Research Article |
Corresponding author: Zigmantas Gudžinskas ( zigmantas.gudzinskas@gamtc.lt ) Academic editor: Franz Essl
© 2020 Zigmantas Gudžinskas, Lukas Petrulaitis, Egidijus Žalneravičius.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gudžinskas Z, Petrulaitis L, Žalneravičius E (2020) Emerging invasion threat of the liana Celastrus orbiculatus (Celastraceae) in Europe. NeoBiota 56: 1-25. https://doi.org/10.3897/neobiota.56.34261
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The woody vine Celastrus orbiculatus (Celastraceae), Oriental bittersweet, is an alien species that recently has been found to be spreading in Europe. Many aspects of its biology and ecology are still obscure. This study evaluates the distribution and habitats, as well as size and age of stands of C. orbiculatus in Lithuania. We investigated whether meteorological factors affect radial stem increments and determined seedling recruitment in order to judge the plant’s potential for further spread in Europe. We studied the flower gender of C. orbiculatus in four populations in Lithuania and found that all sampled individuals were monoecious, although with dominant either functionally female or male flowers. Dendrochronological methods enabled us to reveal the approximate time of the first establishment of populations of C. orbiculatus in Lithuania. The youngest recorded individual with fruits was determined to be 10 years old. Analysis of radial increments revealed no reliable correlations with meteorological conditions. Therefore, we conclude that climatic conditions in the region are favourable for the growth, reproduction, and invasion of this species. C. orbiculatus produces viable seeds, successfully reproduces and spreads within and around the established stands. The presence of seedlings and two- to four-year-old saplings in the population confirms constant generative recruitment. Available information on the distribution of C. orbiculatus in Europe revealed its existence in 13 countries. In total, 58 occurrences of this species have been recorded in Europe so far. We consider that the lag period lasted until 2005 and that the exponential population growth phase has now set in. In Lithuania, the invaded area is quite small (0.51 ha); however, the total estimated invaded area in Europe could be about 250 ha. At the current stage of invasion and distribution in Europe, measures for control, management, and eradication of C. orbiculatus have a chance of being effective and economically feasible.
dendrochronology, dioecious plants, distribution, flower gender, habitats, reproduction, seedlings
Lianas are strong competitors with trees for above- and belowground resources (
Implementing the European Union Regulation 1143/2014 on invasive alien species, a horizon scanning for potentially invasive alien species has revealed 98 species that pose particular danger in the EU (
Celastrus orbiculatus was introduced to Europe as an ornamental plant at around 1860. Soon after its introduction, in 1863, it was first offered for sale to the public as an ornamental plant in Siebold’s nursery (The Netherlands) catalogue (Del Tredici 2014). According to
Celastrus orbiculatus was introduced to the United States from Asia in the late 19th century, and has become naturalised in the Eastern States where it occupies large areas of disturbed temperate forests, alluvial woods, and roadsides (
Quite recently a risk assessment of C. orbiculatus for the EU, with special focus on the Netherlands, has been performed (
Despite the increased number of studies on C. orbiculatus, many of its biological and ecological properties, which have an important effect on the spread and emerging invasion, were insufficiently known in Europe. Therefore, we started to investigate established populations of C. orbiculatus in Lithuania, which are at the north-eastern limit of current naturalisation of this species in the southern part of the boreal biogeographic region of Europe. The aims of this study were (a) to analyse the current distribution of C. orbiculatus in Lithuania and Europe, (b) to evaluate the size of C. orbiculatus populations and occupied habitats in Lithuania, (c) to assess allocation of flowers by the gender; (d) to evaluate the age of individuals and approximate time of population establishment, (e) to study radial increment rates and possible relationships with meteorological conditions, (f) to investigate generative reproduction and sapling density in an invaded habitat. The results of this study provide information for a better understanding of C. orbiculatus reproduction, spread and invasion to natural, seminatural and human-made habitats.
Celastrus orbiculatus Thunb. (Celastraceae R. Br.) is a deciduous woody vine that climbs by means of twining about a support. Cane-forming stems are located just above the ground and liana-forming stems are in the canopy layer where it climbs through the tree trunk and branches. (
A distribution map of C. orbiculatus in Lithuania was compiled applying a system of grid cells, which were arranged according to geographical coordinates with sides of 6' of latitude and 10' of longitude (
We performed field studies on four populations of C. orbiculatus in Lithuania during the growth season of 2016 and flower gender was studied in June of 2018. We also surveyed a newly discovered population in Vandžiai (Raseiniai distr.) in September of 2018. Therefore, population size and habitats of C. orbiculatus were evaluated in five sites in Lithuania (Table
Geographical characteristics of the studied Celastrus orbiculatus sites in Lithuania.
Site | Latitude (N) / Longitude (E) | Altitude (m a.s.l.) |
Paneriai (Vilnius city) | 54.6344, 25.1526 | 150 |
Visoriai (Vilnius city) | 54.7403, 25.2606 | 174 |
Babrungėnai (Plungė distr.) | 55.9986, 21.8699 | 158 |
Palanga city | 55.9591, 21.0915 | 10 |
Vandžiai (Raseiniai distr.) | 55.3017, 23.4276 | 58 |
Population size and habitat characteristics
The areas of small populations were measured using measurement tape, whereas the size of large populations with complicated configuration was measured by applying geographical coordinates established at certain points of the stand perimeter. The area occupied by C. orbiculatus was calculated using online software provided from the Spatial Information Portal of Lithuania (www.geoportal.lt). The quantity of fruits was estimated visually. When solitary fruits were found on the lateral branches of a C. orbiculatus individual, its fruit yield was classified as poor, whereas for individuals having one or more fruits on most of their axillary cymes on a lateral branch, its fruit yield was classified as abundant.
The height of trees in the habitats and height of C. orbiculatus in the trees were measured with a height measurement device (Haglöf EC II, Sweden). Coverage of plants (in per cent) of different vegetation layers was estimated visually. We identified habitat types following the Interpretation Manual of European Union Habitats (
Flower gender of Celastrus orbiculatus was studied in four populations (Babrungėnai, Palanga, Paneriai and Visoriai) at the beginning of June of 2018. We sampled from two to five mature and clearly identifiable separate individuals from each population, 12 individuals in total. Ten lateral branches (usually 15–20 cm long) at 1–3 m high from the ground with several inflorescences were taken from each plant, labelled and placed into separate plastic bags and brought to the laboratory for further analysis. At the laboratory, the number of inflorescences and number of flowers in each inflorescence was counted starting from the base of branch and inflorescence. Each flower was studied under the binocular microscope and dissected, when necessary, to estimate its gender. Flowers with developed anthers and undeveloped pistil and ovary were treated as functionally male, flowers with developed pistil and ovary but undeveloped anthers were treated as functionally female flowers (Figure
Flowers of Celastrus orbiculatus. A Functionally female flower with developed pistil (a) and staminodes (b) B longitudinal section of functionally female flower (petals removed) with developed pistil (a), staminodes (b) and ovules in the ovary (c) C functionally male flower with developed stamens (d) D longitudinal section of functionally male flower with filaments of developed stamens (e), pistillode (undeveloped pistil, f) and ovary with aborted ovules (g). Scale bars: 1 mm (A–C); 2 mm (D).
The density of C. orbiculatus seedlings and saplings was estimated in three parallel 20-meter-long transects in Paneriai Forest in early September 2016. The transects were at a distance of approx. 30 m between the previous transect. Transects were selected in an area free of ground cover of C. orbiculatus shoots. The first transect was arranged in the central part of the C. orbiculatus stand with the densest and highest mature individuals, the second transect was selected at the edge of the stand with mature individuals, and the third transect was in an area free of mature C. orbiculatus individuals. The transects were divided into 20 sampling plots of 1 m2 using a frame with all sides of 1 m. The number of seedlings and saplings grown from seeds was counted. Individuals grown from seeds in the year of the study were ascribed to the group of seedlings, whereas elder individuals (two or more years old) were identified as saplings. We distinguished seedlings from saplings by the level of stem lignification. Saplings and individuals grown from underground shoots were distinguished by the shape of their leaves and the character of the shoot. In each transect, we also measured the height of 10 randomly selected seedlings and 10 saplings.
Stems of C. orbiculatus for dendrochronological analysis were sampled in three populations (Babrungėnai, Visoriai and Paneriai) at the end of the growth period in September 2016. The population of Palanga was not sampled the same way because of an insufficient number of primary stems. Two series of samples were collected in the population of Paneriai Forest: the first series was collected in the - presumably - oldest part of the population, another series in the peripheral part of the stand, at 40–50 m from the first sampling site. Ten of the largest individuals from each sampling area were selected and 15–20-cm-long stem sections were cut with a saw at ground level. Samples were numbered, diameters measured with a caliper and placed into paper bags for drying at ambient temperature for three months.
Before counting annual rings, sections of ca. 5 cm were cut from each dried sample of the stem and one of its surfaces sanded and polished. Polished surfaces of the samples were stained with original stain prepared from the powder of Curcuma rhizomes boiled in vegetable oil (5 g of powder and 20 ml of vegetable oil). Prepared cross-sections of Celastrus orbiculatus stems were examined under a binocular microscope (LEICA EZ4), the annual rings of the xylem were counted and the width of annual rings was measured with an accuracy of 0.1 mm.
Information on the monthly minimum temperatures and the amount of precipitation in the period from 1989 to 2016 was provided by the Hydrometeorological Service under the Ministry of Environment of Lithuania. The data was used to evaluate the impact of meteorological factors on the radial stem growth of C. orbiculatus.
The normality of data distribution was evaluated using the Shapiro-Wilk test. Comparison of normally distributed data sets (pooled seedling and sapling density, width of annual rings) was performed applying ANOVA several-sample F-test and Tukey’s post-hoc pairwise comparisons. Two normally distributed data sets of seedling and sapling height were compared applying Student’s t-test. Because seedling and sapling density in the studied plots were distributed non-normally, comparison of their densities was analysed applying non-parametric Kruskal-Wallis H-test and Mann-Whitney post-hoc pairwise comparisons. Correlations between normally distributed data sets of annual ring width and meteorological parameters were calculated applying Pearson’s rank correlation test. The significance level of statistical tests was set at p < 0.05. Dependence of stem age and its diameter were evaluated using linear bivariate model. Descriptive statistical analysis results include mean values and standard deviations (mean ± SD), in analyses of seedling and sapling density also including minimum, maximum and median. All calculations were performed using PAST 3.20 (
Currently, C. orbiculatus occurs in six localities in southeastern (Paneriai and Visoriai, Vilnius city), central (Vandžiai, Raseiniai distr. and Girionys, Kaunas distr.) and western (Babrungėnai, Plungė distr. and environs of Palanga city) parts of Lithuania (Figure
Information on the distribution of C. orbiculatus in Europe was quite dispersed over numerous publications and other sources of information. According to these sources C. orbiculatus has been recorded in 13 European countries: Austria (
Analysis of the spread dynamics of registered C. orbiculatus populations in Europe revealed very slow increase of sites of occurrence (on average three new sites per 10 years) from its first report in Europe in 1953 (
Distribution of Celastrus orbiculatus in Europe mapped on to the grid of the Atlas Florae Europaeae.
Four of the studied populations (Paneriai, Visoriai, Babrungėnai and Vandžiai) were in forest habitats and one population (Palanga) on the edge of a forest. In Paneriai Forest, the plants occupied an approximately 70-year-old Pinus sylvestris stand with admixture of Acer platanoides and Quercus robur in the second tree layer (Table
Character | Site | ||||
---|---|---|---|---|---|
Paneriai | Visoriai | Babrungėnai | Palanga | Vandžiai | |
Area occupied by the stand of Celastrus orbiculatus (m2) | 3640 | 480 | 90 | 20 | 880 |
Age of the dominant trees (years) | 70 | 60 | 50 | – | 20 |
Maximum height of the tree layer (m) | 24 | 17 | 15 | – | 15 |
Coverage of the tree layer (%) | 60 | 50 | 50 | – | 40 |
Coverage of the shrub layer (%) | 60 | 30 | 60 | 20 | 50 |
Coverage of the herb layer (%) | 40 | 30 | 30 | 60 | 20 |
Coverage of the bryophyte layer (%) | 60 | 10 | 5 | 20 | 5 |
Maximum height of Celastrus orbiculatus (m) | 18 | 14 | 10 | 3 | 14 |
Coverage of Celastrus orbiculatus in the tree layer (%) | 15 | 20 | 10 | – | 20 |
Coverage of Celastrus orbiculatus in the shrub layer (%) | 50 | 40 | 30 | 10 | 60 |
Coverage of Celastrus orbiculatus in the herb layer (%) | 60 | 30 | 60 | 50 | 60 |
Total coverage of Celastrus orbiculatus in all layers (%) | 70 | 60 | 70 | 50 | 70 |
The studied forest habitats invaded by C. orbiculatus had characteristic vertical structure of the tree and shrub layers and their cover (excluding C. orbiculatus) ranged from 40% to 60% and from 30% to 60%, respectively (Table
In Lithuania, C. orbiculatus occurs in quite different types of habitats. The population in the environs of Palanga occupies a transitional area between a habitat of wooded dunes (2180 Wooded dunes of the Atlantic, Continental and Boreal region) and sand grasslands (6120* Xeric sand calcareous grasslands). In Paneriai, this species occupies mature pine forest (9010* Western Taiga) and the transitional zone to spruce forest (9050 Fennoscandian herb-rich forests with Picea abies). In Visoriai, C. orbiculatus grows in stadial forest dominated by Betula pendula with an admixture of Picea abies, which according to the ecological conditions and species composition is close to the herb-rich spruce forest habitat (9050 Fennoscandian herb-rich forests with Picea abies). The stand of C. orbiculatus in Vandžiai invaded a young Betula pendula stand situated in the transitional zone between alluvial forest (91E0 Alluvial forests with Alnus glutinosa and Fraxinus excelsior) and broad-leaved forest (9020* Fennoscandian hemiboreal natural old broad-leaved deciduous forests).
In Paneriai Forest, a dense stand of this species with mature individuals occupies an area of 2600 m2. The total area of the stand, including recorded seedlings and saplings, comprises 3640 m2. At the Vandžiai site (Raseiniai distr., Central Lithuania) the species occupies 880 m2 while the other studied stands of C. orbiculatus were significantly smaller (Table
In all studied sites, C. orbiculatus shoots were distributed over all vegetation layers and in all cases it was the dominant species in the entire community. Its total coverage ranged from 50% to 70% (Table
Analysis of flower (n = 1913) gender of 12 individuals revealed that all plants were monoecious, whereas dioecious or polygamo-dioecious individuals were not found. In all studied individuals either functionally female or male flowers prevailed, from 74.2% to 84.3% and from 76.5% to 84.9%, respectively (Table
We noted that functionally female flowers of individuals with prevailing male flowers were usually arranged in the proximal part of lateral branches, at the first to the third node, and in the proximal part of the inflorescence. In individuals with prevailing female flowers, functionally male flowers were usually arranged in inflorescences at the distal part of a branch or at the apex of the inflorescence.
The existence of monoecious individuals in all studied sites explains the pattern of C. orbiculatus fructification. At the Babrungėnai and Visoriai sites, the set of C. orbiculatus fruits in 2016 was poor because dioecious individuals bearing only dominant male flowers were recorded. In Palanga and in Paneriai, some individuals produced abundant fruits in 2016. In both these sites individuals with dominant functionally female and dominant functionally male flowers were registered.
Distribution of flowers by gender in the studied individuals of Celastrus orbiculatus from four populations in 2018.
Site | Number of individuals | Number of studied flowers | Prevailing gender of flowers | Gender of flowers | |||
---|---|---|---|---|---|---|---|
female | male | ||||||
number | % | number | % | ||||
Paneriai | 1 | 138 | male | 25 | 18.1 | 113 | 81.9 |
2 | 147 | female | 116 | 78.9 | 31 | 21.1 | |
3 | 124 | female | 92 | 74.2 | 32 | 25.8 | |
4 | 161 | female | 135 | 83.9 | 26 | 16.1 | |
5 | 235 | female | 198 | 84.3 | 37 | 15.7 | |
Visoriai | 1 | 156 | male | 27 | 17.3 | 129 | 82.7 |
2 | 131 | male | 23 | 17.6 | 108 | 82.4 | |
3 | 166 | male | 39 | 23.5 | 127 | 76.5 | |
Babrungėnai | 1 | 152 | male | 30 | 19.7 | 122 | 80.3 |
2 | 126 | male | 19 | 15.1 | 107 | 84.9 | |
Palanga | 1 | 170 | male | 32 | 18.8 | 138 | 81.2 |
2 | 207 | female | 171 | 82.6 | 26 | 17.4 |
In all sampling plots of the central part of the C. orbiculatus stand, we recorded 162 seedlings and saplings, and their mean density was 8.10 ± 1.94 individuals/m2 (Table
Analysis of the proportions of seedlings and saplings revealed that in the central part of the stand saplings prevailed (55.55%), whereas on the periphery and outside the main stand seedlings prevailed (54.10% and 52.63%, respectively). Density of seedlings and saplings in the centre of the stand was statistically significantly higher than in the other two transects (Table
Two- and three-year-old saplings were almost equally presented in all transects. Four-year-old saplings were recorded in the central part of the stand only. The height of the measured seedlings ranged from 4 cm to 10 cm. Mean height of seedlings was 6.70 ± 1.58 cm. The height of saplings ranged from 9 cm to 19 cm and their mean height was 13.03 ± 2.70 cm. Saplings were significantly taller than seedlings (Student’s t-test, t = 11.07, n = 60, p < 0.001). Two four-year-old saplings were 18 cm and 19 cm in height.
Total number of Celastrus orbiculatus seedlings and saplings in the studied transects (Paneriai Forest, Vilnius), their density (mean ± SD), minimum and maximum number in sampling plot and median. The same letter indicates statistically significant differences according to Tukey’s pairwise comparison (p < 0.001).
Transect location | Sampling plots (n) | Total number | Density (individuals/m2) | Minimum | Maximum | Median |
---|---|---|---|---|---|---|
Centre | 20 | 162 | 8.10 ± 1.94ab | 5 | 12 | 8 |
Periphery | 20 | 61 | 3.05 ± 1.50a | 0 | 6 | 3 |
Outside | 20 | 38 | 1.90 ± 1.41b | 0 | 5 | 2 |
Number, percentage and density (mean ± SD, individuals/m2) of Celastrus orbiculatus seedlings and saplings in three transects in Paneriai Forest (Vilnius). The same letter indicates statistically significant differences according to Mann-Whitney pairwise comparison (p < 0.001).
Transect location | Seedlings | Saplings | ||||
---|---|---|---|---|---|---|
Number | % | Density | Number | % | Density | |
Centre | 72 | 44.45 | 3.60 ± 1.67ab | 90 | 55.55 | 4.50 ± 1.15ab |
Periphery | 33 | 54.10 | 1.65 ± 1.26a | 28 | 45.90 | 1.4 ± 0.88a |
Outside | 20 | 52.63 | 1.00 ± 0.97b | 18 | 47.37 | 0.90 ± 0.91b |
According to the number of annual rings, the oldest analysed population of C. orbiculatus was that at Visoriai, in which the oldest recorded living stem was 21 years old; the oldest dead stem was 30 years old, and mean age of sampled stems was 11.9 ± 4.7 years (Table
Age and stem diameter of studied Celastrus orbiculatus stems (n = 10 in each stand) and estimated year of the stand initiation.
Features | Visoriai | Paneriai | Babrungėnai | |
---|---|---|---|---|
Centre | Periphery | |||
Age of the oldest living stem (years) | 21 | 18 | 11 | 11 |
Age of the youngest stem (years) | 7 | 11 | 6 | 3 |
Age of the oldest dead stem (years) | 30 | – | – | – |
Estimated year of the stand initiation | 1987 | 1999 | 2006 | 2006 |
Diameter of the oldest living stem (mm) | 47.5 | 42.1 | 18.0 | 15.7 |
Mean age of stems (years) | 11.9 ± 4.7 | 13.2 ± 2.3 | 9.6 ± 1.9 | 6.9 ± 3.0 |
Mean diameter of stems (mm) | 20.4 ± 12.8 | 25.8 ± 8.7 | 14.6 ± 2.7 | 11.99 ± 2.4 |
The diameter of sampled stems of C. orbiculatus was strongly correlated with their age (R2 = 0.73, p < 0.001, n = 40). However, analysis of same age stem diameters revealed a significant variation. The diameter of 11-year-old stems (n = 10), comprising the largest age group among the sampled stems, ranged from 11.2 mm to 25.4 mm (mean 17.7 ± 3.9 mm). Variation of diameter of same aged stems in the same habitat suggests a significant effect of the stem position and competition with other stems, height of foliage, light availability, etc.
The mean width of annual rings (n = 376) of the xylem of C. orbiculatus of all studied stems (n = 40) was 0.77 ± 0.26 mm. The minimum width of annual rings in the studied plants was 0.3 mm, the maximum width was 1.7 mm, and the median width of annual rings was 0.7 mm. We compared the width of annual rings formed from 2006–2016 and found no statistically significant differences among years (F (10, 99) = 0.56, p = 0.84). Also, differences could not be detected by pairwise comparison between years. However, we found statistically significant differences among annual ring width of the same age (11-year-old) stems (F (9, 100) = 16.93, p < 0.001). Pairwise comparison also revealed significant differences between annual ring width in stems both from the same site and from different sites.
Results from the analysis of the relationships of annual ring width and radial increment of the stem diameter with meteorological factors were quite unexpected. No statistically reliable correlations were found between annual ring widths and mean annual temperature (r = 0.16, p = 0.64), mean winter temperature (r = –0.01, p = 0.84), mean summer temperature (r = 0.21, p = 0.53), minimum winter temperature (r = –0.10; p = 0.43), and annual precipitation (r = 0.12; p = 0.72).
The currently recorded localities of C. orbiculatus in Lithuania are in different regions of the country and separated by distances ranging from 10 km to 130 km. Therefore, we suppose that escaped populations of C. orbiculatus have originated from different sources. Considering the common cultivation of this species in gardens and for landscaping in Lithuania (
The map of its current distribution (Figure
In its native range in East Asia, C. orbiculatus grows mainly in mixed forests, along forest margins and on grassy slopes (
In Europe, C. orbiculatus most frequently has been found in areas close to urban environments and occupying human-made or disturbed habitats; however, in Austria, Germany and Poland this species has been found occurring in natural or seminatural habitats located at significant distances from urban areas (
Celastrus orbiculatus affects all layers of the vegetation in the invaded habitats. Its coverage in the tree canopy layer was quite low in Lithuania, although it substantially reduces light availability in the lower vegetation layers. We detected the highest coverage of Celastrus orbiculatus in the shrub and herb layers. Even higher total coverage, ranging from 80% to 100%, of this species has been recorded in Hessen, Germany (
Registered populations of C. orbiculatus currently occupy an area of 0.51 ha in Lithuania. However, information on the area occupied by this species in other European countries is incomplete. Probably the largest area occupied by C. orbiculatus is in Poland, Lubuskie province. There, about 170 ha in the environs of World War II fortifications are occupied by C. orbiculatus with very dense stands on about 9 ha (
Individuals of C. orbiculatus are known to be either functionally dioecious (
The available information on the generative reproduction of C. orbiculatus in Europe is controversial.
The existence of monoecious individuals of C. orbiculatus in Lithuania supports the assumption that monoecious or polygamo-dioecious plants in other populations in Europe also exist. Thus, when monoecious or polygamo-dioecious individuals occur in an area, there are no obstacles for their sexual reproduction (
Celastrus orbiculatus does not form a persistent seedbank; almost all seeds germinate in spring and the light intensity does not affect their germination (
Our research revealed significant densities of C. orbiculatus seedlings and saplings within limits of its dense stand with mature individuals, on the periphery and in proximity of the stand without mature individuals. The quite slow growth rate during the first three to four years, as revealed by our study, confirms that saplings form so-called seedling banks (
There is insufficient information regarding the time point when individuals of C. orbiculatus grown from seeds reach maturity.
Propagule pressure is among the most significant factors affecting the spread and invasiveness of plant species (
The dendrochronological methods employed in this study enabled us to determine the approximate time of establishment of populations of C. orbiculatus and the age of their generative maturity.
Studies on the age structure of C. orbiculatus in the area of the Hudson River Estuary (New York State, USA) revealed that the oldest individuals were about 20 years old and their diameter was up to 70 mm, while the mean stem diameter of 5–7-year-old individuals was 17.5 mm (
We found that the oldest living stem in the studied populations in Lithuania was 21 years old, and a dead stem (most probably quite recently strangled by other stems of C. orbiculatus) was 30 years old. The diameter of the oldest living stem was 47.5 mm. Significant differences in stem diameters in the area of the Hudson River Estuary, USA (
We analysed the effect of the meteorological factors in Lithuania on the radial growth of C. orbiculatus stems and found no specific relationships. The absence of reliable correlations between the width of annual rings and the mean annual temperature, mean winter temperatures, mean summer temperatures, minimum winter temperatures and annual precipitation suggests that climatic conditions in Lithuania are optimal for this species. Surprisingly, even prolonged very cold winter temperatures below –25 °C (in 2003, 2006, 2010, 2012) and occasionally below –30 °C (in 1997) did not damage C. orbiculatus shoots significantly to inhibit radial increments in the following growth period. The sites of the three largest and oldest populations studied in Lithuania fall in the first hardiness zone (H1) (
Celastrus orbiculatus is spreading in Europe, although its range is still restricted. Climatic conditions are suitable for this species in the Atlantic and continental biogeographic regions, as well as in the southern part of the boreal biogeographic region of Europe. It invades a wide range of forest habitats and poses a threat to dry grasslands, dunes, forest fringe communities of high conservation value, as well as diverse anthropogenic habitats.
Celastrus orbiculatus, at least in Lithuania, but probably also in other European countries, is represented not only by dioecious, but also by monoecious individuals. Thus, even in cases of the occurrence of solitary individuals or small groups of monoecious plants, it produces viable seeds and successfully reproduces sexually. Considering its current distribution and the quite frequent cultivation of C. orbiculatus in gardens, most likely it will steadily spread into new areas. Nevertheless, it is still possible to stop the spread of this liana and to reduce the risk of its further invasion.
The effectiveness of control and eradication measures of invasive plant populations is highest when the occupied area is rather small (
Increasing public awareness about the threat of C. orbiculatus invasion is among the most important tasks aiming to reduce the possibility of its spread from gardens. Further surveillance of exiting stands of C. orbiculatus and search for invaded but still overlooked areas should be performed involving citizens, amateur botanists, and specialists.
We are grateful to the Committee for Mapping the Flora of Europe (Helsinki University, Finland) for kind permission to use their software and base-map for preparing our distribution map. We also express our gratitude to Prof. Sergei L. Mosyakin (M.H. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, Kiev) for providing unpublished information on species distribution in Ukraine and to Prof. Dr. Klaus Adolphi (Rossbach, Germany) for supplying published information on C. orbiculatus in Germany. We also thank the reviewers Dr. Swen Follak and Dr. Barbara Tokarska-Guzik for valuable comments and suggestions. Special thanks go to Theodor C.H. Cole (FU Berlin) for English-language editing and for valuable comments and suggestions.