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9HETEROGENEOUS VARIABILITY OF CENOTIC CHARACTERISTICS OFTHE FALLOW WITH THE PARTICIPATION OF INVASIVE SPECIES
O.V. Sozinov1*, K.V. Shchukina2**, D.S. Kessel2, N.S. Liksakova2, A.G. Khmarik2
1 Grodno State University named after Yanka Kupala E. Ozheshko, Belarus *e-mail: ledum@list.ru; o.sozinov@grsu.by 2V.L. Komarov Botanical Institute RAS, Russia **e-mail: schukina@binran.ru
Received: 22.08.2022. Corrected: 28.09.2022. Accepted for publication: 03.10.2022.
This study focuses on the results of attendance of changes in the vegetation structure of the fallow after plowing, grass mixture's sowing and measures to root away Heracleum sosnowskyi Manden. over two growing seasons (2021-2022). The work was carried out on the territory of the scientific and experimental station Otradnoye (Leningrad Region) on a three-year fallow and it revealed the formation of a grass-forb meadow. The number of plant species in each season was approximately similar (57-60) with a strong multi-year fluctuation in the occurrence of assectators. The research enables to note a significant decrease in the level of species dominance. In terms of species diversity, vegetation has maintained relative stability. There are 8 species that define 75% of species differences between the communities of the fallow of two seasons, and 2/3 of that groupe of species are cereals. Lupinus polyphyllus Lindl. make a significant contribution to the multi-year differentiation of vegetation (21%). The abundance of Sosnovsky's hogweed tends to decrease (from 2% to 1.3%). The proposed and tested methodological approach for restoration of meadow vegetation on fallows under the conditions of artificial elimination of Heracleum sosnowskyi Manden. has shown its efficiency and perspective of use.
Key words: invasive species, Heraculum sosnowskyi Manden., Lupinus polyphyllus Lindl., fallow, species diversity, Karelian isthmus, Otradnoye
https://dx.doi.org/10.24412/cl-31646-2686-7117-2022-31-234-241
Introduction
There are between 20 and 30 invasive plant species in the Leningrad Region and Karelia (Invasive plants..., 2021; Tkachenko, 2020). Lupinus polyphyllus Lindl. and Heracleum sosnowskyi Manden are among the most active invasive species in Europe. Lupinus polyphyllus is a North American species that has been a "refugee" from culture in Fennoscandia since the 1940s and prefers poor acidic and weakly acidic soils (Vinogradova et al., 2014; Kostina, 2020; Invasive plants..., 2021). Lupinus polyphyllus is an ecotone species (distributed along roadsides, forest edges, fallow lands, ruderal sites, meadows, etc.) and due to its ability to fix nitrogen, large leaf surface area, high shoot density, and seed productivity contributes to changes in plant communities (Meier et al., 2013; Black Book of Flora of Belarus..., 2020). Heracleum sosnowskyi was introduced into cultivation in Russia in the 1940s. The studies were conducted on its selection in the 1950s and 1970s as well as on its characteristics as a forage including at the Otradnoe Research Station of the BIN RAS (Geltman, 2007; Tkachenko, 2014). Heracleum sosnowskyi is a species with high adaptive plasticity, short vegetation period. It prefers well-lit places with fertile
soil and has high seed productivity (Dalke et al., 2018; Invasive plants..., 2021). Heracleum sosnowskyi actively displaces native species and forms new communities (Panasenko et al., 2014; Arepieva, 2022).
The active distribution of Lupinus polyphyllus and Heracleum sosnowskyi in the Northwest, as well as in other regions of Russia, Central and Western Europe, is a consequence of the termination of economic activities on agricultural land, the lack of control and regulation over their condition and the presence of a large number of disturbed habitats (Geltman, 2007; Afonin et al., 2017; Invasive species..., 2020).
Areas removed from economic use undergo successional changes. A complex set of both abiotic and biotic factors influence the rate and sequence. At the same time, habitats with disturbed vegetation cover are most susceptible to invasion by alien species. Their dispersal into natural and semi-natural phytocenoses starts there (Gusev, 2012; Broback, 2015; Hensgen & Wachendorf, 2016; Swierszcz et al., 2017). One of the most important consequences of the introduction of both Heracleum sosnowskyi and Lupinus polyphyllus into plant communities is the impact of these species on the composition and structure of phytocenoses, leading to a significant decrease in autochthonous biodiversity and a change in natural successional processes (Hejda et al., 2009; Konechnaya, Krupkina, 2011; Prass et al., 2022).
This study is aimed at the identification of year-to-year variability of phytocenotic characteristics of the fallow with the participation of invasive plant species for the current evaluation of methodological techniques for meadow vegetation restoration during artificial elimination of Heracleum sosnowskyi.
Material and methods
The studied fallow (about 2 ha) is located on the territory of the Otradnoe Research Station of the BIN RAS (Leningrad Region, Russian Federation; Karelian Isthmus, field center: 60.811986 N, 30.236111 E). In the 1970s-1980s, this site served as an experimental field for cultivation of medicinal plants and other crops. Then, for the past 30 years, the field has not been used and, consequently, has undergone intense expansion of Heracleum sosnowskyi, which was fought with herbicides in 2016-2018. In 2019, there was an experiment of total mechanized treatment of this field (plowing, stone collection, discing, cultivation). In August 2019, cereal grass mixtures were sown: Lolium perenne L. >50%, Poa trivialis L., Phleum pratense L., Festuca sp. (Khmarik & Khmarik, 2021). After these measures, chemical and mechanical control of Heracleum sosnowskyi continued. Monitoring observations in July 2020 showed a predominance of forage grasses in the study area, with Lolium perenne dominating (77%). At the same time, Lupinus polyphyllus dispersed from the southern periphery of the field, forming up to 20% of the projective cover at the time of observations. The abundance of Heracleum sosnowskyi significantly decreased, amounting to 305 specimens (Khmarik & Khmarik, 2021). Also, in the field there was a local appearance of undergrowth of Alunas glutinosa (L.) Gaertn. (h=1.5-2 m) (Fig. 1).
Fig. 1. The fallow with the participation of Heracleum sosnowskyi and Lupinus polyphyllus (July, 2022).
Relatively high density of Heracleum sosnowskyi is preserved in separate loci outside the fallow - on forest edges and glades, where a set of chemical and mechanical control measures against this species is carried out annually.
In the future, the most valuable in terms of biodiversity areas of the scientific experimental station Otradnoe are planned to be given the status of a specially protected natural area. For this purpose, it is necessary to make an inventory of species composition of plant communities and groupings growing there, including participation of invasive species in plant cover formation.
Geobotanical descriptions of meadow vegetation were performed with ruderal and segmental elements in July 2021 on 30 sample plots (SP; 25 m2), relatively evenly distributed throughout the field. In 2022, we repeated the description of sample plots. We recorded species composition and abundance (projective coverage of each species in %). Critical taxa were herbarized with further determination of species affiliation (Tsvelev, 2000). Cenotic classification of the entire dataset was not carried out due to the high mosaicism and variability of vegetation on the fallow. However, it should be noted that in all descriptions, Lupinus polyphyllus dominated during both seasons (Fig. 1).
The Simpson (dominance), Shannon (species diversity), Margalef and Menchinik (species richness), Pielu (alignment), and Berger-Parker (dominance)
indices were calculated based on the abundance of all identified herbaceous plant species (Magurran, 2004).
Analysis of similarities and differences in the entire array of geobotanical descriptions over two growing seasons was performed using Analysis of similarities (ANOSIM) and Permutational multivariate analysis of variance (PERMANOVA) based on the Bray-Curtis distance (Sushko, 2021). The SIMPER test (Similarity Percentage) using the Bray-Curtis similarity measure (Sushko, 2021) was used to estimate the contribution of species to the observed difference in the year-to-year variability of meadow vegetation.
Statistical processing of the geobotanical descriptions was performed using PAST 4.10 (Hammer et al., 2001).
Results and discussion
Analysis of the obtained data on meadow vegetation in the fallow during two vegetation seasons revealed the presence of 78 vascular plant species in the communities. The number of plant species in each season was approximately similar: 57 and 60 species, respectively, but the species commonality was only 47%, indicating a high rate of variability in the species composition during the first stages of secondary succession. The main differences are observed in the composition of sporadically recorded species (assemblages). Indicators of a-diversity of vegetation show varying degrees of year-to-year variability (Table 1). Significant changes occurred in the level of dominance (indices: Simpson and Berger-Parker) in the direction of reducing the role of dominant species in meadow ecosystem functioning, while in terms of species diversity and richness values are relatively stable except for Menchinik index (Table 1).
Table 1. Variability of a-diversity of meadow vegetation on fallow during two vegetative seasons
Index Vegetative season P
2021 2022
Simpson 0.4028 0.2522 0.007*
Shannon 2.279 2.202 0.6727
Pielu 0.1714 0.1507 0.7806
Menchinik 6.076 5.324 0.0007
Margalef 12.51 12.18 0.5186
Berger-Parker 0.6023 0.4016 0.0051
Note: * - p<0.05 values are in bold.
While in 2021, Lupinus polyphyllus was the main dominant species in the cenosis with an average projective cover of 53.8% on the whole field, and the contribution of the most abundant of the other species, Lolium perenne, did not exceed 10%, in 2022, there was a shift towards the polydominant community model (Table 2). At the same time, the coverage of Lupinus polyphyllus remained almost at the same level, but the abundance of cereals: Phleum pratense, Festuca arundinaceae, Elytrigia repens and especially Agrostis tenuis significantly increased. Average projective cover of Agrostis tenuis increased by more than 5 times (from 6.57% to 33.1%).
A pairwise comparison of the geobotanical descriptions of each sample area during two seasons showed a more distinct differentiation of communities by
a-diversity. There were 26.6% of comparison pairs that showed no reliable differences in all indices (p>0.05). There were 20% of descriptions that showed an increase in species diversity and only 7% showed a decrease (by Shannon index, p<0.05). In general, multidirectional, cenotic processes of meadow vegetation formation often take place on the whole area of the fallow complex.
Table 2. Indicator species contributing to the greatest annual differences of meadow communities on fallow (SIMPER test)_
Indicator species Average difference, % Contribution to differences, % Accumulated differences, % Abundance*, %
2021 г. 2022 г.
Agrostis tenuis 12.07 22.95 22.95 6.57 33.1
Lupinus polyphyllus 10.99 20.91 43.86 53.8 51.2
Lolium perenne 3.881 7.384 51.24 9.73 4.48
Festuca arundinaceae 3.843 7.311 58.56 3.73 11.2
Phleum pratense 3.507 6.672 65.23 4.47 12.2
Elytrigia repens 1.919 3.65 68.88 1.17 4.33
Poa trivialis 1.792 3.408 72.29 3.6 2.85
Trifolium hybridum 1.657 3.153 75.44 3.9 1.42
Note: * - projective cover.
Comparison of the species composition of the herbaceous vegetation of the fallow during two years showed reliable differences: by ANOSIM (R=0.3) and PERMANOVA (F=11.9) p=0.0001, which illustrates the rapid cardinal cenotic restructuring of communities at the stage of initial formation.
Seventy-five percent of the difference between the vegetation of the fallow of the two seasons is determined by 8 species, of which 2/3 are cereals (Table 2). There was an increase in participation in of short-rooted grasses forming loose tussocks, which in perspective may lead to formation of loose tussocky meadow, the stage of "maturity" (Fedoruk, 1976). Lupinus polyphyllus and Festuca arundinaceae make a significant contribution to differential vegetation differentiation - 21% and 7.3%, respectively. Heracleum sosnowskyi is in the tenth place by its contribution to the observed cenotic difference and determines only 2.34% of differences (its average abundance decreased from 2% to 1.3%). This indicates the effectiveness of control measures and regulation of the undesirable species in this habitat. Significant correlation between abundance (%/25m2) of Heracleum sosnowskyi and Lupinus polyphyllus was not revealed (p=0.76).
Conclusion
Thus, in the fallow of the Otradnoe experimental station (Leningrad Region, Russian Federation), a cereal-grass meadow begins to form in the second or third year after plowing and using cereal grass mixtures. This is possible with the systemic control of Heracleum sosnowskyi. It is also likely that Lupinus polyphyllus plays an important cenotic role in meadow formation under conditions of artificial elimination of Heracleum sosnowskyi. The extent of its contribution to successional processes in meadows of North-West Russia requires further study. During two vegetation seasons, 78 species of vascular plants were identified in the communities. For the two
years of observations, a low species commonality (about 50%) and a significant difference in the cenotic structure of vegetation were noted. Cereals as well as the adventive species Lupinus polyphyllus play the greatest role in differentiating the phytocenoses of the fallow. A general downward trend in the role of dominant species in community composition was revealed. According to other parameters of species' diversity during two vegetation seasons, the studied communities are relatively stable against the background of various multidirectional cenotic processes in vegetation. The role of Heracleum sosnowskyi, under system control and regulation, is insignificant and tends to decrease with the further prospect of elimination of the species from the cenosis composition.
The proposed and tested methodological approach to the restoration of meadow vegetation on the fallow with artificial elimination of Heracleum sosnowskyi has shown its effectiveness and promising use.
Acknowledgements
This study was performed within the framework of the planned theme of the V.L. Komarov Botanical Institute of the Russian Academy of Sciences "Vegetation of European Russia and Northern Asia: Diversity, Dynamics, Principles of Organization", No. 121032500047-1.
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РАЗНОГОДИЧНАЯ ИЗМЕНЧИВОСТЬ ЦЕНОТИЧЕСКИХ ХАРАКТЕРИСТИК ЗАЛЕЖИ С УЧАСТИЕМ ИНВАЗИВНЫХ ВИДОВ
О.В. Созинов1*, К.В. Щукина2**, Д.С. Кессель2, Н.С. Ликсакова2,
А.Г. Хмарик2
1 Гродненский государственный университет им. Янки Купалы, Республика Беларусь
*e-mail: ledum@list.ru,_ o.sozinov@grsu.by 2Ботанический институт им. В.Л. Комарова РАН, Россия **e-mail: schukina@binran.ru
Приведены результаты наблюдений за изменением строения растительности залежи после вспашки, посева травосмеси и мероприятий по уничтожению Heracleum sosnowskyi за два вегетационных сезона (2021-2022 гг.). Работы осуществлялись на территории научно-опытной станции Отрадное (Ленинградская область) на трехлетней залежи. Выявлено формирование злаково-разнотравного луга. Количество видов растений в каждом сезоне было примерно сходным (57-60) при сильной разногодичной флуктуации встречаемости ассектаторов. Отмечено значимое уменьшение уровня доминирования видов. По показателям видового разнообразия растительность сохранила относительную стабильность. 75% видовых различий между сообществами залежи двух сезонов определяет 8 видов, из которых 2/3 - злаки. Значимый вклад в разногодичную дифференциацию растительности дает Lupinus polyphyllus (21%). Heracleum sosnowskyi имеет тенденцию к снижению обилия (с 2% до 1.3%). Предложенный и апробированный методический подход по восстановлению луговой растительности на залежи при искусственной элиминации Heracleum sosnowskyi показал свою эффективность и перспективность использования.
Ключевые слова: инвазивные виды, Heracleum sosnowskyi Manden., Lupinus polyphyllus Lindl., залежь, видовое разнообразие, Карельский перешеек, Отрадное
