James R. Kellner

Poleward expansion of mangroves is a threshold response to decreased frequency of extreme cold events

Significance Coastal mangrove forests support a diverse array of associated species and provide ecosystem services to human communities. Mangroves cannot tolerate extreme freezing temperatures and so are generally limited to tropical environments. However, climate change in the form of increasing temperatures has the potential to facilitate increases in mangrove abundance near tropical–temperate transition zones. Here, we use 28 y of satellite imagery to demonstrate that increases in mangrove area have already occurred along the northeast coast of Florida. These increases correspond to decreases in the frequency of extreme cold events in this region. We also identify a temperature-related ecological threshold of −4°C. These results suggest that landscape-scale increases in mangrove area may occur in other regions where this threshold is crossed. Regional warming associated with climate change is linked with altered range and abundance of species and ecosystems worldwide. However, the ecological impacts of changes in the frequency of extreme events have not been as well documented, especially for coastal and marine environments. We used 28 y of satellite imagery to demonstrate that the area of mangrove forests has doubled at the northern end of their historic range on the east coast of Florida. This expansion is associated with a reduction in the frequency of “extreme” cold events (days colder than −4 °C), but uncorrelated with changes in mean annual temperature, mean annual precipitation, and land use. Our analyses provide evidence for a threshold response, with declining frequency of severe cold winter events allowing for poleward expansion of mangroves. Future warming may result in increases in mangrove cover beyond current latitudinal limits of mangrove forests, thereby altering the structure and function of these important coastal ecosystems.

How many tree species are there in the Amazon and how many of them will go extinct

New roads, agricultural projects, logging, and mining are claiming an ever greater area of once-pristine Amazonian forest. The Millennium Ecosystems Assessment (MA) forecasts the extinction of a large fraction of Amazonian tree species based on projected loss of forest cover over the next several decades. How accurate are these estimates of extinction rates? We use neutral theory to estimate the number, relative abundance, and range size of tree species in the Amazon metacommunity and estimate likely tree-species extinctions under published optimistic and nonoptimistic Amazon scenarios. We estimate that the Brazilian portion of the Amazon Basin has (or had) 11,210 tree species that reach sizes >10 cm DBH (stem diameter at breast height). Of these, 3,248 species have population sizes >1 million individuals, and, ignoring possible climate-change effects, almost all of these common species persist under both optimistic and nonoptimistic scenarios. At the rare end of the abundance spectrum, however, neutral theory predicts the existence of ≈5,308 species with <10,000 individuals each that are expected to suffer nearly a 50% extinction rate under the nonoptimistic deforestation scenario and an ≈37% loss rate even under the optimistic scenario. Most of these species have small range sizes and are highly vulnerable to local habitat loss. In ensembles of 100 stochastic simulations, we found mean total extinction rates of 20% and 33% of tree species in the Brazilian Amazon under the optimistic and nonoptimistic scenarios, respectively.

Macrosystems ecology: understanding ecological patterns and processes at continental scales

Macrosystems ecology is the study of diverse ecological phenomena at the scale of regions to continents and their interactions with phenomena at other scales. This emerging subdiscipline addresses ecological questions and environmental problems at these broad scales. Here, we describe this new field, show how it relates to modern ecological study, and highlight opportunities that stem from taking a macrosystems perspective. We present a hierarchical framework for investigating macrosystems at any level of ecological organization and in relation to broader and finer scales. Building on well-established theory and concepts from other subdisci- plines of ecology, we identify feedbacks, linkages among distant regions, and interactions that cross scales of space and time as the most likely sources of unexpected and novel behaviors in macrosystems. We present three examples that highlight the importance of this multiscaled systems perspective for understanding the ecology of regions to continents.

Convergent structural responses of tropical forests to diverse disturbance regimes

Size frequency distributions of canopy gaps are a hallmark of forest dynamics. But it remains unknown whether legacies of forest disturbance are influencing vertical size structure of landscapes, or space-filling in the canopy volume. We used data from LiDAR remote sensing to quantify distributions of canopy height and sizes of 434,501 canopy gaps in five tropical rain forest landscapes in Costa Rica and Hawaii. The sites represented a wide range of variation in structure and natural disturbance history, from canopy gap dynamics in lowland Costa Rica and Hawaii, to stages and types of stand-level dieback on upland Mauna Kea and Kohala volcanoes. Large differences in vertical canopy structure characterized these five tropical rain forest landscapes, some of which were related to known disturbance events. Although there were quantitative differences in the values of scaling exponents within and among sites, size frequency distributions of canopy gaps followed power laws at all sites and in all canopy height classes. Scaling relationships in gap size at different heights in the canopy were qualitatively similar at all sites, revealing a remarkable similarity despite clearly defined differences in species composition and modes of prevailing disturbance. These findings indicate that power-law gap-size frequency distributions are ubiquitous features of these five tropical rain forest landscapes, and suggest that mechanisms of forest disturbance may be secondary to other processes in determining vertical and horizontal size structure in canopies.

Pervasive canopy dynamics produce short‐term stability in a tropical rain forest landscape

A fundamental property of all forest landscapes is the size frequency distribution of canopy gap disturbances. But characterizing forest structure and changes at large spatial scales has been challenging and most of our understanding is from permanent inventory plots. Here we report the first application of light detection and ranging remote sensing to measurements of canopy disturbance and regeneration in an old-growth tropical rain forest landscape. Pervasive local height changes figure prominently in the dynamics of this forest. Although most canopy gaps recruited to higher positions during 8.5 years, size frequency distributions were similar at two points in time and well-predicted by power-laws. At larger spatial scales (hundreds of ha), height increases and decreases occurred with similar frequency and changes to canopy height that were analysed using a height transition matrix suggest that the distribution of canopy height at the beginning of the study was close to the projected steady-state equilibrium under the recent disturbance regime. Taken together, these findings show how widespread local height changes can produce short-term stability in a tropical rain forest landscape.

Forest Canopy Gap Distributions in the Southern Peruvian Amazon

Canopy gaps express the time-integrated effects of tree failure and mortality as well as regrowth and succession in tropical forests. Quantifying the size and spatial distribution of canopy gaps is requisite to modeling forest functional processes ranging from carbon fluxes to species interactions and biological diversity. Using high-resolution airborne Light Detection and Ranging (LiDAR), we mapped and analyzed 5,877,937 static canopy gaps throughout 125,581 ha of lowland Amazonian forest in Peru. Our LiDAR sampling covered a wide range of forest physiognomies across contrasting geologic and topographic conditions, and on depositional floodplain and erosional terra firme substrates. We used the scaling exponent of the Zeta distribution (λ) as a metric to quantify and compare the negative relationship between canopy gap frequency and size across sites. Despite variable canopy height and forest type, values of λ were highly conservative (λ mean  = 1.83, s  = 0.09), and little variation was observed regionally among geologic substrates and forest types, or at the landscape level comparing depositional-floodplain and erosional terra firme landscapes. λ-values less than 2.0 indicate that these forests are subjected to large gaps that reset carbon stocks when they occur. Consistency of λ-values strongly suggests similarity in the mechanisms of canopy failure across a diverse array of lowland forests in southwestern Amazonia.

Approaches to advance scientific understanding of macrosystems ecology

The emergence of macrosystems ecology (MSE), which focuses on regional- to continental-scale ecological patterns and processes, builds upon a history of long-term and broad-scale studies in ecology. Scientists face the difficulty of integrating the many elements that make up macrosystems, which consist of hierarchical processes at interacting spatial and temporal scales. Researchers must also identify the most relevant scales and variables to be considered, the required data resources, and the appropriate study design to provide the proper inferences. The large volumes of multi-thematic data often associated with macrosystem studies typically require validation, standardization, and assimilation. Finally, analytical approaches need to describe how cross-scale and hierarchical dynamics and interactions relate to macroscale phenomena. Here, we elaborate on some key methodological challenges of MSE research and discuss existing and novel approaches to meet them.

Operational Tree Species Mapping in a Diverse Tropical Forest with Airborne Imaging Spectroscopy.

Remote identification and mapping of canopy tree species can contribute valuable information towards our understanding of ecosystem biodiversity and function over large spatial scales. However, the extreme challenges posed by highly diverse, closed-canopy tropical forests have prevented automated remote species mapping of non-flowering tree crowns in these ecosystems. We set out to identify individuals of three focal canopy tree species amongst a diverse background of tree and liana species on Barro Colorado Island, Panama, using airborne imaging spectroscopy data. First, we compared two leading single-class classification methods—binary support vector machine (SVM) and biased SVM—for their performance in identifying pixels of a single focal species. From this comparison we determined that biased SVM was more precise and created a multi-species classification model by combining the three biased SVM models. This model was applied to the imagery to identify pixels belonging to the three focal species and the prediction results were then processed to create a map of focal species crown objects. Crown-level cross-validation of the training data indicated that the multi-species classification model had pixel-level producer’s accuracies of 94–97% for the three focal species, and field validation of the predicted crown objects indicated that these had user’s accuracies of 94–100%. Our results demonstrate the ability of high spatial and spectral resolution remote sensing to accurately detect non-flowering crowns of focal species within a diverse tropical forest. We attribute the success of our model to recent classification and mapping techniques adapted to species detection in diverse closed-canopy forests, which can pave the way for remote species mapping in a wider variety of ecosystems.

Integrating physiological threshold experiments with climate modeling to project mangrove species’ range expansion

Predictions of climate-related shifts in species ranges have largely been based on correlative models. Due to limitations of these models, there is a need for more integration of experimental approaches when studying impacts of climate change on species distributions. Here, we used controlled experiments to identify physiological thresholds that control poleward range limits of three species of mangroves found in North America. We found that all three species exhibited a threshold response to extreme cold, but freeze tolerance thresholds varied among species. From these experiments, we developed a climate metric, freeze degree days (FDD), which incorporates both the intensity and the frequency of freezes. When included in distribution models, FDD accurately predicted mangrove presence/absence. Using 28 years of satellite imagery, we linked FDD to observed changes in mangrove abundance in Florida, further exemplifying the importance of extreme cold. We then used downscaled climate projections of FDD to project that these range limits will move northward by 2.2-3.2 km yr(-1) over the next 50 years.

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.