A hierarchical model of plumage: morphology,development, and evolution

Plumage is a complex component of the avian phenotype. The plumage of an individual is composed of numerous hierarchically arranged developmental and morphological modules. We present a hierarchical model of plumage that provides an intellectual framework for understanding the development and evolution of feathers. Independence, covariation, and interaction among plumage modules create numerous opportunities for developmental and evolutionary diversification of feather complexity and function. The hierarchical relationships among plumage modules are characterized by both top-down and bottom-up effects in which properties of modules at one level of the hierarchy determine or influence the properties of modules at lower or higher levels of the hierarchy. Plumage metamodules are created by covariation or interaction among modules at different levels of the hierarchy.     

Biological hierarchies and the nature of extinction

Hierarchy theory recognises that ecological and evolutionary units occur in a nested and interconnected hierarchical system, with cascading effects occurring between hierarchical levels. Different biological disciplines have routinely come into conflict over the primacy of different forcing mechanisms behind evolutionary and ecological change. These disconnects arise partly from differences in perspective (with some researchers favouring ecological forcing mechanisms while others favour developmental/historical mechanisms), as well as differences in the temporal framework in which workers operate. In particular, long‐term palaeontological data often show that large‐scale (macro) patterns of evolution are predominantly dictated by shifts in the abiotic environment, while short‐term (micro) modern biological studies stress the importance of biotic interactions. We propose that thinking about ecological and evolutionary interactions in a hierarchical framework is a fruitful way to resolve these conflicts. Hierarchy theory suggests that changes occurring at lower hierarchical levels can have unexpected, complex effects at higher scales due to emergent interactions between simple systems. In this way, patterns occurring on short‐ and long‐term time scales are equally valid, as changes that are driven from lower levels will manifest in different forms at higher levels. We propose that the dual hierarchy framework fits well with our current understanding of evolutionary and ecological theory. Furthermore, we describe how this framework can be used to understand major extinction events better. Multi‐generational attritional loss of reproductive fitness (MALF) has recently been proposed as the primary mechanism behind extinction events, whereby extinction is explainable solely through processes that result in extirpation of populations through a shutdown of reproduction. While not necessarily explicit, the push to explain extinction through solely population‐level dynamics could be used to suggest that environmentally mediated patterns of extinction or slowed speciation across geological time are largely artefacts of poor preservation or a coarse temporal scale. We demonstrate how MALF fits into a hierarchical framework, showing that MALF can be a primary forcing mechanism at lower scales that still results in differential survivorship patterns at the species and clade level which vary depending upon the initial environmental forcing mechanism. Thus, even if MALF is the primary mechanism of extinction across all mass extinction events, the primary environmental cause of these events will still affect the system and result in differential responses. Therefore, patterns at both temporal scales are relevant.     

SCALE AND HIERARCHY IN MACROEVOLUTION

Abstract: Scale and hierarchy must be incorporated into any conceptual framework for the study of macroevolution, i.e. evolution above the species level. Expansion of temporal and spatial scales reveals evolutionary patterns and processes that are virtually inaccessible to, and unpredictable from, short‐term, localized observations. These larger‐scale phenomena range from evolutionary stasis at the species level and the mosaic assembly of complex morphologies in ancestral forms to the non‐random distribution in time and space of the origin of major evolutionary novelties, as exemplified by the Cambrian explosion and post‐extinction recoveries of metazoans, and the preferential origin of major marine groups in onshore environments and tropical waters. Virtually all of these phenomena probably involve both ecological and developmental factors, but the integration of these components with macroevolutionary theory has only just begun. Differential survival and reproduction of units can occur at several levels within a biological hierarchy that includes DNA sequences, organisms, species and clades. Evolution by natural selection can occur at any level where there is heritable variation that affects birth and death of units by virtue of interaction with the environment. This dynamic can occur when selfish DNA sequences replicate disproportionately within genomes, when organisms enjoy fitness advantages within populations (classical Darwinian selection), when differential speciation or extinction occurs within clades owing to organismic properties (effect macroevolution), and when differential speciation or extinction occurs within clades owing to emergent, species‐level properties (in the strict sense species selection). Operationally, emergent species‐level properties such as geographical range can be recognized by testing whether their macroevolutionary effects are similar regardless of the different lower‐level factors that produce them. Large‐scale evolutionary trends can be driven by transformation of species, preferential production of species in a given direction, differential origination or extinction, or any combination of these; the potential for organismic traits to hitch‐hike on other factors that promote speciation or damp extinction is high. Additional key attributes of macroevolutionary dynamics within biological hierarchies are that (1) hierarchical levels are linked by upward and downward causation, so that emergent properties at a focal level do not impart complete independence; (2) hierarchical effects are asymmetrical, so that dynamics at a given focal level need not propagate upwards, but will always cascade downwards; and (3) rates are generally, although not always, faster at lower hierarchical levels. Temporal and spatial patterns in the origin of major novelties and higher taxa are significantly discordant from those at the species and genus levels, suggesting complex hierarchical effects that remain poorly understood. Not only are many of the features promoting survivorship during background times ineffective during mass extinctions, but also they are replaced in at least some cases by higher‐level, irreducible attributes such as clade‐level geographical range. The incorporation of processes that operate across hierarchical levels and a range of temporal and spatial scales has expanded and enriched our understanding of evolution.     

Analyzing nested variation in the body form of Lepomid sunfishes

Phylogenetic hierarchies are often composed of younger diverging lineages nested within older diverging lineages. Comparing phenotypic variation among several hierarchical levels can be used to test hypotheses about selection, phenotypic evolution and speciation. Such hierarchical comparisons have only been performed in threespine stickleback, and so here we use a hierarchical pattern of divergences between near-shore littoral and off-shore pelagic habitats to test for selection on the evolution of body form in Lepomis sunfish in lakes. We compare variation in external body form between fish from littoral and pelagic habitats at three levels: among ecomorphs within individual lake populations (intrapopulation), among populations of the same species in different lakes (interpopulation), and between bluegill and pumpkinseed sunfish species (interspecifically). Using geometric morphometric methods, we first demonstrate that interpopulation variation in mean body form of pumpkinseed sunfish varies with the presence of pelagic habitat. We then incorporate these results with existing data in order to test the similarity of phenotypic divergence between littoral and pelagic habitats at different hierarchical levels. Parallel relationships between certain body form traits (head length, caudal length and pectoral length) and habitat occur at all three levels suggesting that selection persistently acts at all levels to diversify these traits and so may contribute to species formation. For other traits (caudal depth and pectoral altitude), divergence between habitats is inconsistent at different hierarchical levels. Thus, nested biological variation in Lepomid body form reflects a history of deterministic selection and historical contingency, and also identifies traits that likely have likely influenced fitness and so serve important functions.     

Investigating Morphospace Occupation in Multi-Scale Ecological and Evolutionary Data Using Regression Tree: Case Studies and Perspectives

A key challenge in ecology and evolutionary biology is to explain the origin, structure and temporal patterns of phenotypic diversity. With regard to the potentially complex determinism of phenotypic differences, the issue should be comprehended in a general view, across multiple scales and an increasing number of phenomic studies investigate shape variation through large taxonomic, biogeographic or temporal scales. In this context, there is an ever-increasing need to develop new tools for a coherent understanding of morphospace occupation by disentangling and quantifying the main determinants of phenotypic changes. The present study briefly introduce the possibility to use multivariate regression tree technique to cope with morphological data, as embedded in a geometric morphometric framework. It emphasizes that hierarchical partitioning methods produce a hierarchy between causal variables that may help analyzing complexity in multi-scale ecological and evolutionary data. I therefore suggest that morphological studies would benefit from the combined use of the classical statistical models with rapidly emerging and diversifying methods of machine-learning. Doing so allows one to primary explore in an extensive exploratory manner the hierarchy of nested organisational levels underlying morphological variation, and then conduct hypothesis-driven analysis by focusing on a relevant scale or by investigating the appropriate model that reflects hypothesized nested influence of explanatory variables. The outlined approach may help investigating morphospace occupation in an explicitly hierarchical quantitative context.     

Weighted genome trees: refinements and applications

There are many ways to group completed genome sequences in hierarchical patterns (trees) reflecting relationships between their genes. Such groupings help us organize biological information and bear crucially on underlying processes of genome and organismal evolution. Genome trees make use of all comparable genes but can variously weight the contributions of these genes according to similarity, congruent patterns of similarity, or prevalence among genomes. Here we explore such possible weighting strategies, in an analysis of 142 prokaryotic and 5 eukaryotic genomes. We demonstrate that alternate weighting strategies have different advantages, and we propose that each may have its specific uses in systematic or evolutionary biology. Comparisons of results obtained with different methods can provide further clues to major events and processes in genome evolution.     

Application of ITS sequences of nuclear rDNA in phylogenetic and evolutionary studies of angiosperms

Nuclear rRNA genes (rDNA) in angiosperms are arranged in long tandem repeat ing units, much like those of other higher eukaryotes. Owing to rapid concerted evolution, the repeat units have homogenized or nearly so in most species. The internal transcribed spacer (ITS) of nuclear rDNA is composed of ITS1 and ITS2, which are seperated by 5.8S rDNA. The two spacers, ITS1 (187～298 bp) and ITS2 (187～252 bp), can be readily amplified by PCR and sequenced using universal primers. The sequences contain many vari able sites and potential informative sites among related species, and have been proven to be a useful molecular marker in phylogenetic and evolutionary studies of many angiosperm taxa. It can be used not only in classification and phylogenetic inferences at the levels of family, subfamily, tribe, genus and section, but also in reconstruction of reticulate evolution and de tection of the speciation via hybridization and polyploidization. But this region may not be useful for resolving phylogenetic relationships among families or taxa of higher hierarchy ow- ing to the rapid variation of the ITS sequences.     

Guest-Editorial Introduction: Converging Evolutionary Patterns in Life and Culture

The natural world demonstrates signs of spatial–temporal order, an order that appears to us through a series of recognizable, recurring and consecutive patterns, i.e. regularities in forms, functions, behaviors, events and processes. These patterns lend insight into the modes and tempos of evolution and thus into the units, levels, and mechanisms that underlie the evolutionary hierarchy. Contributors to this special issue analyze converging patterns in the biological and sociocultural realm across and beyond classic divisions between micro- and macro-evolution; horizontal/reticulate and vertical evolution; phylogeny, ontogeny and ecology; synchronic and diachronic sociocultural and linguistic research; and tree and network diagrams. Explanations are sought in complexity theory, major transitions of evolution, and process and mechanism approaches to change; and consequences for notions such as “life”, “species”, “biological individuality”, “units” and “levels” of evolution are given.     

On the thermodynamics of multilevel evolution

Biodiversity is hierarchically structured both phylogenetically and functionally. Phylogenetic hierarchy is understood as a product of branching organic evolution as described by Darwin. Ecosystem biologists understand some aspects of functional hierarchy, such as food web architecture, as a product of evolutionary ecology; but functional hierarchy extends to much lower scales of organization than those studied by ecologists. We argue that the more general use of the term “evolution” employed by physicists and applied to non-living systems connects directly to the narrow biological meaning. Physical evolution is best understood as a thermodynamic phenomenon, and this perspective comfortably includes all of biological evolution. We suggest four dynamical factors that build on each other in a hierarchical fashion and set the stage for the Darwinian evolution of biological systems: (1) the entropic erosion of structure; (2) the construction of dissipative systems; (3) the reproduction of growing systems and (4) the historical memory accrued to populations of reproductive agents by the acquisition of hereditary mechanisms. A particular level of evolution can underpin the emergence of higher levels, but evolutionary processes persist at each level in the hierarchy. We also argue that particular evolutionary processes can occur at any level of the hierarchy where they are not obstructed by material constraints. This theoretical framework provides an extensive basis for understanding natural selection as a multilevel process. The extensive literature on thermodynamics in turn provides an important advantage to this perspective on the evolution of higher levels of organization, such as the evolution of altruism that can accompany the emergence of social organization.     

Mapping the evolutionary twilight zone: molecular markers, populations and geography

Since evolutionary processes, such as dispersal, adaptation and drift, occur in a geographical context, at multiple hierarchical levels, biogeography provides a central and important unifying framework for understanding the patterns of distribution of life on Earth. However, the advent of molecular markers has allowed a clearer evaluation of the relationships between microevolutionary processes and patterns of genetic divergence among populations in geographical space, triggering the rapid development of many research programmes. Here we provide an overview of the interpretation of patterns of genetic diversity in geographical and ecological space, using both implicit and explicit spatial approaches. We discuss the actual or potential interaction of phylogeography, molecular ecology, ecological genetics, geographical genetics, landscape genetics and conservation genetics with biogeography, identifying their respective roles and their ability to deal with ecological and evolutionary processes at different levels of the biological hierarchy. We also discuss how each of these research programmes can improve strategies for biodiversity conservation. A unification of these research programmes is needed to better achieve their goals, and to do this it is important to develop cross‐disciplinary communication and collaborations among geneticists, ecologists, biogeographers and spatial statisticians.     

Inferring evolutionary processes from phylogenies

Evolutionary processes shape the regular trends of evolution and are responsible for the diversity and distribution of contemporary species. They include correlated evolutionary change and trajectories of trait evolution, convergent and parallel evolution, differential rates of evolution, speciation and extinction, the order and direction of change in characters, and the nature of the evolutionary process itself—does change accumulate gradually, episodically, or in punctuational bursts. Phylogenies, in combination with information on species, contain the imprint of these historical evolutionary processes. By applying comparative methods based upon statistical models of evolution to well resolved phylogenies, it is possible to infer the historical evolutionary processes that must have existed in the past, given the patterns of diversity seen in the present. I describe a set of maximum likelihood statistical methods for inferring such processes. The methods estimate parameters of statistical models for inferring correlated evolutionary change in continuously varying characters, for detecting correlated evolution in discrete characters, for estimating rates of evolution, and for investigating the nature of the evolutionary process itself. They also anticipate the wealth of information becoming available to biological scientists from genetic studies that pin down relationships among organisms with unprecedented accuracy.     

Cave-adapted evolution in the North American amblyopsid fishes inferred using phylogenomics and geometric morphometrics

Cave adaptation has evolved repeatedly across the Tree of Life, famously leading to pigmentation and eye degeneration and loss, yet its macroevolutionary implications remain poorly understood. We use the North American amblyopsid fishes, a family spanning a wide degree of cave adaptation, to examine the impact of cave specialization on the modes and tempo of evolution. We reconstruct evolutionary relationships using ultraconserved element loci, estimate the ancestral histories of eye-state, and examine the impact of cave adaptation on body shape evolution. Our phylogenomic analyses provide a well-supported hypothesis for amblyopsid evolutionary relationships. The obligate blind cavefishes form a clade and the cave-facultative eyed spring cavefishes are nested within the obligate cavefishes. Using ancestral state reconstruction, we find support for at least two independent subterranean colonization events within the Amblyopsidae. Eyed and blind fishes have different body shapes, but not different rates of body shape evolution. North American amblyopsids highlight the complex nature of cave-adaptive evolution and the necessity to include multiple lines of evidence to uncover the underlying processes involved in the loss of complex traits.     

The evolution of diet breadth: Monophagy and polyphagy in swallowtail butterflies

Much of the world's biodiversity has resulted from specialization of insect populations onto different plant species, partially through evolution of preference in ovipositing females. Here I report an experimental analysis of how an oviposition preference hierarchy has evolved during the evolutionary diversification of an insect group to produce taxa ranging from monophagous to polyphagous. Tests on the Papilio machaon group of swallowtail butterflies show that the preference hierarchy for plant species is evolutionarily dynamic within this species complex, yet constrained among most populations within species, creating a geographic mosaic of populations differing to various degrees in patterns of host preference. The results indicate that different diet breadths can evolve within a group of closely-related species through a combination of conservatism in preference hierarchy among some populations, occasional but rare rearrangements in preference among others, correlations in preference for some plant species, and availability of similarly ranked hosts.     

Evolving Concepts of Bacterial Species

The same evolutionary forces that cause diversification in sexual eukaryotes are expected to cause diversification in bacteria. However, in bacteria, the wider variety of mechanisms for gene exchange (or lack thereof) increases the range of expected diversity patterns compared to those of sexual organisms. Two parallel concepts for bacterial speciation have developed, based on ecological divergence or barriers to recombination in turn. Recent evidence from DNA sequence data shows that both processes can generate independently evolving groups that are equivalent to sexual species and that represent separate arenas within which recombination (when it occurs), selection and drift occur. It remains unclear, however, how often different processes act in concert to generate simple units of diversity, or whether a more complex model of diversity is required, specifying hierarchical levels at which different cohesive processes operate. We advocate an integrative approach that evaluates the effects of multiple evolutionary forces on diversity patterns. There is also great potential for laboratory studies of bacterial evolution that test evolutionary mechanisms inferred from population genetic analyses of multi-locus and genome sequence data.     

Hierarchies and the Sloshing Bucket: Toward the Unification of Evolutionary Biology

Evolutionary biology presents a bewildering array of phenomena to scientists and students alike—ranging from molecules to species and ecosystems; and embracing 3.8 billion years of life’s history on earth. Biological systems are arranged hierarchically, with smaller units forming the components of larger systems. The evolutionary hierarchy, based on replication of genetic information and reproduction, is a complex of genes/organisms/demes/species and higher taxa. The ecological hierarchy, based on patterns of matter–energy transfer, is a complex of proteins/organisms/avatars/local ecosystems/regional ecosystems. All organisms are simultaneously parts of both hierarchical systems. Darwin’s original formulation of natural selection maps smoothly onto a diagram where the two hierarchical systems are placed side-by-side. The “sloshing bucket” theory of evolution emerges from empirical cases in biological history mapped onto this dual hierarchy scheme: little phenotypically discernible evolution occurs with minor ecological disturbance; conversely, greatest concentrations of change in evolutionary history follow mass extinctions, themselves based on physical perturbations of global extent. Most evolution occurs in intermediate-level regional “turnovers,” when species extinction leads to rapid evolution of new species. Hierarchy theory provides a way of integrating all fields of evolutionary biology into an easily understood—and taught—rubric.

Changing conceptions of species

Species are thought by many to be important units of evolution. In this paper, I argue against that view. My argument is based on an examination of the role of species in the synthetic theory of evolution. I argue that if one adopts a gradualist view of evolution, one cannot make sense of the claim that species are units in the minimal sense needed to claim that they are units of evolution, namely, that they exist as discrete entities over time. My second argument is directed against an appeal to Eldredge and Gould's theory of punctuated equilibria to support the claim that species are units of evolution. If one adopts their view, it may be possible to identify discrete temporal entities that can plausibly be termed species, but there is no reason to claim that those entities are units of evolution. Thus, on two plausible interpretations of the role of natural selection in the process of evolution, species are of no special importance. I then consider some of the reasons why species have been thought to be important evolutionary units by many contemporary evolutionary biologists. Finally, I discuss briefly the implications of this conclusion for evolutionary biology.     

A comparative analysis of Y chromosome and mtDNA phylogenies of the Hylobates gibbons

ABSTRACT: BACKGROUND: The evolutionary relationships of closely related species have long been of interest to biologists since these species experienced different evolutionary processes in a relatively short period of time. Comparison of phylogenies inferred from DNA sequences with differing inheritance patterns, such as mitochondrial, autosomal, and X and Y chromosomal loci, can provide more comprehensive inferences of the evolutionary histories of species. Gibbons, especially the genus Hylobates, are particularly intriguing as they consist of multiple closely related species which emerged rapidly and live in close geographic proximity. Our current understanding of relationships among Hylobates species is largely based on data from the maternally-inherited mitochondrial DNAs (mtDNAs). RESULTS: To infer the paternal histories of gibbon taxa, we sequenced multiple Y chromosomal loci from 26 gibbons representing 10 species. As expected, we find levels of sequence variation some five times lower than observed for the mitochondrial genome (mtgenome). Although our Y chromosome phylogenetic tree shows relatively low resolution compared to the mtgenome tree, our results are consistent with the monophyly of gibbon genera suggested by the mtgenome tree. In a comparison of the molecular dating of divergences and on the branching patterns of phylogeny trees between mtgenome and Y chromosome data, we found: 1) the inferred divergence estimates were more recent for the Y chromosome than for the mtgenome, 2) the species H. lar and H. pileatus are reciprocally monophyletic in the mtgenome phylogeny but a H. pileatus individual falls into the H. lar Y chromosome clade. CONCLUSIONS: Based on the ~6.4 kb of Y chromosomal DNA sequence data generated for each of the 26 individuals in this study, we provide molecular inferences on gibbon and particularly on Hylobates evolution complementary to those from mtDNA data. Overall, our results illustrate the utility of comparative studies of loci with different inheritance patterns for investigating potential sex specific processes on the evolutionary histories of closely related taxa, and emphasize the need for further sampling of gibbons of known provenance.     

A hierarchical view of convergent evolution in microbial eukaryotes

Distinguishing convergent evolution from other causes of similarity in organisms is necessary for reconstructing phylogenetic relationships, inferring patterns of character evolution, and investigating the forces of natural selection. In contrast to animals and land plants, the pervasiveness and adaptive significance of convergent evolution in microbes has yet to be systematically explored or articulated. Convergent evolution in microbial eukaryotes, for instance, often involves very distantly related lineages with relatively limited repertoires of morphological features. These large phylogenetic distances weaken the role of ancestral developmental programs on the subsequent evolution of morphological characters, making convergent evolution between very distantly related lineages fundamentally different from convergent evolution between closely related lineages. This suggests that examples of convergence at different levels in the phylogenetic hierarchy offer different clues about the causes and processes of macroevolutionary diversification. Accordingly (and despite opinions to the contrary), I recognize three broad and overlapping categories of phenotypic convergence-"parallel", "proximate" and "ultimate"-that represent either (1) subcellular analogues, (2) subcellular analogues to multicellular systems (and vice versa), or (3) multicellular analogues. Microbial eukaryotes living in planktonic environments, interstitial environments, and the intestinal environments of metazoan hosts provide compelling examples of ultimate convergence. After describing selected examples in microbial eukaryotes, I suggest some future directions needed to more fully understand the hierarchical structure of convergent evolution and the overall history of life.     

Functional morphology and phylogenetic testing within the framework of symecomorphosis

On the basis of the contrasting evolutionary patterns of the Teleostei and the "Chondrostei" the merit of phylogenetic testing is summarized as a non-arbitrary method for assessing the possible role of various designs in producing differential morphological diversity in different lineages. Arguments are presented for the recognition of a genealogical (reproductive, informational) and an ecological hierarchy. Various levels are proposed within hierarchies, because there are processes intrinsic to each level that are not reducible to those of lower levels or subsumed by higher levels. Mutual influences exist between successive levels within a hierarchy and possible interhierarchical mutual influences are hypothesized between organisms, demes, and avatars, and from the germ line to functional units. The term symecomorphosis is proposed to denote the balanced symmetry of the co-existing and mutually interdependent ecological and genealogical hierarchies. Symecomorphosis predicts that a disturbance in environmental systems can destroy this balance with profound effects on the genealogical hierarchy. Using the evolutionary differentiation of four lineages of air breathing teleosts as an example, it is demonstrated how the principle of symecomorphosis can be included in tests establishing a causal relationship between design and differential diversity among lineages.