz., Sem., Tehr., (Map 195)550–320028 (4)AHCham./Hem.964410,80029.7VU B1b(i)LCVCF. Iranica 9: 17; F. Iran 25: 25; (A. 13,757)
198Primula heterochroma Stapf, (Fig. 4A)Gil., Gol., Maz., Sem. Talish, (Map 196)60–240055 (23)OHHem.115,74850,00066.9LCLCCLF, CMF, CSFAkhani (1998); F. Iranica 9: 5; F. Iran 25: 11–13; (A. & al. 21,062)
Ranunculaceae
199Aconitum iranshahrii H. RiedlMaz., (Map 197)2000–25001 (1)CHGeo.EN A4cMFF. Iranica 171: 38; (Ir.&Re. 16,765 (IRAN))
200Anemone caucasica Willd. ex Rupr., (Fig. 4B)Cauc., Central Alborz, E Anatolia, Gil., Gol., Maz., Sem., Talish, (Map 198)700–290053 (9)XHGeo.378,69687,500152.4LCLCGOWO, LFAkhani (1998); F. Iranica 171: 224–225; Grossheim (1950); (A. 14,573)
201Delphinium syncarpum FreynMaz., (Map 199)1CHTher.DDCSD ?F. Iranica 171: 85–86
202Delphinium ursinum Rech. f., (Akhani, 2005: Figs. 519–520)Alborz, Gol., Maz., (Map 200)700–220016 (3)CEHGeo.37,16525,34448.4NTNTAZ, FO, MWOAkhani (1998); F. Iranica 171: 70–71; (A. 11,676)
203Paraquilegia caespitosa (Boiss. & Hohen.) Drumm. & Hutch.Maz., (Map 201)2500–480010AHHem.3096264620.7EN B1b(i,ii)ENAZ, ROF. Iranica 171: 8
204Ranunculus cicutarius Schlechtend., (Akhani, 2005: Figs. 522–523)Gil., Gol., Maz., Khor.?, Sem., Talish, (Map 202)−26–220055 (4)OHHem.377,29172,500104.7LCLCFM, LF, MF, MH, OSC, SFAkhani (1998); F. Iranica 171: 159–160; Grossheim (1950); (A. 10,347)
205Ranunculus dolosus Fisch. & C. A. Mey.Gil., Gol., Maz., Talish, (Map 203)−26–120013 (1)OHGeo./Hem.92,64127,50064.3LCLCMHF. Iranica 171:137; Grossheim (1950); (A. 13,274)
Rhamnaceae
206Frangula grandifolia (Fisch. & C. A. Mey.) GrubovGil., Gol., Maz., Talish, (Map 204)200–130033 (8)OHPhan.96,70335,00066.8LCLCLFBrowicz (1984); F. Iranica 125: 26–27; F. Iran 55: 19; (A.20,271)
Rosaceae
207Alchemilla amardica Rothm.Gil. (unknown point), Maz., (Map 205)2000–29004AHHem.13222409EN B1+B2ab (i,ii)ENAMW, SMWF. Iranica 66: 145–146; F. Iran 6: 176; Naqinezhad et al. (2010)
208Alchemilla citrina FröhnerArdabil, Gil., Gol., Maz., (Map 206)2000–320010OHHem.43,81412,50060.2NTNTAMW, MF, SMWF. Iranica 66: 142–143; F. Iran 6: 173–174; Naqinezhad et al. (2010)
209Alchemilla condensa FröhnerGil., Maz., (Map 207)1800–21004OHHem.9899692441.7VU B1ab (i,ii)VUMMWF. Iranica 66: 143–144; F. Iran 6: 175
210Alchemilla gigantodus FröhnerAras., Gil., Gol., Maz., (Map 208)1300–34009OHHem.57,83112,50072.7LCLCAZ, SZF. Iranica 66: 141–142; F. Iran 6: 171–172; Naqinezhad et al. (2010)
211Alchemilla hyrcana (Buser) Juz.Aras., Gil., Maz., Talish, (Map 209)1200–36009WHHem.38,23220,00077.3NTNTAMW, SMWF. Iranica 66: 137–138; F. Iran 6: 162–163; Grossheim (1952); Hamzeh'ee et al., 2010
212Alchemilla mazandarana Naqinezhad & S.E. FröhnerMaz., (Map 210)2636–29507AHHem.303.290755.4EN B1+B2b(i,ii)ENRO, MMW, GOWONaqinezhad et al. (2017)
213Alchemilla microscopica FröhnerGil., Maz., (Map 211)1800–23004OHHem.20,958661847NTNTMMWF. Iranica 66: 138–139; F. Iran 6: 167–168
214Alchemilla pectiniloba FröhnerGil., Maz., (Map 212)1800–32005WHHem.1085235228.4EN B1ab (i,ii)ENAZ, SZF. Iranica 66: 139; F. Iran 6: 168–169
215Alchemilla plicatissima FröhnerGil., Gol., Maz., (Map 213)1800–26004OHHem.5827750052.3VU B1ab (i,ii)VUMMWF. Iranica 66: 143; F. Iran 6: 174-175
216Cydonia oblonga MillerAras., Gil., Gol., Maz., Talish, Turk., (Map 214)40–170053 (2)XH (widely cultivated in other parts)Phan.2,350,436107,500123.5LCVU A4acCLF, CSF, OLF, OSFAkhani (1998); Browicz (1996); F. Iranica 66: 26–27; F. Iran 6: 180–181; Grossheim (1952); (A. 13,671)
217Potentilla petraea Willd. ex Schlecht.Alborz, Ardabil/Gil., Gil., Maz., Talish, (Map 215)2300–310010WHHem.12,196713537.8VU B1ab(i)NTRO, SZF. Iranica 66: 107–108; F. Iran 6: 135–136; Grossheim (1952)
218Pyrus boissieriana Buhse (Pro. Syn. P. ghahremanii Attar & Zamani, P.longipedicellata Zamani & Attar, P. kandevanica Ghahreman & Khatamsaz, P. cordifolia Zamani & Attar)15, (Fig. 4C and D)Gil., Gol., Maz., Sem./Gol., Talish, Turk., (Map 216)400–240057 (16)OHPhan.183,34355,00070LCLCCMF, MST, OMF, SC, ROAkhani (1998); Browicz (1982); F. Iranica 66: 29; F. Iran 6: 183–185; (A. 12,317)
219Pyrus grossheimii Fedor.Gil., Maz., Talish, (Map 217)700–24008WHPhan.9285950739.9VU B1ab (i,ii)NTMF, MSC, SF, SMSCF. Iranica 66: 30; F. Iran 6: 186–188; Grossheim (1952)
220Pyrus hyrcana Fedor.Aras., Gil., Maz., Talish, (Map 218)40–245011WHPhan.28,40115,00050.3NTNTLF, LSC, LWO, MF, MSC, MWOF. Iranica 66: 30–31; F. Iran 6: 188; Grossheim (1952)
221Pyrus mazanderanica Schönbeck-TemesyAlborz, Maz., (Map 219)1900–24005CHPhan.324.4CR B1b(i,ii)DDMSCF. Iranica 66: 32; F. Iran 6: 194
222Rubus persicus Boiss. (= R. hyrcanus Juz.)Gil., Gol., Maz., Talish, (Map 220)30–160029OHPhan.164,04132,50072.7LCLCDH, FM, LF, RH, SFF. Iranica 66: 70–71; F. Iran 6: 30–31; Grossheim (1952); Zieliński (1978)
223Rubus raddeanus FockeAras., Gil., Gol., Maz., Talish, (Map 221)50–200014 (1)OHPhan.97,30822,50079.1LCLCLF, RHAkhani (1998); F. Iranica 66: 70–71; F. Iran 6: 28–30; Grossheim (1952); Zieliński (1978); (A. 11,822)
224Spiraea sheikhii ZareMaz., (Map 222)200–2502CHPhan.DDLFZare (2002)
Rubiaceae
225Asperula gorganica Schönb.-Tem. & Ehrend.Gol., (Map 223)600–208015 (8)EH±Cham.4136225916.8EN B1b(i,ii)VU A4aCRO, SC, VC, WOAkhani (1998); F. Iranica 176: 149; (A. 11,143)
226Asperula mazanderanica Ehrend.Gol., Maz., (Map 224)650–27009EHCham./Hem.6015240020.1VU B1ab (i,ii)VUMSC, RO,F. Iranica 176: 148
227Asperula microphylla Boiss.Gil., Gol., Maz., (Map 225)300–340016 (1)AHCham./Hem.14,58410,08641.4VU B1b(i)VUCRO, WOF. Iranica 176: 147–148; (J. 1723a)
228Asperula sherardioides (Boiss.) Jaub. & SpachGil., (Map 226)400–6002MRTher.CR C2bSMXF. Iranica 176: 155–156; (Aucher 4675 (type-Photo-G))
229Crucianella gilanica Trin. subsp. hirsuta (Ehrend.) Ehrend. & Schönb.-Tem.Alborz, Maz., Tehr., (Map 227)850–410019AHHem.4891284.4EN B1+B2b(i,ii)VU A4cAZ, CRO, MF, MSC, SRO, VROF. Iranica 176: 85–86
230Crucianella gilanica Trin. subsp. nezvensis Ehrend. & Schönb.-Tem.Maz., Sem., (Map 228)2000–32005AHHem.202362.7EN B1+B2ab (i,ii)ENMF, RO, SASC, SZF. Iranica 176: 83–84
231Crucianella gilanica Trin. subsp. suleimanica Ehrend. & Schönb.-Tem.Maz., (Map 229)2000–35202AHHem.81.8CR B1+B2b(i,ii)EN A4cAMW, ROF. Iranica 176: 84
232Crucianella platyphylla Ehrend. & Schönb.-Tem.Gol., Sem., (Map 230)1800–25006 (5)EHCham./Hem.3959162017.6EN B1b(i,ii)VU A4LVC, MWOAkhani (1998); F. Iranica 176: 79; (A. 12,026)
233Galium aucheri Boiss.Gol., Maz., Sem., (Map 231)3000–41008AHHem.6971684537.3VU B1ab (i,ii)NTAZ, CRO, SPF. Iranica 176: 192
234Galium caspicum StevenGil., Maz., Talish, (Map 232)700–18507WHHem.26,349921648NTNTMH, SFF. Iranica 176: 198
235Galium elbursense Bornm. & Gauba ex Bornm.Gil., Maz., (Map 233)1700–18002AHCham./Hem.64818.1CR B1b(i,ii)CRRO, SC, SFF. Iranica 176: 229–230
236Galium kuetzingii Boiss. & BuhseGil., Talish, (Map 234)250–4005WHCham./Hem.796.529172824.5EN B1ab (i,ii)ENSAMXF. Iranica 176: 204–205
237Galium wendelboi Ehrend. & Schönb.-Tem.Maz., (Map 235)450–18004 (1)CHHem.21.1CR B1+B2b(i,ii)CRMF, ROF. Iranica 176: 228–229; (A. 13,858)
238Phuopsis stylosa (Trin.) Hook. f., (Fig. 4E)Gil., Gil./Ardabil, Maz., Talish, (Map 236)200–264041 (19)WHGeo.34,77924,29946.8NTLCFO, MW, SCF. Iranica 176: 73–74; (Gho. 4489)
Salicaceae
239Populus caspica Bornm., (Fig. 4F)Aras., Gil., Gol., Maz., N Khor., Talish, Turk., (Map 237)300–140032 (15)OH (widely cultivated outside of Caspian area)Phan.1,852,14285,00085.1LCLCALF, LF, RHAkhani (1998); Browicz (1982); F. Iranica 65: 10–11; (A. 11,554)
Sapindaceae
240Acer iranicum M. Mohtashamian & A. Rastegar/A. mazandaranicum Amini, Zare & Assadi 17Gil., (Map 238)950–11505WHPhan.0.1230.0380.1CR B1+B2b(i,ii)DDCSFAmini et al. (2008); Mohtashamian et al. (2020)
241Acer velutinum Boiss. [incl. var. velutinum & var. glabrescens (Boiss. & Buhse) E. Murray], (Fig. 4G)Gil., Gol., Gol./N Khor., Maz., Talish, (Map 239)0–236468 (31)OHPhan.235,49462,500100.4LCLCCLF, CMF, CSF, RHAkhani (1998); Browicz (1982); F. Iranica 61: 4–5; (A. 16,614)
Saxifragaceae
242Saxifraga iranica Bornm.Maz., (Map 240)2800–42008AHCham./Hem.568.867124.5035EN B1+B2b(i,ii)ENAZ, ROF. Iranica 42: 13; F. Iran 12: 10–11
243Saxifraga mazanderanica Rech. f.Gil., Maz., (Map 241)1100–30009 (1)AHCham./Hem.8482530032.6VU B1ab(i)EN C1ROF. Iranica 42: 10; F. Iran 12: 16–18; (Ma. 38,628 (IRAN))
244Saxifraga ramsarica JamzadMaz., (Map 242)27502 (1)AHCham./Hem.0.720.6CR B1+B2b(i,ii)CRCROF. Iran 12: 11–12; (R.&Ma. 20,903 (holotype-TARI))
245Saxifraga wendelboi Schönb.-Tem., (Fig. 4H)Gol., Maz., Sem., (Map 243)1900–30009 (4)EHCham./Hem.1879137213.9EN B1b(i,ii)VU A4VCF. Iranica 42: 12; F. Iran 12: 9; (A. 13,611)
Scrophulariaceae
246Scrophularia gaubae Bornm.Gil., Gol., Maz., Khor., (Map 244)400–300029 (8)CEHHem.54,11932,50064.1LCLCCLF, CMF, CSF, FO, RH SCAkhani (1998); F. Iranica 147: 242–243; F. Iran 68: 286; Joharchi et al. (2007); (A. 17,098)
247Scrophularia megalantha Rech. f.Maz., (Map 245)40–60017CHHem.3459123910.6EN B1b(i,ii)ENLFF. Iranica 147: 231; F. Iran 68: 225
248Scrophularia rostrata Boiss. & BuhseGil., Gil./Ardabil, Maz., Talish, (Map 246)50–301023 (1)WHHem.36,27324,95647.7NTLCLF, MF, SC, SF, SZF. Iranica 147: 241–242; F. Iran 68: 289; Grossheim (1967); (A. & al. 20,933)
249Scrophularia vernalis L. subsp. clausii (Boiss. & Buhse) Grau, (Fig. 4I)Gil., Gol., Maz., Sem., Talish, W Azar.?, (Map 247)400–264041 (3)OHHem.220,92245,00078.5LCLCMF, RO, SCF. Iranica 147: 229–230; F. Iran 68: 219; Grossheim (1967); (A. 15,016)
250Verbascum parsana Sotoodeh, Attar & CiveyrelGil., (Map 248)18761WHHem. (Bien.)DDOMFSotoodeh et al. (2016); (Moazzeni & Keshvari 34,941 (holotype-TUH))
251Verbascum shahsavarensis Sotoodeh, Attar & CiveyrelMaz., (Map 249)15701CHHem. (Bien. Peren.)EN C2bOMFSotoodeh et al. (2015)
252Verbascum stachydiforme Boiss. & BuhseGil./Ardabil, Maz., Talish, (Map 250)1200–230010 (1)WHHem.26,10810,00050.9NTVU A4cMF, WOF. Iranica 147: 43; F. Iran 68: 62; (Manutshehri 561)
253Verbascum sublobatum Murb.Gil., Gol., Maz., N Khor., (Map 251)500–210015 (4)CEHHem.51,66917,50059LCVU A4cFO, LF, MF, SC, SF, WOAkhani (1998); F. Iranica 147: 42; F. Iran 68: 59; (A. 16,323)
Solanaceae
254Atropa pallidiflora Schönbeck-TemesyGol., Maz., (Map 252)35–250024CEHHem.30,20023,04048.1NTLCLF, MF, SF,F. Iranica 100: 41–42; F. Iran 24: 54–56
255Solanum kieseritzkii C. A. Mey., (Fig. 4J)Gil., Gol., Maz., Talish, (Map 253)90–240025 (8)OHCham.105,43232,50067.5LCLCCLF, CMF, CSFAkhani (1998); F. Iranica 100: 13; F. Iran 24: 15; Grossheim (1967); (A. & al. 21,294)
Thymelaeaceae
256Daphne mezereum L. subsp. rechingeri (Wendelbo) HaldaGil., Maz., (Map 254)500–17508 (1)WHPhan.12,109571033.8VU B1ab (i,ii)VULF, SC, SFF. Iranica 95: 2–3; F. Iran 15: 4–6; (Sa.&Ga. 42,162 (IRAN))

Appendix 2. 

Taxonomic notes including doubtful or excluded species

Apiaceae

  • 1.

    Bupleurum gilanicum Mozaff. Iran. J. Bot. 21 (1): 83. 2015. The type specimen was not available for evaluation. The original description shows its close affinity with the highly polymorphic complex B. exaltatum B. Bieb.

  • 2.

    Peucedanum pimenovii Mozaff. Bot. Zhurn. 88 (4): 116. 2003. Mozaffarian (2003) described P. pimenovii from Gorgan: Between Tuskestan and Chaharbagh, at the beginning of Bagheshah gorge of Pakotal, ca. 940 m, 30.09.1997, V. Mozaffarian 77,840 (TARI!). Peucedanum glaucopruinosum Rech. f. was described from Mazandaran: Robate Qozloq (Gozlu), in silvis, Koelz 16,206 (Isotype W!) (Rechinger, 1952). Both localities are indeed the same place (http://www.iranvillage.ir/article-print-8957.html). Akhani's study of the type specimens of both names and field studies in the area confirmed that there is no difference between them. P. glaucopruinosum was transferred to the genus Leutea by Akhani and Salimian (Yassa et al., 2003).

  • 3.

    Leutea translucens (Rech.f.) Akhani & Salimian comb. nov. Basionym: Peucedanum translucens Rech.f. Anz. Math.-Nat. Kl. Osterr. Akad. Wiss. 89: 243 (1952) = P. hyrcanicum Golizadeh, Naqinezhad & Mozaff. Ann. Bot. Fenn. 54: 292. 2017. syn. nov.

  • Specimens examined: Golestan: In declivibus borealibus M. Shahvar prope Hajilang, 2400–2600 m, in juniperetis saxosis, K. H. Rechinger 6154 (W); Shahvar mountain, between Tash and Siahmarzkuh, ca. 5 km S. Siahmarzkuh, northern steep scree slope, with open Juniperus sabina community, 2750 m, 36°38′N, 54°44′E, 15.7.2001, H. Akhani & M. Salimian 15,359 (Hb. Akh.); Northern slopes of Shahvar mountain, ca. 2 km from Siahmarzkuh towards Kholindarreh, deep valley along the road, steep-cliffs, 1750 m, 36°39′27″N, 54°45′18″E, 15.7.2001. H. Akhani & M. Salimian 15,362 (Hb. Akh.); ca. 30 km SES of Gorgan, ca. 10 km NE Chaharbagh, Pir-Gerdeh Kuh, 36°39′19″N, 54°34′35″E, rocky steep to subvertical N-facing slopes, ca. 2500–2600 m, 10.9.1999, H. Akhani 13,711 (Hb. Akh.).

  • L. translucens was not accounted by Pimenov in Flora Iranica (Pimenov, 1987) and therefore was treated in its original description by Rechinger from the type locality. We were succeeded to rediscover this plant from area of its locus classicus (Akhani & Salimian 15,359). Information on fruit morphology, fruit anatomy and pollen morphology (Salimian, 2002) and ITS sequences (Akhani, unpublished data) clearly confirmed its position within the genus Leutea as recently accepted by Panahi et al. (2015). The mericarps in L. transulens are elliptic, 9-11 X 4–5 mm, rounded at apex and base having slightly prominent dorsal ribs, vittae solitary in valleculae occupying nearly whole of vallecular region, mesocarp cells above vittae absent, vittae in wings absent, commissural vittae 2, small, located between wing and raphe. The pollen grains are 3-zonocolporate and distinctly rhombic typical for the genus Leutea (Salimian, 2002).

  • The newly described Peucedanum hyrcanicum from Mazandaran: Sangdeh, Kheru-Neru mountain, 35°59′2.6″N, 53°12′48.4″E, 2700 m a.s.l., 10.08.2016, H. Gholizadeh & A. Naqinezhad 7610 (holotype HUMZ; isotype TARI, n.v.) by Gholizadeh et al. (2017) was collected from 155 km aerial distance. Although the type specimen was not seen by the authors, checking the description, illustration and data on habitat and locality remain no doubt that P. hyrcanicum is identical with L. translucens.

Asteraceae

  • 4.

    Artemisia kulbadica Boiss. & Buhse. Nouv. Mém. Soc. Imp. Naturalistes Moscou 12: 120. 1860.

  • This species was described from Gorgan: In pratis ad mare Caspium prope Kulbad non procul ab urbe, Buhse (Photo available in http://www.villege.ch/musinfo/bd/cjb/chg/adetail.php?id=274814&base=img&lang=en). The illustration of the type and own field observations in the type locality revealed that this is indeed a synonym of A. absinthium L., commonly growing in the area.

Betulaceae

  • 5.

    Alnus djavanshirii Zare and A. dolichocarpa Zare, Amini & Assadi have been described from Mazandaran province as new species from N Iran (Zare and Amini, 2012). Both species are doubtful and require future studies.

Boraginaceae

  • 6.

    Heliotropium sp. nov. This species is a local endemic belonging to H. dissitiflorum Boiss. complex. The species will be named and validated in an on-going publication.

Gentianaceae

  • 7.

    Gentiana grossheimii Doluch. Zametki Sist. Geogr. Rast. 14: 51. 1948. Fig. 3H.

  • Examined specimen: Iran: Golestan province, Jahan Nama Protected Area, ca. 2–3 km E of Jahan Nama Protection Station, Chekele Shahpasand, uppermost parts of cliffs, 36°40′12″N, 54°19′54″E, 2044–2067 m.a.s.l., H. Akhani 23,291 (Hb. Akh.).

  • This is a new report from Iran. This chasmophytic species belongs to Gentiana septemfida Pall. complex but evidently differs not only by its habitat but also with a number of characters including smaller habit (up to 20 cm tall, not up to 40 cm), smaller leaves (8–15 mm, not 20–35 (−50) mm), shorter corolla (29–35 mm long, not 35–45 mm long), the presence of yellowish spots on corolla and flowering time in autumn (not in summer).

  • In spite of our intention to describe this as a new species, comparison of Iranian collection with the type specimen and other identical specimens of G. grossheimii in LE clearly proved their extreme similarity and an interesting disjunction between east Hyrcanian forests and Dagestan in the Caucasus (Supplemental file 2: Map 148).

Geraniaceae

  • 8.

    Geranium montanum Hablitz ex Pall. in Neue Nord. Beytr. Phys. Geogr. Erd-Völkerbeschreib. 4: 51. 1784. was described from N Iran, “In alpib. Samamisius”, Aug, Hablitz (LE). This is recorded as a synonym of G. ibericum Cav. which its distribution concentrated on the eastern parts of the Black Sea and the Caucasus (Aedo, 2005).

Malvaceae

  • 9.

    The genus Tilia in Iran was simultaneously reviewed by two working groups (Yousefzadeh et al., 2012; Zare et al., 2012). Rather surprising is that M. Assadi is the co-author of both papers and both papers have been accepted in less than a month of interval (26.01.2012 and 21.02.2012). Following species are reported for Iran: T. begonifolia Steven, T. dasystyla Steven (wrongly cited T. dastyla in the whole paper), T. rubra DC. subsp. caucasica (Rupr.) V. Engler and T. hyrcana Tabari and Colagar as a new endemic Hyrcanian species (Yousefzadeh et al., 2012). In Zare et al. (2012) T. cordata Mill., T. dasystyla, T. rubra subsp. caucasica, T. begonifolia and two new species Tilia sabetii H. Zare and Tilia stellato-pilosa H. Zare, Amini & Assadi were recorded. We prefer to keep the nomenclature of Flora Iranica by accepting only Tilia platyphyllos Scop. subsp. caucasica (Rupr.) Loria in this paper before future studies prove reliability of these names (Browicz, 1981).

Orobanchaceae

  • 10.

    The taxonomic status of species including Orobanche eriophora Bornm. & Gauba, O. bungeana G. Beck, O. caucasica G. Beck and O. schwingenschussii Gilli needs detailed investigation (Gilli, 1979; Iranshahr, 2008). Recently Saeidi Mehrvarz et al. (2010) pointed out that these are acceptable species.

Paeoniaceae

  • 11.

    Paeonia tomentosa (Lomak.) N. Busch

  • The name of P. tomentosa is replaced the frequently used P. wittmanniana Stev. in most of botanical references before 2014, when Rukšãns and Zetterlund (2014) provided evidences regarding to correct name of the Iranian plants.

Plantaginaceae

  • 12.

    Linaria golestanensis Hamdi & Assadi and L. mazandaranensis Hamdi & Assadi described from Golestan and Mazandaran provinces, respectively (Hamdi et al., 2006) are closely related or conspecific with L. dalmatica (L.) Miller and L. grandiflora Desf. Further studies are required for a convincing decision (Hamdi et al., 2006).

Polygonaceae

  • 13.

    Polygonum lencoranicum Komar. is synonymous of P. litorale Meisn., a species with larger distribution range (Grossheim, 1945).

Rosaceae

  • 14.

    Type specimen of Cotoneaster assadii Khatamsaz seen in TARI herbarium (Khatamsaz & Assadi 43,257-b!) seems to be a variant of C. ovatus Pojark. with orange fruits.

  • 15.

    Pyrus kandevanica Ghahreman & Khatamsaz, P. ghahremanii Attar & Zamani, P. longipedicellata Zamani & Attar and P. cordifolia Zamani & Attar have been described from near localities “ca. 20 km after Gachsar towards Chalus and Siahbisheh/ca. 32 km after Gachsar to Chalus” (Khatamsaz, 1991; Zamani et al., 2009, 2016; Zamani and Attar, 2010). Several times we have tried to study the type specimens of them in TUH herbarium but our request was rejected by the curator of the herbarium who is the corresponding author of three latter papers. According to descriptions, it seems these are either morphotypes of the variable ‘P. boissieriana’ or hybrids.

Salicaceae

  • 16.

    The diagnosis of Salix baladehensis Maassoumi, Moeeni & Rahiminejad separated from S. aegyptiaca L. is not convincing for a new species (Maassoumi et al., 2008).

Sapindaceae

Appendix A. Supplementary data

The following are the Supplementary data to this article:

Multimedia component 1:
Click here to view.(79K, pdf)Multimedia component 1
Multimedia component 2:
Click here to view.(5.2M, pdf)Multimedia component 2

References

  • Aedo C. (1680) Proposal to conserve the name Geranium ibericum against G. montanum (Geraniaceae) with a conserved type. Taxon. 2005;54:207–208. [Google Scholar]
  • Akhani H. Studies on the flora and vegetation of the Golestan National Park, NE Iran, I: a new species and some new plant records. Ann. Nat. Hist. Mus. Wien. 1996;98B(Suppl. l):97–105. [Google Scholar]
  • Akhani H. Plant biodiversity of golestan national Park, Iran. Stapfia. 1998;53:1–411. [Google Scholar]
  • Akhani H. Studies on the flora and vegetation of the Golestan National Park, NE Iran III. Three new species, one new subspecies and fifteen new records for Iran. Edinb. J. Bot. 1999;56:1–31. [Google Scholar]
  • Akhani H. vol. 1. Tehran University Press; Tehran: 2005. (The Illustrated Flora of Golestan National Park, Iran). [Google Scholar]
  • Akhani H., Djamali M., Ghorbanalizadeh A., et al. Plant biodiversity of Hyrcanian relict forests, N Iran: an overview of the flora, vegetation, palaeoecology and conservation. Pakistan J. Bot. 2010;42:231–258. [Google Scholar]
  • Akhani H., Khoshravesh R., Malekmohammadi M. Taxonomic novelties from Irano-Turanian region and NE Iran: Oreosalsola, a new segregate from Salsola s.l., two new species in Anabasis and Salvia, and two new combinations in Caroxylon and Seseli. Phytotaxa. 2016;249:159–180. [Google Scholar]
  • Akhani H., Mahdavi P., Noroozi J., et al. Vegetation patterns of the Irano-Turanian steppe along a 3,000 m altitudinal gradient in the Alborz Mountains of northern Iran. Folia Geobot. 2013;48:229–255. [Google Scholar]
  • Akhani H., Scholz H. Studies on the flora and vegetation of the Golestan National Park, NE Iran II: a new Poa and some new and noteworthy grass records for Iran. Edinb. J. Bot. 1998;55:443–453. [Google Scholar]
  • Akhani H., Ziegler H. Photosynthetic pathways and habitats of grasses in Golestan National Park (NE Iran), with an emphasis on the C4-grass dominated rock communities. Phytocoenologia. 2002;32:455–501. [Google Scholar]
  • Amini T., Zare H., Assadi M. Acer mazandaranicum (Aceraceae), a new species from northern Iran. Iran. J. Bot. 2008;14:81–86. [Google Scholar]
  • Anonymous . Azerbaijan E.C.O. & Michael Succow Stiftung zum Schutz der Natur; Greifswald: 2010. [Google Scholar]
  • Assadi M. The genus Dianthus L. (Caryophyllaceae) in Iran. Iran. J. Bot. 1985;3:9–54. [Google Scholar]
  • Assadi M. Plants of Arasbaran protected area, NW. Iran (Part I). Iran. J. Bot. 1987;3:129–175. [Google Scholar]
  • Assadi M. Plants of Arasbaran protected area, NW Iran. (Part II). Iran. J. Bot. 1988;4:1–59. [Google Scholar]
  • Assadi M., Maassoumi A.A., Khatamsaz M., Mozaffarian V., editors. Flora of Iran. Ministry of Jihad-e-Agriculture, Research Institute of Forests and Rangelands; Tehran: 1992-2016. [Google Scholar]
  • Bachman S., Moat J., Hill A.W., et al. Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. Zookeys. 2011;150:117–126. [Europe PMC free article] [Abstract] [Google Scholar]
  • Bidarlord M., Ghahremaninejad F. Ornithogalum boissieri (Asparagaceae), a new species from the Talesh mountains, Iran. Ann. Bot. Fenn. 2016;53:69–72. [Google Scholar]
  • Binka K., Nitychoruk J., Dzierzek J. Parrotia persica CAM (Persian witch hazel, Persian ironwood) in the Mazovian (Holsteinian) Interglacial of Poland. Grana. 2003;42:227–233. [Google Scholar]
  • Browicz K. In: Flora Iranica. Rechinger K.H., editor. Graz; Austria: 1981. [Google Scholar]
  • Browicz K. Institute of Dendrology; Poznan: 1982. [Google Scholar]
  • Browicz K. vol. 2. Polish Academy of Sciences, Institute of Dendrology; Poznan: 1983. (Chorology of Trees and Shrubs in South-West Asia and Adjacent Regions). [Google Scholar]
  • Browicz K. vol. 3. Polish Academy of Sciences, Institute of Dendrology; Poznan: 1984. (Chorology of Trees and Shrubs in South-West Asia and Adjacent Regions). [Google Scholar]
  • Browicz K. Institute of Dendrology; Poznan: 1986. [Google Scholar]
  • Browicz K. vol. 6. Polish Academy of Sciences, Institute of Dendrology; Poznan: 1988. (Chorology of Trees and Shrubs in South-West Asia and Adjacent Regions). [Google Scholar]
  • Browicz K. Chorology of the Euxinian and hyrcanian element in the woody flora of Asia. Plant Systemat. Evol. 1989;162:305–314. [Google Scholar]
  • Browicz K. vol. 8. Polish Academy of Sciences, Institute of Dendrology; Poznan: 1991. (Chorology of Trees and Shrubs in South-West Asia and Adjacent Regions). [Google Scholar]
  • Browicz K. vol. 9. Polish Academy of Sciences, Institute of Dendrology; Poznan: 1992. (Chorology of Trees and Shrubs in South-West Asia and Adjacent Regions). [Google Scholar]
  • Browicz K. Institute of Dendrology; Poznan: 1994. [Google Scholar]
  • Browicz K. Academy of Sciences, Institute of Dendrology; Poznan: 1996. [Google Scholar]
  • Browicz K., Zieliński J. vol. 4. Polish Academy of Sciences, Institute of Dendrology; Poznan: 1984. (Chorology of Trees and Shrubs in South-West Asia and Adjacent Regions). [Google Scholar]
  • Burbidge R.B., Richardson J.B.K. A revision of the genus Rhynchocorys. Notes R. Bot. Gard. Edinb. 1970;30:97–107. [Google Scholar]
  • Byng J.W., Chase M.W., Christenhusz M.J.M., et al. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Bot. J. Linn. Soc. 2016;181:1–20. [Google Scholar]
  • Callmander M.W., Schatz G.E., Lowry P.P. IUCN red list assessment and the global strategy for plant conservation: taxonomists must act now. Taxon. 2005;54:1047–1050. [Google Scholar]
  • Christenhusz M.J.M., Govaerts R., David J.C., et al. Tiptoe through the tulips – cultural history, molecular phylogenetics and classification of Tulipa (Liliaceae) Bot. J. Linn. Soc. 2013;172:280–328. [Google Scholar]
  • Clennett J.C.B. An analysis and revision of Cyclamen L. with emphasis on subgenus Gyrophoebe O. Schwarz. Bot. J. Linn. Soc. 2002;138:473–481. [Google Scholar]
  • Djamali M., Akhani H., Khoshravesh R., et al. Application of the Global Bioclimatic Classification to Iran: implications for understanding the modern vegetation and biogeography. Ecol. Medit. 2011;37:91–114. [Google Scholar]
  • Djamali M., Brewer S., Breckle S.W., et al. Climatic determinism in phytogeographic regionalization: a test from the Irano-Turanian region, SW and Central Asia. Flora. 2012;207:237–249. [Google Scholar]
  • Dolukhanof A. De Gentianis Caucasicis e Subsectione Septemfidae Kusn. nutulae criticae. Notulae Systematicae ae Geographicae Instituti Botanici Tphilisiensis. 1948;14:38–60. [Google Scholar]
  • Dvořák F. Some remarks on Hesperis series Matronales in Caucasia and Transcaucasia. Phyton (Horn) 1964;11:93–101. [Google Scholar]
  • Edmondson J.R. In: Flora of Turkey and the East Aegean Islands. Davis P.H., editor. Edinburgh University Press; Endinburgh: 1985. pp. 470–486. [Google Scholar]
  • Ehrendorfer F., Schönbeck-Temesy E. In: Rechinger W., editor. Naturhistorisches Museum Wien; Wien: 2005. pp. 76–105. (Flora Iranica). [Google Scholar]
  • Ferguson D.K., Knobloch E. A fresh look at the rich assemblage from the Pliocene sink-hole of Willershausen, Germany. Rev. Palaeobot. Palyno. 1998;101:271–286. [Google Scholar]
  • Frey W., Kürschner H., Probst W. In: Yarshater E., editor. The Encyclopædia Iranica Foundation, Inc.; New York: 1999. pp. 43–63. (Encyclopaedia Iranica). [Google Scholar]
  • Frey W., Probst W. In: Contributions to the Vegetation of Southwest Asia. Kürschner H., editor. Dr. Ludwig Reichert; Wiesbaden: 1986. pp. 9–43. [Google Scholar]
  • Fritsch R.M., Abbasi M. IPK Gatersleben; 2013. p. 244. [Google Scholar]
  • Gholipour A., Sheidai M. A new record and some rediscovered endemic Silene (Caryophyllaceae) species in Iran. Rostaniha. 2009;10:212–220. (In Persian, with English Abstract 299–100) [Google Scholar]
  • Gholizadeh H., Mehrvarz S.S., Naqinezhad A., et al. Peucedanum hyrcanicum (Apiaceae), a new species of Peucedanum s. l. from northern Iran. Ann. Bot. Fenn. BioOne. 2017:291–296. [Google Scholar]
  • Gholizadeh H., Naqinezhad A., Chytrý M. Classification of the hyrcanian forest vegetation, northern Iran. Appl. Veg. Sci. 2020;23:107–126. [Google Scholar]
  • Gilli A. Die Orobanchaceen der 'Flora Iranica' (Additamenta ad floram iranicam 2.) Candollea. 1979;34:279–305. [Google Scholar]
  • Gölz P., Gämperle R., Gügel E., et al. Über die Orchideenflora des Iran. J. Eur. Orch. 2006;38:79–104. [Google Scholar]
  • Gölz P., Gügel E., Wagner R., et al. Über die Orchideenflora des Iran (2. Teil) und Turkmenistan. J. Eur. Orch. 2007;39:297–322. [Google Scholar]
  • Green A.F., Ramsey T.S., Ramsey J. Phylogeny and biogeography of Ivies (Hedera spp., Araliaceae), a polyploid complex of woody vines. Syst. Bot. 2011;36:1114–1127. [Google Scholar]
  • Grossheim A.A. vol. I. VII. Nauk; Leningrad: 1939-1967. (Flora Kavkaza). [Google Scholar]
  • Hamdi S.M.M., Assadi M., Fallahian F., et al. Linaria mazandaranensis and L. golestanensis (Scrophulariaceae), two new species from Iran. Iran. J. Bot. 2006;11:251–258. [Google Scholar]
  • Hamzeh’ee B., Safavi S.R., Asri Y., et al. Floristic analysis and a preliminary vegetation description of Arasbarann Biosphere Reserve. Rostaniha. 2010;11:1–16. [Google Scholar]
  • Hedge I.C., Wendelbo P. Patterns of distribution and endemism in Iran. Notes Roy. Bot. Gard. Edind. 1978;36:441–464. [Google Scholar]
  • Hong D.-Y., Zhou S.-L. Paeonia (Paeoniaceae) in the Caucasus. Bot. J. Linn. Soc. 2003;143:135–150. [Google Scholar]
  • Iranshahr M. Parasitic and semiparasitic flowering plants of Iran, Volume 2. Rostaniha. 2008;9:1–79. (Persian) [Google Scholar]
  • Iranshahr M., Wendelbo P. A new subspecies of Cortusa matthioli (Primulaceae) from N. Iran. Iran. J. Bot. 1976;1:57–60. [Google Scholar]
  • IUCN . 2019. [Google Scholar]
  • Iversen J. Viscum, Hedera and Ilex as climate indicators. Förh. Geol. Fören. Stockholm. 1944;66(3):463–483. [Google Scholar]
  • Jafari A., Maassoumi A.A. Synopsis of Leopoldia, Muscari and Pseudomuscari (Hyacinthaceae) in Iran, with Leopoldia ghouschtchiensis sp. nova. Ann. Bot. Fenn. 2011;48:396–400. [Google Scholar]
  • Jafari S.M., Akhani H. Plants of Jahan Nama protected area, golestan province, N. Iran. Pakistan J. Bot. 2008;40:1533–1554. [Google Scholar]
  • Jamzad Z. Eremostachys lanata and Mentha mozaffarianii, two new Labiatae from Iran. Iran. J. Bot. 1987;3:111–116. [Google Scholar]
  • Jamzad Z. The genus Lagochilus (Labiatae) in Iran. Iran. J. Bot. 1988;4:91–103. [Google Scholar]
  • Jamzad Z. Nepeta menthoides Boiss. & Buhse and species allied to it in Iran. Iran. J. Bot. 1991;5:17–27. [Google Scholar]
  • Jamzad Z., Assadi M. New species of the genera Nepeta and Ajuga (Labiatae) from Iran. Iran. J. Bot. 1984;2:95–102. [Google Scholar]
  • Jalili A., Jamad Z. Research Institute of Forests and Rangelands; 1999. [Google Scholar]
  • Joharchi M., Ghahremaninejad F., Vitek E. New plant records for Khorassan province, Iran. Ann. Naturhist. Mus. Wien. B. 2007;108:277–301. [Google Scholar]
  • Khatamsaz M. Two new species of the Rosaceae from Iran. Iran. J. Bot. 1991;5:1–5. [Google Scholar]
  • Khoshravesh R., Akhani H., Eskandari M., et al. Fern and fern allies of Iran. Rostaniha. 2009;10:1–130. [Google Scholar]
  • Klein J.C. Institute Français de Recherche en Iran; Téhéran: 1994. [Google Scholar]
  • Leroy S.A.G., Roiron P. Latest Pliocene pollen and leaf floras from Brenasso paleolake (Escandorgue Massif, Hérault, France) Rev. Palaeobot. Palyno. 1996;94:295–328. [Google Scholar]
  • Li J., Del Tredici P. The Chinese Parrotia: a sibling species of the Persian Parrotia. Arnoldia. 2008;66:2–9. [Google Scholar]
  • Maassoumi A., Moeeni F., Rahiminejad M.R. New species and new records of the genus Salix (Salicaceae) from Iran. Iran. J. Bot. 2008;14:1–6. [Google Scholar]
  • Mahdavi P., Akhani H., Van der Maarel E. Species diversity and life form patterns in steppe vegetation along a 3000 m altitudinal gradient in the Alborz Mountains, Iran. Folia Geobot. 2013;48:7–22. [Google Scholar]
  • Manafzadeh S., Salvo G., Conti E. A tale of migrations from east to west: the Irano-Turanian floristic region as a source of Mediterranean xerophytes. J. Biogeogr. 2014;41:366–379. [Google Scholar]
  • Manafzadeh S., Staedler Y.M., Conti E. Visions of the past and dreams of the future in the Orient: the Irano-Turanian region from classical botany to evolutionary studies. Biol. Rev. 2017;92:1365–1388. [Abstract] [Google Scholar]
  • Manen J.F., Barriera G., Loizeau P.A., et al. The history of extant Ilex species (Aquifoliaceae): evidence of hybridization within a Miocene radiation. Molec. Phylogen. Evol. 2010;57:961–977. [Abstract] [Google Scholar]
  • Martins L. Systematics and biogeography of Klasea (Asteraceae-Cardueae) and a synopsis of the genus. Bot. J. Linn. Soc. 2006;152:435–464. [Google Scholar]
  • Mathew B. A taxonomic and horticultural review of Erythronium L (Liliaceae) Bot. J. Linn. Soc. 1992;109:453–471. [Google Scholar]
  • Memariani F., Akhani H., Joharchi M.R. Endemic plants of Khorassan-Kopet Dagh floristic province in Irano-Turanian region: diversity, distribution patterns and conservation status. Phytotaxa. 2016;249:31–117. [Google Scholar]
  • Meusel H., Jäger E., Weinert E. Gustav Fischer Verlag Jena; 1965. [Google Scholar]
  • Moat J. GIS Unit, Royal Botanic Gardens; Kew: 2007. http://www.rbgkew.org.uk/gis/cats Available at: [Google Scholar]
  • Mohtashamian M., Chatrenoor T., Fatehi F., et al. Acer iranicum (Sapindaceae), a new species of Maple from northern Iran. Syst. Bot. 2020;45:163–172. [Google Scholar]
  • Morton A. The author; Berkshire: 2001. [Google Scholar]
  • Mozaffarian M. New species and new records of Iranian Umbelliferae. Bot. Zhurn. 2003;88:104–124. [Google Scholar]
  • Mucina L., Bültmann H., Dierßen K., et al. Vegetation of Europe: hierarchical floristic classification system of vascular plant, bryophyte, lichen, and algal communities. Appl. Veg. Sci. 2016;19:3–264. [Google Scholar]
  • Naqinezhad A., Frohner S.E., Esmailpoor A. Alchemilla mazandarana (Rosaceae), a new species from high mountainous areas of the Hyrcanian relict region, N. Iran. Phytotaxa. 2017;331:93–100. [Google Scholar]
  • Naqinezhad A., Hosseini S., Rajamand M.A., et al. A floristic study on Mazibon and Sibon protected forests, Ramsar, across the altitudinal gradient (300-2300 m) Taxon. Biosyst. 2010;5:93–114. of Per. pages (Per.), 117 of Eng. pages (Eng. Abs.) [Google Scholar]
  • Nath J., Nielsen E.L. Cytology of Phleum sp. affin. P. montanum C. KOCH. Euphytica. 1963;12:161–166. [Google Scholar]
  • Naqinezhad A., Zare-Maivan H., Gholizadeh H. A floristic survey of the Hyrcanian forests in Northern Iran, using two lowland-mountain transects. J. For. Res. 2015;26:187–199. [Google Scholar]
  • Nejhad-Falatoury A., Pakravan M., Tavassoli A. A new species and some notes on the genus Nonea (Boraginaceae) in Iran. Feddes Repert. 2011;122:425–432. [Google Scholar]
  • Noroozi J., Akhani H. The alpine plant diversity and vegetation of Tuchal Mountains (N. Tehran, Iran) Colloq. Phytosociol. 2013;29:431–449. [Google Scholar]
  • Noroozi J., Akhani H., Breckle S.W. Biodiversity and phytogeography of the alpine flora of Iran. Biodiv. Cons. 2008;17:493–521. [Google Scholar]
  • Noroozi J., Moser D., Essl F. Diversity, distribution, ecology and description rates of alpine endemic plant species from Iranian mountains. Alp. Bot. 2016;126:1–9. [Google Scholar]
  • Pahlevani A.H., Liede-Schumann S., Akhani H. Diversity, distribution, endemism and conservation status of Euphorbia (Euphorbiaceae) in SW Asia and adjacent countries. Plant Systemat. Evol. 2020;306:1–26. [Google Scholar]
  • Pahlevani A.H., Riina R. Synopsis of Euphorbia subgen. Esula sect. Helioscopia (Euphorbiaceae) in Iran with the description of Euphorbia mazandaranica sp. nov. Nord. J. Bot. 2014;32:257–278. [Google Scholar]
  • Panahi M., Banasiak L., Piwczynski M., et al. Phylogenetic relationships among Dorema, Ferula and Leutea (Apiaceae: scandiceae: Ferulinae) inferred from nrDNA ITS and cpDNA noncoding sequences. Taxon. 2015;64:770–783. [Google Scholar]
  • Pimenov M.G. In: Flora Iranica. Rechinger K.H., editor. Graz; Austria: 1987. pp. 457–462. [Umbelliferae] [Google Scholar]
  • Probst W. In: Beiträge zur Umweltgeschichte des Vorderen Orients. Frey W., Uerpman H.-P.H., editors. Dr. Ludwig Reichert; Wiesbaden: 1981. pp. 26–39. [Google Scholar]
  • Rechinger K.H. Umbelliferae novae iranicae, II. (Rechingeri iter iranicum secundum No. 25) Anzeiger der mathematisch-naturwissenschaftlichen Klasse der Österreichischen Akademie der Wissenschaften. 1952;89:195–204. [Google Scholar]
  • Rechinger K.H. Akademsiche Druck- u. Verlagsanstalt und Naturhistorisches Museum Wien, Graz & Wien; 1963-2015. [Google Scholar]
  • Renz J. In: Flora Iranica. Rechinger K.H., editor. Graz; Austria: 1978. [Google Scholar]
  • Rudov A., Mashkour M., Djamali M., Akhani H. A review of C4 plants in Southwest Asia: an ecological, geographical and taxonomical analysis of a region with high diversity of C4 eudicots. Front. Plant Sci. 2020;11:546518. 10.3389/fpls.2020.546518. [Europe PMC free article] [Abstract] [CrossRef] [Google Scholar]
  • Rukšãns J., Zetterlund H. An Iranian peony to honour Per Wendelbo. Alp. Gard. 2014;82:230–237. [Google Scholar]
  • Saeidi-Mehrvarz S. University of Tehran; Tehran: 1999. [Google Scholar]
  • Saeidi-Mehrvarz S., Torabi A., Aghabeigi F. Notes on the genus Orobanche (Orobanchaceae) in Iran. Iran. J. Bot. 2010;16:107–113. [Google Scholar]
  • Salimian M. University of Tehran; 2002. [Google Scholar]
  • Salmaki Y., Zarre S., Govaerts R., et al. A taxonomic revision of the genus Stachys (Lamiaceae: Lamioideae) in Iran. Bot. J. Linn. Soc. 2012;170:573–617. [Google Scholar]
  • Schönbeck-Temesy E., Ehrendorfer F. The perennial taxa of Crucianella (Rubiaceae) in SW. Asia and their eco-geographical differentiation. Plant Systemat. Evol. 1989;165:101–136. [Google Scholar]
  • Sotoodeh A., Attar F., Civeyrel L. Verbascum shahsavarensis (Scrophulariaceae), a new species for Flora of Iran. Phytotaxa. 2015;203:76–80. [Google Scholar]
  • Sotoodeh A., Attar F., Civeyrel L. A new species of Verbascum L. (Scrophulariaceae) from the Gilan province (Iran), based on morphological and molecular evidences. Adansonia. 2016;38:127–132. [Google Scholar]
  • Speta F. Systematic analysis of the genus Scilla L. s.l. (Hyacinthaceae) (Gem.) Phyton (Horn) 1998;38:1–141. [Google Scholar]
  • Takhtajan A. University of California Press; California: 1986. [Google Scholar]
  • Tavakkoli S., Osaloo S.K., Mozaffarian V., et al. Atraphaxis radkanensis (Polygonaceae), a new species from Iran. Ann. Bot. Fenn. 2013;50:372–374. [Google Scholar]
  • Toro-Nunez O., Al-Shehbaz I.A., Mort M.E. Phylogenetic study with nuclear and chloroplast data and ecological niche reveals Atacama (Brassicaceae), a new monotypic genus endemic from the Andes of the Atacama Desert, Chile. Plant Systemat. Evol. 2015;301:1377–1396. [Google Scholar]
  • Tralau H. Asiatic Dicotyledonous affinities in the Cainozoic flora of Europe Kungl. Svenska Vetenskapsakad. Handl. 1963;4 Ser. 9, 3, 1–87 + 5 plates. [Google Scholar]
  • Tzvelev N.N. A new species of the genus Festuca (Poaceae) from Iran. Bot. Zhurn. 1997;82:118–119. [Google Scholar]
  • Valcárcel V., Guzman B., Medina N.G., et al. Phylogenetic and paleobotanical evidence for late Miocene diversification of the Tertiary subtropical lineage of ivies (Hedera L., Araliaceae) BMC Evol. Biol. 2017;17:146. 10.1186/s12862-017-0984-1. [Europe PMC free article] [Abstract] [CrossRef] [Google Scholar]
  • Valiejo-Roman C.M., Terentieva E.I., Samigullin T.H., et al. Molecular data (nrITS-sequencing) reveal relationships among Iranian endemic taxa of the Umbelliferae. Feddes Repert. 2006;177:367–388. [Google Scholar]
  • Van Laere K., Hermans D., Leus L., et al. Genetic relationships in European and Asiatic Buxus species based on AFLP markers, genome sizes and chromosome numbers. Plant Systemat. Evol. 2011;293:1–11. [Google Scholar]
  • Vargas P., McAllister H.A., Morton C., et al. Polyploid speciation in Hedera (Araliaceae): phylogenetic and biogeographic insights based on chromosome counts and ITS sequences. Plant Systemat. Evol. 1999;219:165–179. [Google Scholar]
  • Warwick S.I., Mummenhoff K., Sauder C.A., et al. Closing the gaps: phylogenetic relationships in the Brassicaceae based on DNA sequence data of nuclear ribosomal ITS region. Plant Systemat. Evol. 2010;285:209–232. [Google Scholar]
  • White F., Leonard J. Phytogeographical links between Africa and southwest Asia. Flora Veg. Mundi. 1991;9:229–246. [Google Scholar]
  • Yassa N., Akhani H., Aqaahmadi M., et al. Essential oils from two endemic species of Apiaceae from Iran. Z. Naturforsch. C J. Biosci. 2003;58:459–463. [Abstract] [Google Scholar]
  • Yousefzadeh H., Hosseinzadeh Colagar A., Tabari M., et al. Utility of ITS region sequence and structure for molecular identification of Tilia species from Hyrcanian forests, Iran. Plant Systemat. Evol. 2012;298:947–961. [Google Scholar]
  • Zamani A., Attar F. Pyrus longipedicellata sp. nov. (Rosaceae) from central Alborz, Iran. Nord. J. Bot. 2010;28:484–486. [Google Scholar]
  • Zamani A., Attar F., Civeyrel L., et al. Pyrus cordifolia sp nov and two new records of Pyrus (Rosaceae) from Iran. Nord. J. Bot. 2016;34:739–743. [Google Scholar]
  • Zamani A., Attar F., Joharchi M.R. Pyrus ghahremanii spec. nova and P. giffanica spec. nova (Rosaceae-Maloideae) from Iran. Phyton (Horn) 2009;49:105–115. [Google Scholar]
  • Zare H. Spiraea sheikhii (Rosaceae), a new species from Iran. Iran. J. Bot. 2002;9:253–255. [Google Scholar]
  • Zare H., Amini Eshkevari T., Assadi M. A review of the genus Tilia L.(Tiliaceae) in Iran, new records and new species. Iran. J. Bot. 2012;18:175–190. [Google Scholar]
  • Zare H., Amini T. A review of the genus Alnus Gaertn. in Iran, new records and new species. Iran. J. Bot. 2012;18:10–21. [Google Scholar]
  • Zieliński J. New data on the taxonomy and distribution of species of the genus Rubus L. in Iran. Fragm. Florist. Geobot. 1978;24:427–437. [Google Scholar]
  • Zohary M. Gustav Fische Verlag; Stuttgart, Amsterdam: 1973. [Google Scholar]

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