Patterns, distribution, ecological features, and conservation issues of endemic plants in Armenia

Patterns, distribution, ecological features, and conservation issues of endemic plants in Armenia George M. Fayvush1, Karine Z. Janjughazyan1, Alla S. Aleksanyan1,2,3 1 Department of Geobotany and Ecological Physiology, A.L. Takhtajan Institute of Botany, National Academy of Sciences, Yerevan, Armenia 2 Armenian State University of Economics, Yerevan, Armenia 3 InternationalScientificEducationalCenterofNASRA,Yerevan,Armenia Corresponding author: George M. Fayvush ([email protected]) Editor Janet Franklin Received 29 July 2025♦ Accepted 24 September 2025♦ Published 10 October 2025 Frontiers of Biogeography 18, 2025, e167093|DOI 10.21425/fob.18.167093 Copyright George M. Fayvush et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. RESEARCH ARTICLE FRONTIERS OF BIOGEOGRAPHY Thescientificjournalof The International Biogeography Society Abstract This study aims to analyse plant endemism in Armenia. The richness of the republic’s flora was assessed in terms of the representation of endemic taxa. The distribution of endemic plant species was identified and analysed both within the republic and across altitudinal belts and major ecosystems. The greatest number of endemic taxa belongs to the families Rosaceae, Asteraceae, Fabaceae, and Poaceae, with the richest genera being Pyrus, Astragalus, and Psephellus. Most endemic species are confined to the arid regions of the republic, particularly the middle mountain belt and grassy ecosystems. Several “centres of endemism” were identified, where populations of multiple endemic species coexist in a small area. The relevant government organisations must prioritise these centres when creating new protected natural areas. Highlights • We identified 139 endemic taxa (species and/or subspecies) of vascular plants belonging to 27 families and 69 genera in modern Armenia. For Armenia, we calculated Bykov’s index (le = Ef/En, where Ef represents the real percentage of endemic taxa, and En denotes the expected or normal percentage of endemic taxa in the basic graph) to be 3.7, which is 3.7 times higher than the norm. This index surpasses that of renowned countries such as South Africa (3.6), Greece (2.1), Turkey, Cyprus (~2), and the Caucasus as a whole (1.3). This finding suggests that Armenia is one of the most significant territories in terms of plant diversity within the global biodiversity hotspot of the Caucasus. • The greatest number of endemics belongs to the Rosaceae family, particularly due to a high diversity within the genus Pyrus (10 species), followed by the Asteraceae, Fabaceae, and Poaceae families. The Caryophyllaceae (10 species) and Scrophulariaceae (eight species) families follow. At the genus level, Pyrus has 10 local endemic species, while the largest genus in Armenia, Astragalus, has seven endemic species, as does Psephellus, followed by Centaurea (six species). • The highest number of endemic vascular plant taxa is found in the Yerevan and Darelegis floristic regions, with fewer in the Aparan, Shirak, and Meghri regions. The majority of these taxa thrive in the arid regions of the Armeno-Iranian province of the Ancient Mediterranean subkingdom. Most of these species should be classified as neoendemics, formed in arid conditions. In contrast, the majority of endemics associated with the “Caucasian root,” which flourish in the more humid conditions of the remaining floristic regions of Armenia, are likely paleoendemics, preserved from glacial and postglacial periods. • The overwhelming majority of endemic vascular plant taxa in Armenia are confined to herbaceous habitats—84 species. Significantly fewer are found in petrophyte habitats and those with no or poorly developed vegetation—29 species, in forest and shrub habitats—25 species each, and in aquatic habitats, where only seven taxa are present. No endemic species or subspecies have been recorded in marsh habitats. This distribution can be attributed to the fact that herbaceous habitats occupy nearly two-thirds of Armenia’s territory. Furthermore, most endemics are associated with arid ecosystems, which are predominantly represented by habitats of this category. Keywords Armenia, centres of endemism, conservation of rare species, distribution of endemic plants, endemic plants Frontiers of Biogeography 18, 2025, e167093 George M. Fayvush et al. 2 Introduction Endemism is a crucial concept in modern biogeography. The rank and number of endemic taxa are key factors in biogeographical, floristic, and faunal zoning (Takhtajan 1986). The abundance of endemics is also an important criterion for identifying “hotspots of biodiversity” and developing nature conservation measures and systems of protected natural areas (Myers et al. 2000; Riemann and Ezcurra 2005; Lamoreux et al. 2006). However, as Anderson (1994) points out, researchers studying endemism face several challenges. The question arises: what is considered endemic? The simplest definition is as follows: “An endemic taxon is one that exists only in a specific territory.” However, this definition is insufficient. The range of any taxon, influenced by both biotic and abiotic factors, tends to change—either expanding or declining. Additionally, it is essential to consider the knowledge of the territory in which the taxon currently exists, as well as that of adjacent territories. On the basis of our experience, studying “foreign” territories often reveals that a taxon can no longer be considered endemic to its original area of distribution. Therefore, calling a taxon endemic without specifying the exact habitat is meaningless. For the first time, we calculated and analysed the number of endemic plant species growing exclusively in the Republic of Armenia (Fayvush 2007). The endemic taxa (species and subspecies) of vascular plants numbered 124. This figure was likely underestimated, as the processing of one of the largest families (Poaceae) for the multi-volume publication “Flora of Armenia” had not yet been completed. The eleventh volume of this publication, dedicated to this family, was published in 2010 (Gabrielian and Oganesian 2010). The Fifth National Report on Biodiversity (FNC 2014) listed 142 taxa of vascular plants endemic to Armenia. In recent years, botanical research, including floristic and taxonomic investigations, has continued in Armenia despite the challenging political and economic situation. Consequently, the composition of endemic taxa of vascular plants has changed significantly, and the distribution of many of these taxa has been clarified. The general results of these studies and their analysis are presented in this work. At present, we count 139 taxa as endemic to the flora of Armenia. Material and methods Study area Armenia is a republic in the South Caucasus, bordering Georgia, Azerbaijan, Turkey, and Iran. Its total area is 29,740 km², and it lies between 38°50' and 41°18'N latitude and between 43°27' and 46°37'E longitude. The distance from the northern to the southeast along the main axis is approximately 400 km. Armenia is predominantly mountainous, with an average altitude of 1850 m above sea level; the lowest point is 375 m (the extreme southeast and north of the Republic – Meghri and Idjevan floristic regions), and the highest point (mnt. Aragats in Central Armenia) reaches 4095 m above sea level (Fayvush 2023). Armenia is at the intersection of two biodiversity hotspots (Caucasian and Irano-Anatolian; Mittermeier et al. 2011) and two phytogeographical subkingdoms: Boreal (Euro-Siberian) and Ancient Mediterranean (Takhtadjan 1986). The country exhibits a wide range of climatic zones, characterised by a pronounced vertical succession of six basic climate types—from dry subtropical to alpine. The average annual temperature varies from -8°C in high-altitude mountainous regions (2500 m above sea level and higher) to 12–14°C in low-lying valleys (Fayvush 2023). The diversity of geological, soil and climatic conditions significantly influences Armenia’s varied vegetation. The lower mountain belt (480–1200 m) is predominantly covered by semi-desert formations, gypsophilous, or halophilous vegetation. The middle and upper mountain belts (1200–2200 m) are characterised by various types of steppe and forest vegetation; the subalpine and alpine belts (2200–4000 m) are primarily covered by meadows and alpine carpets (Magakyan 1941; Takhtadjan 1941; Fayvush 2006; Fayvush and Aleksanyan 2016). In 1954, A. L. Takhtadzhyan proposed a scheme of floristic zoning of Armenia; in this work we utilize a refined and revised scheme of this floristic division to indicate the distribution of plant species throughout the republic (Tamanyan and Fayvush 2009a). It is noteworthy that, according to our previous studies, the Shirak, Aparan, Yerevan, Darelegis, and Meghri floristic regions belong to the arid Armeno-Iranian floristic province (Ancient Mediterranean subkingdom), while the remaining regions are classified under the more mesophilic Caucasian province (Boreal subkingdom; Tamanyan and Fayvush 2009a; Fayvush et al. 2023a). Methods The basis for this study was the herbarium material stored in the Institute of Botany NAS RA (ERE).Its first collections date back to the beginning of the 20th century, and currently, it contains about 200,000 specimens of Armenian flora. We joined the research of the flora and vegetation of Armenia by leading botanists of Armenia and the Soviet Union more than 45 years ago. Field research was conducted using the route-expedition method, visiting all regions of Armenia during various periods of the growing season. These studies were conducted particularly intensively in the period 2002–2012, when Dr. E. Vitek (Natural History Museum Vienna) annually organised targeted expeditions to different regions of Armenia and Nagorno-Karabakh (Vitek et al. 2015) (Fig. 1). Herbarium material collected during these expeditions is currently stored in the herbariums of the Institute of Botany of the National Academy of Sciences of the Republic of Armenia (ERE) and the Natural History Museum Vienna (W). Geobotanical descriptions of plant communities Frontiers of Biogeography 18, 2025, e167093 Endemic plants in Armenia 3 were conducted, ecosystems and habitats of individual species were clarified, and both individual plants and ecosystems were photographed in their entirety. For the taxonomic analysis, we relied on the multi-volume publication “Flora of Armenia” (1954–2010) and recent works published by researchers from the Department of Taxonomy of Higher Plants at the Institute of Botany after A. Takhtajyan NAS RA. Additionally, all plant names were verified against the “Plants of the World Online” database (Royal Botanical Gardens Kew - https://powo. science.kew.org). In a few cases, more recent information was utilised; for example, Martin etal. (2023) led us to exclude Gundelia armeniaca from the list of Armenian endemics, as it was found in the Mush province of Turkey. For the analysis of the geographic distribution of endemic vascular plant taxa across Armenia, we employed our floristic zoning (Tamanyan and Fayvush 2009a, Fig. 2). It is important to note that only taxa growing exclusively within the administrative boundaries of Armenia were included in this analysis. If we considered species growing in Armenia and near its borders in neighbouring countries, their number could increase by more than 2–3 times. In determining the habitats of individual taxa, we utilised the EUNIS (European Nature Information System) habitats scheme (2022), which was adapted to the conditions of Armenia and published by us (Fayvush and Aleksanyan 2016; Fayvush et al. 2023b). It describes both habitats characteristic of Europe and new ones previously undescribed and specific to our republic. To assess the territory’s importance for biodiversity conservation, including the identification of protected areas, we decided to use Bykov’s index (Bykov 1983). This index (le) is calculated as Ef/En, where Ef represents the real percentage of endemic taxa, and En denotes the expected or normal percentage of endemic taxa as manifested in the basic nomogram (Fig. 3; Hawksworth and Kalin-Arroyo 1995: 173; Kier and Barthlott 2001; Georghiou and Delipetrou 2010; Rankou et al. 2013). For a unit area of 625 km², the percentage of endemism is set at 1. This endemism index has been calculated for countries such as Austria, Guinea, Libya, and the USA, while for countries like Denmark, Ireland, and Niger, it is significantly less than one (Cowling and Samways 1995). Conversely, New Caledonia and New Zealand have an “le” exceeding 80 (Nikolic 2001). For most European territories (islands and rather small countries), “le” is negative or close to one, with exceptions including the Canary Islands (10.68), Madeira Archipelago (10.36), Azores Archipelago (3.29), Crete (3.08, Bruchmann 2011), and Croatia (Nikolic 2001). Figure 1. Excursions Vitek et al. 2002–2012, visited localities in Armenia. Frontiers of Biogeography 18, 2025, e167093 George M. Fayvush et al. 4 Results As a result of this study, we identified 139 taxa (species and/or subspecies) of vascular plants belonging to 27 families (Table 1) and 69 genera (Table 2) in modern Armenia (see also Suppl. material 1). On the map of Armenia, with grid cells measuring 10 × 10 km, all squares containing endemic plant species have been highlighted, and their respective numbers noted (Fig. 4). Discussion The flora of the entire Caucasian ecoregion (including Armenia, Georgia, Azerbaijan, the Russian part of the Greater Caucasus, and neighbouring parts of Turkey and Iran) is estimated to comprise approximately 6,000–6,500 species of vascular plants, with 47% (2,791 species) being endemics (Solomon et al. 2013). Taxonomic analysis reveals 21 genera of vascular plants endemic to the Caucasus region. These genera are predominantly monotypic or oligotypic, with some having unclear relationships. The majority of these genera are endemic to the Greater Caucasus, while only four endemic genera exist in the Lesser Caucasus (including Armenia): Agasyllis (Apiaceae, one species), Grossheimia (Asteraceae, six species), Pseudovesicaria (Brassicaceae, one species), and Zuvanda (Brassicaceae, one species). Species of these genera primarily inhabit highland areas and are confined to stony habitats, screes, rocks, and subalpine meadows (Takhtajan 1986; Solomon et al. 2013). Figure 2. Floristic regions of Armenia* (Tamanyan and Fayvush 2009a): 1–Upper Akhuryan (five endemic taxa), 2–Shirak (20), 3– Lori (9), 4–Idjevan (14), 5–Aparan (26), 6–Yerevan (37), 7–Sevan (29), 8–Areguni (16), 9–Darelegis (39), 10–North Zangezur (19), 11–South Zangezur (17), 12–Meghri (22). *The intensity of colouration of floristic regions reflects the number of endemic taxa of vascular plants growing in them. Figure 3. Bykov’s (1983) nomogram for determining average endemism. Points above the diagonal line have lower than average endemism; points below have more (WCMC 1992). Table 1. Number of endemic taxa in the families of the flora of Armenia. N° Family Number of endemic taxa (species and/or subspecies) 1. Rosaceae 28 2. Asteraceae 22 3. Poaceae 12 4. Fabaceae 10 5. Caryophyllaceae 9 6. Scrophulariaceae 8 7. Brassicaceae 6 8. Orobanchaceae 6 9. Papaveraceae 5 10. Apiaceae 4 11. Colchicaceae 4 12. Alliaceae 3 13. Boraginaceae 2 14. Euphorbiaceae 2 15. Grossulariaceae 2 16. Iridaceae 2 17. Polygalaceae 2 18. Campanulaceae 1 19. Cyperaceae 1 20. Geraniaceae 1 21. Hyacinthaceae 1 22. Hypericaceae 1 23. Lamiaceae 1 24. Linaceae 1 25. Malvaceae 1 26. Plumbaginaceae 1 27. Ranunculaceae 1 Frontiers of Biogeography 18, 2025, e167093 Endemic plants in Armenia 5 Although generic endemism is characteristic of the entire Irano-Turanian region, it is particularly pronounced in its eastern part (Kamelin 1965; Hedge and Wendelbo 1970, 1978; Noroozi 2018). In the north-eastern part of the Armenian Highlands, Ancient Mediterranean roots give rise to eight genera: Peltariopsis, Pseudoanastatica, Takhtajaniella (all from Brassicaceae), Smyrniopsis, Stenotaenia, Szovitsia (all from Apiaceae), Huynhia (Boraginaceae), and Callicephalus(Asteraceae; Fayvush and Aleksanyan 2020). In Armenia, 139 species of flora (3.7%) are classified as local endemics. This paper does not delve into the taxonomic nuances of endemic species and subspecies of vascular plants in Armenian flora. However, among the 139 taxa, 23 (16%) are known only from type herbarium specimens collected over 30–40 years ago. Since then, information regarding their further existence is absent. From our perspective, these species are questionable; nonetheless, we include them in our analysis without awaiting future taxonomic conclusions regarding their status. Considering that the flora of Armenia comprises approximately 3,800 taxa of vascular plants (FNC 2014), the number of endemic taxa (species and subspecies) constitutes3.7% of the total flora. This percentage is quite high and exceeds that of many countries worldwide (WCMC 1992) but cannot be compared to that of countries such as Australia, New Zealand, Indonesia, South Africa, and Madagascar, where the expected percentage of endemism exceeds 66.7% (Callmander et al. 2011; Erenso and Maryo 2014). Furthermore, as demonstrated in numerous studies (Kier and Barthlott 2001; Berisha and Bytyqi 2021; Hobohm and Barker 2023, etc.), the percentage of endemism is significantly influenced by the area of the territory under study and its geographical context (island, mountainous country, and lowland Table 2. Number of endemic taxa in the genera of the flora of Armenia. N° Genus Number of endemic species and/or subspecies 1. PyrusL. 10 2. Astragalus L. 7 3. Psephellus Cass. 7 4. CentaureaL. 6 5. Papaver L. 5 6. Verbascum L. 5 7. Alchemilla L. 4 8. CrataegusL. 4 9. RosaL. 4 10. Allium L. 3 11. CotoneasterMedik. 3 12. Dianthus L. 3 13. Isatis L. 3 14. Onobrychis Hill 3 15. PhelipanchePomel 3 16. ColchicumL. 2 17. Euphorbia L. 2 18. Gladiolus L. 2 19. Gundelia L. 2 20. Merendera Ramond 2 21. Orobanche L. 2 22. Polygala L. 2 23. RibesL. 2 24. RosaL. 2 25. Scrophularia L. 2 26. Sonchus L. 2 27. Stipa L. 2 28. Tragopogon L. 2 29. Acantholimon Boiss. 1 30. Agrostis L. 1 31. Alcea L. 1 32. Allochrusa Bunge 1 33. Bromopsis (Dumort.) Fourr. 1 34. Bufonia L. 1 N° Genus Number of endemic species and/or subspecies 35. Bupleurum L. 1 36. CampanulaL. 1 37. CarexL. 1 38. CarthamusL. 1 39. CistancheHoffmanns. & Link 1 40. CousiniaCass. 1 41. Erodium L’Her. 1 42. Erysimum L. 1 43. Ferula L. 1 44. Festuca L. 1 45. Gaudinopsis (Boiss.) Eig 1 46. Grossheimia Sosn. & Takht. 1 47. Gypsophila L. 1 48. Hyalopoa (Tzvelev) Tzvelev 1 49. Hypericum L. 1 50. Koeleria Pers. 1 51. Linum L. 1 52. Minuartia L. 1 53. MyosotisL. 1 54. Nepeta L. 1 55. Ornithogalum L. 1 56. Paracolpodium (Tzvelev) Tzvelev 1 57. Poa L. 1 58. Potentilla L. 1 59. Pseudolysimachion Opiz 1 60. RanunculusL. 1 61. RorippaScop. 1 62. Scorzonera L. 1 63. Seseli L. 1 64. Silene L. 1 65. Smyrniopsis Boiss. 1 66. Symphytum L. 1 67. Thlaspi L. 1 68. Trisetum Pers. 1 69. Zingeria P. A. Smirn. 1 Frontiers of Biogeography 18, 2025, e167093 George M. Fayvush et al. 6 country). In this regard, for a realistic assessment of the territory of Armenia in terms of the richness of plant diversity and the representation of endemic taxa, we used Bykov’s index (Bykov 1983). For Armenia, we calculated the endemism index to be 3.7, which is 3.7 times higher than the norm. This index surpasses that of renowned countries such as South Africa (3.6), Greece (2.1), Turkey, and Cyprus (~2) (WCMC 1992; Georghiou and Delipetrou 2010; Rankou et al. 2013). Notably, our results, based on data presented in the work of Solomon et al. (2013), indicate that “le” for the Caucasus as a whole is 1.3 (with a percentage of endemism at 47% and an area of 170,405 km²). In terms of family spectrum, the largest families of Angiosperms in Armenia occupy the top four positions, with the greatest number of endemics belonging to the Rosaceae family (Table 1), particularly due to a high diversity within the genus Pyrus (10 species). This is followed by Asteraceae, Fabaceae, and Poaceae. The Caryophyllaceae (10 species) and Scrophulariaceae (eight species) families follow. At the genus level, Pyrus has 10 local endemic species, while the largest genus in Armenia, Astragalus, has seven endemic species, as does Psephellus, followed by Centaurea (six species). The flora of Armenia includes more than 30 species of the genus Pyrus, with nearly one-third being endemic to our republic (Akopian 2007). Intensive speciation processes in this genus are characteristic of the Lesser Caucasus (Fayvush et al. 2023b). Species of this genus can be found both as individual specimens in forest ecosystems and in “pear forests,” sparse forests dominated by several species of pears (according to Fayvush and Aleksanyan 2016–habitat F5.345). The genus Astragalus is typical of the entire Irano-Turanian region and is the largest genus in Armenia and the Lesser Caucasus. However, narrowly endemic species are relatively few (only 5%); the distribution ranges of most species are not confined to one country or floristic region. This suggests that species of this genus have better opportunities for dispersal and adaptation to new conditions (Fayvush et al. 2023b). If we consider the genus Centaurea sensu lato, which includes the species Centaurea, Psephellus, Grossheimia, and others, Armenia emerges as the most significant centre of species diversity for this genus across Eurasia (Wagenitz 1986; Gabrielian and Fayvush 1989; Fayvush and Aleksanyan 2020; Fayvush and Ghukasyan 2023; Fayvush et al. 2023b). Figure 4. Number of endemic taxa in Armenia (grid cells 10 × 10 km). Frontiers of Biogeography 18, 2025, e167093 Endemic plants in Armenia 7 Speciation within the genus Verbascum is also characteristic of the arid regions of the Ancient Mediterranean; however, most species endemic to Armenia are evidently of hybridogenic origin (four hybrid species from five endemics). Hybridisation plays an important role in the evolution of many taxonomic groups, but large-scale phylogenetic patterns of hybridisation are poorly understood (Whitney et al. 2010). The hybrid origin of these four species is indicated only by morphological features. We hope that DNA studies of the Verbascum species will help to more accurately identify their relationships, origin, and taxonomical status. Despite the lack of study, we nevertheless decided to include these species in our analysis of endemism. The distribution of endemic richness in different floristic regions is illustrated on the Armenia map (Fig. 2). On the basis of the premise that alloand autochthonous trends in the evolution of the Armenian flora are approximately equivalent (Fayvush 1990; Fayvush et al. 2023b), it is recognised that roughly half of the species migrated to Armenia from other regions, while the other half originated within the Armenian Highlands and partially in the Caucasus and Armenia itself. The highest number of endemic vascular plant taxa is found in the Yerevan and Darelegis floristic regions, with fewer in the Aparan, Shirak, and Meghri regions (Fig. 2, Suppl. material 1). The majority of these taxa thrive in the arid regions of the Armeno-Iranian province of the Ancient Mediterranean subkingdom. Most of these species should be classified as neoendemics, formed in arid conditions as a result of anexplosive diversification (Givnish 2015). In contrast, the majority of endemics associated with the “Caucasian root,” whichflourish in the more humid conditions of the remaining floristic regions of Armenia, are likely paleoendemics, preserved from glacial and postglacial periods. Regarding the distribution of endemic taxa along high-altitude belts, they are notably more concentrated in the middle and upper montane belts, which belts provide the greatest variety of habitats and are most conducive to speciation. Unlike in Iran, where the majority of endemic species are found in the highlands (Noroozi et al. 2018, 2019, 2021), the number of endemics in Armenia’s subalpine and alpine belts is minimal, likely due to the challenges of adapting new taxa to the conditions of closed plant communities in these belts (meadows, carpets, tall grasses), as nearly all highland endemics are confined to open habitats with sparse vegetation (category H according to the EUNIS system 2022). Many authors have directly correlated the diversity in a given territory to the richness of the overall taxonomic composition of plants, particularly endemics (Hobohm 2000; Guilhaumon et al. 2008; Alsterberg et al. 2017; Vladimirov et al. 2020; Al-sodany et al. 2024). This situation is especially relevant for narrow-local endemics, which often adapt to only one specific type of habitat (Hannus and Numers 2008; Sfenthourakis and Triantis 2009; Stein et al. 2014). In this context, Armenia is distinguished by not only the richness of its flora but also the diversity of its habitats. All the main habitat types of the EUNIS system are represented here, except for categories A and B (due to the absence of seas; Fayvush and Aleksanyan 2016; Fayvush et al. 2023a). It should also be noted that all the primary vegetation types and ecosystems common throughout the Caucasus are present in Armenia, with the exception of humid subtropical ecosystems. The overwhelming majority of endemic vascular plant taxa in Armenia are confined to herbaceous habitats (category E)—84 species. Significantly fewer are found in petrophyte habitats and those with no or poorly developed vegetation (H)—29 species, in forest and shrub habitats (F and G)—25 species each, and in aquatic habitats (C), where only seven taxa are present. No endemic species or subspecies have been recorded in marsh habitats (D) (Suppl. material 1). This distribution can be attributed to the fact that herbaceous habitats occupy nearly two-thirds of Armenia’s territory. Furthermore, most endemics are associated with arid ecosystems, which are predominantly represented by habitats of this category. Marsh habitats (D) are limited in area and highly fragmented. These habitats, along with aquatic ones (C), host a large number of rare relict species that have survived since the Ice Age. The conditions in these habitats are stable and conservative, which do not facilitate speciation processes. It has to be noted that two species are dominant in natural ecosystems: Ranunculus aragazi, a dominant species of alpine carpets on Mount Aragats (E4.3A29), and Smyrniosis armena, a dominant species of grass-forb steppes (E1.2E24). Conservation Endemism, defined as the restricted distribution of a taxon to a specific geographical area, is central to the study of biogeography. Understanding endemism not only addresses various evolutionary and biogeographical issues but also plays a crucial role in maintaining biodiversity and identifying priority areas for conservation (Meddour et al. 2023). Due to their genetic characteristics, most neoand paleoendemics of Armenia are narrow-range stenotopic species. Consequently, the populations of many of these species are threatened and may face extinction for various reasons, primarily due to anthropogenic factors such as habitat destruction from economic activities. Global climate change also poses a significant threat, as it may adversely affect endemics with narrow ecological amplitudes (Tamanyan et al. 2010; FNC 2014). In 2010, the Red Book of Plants of Armenia was published (Tamanyan et al. 2010), during which the status of populations of most endemic species was assessed according to International Union for Conservation of Nature (IUCN) criteria. Eighteen species were classified as Critically Endangered, 26 as Endangered, and two species as Vulnerable. All “doubtful” species were categorised as Data Deficient. A small number of species were assessed as Near Threatened and Least Concern. In recent years, botanists have described several new taxa from Armenia, almost all of which are endemic to the country and Frontiers of Biogeography 18, 2025, e167093 George M. Fayvush et al. 8 included in our analysis, but they have not yet been officially assessed regarding their extinction risk. Armenia has a well-developed network of specially protected natural areas, which includes three reserves, four national parks, and approximately 30 wildlife sanctuaries. According to our estimates, only 37 out of 139 endemic species are found within these protected areas. The remaining species require further research; the “questionable” species are pending a decision regarding their taxonomic status, newly described species need assessment according to IUCN criteria, and species classified as Data Deficient require additional field research to determine their presence or absence in nature. In general, all endemic and narrow-ranged species necessitate special measures to protect their habitats and clarify the possibilities for their ex-situ conservation. Fig. 4 shows a map of Armenia with 10 × 10 km grid cells, where the number of endemic plant species found here is shown in shades of different colors. The square in the South Zangezur floristic region stands out, as it hosts 16 endemic plant species. Unfortunately, this square does not encompass any protected areas, highlighting the need to expand the nearby Zangezur State Nature Sanctuary to preserve the ecosystems supporting these endemic plant species. The next centres rich in endemic plant species (9–12 taxa) are located in the Darelegis and Yerevan floristic regions, which are generally noted for their abundance of endemic species. As previously mentioned, these predominantly arid territories are primarily represented by neoendemics. A small portion of the endemism centres in these areas falls within the “Khosrov Forest” State Reserve, while the majority are situated outside existing protected areas, indicating the necessity to create new protected areas and expand those that currently exist. Less rich centres (6–9 taxa) are found in the Lake Sevan basin (Sevan and Areguni floristic regions), in the northeast (Ijevan floristic region), and in the southern part of the republic (Darelegis and South Zangezur floristic regions), all of which require attention to implement appropriate conservation measures. However, other less rich centres of endemic taxa accumulation (1–6) should not be overlooked. It is noteworthy that the most significant centres of plant endemism in Armenia coincide with the “biodiversity hotspots” identified earlier (Fayvush et al. 2013) based on the distribution of rare plant and animal species included in the Red Book of Armenia (Aghasyan and Kalashyan 2010; Tamanyan et al. 2010), taking into account the Important Plant Areas (Tamanyan and Fayvush 2009b; Asatryan and Fayvush 2013). Conclusion It has been established that 139 endemic species and subspecies of vascular plants from 27 families and 69 genera grow within the Republic of Armenia. The families Rosaceae, Asteraceae, Poaceae, and Fabaceae are the richest in endemics. The largest number of endemics is recorded in the genera Pyrus and Astragalus. The majority of endemic species are neoendemics, thriving in floristic regions belonging to the Ancient Mediterranean subkingdom of flora, particularly in grassy, arid habitats. Species from more humid habitats are likely paleoendemics, with their distribution primarily limited to floristic regions belonging to the Boreal subkingdom of flora. In contrast to Northern Iran, the Armenian highlands are relatively poor in endemic species formed and adapted to closed grass ecosystems (meadows, carpets, and tall grasses). In the alpine and subalpine belts, endemics are predominantly found in open communities of rocks and screes. Endemic plant species require further in-depth studies. Some of these “questionable” species require additional taxonomic and floristic research. In general, all endemic species require the development of specialised measures for their conservation in both natural and ex-situ conditions. The identified centres of endemism—territories with the greatest wealth of endemic taxa—should attract the attention of environmental agencies (first of all, the Ministry of Environment of RA, as well as World Wildlife Fund (WWF), Foundation for the Preservation of Wildlife and Cultural Assets (FPWC), etc.) to organise protected areas for preserving the habitats of these endemics. Acknowledgements The research was partly supported by the Higher Education and Science Committee of MESCS RA (Research project № 21AG-1F004). Author contributions All authors contributed to the study’s conception and design. Material preparation and data collection were conducted by Karine Janjughazyan, while analysis was performed by George Fayvush and Alla Aleksanyan. The first draft of the manuscript was written by George Fayvush, and all authors provided feedback on previous versions. All authors read and approved the final manuscript. 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