Functional trait-based restoration alters nutrient cycling and invasion rates in Hawaiian lowland wet forest.

Many degraded ecosystems have altered nutrient dynamics due to invaders' possessing a suite of traits that allow them to both outcompete native species and alter the environment. In ecosystems where invasive species have increased nutrient turnover rates, it can be difficult to reduce nutrient availability. This study examined whether a functional trait-based restoration approach involving the planting of species with conservative nutrient-use traits could slow rates of nutrient cycling and consequently reduce rates of invasion. We examined a functional trait restoration initiative in a heavily invaded lowland wet forest site in Hilo, Hawai'i. Native and introduced species were chosen to create four experimental hybrid forest communities, in comparison to the invaded forest, with a factorial design in which communities varied in rates of carbon turnover (slow or moderate [SLOW, MOD]), and in the relationship of species in trait space (redundant or complementary [RED, COMP]). After the first 5 years, we evaluated community-level outcomes related to nutrient cycling: carbon (C), nitrogen (N), and phosphorus (P) via litterfall, litter decomposition, and outplant productivity and rates of invasion. We found that (1) regardless of treatment, the experimental communities had low rates of nutrient cycling through litterfall relative to the invaded reference forest, (2) the MOD communities had greater nutrient release via litterfall than the SLOW communities, (3) introduced species had greater nutrient release than native species in the two MOD experimental communities, and (4) within treatments, there was a positive relationship between nutrient levels and outplant basal area, but outplant basal area was negatively associated with rates of invasion. The negative relationships among basal area and weed invasion, particularly for the two COMP treatments, suggest species existing in different parts of trait space may help confer some degree of invasion resistance. The diversification of trait space was facilitated by the use of introduced species, a new concept in Hawaiian forest management. Although challenges remain in endeavors to restore this heavily degraded ecosystem, this study provides evidence that functional trait-based restoration approaches using carefully crafted hybrid communities can reduce rates of nutrient cycling and invasion in order to reach management goals.

Distribution of biomass dynamics in relation to tree size in forests across the world.

Tree size shapes forest carbon dynamics and determines how trees interact with their environment, including a changing climate. Here, we conduct the first global analysis of among-site differences in how aboveground biomass stocks and fluxes are distributed with tree size. We analyzed repeat tree censuses from 25 large-scale (4-52 ha) forest plots spanning a broad climatic range over five continents to characterize how aboveground biomass, woody productivity, and woody mortality vary with tree diameter. We examined how the median, dispersion, and skewness of these size-related distributions vary with mean annual temperature and precipitation. In warmer forests, aboveground biomass, woody productivity, and woody mortality were more broadly distributed with respect to tree size. In warmer and wetter forests, aboveground biomass and woody productivity were more right skewed, with a long tail towards large trees. Small trees (1-10 cm diameter) contributed more to productivity and mortality than to biomass, highlighting the importance of including these trees in analyses of forest dynamics. Our findings provide an improved characterization of climate-driven forest differences in the size structure of aboveground biomass and dynamics of that biomass, as well as refined benchmarks for capturing climate influences in vegetation demographic models.

The role of microtopography and resident species in post-disturbance recovery of arid habitats in Hawai'i.

Habitat-suitability indices (HSI) have been employed in restoration to identify optimal sites for planting native species. Often, HSI are based on abiotic variables and do not include biotic interactions, even though similar abiotic conditions can favor both native and nonnative species. Biotic interactions such as competition may be especially important in invader-dominated habitats because invasive species often have fast growth rates and can exploit resources quickly. In this study, we test the utility of an HSI of microtopography derived from airborne LiDAR to predict post-disturbance recovery and native planting success in native shrub-dominated and nonnative, invasive grass-dominated dryland habitats in Hawai'i. The HSI uses high-resolution digital terrain models to classify sites' microtopography as high, medium, or low suitability, based on wind exposure and topographic position. We used a split-plot before-after-control-impact design to implement a disturbance experiment within native shrub (Dodonaea viscosa) and nonnative, invasive grass (Cenchrus clandestinus)-dominated ecosystems across three microtopography categories. In contrast to previous studies using the same HSI, we found that microtopography was a poor predictor of pre-disturbance conditions for soil nutrients, organic matter content, or foliar C:N, within both Dodonaea and Cenchrus vegetation types. In invader-dominated Cenchrus plots, microtopography helped predict cover, but not as expected (i.e., highest cover would be in high-suitability plots): D. viscosa had the greatest cover in low-suitability and C. clandestinus had the greatest cover in medium-suitability plots. Similarly, in native-dominated Dodonaea plots, microtopography was a poor predictor of D. viscosa, C. clandestinus, and total plant cover. Although we found some evidence that microtopography helped inform post-disturbance plant recovery of D. viscosa and total plant cover, vegetation type was a more important predictor. Important for considering the success of plantings, percent cover of D. viscosa decreased while percent cover of C. clandestinus increased within both vegetation types 20 months after disturbance. Our results are evidence that HSIs based on topographic features may prove most useful for choosing planting sites in harsh habitats or those already dominated by native species. In more productive habitats, competition from resident species may offset any benefits gained from "better" suitability sites.

CTFS-ForestGEO: a worldwide network monitoring forests in an era of global change.

Global change is impacting forests worldwide, threatening biodiversity and ecosystem services including climate regulation. Understanding how forests respond is critical to forest conservation and climate protection. This review describes an international network of 59 long-term forest dynamics research sites (CTFS-ForestGEO) useful for characterizing forest responses to global change. Within very large plots (median size 25 ha), all stems ≥ 1 cm diameter are identified to species, mapped, and regularly recensused according to standardized protocols. CTFS-ForestGEO spans 25 °S-61 °N latitude, is generally representative of the range of bioclimatic, edaphic, and topographic conditions experienced by forests worldwide, and is the only forest monitoring network that applies a standardized protocol to each of the world's major forest biomes. Supplementary standardized measurements at subsets of the sites provide additional information on plants, animals, and ecosystem and environmental variables. CTFS-ForestGEO sites are experiencing multifaceted anthropogenic global change pressures including warming (average 0.61 °C), changes in precipitation (up to ± 30% change), atmospheric deposition of nitrogen and sulfur compounds (up to 3.8 g N m(-2) yr(-1) and 3.1 g S m(-2) yr(-1)), and forest fragmentation in the surrounding landscape (up to 88% reduced tree cover within 5 km). The broad suite of measurements made at CTFS-ForestGEO sites makes it possible to investigate the complex ways in which global change is impacting forest dynamics. Ongoing research across the CTFS-ForestGEO network is yielding insights into how and why the forests are changing, and continued monitoring will provide vital contributions to understanding worldwide forest diversity and dynamics in an era of global change.

Mapping habitat suitability for at-risk plant species and its implications for restoration and reintroduction.

The conservation of species at risk of extinction requires data to support decisions at landscape to regional scales. There is a need for information that can assist with locating suitable habitats in fragmented and degraded landscapes to aid the reintroduction of at-risk plant species. In addition, desiccation and water stress can be significant barriers to the success of at-risk plant reintroduction programs. We examine how airborne light detection and ranging (LiDAR) data can be used to model microtopographic features that reduce water stress and increase resource availability, providing information for landscape planning that can increase the success of reintroduction efforts for a dryland landscape in Hawaii. We developed a topographic habitat-suitability model (HSM) from LiDAR data that identifies topographic depressions that are protected from prevailing winds (high-suitability sites) and contrasts them with ridges and other exposed areas (low-suitability sites). We tested in the field whether high-suitability sites had microclimatic conditions that indicated better-quality habitat compared to low-suitability sites, whether plant-response traits indicated better growing conditions in high-suitability sites, whether the locations of individuals of existing at-risk plant species corresponded with our habitat-suitability classes, and whether the survival of planted individuals of a common native species was greater in high-suitability, compared to low-suitability, planting sites. Mean wind speed in a high-suitability field site was over five times lower than in a low-suitability site, and soil moisture and leaf wetness were greater, indicating less stress and greater resource availability in high-suitability areas. Plant height and leaf nutrient content were greater in high-suitability areas. Six at-risk species showed associations with high-suitability areas. The survival of planted individuals was less variable among high-suitability plots. These results suggest that plant establishment and survival is associated with the habitat conditions identified by our model. The HSM can improve the survival of planted individuals, reduce the cost of restoration and reintroduction programs through targeted management activities in high-suitability areas, and expand the ability of managers to make landscape-scale decisions regarding land-use, land acquisition, and species recovery.

Invasive-dominated grasslands in Hawai'i are resilient to disturbance.

Non-native-dominated landscapes may arise from invasion by competitive plant species, disturbance and invasion of early-colonizing species, or some combination of these. Without knowing site history, however, it is difficult to predict how native or non-native communities will reassemble after disturbance events. Given increasing disturbance levels across anthropogenically impacted landscapes, predictive understanding of these patterns is important. We asked how disturbance affected community assembly in six invaded habitat types common in dryland, grazed landscapes on Island of Hawai'i. We mechanically disturbed 100 m2 plots in six vegetation types dominated by one of four invasive perennial grasses (Cenchrus ciliaris, Cenchrus clandestinus, Cenchrus setaceus, or Melinis repens), a native shrub (Dodonaea viscosa), or a native perennial bunchgrass (Eragrostis atropioides). We censused vegetation before disturbance and monitored woody plant colonization and herbaceous cover for 21 months following the disturbance, categorizing species as competitors, colonizers, or a combination, based on recovery patterns. In addition, we planted individuals of the native shrub and bunchgrass and monitored survival to overcome dispersal limitation of native species when exploring these patterns. We found that the dominant vegetation types showed variation in post-disturbance syndrome, and that the variation in colonizer versus competitor syndrome occurred both between species, but also within species among different vegetation types. Although there were flushes of native shrub seedlings, these did not survive to 21 months within invaded habitats, probably due to regrowth by competitive invasive grasses. Similarly, survival of planted native individuals was related to the rate of regrowth by dominant species. Regardless of colonization/competitor syndrome, however, all dominant vegetation types were relatively resilient to change. Our results highlight that the altered post-agricultural, invaded grassland landscapes in Hawai'i are stable states. More generally, they point to the importance of resident communities and their effects on species interactions and seed availability in shaping plant community response to disturbance.

Mycorrhizal feedbacks influence global forest structure and diversity.

One mechanism proposed to explain high species diversity in tropical systems is strong negative conspecific density dependence (CDD), which reduces recruitment of juveniles in proximity to conspecific adult plants. Although evidence shows that plant-specific soil pathogens can drive negative CDD, trees also form key mutualisms with mycorrhizal fungi, which may counteract these effects. Across 43 large-scale forest plots worldwide, we tested whether ectomycorrhizal tree species exhibit weaker negative CDD than arbuscular mycorrhizal tree species. We further tested for conmycorrhizal density dependence (CMDD) to test for benefit from shared mutualists. We found that the strength of CDD varies systematically with mycorrhizal type, with ectomycorrhizal tree species exhibiting higher sapling densities with increasing adult densities than arbuscular mycorrhizal tree species. Moreover, we found evidence of positive CMDD for tree species of both mycorrhizal types. Collectively, these findings indicate that mycorrhizal interactions likely play a foundational role in global forest diversity patterns and structure.

Remote analysis of biological invasion and the impact of enemy release.

Escape from natural enemies is a widely held generalization for the success of exotic plants. We conducted a large-scale experiment in Hawaii (USA) to quantify impacts of ungulate removal on plant growth and performance, and to test whether elimination of an exotic generalist herbivore facilitated exotic success. Assessment of impacted and control sites before and after ungulate exclusion using airborne imaging spectroscopy and LiDAR, time series satellite observations, and ground-based field studies over nine years indicated that removal of generalist herbivores facilitated exotic success, but the abundance of native species was unchanged. Vegetation cover <1 m in height increased in ungulate-free areas from 48.7% +/- 1.5% to 74.3% +/- 1.8% over 8.4 years, corresponding to an annualized growth rate of lambda = 1.05 +/- 0.01 yr(-1) (median +/- SD). Most of the change was attributable to exotic plant species, which increased from 24.4% +/- 1.4% to 49.1% +/- 2.0%, (lambda = 1.08 +/- 0.01 yr(-1)). Native plants experienced no significant change in cover (23.0% +/- 1.3% to 24.2% +/- 1.8%, lambda = 1.01 +/- 0.01 yr(-1)). Time series of satellite phenology were indistinguishable between the treatment and a 3.0-km2 control site for four years prior to ungulate removal, but they diverged immediately following exclusion of ungulates. Comparison of monthly EVI means before and after ungulate exclusion and between the managed and control areas indicates that EVI strongly increased in the managed area after ungulate exclusion. Field studies and airborne analyses show that the dominant invader was Senecio madagascariensis, an invasive annual forb that increased from < 0.01% to 14.7% fractional cover in ungulate-free areas (lambda = 1.89 +/- 0.34 yr(-1)), but which was nearly absent from the control site. A combination of canopy LAI, water, and fractional cover were expressed in satellite EVI time series and indicate that the invaded region maintained greenness during drought conditions. These findings demonstrate that enemy release from generalist herbivores can facilitate exotic success and suggest a plausible mechanism by which invasion occurred. They also show how novel remote-sensing technology can be integrated with conservation and management to help address exotic plant invasions.

Arbuscular mycorrhizal trees influence the latitudinal beta-diversity gradient of tree communities in forests worldwide.

Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity.

OpenNahele: the open Hawaiian forest plot database.

BACKGROUND: This data paper provides a description of OpenNahele, the open Hawaiian forest plot database. OpenNahele includes 530 forest plots across the Hawaiian archipelago containing 43,590 individuals of 185 native and alien tree, shrub and tree fern species across six islands. We include estimates of maximum plant size (D950.1 and Dmax3) for 58 woody plant species, a key functional trait associated with dispersal distance and competition for light. OpenNahele can serve as a platform to test key ecological, evolutionary and conservation questions in a hotspot archipelago. NEW INFORMATION: OpenNahele is the first database that compiles data from a large number of forest plots across the Hawaiian archipelago to allow broad and high resolution studies of biodiversity patterns.Keywords: Hawaii, forests, islands, biodiversity, community ecology, evolutionary ecology.

Compilation of climate data from heterogeneous networks across the Hawaiian Islands.

Long-term, accurate observations of atmospheric phenomena are essential for a myriad of applications, including historic and future climate assessments, resource management, and infrastructure planning. In Hawai'i, climate data are available from individual researchers, local, State, and Federal agencies, and from large electronic repositories such as the National Centers for Environmental Information (NCEI). Researchers attempting to make use of available data are faced with a series of challenges that include: (1) identifying potential data sources; (2) acquiring data; (3) establishing data quality assurance and quality control (QA/QC) protocols; and (4) implementing robust gap filling techniques. This paper addresses these challenges by providing: (1) a summary of the available climate data in Hawai'i including a detailed description of the various meteorological observation networks and data accessibility, and (2) a quality controlled meteorological dataset across the Hawaiian Islands for the 25-year period 1990-2014. The dataset draws on observations from 471 climate stations and includes rainfall, maximum and minimum surface air temperature, relative humidity, wind speed, downward shortwave and longwave radiation data.

Climate sensitive size-dependent survival in tropical trees.

Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four 'survival modes' that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Hülsmann and Hartig suggest that ecological mechanisms other than specialized natural enemies or intraspecific competition contribute to our estimates of conspecific negative density dependence (CNDD). To address their concern, we show that our results are not the result of a methodological artifact and present a null-model analysis that demonstrates that our original findings-(i) stronger CNDD at tropical relative to temperate latitudes and (ii) a latitudinal shift in the relationship between CNDD and species abundance-persist even after controlling for other processes that might influence spatial relationships between adults and recruits.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Chisholm and Fung claim that our method of estimating conspecific negative density dependence (CNDD) in recruitment is systematically biased, and present an alternative method that shows no latitudinal pattern in CNDD. We demonstrate that their approach produces strongly biased estimates of CNDD, explaining why they do not detect a latitudinal pattern. We also address their methodological concerns using an alternative distance-weighted approach, which supports our original findings of a latitudinal gradient in CNDD and a latitudinal shift in the relationship between CNDD and species abundance.

Plant diversity increases with the strength of negative density dependence at the global scale.

Theory predicts that higher biodiversity in the tropics is maintained by specialized interactions among plants and their natural enemies that result in conspecific negative density dependence (CNDD). By using more than 3000 species and nearly 2.4 million trees across 24 forest plots worldwide, we show that global patterns in tree species diversity reflect not only stronger CNDD at tropical versus temperate latitudes but also a latitudinal shift in the relationship between CNDD and species abundance. CNDD was stronger for rare species at tropical versus temperate latitudes, potentially causing the persistence of greater numbers of rare species in the tropics. Our study reveals fundamental differences in the nature of local-scale biotic interactions that contribute to the maintenance of species diversity across temperate and tropical communities.

Home range use and movement patterns of non-native feral goats in a tropical island montane dry landscape.

Advances in wildlife telemetry and remote sensing technology facilitate studies of broad-scale movements of ungulates in relation to phenological shifts in vegetation. In tropical island dry landscapes, home range use and movements of non-native feral goats (Capra hircus) are largely unknown, yet this information is important to help guide the conservation and restoration of some of the world's most critically endangered ecosystems. We hypothesized that feral goats would respond to resource pulses in vegetation by traveling to areas of recent green-up. To address this hypothesis, we fitted six male and seven female feral goats with Global Positioning System (GPS) collars equipped with an Argos satellite upload link to examine goat movements in relation to the plant phenology using the Normalized Difference Vegetation Index (NDVI). Movement patterns of 50% of males and 40% of females suggested conditional movement between non-overlapping home ranges throughout the year. A shift in NDVI values corresponded with movement between primary and secondary ranges of goats that exhibited long-distance movement, suggesting that vegetation phenology as captured by NDVI is a good indicator of the habitat and movement patterns of feral goats in tropical island dry landscapes. In the context of conservation and restoration of tropical island landscapes, the results of our study identify how non-native feral goats use resources across a broad landscape to sustain their populations and facilitate invasion of native plant communities.

Primary Succession on a Hawaiian Dryland Chronosequence.

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.

Native trees show conservative water use relative to invasive trees: results from a removal experiment in a Hawaiian wet forest.

While the supply of freshwater is expected to decline in many regions in the coming decades, invasive plant species, often 'high water spenders', are greatly expanding their ranges worldwide. In this study, we quantified the ecohydrological differences between native and invasive trees and also the effects of woody invasive removal on plot-level water use in a heavily invaded mono-dominant lowland wet tropical forest on the Island of Hawaii. We measured transpiration rates of co-occurring native and invasive tree species with and without woody invasive removal treatments. Twenty native Metrosideros polymorpha and 10 trees each of three invasive species, Cecropia obtusifolia, Macaranga mappa and Melastoma septemnervium, were instrumented with heat-dissipation sap-flux probes in four 100 m(2) plots (two invaded, two removal) for 10 months. In the invaded plots, where both natives and invasives were present, Metrosideros had the lowest sap-flow rates per unit sapwood, but the highest sap-flow rates per whole tree, owing to its larger mean diameter than the invasive trees. Stand-level water use within the removal plots was half that of the invaded plots, even though the removal of invasives caused a small but significant increase in compensatory water use by the remaining native trees. By investigating the effects of invasive species on ecohydrology and comparing native vs. invasive physiological traits, we not only gain understanding about the functioning of invasive species, but we also highlight potential water-conservation strategies for heavily invaded mono-dominant tropical forests worldwide. Native-dominated forests free of invasive species can be conservative in overall water use, providing a strong rationale for the control of invasive species and preservation of native-dominated stands.

Forest structure in low-diversity tropical forests: a study of Hawaiian wet and dry forests.

The potential influence of diversity on ecosystem structure and function remains a topic of significant debate, especially for tropical forests where diversity can range widely. We used Center for Tropical Forest Science (CTFS) methodology to establish forest dynamics plots in montane wet forest and lowland dry forest on Hawai'i Island. We compared the species diversity, tree density, basal area, biomass, and size class distributions between the two forest types. We then examined these variables across tropical forests within the CTFS network. Consistent with other island forests, the Hawai'i forests were characterized by low species richness and very high relative dominance. The two Hawai'i forests were floristically distinct, yet similar in species richness (15 vs. 21 species) and stem density (3078 vs. 3486/ha). While these forests were selected for their low invasive species cover relative to surrounding forests, both forests averaged 5->50% invasive species cover; ongoing removal will be necessary to reduce or prevent competitive impacts, especially from woody species. The montane wet forest had much larger trees, resulting in eightfold higher basal area and above-ground biomass. Across the CTFS network, the Hawaiian montane wet forest was similar to other tropical forests with respect to diameter distributions, density, and aboveground biomass, while the Hawai'i lowland dry forest was similar in density to tropical forests with much higher diversity. These findings suggest that forest structural variables can be similar across tropical forests independently of species richness. The inclusion of low-diversity Pacific Island forests in the CTFS network provides an ∼80-fold range in species richness (15-1182 species), six-fold variation in mean annual rainfall (835-5272 mm yr(-1)) and 1.8-fold variation in mean annual temperature (16.0-28.4°C). Thus, the Hawaiian forest plots expand the global forest plot network to enable testing of ecological theory for links among species diversity, environmental variation and ecosystem function.

Trade-offs in seedling growth and survival within and across tropical forest microhabitats.

For niche differences to maintain coexistence of sympatric species, each species must grow and/or survive better than each of the others in at least one set of conditions (i.e., performance trade-offs). However, the extent of niche differentiation in tropical forests remains highly debated. We present the first test of performance trade-offs for wild seedlings in a tropical forest. We measured seedling relative growth rate (RGR) and survival of four common native woody species across 18 light, substrate, and topography microhabitats over 2.5 years within Hawaiian montane wet forest, an ideal location due to its low species diversity and strong species habitat associations. All six species pairs exhibited significant performance trade-offs across microhabitats and for RGR versus survival within microhabitats. We also found some evidence of performance equivalence, with species pairs having similar performance in 26% of comparisons across microhabitats. Across species, survival under low light was generally positively associated with RGR under high light. When averaged over all species, topography (slope, aspect, and elevation) explained most of the variation in RGR attributable to microhabitat variables (51-53%) followed by substrate type (35-37%) and light (11-12%). However, the relative effects of microhabitat differed among species and RGR metric (i.e., RGR for height, biomass, or leaf area). These findings indicate that performance trade-offs among species during regeneration are common in low-diversity tropical forest, although other mechanisms may better explain the coexistence of species with small performance differences.