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Distribution of Ambrosia artemisiifolia (Asteraceae) invasive plant species in Azerbaijan (South Caucasus)

Authors:
Pak. J. Bot., 57(1): 319-325, 2025. DOI: http://dx.doi.org/10.30848/PJB2025-1(35)
DISTRIBUTION OF AMBROSIA ARTEMISIIFOLIA (ASTERACEAE) INVASIVE PLANT
SPECIES IN AZERBAIJAN (SOUTH CAUCASUS)
ABDIYEVA RENA1*, IBRAHIMOVA AIDA2, ASADOVA KAMALA1 AND LITVINSKAYA SVETLANA3,4
1Department of Geobotany, Institute of Botany, Ministry of Science and Education of the Republic of Azerbaijan,
Baku, Azerbaijan
2Department of Biomorphology and Phytointroduction, Seed Bank, Institute of Botany, Ministry of Science and
Education of the Republic of Azerbaijan, Baku, Azerbaijan
3Department of Botany, Southern Federal University, Rostov-on-Don, Russia; 4Kuban State University, Krasnodar, Russia
*
Corresponding author's email: abdiyeva.rena@mail.ru
Abstract
Plants with an alien native range are today a serious problem for local floras. Invasive plants pose a threat not only of an
ecological nature; also many of them hybridize with native plants and also cause allergic reactions in humans. In recent years,
an increase in the activity of invasive plants has been observed in the world flora, including the flora of Azerbaijan. Researchers
emphasize that climate change is one of the factors in this process. In comparison with previous years, an increase in the
activity of the invasive species Ambrosia artemisiifolia L. (Asteraceae) in Azerbaijan has been established. In this study, the
modern habitats of A. artemisiifolia in the country were studied, plant communities with the participation of the species were
identified, and ecological niche models were created to represent the potential distribution of this species in the territory of
Azerbaijan. These results indicate that predictive modeling can be an important resource for early detection of the
aggressiveness of potentially invasive species, which will play a positive role in preventing their undesirable spread and
conserving biodiversity. Future predictions were made based on the MRI-CGCM3 (MG) scenario for four representative
concentration pathways (RCPs) based on Intergovernmental Panel on Climate Change (IPCC). The main findings of the study
are that A. artemisiifolia is now beginning to invade the forest and grassland ecosystems of Azerbaijan.
Key words: Plant invasion, Ecological niche modelling, Natural ecosystems, Conservation.
Introduction
Invasive alien species (IAS) are known to be one of the
main causes of the decline in the biodiversity and floristic
and faunistic gene pool of local ecosystems, as well as
having a negative impact on sectors of the economy
(agricultural, etc.) and human health (Barrett & Husband,
1989; Froud-Williams, 1997; Pimental et al., 2001;
Richardson & Pysek, 2006; Palmer & Nursey-Bray, 2007).
Introduction of IAS into natural and semi-natural
communities can lead to significant changes in ecosystems
or their complete transformation (Elton, 1958; Reichard &
Hamilton, 1997; Simberloff, 2011; Sax & Brown, 2000;
Meekins & McCarthy, 2001; Réjmánek et al., 2000). In the
world practice of effective prevention of the spread of IAS,
first of all, it includes the creation of various databases
(Global Naturalized Alien Flora (GloNAF); Global Invasive
Species Database, (GISD); Hulme et al., 2009; Lambdon et
al., 2008), as well as ecological and geographical modeling
of niches, which allows the potential distribution of
biological objects to be determined with high accuracy (Nix,
1986; Solano & Feria, 2007; Suárez-Mota et al., 2016).
The ecological niche model (ENM) is currently one of
the most widely used methods for predicting the potential
spread of IAS (Guisan & Thuiller, 2005; Barve, 2011). The
Maxent (maximum entropy) approach of the ENM is
commonly used by scientists in conservation biology, ecology,
and evolution research (Kurpis, 2019; Kariyawasam et al.,
2019). Various mathematical and statistical aspects make the
MaxEnt modeling approach well-suited for ENM. In recent
years, researchers have begun to predict the distribution of
invasive species in the Caucasus region (Pshegusov, 2020).
For Georgia, in the South Caucasus, predictions have been
made for several invasive species in the country (Slodowicz et
al., 2018; Thalmann et al., 2014). The study on plant invasions
started in Azerbaijan a few years ago, and currently the focus
is on identifying and studying the phytocenotic role in the
areas of discovery (Abdiyeva, 2019; Abdiyeva et al., 2023).
Along with the research of phytocenotic features of invasions,
we also aim to use ENM to predict the potential distribution
of IAS in Azerbaijan in order to control the spread of IAS in
the country (Abdiyeva et al., 2021).
Ambrosia artemisiifolia L. (Asteraceae) is an annual herb
with a North American native range (Lambdon et al., 2008).
In Europe, this plant species was accidentally introducedin the
18th century, along with soil and seed agricultural material
delivered from North America (Brandes & Nitzsche 2006;
Bullock et al., 2012). Researchers who studied the biological
and phytocenotic characteristics of the species in the local
floras of Western and Eastern Europe noted that the activity of
A. artemisiifolia in Europe began to increase significantly in
the middle of the 20th century, and one of the main reasons for
this was climate change (Brandes & Nitzsche, 2006; Bullock
et al., 2012). The first outbreaks of A. artemisiifolia appeared
in 1918 in Russia; in the North Caucasus, the species was
discovered in 1930 (Vinogradova et al., 2010). From the
North Caucasus the plant migrated to Transcaucasia. In
Azerbaijan, A. artemisiifolia was discovered in the middle of
the 20th century in areas located on the border with Georgia
(Akhundov, 1961). Therefore, the introduction of A.
artemisiifolia into Azerbaijan most likely originated in
Georgia, a neighboring country situated in the South Caucasus
contact zone. Transference of A. artemisiifolia from Georgia
occurred with fruits, which could have been brought with
seed, and food material, on the wheels of vehicles, on the
fur/wool of transported livestocks, with water flows and wind
from the territory. In local ecosystems, A. artemisiifolia spread
naturally - by water flows (mudflows, river network), wind
flows, with littered waste and on animal fur.
ABDIYEVA RENAET AL.,
320
A. artemisiifolia has established itself in recent years
and transformed the ecosystems of Azerbaijan. It was
regarded as a common weed in the gardens and orchards of
rural residents. Additionally, the species has become a
malicious biological pollutant of agro-phytocenoses
resulting in the decrease of the quality of the crop. The
range of A. artemisiifolia was limited to the Steppe plateau
botanical-geographical region in northwest Azerbaijan by
the middle of the 20th century (Akhundov, 1961). It started
to expand to the neighboring regions (the Greater Caucasus
Mountains) around the beginning of the 21st century,
where some of them came in touch with protected areas
(Abdiyeva 2019; Abdiyeva, Litvinskaya 2023). Over the
past ten years, this plant has started to spread to other
districts of the country. Therefore, the objectives of our
research were to study the phytocenotic role of A.
artemisiifolia in local natural ecosystems, as well as the
possible distribution of the species in Azerbaijan under
current and future climatic conditions. This will give an
idea of which areas of Azerbaijan, as well as which
protected areas, are at particular risk of invasion.
Material and Method
Study area: Azerbaijan is one of the countries of the
Caucasus region, which is located in the east of the South
Caucasus, at the crossroads of Eastern Europe and
Southwestern Asia, between latitudes 38°–42° N, and
longitudes 44°52° E. It is area is 86,600 km2. The area is
situated between the Caspian and Black seas. It is bounded
by the Caspian Sea to the east, Russia to the north, Georgia
to the northwest, Armenia to the west, Iran to the south and
has a short border with Turkey to the northwest through the
Nakhchivan. The great geopolitical significance of
Azerbaijan’s position once derived from caravan routes
connecting distant countries of the West in ancient times,
as well as the famous Silk Road stretching from China to
Europe. Azerbaijan is mainly mountainous country, and it
is surrounded by the Greater Caucasus (Bazarduzu
mountain peak, 4466 m above sea level (asl)), Lesser
Caucasus (Qamishdag, 3724 m asl), and Talysh Mountains
(Komurkoy, 2492 m asl). The Kur-Araz lowland, with
plains and low mountainous reefs, is situated between the
Greater and Lesser Caucasus and stretches to the Caspian
Sea in the central-southern part of the country. Gobustan
lies in the eastern part with its numerous mud volcanoes.
Lankaran lowland is situated in the south-eastern part,
stretching along the seacoast. The elevation changes over a
relatively short distance from lowlands to highlands,
between –27 meters below sea level up to 4466
(Museyibov, 1998). The complex natural conditions of the
country form 11 landscapes. There are representatives of
the ancient forest, boreal, steppe, xerophilic, desert,
caucasian, as well as adventitious (alien) geographic types
in the flora (Grossheim, 1940-1948). The climate of
Azerbaijan is the Northern borders of subtropical climate
zone. The high mountain plays an important role in
preventing cold air masses coming from the north, making
the air condition colder in the northern slopes and relatively
mild in the southern slopes. Eight out of 11 main climate
types are represented in the country (Museyibov, 1998).
These are semidesert and dry steppe, moderately warm
climate with dry winters, moderately warm climate with
dry summers, cold climate with dry winters, cold climate
with dry winters, temperate warm climate with equal
precipitation, cold climate with rains in all seasons and
mountain tundra climates.
There are 44 Protected Areas (PA) organized on the
territory of Azerbaijan to preserve the biodiversity of the
country (www.eco.gov.az). The PA covers 8,93km2, which
is 10.31% of the country's territory. The main protected
areas are concentrated along the Caspian Sea (Samur-
Yalama, Absheron, and Gizilagach National Parks), in the
Greater Caucasus (Shahdag National Park, Zagatala and
Ilisu State Nature Reserves), in the Lesser Caucasus
(Goygol National Park and Eldar Pine State Nature
Reserve) and in Talysh (Hirkan National Park). The global
(Protected Planet, 2014-2024) data sources provide
information about 35 of the 45 PA. Since it was not possible
to obtain information on the territories covered by the
remaining protected areas, we were only able to use 35 PA
for the analysis in the study.
Data collection: Data on the past and current distribution
of A. artemisiifolia were extracted from the literature,
herbaria, and our field surveys. To identify current
distribution areas of A. artemisiifolia monitoring was
carried out throughout the territory of Azerbaijan in the
period 2012-2021. Field studies were carried out in various
types of habitats: anthropogenic (settlement, garden, park,
farm, railways, rural roads, highways), semi-natural
(abandoned sites connecting anthropogenic and natural
areas) and natural (meadow, forest, pond, riverbank, etc.).
In total, 277 distribution records were collected and used
in our ecological niche modeling. Herbarium entries
collected during field works are stored in the Herbarium
Foundation (BAK) at the Institute of Botany of the
Ministry of Science and Education of the Republic of
Azerbaijan. Species abundance is determined using Braun-
Blanquet scale (Braun-Blanquet, 1964).
Ecological niche modeling: The ecological niche
modeling was performed using the species distribution
modeling (SDM) approach with the Maximum Entropy
Method (Maxent) (Phillips et al., 2006;). We used
"dismo" package of "R" software (R Core Team, 2019).
The current climate conditions (Fick et al., 2017;
http://worldclim.org/version2, accessed 2017) and future
climate projections (the resolution of 30 seconds of arc)
(Hijmans et al., 2005; Hijmans et al., 2011; Hijmans,
2012; http://www.worldclim.org/cmip5_30s, accessed
2017) were downloaded from the WorldClim database, as
a set of 19 bioclimatic variables. Future climate
projections were constructed based on Scenario MRI-
CGCM3 (MG) (Yukimoto et al., 2012) for 4
representative concentration pathways (RCP 2.6; 4.5; 6;
8.5) from the Intergovernmental Panel on Climate
Change (IPCC), which is part of simulations in the
Coupled Model Intercomparison Project Phase 5
(CMIP5) (Meinshausen, 2011; IPCC, 2014). First, we
analyzed the importance of all bioclimatic variables for
the probability of species occurrence and selected the
DISTRIBUTION OF AMBROSIA ARTEMISIIFOLIA (ASTERACEAE) IN AZERBAIJAN
321
most sensitive climate conditions to variation in the
distribution. Ultimately, 11 climatic variables (Bio 1; 6;
7; 9; 11; 12; 14; 15; 17; 18; 19) were used as input data.
To evaluate the predictive accuracy of the model, we
selected the receiver operator characteristic (ROC) by the
area under the curve (AUC) statistical test (Hirzel et al.,
2006; Lobo et al., 2008). Finally, we overlaid the
shapefile of Azerbaijan and its protected areas. Shapefiles
of protected areas were downloaded from the Protected
Planet website (Protected Planet, 2014-2024) and
shapefile of the country from DIVA-GIS repository
(DIVA-GIS, 2017). The protected areas represented with
blue lines on the predicting maps.
Results
Modeling approach: The final model had the AUC value
of 0.98, which indicated excellent performance of the
model and high reliability of the results.
Before using the model to project the potential
distribution of A. artemisiifolia, we assessed how well the
model could predict occurrence of the species. Firstly, we
analyzed all bioclimatic variables in order to select the
necessary climate determinants. We then created a model
using only the combinations of bioclimatic variables that
best predicted the species’ occurrence (Table 1). While all
variables had corresponding probability percentages, some
made more significant contributions to the model.
Phenological observations and habitat studies of A.
artemisiifolia in the study area indicated that temperature
factors and precipitation affected its distribution equally,
as indicated by the highest percentage contributions of
Bio6 (34.3%), Bio9 (17.1%), Bio14 (23.8%), and Bio19
(13.4%) (Table 1). The estimated relative contributions of
environmental variables to the model for IAS were
generally consistent with field observations of these
species in nature.
Current distribution range: During field surveys (2012-
2021) on the territory of Azerbaijan, we discovered new
distribution areas of A. artemisiifolia. It was more common
in the north-west of Azerbaijan (Steppe plateau, Alazan-
Ayrichay valley, Greater Caucasus Mountains). Research
on the phytocenotic characteristics of plant species in the
investigated area indicated that agrophytocenoses had a
higher prevalence of A. artemisiifolia than natural habitats.
It tends to grow more in lowland and foothills agricultural
areas (Steppe plateau, Alazan-Ayrichay valley) at a height
of 200–400 m above sea level. However, in the northwest
part of the country (the Greater Caucasus Mountains), the
species had already begun to expand into natural
ecosystems such as forests and meadows in the lower and
middle mountain belts (800–950 m asl) in recent years. In
forest ecosystems, it was found in the herbaceous layer
dominated by Acer campestre, Corylus colurna, Acer
laetum, and Quercus iberica (Table 2).
Over the 7-8 years, the spread of A. artemisiifolia has
been found in a small area of the southern (Lankaran
lowland) and central (Absheron) coast of the Caspian Sea
(within Azerbaijan). These new habitats of the species are
very different from the habitats found in northwestern
Azerbaijan. In all studied natural and anthropogenic
territories of the north-west, the species was in satisfactory
condition. On the contrary, in the eastern part it was found
in the form of single, crushed individuals in summer
cottages. We assessed the appearance of the species in these
areas as an unintentional introduction of seed material into
the soil. The soil was brought from the territory of the
Azerbaijani part of the Greater Caucasus in order to enrich
the sandy soil of garden plots, which was poor in
microelements. However, at present A. artemisiifolia does
not show activity or high vitality.
Current niche modeling: Current niche modeling showed
that, under the current climate conditions, 21.5% of total
area across the country was suitable for potential invasion
by A. artemisiifolia. Within the respective total suitable
area, 8.99% was highly suitable for A. artemisiifolia (Fig.
1a). The optimal environmental conditions were identified
in lowland, foothill, and mountainous areas in the
northwest of Azerbaijan.
Future niche modeling: A. artemisiifolia will likely
continue to spread into the same areas that it has occupied
in the past and the present, following the model analysis
performed under future climatic conditions (Fig. 1b-e).
Furthermore, the suitable habitat area will increase in the
future, especially under scenarios RCP 4.5, RCP 6 and RCP
8.5 (Fig. 1c-e). The migration vector is expected to move
from the northwest region of Azerbaijan to the central and
western regions. According to RCP 2.6 and RCP 8.5, there
is a possibility that the species will spread to new regions
of the Greater Caucasus in the northeastern part of the
country (Fig. 1b, e), and from there to regions that are in
contact with the Caspian Sea coast. It is quite possible that
this kind of migration will occur as a result of migrating
birds and animals carrying achenes or because soil is richer
in microelements is being transported from the country's
northwest to the coast, where it will be used to introduce
ornamental plants and fruit trees. Fertile soil is still being
transported to Azerbaijan's lowlands in a similar way.
Consequently, isolated individuals of A. artemisiifolia have
occasionally been discovered in summer cottages along the
coast, particularly in Absheron.
Thus, under emission scenarios RCP 2.6, 4.5 and 6.0,
the potential geographic distribution of A. artemisiifolia
will expand into the plains and foothills of the Greater
Caucasus. From there, this species will spread into the flat
regions of the northern part of the Lesser Caucasus, as well
as into the foothill and low-mountain zones of the central
part. Under the predicted conditions of RCP 6 and RCP 8.,
the emergence of A. artemisiifolia in the Caspian Sea coast
will become possible.
Predicted distribution of target species in protected
areas: The distribution of A. artemisiifolia in Azerbaijan
raises a natural question: what is the threat of this plant
intrusion into protected areas? To do this, we overlay maps
of the predicted distribution of A. artemisiifolia onto a map
ABDIYEVA RENAET AL.,
322
of protected areas of Azerbaijan. This allowed us to
determine protected areas already subjected to colonization
by the species, as well as areas with the greatest risk of
invasion in the future. The analysis showed that in front of
some protected areas there was a high probability of the
introduction of A. artemisiifolia (Fig. 1b-e). In particular,
the analysis of model indicates a high risk (62%) for the
Zagatala and Ilisu State Reserves. These protected areas are
located in the zone of contact of current habitats of A.
artemisiifolia. Therefore, the expected invasioncan be
considered quite natural. Furthermore, it is quite possible
that A. artemisiifolia species will spread to six more
protected areas around the country. These are the
following: Turyanchay and Garayazi State Nature
Reserves; Korchay, Shamkir, Ismayilli, and Gabala State
Nature Sanctuaries (Fig. 1c-e).
Table 1. Percentage contribution (%) of the most influential climatic factors in the current
distribution of Ambrosia artemisiifolia.
Code of variables
Bioclimate variable
Percentage
contribution (%)
Bio1
3.3
Bio6
34.3
Bio7
2.9
Bio9
17.1
Bio11
2.4
Bio12
0.5
Bio14
23.8
Bio15
1.7
Bio17
0.2
Bio18
0.4
Bio19
13.4
Table 2. The main composition of plant communities with the participation of Ambrosia artemisiifolia
in the ecosystems of Azerbaijan.
Distribution area
Plant community
Species composition in the community
(with Braun-Blankuet scale) on a model plot of 20x20 m
Natural phytocenoses
The Greater Caucasus
Mountains; forest; 850-950
m above sea level
Acer campestre + Corylus colurna
Woody layer: Acer campestre (4), Acer laetum (2), Corylus
colurna (3), Quercus iberica (2),
Herbaceous layer: Ambrosia artemisiifolia (3), Equisetum
arvense (3), Dryopteris filix-mas (2), Phytolacca americana (2),
Lythrum salicaria (2) Plantago major (2), Prunella vulgaris (2),
Urtica dioica (1),Sambucus ebulus (1)
The Greater Caucasus
Mountains; meadow; 800-
870 m above sea level
Daucus carota + Plantago major +
Achillea millefolium
Ambrosia artemisiifolia (3), Daucus carota (3), Plantago major
(3), Carum carvi (2), Lamium album (2), Rumex crispus (2),
Phytolacca americana (1), Urtica dioica (1), Cichorium intybus
(1), Salvia verticillata (1), Euphorbia seguiriana (1)
Lankaran lowland; -20 m
above sea level
In the forb groups of the coastal zone
Xanthium strumarium + Amaranthus
retroflexus +Juncus acutus
Ambrosia artemisiifolia (1), Limonium mayeri (2), Convolvulus
arvensis (2),Amaranthus spinosus (2), A. retroflexus (3), Juncus
acutus (2), Xanthium strumarium (3), Daucus carota (1),
Erigeron canadensis (1), Hordeum leporinum (1), Persicaria
lapathifolia (1), Verbascum pyramidatum (1)
Agricultural, semi-disturbed, disturbed phytocenoses
Alazan-Ayrichay valley;
300-400 m above sea level
In the herbaceous layer of hazelnut
orchards (Coryllus avellana)
Ambrosia artemisiifolia (3), Trifolium repens (3), Alkekengi
officinarum (2), Plantago major (2),Prunella vulgaris (2),
Asparagus officinalis (1), Inula helenium (1), Erigeron annuus
(1), Urtica dioica (1)
Absheron; 10 m above sea
level
In summer cottages, in the herbaceous
layer of an orchard
Woody layer: Ficus carica (3).
Herbaceous layer: Ambrosia artemisiifolia (1), Chenopodium
hybridum (2), Urtica dioica (2), Vitis vinifera (2)
Steppe plateau; 200-300 m
above sea level
In littered and abandoned areas in forb
groups
Ambrosia artemisiifolia (2), Urtica dioica (2), Amaranthus
retroflexus (2), Convolvulus arvensis (3)
DISTRIBUTION OF AMBROSIA ARTEMISIIFOLIA (ASTERACEAE) IN AZERBAIJAN
323
Fig. 1. Predicted potential distribution of Ambrosia artemisiifolia in Azerbaijan, based on current (a) and future (b - RCP 2.6; c - RCP 4.5; d - RCP 6;
e - RCP 8.5) climate scenarios. The prediction map for current conditions agreed with occurrences recorded in field surveys (2012-2021). Different
colors and shades show predicted habitat suitability. Blue frames indicate 35 Protected Areas (ef. text for details).
Discussion
Due to their high adaptability, IAS can occupy new habitats
and ecological niches that differ from their natural ranges
(Petitpierre et al., 2012; Early & Sax, 2014; Wan et al.,
2016). Climate change can create and expand suitable
habitats for IAS (Vicente et al., 2013; Thalmann et al.,
2014; Foxcroft et al., 2017). Therefore, modeling
approaches that can predict the distribution of IAS in future
habitats are of critical importance.
ABDIYEVA RENAET AL.,
324
In this study, the current and future distribution of A.
artemisiifolia in Azerbaijan was modeled for the first time
at national scale using the Maximum entropy modeling
approach in SDM. Maps of projections of the distribution
of species showed that the species will penetrate from the
northwest of Azerbaijan.
Mountain forest regions of Azerbaijan will be at the
greatest risk of invasion, according to the projections. Most
of the protected areas in Azerbaijan are located in
mountainous parts of the country, and therefore, the target
invasive species will experience stability since their current
distribution range is in these areas. A. artemisiifolia is the
most “unpredictable” species of the other invasive species оf
Azerbaijan. It is interesting to note that this invasive plant,
which was accidentally introduced about 60 years ago, is
displaying high aggressiveness in its current narrow area in
the foothills region of Azerbaijan (Shaki-Zagatala
economic-geographical region), but for some reason, has not
yet expanded beyond this area. Habitat predictions under
future climate scenarios showed that its distribution will not
remain stable in the foothills region as the habitat there will
become less suitable, with increasing CO2 content and air
temperature, reducing the range of A. artemisiifolia, while
its occurrence may rise in the lower and possibly middle
mountain belts. However, the prediction maps showed that
A. artemisiifolia will not have a tendency to expand its range
overall in Azerbaijan. This interesting finding makes it
necessary to consider the biological and ecological
characteristics of A. artemisiifolia. In the past, A.
artemisiifolia was found exclusively as a weed along narrow
water channels, on abandoned farms, and in areas with
agricultural activities (garden plots, tea, tobacco, and cotton
plantations), where it formed stable groups. Over the past
three years, we have observed occurrence of A. artemisiifolia
in native forest and meadow communities, such as Acer
campestre + Corylus colurna+ and Daucus carota+
Plantago major +Achillea millefolium. The model applied in
this study revealed how actively the invasive species will
spread under current and future climate scenarios. Therefore,
potential habitats for IAS should be monitored in both
protected and unprotected areas. The work should include
field surveys for A. artemisiifolia, campaigns to raise
awareness of these species among the local population, and
studies on the bioecological, phytocenotic and reproductive
characteristics of the species. There are already 70 species of
IAS in Azerbaijan, 10 of which are the most dangerous form,
transformers (Abdiyeva, 2019). The ecological niche and
species distribution modelling approach developed here can
be applied to other invasive species in Azerbaijan, and to
IAS in other countries, in future research to resolve this
global issue.
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