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Silvicultural management does not affect biotic communities in conifer plantations in the short-term: A multi-taxon assessment using a BACI approach

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Biodiversity maintenance is a key strategy for sustainable forestry in both above-ground and below-ground biotic communities. However, few studies applied continuous monitoring to analyse the responses of different taxonomic groups to silvicultural treatments. We studied the short-term effects of three silvicultural treatments (no thinning, thinning from below, and selective thinning) on taxonomic richness and composition in two Pinus nigra J.F. Arnold plantations in Tuscany (Italy). We conducted a 1 year before–3 years after control-impact (BACI) experiment with a complete randomized design and analysed the responses of five different taxonomic groups (bacteria, nematodes, microarthropods, mushrooms and vascular plants (overstorey and understorey), along with the patterns of different structural variables. The silvicultural treatments induced a sudden decrease of many parameters such as number of trees per hectare, basal area, and standing volume, with a direct impact on the Photosynthetic Active Radiation on the ground (PAR). Despite this, biological communities showed a high resistance to the tree thinning intensities. Indeed, none of the thinning treatments significantly affected the entire community in the short-term, neither regarding taxonomic richness nor composition. The different taxonomic groups showed a similar, low or null, sensitivity to forest management, and thus a high congruence in their responses.
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... Since different taxonomic communities can have contrasting responses to anthropogenic and environmental drivers, multi-taxonomic assessments are important for better understanding humans' impacts on biodiversity [45][46][47]. Although vascular plants and springtails are recognized as effective biological indicators in agroecosystems [48,49], their combined response to agricultural management practices remains largely unexplored. ...
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Black pine has been used often in central and southern Italy to reforest mountainous areas depleted by the intensive use of natural resources. The main purpose of establishing pine forests in Italy was to protect the soil from excessive erosion, and also to facilitate the natural succession toward mixed forests with deciduous species. The most common silvicultural treatments in Europe currently aim at maximizing the stability of the stands and facilitating the transition from pure to mixed stands comprising a larger component of native tree species. In this work, we investigated the relationships between the living whorls number and four indexes of individual tree stability: the slenderness ratio, the crown depth, the crown projection, and an eccentricity index of the canopy. The data set used was composed of 1098 trees from ten black pine plantations located in central Italy. Our results demonstrate that the living whorls number can be handily used to predict the slenderness ratio with an error of 18%. A non-parametric model based on a reduced number of field measures was obtained as a support for thinning operations aimed at improving single tree stability.
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Accettato il 29 settembre 2010 Riassunto – Si riportano i risultati di una sperimentazione impostata nel 1978 sul trattamento selvicolturale di una pineta di impianto di pino nero nel territorio del Pratomagno casentinese (Arezzo). Le tesi poste a confronto sono state: (i) diradamento dal basso moderato; (ii) diradamento dal basso forte; (iii) controllo. Il trattamento ha previsto tre diradamenti nel 1978, nel 1999 e nel 2009 alle età di 24, 45 e 55 anni. Si analizzano gli interventi attuati e gli effetti nel tempo sui parametri dendrometrici e di stabilità del soprassuolo H/D (rapporto ipsodiametrico medio) e H/D dom (rapporto ipsodiametrico dominante). Sono analizzati poi i parametri: rapporto ipsodiametrico, profondità, area di insidenza ed eccentricità della chioma su un campione di alberi stratificati per classe sociale e per tesi di trattamento in confronto con uno cresciuto in assenza di concorrenza. I risultati confermano che i diradamenti sono il mezzo più incisivo per accrescere la stabilità di popolamenti forestali artificiali monoplani. Tipo, regime e grado dei singoli interventi in rapporto allo stadio di sviluppo sono gli elementi determinanti. Nei diradamenti dal basso l'intensità dell' intervento influisce direttamente sull'incremento dei parametri di stabilità nel tempo soltanto se gli interventi interessano anche il piano dominante. La sperimentazione ha dimostrato che tutti i parametri di stabilità sono migliorati nei dieci anni successivi al diradamento intermedio eseguito a 45 anni, in misura maggiore nella tesi di diradamento forte. Abstract – Effects of silvicultural treatment on the stability of black pine plantations. Black pine plantations have been established at the purpose of recovering a forest cover to marginal soils, mostly throughout the Apennines range in Italy, since the end of the eighteenth century and up to the mid 1900. Both the decay of forest cover and soil erosion were the outcome of the long-lasting overuse through the intensive forest exploitation practices, grazing of the forest floor and wildfires, occurring since many centuries ago. The primary function of pine reafforestation was therefore to re-establish a first cover with a pioneer species, preparatory to future mixed forest types based on the natural reintroduction of broadleaves originally living in the same areas, mainly deciduous oaks and beech in the upper part. These goals have been partly met over the wide reafforestation area; the key functions of pine stands are today the protection against soil erosion and the hydrological regulation of catchments. The pine stands have been assuming today also a scenic role because they have been incorporated in the landscape physiognomy. A series of thinning up to the regeneration phase had been planned by foresters since the design of these plantations, but many stands have grown unthinned and fully stocked for a long time, this condition contributing a less mechanical stability of trees. Alternative forms of regeneration in grown-up stands have been and are being tested to improve both the natural and artificial establishment of indigenous species, but thinnings remain, even if a tardy measure, the main practice enforceable to these pine forests. The results of experimental trials undertaken in the black pine forest stand located in Pratomagno casentinese (Arezzo) are being reported in the paper. The study started in 1978 and the following theses were tested (A) heavy thinning from below; (B) moderate thinning from below; (C) control. Three thinnings were carried out in 1978, 1999 and 2009 at the ages of 24, 45 and 55. The action over time of each intervention on the mensurational and mechanical stability parameters (H/D ratio H/D dom ratio), are being analyzed. The stability parameters of the pine forest after three interventions (H/D ratio, crown ratio, horizontal crown projection and eccentricity of the crown) have been also analyzed on a sample of trees per thesis and compared with those of trees grown in absence of competition. Sampled trees were sorted per social class. Results proved that type, interval and intensity of thinning related to the age of intervention are the main determinants of a successful treatment. Thinnings from below increase individual stability over time only when also trees living in the main crop layer are being removed. The trials showed the improvement of stability parameters following the intermediate thinning applied at the age of 45. The thesis of heavy thinning had the best performance as for the awaited purpose.
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A holy grail of conservation is to find simple but reliable measures of environmental change to guide management. For example, particular species or particular habitat attributes are often used as proxies for the abundance or diversity of a subset of other taxa. However, the efficacy of such kinds of species-based surrogates and habitat-based surrogates is rarely assessed, nor are different kinds of surrogates compared in terms of their relative effectiveness. We use 30-year datasets on arboreal marsupials and vegetation structure to quantify the effectiveness of: (1) the abundance of a particular species of arboreal marsupial as a species-based surrogate for other arboreal marsupial taxa, (2) hollow-bearing tree abundance as a habitat-based surrogate for arboreal marsupial abundance, and (3) a combination of species- and habitat-based surrogates. We also quantify the robustness of species-based and habitat-based surrogates over time. We then use the same approach to model overall species richness of arboreal marsupials. We show that a species-based surrogate can appear to be a valid surrogate until a habitat-based surrogate is co-examined, after which the effectiveness of the former is lost. The addition of a species-based surrogate to a habitat-based surrogate made little difference in explaining arboreal marsupial abundance, but altered the co-occurrence relationship between species. Hence, there was limited value in simultaneously using a combination of kinds of surrogates. The habitat-based surrogate also generally performed significantly better and was easier and less costly to gather than the species-based surrogate. We found that over 30 years of study, the relationships which underpinned the habitat-based surrogate generally remained positive but variable over time. Our work highlights why it is important to compare the effectiveness of different broad classes of surrogates and identify situations when either species- or habitat-based surrogates are likely to be superior.
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Monocultural coniferous plantations have prevailed worldwide in recent decades, which supplied much of the world’s timber, but also exerted some negative effects on local ecologies and environmental systems. Continuous development has increased the various demands of human society for forests and it is necessary to balance concerns for the ecological and economic functions. Ample evidence indicates that mixed forests are an ideal option for providing more diversified ecological services and forest goods. Converting pure forests into mixed forests by introducing broad-leaved hardwood species below coniferous plantations has become an increasingly popular forest management strategy. Yet, there has been not enough research to date on suitable management methods for enhancing forest diversity and resilience in the context of forest conversion. To comprehensively examine how the intensity in Masson pine thinning influences forest evolution, seedlings of two native hardwood species were introduced below unthinned and thinned (varying intensity) Masson pine plantations. The effects of thinning on residual tree growth, seedling survival and growth, and understory vegetation development were analyzed using a generalized additive mixed model (GAMM). Monitoring results over 10 years indicate that thinning is a necessary management measure to accelerate forest succession in the conversion process; thinning exerts significant effects on the growth of residual trees, the survival and growth of seedlings, and the development of understory vegetation. Intense thinning results in more residual tree growth and enhances the richness and diversity of herbaceous species. However, excessive thinning can reduce the likelihood of seedling survival and growth as well as the richness and diversity of shrubs. Optimal thinning intensities appear to fall between 50% and 60% depending on the specific introduced species; light-demanding species may need higher overstory thinning intensity than shade-tolerant species. The biological characteristics of the introduced species must be taken into account to design an effective thinning strategy for pine plantations conversion.
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The influence of forest management on fungal diversity and community composition has been the subject of a wide number of studies over the last two decades. However, the difficulty of studying the complex kingdom of fungi under real forest conditions has led to rather scattered scientific knowledge. Here, we provide the current state of knowledge suggesting future research directions regarding (i) stand structure attributes (age, tree cover, stand density, tree species composition), (ii) management history (managed vs unmanaged), (iii) silvicultural treatments (thinning, clearcutting, shelterwood methods, selective cutting) and (iv) other anthropogenic disturbances (mushroom picking, salvage logging, prescribed burning, fertilization) affecting fungal diversity and community composition. The reviewed studies reported a positive correlation between fungal diversity and stand structure variables such as canopy cover, basal area of the stand and tree species diversity, particularly for mycorrhizal species. Abundance and diversity in size, tree species and decomposition stage of deadwood are reported as positively related to richness of wood-inhabiting fungi. The main findings about the effects of silvicultural practices suggest that the higher is the management intensity the lower is the diversity of ectomycorrhizal and wood-inhabiting species, at least in the short term. We have therefore reported those silvicultural practices which may reduce trade-offs between timber harvesting and fungal diversity conservation. Indeed, fungal diversity can be conserved in managed forests if (i) low impact logging operations are performed; (ii) stand structural complexity and late-successional forest characteristics are enhanced; (iii) deadwood amount and diversity is promoted, (iv) landscape heterogeneity and connectivity is improved or maintained.
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Soil plays a fundamental role in many ecological processes, throughout a complex network of above- and below-ground interactions. This has aroused increasing interest in the use of correlates for biodiversity assessment and has demonstrated their reliability with respect to proxies based on environmental data alone. Although co-variation of species richness and composition in forests has been discussed in the literature, only a few studies have explored these elements in forest plantations, which are generally thought to be poor in biodiversity, being aimed at timber production. Based on this premise our aims were 1) to test if cross-taxon congruence across different groups of organisms (bacteria, vascular plants, mushrooms, ectomycorrhizae, mycelium, carabids, microarthropods, nematodes) is consistent in artificial stands; 2) to evaluate the strength of relationships due to the existing environmental gradients as expressed by abiotic and biotic factors (soil, spatial-topographic, dendrometric variables). Correlations between groups were studied with Mantel and partial Mantel tests while variance partition analysis was applied to assess the relative effect of environmental variables on the robustness of observed relationships. Significant cross-taxon congruence was observed across almost all taxonomic groups pairs. However, only bacteria/mycelium and mushrooms/mycelium correlations remained significant after removing the environmental effect, suggesting that a strong abiotic influence drives species composition. Considering variation partitioning, the results highlighted the importance of bacteria as a potential indicator: bacteria were the taxonomic group with the highest compositional variance explained by the predictors used; furthermore, they proved to be involved in the only cases where the variance attributed solely to the pure effect of biotic or abiotic predictors was significant. Remarkably, the co-dependent effect of all predictors always explained the highest portion of total variation in all dependent taxa, testifying the intricate and dynamic interplay of environmental factors and biotic interactions in explaining cross-taxon congruence in forest plantations.
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Monitoring restoration effectiveness implies that restoration measures should be evaluated using multiscale analysis and modelling approaches. In this way, the scale-dependent nature of ecosystem processes and dynamics can be adequately described. Here, we evaluated how grain size (i.e. the dimension of sampling units) and type of data (abundance vs. presence/absence) impact the analysis of the response to restoration management activities aiming at halting the loss of European beech forests with Abies alba in the central Apennines (Italy). The proposed analysis relies on forests sampled in three Sites of Community Importance (SCIs) located in central Italy. For each treatment (reference, control and impact), randomly located multi-scale plots were selected. Vascular plant occurrences were recorded twice in every plot during summer 2010 or 2011 (before restoration) and in 2013 or 2014 (after restoration). Analysis of species abundances, recorded at the largest grain size available (20 X 20 m plot), showed a significant change in plant species composition in restored sites just after two years from the restoration management. When presence/absence data were used previous results were less visible, for all the examined grains, in two out of the three sites, suggesting a site-dependent response.
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The impacts of climate extremes on species, communities and ecosystems have become critical concerns to science and society. Under a changing climate, how restoration outcomes are affected by extreme climate variables is a largely unknown topic. We analyzed the effects of experimental factors (grazing and sowing of native species), extreme climate events (intense precipitation and extreme temperatures indexes) and their combination on the restoration progress of a dry, calcareous grassland in Tuscany (Italy) with a 1 year before/15 years continuous annual monitoring after, control/impact (BACI) experiment. Grazing had a beneficial effect on the diversity of the grassland, while sowing had a limited impact. The climatic index that most affected the entire plant community composition was the number of very heavy precipitation days. The interaction of grazing and extreme climatic indexes had a significant detrimental effect on restoration outcomes, increasing the cover of synanthropic and cosmopolitan-subcosmopolitan generalist species and decreasing the cover of more valuable species such endemic species. In the richer grazed plots, species richness showedn a lower sensitivity to the average precipitation per wet day.: this results suggest that dealing with climate extremes, the richest communities might not be the more resistant. In grazed site restoration outcomes can be negatively influenced by the intensification of precipitation and temperature extremes. In a context of progressive tropicalization of the Mediterranean area, to assist managers setting achievable restoration goals, restoration practitioners should consider that climate extremes might interfere with the beneficial effects of restoration practices.
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Although bacteria within the Verrucomicrobia phylum are pervasive in soils around the world, they are under-represented in both isolate collections and genomic databases. Here, we describe a single verrucomicrobial group within the class Spartobacteria that is not closely related to any previously described taxa. We examined more than 1,000 soils and found this spartobacterial phylotype to be ubiquitous and consistently one of the most abundant soil bacterial phylotypes, particularly in grasslands, where it was typically the most abundant. We reconstructed a nearly complete genome of this phylotype from a soil metagenome for which we propose the provisional name 'Candidatus Udaeobacter copiosus'. The Ca. U. copiosus genome is unusually small for a cosmopolitan soil bacterium, estimated by one measure to be only 2.81 Mbp, compared to the predicted effective mean genome size of 4.74 Mbp for soil bacteria. Metabolic reconstruction suggests that Ca. U. copiosus is an aerobic heterotroph with numerous putative amino acid and vitamin auxotrophies. The large population size, relatively small genome and multiple putative auxotrophies characteristic of Ca. U. copiosus suggest that it may be undergoing streamlining selection to minimize cellular architecture, a phenomenon previously thought to be restricted to aquatic bacteria. Although many soil bacteria need relatively large, complex genomes to be successful in soil, Ca. U. copiosus appears to use an alternative strategy, sacrificing metabolic versatility for efficiency to become dominant in the soil environment. © 2016 Macmillan Publishers Limited, part of Springer Nature. All rights reserved.
Chapter
This chapter draws a link between the volume-oriented forestry measures and the biomass measures used in production ecology. Tools and rules of thumb will be provided to convert the forestry growth and yield values into primary productivity and production efficiencies, and vice versa. In ecological studies in the temperate zone of central Europe, direct physiological measurements of the gas exchange in plants, and indirect estimations based on energy balances, litter production or evaporation rates provide approximate values for gross primary production (GPP) of 20–40 tons ha-1 year-1 or 2–4 kg m-2 year-1 and net primary production (NPP) of 10–20 tons ha-1 year-1 or 1–2 kg m-2 year-1 respectively. Forest inventories and long-term experimental plots, on the other hand, report an average growth of merchantable wood volume of 10–20 m3 ha-1 year-1. This chapter explains how to convert wood volume growth data to NPP through multiplication with the specific wood density, biomass expansion factors and turnover rates, and to GPP by adding respiration.
Chapter
The introduction points out the very specific system characteristics of trees and forest stands which determine the approach and methods to analyse and model forest stand dynamics, growth and yield. Like other disciplines, forest science looks with an ever-increasing spatial and temporal resolution on the functions and structures of woodlands, forest stands, trees, tree organs, plant cells…. This makes it necessary to discuss concepts for bridging the growing gap between an increasing amount of knowledge on structures and processes on temporarily and spatially highly resolved scales (chemical, biochemical, molecular-biological processes), and an information deficit on more strongly aggregated system levels (stand, ecosystem, landscape scale). The reconciliation between progressing reductionism on the one hand and the demand for holistic system knowledge for forest ecosystem management on the other is a general challenge to forest science and other “green sciences”.
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: We examined the concordance between vascular plants and macrofungi (grouped into trophic groups) in Mediterranean forest habitats (central Italy). Our goal was to test how consistently plant and fungi groups classify plots in a broadleaf deciduous forest dominated by Quercus cerris. Our hypothesis was that groups of plants can be used as surrogates for the classification of macrofungal communities. The test of concordance comprised two steps: 1) the plant species data sets were subjected to cluster analysis, to obtain three classifications based on presence of all plants, presence and frequency of only woody species; 2) Multiple Response Permutation Procedures (MRPP) was used to test the performance of each plant classification applied to the fungi data sets. The sample scores along the first PCA axis was used to investigate the relationships between compositional patterns. In the concordance analysis, the classification of only woody plants provided better results than the classification obtained with herbaceous and woody plants together. Cross-tests gave the best results when the “woody plants” classification was applied to ectomycorrhizal fungi (EMF) and, to a certain extent, to humicolous saprotrophs (Sh). The ordination analysis indicated that the frequency of woody plants follows a similar spatial distribution to EMF and Sh fungal groups and is therefore expected to discriminate the same environmental gradients. Many EMF exhibit preferences for few (one or two) hosts. Significant associations were found among a greater number of EMF and woody plant species. Woody plants such as Sorbus domestica and Prunus spinosa appear to be associated with many EMF. The combination of a high frequency of Fraxinus oxycarpa and Quercus petraea seems to promote distinct assemblages of EMF and Sh fungi. Then, characteristic assemblages of fungi were found in association with certain tree and shrub combinations.
Article
Anthropogenic disturbances are detrimental to the functioning and stability of natural ecosystems. Critical ecosystem processes driven by microbial communities are subjected to these disturbances. Here, we examine the stabilizing role of bacterial diversity on community biomass in the presence of abiotic perturbations such as addition of heavy metals, NaCl and warming. Bacterial communities with a diversity gradient of 1-12 species were subjected to the different treatments, and community biomass (OD600) was measured after 24 h. We found that initial species richness and phylogenetic structure impact the biomass of communities. Under abiotic perturbations, the presence of tolerant species in community largely contributed in community biomass production. Bacterial diversity stabilized the biomass across the treatments, and differential response of bacterial species to different perturbations was the key reason behind these effects. The results suggest that biodiversity is crucial for maintaining the stability of ecosystem functioning and acts as ecological insurance under abiotic perturbations. Biodiversity in natural ecosystems may also uphold the ecosystem functioning under anthropogenic disturbance.The ISME Journal advance online publication, 13 June 2014; doi:10.1038/ismej.2014.91.
Article
The most appropriate strategy to be used to create a permutation distribution for tests of individual terms in complex experimental designs is currently unclear. There are often many possibilities, including restricted permutation or permutation of some form of residuals. This paper provides a summary of recent empirical and theoretical results concerning available methods and gives recommendations for their use in univariate and multivariate applications. The focus of the paper is on complex designs in analysis of variance and multiple regression (i.e., linear models). The assumption of exchangeability required for a permutation test is assured by random allocation of treatments to units in experimental work. For observational data, exchangeability is tantamount to the assumption of independent and identically distributed errors under a null hypothesis. For partial regression, the method of permutation of residuals under a reduced model has been shown to provide the best test. For analysis of variance, one must first identify exchangeable units by considering expected mean squares. Then, one may generally produce either (i) an exact test by restricting permutations or (ii) an approximate test by permuting raw data or some form of residuals. The latter can provide a more powerful test in many situations.La stratégie la plus appropriée pour générer une distribution de permutation en vue de tester les termes individuels d'un plan expérimental complexe n'est pas évidente à l'heure actuelle. Il y a souvent plusieurs options, dont la permutation restreinte et la permutation d'une quelconque forme des résiduels. On trouvera ici un résumé d'informations récentes empiriques et théoriques sur les méthodes disponibles, ainsi que des recommandations pour leur utilisation dans des applications unidimensionnelles et multidimensionnelles. L'emphase est mise sur les plans complexes d'analyse de variance et de régression multiple (i.e. les modèles linéaires). Dans un travail expérimental, la supposition d'échangeabilité requise pour un test par permutation est assurée par l'assignation au hasard à des unités des divers traitements. Dans le cas d'observations, l'échangeabilité équivaut à supposer que les erreurs, dans une hypothèse nulle, sont indépendantes et distribuées de façon identique. Pour la régression partielle, la méthode de permutation des résiduels dans un modèle réduit s'est avérée la meilleure. Pour l'analyse de variance, il faut d'abord identifier les unités échangeables à l'examen des carrés moyens attendus. Ensuite, il est généralement possible de produire (i) un test exact en restreignant les permutations ou alors (ii) un test approximatif en permutant les données brutes ou une forme quelconque des résiduels. Cette dernière méthode fournit, dans plusieurs situations, un test plus puissant.[Traduit par la Rédaction]
Article
Forest management activities influence forest structure and development, and thereby influence biodiversity. We investigated diversity and function of macrofungi and soil microbes in a Chamaecyparis formosensis forest with light and heavy thinning treatments and control plots, and we compared them with an adjacent broadleaf forest. Principal component analysis was used to separate macrofungal species components exposed to thinning treatments and those in the broadleaf forest. The macrofungal diversity in the broadleaf forest was higher than in the plantations, and the diversity of fungi from the lightly thinned plantation was higher than those from the heavily thinned and unthinned plantations. Denaturing gradient gel electrophoresis profiles showed that soil fungal and bacterial communities were significantly different among the treatments. However, the carbon utilisation patterns of soil microbes did not differ significantly among treatments. Thinning increased the diversity of saprotrophic macrofungi and changed the soil microbial community of the plantation. This finding might be due to the functional redundancy of soil microbes, as there was no significant thinning effect on the soil microbial function through the fourth year after thinning.
Article
Several permutation strategies are often possible for tests of individual terms in analysis-of-variance (ANOVA) designs. These include restricted permutations, permutation of whole groups of units, permutation of some form of residuals or some combination of these. It is unclear, especially for complex designs involving random factors, mixed models or nested hierarchies, just which permutation strategy should be used for any particular test. The purpose of this paper is two-fold: (i) we provide a guideline for constructing an exact permutation strategy, where possible, for any individual term in any ANOVA design; and (ii) we provide results of Monte Carlo simulations to compare the level accuracy and power of different permutation strategies in two-way ANOVA, including random and mixed models, nested hierarchies and tests of interaction terms. Simulation results showed that permutation of residuals under a reduced model generally had greater power than the exact test or alternative approximate permutation methods (such as permutation of raw data). In several cases, restricted permutations, in particular, suffered more than other procedures, in terms of loss of power, challenging the conventional wisdom of using this approach. Our simulations also demonstrated that the choice of correct exchangeable units under the null hypothesis, in accordance with the guideline we provide, is essential for any permutation test, whether it be an exact test or an approximate test. For reference, we also provide appropriate permutation strategies for individual terms in any two-way or three-way ANOVA for the exact test (where possible) and for the approximate test using permutation of residuals.