Holocene shifts in the assembly of plant and animal communities implicate human impacts.

Understanding how ecological communities are organized and how they change through time is critical to predicting the effects of climate change. Recent work documenting the co-occurrence structure of modern communities found that most significant species pairs co-occur less frequently than would be expected by chance. However, little is known about how co-occurrence structure changes through time. Here we evaluate changes in plant and animal community organization over geological time by quantifying the co-occurrence structure of 359,896 unique taxon pairs in 80 assemblages spanning the past 300 million years. Co-occurrences of most taxon pairs were statistically random, but a significant fraction were spatially aggregated or segregated. Aggregated pairs dominated from the Carboniferous period (307 million years ago) to the early Holocene epoch (11,700 years before present), when there was a pronounced shift to more segregated pairs, a trend that continues in modern assemblages. The shift began during the Holocene and coincided with increasing human population size and the spread of agriculture in North America. Before the shift, an average of 64% of significant pairs were aggregated; after the shift, the average dropped to 37%. The organization of modern and late Holocene plant and animal assemblages differs fundamentally from that of assemblages over the past 300 million years that predate the large-scale impacts of humans. Our results suggest that the rules governing the assembly of communities have recently been changed by human activity.

Large, connected floodplain forests prone to flooding best sustain plant diversity.

Dams, levees, and water withdrawals disrupt hydrologic regimes and associated floodplain forests. Because these forests are also responding to changes in land use, species invasions, and climate change, the relative effects of these factors are hard to disentangle. Most studies of floodplain forests lack historic data, requiring us to rely on recent data or contemporary spatial relationships to these drivers to infer those causes of vegetation dynamics. Here, we use survey data from the 1950s to reconstruct plant community changes across 40 floodplain forests in Wisconsin. We applied two partial least squares regression (PLS) models to evaluate how current site and landscape scale conditions and changes in these conditions since the 1950s influence contemporary patterns of community diversity and composition. Local site variables were among the most important in explaining current composition metrics and their changes, but historic landscape variables and changes in these were also important. Current local diversity (α) was the highest at sites prone to frequent flooding, even at sites in fragmented landscapes. Sites along sinuous rivers in large watershed areas with more contiguous forest had the highest abundance of wetland indicator plants in the re-survey and had the largest increases in α diversity since the 1950s, despite having the highest presence of exotic species then. These same sites have converged in composition, reflecting increases in wetland indicator plants and common native species. These patterns of increasing α diversity coupled with declines in community distinctiveness are uncommon among long-term studies. Increases in wetland plants may indicate that sites have become wetter with hydrologic changes, but these increases may also reflect improved colonization and establishment processes involving a robust regional pool of generalist wetland taxa. Woody and exotic plants typical of upland forests increased at rarely flooded sites in fragmented and urbanizing landscapes, indicating shifts towards a later-successional conditions and a dampened disturbance regime. This has reduced local species diversity and increased regional distinctness at some sites. As hydrologic connections appear to best maintain native species diversity and composition, even in fragmented landscapes, managers should seek to recreate these whenever feasible.

Trait--environment relationships remain strong despite 50 years of trait compositional change in temperate forests.

Temperate North American forest communities have changed considerably in response to logging, fragmentation, herbivory, and other global change factors. Significant changes in the structure and composition of seemingly undisturbed Wisconsin forest communities have occurred over the past 50 years, including widespread declines in alpha and beta species diversity. To investigate how shifts in species composition have affected distributions of plant functional traits, we first compiled extensive data on understory plant species traits. We then computed community-weighted trait means and functional diversity metrics for communities in both the 1950s and 2000s. We examined how trait values and diversity varied across environmental gradients and among Wisconsin's four main ecoregions. Trait means and diversity values reflect conspicuous gradients in species composition, soils, and climatic conditions. Over the past 50 years, values of most traits have changed as communities shifted toward species with higher leaf nutrient levels and specific leaf area, particularly in the southern ecoregions. Trait richness and diversity have declined, particularly in historically species- and trait-rich unglaciated southwestern Wisconsin. Reductions in within-site trait diversity may be diminishing the ability of these forest communities to resist or resiliently respond to shifts in environmental conditions. Despite changes in trait and community composition, trait-environment relationships measured directly via fourth-corner analysis remain strong for most plant traits. Nevertheless, accelerating ecological change (including climate change) could outstrip the ability of plant species and traits to match their environment, particularly in more fragmented landscapes.

Physiological mechanisms drive differing foliar calcium content in ferns and angiosperms.

Recent evidence points to ferns containing significantly lower contents of foliar calcium and other cations than angiosperms. This is especially true of more ancient 'non-polypod' fern lineages, which predate the diversification of angiosperms. Calcium is an important plant nutrient, the lack of which can potentially slow plant growth and litter decomposition, and alter soil invertebrate communities. The physiological mechanisms limiting foliar calcium (Ca) content in ferns are unknown. While there is a lot we do not know about Ca uptake and transport in plants, three physiological processes are likely to be important. We measured transpiration rate, cation exchange capacity, and leaching loss to determine which process most strongly regulates foliar Ca content in a range of fern and co-occurring understory angiosperm species from a montane Hawaiian rainforest. We found higher instantaneous and lifetime (corrected for leaf lifespan) transpiration rates in angiosperms relative to ferns. Ferns preferentially incorporated Ca into leaves relative to strontium, which suggests that root or stem cation exchange capacity differs between ferns and angiosperms, potentially affecting calcium transport in plants. There were no differences in foliar Ca leaching loss between groups. Among the physiological mechanisms measured, foliar Ca was most strongly correlated with leaf-level transpiration rate and leaf lifespan. This suggests that inter-specific differences in a leaf's lifetime transpiration may play a significant role in determining plant nutrition.

Functional trait data for vascular plant species from northeastern North America.

Wisconsin's plant communities are responding to shifting disturbance regimes, habitat fragmentation, aerial nitrogen deposition, exotic species invasions, ungulate herbivory, and successional processes. To better understand how plant functional traits mediate species' responses to changing environmental conditions, we collected a large set of functional trait data for vascular plant species occupying Wisconsin forests and grasslands. We used standard protocols to make 76,213 measurements of 34 quantitative traits. These data provide rich information on genome size, physical leaf traits (length, width, circularity, thickness, dry matter content, specific leaf area, etc.), chemical leaf traits (carbon, nitrogen, phosphorus, potassium, calcium, magnesium, ash), life history traits (vegetative and flower heights, seed mass), and traits affecting plant palatability (leaf fiber, fat, and lignin). These trait values derive from replicate measurements on 12+ individuals of each species from multiple sites and 45+ individuals for a selected subset of species. Measurements typically reflect values for individuals although some chemical traits involved composite samples from several individuals at the same site. We also qualitatively characterized each species by plant family, woodiness, functional group, and Raunkiaer lifeform. These data allow us to characterize trait dimensionality, differentiation, and covariation among temperate plant species (e.g., leaf and stem economic syndromes). We can also characterize species' responses to environmental gradients and drivers of ecological change. With survey and resurvey data available from >400 sites in Wisconsin, we can analyze variation in community trait distributions and diversity over time and space. These data therefore allow us to assess how trait divergence vs. convergence affects community assembly and how traits may be related to half-century shifts in the distribution and abundance of these species. The data set can be used for non-commercial purposes. The data set is licensed as follows: CC-By Attribution 4.0 International. We request users cite both the OSF data set and this Ecology data paper publication.

Community assembly of coral reef fishes along the Melanesian biodiversity gradient.

The Indo-Pacific is home to Earth's most biodiverse coral reefs. Diversity on these reefs decreases from the Coral Triangle east through the islands of Melanesia. Despite this pattern having been identified during the early 20th century, our knowledge about the interaction between pattern and process remains incomplete. To evaluate the structure of coral reef fish communities across Melanesia, we obtained distributional records for 396 reef fish species in five taxa across seven countries. We used hierarchical clustering, nestedness, and multiple linear regression analyses to evaluate the community structure. We also compiled data on life history traits (pelagic larval duration, body size and schooling behavior) to help elucidate the ecological mechanisms behind community structure. Species richness for these taxa along the gradient was significantly related to longitude but not habitat area. Communities are significantly nested, indicating that species-poor communities are largely composed of subsets of the species found on species rich reefs. These trends are robust across taxonomic groups except for the Pomacentridae, which exhibit an anti-nested pattern, perhaps due to a large number of endemic species. Correlations between life history traits and the number of reefs on which species occurred indicate that dispersal and survival ability contribute to determining community structure. We conclude that distance from the Coral Triangle dominates community structure in reef fish; however, conservation of the most species-rich areas will not be sufficient alone to conserve the vivid splendor of this region.

Quantifying the Human Impacts on Papua New Guinea Reef Fish Communities across Space and Time.

Describing the drivers of species loss and of community change are important goals in both conservation and ecology. However, it is difficult to determine whether exploited species decline due to direct effects of harvesting or due to other environmental perturbations brought about by proximity to human populations. Here we quantify differences in species richness of coral reef fish communities along a human population gradient in Papua New Guinea to understand the relative impacts of fishing and environmental perturbation. Using data from published species lists we categorize the reef fishes as either fished or non-fished based on their body size and reports from the published literature. Species diversity for both fished and non-fished groups decreases as the size of the local human population increases, and this relationship is stronger in species that are fished. Additionally, comparison of modern and museum collections show that modern reef communities have proportionally fewer fished species relative to 19th century ones. Together these findings show that the reef fish communities of Papua New Guinea experience multiple anthropogenic stressors and that even at low human population levels targeted species experience population declines across both time and space.

Stoichiometry of ferns in Hawaii: implications for nutrient cycling.

We asked if element concentrations in ferns differ systematically from those in woody dicots in ways that could influence ecosystem properties and processes. Phylogenetically, ferns are deeply separated from angiosperms; for our analyses we additionally separated leptosporangiate ferns into polypod ferns, a monophyletic clade of ferns which radiated after the rise of angiosperms, and all other leptosporangiate (non-polypod) ferns. We sampled both non-polypod and polypod ferns on a natural fertility gradient and within fertilized and unfertilized plots in Hawaii, and compared our data with shrub and tree samples collected previously in the same plots. Non-polypod ferns in particular had low Ca concentrations under all conditions and less plasticity in their N and P stoichiometry than did polypod ferns or dicots. Polypod ferns were particularly rich in N and P, with low N:P ratios, and their stoichiometry varied substantially in response to differences in nutrient availability. Distinguishing between these two groups has the potential to be useful both in and out of Hawaii, as they have distinct properties which can affect ecosystem function. These differences could contribute to the widespread abundance of polypod ferns in an angiosperm-dominated world, and to patterns of nutrient cycling and limitation in sites where ferns are abundant.