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Lianas escape self-thinning: Experimental evidence of positive density dependence in temperate lianas Celastrus orbiculatus and C. scandens

The neighborhood density of plants strongly affects their growth, reproduction, and survival. In most cases, high density increases competition and negatively affects a focal plant in predictable ways, leading to the self-thinning law. There are, however, situations in which high densities of plants facilitate focal plant performance, resulting in positive density dependence. Despite their importance in forest gap dynamics and distinctive growth form, there have been very few studies of the effect of density on lianas or vines. We grew an invasive ( Celastrus orbiculatus ) and a native ( Celastrus scandens ) liana species together in three different density treatments, while also manipulating the light and support availability. We found that across treatment conditions, C. orbiculatus always out-performed C. scandens , showing greater relative growth rate in height and diameter, greater biomass and higher survival. Both species responded similarly to the density treatments: with plants in high density not showing a decrease in relative height growth rate compared to medium density. Aboveground biomass for C. scandens was not affected by density, while for C. orbiculatus , the most massive plants were growing in medium density without support. More surprisingly, survival analysis indicated that the two species both had significantly lower mortality rates in the highest density treatment; this trend held true across the other treatments of light and supports. More generally, this study demonstrates that these lianas can escape the consequences of high density and thus the self-thinning law that affects self-supporting plants. This suggests a broader hypothesis about lianas in general: their greater flexibility in allocating growth resources allows them to grow taller and thinner without collapsing and thereby potentially escape shading and mortality even at high densities.

Perspectives in Plant Ecology, Evolution and Syste

Hawai‘i forest review: Synthesizing the ecology, evolution, and conservation of a model system

As the most remote archipelago in the world, the Hawaiian Islands are home to a highly endemic and disharmonic biota that has fascinated biologists for centuries. Forests are the dominant terrestrial biome in Hawai‘i, spanning complex, heterogeneous climates across substrates that vary tremendously in age, soil structure, and nutrient availability. Species richness is low in Hawaiian forests compared to other tropical forests, as a consequence of dispersal limitation from continents and adaptive radiations in only some lineages, and forests are dominated by the widespread Metrosideros species complex. Low species richness provides a relatively tractable model system for studies of community assembly, local adaptation, and species interactions. Moreover, Hawaiian forests provide insights into predicted patterns of evolution on islands, revealing that while some evidence supports “island syndromes,” there are exceptions to them all. For example, Hawaiian plants are not as a whole less defended against herbivores, less dispersible, more conservative in resource use, or more slow-growing than their continental relatives. Clearly, more work is needed to understand the drivers, sources, and constraints on phenotypic variation among Hawaiian species, including both widespread and rare species, and to understand the role of this variation for ecological and evolutionary processes, which will further contribute to conservation of this unique biota. Today, Hawaiian forests are among the most threatened globally. Resource management failures – the proliferation of non-native species in particular – have led to devastating declines in native taxa and resulted in dominance by novel species assemblages. Conservation and restoration of Hawaiian forests now rely on managing threats including climate change, ongoing species introductions, novel pathogens, lost mutualists, and altered ecosystem dynamics through the use of diverse tools and strategies grounded in basic ecological, evolutionary, and biocultural principles. The future of Hawaiian forests thus depends on the synthesis of ecological and evolutionary research, which will continue to inform future conservation and restoration practices.

Hawaii

Causes, consequences, and conservation of ungulate migration

Our understanding of ungulate migration is advancing rapidly due to innovations in modern animal tracking. Herein, we review and synthesize nearly seven decades of work on migration and other long-distance movements of wild ungulates. Although it has long been appreciated that ungulates migrate to enhance access to forage, recent contributions demonstrate that their movements are fine tuned to dynamic landscapes, where forage, snow, and drought change seasonally. Researchers are beginning to understand how ungulates navigate migrations, with the emerging view that animals blend gradient tracking with spatial memory, some of which is socially learned. Although migration often promotes abundant populations—with broad effects on ecosystems—many migrations around the world have been lost or are currently threatened by habitat fragmentation, climate change, and barriers to movement. Fortunately, new efforts that use empirical tracking data to map migrations in detail are facilitating effective conservation measures needed to maintain ungulate migration.

Annual Review of Ecology, Evolution, and Systemati

Characterizing species interactions to understand press perturbations: What is the community matrix?

The community matrix is among ecology's most important mathematical abstractions, formally encapsulating the interconnected network of effects that species have on one another's populations. Despite its importance, the term `community matrix' has been applied to matrices having differing interpretations. This has hindered the application of theory for understanding community structure and perturbation responses, particularly in the contexts of ecosystem-based management and conservation. Here we clarify the correspondence and distinctions between the Interaction matrix, the Alpha matrix and the Jacobian matrix, terms which are frequently used interchangeably and have numerous synonyms, including the term Community matrix. We illustrate how these matrices correspond to different ways of characterizing interaction strengths, how they permit insights regarding different types of press perturbations of species growth rates or abundances, and how these are related by a simple scaling relationship. Connections to additional interaction strength characterizations encapsulated by the Beta matrix, the Gamma matrix, and the Removal matrix are also discussed. Our synthesis highlights the empirical challenges that remain in using these mathematical tools to understand actual communities.

Ecology, Evolution, and Systematics

Evolutionary ecology of fire

Fire has been an ecosystem process since plants colonized land over 400 million years ago. Many diverse traits provide a fitness benefit following fires, and these adaptive traits vary with the fire regime. Some of these traits enhance fire survival, while others promote recruitment in the postfire environment. Demonstrating that these traits are fire adaptations is challenging, since many arose early in the paleontological record, although increasingly better fossil records and phylogenetic analysis make timing of these trait origins to fire more certain. Resprouting from the base of stems is the most widely distributed fire-adaptive trait, and it is likely to have evolved under a diversity of disturbance types. The origins of other traits like serotiny, thick bark, fire-stimulated germination, and postfire flowering are more tightly linked to fire. Fire-adaptive traits occur in many environments: boreal and temperate forests, Mediterranean-type climate (MTC) shrublands, savannas, and other grasslands. MTC ecosystems are distinct in that many taxa in different regions have lost the resprouting ability and depend solely on postfire recruitment for postfire recovery. This obligate seeding mode is perhaps the most vulnerable fire-adaptive syndrome in the face of current global change, particularly in light of increasing anthropogenic fire frequency.

Annual Review of Ecology, Evolution, and Systemati

Functional ecology of free-living nitrogen fixation: A contemporary perspective

Nitrogen (N) availability is thought to frequently limit terrestrial ecosystem processes, and explicit consideration of N biogeochemistry, including biological N 2 fixation, is central to understanding ecosystem responses to environmental change. Yet, the importance of free-living N 2 fixation—a process that occurs on a wide variety of substrates, is nearly ubiquitous in terrestrial ecosystems, and may often represent the dominant pathway for acquiring newly available N—is often underappreciated. Here, we draw from studies that investigate free-living N 2 fixation from functional, physiological, genetic, and ecological perspectives. We show that recent research and analytical advances have generated a wealth of new information that provides novel insight into the ecology of N 2 fixation as well as raises new questions and priorities for future work. These priorities include a need to better integrate free-living N 2 fixation into conceptual and analytical evaluations of the N cycle's role in a variety of global change scenarios.

Annual Review of Ecology, Evolution, and Systemati

Observed and potential range shifts of native and non-native species with climate change

There is broad concern that the range shifts of global flora and fauna will not keep up with climate change, increasing the likelihood of population declines and extinctions. Many populations of nonnative species already have advantages over native species, including widespread human-aided dispersal and release from natural enemies. But do nonnative species also have an advantage with climate change? Here, we review observed and potential range shifts for native and nonnative species globally. We show that nonnative species are expanding their ranges orders of magnitude faster than native species, reflecting both traits that enable rapid spread and ongoing human-mediated introduction. We further show that nonnative species have large potential ranges and range expansions with climate change, likely due to a combination of widespread introduction and broader climatic tolerances. With faster spread rates and larger potential to persist or expand, nonnative populations have a decided advantage in a changing climate.

Annual Review of Ecology, Evolution, and Systemati

Interactions among invasive plants: Lessons from Hawai‘i

Most ecosystems have multiple-plant invaders rather than single-plant invaders, yet ecological studies and management actions focus largely on single invader species. There is a need for general principles regarding invader interactions across varying environmental conditions, so that secondary invasions can be anticipated and managers can allocate resources toward pretreatment or postremoval actions. By reviewing removal experiments conducted in three Hawaiian ecosystems (a dry tropical forest, a seasonally dry mesic forest, and a lowland wet forest), we evaluate the roles environmental harshness, priority effects, productivity potential, and species interactions have in influencing secondary invasions, defined here as invasions that are influenced either positively (facilitation) or negatively (inhibition/priority effects) by existing invaders. We generate a conceptual model with a surprise index to describe whether long-term plant invader composition and dominance is predictable or stochastic after a system perturbation such as a removal experiment. Under extremely low resource availability, the surprise index is low, whereas under intermediate-level resource environments, invader dominance is more stochastic and the surprise index is high. At high resource levels, the surprise index is intermediate: Invaders are likely abundant in the environment but their response to a perturbation is more predictable than at intermediate resource levels. We suggest further testing across environmental gradients to determine key variables that dictate the predictability of postremoval invader composition.

Hawai'i

Threshold changes in winter temperature and precipitation drive threshold responses across nine global climate zones and associated biomes

Globally, winter temperatures are rising, and snowpack is shrinking or disappearing entirely. Despite previous research and published literature reviews, it remains unknown whether biomes across the globe will cross important thresholds in winter temperature and precipitation that will lead to significant ecological changes. Here, we combine the widely used Köppen–Geiger climate classification system with worst-case-scenario projected changes in global monthly temperature and precipitation to illustrate how multiple climatic zones across Earth may experience shifting winter conditions by the end of this century. We then examine how these shifts may affect ecosystems within corresponding biomes. Our analysis demonstrates potential widespread losses of extreme cold (<−20°C) in Arctic, boreal, and cool temperate regions. We also show the possible disappearance of freezing temperatures (<0°C) and large decreases in snowfall in warm temperate and dryland areas. We identify important and potentially irreversible ecological changes associated with crossing these winter climate thresholds.

Annual Review of Ecology, Evolution, and Systemati

A multiscale, hierarchical model of pulse dynamics in arid-land ecosystems

Ecological processes in arid lands are often described by the pulse-reserve paradigm, in which rain events drive biological activity until moisture is depleted, leaving a reserve. This paradigm is frequently applied to processes stimulated by one or a few precipitation events within a growing season. Here we expand the original framework in time and space and include other pulses that interact with rainfall. This new hierarchical pulse-dynamics framework integrates space and time through pulse-driven exchanges, interactions, transitions, and transfers that occur across individual to multiple pulses extending from micro to watershed scales. Climate change will likely alter the size, frequency, and intensity of precipitation pulses in the future, and arid-land ecosystems are known to be highly sensitive to climate variability. Thus, a more comprehensive understanding of arid-land pulse dynamics is needed to determine how these ecosystems will respond to, and be shaped by, increased climate variability.

Annual Review of Ecology, Evolution, and Systemati

Marine infectious disease ecology

To put marine disease impacts in context requires a broad perspective on the roles infectious agents have in the ocean. Parasites infect most marine vertebrate and invertebrate species, and parasites and predators can have comparable biomass density, suggesting they play comparable parts as consumers in marine food webs. Although some parasites might increase with disturbance, most probably decline as food webs unravel. There are several ways to adapt epidemiological theory to the marine environment. In particular, because the ocean represents a three-dimensional moving habitat for hosts and parasites, models should open up the spatial scales at which infective stages and host larvae travel. In addition to open recruitment and dimensionality, marine parasites are subject to fishing, filter feeders, dosedependent infection, environmental forcing, and death-based transmission. Adding such considerations to marine disease models will make it easier to predict which infectious diseases will increase or decrease in a changing ocean.

Annual Review of Ecology, Evolution, and Systemati

Mediterranean biomes: Evolution of their vegetation, floras and climate

Mediterranean-type ecosystems (MTEs) possess the highest levels of plant species richness in the world outside of the wet tropics. Sclerophyll vegetation similar to today’s mediterranean-type shrublands was already present on oligotrophic soils in the wet and humid climate of the Cretaceous, with fire-adapted Paleogene lineages in southwestern Australia and the Cape Region. The novel MTC seasonality present since the mid-Miocene has allowed colonization of MTEs from a regional species pool with associated diversification. Fire persistence has been a primary driving factor for speciation in four of the five regions. Understanding the regional patterns of plant species diversity among the MTEs involves complex interactions of geologic and climatic histories for each region as well as ecological factors that have promoted diversification in the Neogene and Quaternary. A critical element of species richness for many MTE lineages has been their ability to speciate and persist at fine spatial scales, with low rates of extinction.

Annual Review of Ecology, Evolution, and Systemati

Are diseases increasing in the ocean?

Many factors (climate warming, pollution, harvesting, introduced species) can contribute to disease outbreaks in marine life. Concomitant increases in each of these makes it difficult to attribute recent changes in disease occurrence or severity to any one factor. For example, the increase in disease of Caribbean coral is postulated to be a result of climate change and introduction of terrestrial pathogens. Indirect evidence exists that ( a ) warming increased disease in turtles; ( b ) protection, pollution, and terrestrial pathogens increased mammal disease; ( c ) aquaculture increased disease in mollusks; and ( d ) release from overfished predators increased sea urchin disease. In contrast, fishing and pollution may have reduced disease in fishes. In other taxa (e.g., sea grasses, crustaceans, sharks), there is little evidence that disease has changed over time. The diversity of patterns suggests there are many ways that environmental change can interact with disease in the ocean.

Annual Review of Ecology, Evolution, and Systemati

Individual-based modeling of ecological and evolutionary processes

Individual-based models (IBMs) allow the explicit inclusion of individual variation in greater detail than do classical differential-equation and difference-equation models. Inclusion of such variation is important for continued progress in ecological and evolutionary theory. We provide a conceptual basis for IBMs by describing five major types of individual variation in IBMs: spatial, ontogenetic, phenotypic, cognitive, and genetic. IBMs are now used in almost all subfields of ecology and evolutionary biology. We map those subfields and look more closely at selected key papers on fish recruitment, forest dynamics, sympatric speciation, metapopulation dynamics, maintenance of diversity, and species conservation. Theorists are currently divided on whether IBMs represent only a practical tool for extending classical theory to more complex situations, or whether individual-based theory represents a radically new research program. We feel that the tension between these two poles of thinking can be a source of creativity in ecology and evolutionary theory.

Annual Review of Ecology, Evolution, and Systemati

Nearshore energy subsidies support Lake Michigan fishes and invertebrates following major changes in food web structure

Aquatic food webs that incorporate multiple energy channels (e.g. nearshore benthic or pelagic) with varying productivity and turnover rates convey stability to biological communities by providing multiple independent energy sources. Within the Lake Michigan food web, invasive dreissenid mussels have caused rapid changes to food web structure and potentially altered the channels through which consumers acquire energy. We used stable C and N isotopes to determine how Lake Michigan food web structure has changed in the past decade, coincident with the expansion of dreissenid mussels, decreased pelagic phytoplankton production and increased nearshore benthic algal production. Fish and invertebrate samples collected from sites around Lake Michigan were analyzed to determine taxa-specific 13C:12C (delta 13C) and 15N:14N (delta 15N) ratios. Sampling took place during two distinct periods, 2002-2003 and 2010-2012, that spanned the period of dreissenid expansion, and included nearshore, pelagic and profundal fish and invertebrate taxa. Magnitude and direction of the 13C shift indicated significantly greater reliance upon nearshore benthic energy sources among nearly all fish taxa as well as profundal invertebrates. Although the mechanisms underlying this 13C shift likely varied among species, possible causes include the transport of benthic algal production to offshore waters and an increased reliance on nearshore prey items. Delta 15N shifts were more variable and of smaller magnitude across taxa although declines in delta 15N among some pelagic fishes may indicate a shift to alternative prey resources. Lake Michigan fishes and invertebrates appear to have responded to dreissenid induced changes in nutrient and energy pathways by switching from pelagic to alternative nearshore energy subsidies. Although large shifts in energy allocation (i.e. pelagic to nearshore benthic) resulting from invasive species appear to have affected total production at upper trophic levels, changes in trophic structure and utilization of novel energy pathways may help to stabilize food webs following species invasions.

Lake Michigan

Scale dependence of disease impacts on quaking aspen (Populus tremuloides) mortality in the southwestern United States

Depending on how disease impacts tree exposure to risk, both the prevalence of disease and disease effects on survival may contribute to patterns of mortality risk across a species' range. Disease may accelerate tree species' declines in response to global change factors, such as drought, biotic interactions, such as competition, or functional traits, such as allometry. To assess the role of disease in mediating mortality risk in quaking aspen ( Populus tremuloides ), we developed hierarchical Bayesian models for both disease prevalence in live aspen stems and the resulting survival rates of healthy and diseased aspen near the species' southern range limit using 5088 individual trees on 281 United States Forest Service Forest Inventory and Analysis plots in the southwestern United States. We found that disease prevalence depended primarily on tree size, tree allometry, and spatial variation in precipitation, while mortality depended on tree size, allometry, competition, spatial variation in summer temperature, and both temporal and spatial variation in summer precipitation. Disease prevalence was highest in large trees with low slenderness found on dry sites. For healthy trees, mortality decreased with diameter, slenderness, and temporal variation in summer precipitation, but increased with competition and spatial variation in summer temperature. Mortality of diseased trees decreased with diameter and aspen relative basal area and increased with mean summer temperature and precipitation. Disease infection increased aspen mortality, especially in trees of intermediate size and trees on plots at climatic extremes (i.e., cool, wet and warm, dry climates). By examining variation in disease prevalence, mortality of healthy trees, and mortality of diseased trees, we showed that the role of disease in aspen tree mortality depended on the scale of inference. For variation among individuals in diameter, disease tended to expose intermediate-size trees experiencing moderate risk to greater risk. For spatial variation in summer temperature, disease exposed lower risk populations to greater mortality probabilities, but the magnitude of this exposure depended on summer precipitation. Furthermore, the importance of diameter and slenderness in mediating responses to climate supports the increasing emphasis on trait variation in studies of ecological responses to global change.

Ecology, Evolution, and Systematics

Evaluating potential conservation conflicts between two listed species: Sea otters and black abalone

Population consequences of endangered species interacting as predators and prey have been considered theoretically and legally, but rarely investigated in the field. We examined relationships between spatially variable populations of a predator, the California sea otter, Enhydra lutris nereis , and a prey species, the black abalone, Haliotis cracherodii . Both species are federally listed under the Endangered Species Act and co-occur along the coast of California. We compared the local abundance and habitat distribution of black abalone at 12 sites with varying densities of sea otters. All of the populations of abalone we examined were in the geographic area currently unaffected by withering disease, which has decimated populations south of the study area. Surprisingly, our findings indicate that sea otter density is positively associated with increased black abalone density. The presence of sea otters also correlated with a shift in black abalone to habitat conferring greater refuge, which could decrease illegal human harvest. These results highlight the need for a multi-species approach to conservation management of the two species, and demonstrate the importance of using field-collected data rather than simple trophic assumptions to understand relationships between jointly vulnerable predator and prey populations.

California

Tick control: Trapping, bio-control, host management and other alternative strategies

Biology of Ticks is the most comprehensive work on tick biology and tick-borne diseases. This second edition is a multi-authored work, featuring the research and analyses of renowned experts across the globe. Spanning two volumes, the book examines the systematics, biology, structure, ecological adaptations, evolution, genomics and the molecular processes that underpin the growth, development and survival of these important disease-transmitting parasites. Also discussed is the remarkable array of diseases transmitted (or caused) by ticks, as well as modern methods for their control. This book should serve as a modern reference for students, scientists, physicians, veterinarians and other specialists. Volume II includes chapters on the ecology of non-nidicolous and nidicolous ticks, genetics and genomics (including the genome of the Lyme disease vector Ixodes scapularis) and immunity, including host immune responses to tick feeding and tick-host interactions, as well as the tick's innate immune system that prevents and/or controls microbial infections. Six chapters cover in depth the many diseases caused by the major tick-borne pathogens, including tick-borne protozoa, viruses, rickettsiae of all types, other types of bacteria (e.g., the Lyme disease agent) and diseases related to tick paralytic agents and toxins. The remaining chapters are devoted to tick control using vaccines, acaricides, repellents, biocontrol, and, finally, techniques for breeding ticks in order to develop tick colonies for scientific study.

Book chapter