Changes In Species Composition Research Articles

Proportions of taxa belonging to the gut core microbiome change throughout the life cycle and season of the bark beetle Ips typographus.

The European spruce bark beetle, Ips typographus, is a serious pest of spruce forests in Europe, and its invasion and development inside spruce tissues are facilitated by microorganisms. We investigated the core gut bacterial and fungal microbiomes of I. typographus throughout its life cycle in spring and summer generations. We used cultivation techniques and molecular identification in combination with DNA and RNA metabarcoding. Our results revealed that communities differ throughout their life cycle and across generations in proportion of dominantly associated microbes, rather than changes in species composition. The bacteriome consisted mostly of the phylum Gammaproteobacteria, with the most common orders and genera being Enterobacteriales (Erwinia and Serratia), Pseudomonadales (Pseudomonas), and Xanthomonadales. The fungal microbiome was dominated by yeasts (Saccharomycetes-Wickerhamomyces, Kuraishia, and Nakazawaea), followed by Sordariomycetes (Ophiostoma bicolor and Endoconidiophora polonica). We did not observe any structure ensuring long-term persistence of microbiota on any part of the gut epithelium, suggesting that microbial cells are more likely to pass through the beetle's gut with chyme. The most abundant taxa in the beetle's gut were also identified as dominant in intact spruce phloem. Therefore, we propose that these taxa are acquired from the environment rather than specifically vectored between generations.

Associations between ecological diversity and rodent plague circulation in Yunnan Province, China, 1983-2020: A data-informed modelling study.

Following its resurgence in 1982, rodent plague has been linked to a wide range of circulation risks in Yunnan Province. The most serious public health concern associated with effective plague control is determining how various ecological variables influence the differential risk of transmission. We investigated the population dynamics of the hosts and vectors using large-scale epidemiological surveillance data. In a seasonal eco-epidemiological model, we evaluated the impact of ecological conditions on the vectored flea index (VFI) to determine the rate of plague transmission. The findings revealed a changing species composition in natural foci over time. Additionally, shifting distributional ranges of species by elevation may be vital in modulating the VFI. The model estimates indicate that the dynamic VFI contributes to spatiotemporal variance in transmission. The VFI could be a critical ecological indicator, allowing for real-time tracking and prompt intervention in the circulation of rodent plague. Understanding eco-epidemiological diversity can provide essential insights into effective responses to future plague resurgence.

ZOOPLANKTON CURRENT STATE AND TROPHIC RELATIONSHIPS IN THE POSOLSKY SOR OF BAIKAL LAKE IN 2022

The present study provides the original results of research of zooplankton in the most typical shallow area of Lake Baikal (Posolsky Sor) in 2022. Changes in species composition, quantitative indicators of development and the structure of zooplankton are analyzed. Since this area of Baikal is considered a feeding ground for juvenile omul, special attention is paid to assessing the productivity of zooplankton as a food supply for fi sh.

Half a century of herb layer changes in Quercus-dominated forests of the Western Carpathians

Species composition of European temperate Quercus forests have undergone dramatic changes in the past century. Observed changes have largely been attributed to anthropogenic environmental changes that drove the processes of thermophilization, eutrophication, or biotic homogenization. However, ecological processes behind these changes can be difficult to explain considering the variability among regions: their history, management, landscape patterns, site properties, and vegetation types. To reveal vegetation change trends and their drivers in the subcontinental Western Carpathians, we analysed vegetation and environmental data from 158 quasi-permanent plots established between 1961 and 1980 in Quercus petraea- dominated forests on the andesite bedrock of the Slovenské stredohorie Mountains in Slovakia (Central Europe), and we resurveyed them again between 2005 and 2020.Among alpha and gamma diversity measures, we found only slight species richness decreases. A significant shift in species composition associated with very high species turnover (64%) and a decrease in herb layer cover (by 24%) resulted in higher spatial heterogeneity of herb understoreys in the resurvey. Winning species were more thermophilous, eutrophic, and mesic/oceanic plants of lower latitudes, replacing original species of open Quercus forests. Increased temperatures was the most significant environmental driver. Climate warming corresponded not only to thermophilization, but, after interaction with management, it was also related to eutrophication of the herb layer. Eutrophication was identified as the main trend of the species composition changes (mean EIV for nutrients of winners was 5.9 compared to 4.1 of losers) even though the effect of anthropic nitrogen depositions was negligible. Successive undergrowing of Quercus canopies by more shade-casting trees (Acer campestre, Carpinus betulus, and Fagus sylvatica) in the period of decreasing management intensity was another major driver. Their increase by 8.6% in the resurveyed plots contributed to decreased species richness and they supported shade-tolerant eutrophic understorey species typical of mesic Carpinus betulus and Fagus sylvatica forests.We concluded that Quercus forests in the Western Carpathians underwent thermophilization and eutrophication, similar to the forests in densely inhabited Western Europe; however, their plant understoreys were not homogenized, and eutrophication was not driven by nitrogen depositions. Eutrophication should be recognised as the next important negative impact of global warming on ecosystems. Dramatic changes in species composition and loss of original characteristic species can only be partly halted by reducing shade-casting trees from lower tree layers in managed stands.

Ecological Factors Driving Tree Diversity across Spatial Scales in Temperate Forests, Northeast China

Species diversity has been shown to be influenced by environmental factors, but the mechanism underlying their relationship remains unclear across spatial scales. Based on field investigation data collected from 3077 sample plots in temperate forest ecosystems, we compared tree species richness, evenness and dominance at 10 km × 10 km, 30 km × 30 km and 90 km × 90 km spatial scales. Then, we detected the scale dependence of changes in tree species composition on climate, topography and forest structure using variation partitioning and quantified their contribution to tree diversity with gradient–boosted models (GBMs) and fitted their relationships. The magnitude of tree richness, evenness and dominance significantly increased with spatial scale. Ecological factors jointly accounted for 24.3%, 26.5% and 38.5% of the variation in tree species composition at the three spatial scales, respectively. The annual mean temperature had a strong impact on tree richness, evenness and dominance and peaked at an intermediate scale. Tree evenness and dominance increased with the variation of temperature but had upper and lower limits. Tree richness obviously increased with annual precipitation on multiple scales and decreased with annual sunshine duration at large spatial scales. Tree richness, evenness and dominance obviously increased with the variation in elevation and diameter at breast height at large scales and small scales, respectively. Tree dominance decreased with tree height at a small scale. The dependence of tree diversity on ecological factors increased with spatial scales. Furthermore, different factors exert various controls on tree diversity at different spatial scales, representing a comprehensive mechanism regulating tree diversity.

The biogeography of Gabonese savannas: Evidence from termite community richness and composition

AbstractAimThe mosaic of savannas that persists in the forest‐dominant Congo Basin is thought to be palaeoclimatic relics, but past biogeographical processes that have formed and maintained these systems are poorly understood. Here, we explored the post‐Pleistocene biogeography of Gabon's savannas using termites as biological indicators to understand historical and mechanistic factors influencing present‐day termite communities in the country's extant savannas.LocationGabon, Central Africa.TaxonBlattodea: Termitoidae.MethodsUsing standardised transect methods, we sampled termite communities in four disjunct modern savanna areas of Gabon: the centre (Lopé), the southeast (Batéké) and the south (Mayombe North and South). Termites at Lopé were collected in three habitats (annually burned savannas, savannas with a depressed fire regime and forest). We used DNA barcoding of the COII region to identify termite species and compared abundance, species richness and community composition across areas and habitats.ResultsCommunity composition differed greatly between Lopé and both Batéké and Mayombe savannas with Lopé being exceptionally depauperate and lacking characteristic savanna species. Within Lopé, termite abundance and diversity was highest in forests and lowest in annually burned savannas, with a gradual change in species composition across the forest–savanna gradient associated with fire history.Main ConclusionsThe absence of savanna typical species in Lopé savannas challenges current assumptions that these savannas were linked to the south/southeastern savannas during the Pleistocene and suggests a different evolutionary history. Lopé savannas may instead have opened as an isolated grassland and never have been contiguous with neighbouring savannas, or were isolated soon after forest expansion began and have now lost savanna‐typical species. Furthermore, the patterns of termite community composition in fire suppressed savannas support a hypothesis of rapid change driven by fire frequency where either fire suppression or infrequent burning over 23 years has meant savannas have become ecologically much more forest‐like.

Moderate chlorophyll-a environments reduce coral bleaching during thermal stress in Yap, Micronesia

Thermal-stress events on coral reefs lead to coral bleaching, mortality, and changes in species composition. The coral reefs of Yap, in the Federated States of Micronesia, however, remained largely unaffected by major thermal-stress events until 2020, when temperatures were elevated for three months. Twenty-nine study sites were examined around Yap to determine geographical and taxonomic patterns of coral abundance, bleaching susceptibility, and environmental predictors of bleaching susceptibility. Island-wide, 21% (± 14%) of the coral cover was bleached in 2020. Although inner reefs had a greater proportion of thermally-tolerant Porites corals, the prevalence of bleaching was consistently lower on inner reefs (10%) than on outer reefs (31%) for all coral taxa. Corals on both inner and outer reefs along the southwestern coast exhibited the lowest prevalence of coral bleaching and had consistently elevated chlorophyll-a concentrations. More broadly, we revealed a negative relationship between bleaching prevalence and (moderate) chlorophyll-a concentrations that may have facilitated resistance to thermal stress by reducing irradiance and providing a heterotrophic energy source to benefit some corals exposed to autotrophic stress. Southwestern reefs also supported a high but declining fish biomass, making these bleaching-resistant and productive reefs a potential climate-change refuge and a prime target for conservation.

Changes of Key Soil Factors, Biochemistry and Bacterial Species Composition during Seasons in the Rhizosphere and Roots of Codonopsis pilosula (tangshen)

Codonopsis pilosula is a medicinal and edible herb with a rich nutritional value. In Gansu Province, China, its production quality and yield differ during the four seasons. Here, we investigated the differences in the microfloral composition and metabolic functions in the rhizospheric soil and roots of C. pilosula during the four seasons, and we also analyzed their dynamic and synergistic effects on C. pilosula growth and carbohydrate content change. The C. pilosula samples were analyzed for plant physiology, microfloral composition and metabolic functions in the rhizospheric soil and roots using high-throughput sequencing technology. Environmental indices including soil physiochemistry and meteorological conditions were also determined by the coupled chromatography–spectroscopy technique. The results revealed that the C. pilosula growth was affected by temperature, precipitation and light intensity, with the bacterial structures and functions of the soil and root samples showing obvious seasonal changes. Due to the diversity of microbial composition and community metabolic function, and the synergistic effect of microbial and environmental factors, there are significant differences in stress resistance, physiological status and metabolites of C. pilosula in different seasons. Furthermore, the change in seasons was significantly correlated with the quality and yield of C. pilosula. This study provides a scientific basis for soil improvement and the refinement of local Radix C. pilosula cultivation methods.

Consumers decrease variability across space and turnover through time during coral reef succession.

Consumers play an integral role in mediating ecological succession-the change in community composition over time. As consumer populations are facing rapid decline in ecosystems around the world, understanding of their ecological role is becoming increasingly urgent. Increased understanding of how changes in consumer populations may influence community variability across space and turnover through time during succession is particularly important for coral reefs, which are among the most threatened ecosystems globally, and where fishes play vital roles in structuring benthic succession. Here, we examine how consumers influence coral reef succession by deploying 180 paired settlement tiles, caged (to exclude fishes larger than approximately 15cm) and uncaged, within Palmyra Atoll, a remote marine wildlife refuge with previously documented high fish abundance, and monitored benthic community development one and three years after deployment. We found that excluding large fishes lead to lower alpha diversity and divergent community states across space (i.e.,, high beta diversity among caged tiles), suggesting that benthic fish feeding maintains local diversity but tends to homogenize community composition with dominance by crustose coralline algae. In addition, when fish were experimentally excluded, the developing benthic community exhibited a greater change in species composition over time (i.e., high temporal beta diversity), indicating that fish feeding tends to canalize community successional trajectories. Finally, the caged and uncaged tiles became more similar over time, suggesting that fish feeding plays a more important role during early succession. Our results demonstrate that the loss of large fishes, for example from overfishing, may result in benthic communities that are more variable across space and time. Increased variability could have important implications for ecosystem function and coral reef resilience in the face of escalating global stressors.

Changes in Species Composition and Growth in a Restored Maple-Basswood Forest after 30 Years

Changes in Species Composition and Growth in a Restored Maple-Basswood Forest after 30 Years

Indirect interactions between pollinators drive interaction rewiring through space

AbstractIn recent years, an extended body of literature has focused on the importance of either temporal or spatial dynamics in shaping the structure of interacting plant and pollinator communities. This improvement from a previously static and aggregated perspective has allowed us to understand many of the ecological processes that shape community assembly. However, fewer are the studies that have simultaneously focused on spatial and temporal dynamics, and even fewer are those that collect data across different habitat types to assess the generality of their findings. Here, we used a dataset collected weekly throughout the full flowering season for two consecutive years and within two contrasting habitat types in N and SW Spain: a mountain grassland area and the understory of sparse pine forests. We evaluated species and interaction persistence through space and time, pollinator fidelity, and turnover patterns in interaction composition while providing a potential mechanistic explanation for the patterns observed. Our results show that although species generalization does not explain species or interaction persistence, moderately generalist species are those showing the greatest fidelity to the subset of plant species they visit through space and time. Further, we find that interaction turnover through time is mostly driven by changes in species composition, while through space it is mostly driven by interaction rewiring resulting from indirect competitive interactions between pollinator species. Our results help to shed light on the potential mechanisms driving community assembly patterns beyond niche or neutral processes by adding within‐trophic‐level interactions that can modify pollinator preferences.

Macrophyte communities as bioindicator of stormwater pollution in rivers: a quantitative analysis.

Macrophytes are one of the important indicators used in assessing the anthropic impact on aquatic ecosystems. The structure of macrophyte communities of two rivers were compared by species composition, dominant species and projective cover using statistical methods. It is shown that the influence of storm runoff on these rivers is manifested in the form of a change in the dominant species composition. Based on the statistical analysis carried out, it can be argued that, despite the peculiarities of the flora composition of each of the rivers, the influence of storm runoffs largely neutralizes this specificity, determining the situation in local areas immediately below the runoff. In the area of the effluent discharge the dominance of individual species and an increase in the area overgrown with macrophytes was observed. In the area of stormwater discharge on the Psel River, species were usually present: Nuphar lutea, Ceratophyllum demersum, Myriophyllum spicatum and on the Bystrica River-Glyceria maxima, Sagitaria sagittiformis, Stuckenia pectinata and Potamogeton crispus. The use of the NMDS method has been found to provide good insight into the structural rearrangements in macrophyte communities affected by runoff from stormwater systems.

Variations in soil microbial community structure and extracellular enzymatic activities along a forest-wetland ecotone in high-latitude permafrost regions.

Permafrost degradation by global warming is expected to alter the hydrological processes, which results in changes in vegetation species composition and gives rise to community succession. Ecotones are sensitive transition areas between ecosystem boundaries, attract particular interest due to their ecological importance and prompt responses to the environmental variables. However, the characteristics of soil microbial communities and extracellular enzymes along the forest-wetland ecotone in high-latitude permafrost region remain poorly understood. In this study, we evaluated the variations of soil bacterial and fungal community structures and soil extracellular enzymatic activities of 0-10 cm and 10-20 cm soil layers in five different wetland types along environmental gradients, including Larix gmelinii swamp (LY), Betula platyphylla swamp (BH), Alnus sibirica var. hirsute swamp (MCY), thicket swamp (GC), and tussock swamp (CC). The relative abundances of some dominant bacterial (Actinobacteria and Verrucomicrobia) and fungal (Ascomycota and Basidiomycota) phyla differed significantly among different wetlands, while bacterial and fungal alpha diversity was not strongly affected by soil depth. PCoA results showed that vegetation type, rather than soil depth explained more variation of soil microbial community structure. β-glucosidase and β-N-acetylglucosaminidase activities were significantly lower in GC and CC than in LY, BH, and MCY, while acid phosphatase activity was significantly higher in BH and GC than LY and CC. Altogether, the data suggest that soil moisture content (SMC) was the most important environmental factor contributing to the bacterial and fungal communities, while extracellular enzymatic activities were closely related to soil total organic carbon (TOC), nitrate nitrogen () and total phosphorus (TP).

Correlated biodiversity change between plant and insect assemblages resurveyed after 80 years across a dynamic habitat mosaic.

Historical data on co-occurring taxa are extremely rare. As such, the extent to which distinct co-occurring taxa experience similar long-term patterns in species richness and compositional change (e.g., when exposed to a changing environment) is not clear. Using data from a diverse ecological community surveyed in the 1930s and resurveyed in the 2010s, we investigated whether local plant and insect assemblages displayed cross-taxon congruence-that is, spatiotemporal correlation in species richness and compositional change-across six co-occurring taxa: vascular plants, non-vascular plants, grasshoppers and crickets (Orthoptera), ants (Hymenoptera: Formicinae), hoverflies (Diptera: Syrphidae), and dragonflies and damselflies (Odonata). All taxa exhibited high levels of turnover across the ca. 80-year time period. Despite minimal observed changes at the level of the whole study system, species richness displayed widespread cross-taxon congruence (i.e., correlated temporal change) across local assemblages within the study system. Hierarchical logistic regression models suggest a role for shared responses to environmental change underlying cross-taxon correlations and highlight stronger correlations between vascular plants and their direct consumers, suggesting a possible role for biotic interactions between these groups. These results provide an illustration of cross-taxon congruence in biodiversity change using data unique in its combination of temporal and taxonomic scope, and highlight the potential for cascading and comparable effects of environmental change (abiotic and biotic) on co-occurring plant and insect communities. However, analyses of historical resurveys based on currently available data come with inherent uncertainties. As such, this study highlights a need for well-designed experiments, and monitoring programs incorporating co-occurring taxa, to determine the underlying mechanisms and prevalence of congruent biodiversity change as anthropogenic environmental change accelerates apace.

Looking back on biodiversity change: lessons for the road ahead.

Estimating biodiversity change across the planet in the context of widespread human modification is a critical challenge. Here, we review how biodiversity has changed in recent decades across scales and taxonomic groups, focusing on four diversity metrics: species richness, temporal turnover, spatial beta-diversity and abundance. At local scales, change across all metrics includes many examples of both increases and declines and tends to be centred around zero, but with higher prevalence of declining trends in beta-diversity (increasing similarity in composition across space or biotic homogenization) and abundance. The exception to this pattern is temporal turnover, with changes in species composition through time observed in most local assemblages. Less is known about change at regional scales, although several studies suggest that increases in richness are more prevalent than declines. Change at the global scale is the hardest to estimate accurately, but most studies suggest extinction rates are probably outpacing speciation rates, although both are elevated. Recognizing this variability is essential to accurately portray how biodiversity change is unfolding, and highlights how much remains unknown about the magnitude and direction of multiple biodiversity metrics at different scales. Reducing these blind spots is essential to allow appropriate management actions to be deployed. This article is part of the theme issue 'Detecting and attributing the causes of biodiversity change: needs, gaps and solutions'.

Intensive Meadows on Organic Soils of Temperate Climate–Useful Value of Grass Mixtures after the Regeneration

Meadows on organic soils perform an essential role as a source of fodder and biomass for energy purposes. In the case of intensive meadows, it is crucial to use grass mixtures that enable obtaining high yields of good quality; however, on organic soils, the grass species composition changes dynamically. We carried out the full cultivation (ploughing) for grassland restoration. The floristic composition of meadow sward in the first year of full use (2013) was similar to the composition of mixtures used for sowing (2012) individual plots. Festuca arundinacea and Phleum pratense showed greater resistance to low temperatures in winter and excess water in spring compared to Lolium perenne. In comparison, we obtained the highest yield (fresh and dry mass) from mixtures with Festuca arundinacea and Lolium perenne. We obtained the best quality forage from the first swath and the object with the highest share of Lolium perenne. For grassland restoration in the analysed habitat, it is reasonable to use grass mixtures with varied compositions, in which the share of Lolium perenne is between 25–50%.

Cumulative effects of biomass harvesting and herbicide application on litter-dwelling arthropod communities in jack pine-dominated forests: 7th year postharvest assessment

Forest biomass harvest has the potential to provide feedstocks for energy production to offset fossil fuel consumption. However, concerns have been raised regarding the ecological sustainability of removing additional biomass from forests, in particular the impacts on biodiversity. In this paper, we used a suite of ground-dwelling arthropod taxa (ground beetles, spiders, and rove beetles) to measure community compositional changes along a gradient of biomass removal treatments 7 years postharvest, and compared against reference. Based on multivariate regression trees, changes in species composition reflected the intensity gradient of the biomass removal treatments or stand attributes associated with the level of forest floor disturbance across all arthropod groups. For each arthropod group, changes in composition were defined primarily by reductions or loss of abundant forest associated species and increases in the number and abundance of species associated with more xeric conditions and increased disturbance intensity. There were no differences between full-tree and tree-length treatments. Overall, results indicated a strong arthropod response to the removal of overstory, forest floor disturbance, and reductions in understory cover mostly resulting from the glyphosate applications. Arthropod recovery would benefit from overstory retention, reduction in forest floor disturbance, and judicious use of glyphosate.

Anchoring grassland sustainability with a nature-based small burrowing mammal control strategy.

Over the last 40 years, a burrowing mammal eradication policy has been prevalent on the Qinghai-Tibetan Plateau (QTP). This policy is based on similar burrowing mammal eradication programs in other areas and is justified on the assumptions that burrowing mammals compete with livestock for forage and contribute to grassland degradation. However, there is no clear theoretical or experimental evidence supporting these assumptions. This paper synthesizes the ecological functioning of small burrowing mammals in natural grasslands and discusses the irrationality and consequences of burrowing mammal eradication for sustainable livestock grazing and grassland degradation. Past burrowing mammal eradication efforts have failed because increased food availability for the remaining rodents and reduced predator populations led to rapid population rebounds. Herbivores differ in diet, and there is clear evidence that burrowing mammals, especially plateau zokors Myospalax baileyi, have a different diet than livestock. In QTP meadows, burrowing mammal eradication induces a shift towards plant communities with fewer species preferred by livestock and more species preferred by burrowing mammals. Thus, eradicating burrowing mammals has the opposite effect, a reduction in livestock preferred vegetation. We suggest that the policy of poisoning burrowing mammals needs to be reconsidered and revoked as soon as possible. We argue that incorporating density-dependent factors such as predation and food availability are essential for maintaining a low burrowing mammal density. For degraded grasslands, we suggest that the optimal sustainable approach is to decrease the intensity of livestock grazing. Lower grazing induces changes in vegetation structure and plant species composition that increases predation on burrowing mammals and decreases the abundance of plants preferred by burrowing mammals. Such a nature-based grassland management system maintains the density of burrowing mammals at a low stable density while minimizing human management and interventions.

Identifying high snakebite risk area under climate change for community education and antivenom distribution

Snakebite is one of the largest risks from wildlife, however little is known about venomous snake distribution, spatial variation in snakebite risk, potential changes in snakebite risk pattern due to climate change, and vulnerable human population. As a consequence, management and prevention of snakebite is hampered by this lack of information. Here we used habitat suitability modeling for 10 medically important venomous snakes to identify high snakebite risk area under climate change in Iran. We identified areas with high snakebite risk in Iran and showed that snakebite risk will increase in some parts of the country. Our results also revealed that mountainous areas (Zagros, Alborz, Kopet–Dagh mountains) will experience highest changes in species composition. We underline that in order to improve snakebite management, areas which were identified with high snakebite risk in Iran need to be prioritized for the distribution of antivenom medication and awareness rising programs among vulnerable human population.

Restoration of woody plant communities in red pine plantations on abandoned agricultural land in northern Minnesota, USA

While tree plantations have increased globally, there is growing interest in restoring plantations to native ecosystems. Restoration efforts may be more successful when tree species native to original ecosystems are planted instead of species that are either introduced or native, but off site.Plantations of red pine (Pinus resinosa Ait.) in the western Great Lakes region of North America were commonly established on red pine sites, often after agricultural clearing and abandonment. Red pine is the only species in the overstory, compared to 10 or more species in red pine woodlands. Plantations differ in structure from woodlands, being single-aged versus multi-cohort and having high and spatially-uniform density versus lower and variable density.Our aim was to contrast changes in structure and woody species composition over a 15-year period in response to two silvicultural approaches in red pine plantations on the Red Lake Wildlife Management Area in northwestern Minnesota, USA. Approaches included: 1) uniform thinning followed by a dispersed retention harvest (dispersed treatment) and 2) variable density thinning followed by an expanding gap irregular shelterwood (aggregated treatment).Basal area of trees ≥10 cm dbh was reduced similarly in both treatments to 9 m2 ha−1, but there was high within-stand variability (coefficient of variation) in the aggregated treatment (>100%) compared to the dispersed treatment (20%). Understory woody species abundance increased with both treatments, including sapling, large regeneration, and small regeneration layers, with >100% increases. In the sapling layer, the increased abundance (to 2.5 m2 ha−1) reflected new recruitment into this layer, as there were few saplings before treatment (<0.5 m2 ha−1). By year 15, the majority of saplings were canopy capable species, while lower regeneration layers contained mixtures of canopy trees, sub-canopy trees, and shrubs.Composition in the understory changed, but was similar between treatments, including increases in woody shrubs in the small regeneration layer and increases in red pine regeneration in the large regeneration layer. The sapling layer had increased importance of several native tree species, especially in the aggregated treatment.Both of our silvicultural approaches moved structurally simple, species poor red pine plantations toward a condition reflecting native red pine woodlands, which had mixed-species composition and variably open canopy cover. The treatment using variable density thinning and an irregular shelterwood resulted in greater spatial variability of structure and greater importance of shade-intolerant tree species, both characteristics of native woodlands. While focused on red pine, this approach should have application to conifer restoration more generally.