Preface to the Princeton Landmarks in Biology Edition vii Preface xi Symbols Used xiii 1. The Importance of Islands 3 2. Area and Number of Speicies 8 3. Further Explanations of the Area-Diversity Pattern 19 4. The Strategy of Colonization 68 5. Invasibility and the Variable Niche 94 6. Stepping Stones and Biotic Exchange 123 7. Evolutionary Changes Following Colonization 145 8. Prospect 181 Glossary 185 References 193 Index 201

The Theory of Island Biogeography

Hybrid zones are, roughly speaking, narrow regions in which genetically distinct populations meet, mate, and produce hybrids. They are often only a few hundred meters wide and yet may be several hundred kilometers long. They are found in a wide variety of organisms: Tables 1 and 2 list about 150 reasonably clear examples, in which there is a spatial transition between two hybridizing forms. Such a widespread and striking phenomenon requires explanation; more importantly, it offers us several ways of understanding the nature and origin of species. First, hybrid zones pose interesting questions for the taxonomist, for they contrast two views of the species: as a set of populations delimited by genetic barriers to gene exchange; and as a set of populations maintained in a particular stable equilibrium by selection. Second, the wide range of genotypes found in a hybrid zone can be used to analyze the genetic differences and selective forces that separate the taxa involved. This may allow some inferences about the way these differences evolved and, by extrapolation, about the way fully isolated species diverge from each other. Finally, models of parapatric speciation, and of Wright's "shifting balance," involve the formation, move­ ment, and modification of hybrid zones. Hybrid zones must be understood before the plausibility of these models can be judged. After clarifying our terminology, we will summarize the relevant theory and then see how this can be used to make inferences from field data. A notable increase in the number of detailed genetic studies of hybrid zones occurred over

Analysis of Hybrid Zones

http://aerg.canberra.edu.au/library/sex_general/2002_Devlin_Nagahama_sex_determination_in_fish_review.pdf

Sex determination and sex differentiation in fish: an overview of genetic, physiological, and environmental influences

With these words Darwin opened a brief paragraph citing observations antithetical to his supposition: anecdotal reports of the inheritance of characters missing from one side of the body. His initial hunch, however, has stood the test of time: Genetic studies have confirmed that where only small, random deviations from bilateral symmetry exist, the deviations in a particular direction have little or no measurable heritability (17, 47, 51, 65a, 73a, 74, 91). The genetic basis of bilateral symmetry thus appears to differ fundamentally from that of virtually all other morphological features. Why, then, all the recent (and not so recent) interest in such minor, nondirectional deviations from bilateral symmetry [fluctuating asymmetry (FA); 60 cited in 99]? Four reasons. First, FA relates in a curious way to what is perhaps the major unsolved general problem in modem biology: the orderly expression of genotypes as complex, three-dimensional phenotypes. As was emphasized in a flurry of activity in the mid to late 1950s, and many times since, FA provides an appealing measure of 'developmental noise,' or minor environmentally induced departures from some ideal developmental program (101). Its appeal derives from an a priori knowledge of the ideal: perfect bilateral symmetry. For unilateral characters, the ideal is rarely known (but see 1, 2, and 59 for one possible approach). A second reason for interest in

Fluctuating Asymmetry: Measurement, Analysis, Patterns

The ecological hypothesis of speciation is that reproductive isolation evolves ultimately as a consequence of divergent natural selection on traits between environments. Ecological speciation is general and might occur in allopatry or sympatry, involve many agents of natural selection, and result from a combination of adaptive processes. The main difficulty of the ecological hypothesis has been the scarcity of examples from nature, but several potential cases have recently emerged. I review the mechanisms that give rise to new species by divergent selection, compare ecological speciation with its alternatives, summarize recent tests in nature, and highlight areas requiring research.

/pdf/ecology-and-the-origin-of-species-38r7yix3b6.pdf

Ecology and the origin of species

Fluctuating asymmetry consists of random deviations from perfect symmetry in populations of organisms. It is a measure of developmental noise, which reflects a population’s average state of adaptation and coadaptation. Moreover, it increases under both environmental and genetic stress, though responses are often inconsistent. Researchers base studies of fluctuating asymmetry upon deviations from bilateral, radial, rotational, dihedral, translational, helical, and fractal symmetries. Here, we review old and new methods of measuring fluctuating asymmetry, including measures of dispersion, landmark methods for shape asymmetry, and continuous symmetry measures. We also review the theory, developmental origins, and applications of fluctuating asymmetry, and attempt to explain conflicting results. In the process, we present examples from the literature, and from our own research at “Evolution Canyon” and elsewhere.

https://www.mdpi.com/2073-8994/2/2/466/pdf

Fluctuating Asymmetry: Methods, Theory, and Applications

Fluctuating asymmetry is the most commonly used measure of developmental instability. Some authors have claimed that antisymmetry and directional asymmetry may have a significant genetic basis, thereby rendering these forms of asymmetry useless for studies of developmental instability. Using a modified Rashevsky-Turing reaction-diffusion model of morphogenesis, we show that both antisymmetry and directional asymmetry can arise from symmetry-breaking phase transitions. Concentrations of morphogen on right and left sides can be induced to undergo transitions from phase-locked periodicity, to phase-lagged periodicity, to chaos, by simply changing the levels of feedback and inhibition in the model. The chaotic attractor has two basins of attraction-right sidedominance and left side dominance. With minor disturbance, a developmental trajectory settles into one basin or the other. With increasing disturbance, the trajectory can jump from basin to basin. The changes that lead to phase transitions and chaos are thoseexpected to occur with either genetic change or stress. If we assume that the morphogen influences the behavior of cell populations, then a transition from phase-locked periodicity to chaos in the morphogen produces a corresponding transition from fluctuating asymmetry to antisymmetry in both morphogen concentrations and cell populations. Directional asymmetry is easily modeled by introducing a bias in the conditions of the simulation. We discuss the implications of this model for researchers using fluctuating asymmetry as an indicator of stress.

Antisymmetry, directional asymmetry, and dynamic morphogenesis

Does endogenous or exogenous selection stabilize the big sagebrush (Artemisia tridentata) hybrid zone? After two years of study, our reciprocal transplant experiments showed significant genotype by environment interactions for a number of fitness components, including germination, growth, and reproduction. Hybrids were the most fit within the hybrid garden. In the parental gardens, the native parental taxon was more fit than either the alien parental or hybrids. These results are consistent with the bounded hybrid superiority model, which assumes exogenous selection, but are clearly at odds with the dynamic equilibrium model, which assumes endogenous selection and universal hybrid unfitness.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1558-5646.1997.tb02391.x

Narrow hybrid zone between two subspecies of big sagebrush (artemisia tridentata: asteraceae). iv. reciprocal transplant experiments

Plant developmental stability has received little attention in the past three or four decades. Here we review differences in plant and animal development, and discuss the advantages of using plants as experimental subjects in exploring developmental stability. We argue that any type of developmental invariant may be used to assess developmental stability and review the use of fluctuating asymmetry in studies of plant developmental stability. We also examine the use of deviations from translatory, radial, and self-symmetry as measures of developmental instability. The role of nonlinear dynamics and epigenesis in the production of the phenotype is also discussed.

Developmental stability in plants: symmetries, stress and epigenesis

Three widely used methods of estimating fluctuating asymmetry may yield serious overestimates if directional asymmetry is present. When two sides of a bilateral trait grow at different rates, then the asymmetry variance (Var[ l−r ]) increases with size, even when developmental noise is nil. But the residual variance around a population's mean developmental trajectory is invariant with respect to size. Thus, it can be used as a measure of developmental instability. We introduce a measure of developmental instability, the residual variance ( s 2 δ ), obtainable from either a major axis regression, which is equivalent to a principal component analysis on l and r , or a general structural model. This residual variance can be estimated from directionally asymmetric or even antisymmetric traits. We present examples of developmental instability estimated from directionally asymmetric mandibles (house mouse) and leaves (soybean), and antisymmetric claws (fiddler crab).

John H. Graham

Papers

Fluctuating Asymmetry: Methods, Theory, and Applications

Antisymmetry, directional asymmetry, and dynamic morphogenesis

Narrow hybrid zone between two subspecies of big sagebrush (artemisia tridentata: asteraceae). iv. reciprocal transplant experiments

Developmental stability in plants: symmetries, stress and epigenesis

Directional asymmetry and the measurement of developmental instability