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

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.

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.

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.

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?

Material and methods

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).

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:

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.

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.

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

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.

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.

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).

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. 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 categories	No. of taxa	Percent (%)
CR	30	11.7
EN	52	20.3
VU	30	11.7
NT	25	9.8
LC	81	31.6
No category/DD	38	14.8
Total	256	99.9

Fig. 8

Richness map of all Hyrcanian endemic and near endemic taxa.

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 8).

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).

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).

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).

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).

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.

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).

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.

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.

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).

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).

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.

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.

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.

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.

No.	Family and species (Figure number)	Distribution (Map number)	Elev. range (m)	Rec. (Herb.)	Chorotype	Life form	EOO (km2)	AOO (km2)	10% max. dist. (km)	Threat category		Habitat type	References and (selected voucher)
										GeoCat	Imp. ass.
Magnoliids and Monocots
Aristolochiaceae
1	Aristolochia hyrcana Davis & M. S. Khan, (Fig. 2A)	Gil., (Map 1)	200–1100	7 (4)	MR	Geo.	406	150	4.8	EN B1+B2b (i,ii,iii)	EN	SWMX	F. Iranica 26: 1–2; F. Iran 29: 4–6; (A. & al. 20,943)
Amaryllidaceae
2	Allium grande Lipsky (= A. chelotum Wendelbo), (Akhani, 2005: Figs. 171–172)	Gol., Maz., Sem./Gol. (Map 2)	1000–3100	19 (8)	CEH	Geo.	26,409	17,672	46.9	NT	VU C2b	MSC, SMSC	Akhani (1998); F. Iranica 76: 87–88; Fritsch and Abbasi (2013); (A. 11,305)
3	Allium lenkoranicum Miscz., (Akhani, 2005: Fig. 180)	Gol., Maz., Talish, Turk., (Map 3)	600–1600	20 (5)	OH	Geo.	127,732	17,500	80.4	LC	LC	CSF, LMW	Akhani (1998); F. Iranica 76: 61; Grossheim (1940); (A. 17,172)
4	Allium subvineale Wendelbo, (Akhani, 2005: Fig. 186)	Gil., Gol., Maz., (Map 4)	1900–2060	4 (2)	AH	Geo.	26,938	7500	54.8	NT	EN C2b	SMW	Akhani (1998); F. Iranica 76:53; (A. 11,311)
5	Allium talyschense Miscz.	Talish, (Map 5)	1800–2400	4	WH	Geo.	1817	3844	31.1	EN B1ab (i,ii)	EN	RO	F. Iranica 76: 56–57; Grossheim (1940)
Asparagaceae
6	Danaë racemosa (L.) Moench, (Akhani, 2005: Figs. 211–212)	Gil., Gol., Maz., Talish, (Map 6)	50–1923	69 (15)	OH (disjunctly in East Mediterranean)	Phan.	59,964	60,000	179.3	LC	NT	CLF, CSF, RO, VC	Akhani (1998); Browicz (1988); F. Iranica 165: 177; (A. 13,387)
7	Fessia gorganica (Speta) Speta (=Scilla gorganica Speta), (Akhani, 2005: Fig. 225)	Gol., Maz., (Map 7)	100–2472	14 (5)	EH	Geo.	9657	13,690	36.6	VU B1b(i,ii)	LC	LF, MF, SC, SF	Akhani (1998); F. Iranica 165: 114–115; Speta (1998); (A. 13,393)
8	Fessia greilhuberi (Speta) Speta (=Scilla greilhuberi Speta)	Gil., Maz., (Map 8)	50–1150	13 (3)	WH	Geo.	36,869	26,411	48.7	NT	NT	LF	F. Iranica 165: 113; Speta (1998); (A. 14,566)
9	Fessia hohenackeri (Fisch. & C.A.Mey.) Speta (=Scilla hohenackeri Fisch. & C. A. Mey.)	Talish, (Map 9)	–	9	WH	Geo.	3996	3087	21.3	EN B1b(i,ii)	EN	LF (Possibly)	F. Iranica 165: 112–113; Grossheim (1940); Speta (1998)
10	Hyacinthella persica (Boiss. & Buhse) Chouard	Gil., Gil./Qazv., (Map 10)	390–700	4	MR	Geo.	4.173	12	2.2	CR B1b(i,ii,iii)	CR	SMX	F. Iranica 165: 137–138
11	Ornithogalum boissieri Bidarlord & F. Ghahrem.	Ardabil/Gil., (Map 11)	2400	1	AH	Geo.					DD	MMW	Bidarlord and Ghahremaninejad (2016)
12	Ornithogalum bungei Boiss., (Akhani, 2005: Figs. 221–223)	Gol., Maz., Sem./Gol., Talish, (Map 12)	150–2500	26 (5)	OH	Geo.	95,808	27,500	72.1	LC	LC	CMF, FM	Akhani (1998); F. Iranica 165: 124–125; Grossheim (1940); (A. 14,570)
13	Ornithogalum sintenisii Freyn, (Akhani, 2005: Fig. 224)	Gil., Gol., Maz., Talish, (Map 13)	50–2452	56 (8)	OH	Geo.	294,976	60,000	97.2	LC	LC	LF, MWO, RO	Akhani (1998); F. Iranica 165: 129; Grossheim (1940); (A. 13,943)
14	Polygonatum glaberrimum C. Koch	Gol., Maz., (Map 14)	800–2200	62	XH	Geo.	511,431	127,500	171.8	LC	LC	LF, MF, SF	F. Iranica 165: 179; Grossheim (1940)
15	Pseudomuscari chalusicum (DC. Stuart) Garbari (= Muscari pseudomuscari (Boiss. & Buhse) Wendelbo)	Gil. (without point in map), Gol., Maz., Sem./Gol., (Map 15)	1100–1800	6 (1)	CEH	Geo.	2953	3468	34.5	EN B1b(i,ii)	EN	GSC, RO	F. Iranica 165: 148; Jafari and Maassoumi (2011); (A. 17,505)
16	Ruscus hyrcanus Woron., (Fig. 2B)	Gil., Gol., Maz., Talish, (Map 16)	0–1200	70 (25)	OH	Cham.	90,256	50,000	58.4	LC	LC	LF, SF	F. Iranica 165: 178; (A. 14,139)
Iridaceae
17	Crocus almehensis Brickell & Mathew, (Akhani, 2005: Fig. 226)	Gol. (Golestan National Park), (Map 17)	1470–2000	5 (2)	EH	Geo.	79.127	3	1.5	CR B1+B2ab (i,ii)	CR	MMW, SAST	Akhani (1998); F. Iranica 112: 5; F. Iran 31: 6–7; (A. & Sh. 6107)
18	Crocus caspius Fisch. & C. A. Mey. (Fig. 2C)	Gil., Gol., Maz., Talish, (Map 18)	−20–1300	50 (21)	OH	Geo.	69,078	45,000	53.6	LC	LC	LF	F. Iranica 112: 8 & 75; F. Iran 31: 9–10; (A. 13,774)
19	Crocus gilanicus Mathew	Gil., Maz., (Map 19)	1500–2400	4	WH	Geo.	4063	2352	27.9	EN B1ab (i,ii)	EN	SZ	F. Iranica 112: 11 & 75; F. Iran 31: 16
Liliaceae
20	Erythronium caucasicum Woron.	Gil., Maz., (Map 20)	600–1400	17	XH	Geo.	220,636	40,000	146.5	LC	NT	LF, SF	F. Iranica 165: 103; Grossheim (1940); Mathew (1992)
21	Fritillaria kotschyana Herbert [incl. subsp. kotschyana & subsp. grandiflora (Grossh.) Rix], (Akhani, 2005: Figs. 279–280)	E Azar.?, Gil., Gol., Maz., Talish, Tehr., (Map 21)	1000–3700	41 (19)	AH	Geo.	129,527	25,000	77.8	LC	LC	AMW, MF	Akhani (1998); F. Iranica 165: 72; (A. 13,386)
22	Lilium ledebourii (Baker) Boiss., (Fig. 2D)	Ardabil, Gil., Maz., Talish, (Map 22)	1350–2250	9 (4)	WH	Geo.	10,238	4900	34.6	VU B1ab (i,ii,iii)	EN C1	FM, SGR	F. Iranica 165: 58–59; (T. & D. 30,061 (IRAN))
23	Tulipa harazensis Rech. f.	Maz., (Map 23)	700	1	AH	Geo.					DD	ST	Christenhusz et al. (2013); F. Iranica 165: 91–92
24	Tulipa ulophylla Wendelbo	Gol./Sem., Maz., (Map 24)	600–2500	3	AH	Geo.		1250	25	CR B1b(i,ii,iii)	DD	SC	F. Iranica 165: 95–96
Orchidaceae
25	Cephalanthera caucasica Kränzl., (Fig. 2E)	Aras., Gil., Gol., Maz., Talish, (Map 25)	50–2003	48 (7)	OH	Geo.	280,761	55,000	92.6	LC	LC	CLF, CMF	Akhani (1998); F. Iranica 126: 29–31; F. Iran 57: 15–16; Grossheim (1940); Hamzeh'ee et al., 2010; (A. 11,253)
26	Epipactis rechingeri Renz, (Akhani, 2005: Figs. 253–254)	Gol., Maz., (Map 26)	950–2200	15 (4)	CEH	Geo.	27,703	14,175	45.5	NT	VU C1	MF, SF	Akhani (1998); F. Iranica 126: 36–37; F. Iran 57: 19–20; (A. 11,982)
27	Himantoglossum formosum (Stev.) C. Koch	Talish, (Map 27)	150	9	WH	Geo.	22,076	7776	35.6	NT	EN C1	LF, MF	F. Iranica 126: 90–91; Grossheim (1940)
28	Ophrys × aghemani Renz (Ophrys scolopax × transhyrcana)	Gol., (Map 28)	1250	1	EH	Geo.					CRD	MW	F. Iranica 126: 86
29	Ophrys kojurensis Gölz	Maz., (Map 29)	650–1020	7	CH	Geo.		50	5.4	CR B1b(i,ii,iii)	CR	GSC	Gölz et al. (2006); Gölz et al. (2007)
30	Ophrys sintenisii Fleischamn. & Bornm.	Gol., Maz., (Map 30)	70–650	6	EH	Geo.	310.946	900	15.5	EN B1b(i,ii,iii)	EN	LMW	Gölz et al. (2007)
31	Orchis adenocheila Czerniak. (= O. stevenii auct Fl. Iranica non Rchb. f.), (Akhani, 2005: Fig. 262)	Aras., Gol., Maz., Talish, Turk., (Map 31)	150–2619	76 (12)	XH	Geo.	359,593	67,500	146.1	LC	LC	FM, LF, MF, SC	Akhani (1998); F. Iranica 126: 115–116; F. Iran 57: 57–58; Grossheim (1940); (A. 13,511)
32	Orchis caspia Trautv.	Gol., Talish, (Map 32)	–	18	OH	Geo.	157,685	35,000	109.6	LC	ENC1	MW, SC	F. Iranica 126: 105–106; Grossheim (1940)
Poaceae
33	Agropyron bulbosum Boiss.	Maz., (Map 33)	2300	2	AH	Hem.					DD	GR	F. Iranica 70: 156–157
34	Festuca akhanii Tzvelev	Gol., Maz., (Map 34)	1300–1750	2 (2)	CEH	Hem.		4050	44.8	CR B1b(i,ii)	NT	MST	Akhani (1998); Tzvelev (1997); (A. 10,806)
35	Phleum iranicum Bornm. & Gauba	Maz., Maz./Qazv., Maz./Sem., Maz./Tehr., (Map 35)	2130–2800	5	AH	Hem.	1368	1728	24.4	EN B1ab (i,ii)	EN	MMW	F. Iranica 70: 289; Nath and Nielsen (1963)
36	Poa golestanensis H. Scholz & Akhani, (Fig. 2F)	Gol., (Map 36)	1250–2380	10 (10)	EH	Hem.	2099	1444	18.7	EN B1b(i,ii)	EN	MF, MMW, MSC	Akhani (1998); (A. 11,329)
37	Poa masenderana Freyn & Sint., (Akhani, 2005: Fig. 438)	Gil., Gol., Iraq, Maz., Sem., Talish, Tehr., Turkey, (Map 37)	75–2600	33 (8)	OH (disjunctly it occurs in Zagros)	Hem.	524,969	47,500	115.9	LC	LC	CLF, CMF, RH	Akhani (1998); Edmondson (1985); F. Iranica 70: 38; Grossheim (1939); (A. 11,912)
38	Trisetum bungei Boiss.	Gol., Sem., (Map 38)	2600	3	AH	Hem.	653.836	450	14.6	EN B1+B2ab (i,ii)	DD	GR	F. Iranica 70: 318
Typhaceae
39	Typha caspica Pobed.	Gil., Talish, (Map 39)	–	2	WH	Hem.		18	3.4	CR B1b(i,ii)	DD	MH	F. Iranica 71: 3; F. Iran 42: 9
Eudicots
Apiaceae
40	Bunium scabrellum Korov.	Talish, (Map 40)	–	1	WH	Geo.					DD	Unknown	F. Iranica 162: 251–252
41	Bupleurum flexile Bornm. & Gauba	Gil., Gol., Maz., (Map 41)	1200–2700m	15 (7)	OH	Hem.	52,699	22,500	56.6	LC	LC	MF, RO	F. Iranica 162: 290; F. Iran 54: 295; (A. 13,709)
42	Bupleurum ghahremanii Mozaff.	Maz., (Map 42)	1850–2950	3 (3)	CH	Hem.		2	1.2	CR B1+B2b(i,ii)	CR	MF, SP	F. Iran 54: 295–296; (G.&M. 8436 (TUH))
43	Cervaria cervariifolia (C. A. Mey.) M. Pimen.	E Azar., Gil., Gol., Maz., Qazv., Talish, Tehr., Turk., (Map 43)	500–3500	82 (8)	OH	Hem.	223,063	60,000	95	LC	LC	JWO, MF, SC	Akhani (1998); F. Iranica 162: 452–453; F. Iran 54: 449–451; Joharchi et al. (2007); (A. 11,157)
44	Daucus littoralis Smith subsp. hyrcanicus Rech. f.	Gil., Gil./Ardabil, Maz., (Map 44)	−20–1370	16 (1)	WH	Hem.	20,245	13,448	40.9	NT	VU A1c	CSD	F. Iranica 162: 139–140; F. Iran 54: 181–183; (A. 15,455)
45	Dichoropetalum ramosissimum (Mozaff.) Pimenov &Kljuykov (=Johrenia ramosissima Mozaff.)	Maz., (Map 45)	200–1700	5 (4)	CH	Hem.	228.368	36	2.9	EN B1+B2ab (i,ii)	EN	LF, MF, SF	F. Iran 54: 367–368; (M. 72,066 (TARI))
46	Eriocycla ghafooriana Akhani	Gol., (Map 46)	550–1015	4 (3)	EH	Cham.		392	13.8	CR B1b(i,ii)	CR	LVC	Akhani (1998); F. Iran 54: 343–344; (A. 12,322)
47	Froriepia subpinnata (Ledeb.) Baill.	Gil., Gol., Maz., Talish, (Map 47)	0–1650	41 (11)	XH	Hem.	354,024	60,000	153.3	LC	LC	FO, SC	Akhani (1998); F. Iranica 162: 297–298; F. Iran 54: 296–297; (A. 12,394)
48	Heracleum gorganicum Rech. f.	Gol., Maz., (Map 48)	1220–2600	8 (4)	CEH	Hem.	10,533	11,094	43.1	VU B1ab (i,ii,iii)	LC	CMF	Akhani (1998); F. Iranica 162: 497; F. Iran 54: 483; (A. 10,591)
49	Heracleum rechingeri Manden	Gil., Maz., Tehr., (Map 49)	1250–2950	6	WH	Hem.	3760	2700	29.7	EN B1b(i,ii)	VU A4	MF	F. Iranica 162: 496; F. Iran 54: 482–483
50	Johrenia golestanica Rech. f.	Gol., (Map 50)	220–2000	21 (18)	EH	Hem.	680.062	504	6.3	EN B1b(i,ii)	EN	FO, RO, SC	Akhani (1998); F. Iranica 162: 376–377; F. Iran 54: 365–366; (A.12,302)
51	Leutea glaucopruinosa (Rech.f.) Akhani & Salimian (=Peucedanum pimenovii Mozaff.)2	Gol., Maz., (Map 51)	1460–1890	6 (2)	CEH	Hem.	447.061	2048	31.6	EN B1b(i,ii)	EN	MF, RO	F. Iranica 162: 443–444; F. Iran 54: 437–438; (A. 15,015)
52	Leutea gracillima M. Pimenov	Gol. (Golestan National Park), (Map 52)	750–2080	3 (2)	EH	Hem.	71.052	12	1.9	CR B1b(i,ii)	EN A4a	CRO, JWO	Akhani (1998); F. Iranica 162: 448–449; (A. 12,060)
53	Leutea laseroides Akhani	Gol. (Golestan National Park), (Map 53)	700–2010	6 (6)	EH	Hem.	33.795	20	1.6	CR B1b(i,ii)	EN A4a	LVC	Akhani (1998); (A. 12,361)
54	Leutea nematoloba (Rech. f.) M. Pimen.	Maz., Sem., (Map 54)	1000–2400	7 (1)	AH	Hem.	461.596	1764	20.6	EN B1b(i,ii)	EN	CRO	F. Iranica 162: 447; F. Iran 54: 445; (A.&Gh. 15,473)
55	Leutea polyscias (Boiss.) M. Pimen.	Gil., Qazv., (Map 55)	200–1650	16 (4)	MR	Hem.	1409	700	10.2	EN B1b(i,ii)	EN	VC	F. Iranica 162: 446–447; F. Iran 54: 441–442
56	Leutea translucens (Rech.f.) Akhani & Salimian (= Peucedanum translucens Rech. f.)3, (Fig. 2G)	Maz., Gol., (Map 56)	1750–2750	4 (3)	EH	Hem.	687.362	768	15.8	EN B1ab (i,ii)	CR C1	JWO	F. Iranica 162: 444; (A. 15,359)
57	Polylophium involucratum (Pall.) Boiss.	Maz., (Map 57)	2000–3000	6	AH	Hem.	699.317	245	7.1	EN B1+B2b(i,ii)	VU A4	AZ (MW, SMW)	F. Iranica 162: 525; F. Iran 54: 507
58	Seseli elbursense Pimenov & Kljuykov (=Trachydium eriocarpum Bornm. & Gauba)	Maz., (Map 58)	3200–3400	2	AH	Hem.		8	2.4	CR B1+B2b(i,ii)	EN A4	AZ, CRO	F. Iranica 162: 181–182
59	Seseli kiabii (Akhani) Akhani	Gol., Maz., (Map 59)	300–2000	10 (8)	EH	Hem.	7664	3600	30.1	VU B1ab (i,ii)	VU	OSC, RO	Akhani (1998); Akhani et al. (2016); F. Iran 54: 349–350; (A. 12,370)
Aquifoliaceae
60	Ilex spinigera (Loes.) Loes., (Fig. 2H)	Gil., Gol., Maz., Talish, (Map 60)	30–2018	68 (31)	OH	Phan.	117,529	47,500	68.6	LC	LC	LF, MF, SF	Akhani (1998); Browicz (1984); F. Iranica 25: 1; F. Iran 130: 3–5; (A. 14,134)
Araliaceae
61	Hedera pastuchovii Woron. ex Grossh., (Fig. 2I)	Gil., Gol., Maz., Talish, (Map 61)	50–1800m	76 (16)	OH	Phan. (Liana)	247,810	67,500	102.4	LC	LC	LF, MF, SF	Browicz (1986); F. Iranica 102: 2–3; (A. 13,661)
Asteraceae
62	Amblyocarpum inuloides Fisch. & C. A. Mey.	Gil., Gol., Maz., Talish, (Map 62-Inc.)	Sea level–680m	9 (2)	OH	Ther.	53,038	12,500	57.6	LC	VU A4c	MH, RH	F. Iranica 145: 126; (Z. 7162 (IRAN))
63	Anthemis mazandaranica Iranshahr	Gil., Maz., (Map 63)	900–1520	5	WH	Ther.	4325	3888	36	EN B1ab (i,ii)	EN	Unknown	F. Iranica 158: 21; F. Iran 59: 26
64	Anthemis talyschensis A. Fedor.	Gil., Gil./Ardabil, Maz., Talish, (Map 64)	1250–4800	10	WH	Hem.	16,117	8000	40.1	VU B1ab(i)	VU	AZ, MSC	F. Iranica 158: 26; F. Iran 59: 34
65	Anthemis triumfettii (L.) All. subsp. khorasanica (Rech. f.) Iranshahr	Gil., Gol., Gol./N Khor., Gol./Semn, Maz., Turk., (Map 65)	400–3000	29 (4)	CEH	Hem.	83,123	16,000	64.2	LC	LC	LSC, LWO, MSC, MWO	Akhani (1998); F. Iranica 158: 20; F. Iran 59: 22–26; (A. 10,684)
66	Carduus transcaspicus Gandog. subsp. transcaspicus	Gol., Turk., (Map 66)	780–1700	7 (4)	EH	Ther.	9961	4205	28.9	VU B1ab (i,ii)	LC	DH, FM, RO	Akhani (1998); F. Iranica 139a: 223; (A. 11,476)
67	Centaurea golestanica Akhani & Wagenitz	Gol. (Golestan National Park), (Map 67)	1200–1550	4 (4)	EH	Hem.	4.094	1.080	0.6	CR B1+B2b(i,ii,iii)	CR	GOWO	Akhani (1998); (A. 11,704)
68	Centaurea hyrcanica Bornm., (Fig. 2J)	Aras., Gil., Gil./Ardabil, Gol., Maz., Talish, (Map 68)	50–2810	101 (57)	OH	Hem.	130,026	65,000	81.5	LC	LC	LF, MF, RO, SC	Akhani (1998); F. Iranica 139b: 345–346; (A. & al. 21,281)
69	Centaurea kandavanensis Wagenitz	Alborz, Maz., (Map 69)	270–2500	15 (3)	AH	Hem.	3562	1296	12.3	EN B1b(i,ii)	EN	RO, SC	F. Iranica 139b: 392; (T. & al. 8182 (IRAN))
70	Centaurea meyerana Tzvel.	Talish, (Map 70)	1200	2	WH	Hem.					DD	Unknown	F. Iranica 139b: 404–405
71	Centaurea zuvandica (Sosn.) Sosn.	Ardabil, Aras., Gil., Gol., Maz., Sem., Talish, Tehr., (Map 71)	0–3000	90 (49)	OH	Hem.	143,494	50,000	81.1	LC	LC	CRO, VC	Akhani (1998); F. Iranica 139b: 403–404; (A. & al. 21,044)
72	Cephalorrhynchus gorganicus (Rech. f. & Esfand.) Tuisl	Gol., Maz., Talish, (Map 72)	2700	3	OH	Hem.	58,033	7500	62.5	LC	DD	SWO	F. Iranica 122: 212–213
73	Cirsium gadukense Petrak	Maz., (Map 73)	2150	1	AH	Hem.					DD	MST, OSC	F. Iranica 139a: 257
74	Cousinia alfredii Bornm. & Gauba	Alborz, Maz., (Map 74)	1300–1500	2	AH	Hem.		98	6.8	CR B1b(i,ii)	CR	SF (Xeric slopes)	F. Iranica 90: 92–93; F. Iranica 139a: 117
75	Cousinia decipiens Boiss. & Buhse	Gol., Gol./N Khor., Gol./Sem., Maz., (Map 75)	1000–2810	18 (8)	EH	Hem.	5440	5103	27.4	VU B1b(i)	LC	AMW, JWO, MST, SC	Akhani (1998); F. Iranica 90: 158–159; F. Iranica 139a: 126; (A. 16,844)
76	Cousinia erinacea Jaub. & Spach	Gil., (Map 76)	300–1000	6	MR	Hem.	716.516	125	4.6	EN B1+B2b(i,ii,iii)	EN	STMX	F. Iranica 90: 307–308; F. Iranica 139a: 150
77	Cousinia gmelini C. Winkl.	Gil., Maz., (Map 77)	2600–3300	4	AH	Hem.	1715	432	12.4	EN B1+B2ab (i,ii)	EN	AMW, SMW	F. Iranica 90: 121–122; F. Iranica 139a: 121
78	Cousinia hypochionea Bornm.	Gil., (Map 78)	400–450	1	MR	Hem.					DD	SEMX	F. Iranica 90: 316
79	Cousinia rechingerae Bornm.	Gol., N Khor., (Map 79)	740–1300	11 (3)	EH	Hem.	2166	448	8.5	EN B1+B2b(i,ii)	VU A4a	OSC	F. Iranica 90: 93–94; F. Iranica 139a: 117; (A. 14,111)
80	Cousinia wendelboi Rech. f.	Maz., (Map 80)	2000	1	AH	Hem.					DD	MSC	F. Iranica 90: 167
81	Crepis alfredi Bornm.	Gil., (Map 81)	300	1	MR	Hem.					DD	Unknown	F. Iranica 122: 338
82	Crepis papposissima Babcock	Gol., (Map 82)	–	1	EH	Ther.					DD	Unknown	F. Iranica 122: 319
83	Crepis willemetioides Boiss.	Gol., Maz., Turk., (Map 83)	50–2900	20 (10)	CEH	Geo.	43,886	15,000	54.5	NT	LC	LF, MSC, MWO	Akhani (1998); F. Iranica 122: 302–303; (A. 16,667)
84	Dolichorrhiza persica (Boiss.) B. Nord.	Gil., Maz., (Map 84)	2600–3400	4	AH	Hem.		648	17.7	CR B1b(i,ii)	CR	AZ	F. Iranica 164: 52–53
85	Doronicum wendelboi Edmondson	Gol., Maz., Sem., (Map 85)	2000–2600	4	EH	Hem.	2499	1296	18.1	EN B1ab (i,ii)	EN	MF	F. Iranica 164: 46
86	Echinops koelzii Rech. f.	Ardabil/Gil., Gol., Kho., Maz., Maz./Tehr., Sem., W Azar.?, (Map 86)	500–3000	19 (3)	AH	Hem.	153,766	27,500	74.8	LC	LC	CMF, CSF, OMF, OSF, SC	Akhani (1998); F. Iranica 139a: 81; Joharchi et al. (2007); (A. 9827)
87	Hieracium hoppeanum Schultes	Aras., Gil., (Map 87)	1200–2400	5	WH	Hem.	6363	6480	35.7	VU B1ab(i)	VU	MMW	F. Iranica 122: 172–173
88	Hieracium kandawanicum (Rech. f. & Zahn) Rech. f.	Maz./Alborz, (Map 88)	2100–2700	3	AH	Hem.		1.28	0.8	CR B1+B2b(i,ii)	CR	SZ	F. Iranica 122: 173–174
89	Klasea quinquefolia (Willd.) Greuter & Wagenitz (= Serratula quinquefolia M. B. ex Willd.)	Aras., Ardabil/Gil., Gil., Gol., Maz., Talish, W Azar., (Map 89)	385–2200	36 (10)	XH	Hem.	647,246	57,500	172.2	LC	LC	LF, MF, SF	Akhani (1998); F. Iranica 139b: 298–299; Martins (2006); (A. 13,726)
90	Leontodon stenocalathius Rech. f.	Gol., Gol./Sem., (Map 90)	2400–2600	2	EH	Hem.		50	5.3	CR B1b(i,ii)	CR	CRO	F. Iranica 122: 128
91	Scorzonera persica Boiss. & Buhse	Gil., (Map 91)	–	1	MR	Hem.					DD	STMX	F. Iranica 122: 69; (Buhse 984(Photo-G))
92	Serratula gracillima Rech. f.	Gol., (Map 92)	–	1	EH	Hem.					DD	Unknown	F. Iranica 139b: 291–292
93	Tanacetum archibaldii Podl.	Maz./Alborz, (Map 93)	2190	1	AH	Hem.					DD	CRO	F. Iranica 158: 105
94	Tragopogon gongylorrhizus Rech. f.	Gol., Gol./N Khor., Maz., (Map 94)	450–1820	15 (5)	EH	Hem.	7155	8214	36.7	VU B1b(i)	VU	FO, GR, SC	Akhani (1998); F. Iranica 122: 98; (A. 16,225)
95	Willemetia tuberosa DC. (= Calycocorsus tuberosus (DC.) Rauschert	Gil., Gol., Maz., Talish, (Map 95)	20–2000	48 (20)	OH	Geo.	195,749	45,000	77.4	LC	LC	LF, MH	Akhani (1998); F. Iranica 122: 289–290; (A. 21,075)
Berberidaceae
96	Berberis orthobotrys Bienert ex C. K. Schneider	Gil., Gol., Maz., Turk., (Map 96)	700–2800	43 (2)	CEH	Phan.	93,383	37,500	59.3	LC	LC	SC	Akhani (1998); F. Iranica 111: 13; F. Iran 64: 9–12; (A. 11,625)
97	Epimedium pinnatum Fisch., (Fig. 2K)	Gil., Maz., Talish, (Map 97)	100–1700	47 (9)	WH	Geo.	29,342	22,275	45	NT	NT	LF, MF, SF	F. Iranica 101: 3; F. Iran 56: 4–7; (A. & al. 21,061)
Betulaceae
98	Alnus subcordata C. A. Mey. [incl. var. subcordata & var. villosa (Regel) H. Winkl.]	Gil., Gol., Maz., Talish, (Map 98)	100–1900	90 (19)	OH	Phan.	103,886	45,000	68	LC	LC	LF, RH, SF	Akhani (1998); (Browicz, 1982); F. Iranica 96: 7–8; (A. 11,995)
Boraginaceae
99	Cynoglossum kandavanensis (Bornm. & Gauba) Akhani	Alborz, Aras., Gil., Gol., Maz., (Map 99)	Sea level–2360	16 (8)	OH	Hem.	107,255	12,500	82.1	LC	LC	FO, MMW	Akhani (1998); F. Iranica 48: 141–142; F. Iran 39: 440–442; Hamzeh'ee et al., 2010; (A. 13,521)
100	Echium amoenum Fisch. & C. A. Mey., (Fig. 2L)	Alborz, Ardabil, E Azar.?, Gil., Gol., Maz., Talish, Tehr., (Map 100)	60–3000	110 (42)	OH	Hem.	172,439	57,500	71.6	LC	LC	FO, MMW	Akhani (1998); F. Iranica 48: 215; F. Iran 39: 184–185; Grossheim (1967); (A. & al. 20,932)
101	Eritrichium gracillimum Rech. f.	Gol., (Map 101)	3000	1	AH	Hem.					DD	AZ	F. Iranica 48: 65–66
102	“Heliotropium sp. nov.”6	Gil., Qazvin, (Map 102)	150–850	6 (6)	MR	Ther.	59.498	80	4.2	CR B1b(i,ii,iii)	EN C2b	SMX	Unpublished
103	Lepechiniella persica (Boiss.) H. Riedl	Gol./Sem., Maz., (Map 103)	2400–3900	4	AH	Hem.	10,958	4563	38.9	VU B1ab (i,ii)	DD	AZ	F. Iranica 48: 81–82; F. Iran 39: 335–336
104	Lepechiniella wendelboi H. Riedl	Maz., Sem., Tehr., (Map 104)	1000–4100	8	AH	Hem.	12,330	5766	30.7	VU B1ab (i,ii)	VU	AZ, SZ	F. Iranica 48: 82; F. Iran 39: 336–338
105	Myosotis anomala H. Riedl	Aras., Ardabil ?, Gil., Maz., N Khor., (Map 105)	250–2450	11	OH	Hem.	115,944	17,500	86.6	LC	LC	LF, MF, MH	F. Iranica 48: 266; F. Iran 39: 266–267; Joharchi et al. (2007)
106	Nonea longiflora Wettst. ex Stapf	Gil., (Map 106)	200–500	5	MR	Ther.	30.299	12	1.9	CR B1b(i,ii,iii)	CR	SC, ST	F. Iranica 48: 244; Nejhad-Falatoury et al. (2011)
107	Rindera regia (Gmelin) Kusn.	Gil., Maz., (Map 107)	2500–2750	2	AH	Hem.		0.18	0.3	CR B1+B2b(i,ii)	CR	AZ	F. Iranica 48: 127; F. Iran 39: 428
Brassicaceae
108	Alyssopsis mollis (Jacq.) O. E. Schulz	Ardabil, Gil., Gol., Maz., N Khor., Sem., Talish, Tehr., (Map 108)	400–3200	87 (35)	OH	Hem.	132,142	52,500	80	LC	LC	LF, MF, RO, SC, VC	Akhani (1998); F. Iranica 57: 200; F. Iran 143: 492–495; (A. 13,707)
109	Hesperis hyrcana Bornm. & Gauba, (Fig. 2M)	Aras., Gil., Gol., Maz., Cauc., (Map 109)	Sea level–2750	126 (33)	XH	Hem.	497,196	147,500	169.9	LC	LC	CLF, FM, MF, MH, SF	Akhani (1998); Dvořák (1964); F. Iranica 57: 268; F. Iran 143: 648–651; (A. 9349)
110	Isatis gaubae Bornm.	Gol., N Khor., Maz., Sem., Tehr., (Map 110)	570–3000	39	CEH	Ther.	57,432	37,500	68.9	LC	LC	MSC	F. Iranica 57: 85; F. Iran 143: 541-543
111	Noccaea umbellata (Steven ex DC.) Al-Shehbaz	Gil., Gol., Maz., Talish, (Map 111)	Sea level–1930	43 (2)	OH	Ther.	158,756	50,000	80	LC	LC	LF, MF, SF	F. Iranica 57: 113–114; F. Iran 143: 143; Grossheim (1950); (J. 1866)
112	Noccaea hastulata (Steven ex DC.) Khosravi	Gil., Gol., Maz., Talish, (Map 112)	150–2500	54 (4)	OH	Ther.	89,083	45,000	62.6	LC	LC	FM, MF, MW	F. Iranica 57: 115–116; F. Iran 143: 148–151; Grossheim (1950); (A. 13,746)
Buxaceae
113	Buxus hyrcana Pojark., (Fig. 2N)	Gil., Gol., Maz., Talish, (Map 113)	−20–1250	48 (15)	OH	Phan.	62,517	40,000	54.2	LC	NT	ALF	F. Iranica 27: 4; F. Iran 131, 3–5; (A. 21,266)
Campanulaceae
114	Campanula ghilanensis Pall.	Gil., (unknown point, without Map)	–	1	WH	Hem.					DD	Unknown	F. Iranica 13: 37
115	Campanula lourica Boiss., (Fig. 3A)	Alborz, Gol., Maz., N Khor., Sem., Tehr., (Map 114)	700–3400	41 (8)	CEH	Hem.	34,716	35,000	52.7	NT	LC	CRO	Akhani (1998); F. Iranica 13: 24; F. Iran 66: 50–51; (A. 11,396)
Caprifoliaceae
116	Lonicera floribunda Boiss. & Buhse	Gil., Gol., Maz., Turk., Cauc., (Map 115)	400–2460	72 (22)	OH	Phan.	444,406	65,000	129.7	LC	LC	SC	Akhani (1998); (Browicz, 1991); F. Iranica 10: 13; F. Iran 13: 20–23; (A. 16,209)
117	Lonicera iberica M. Bieb., (Fig. 3B)	Ardabil, Cauc., E Azar., Gil., Gol., Maz., Qazv., Sem., Talish, Tehr., Turkey, W Azar., (Map 116)	500–3000	136 (5)	XH	Phan.	535,599	155,000	135.1	LC	NT	MF, JWO, RO, SASC	Akhani (1998); Browicz (1988); F. Iranica 10: 9–10; F. Iran 13: 15–16; (A.11,350)
118	Scabiosa schimperiana Boiss. & Buhse	Gil., (Map 117)	300–1300	6	MR	Hem.	148.546	75	5.4	EN B1+B2b(i,ii,iii)	EN	PMX	F. Iranica 168: 48–49; F. Iran 8: 91–94
Caryophyllaceae
119	Cerastium microspermum C. A. Mey.	Ardabil, Gil., Talish, (Map 118)	500–2400	7	WH	Ther.	2420	2704	26.2	EN B1b(i,ii)	EN	MW	F. Iranica 163: 94
120	Dianthus agrostolepis Rech. f.	Ardabil/Gil., Gil., (Map 119)	420–2100	4	WH	Hem.	4232	1600	20	EN B1ab (i,ii)	EN	MF	Assadi (1985); F. Iranica 163: 155
121	Dianthus hyrcanicus Rech. f.	Gil., Maz., (Map 120)	0–2250	4	WH	Hem.	179.158	675	15.4	EN B1ab (i,ii)	EN	LF, MF, RO	F. Iranica 163: 144
122	Dianthus mazanderanicus Rech. f.	Maz., (Map 121)	1950–2700	2	AH	Hem.		50	4.6	CR B1b(i,ii)	ENC1	MST	F. Iranica 163: 154–155
123	Dianthus orientalis Adams subsp. gilanicus Rech. f.	Ardabil/E Azar. ?, Gil., Gil./Qazv., Maz., Sem., (Map 122)	200–2320	10	AH	Cham.	19,203	12,500	53.2	VU B1ab(i)	LC	LSC, MSC, ST	F. Iranica 163: 158–159
124	Dianthus orientalis Adams subsp. gorganicus Rech. f., (Akhani, 2005: Figs. 560–561)	Gol., Maz., Sem./Gol., (Map 123)	350–2600	38 (15)	EH	Cham.	14,298	10,890	33.4	VU B1b(i)	LC	RO, SC	Akhani (1998); F. Iranica 163: 162; (A. 13,700)
125	Dianthus rudbaricus Assadi	Gil., (Map 124)	200	2	MR	Hem.		0.08	0.2	CR B1+B2b(i,ii,iii)	DD	SAMX	F. Iranica 163: 188
126	Dianthus talyschensis Boiss. & Buhse	Talish, (Map 125)	900	1	WH	Hem.					DD	LF	F. Iranica 163: 143–144; (Buhse 802 a (Photo-G))
127	Gypsophila modesta Bornm.	Gil., (Map 126)	400–500	2	MR	Ther.		32	3.9	CR B1b(i,ii,iii)	CR	SMX	F. Iranica 163: 246
128	Saponaria bodeana Boiss., (Fig. 3C)	Gol., Maz., (Map 127)	400–2600	20 (12)	EH	Hem.	11,741	13,689	39.3	VU B1b(i)	VU	MF	Akhani (1998); F. Iranica 163: 198; Joharchi et al. (2007); (A. 11,092)
129	Silene lineata Boiss. & Buhse	Gil., (Map 128)	–	1	MR	Hem.					DD	Unknown	F. Iranica 163: 386; (Buhse 984 (Photo-G))
130	Silene schafta Gmel. Jun. ex Hohen.	Gil., Gol., Maz., Talish, (Map 129)	820–2810	33 (12)	OH	Hem.	68,790	32,500	59.2	LC	LC	MSC, MST, RO	F. Iranica 163: 466–467; (A. 13,660)
131	Silene sojakii Melzh.	Gol. (Golestan National Park)?, Sem., (Map 130)	2100–2800	4	EH	Cham.	2180	3072	31.8	EN B1ab (i,ii)	EN	CRO	Akhani (1998); F. Iranica 163: 462; Gholipour and Sheidai (2009)
Celastraceae
132	Evonymus velutina (C. A. Mey.) Fisch. & C. A. Mey., (Fig. 3D)	Gol., Maz., N Khor., Talish, Turk., (Map 131)	300–2000	48 (18)	OH	Phan.	201,896	37,500	90.2	LC	NT	OLF, OMF, OSC, RO	Akhani (1998); F. Iranica 64: 2; Browicz (1986); Joharchi et al. (2007); (A. 14,121)
Crassulaceae
133	Sedum lenkoranicum Grossh.	Alborz, Aras., Ardabil/Gil., Gil., Maz., Sem., Talish, (Map 132)	1300–2500	18	AH	Hem.	96,990	30,000	75.7	LC	LC	MF, MSC, RO	F. Iranica 72: 12; F. Iran 32: 24–26; Grossheim (1950); (A. 12,047)
134	Sempervivum iranicum Bornm. & Gauba, (Akhani, 2005: Figs. 784–785)	Alborz, Gil., Gol., Maz., Sem., (Map 133)	1650–2950	25 (8)	AH	Hem.	49,691	32,500	57	LC	VU A4c	SAST, SP, SWO	Akhani (1998); F. Iranica 72: 4; F. Iran 32: 11–13; (A. 17,001)
Euphorbiaceae
135	Euphorbia mazandaranica Pahlevani	Maz., (Map 134)	Sea level–1450	9	CH	Hem.	4289.5	4375	25	EN B1b(i,ii)	VU A4	LF, SF	Grossheim (1962); Pahlevani and Riina (2014); Pahlevani et al. (2020)
Fabaceae
136	Albizzia julibrissin Durazz., (Fig. 3E)	Gil., Gol., Maz., Talish, (Map 135)	Sea level–620	44 (5)	OH (disjunctly occurs in SE Asia)	Phan.	57,575	30,000	53.6	LC	NT	LF	Browicz (1982); F. Iranica 161: 9; F. Iran 18: 30; (Gho. 4535)
137	Astragalus acutifolius Bunge (= A. ammodendroides Podlech & Zarre)	Gil., (Map 136)	200–360	7 (2)	MR	Cham.	0.141	1.470	0.7	CR B1+B2b(i,ii,iii)	CR	SMX	F. Iranica 178: 379; (Ma.&Sa. 85,976 (TARI))
138	Astragalus lacus-valashti Maassoumi, Podlech & Jalili	Maz., (Map 137)	530–2268	5 (1)	CH	Hem.	94.112	80	4	CR B1b(i,ii)	EN C1	MSC, SMSC, WO	F. Iranica 178: 343; (Ma.&Ja. 82,422 (TARI))
139	Astragalus nurensis Boiss. & Buhse	Alborz, Gol., Maz., Sem., (Map 138)	150–3000	37 (11)	AH	Hem.	25,953	26,460	42	NT	NT	MSC, MST, RO	F. Iranica 178: 44–45; (Ma. 75,471 (TARI))
140	Astragalus rosellus Širj. & Rech. f.	Alborz, Gil., Gol., Maz., Sem., (Map 139)	1500–2750	11 (2)	AH	Cham.	17,825	10,086	41.2	VU B1ab(i)	LC	SASC	F. Iranica 177: 103–104; (Fo. 453 (TARI))
141	Astragalus vereskensis Maassoumi & Podlech	Gil., Gol., Maz., Sem., (Map 140)	200–3000	33 (1)	AH	Hem.	31,527	32,500	53.6	NT	NT	LMW, LSC, LST, MMW, MSC, MST	F. Iranica 174: 236–237; F. Iran 43: 185–186; (Klet 44,087 (IRAN))
142	Astragalus yushensis T. Sabaii, Zarre & Podlech. (=A. submitis Boiss. & Hohen. subsp. maassoumii Tietz & Zarre)	Maz., (Map 141)	500–2500	6	AH	Cham.	62.895	80	3.9	CR B1b(i,ii)	ENC1	MSC, MST, SMSC, SMST	F. Iranica 179: 120-121
143	Colutea buhsei (Boiss.) Shap.	Alborz, Ardabil, Gol., Maz., N Khor., Qazv., R. Khor., Sem., Tehr., Turk., (Map 142)	320–3000	109 (9)	CEH (also in Khorassan-Kopet Dagh)	Phan.	201,475	65,000	97.5	LC	LC	DV, FM, OSC, RO	Akhani (1998); Browicz (1986); F. Iranica 157: 72–73; (A. 16,616)
144	Gleditsia caspica Desf.	Gil., Gol., Maz., Talish, (Map 143)	−25–390	46 (8)	OH	Phan.	63,189	32,500	53.6	LC	LC	LF	Browicz (1982); F. Iranica 160: 10–11; F. Iran 45: 20–21; (A. & al. 21,030)
145	Onobrychis heterophylla C. A. Mey.	Talish, (Map 144)	900–1200	7	WH	Hem.	3488	4500	29.8	EN B1b(i,ii)	EN	FO, SC	F. Iranica 157: 437; Grossheim (1952)
146	Onobrychis mazanderanica Rech. f.	Alborz, Gol., Maz., (Map 145)	500–2400	13 (1)	AH	Hem.	24,924	16,200	44.8	NT	NT	FO, SC	F. Iranica 157: 457; (A. 14,577)
147	Trifolium mazanderanicum Rech. f.	Maz., (Map 146)	400	1	EH	Hem.					DD	FO	F. Iranica 157: 313
Fagaceae
148	Quercus castaneifolia C. A. Mey., (Fig. 3G)	Gil., Gol., Maz., Talish, (Map 147)	Sea level–2400	119 (36)	OH	Phan.	225,250	67,500	77.5	LC	LC	CLF, OLF, CSF, OSF, CMF, OMF, SC	Akhani (1998); Browicz (1982); F. Iranica 77: 10–11; (A. 14,580)
Gentianaceae
149	Gentiana grossheimii Doluch.7, (Fig. 3H)	Daghestan, Gol., (Map 148)	800–2300	13 (2)	OH	Hem.	9253 (for Daghesten records)	3249 (for Daghesten records)	19 (for Daghesten records)	VU B1b(i,ii) (Daghestan)	CR D (Iran)	LVC, CRO, OSC	Dolukhanof (1948); Grossheim (1967); (A. 23,291)
Geraniaceae
150	Erodium dimorphum Wendelbo	Maz., Tehr./Sem., (Map 149)	2600–3300	2	AH	Hem.		18	3.5	CR B1b(i,ii,iii)	CR	RO, AZ	F. Iranica 69: 50; F. Iran 62: 89–91
Hamamelidaceae
151	Parrotia persica (DC.) C. A. Mey., (Fig. 3F)	Gil., Gol., Maz., Talish, (Map 150)	Sea level–1500	102 (36)	OH	Phan.	174,918	55,000	78.6	LC	LC	ALF, LF, RH	Akhani (1998); Browicz (1982); F. Iranica 53: 1–2; F. Iran 136: 3–5; (A. 14,579)
Hypericaceae
152	Hypericum fursei N. Robson	Maz., (Map 151)	1200	1	CH	Hem.					CR D	SC	F. Iranica 49: 16
Lamiaceae
153	Ballota platyloma Rech. f.	Gil., Maz., Sem. Tehr., (Map 152)	400–2500	17 (5)	AH	Hem.	9776	7688	31	VU B1b(i)	VU	LSC, LWO, MSC, MWO, RH	F. Iranica 150: 352–353; F. Iran 76: 411; (A. 13,749)
154	Betonica nivea Stev. subsp. mazandarana (Bornm.) Rech. f.	Maz., Tehr., (Map 153)	2000–4000	13 (6)	AH	Hem.	5576	3200	20.1	VU B1b(i,ii)	LC	AZ, RO	F. Iranica 150: 399–400; (Mo. 14.7.2006, s.n.)
155	Eremostachys lanata Jamzad	Maz., (Map 154)	1900–2350	2 (2)	AH	Hem.					CR C2b	MST	Jamzad (1987); (As.& M. 33,028 (holotype- TARI))
156	Lagochilus quadridentatus Jamzad	Maz., (Map 155)	1900–2300	2 (1)	AH	Hem.					DD	MST	Jamzad (1988); (As.& M. 33,030 (holotype- TARI))
157	Nepeta pogonosperma Jamzad & Assadi	Maz., Zanjan ?, (Map 156)	2900–3600	6 (1)	AH	Hem.	4780	4500	30.3	EN B1b(i,ii)	EN	AZ, RO	Jamzad and Assadi (1984); Jamzad (1991); (R.&Ma. 21,689 (holotype- TARI))
158	Phlomis ghilanensis C. Koch	Gil., (Map 157)	–	1	AH	Hem.					DD	AZ	F. Iranica 150: 301–302
159	Salvia oligophylla Auch. ex Benth.	Gil., Qazv., Zanjan, (Map 158)	450–1470	12	MR	Hem.	1629	1372	14.4	EN B1b(i,ii,iii)	EN	MX	F. Iranica 150: 466; F. Iran 76: 915
160	Salvia shahkuhmahalei Akhani	Gol./N Khor., (Map 159)	1500–1935	6	EH	Hem.	30.014	6	1.3	CR B1+B2b(i,ii)	CR	SC	Akhani et al. (2016)
161	Satureja intermedia C. A. Mey.	Gil., Talish, (Map 160)	1900–2350	7 (1)	WH	Hem.	426.775	432	12.3	EN B1+B2b(i,ii)	EN	RO	F. Iranica 150: 498; F. Iran 76: 675
162	Satureja isophylla Rech. f.	Maz., (Map 161)	1600–3300	13 (1)	CH	Hem.	1213	576	8	EN B1b(i,ii)	EN	MF, MSC, MST,	F. Iranica 150: 498; F. Iran 76: 687
163	Satureja mutica Fisch. & C. A. Mey.	Gil., Gol., Maz., N Khor., Talish, Turk., (Map 162)	550–2080	21 (8)	OH	Hem.	174,541	27,500	89	LC	LC	RO, VC	Akhani (1998); F. Iranica 150: 499; F. Iran 76: 694; (A. 12,263)
164	Scutellaria tournefortii Benth., (Fig. 3I)	Gil., Gol., Maz., Talish, (Map 163)	20–2130	86 (11)	OH	Hem.	124,962	70,000	68	LC	LC	CLF, CMF	Akhani (1998); F. Iranica 150: 52–53; F. Iran 76: 92; Grossheim (1967); (A. 9278)
165	Stachys laxa Boiss. & Buhse	Gol., Maz., Sem., Tehr., (Map 164)	500–2810	43 (8)	CEH	Hem.	36,956	29,584	42.7	NT	LC	RO	Akhani (1998); F. Iranica 150: 389–390; Salmaki et al. (2012); (A. 12,326)
166	Stachys talyschensis Kapeller	Talish, (Map 165)	–	1	WH	Hem.					DD	SP	F. Iranica 150: 379; Grossheim (1967)
167	Teucrium hyrcanicum L.	Gil., Gol., Maz., Talish, (Map 166)	−20–2000	97 (30)	XH	Hem.	495,070	115,000	149.1	LC	LC	LF, MF, OSC, WO	Akhani (1998); F. Iranica 150: 36–37; Grossheim (1967); (A. 11,960)
Linaceae
168	Linum bungei Boiss.	Gol., Maz., Sem., (Map 167)	2100–3200	14 (3)	AH	Hem.	27,044	15,129	41.1	NT	NT	MMW, MSC, MST	F. Iranica 106: 12; F. Iran 34: 29; (Furse 2769 (IRAN))
Lythraceae
169	Peplis hyrcanica Sosn.	Talish, (Map 168)	–	1	WH	Ther.					DD	MH	F. Iranica 51: 6–7
Malvaceae
170	Alcea 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)	100–2050	35 (12)	CEH	Hem.	123,614	21,160	46.2	LC	LC	FM, FO, OSC, WO	Akhani (1998); F. Iranica 120: 72, 76–77, 83; F. Iran 58: 77–80; (A. 11,197)
171	Alcea hyrcana (Grossh.) Grossh.	Gil., Maz., Talish, (Map 170)	100–2100	5	WH	Hem.	20,362	5445	33	NT	NT	LF, MF	F. Iranica 120: 57; F. Iran 58: 108–110
172	Tilia platyphyllos Scop. subsp. caucasica (Rupr.) Loria9, (Fig. 3J)	Gil., Gol., Maz., Talish, (Map 171)	100–2400	141 (4)	XH	Phan.	1,452,474	217,500	252.9	LC	NT	CLF, CMF	Akhani (1998); F. Iranica 148: 6–7
Orobanchaceae
173	Orobanche eriophora Bornm. & Gauba10	Maz., (Map 172)	2300	1	AH	Geo.					DD	MSC?	F. Iranica 5: 10
174	Orobanche schwingenschussii Gilli	Maz./Alborz, (Map 173)	2860	2	AH	Geo.		0.08	0.2	CR B1+B2b(i,ii)	DD	MSC?	F. Iranica 5: 10; F. Iran 86: 50-51
175	Rhynchocorys maxima C. Richter, (Fig. 3K)	Alborz, Gil., Gol., Maz., Talish, Tehr., (Map 174)	−26–2750	70 (35)	OH	Hem.	185,771	62,500	88.8	LC	LC	DH, FM, FO, LF, MF,	Akhani (1998); Burbidge and Richardson (1970); F. Iranica 147: 209; (A. & al. 20,929)
Paeoniaceae
176	Paeonia tomentosa (Lomak.) N. Busch11, (Fig. 3L)	Gil., Gol., Maz., Talish, (Map 175)	1000–3000	56 (6)	OH	Geo.	119,308	40,000	68.8	LC	LC	CMF, FM, FO, SC	Akhani (1998); F. Iranica 60: 4; F. Iran 137: 4–6; Hong and Zhou (2003); Rukšãns and Zetterlund (2014); (A. 10,310)
177	Paeonia wendelboi Rukšāns & Zetterlund	Ardabil-Gil., (Map 176)	2000–2300	5	WH	Geo.	9.727	3	1	CR B1+B2b(i,ii,iii)	CR	SMW, RO	Rukšāns and Zetterlund (2014)
Papaveraceae
178	Corydalis chionophila Czernjak.	Gol., N Khor., R. Khor., Turk., (Map 177)	1650–2340	12 (11)	EH KHORASSAN-KOPET DAGH (Memariani)	Geo.	32,757	12,943	43.3	NT	NT	MMW, WO	Akhani (1998); F. Iranica 110: 20; (A. 10,203)
179	Corydalis hyrcana Wendelbo	Ardabil ?, Gol., Maz., (Map 178)	1000–2450	5 (3)	OH	Geo.	32,704	7500	60	NT	VU A4c	MMW, WO	F. Iranica 110: 14–15; (J. 1590)
180	Papaver chelidoniifolium Boiss & Buhse, (Fig. 3M)	Gil., Maz., Talish, (Map 179)	−22-1800	49 (2)	OH	Ther.	57,919	39,762	47.2	LC	LC	FM, LF, SC, SF	F. Iranica 34: 19; F. Iran 127: 99–102; (A. 5335)
Plantaginaceae
181	Digitalis nervosa Steud. & Hochst. ex Benth., (Fig. 3N)	Aras.,Ardabil/Gil., Gil., Gol., Maz., Sem., Talish, (Map 180)	200–3200	88 (32)	OH	Hem.	262,974	75,000	93.6	LC	LC	FM, FO, LF, MF, RO, SC	Akhani (1998); F. Iranica 147: 169–170; F. Iran 68: 152; Grossheim (1967); (A. 13,704)
182	Plantago podlechii Akhani	Gol. (Golestan National Park), (Map 181)	1600–1700	2 (2)	EH	Hem.		0.98	0.7	CR B1+B2b(i,ii)	CR	MST, OSC	Akhani (1998); (A. 17,084)
183	Veronica bungei Boiss.	Maz./Sem., (Map 182)	Ca. 2300	1	EH	Ther.					DD	MF	F. Iranica 147: 97–98; (Bunge 15 (holotype- Photo-G-Boiss.))
184	Veronica euphrasiifolia Link	Gil. (unknown point, without map)	–	1	AH	Hem.					DD	AZ	F. Iranica 147: 124; (Willdenow 203 (type-B-Willd.))
185	Veronica francispetae M. A. Fischer	Gil., Gol., Maz., (Map 183)	20–1200	9	OH	Ther.	15,789	10,647	39.1	VU B1ab(i)	VU	LF, SF	F. Iranica 147: 104–105; Saeidi-Mehrvarz (1999)
186	Veronica gaubae Bornm.	Gil., Gol., Maz., Khor. Sem., Tehr., Turk., (Map 184)	2000–3200	35 (1)	AH	Ther.	103,502	37,500	84.5	LC	LC	MF, MST, RO	Akhani (1998); F. Iranica 147: 112–113; F. Iran 68: 115; (A. 12,036)
187	Veronica mazanderanae Wendelbo	Ardabil, Gil., Gol., Maz., (Map 185)	520–2619	13 (4)	AH	Ther.	89,212	17,500	63.9	LC	LC	LST, MST	F. Iranica 147: 78–79; Saeidi-Mehrvarz (1999); (J. 1779)
188	Veronica paederotae Boiss.	Maz., Tehr., (Map 186)	2700–4100	7 (1)	AH	Hem.	1008	486	9.1	EN B1+B2 b (i,ii)	EN	AZ, MST, SP	F. Iranica 147: 144–145; Saeidi-Mehrvarz (1999); (N. 942)
189	Veronica siaretensis Lehmann	Gol., Maz., Talish, (Map 187)	400–1500	17 (3)	OH	Ther.	84,722	15,000	62.6	LC	LC	LF, MF	Akhani (1998); F. Iranica 147: 98–99; Saeidi-Mehrvarz (1999); (A. 10,403)
Polygalaceae
190	Polygala platyptera Bornm. & Gauba	Maz., (Map 188)	Sea level–2500	42 (4)	CH	Hem.	19,833	11,760	28.4	VU B1b(i)	LC	LF, MF, SF,	F. Iranica 124: 7–8; F. Iran 49: 15–17; (Abai 12,645 (IRAN))
Polygonaceae
191	Atraphaxis aucheri Jaub. & Spach	Gil., (Map 189)	300–600	4	MR	Cham.	55.222	27	3.2	CR B1b(i,ii,iii)	CR	STMX	F. Iranica 56: 35
192	Atraphaxis radkanensis S.Tavakkoli, Kaz.Osaloo & Mozaff.	Gol., (Map 190)	1400–1762	3	EH	Cham.	11.359	3	1.1	CR B1+B2b(i,ii)	CR	CRO, MRO	Tavakkoli et al. (2013); (A. 23,283)
193	Polygonum hyrcanicum Rech. f., (Akhani, 2005: Figs. 747–748)	Ardabil, Gil., Gol., Maz., N Khor., Qazv., Sem., (Map 191)	10–1750	33 (11)	OH	Hem.	179,483	40,000	90.3	LC	LC	DH, LMW, MMW	Akhani (1998); F. Iranica 56: 64–65; (A. 12,307)
194	Rumex kandavanicus (Rech. f.) Rech. f.	Maz./Alborz, (Map 192)	2800	1	AH	Hem.					DD	MH, SZ	F. Iranica 56: 9–10
Primulaceae
195	Cortusa matthioli L. subsp. iranica Iranshahr & Wendelbo	Maz., (Map 193)	2500–3000	2 (1)	AH	Hem.					CR C2b	MF?	F. Iran 25: 59; Iranshahr and Wendelbo (1976); (Re.&Ir. 16,804 (type-IRAN))
196	Cyclamen elegans Boiss. & Buhse (= C. coum auct. Flora Iranica 9: 30, 1965 non Miller)	Gil., Gol., Maz., Talish, (Map 194)	50–1750	33 (7)	OH	Geo.	62,663	35,000	55.3	LC	LC	CLF, CSF	Clennett (2002); F. Iranica 9: 30–31; F. Iran 25: 67–69; Grossheim (1967); (A. 13,795)
197	Dionysia aretioides (Lehm.) Boiss.	Gil., Maz., Sem., Tehr., (Map 195)	550–3200	28 (4)	AH	Cham./Hem.	9644	10,800	29.7	VU B1b(i)	LC	VC	F. Iranica 9: 17; F. Iran 25: 25; (A. 13,757)
198	Primula heterochroma Stapf, (Fig. 4A)	Gil., Gol., Maz., Sem. Talish, (Map 196)	60–2400	55 (23)	OH	Hem.	115,748	50,000	66.9	LC	LC	CLF, CMF, CSF	Akhani (1998); F. Iranica 9: 5; F. Iran 25: 11–13; (A. & al. 21,062)
Ranunculaceae
199	Aconitum iranshahrii H. Riedl	Maz., (Map 197)	2000–2500	1 (1)	CH	Geo.					EN A4c	MF	F. Iranica 171: 38; (Ir.&Re. 16,765 (IRAN))
200	Anemone caucasica Willd. ex Rupr., (Fig. 4B)	Cauc., Central Alborz, E Anatolia, Gil., Gol., Maz., Sem., Talish, (Map 198)	700–2900	53 (9)	XH	Geo.	378,696	87,500	152.4	LC	LC	GOWO, LF	Akhani (1998); F. Iranica 171: 224–225; Grossheim (1950); (A. 14,573)
201	Delphinium syncarpum Freyn	Maz., (Map 199)	–	1	CH	Ther.					DD	CSD ?	F. Iranica 171: 85–86
202	Delphinium ursinum Rech. f., (Akhani, 2005: Figs. 519–520)	Alborz, Gol., Maz., (Map 200)	700–2200	16 (3)	CEH	Geo.	37,165	25,344	48.4	NT	NT	AZ, FO, MWO	Akhani (1998); F. Iranica 171: 70–71; (A. 11,676)
203	Paraquilegia caespitosa (Boiss. & Hohen.) Drumm. & Hutch.	Maz., (Map 201)	2500–4800	10	AH	Hem.	3096	2646	20.7	EN B1b(i,ii)	EN	AZ, RO	F. Iranica 171: 8
204	Ranunculus cicutarius Schlechtend., (Akhani, 2005: Figs. 522–523)	Gil., Gol., Maz., Khor.?, Sem., Talish, (Map 202)	−26–2200	55 (4)	OH	Hem.	377,291	72,500	104.7	LC	LC	FM, LF, MF, MH, OSC, SF	Akhani (1998); F. Iranica 171: 159–160; Grossheim (1950); (A. 10,347)
205	Ranunculus dolosus Fisch. & C. A. Mey.	Gil., Gol., Maz., Talish, (Map 203)	−26–1200	13 (1)	OH	Geo./Hem.	92,641	27,500	64.3	LC	LC	MH	F. Iranica 171:137; Grossheim (1950); (A. 13,274)
Rhamnaceae
206	Frangula grandifolia (Fisch. & C. A. Mey.) Grubov	Gil., Gol., Maz., Talish, (Map 204)	200–1300	33 (8)	OH	Phan.	96,703	35,000	66.8	LC	LC	LF	Browicz (1984); F. Iranica 125: 26–27; F. Iran 55: 19; (A.20,271)
Rosaceae
207	Alchemilla amardica Rothm.	Gil. (unknown point), Maz., (Map 205)	2000–2900	4	AH	Hem.	1322	240	9	EN B1+B2ab (i,ii)	EN	AMW, SMW	F. Iranica 66: 145–146; F. Iran 6: 176; Naqinezhad et al. (2010)
208	Alchemilla citrina Fröhner	Ardabil, Gil., Gol., Maz., (Map 206)	2000–3200	10	OH	Hem.	43,814	12,500	60.2	NT	NT	AMW, MF, SMW	F. Iranica 66: 142–143; F. Iran 6: 173–174; Naqinezhad et al. (2010)
209	Alchemilla condensa Fröhner	Gil., Maz., (Map 207)	1800–2100	4	OH	Hem.	9899	6924	41.7	VU B1ab (i,ii)	VU	MMW	F. Iranica 66: 143–144; F. Iran 6: 175
210	Alchemilla gigantodus Fröhner	Aras., Gil., Gol., Maz., (Map 208)	1300–3400	9	OH	Hem.	57,831	12,500	72.7	LC	LC	AZ, SZ	F. Iranica 66: 141–142; F. Iran 6: 171–172; Naqinezhad et al. (2010)
211	Alchemilla hyrcana (Buser) Juz.	Aras., Gil., Maz., Talish, (Map 209)	1200–3600	9	WH	Hem.	38,232	20,000	77.3	NT	NT	AMW, SMW	F. Iranica 66: 137–138; F. Iran 6: 162–163; Grossheim (1952); Hamzeh'ee et al., 2010
212	Alchemilla mazandarana Naqinezhad & S.E. Fröhner	Maz., (Map 210)	2636–2950	7	AH	Hem.	303.290	75	5.4	EN B1+B2b(i,ii)	EN	RO, MMW, GOWO	Naqinezhad et al. (2017)
213	Alchemilla microscopica Fröhner	Gil., Maz., (Map 211)	1800–2300	4	OH	Hem.	20,958	6618	47	NT	NT	MMW	F. Iranica 66: 138–139; F. Iran 6: 167–168
214	Alchemilla pectiniloba Fröhner	Gil., Maz., (Map 212)	1800–3200	5	WH	Hem.	1085	2352	28.4	EN B1ab (i,ii)	EN	AZ, SZ	F. Iranica 66: 139; F. Iran 6: 168–169
215	Alchemilla plicatissima Fröhner	Gil., Gol., Maz., (Map 213)	1800–2600	4	OH	Hem.	5827	7500	52.3	VU B1ab (i,ii)	VU	MMW	F. Iranica 66: 143; F. Iran 6: 174-175
216	Cydonia oblonga Miller	Aras., Gil., Gol., Maz., Talish, Turk., (Map 214)	40–1700	53 (2)	XH (widely cultivated in other parts)	Phan.	2,350,436	107,500	123.5	LC	VU A4ac	CLF, CSF, OLF, OSF	Akhani (1998); Browicz (1996); F. Iranica 66: 26–27; F. Iran 6: 180–181; Grossheim (1952); (A. 13,671)
217	Potentilla petraea Willd. ex Schlecht.	Alborz, Ardabil/Gil., Gil., Maz., Talish, (Map 215)	2300–3100	10	WH	Hem.	12,196	7135	37.8	VU B1ab(i)	NT	RO, SZ	F. Iranica 66: 107–108; F. Iran 6: 135–136; Grossheim (1952)
218	Pyrus 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)	400–2400	57 (16)	OH	Phan.	183,343	55,000	70	LC	LC	CMF, MST, OMF, SC, RO	Akhani (1998); Browicz (1982); F. Iranica 66: 29; F. Iran 6: 183–185; (A. 12,317)
219	Pyrus grossheimii Fedor.	Gil., Maz., Talish, (Map 217)	700–2400	8	WH	Phan.	9285	9507	39.9	VU B1ab (i,ii)	NT	MF, MSC, SF, SMSC	F. Iranica 66: 30; F. Iran 6: 186–188; Grossheim (1952)
220	Pyrus hyrcana Fedor.	Aras., Gil., Maz., Talish, (Map 218)	40–2450	11	WH	Phan.	28,401	15,000	50.3	NT	NT	LF, LSC, LWO, MF, MSC, MWO	F. Iranica 66: 30–31; F. Iran 6: 188; Grossheim (1952)
221	Pyrus mazanderanica Schönbeck-Temesy	Alborz, Maz., (Map 219)	1900–2400	5	CH	Phan.		32	4.4	CR B1b(i,ii)	DD	MSC	F. Iranica 66: 32; F. Iran 6: 194
222	Rubus persicus Boiss. (= R. hyrcanus Juz.)	Gil., Gol., Maz., Talish, (Map 220)	30–1600	29	OH	Phan.	164,041	32,500	72.7	LC	LC	DH, FM, LF, RH, SF	F. Iranica 66: 70–71; F. Iran 6: 30–31; Grossheim (1952); Zieliński (1978)
223	Rubus raddeanus Focke	Aras., Gil., Gol., Maz., Talish, (Map 221)	50–2000	14 (1)	OH	Phan.	97,308	22,500	79.1	LC	LC	LF, RH	Akhani (1998); F. Iranica 66: 70–71; F. Iran 6: 28–30; Grossheim (1952); Zieliński (1978); (A. 11,822)
224	Spiraea sheikhii Zare	Maz., (Map 222)	200–250	2	CH	Phan.					DD	LF	Zare (2002)
Rubiaceae
225	Asperula gorganica Schönb.-Tem. & Ehrend.	Gol., (Map 223)	600–2080	15 (8)	EH	±Cham.	4136	2259	16.8	EN B1b(i,ii)	VU A4a	CRO, SC, VC, WO	Akhani (1998); F. Iranica 176: 149; (A. 11,143)
226	Asperula mazanderanica Ehrend.	Gol., Maz., (Map 224)	650–2700	9	EH	Cham./Hem.	6015	2400	20.1	VU B1ab (i,ii)	VU	MSC, RO,	F. Iranica 176: 148
227	Asperula microphylla Boiss.	Gil., Gol., Maz., (Map 225)	300–3400	16 (1)	AH	Cham./Hem.	14,584	10,086	41.4	VU B1b(i)	VU	CRO, WO	F. Iranica 176: 147–148; (J. 1723a)
228	Asperula sherardioides (Boiss.) Jaub. & Spach	Gil., (Map 226)	400–600	2	MR	Ther.					CR C2b	SMX	F. Iranica 176: 155–156; (Aucher 4675 (type-Photo-G))
229	Crucianella gilanica Trin. subsp. hirsuta (Ehrend.) Ehrend. & Schönb.-Tem.	Alborz, Maz., Tehr., (Map 227)	850–4100	19	AH	Hem.	489	128	4.4	EN B1+B2b(i,ii)	VU A4c	AZ, CRO, MF, MSC, SRO, VRO	F. Iranica 176: 85–86
230	Crucianella gilanica Trin. subsp. nezvensis Ehrend. & Schönb.-Tem.	Maz., Sem., (Map 228)	2000–3200	5	AH	Hem.	202	36	2.7	EN B1+B2ab (i,ii)	EN	MF, RO, SASC, SZ	F. Iranica 176: 83–84
231	Crucianella gilanica Trin. subsp. suleimanica Ehrend. & Schönb.-Tem.	Maz., (Map 229)	2000–3520	2	AH	Hem.		8	1.8	CR B1+B2b(i,ii)	EN A4c	AMW, RO	F. Iranica 176: 84
232	Crucianella platyphylla Ehrend. & Schönb.-Tem.	Gol., Sem., (Map 230)	1800–2500	6 (5)	EH	Cham./Hem.	3959	1620	17.6	EN B1b(i,ii)	VU A4	LVC, MWO	Akhani (1998); F. Iranica 176: 79; (A. 12,026)
233	Galium aucheri Boiss.	Gol., Maz., Sem., (Map 231)	3000–4100	8	AH	Hem.	6971	6845	37.3	VU B1ab (i,ii)	NT	AZ, CRO, SP	F. Iranica 176: 192
234	Galium caspicum Steven	Gil., Maz., Talish, (Map 232)	700–1850	7	WH	Hem.	26,349	9216	48	NT	NT	MH, SF	F. Iranica 176: 198
235	Galium elbursense Bornm. & Gauba ex Bornm.	Gil., Maz., (Map 233)	1700–1800	2	AH	Cham./Hem.		648	18.1	CR B1b(i,ii)	CR	RO, SC, SF	F. Iranica 176: 229–230
236	Galium kuetzingii Boiss. & Buhse	Gil., Talish, (Map 234)	250–400	5	WH	Cham./Hem.	796.529	1728	24.5	EN B1ab (i,ii)	EN	SAMX	F. Iranica 176: 204–205
237	Galium wendelboi Ehrend. & Schönb.-Tem.	Maz., (Map 235)	450–1800	4 (1)	CH	Hem.		2	1.1	CR B1+B2b(i,ii)	CR	MF, RO	F. Iranica 176: 228–229; (A. 13,858)
238	Phuopsis stylosa (Trin.) Hook. f., (Fig. 4E)	Gil., Gil./Ardabil, Maz., Talish, (Map 236)	200–2640	41 (19)	WH	Geo.	34,779	24,299	46.8	NT	LC	FO, MW, SC	F. Iranica 176: 73–74; (Gho. 4489)
Salicaceae
239	Populus caspica Bornm., (Fig. 4F)	Aras., Gil., Gol., Maz., N Khor., Talish, Turk., (Map 237)	300–1400	32 (15)	OH (widely cultivated outside of Caspian area)	Phan.	1,852,142	85,000	85.1	LC	LC	ALF, LF, RH	Akhani (1998); Browicz (1982); F. Iranica 65: 10–11; (A. 11,554)
Sapindaceae
240	Acer iranicum M. Mohtashamian & A. Rastegar/A. mazandaranicum Amini, Zare & Assadi 17	Gil., (Map 238)	950–1150	5	WH	Phan.	0.123	0.038	0.1	CR B1+B2b(i,ii)	DD	CSF	Amini et al. (2008); Mohtashamian et al. (2020)
241	Acer velutinum Boiss. [incl. var. velutinum & var. glabrescens (Boiss. & Buhse) E. Murray], (Fig. 4G)	Gil., Gol., Gol./N Khor., Maz., Talish, (Map 239)	0–2364	68 (31)	OH	Phan.	235,494	62,500	100.4	LC	LC	CLF, CMF, CSF, RH	Akhani (1998); Browicz (1982); F. Iranica 61: 4–5; (A. 16,614)
Saxifragaceae
242	Saxifraga iranica Bornm.	Maz., (Map 240)	2800–4200	8	AH	Cham./Hem.	568.867	124.503	5	EN B1+B2b(i,ii)	EN	AZ, RO	F. Iranica 42: 13; F. Iran 12: 10–11
243	Saxifraga mazanderanica Rech. f.	Gil., Maz., (Map 241)	1100–3000	9 (1)	AH	Cham./Hem.	8482	5300	32.6	VU B1ab(i)	EN C1	RO	F. Iranica 42: 10; F. Iran 12: 16–18; (Ma. 38,628 (IRAN))
244	Saxifraga ramsarica Jamzad	Maz., (Map 242)	2750	2 (1)	AH	Cham./Hem.		0.72	0.6	CR B1+B2b(i,ii)	CR	CRO	F. Iran 12: 11–12; (R.&Ma. 20,903 (holotype-TARI))
245	Saxifraga wendelboi Schönb.-Tem., (Fig. 4H)	Gol., Maz., Sem., (Map 243)	1900–3000	9 (4)	EH	Cham./Hem.	1879	1372	13.9	EN B1b(i,ii)	VU A4	VC	F. Iranica 42: 12; F. Iran 12: 9; (A. 13,611)
Scrophulariaceae
246	Scrophularia gaubae Bornm.	Gil., Gol., Maz., Khor., (Map 244)	400–3000	29 (8)	CEH	Hem.	54,119	32,500	64.1	LC	LC	CLF, CMF, CSF, FO, RH SC	Akhani (1998); F. Iranica 147: 242–243; F. Iran 68: 286; Joharchi et al. (2007); (A. 17,098)
247	Scrophularia megalantha Rech. f.	Maz., (Map 245)	40–600	17	CH	Hem.	3459	1239	10.6	EN B1b(i,ii)	EN	LF	F. Iranica 147: 231; F. Iran 68: 225
248	Scrophularia rostrata Boiss. & Buhse	Gil., Gil./Ardabil, Maz., Talish, (Map 246)	50–3010	23 (1)	WH	Hem.	36,273	24,956	47.7	NT	LC	LF, MF, SC, SF, SZ	F. Iranica 147: 241–242; F. Iran 68: 289; Grossheim (1967); (A. & al. 20,933)
249	Scrophularia vernalis L. subsp. clausii (Boiss. & Buhse) Grau, (Fig. 4I)	Gil., Gol., Maz., Sem., Talish, W Azar.?, (Map 247)	400–2640	41 (3)	OH	Hem.	220,922	45,000	78.5	LC	LC	MF, RO, SC	F. Iranica 147: 229–230; F. Iran 68: 219; Grossheim (1967); (A. 15,016)
250	Verbascum parsana Sotoodeh, Attar & Civeyrel	Gil., (Map 248)	1876	1	WH	Hem. (Bien.)					DD	OMF	Sotoodeh et al. (2016); (Moazzeni & Keshvari 34,941 (holotype-TUH))
251	Verbascum shahsavarensis Sotoodeh, Attar & Civeyrel	Maz., (Map 249)	1570	1	CH	Hem. (Bien. Peren.)					EN C2b	OMF	Sotoodeh et al. (2015)
252	Verbascum stachydiforme Boiss. & Buhse	Gil./Ardabil, Maz., Talish, (Map 250)	1200–2300	10 (1)	WH	Hem.	26,108	10,000	50.9	NT	VU A4c	MF, WO	F. Iranica 147: 43; F. Iran 68: 62; (Manutshehri 561)
253	Verbascum sublobatum Murb.	Gil., Gol., Maz., N Khor., (Map 251)	500–2100	15 (4)	CEH	Hem.	51,669	17,500	59	LC	VU A4c	FO, LF, MF, SC, SF, WO	Akhani (1998); F. Iranica 147: 42; F. Iran 68: 59; (A. 16,323)
Solanaceae
254	Atropa pallidiflora Schönbeck-Temesy	Gol., Maz., (Map 252)	35–2500	24	CEH	Hem.	30,200	23,040	48.1	NT	LC	LF, MF, SF,	F. Iranica 100: 41–42; F. Iran 24: 54–56
255	Solanum kieseritzkii C. A. Mey., (Fig. 4J)	Gil., Gol., Maz., Talish, (Map 253)	90–2400	25 (8)	OH	Cham.	105,432	32,500	67.5	LC	LC	CLF, CMF, CSF	Akhani (1998); F. Iranica 100: 13; F. Iran 24: 15; Grossheim (1967); (A. & al. 21,294)
Thymelaeaceae
256	Daphne mezereum L. subsp. rechingeri (Wendelbo) Halda	Gil., Maz., (Map 254)	500–1750	8 (1)	WH	Phan.	12,109	5710	33.8	VU B1ab (i,ii)	VU	LF, SC, SF	F. Iranica 95: 2–3; F. Iran 15: 4–6; (Sa.&Ga. 42,162 (IRAN))