A busca de padrões consistentes na natureza tem sido a principal meta dos ecólogos. Essa dissertação teve como objetivo usar abordagens filogenéticas na tentativa de compreender melhor o processo ecológico da expansão florestal sobre áreas abertas. O uso da informação filogenética em análises ecológicas considera as espécies não independentes umas das outras, pois estas compartilham grande parte da história evolutiva. Essa hierarquia de organização das espécies é muito importante para determinar as regras que governam os processos de montagem das comunidades locais. Nesta dissertação foram avaliados padrões filogenéticos de estruturação da vegetação lenhosa florestal ocorrente em ecótonos de áreas abertas com vegetação florestal distribuídos em diferentes regiões do extremo sul do Brasil. Estes ecótonos são formados por diversas formações florestais, as quais tendem a expandir sobre as áreas abertas. Dados sobre composição de espécies provieram de estudos já realizados e de amostragens em alguns sítios. Duas métricas filogenéticas complementares foram usadas para avaliar a estrutura filogenética em cada categoria de habitat nos ecótonos: índice de parentesco líquido (NRI) e coordenadas principais da estrutura filogenética (PCPS). As análises dos valores de NRI não mostraram um padrão nítido de estruturação filogenética das comunidades. Já a análise dos PCPS mostrou padrões consistentes nas três escalas espaciais abordadas e independente da composição de espécies. Clados basais associaram-se às áreas florestais, enquanto clados de diversificação recente associaram-se às áreas abertas. Estes resultados indicam que áreas abertas atuam como um filtro filogenético de habitat para as espécies lenhosas florestais em todos os locais analisados, independentemente da escala e da composição de espécies de cada local. Os resultados sugerem que os clados de Rosanae e Asteranae estão na linha de frente do processo de expansão florestal sobre as áreas abertas, e o clado de Magnolianae restrito às áreas florestais. A busca por padrões gerais de organização das comunidades ecológicas a partir de sua estrutura filogenética parece consistir numa ferramenta útil para a exploração e entendimento sobre o funcionamento de sistemas ecológicos. Estas abordagens poderiam beneficiar estratégias de gerenciamento e conservação destes sistemas, por simplificarem sistemas ecológicos complexos e por mostrarem padrões gerais independentes da escala espacial analisada. / The search for consistent patterns in nature has been a major goal of ecologists. This study aimed to employ phylogenetic analyses to improve the understanding of an ecological process, the expansion of forest expansion over open areas. The use of evolutionary information considers species as not independent units in relation to each other, as they share their evolutionary history. Such hierarchical organization of species is very important to determine the rules governing assembly processes in local communities. Phylogenetic approaches were used to evaluate phylogenetic patterns in forest woody vegetation occurring in ecotones comprising open areas and forest vegetation, and distributed across different regions in the southernmost Brazilian region. Those ecotones are composed by different forest vegetation types, which tend to expand over open areas. Data on species composition were compiled from previous studies, and vegetation sampling was carried out in sites without available information on species composition in ecotones. Two complementary phylogenetic metrics were used to evaluate the phylogenetic structure in each habitat type occurring in ecotones: net relatedness index (NRI) and principal coordinates of phylogenetic structure (PCPS). Analyses of NRI values did not show any clear pattern of phylogenetic structuring of the communities. Nonetheless, PCPS analysis showed consistent patterns across the threes spatial scales evaluated, which were independent of the species composition of the sites. Basal clades were associated with forest areas, while late-divergence clades were associated with open areas. These findings indicate that open areas act as a phylogenetic habitat filtering to forest woody species throughout the region, independently of the spatial scale and of the species composition in each site. The results suggest that the clades Rosanae and Asteranae represent the vanguard in theforest expansion process over open areas, while the distribution of the basal clade Magnolianae is restricted to forest sites. The search for general organization patterns in ecological communities based on their phylogenetic structure seems to be a useful tool for exploring and understanding the functioning of ecological systems. Such approach might benefit ecosystem managing and conservation strategies, as it simplifies complex ecological systems, and shows general patterns independently of the scale analyzed.

A busca de padrões consistentes na natureza tem sido a principal meta dos ecólogos. Essa dissertação teve como objetivo usar abordagens filogenéticas na tentativa de compreender melhor o processo ecológico da expansão florestal sobre áreas abertas. O uso da informação filogenética em análises ecológicas considera as espécies não independentes umas das outras, pois estas compartilham grande parte da história evolutiva. Essa hierarquia de organização das espécies é muito importante para determinar as regras que governam os processos de montagem das comunidades locais. Nesta dissertação foram avaliados padrões filogenéticos de estruturação da vegetação lenhosa florestal ocorrente em ecótonos de áreas abertas com vegetação florestal distribuídos em diferentes regiões do extremo sul do Brasil. Estes ecótonos são formados por diversas formações florestais, as quais tendem a expandir sobre as áreas abertas. Dados sobre composição de espécies provieram de estudos já realizados e de amostragens em alguns sítios. Duas métricas filogenéticas complementares foram usadas para avaliar a estrutura filogenética em cada categoria de habitat nos ecótonos: índice de parentesco líquido (NRI) e coordenadas principais da estrutura filogenética (PCPS). As análises dos valores de NRI não mostraram um padrão nítido de estruturação filogenética das comunidades. Já a análise dos PCPS mostrou padrões consistentes nas três escalas espaciais abordadas e independente da composição de espécies. Clados basais associaram-se às áreas florestais, enquanto clados de diversificação recente associaram-se às áreas abertas. Estes resultados indicam que áreas abertas atuam como um filtro filogenético de habitat para as espécies lenhosas florestais em todos os locais analisados, independentemente da escala e da composição de espécies de cada local. Os resultados sugerem que os clados de Rosanae e Asteranae estão na linha de frente do processo de expansão florestal sobre as áreas abertas, e o clado de Magnolianae restrito às áreas florestais. A busca por padrões gerais de organização das comunidades ecológicas a partir de sua estrutura filogenética parece consistir numa ferramenta útil para a exploração e entendimento sobre o funcionamento de sistemas ecológicos. Estas abordagens poderiam beneficiar estratégias de gerenciamento e conservação destes sistemas, por simplificarem sistemas ecológicos complexos e por mostrarem padrões gerais independentes da escala espacial analisada. / The search for consistent patterns in nature has been a major goal of ecologists. This study aimed to employ phylogenetic analyses to improve the understanding of an ecological process, the expansion of forest expansion over open areas. The use of evolutionary information considers species as not independent units in relation to each other, as they share their evolutionary history. Such hierarchical organization of species is very important to determine the rules governing assembly processes in local communities. Phylogenetic approaches were used to evaluate phylogenetic patterns in forest woody vegetation occurring in ecotones comprising open areas and forest vegetation, and distributed across different regions in the southernmost Brazilian region. Those ecotones are composed by different forest vegetation types, which tend to expand over open areas. Data on species composition were compiled from previous studies, and vegetation sampling was carried out in sites without available information on species composition in ecotones. Two complementary phylogenetic metrics were used to evaluate the phylogenetic structure in each habitat type occurring in ecotones: net relatedness index (NRI) and principal coordinates of phylogenetic structure (PCPS). Analyses of NRI values did not show any clear pattern of phylogenetic structuring of the communities. Nonetheless, PCPS analysis showed consistent patterns across the threes spatial scales evaluated, which were independent of the species composition of the sites. Basal clades were associated with forest areas, while late-divergence clades were associated with open areas. These findings indicate that open areas act as a phylogenetic habitat filtering to forest woody species throughout the region, independently of the spatial scale and of the species composition in each site. The results suggest that the clades Rosanae and Asteranae represent the vanguard in theforest expansion process over open areas, while the distribution of the basal clade Magnolianae is restricted to forest sites. The search for general organization patterns in ecological communities based on their phylogenetic structure seems to be a useful tool for exploring and understanding the functioning of ecological systems. Such approach might benefit ecosystem managing and conservation strategies, as it simplifies complex ecological systems, and shows general patterns independently of the scale analyzed.

Les changements d'intensité dans l'utilisation des sols et la dégradation de l'environnement en raison de activités humaines ont entraîné une forte altération des écosystèmes tropicaux, en particulier dans les cours d'eau de tête de bassin de la savane néotropicale. Les pressions humaines liées à l'expansion agricole et à l'urbanisation ont conduit à une réduction drastique de la couverture végétale indigène, affectant les zones riveraines et altérant le fonctionnement des écosystèmes aquatiques. Il est urgent de quantifier et de prévoir comment les communautés aquatiques répondent aux changements de l'utilisation des sols pour guider les efforts de conservation et de gestion des ressources naturelles. Dans ce contexte, cette thèse visait à évaluer à quelles échelles spatiales la relation entre habitat et communautés de macroinvertébrés benthiques s'exprimait le plus fortement et comment l'intensité d'utilisation des sols affectait les relations trophiques et la composition en traits biologiques des communautés de macroinvertébrés benthiques. Dans le chapitre 1, j'évalue comment l'intensité d'utilisation des sols (représentée par un gradient depuis la végétation autochtone, au pâturage, et à la plantation de canne à sucre et la végétation indigène) influe sur les flux d'énergie et les niches trophiques des macroinvertébrés benthiques. Ensuite, j'ai évalué à quelle échelle spatiale (bassin versant, localité) et pour quelles variables, le lien entre la composition taxonomique et fonctionnelle des assemblages de macroinvertébrés benthiques était le mieux exprimé (chapitre 2). Enfin, j'ai montré que les perturbations anthropiques avait unj impact sur la diversité fonctionnelle des assemblages de macroinvertébrés benthiques (chapitre 3). Les résultats présentés ici montrent que les changements d'intensité de l'utilisation des sols conduisent les assemblages de macroinvertébrés benthiques vers des comportements d'alimentation plus généralistes avec un chevauchement des niches trophiques (chapitre 1). De plus, les variables environnementales à l'échelle locale et du bassin versant expliquent de façon significative les variations de la composition taxonomique et fonctionnelle des assemblages d'Ephéméroptères, Plécoptères et Trichoptères, mais les variables décrivant l'intensité d'utilisation des sols expliquent le mieux les différences de composition fonctionnelle entre les différents sites (chapitre 2). Enfin, J'ai montré que les sites quasi-naturels (dans des conditions de référence) ont des assemblages de macroinvertébrés plus spécialisés comparés aux sites perturbés (chapitre 3). Les résultats corroborent le fait que la biodiversité doit être évaluée en tenant compte de variables agissant à de multiples échelles spatiales et que les éléments fonctionnels des communautés biologiques doivent être considérés, en vue de la conservation et du développement d'outils prédictifs. Cette étude contribue à une meilleure compréhension de la structure et du fonctionnement des cours d'eau dans la savane néotropicale dans le contexte du développement d'outils d'évaluation environnementale. Ces approches contribueront à l'élaboration de mesures de gestion et de conservation plus appropriées et permettront d'examiner les conséquences futures d'une poursuite de la dégradation des conditions écologiques dans les cours d'eau tropicaux / Changes in land cover and use and the associated environmental degradation due to human activities have resulted in extreme alterations of tropical ecosystems, especially in headwater streams and their watersheds in the neotropical savanna. Human pressures related to agricultural expansion and urbanization have led to drastic reductions in native vegetation cover, affecting riparian zones and degrading aquatic ecosystem functioning. There is an urgent need to quantify and predict how aquatic communities respond to different intensities of land use to guide conservation and natural resource management efforts. This thesis aims to evaluate how spatial scales influence the relationship between habitat and benthic macroinvertebrate communities and how land use intensity affects the trophic relationships and biological traits of macroinvertebrates. In Chapter 1, we evaluated how the intensity of land use (represented by a gradient moving from native vegetation toward pasture and sugar cane plantations) influences the energy flow and trophic niches of macroinvertebrates. In Chapter 2, we investigated the spatial scales (e.g., catchment, local) that most influence the taxonomic and functional composition of macroinvertebrate assemblages. Finally, in Chapter 3, we examined the impacts of human pressures on the functional diversity of macroinvertebrate assemblages. we showed that the intensity of land use affects benthic macroinvertebrate assemblages, yielding more generalist feeding behaviors with greater overlap of trophic niches (Chapter 1). In addition, environmental variables at the local and catchment scales significantly explained the variations in the taxonomic and functional composition of Ephemeroptera, Plecoptera and Trichoptera assemblages, but land use variables best explained the differences in functional composition among sites (Chapter 2). Finally, we showed that less impacted sites (under reference conditions) had more specialized and more functional diverse macroinvertebrate assemblages compared to disturbed sites (Chapter 3). These results corroborate the idea that biodiversity should be evaluated at multiple spatial scales and that the functional elements of biological communities should be considered when aiming for conservation and the development of predictive tools. This study contributes to a better understanding of the structure and functioning of streams in the neotropical savanna by subsidizing the development of environmental assessment tools. Such approaches will contribute to the development of more appropriate management and conservation measures that will allow for evaluation of the impacts on biota of further degradation of the ecological conditions in tropical streams

La vallée de la Punaru’u est le second bassin versant, par sa taille, de la Polynésie française. Abritant la plusgrande zone industrielle du Pays, la vallée de la Punaru’u est emblématique des impacts de l’anthropisation sur le milieu naturel. L’exploitation de sa ressource en eau est au centre des préoccupations des industriels et des collectivités locales. L’objectif du projet était de mieux définir et de mieux comprendre les transferts d’eau dans le sol sous la zone industrielle et d’estimer l’impact des activités humaines sur la ressource en eau. En s’appuyant sur la définition des caractéristiques hydrodynamiques du sol une première partie du travail s’est concentrée à faire le lien avec les propriétés texturales et structurales du sol. Cette approche physique et analytique a permis d’ouvrir la voie à un protocole d’acquisition de données novateur basé sur les propriétés texturales du sol. Ce résultat a l’avantage de résoudre les problèmes d’échelle et de s’affranchir d’une dépendance aux conditions initiales et limites qui habituellement handicape l’interprétation des mesures des paramètres hydrodynamiques du sol.Une seconde phase de travail s’est concentrée sur l’analyse statistique des données hydrologiques. Il en résulte un schéma rénové des dynamiques hydrologiques et hydrogéologiques dans la basse vallée de la Punaru’u. D’un point de vue quantitatif, la ressource en eau s’épuise beaucoup plus en surface et dans les premiers mètres du sol que dans les aquifères plus profonds.L’approche analytique et l’interprétation hydrogéologique ont permis finalement de paramétrer une modélisation du sous-sol de la zone industrielle basée sur le solver ModFlow. / Punaru'u’s valley is the second watershed by size of French Polynesia. The valley hosts the most important industrial area of the country. The area is becoming an emblematic area because of urbanization impacts on natural environment. Exploitation of water resources is a central preoccupation.The aim of the present study is to analyze and to understand the surface and soil water transfer processes allowing for quantifying its vulnerability to the impact of human activities. Using a theoretical methodology especially developed for the characterization of hydrodynamic soil characteristics, the first part of this work is focused on the relation between textural and structural soil properties. Physical and analytical concepts are used to find a way to open a new data provision strategy based on the knowledge of measurable textural soil properties. The approach provides an original solution for tackling the important scale problem that affects most integrated modeling schemes used to describe hydrological flow processes at watershed scale. Second part of this work is focused on the statistical analysis of the hydrological field data. The results give a new insight in the hydrological and hydrogeological flow processes of the lower part of the Punaru'u valley. A quantitative interpretation shows how soil water resources are affect by human activities. The three alluvial aquifers situated in the first 40 m from the soil surface are all connected. The analytical approach and the hydrogeological interpretation allow for setting up a modeling scheme based on the “ModFlow” solver; the model is able to reproduce the hydrological flow processes of the lower part of thePunaru’u valley.

Abstract Understanding of natural and human induced changes to freshwater biodiversity patterns is critical for efficiently conserving and managing these highly impacted ecosystems. In this doctoral thesis, I study the natural spatial and temporal variability of littoral macroinvertebrate communities, as well as aquatic biodiversity responses (of bryophytes, macroinvertebrates and diatoms) to human impact on forested headwater streams. I also study the temporal variability of natural and human-impacted stream communities. I show that littoral macroinvertebrate communities are most variable between individual samples and to some extent between littorals, and that the diversity patterns of rare and common littoral macroinvertebrates vary at different spatial scales. Temporal (interannual) variability is of minor importance, particularly for the core species of the community. For rare species, however, the temporal component was generally more important than spatial turnover. For headwater streams, I show that most taxonomic groups responded negatively to human impact on the adjacent riparian forest. Nevertheless, pristine riparian forest is not a sufficient surrogate for stream biodiversity as also streams with relatively disturbed riparian forests supported diverse benthic communities. A rapid classification method based on the physical structure of the stream channel and its riparian forest is able to identify the reference-state communities, as well as the most severely altered communities, but a reliable identification of the intermediately-disturbed sites would need more precise information about the key stressors of stream communities. I found that fine sediments originating from forest drainage were the main factor causing reduction of stream biodiversity. I also found that forestry impacted streams varied temporally more than pristine streams, emphasizing the importance of interannual sampling of impacted streams for a reliable status assessment. / Tiivistelmä Luonnollisen ja ihmisen aiheuttaman biodiversiteettivaihtelun ymmärtäminen hoito- ja suojelutoimissa on tärkeää pienvesien monimuotoisuuden turvaamiseksi. Tarkastelen väitöskirjassani järvien rantavyöhykkeen pohjaeläinten ajallista ja paikallista vaihtelua sekä latvapurojen eliöyhteisöjen (pohjaeläimet, sammalet, piilevät) vasteita ihmisen aiheuttamaan fysikaaliseen muutokseen rantametsässä ja purouomassa. Tarkastelen myös ajallisen vaihtelun eroja luonnontilaisten ja ihmisvaikutteisten latvapurojen eliöyhteisöjen välillä. Osoitan, että järven rantavyöhykkeen pohjaeläinyhteisöt vaihtelevat eniten yksittäisten näytteiden välillä ja pienessä määrin eri rantojen välillä, mutta ajallinen vaihtelu ei heikennä tulosten luotettavuutta muiden kuin harvinaisten lajien osalta yhteen vuoteen rajoittuvassa näytteenotossa. Purotutkimuksissa osoitan, että useiden vesieliöryhmien lajirunsaus muuttuu ihmisen aiheuttaman rantametsän muutoksen seurauksena, mutta luonnontilaista rantametsää ei silti voida pitää yksiselitteisesti myös puron suojelutason mittarina, koska se ei välttämättä turvaa eliöiden lajistollista eheyttä suhteessa alkuperäiseen alueelliseen lajistoon. Rantametsän ja uoman fysikaalisiin rakennepiirteisiin perustuvaa nopeaa luokittelumenetelmää voidaan käyttää eliöyhteisöltään monimuotoisimpien ja metsätalouden voimakkaimmin rasittamien purojen tunnistamisessa. Eliöyhteisöltään lievemmin muutettujen purojen luotettavaan erotteluun luokittelun pitäisi kuitenkin sisältää tarkempaa tietoa metsäojitusten aiheuttamasta liettymisestä, joka on tulosteni perusteella haitallisin metsätaloudesta johtuva puroyhteisöjä rasittava tekijä. Havaitsin myös metsätalouden rasittamien purojen tilanarvioinnin toistettavuuden olevan luonnontilaisia puroja heikompaa, joten metsätalouden rasittamien purojen pidempiaikainen seuraaminen tuntuu välttämättömältä luotettavien tilanarvioiden tuottamiseksi.

The current ecological hypothesis states that the soil type (e.g., chemical and physical properties) determines which microbes occupy a particular soil and provides the foundation for soil provenance studies. As human profiles are used to determine a match between evidence from a crime scene and a suspect, a soil microbial profile can be used to determine a match between soil found on the suspect’s shoes or clothing to the soil at a crime scene. However, for a robust tool to be applied in forensic application, an understanding of the uncertainty associated with any comparisons and the parameters that can significantly influence variability in profiles needs to be determined. This study attempted to address some of the most obvious uncertainties of soil provenance applications such as spatial variability, temporal variability, and marker selection (i.e., taxa discrimination). Pattern analysis was used to validate the ecological theories driving the soil microbial biogeography. Elucidating soil microbial communities’ spatial and temporal variability is critical to improve our understanding of the factors regulating their structure and function. Microbial profiling and bioinformatics analyses of the soil community provided a rapid method for soil provenance that can be informative, easier to perform, and more cost effective than approaches using traditional physico-chemical data. This study also showed that stable profiles may allow comparison between evidence and a possible crime scene despite the time lapse (4 years) between sample collections, however, this is dependent on the analysis method, site, vegetation, and level of disturbance. Marker selection was also an important consideration for profiling. Even though Fungi look promising for single taxon soil discrimination, the additional markers can help discriminate between a wide variety of soil types. As in human identification, the more DNA markers queried the greater the discrimination power. Lastly, this study illustrated a novel method to query the iron relating genes and ability to design a novel marker that can easily be used to profile the functional diversity of a soil community to enhance soil classification. Overall this research demonstrated the potential and effectiveness of using microbial DNA from soil, not just for comparison, but also for intelligence gathering to pinpoint the geographic origin of the soil.

Invasive plants can outcompete native plants, replacing diverse plant communities with monocultures, which can negatively impact the whole ecosystem. One invasive plant, Phragmites australis, has invaded wetlands across North America. In Utah’s Great Salt Lake, it has greatly reduced the area of native plants that are important habitat for migratory birds. Here we describe experiments that assess multiple treatments for Phragmites removal and evaluate the return of native plants after Phragmites management. The treatments were applied to Phragmites patches at two scales (small 1/4-acre plots and large 3-acre plots) and across multiple sites to evaluate how patch size and environmental differences can influence the plants that return after Phragmites removal. The treatments (applied over 3 years and monitored two more) compared two different herbicides (glyphosate and imazapyr) and different herbicide and mowing timings. The treatments evaluated in the large patch study were 1.) untreated control 2.) fall glyphosate, winter mow, 3.) summer imazapyr, winter mow, 4.) summer glyphosate, winter mow. The treatments evaluated in the small patch study included treatments 1-4 above plus 5.) summer mow, fall glyphosate, 6.) summer mow, then black plastic solarization. In the small patches, we also monitored the seeds in the soil to assess how Phragmites management treatments can change the densities of Phragmites and native seeds. Fall glyphosate treatments were superior for Phragmites cover reduction. After the initial treatment, summer herbicide and mow treatments reduced Phragmites seed production, while fall glyphosate did not. Phragmites seeds were plentiful in the soil but were reduced following three years of all herbicide treatments. Native plant recovery following Phragmites management was extremely variable across sites. Sites with high soil moisture had better Phragmites removal and more native plants. But when flooding was deep, native plants were rare. Native seed density in the soil did not change due to Phragmites management, but soil seed densities were different across sites, which influenced native plant recruitment. Phragmites was removed more effectively and native plants returned in greater numbers in small patches compared with large. This was because small patches were typically near established native plant communities, which likely provided more native plant seeds and had hydrology that was less disturbed by human activity. In sites where native plants do not return after Phragmites management, practitioners may need to try revegetation with native plant seeds to restore important native plant communities.

No description available.

Species-species and species-habitat interactions have been demonstrated to be important in influencing diversity across a variety of ecosystems. Despite generalities in the importance of these interactions, appropriate mechanisms to explain them are absent in many systems. In sandhill systems of the southeast U.S., gopher tortoises have been hypothesized to be a crucial species in the maintenance of diversity and function. However, the mechanisms and magnitude in which they influence their communities and habitats have rarely been empirically quantified. I examined how habitat structure influences tortoise abandonment of burrows and how tortoise densities influence nonvolant vertebrate community diversity. Tortoise burrow abandonment is directly influenced by canopy closure, with each percent increase in canopy cover relating to a ~2% increase in the probability of burrow abandonment. In addition, tortoise burrow density was positively correlated with diversity and evenness, but not species richness. This influence was directly proportional to burrow density, supporting a dominance role for this species and rejecting the commonly asserted keystone species mechanism. I also quantified the influence of tortoises in influencing diversity relative to other environmental and habitat variables. Through this research, I have demonstrated that disturbance and habitat structure are important, but diversity responds most to density of burrows in the habitat. These findings demonstrate the intricate relationships interacting to maintaining diversity in sandhill systems. In particular, habitat change leading to declines of gopher tortoises may have drastic negative impacts on vertebrate species diversity.

The observational study on natural regeneration of Norway spruce (Picea abies) was provided during seasons 2010 and 2011 in the windthrow area of High Tatra National Park. The main aim of this master thesis was to describe the distribution pattern in Norway spruce regeneration as well as to estimate the spatial scale, which most preciselly describes the differences in its phenotypic variability. Following step was to determine the relative influence of abiotic and stand factors on the growth characteristics and abundance of spruce seedlings. The Main focus lied on stressing the role of biotic interactions, like intraspecific and interspecific competition, leading to certain discrepancies in the phenotype, as well as in abundance of spruce seedlings.The reponse in variability in spruce traits was recognized at each spatial level. The results showed clear causality between soil moisture and vegetation composition, which both directly modified not only the phenotype of spruce seedlings and plants, but have even influenced their counts on particular stands.