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Structural Equation Modeling: Applications in ecological and evolutionary biology

This book presents an introduction to the methodology of structural equation modeling, illustrates its use, and goes on to argue that it has revolutionary implications for the study of natural systems. A major theme of this book is that we have, up to this point, attempted to study systems primarily using methods (such as the univariate model) that were designed only for considering individual processes. Understanding systems requires the capacity to examine simultaneous influences and responses. Structural equation modeling (SEM) has such capabilities. It also possesses many other traits that add strength to its utility as a means of making scientific progress. In light of the capabilities of SEM, it can be argued that much of ecological theory is currently locked in an immature state that impairs its relevance. It is further argued that the principles of SEM are capable of leading to the development and evaluation of multivariate theories of the sort vitally needed for the conservation of natural systems. Supplementary information can be found at the authors website, http://www.jamesbgrace.com/ . • Details why multivariate analyses should be used to study ecological systems • Exposes unappreciated weakness in many current popular analyses • Emphasizes the future methodological developments needed to advance our understanding of ecological systems.

Book

Opportunities and challenges of macrogenetic studies

The rapidly emerging field of macrogenetics focuses on analysing publicly accessible genetic datasets from thousands of species to explore large-scale patterns and predictors of intraspecific genetic variation. Facilitated by advances in evolutionary biology, technology, data infrastructure, statistics and open science, macrogenetics addresses core evolutionary hypotheses (such as disentangling environmental and life-history effects on genetic variation) with a global focus. Yet, there are important, often overlooked, limitations to this approach and best practices need to be considered and adopted if macrogenetics is to continue its exciting trajectory and reach its full potential in fields such as biodiversity monitoring and conservation. Here, we review the history of this rapidly growing field, highlight knowledge gaps and future directions, and provide guidelines for further research.

Nature Reviews Genetics

Genetic implications of bottleneck effects of differing severities on genetic diversity in naturally recovering populations: An example from Hawaiian coot and Hawaiian gallinule

The evolutionary trajectory of populations through time is influenced by the interplay of forces (biological, evolutionary, and anthropogenic) acting on the standing genetic variation. We used microsatellite and mitochondrial loci to examine the influence of population declines, of varying severity, on genetic diversity within two Hawaiian endemic waterbirds, the Hawaiian coot and Hawaiian gallinule, by comparing historical (samples collected in the late 1800s and early 1900s) and modern (collected in 2012–2013) populations. Population declines simultaneously experienced by Hawaiian coots and Hawaiian gallinules differentially shaped the evolutionary trajectory of these two populations. Within Hawaiian coot, large reductions (between −38.4% and −51.4%) in mitochondrial diversity were observed, although minimal differences were observed in the distribution of allelic and haplotypic frequencies between sampled time periods. Conversely, for Hawaiian gallinule, allelic frequencies were strongly differentiated between time periods, signatures of a genetic bottleneck were detected, and biases in means of the effective population size were observed at microsatellite loci. The strength of the decline appears to have had a greater influence on genetic diversity within Hawaiian gallinule than Hawaiian coot, coincident with the reduction in census size. These species exhibit similar life history characteristics and generation times; therefore, we hypothesize that differences in behavior and colonization history are likely playing a large role in how allelic and haplotypic frequencies are being shaped through time. Furthermore, differences in patterns of genetic diversity within Hawaiian coot and Hawaiian gallinule highlight the influence of demographic and evolutionary processes in shaping how species respond genetically to ecological stressors.

Ecology and Evolution

Tadpoles: the Biology of Anuran Larvae

The recent alarming declines in amphibian populations worldwide and the suitability of amphibians for use in answering research questions in disciplines as diverse as molecular systematics, animal behavior, ecology, and evolutionary biology have focused enormous attention on tadpoles. Yet despite this growing interest, relatively little is known about these fascinating creatures. n this invaluable reference, leading experts on tadpole biology relate what we currently know about tadpoles and what we might learn from them in the future. Tadpoles provides detailed summaries of tadpole morphology, development, behavior, ecology, and environmental physiology; explores the evolutionary consequences of the tadpole stage; synthesizes available information on their biodiversity, and presents a standardized terminology and an exhaustive literature review of tadpole biology.

Book

Ecological community integration increases with added trophic complexity

The existence of functional biological organization at the level of multi-species communities has long been contested in ecology and evolutionary biology. I found that adding a trophic level to simulated ecological communities enhanced their ability to compete at the community level, increasing the likelihood of one community forcing all or most species in a second community to extinction. Community-level identity emerged within systems of interacting ecological networks, while competitive ability at the community level was enhanced by intense within-community selection pressure. These results suggest a reassessment of the nature of biological organization above the level of species, indicating that the drive toward biological integration, so prominent throughout the history of life, might extend to multi-species communities.

Ecological Complexity

Genomes & islands & evolution: Oh my!

A central question in evolutionary biology is how lineages quickly diversify to occupy different ecological niches, along with determining genomic factors that facilitate evolutionary change. Isolated, oceanic archipelagos are famous for adaptive radiations characterized by endemic, species-rich clades with substantial ecological variation, yet genome resources key to determining eco-evo processes are generally lacking. Here I present a comparison of the number of genome reference assemblies available (as of May 31, 2023) for three major eukaryotic lineages, briefly describe genome sequencing and benchmarking strategies, and highlight as a case study a genome assembly project for Bidens hawaiensis (Koʻokoʻolau, Asteraceae or Compositae; Coreopsidae), a member of a hexaploid Hawaiian plant adaptive radiation. The total number of plant genome references (1,394) was found to substantially lag the total number of genome references for animal (6,003) and fungi (4,400). Improvements to the quality of de novo assembled genomes are fueled by second- and third-generation long-read sequencing advancements, among other sequencing approaches. In conjunction, strategies to improve genome contiguity include optical maps, Hi-C chromatin capture, or trio binning. Continual improvements to genome sequencing and assembly algorithms have brought within reach telomere-to-telomere genome assemblies, albeit this level of sequencing has to date only been achieved in a few cases. With improvements in sequencing techniques and per-base pair costs that continue to trend downward, the number of high-quality genomes is anticipated to continue to increase, leading to the filling in of taxonomic gaps and sampling of groups of taxa from under sampled geographic areas. Increasing the number of plant genome resources available for the study of island endemism could help to shed light on genome-phenome relationships and genome characteristics that have produced the stunning biological diversity that we now observe across the globe.

Capitulum

A common garden super-experiment: An impossible dream to inspire possible synthesis

Global change threatens plant diversity and disrupts its interrelationship with ecosystem structure and function. This disruption in turn undermines confidence in the knowledge ecologists produce, and whether it will translate into multidisciplinary research settings or guide the effective management of natural lands. To address this challenge, ecology needs to consider the interactions between different levels of biological hierarchy, especially how they feedback on, and are mediated by, plant diversity. Doing so will require conducting empirical work and developing theory that simultaneously considers multiple disciplinary perspectives and units of study. Here we advocate the use of common gardens to integrate ecology, evolutionary biology, and ecosystem science through an explicit focus on simultaneous measurement of response variables at multiple levels of biological organization. This approach will provide opportunities to evaluate assumptions important to prediction, such as space-for-time substitution, and tackle the integration of physicochemical and eco-evolutionary foundations to understanding plants and ecosystems. Synthesis: We summarize the large body of research on Sonoran Desert winter annuals to demonstrate how experimental designs that employ common gardens to integrate processes across scales hold special promise. This includes refining trait-based theories of plant strategies, providing insight into ecosystem responses to global change, and collaborating effectively with other scientific disciplines.

Journal of Ecology

Evolutionary fire ecology: An historical account and future directions

The idea that fire acts as an evolutionary force contributing to shaping species traits started a century ago, but had not been widely recognized until very recently. Among the first to realize this force were E dward B . Poulton, R. D ale Guthrie, and E dwin V . Komarek in animals and W illis L . Jepson, W alter W . Hough, T om M . Harris, P hilip V . Wells, and R obert W . Mutch in plants. They were all ahead of their time in their evolutionary thinking. Since then, evolutionary fire ecology has percolated very slowly into the mainstream ecology and evolutionary biology; in fact, this topic is still seldom mentioned in textbooks of ecology or evolution. Currently, there is plenty of evidence suggesting that we cannot understand the biodiversity of our planet without considering the key evolutionary role of fire. But there is still research to be done in order to fully understand fire's contribution to species evolution and to predicting species responses to rapid global changes.

BioScience

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

Signatures of adaptive divergence among populations of an avian species of conservation

Understanding the genetic underpinning of adaptive divergence among populations is a key goal of evolutionary biology and conservation. Gunnison sage‐grouse ( Centrocercus minimus ) is a sagebrush obligate species with a constricted range consisting of seven discrete populations, each with distinctly different habitat and climatic conditions. Though geographically close, populations have low levels of natural gene flow resulting in relatively high levels of differentiation. Here, we use 15,033 SNP loci in genomic outlier analyses, genotype–environment association analyses, and gene ontology enrichment tests to examine patterns of putatively adaptive genetic differentiation in an avian species of conservation concern. We found 411 loci within 5 kbp of 289 putative genes associated with biological functions or pathways that were overrepresented in the assemblage of outlier SNPs. The identified gene set was enriched for cytochrome P450 gene family members (CYP4V2, CYP2R1, CYP2C23B, CYP4B1) and could impact metabolism of plant secondary metabolites, a critical challenge for sagebrush obligates. Additionally, the gene set was also enriched with members potentially involved in antiviral response (DEAD box helicase gene family and SETX). Our results provide a first look at local adaption for isolated populations of a single species and suggest adaptive divergence in multiple metabolic and biochemical pathways may be occurring. This information can be useful in managing this species of conservation concern, for example, to identify unique populations to conserve, avoid translocation or release of individuals that may swamp locally adapted genetic diversity, or guide habitat restoration efforts.

Arizona, Colorado, New Mexico, Utah

Assessing models of speciation under different biogeographic scenarios; An empirical study using multi-locus and RNA-seq analyses

Evolutionary biology often seeks to decipher the drivers of speciation, and much debate persists over the relative importance of isolation and gene flow in the formation of new species. Genetic studies of closely related species can assess if gene flow was present during speciation, because signatures of past introgression often persist in the genome. We test hypotheses on which mechanisms of speciation drove diversity among three distinct lineages of desert tortoise in the genus Gopherus . These lineages offer a powerful system to study speciation, because different biogeographic patterns (physical vs. ecological segregation) are observed at opposing ends of their distributions. We use 82 samples collected from 38 sites, representing the entire species' distribution and generate sequence data for mtDNA and four nuclear loci. A multilocus phylogenetic analysis in *BEAST estimates the species tree. RNA‐seq data yield 20,126 synonymous variants from 7665 contigs from two individuals of each of the three lineages. Analyses of these data using the demographic inference package ∂a∂i serve to test the null hypothesis of no gene flow during divergence. The best‐fit demographic model for the three taxa is concordant with the *BEAST species tree, and the ∂a∂i analysis does not indicate gene flow among any of the three lineages during their divergence. These analyses suggest that divergence among the lineages occurred in the absence of gene flow and in this scenario the genetic signature of ecological isolation (parapatric model) cannot be differentiated from geographic isolation (allopatric model).

Ecology and Evolution

Sight and blindness: The relationship between ostracod eyes, water depth, and light availability in the Arctic Ocean

Eye loss has been a long-standing interest in evolutionary biology. Many organisms that inhabit environments without light penetration, for example the deep sea, exhibit eye loss and thus become blind. However, water-depth distribution of eyes in marine organisms is poorly understood. Ostracods are widely distributed crustaceans, and many sighted marine ostracods have eye tubercles (lenses) on their shells. Since eye tubercles are visible on the shells illustrated in much literature, it is easy to determine their presence or absence via a literature survey. Here, we used a large Arctic-wide ostracod census dataset (Arctic Ostracode Database) to calculate the eye index (the percentage of species with eyes), and compare them with water depth and light availability. As water depth increases, eye index values decrease and become constantly zero in water deeper than 1000 m. Similar decline of sighted species with increasing depth is also known in isopods and amphipods, suggesting that it may be common in other crustaceans and perhaps in deep-sea organisms in general. We also show that eye index values increase as light availability increases. This study is the first to quantify how distributions of sighted and blind species change with light availability, giving baseline information on vision in the deep sea.

Limnology and Oceanography

Individual heterogeneity in growth and age at sexual maturity: A gamma process analysis of capture–mark–recapture data

Knowledge of organisms’ growth rates and ages at sexual maturity is important for conservation efforts and a wide variety of studies in ecology and evolutionary biology. However, these life history parameters may be difficult to obtain from natural populations: individuals encountered may be of unknown age, information on age at sexual maturity may be uncertain and interval-censored, and growth data may include both individual heterogeneity and measurement errors. We analyzed mark–recapture data for Red-backed Salamanders ( Plethodon cinereus ) to compare sex-specific growth rates and ages at sexual maturity. Aging of individuals was made possible by the use of a von Bertalanffy model of growth, complemented with models for interval-censored and imperfect observations at sexual maturation. Individual heterogeneity in growth was modeled through the use of Gamma processes. Our analysis indicates that female P. cinereus mature earlier and grow more quickly than males, growing to nearly identical asymptotic size distributions as males.

Journal of Agricultural, Biological, and Environme

Excursions in fluvial (dis)continuity

Lurking below the twin concepts of connectivity and disconnectivity are their first, and in some ways, richer cousins: continuity and discontinuity. In this paper we explore how continuity and discontinuity represent fundamental and complementary perspectives in fluvial geomorphology, and how these perspectives inform and underlie our conceptions of connectivity in landscapes and rivers. We examine the historical roots of continuum and discontinuum thinking, and how much of our understanding of geomorphology rests on contrasting views of continuity and discontinuity. By continuum thinking we refer to a conception of geomorphic processes as well as geomorphic features that are expressed along continuous gradients without abrupt changes, transitions, or thresholds. Balance of forces, graded streams, and hydraulic geometry are all examples of this perspective. The continuum view has played a prominent role in diverse disciplinary fields, including ecology, paleontology, and evolutionary biology, in large part because it allows us to treat complex phenomena as orderly progressions and invoke or assume equilibrium processes that introduce order and prediction into our sciences. In contrast the discontinuous view is a distinct though complementary conceptual framework that incorporates non-uniform, non-progressive, and non-equilibrium thinking into understanding geomorphic processes and landscapes. We distinguish and discuss examples of three different ways in which discontinuous thinking can be expressed: 1) discontinuous spatial arrangements or singular events; 2) specific process domains generally associated with thresholds, either intrinsic or extrinsic; and 3) physical dynamics or changes in state, again often threshold-linked. In moving beyond the continuous perspective, a fertile set of ideas comes into focus: thresholds, non-equilibrium states, heterogeneity, catastrophe. The range of phenomena that is thereby opened up to scientific exploration similarly expands: punctuated episodes of cutting and filling, discretization of landscapes into hierarchies of structure and control, the work of extreme events. Orderly and progressive evolution towards a steady or ideal state is replaced by chaotic episodes of disturbance and recovery. Recent developments in the field of geomorphology suggest that we may be on the cusp of a new paradigm that recognizes that both continuous and discontinuous processes and mechanisms play a role in fluvial processes and landscape evolution with neither holding sway over the other and both needed to see rivers as they are.

Geomorphology

Global physical controls on estuarine habitat distribution during sea levelchange: Consequences for genetic diversification through time

Determining the extrinsic (physical) factors controlling speciation and diversification of species through time is of key interest in paleontology and evolutionary biology. The role of sea-level change in shaping species richness patterns of marginal marine species has received much attention, but with variable conclusions. Recent work combining genetic data and Geographical Information Systems (GIS)-based habitat modeling yielded a framework for how geomorphology of continental margins mediates genetic connectivity of populations during sealevel change. This approach may ultimately yield insights on how distinct lineages, species, and biodiversity accumulate in coastal settings. Here, we expand this GIS work globally to different geomorphic settings to model estuarine habitat in a larger geographic framework and test how tectonic setting, oceanographic setting, climate, and margin age affect habitat distribution during sea-level change. In addition, independent of estuaries we explore paleobiologic (e.g. Olsson, 1961) and neontolologic effects of sea-level change on evolution, and test the relation between overall shelf area and species richness using data of 1721 fish species. We find 82% global reduction of estuarine habitat abundance at lowstand relative to highstand, and find large habitats change in size much more than small habitats. Consistent with prior work, narrow continental margins have significantly less habitat at highstand and lowstand than wide margins, and narrow margins significantly associate with fore-arc settings, effectively linking tectonic setting to habitat abundance. Surprisingly, narrow margins host greater species richness, a finding which violates the canonical species-area relation. This finding can be explained if: 1) the physical isolation imposed by narrow margins facilitates the formation of new species over time; 2) the sizestability of small habitats, which disproportionately occur on narrow margins, accumulate and retain species extirpated in the more variable habitats on wide margins; or 3) the smaller habitats on narrow margins facilitate greater species richness through greater habitat heterogeneity. These results are generally at odds with prior interpretations, but the combination of richness data and population genetic principles offer a different perspective on these long-studied questions. Finally, we emphasize that the nuance of Pleistocene-Holocene sea level oscillations should be more explicitly considered in genetic studies.

Global and Planetary Change

The airspace is habitat

A preconception concerning habitat persists and has gone unrecognized since use of the term first entered the lexicon of ecological and evolutionary biology many decades ago. Specifically, land and water are considered habitats, while the airspace is not. This might at first seem a reasonable, if unintended, demarcation, since years of education and personal experience as well as limits to perception predispose a traditional view of habitat. Nevertheless, the airspace satisfies the definition and functional role of a habitat, and its recognition as habitat may have implications for policy where expanding anthropogenic development of airspace could impact the conservation of species and subject parts of the airspace to formalized legal protection.

Trends in Ecology and Evolution

Testing for variation in taxonomic extinction probabilities: A suggested methodology and some results

Several important questions in evolutionary biology and paleobiology involve sources of variation in extinction rates. In all cases of which we are aware, extinction rates have been estimated from data in which the probability that an observation (e.g., a fossil taxon) will occur is related both to extinction rates and to what we term encounter probabilities. Any statistical method for analyzing fossil data should at a minimum permit separate inferences on these two components. We develop a method for estimating taxonomic extinction rates from stratigraphic range data and for testing hypotheses about variability in these rates. We use this method to estimate extinction rates and to test the hypothesis of constant extinction rates for several sets of stratigraphic range data. The results of our tests support the hypothesis that extinction rates varied over the geologic time periods examined. We also present a test that can be used to identify periods of high or low extinction probabilities and provide an example using Phanerozoic invertebrate data. Extinction rates should be analyzed using stochastic models, in which it is recognized that stratigraphic samples are random variates and that sampling is imperfect.

Paleobiology

Spatial personalities: A meta-analysis of consistent individual differences in spatial behavior

Individual variation in behavior, particularly consistent among-individual differences (i.e., personality), has important ecological and evolutionary implications for population and community dynamics, trait divergence, and patterns of speciation. Nevertheless, individual variation in spatial behaviors, such as home range behavior, movement characteristics, or habitat use has yet to be incorporated into the concepts or methodologies of ecology and evolutionary biology. To evaluate evidence for the existence of consistent among-individual differences in spatial behavior – which we refer to as “spatial personality” – we performed a meta-analysis of 200 repeatability estimates of home range size, movement metrics, and habitat use. We found that the existence of spatial personality is a general phenomenon, with consistently high repeatability (r) across classes of spatial behavior (r = 0.67–0.82), taxa (r = 0.31–0.79), and time between repeated measurements (r = 0.54–0.74). These results suggest: 1) repeatable spatial behavior may either be a cause or consequence of the environment experienced and lead to spatial personalities that may limit the ability of individuals to behaviorally adapt to changing landscapes; 2) interactions between spatial phenotypes and environmental conditions could result in differential reproduction, survival, and dispersal, suggesting that among-individual variation may facilitate population-level adaptation; 3) spatial patterns of species' distributions and spatial population dynamics may be better understood by shifting from a mean field analytical approach towards methods that account for spatial personalities and their associated fitness and ecological dynamics.

Behavioral Ecology