Isolation of habitat patches limits colonisation by moorland Hemiptera

There has been a significant loss in the UK, and elsewhere in Europe, of moorland dominated by heather Calluna vulgaris. One response to this has been the initiation of moorland vegetation restoration projects. Heather moorland has distinct assemblages of invertebrates and the ability of these to colonise newly created habitat patches has not previously been experimentally tested. We established a dispersal and colonisation experiment by transplanting C. vulgaris-dominated turfs within grassland at different distances (up to 40 m) from heather moorland vegetation. Hemiptera were cleared from these turfs by the use of an insecticide and were sampled 1 year later to investigate re-colonisation rates. Hemiptera assemblages on transplanted turfs were most dissimilar to those of heather moor at the greatest distances of these turfs from heather moor. Colonisation rates of heathland-indicator Hemiptera declined exponentially with distance. The number of individual heathland-indicator Hemiptera was higher on turfs 5 and 10 m from heather moor than on turfs transplanted back into the heather moor, possibly due to a crowding effect. Our findings indicate that moorland Hemiptera assemblages may be limited by dispersal ability. We recommend that moorland restoration schemes should be prioritised on ground as close as possible to existing heather moors.     

Intra-specific variation in the inhibitory effects of Thymus vulgaris (Labiatae) monoterpenes on seed germination

The ecological role of plant secondary compounds has received a great deal of attention yet little is known regarding variation in the ecological significance of different compounds produced by a single species. This is particularly pertinent to species where variation in compound presence is under genetic control. In this study we have quantified variation in the inhibitory effects of the six dominant monoterpene oils produced by different genotypes (or chemotypes) of Thymus vulgaris on achene germination of an associated species (Brachypodium phoenicoides), and seeds of T. vulgaris itself, in controlled experimental conditions. The experiments involved the germination of seeds of the two species with crushed leaves or pure essences of the six chemotypes, i.e. four experimental trials. A significant inhibitory effect against B. phoenicoides was found for both crushed leaves and the range of concentrations of pure oils used in the trial. The two phenolic compounds had a greater effect than the non-phenolic leaves and pure oils. The significant differences between non-phenols and the controls declined faster over time for crushed leaves and low concentrations of the pure essences than did that between the phenolics and the controls. There was a significant inhibitory effect against the germination of T. vulgaris seeds for the crushed leaves and the pure oils, although the contrast between the non-phenols and the phenols was not significant. The effect of the different oils varied depending on the seed source of the thyme seeds used in the trial. This is suggested to be due to differences in the rates of outcrossing among the sampled populations. The effects of the oils on the associated species, B. phoenicoides, were not found to be greater than their effects on the germination of thyme seeds. The results are discussed in the context of a potential role for such inhibitory effects in relation to the onset of rainfall and suitable germination conditions.     

The effect of nutrient supply and light intensity on tannins and mycorrhizal colonisation in Dutch heathland ecosystems

(1) Increased atmospheric nitrogen deposition has shifted plant dominance from ericaceous plants to grass species. To elucidate the reduced competitiveness of heather, we tested the hypothesis that additions of nitrogen reduce the concentrations of phenolics and condensed tannins in ericaceous leaves and retard mycorrhizal colonisation in ericaceous plants. We also tested the negative effects of reduced light intensity on carbon-based secondary compounds and mycorrhizal colonisation in ericaceous plants. (2) We performed a field inventory at three heathland sites in the Netherlands varying in nutrient supply and light intensity. Leaves of ericaceous plants and grasses were collected and analysed for concentrations of tannins, phenolics and nutrients. Similarly, we took root samples to record mycorrhizal colonisation and soil samples to measure the soil mineralisation. In addition, we conducted two-factorial experiments with Calluna vulgaris plants, in which we varied fertiliser and shade levels under greenhouse and field conditions. (3) The field inventory revealed that nitrogen addition and shading both negatively affected the concentration of total phenolics. The total phenolics and condensed tannin concentrations were positively correlated (P < 0.001), but in the field experiment, the condensed tannins were not significantly affected by the treatments. Our results provide the first evidence that the carbon nutrient balance can be used to predict the amount of total phenolics in the dwarf shrub C. vulgaris. (4) In the field experiments, shading of plants resulted in significantly less mycorrhizal colonisation. Only in the greenhouse experiment did addition of nitrogen negatively affect mycorrhizal colonisation. (5) Our results imply that increased atmospheric nitrogen deposition can depress the tannin concentrations in ericaceous plants and the mycorrhizal colonisation in roots, thereby reducing the plants’ competitiveness with respect to grasses. Additionally, if ericaceous plants are shaded by grasses that have become dominant due to increased nitrogen supply, these effects will be intensified and competitive replacement will be accelerated.     

A global change-induced biome shift in the Montseny mountains (NE Spain)

Shifts in plant species and biome distribution in response to warming have been described in past climate changes. However, reported evidence of such shifts under current climate change is still scarce. By comparing current and 1945 vegetation distribution in the Montseny mountains (Catalonia, NE Spain), we report here a progressive replacement of cold‐temperate ecosystems by Mediterranean ecosystems. Beech (Fagus sylvatica) forest has shifted altitudinally upwards by ca. 70 m at the highest altitudes (1600–1700 m). Both the beech forests and the heather (Calluna vulgaris) heathlands are being replaced by holm oak (Quercus ilex) forest at medium altitudes (800–1400 m). This beech replacement has been observed to occur through a progressive isolation and degradation of beech stands. In ‘isolated’ (small and surrounded by holm oaks) beech stands, beech trees are 30% more defoliated, beech recruitment is 41% lower, and holm oak recruitment is three times higher than in ‘continental’ (large and continuous) beech stands. The progressively warmer conditions, complemented by the land use changes (mainly the cessation of traditional land management) are the apparent causes, providing a paradigmatic example of global change affecting distributions of plant species and biomes.     

Phytochemical Analysis and Total Antioxidant Capacity of Rhizome,Above‐Ground Vegetative Parts and Flower of Three Iris Species

This study was aimed at investigating the phytochemical composition and antioxidant capacity of rhizomes, above‐ground vegetative parts and flowers of three Iris species: Iris humilis Georgi , Iris pumila L. and Iris variegata L. UHPLC‐Orbitrap MS analysis was used for determination of phytochemical profile. Total pigments, phenolics, flavonoids, soluble sugars and starch content as well as ABTS antioxidant capacity were also determined. In total, 52 phenolics compounds were identified with 9 compounds (derivatives of iriflophenone, apigenin C‐glycosides, luteolin O‐glycoside, isoflavones derivatives of iristectorigenin, dichotomitin, nigracin and irilone) never reported before in Iris spp. Differences in phenolic composition profile, pigments, soluble sugar, starch, total phenolics and flavonoids content and total antioxidant capacity were found among Iris species and different part of plants. Significant correlation between total phenolic content and antioxidant capacity was determined. The obtained results are comparable with those obtained for medical plants. These findings could be useful for fingerprinting characterization of Iris species and estimation of possible use in pharmaceutical industries.     

Chloroplast DNA and isozyme evidence on the evolution ofSenecio vulgaris (Asteraceae)

Chloroplast DNA (cpDNA) and isozyme variation were analyzed over a range of populations of two infraspecific taxa of the tetraploidSenecio vulgaris. The isozyme data were supportive of the hypothesis that the weedy and cosmopolitanS. vulgaris var.vulgaris is an evolutionary derivative ofS. vulgaris subsp.denticulatus from the coasts of W Europe and montane altitudes in S Spain and Sicily. The two taxa exhibited a very high genetic identity with subsp.denticulatus containing slightly more isozyme diversity than was found in var.vulgaris. — Three cpDNA haplotypes (A, B, C) already known from other Mediterranean diploid species ofSenecio were resolved in var.vulgaris, and an additional fourth haplotype (E) was found in subsp.denticulatus. Two alternative hypotheses were chosen to account for the origin and maintenance of the observed cpDNA composition ofS. vulgaris. It either reflects (1) the retention of an ancestral polymorphism which stems from the recurrent and polytopic formation of ancestral tetraploid lineages; or (2)S. vulgaris originally was characterized by haplotype E, and haplotypes A, B and C were acquired through repeated introgressive hybridization with related diploid species. The finding that very low levels of nuclear (isozyme) diversity were present in both taxa ofS. vulgaris examined supports the second of these two hypotheses; however, more detailed analysis of nuclear genetic diversity is required before a firm conclusion can be reached on this matter.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Altitudinal variation of secondary metabolites in flowering heads of the Asteraceae: trends and causes

Plants in alpine habitats are exposed to severe environmental stresses including temperature and radiation extremes. The observation that flowering heads from high altitude populations of Scorzoneroides helvetica (Mérat) J.Holub (synonym: Leontodon helveticus Mérat) contained higher amounts of luteolin derivatives than conspecific populations in lower altitudes prompted further investigations. An elevational increase of phenolics was confirmed in flowering heads of neophytic populations of three additional taxa of the Cichorieae tribe in New Zealand. A solely genetic basis of the altitudinal trends of flavonoids and phenolic acids was eliminated by field experiments with cultivars of the medicinal herbs Arnica montana L. and Matricaria chamomilla L. planted at nine different altitudes ranging from 600 to 2,200 m in the Tyrolean Alps. Parallel experiments on potted plants excluded soil characteristics as the factor causing the observed variation. The initial hypotheses that enhanced UV-B radiation in higher altitudes was triggering an increase in the ratio of B-ring-ortho-diphenolic versus B-ring-monophenolic flavonols in flowering heads of Arnica was disproved by climate chamber experiments resulting in no significant difference between plants grown in ambient and threefold ambient UV-B radiation regimes. In contrast, an increase of this ratio similar to the changes observed in higher altitudes resulted from a decrease in temperature by 5°C in a second climate chamber experiment. Conclusively, enhanced UV-B radiation is probably not the key factor inducing shifts in the phenolic composition in Asteraceae growing at higher altitudes but it is rather the temperature which decreases with altitude.     

THE FLAVONOIDS OF LASTHENIA (COMPOSITAE)

Lasthenia (Compositae: Helenieae), a western North American-Chilean genus of 16 species, produces 22 flavonoid glycosides. Flavonoids are the chalcones butein and okanin, the aurones maritimetin and sulfuretin, the flavone luteolin, and the flavonols kaempferol, quercetin, and patuletin. The presence or absence of various of these classes of compounds in general follows sectional alignments in the genus, confirms affinities based on morphological and cytological evidence, and suggests relationships of problematical species. Intraspecific variation in flavonoid constituents occurs in several species, and in one taxon intrapopulation variation seems to exist as well. Evolution within Lasthenia has been associated with a loss of the ability to produce or accumulate luteolin, chalcones, and aurones; an increase in diversity of quercetin glycosides; acquisition of the ability to produce patuletin; and an elaboration of glycosylation patterns of patuletin.     

Management and diversity of arthropods inCalluna heathland

Heathlands, dominated byCalluna vulgaris, are located along the western seaboard of Europe. Most undergo a cyclical succession as the heather plants pass through four definable growth-phases over a period of about 30 years. Examples of upland heathlands are drawn from the North York Moors National Park, an area of Jurassic rocks in the north of England (approximately 54°25N, 0°55W). Botanical diversity is low. In a survey of 10 heathlands selected to encompass the botanical variation of the heathlands of the National Park, only 40 species of higher plants and 31 species of mosses were recorded (2.2% and 4.5% of the British flora, respectively).However, diversity of some invertebrate groups is high. Totals of 54 species of carabid beetles and 127 species of spiders were recorded (15.3% and 20.4% of the British fauna, respectively). At least two environmental gradients appear to influence the species composition of the invertebrate assemblages: a wet-dry gradient and the height of the vegetation (corresponding to the growth-phase of the heather). Experimental areas on three heathlands were burnt and cut, the traditional and more modern methods of heather management. Analysis of both the spider and beetle data indicated that the species assemblages were primarily influenced by the growth-phase of the heather. Some of the nationally rarer species are associated with the open conditions of recently-cut or burnt heathland. An altitudinal effect was also inferred since heathland at 410 m above sea level tended to be less species-rich than heathlands at ca 260 m above sea level.These results present a paradox. Although botanical diversity is extremely low, and although it is a non-natural ecosystem maintained by fire and grazing management to prevent a succession to woodland, arthropod diversity can be extremely high. The discussion focuses on the importance of upland heathlands in the conservation of insect and spider diversity, as well as on the value of these upland areas for conservation more generally.C. vulgaris is seen to be a key species on which many other species depend. One conclusion is that there is insufficient effort, nationally and especially internationally, to conserve the heathlands of Europe.     

Thermal ecology of allopatric lizards (Sphenomorphus) in southeast Australia

Summary The Sphenomorphus quoyi species complex (Lacertilia: Scincidae) of southeast Australia consists of three lizard species, distributed as continuous but largely allopatric populations, and a fourth species which has a disjunct distribution at high altitudes within the range of the other three. The four species have similar Voluntary temperatures (range of body temperatures within which normal activity occurs) despite the fact that they occupy different geographical areas.Altitudinal zonation is shown in the distribution pattern of the four species with S. quoyi and S. tympanum (warm temperate) restricted to the lower altitudes. The level of the mean daily minimum and maximum temperatures are different in each of the four species distribution areas. Similarly, the area occupied by each species is characterized by different daily sunlit and shade surface temperatures and different duration of ground surface subzero temperatures. Floristic composition of the four areas and the basking sites utilized by each species are also dissimilar.The lower temperature tolerances and the acclimation range of the Critical Minimum temperatures of S. quoyi and S. tympanum (warm temperate) exclude these species from the higher altitudes with the relatively lower environmental temperature levels. The Critical temperatures of the species which are found at high altitudes, do not explain why sympatry is not found with the lowland species.     

Altitudinal flower size variation correlates with local pollinator size in a bumblebee‐pollinated herb,Prunella vulgaris L. (Lamiaceae)

The influence of locally different species interactions on trait evolution is a focus of recent evolutionary studies. However, few studies have demonstrated that geographically different pollinator‐mediated selection influences geographic variation in floral traits, especially across a narrow geographic range. Here, we hypothesized that floral size variation in the Japanese herb Prunella vulgaris L. (Lamiaceae) is affected by geographically different pollinator sizes reflecting different pollinator assemblages. To evaluate this hypothesis, we posed two questions. (1) Is there a positive correlation between floral length and the proboscis length of pollinators (bumblebees) across altitude in a mountain range? (2) Does the flower–pollinator size match influence female and male plant fitness? We found geographic variation in the assemblage of pollinators of P. vulgaris along an altitudinal gradient, and, as a consequence, the mean pollinator proboscis length also changed altitudinally. The floral corolla length of P. vulgaris also varied along an altitudinal gradient, and this variation strongly correlated with the local pollinator size but did not correlate with altitude itself. Furthermore, we found that the size match between the floral corolla length and bee proboscis length affected female and male plant fitness and the optimal size match (associated with peak fitness) was similar for the female and male fitness. Collectively, these results suggest that pollinator‐mediated selection influences spatial variation in the size of P. vulgaris flowers at a fine spatial scale.     

Heartwood extractives and lignin content of different larch species (<Emphasis Type="Italic">Larix </Emphasis>sp.) and relationships to brown-rot decay-resistance

The extractive content of lignin and the brown-rot decay-resistance against Coniophora puteana and Poria placenta were studied in larch heartwood from different species and origin (Larix decidua var. decidua, L. decidua var. sudetica, L. kaempferi , L. × eurolepis). The study material consisted of 106 trees from a 39-year old provenance trial in France. The hot-water-soluble extractives were very variable (from5.66% to 20.50% of dry weight), but there was no significant variation between the investigated species and origins. In contrast, acetone extractives, the total amount of phenolics and lignin showed significant differences. The concentration of phenolics and lignin was significantly higher in L. kaempferi and in L. × eurolepis than in L. decidua. The total phenolics content was strongly correlated with decay-resistance in all investigated larch origins. A higher concentration of phenolics goes hand in hand with higher decay resistance and phenolics might therefore be a promising parameter to rapidly evaluate the level of decay-resistance in larch.     

Effects of human disturbance on the diet composition of wild red deer (Cervus elaphus)

Disturbance from human recreational activities may affect the nutrition of free-ranging herbivores due to trade-offs between feeding in preferred habitats and perceived predation risk. To test this hypothesis, we estimated diet composition for red deer in the Scottish highlands in spring, when recreational activity tends to be high, and in winter when it is lower. We analysed faecal samples from three habitat types (grassland, heather moorland and woodland) collected at sites close to a busy track (disturbed) and at a distance from it (less disturbed). The diet consisted of 39% grasses, sedges, herbs and rushes (GSHR) and 58% Calluna vulgaris and Erica spp. (heather) in spring, compared with 14% grasses and 77% heather in winter, with small quantities of Vaccinium spp. (berry) and Pinus sylvestris (tree) in both seasons. In spring, faeces from disturbed grass and woodland sites indicated a diet with less GSHR and more heather and tree than faeces from less-disturbed sites, which could be due to an increased need for vigilance in exposed grassland and the need to seek cover. In contrast, faeces from all disturbed sites in winter indicated a diet with more GSHR and less heather than faeces from less-disturbed sites. This could be due to a seasonal decline in recreation and increase in hunting activity reversing the disturbance levels at the different sites, since hunting is not normally carried out in areas used by the public for recreation. We conclude that there may be nutritional benefits to deer of reducing disturbance near open grassland.     

Polyphenolic compounds on leaves limit iron availability and affect growth of epiphytic bacteria

Polyphenolic compounds produced by plants can chelate iron, reducing its bioavailability to plant‐associated bacteria. In response to limited iron levels, most bacteria produce siderophores to acquire needed iron quantities. The amount of phenolic compounds detected in methanolic washings of leaves of different plant species varied greatly, being nearly sevenfold higher in Viburnum tinus than in Phaseolus vulgaris. In species with high levels of total phenolics (e.g. Pelargonium hortorum), tannin concentration of leaf washings was also high and accounted for up to 85% of total phenolics. Both stimulation of production of the siderophore pyoverdine in Pseudomonas syringae strain B728a and inhibition of growth of an isogenic mutant I‐1, deficient in pyoverdine production were associated with plants harbouring high levels of leaf surface phenolics. Levels of tannic acid sufficient to inhibit growth of the pyoverdine mutant in culture in an iron‐reversible fashion were similar to tannin levels found on leaves of plants such as P. hortorum. Additionally, the amount of pyoverdines produced by P. syringae and quantified in leaf washings from a variety of plants was directly related to the concentration of tannins released from the leaf, indicating that tannins were responsible for sequestering iron. Phenolic compounds, principally tannins, may thus play an important role in plant–microbe interactions.     

Limitations on reproductive success in endemic <Emphasis Type="Italic">Aquilegia viscosa</Emphasis> (Ranunculaceae) relative to its widespread congener <Emphasis Type="Italic">Aquilegia vulgaris</Emphasis>: the interplay of herbivory and pollination

Plant reproduction can be strongly affected by herbivory and different features of pollination ecology, such as pollinator visitation rates and capacity for self-pollination. The purpose of this study is to compare the relative impact of herbivory and pollination on maternal reproductive success in endemic Aquilegia viscosa and its widespread congener Aquilegia vulgaris. We conducted herbivore exclusion experiments in two populations of each species in 2 different years and showed that the maternal fertility of A. viscosa was significantly more limited by floral predation and pre-dispersal seed predation than its widespread congener. In the absence of herbivory, A. viscosa retained significantly lower maternal fertility than A. vulgaris. Experimental pollinations in an insect-free glasshouse showed that the two species have an equal seed/ovule ratio both in the absence of pollinators and in the presence of non-limiting outcross pollination. Pollinator visitation rates were significantly higher in populations of A. vulgaris than in populations of A. viscosa. In addition, path analyses showed that spur length, an important trait for pollinator attraction in Aquilegia, and, indirectly sepal and petal width, contribute positively to the seed/ovule ratio in A. vulgaris, but not in A. viscosa. These results indicate that maternal fertility of endemic A. viscosa is strongly reduced by flower and seed predation despite low rates of pollinator visitation, and that pollen or resource limitation in the wild may further reduce maternal fertility. Finally, floral trait variation appears to be decoupled from fertility variation in endemic A. viscosa, which possibly constrains the evolution of reproductive traits in this species.     

THE FLAVONOIDS OF THE DECIDUOUS RHODODENDRON OF NORTH AMERICA (ERICACEAE)

The native Azaleas of North America (Rhododendron: subgenus Pentanthera) produce 58 flavonoids in five aglycone classes: flavonols, dihydroflavonols, flavanones, dihydrochalcones and chalcones. A comparison of the flavonoids of selected samples of these species indicated that the compounds generally occur as species specific ensembles. Species were grouped into “alliances” based on common flavonoid constituents and a phylogenetic treatment of the group was developed. Evolutionary trends of flavonoids within subgenus Pentanthera are not well-defined but appear to be associated with a loss of the ability to synthesize some compounds and a decrease in diversity of glycosides and methoxylated flavonoids. Intraspecific variation in flavonoids was found in Rhododendron canescens, R. alabamense and R. austrinum when these were sampled on a population basis.     

紫翅猪毛菜、钠猪毛菜不同个体大小繁殖分配差异及随海拔的变化

以新疆准噶尔盆地藜科猪毛菜属植物紫翅猪毛菜(Salsola affinis C.A.Mey)、钠猪毛菜(Salsola nitraria Pall)为研究对象,用繁殖分配比例的方法对比分析了两种猪毛菜不同海拔同一种群内不同个体大小繁殖分配的特点,并用异速生长模型分析了不同海拔繁殖生物量与营养生物量之间分配与个体大小的依赖关系。结果发现:1)不同海拔繁殖生物量(R)与营养生物量(V)呈不同程度的异速生长。紫翅猪毛菜随海拔的升高R-V的异速生长斜率显著升高,截距随海拔的升高没有显著增加;而钠猪毛菜的斜率随海拔升高显著降低,截距则显著升高。2)紫翅猪毛菜在较低海拔个体大小与繁殖分配呈负相关,在较高海拔呈正相关;钠猪毛菜在较低海拔个体大小与繁殖分配呈正相关,在较高海拔呈负相关;两种猪毛菜繁殖分配的适应对策相反。3)将同一种群个体大小分成大、中、小3种类型,多重比较发现紫翅猪毛菜在较低海拔,中小个体的繁殖分配显著高于大个体的繁殖分配;在较高海拔,大个体的繁殖分配显著高于中小个体的繁殖分配。钠猪毛菜在较低海拔,大个体的繁殖分配显著高于中、小个体的繁殖分配;在较高海拔,小个体的繁殖分配显著高于大、中个体的繁殖分配。综合分析认为:两个物种随海拔变化产生不同的繁殖分配策略,除遗传效应外,环境和个体大小对钠猪毛菜繁殖分配的变化均产生重要影响,而紫翅猪毛菜繁殖分配的变化主要由海拔差异导致。由于微生境对同一种群的个体大小产生影响,进而产生不同的繁殖分配模式,所以在干旱区更应重视个体大小对繁殖分配的影响。     

The influence of vegetation type,soil properties and precipitation on the composition of soil mite and microbial communities at the landscape scale

Aim We used a landscape‐scale study of birch invasion onto heather moorland to determine the consistency of changes in vegetation type and soil properties and in the community composition of five soil organism groups. Our aim was to determine whether the degree to which soil organisms respond to natural changes and/or induced changes (e.g. changes in land‐use type and climate) in habitat is consistent across trophic and taxonomic groups in the context of conservation policies for birch woodland and heather moorland. Location Mainland Scotland. Methods We sampled mesostigmatid mites, oribatid mites, fungi, bacteria and archaea in adjacent patches of birch woodland (dominated by Betula pubescens) and heather moorland (dominated by Calluna vulgaris) at 12 sites for which annual rainfall ranged between 713 and 2251 mm. Differences in community composition were visualized using non‐metric multidimensional scaling based on Bray–Curtis dissimilarities. The factors contributing to differences between habitats within sites were explored using general linear models and those among sites using redundancy analysis. Results The communities of all groups differed between habitats within sites, but only the oribatid mites and fungi differed consistently between habitats across sites. Within sites, dissimilarity in fungal communities was positively related to the difference in C. vulgaris cover between habitats, whereas dissimilarities in bacteria and archaea were positively related to differences in soil pH and C:N ratio between habitats, respectively. Main conclusions The influence of vegetation type and soil properties differed between groups of soil organisms, albeit in a predictable manner, across the 12 sites. Organisms directly associated with plants (fungi), and organisms with microhabitat and resource preferences (Oribatida) were strongly responsive to changes in habitat type. The response of organisms not directly associated with plants (bacteria, archaea) depended on differences in soil properties, while organisms with less clear microhabitat and resource preferences (Mesostigmata) were not strongly responsive to either vegetation type or soil properties. These results show that it is possible to predict the impact of habitat change on specific soil organisms depending on their ecology. Moreover, the community composition of all groups was related to variation in precipitation within the study area, which shows that external factors, such as those caused by climate change, can have a direct effect on belowground communities.