Pa e ns, dis ibu ion, ecological ea u es, and conse a ion issues
o endemic plan s in A menia
Geo ge M. Fay ush1, Ka ine Z. Janjughazyan1, Alla S. Aleksanyan1,2,3
1 Depa men o Geobo any and Ecological Physiology, A.L. Takh ajan Ins i u e o Bo any, Na ional Academy o Sciences, Ye e an, A menia
2 A menian S a e Uni e si y o Economics, Ye e an, A menia
3 In e na ionalScien i icEduca ionalCen e o NASRA,Ye e an,A menia
Co esponding au ho : Geo ge M. Fay ush ([email p o ec ed])
Edi o Jane F anklin
Recei ed
29 July 2025♦
Accep ed
24 Sep embe 2025♦
Published
10 Oc obe 2025
F on ie s o Biogeog aphy 18, 2025,
e167093|DOI 10.21425/ ob.18.167093
Copy igh Geo ge M. Fay ush e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion
License (CC BY 4.0), which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho
and sou ce a e c edi ed.
RESEARCH ARTICLE
FRONTIERS OF
BIOGEOGRAPHY
Thescien i icjou nalo
The In e na ional Biogeog aphy Socie y
Abs ac
This s udy aims o analyse plan endemism in A menia.
The ichness o he epublic’s lo a was assessed in e ms
o he ep esen a ion o endemic axa. The dis ibu ion o
endemic plan species was iden i ied and analysed bo h
wi hin he epublic and ac oss al i udinal bel s and majo
ecosys ems. The g ea es numbe o endemic axa be-
longs o he amilies Rosaceae, As e aceae, Fabaceae,
and Poaceae, wi h he iches gene a being Py us, As a-
galus, and Psephellus. Mos endemic species a e con ined
o he a id egions o he epublic, pa icula ly he middle
moun ain bel and g assy ecosys ems. Se e al “cen es o
endemism” we e iden i ied, whe e popula ions o mul iple
endemic species coexis in a small a ea. The ele an go -
e nmen o ganisa ions mus p io i ise hese cen es when
c ea ing new p o ec ed na u al a eas.
Highligh s
• We iden i ied 139 endemic axa (species and/o sub-
species) o ascula plan s belonging o 27 amilies
and 69 gene a in mode n A menia. Fo A menia, we cal-
cula ed Byko ’s index (le = E /En, whe e E ep esen s
he eal pe cen age o endemic axa, and En deno es
he expec ed o no mal pe cen age o endemic axa
in he basic g aph) o be 3.7, which is 3.7 imes highe
han he no m. This index su passes ha o enowned
coun ies such as Sou h A ica (3.6), G eece (2.1), Tu -
key, Cyp us (~2), and he Caucasus as a whole (1.3).
This inding sugges s ha A menia is one o he mos
signi ican e i o ies in e ms o plan di e si y wi hin
he global biodi e si y ho spo o he Caucasus.
• The g ea es numbe o endemics belongs o he
Rosaceae amily, pa icula ly due o a high di e si y
wi hin he genus Py us (10 species), ollowed by he
As e aceae, Fabaceae, and Poaceae amilies. The
Ca yophyllaceae (10 species) and Sc ophula iaceae
(eigh species) amilies ollow. A he genus le el, Py us
has 10 local endemic species, while he la ges genus
in A menia, As agalus, has se en endemic species, as
does Psephellus, ollowed by Cen au ea (six species).
• The highes numbe o endemic ascula plan axa
is ound in he Ye e an and Da elegis lo is ic egions,
wi h ewe in he Apa an, Shi ak, and Megh i egions.
The majo i y o hese axa h i e in he a id egions
o he A meno-I anian p o ince o he Ancien Med-
i e anean subkingdom. Mos o hese species
should be classi ied as neoendemics, o med in a id
condi ions. In con as , he majo i y o endemics as-
socia ed wi h he “Caucasian oo ,” which lou ish in
he mo e humid condi ions o he emaining lo is ic
egions o A menia, a e likely paleoendemics, p e-
se ed om glacial and pos glacial pe iods.
• The o e whelming majo i y o endemic ascula
plan axa in A menia a e con ined o he baceous
habi a s—84 species. Signi ican ly ewe a e ound in
pe ophy e habi a s and hose wi h no o poo ly de-
eloped ege a ion—29 species, in o es and sh ub
habi a s—25 species each, and in aqua ic habi a s,
whe e only se en axa a e p esen . No endemic spe-
cies o subspecies ha e been eco ded in ma sh hab-
i a s. This dis ibu ion can be a ibu ed o he ac
ha he baceous habi a s occupy nea ly wo- hi ds o
A menia’s e i o y. Fu he mo e, mos endemics a e
associa ed wi h a id ecosys ems, which a e p edom-
inan ly ep esen ed by habi a s o his ca ego y.
Keywo ds
A menia, cen es o endemism, conse a ion o a e species,
dis ibu ion o endemic plan s, endemic plan s
F on ie s o Biogeog aphy 18, 2025, e167093
Geo ge M. Fay ush e al.
2
In oduc ion
Endemism is a c ucial concep in mode n biogeog aphy.
The ank and numbe o endemic axa a e key ac o s in
biogeog aphical, lo is ic, and aunal zoning (Takh ajan
1986). The abundance o endemics is also an impo an
c i e ion o iden i ying “ho spo s o biodi e si y” and
de eloping na u e conse a ion measu es and sys ems
o p o ec ed na u al a eas (Mye s e al. 2000; Riemann
and Ezcu a 2005; Lamo eux e al. 2006). Howe e , as
Ande son (1994) poin s ou , esea che s s udying ende-
mism ace se e al challenges. The ques ion a ises: wha
is conside ed endemic? The simples de ini ion is as ol-
lows: “An endemic axon is one ha exis s only in a spe-
ci ic e i o y.” Howe e , his de ini ion is insu icien . The
ange o any axon, in luenced by bo h bio ic and abio ic
ac o s, ends o change—ei he expanding o declining.
Addi ionally, i is essen ial o conside he knowledge o
he e i o y in which he axon cu en ly exis s, as well as
ha o adjacen e i o ies. On he basis o ou expe ience,
s udying “ o eign” e i o ies o en e eals ha a axon
can no longe be conside ed endemic o i s o iginal a ea
o dis ibu ion. The e o e, calling a axon endemic wi hou
speci ying he exac habi a is meaningless.
Fo he i s ime, we calcula ed and analysed he num-
be o endemic plan species g owing exclusi ely in he
Republic o A menia (Fay ush 2007). The endemic axa
(species and subspecies) o ascula plan s numbe ed
124. This igu e was likely unde es ima ed, as he p o-
cessing o one o he la ges amilies (Poaceae) o he
mul i- olume publica ion “Flo a o A menia” had no ye
been comple ed. The ele en h olume o his publica ion,
dedica ed o his amily, was published in 2010 (Gab ie-
lian and Oganesian 2010). The Fi h Na ional Repo on
Biodi e si y (FNC 2014) lis ed 142 axa o ascula plan s
endemic o A menia.
In ecen yea s, bo anical esea ch, including lo is ic and
axonomic in es iga ions, has con inued in A menia despi e
he challenging poli ical and economic si ua ion. Conse-
quen ly, he composi ion o endemic axa o ascula plan s
has changed signi ican ly, and he dis ibu ion o many o
hese axa has been cla i ied. The gene al esul s o hese
s udies and hei analysis a e p esen ed in his wo k. A p es-
en , we coun 139 axa as endemic o he lo a o A menia.
Ma e ial and me hods
S udy a ea
A menia is a epublic in he Sou h Caucasus, bo de -
ing Geo gia, Aze baijan, Tu key, and I an. I s o al a ea is
29,740 km², and i lies be ween 38°50' and 41°18'N la i-
ude and be ween 43°27' and 46°37'E longi ude. The dis-
ance om he no he n o he sou heas along he main
axis is app oxima ely 400 km. A menia is p edominan ly
moun ainous, wi h an a e age al i ude o 1850 m abo e
sea le el; he lowes poin is 375 m ( he ex eme sou heas
and no h o he Republic – Megh i and Idje an lo is ic
egions), and he highes poin (mn . A aga s in Cen al A -
menia) eaches 4095 m abo e sea le el (Fay ush 2023).
A menia is a he in e sec ion o wo biodi e si y
ho spo s (Caucasian and I ano-Ana olian; Mi e meie e
al. 2011) and wo phy ogeog aphical subkingdoms: Bo e-
al (Eu o-Sibe ian) and Ancien Medi e anean (Takh adjan
1986). The coun y exhibi s a wide ange o clima ic zones,
cha ac e ised by a p onounced e ical succession o six
basic clima e ypes— om d y sub opical o alpine. The
a e age annual empe a u e a ies om -8°C in high-al i-
ude moun ainous egions (2500 m abo e sea le el and
highe ) o 12–14°C in low-lying alleys (Fay ush 2023).
The di e si y o geological, soil and clima ic condi-
ions signi ican ly in luences A menia’s a ied ege a ion.
The lowe moun ain bel (480–1200 m) is p edominan -
ly co e ed by semi-dese o ma ions, gypsophilous, o
halophilous ege a ion. The middle and uppe moun ain
bel s (1200–2200 m) a e cha ac e ised by a ious ypes
o s eppe and o es ege a ion; he subalpine and alpine
bel s (2200–4000 m) a e p ima ily co e ed by meadows
and alpine ca pe s (Magakyan 1941; Takh adjan 1941;
Fay ush 2006; Fay ush and Aleksanyan 2016).
In 1954, A. L. Takh adzhyan p oposed a scheme o lo-
is ic zoning o A menia; in his wo k we u ilize a e ined
and e ised scheme o his lo is ic di ision o indica e
he dis ibu ion o plan species h oughou he epublic
(Tamanyan and Fay ush 2009a).
I is no ewo hy ha , acco ding o ou p e ious s udies,
he Shi ak, Apa an, Ye e an, Da elegis, and Megh i lo is ic
egions belong o he a id A meno-I anian lo is ic p o -
ince (Ancien Medi e anean subkingdom), while he e-
maining egions a e classi ied unde he mo e mesophilic
Caucasian p o ince (Bo eal subkingdom; Tamanyan and
Fay ush 2009a; Fay ush e al. 2023a).
Me hods
The basis o his s udy was he he ba ium ma e ial
s o ed in he Ins i u e o Bo any NAS RA (ERE).I s i s
collec ions da e back o he beginning o he 20 h cen u-
y, and cu en ly, i con ains abou 200,000 specimens o
A menian lo a. We joined he esea ch o he lo a and
ege a ion o A menia by leading bo anis s o A menia
and he So ie Union mo e han 45 yea s ago. Field e-
sea ch was conduc ed using he ou e-expedi ion me h-
od, isi ing all egions o A menia du ing a ious pe iods
o he g owing season. These s udies we e conduc ed
pa icula ly in ensi ely in he pe iod 2002–2012, when D .
E. Vi ek (Na u al His o y Museum Vienna) annually o gan-
ised a ge ed expedi ions o di e en egions o A menia
and Nago no-Ka abakh (Vi ek e al. 2015) (Fig. 1). He -
ba ium ma e ial collec ed du ing hese expedi ions is cu -
en ly s o ed in he he ba iums o he Ins i u e o Bo any
o he Na ional Academy o Sciences o he Republic o
A menia (ERE) and he Na u al His o y Museum Vienna
(W). Geobo anical desc ip ions o plan communi ies
F on ie s o Biogeog aphy 18, 2025, e167093
Endemic plan s in A menia
3
we e conduc ed, ecosys ems and habi a s o indi idual
species we e cla i ied, and bo h indi idual plan s and eco-
sys ems we e pho og aphed in hei en i e y.
Fo he axonomic analysis, we elied on he mul i- ol-
ume publica ion “Flo a o A menia” (1954–2010) and e-
cen wo ks published by esea che s om he Depa men
o Taxonomy o Highe Plan s a he Ins i u e o Bo any
a e A. Takh ajyan NAS RA. Addi ionally, all plan names
we e e i ied agains he “Plan s o he Wo ld Online”
da abase (Royal Bo anical Ga dens Kew - h ps://powo.
science.kew.o g). In a ew cases, mo e ecen in o ma-
ion was u ilised; o example, Ma in e al. (2023) led us
o exclude Gundelia a meniaca om he lis o A menian
endemics, as i was ound in he Mush p o ince o Tu key.
Fo he analysis o he geog aphic dis ibu ion o endemic
ascula plan axa ac oss A menia, we employed ou lo-
is ic zoning (Tamanyan and Fay ush 2009a, Fig. 2). I is
impo an o no e ha only axa g owing exclusi ely wi hin
he adminis a i e bounda ies o A menia we e included in
his analysis. I we conside ed species g owing in A menia
and nea i s bo de s in neighbou ing coun ies, hei num-
be could inc ease by mo e han 2–3 imes.
In de e mining he habi a s o indi idual axa, we u i-
lised he EUNIS (Eu opean Na u e In o ma ion Sys em)
habi a s scheme (2022), which was adap ed o he condi-
ions o A menia and published by us (Fay ush and Alek-
sanyan 2016; Fay ush e al. 2023b). I desc ibes bo h
habi a s cha ac e is ic o Eu ope and new ones p e iously
undesc ibed and speci ic o ou epublic.
To assess he e i o y’s impo ance o biodi e si y
conse a ion, including he iden i ica ion o p o ec ed a -
eas, we decided o use Byko ’s index (Byko 1983). This
index (le) is calcula ed as E /En, whe e E ep esen s he
eal pe cen age o endemic axa, and En deno es he ex-
pec ed o no mal pe cen age o endemic axa as mani-
es ed in he basic nomog am (Fig. 3; Hawkswo h and
Kalin-A oyo 1995: 173; Kie and Ba hlo 2001; Geo gh-
iou and Delipe ou 2010; Rankou e al. 2013). Fo a uni
a ea o 625 km², he pe cen age o endemism is se a
1. This endemism index has been calcula ed o coun-
ies such as Aus ia, Guinea, Libya, and he USA, while
o coun ies like Denma k, I eland, and Nige , i is sig-
ni ican ly less han one (Cowling and Samways 1995).
Con e sely, New Caledonia and New Zealand ha e an “le”
exceeding 80 (Nikolic 2001). Fo mos Eu opean e i o-
ies (islands and a he small coun ies), “le” is nega i e
o close o one, wi h excep ions including he Cana y Is-
lands (10.68), Madei a A chipelago (10.36), Azo es A chi-
pelago (3.29), C e e (3.08, B uchmann 2011), and C oa ia
(Nikolic 2001).
Figu e 1. Excu sions Vi ek e al. 2002–2012, isi ed locali ies in A menia.
F on ie s o Biogeog aphy 18, 2025, e167093
Geo ge M. Fay ush e al.
4
Resul s
As a esul o his s udy, we iden i ied 139 axa (spe-
cies and/o subspecies) o ascula plan s belonging o
27 amilies (Table 1) and 69 gene a (Table 2) in mode n
A menia (see also Suppl. ma e ial 1).
On he map o A menia, wi h g id cells measu ing 10 ×
10 km, all squa es con aining endemic plan species ha e
been highligh ed, and hei espec i e numbe s no ed (Fig. 4).
Discussion
The lo a o he en i e Caucasian eco egion (including A -
menia, Geo gia, Aze baijan, he Russian pa o he G ea e
Caucasus, and neighbou ing pa s o Tu key and I an) is
es ima ed o comp ise app oxima ely 6,000–6,500 species
o ascula plan s, wi h 47% (2,791 species) being endem-
ics (Solomon e al. 2013). Taxonomic analysis e eals 21
gene a o ascula plan s endemic o he Caucasus egion.
These gene a a e p edominan ly mono ypic o oligo ypic,
wi h some ha ing unclea ela ionships. The majo i y o
hese gene a a e endemic o he G ea e Caucasus, while
only ou endemic gene a exis in he Lesse Caucasus (in-
cluding A menia): Agasyllis (Apiaceae, one species), G oss-
heimia (As e aceae, six species), Pseudo esica ia (B as-
sicaceae, one species), and Zu anda (B assicaceae, one
species). Species o hese gene a p ima ily inhabi highland
a eas and a e con ined o s ony habi a s, sc ees, ocks, and
subalpine meadows (Takh ajan 1986; Solomon e al. 2013).
Figu e 2. Flo is ic egions o A menia* (Tamanyan and Fay ush
2009a): 1–Uppe Akhu yan ( i e endemic axa), 2–Shi ak (20), 3–
Lo i (9), 4–Idje an (14), 5–Apa an (26), 6–Ye e an (37), 7–Se an
(29), 8–A eguni (16), 9–Da elegis (39), 10–No h Zangezu (19),
11–Sou h Zangezu (17), 12–Megh i (22). *The in ensi y o col-
ou a ion o lo is ic egions e lec s he numbe o endemic axa
o ascula plan s g owing in hem.
Figu e 3. Byko ’s (1983) nomog am o de e mining a e age en-
demism. Poin s abo e he diagonal line ha e lowe han a e age
endemism; poin s below ha e mo e (WCMC 1992).
Table 1. Numbe o endemic axa in he amilies o he lo a o
A menia.
N° Family Numbe o endemic axa
(species and/o subspecies)
1. Rosaceae 28
2. As e aceae 22
3. Poaceae 12
4. Fabaceae 10
5. Ca yophyllaceae 9
6. Sc ophula iaceae 8
7. B assicaceae 6
8. O obanchaceae 6
9. Papa e aceae 5
10. Apiaceae 4
11. Colchicaceae 4
12. Alliaceae 3
13. Bo aginaceae 2
14. Eupho biaceae 2
15. G ossula iaceae 2
16. I idaceae 2
17. Polygalaceae 2
18. Campanulaceae 1
19. Cype aceae 1
20. Ge aniaceae 1
21. Hyacin haceae 1
22. Hype icaceae 1
23. Lamiaceae 1
24. Linaceae 1
25. Mal aceae 1
26. Plumbaginaceae 1
27. Ranunculaceae 1
F on ie s o Biogeog aphy 18, 2025, e167093
Endemic plan s in A menia
5
Al hough gene ic endemism is cha ac e is ic o he en-
i e I ano-Tu anian egion, i is pa icula ly p onounced in
i s eas e n pa (Kamelin 1965; Hedge and Wendelbo 1970,
1978; No oozi 2018). In he no h-eas e n pa o he A -
menian Highlands, Ancien Medi e anean oo s gi e ise
o eigh gene a: Pel a iopsis, Pseudoanas a ica, Takh a-
janiella (all om B assicaceae), Smy niopsis, S eno aenia,
Szo i sia (all om Apiaceae), Huynhia (Bo aginaceae), and
Callicephalus(As e aceae; Fay ush and Aleksanyan 2020).
In A menia, 139 species o lo a (3.7%) a e classi ied
as local endemics. This pape does no del e in o he ax-
onomic nuances o endemic species and subspecies o
ascula plan s in A menian lo a. Howe e , among he
139 axa, 23 (16%) a e known only om ype he ba ium
specimens collec ed o e 30–40 yea s ago. Since hen, in-
o ma ion ega ding hei u he exis ence is absen . F om
ou pe spec i e, hese species a e ques ionable; none he-
less, we include hem in ou analysis wi hou awai ing u-
u e axonomic conclusions ega ding hei s a us.
Conside ing ha he lo a o A menia comp ises ap-
p oxima ely 3,800 axa o ascula plan s (FNC 2014),
he numbe o endemic axa (species and subspecies)
cons i u es3.7% o he o al lo a. This pe cen age is
qui e high and exceeds ha o many coun ies wo ldwide
(WCMC 1992) bu canno be compa ed o ha o coun-
ies such as Aus alia, New Zealand, Indonesia, Sou h
A ica, and Madagasca , whe e he expec ed pe cen age
o endemism exceeds 66.7% (Callmande e al. 2011; E -
enso and Ma yo 2014). Fu he mo e, as demons a ed
in nume ous s udies (Kie and Ba hlo 2001; Be isha
and By yqi 2021; Hobohm and Ba ke 2023, e c.), he
pe cen age o endemism is signi ican ly in luenced by
he a ea o he e i o y unde s udy and i s geog aph-
ical con ex (island, moun ainous coun y, and lowland
Table 2. Numbe o endemic axa in he gene a o he lo a o A menia.
N° Genus
Numbe o endemic
species and/o
subspecies
1. Py usL. 10
2. As agalus L. 7
3. Psephellus Cass. 7
4. Cen au eaL. 6
5. Papa e L. 5
6. Ve bascum L. 5
7. Alchemilla L. 4
8. C a aegusL. 4
9. RosaL. 4
10. Allium L. 3
11. Co oneas e Medik. 3
12. Dian hus L. 3
13. Isa is L. 3
14. Onob ychis Hill 3
15. PhelipanchePomel 3
16. ColchicumL. 2
17. Eupho bia L. 2
18. Gladiolus L. 2
19. Gundelia L. 2
20. Me ende a Ramond 2
21. O obanche L. 2
22. Polygala L. 2
23. RibesL. 2
24. RosaL. 2
25. Sc ophula ia L. 2
26. Sonchus L. 2
27. S ipa L. 2
28. T agopogon L. 2
29. Acan holimon Boiss. 1
30. Ag os is L. 1
31. Alcea L. 1
32. Alloch usa Bunge 1
33. B omopsis (Dumo .) Fou . 1
34. Bu onia L. 1
N° Genus
Numbe o endemic
species and/o
subspecies
35. Bupleu um L. 1
36. CampanulaL. 1
37. Ca exL. 1
38. Ca hamusL. 1
39. Cis ancheHo manns. & Link 1
40. CousiniaCass. 1
41. E odium L’He . 1
42. E ysimum L. 1
43. Fe ula L. 1
44. Fes uca L. 1
45. Gaudinopsis (Boiss.) Eig 1
46. G ossheimia Sosn. & Takh . 1
47. Gypsophila L. 1
48. Hyalopoa (Tz ele ) Tz ele 1
49. Hype icum L. 1
50. Koele ia Pe s. 1
51. Linum L. 1
52. Minua ia L. 1
53. Myoso isL. 1
54. Nepe a L. 1
55. O ni hogalum L. 1
56. Pa acolpodium (Tz ele ) Tz ele 1
57. Poa L. 1
58. Po en illa L. 1
59. Pseudolysimachion Opiz 1
60. RanunculusL. 1
61. Ro ippaScop. 1
62. Sco zone a L. 1
63. Seseli L. 1
64. Silene L. 1
65. Smy niopsis Boiss. 1
66. Symphy um L. 1
67. Thlaspi L. 1
68. T ise um Pe s. 1
69. Zinge ia P. A. Smi n. 1
F on ie s o Biogeog aphy 18, 2025, e167093
Geo ge M. Fay ush e al.
6
coun y). In his ega d, o a ealis ic assessmen o he
e i o y o A menia in e ms o he ichness o plan di-
e si y and he ep esen a ion o endemic axa, we used
Byko ’s index (Byko 1983). Fo A menia, we calcula ed
he endemism index o be 3.7, which is 3.7 imes highe
han he no m. This index su passes ha o enowned
coun ies such as Sou h A ica (3.6), G eece (2.1), Tu -
key, and Cyp us (~2) (WCMC 1992; Geo ghiou and De-
lipe ou 2010; Rankou e al. 2013). No ably, ou esul s,
based on da a p esen ed in he wo k o Solomon e al.
(2013), indica e ha “le” o he Caucasus as a whole is
1.3 (wi h a pe cen age o endemism a 47% and an a ea
o 170,405 km²).
In e ms o amily spec um, he la ges amilies o
Angiospe ms in A menia occupy he op ou posi ions,
wi h he g ea es numbe o endemics belonging o he
Rosaceae amily (Table 1), pa icula ly due o a high di e -
si y wi hin he genus Py us (10 species). This is ollowed by
As e aceae, Fabaceae, and Poaceae. The Ca yophyllace-
ae (10 species) and Sc ophula iaceae (eigh species) am-
ilies ollow. A he genus le el, Py us has 10 local endemic
species, while he la ges genus in A menia, As agalus,
has se en endemic species, as does Psephellus, ollowed
by Cen au ea (six species).
The lo a o A menia includes mo e han 30 species o
he genus Py us, wi h nea ly one- hi d being endemic o
ou epublic (Akopian 2007). In ensi e specia ion p ocess-
es in his genus a e cha ac e is ic o he Lesse Cauca-
sus (Fay ush e al. 2023b). Species o his genus can be
ound bo h as indi idual specimens in o es ecosys ems
and in “pea o es s,” spa se o es s domina ed by se e al
species o pea s (acco ding o Fay ush and Aleksanyan
2016–habi a F5.345).
The genus As agalus is ypical o he en i e I ano-Tu -
anian egion and is he la ges genus in A menia and he
Lesse Caucasus. Howe e , na owly endemic species a e
ela i ely ew (only 5%); he dis ibu ion anges o mos
species a e no con ined o one coun y o lo is ic egion.
This sugges s ha species o his genus ha e be e op-
po uni ies o dispe sal and adap a ion o new condi ions
(Fay ush e al. 2023b).
I we conside he genus Cen au ea sensu la o, which
includes he species Cen au ea, Psephellus, G ossheimia,
and o he s, A menia eme ges as he mos signi ican cen-
e o species di e si y o his genus ac oss Eu asia (Wa-
geni z 1986; Gab ielian and Fay ush 1989; Fay ush and
Aleksanyan 2020; Fay ush and Ghukasyan 2023; Fay ush
e al. 2023b).
Figu e 4. Numbe o endemic axa in A menia (g id cells 10 × 10 km).
F on ie s o Biogeog aphy 18, 2025, e167093
Endemic plan s in A menia
7
Specia ion wi hin he genus Ve bascum is also cha ac-
e is ic o he a id egions o he Ancien Medi e anean;
howe e , mos species endemic o A menia a e e iden ly o
hyb idogenic o igin ( ou hyb id species om i e endem-
ics). Hyb idisa ion plays an impo an ole in he e olu ion
o many axonomic g oups, bu la ge-scale phylogene ic
pa e ns o hyb idisa ion a e poo ly unde s ood (Whi ney
e al. 2010). The hyb id o igin o hese ou species is indi-
ca ed only by mo phological ea u es. We hope ha DNA
s udies o he Ve bascum species will help o mo e accu-
a ely iden i y hei ela ionships, o igin, and axonomical
s a us. Despi e he lack o s udy, we ne e heless decided
o include hese species in ou analysis o endemism.
The dis ibu ion o endemic ichness in di e en lo is-
ic egions is illus a ed on he A menia map (Fig. 2).
On he basis o he p emise ha allo- and au och ho-
nous ends in he e olu ion o he A menian lo a a e
app oxima ely equi alen (Fay ush 1990; Fay ush e al.
2023b), i is ecognised ha oughly hal o he species
mig a ed o A menia om o he egions, while he o he
hal o igina ed wi hin he A menian Highlands and pa ial-
ly in he Caucasus and A menia i sel .
The highes numbe o endemic ascula plan axa is
ound in he Ye e an and Da elegis lo is ic egions, wi h
ewe in he Apa an, Shi ak, and Megh i egions (Fig. 2, Sup-
pl. ma e ial 1). The majo i y o hese axa h i e in he a id
egions o he A meno-I anian p o ince o he Ancien Med-
i e anean subkingdom. Mos o hese species should be
classi ied as neoendemics, o med in a id condi ions as a
esul o anexplosi e di e si ica ion (Gi nish 2015). In con-
as , he majo i y o endemics associa ed wi h he “Cauca-
sian oo ,” which lou ish in he mo e humid condi ions o
he emaining lo is ic egions o A menia, a e likely paleo-
endemics, p ese ed om glacial and pos glacial pe iods.
Rega ding he dis ibu ion o endemic axa along
high-al i ude bel s, hey a e no ably mo e concen a ed in
he middle and uppe mon ane bel s, which bel s p o ide
he g ea es a ie y o habi a s and a e mos conduci e o
specia ion. Unlike in I an, whe e he majo i y o endemic
species a e ound in he highlands (No oozi e al. 2018,
2019, 2021), he numbe o endemics in A menia’s subal-
pine and alpine bel s is minimal, likely due o he challeng-
es o adap ing new axa o he condi ions o closed plan
communi ies in hese bel s (meadows, ca pe s, all g ass-
es), as nea ly all highland endemics a e con ined o open
habi a s wi h spa se ege a ion (ca ego y H acco ding o
he EUNIS sys em 2022).
Many au ho s ha e di ec ly co ela ed he di e si y in
a gi en e i o y o he ichness o he o e all axonomic
composi ion o plan s, pa icula ly endemics (Hobohm
2000; Guilhaumon e al. 2008; Als e be g e al. 2017; Vlad-
imi o e al. 2020; Al-sodany e al. 2024). This si ua ion
is especially ele an o na ow-local endemics, which
o en adap o only one speci ic ype o habi a (Hannus
and Nume s 2008; S en hou akis and T ian is 2009; S ein
e al. 2014). In his con ex , A menia is dis inguished by
no only he ichness o i s lo a bu also he di e si y
o i s habi a s. All he main habi a ypes o he EUNIS
sys em a e ep esen ed he e, excep o ca ego ies A and
B (due o he absence o seas; Fay ush and Aleksanyan
2016; Fay ush e al. 2023a). I should also be no ed ha
all he p ima y ege a ion ypes and ecosys ems common
h oughou he Caucasus a e p esen in A menia, wi h he
excep ion o humid sub opical ecosys ems.
The o e whelming majo i y o endemic ascula plan
axa in A menia a e con ined o he baceous habi a s (ca -
ego y E)—84 species. Signi ican ly ewe a e ound in pe -
ophy e habi a s and hose wi h no o poo ly de eloped
ege a ion (H)—29 species, in o es and sh ub habi a s
(F and G)—25 species each, and in aqua ic habi a s (C),
whe e only se en axa a e p esen . No endemic species
o subspecies ha e been eco ded in ma sh habi a s (D)
(Suppl. ma e ial 1). This dis ibu ion can be a ibu ed o
he ac ha he baceous habi a s occupy nea ly wo- hi ds
o A menia’s e i o y. Fu he mo e, mos endemics a e as-
socia ed wi h a id ecosys ems, which a e p edominan ly
ep esen ed by habi a s o his ca ego y. Ma sh habi a s
(D) a e limi ed in a ea and highly agmen ed. These hab-
i a s, along wi h aqua ic ones (C), hos a la ge numbe o
a e elic species ha ha e su i ed since he Ice Age. The
condi ions in hese habi a s a e s able and conse a i e,
which do no acili a e specia ion p ocesses. I has o be
no ed ha wo species a e dominan in na u al ecosys-
ems: Ranunculus a agazi, a dominan species o alpine
ca pe s on Moun A aga s (E4.3A29), and Smy niosis a -
mena, a dominan species o g ass- o b s eppes (E1.2E24).
Conse a ion
Endemism, de ined as he es ic ed dis ibu ion o a axon
o a speci ic geog aphical a ea, is cen al o he s udy o bio-
geog aphy. Unde s anding endemism no only add esses
a ious e olu iona y and biogeog aphical issues bu also
plays a c ucial ole in main aining biodi e si y and iden i-
ying p io i y a eas o conse a ion (Meddou e al. 2023).
Due o hei gene ic cha ac e is ics, mos neo- and
paleoendemics o A menia a e na ow- ange s eno opic
species. Consequen ly, he popula ions o many o hese
species a e h ea ened and may ace ex inc ion o a i-
ous easons, p ima ily due o an h opogenic ac o s such
as habi a des uc ion om economic ac i i ies. Global
clima e change also poses a signi ican h ea , as i may
ad e sely a ec endemics wi h na ow ecological ampli-
udes (Tamanyan e al. 2010; FNC 2014).
In 2010, he Red Book o Plan s o A menia was pub-
lished (Tamanyan e al. 2010), du ing which he s a us
o popula ions o mos endemic species was assessed
acco ding o In e na ional Union o Conse a ion o Na-
u e (IUCN) c i e ia. Eigh een species we e classi ied as
C i ically Endange ed, 26 as Endange ed, and wo species
as Vulne able. All “doub ul” species we e ca ego ised as
Da a De icien . A small numbe o species we e assessed
as Nea Th ea ened and Leas Conce n. In ecen yea s,
bo anis s ha e desc ibed se e al new axa om A me-
nia, almos all o which a e endemic o he coun y and
F on ie s o Biogeog aphy 18, 2025, e167093
Geo ge M. Fay ush e al.
8
included in ou analysis, bu hey ha e no ye been o icial-
ly assessed ega ding hei ex inc ion isk.
A menia has a well-de eloped ne wo k o specially p o-
ec ed na u al a eas, which includes h ee ese es, ou
na ional pa ks, and app oxima ely 30 wildli e sanc ua ies.
Acco ding o ou es ima es, only 37 ou o 139 endemic spe-
cies a e ound wi hin hese p o ec ed a eas. The emaining
species equi e u he esea ch; he “ques ionable” species
a e pending a decision ega ding hei axonomic s a us,
newly desc ibed species need assessmen acco ding o
IUCN c i e ia, and species classi ied as Da a De icien e-
qui e addi ional ield esea ch o de e mine hei p esence o
absence in na u e. In gene al, all endemic and na ow- anged
species necessi a e special measu es o p o ec hei habi-
a s and cla i y he possibili ies o hei ex-si u conse a ion.
Fig. 4 shows a map o A menia wi h 10 × 10 km g id cells,
whe e he numbe o endemic plan species ound he e is
shown in shades o di e en colo s. The squa e in he Sou h
Zangezu lo is ic egion s ands ou , as i hos s 16 endemic
plan species. Un o una ely, his squa e does no encom-
pass any p o ec ed a eas, highligh ing he need o expand
he nea by Zangezu S a e Na u e Sanc ua y o p ese e
he ecosys ems suppo ing hese endemic plan species.
The nex cen es ich in endemic plan species (9–12 axa)
a e loca ed in he Da elegis and Ye e an lo is ic egions,
which a e gene ally no ed o hei abundance o endemic
species. As p e iously men ioned, hese p edominan ly a id
e i o ies a e p ima ily ep esen ed by neoendemics.
A small po ion o he endemism cen es in hese a eas
alls wi hin he “Khos o Fo es ” S a e Rese e, while he
majo i y a e si ua ed ou side exis ing p o ec ed a eas, in-
dica ing he necessi y o c ea e new p o ec ed a eas and
expand hose ha cu en ly exis . Less ich cen es (6–9
axa) a e ound in he Lake Se an basin (Se an and A eguni
lo is ic egions), in he no heas (Ije an lo is ic egion),
and in he sou he n pa o he epublic (Da elegis and
Sou h Zangezu lo is ic egions), all o which equi e a -
en ion o implemen app op ia e conse a ion measu es.
Howe e , o he less ich cen es o endemic axa accumu-
la ion (1–6) should no be o e looked. I is no ewo hy ha
he mos signi ican cen es o plan endemism in A menia
coincide wi h he “biodi e si y ho spo s” iden i ied ea lie
(Fay ush e al. 2013) based on he dis ibu ion o a e plan
and animal species included in he Red Book o A menia
(Aghasyan and Kalashyan 2010; Tamanyan e al. 2010),
aking in o accoun he Impo an Plan A eas (Tamanyan
and Fay ush 2009b; Asa yan and Fay ush 2013).
Conclusion
I has been es ablished ha 139 endemic species and
subspecies o ascula plan s om 27 amilies and 69
gene a g ow wi hin he Republic o A menia. The amilies
Rosaceae, As e aceae, Poaceae, and Fabaceae a e he
iches in endemics. The la ges numbe o endemics is
eco ded in he gene a Py us and As agalus. The majo i y
o endemic species a e neoendemics, h i ing in lo is ic
egions belonging o he Ancien Medi e anean subking-
dom o lo a, pa icula ly in g assy, a id habi a s. Species
om mo e humid habi a s a e likely paleoendemics, wi h
hei dis ibu ion p ima ily limi ed o lo is ic egions be-
longing o he Bo eal subkingdom o lo a. In con as o
No he n I an, he A menian highlands a e ela i ely poo
in endemic species o med and adap ed o closed g ass
ecosys ems (meadows, ca pe s, and all g asses). In he
alpine and subalpine bel s, endemics a e p edominan ly
ound in open communi ies o ocks and sc ees.
Endemic plan species equi e u he in-dep h s udies.
Some o hese “ques ionable” species equi e addi ional
axonomic and lo is ic esea ch. In gene al, all endemic
species equi e he de elopmen o specialised measu es
o hei conse a ion in bo h na u al and ex-si u condi ions.
The iden i ied cen es o endemism— e i o ies wi h
he g ea es weal h o endemic axa—should a ac he
a en ion o en i onmen al agencies ( i s o all, he Min-
is y o En i onmen o RA, as well as Wo ld Wildli e Fund
(WWF), Founda ion o he P ese a ion o Wildli e and
Cul u al Asse s (FPWC), e c.) o o ganise p o ec ed a eas
o p ese ing he habi a s o hese endemics.
Acknowledgemen s
The esea ch was pa ly suppo ed by he Highe Educa-
ion and Science Commi ee o MESCS RA (Resea ch p oj-
ec № 21AG-1F004).
Au ho con ibu ions
All au ho s con ibu ed o he s udy’s concep ion and de-
sign. Ma e ial p epa a ion and da a collec ion we e con-
duc ed by Ka ine Janjughazyan, while analysis was pe -
o med by Geo ge Fay ush and Alla Aleksanyan. The i s
d a o he manusc ip was w i en by Geo ge Fay ush,
and all au ho s p o ided eedback on p e ious e sions.
All au ho s ead and app o ed he inal manusc ip .
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