Abstract
The introduction of tree and shrub species represents a key instrument for establishing resilient plantations in arid regions. The present study aimed to provide a comprehensive assessment of the adaptive potential of 42 introduced species cultivated under semi-desert conditions in Western Kazakhstan. Adaptation was evaluated using a multi-criteria framework including winter hardiness, frost resistance, drought tolerance, resistance to pests and diseases, phenological traits, and reproductive capacity. All parameters were scored using a standardized rating system, and an integral adaptation coefficient was calculated as the ratio of the sum of actual scores to the maximum possible value. The results revealed substantial interspecific variability in adaptive performance. The highest adaptive potential was observed in species combining tolerance to abiotic stresses, early cessation of shoot growth, and stable flowering and fruiting. The application of an integral index enabled objective ranking of introduced taxa and identification of promising species for shelterbelts and forest reclamation plantings under arid climatic conditions.
Keywords:
plant introduction; adaptation coefficient; tree species; drought tolerance; winter hardiness; phenology; arid ecosystems
Resumo
A introdução de espécies arbóreas e arbustivas representa um instrumento fundamental para o estabelecimento de plantações resilientes em regiões áridas. O presente estudo teve como objetivo fornecer uma avaliação abrangente do potencial adaptativo de 42 espécies introduzidas, cultivadas sob condições semiáridas no oeste do Cazaquistão. A adaptação foi avaliada utilizando uma estrutura multicritério, incluindo resistência ao frio, resistência à geada, tolerância à seca, resistência a pragas e doenças, características fenológicas e capacidade reprodutiva. Todos os parâmetros foram pontuados utilizando um sistema de classificação padronizado, e um coeficiente de adaptação integral foi calculado como a razão entre a soma das pontuações reais e o valor máximo possível. Os resultados revelaram uma variabilidade interespecífica substancial no desempenho adaptativo. O maior potencial adaptativo foi observado em espécies que combinam tolerância a estresses abióticos, cessação precoce do crescimento dos brotos e floração e frutificação estáveis. A aplicação de um índice integral permitiu a classificação objetiva dos táxons introduzidos e a identificação de espécies promissoras para quebra-ventos e plantios de recuperação florestal em condições climáticas áridas.
Palavras-chave:
introdução de plantas; coeficiente de adaptação; espécies arbóreas; tolerância à seca; resistência ao inverno; fenologia; ecossistemas áridos
1. Introduction
Sparsely forested and treeless territories of arid and semi-arid zones are among the most vulnerable landscapes under contemporary climate change and increasing anthropogenic pressure. These regions are characterized by limited precipitation, pronounced continentality, extreme temperature fluctuations, and widespread saline soils, all of which severely constrain the natural regeneration of woody vegetation (IPBES, 2018; Reynolds et al., 2007). In this context, the introduction of stress-tolerant tree and shrub species is considered one of the principal strategies for increasing forest cover, stabilizing ecosystems, and conserving biodiversity. At the same time, expansion of species pools requires careful consideration of risks associated with naturalization and potential invasiveness (Kowarik, 2010; Richardson and Rejmánek, 2011; Pyšek et al., 2020). Western Kazakhstan is characterized by historically low forest cover and high susceptibility to land degradation processes. In the arid zones of Kazakhstan, restoration and establishment of protective plantations, including forest reclamation and shelterbelt systems, are particularly important for reducing wind erosion (deflation) and soil degradation. Reforestation practices in comparable arid landscapes demonstrate that the success of such interventions depends not only on species selection but also on moisture regimes and silvicultural techniques (Utebekova et al., 2021; Rakymbekov et al., 2023).
Water deficit represents the primary limiting factor for the introduction of woody plants in semi-desert environments, and this constraint is intensifying under rising temperatures and increasing vapor pressure deficit (Grossiord et al., 2020). In arid regions of Kazakhstan, the performance of xerophytic tree species can be substantially improved through soil moisture conservation practices. For example, deep non-moldboard tillage has been shown to enhance survival and growth of black saxaul seedlings compared to untreated controls, while the application of water-retaining materials and growth stimulants improves seedling survival and biometric parameters in plantations (Dosmanbetov et al., 2020). Hydrological regime is another critical determinant of woody plant persistence. In floodplain forests (tugai ecosystems), disruption of natural flow dynamics due to river regulation and other anthropogenic impacts leads to desiccation and soil salinization. Sustainable regeneration in these systems is strongly linked to surface flooding regimes and the synchronization between sediment deposition and seed dispersal. Dendrochronological analyses of turanga indicate that radial growth depends on the interaction between flooding intensity and climatic variables, with precipitation and temperature effects mediated by river hydrology (Dukenov et al., 2023b). These findings highlight the necessity of a multidimensional assessment of introduced taxa, incorporating tolerance to abiotic stress, phenological synchronization, and reproductive capacity.
Despite a substantial body of introduction research in Kazakhstan, most studies have focused on forest-steppe and steppe zones, whereas the semi-desert territories of the Northern Caspian region remain insufficiently investigated. In particular, comprehensive adaptive assessments of introduced tree and shrub species, integrating phenological characteristics, winter hardiness, drought tolerance, and resistance to biotic stressors, are limited.
Accordingly, systematic evaluation of introduced woody species under semi-desert conditions in Western Kazakhstan is both timely and necessary to identify the most resilient and promising taxa for afforestation, protective plantations, and urban greening. The present study provides an integrated assessment of the adaptive potential of introduced species at the Zhanibek experimental station, thereby contributing to the development of science-based approaches for expanding the range of woody plants in extreme arid landscapes.
2. Materials and Methods
2.1. Study area
The study was conducted at the Zhanibek experimental station (Northern Caspian region, Western Kazakhstan), located within the clay semi-desert zone. The climate is sharply continental and arid. The mean annual air temperature is 6.9 °C, with an absolute minimum of -29.8 °C and an absolute maximum of 39.4 °C. Mean annual precipitation is approximately 280 mm. Soils are represented by heavy loams with varying degrees of salinity. Groundwater depth ranges from 6 to 7 m.
2.2. Study objects
The study focused on introduced trees and shrubs cultivated in dendrological collections and protective plantations at the Zhanibek station. A total of 42 species were included, comprising 16 tree species and 26 shrub species of diverse ecological and geographical origins.
2.3. Study design
Field investigations were conducted from 2022 to 2025. Introduced species were evaluated under contrasting microhabitat conditions, including solonetz soil complexes and lower topographic positions. Observations were performed annually throughout the growing season on representative individuals of each species.
2.4. Assessment of winter hardiness and frost resistance
Winter hardiness was assessed in spring following overwintering using a five-point scale based on the degree of shoot and bud damage (from no visible injury to complete dieback of the aboveground part).
The scale of N.A. Bolotov was interpreted as follows:
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5 points (1.0) – no damage under winter conditions;
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4 points (0.8) – occasional damage to current-year shoots;
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3 points (0.6) – complete dieback of current-year shoots and partial damage to older branches;
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2 points (0.4) – dieback of skeletal branches;
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1 point (0.2) – complete dieback of the aboveground part;
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0 points (0.0) – complete plant mortality.
Frost resistance was evaluated after spring frost events using a three-point scale based on damage to leaves, shoots, and reproductive organs:
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3 points – no damage;
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2 points – partial damage to leaves, shoots, inflorescences, flowers, or buds;
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1 point – complete damage to leaves, shoots, and flowers.
2.5. Assessment of drought tolerance
Drought tolerance was visually assessed during dry periods of the growing season using a five-point scale accounting for loss of turgor, leaf necrosis, shoot damage, and plant mortality. For each species, a mean drought tolerance score was calculated. The scale was defined as follows:
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5 points (1.0) – no visible response to drought stress;
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4 points (0.8) – no structural damage to leaves and shoots; temporary loss of turgor and wilting observed;
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3 points (0.6) – marginal leaf damage in the majority of leaves;
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2 points (0.4) – most leaves desiccated; apical shoots damaged;
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1 point (0.2) – leaf abscission; young shoots severely damaged;
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0 points (0.0) – complete plant mortality.
2.6. Resistance to pests and diseases
Resistance to pests and diseases was evaluated throughout the growing season using a five-point scale, taking into account the incidence and severity of damage to vegetative and generative organs The assessment of the resistance of introduced plants to pests and diseases is carried out throughout the growing season on the scale of A.Y. Ogorodnikov:
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5 points (1.0) – no or isolated damage;
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4 points (0.8) – damage to no more than ¼ of plants;
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3 points (0.6) – moderate and periodic damage, primarily affecting vegetative organs;
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2 points (0.4) – severe damage to at least half of plants;
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1 point (0.2) – very severe damage to at least 75% of plants, suppressing growth and development;
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0 points (0.0) – plant mortality.
2.7. Phenological observations
Phenological monitoring included the recording of key developmental stages: bud swelling, bud break, leaf expansion, flowering, fruiting, and leaf senescence.
The duration of the growing season was defined as the interval from bud break to complete leaf yellowing. To assess fruiting, a modified version of the V.G. Kaper scale was applied. The modification consisted of linking fruiting evaluation to flowering intensity and applying it only to species that flowered during the given season but failed to set seed.
During the flowering period:
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5 points – abundant flowering;
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4 points – good flowering;
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3 points – moderate flowering;
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2 points – weak flowering;
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1 point – very weak flowering.
During the fruiting period:
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5 points – very high yield; abundant fruiting;
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4 points – good yield; fruiting observed in most individuals;
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3 points – moderate yield; fruits present in moderate quantities on many individuals;
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2 points – low yield; fruits and seeds present in small quantities;
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1 point – very poor yield; fruits/seeds present only on isolated individuals;
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0 points – no fruit set despite flowering.
2.8. Assessment of generative capacity
Generative capacity was evaluated based on flowering and fruiting intensity using the five-point scale described above, allowing assessment of reproductive performance under semi-desert conditions.
2.9. Integrated adaptive assessment
The overall success of introduction was quantified by calculating an integral adaptation coefficient (K), which was defined as the ratio of the sum of actual scores obtained to the sum of the maximum possible scores for a set of indicators (growth performance, winter hardiness, drought tolerance, resistance to pests and diseases, and generative capacity).
2.10. Statistical analysis
Statistical analyses were performed using Statistica 10.0 (StatSoft Inc., USA). For each indicator, mean values and standard deviations were calculated (mean ± SD). Differences in adaptive performance among habitat conditions were tested using the non-parametric Kruskal-Wallis test due to the ordinal (scored) nature of the data. Relationships among traits were assessed using Spearman’s rank correlation coefficient (Spearman’s ρ). Differences were considered statistically significant at p < 0.05.
3. Results
Under the semi-desert conditions of Western Kazakhstan, introduced tree and shrub species exhibited pronounced species-specific responses to winter and spring temperature stresses.
Among tree species, the highest winter hardiness scores (mean values > 4.0) were recorded for Ulmus pumila, Quercus robur, Acer tataricum, Betula pendula, and Populus alba. These taxa were characterized by minimal damage to shoots and buds after overwintering.
Among shrubs, the greatest winter hardiness was observed in Caragana arborescens, Lonicera tatarica, Elaeagnus angustifolia, Ribes aureum, and Syringa vulgaris.
Frost resistance generally correlated with winter hardiness. Species exhibiting earlier cessation of vegetative growth showed reduced damage to generative organs during spring frost events, indicating phenological synchronization with local climatic constraints. Winter hardiness and frost resistance values are presented in Table 1.
Under conditions of pronounced moisture deficit, the highest drought tolerance scores were recorded for species exhibiting xeromorphic traits and broad ecological amplitude. Among tree species, elevated drought tolerance was observed in Ulmus pumila, Acer tataricum, Fraxinus pennsylvanica, and Quercus robur.
Among shrubs, the most drought-resistant taxa included Elaeagnus angustifolia, Tamarix spp., Caragana arborescens, and Ribes aureum.
In lower topographic positions, mean drought tolerance scores were lower than in the solonetz soil complex. This pattern is likely associated with prolonged vegetative growth and higher tissue hydration levels in moister microhabitats, potentially reducing physiological hardening against water deficit. The drought tolerance of introduced tree species, calculated on the basis of the average values for three replications, is presented in Table 2. The highest indicators of drought resistance are found in species characterized by xeromorphic features and a wide ecological range.
Most introduced species demonstrated high or moderate resistance to pests and diseases. Minimal damage was recorded in Robinia pseudoacacia, Quercus robur, Acer tataricum, and Acer platanoides.
Lower resistance was observed in Betula pendula, Pyrus communis, and Quercus rubra, where periodic damage to the leaf apparatus was recorded.
Indicators of resistance of introduced species to pests and diseases are presented in Table 3. Overall, the results indicate that most introduced taxa were characterized by moderate to high resistance, suggesting the absence of pronounced phytopathological pressure under the environmental conditions of the study area.
Phenological observations revealed considerable variation in the timing of the beginning and end of the growing season among species. Species characterized by earlier cessation of shoot growth generally demonstrated higher levels of winter and drought resistance.
The duration of the growing season in highly resistant species was shorter compared with less adapted introduced taxa, indicating better synchronization of developmental rhythms with the climatic conditions of the region. Phenological characteristics of introduced species, particularly the timing of shoot growth cessation, are presented in Table 4. Earlier completion of shoot growth was typical for species showing increased tolerance to abiotic stresses.
The intensity of flowering and fruiting varied significantly among species. The highest generative capacity was observed in Ulmus pumila, Quercus robur, Acer platanoides, Ribes aureum, and Lonicera tatarica.
In several species, the absence of fruiting despite flowering indicates the influence of unfavorable weather conditions during the flowering period. Indicators of generative capacity of introduced species, expressed through flowering and fruiting intensity, are presented in Table 5. High values of these parameters indicate successful adaptation of the reproductive cycle to the environmental conditions of the semi-desert zone.
The integrated analysis demonstrated significant interspecific variation in the adaptation coefficient (K). The highest values (K > 0.80) were recorded for Ulmus pumila, Quercus robur, Acer tataricum, Betula pendula, and Syringa vulgaris.
Species with lower values of the adaptation coefficient were characterized by increased sensitivity to winter and summer stresses (Table 6).
4. Discussion
The obtained results confirm that the success of introducing tree and shrub species in the semi-desert zone of Western Kazakhstan is determined by the combined influence of abiotic environmental factors and adaptive biological traits of plants. The highest adaptive potential is formed through the combination of drought and temperature tolerance, synchronization of phenological phases with the regional climatic rhythm, and the stability of the generative cycle. Contemporary studies indicate that under conditions of increasing aridization and more frequent extreme weather events, the resilience of tree species depends on the complex interaction of hydraulic, physiological, and morphological mechanisms that determine their ability to withstand water and thermal stress (Allen et al., 2010; Choat et al., 2012; Huang et al., 2016).
High winter hardiness and frost resistance observed in some introduced species are consistent with the fact that, in strongly continental climates, early cessation of shoot growth and timely tissue lignification are critical factors reducing the risk of damage caused by low winter temperatures and spring return frosts (Millar and Stephenson, 2015). In the present study, more resilient species were characterized by earlier termination of shoot growth, reflecting an adaptive phenological strategy. The shortening of the active growth period promotes timely lignification of tissues and reduces the risk of damage from late spring and early autumn temperature fluctuations.
Under conditions of limited water availability, drought tolerance acts as a central environmental filter determining species survival. Physiological mechanisms of resistance to water stress have been extensively described for woody plants and are considered increasingly important under ongoing aridization processes (Jump and Peñuelas, 2005; McDowell et al., 2013). The practical importance of water availability is also confirmed by regional data on afforestation practices: during the cultivation of saxaul in arid regions of Kazakhstan, deep soil preparation significantly increased survival rates and growth intensity compared to untreated soils. In contrast, moisture deficit in plots without soil preparation was associated with the suppression of growth processes as part of adaptive adjustments in plant water balance (Akhmetov et al., 2023).
Similarly, the use of moisture-retaining materials (e.g., hydrogels) and certain growth stimulants has been shown to increase survival rates and improve growth performance in saxaul plantations (Yessimbek et al., 2022).
The obtained results indicate that the most resilient introduced species belong to taxa characterized by pronounced xeromorphic traits and a wide ecological amplitude, which is consistent with the concept of ecological preadaptation (Niinemets and Valladares, 2006).
In most cases, the introduced species demonstrated high or moderate resistance to pests and diseases, which may indicate relatively low phytosanitary pressure in the studied conditions. However, the observed interspecific differences suggest that certain taxa remain vulnerable to local phytophages and pathogens or may establish new biotic interactions within the recipient ecosystems (Richardson et al., 2000). Therefore, even the most adapted species require monitoring of potential biotic risks when plantation areas are expanded.
Phenological synchronization is one of the key mechanisms enabling plants to adapt to extreme environmental conditions and climatic variability (Parmesan, 2006). In the present study, species with shorter vegetation periods and earlier termination of shoot growth demonstrated higher levels of winter and drought tolerance, confirming the functional role of phenology as an integrator of abiotic constraints. Additionally, studies of riparian woody communities indicate that successful regeneration may depend not only on temperature and precipitation but also on the hydrological regime of habitats. In the absence of surface flooding, the natural formation of young tugai forest stands becomes sharply limited, and seed regeneration is strongly associated with the coincidence of sediment accumulation periods and the dispersal of mature seeds. This highlights that the phenology and reproductive success of introduced species in arid regions should be evaluated with consideration of critical windows of favorable weather and hydrological conditions (Dukenov et al., 2023a).
Generative capacity represents one of the key indicators of successful plant introduction, as it reflects the completion of the life cycle under new environmental conditions. High flowering and fruiting intensity indicate favorable adaptation of reproductive processes, whereas cases of flowering without fruit formation may be associated with stressful weather conditions during pollination and fruit set. The importance of reproductive traits in assessing species adaptive potential has been emphasized in studies of plant invasiveness and life-history strategies (Rejmánek and Richardson, 1996), as well as in research on the functioning of woody ecosystems under climatic stress (Reich, 2014; Zhang et al., 2017). From a practical perspective, this implies that species with high values of the integral adaptation coefficient (K) and stable fruiting are preferable for long-term protective plantations, as they provide the potential for population self-maintenance. The use of the integral adaptation coefficient (K) allowed introduced species to be compared based on a combination of traits and reduced the subjectivity of individual scoring assessments. Similar indices are widely used in ecological and forestry studies for ranking species under multifactorial adaptation conditions (Bastin et al., 2019; Brooker et al., 2008). In this study, species with K > 0.80 combined high tolerance to abiotic stresses with more favorable phenological and reproductive characteristics, making them promising candidates for protective and forest reclamation plantations in the semi-desert zone.
From a practical perspective, the obtained results justify a two-level approach:
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preliminary selection of species based on the integral adaptation coefficient (K) with mandatory evaluation of the stability of their generative capacity;
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subsequent optimization of growing conditions, taking into account the soil and climatic characteristics of the site in order to minimize water and temperature stress.
When expanding the range of introduced species, it is necessary to consider their high ecological plasticity and capacity for rapid physiological adaptation under changing climatic conditions. Species characterized by a broad spectrum of functional traits and flexible resource-use strategies often demonstrate increased competitiveness and stress tolerance, which necessitates systematic monitoring when they are introduced into new ecosystems (Wright et al., 2004; Lambers et al., 2008; Pretzsch et al., 2017). Under intensifying climate change, such species may exhibit altered growth and survival dynamics, highlighting the importance of long-term assessments of their ecological behavior.
5. Conclusion
The conducted study provided a comprehensive assessment of the adaptive potential of introduced tree and shrub species under the semi-desert conditions of Western Kazakhstan. The results demonstrate that the success of plant introduction is determined by a combination of factors, including winter hardiness, drought tolerance, resistance to pests and diseases, phenological synchronization, and generative capacity.
The pronounced interspecific variability of these indicators confirms the necessity of a multifactorial evaluation of introduced species under extreme climatic conditions. The application of the integral adaptation coefficient (K) enabled objective ranking of the studied taxa and identification of the most promising species for use in protective, forest reclamation, and urban greening plantations in the region.
The results indicate that species characterized by early termination of shoot growth, high tolerance to abiotic stresses, and stable generative capacity possess the greatest adaptive potential under arid climatic conditions. The integrated approach applied in this study can be recommended for the practical selection of woody and shrub species in regions characterized by strong continental climates and limited water availability.
Data Availability Statement
The entire data set that supports the results of this study will be published in the article itself.
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