Dry grasslands and thorn-cushion communities of Armenia: a first syntaxonomic classification

D y g asslands and ho n-cushion communi ies
o A menia: a i s syn axonomic classi ica ion
Denys Vynoku o 1,2, Alla Aleksanyan3, Thomas Becke 4, Idoia Biu un1, Da iia Bo o yk2,5,
Geo ge Fay ush3, I zia Ga cía-Mijangos1, Ma in Magnes6, Salza Palpu ina7, U e Becke 8,
Asun Be as egi9, Bea a Cykowska-Ma zencka10, Iwona Dembicz11, Die e F ank12, And eas Hilpold13,
Philipp Ki schne 14, Helmu May ho e 6, Ma ine Oganesian15, Iuliia Vasheniak2, Jü gen Dengle 16,17
1 Depa men o Plan Biology and Ecology, Uni e si y o he Basque Coun y UPV/EHU, Bilbao, Spain
2 Depa men o Geobo any and Ecology, M.G. Kholodny Ins i u e o Bo any Na ional Academy o Sciences o Uk aine, Kyi , Uk aine
3 Depa men o Geobo any and Ecophysiology, Ins i u e o Bo any a . A. Takh ajyan Na ional Academy o Sciences, Ye e an, A menia
4 Depa men o Geobo any, Facul y o Regional and En i onmen al Sciences, Uni e si y o T ie , T ie , Ge many
5 Depa men o Bo any and Zoology, Facul y o Science, Masa yk Uni e si y, B no, Czech Republic
6 Depa men o Plan Sciences, Ins i u e o Biology, Uni e si y o G az, G az, Aus ia
7 Na ional Museum o Na u al His o y, Bulga ian Academy o Sciences, So ia, Bulga ia
8 Bo anic Ga den o he Johannes Gu enbe g-Uni e si y, Mainz, Ge many
9 Biodi e si y Depa men , En i onmen al Managemen o Na a e, Pamplona-I uña, Na a oa, Spain
10 Wädenswil, Swi ze land
11 Ins i u e o En i onmen al Biology, Facul y o Biology, Uni e si y o Wa saw, Wa saw, Poland
12 En i onmen al P o ec ion Agency Saxony-Anhal (LAU), Halle, Ge many
13 Ins i u e o Alpine En i onmen , Eu ac Resea ch, Bolzano/Bozen, I aly
14 Depa men o Bo any, Uni e si y o Innsb uck, Innsb uck, Aus ia
15 Depa men o Highe Plan s Taxonomy and Geog aphy, Ins i u e o Bo any a e A. Takh ajyan o he Na ional Academy o Sciences o A menia,
Ye e an, A menia
16 Vege a ion Ecology Resea ch G oup, Ins i u e o Na u al Resou ce Sciences (IUNR), Zu ich Uni e si y o Applied Sciences (ZHAW), Wädenswil, Swi ze land
17 Plan Ecology, Bay eu h Cen e o Ecology and En i onmen al Resea ch (BayCEER), Uni e si y o Bay eu h, Bay eu h, Ge many
Co esponding au ho : Jü gen Dengle (jue gen.dengle @zhaw.ch)
Academic edi o : Vic o Chepinoga♦Linguis ic edi o : Michael Glase
Recei ed 22 Janua y 2024♦Accep ed 15 Ma ch 2024♦Published 9 May 2024
Abs ac
Aim: To p o ide he i s syn axonomic, plo -based classi ica ion o he d y g asslands and ho n-cushion communi ies
in A menia. S udy a ea: A menia. Me hods: We sampled 111 ege a ion plo s (10 m2) and eco ded en i onmen al and
s uc u al pa ame e s. We collec ed addi ional 487 ele és om su ounding coun ies o a b oad-scale compa ison.
We used modi ied TWINSPAN o de i e a syn axonomic classi ica ion sys em, whose uni s we e hen compa ed among
each o he ega ding species composi ion, s uc u e, si e condi ions and dis ibu ion. Resul s: The classi ica ion o A -
menian ege a ion plo s esul ed in a 12-clus e solu ion. Unsupe ised classi ica ion o he b oad-scale da ase yielded
i e main g oups, which we e used o he high-le el syn axonomic assignmen s o he A menian da a. We assigned
abou hal o he plo s o he A menian da ase o he Fes uco-B ome ea, while he emaining ep esen ed a po en ial
new class, p elimina ily called “Zizipho a enuio -S ipa a abica g asslands”. Mos o he syn axa below class le el a e new
o science, he e o e we p o ide o mal desc ip ions o h ee o de s (Plan agini a a ae-B omopsie alia a iega ae, On-
ob ychido anscaucasicae-S ipe alia pulche imae, Cousinio b achyp e ae-S ipe alia a abicae), ou alliances (Acan holi-
mono ca yophyllacei-S ipion holose iceae, A emision ag an is, Onob ychido michauxii-S ipion capilla ae, Onob ychido
anscaucasicae-S ipion pulche imae) and six associa ions. We ound signi ican di e ences in he opog aphic, clima ic
and soil cha ac e is ics, and s uc u al pa ame e s, species li e o ms and dis ibu ion ange ypes be ween he g assland
ypes a di e en syn axonomic le els. The mean species ichness was 47.3 ( ascula plan s: 46.8, b yophy es: 0.4, lichens:
0.1). Conclusions: We ound ema kable di e ences o he A menian d y g asslands om he p e iously known uni s
Copy igh Denys Vynoku o 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.
Vege a ion Classi ica ion and Su ey 5: 39–73
doi: 10.3897/VCS.119253
RESEARCH PAPER
In e na ional Associa ion o Vege a ion Science (IAVS)
GRASSLANDS OF ASIA
Denys Vynoku o e al.: D y g asslands o A menia40
and desc ibed mos o he highe syn axa and all he associa ions as new o science. Ou s udy p o ides a gumen s o a
po en ial new class o Zizipho a enuio -S ipa a abica g asslands sepa a e bo h om he Eu o-Sibe ian Fes uco-B ome ea
and he Ana olian As agalo-B ome ea. Finally, we ound plo scale ichness o ascula plan s clea ly abo e he Palaea c-
ic a e age o d y g asslands and ha o non- ascula plan s clea ly below, which calls o u he biodi e si y analyses.
Taxonomic e e ence: Eu o+Med (2023) o ascula plan s, Hodge s e al. (2020) o b yophy es, Nimis e al. (2018)
o lichens excep o Xan hopa melia cam schadalis (Ach.) Hale.
Abb e ia ions: EDGG = Eu asian D y G assland G oup; DCA = de ended co espondence analysis; ICPN = In e na-
ional Code o Phy osociological Nomencla u e (Theu illa e al. 2021); TWINSPAN = wo-way indica o species analysis.
Keywo ds
A menia, classi ica ion, d y g assland, Fes uco-B ome ea, I ano-Tu anian, moun ain s eppe, Onob ychido anscaucasi-
cae-S ipe alia pulche imae, Plan agini a a ae-B omopsie alia a iega ae, species ichness, syn axonomy, ho n-cushion
communi y, Cousinio b achyp e ae-S ipe alia a abicae
In oduc ion
A menia is a land-lock coun y loca ed in he sou he n
pa o he Lesse Caucasus, belonging o wo global bi-
odi e si y ho spo s: he Caucasian and he I ano-Ana o-
lian (Mi e meie e al. 2004). Wi h 3,800 ascula plan
species, including 144 coun y endemics, i has an ou -
s andingly ich ascula plan lo a o a e i o y o less
han 30,000 km2 (Biodi e si y and Landscape Conse a-
ion Union 2014). The speci ici y o he ege a ion o A -
menia is caused by he ac ha he coun y is loca ed on
he bo de be ween he Eu o-Sibe ian (o Ci cumbo eal)
and I ano-Tu anian biogeog aphical egions (Takh ajan
1986; Mana zadeh e al. 2017; Loidi e al. 2022), which a e
belonging o di e en subkingdoms: Bo eal and Ancien
Medi e anean (Takh ajan 1986).
G asslands and sh ublands in he Middle Eas and he
Caucasus a eas a e o g ea in e es as hey hos a high
biodi e si y o species and habi a s (Amba lı e al. 2020),
and ha e been shown o be he a ea o o igin o impo an
elemen s o Wes e n Asian and Eu opean d y g asslands
(e.g. Eupho bia seguie iana, F ajman e al. 2019; Eupho -
bia nicaeensis g oup, S ojilko ič e al. 2022). G asslands
a e linked o human ac i i ies, so can be hea ily impac ed
by changes in land use ha a e leading o he loss and dis-
u bance o hese habi a s, wi h he consequen loss o bio-
di e si y (Tö ök and Dengle 2018). In his a ea, un eg-
ula ed (unsus ainable) g azing, con e sion o c opland,
a o es a ion wi h non-na i e ee species, and mining and
ene gy p oduc ion p ojec s a e he majo h ea s (Amba lı
e al. 2020). The knowledge o hese habi a s in A menia
is c ucial o unde s anding hei biodi e si y and ac ual
dis ibu ion, which will make i possible o es ablish he
necessa y managemen measu es o conse a ion (Am-
ba lı e al. 2018).
Due o he abo emen ioned impo ance o he ypology
o habi a s and ege a ion, he e is a g owing in e na ional
consensus on he need o cohe en ege a ion classi ica-
ion sys ems based on he analysis o ege a ion-plo da a
(De Cáce es e al. 2015). The e a e di e en plo -based
ege a ion classi ica ion app oaches (De Cáce es e al.
2018), he mos impo an globally being he phy osocio-
logical app oach (Dengle e al. 2008) and he EcoVeg ap-
p oach (Fabe -Langendoen e al. 2014). The la e , which
is applied by he In e na ional Vege a ion Classi ica ion
(IVC, Fabe -Langendoen e al. 2020) o mally inco po-
a es wo le els abo e he class le el, which is he highes
o mal le el in he phy osociological app oach. The o -
ma ion is based on physiognomic-s uc u al ea u es o
he ege a ion and hus i is e y use ul o de ine biomes.
Recen ly Willne and Fabe -Langendoen (2021) made a
i s a emp o link he Eu opean classi ica ion sys em,
based on he phy osociological app oach (EVC, Mucina e
al. 2016), wi h he In e na ional Vege a ion Classi ica ion.
Up o da e, he e is no o malised plo -based classi ica-
ion sys em o A menia, which was a So ie Socialis Re-
public un il 1991. As he B aun-Blanque app oach o eg-
e a ion classi ica ion (B aun-Blanque 1964; Gua ino e al.
2018) de eloped in Cen al and Wes e n Eu ope, i did no
play a ole in he USSR o poli ical easons (Masing 1991).
Hie a chical classi ica ions based on syn axa we e i ual-
ly non-exis en in A menia, a leas no o g asslands, as
ege a ion scien is s ollowed he so-called dominance ap-
p oach, which ca ego ised ege a ion acco ding o domi-
nan and ecologically signi ican species (Sukache 1928).
This app oach does no usually p oduce comple e species
lis s wi h abundance da a o small sample a eas o de ined
size, i.e. i does no p oduce ele és o ege a ion plo s. As
a esul , he i s classi ica ion a emp o g assland habi a s
in A menia iden i ied h ee ege a ion ypes (G ossgeim
1928): semi-dese s, moun ain-xe ophilous ege a ion
and moun ain s eppes. A e wa ds, di e en classi ica-
ions we e de eloped o d y g asslands and s eppe ege a-
ion. Fi s , Makagian (1941) de ined ou ypes o s eppes
and s eppe-like ege a ion (s ony semi-dese s, s eppes,
meadow-s eppes and highland xe ophy es). La e , Zi oy-
an (1989) ca ego ised na u al d y g asslands in A menia
in o i e ege a ion ypes ela ed o zonal and al i udinal
dis ibu ion (dese s, semi-dese s, highland xe ophy es,
Vege a ion Classi ica ion and Su ey 41
moun ain s eppes and moun ain meadow s eppes). Final-
ly, Fay ush (1992) ecognised ou s eppe sub ypes ( ue
s eppes, ho ny-cushion s eppes, sh ubby s eppes and
meadow s eppes) wi h 12 classes o o ma ions. The lack
o a uni ied amewo k o g assland ypology impedes ac-
qui ing knowledge abou hei dis ibu ion and di e si y,
which is ha m ul o hei conse a ion.
Al hough he phy osociological app oach has no been
applied o he su ey o A menian g asslands, hose o
neighbou ing coun ies o he Sou he n Caucasus ha e
been a leas agmen a ily s udied in T anscaucasia
(Aze baijan: Pepe e al. 2010; E zold e al. 2016; Jabba o
e al. 2020; Geo gia: Pyšek and Š ů ek 1989; Nakhu s i-
sh ili e al. 2022, e c.), Eas e n Ana olia in Tu key (Çe-
ik and Ta lı 1975; Ta lı 1991; Gümüs 1992; Gümüs e al.
2003; Hamzaoğlu 2006, e c.) and No h I an (Klein 1982,
1987; Klein and Lacos e 1994; No oozi e al. 2010, 2014,
2017, e c.).
Many o he da a used o hese phy osociological su -
eys a e s o ed in ege a ion-plo da abases. In he las de-
cades small egional and/o pe sonal da abases ha e been
compiled in la ge ege a ion plo s da abases. The Eu ope-
an Vege a ion A chi e (EVA, Chy ý e al. 2016) was he
pionee , ollowed by he global ege a ion da abase (sPlo ,
B uelheide e al. 2019) and by G assPlo (Dengle e al.
2018). These da abases a e he basis o la ge-scale ege-
a ion classi ica ion s udies (No ák e al. 2023b; Pe e ka e
al. 2023), bu hey also pe mi mac oecological s udies o
g ea scales leading o a b oad unde s anding o he dis-
ibu ion and di e si y pa e ns (G aco-Roza e al. 2022;
Saba ini e al. 2022; Večeřa e al. 2023). Especially o
g asslands, he G assPlo da abase hos s high-quali y da a
om he Palaea c ic ealm, sampled on p ecisely delim-
i ed plo s, including ascula plan s and c yp ogam da a
(Dengle e al. 2018; Biu un e al. 2019). A he egional
scale, he T anscaucasian Vege a ion Da abase, a phy oso-
ciological da abase o he Sou he n Caucasus, was ecen -
ly es ablished (No ák e al. 2023a). Howe e , i cu en ly
lacks ege a ion plo s o d y g assland and ho n-cushion
communi ies in A menia.
I is he e o e bo h an oppo uni y and a challenge o
eco d ege a ion plo s and apply he B aun-Blanque
classi ica ion app oach o A menia – as i s ege a ion, o
ou knowledge, has ne e been s udied acco ding o his
app oach. The e o e, he Eu asian D y G assland G oup
(EDGG; www.edgg.o g) conduc ed an in e na ional e-
sea ch expedi ion (called “Field Wo kshop”) in A menia
o collec s anda dised, high-quali y ege a ion-plo da a
om d y g asslands and ho n-cushion communi ies
h oughou he coun y. In p inciple he EDGG Field
Wo kshops aim a collec ing such da a o egional s udies
on biodi e si y pa e ns (Kuzemko e al. 2016; Dembicz
e al. 2021b; Be gaue e al. 2022) and phy osociology
(Ga cía-Mijangos e al. 2021; Magnes e al. 2021). Mo e-
o e , hese da a a e p o ided o he G assPlo da abase
(Dengle e al. 2018) o biodi e si y s udies ac oss all Pa-
laea c ic g asslands and o he open habi a s (Biu un e
al. 2021; Dembicz e al. 2021a; Zhang e al. 2021) as well
as o he Eu opean Vege a ion A chi e (EVA; Chy ý e al.
2016) and he global ege a ion-plo da abase sPlo (B u-
elheide e al. 2019) o ill impo an da a gaps in con inen-
al and global s udies o biodi e si y, global change and
syn axonomy.
In his pape , we used he plo da a sampled du ing he
Field Wo kshop o p o ide he i s syn axonomic classi-
ica ion scheme o he d y g asslands and ho n-cushion
communi ies o he coun y, using nume ical me hods o
unsupe ised classi ica ion and de e mina ion o diagnos-
ic species. Speci ically, we asked:
(1) Which associa ion- ank communi ies can be dis in-
guished, and o which highe - ank syn axa do hey
belong?
(2) How do he syn axa in A menia compa e o hose in
he neighbou ing coun ies o Wes e n Asia?
(3) How a e he syn axa in A menia di e en ia ed om
each o he in e ms o species composi ion, species
ichness, s uc u e, si e condi ions and dis ibu ion?
S udy a ea
Physiogeog aphy, clima e, soils, geology
A menia is a Sou h Caucasian epublic, bo de ing Geo -
gia, Aze baijan, Tu key, and I an. I is a landlocked coun-
y wi h a o al a ea o 29,740 km2, a abou 145 km om
he Black Sea and 175 km om he Caspian Sea. I lies
be ween 38°50' and 41°18' no he n la i ude and be ween
43°27' and 46°37' eas e n longi ude, and measu es 400 km
along i s main axis (no h-wes o sou h-eas ). A menia is
gene ally a moun ainous coun y, ha ing i s lowes poin
a 375 m a.s.l. and culmina ing a 4,095 m a.s.l. in he A -
aga s, wi h an a e age ele a ion o 1,850 m a.s.l.
The loca ion o A menia a he in e sec ion o wo phy-
ogeog aphical subkingdoms (Bo eal and Ancien Medi-
e anean), oge he wi h he di e si y in clima ic condi-
ions and he ac i e geological p ocesses, ha e esul ed in
he o ma ion o di e se ecosys ems and high biodi e si y
wi h a high le el o endemism (Fay ush and Aleksanyan
2016). On he small e i o y o he coun y, he e a e
abou 3,800 species o ascula plan s, 497 species o soil
and wa e algae, 433 species o b yophy es, 4,577 species
o non-lichenized ungi, 619 species o lichens, 567 spe-
cies o e eb a es and abou 17,000 species o in e e-
b a es (Fay ush 2023).
A wide ange o clima ic zones a e dis inguished wi hin
A menia, which expe iences la ge clima ic con as s be-
cause o i s in ica e e ain and he big clima ic g adien s
(Minis y o Na u e P o ec ion o he Republic o A menia
2015). The basic clima e ypes mainly ollow he ele a ion-
al g adien , om d y sub opical up o se e e alpine. The
a e age annual empe a u e anges om -8°C in high-al-
i ude moun ainous egions (2,500 m a.s.l. and highe ) o
12–14°C in low- aced alleys. The o e all clima e is bes
cha ac e ised as d y con inen al, in some a eas wi h an
Denys Vynoku o e al.: D y g asslands o A menia42
annual hy hm like he Medi e anean clima e egime.
The a e age annual p ecipi a ion in A menia is 592 mm.
The mos a id egions a e he A a a alley and he egion
o Megh i wi h annual p ecipi a ion o 200–250mm. The
highes annual p ecipi a ion o 800–1000 mm is obse ed
in high-al i ude moun ain egions.
F om he o og aphical and physico-geog aphical
poin s o iew, A menia o ms he no he n edge o he
sys em o olded-block moun ains o he A menian High-
land. Unlike he G ea e Caucasus, A menia and he Less-
e Caucasus a e no a single, dis inc wa e shed idge. I
is a sys em o coulisse-spaced idges ha me ge wi h he
moun ain o ma ions o he inne pa s o he A menian
Highland and adjacen high a eas (Aslanyan 1958, 1985).
Fou main geomo phological egions can be ecognized
wi hin A menia. (1) Moun ain idges and alleys in he
no h-eas o he coun y which bea wi ness o ex en-
si e e osion. (2) A eas co e ed by la a o ela i ely ecen
(uppe Pliocene) o igin wi hin Asia Mino a e cha ac e -
ised by gen le slopes wi h li le e idence o e osion bu , in
which la ge i e s ha e ca ed ou deep go ges and can-
yons. (3) A se ies o idged moun ains in he sou h o A -
menia, which cons i u e he Lesse Caucasus sys em and
show in ense e osion. (4) The A a a Valley ep esen s he
lowes pa o he A a a dep ession co e ed wi h allu ial
and p olu ial sedimen s (Aslanyan 1958; Gab ielyan 1962;
Dumi ashko 1979).
In ou s udy we ied o co e as much o he coun y’s
d y g assland di e si y as possible wi hin 11 days, wi h a
ocus on he no hwes e n and cen al pa s (Figu e 1). In
o al, we sampled in i e o he 11 adminis a i e p o inces
o he coun y (A aga so n, A a a , Gegha kunik, Shi ak,
Vayo s Dzo ) and se en o he 12 lo is ic egions. We co -
e ed an ele a ional g adien om 1,338 o 2,350 m a.s.l.
Na ional ypologies o A menian d y g asslands
So a , he syn axonomy o g assland o ho n-cushion
ege a ion o A menia hasn’ been de eloped ye . The only
exis ing ege a ion ypologies a e based on he dominance
app oach. The i s o e iew o he A menian ege a ion
ypes was pe o med by G ossgeim (1928). He dis in-
guished eigh main ypes o ege a ion: (1) aqua ic and
bog ege a ion; (2) o es ege a ion; (3) semi-dese s; (4)
moun ain-xe ophilous ege a ion (moun ain semi-dese );
(5) moun ain s eppes; (6) meadow-like ege a ion; (7) so-
lone zs and solonchaks; (8) weeds. In he semi-dese ype,
he selec ed h ee sub ypes: (a) alkaline-loamy semi-dese
on he allu ium o he A as Ri e ; (b) wo mwood semi-de-
se on e up i e ocks; (c) sandy semi-dese .
Takh ajan (1941) explo ed he phy ogeog aphic pa -
e ns o A menia, including he di ision o he egion in o
phy ogeog aphic p o inces and dis ic s, he his o ical
Figu e 1. Map o A menia indica ing zonal ege a ion ypes based on G ossgeim (1928). Black do s indica e he sam-
pling locali ies. The ligh blue a ea e e s o he Lake Se an.
Vege a ion Classi ica ion and Su ey 43
de elopmen o i s ege a ion, and he classi ica ion o
ege a ion in o b oad ypes. Among hese ypes, subalpine
ege a ion, moun ain-s eppes, xe ophilous ege a ion o
skele on moun ains, and wo mwood semi-dese s could
be conside ed as he scope o ou s udy.
Makagian (1941) de eloped he ege a ion ypology o
A menia in mo e de ail. The s eppe and s eppe-like ege-
a ion included in his scheme was classi ied as:
• S ony semi-dese s (wo mwood semi-dese ,
wo mwood-epheme al semi-dese , wo mwood
semi-dese wi h pe ennial g asses, e c.)
• S eppes (g ass- o b and d y o b-g ass s eppe, ea h-
e -g ass s eppe, bea dg ass s eppe, escue and es-
cue-juneg ass s eppe, mixed-g ass s eppe, o b and
legume s eppe)
• Meadow-s eppes (g ass meadow-s eppe, o b and
o b-g ass meadow-s eppe, legume meadow-s eppe,
dwa -sedge meadow-s eppe)
• Highland xe ophy ic ege a ion (Mino -Asian
ho n-cushion sh ubs o As acan ha, Acan holimon
e c., xe omo phic ege a ion o sc ees and ocks)
A e wa ds, he classi ica ion o na u al g asslands o
A menia was done by Zi oyan (1989) on he p inciples
o he dominan app oach. The au ho dis inguished i e
ege a ion ypes o dese and s eppe ege a ion which
ha e s ong zonal cha ac e and mainly a e cha ac e is-
ic o a pa icula al i udinal bel : dese s ( wo classes o
o ma ions, 16 o ma ions) in he lowes ele a ions up o
1,000 m a.s.l.; semi-dese s ( wo classes o o ma ions, 12
o ma ions) a he ele a ions 1,000–1,300 m a.s.l.; high-
land xe ophy ic ege a ion (one class o o ma ions, 12
o ma ions), 1,200–1,600 m a.s.l.; moun ain s eppes ( wo
classes o o ma ions, 12 o ma ions), 1,600–2,000 m a.s.l.;
moun ain meadow s eppes ( wo classes o o ma ions, 12
o ma ions), 2,000–2,400 (occasionally up o 2,800) m a.s.l.
Fay ush (1992) p esen s a de ailed classi ica ion o he
ypes o moun ain s eppes o A menia based on he dom-
inance app oach. The au ho dis inguishes ou sub ypes
( ue s eppe, ho ny-cushion s eppe, sh ubby s eppe and
meadow s eppe) and 12 classes o o ma ions wi hin i .
O e iew on he desc ibed syn axa om he
su ounding coun ies
In he T anscaucasus egion, he i s wo k wi h a de-
sc ip ion o syn axa ollowing he B aun-Blanque ap-
p oach was done by M. Guinoche in Aze baijan and
Geo gia. Guinoche (1984) desc ibed wo associa ions
o s eppe-like communi ies wi hou an assignmen o
highe syn axonomic uni s. The associa ion Zizipho o
se pyllaceae-Scu ella ie um o ien alis Guinoche 1984 is
cha ac e ized by he p esence o many he ophy es and is
desc ibed om he lowe ele a ions in Aze baijan. Guin-
oche concludes ha his ege a ion ype could be like he
class The o-B achypodie ea and emphasizes ha a unique
class and o de should be desc ibed o include his asso-
cia ion. The o he associa ion is desc ibed om highe el-
e a ions in he Pi qulu S a e Rese e (Aze baijan), om
he subalpine bel : Onob ychie o cy i-Fes uce um sulca ae
Guinoche 1984 nom. in al. (A icle 2b ICPN, Theu illa
e al. 2021). This associa ion comp ises moun ain s eppes
and is simila , acco ding o M. Guinoche , o he concep
o he subalpine s eppe o Gadzhie (1962) om he na-
ional ypology o Aze baijan. Addi ionally, Guinoche
(1984) desc ibed in Geo gia ano he associa ion om he
alpine bel no belonging o he s eppic ege a ion and
assigned o he class Ca ici upes is-Kob esie ea bella dii:
Alchemillo caucasicae-Fes uce um supinae, oge he wi h
he new o de Campanulo- iden a ae-Ca ice alia is-
is and he new alliance Alchemillo-Fes ucion supinae, all
h ee being in alidly published due o insu icien o iginal
diagnosis (A icle 2b ICPN, Theu illa e al. 2021). La e-
ly, he e was an a emp o classi y he ph yganoid ege-
a ion o he Nakhchi an egion o Aze baijan (Jabba o
e al. 2020). The au ho s ou lined se e al associa ion-le el
uni s wi hou a o mal desc ip ion (“Thyme o-Acan holi-
mone um b ac ea ae”, “Thyme o-Onob yche um co nu-
ae”), and assigned hem o he o de As agalo-B ome a-
lia Quézel 1973 and he class As agalo-B ome ea Quézel
1973. Recen ly, a new s udy on he syn axonomy o alpine
and subalpine g asslands has been conduc ed in Geo gia
(Nakhu s ish ili e al. 2022). The au ho s p oposed a new
class B omopsio a iega ae-Fes uce ea o inae o uni e sub-
alpine meso-xe ic and mesic g asslands, including one
new o de , h ee alliances and se en associa ions. None o
he sugges ed uni s we e published alidly due o insu i-
cien o iginal diagnoses.
Fo he No he n Caucasus, Tsepko a (1987) p oposed
a new class o high-moun ain a id g asslands wi h he
p o isional name Bo h yochloo-Sal ie ea, which acco d-
ing o Vynoku o e al. (2021) is a syn axonomic synonym
o he Fes uco-B ome ea. O he s eppic g asslands o he
No he n Caucasus ha e been adi ionally assigned o
he class Fes uco-B ome ea (Tsepko a 2005; Demina e al.
2017; Vynoku o e al. 2021).
In Tu key, se e al high-le el syn axonomic uni s ha e
been es ablished o d y g asslands and ho n-cushion
ege a ion. Zoha y (1973) uni ed semi-dese and wo m-
wood s eppe g asslands in o he class “A emisie ea a-
g an is ana olica” Zoha y 1973 nom. in . (A . 2b ICPN),
and p oposed he class “As agale ea a meno- u cica” Zo-
ha y 1973 nom. in . (A . 2b ICPN) o subalpine aga-
can hic ege a ion in his egion. Simul aneously, Quézel
(1973) desc ibed ano he uni o hedgehog plan com-
muni ies in he subalpine zone o he Tau us Moun ains in
Tu key, occu ing a ele a ions 2000–2500 m a.s.l., beyond
he ee line – As agalo-B ome ea Quézel 1973. In he
same publica ion, Quézel (1973) also published ano he
class o xe o-mesophy ic and mesophy ic subalpine g ass-
lands, occupying highe al i udes han agacan h ege a-
ion communi ies: T i olio ana olici-Polygone ea a enas i
Quézel 1973. In he eas e n pa o Tu key, in Eas e n
Ana olia, he e we e some syn axonomic in es iga ions o

Denys Vynoku o e al.: D y g asslands o A menia44
s eppe ege a ion (Çe ik and Ta lı 1975; Ta lı 1991; Oca-
k e di 1992; Hamzaoğlu 2006; Öz ü k e al. 2015). Oca-
k e di (1992) su eyed he ege a ion (including s eppe
ege a ion) in he egion o Tu key bo de ing wi h A me-
nia. He dis inguished wo physiognomic ypes o s eppe
ege a ion (g ass s eppe and agacan h s eppe) and h ee
al i udinal a ian s: “plain” s eppe (1,675–1,725 m a.s.l.,
mainly g ass s eppe ege a ion), lowe moun ain s eppe
(1,750–1,850 m a.s.l., bo h g ass s eppe and agacan h
s eppe) and high-moun ain s eppe (1,950–2,696 m a.s.l.,
g ass s eppe). He p oposed ou associa ions, none o
hem published alidly (A . 1 ICPN). La e , Ocak e di e
al. (2009) desc ibed 10 new associa ions om he same e-
gion (Kısı Moun ain). Hamzaoğlu (2006) s udied s eppe
communi ies o Eas Ana olia. He uni ed he s udied eg-
e a ion o a new o de Fes uco o eophilae-Ve onice alia
o ien alis Hamzaoğlu 2006, subo dina ed o he class As-
agalo-B ome ea Quézel 1973.
In ano he bo de ing egion, I an, xe ophilous g ass-
land and sc ub communi ies we e i s delinea ed by
Zoha y (1973), who p oposed se e al ege a ion class-
es: “A emisie ea ag an is ana olica” Zoha y 1973 nom.
in . (A . 2b ICPN) o wo mwood s eppe g asslands in
No hwes e n I an and Ana olia, “A emisie ea he bae-al-
bae i anica” Zoha y 1973 nom. in . (A . 2b ICPN) g oup-
ing wo mwood semidese s in he Cen al Pla eu o I an,
and “As agale ea i anica” Zoha y 1973 nom. in . (A . 2b
ICPN) o agacan h communi ies in I an and I aq. La e ,
he ege a ion o he subalpine and alpine bel s o he Al-
bo z Moun ains ha e been s udied by Klein (1982, 1987).
He desc ibed se e al new classes: Onob ychide ea co nu-
ae Klein 1987 nom. in al. (A . 2b ICPN) om he lowe
alpine bel o Albo z (3,200–3,500 m a.s.l.) and P ange ea
ulop e ae Klein 1987 nom. in al. (A . 2b ICPN) om he
subalpine bel o Albo z (2,500–3,200 m a.s.l.), aimed o
uni e high-moun ain hedgehog communi ies and xe ic
all-he b ege a ion espec i ely. Also, om he subalpine
bel o he no he n mac oslope o Albo z Moun ains,
Klein and Lacos e (1994) desc ibed one associa ion sub-
o dina ed o he Fes uco-B ome ea, Alchemille um plica is-
simae Klein e Lacos e 1994, bu did no assign i o an
alliance o o de .
In gene al, he class As agalo-B ome ea is he mos
widely used name o uni e he d y g asslands and
ho n-cushion ege a ion in he wes e n pa o he I a-
no-Tu anian egion. Howe e , i s concep ual bounda -
ies ha e unde gone signi ican changes o e ime, bo h
geog aphically and physiognomically. Many esea che s
now ex end i s scope o include agacan h ege a ion no
only om he subalpine bel bu also om he lowe el-
e a ions, as well as chamaephy e-domina ed ph yganoid
ege a ion, non- agacan h d y g asslands a lowe ele a-
ions, saline s eppes, and gypsophilous ocky g asslands
(Ke enoğlu e al. 1983; Akman e al. 1984; Aydoğdu e al.
1994; Aydoğdu e a. 2004; Hamzaoğlu e al. 2004; Kaya
2011). Addi ionally, xe o-mesophy ic and mesophy ic
g asslands a highe ele a ions, which we e classi ied by
Quézel (1973) as a sepa a e class T i olio-Polygone ea, a e
some imes included wi hin he As agalo-B ome ea (E en
e al. 2004; Pa olly 2004). Some au ho s ha e p oposed
ex ending he concep o he class also o he no he n
pa o I an, synonymizing he Onob ychide ea co nu ae
and P ange ea ulop e ae desc ibed he e (Pa olly 2004).
Recen ly, he d y ea he -g ass s eppes o Tajikis an also
we e p o isionally included in o he As agalo-B ome ea
(Nowak e al. 2016, 2018).
Me hods
Field sampling in A menia
We sampled 111 plo s o 10 m2- size (Suppl. ma e ials 1, 2)
du ing he 13 h Field Wo kshop o he Eu asian D y G ass-
land G oup (EDGG) in A menia, om 26 June o 6 July
2019 (Aleksanyan e al. 2020; o dis ibu ion o si es, see
Figu e 1). Wi hin each plo , we eco ded ascula plan s
and e icolous b yophy es and lichens wi h he shoo
p esence me hod (Dengle 2008). Besides, we es ima ed
hei pe cen age co e on a con inuous scale ( o discus-
sion o ad an ages o his me hod compa ed o o dinal
scales, see Dengle and Dembicz 2023). Specimens ha
could no be de e mined in he ield we e d ied and aken
o he lab o u he de e mina ion.
O he en i onmen al and s uc u al pa ame e s ha
we e eco ded in si u ollowing he EDGG sampling me h-
odology (Dengle e al. 2016), included: geog aphical po-
si ion (la i ude, longi ude), ele a ion (m), slope aspec
(°), slope inclina ion (°), maximum mic o elie (cm), soil
dep h (cm, mean and SD o i e measu emen s pe plo ),
ege a ion co e s (%; o al ege a ion, sh ub laye , he b
laye , c yp ogam laye ), co e o li e (%), co e s o s ones
and ocks, g a el, and ine soil (all h ee ac ions summing
up o 100%), maximum heigh o sh ubs (m), maximum
heigh o he bs (cm), heigh o he b laye (cm, mean and
SD o i e measu emen s wi h a alling disc pe plo ), and
land use de ails (g azing, mowing, bu ning, abandonmen ).
Soil was collec ed as mixed samples om he uppe -
mos 15 cm o he soil in i e andom poin s inside each
plo . A e ai d ying and sie ing o he ine-soil ac ion,
he ollowing pa ame e s we e measu ed in he lab: pH (in
H2O), elec ical conduc i i y (μS cm-1), o ganic C con en
(%), humus con en (%), N con en (%), and C/N a io.
Sou hing was calcula ed om aspec as -cos (aspec ).
The nomencla u e o ascula plan s was s anda dised
o Eu o+Med (2023) o ascula plan s, Hodge s e al.
(2020) o b yophy es, Nimis e al. (2018) o lichens.
Fo some g oups o closely ela ed species ha could no
always be dis inguished, we de ined addi ional species
agg ega es (“agg .”; see Suppl. ma e ial 3). The alue dis-
ibu ion o all eco ded and analysed nume ical en i-
onmen al, s uc u al and biodi e si y a iables is gi en
in Suppl. ma e ial 4. The comple e da a a e s o ed in and
a ailable om he G assPlo da abase (Dengle e al. 2018;
Biu un e al. 2019; h ps://edgg.o g/da abases/G assPlo )
as da ase “AM_B”.
Vege a ion Classi ica ion and Su ey 45
Da a om ex e nal sou ces
Plo da a om he su ounding coun ies
To be able o iden i y he high-le el syn axonomic uni s,
we digi ised om li e a u e and used o compa ison
ele és om he bo de ing egions, ocusing on he o ig-
inal diagnosis o he high-le el uni s o simila ege a ion
ypes: 230 plo s om Ana olia (Tu key) and 51 plo s om
No he n I an. Among hem, he o iginal diagnosis o he
class As agalo-B ome ea Quézel 1973 wi h he ype o de
As agalo-B ome alia Quézel 1973 and he o he o de s
D abo-And osace alia Quézel 1973, Hype ico lina ioid-
is-Thyme alia sco pilii Akman e al. 1987, Onob ychido a -
menae-Thyme alia leucos omi Akman e al. 1985, Fes uco
o eophilae-Ve onice alia o ien alis Hamzaoğlu 2006, and
o he lowe ank uni s (Quézel 1973; Akman e al. 1984,
1987; Gümüs 1992; Klein and Lacos e 1994; Hamzaoğlu
2006; Ocak e di e al. 2009). Also, 206 plo s om No h-
e n Caucasus (Russia) we e aken om he Eas e n Eu o-
pean S eppe Da abase (Vynoku o e al. 2020). Combined
wi h ou own plo s om A menia, a da ase o 598 ele és
esul ed (Suppl. ma e ial 5). A e he uni ica ion o he
species axonomy, emo ing he axa de e mined o he
genus le el, and combining he agg ega es, he inal da a-
se con ained 1,556 axa.
En i onmen al da a om ex e nal sou ces
Clima ic da a we e ex ac ed om he CHELSA Clima e
da abase (Ka ge e al. 2017, 2018). We used he ollowing
a iables: BIO01 – mean annual ai empe a u e, BIO07
– annual ange o ai empe a u e, BIO12 – annual p e-
cipi a ion amoun , BIO17 – mean mon hly p ecipi a ion
amoun o he d ies qua e .
Maps we e c ea ed using QGIS so wa e (QGIS De el-
opmen Team 2009). As basemaps, we used he map o
main ege a ion ypes o A menia by G ossgeim (1928),
and he SRTM (Shu le Rada Topog aphy Mission) ele a-
ion model o A menia (EROS Cen e 2017).
A ibu es o he A menian species
In o de o assess he dis ibu ion anges o species, we
analysed hei dis ibu ion maps (acco ding o GBIF 2023
and POWO 2023) and classi ied hei dis ibu ion anges
in o he ollowing b oad ca ego ies: Eu opean (occu ing
mainly in he empe a e egions o Eu ope and Wes e n
Sibe ia), Medi e anean (co e ing he sou he n pa o
Eu ope in a eas wi h Medi e anean clima e), I ano-Tu a-
nian (occu ing in Wes e n and Cen al Asia), T anscau-
casian (na ow endemics o A menia and su ounding
egions o T anscaucasia), Caucasian (b oade endemics
o he whole Caucasus Range, including he No he n
Caucasus and adjacen pa s o Eas e n Ana olia in Tu -
key and No h-Wes e n I an). I he species occu ed p e-
dominan ly in one o he men ioned egions, we assigned
a alue o one (1) o he co esponding ca ego y. In case
ha he species occu ed in wo egions simul aneously
(Eu opean and I ano-Tu anian, Eu opean and Medi e a-
nean, o Medi e anean and I ano-Tu anian), we assigned
a alue o 0.5 o he co esponding ca ego ies. I he spe-
cies dis ibu ion co e ed h ee egions (Eu opean, Medi-
e anean and I ano-Tu anian), we assigned a alue ‘0.3’
o each o hese ca ego ies. In all o he cases, when he
species dis ibu ion a ea was g ea e han he men ioned
ca ego ies (e.g., Palaea c ic, Hola c ic, e c.), we classi ied
hese species in o he ca ego y ‘O he ’. The assigned dis i-
bu ion anges o species a e a ailable in Suppl. ma e ial 6.
In addi ion, we classi ied all he species in o one o he
Raunkiæ plan li e o ms: he ophy es, geophy es, hemic-
yp ophy es, chamaephy es, and phane ophy es. The da a
a e also a ailable in he Suppl. ma e ial 6.
S a is ical analysis
Unsupe ised classi ica ion
Unsupe ised classi ica ion o bo h he Wes Asian and
A menian da ase was done in JUICE 7.0 (Tichý 2002) us-
ing he modi ied TWINSPAN me hod (Roleček e al. 2009)
wi h h ee pseudospecies cu le els (0, 5, and 15), and
Whi ake ’s be a-di e si y index as a measu e o in e nal
clus e he e ogenei y. Diagnos ic species we e de e mined
based on phi alues (Chy ý e al. 2002), s anda dised o
equal plo numbe s a associa ion le el (Tichý and Chy ý
2006). This was done hie a chically a he ou syn axo-
nomic le els om associa ion o class (Ga cía-Mijangos e
al. 2021). Since his app oach is no implemen ed in JUICE
(Tichý 2002) ye , we had o do i in Mic oso Excel, which
p ecluded he use o Fishe ’s exac es o signi icance. To
a oid selec ing non-signi ican diagnos ic species, we pu
he h esholds o phi alues a he high. Fo associa ions
and alliances, we used > 0.4 o diagnos ic species and > 0.6
o highly diagnos ic species, while o o de s and classes
he h esholds we e > 0.3 and > 0.5, espec i ely. Mo eo e ,
we also ensu ed ha he phi alues we e a leas 0.2 highe
in he a ge syn axon han he syn axon o he same ank
wi h he nex -lowe phi alue (see Ga cía-Mijangos e al.
2021). I a species ul illed he c i e ia o be diagnos ic a
se e al hie a chical le els, i was assigned o he le el wi h
he highes phi alue. In case o mono ypic syn axa, diag-
nos ic species we e only assessed a he highe le el.
O dina ion
DCA-O dina ion was pe o med wi h Canoco 5 ( e B aak
and Šmilaue 2012) wi h log- ans o med pe cen co e al-
ues o species and downweigh ing a e species and pos hoc
i ed a iables (en i onmen al, calcula ed o measu ed).
Compa ison o syn axon cha ac e is ics
Di e ences in a iables be ween syn axa we e es ed by
uni a ia e ANOVA using SPSS 22 (IBM, A monk, NY,
US). We es ed whe he he assump ions o ANOVA
(no mal dis ibu ion, equal a iance) we e su icien ly
me by isually inspec ing he equency dis ibu ion o
he esiduals and by es ing o homogenei y o a iance
acco ding o Le ene (Quinn and Keough 2002). Whe e
ANOVA e ealed a signi ican pa e n, Tukey’s pos -hoc
Denys Vynoku o e al.: D y g asslands o A menia46
es a p < 0.05 was used o iden i y homogeneous g oups
o syn axa. Resul s we e p esen ed as box-whiske plo s
wi h median and mean, and 25 h/75 h (boxes) and 10 h/90 h
(whiske s) pe cen iles as well as ou lie s.
Syn axonomic assignmen
We selec ed a TWINSPAN esolu ion whe e he e minal
clus e s we e lo is ically s ill well-cha ac e ised and no
oo small. These clus e s we e hen assigned o he ank o
associa ion. Alliance, o de and class le els we e assigned
o highe cu le els o he dend og am, wi h he double
aim o ha e lo is ically well di e en ia ed and ecological-
ly and cho ologically in e p e able uni s on a compa able
le el as hese hie a chies ha e in Mucina e al. (2016).
A e de ining he hie a chical uni s, we ca e ully
checked he syn axonomic li e a u e o he neighbou ing
coun ies o de e mine whe he syn axa wi h his con en
al eady exis ed. I his was he case, we ook he es ab-
lished name. I no , we o mally desc ibed new syn axa
acco ding o he ICPN (Theu illa e al. 2021). Following
Recommenda ion 7A o he ICPN, we e ained om es-
ablishing new associa ions when we had ewe han 10
ele és and ins ead ea ed he espec i e clus e as an
in o mal communi y a associa ion ank. Likewise, we
e ained om a o mal desc ip ion o he new class ha
was suppo ed by ou analyses, sugges ing ha his should
i s be “ alida ed” in a b oade -scale analysis in ol ing
he neighbou ing coun ies o A menia.
Resul s
Unsupe ised classi ica ion and o dina ion o
he Wes Asian and Caucasian da ase
Modi ied TWINSPAN esul ed in o 16 clus e s wi h i e
main g oups o clus e s: A (clus e s 1–3), B (4–6), C (7), D
(8–10) and E (11–16) (Figu e 2). The synop ic able buil
wi h hese i e main g oups is shown in Suppl. ma e ial 7.
The g oup A (clus e s 1–3) comple ely consis ed
o he ele és o iginally assigned o he class As aga-
lo-B ome ea, including he ype o de As agalo-B ome-
alia Quézel 1973 (in he clus e 2), desc ibed om he
Tau us Moun ains in Sou h-Wes e n Tu key (Quézel
1973). The plo s assigned o he o he o de s o his
class wi h hei co esponding ypes we e also included
in his g oup o clus e s: D abo-And osace alia Qué-
zel 1973 (clus e 2) om he same Tau us Moun ain
ange; Hype ico lina ioidis-Thyme alia sco pilii Ak-
man e al. 1987 (clus e 1) om he Ilgaz Moun ains
in No h-Wes e n Tu key; Onob ychido a menae-Thy-
me alia leucos omi Akman e al. 1985 (clus e 3) de-
sc ibed om Cen al Ana olia. Plo s om A menia did
no i in o his g oup.
The second g oup B (clus e s 4–6) comp ised plo s
om he high-moun ain s eppe ege a ion. Clus e 4
consis ed mainly o ele és om he no he n slope o
he Albo z Moun ains in I an, assigned o he associa-
ion Alchemille um plica issimae Klein e Lacos e 1994.
Plo s om he No he n Caucasus egion assigned o he
alliance A emisio chamaemeli oliae-B omopsion a iega-
ae Vynoku o in Vynoku o e al. 2021 we e classi ied
in o clus e 5, oge he wi h some plo s om A menia.
Clus e 6 comp ised plo s sampled nea he Kısı Moun-
ain in Tu key, in Eas e n Ana olia. They we e o iginal-
ly assigned o se e al associa ions o he class As aga-
lo-B ome ea bu wi hou placemen in alliances and
o de s (Ocak e di e al. 2009).
Tho n-cushion communi ies om Eas e n Ana o-
lia a e combined in he g oup C (clus e 7). They we e
o iginally assigned o he o de Fes uco o eophilae-Ve-
onice alia o ien alis Hamzaoğlu 2006 o he class As-
agalo-B ome ea, oge he wi h i s ype alliance Fes uco
o eophilae-Ve onicion o ien alis Hamzaoğlu 2006 and he
espec i e associa ion As agalo-Onob ychide um co nu-
ae Gümüs 1992. E en hough his clus e was no placed
in he co esponding g oup A in he dend og am (Figu e
2), hey seem closely ela ed acco ding o he o dina ion
(Figu e 3).
Clus e s 8–10 o med g oup D. I consis ed exclusi ely
o plo s om A menia. Among hem, clus e 8 con ained
he mos xe ic communi ies sampled in he d ies pa s o
A menia, ollowed by clus e 10. Clus e 9 was ansi ional
be ween he g oups D and E.
The g oup E (clus e s 11–16) was o med by plo s con-
aining he mo e ‘ ypical’ Fes uco-B ome ea species, mos -
ly om he egion o he No he n Caucasus. Clus e 11
was comp ised mainly o ocky g asslands belonging o
he o de Asphodelino au icae-Eupho bie alia pe ophi-
lae Vynoku o in Vynoku o e al. 2021; clus e 12 con-
ained g ass s eppes mos ly o he Fes uce alia alesiacae.
Clus e 13 combined moun ain s eppes exclusi ely om
A menia. Clus e s 14–16 comp ised mos ly meso-xe ic
communi ies o he o de B achypodie alia pinna i om
he No he n Caucasus. A synop ic able wi h he i e dis-
inguished g oups o clus e s (A–E) is p o ided in Suppl.
ma e ial 7.
Figu e 2. Resul s o he Modi ied TWINSPAN classi i-
ca ion o he b oad-scale compa ison in ol ing plo s
om he bo de ing coun ies (n = 598). The wid h o he
ba s is p opo ional o he numbe o included plo s. The
main g oups (le e s) and e minal clus e s (numbe s)
a e desc ibed in he ex . Blue colou indica es clus e s
ha p edominan ly o comple ely consis ed o A meni-
an plo s.
Vege a ion Classi ica ion and Su ey 47
Gene al o e iew o he A menian plo s
In ou 111 plo s o 10 m2, we eco ded a o al o 739 as-
cula plan , 40 b yophy e and 13 lichen axa (subspecies,
species, agg ega es and sec ions, u he as ‘species’). The
species ichness pe plo anged om 21 o 85, wi h a
mean o 47.3. On a e age he e we e 46.8 ascula plan ,
0.4 b yophy e and 0.1 lichen species pe plo . The mos
equen ascula plan was Galium e um (in 72% o
all plo s), ollowed by Thymus ko schyanus (59%), Teu-
c ium capi a um (58%), Poa bulbosa (55%), Dac ylis
glome a a (54%), Scu ella ia o ien alis agg . (53%), Koe-
le ia mac an ha (51%), S achys ec a (50%) and Po en il-
la ec a agg . (50%). The mos equen b yophy es we e
Syn ichia u alis (27%), P ychos omum imb ica ulum
(19%) and Syn ichia mon ana (14%). Lichens we e absen
in mos plo s, wi h he mos equen one (Cladonia olia-
cea) eaching jus 4%.
Classi ica ion o he A menian da ase
The mos meaning ul modi ied TWINSPAN classi ica-
ion o he plo s om A menia esul ed in he 12-clus-
e solu ion (Figu e 4). The i s clus e (X) had only a
single ele é o sc ee ege a ion eco ded in he Vayo s
Dzo P o ince nea He mon. The es o he clus e s a e
in e p e ed a he communi y o associa ion le el. Clus-
e s 1.1.1.1–1.1.3.2 consis ed o he mos xe ic plo s o
he semi-dese , ocky and ho n-cushion ege a ion in
he lowe ele a ions. Clus e s 2.1.1.1 and 2.1.1.2 ep e-
sen plo s o moun ain meadow s eppes om he highes
ele a ions. Clus e s 2.2.1.1–2.2.1.4 consis ed o plo s o
ho n-cushion and s eppic g asslands o so-called moun-
ain s eppes.
O dina ion o he A menian da ase
O dina ion o he plo s wi h he assignmen o one o
hese clus e s e ealed ha he i s axis o he DCA
g aph (Figu e 5) co esponds o a g adien o mois u e
and empe a u e connec ed wi h he ele a ion ange.
The mos mesic plo s occupied highe al i udes (clus e
2.1.1.1), while he mos xe ic ones we e dis ibu ed in he
lowe ele a ions (clus e s 1.1.2.1–1.1.3.2). The clima -
ic-ele a ion g adien and communi y pa ame e s co e-
la ed wi h he di e en ia ion o wo highe classi ica ion
uni s – Zizipho a enuio -S ipa a abica g asslands and
he class Fes uco-B ome ea.
Syn axonomic scheme
Resul ing om ou analyses o he A menian da a and
he compa ison wi h he syn axa o neighbou ing e i-
o ies, we p opose he ollowing syn axonomic scheme
Figu e 3. DCA o he Wes Asian da ase (DCA wi h sup-
plemen a y a iables, eigen alues/g adien leng hs/cu-
mula i e explained a ia ion o axis 1: 0.6313/5,37/4.02,
axis 2: 0.4207/4.94/6.69). Vec o s (en i onmen al a ia-
bles): BIO1: annual mean empe a u e; BIO7: empe a-
u e annual ange; BIO12: annual p ecipi a ion; ele a ion:
ele a ion (m a.s.l.); N species: ascula plan ichness.
Figu e 4. Resul s o he Modi ied TWINSPAN classi ica ion o he A menian plo s (n = 111). The e minal clus e s we e
in e p e ed as associa ions o , i ep esen ed by oo ew plo s, as in o mal communi ies a associa ion ank. The i s
clus e (X) consis ed only o one plo o sc ee ege a ion. Codes a he ips o he o he clus e s co espond o Table 1.
Denys Vynoku o e al.: D y g asslands o A menia54
Syn axon All 1 2 1.1 2.1 2.2 1.1.1 1.1.2 1.1.3 2.1.1 2.2.1 1.1.1.1 1.1.1.2 1.1.2.1 1.1.3.1 1.1.3.2 2.1.1.1 2.1.1.2 2.2.1.1 2.2.1.2 2.2.1.3 2.2.1.4
Numbe o plo s 110 47 63 47 21 42 17 13 17 21 42 7 10 13 10 7 13 8 8 7 16 11
Mean species ichness in 10 m2 (all) 47.4 45.0 49.2 45.0 50.1 48.8 43.5 45.0 46.6 50.1 48.8 42.4 44.2 45.0 45.3 48.4 56.8 39.3 49.1 52.3 42.3 55.8
Mean species ichness in 10 m2 ( ascula plan s) 46.9 44.6 48.7 44.6 49.3 48.3 43.2 44.9 45.9 49.3 48.3 41.9 44.1 44.9 44.6 47.7 55.6 39.1 48.9 51.4 41.9 55.3
Mean species ichness in 10 m2 (byophy es) 0.4 0.2 0.5 0.2 0.6 0.4 0.3 0.1 0.3 0.6 0.4 0.6 0.1 0.1 0.3 0.3 0.9 0.0 0.3 0.7 0.3 0.5
Mean species ichness in 10 m2 (lichens) 0.1 0.1 0.1 0.1 0.2 0.0 0.0 0.0 0.4 0.2 0.0 0.0 0.0 0.0 0.4 0.4 0.2 0.1 0.0 0.1 0.1 0.0
Rhinan hus subula us 2 . 3 . . 5 . . . . 5 . . . . . . . . 29 . .
La hy us la i olius 2 . 3 . . 5 . . . . 5 . . . . . . . . 29 . .
Vale iana o icinalis agg . 2 . 3 . . 5 . . . . 5 . . . . . . . . 29 . .
Campanula bononiensis 2 . 3 . . 5 . . . . 5 . . . . . . . . 29 . .
Melampy um sp. 2 . 3 . . 5 . . . . 5 . . . . . . . . 29 . .
Seseli libano is 8 . 14 . 14 14 . . . 14 14 . . . . . 15 13 13 57 6 .
Assoc. 2.2.1.4
Linum enui olium 15 . 25 . . 38 . . . . 38 . . . . . . . 13 . 44 73
Co onilla co ona a 4 . 6 . . 10 . . . . 10 . . . . . . . . . . 36
Asphodeline au ica 4 . 6 . . 10 . . . . 10 . . . . . . . . . . 36
And osace chamaejasme 4 . 6 . . 10 . . . . 10 . . . . . . . . . . 36
B Hypnum cup essi o me 13 4 19 4 14 21 . 8 6 14 21 . . 8 . 14 15 13 13 . . 73
Campanula sibi ica 9 6 11 6 . 17 18 . . . 17 43 . . . . . . . . . 64
Sesle ia phleoides 6 2 10 2 5 12 . 8 . 5 12 . . 8 . . 8 . . . . 45
Aspe ula a inis 5 . 8 . . 12 . . . . 12 . . . . . . . . . 6 36
Campanula apunculoides 11 . 19 . 5 26 . . . 5 26 . . . . . 8 . 13 43 . 64
Pon echium macula um 7 . 13 . 14 12 . . . 14 12 . . . . . 23 . . . . 45
F i illa ia caucasica 3 . 5 . . 7 . . . . 7 . . . . . . . . . . 27
Di e en ial species in mo e han one associa ion
A ena ia dian hoides 5 . 8 . 14 5 . . . 14 5 . . . . . . 38 25 . . .
Gagea ca oli-kochii 5 . 8 . 10 7 . . . 10 7 . . . . . . 25 38 . . .
Silene cephalan ha 5 . 8 . 10 7 . . . 10 7 . . . . . . 25 38 . . .
Companion species
Thymus ko schyanus 60 66 56 66 48 60 59 69 71 48 60 43 70 69 60 86 31 75 50 71 56 64
Scu ella ia o ien alis agg . 54 62 48 62 43 50 71 54 59 43 50 100 50 54 50 71 15 88 50 57 44 55
Koele ia mac an ha 52 55 49 55 43 52 59 77 35 43 52 57 60 77 40 29 62 13 38 29 63 64
Ely igia in e media agg . 47 49 46 49 19 60 65 46 35 19 60 71 60 46 50 14 8 38 88 57 44 64
E yngium billa die ei 38 43 35 43 5 50 24 46 59 5 50 14 30 46 60 57 . 13 50 43 69 27
Con ol ulus linea us 36 51 25 51 . 38 71 62 24 . 38 43 90 62 40 . . . 25 . 50 55
Medicago sa i a 35 34 37 34 14 48 29 46 29 14 48 . 50 46 50 . 15 13 38 71 44 45
Alyssum alyssoides 35 43 29 43 19 33 41 38 47 19 33 43 40 38 40 57 23 13 50 . 56 9
Fes uca alesiaca agg . 35 23 43 23 48 40 41 8 18 48 40 14 60 8 20 14 54 38 75 . 38 45
Eupho bia seguie iana 32 40 25 40 10 33 24 62 41 10 33 43 10 62 50 29 8 13 13 29 56 18
Cen au ea o ina agg . 27 38 19 38 . 29 47 46 24 . 29 . 80 46 30 14 . . . 14 56 18
B Syn ichia u alis 27 21 32 21 24 36 6 31 29 24 36 . 10 31 50 . 23 25 38 57 19 45
Dian hus c e aceus 26 21 30 21 52 19 29 31 6 52 19 14 40 31 . 14 54 50 13 29 6 36
Medicago x a ia 26 26 27 26 29 26 41 23 12 29 26 43 40 23 20 . 8 63 13 . 44 27
Zizipho a clinopodioides 25 19 30 19 14 38 24 . 29 14 38 . 40 . 40 14 . 38 13 43 56 27
Hype icum scab um 25 32 19 32 . 29 41 .47 . 29 29 50 . 60 29 . . 13 . 56 18
Leon odon aspe imus 25 13 33 13 33 33 29 . 6 33 33 14 40 . 10 . 23 50 25 14 38 45
Thesium amosum 22 36 11 36 5 14 53 23 29 5 14 43 60 23 30 29 8 . 13 29 6 18
Plan ago lanceola a 22 15 27 15 10 36 35 . 6 10 36 14 50 . 10 . 8 13 63 43 25 27
Onob ychis co nu a 21 19 22 19 . 33 29 8 18 . 33 . 50 8 20 14 . . . 29 44 45
Falca ia ulga is 20 17 22 17 10 29 18 8 24 10 29 14 20 8 40 . 8 13 13 57 38 9

Vege a ion Classi ica ion and Su ey 55
Po en ial class 1: Zizipho a enuio -S ipa a abica g ass-
lands – Wes e n Asian d y g asslands and xe ic ho n
cushion communi ies
Diagnos ic species: Achillea a abica, Aegilops cylind i-
ca, Ag opy on c is a um, Allium pseudo la um, Alyssum
u kes anicum, And osace maxima, Anisan ha ec o um,
A abis au icula a agg ., A ena ia se pylli olia agg ., A e-
misia ag ans, Aspe ula a ensis, B omus dan honiae, B.
japonicus, B. squa osus, Cen au ea agg ega a, Cha dinia
o ien alis, C epis sanc a, C upina ulga is, Dian hus o ien-
alis, Helian hemum ledi olium, Helich ysum plica um, Ho-
los eum ma gina um, H. umbella um, Ma ubium pa -
i lo um, Meniocus lini olius, Minua ia hama a, Noaea
muc ona a, Poa bulbosa, Side i is mon ana, S achys in la a,
S. la anduli olia, S ipa a abica, S. capilla a, S. holose icea,
Taenia he um capu -medusae subsp. c ini um, Tanace-
um au eum, Teuc ium capi a um, Thymelaea passe ina,
Xe an hemum squa osum, Zizipho a enuio .
Ecology and dis ibu ion. Communi ies o he po-
en ial new class occu in he lowe ele a ions in d y
condi ions and include semi-dese ege a ion, xe ic
ho n-cushion communi ies, and xe ic g asslands. Wi hin
A menia, i is ep esen ed by one o de and h ee alliances.
O de 1.1: Cousinio b achyp e ae-S ipe alia a abicae –
Wes e n Asian d y g asslands and xe ic ho n cushion
communi ies
Diagnos ic species: Iden ical wi h hose o he mono ypic
class.
Ecology and dis ibu ion. Communi ies o his o -
de a e dis ibu ed in T anscaucasia and possibly e en
b oade wi hin Wes e n Asia. We expec hem o occu
h oughou Wes e n Asia in d y condi ions (semi-dese s,
d y s eppe-like communi ies, low-ele a ion ho n-cush-
ion communi ies). Also, acco ding o ou analysis, com-
muni ies o his o de may occu e en in highe ele a ions
on ocky subs a es.
Alliance 1.1.1: Onob ychido michauxii-S ipion capilla-
ae – T anscaucasian ocky d y g asslands
Diagnos ic species: As acan ha condensa a, Onob ychis
michauxii, Sal ia ae hiopis, S achys la anduli olia, Teuc i-
um capi a um, Ve onica mul i ida (mainly nega i ely di -
e en ia ed cen al alliance).
Ecology and dis ibu ion. Communi ies o his alli-
ance a e dis ibu ed in highe ele a ions han hose o he
o he wo alliances included in his o de . This alliance is
a ansi ional uni be ween his o de and he o de On-
ob ychido anscaucasicae-S ipe alia pulche imae, com-
p ising T anscaucasian moun ain s eppes (see below).
1.1.1.1: S achys la anduli olia-As acan ha condensa a
communi y (Figu e 7B)
Diagnos ic species: Aspe ula a ensis, As acan ha con-
densa a, Cen au ea ph ygia subsp. abb e ia a, C epis cil-
ia a, Eupho bia o ien alis, Gypsophila elegans, He nia ia
hi su a, Lep unis ichodes, Melica au ica, Nepe a acemo-
sa, Onob ychis michauxii, Onosma se osa, Sal ia ae hiopis,
Sempe i um anscaucasicum, S achys la anduli olia,
Tanace um au eum, Teuc ium o ien ale, T agopogon dubi-
us, Viola occul a, Zosima absin hii olia.
S uc u e, ecology and dis ibu ion. We sampled his
ege a ion ype in he Gegha kunik ( icini y o he own
o Se an, Sho ja) and Vayo s Dzo (He mon, icini y o
Gnishik and Khachik) p o inces. These communi ies
we e loca ed a he mos sou h- acing ocky slopes wi h
shallow soil and low humus con en . The he b laye was
spa se and wi h a high ep esen a ion o I ano-Tu ani-
an species, e.g. As acan ha condensa a, A. mic ocephala,
S achys la anduli olia, Teuc ium o ien ale.
1.1.1.2: Ma ubio pa i lo ae-S ipe um capilla ae (Fig-
u e 7C)
Diagnos ic species: Allium ca dios emon, Cen au ea
o ina agg ., Eupho bia condyloca pa, Ma ubium pa i-
lo um, S ipa capilla a.
S uc u e, ecology and dis ibu ion: These communi-
ies we e sampled on slopes wi h shallow ocky subs a es
in Gegha kunik (A danish), Lo i (nea Shi akamu ) and
Vayo s Dzo ( icini y o Gnishik and Khachik) p o inces.
The associa ion di e ed by a highe he b laye co e com-
pa ed o he p e ious communi y and he highes pa ici-
pa ion o hemic yp ophy es among all associa ions o he
class. The dominan species we e Fes uca alesiaca agg .,
Onob ychis co nu a and Teuc ium capi a um.
Alliance 1.1.2: A emision ag an is – T anscaucasian
wo mwood semi-dese s
Diagnos ic species: Ag opy on c is a um, Allium pseudo-
la um, Alyssum u kes anicum, And osace albana, A e-
na ia se pylli olia agg ., A emisia ag ans, As agalus
hyalolepis, B omopsis ipa ia, Ce a ocephala alca a,
Cousinia b achyp e a, C upina ulga is, Cuscu a a a-
a ica, Consolida hispanica, Didymodon ophaceus (B),
Meniocus lini olius, Minua ia hama a, Noaea muc ona-
a, Peganum ha mala, Polygala hohenacke iana, Scle oca -
yopsis spinoca pos, Syn ichia canine is (B).
Ecology and dis ibu ion: A emisia ag ans
semi-dese s in A menia a e dis ibu ed in he lowes ele-
a ions in he coun y. We did no sample o he semi-des-
e ypes, bu we can expec ha A menian loamy and
sandy semi-dese s will also be p obably included in his
uni . In ou da ase , his alliance is ep esen ed by a single
associa ion.
1.1.2.1: Noaeo muc ona ae-A emisie um ag an is
(Figu e 7D)
Diagnos ic species: iden ical wi h hose o he mono ypic
alliance.
S uc u e, ecology and dis ibu ion: This associa ion
is ypical o he A aga so n p o ince ( icini y o Dash a-
dem and Ta ool). The sampled plo s we e dis ibu ed a
he lowes ele a ions wi h he highes mean annual em-
pe a u e and lowes mean annual p ecipi a ion compa ed
o he o he s udied associa ions. The communi ies we e
domina ed by A emisia ag ans, Poa bulbosa, and Taeni-
Denys Vynoku o e al.: D y g asslands o A menia56
a he um capu -medusae subsp. c ini um. The he b laye is
ela i ely spa se and wi h a high ep esen a ion o he o-
phy es (e.g. Alyssum u kes anicum, B omus squa osus,
Ce a ocephala alca a, C upina ulga is, Scle oca yopsis
spinoca pos) and cha ac e is ic chamaephy es (A emisia
ag ans, Noaea muc ona a).
Alliance 1.1.3: Acan holimono ca yophyllacei-S ipion
holose iceae – T anscaucasian d y g asslands and xe ic
ho n-cushion communi ies
Diagnos ic species: Acan holimon edicum, Aegilops cy-
lind ica, Ae hionema ca neum, A abis au icula a agg .,
B omus japonicus, Bunium mic oca pum, Cha dinia o i-
en alis, C epis sanc a, C upina ulga is, Ephed a p oce a,
Galium e icilla um, Gaudiniopsis mac a, Helian hemum
ledi olium, Noccaea pe olia a, Papa e minus, Roeme -
ia hyb ida, S achys in la a, S ipa a abica, S. holose icea,
Taenia he um capu -medusae subsp. c ini um, Pe o hagia
c e ica, Zizipho a enuio .
Ecology and dis ibu ion: This uni comp ises ege a-
ion adi ionally known as ‘highland xe ophy ic ege a-
ion’. I includes d y g asslands and xe ic agacan h com-
muni ies dis ibu ed abo e he semi-dese bel and below
he moun ain s eppe al i udinal bel . We dis inguish one
associa ion and one communi y wi hin his alliance.
1.1.3.1: Acan holimono ca yophyllacei-S ipe um holos-
e iceae (Figu e 7E)
Diagnos ic species: Acan holimon ca yophyllaceum,
Aegilops iuncialis, Ca duus hamulosus, C epis sanc a,
Ge anium lucidum, Lomelosia o a a, Noccaea annua, S i-
pa zalesskii subsp. pon ica, To ilis a ensis, Xe an hemum
squa osum.
S uc u e, ecology and dis ibu ion: We sampled his
associa ion mainly in he Vayo s Dzo p o ince (He mon,
icini ies o A eni, Gnishik and Khachik), and also in one
locali y in A aga so n p o ince (nea Ta ool). The com-
muni ies we e dis ibu ed on shallow soils, bu wi h high-
e humus con en and lowe g a el co e compa ed o he
o he associa ions o his class. Acan holimon ca yophylla-
ceum, S ipa holose icea, and Taenia he um capu -medusae
subsp. c ini um a e he dominan species in his associa-
ion. Among o he species, I ano-Tu anian elemen s o -
en occu , such as Achillea a abica, E yngium billa die ei,
Hype icum scab um, and Thymus ko schyanus.
1.1.3.2: S achys in la a-Acan holimon edicum commu-
ni y (Figu e 7F)
Diagnos ic species: Acan holimon edicum, Aegilops bi-
uncialis, A. cylind ica, Ae hionema ca neum, And osace
maxima, A abis au icula a agg ., A emisia ag ans,
Aspicilia hispida (L), As agalus o ni hopodioides, Buni-
um mic oca pum, Callipel is cuculla ia, Camelina laxa,
Cha dinia o ien alis, Cousinia da alaghezica, C ossidi-
um squami e um (B), C ucianella exaspe a a, C upina
ulga is, Cuscu a pedicella a, Ephed a p oce a, Galium
e icilla um, Gaudiniopsis mac a, Helian hemum ledi-
olium, Holos eum ma gina um, Lac uca ube osa, Lami-
um amplexicaule, Onob ychis a opa ana, Papa e mi-
nus, Pe o hagia c e ica, S achys in la a, S ipa a abica,
Tanace um au eum, T inia glauca, Vale ianella co ona a,
Zizipho a enuio .
S uc u e, ecology and dis ibu ion: We sampled his
ege a ion ype in A a a ( icini y o Tig anashen) and
Vayo s Dzo ( icini y o Gnishik) p o inces. Communi-
ies we e dis ibu ed a lowe ele a ions wi h high mean
annual empe a u e and low mean annual p ecipi a ion.
The subs a e di e ed by he mos alkaline soil eac ion
(mean pH: 8). The he b laye was spa se and wi h a high
ep esen a ion o I ano-Tu anian and Medi e anean
he ophy e species (Aegilops spp., C upina ulga is, Pe o -
hagia c e ica, Zizipho a enuio ), while he co e o hemic-
yp ophy es was he lowes among all s udied communi-
ies. These communi ies did no ha e clea dominan s, bu
Cha dinia o ien alis, S achys in la a, S ipa a abica, and S.
sa ep ana subsp. aniso icha occu ed wi h highe co e
han he o he species. The species ichness o ascula
plan s, b yophy es and lichens was highe compa ed o
he o he associa ions o he class.
Class 2: Fes uco-B ome ea – Mesoxe ic and xe ic ba-
siphilous g asslands o empe a e Eu ope and adjacen
egions
Diagnos ic species: Abie inella abie ina (B), Achillea
mille olium agg ., A emisia absin hium, Bupleu um al-
ca um agg ., Campanula glome a a agg ., C. apunculoi-
des, C. s e enii, Ci sium leucocephalum, Dac ylis glome -
a a, Galium e um, Koele ia albo ii, Leon odon hispidus,
Linum ne osum, L. enui olium, Lo us co nicula us,
Onob ychis anscaucasica, O iganum ulga e, Phleum
phleoides, Pimpinella saxi aga agg ., Plan ago a a a, Poa
p a ensis agg ., Polygala ana olica, Polygonum cogna um,
Po en illa ec a agg ., Scabiosa bipinna a, Secu ige a a ia,
S achys mac os achys, Ta axacum sec . Ta axacum, Thalic-
um minus, T i olium alpes e, T. ambiguum, T. ichoce-
phalum, T ise um la escens.
Ecology and dis ibu ion: Wi hin A menia, his class
comp ises meso-xe ic g asslands and moun ain s eppes a
highe ele a ions. We dis inguish wo o de s ep esen ing
di e en al i udinal bel s.
O de 2.1: Plan agini a a ae-B omopsie alia a iega ae
– High-moun ain meso-xe ic g asslands o he Caucasus
Diagnos ic species: Achillea mille olium agg ., Ajuga o i-
en alis, Alchemilla se icea, A ena ia blepha ophylla agg .,
A. gypsophiloides, As e alpinus, B achypodium pinna um,
B omopsis a iega a, Campanula collina, C. s e enii,
Ci sium leucocephalum, Fes uca o ina agg ., Schedono us
p a ensis, Fes uca ub a agg ., Filipendula ulga is, Gagea
glacialis, Galium co da um, Gen iana sep em ida, Huynhia
pulch a, Koele ia albo ii, La hy us digi a us, Lomelosia
caucasica, Lo us co nicula us, Luzula mul i lo a, Medi-
cago papillosa, Musca i a meniacum, Myoso is alpes is,
O ni hogalum sigmoideum, Papa e o ien ale, Pedicula is
condensa a, Phleum alpinum, Pilosella o icina um agg .,
Pimpinella saxi aga agg ., Plan ago a a a, Poa p a ensis
Vege a ion Classi ica ion and Su ey 57
agg ., Pohlia nu ans (B), Polygonum cogna um, Po en illa
a gen ea, Psephellus xan hocephalus, Pseudoleskella ec o-
um (B), Pulsa illa albana, Ranunculus caucasicus, Rumex
ace osella, Senecio pseudo-o ien alis, S achys mac an ha,
S ipa i sa, Ta axacum sec . Ta axacum, Thymus collinus,
To ula acaulon (B), T i olium ambiguum, T. spadiceum,
T. ichocephalum, Ve bascum speciosum, Ve onica denu-
da a, V. gen ianoides.
Ecology and dis ibu ion: Communi ies o his o de
occupy he highes sampled ele a ions in A menia: uppe
subalpine and lowe alpine bel s. They o m a pa icu-
la uni ecognized in he dominan app oach ypology:
moun ain meadow s eppes. Beyond his ele a ion bel ,
hey a e eplaced by he alpine g asslands which possibly
belong o he class Junce ea i idi Hadač in Klika e Hadač
1944 (Fes uce alia wo onowii Tsepko a 1987).
Alliance 2.1.1: A emisio chamaemeli oliae-B omopsion
a iega ae – Caucasian subalpine and lowe -alpine me-
so-xe ic g asslands
Diagnos ic species: Achillea mille olium agg ., Alche-
milla se icea, A ena ia gypsophiloides, B omopsis a iega-
a, Campanula collina, C. s e enii, Fes uca o ina agg ., F.
ub a agg ., Huynhia pulch a, Koele ia albo ii, La hy us
digi a us, Lomelosia caucasica, Lo us co nicula us, Myo-
so is alpes is, Plan ago a a a, Polygonum cogna um, Po-
en illa a gen ea, Ranunculus caucasicus, Ta axacum sec .
Ta axacum, T i olium ambiguum, T. ichocephalum, Ve-
onica denuda a, V. gen ianoides.
Ecology and dis ibu ion: This uni was desc ibed
om he Main Range o he No h Caucasus (Vynoku o
e al. 2021) in he ele a ions o 1,800–2,200 m a.s.l. In A -
menia, i occu s mainly highe han 2,000 m a.s.l., and
sha es mul iple species wi h he No he n Caucasus uni .
Thus, we a e classi ying moun ain meadow s eppes o A -
menia wi hin he same alliance. He e we dis inguish one
associa ion and one in o mal communi y wi hin i .
2.1.1.1: Ranunculo caucasici-B omopsie um a iega ae
(Figu e 8A)
Diagnos ic species: Alchemilla se icea, A ena ia bleph-
a ophylla agg ., A emisia chamaemeli olia, As e alpinus,
A enula pubescens, B achypodium pinna um, B omopsis
a iega a, Campanula collina, C. s e enii, Ca ex ca yo-
phyllea, Ci sium leucocephalum, Schedono us p a ensis,
Galium co da um, Huynhia pulch a, Lomelosia caucasi-
ca, Luzula mul i lo a, Medicago papillosa, Musca i a meni-
acum, Myoso is alpes is, Pedicula is condensa a, Phascum
cuspida um (B), Phleum alpinum, Pilosella o icina um
agg ., Pimpinella saxi aga agg ., Plan ago a a a, Polygo-
num cogna um, Psephellus xan hocephalus, Pulsa illa al-
bana, Ranunculus caucasicus, Rumex ace osella, S achys
mac an ha, S ipa i sa, T agopogon g amini olius, T i oli-
um ichocephalum, Ve onica gen ianoides.
S uc u e, ecology and dis ibu ion: We sampled his
ege a ion ype a he s eep no h- acing slopes a ele a-
ions a ound 2,100 m, mainly in he Shi ak p o ince ( i-
cini ies o Amasia and Zo ake ), and also in Gegha kunik
p o ince (A danish). The locali ies we e cha ac e ised by
a high mean annual p ecipi a ion (a ound 700–900 mm).
The soil eac ion was sligh ly acidic (mean pH: 6.5). The
associa ion di e ed by high species ichness and he
highes pa icipa ion o Caucasian species, e.g. Dian hus
c e aceus, Lomelosia caucasica, S achys mac an ha, and
T i olium ichocephalum, among all s udied communi-
ies. G aminoids we e dominan , pa icula ly B achypo-
dium pinna um, B omopsis a iega a, Ca ex humilis, and
Phleum alpinum.
2.1.1.2: T agopogon e icula us-As acan ha au ea com-
muni y (Figu e 8B)
Diagnos ic species: A ena ia dian hoides, A. gypsophiloi-
des, As acan ha au ea, Campanula s e enii, Ely igia e-
pens, Gagea glacialis, Koele ia albo ii, La hy us digi a us,
Papa e o ien ale, Plan ago a a a, Senecio pseudo-o ien a-
lis, T agopogon e icula us, T i olium ambiguum, T. spadi-
ceum, T ise um la escens, Ve bascum speciosum.
S uc u e, ecology and dis ibu ion: We sampled his
communi y mainly in he Gegha kunik p o ince (Selim
pass), and also in he A aga so n p o ince (nea he o -
ess o Ambe d). Mos o he locali ies we e si ua ed a
2,300–2,400 m a.s.l. and ep esen ed he highes ele a ions
among all s udied si es. The soil eac ion was sligh ly acid-
ic. These communi ies a e cha ac e ised by low species
ichness wi h a high pa icipa ion o T anscaucasian and
Caucasian species, e.g. A ena ia dian hoides, As acan ha
au ea, Koele ia albo ii. Fes uca o ina agg . and Plan ago
a a a we e he dominan species.
O de 2.2: Onob ychido anscaucasicae-S ipe alia pul-
che imae – T anscaucasian moun ain s eppes
Diagnos ic species: A emisia absin hium, Bupleu um al-
ca um agg ., Campanula glome a a agg ., C. apunculoides,
Ce in he mino , Dac ylis glome a a, Euph asia pec ina a,
Galium e um, Globula ia ichosan ha, Helic ochloa a -
meniaca, Hype icum pe o a um, Klasea adia a, Linum
ne osum, L. enui olium, Lo us co nicula us, Nepe a nuda,
Onob ychis anscaucasica, O iganum ulga e, Phlomis u-
be osa, Polygala ana olica, Rosa spinosissima, Sal ia e i-
cilla a, Scabiosa bipinna a, Secu ige a a ia, S achys mac-
os achys, S. ec a, S ipa penna a, S. pulche ima, Teuc ium
chamaed ys, Thalic um minus, Vicia canescens subsp.
a iega a, Viola ambigua.
Ecology and dis ibu ion: Moun ain s eppes in he
T anscaucasus o m a dis inc al i udinal bel abo e high-
land xe ophy e ege a ion (Cousinio b achyp e ae-S ipe a-
lia a abicae) and below moun ain meadow s eppes (Plan-
agini a a ae-B omopsie alia a iega ae). In ou da ase ,
he o de is ep esen ed by one alliance.
Alliance 2.2.1: Onob ychido anscaucasicae-S ipion
pulche imae – T anscaucasian moun ain s eppes
Diagnos ic species: Campanula apunculoides, Ce in he
mino , Dac ylis glome a a, Linum ne osum, L. enui olium,
Onob ychis anscaucasica, O iganum ulga e, Scabiosa bi-
pinna a, Secu ige a a ia, S achys mac os achys, S. ec a.
Denys Vynoku o e al.: D y g asslands o A menia58
Ecology and dis ibu ion: Despi e i s high lo is ic
he e ogenei y, we uni e all moun ain s eppes in o one alli-
ance. We dis inguish wo in o mal communi ies and wo
associa ions.
2.2.1.1: T ise um la escens-S achys mac os achys com-
muni y (Figu e 8C)
Diagnos ic species: A ena ia g aminea, A emisia absin-
hium, Chae ophyllum oseum, Gagea ca oli-kochii, Silene
cephalan ha, S achys mac os achys, T ise um la escens,
Ve bascum chei an hi olium.
S uc u e, ecology and dis ibu ion: We sampled
his ege a ion ype a he ele a ions 1,950–2,300 m
a.s.l. in A aga so n (nea Ambe d o ess), Ge-
gha kunik (Selim pass, Sho ja) and Shi ak ( icini y
o Amasia) p o inces. These communi ies de elop on
soils wi h high humus con en . The he b laye is el-
a i ely dense wi h dominance o g asses (Ely igia in-
e media agg ., Phleum nodosum, Poa p a ensis agg .,
T ise um la escens) and legumes (Secu ige a a ia,
T i olium alpes e, Vicia enui olia subsp. a iabilis, V.
canescens subsp. a iega a).
Figu e 7. D y g assland and ho n-cushion communi ies o A menia ha do no belong o he Fes uco-B ome ea.
A. Eupho bia o ien alis-Melilo us o icinalis sc ee communi y nea He mon (Vayo s Dzo P o ince); B. S achys la an-
duli olia-As acan ha condensa a communi y (alliance 1.1.1); C. Ma ubio pa i lo ae-S ipe um capilla ae (alliance
1.1.1); D. Semi-dese s o he associa ion Noaeo muc ona ae-A emisie um ag an is (alliance 1.1.2); E. Highland
xe ophy ic ege a ion o he Acan holimono ca yophyllacei-S ipe um holose iceae (alliance 1.1.3); F. S achys in la-
a-Acan holimon edicum communi y (alliance 1.1.3) (Pho os: A, C, E: Jü gen Dengle ; B: Thomas Becke ; D: Denys
Vynoku o ; F: Da iia Bo o yk).
Vege a ion Classi ica ion and Su ey 59
2.2.1.2: Onob ychis anscaucasica-Vicia canescens sub-
sp. a iega a communi y (Figu e 8D)
Diagnos ic species: A abis hi su a, Campanula bononien-
sis, Campyliadelphus ch ysophyllus (B), Chae ophyllum
au eum, Daphne oleoides subsp. ku dica, Helic ochloa a -
meniaca, Klasea adia a, La hy us la i olius, Linum ne o-
sum, Nepe a nuda, Onob ychis anscaucasica, O iganum
ulga e, Phlomis ube osa, P imula e is subsp. mac o-
calyx, Rhinan hus subula us, Sal ia nemo osa, Secu ige a
a ia, Seseli libano is, S achys mac os achys, S ipa zalesskii
subsp. canescens, Thalic um minus, Vale iana o icinalis
agg ., Vicia canescens subsp. a iega a, Viola ambigua.
S uc u e, ecology and dis ibu ion: We eco ded ele-
és o his communi y only in he Vayo s Dzo p o ince, a
no h- acing slopes (inclina ion 10–40°) in he icini y o
Gnishik and be ween Khachik and A eni. Communi ies
di e ed by closed he b laye and high li e co e . The spe-
cies composi ion is cha ac e ised by a high pa icipa ion
o o bs wi h Eu opean dis ibu ion, such as Campanu-
la bononiensis, Klasea adia a, and Secu ige a a ia. The
dominan species is Vicia canescens subsp. a iega a.
2.2.1.3: Globula io ichosan hae-S ipe um pulche i-
mae (Figu e 8E)
Diagnos ic species: cen al associa ion (no diagnos ic
species)
S uc u e, ecology and dis ibu ion: We sampled his
ege a ion ype a ele a ions a ound 1,900–2,200 m a.s.l.
in he p o inces o Gegha kunik (A danish and icini y
o he own o Se an), Shi ak (Jaju pass) and Vayo s Dzo
( icini ies o Gnishik, Khachik and A eni). In he species
composi ion, p e ailing g oups o species we e Eu opean
(Po en illa ec a agg ., S achys ec a, S ipa pulche ima)
and I ano-Tu anian (E yngium billa die ei, Onob ychis
michauxii, Thymus ko schyanus, Zizipho a clinopodioides),
ollowed by Caucasian endemics (As agalus cancella us,
Cen au ea pseudoscabiosa, Scabiosa bipinna a). The domi-
nan species o he associa ion was S ipa pulche ima.
2.2.1.4: Sesle io phleoidis-Onob ychide um co nu ae
(Figu e 8F)
Diagnos ic species: Abie inella abie ina (B), Adonis olgen-
sis, And osace chamaejasme, Aspe ula a inis, Asphodeline
au ica, B iza media, Campanula apunculoides, C. sibi ica,
Co onilla co ona a, Eupho bia esula agg ., Euph asia pec -
ina a, F i illa ia caucasica, Homalo hecium lu escens (B),
Hypnum cup essi o me (B), Leucan hemum ulga e, Linum
enui olium, Pimpinella saxi aga agg ., Pinus syl es is,
Pon echium macula um, Psephellus ka abaghensis, Sesle ia
phleoides, Spi aea c ena a, Thalic um oe idum, Viola alba.
S uc u e, ecology and dis ibu ion: We sampled
his associa ion a ele a ions 1,940–2,070 m a.s.l. in Ge-
gha kunik (A danish, Sho ja, icini y o he own o Se-
an) and Shi ak (Jaju pass) p o inces. The associa ion
di e ed by he highes mean o al species ichness and
ichness o b yophy es ac oss all s udied communi ies.
The species composi ion is ep esen ed by a high pa ic-
ipa ion o Eu opean species o g asses and o bs, such as
B iza media, Campanula apunculoides, Galium e um,
Pimpinella saxi aga agg ., and S achys ec a. Ca ex humi-
lis, Onob ychis co nu a and Teuc ium chamaed ys a e he
dominan species.
Di e en ia ion o he syn axa wi h espec o
ecology, s uc u e and biodi e si y
Topog aphy, clima e and soil
Communi ies wi hin he Fes uco-B ome ea p edominan -
ly h i ed a highe ele a ions, especially hose o Plan-
agini a a ae-B omopsie alia a iega ae, eaching up o
2,400 m a.s.l. (Figu e 9A). Gene ally, hey occu ed wi h-
in he ele a ion ange o 1,900–2,400 m a.s.l., wi h some
excep ions, like in he case o he Onob ychis anscauca-
sica-Vicia canescens subsp. a iega a communi y, which
we e ound be ween 1,700 o 2,100 m a.s.l. These low ele-
a ion occu ences we e compensa ed by local opog aph-
ic p e e ences, pa icula ly a oidance o d ie slopes wi h
a sou he ly aspec . This communi y exhibi ed he lowes
sou he ness index among all syn axa (Figu e 9B).
In con as , communi ies o he Zizipho a enuio -S i-
pa a abica g asslands wi h i s o de Cousinio b achyp e -
ae-S ipe alia a abicae occu ed a lowe ele a ions, p i-
ma ily below 2,000 m a.s.l. Among i s h ee alliances,
A emision ag an is, which comp ises wo mwood
semi-dese s, h i ed a he lowes al i udes, anging om
1,300 o 1,600m a.s.l. Addi ionally, he S achys in la a-Ac-
an holimon edicum communi y, which belongs o he
alliance Acan holimono ca yophyllacei–S ipion holose i-
ceae, also occupied compa ably low ele a ions, a abou
1,600m a.s.l. O he uni s o his o de gene ally occupy
ele a ions no exceeding 2,000 m a.s.l.
The al i udinal zona ion e lec ed he clima ic p e e -
ences o he communi ies. The Cousinio b achyp e ae-S i-
pe alia a abicae communi ies ended o p e e wa me and
d ie condi ions, while he class Fes uco-B ome ea h i ed
in mo e mesic and coole en i onmen s (Figu e 9C, D).
Rega ding soil cha ac e is ics, mos communi ies p e-
e ed neu al o sligh ly alkaline soils, excep o he o de
Plan agini a a ae-B omopsie alia a iega ae, which oc-
cu ed in sligh ly acidic soil condi ions wi h a pH a ound
6.5 (Figu e 10A). This a ia ion aligns wi h he al i udinal
zona ion, e lec ing di e en soil composi ions and eg-
e a ion ypes ac oss ele a ion le els. Lowe and medium
ele a ions (mon ane and lowe subalpine bel ) we e cha -
ac e ised by kas anozems and che nozems, which hos ed
semi-dese and s eppe ege a ion, and had neu al o
sligh ly alkaline eac ion. We classi ied hem wi hin he
o de s Cousinio b achyp e ae-S ipe alia a abicae and On-
ob ychido anscaucasicae-S ipe alia pulche imae espec-
i ely. Uppe subalpine and lowe alpine bel s we e home
o meadow-s eppe soils and moun ain-meadow soils,
which we e cha ac e ised by sligh ly acidic o acidic eac-
ion. He e, moun ain meadow s eppe ege a ion h i ed,
which we summa ised in he o de Plan agini a a ae-B o-
mopsie alia a iega ae. A e en highe al i udes, moun ain
s eppes we e eplaced by acidophilous alpine g asslands,
which we e no co e ed in ou s udy.

Denys Vynoku o e al.: D y g asslands o A menia60
In e ms o skele on con en , communi ies o he
Zizipho a enuio -S ipa a abica g asslands gene ally
ended o occu on si es wi h shallowe soils and highe
skele on con en (Figu e 10B, C). Among hem, S achys
la anduli olia-As acan ha condensa a communi y and
S achys in la a-Acan holimon edicum communi y ep-
esen ocky g asslands, and occu ed on si es wi h
he highes p opo ion o g a el in he uppe soil le el,
eaching up o 80% and 90% espec i ely. Among he
communi ies classi ied in o he class Fes uco-B ome-
ea, we obse ed highe he e ogenei y. Bo h uni s o
he o de Plan agini a a ae-B omopsie alia a iega-
Figu e 8. D y g asslands o A menia classi ied wi hin he Fes uco-B ome ea. A. Moun ain meadow s eppes in
he Lake A pi Na ional Pa k wi h he associa ion Ranunculo caucasici-B omopsie um a iega ae (alliance 2.1.1);
B. T agopogon e icula us-As acan ha au ea communi y (alliance 2.1.1); C. T ise um la escens-S achys mac os-
achys communi y (alliance 2.2.1); D. Onob ychis anscaucasica-Vicia canescens subsp. a iega a communi y (alli-
ance 2.2.1); E. Globula io ichosan hae-S ipe um pulche imae (alliance 2.2.1); F. moun ain s eppes nea he Se an
Lake wi h he associa ion Sesle io phleoidis-Onob ychide um co nu ae (alliance 2.2.1). (Pho os: A: Philipp Ki schne ;
B, C: Da iia Bo o yk; D, F: Jü gen Dengle ; E: Denys Vynoku o ).
Vege a ion Classi ica ion and Su ey 61
ae occu ed on si es wi h low g a el con en . Among
hem, Ranunculo caucasici-B omopsie um a iega ae
was dis ibu ed on deepe soils. Moun ain s eppes o
he o de Onob ychido anscaucasicae-S ipe alia pul-
che imae seemed o occu on si es wi h a ying skel-
e on p opo ion. While he T ise um la escens-S achys
mac os achys and Onob ychis anscaucasica-Vicia ca-
nescens subsp. a iega a communi ies occu ed on si es
wi h low g a el p opo ions, he associa ions Globu-
la io ichosan hae-S ipe um pulche imae and Sesle io
phleoidis-Onob ychide um co nu ae we e dis ibu ed
mainly on ocky si es.
All communi ies o he class Fes uco-B ome ea end-
ed o occu on si es wi h high humus con en , wi h mean
alues wi hin a na ow ange o 7.5–8.1%. The en a i e
Zizipho a enuio -S ipa a abica g asslands occu ed on
si es wi h signi ican ly lowe humus con en , wi h mean
alues o 3.9–4.5% (Figu e 10D).
Figu e 9. Si e cha ac e is ics o he g assland ypes a di e en syn axonomic le els. Box plo s (median, in e qua -
ile ange, ange and ou lie s) as well as a i hme ic means (black poin s) a e shown. Fo he codes o syn axa, see
Table 1. Di e en le e s wi hin one syn axonomic le el indica e signi ican di e ences a p < 0.05 acco ding o
Tukey’s es ollowing a signi ican ANOVA. Fo o de s, he wo-digi codes a e he slashes and he le e s below
he boxplo s apply.
Figu e 10. Soil cha ac e is ics o he g assland ypes a di e en syn axonomic le els. Box plo s (median, in e -
qua ile ange, ange and ou lie s) as well as a i hme ic means (black poin s) a e shown. Fo he codes o syn axa,
see Table 1. Di e en le e s wi hin one syn axonomic le el indica e signi ican di e ences a p < 0.05 acco ding o
Tukey’s es ollowing a signi ican ANOVA. Fo o de s, he wo-digi codes a e he slashes and he le e s below
he boxplo s apply.
Denys Vynoku o e al.: D y g asslands o A menia62
S uc u e and species composi ion
He b laye co e and li e co e , p oxies o ecosys em
p oduc i i y, we e no ably highe in he communi ies be-
longing o he class Fes uco-B ome ea (Figu e 11A, B).
Rega ding he dominan li e o ms, we obse ed a s ong
di e en ia ion be ween he wo classes. Communi ies be-
longing o he class Fes uco-B ome ea had a signi ican ly
highe p opo ion o hemic yp ophy es, which was highes in
he case o he Ranunculo caucasici-B omopsie um a iega ae
associa ion (Figu e 11D). On he con a y, communi ies o
he en a i e new class ‘Zizipho a enuio -S ipa a abica g ass-
lands’ had much lowe p opo ions o hemic yp ophy es, bu
a much highe p opo ion o he ophy es (Figu e 11C). The
p opo ion o he ophy es was pa icula ly high in he case
o he alliances A emision ag an is and Acan holimono
ca yophyllacei-S ipion holose iceae, which ep esen wo m-
wood dese s eppes and xe ophy ic ho n-cushions espec-
i ely. The p opo ion o chamaephy es was also signi ican ly
highe in he case o he la e class (no shown).
The p opo ion o species’ ange ypes also showed a
s ong di e en ia ion be ween he wo classes (Figu e 12).
The class Fes uco-B ome ea was cha ac e ised by a signi i-
can ly highe p opo ion o species wi h Eu opean dis ibu-
ion (Figu e 12A). This sugges s ha he class Fes uco-B om-
e ea comp ises exclusi ely Eu o-Sibe ian s eppe ege a ion
and ha i s dis ibu ion ange eaches a limi in A menia.
This is also well e lec ed by he co-occu ence o wo bio-
geog aphic egions: he Eu o-Sibe ian and he I ano-Tu -
anian. On he con a y, he new en a i e class ‘Zizipho a
enuio -S ipa a abica g asslands’ ep esen s communi ies
wi h a lowe p esence o species wi h Eu opean dis ibu ion,
and signi ican ly highe p opo ions o species wi h Medi-
e anean and I ano-Tu anian dis ibu ion (Figu e 12B, E).
The p esence o endemic species in he s udied
communi ies was also ema kable. While he e we e
no signi ican di e ences de ec ed be ween he wo
classes conce ning na ow T anscaucasian endemics
(Figu e 12D), a dis inc pa e n eme ged o he b oade
Caucasian endemics. These species, anging ac oss he
Caucasus egion, including he No h Caucasus, we e
mo e p e alen in he Fes uco-B ome ea compa ed o he
Zizipho a enuio -S ipa a abica g asslands (Figu e 12C).
Among he wo ecognized o de s o he Fes uco-B ome-
ea, he p opo ion o Caucasian endemics was highe in
he Plan agini a a ae-B omopsie alia a iega ae, which
comp ises moun ain meadow s eppes o highe ele a-
ions. O e all, we obse ed an inc easing p opo ion o
endemic species alongside ele a ion. In e es ingly, he
p opo ion o species wi h o he dis ibu ion anges,
namely, b oade han he lis ed abo e, was also signi i-
can ly highe in he case o he Fes uco-B ome ea plan
communi ies (Figu e 12F).
Plo -scale species ichness
To al species ichness in 10 m2 did no di e signi ican ly
among he highe syn axa (Figu e 13). A he associa ion
le el, he e was no s ong di e en ia ion ei he , wi h only
he Ranunculo caucasici-B omopsie um a iega ae and he
Sesle io phleoidis-Onob ychide um co nu ae being abo e
a e age and T agopogon e icula us-As acan ha au ea
communi y below. Fo ascula plan species ichness,
he pa e n la gely ma ched ha o o al species ichness,
while o lichens he e we e no signi ican di e ences a
any le el. Only b yophy es showed a weak ichness pa -
e n a he o de and alliance le el, wi h he mono ypic
o de Plan agini a a ae-B omopsie alia a iega ae being
he iches .
Figu e 11. S uc u e and dominan li e o ms o he g assland ypes a di e en syn axonomic le els. Box plo s (me-
dian, in e qua ile ange, ange and ou lie s) as well as a i hme ic means (black poin s) a e shown. Fo he codes
o syn axa, see Table 1. Di e en le e s wi hin one syn axonomic le el indica e signi ican di e ences a p < 0.05
acco ding o Tukey’s es ollowing a signi ican ANOVA. Fo o de s, he wo-digi codes a e he slashes and he
le e s below he boxplo s apply.
Vege a ion Classi ica ion and Su ey 63
Figu e 12. P opo ions o ange ypes o he g assland ypes a di e en syn axonomic le els. Box plo s (median,
in e qua ile ange, ange and ou lie s) as well as a i hme ic means (black poin s) a e shown. Fo he codes o syn-
axa, see Table 1. Di e en le e s wi hin one syn axonomic le el indica e signi ican di e ences a p < 0.05 acco d-
ing o Tukey’s es ollowing a signi ican ANOVA. Fo o de s, he wo-digi codes a e he slashes and he le e s
below he boxplo s apply.
Figu e 13. Species ichness in 10 m2-plo s o di e en axonomic g oups compa ed a di e en syn axonomic le els.
Box plo s (median, in e qua ile ange, ange and ou lie s) as well as a i hme ic means (black poin s) a e shown. Fo
he codes o syn axa, see Table 1. Di e en le e s wi hin one syn axonomic le el indica e signi ican di e ences a
p < 0.05 acco ding o Tukey’s es ollowing a signi ican ANOVA. Fo o de s, he wo-digi codes a e he slashes
and he le e s below he boxplo s apply.
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Appendix 1
Fo mal desc ip ions o he new syn axa acco ding o he
ICPN
Fo he diagnos ic species we e e o he main ex as well
as Table 2 and Suppl. ma e ial 2.
1.1 Cousinio b achyp e ae-S ipe alia a abicae o d. no .
hoc loco
Holo ypus hoc loco: A emision ag an is Vynoku o e al.
2024 ( his pape )
2.1 Plan agini a a ae-B omopsie alia a iega ae o d
no . hoc loco
Holo ypus hoc loco: A emisio chamaemeli oliae-B omopsion
a iega ae Vynoku o in Vynoku o e al. 2021 (page 186)

Vege a ion Classi ica ion and Su ey 71
2.2 Onob ychido anscaucasicae-S ipe alia pulche i-
mae o d. no . hoc loco
Holo ypus hoc loco: Onob ychido anscaucasicae-S ipion
pulche imae Vynoku o e al. 2024 ( his pape )
1.1.1 Onob ychido michauxii-S ipion capilla ae all. no .
hoc loco
Holo ypus hoc loco: Ma ubio pa i lo ae-S ipe um capilla ae
Vynoku o e al. 2024 ( his pape )
1.1.2 A emision ag an is all. no . hoc loco
Holo ypus hoc loco: Noaeo muc ona ae-A emisie um a-
g an is Vynoku o e al. 2024 ( his pape )
1.1.3 Acan holimono ca yophyllacei-S ipion holose ice-
ae all. no . hoc loco
Holo ypus hoc loco: Acan holimono ca yophyllacei-S ipe um
holose iceae Vynoku o e al. 2024 ( his pape )
2.2.1 Onob ychido anscaucasicae-S ipion pulche i-
mae all. no . hoc loco
Holo ypus hoc loco: Sesle io phleoidis-Onob ychide um co -
nu ae Vynoku o e al. 2024 ( his pape )
1.1.1.2 Ma ubio pa i lo ae-S ipe um capilla ae ass.
no . hoc loco
Holo ypus hoc loco: plo ID 81 in Suppl. ma e ial 2 (o igi-
nal code AMR008); A menia, Lo i P o ince, icini y o Shi-
akamu , 40.85801°N, 44.19223°E, al i ude: 1,662 m a.s.l.,
aspec : 210°, inclina ion: 40°, 27 June 2019, au ho s o he
ele é: Die e F ank, Salza Palpu ina, Denys Vynoku o . Flo-
is ic composi ion (species so ed by hei co e in pe cen ):
Ely igia in e media agg . 10, Ma ubium pa i lo um 10,
Achillea a abica 5, Teuc ium polium 5, Onob ychis michauxii
4, A emisia aus iaca 3, Thymus sipyleus 3, Medicago x a ia
2, Co a melanoloma 1, Fes uca alesiaca agg . 1, I is pumila
1, Isa is s e eniana 1, Psephellus zu andicus 1, Xe an hemum
longepapposum 1, Ajuga chamaepi ys subsp. chia 0.5, Aspe u-
la a ensis 0.5, Dac ylis glome a a 0.5, Odon a hena mu alis
0.5, S achys ec a 0.5, Viola ambigua 0.5, Holos eum umbella-
um 0.3, Nonea pulla 0.3, Scle an hus annuus 0.3, Con ol ulus
linea us 0.2, Eupho bia condyloca pa 0.2, S ipa capilla a 0.2,
Ze a schania pauci adia a 0.2, Cen au ea o ina agg . 0.1, Sal-
ia nemo osa 0.1, S ipa a abica 0.1, Thesium a ense 0.1, Fal-
ca ia ulga is 0.01, Meniocus lini olius 0.01, Nepe a acemosa
0.01, Noccaea pe olia a 0.01, Reseda lu ea 0.01, Ve onica mul-
i ida 0.01, Viola a ensis agg . 0.01.
1.1.2.1 Noaeo muc ona ae-A emisie um ag an is ass.
no . hoc loco
Holo ypus hoc loco: plo ID 65 in Suppl. ma e ial 2 (o iginal
code AM11NE); A menia, A aga so n P o ince, icini y o
Dash adem, 40.32428°N, 43.85075°E, al i ude: 1,357 m a.s.l.,
aspec : 265°, inclina ion: 7°, 29 June 2019, au ho s o he ele-
é: Alla Aleksanyan, Jü gen Dengle , Denys Vynoku o . Flo-
is ic composi ion (species so ed by hei co e in pe cen ):
Poa bulbosa 20, A emisia ag ans 17, Peganum ha mala 10,
Taenia he um capu -medusae subsp. c ini um 10, S ipa a a-
bica 1, And osace albana 0.5, Koelpinia linea is 0.5, S ipa x
kolako skyi 0.5, Allium pseudo la um 0.2, C upina ulga is
0.2, Minua ia hama a 0.2, Noaea muc ona a 0.2, A ena ia
se pylli olia agg . 0.1, Alyssum u kes anicum 0.1, As agalus
hyalolepis 0.1, Cousinia b achyp e a 0.1, E ysimum gelidum
0.1, Odon a hena o uosa 0.1, Scle oca yopsis spinoca pos
0.1, Syn ichia canine is 0.1, Xe an hemum squa osum 0.1,
Aegilops cylind ica 0.01, Ce a ocephala alca a 0.01, Co a
ium e ii 0.01, Dian hus c ini us 0.01, D aba e na 0.01,
Eupho bia seguie iana 0.01, Helich ysum plica um 0.01, Ho-
henacke ia exscapa 0.01, Holos eum umbella um 0.01, Lina ia
simplex 0.01, Medicago medicaginoides 0.01, Meniocus lini oli-
us 0.01, Minua ia meye i 0.01, Polygala hohenacke iana 0.01,
Thesium szo i sii 0.01, Thymus collinus 0.01.
1.1.3.1 Acan holimono ca yophyllacei-S ipe um holos-
e iceae ass. no . hoc loco
Holo ypus hoc loco: plo ID 71 in Suppl. ma e ial 2 (o igi-
nal code AMR035); A menia, Vayo s Dzo P o ince, icin-
i y o Khachik, 39.659529°N, 45.201325°E, al i ude: 2,017 m
a.s.l., aspec : 315°, inclina ion: 9°, 4 July 2019, au ho s o he
ele é: Alla Aleksanyan, Idoia Biu un, Da iia Bo o yk. Flo-
is ic composi ion (species so ed by hei co e in pe cen ):
Cha dinia o ien alis 35, Acan holimon ca yophyllaceum 25,
Taenia he um capu -medusae subsp. c ini um 20, Poa bulbo-
sa 15, Anisan ha ec o um 10, Syn ichia u alis 6, E yngium
billa die ei 5, Onob ychis co nu a 3, Medicago sa i a 2, S ipa
eh enbe giana 0.5, S ipa holose icea 2, Teuc ium polium 2,
Achillea a abica 0.5, Con ol ulus linea us 0.5, Gagea ge main-
ae 0.5, Xe an hemum squa osum 0.3, Zizipho a capi a a 0.3,
Cen au ea i ga a 0.2, Hype icum scab um 0.2, Rochelia dis-
pe ma 0.2, Alyssum u kes anicum 0.1, Aspe ula a ensis 0.1,
B omus dan honiae 0.1, B omus japonicus 0.1, Ce a ocephala
alca a 0.1, C epis sanc a 0.1, Eupho bia ibe ica 0.1, Ho deum
bulbosum 0.1, Ma ubium pa i lo um 0.1, Odon a hena o -
uosa 0.1, Po en illa ec a agg . 0.1, Thymus ko schyanus 0.1,
Alyssum alyssoides 0.01, A ena ia se pylli olia agg . 0.01, Cau-
calis pla yca pos 0.01, Holos eum umbella um 0.01, Minua ia
meye i 0.01, Noccaea pe olia a 0.01, Ve onica denuda a 0.01.
2.1.1.1 Ranunculo caucasici-B omopsie um a iega ae
ass. no . hoc loco
Holo ypus hoc loco: plo ID 18 in Suppl. ma e ial 2 (o iginal
code AM06NW); A menia, Shi ak P o ince, icini y o Zo -
ake , 41.10135°N, 43.66933°E, al i ude: 2,114 m a.s.l., aspec :
255°, inclina ion: 35°, 28 June 2019, au ho s o he ele é:
Elena Belono skaya, Jü gen Dengle , Denys Vynoku o . Flo-
is ic composi ion (species so ed by hei co e in pe cen ):
Thymus sipyleus 25, Achillea mille olium agg . 15, Phleum
alpinum 15, Koele ia mac an ha 12, Fes uca o ina agg . 10,
As agalus ince us 8, Ca ex humilis 8, T i olium ambiguum 7,
B omopsis a iega a 5, Hypnum cup essi o me 5, Lo us co nic-
ula us 5, Medicago alca a 5, Abie inella abie ina 2, Myoso is
alpes is 2, Poa p a ensis agg . 2, B yum caespi icium 1, Plan-
ago a a a 1, Secu ige a a ia 1, Weissia b achyca pa 1, A -
enula pubescens 0.5, S eblo ichum con olu um 0.5, Ci sium
leucocephalum 0.5, Gelasia igida 0.5, Huynhia pulch a 0.5,
Lomelosia caucasica 0.5, Rumex ace osella 0.5, Scabiosa bip-
Denys Vynoku o e al.: D y g asslands o A menia72
inna a 0.5, S achys ec a 0.5, Ca ex ca yophyllea 0.3, Hype i-
cum lina ioides 0.3, Poa badensis 0.3, Po en illa a gen ea 0.3,
Schedono us p a ensis 0.3, Ta axacum sec . Ta axacum 0.3,
Teuc ium o ien ale 0.3, Ve onica denuda a 0.3, Galium e -
um 0.2, Papa e monan hum 0.2, Pimpinella saxi aga agg .
0.2, Po en illa ec a agg . 0.2, Ajuga o ien alis 0.1, Alyssum
alyssoides 0.1, A ena ia blepha ophylla agg . 0.1, A ena ia
gypsophiloides 0.1, A emisia absin hium 0.1, Bupleu um al-
ca um agg . 0.1, Campanula s e enii 0.1, Ci sium ob alla um
0.1, Colchicum igynum 0.1, Galium co da um 0.1, Lina ia
schelkownikowii 0.1, Pedicula is condensa a 0.1, Phascum cus-
pida um 0.1, Ranunculus caucasicus 0.1, Scle an hus pe ennis
0.1, Silene bupleu oides 0.1, Silene chlo an ha 0.1, Syn ichia
mon ana 0.1, B yum a gen eum 0.01, C epis alpina 0.01, D a-
ba nemo osa 0.01, Encalyp a ulga is 0.01, He nia ia incana
0.01, Musca i a meniacum 0.01, Polygala alpicola 0.01, Polyg-
onum cogna um 0.01, Sedum ac e 0.01.
2.2.1.3 Globula io ichosan hae-S ipe um pulche imae
ass. no . hoc loco
Holo ypus hoc loco: plo ID 93 in Suppl. ma e ial 2 (o igi-
nal code AM42SE); A menia, Vayo s Dzo P o ince, icin-
i y o Gnishik, 39.675269°N, 45.30616°E, al i ude: 2,136 m
a.s.l., aspec : 80°, inclina ion: 23°, 3 July 2019, au ho s o he
ele é: Asun Be as egi, Idoia Biu un, Denys Vynoku o . Flo-
is ic composi ion (species so ed by hei co e in pe cen ):
Zizipho a clinopodioides 15, Koele ia mac an ha 7, S ipa cap-
illa a 7, Dac ylis glome a a 5, S ipa pulche ima 5, Hype icum
scab um 4, Onob ychis co nu a 4, Helich ysum g a eolens
3, Lo us co nicula us 2, Odon a hena o uosa 2, Teuc ium
chamaed ys 2, Teuc ium polium 2, Cen au ea o ina agg . 1,
Daphne oleoides subsp. ku dica 1, E yngium billa die ei 1,
Galium e icilla um 1, Galium e um 1, Linum enui olium
1, Medicago sa i a 1, S ipa penna a 1, Cicho ium in ybus 0.5,
C ucia a lae ipes 0.5, Cuscu a epi hymum 0.5, Eupho bia
seguie iana 0.5, Hype icum lina ioides 0.5, Plan ago lance-
ola a 0.5, Scabiosa bipinna a 0.5, Secu ige a a ia 0.5, S ipa
i sa 0.5, Tanace um polycephalum subsp. a gy ophyllum 0.5,
Globula ia ichosan ha 0.3, Leon odon hispidus 0.3, S achys
la anduli olia 0.3, S achys ec a 0.3, Ajuga chamaepi ys subsp.
chia 0.2, Ce in he mino 0.2, Leon odon aspe imus 0.2, Linum
ne osum 0.2, Phleum phleoides 0.2, Poa bulbosa 0.2, T agopo-
gon sosnowskyi 0.2, Ve bascum chaixii subsp. aus iacum 0.2,
As agalus cancella us 0.1, Bupleu um alca um agg . 0.1, Ca -
lina ulga is 0.1, Sanguiso ba mino 0.1, Silene bupleu oides
0.1, Thesium a ense 0.1, T inia glauca 0.1, Ve onica mic oca -
pa 0.1, Achillea mille olium agg . 0.01, Ag imonia eupa o ia
0.01, C epis pulch a 0.01, Euph asia pec ina a 0.01, Odon i es
auche i 0.01, Polygala alpicola 0.01.
2.2.1.4 Sesle io phleoidis-Onob ychide um co nu ae ass.
no . hoc loco
Holo ypus hoc loco: plo ID 13 in Suppl. ma e ial 2 (o igi-
nal code AMR076); A menia, Gegha kunik P o ince, icin-
i y o Shoghaka , 40.49547°N, 45.29575°E, al i ude: 1,959 m
a.s.l., aspec : 305°, inclina ion: 30°, 1 July 2019, au ho s o
he ele é: Geo ge Fay ush, Salza Palpu ina, Iulia Vasheniak.
Flo is ic composi ion (species so ed by hei co e in pe -
cen ): Ca ex humilis 20, Onob ychis co nu a 15, Ely igia in-
e media agg . 8, P imula e is subsp. mac ocalyx 7, Thymus
ko schyanus 7, S ipa pulche ima 6, Onob ychis anscaucasica
5, Scu ella ia o ien alis agg . 5, Galium e um 4, Inula aspe a
4, Helian hemum nummula ium 2, Sesle ia phleoides 2, Teu-
c ium chamaed ys 2, Abie inella abie ina 1, B omus scopa ius
1, Campanula sibi ica 1, Eupho bia ibe ica 1, Homalo hecium
lu escens 1, Hypnum cup essi o me 1, Linum enui olium 1,
Lo us co nicula us 1, Medicago sa i a 1, Syn ichia u alis 1,
Zizipho a clinopodioides 1, Achillea mille olium agg . 0.5, As-
acan ha s enonychioides 0.5, Campanula apunculoides 0.5,
Dian hus c e aceus 0.5, Filipendula ulga is 0.5, Linum ne o-
sum 0.5, Plan ago media 0.5, Polygala ana olica 0.5, Scabiosa
bipinna a 0.5, S achys ec a 0.5, Thalic um oe idum 0.5, Ju-
inea squa osa 0.3, Viola ambigua 0.3, Phleum phleoides 0.2,
Poa badensis 0.2, Psephellus ka abaghensis 0.2, Tanace um
au eum 0.2, Con ol ulus linea us 0.1, Fes uca alesiaca agg .
0.1, Pon echium macula um 0.1, T agopogon e icula us 0.1,
Xan hopa melia cam schadalis 0.1, Aspe ula p os a a 0.01,
Ca um caucasicum 0.01, Ce a ia e ice o um 0.01, Euph asia
se anensis 0.01, Hype icum scab um 0.01, O obanche alsa ica
0.01, Teuc ium o ien ale 0.01.
E-mail and ORCID
Denys Vynoku o (denys. ynoku o @gmail.com), ORCID: h ps://o cid.o g/0000-0001-7003-6680
Alla Aleksanyan (alla.alexan[email p o ec ed]m), ORCID: h ps://o cid.o g/0000-0003-4073-1812
Thomas Becke (becke h@uni- ie .de), ORCID: h ps://o cid.o g/0000-0002-7942-5575
Idoia Biu un (idoia.b[email p o ec ed]), ORCID: h ps://o cid.o g/0000-0002-1454-0433
Da iia Bo o yk (da iiabo o [email p o ec ed]), ORCID: h ps://o cid.o g/0000-0001-7140-7201
Geo ge Fay ush (g ay [email protected]), ORCID: h ps://o cid.o g/0000-0002-9710-2200
I zia Ga cía-Mijangos (i zia .ga [email protected]), ORCID: h ps://o cid.o g/0000-0002-6642-7782
Ma in Magnes (ma[email p o ec ed]), ORCID: h ps://o cid.o g/0000-0002-5453-5735
Salza Palpu ina (salza.palpu [email p o ec ed]m), ORCID: h ps://o cid.o g/0000-0003-0416-5622
U e Becke ([email p o ec ed])
Asun Be as egi ([email p o ec ed]), ORCID: h ps://o cid.o g/0000-0003-0456-3305
Bea a Cykowska-Ma zencka (bea acyko[email p o ec ed]), ORCID: h ps://o cid.o g/0000-0002-5468-4909
Iwona Dembicz ([email protected]), ORCID: h ps://o cid.o g/0000-0002-6162-1519
Die e F ank (die e . [email protected] .de), ORCID: h ps://o cid.o g/0000-0002-8602-9570
Vege a ion Classi ica ion and Su ey 73
And eas Hilpold (and eas.hil[email p o ec ed]), ORCID: h ps://o cid.o g/0000-0003-0262-8080
Philipp Ki schne (philipp[email p o ec ed]), ORCID: h ps://o cid.o g/0000-0002-9263-5112
Helmu May ho e (helmu .may ho e @uni-g az.a ), ORCID: h ps://o cid.o g/0000-0002-9433-3847
Ma ine Oganesian ([email protected]), ORCID: h ps://o cid.o g/0000-0002-3297-8519
Iuliia Vasheniak (a hena [email p o ec ed]), ORCID: h ps://o cid.o g/0000-0003-1020-3007
Jü gen Dengle (Co esponding au ho , jue gen.dengle @zhaw.ch), ORCID: h ps://o cid.o g/0000-0003-3221-660X
Supplemen a y ma e ial
Supplemen a y ma e ial 1
Comple e heade da a o he A menian plo s (*.xlsx)
Link: h ps://doi.o g/10.3897/VCS.119253.suppl1
Supplemen a y ma e ial 2
Comple e synop ic able o 110 classi ied A menian plo s wi h pe cen cons ancies and phi alues o he syn axa
o all le els as well as he indi idual plo s (*.xlsx)
Link: h ps://doi.o g/10.3897/VCS.119253.suppl2
Supplemen a y ma e ial 3
De ini ion o addi ional species agg ega es (*.pd )
Link: h ps://doi.o g/10.3897/VCS.119253.suppl3
Supplemen a y ma e ial 4
Value dis ibu ion o all eco ded and analysed nume ical en i onmen al, s uc u al and biodi e si y a iables (*.pd )
Link: h ps://doi.o g/10.3897/VCS.119253.suppl4
Supplemen a y ma e ial 5
Da a sou ces o he Wes Asian and Caucasian da ase (*.pd )
Link: h ps://doi.o g/10.3897/VCS.119253.suppl5
Supplemen a y ma e ial 6
Lis o he ascula plan s om he A menian da ase wi h assignmen o li e o ms and dis ibu ion anges (*.pd )
Link: h ps://doi.o g/10.3897/VCS.119253.suppl6
Supplemen a y ma e ial 7
Synop ic able o he Wes Asian and Caucasian da ase wi h he esul s o he b oad-scale compa ison wi h i e
dis inguished g oups o clus e s (A–E) (*.pd )
Link: h ps://doi.o g/10.3897/VCS.119253.suppl7
Supplemen a y ma e ial 8
Planned applica ions o he Eu oVegChecklis Commi ee (EVCC) (*.pd )
Link: h ps://doi.o g/10.3897/VCS.119253.suppl8