Habitat fragmentation is widespread across ecosystems, detrimentally affecting biodiversity. Although most habitat fragmentation studies have been conducted on macroscopic organisms, microbial communities and fungal processes may also be threatened by fragmentation. This study investigated whether fragmentation, and the effects of fragmentation on plants, altered fungal diversity and function within a fragmented shrubland in southern California. Using fluorimetric techniques, we assayed enzymes from plant litter collected from fragments of varying sizes to investigate enzymatic responses to fragmentation. To isolate the effects of plant richness from those of fragment size on fungi, we deployed litter bags containing different levels of plant litter diversity into the largest fragment and incubated in the field for one year. Following field incubation, we determined litter mass loss and conducted molecular analyses of fungal communities. We found that leaf-litter enzyme activity declined in smaller habitat fragments with less diverse vegetation. Moreover, we detected greater litter mass loss in litter bags containing more diverse plant litter. Additionally, bags with greater plant litter diversity harbored greater numbers of fungal taxa. These findings suggest that both plant litter resources and fungal function may be affected by habitat fragmentation's constraints on plants, possibly because plant species differ chemically, and may thus decompose at different rates. Diverse plant assemblages may produce a greater variety of litter resources and provide more ecological niche space, which may support greater numbers of fungal taxa. Thus, reduced plant diversity may constrain both fungal taxa richness and decomposition in fragmented coastal shrublands. Altogether, our findings provide evidence that even fungi may be affected by human-driven habitat fragmentation via direct effects of fragmentation on plants. Our findings underscore the importance of restoring diverse vegetation communities within larger coastal sage scrub fragments and suggest that this may be an effective way to improve the functional capacity of degraded sites.

Tropical forest conversion to pasture, which drives greenhouse gas emissions, soil degradation, and biodiversity loss, remains a pressing socio-ecological challenge. This problem has spurred increased interest in the potential of small-scale agroforestry systems to couple sustainable agriculture with biodiversity conservation, particularly in rapidly developing areas of the tropics. In addition to providing natural resources (i.e. food, medicine, lumber), agroforestry systems have the potential to maintain higher levels of biodiversity and greater biomass than lower diversity crop or pasture systems. Greater plant diversity may also enhance soil quality, further supporting agricultural productivity in nutrient-limited tropical systems. Yet, the nature of these relationships remains equivocal. To better understand how different land use strategies impact ecosystem services, we characterized the relationships between plant diversity (including species richness, phylogenetic diversity, and natural resource diversity), and soil quality within pasture, agroforests, and secondary forests, three common land use types maintained by small-scale farmers in the Pearl Lagoon Basin, Nicaragua. The area is undergoing accelerated globalization following the 2007 completion of the region's first major road; a change which is expected to increase forest conversion for agriculture. However, farmer agrobiodiversity maintenance in the Basin was previously found to be positively correlated with affiliation to local agricultural NGOs through the maintenance of agroforestry systems, despite these farmers residing in the communities closest to the new road, highlighting the potential for maintaining diverse agroforestry agricultural strategies despite heightened globalization pressures. We found that agroforestry sites tended to have higher surface soil %C, %N, and pH relative to neighboring to secondary forest, while maintaining comparable plant diversity. In contrast, pasture reduced species richness, phylogenetic diversity, and natural resource diversity. No significant relationships were found between plant diversity and the soil properties assessed; however higher species richness and phylodiversity was positively correlated with natural resource diversity. These finding suggest that small, diversified agroforestry systems may be a viable strategy for promoting both social and ecological functions in eastern Nicaragua and other rapidly developing areas of the tropics.

We present gridded 8 km-resolution data products of the estimated stem density, basal area, and biomass of tree taxa at Euro-American settlement of the midwestern United States during the middle to late 19th century for the states of Minnesota, Wisconsin, Michigan, Illinois, and Indiana. The data come from settlement-era Public Land Survey (PLS) data (ca. 0.8-km resolution) of trees recorded by land surveyors. The surveyor notes have been transcribed, cleaned, and processed to estimate stem density, basal area, and biomass at individual points. The point-level data are aggregated within 8 km grid cells and smoothed using a generalized additive statistical model that accounts for zero-inflated continuous data and provides approximate Bayesian uncertainty estimates. The statistical modeling smooths out sharp spatial features (likely arising from statistical noise) within areas smaller than about 200 km2. Based on this modeling, presettlement Midwestern landscapes supported multiple dominant species, vegetation types, forest types, and ecological formations. The prairies, oak savannas, and forests each had distinctive structures and spatial distributions across the domain. Forest structure varied from savanna (averaging 27 Mg/ha biomass) to northern hardwood (104 Mg/ha) and mesic southern forests (211 Mg/ha). The presettlement forests were neither unbroken and massively-statured nor dominated by young forests constantly structured by broad-scale disturbances such as fire, drought, insect outbreaks, or hurricanes. Most forests were structurally between modern second growth and old growth. We expect the data product to be useful as a baseline for investigating how forest ecosystems have changed in response to the last several centuries of climate change and intensive Euro-American land use and as a calibration dataset for paleoecological proxy-based reconstructions of forest composition and structure for earlier time periods. The data products (including raw and smoothed estimates at the 8-km scale) are available at the LTER Network Data Portal as version 1.0.

There is a debate concerning the definition and extent of tropical dry forest biome and vegetation type at a global spatial scale. We identify the potential extent of the tropical dry forest biome based on bioclimatic definitions and climatic data sets to improve global estimates of distribution, cover, and change. We compared four bioclimatic definitions of the tropical dry forest biome-Murphy and Lugo, Food and Agriculture Organization (FAO), DryFlor, aridity index-using two climatic data sets: WorldClim and Climatologies at High-resolution for the Earth's Land Surface Areas (CHELSA). We then compared each of the eight unique combinations of bioclimatic definitions and climatic data sets using 540 field plots identified as tropical dry forest from a literature search and evaluated the accuracy of World Wildlife Fund tropical and subtropical dry broadleaf forest ecoregions. We used the definition and climate data that most closely matched field data to calculate forest cover in 2000 and change from 2001 to 2020. Globally, there was low agreement (< 58%) between bioclimatic definitions and WWF ecoregions and only 40% of field plots fell within these ecoregions. FAO using CHELSA had the highest agreement with field plots (81%) and was not correlated with the biome extent. Using the FAO definition with CHELSA climatic data set, we estimate 4,931,414 km2 of closed canopy (≥ 40% forest cover) tropical dry forest in 2000 and 4,369,695 km2 in 2020 with a gross loss of 561,719 km2 (11.4%) from 2001 to 2020. Tropical dry forest biome extent varies significantly based on bioclimatic definition used, with nearly half of all tropical dry forest vegetation missed when using ecoregion boundaries alone, especially in Africa. Using site-specific field validation, we find that the FAO definition using CHELSA provides an accurate, standard, and repeatable way to assess tropical dry forest cover and change at a global scale.

In many parts of the world, the combined effects of habitat fragmentation and altered disturbance regimes pose a significant threat to biodiversity. This is particularly true in Mediterranean-type ecosystems (MTEs), which tend to be fire-prone, species rich, and heavily impacted by human land use. Given the spatial complexity of overlapping threats and species' vulnerability along with limited conservation budgets, methods are needed for prioritizing areas for monitoring and management in these regions. We developed a multi-criteria Pareto ranking methodology for prioritizing spatial units for conservation and applied it to fire threat, habitat fragmentation threat, species richness, and genetic biodiversity criteria in San Diego County, California, USA. We summarized the criteria and Pareto ranking results (from west to east) within the maritime, coastal, transitional, inland climate zones within San Diego County. Fire threat increased from the maritime zone eastward to the transitional zone, then decreased in the mountainous inland climate zone. Number of fires and fire return interval departure were strongly negatively correlated. Fragmentation threats, particularly road density and development density, were highest in the maritime climate zone, declined towards the east, and were positively correlated. Species richness criteria showed distributions among climate zones similar to those of the fire threat variables. When using species richness and fire threat criteria, most lower-ranked (higher conservation priority) units occurred in the coastal and transitional zones. When considering genetic biodiversity, lower-ranked units occurred more often in the mountainous inland zone. With Pareto ranking, there is no need to select criteria weights as part of the decision-making process. However, negative correlations and larger numbers of criteria can result in more units assigned to the same rank. Pareto ranking is broadly applicable and can be used as a standalone decision analysis method or in conjunction with other methods.

Studies of forest dynamics plots (FDPs) have revealed a variety of negative density-dependent (NDD) demographic interactions, especially among conspecific trees. These interactions can affect growth rate, recruitment and mortality, and they play a central role in the maintenance of species diversity in these complex ecosystems. Here we use an equal area annulus (EAA) point-pattern method to comprehensively analyze data from two tropical FDPs, Barro Colorado Island in Panama and Sinharaja in Sri Lanka. We show that these NDD interactions also influence the continued evolutionary diversification of even distantly related tree species in these FDPs. We examine the details of a wide range of these interactions between individual trees and the trees that surround them. All these interactions, and their cumulative effects, are strongest among conspecific focal and surrounding tree species in both FDPs. They diminish in magnitude with increasing phylogenetic distance between heterospecific focal and surrounding trees, but do not disappear or change the pattern of their dependence on size, density, frequency or physical distance even among the most distantly related trees. The phylogenetic persistence of all these effects provides evidence that interactions between tree species that share an ecosystem may continue to promote adaptive divergence even after the species' gene pools have become separated. Adaptive divergence among taxa would operate in stark contrast to an alternative possibility that has previously been suggested, that distantly related species with dispersal-limited distributions and confronted with unpredictable neighbors will tend to converge on common strategies of resource use. In addition, we have also uncovered a positive density-dependent effect: growth rates of large trees are boosted in the presence of a smaller basal area of surrounding trees. We also show that many of the NDD interactions switch sign rapidly as focal trees grow in size, and that their cumulative effect can strongly influence the distributions and species composition of the trees that surround the focal trees during the focal trees' lifetimes.

We used measurements from airborne imaging spectroscopy and LiDAR to quantify the biophysical structure and composition of vegetation on a dryland substrate age gradient in Hawaii. Both vertical stature and species composition changed during primary succession, and reveal a progressive increase in vertical stature on younger substrates followed by a collapse on Pleistocene-aged flows. Tall-stature Metrosideros polymorpha woodlands dominated on the youngest substrates (hundreds of years), and were replaced by the tall-stature endemic tree species Myoporum sandwicense and Sophora chrysophylla on intermediate-aged flows (thousands of years). The oldest substrates (tens of thousands of years) were dominated by the short-stature native shrub Dodonaea viscosa and endemic grass Eragrostis atropioides. We excavated 18 macroscopic charcoal fragments from Pleistocene-aged substrates. Mean radiocarbon age was 2,002 years and ranged from < 200 to 7,730. Genus identities from four fragments indicate that Osteomeles spp. or M. polymorpha once occupied the Pleistocene-aged substrates, but neither of these species is found there today. These findings indicate the existence of fires before humans are known to have occupied the Hawaiian archipelago, and demonstrate that a collapse in vertical stature is prevalent on the oldest substrates. This work contributes to our understanding of prehistoric fires in shaping the trajectory of primary succession in Hawaiian drylands.

The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher's alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼ 40,000 and ∼ 53,000, i.e., at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼ 19,000-25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼ 4,500-6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa.

Knowledge about the biogeographic affinities of the world's tropical forests helps to better understand regional differences in forest structure, diversity, composition, and dynamics. Such understanding will enable anticipation of region-specific responses to global environmental change. Modern phylogenies, in combination with broad coverage of species inventory data, now allow for global biogeographic analyses that take species evolutionary distance into account. Here we present a classification of the world's tropical forests based on their phylogenetic similarity. We identify five principal floristic regions and their floristic relationships: (i) Indo-Pacific, (ii) Subtropical, (iii) African, (iv) American, and (v) Dry forests. Our results do not support the traditional neo- versus paleotropical forest division but instead separate the combined American and African forests from their Indo-Pacific counterparts. We also find indications for the existence of a global dry forest region, with representatives in America, Africa, Madagascar, and India. Additionally, a northern-hemisphere Subtropical forest region was identified with representatives in Asia and America, providing support for a link between Asian and American northern-hemisphere forests.