2022, vol. 88, 16–36
https://doi.org/10.12657/denbio.088.002
Arkadiusz Nowak, Marcin Nobis, Sylwia Nowak, Marcin Kotowski,
Sebastian Świerszcz*
Phytosociological survey of juniper wood
vegetation in Tajikistan (Middle Asia)
Reveived: 7 March 2022; Accepted: 16 May 2022
Abstract
The paper presents the first syntaxonomic classification for juniper wood communities in Tajikistan with
some remarks on their environmental gradients. A total of 119 relevés were sampled between 2015 and
2021 using the seven-degree Braun-Blanquet cover-abundance scale. They were classified by a modified
TWINSPAN method. Diagnostic species were identified using phi coefficient as a fidelity measure. Detrended Correspondence Analysis (DCA) was used to determine relationships between samples, vegetation units
and major gradients in species composition. Plant communities have been divided into two main groups.
The first one is a west Irano-Turanian wood of Juniperus polycarpos var. seravschanica of the Pamir-Alai mountane belt. It occurs in two variants depending mainly on the aspect and precipitation. The second juniper
wood type in Tajikistan is a stand of Juniperus pseudosabina. It prefers the northern slopes and north-eastern
ranges of the Pamir-Alai and slightly lower elevations. Both juniper wood types are highly distinct in terms
of species composition, especially in the moss layer, and have therefore been assigned to different classes:
Pino-Juniperetea (Juniperetum seravschanicae) and Juniperetea pseudosabinae (Carici turkestanicae-Juniperetum pseudosabinae). The main factors determining the species composition of the studied associations are latitude,
growing season precipitation, annual range of air temperature and precipitation of coldest quarter. Our
study has shown that there are two very distinct vegetation types of the juniper wood groves in Tajikistan,
which reflect the main phytogeographical division between the provinces of Turkestan and Central Asia.
Keywords: Juniperetalia seravschanicae, Pamir-Alai Mts., Pino-Juniperetea, syntaxonomy, vegetation classification
Addresses: A. Nowak, M. Kotowski, S, Świerszcz,Botanical Garden, Center for Biological Diversity
Conservation, Polish Academy of Sciences; Prawdziwka 2, 02-973 Warszawa, Poland,
e-mail: s.swierszcz@obpan.pl;
A. Nowak, Department of Botany and Nature Protection, University of Warmia and Mazury in Olsztyn,
Łódzki Plac 1, 10-728, Olsztyn, Poland;
A. Nowak, S. Nowak, S. Świerszcz, Institute of Biology, University of Opole, Oleska 22, 45-052 Opole,
Poland;
M. Nobis, Department of Taxonomy, Phytogeography and Palaeobotany, Institute of Botany,
Jagiellonian University; Gronostajowa 3, 30-387 Kraków, Poland;
S. Świerszcz, The Franciszek Górski Institute of Plant Physiology, Polish Academy of Sciences;
Niezapominajek 21, 30-239 Kraków, Poland;
*Corresponding author
Phytosociological survey of juniper wood vegetation in Tajikistan (Middle Asia)
Introduction
Tajikistan is a mountainous country with an extremely diverse landscape, climate and habitat conditions. It is located in the central part of the Middle
Asia on the borderland of large phytogeographical
units (Central Asian, Turkestanian and Western
Himalayan). In its western part, the vegetation is
controlled by the subhumid Irano-Turanian or humid
alpine climate and consists of typical Irano-Turanian
species. The montane belt of these ranges offer suitable conditions for juniper wood vegetation, which
tolerates summer droughts and high climate continentality (Nowak et al., 2020a).
The juniper woods are one of the most important
and widespread type of forest in Tajikistan, being apparently distinct from mesophilous broad-leaved forests (chernolese), river carrs (belolese), termophilous open woods and scrubs (šhiblyak and pistachio
groves) and subtropical tugai gallery forests (Nowak
& Nobis, 2013; Nowak et al., 2015, 2022; Nowak
et al., 2017a; Świerszcz et al., 2022). According to
Zapryagaeva (1976), the dendroflora of Tajikistan
counts 268 species. Our analyses show that the native dendroflora of Tajikistan counts 102 tree species
and 211 shrub species. Our analyses show that the
forest habitats of Tajikistan are inhabited by ca. 800
native plant species and its dendroflora counts 102
tree and 211 shrub species. Juniper forests harbour
382 species, including 281 endemic and 171 subendemic taxa (Nowak et al., 2020b).
The natural lower treeline, which is visible in the
south-western Pamir-Alai, is about 500-700 m and
is marked by a line of open pistachio woods. However, due to centuries of pastoralism and deforestation
caused by mismanagement of timber resources, it is
now at an altitude of 1,200–1,500 m a.s.l. in most of
the western Tajikistan (Zapryagaeva, 1976). The upper forest limit is at an altitude of about 3,200–3,400
m a.s.l. (rarely 3,700 m a.s.l.) and is bounded by a
line of open juniper woods and azonal forests composed of various birch species. Further east, in southeastern Tibet, it is elevated much higher, to about
4,900 m a.s.l., and marked by juniper stands (Miehe
et al., 2007).
One of the most important synthetic studies on
the forests of Tajikistan is the work of Zapryagaeva
(1976). The author divides juniper woods into two
main types that are controlled by climate, i.e. thermophilous (with J. polycarpos var. seravschanica [= J.
seravschanica]) and cryophilous (with J. pseudosabina
[=J. turkestanica] and J. semiglobosa). Thermophilous juniper woods were reported from altitude of
(1,300)1,700–2,200(2,700) m a.s.l. in Pamir-Alai
with an average precipitation of 600 mm/year and
average annual temperature of 14 °C. This type of juniper stand occurs on chestnut-brown soils in areas
17
with a typically continental climate, where during the
vegetation season the daily amplitude can reach more
than 40 °C and at higher altitudes frost occur even in
mid-summer. In winter, the temperature often drops
to −25 °C. Zapryagaeva ambiguously describes the
dynamic relationship of thermophilus juniper woods
with other vegetation, suggesting a close relationship with mesophilous broad-leaved forests or thermophilous woodlands with Pistacia vera or Crataegus
spp. and Prunus spp. This type of thermophilous juniper groves is known to have a wide range reaching the Atlas Mountains in North Africa, the entire
Mediterrenian region and the Zagros, Alborz, Kopet-dagh, Kugitang and Hindukush Mountains in the
east (e.g. Zohary, 1973; Kamelin, 1979; Fet, 1994;
Popov, 1994; Tsiourlis et al., 2007; Douaihy et al.,
2013; Memariani et al., 2016b; Mucina et al., 2016).
However, it is worth noting that Middle Asian thermophilous juniper stands are distinguished by the
absence of Quercus and Arbutus species.
Following Zapryagaeva (1976), the second type of
juniper stands in Tajikistan occupies wetter and colder habitats in the upper montane belt at 2,200–3,400
m a.s.l., and consists mainly of Juniperus pseudosabina and J. semiglobosa. It is called “microtherm” – cryophilous juniper forest. This zone is characterized
by a high degree of continentalism and temperature
differences between winter and summer reaching up
to 65 °C, long-lasting snow cover (up to 8 months)
and deep freezing of the soil profile (up to 130 cm).
They occur in a mosaic of cryophilous steppes, alpine
tall-forbs and grassland. Significant species co-occurring in these woods are Sorbus tianschanica, Betula
tianshanica, Berberis integerrima, Lonicera simulatrix and
Astragalus tibetanus. These woods reaching the upper
subalpine belt in Middle Asia, are undoubtedly similar to the juniper forests of Tibet and the Himalayas in terms of climatic conditions, mainly built by J.
convallium (=J. mekongensis), J. indica, J. przewalskii, J.
recurva and J. tibetica (e.g. Ghimire et al., 2008; Miehe
et al., 2008; Tambe & Rawat, 2010).
Unlike Zapryagaeva (1976), Safarov (2018) and
Zakirov (1984) distinguished many more juniper forest types depending on the proportion of plant species
recruited from neighbouring vegetation types. These
divisions follow the work of Kamelin (1979) and
Ovchinnikov (1957), who classified juniper woods
into three main types depending on the humidity of
the climate and the phytogeographical origin of the
main floristic components of the communities (Middle Asian, ancient Mediterranean and Eastern Mediterranean). Safarov (2018) points out that juniper
forests in Tajikistan cover approx. 50% of the total
forested area and occur mainly in the mountains of
Zeravshan, Turkestan, Alaian, Darvazian, Hissar, Peter the First, Sarsarak, Vakhsh and Hazratishoh. He
includes in the group of cryophilous juniper forests
18
Arkadiusz Nowak et al.
two main types within Peucedrymion holarcticum – a
taiga-like group of sparse coniferous forests built by
species such as Larix spp., Pinus spp., Picea spp. This
group also includes forests of Picea schrenkiana and
Abies semenovii, found just north of the Pamir-Alai, in
Kyrgyzstan. The cryophilous juniper stands are presented in contrast to birch and poplar forests, which
are also found in the taiga zone, and in Middle Asia
most often form azonal riparian forests along mountain streams, reaching up to 3,800 m above sea level.
(Nowak et al., 2017a). The main formation is a stand
of Juniperus pseudosabina growing between 2,900 and
3,500 m a.s.l. Despite the main diagnostic tree, the
most frequent contributors are Thalictrum minus,
Phleum phleoides, Asyneuma argutum, Ephedra equisetina, Delphinium oreophillum, Dracocephalum bipinnatum,
Roegneria ugamica, Oxytropis lehmanni, Geranium regelii.
In the upper limit of this vegetation, Juniperus sibirica
(=J. communis var. saxatilis Pall.) also plays an important role, especially in central Tajikistan (Obighingou
valley, Rasht and Tavildara districts). The main subgroups were distinguished according to the proportion of tall-forb (Ligularia alpigena and Codonopsis clematidifolia) and grassland species share (Festuca sulcata,
F. valesiaca. Elymus alaicus).
In the second group, which is called Arceuthodrymion mediterraneum, Safarov (2018) lists a number of
thermophilous juniper wood types that occupy the
montane belt from 600–800 to 3,000 m a.s.l. Typical
components of these communities are Artemisia baldshuanica, Rosa kokanica, R. maracandica, Lonicera nummulariifolia, Crataegus pontica, Pyrus korshinskyi, Agropyrum
trichophorum, Poa bulbosa, Carex pachystylis, Artemisia
tenuisecta, Acer turkestanicum, A. regelii, Cotoneaster
hissaricus. This species composition indicates a close
relationship of thermophilous juniper forests with
mesophilous scrub (šhiblyak) and open pistachio
woods (Nowak et al., 2022; Świerszcz et al., 2022).
Recent research on juniper woods of Tajikistan
was conducted by Konnov (1973). He shares the position of earlier researchers on the division of juniper
forests. As the northernmost refuge of J. seravschanica woods, he indicates the Sary-Chelek region in the
Khatkal range in Kyrgyzstan. He points out that J.
semiglobosa is more common in the thermophilous
type of juniper woods and forms its own stands in
Central Tian Shan, Kyrgyzstan. In contrast, he distinguished stands of J. schugnanica in the Western Pamirs, which is now synonymised with J. semiglobosa.
These stands in the subalpine belt (up to 3,700 m) in
the Shugnan, Ishkashim and Shahdarian mountains
are dwarf shrubby vegetation with a J. semiglobosa
cover of about 20–30%.
In the western part of Middle Asia, in the Kopet-dagh Mts., open woodland of Juniperus turcomanica (=Juniperus polycarpos var. turcomanica) have been
recorded at 800–2,400 m altitude. This vegetation at
the upper limit is accompanied by scrubs of Juniperus
communis and J. sabina, which form carpet-like formations on exposed rocks in subalpine areas, especially
on the northern slopes of Aladagh and Ghorkhod Mts.
at altitudes of 1,600–2,600 m a.s.l. (Memariani et al.,
2016a). These woods have 30–40% cover in the tree
layer and extend mainly in territories of southwestern Kopet-dagh, the watershed plateaus between the
Sumbar and Arvaz rivers, the Kurydere and Kalymkhoz valleys and within the Syunt Khasardagh reserve.
They occupy well-humidified northern slopes at an
altitude of about 1,300 to 1,600 m a.s.l. (Fet, 1994).
This endemic to Turkmenistan juniper wood type
has also been observed at an altitude of 400 m a.s.l.,
where annual precipitation is about 200 mm and the
temperature rises to 48 °C in summer (Popov, 1994).
This is very close to the drought line in Juniperus forests in southern Tibet, where relict forests of J. convallium and J. tibetica were formerly common up to
4,800 m a.s.l. (Miehe et al., 2008).
Unfortunately, the area of these forests has been
steadily declining and has decreased by 50% between
1930 and 1960 alone. The same applies to forests
with J. seravschanica, in the Kugitang Mts. on the border with Uzbekistan and Afghanistan. Due to human impact, the lower limit of this juniper stand has
shifted from approx. 800 to as high as 1,300 to 1,700
m a.s.l. (Popov, 1994).
Along the southern edge of Middle Asia, two types
of juniper vegetation occur in the Afghan Hindukush.
The northern with J. excelsa, J. semiglobosa, J. seravschanica and the eastern with J. semiglobosa and J. seravschanica (Freitag, 1971). Additionally, the high-altitude
subalpine scrub built by J. squamata accompanied by J.
nana, Ribes alpestre and R. villosum is mentioned (Freitag, 1971). Additionally, one of the largest surviving
refugium of J. excelsa forests has been reported in the
Balochistan province of Pakistan. It covers an area of
approx. 141,000 hectares between altitudes of 2000–
3350 m a.s.l. (Sarangzai et al., 2012).
The aim of this study is to answer the following
questions: (i) which juniper wood communities can
be distinguished in Tajikistan and how should they
be arranged in the syntaxonomic system of Eurasia?
(ii) what are the compositional, ecological and chorological characteristics of the distinguished juniper
woods? (iii) how the distinguished syntaxa are related to others known from adjacent areas of Asia and
Europe?
Material and methods
Study area
The study area covered the northwestern phytogeographic regions of Tajikistan (Fig. 1): Zeravshanian
Phytosociological survey of juniper wood vegetation in Tajikistan (Middle Asia)
19
Fig. 1. Map showing study area and distribution of the relevés (a) assigned the particular vegetation associations (n =
110), and more detailed maps of relevés distribution in the Zeravshan and Turkestan Mts. (b), and Peter the First Mts.
(c). Note that due to the scale, some points may overlap. The exact locations of the relevés can be found in Table S1
20
Arkadiusz Nowak et al.
B, Zeravshanian C, Turkestanian A and East Tajikistanian A. The plots in Zeravshanian regions were
located along the river valleys of Chapdara, Iskanderkul (west of Lake Iskanderkul), Mura and Nofin (east
of Lake Nofin). In the Turkestanian region, research
was conducted in the Archazor forest along the Aktengi river valley and the Byurogan river valley, and
then further south along the Altykul and the Zaravshan river valley with its two tributaries – Pakhurd
and Rarz. In the eastern part of Tajikistan, the relevés
were located along the Surkhob river valley. The
studied sites were located on hillsides with different
aspects, slopes and altitude, and only ranker soil was
a stable factor. The vegetation plots were located between 1,803 and 3,100 m a.s.l. The annual precipitacion in studied area varies from 405 to 1,246 mm,
and mean annual temperature varies from -1.05 to
6.75 °C.
The Zeravshan region is a typical mountainous
area, located between 850 (Zeravshan river valley
on the border with Uzbekistan) and 5,489 m a.s.l.
(Chimtarga peak). The main mountain subranges
within the Zeravshan Mountains are Dukdon, Fann,
Zinach, Kugibodrawak and Ljangar. Due to its relief and altitude, the alpine character of the climate
strongly influences the area. Along the Zeravshanian
and Turkestanian ranges, in the outer Pamir, Cambrian and Silurian sediments dominate. The rocks
are generally limestone, dolomite, marble, clay shale,
phyllitic shale, dolomite and argillaceous shale. In
general, Tajikistan experiences high insolation, as
well as low cloud cover, high annual temperature amplitudes, low humidity and precipitation. The most
extensive and dense stands of seravshan juniper are
found in the Zeravshan, Turkestan, Hissar and Darvaz ranges in the western Pamir-Alai at an altitude of
about 1,000–2,500 m a.s.l.
Taxonomic treatment
The taxonomy of the genus Juniperus is very complex and, despite decades of research, still vague.
For example, J. seravschanica as well as J. turcomanica
are merged with J. excelsa subsp. polycarpos (Farjon,
1992). Adams (2016), after analysis of nrDNA and
cpDNA (petN-psbM, trnS-trnG, trnD-trnT, trnLtrnF), includes J. seravschanica in J. excelsa var. seravschanica. However, the same author previously includes J. seravschanica in J. polycarpos var. seravschanica
(Adams, 2001, 2004) and this approach is adopted
in the Plant List (The Plant List, 2020). Some authors have synonymised J. seravschanica with J. excelsa
(e.g. Mazur et al., 2004). Juniperus turkestanica, after
examining its relationship to J. centrasiatica, J. pseudosabina and J. indica was synonymised with J. pseudosabina (Adams & Turuspekov, 1998). Due to all these
inconsistencies, we decided to follow the Plant List
(2020) as the basic source for taxonomic resolution
in this study. Therefore, we treat J. seravschanica as a
variety of J. polycarpos whereas J. turkestanica as conspecific with J. pseudosabina. In this approach, data on
Juniperus seravschanica Kom., Juniperus excelsa subsp.
seravschanica (Kom.) R. Kam. ex Imch., Juniperus kulsaica V.D. Dmitriev, Juniperus polysperma V.D. Dmitriev,
Juniperus zaaminica V.D. Dmitriev and Sabina seravschanica (Kom.) Nevski are treated as Juniperus polycarpos var. seravschanica (Kom.) Kitam. Data on Sabina
pseudosabina var. turkestanica (Kom.) C.Y. Yang and Juniperus turkestanica Kom. are treated as synonyms of
J. pseudosabina.
Data sampling and data analyses
The phytosociological survey was conducted from
2015 to 2021. In total, 119 relevés were collected in
all types of naturally occurring juniper forest phytocoenoses growing in the country. Plant material collected during the field studies is preserved at OPUN
(Opole University, Poland) and KRA (Jagiellonian
University, Poland). The vegetation plot size was delimited to 100 m² in such way to enable providing
homogeneity in terms of structure, species composition and habitat conditions of the phytocoenosis following the Braun-Blanquet approach (Dengler et al.,
2008), globally used geobotanical method (Nowak &
Nowak, 2022). For each vegetation plot all species
of vascular plants and terricolous bryophytes were
recorded with the use of 7-degree cover-abundance
scale (r, +, 1, 2, 3, 4, 5; Braun-Blanquet, 1964). Species were recorded in four layers of the wood or scrub
stands: a2 − lower tree layer, b − shrubs, c − herbs,
d − bryophytes. Geographical coordinates, elevation,
aspect and slope inclination were recorded for each
relevé. Geographical coordinates of plots were obtained using a GPS device with an accuracy of ± 10
m and the WGS-84 grid reference system.
The relevés were stored in TURBOVEG format
(Hennekens & Schaminée, 2001) in the Vegetation of Middle Asia Database (GIVD ID AS-00-003;
Nowak et al., 2017b). They were analysed in the
JUICE software (Tichý, 2002). The Braun-Blanquet
scale was transformed by JUICE to percentage values
as follows: + = 2%, 1 = 3%, 2 = 13%, 3 = 38%, 4 =
68% and 5 = 88%. A modified TWlNSPAN analysis
(Roleček et al., 2009) was performed in order to classify the relevés by using cutoff levels of 0%, 2%, 5%
and 25%. Total inertia was used as a measure of cluster heterogeneity (Roleček et al., 2009). Plant species determined only to the genus level were omitted
before the analysis. Diagnostic species were identified using the phi coefficient (Chytrý et al., 2002)
and cover ratio (Willner, 2006) as a fidelity measure.
The size of all groups was standardised to equal size,
and the Fisher’s exact test (p < 0.05) was applied in
Phytosociological survey of juniper wood vegetation in Tajikistan (Middle Asia)
21
Table 1. Synoptic table of juniper wood vegetation in Tajikistan. The phi coefficient values (in superscript) in the table
are multiplied by 100. All diagnostic species and other plants of frequency ≥ 20% are shown. Main values are species
frequencies (in percentages). Abbreviations in layer column: a2 – lower tree layer, b – shrub layer, c – herb layer. Group
No.: 1 – Ass. Juniperetum seravschanicae variant with Hedysarum denticulatum and Oxytropis glabra, 2 – Ass. Juniperetum seravschanicae variant with Primula lactiflora and Brachythecium collinum, 3 – Ass. Carici turkestanicae-Juniperetum pseudosabinae
Cluster No.
3
Layer
No. of relevés
30
Ass. Juniperetum seravschanicae and All. Juniperion seravschanicae
a2
100
31
Juniperus polycarpos var. seravschanica
100
32
b
Juniperus polycarpos var. seravschanica
50
67
c
Oxytropis glabra
57
44
b
Berberis integerrima
47
44
c
Pedicularis dolichorrhiza
17
37
c
Hedysarum denticulatum
50
32
c
Seseli schrenkianum
40
29
c
Silene tachtensis
d
.
–
Brachythecium collinum
c
30
–
Eremogone griffithii
c
.
–
Primula lactiflora
c
50
–
Poa nemoraliformis
Ass. Carici turkestanicae-Juniperetum pseudosabinae and All. Juniperion pseudosabinae
a2
.
–
Juniperus pseudosabina
b
.
–
Juniperus pseudosabina
c
.
–
Juniperus pseudosabina
c
.
–
Oxytropis ovczinnikovii
c
3
–
Gentianella turkestanorum
c
.
–
Cerastium dichotomum subsp. inflatum
c
.
–
Codonopsis clematidea
c
.
–
Erigeron seravschanicus
c
.
–
Phleum phleoides
c
63
–
Carex turkestanica
c
.
–
Helictotrichon hookeri
d
.
–
Sanionia uncinata
c
7
–
Thalictrum kuhistanicum
d
83
–
Hypnum cupressiforme
Others
27
48
c
Allium weschniakowii
c
10
–
Artemisia dracunculus
c
3
–
Artemisia santolinifolia
c
.
–
Astragalus aksuensis
c
43
–
Astragalus tibetanus
c
37
–
Asyneuma argutum subsp. argutum
c
7
–
Aulacospermum roseum
c
30
–
Berberis integerrima
23
17
d
Bryum caespiticium
c
23
–
Campanula glomerata
33
49
c
Carduus nutans
c
.
–
Cicerbita zeravschanica
37
42
b
Cotoneaster zeravschanicus
c
37
–
Crepis pulchra
c
.
–
Cystopteris fragilis
c
30
–
Draba nemorosa
c
3
–
Draba yunussovii
c
10
–
Ephedra intermedia
c
.
–
Erigeron petroiketes
20
41
c
Erigeron pseudoseravschanicus
c
20
–
Euphrasia pectinata
b
60
–
Festuca rupicola
4
23
5
57
100
100
.
26
26
.
65
43
74
87
57
91
31
32
–
–
–
–
50
34
83
73
71
60
.
.
.
21
4
.
.
5
.
5
.
30
–
–
–
–
–
–
–
–
–
–
–
–
.
.
.
.
9
.
.
.
22
9
13
61
.
61
–
–
–
–
–
–
–
–
–
–
–
–
–
–
100
88
100
63
72
49
39
39
49
82
39
96
30
96
100
.
48
30
30
30
39
30
57
13
30
4
22
17
35
4
74
30
26
.
.
4
30
–
23
44
51
–
–
–
25
–
–
–
38
–
–
–
52
47
35
–
–
–
–
.
.
4
.
54
47
33
39
.
84
.
4
.
58
40
33
.
2
21
.
37
81
–
–
–
–
92
100
76
74
66
58
58
50
49
46
44
43
41
27
–
28
–
–
63
–
–
–
29
38
–
–
–
42
–
38
44
22
Cluster No.
No. of relevés
Galium pamiroalaicum
Galium spurium
Gentiana olivieri
Geranium regelii
Hedysarum flavescens
Hieracium robustum
Juniperus polycarpos var. seravschanica
Juniperus semiglobosa
Koeleria pyramidata
Lactuca soongorica
Ligularia thomsonii
Lonicera nummulariifolia
Lonicera stenantha
Myosotis refracta
Myosotis laxa subsp. caespitosa
Nepeta podostachys
Oxytropis capusii
Oxytropis lehmanni
Pedicularis krylowii
Petrorhagia alpina
Poa bulbosa
Poa fragilis
Poa pratensis
Poa trivialis
Polygonum coriarium
Potentilla mollissima
Psychrogeton pseuderigeron
Ribes meyeri
Rosa kokanica
Rosa kokanica
Rosa webbiana
Syntrichia ruraliformis
Syntrichia ruralis
Taraxacum agg.
Thalictrum minus subsp. maxwellii
Thalictrum sultanabadense
Thymus seravschanicus
Veronica rubrifolia
Arkadiusz Nowak et al.
3
30
Layer
c
c
c
c
c
c
c
b
c
c
c
b
b
c
c
c
c
c
c
c
c
c
c
c
c
c
c
b
b
c
b
d
d
c
c
c
c
c
order to exclude species with non-significant occurrence optimum in a particular cluster. Species with a
phi coefficient higher than 0.30 and cover ratio higher than 2 were considered diagnostic for a specific
cluster, except taxa considered diagnostic for various
and different vegetation types in Middle Asia on the
basis of our expert knowledge. We define alliances
by those species that have a phi coefficient ≥ 0.30 in
at least two clusters within the alliance. Species with
frequency higher than 30% in a particular cluster
were defined as constant species. To show compositional differences between distinguished thermophilous open woodlands and scrub units, Detrended Correspondence Analyses (DCA) was computed
using the ‘vegan’ package version 2.5.4 (Oksanen et
al., 2019) in R version 4.0.5 (R Core Team, 2020).
Species cover data were log-transformed (log(x+1))
27
30
23
77
30
17
20
.
33
10
97
87
17
27
3
33
13
.
27
10
40
23
43
33
37
23
.
3
50
7
60
20
57
43
30
3
10
27
4
23
–
30
4
–
47
–
–
–
–
–
–
8
–
38
–
–
–
–
–
–
13
–
–
11
–
44
–
–
23
–
43
41
–
–
30
–
–
–
43
.
.
87
.
52
35
26
43
26
96
70
22
9
17
52
22
22
17
43
13
17
30
30
70
.
43
22
26
13
35
.
83
22
9
52
52
26
5
57
–
–
–
–
–
54
26
24
–
37
–
–
–
–
–
25
29
–
–
31
–
–
–
–
32
–
51
38
–
–
–
–
16
–
–
34
–
–
37
.
4
93
4
4
.
9
35
.
95
65
7
.
37
.
.
21
18
23
11
74
44
5
2
.
12
.
5
25
23
.
40
56
18
2
68
4
–
–
–
31
–
–
–
–
–
–
–
–
–
–
40
–
–
22
–
–
–
41
–
–
–
–
–
–
–
28
–
–
–
24
–
–
36
–
with down-weighting of rare taxa. For ecological
interpretation of the ordination axes, environmental parameters were plotted onto a DCA ordination
diagram as supplementary variables using the envfit
function of the ‘vegan’ package. Environmental parameters include altitude, inclination, latitude, longitude, and bioclimatic variables (bio1 – mean annual
air temperature, bio7 – annual range of air temperature, bio12 – annual precipitation amount, bio18 –
mean monthly precipitation amount of the warmest
quarter, bio19 – mean monthly precipitation amount
of the coldest quarter and gsp – accumulated precipiation amount on growing season days). We also
calculated differences in environmental factors and
vegetation variables (cover tree, shrub, herb and
moss layer, overall species richness and phytogeographical elements richness) between syntaxonomic
Phytosociological survey of juniper wood vegetation in Tajikistan (Middle Asia)
units using the Kruskal–Wallis rank sum test (function kruskal.test) with multiple comparison based on
Dunn’s test using the dunnTest function in the ‘FSA’
package (Ogle et al., 2018) in R. Climatic data were
extracted from the CHELSA database version 2.1
(http://chelsa-climate.org; Karger et al., 2017).
A synoptic table with the fidelity and relative percentage frequency of all diagnostic species and other
plants of frequency ≥ 20% are shown in Table 1. Analytic table including type relevés and full synoptic
table are given in Supplementary material (Table S1
and Table S2 respectively). Newly presented syntaxa
are proposed according to the ICPN (Theurillat et
al., 2021). All mentioned syntaxa are arranged into a
syntaxonomic overview at the end of the description
in the Results section. The association concept follows Willner (2006) and the alliance follows Willner
(2020). The nomenclature of the vascular plants follows generally Cherepanov (1995) and The Plant List
(2020) Version 1.1. (http://www.theplantlist.org/),
and bryophytes follows Ignatov et al. (2006).
Results
General floristic and physiognomic
features
In our data set for Tajikistan (119 relevés), the juniper woodlands harbour 308 vascular plant species
and 13 mosses. In total, 65 species reached constancy
above 10% and 52 taxa above 20%. The open structure of juniper stands allows settlements and gaining
high frequencies of species typical for neigbouring
vegetation like alpine tall-forbs (e.g. Ligularia thomsonii, Geranium regelii, Polygonum coriarium), cryophilous
steppes (e.g. Festuca rupicola, Koeleria pyramidata, Eremogone griffithii) or deciduous forests and shrubs (e.g.
Astragalus tibetanus, Poa pratensis). Among the typical
woodland taxa, the most frequent were: Lonicera
nummulariifolia (73%), Campanula glomerata (52%),
Juniperus polycarpos var. seravschanica (50.9%), J. pseudosabina (20,3%), Poa fragilis (47%), P. nemoraliformis
(44%), Asyneuma argutum subsp. argutum (44%), Berberis integerrima (38%), Draba nemorosa (37%) and
Seseli schrenkianum (25%). Important components of
Tajik cryophilous juniper stands are species typical
for mires, such as Gentianella turkestanorum (37%)
or Codonopsis clematidea (18.5%). Among trees, Juniperus polycarpos var. seravschanica (50.4%) and Juniperus pseudosabina (48%) were the most frequent. The
most constant species in shrub layer were Lonicera
nummulariifolia (73%), Juniperus polycarpos var. seravschanica (50.4%), J. pseudosabina (42%), Rosa webbiana (32.7%), Berberis integerrima (29.4%), R. kokanica
(22.7%), Cotoneaster zeravschanicus (12.6%), L. stenantha (11.7%) and J. semiglobosa (10.1%). Unlike many
23
other vegetation types in the subarid zones of Middle
Asia, juniper woods are relatively abundant and rich
in moss species (Nowak et al., 2022). The most common bryophyte species were: Hypnum cupressiforme
(80.6%), Sanionia uncinata (62.2%), Syntrichia ruralis
(55.5%), Brachythecium collinum (14.3%) and Bryum
caespiticium (9.2%).
Numerical classification and DCA
ordination
The TWINSPAN classification revealed five clusters (Fig. 2), three of them are interpretable by a
set of diagnostic species. On the left side of the diagram, there are two small groups corresponding to
plots dominated by the zeravshan juniper. Cluster 1
corresponds to the plots sampled in the Sary-Khosor
National Park, which differ in the abundance of tallforb species. Cluster 2 encompasses a highly grazed
plots with a high proportion of pasture and steppe
species and low cover in tree and herb layers. These
communities were surveyed in the intensively grazed
Half-kul Valley. Since clusters 1 and 2 consists only of
4 and 5 relevés respectively, we decided to leave them
rankless. Looking further from the left to the right of
the figure (Fig. 2), two fairly large groups of the plots
with Juniperus polycarpos var. seravschanica can be distinguished. First (cluster 3) encompasses Zeravshan
juniper stands that are located on more isolated,
warmer slopes with dominant western, southern and
rarely north-eastern exposition. The right one (cluster 4), located in the central part of the graphs, groups
stands of J. polycarpos var. seravschanica which grow
mostly on northern and north-western, steep slopes
with relatively abundant moss layer. The most homogeneous group corresponds to the Juniperus pseudosabina community (cluster 5). Despite the large number
of plots with this species, they are clearly distinct and
well separated from the other communities at a high
level of dissimilarity. It suggests a very diverse species composition and the possibility of distinction at
a high level in the hierarchical arrangement.
Vegetation groups defined in the TWINSPAN
classification are clearly separated by the two first
axes of DCA analysis (Fig. 3). The first axis of the
DCA reflects a strong latitudinal, growing season
precipitation, annual temperature range and precipitation of the coldest quarter gradients, which differentiates clusters 3–4 and 5. Plots classified to the
association Carici turkestanicae-Juniperetum pseudosabinae occurs at higher latitudes with more continental climate (higher annual temperature ranges) and
are associated with higher precipitation amount in
the growing season (Fig. 3 and 4). In contrast, Juniperetum seravschanicae were found in localities of
lower latitudes and higher precipitation in winter.
24
Arkadiusz Nowak et al.
The second axis shows the relation to mean annual
temperature gradient and the precipitation amount
of the warmest quarter differentiating clusters 3 and
4 (variants of Juniperetum seravschanicae). Variant with
Hedysarum denticulatum and Oxytropis glabra was found
in an area with the higher mean annual temperature,
and higher sum of annual precipitation, as well as
lower precipitation amount in the warmer quarter, in
comparison to the variant with Primula lactiflora and
Brachythecium collinum (Figs 3 and 4).
Fig. 2. Dendrogram illustrating the assigment of relevé groups identified by TWINSPAN to particular syntaxonomic units:
1 and 2 – communities left rankless, 3 – Ass. Juniperetum seravschanicae variant with Hedysarum denticulatum and Oxytropis glabra, 4 – Ass. Juniperetum seravschanicae variant with Primula lactiflora and Brachythecium collinum, 5 – Ass. Carici
turkestanicae-Juniperetum pseudosabinae
Fig. 3. Detrended Correspondence Analysis diagram of juniper woods vegetation in Tajikistan. (a) Spider plots with vegetation units centroids are plotted. The second diagram (b) shows the same ordination with environmental variables
passively plotted onto DCA diagram. The eigenvalues and lengths of gradients were 0.40, 2.66 (Axis 1) and 0.16, 2.02
(Axis 2), respectively. Abbreviations: Bio1 – mean annual air temperature (°C), Bio7 – annual range of air temperature
(°C), Bio12 – annual precipitation amoun (mm), Bio18 – mean monthly precipitation amount of the warmest quarter (mm), Bio19 – mean monthly precipitation amount of the coldest quarter (mm), gsp – accumulated precipiation
amount on growing season (mm), Slope – inclination of the slope (°)
Phytosociological survey of juniper wood vegetation in Tajikistan (Middle Asia)
25
Fig. 4. Boxplots showing median (line), quartiles, outliers and the range of (a) altitude, (b) inclination of the slope, (c)
latitude, (d) longitude, (e) accumulated precipiation amount in growing season (mm), (f) mean annual temperature
(°C), (g) annual range of air temperature (°C), (h) sum of annual precipitation (mm), (i) mean monthly precipitation
amount of the warmest quarter (mm) and (j) mean monthly precipitation amount of the coldest quarter (mm) for
particular syntaxonomic units. The letters represent homogeneous groups according to Dunn’s post hoc test following
a significant Kruskal–Wallis rank sum test (p < 0.05), and labels on the x-axis correspond to cluster numbers
Synopsis of syntaxa
Based on the analyses, we propose the following
classification for the juniper woods in Tajikistan:
Thermophilous Zeravshan juniper stands in the lower montane belt of Middle Asia
Class: Pino-Juniperetea Rivas-Mart. 1965 (?)
Order: Juniperetalia seravschanicae nom. prov.
I. Alliance: Juniperion seravschanicae Nowak et al.
2022 all. nova hoc loco
I.1 Association: Juniperetum seravschanicae
Nowak et al. 2022 ass. nova hoc loco
– variant with Hedysarum denticulatum and Oxytropis glabra
(group 1)
– variant with Primula lactiflora and Brachythecium collinum
(group 2)
Cryophilous juniper stands in the upper montane
belt of Middle and Central Asia (and probably also
Himalayas)
Class: Juniperetea pseudosabinae Mirkin et al. 1986
26
Arkadiusz Nowak et al.
Order: Juniperetalia pseudosabinae Mirkin et al.
1986
II. Alliance: Juniperion pseudosabinae Mirkin et al.
1986
II.1. Association: Carici turkestanicae-Juniperetum pseudosabinae Nowak et al. 2022 ass. nova
hoc loco (group 3)
Description of syntaxa of the
thermophilous open woodland and
scrub vegetation in Tajikistan
Group I. Thermophilous Zeravshan
juniper stands in the lower montane
belt of Middle Asia
Order: Juniperetalia seravschanicae nom. prov.
Nomenclatural type: Juniperion seravschanicae all.
nov. hoc loco
Diagnostic species: Berberis integerrima, Campanula
glomerata, Juniperus polycarpos var. seravschanica, Oxytropis capusii, Nepeta podostachys, Thalictrum sultanbadense, Pedicularis dolichorhiza, P. krylovii, Poa fragilis,
P. nemoraliformis, P. trivialis, Seseli schrenkianum, Silene
tachtensis, Veronica rubrifolia.
Constant species: Asyneuma argutum subsp. argutum, Geranium regelii, Lonicera nummulariifolia, Ligularia
thompsonii, Poa pratensis, Rosa webbiana, R. kokanica.
Geographical range: The phytocoenoses assigned to
this order occur across western parts of Middle Asia
(western Tian Shan, Pamir-Alai, western Hindukush,
and probably also in Kopet-dagh (Turkmenistan),
Baluchistan (Pakistan), Hazarmaysh Mts. (Afghanistan) and Kuyhitang Mts. (Uzbekistan).
Habitat characteristics: Main vertical range in the
montane belt between 1,500 and 2,700 m a.s.l., occasionally descending to lower altitudes, especially towards the south-west. The communities of the new
order develop on the slopes of the mountain ranges
of intermediate altitude, preferring fairly steep slopes
of various aspects. They develop on chestnut-brown
soils and are adapted to the winter-rain mediterranean-like climate of considerable continentality. Juniperetalia seravschanicae thermophilous juniper woods
are used as a source of firewood and are extensively
grazed, mainly by sheep and goats.
Remarks: The order includes phytocoenoses dominated by the Zeravshan juniper Juniperus polycarpos
var. seravschanica. It should probably also include other thermophilous juniper phytocoenoses from the
entire Irano-Turanian region, at least from the Turkestanian province, such as Juniperus polycarpos var.
turcomanica from the Kopet-dagh Mts. in Turkmenistan and Iran. This vegetation has an open structure
with an average tree canopy cover of ca. 50% and is
composed mainly by Irano-Turanian species. Since
the naming taxon is subjected to different taxonomic
treatments, we decided to take as the name of the
order its local name, which is well-rooted in the geobotanical literature and accepted in The Plant List as
a variety name.
I. Alliance: Juniperion seravschanicae all. nov. hoc
loco
Nomenclatural type: Juniperetum seravschanicae
ass. nov. hoc loco
Diagnostic species: at the current stage of the survey the same as for the order.
Constant species: at the current stage of the survey
the same as for the order.
Geographical range: The phytocoenoses assigned
to the alliance occur across western parts of Middle
Asia (western Tian Shan, Pamir-Alai, western Hindukush, and probably also in Kopet-dagh (Turkmenistan), Baluchistan (Pakistan), Hazarmaysh Mts.
(Afghanistan) and Kuyhitang Mts. (Uzbekistan). The
largest forest complexes of Zeravshan juniper extend
in Zeravschan, Hissar, Babatag, Vakhsh, Sarsarak,
Hazratishoh, Kuraminian in Tajikistan, Hindukush
in Afghanistan and the western Tian Shan ranges in
Kyrgyzstan.
Habitat characteristics: Patches of this community occur in the montane belt of the western part of
Middle Asia, mainly Tajikistan and Kyrgyzstan. Thermophilous juniper woods grow in Pamir-Alai at altitudes of (1,300)1,700–2,200(2,700) m a.s.l. in areas
with an average precipitation of 500–700 mm/year
and average annual temperature of about 12–15°C. In
winter the temperature often falls well below -25°C.
The stands inhabit shallow to moderately deep soils
(sometimes also on screes) with a strongly dried topsoil in the hot summer period. This vegetation type
develops on slopes of different inclinations and aspect
and is not strongly associated with northern exposure, such as Juniperus pseudosabina stands. The plant
communities included in this alliance consist mainly
of species of the Irano-Turanian distributional range.
I.1. Juniperetum seravschanicae ass. nov. hoc loco
Diagnostic species: Berberis integerrima, Brachythecium collinum, Eremogone griffithii, Hedysarum denticulatum, Juniperus polycarpos var. seravschanica, Oxytropis
glabra, Pedicularis dolichorrhiza, Poa nemoraliformis,
Primula lactiflora, Seseli schrenkianum, Silene tachtensis.
Constant species: Hypnum cupressiforme, Juniperus
polycarpos var. seravschanica, Lonicera nummulariifolia;
Geranium regelii, Ligularia thomsonii.
Geographical range: Plots of this community were
sampled in Zeravshan Mts., mainly in Pastrud-daria,
Imat and Iskander-daria River Valleys (Fonn Mts; Fig.
1). It is the core distributional area of the main diagnostic taxon and its locus classicus.
Phytosociological survey of juniper wood vegetation in Tajikistan (Middle Asia)
Floristic composition: The tree and shrub layer is
clearly dominated by Juniperus polycarpos var. seravschanica. Additionally, Berberis integerrima, Lonicera nummulariifolia, L. stenantha, Cotoneaster seravschanicus, C.
oliganthus, Rosa kokanica, R. webbiana, Ribes meyeri and
Juniperus semiglobosa also have a significant share in
the shrub layer (mean cover 33%; Fig. 5b). The herbaceous layer is quite scarce, reaches 30–80% (mean
51%; Fig. 5c) and consists of 18–43 taxa (mean 28;
Fig. 5e), with Carex turkestanica, Hedysarum denticulatum, Ligularia thomsonii and Poa trivialis being the
most abundant (Fig. 7b). Mosses are quite abundant
in this wood having from 1 to 30% coverage with the
mean of about 8% (Fig. 5d).
Habitat characteristics: The community inhabits
relatively dry, often stony slopes in the montane elevations (1,800–2,800 m a.s.l., mean ca. 2,500; Fig.
4a). Plots of the community were sampled within
large juniper complexes on slopes with inclination
up to 40° and only occasionally on flat land (Fig. 4b,
27
7a). Aspects were variable with predominantly western and northern exposures (Fig. 6).
Remarks: The TWINSPAN classification showed the
existence of two clusters that differ insignificantly
with the floristic composition of the plots.
Holotypus: (sequence number in Table S1 – 40) 8
July 2021; 68.36853 °E; 39.22339 °N; 2517m a.s.l.;
aspect N; slope 25°; plot area 100 m2; cover tree layer
60%; cover shrub layer 50%; cover herb layer 30%;
cover moss layer 5%; species composition:
Tree layer: Juniperus polycarpos var. seravschanica 4.
Shrub layer: Juniperus polycarpos var. seravschanica 2,
Lonicera nummulariifolia 2, Berberis integerrima 1, Sorbus
persica 1, Ribes meyeri +, Rosa webbiana +. Herb layer:
Geranium regelii 1, Ligularia thomsonii 1, Polygonum coriarium 1, Primula lactiflora 1, Artemisia dracunculus +,
Artemisia santolinifolia +, Astragalus aksuensis +, Draba
nemorosa +, Ephedra intermedia +, Eremogone griffithii +, Leonurus turkestanicus +, Pedicularis krylowii +,
Poa nemoraliformis +, Poa trivialis +, Ribes meyeri +,
Fig. 5. Boxplots showing median (line), quartiles, outliers and the range of (a) cover of tree layer, (b) cover of shrub layer,
(c) cover of herb layer, (d) cover of moss layer, (e) species richness, (f) species richness of Irano-Turanian element
and (g) species richness of Eastern Irano-Turanian with Euro-Siberian element for particular syntaxonomic units. The
letters represent homogeneous groups according to Dunn’s post hoc test following a significant Kruskal–Wallis rank
sum test (p < 0.05), and labels on the x-axis correspond to cluster numbers
28
Arkadiusz Nowak et al.
Fig. 6. The exposition preferences of the researched plant communities
Seseli schrenkianum +, Silene tachtensis +, Thalictrum
sultanabadense +. Moss layer: Hypnum cupressiforme 1,
Brachythecium collinum +, Syntrichia ruralis +.
– variant with Hedysarum denticulatum and
Oxytropis glabra
This variant is distinguished by slightly higher precipitation (especially in the winter), lower slope inclination (wide glacial valleys) and more diverse aspects, also to the south and west (Figs 4 and 5). The
herb layer is a bit more abundant here, in contrast to
the poorer moss layer (Fig. 5d). There are a number
of species typical for more fertile habitats resistant to
grazing, e.g. Carex turkestanica, Gentiana olivieri, Adonis
turkestanica, Carduus nutans, Erigeron pseudoseravschanicus, Potentilla molissima, Thalictrum minus subsp. maxwellii or Pedicularis krylowii. Plots of this community
were found mainly in the valleys of the Kara-kul and
Iskander-daria rivers (Fig. 1).
– variant with Primula lactiflora and Brachythecium
collinum
The variant with Primula lactiflora and Brachythecium
collinum (Fig. 7c) differs from the previous one by significantly more abundant moss layer, which can cover
up to 30% of the area (Fig. 5d). Patches of this community develop on steeper slopes which usually have
northern exposure (Fig. 6). Grazing is less intensive
here and the soil is less fertile. In the undergrowth
the plants typical for this variant are Astragalus aksuensis, Phleum phleoides, Oxytropis lehmannii, Aulacospermum roseum, Ribes meyeri, Cicerbita seravschanica, Psychrogeton pseuderigeron, Artemisia santoliniifolia, Draba
junusovii, Oxytropis capusii.
In the group of thermophilous juniper woods, the
TWINSPAN algorithm distinguished two clusters
of few plots (nine in total). These are the communities from the Sary-Khosor area with a large share
of tall-forb species and from the Half-kul valley with
intensive goat and sheep grazing. For these reasons,
these plots differ strongly from the other groups and
thus we decided to leave it rankless without further
consideration.
Cryophilous juniper stands in the upper
montane belt of Middle and Central Asia
II. Alliance: Juniperion pseudosabinae Mirkin et al.
1986
Diagnostic species: Carex turkestanica, Cerastium
dichotomum subsp. inflatum, Codonopsis clematidea,
Erigeron seravschanicus, Gentianella turkestanorum, Helictotrichon hookeri, Hypnum cupressiforme, Juniperus
pseudosabina, Oxytropis ovczinnikovii, Phleum phleoides,
Sanionia uncinata, Thalictrum kuhistanicum.
Constant species: Carex turkestanica, Geranium regelii,
Hypnum cupressiforme, Juniperus pseudosabina, Ligularia
thomsonii, Lonicera nummulariifolia, Sanionia uncinata,
Thymus seravschanicus.
Geographical range: The vegetation of this alliance
extends from the northern Pamir-Alai, through a
large territory of the western and central Tian Shan,
Altai, to the Sayan Mts. In Tajikistan, this Juniperus
pseudosabina is distributed in the upper montane belt
mainly in Turkestan, Peter the First, Kuraminian and
Zeravshan Mts. at elevations between 2,000 and
3,600 m a.s.l., but the highest outposts of the community resambles rather subalpine krummholz with
patchy shrub vegetation (Fig. 7e).
Floristic composition: The Juniperion pseudosabinae
forms dense monospecies stands (Fig. 7d) with up to
85% cover of the tree layer (mean ca. 50%; Fig. 5a).
The shrub layer is also abundant (mean ca. 40%, Fig.
5b), and the most frequent species are Lonicera nummulariifolia, L. korolkowii, Rosa webbiana and Berberis
integerrima. The plots are clearly dominated by Juniperus pseudosabina in the canopy and Carex turkestanica, Ligularia thomsonii, Poa nemoralis, Ranunculus aureopetalus, Thalictrum kuhistanicum, Thymus seravschanicus,
Hypnum cupressiforme, Sanionia uncinata in herb and
moss layers. Plots of this vegetation are moderately
rich in species but have fairly abundant moss cover.
Habitat characteristics: The vegetation with the domination of Juniperus pseudosabina occurs in moderately
warm and cold regions of Tajikistan. It develops in
the upper montane to subalpine belts on mainly on
northern slopes.
Phytosociological survey of juniper wood vegetation in Tajikistan (Middle Asia)
29
Fig. 7. Photographs of the juniper wood vegetation in Tajikistan: a) Juniperetum seravschanicae in the area of Alaudin
lakes, b) Juniperetum seravschanicae with Ligularia thompsonii in the herb layer, Iskander-daria River Valley, c) Juniperetum
seravschanicae variant with Primula lactiflora and Brachythecium collinum in Pastrud-daria River Valley, d) stands of the
Carici turkestanicae-Juniperetum pseudosabinae near Buragen, e) patches of the Juniperion pseudosabinae vegetation at upper
treeline near Koshtegirmen, f) high bryophytes abundance in the Carici turkestanicae-Juniperetum pseudosabinae stands,
Buragen (pictures a, d, e were taken by A. Nowak and b, c, f by S. Świerszcz)
II.1. Association: Carici turkestanicae-Juniperetum
pseudosabinae ass. nov. hoc loco
Diagnostic species: Carex turkestanica, Cerastium
dichotomum subsp. inflatum, Codonopsis clematidea,
Erigeron seravschanicus, Gentianella turkestanorum, Helictotrichon hookeri, Hypnum cupressiforme, Juniperus
pseudosabina, Oxytropis ovczinnikovii, Phleum phleoides,
Sanionia uncinata, Thalictrum kuhistanicum.
Constant species: Carex turkestanica, Geranium regelii,
Hypnum cupressiforme, Juniperus pseudosabina, Ligularia
thomsonii, Lonicera nummulariifolia, Sanionia uncinata,
Thymus seravschanicus.
Geographical range: The Juniperus sabina vegetation
was surveyed mainly in the Ohtangi Valley and in the
vicinity of Shahristan Pass in the Turkestan Mountains (Fig. 1). It occupies the upper montane belt
between 2,000 and 3,100 m a.s.l. (mean ca. 2,500
30
Arkadiusz Nowak et al.
m), but few patches of a dwarf form of J. pseudosabina
have also been found at altitudes of about 3,700 m in
the Peter the First Mts.
Floristic composition: The canopy of this wood is
clearly dominated by Juniperus pseudosabina. Unlike the
thermophilous J. polycarpos var. seravschanica stands in
the shrub layer the main diagnostic taxon has a low
proportion. This layer is dominated by species such
as Lonicera nummulariifolia, L. korolkowii and Rosa webbiana. The herb layer has a cover of 20–95% (mean
60%; Fig. 5c) and consists of 17–34 species (mean ca.
26; Fig. 5e, 7f). The herb layer occurs with a number
of Eastern Irano-Turanian and Euro-Siberian species
(Fig. 5g), most commonly Astragalus tibetanus, Brachypodium sylvaticum, Campanula glomerata, Crepis pulchra,
Euphrasia pectinata, Festuca rupicola, Phleum phleoides,
Poa pratensis and Trifolium repens. Juniperetum pseudosabinae is rich in moss species that form an abundant
layer with Hypnum cupressifoliae, Distichum capillaceum,
Sanionia uncinata and Syntrychia ruralis. The moss cover reaches 65%, with the mean of ca. 35% (Fig. 5d).
Habitat characteristics: The tree stands of Juniperus pseudosabina are associated with the upper montane belt of the northern ranges in central Tajikistan
and occur on brown mountain soils with poorly developed profile. It prefers moderately sloping sites
(mean inclination approx. 25º, Fig. 4b) and with exposure from north-east to north-west (Fig. 6).
Remarks: In Middle Asia, there is a gradual transition of dense stands of J. pseudosabina into mosaic dwarf shrub communities in the treeline zone is
observed. Such plots with ca. 20–40% juniper cover
and a rich herb layer composed mainly by cryophilous steppe or alpine tall-forb taxa were observed in
various regions of Tajikistan and Kyrgyzstan, e.g. in
the Kyrgyz, Peter the First and Talass Mts. Due to
the considerable physiognomic distinctiveness and
different species composition, these plots were excluded from the study.
Holotypus: (sequence number in Table S1 – 84) 12
July 2021; 68.74634 °E; 39.60838 °N; 2324 m a.s.l.;
aspect NW; slope 25°; plot area 100 m2; cover tree
layer 80%; cover shrub layer 20%; cover herb layer
55%; cover moss layer 35%; species composition:
Tree layer: Juniperus pseudosabina 5. Shrub layer: Lonicera nummulariifolia 2, Festuca rupicola +, Juniperus pseudosabina +. Herb layer: Carex turkestanica 3, Juniperus
pseudosabina 2, Rosa kokanica 2, Koeleria pyramidata 1,
Ligularia thomsonii 1, Thymus seravschanicus 1, Berberis
integerrima +, Campanula glomerata +, Cerastium dichotomum subsp. inflatum +, Crepis pulchra +, Cystopteris fragilis +, Erigeron seravschanicus +, Gentianella
turkestanorum +, Geranium regelii +, Helictotrichon
hookeri +, Oxytropis ovczinnikovii +, Petrorhagia alpina
+, Phleum phleoides +, Poa fragilis +, Taraxacum agg. +,
Thalictrum kuhistanicum +, Viola suavis +. Moss layer:
Hypnum cupressiforme 3, Sanionia uncinata 2.
Discussion
Comparisons of the Middle Asian
juniper woods to the neigbouring areas
Undoubtedly, the thermophilous juniper woods
of Tajikistan dominated by J. polycarpos var. seravschanica, in terms of structure and habitat conditions,
are similar to those of Iran, Turkmenistan, Afghanistan and Uzbekistan (Freitag, 1971; Popov, 1994).
They probably differ only in their species composition due to the high rate of Pamir-Alai endemism.
Despite the scarcity of phytosociological data, we
can find fairly similar features of J. polycarpos var.
turcomanica stands in western Kopet-dagh to eastern
Khorassan in the Bardu and Bezd mountains in Torbat-e Jam on the Iranian-Turkestanian borderland.
They occupy the montane belt at 800–2,400 m a.s.l.
and form open woodland (the average tree cover 30–
40%) with a rich undergrowth composed of thermophilous, steppe and tall-forb species (Memariani et
al., 2016b). However, unlike the Zeravshan juniper
stands in Pamir-Alai, they are limited in the upper
boundary not by zonal juniper stands (like J. pseudosabina woods in Pamir-Alai) but by shrubby, subalpine
vegetation with J. sabina and J. communis. The exposure of the remnant Turkmen junipers also varies
and, due to the increased drought in this region, they
mainly occupy the northern slopes.
In Afghanistan, a stand of J. polycarpos var. seravschanica forms the upper tree line in the northern
Hindukush at an altitude of about 3,500 m. Due to
harsh conditions and severe drought in this region,
the higher elevations are treeless. They are the only
coniferous stands outside the areas of monsoon influence and due to the highly variable topography,
harbour a wide range of undergrowth species (Freitag, 1971). Juniperus open woods form a montane
belt from ca. 1,400–2,900 in the west and in the
north-east from ca. 3,000–3,100 to about 3500 m
a.s.l. Total precipitation here ranges from 500 to as
much as 1,200 mm, with winter precipitation in the
north and summer monsoon in the south. Juniper
stands inhabit skeleton, rendzina soils and form fairly dense stands with up to 60–80% canopy coverage.
They share many species with Tajik stands such as J.
polycarpos var. seravschanica, J. semiglobosa, Ephedra equisetina, Lonicera nummulariifolia, Amygdalus bucharica,
Crataegus songarica, Phlomis cashmeriana, Thalictrum sultanbadense, Koeleria cristata. A striking feature of this
woods is the great share of šhiblyak taxa like Cercis
griffithii, Crataegus turcestanica, Pyrus korshinskyi, Malus
turkmenorum, Astragalus cisdarvasicus. This proves to
some extent the validity of the hypothesis of a common origin of the thermophilous juniper woods and
xerophytic šhiblyak scrub from a single Tertiary formation, the palaeošhiblyak (Kamelin, 1967). At the
Phytosociological survey of juniper wood vegetation in Tajikistan (Middle Asia)
beginning of the Lower Miocene, significant climatic
fluctuations and gradual aridization caused the replacement of broad-leaf turgay woods (the so called
ancient Mediterranean vegetation) by the more xerophytic palaeošhiblyak. This formation is considered
to be the ancestor formation of juniper groves and
present-day šhiblyak (Kamelin, 1967; Ovchinnikov,
1967, 1971; Pavlov, 1980).
The Afghan stands of J. polycarpos var. seravschanica
are closely related to the North Pakistani sites of J.
excelsa subsp. polycarpos in Chitral (Nüsser & Dickore, 2002). In our opinion, the taxonomic vagueness
of the genus is responsible for the different species
assignments and it is likely that this woody vegetation is formed by the same taxon. Despite the physiognomic similarity of this open woodland, a kind of
steppe-forest vegetation, there are also floristic similarities, as both vegetation types share a number of
species such as Acanthocephalus benthaminanus, Artemisia persica, Crataegus songarica, Gypsophila floribunda,
Rosa beggeriana or Scabiosa olivieri.
In Armenia, juniper stands are included in the
Mediterranean matorral vegetation. The shrubby formation consists of Juniperus polycarpos, J. foetidissima,
J. communis, J. oxycedrus, J. phoenicea, J. lycia, J. drupacea
and J. thurifera. It rather resembles Mediterranean
shrubby vegetation with a distinct species composition (Fayvush & Aleksanyan, 2016). However, there
is also a stand of J. excelsa and J. foetidissima in the
upper montane and subalpine belts of Armenia,
which physionomically is quite similar to typical Irano-Turanian woods in Tajikistan.
Also juniper woods in northern Iran in the Alborz
and Hezarmasjed Mts. reveal close similarities to the
open stands of Middle Asia. They occur at 1,300–
2,100 m altitude and are dominated by Juniperus excelsa or J. polycarpos and share a number of species,
like Lonicera nummulariifolia, Cotoneaster nummularioides, C. numulariifolia, Berberis integerrima, Crataegus
pontica (Kartoolinejad & Moshki, 2014; Ravanbakhsh
et al., 2016). Six associations and five subassociations have been defined in Alborz (Ravanbakhsh et
al., 2016). However, most of them on scarce samples.
These have been assigned to Junipero-Pistacietea Zohary 1973.
The Zeravshan juniper stands in Tajikistan also
show close relationship to other eastern Mediterranean and Western Irano-Turanian open woods dominated by Juniperus excelsa, J. oxycedrus or J. polycarpos in
Turkey, J. excelsa in Al-Hajar Mts. in Oman, Lebanon,
southern Pakistan, Dagestan (eastern Caucasus; e.g.
Fisher & Gardner, 1995; Çolak & Rotherham, 2006;
Sarangzai et al., 2012; Douaihy et al., 2013; Sadykova
et al., 2018; Ambarlı et al., 2020).
In contrast to the woods of J. polycarpos var. seravschanica, the Tajik northern forests of J. pseudosabina display strong relationship with juniper woods of
31
Central Asia, Tibet and the Himalayas. In the Himalayan range, almost pure juniper stands were found
above about 3,500 m a.s.l. in the driest areas. These
stands are dominated by Juniperus recurva in the eastern Himalayas and by J. wallichiana and J. communis in
the western part of the range. Like in Middle Asia,
the shrub and undergrowth layers harbour species of
Caragana spp. and Artemisia spp. In the far northern
Altay Mts. (Gobi Altay), J. sabina scrubs are recorded
at the upper boundary of trees which to some extent
resamble the uppermost scrub-like stands of J. pseudosabina in higher elevations in Peter the First and
western Pamirian Ranges (Vanch, Yazgulem, Rushan;
Wesche et al., 2005). The montane scrubland of the
southern Mongolian Mts. has been included in the
order Juniperetalia pseudosabinae (Mirkin et al., 1986)
and is probably the northernmost refugium of this
vegetation in Central Asia. In Tajikistan, south-western refuges of this vegetation form dense zonal
woods in the upper montane belt.
Towards the east, the cryophilous woods change
their floristic composition and are mainly built by
Juniperus indica, J. convallium and J. tibetica, which
form zonal subalpine scrub or isolated bushy relicts
(Ghimire et al., 2008; Miehe et al., 2008; Tambe &
Rawat, 2010). Their last remnants in southern Tibet
have often been saved only because of their religious
significance, but their syntaxonomic status remains
unknown. This woods were much more widespread
before the deforatation began ca. 600 years ago. Human impact relating to pastoral culture and intensive
grazing are causing severe degradation of Tibetan
woodlands, which support the transformation into
the present degraded pastures (Miehe et al., 2008).
Does altitude, or rather continentality
and precipitation gradients really play a
major role in the classification of juniper
woods in Middle Asia?
Most of the studies on the Juniperus woods of Tajikistan to date pointed the altitude as an important
determinant of the distributional pattern of juniper
forests and thus highlighting the thermophilous-cryophilous division. However, looking more generally
and considering the entire Asian juniper-dominated vegetation, it seems that the primary variable is
higher annual temperature range (continentality)
with its associated strong phytogeographic division
between the Turkestan province and the much more
cold and continental Central Asian province. This is
also reflected in the precipitation pattern with winter (spring) rainfall in Southwest Asia and summer
peak rainfall in Central Asia. Such a strong climatic,
floristic and environmental gradient is also apparent
in many other vegetation types, especially steppes,
32
Arkadiusz Nowak et al.
semi-deserts and scree vegetation (see Nowak et
al., 2016, 2018, 2021) and should perhaps become
a reason for the revision of the phytogeographic
division of Middle & Central Asia. For this reason,
we believe that the division of Tajikistan‘s juniper
forests should take into account this fundamental
difference in climatic conditions and may be reflected even at the level of the phytosociological class.
Therefore, we classify the thermophilous stands as
Mediterranean and Southwest Asian Pino-Juniperetea, and cryophilous ones to Juniperetea pseudosabinae.
Such a division reflects a continentality and precipitation during the growing season rather than an altitudinal gradient, although in Tajikistan these two
main vegetation types overlap and form interrelated zones in the montane belt. This is also proved
by the composition of typical Irano-Turanian and
Eastern Irano-Turanian with Euro-Siberian species
in our plots (Fig. 5f and g). The first group clearly dominates in Zeravshan juniper stands and the
second in the northern J. polycarpos ones. Probably
the origin of these two types of woody vegetation in
Middle Asia is much different. As mentioned above,
south-western (Turkestan) juniper woods are direct
descendants of palaeošhiblyak, which evolved from
the Tertiary Turgay flora and then underwent xerophytisation after climate aridisation (Kamelin, 1967;
Ovchinnikov, 1967, 1971; Pavlov, 1980) while Central Asiatic juniper stands may have a different history (Mao et al., 2010).
Should we classify Zeravshan juniper
woods into a class of Pino-Juniperetea
of Europe and turkestan woods into
Juniperetea pseudosabinae?
Classifying our plots to the Pino-Junipereta Rivas-Martinez 1964 may raise some doubts. This is
a vegetation type coined for juniper and pine dominated woods of the western Mediterranean (Brullo
et al., 2001). The range of this vegetation has been
extended much to the east, where it is represented by
the order Juniperetalia hemisphaericae, which comprises two new alliances: Berberidion aetnensis, restricted
to the central Mediterranean area, and Berberido creticae-Juniperion foetidissimae, distributed in the eastern
Mediterranean (Brullo et al., 2001). However, Mucina et al. (2016) in their fundamental work on the European syntaxa inverted the name into Junipero-Pinetea stating that this vegetation type refers to various
pine-dominated communities. Pine is absent in Tajikistan and Kyrgyzstan, and towards the south, the
first sites of Pinus wallichiana and other species only
begin in Pakistan and Afghanistan in monsoon-influenced areas. We believe that the stands of J. polycarpos
var. seravschanica should be included in this class. The
considerable floristic distinctiveness of Middle and
Central Asian juniper woods can be expressed at the
level of class and order. On the other hand, the class
Pino-Juniperetea refers to mountainous habitats in a
Mediterranan type climate zone with winter rains
and skeletal and calcareous soils. In this respect, it
fits very well with Tajik habitats.
For the Middle East region, the juniper open
woodlands were classified along with pistachio
stands. At a higher rank of phytosociological division, they were included in the Junipero-Pistacietea
class (Zohary, 1973). Apart from the fact that this
class was described incorrectly (Art. 2b, see Mucina et al., 2016), it seems that current knowledge
does not justify the inclusion of both types, juniper
and pistachio groves, in one vegetation class. In the
mountains of Uzbekistan, Tajikistan and Kyrgyzstan,
they are clearly distinguished in the landscape as two
separate belts of vegetation. The juniper stands grow
in the upper montane to lower subalpine belts and
only occasionally overlap with pistachio woodlands
(some plots of J. seravshanica in Hodzhamumin, Babatag and Kugitang Mts.). In our relevés from Tajikistan we have not noticed the occurrence of Pistacia
vera in juniper woods.
Certainly, further studies throughout Southwest
Asia are needed to answer the question to which
class the thermophilous juniper woods of Tajikistan
belong to, or whether there is a need to establish a
separate class for this vegetation in the Irano-Turanian region. These studies should also include an analysis of the origin and distribution of all Juniperus sect.
sabina species (Mao et al., 2010). At the present stage
of the study, we are only certain that for western
Middle Asia we can coin the order for thermophilous Zeravshan juniper woodlands that encompasses
at least the Pamir-Alai, Hindukush and western Tian
Shan montane belt.
The fact that the Central Asian juniper woods belong to the class Juniperetea pseudosabinae also raises
some doubts. This is because of the southernmost,
marginal location of this vegetation in Tajikistan in
relation to the locus classicus in Altai. The vegetation
in Altai on the northern limit has a physiognomy of
mosaic dwarf scrub-grassland, while in Tajikistan it
forms dense forest stands, especially in the Turkestan
range. An additional complication is the fact that the
author of the class synonymised it with the Artemisio santolinifoliae-Berberidetea sibiricae Ermakov, Chytrý
and Valachovič (2006), which was referred to steppe
and forest-steppe vegetation (Mirkin & Naumova,
2012), but originally to petrophytic communities
(Ermakov et al., 2006). Moreover, they did not list
Juniperus pseudosabina in the set of diagnostic species.
In our opinion, these two vegetation types are fundamentally different and we are convinced that the
Juniperetea pseudosabinae class should be preserved.
Phytosociological survey of juniper wood vegetation in Tajikistan (Middle Asia)
Nevertheless, the final arrangement of this class
should take into account the results of further phytosociological studies in southern Tibet and the Himalayas in the communities of J. recurva, J. convallium, J.
indica and J. tibetica.
Conservation
Following long-term timber exploitation by local communities, woodlands are one of the most
threatened ecosystems in Tajikistan (Safarov, 2003).
Approximately 90% of them have been legally or
illegally logged in recent centuries, resulting in extraordinary deforestation and consequent landslides
and soil denudation. Almost all authors describing
juniper woods in Middle Asia indicate that their area
and quality are rapidly decreasing. And it is important to notice that juniper woodlands in Tajikistan
constitute half of the forested area. Their cutting for
heating and construction purposes poses a serious
threat not only to forest ecosystems of Middle Asia,
but also to soil erosion and strong aridization. Secondary vegetation that develops after the tree and
shrub layer has been cleared are tall-forb communities (Nowaket al., 2020c), which are often intensively used by grazing. Despite the implementation
of some conservation measures and the establishment of the Zeravshan Nature Reserves and Tajik
National Park, the process of juniper woods decline
is still ongoing. A similar situation is taking place
in neighbouring countries. In Kopet-dagh the area
of juniper forests is constantly decreasing and only
between 1930 and 1960 it decreased by 50%. The
same applies to forests with J. seravshanica, in Kugitang Mts. on the border with Uzbekistan and Afghanistan. Due to human impact, the lower boundary of this juniper stand has shifted from approx. 800
to as high as 1,300 to 1,700 m a.s.l. (Popov, 1994).
Currently, also due to climate change, we face a dramatic endangerement of woody ecosystems in Middle Asia and immediate action is needed to stop this
degradation processes.
Author contributions
A.N. planned the research, A.N., S.Ś, M.N. conducted the field sampling, A.N. and S.Ś. wrote the
draft of the manuscript, S.Ś. performed the statistical
analyses, while all authors contributed to the writing.
Acknowledgements
The authors wish to thank Firuza Illarionova from
the Nature Protection Team Dushanbe for assistance
and help in organizing expeditions. The research was
partially supported by the National Science Centre,
Poland, grant no. 2020/04/X/NZ8/00032.
33
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