In historical literature, Edouard van Beneden is mostly remembered for his cytological discoveries. Less well known, however, is that he also introduced evolutionary morphology – and indeed evolutionary theory as such – in the Belgian academic world. The introduction of this research programme cannot be understood without taking both the international and the national context into account. It was clearly the German example of the Jena University that inspired van Beneden in his research interests. The actual launch (...) of evolutionary morphology at his University of Liège was, however, also connected with the dynamic of Belgian university reforms and the local rationale of creating a research “school.” Thanks to his networks, his mastering of the rhetoric of the “new” biology, his low ideological profile and his capitalising on the new academic élan in late-19th century Belgium, van Beneden managed to turn his programme into a local success from the 1870s onwards. Two decades later, however, the conceptual underpinnings of evolutionary morphology came under attack and the “Van Beneden School” lost much of its vitality. Despite this, van Beneden’s evolutionary morphology was prototypical for the research that was to come. He was one of the first scientific heavyweights in Belgium to turn the university laboratory into a centre of scientific practice and the hub of a research school. (shrink)

Although the role of morphology in evolutionary theory remains a subject of debate, assessing the contributions of morphological investigation to evolutionary developmental biology (Evo-devo) is a more circumscribed issue of direct relevance to ongoing research. Historical studies of morphologically oriented researchers and the formation of the Modern Synthesis in the Anglo-American context identify a recurring theme: the synthetic theory of evolution did not capture multiple levels of biological organization. When this feature is incorporated into a philosophical framework (...) for explaining the origin of evolutionary innovations and novelties (a core domain of inquiry in Evo-devo) two specific roles for morphology can be described: (1) the conceptualization and operational identification of the targets of explanation; and (2) the elucidation of causal interactions at higher levels of organization during ontogeny and through evolutionary time. These roles are critical components of any adequate explanation of innovation and novelty though not exhaustive of the parts played by morphology in evolutionary investigation. They also invite reflection on what counts as an evolutionary cause in contemporary evolutionary biology. (shrink)

In historical literature, Edouard van Beneden (1846–1910) is mostly remembered for his cytological discoveries. Less well known, however, is that he also introduced evolutionary morphology – and indeed evolutionary theory as such – in the Belgian academic world. The introduction of this research programme cannot be understood without taking both the international and the national context into account. It was clearly the German example of the Jena University that inspired van Beneden in his research interests. The actual (...) launch of evolutionary morphology at his University of Liège was, however, also connected with the dynamic of Belgian university reforms and the local rationale of creating a research “school.” Thanks to his networks, his mastering of the rhetoric of the “new” biology, his low ideological profile and his capitalising on the new academic élan in late-19th century Belgium, van Beneden managed to turn his programme into a local success from the 1870s onwards. Two decades later, however, the conceptual underpinnings of evolutionary morphology came under attack and the “Van Beneden School” lost much of its vitality. Despite this, van Beneden’s evolutionary morphology was prototypical for the research that was to come. He was one of the first scientific heavyweights in Belgium to turn the university laboratory into a centre of scientific practice and the hub of a research school. (shrink)

One foundational question in contemporarybiology is how to `rejoin evolution anddevelopment. The emerging research program(evolutionary developmental biology or`evo-devo) requires a meshing of disciplines,concepts, and explanations that have beendeveloped largely in independence over the pastcentury. In the attempt to comprehend thepresent separation between evolution anddevelopment much attention has been paid to thesplit between genetics and embryology in theearly part of the 20th century with itscodification in the exclusion of embryologyfrom the Modern Synthesis. This encourages acharacterization of evolutionary developmentalbiology as (...) the marriage of evolutionary theoryand embryology via developmental genetics. Butthere remains a largely untold story about thesignificance of morphology and comparativeanatomy (also minimized in the ModernSynthesis). Functional and evolutionarymorphology are critical for understanding thedevelopment of a concept central toevolutionary developmental biology,evolutionary innovation. Highlighting thediscipline of morphology and the concepts ofinnovation and novelty provides an alternativeway of conceptualizing the `evo and the `devoto be synthesized. (shrink)

What gets integrated in integrative scientific practices has been a topic of much discussion. Traditional views focus on theories and explanations, with ideas of reduction and unification dominating the conversation. More recent ideas focus on disciplines, fields, or specialties; models, mechanisms, or methods; phenomena, problems. How integration works looks different on each of these views since the objects of integration are ontologically and epistemically various: statements, boundary conditions, practices, protocols, methods, variables, parameters, domains, laboratories, and questions all have their own (...) structures, functions and logics. I focus on one particular kind of scientific practice, integration of “approaches” in the context of a research system operating on a special kind of “platform.” Rather than trace a network of interactions among people, practices, and theoretical entities to be integrated, in this essay I focus on the work of a single investigator, David Wake. I describe Wake’s practice of integrative evolutionary biology and how his integration of approaches among biological specialties worked in tandem with his development of the salamanders as a model taxon, which he used as a platform to solve, re-work and update problems that would not have been solved so well by non-integrative approaches. The larger goal of the project to which this paper contributes is a counter-narrative to the story of 20th century life sciences as the rise and march of the model organisms and decline of natural history. (shrink)

With Carl Gegenbaur and Ernst Haeckel, inspiredby Darwin and the cell theory, comparativeanatomy and embryology became established andflourished in Jena. This tradition wascontinued and developed further with new ideasand methods devised by some of Haeckelsstudents. This first period of innovative workin evolutionary morphology was followed byperiods of crisis and even a disintegration ofthe discipline in the early twentieth century.This stagnation was caused by a lack ofinterest among morphologists in Mendeliangenetics, and uncertainty about the mechanismsof evolution. Idealistic morphology (...) was stillinfluental in Germany, which prevented a fullappreciation of the importance of Darwinstheory of natural selection for comparativemorphology. Evolutionary morphology andembryology failed to contribute significantlyto the modern synthesis of evolutionarybiology, thereby probably delaying theintegration of developmental biology intomodern evolutionary biology. However, Haeckelsstudent Oscar Hertwig, as well as Victor Franzand Alexej N. Sewertzoff from a youngergeneration, all tried to forge their ownsynthetic approaches in which (inspired byHaeckels work) embryology played an importantrole. Important for all three researchers wereattempts to refine, and sometimes redefine, thebiogenetic law, and to find new scientificexplanations for it (and for the manyexceptions to it). Their research was latermore or less forgotten, and had littleinfluence on the architects of the modernsynthesis. As the relationship betweenevolutionary and developmental biology is nowagain rising in importance in the form ofEvo-Devo, we would like to draw attention tohow this earlier research tradition grappledwith similar questions to those now on theagenda, albeit from sometimes quite differentperspectives. (shrink)

The mouthparts of anuran tadpoles are highly derived compared to those of caecilians or salamanders. The suprarostral cartilages support the tadpole's upper beak; the infrarostral cartilages support the lower beak. Both supra‐ and infrarostral cartilages are absent in other vertebrates. These differences reflect the evolutionary origin of a derived feeding mode in anuran tadpoles. We suggest that these unique cartilages stem from the evolution of new articulations within preexisting cartilages, rather than novel cartilage condensations. We propose testing this hypothesis (...) through a search for similarities in the development of the suprarostral and infrarostral cartilage articulations and of the primary jaw joint. In Xenopus, the gene zax is expressed in a region corresponding to the infrarostral cartilage. This gene is related to the bapx1‐gene, which regulates jaw joint development. Further investigation of these genes, as well as other genes with joint‐related functions, in anuran craniofacial development may provide a connection between the morphological diversity seen in the vertebrate head and the corresponding diversity in genetic regulatory processes. We believe that the evolution of larval jaws in anurans may shed light on the general evolutionary mechanisms of how new articulations, not only in the jaw region, could have arisen in the vertebrate skull. BioEssays 27: 526–532, 2005. © 2005 Wiley periodicals, Inc. (shrink)

In this study the relationship between functional morpholoy and evolutionary biology is analysed by confronting the main concepts in both disciplines.Rather than only discussing this connection theoretically, the analysis is carried out by introducing important practical and experimental studies, which use aspects from both disciplines. The mentioned investigations are methodologically analysed and the consequences for extensions of the relationship are worked out. It can be shown that both disciplines have a large domain of their own and also share a (...) large common ground. Many disagreements among evolutionary biologists can be reduced to differences in general philosophy (idealism vs. realism), selection of phenomenona (structure vs. function), definition of concepts (natural selection) and the position of the concept theory as an explaining factor (neutralists vs. selectionists, random variation, determinate selection, etc.). (shrink)

Bernard Norton's research concentrated on the Biometrical school of Darwinism and the social implications of the hereditarian ideas that began to gain popularity in the closing years of the nineteenth century. In this article I want to look at the previous generation of evolutionists, the evolutionary morphologists against whom the Biometricians (and their great rivals, the early Mendelians) were reacting. Despite the prominence of evolutionary morphology in the post-Darwinian era, comparatively little historical work has been done on (...) it. In helping to fill this gap, I hope to honour Bernard Norton's memory by throwing light on a movement that forms a conceptual bridge linking the original Darwinian debate to the Biometrical – Mendelian controversy. I shall also argue that evolutionary morphology had ideological overtones that helped to shape the cultural environment within which the eugenics movement would emerge. Although originally a product of the Victorian faith in progress, evolutionary morphology seemed to confirm that exposure to an unstimulating environment led to degeneration. It thus fuelled the concern over racial degeneration which the supporters of eugenics would seek to allay through the application of their new hereditarian philosophy. (shrink)

Efforts from diverse disciplines, including evolutionary studies and biomechanical experiments, have yielded new insights into the genetic, signaling, and mechanical control of tooth formation and functions. Evidence from fossils and non‐model organisms has revealed that a common set of genes underlie tooth‐forming potential of epithelia, and changes in signaling environments subsequently result in specialized dentitions, maintenance of dental stem cells, and other phenotypic adaptations. In addition to chemical signaling, tissue forces generated through epithelial contraction, differential growth, and skeletal constraints (...) act in parallel to shape the tooth throughout development. Here recent advances in understanding dental development from these studies are reviewed and important gaps that can be filled through continued application of evolutionary and biomechanical approaches are discussed. (shrink)

The concept that renders morphology a tool for phylogeny reconstruction is homology. The concept of homology is rooted in pre-evolutionary idealistic morphology. The claim that the goal of idealistic morphology was the seriability of form may sound paradoxical given that this discipline proceeded within a framework of strictly delimited types. But the types only demarcate where seriability starts and where it comes to an end. Carl Gegenbaur’s was recognized as a milestone in idealistic morphology. A (...) comparison with the second edition of 1870 illustrates Gegenbaur’s turn to evolutionary morphology. The methodology remained the same–seriability of form–but the series was no longer merely descriptive or conceptual but now a historical, evolutionary one. Gegenbaur emphasized that seriability of form was possible not only between species of the same type, but also between parts of organisms of the same type. Pursuing this project, he found that different parts of organisms evolve at different rates, resulting in an incongruence between the series of parts relative to the series of species under comparison. This incongrence was called chevauchement des spécialisations by Louis Dollo, Spezialisationskreuzungen by Othenio Abel, and heterobathmy of characters by Armen Takhtajan. Willi Hennig, the founder of modern methods in phylogenetic systematics, discovered that the heterobathmy of characters was a precondition for the establishment of the phylogenetic relationships based on shared derived characters. The result was a replacement of the search for ancestors by a search for relative degrees of phylogenetic relationships. (shrink)

The emerging contemporary natural philosophy provides a common ground for the integrative view of the natural, the artificial, and the human-social knowledge and practices. Learning process is central for acquiring, maintaining, and managing knowledge, both theoretical and practical. This paper explores the relationships between the present advances in understanding of learning in the sciences of the artificial, natural sciences, and philosophy. The question is, what at this stage of the development the inspiration from nature, specifically its computational models such as (...) info-computation through morphological computing, can contribute to machine learning and artificial intelligence, and how much on the other hand models and experiments in machine learning and robotics can motivate, justify, and inform research in computational cognitive science, neurosciences, and computing nature. We propose that one contribution can be understanding of the mechanisms of ‘learning to learn’, as a step towards deep learning with symbolic layer of computation/information processing in a framework linking connectionism with symbolism. As all natural systems possessing intelligence are cognitive systems, we describe the evolutionary arguments for the necessity of learning to learn for a system to reach humanlevel intelligence through evolution and development. The paper thus presents a contribution to the epistemology of the contemporary philosophy of nature. (shrink)

The emerging contemporary natural philosophy provides a common ground for the integrative view of the natural, the artificial, and the human-social knowledge and practices. Learning process is central for acquiring, maintaining, and managing knowledge, both theoretical and practical. This paper explores the relationships between the present advances in understanding of learning in the sciences of the artificial (deep learning, robotics), natural sciences (neuroscience, cognitive science, biology), and philosophy (philosophy of computing, philosophy of mind, natural philosophy). The question is, what at (...) this stage of the development the inspiration from nature, specifically its computational models such as info-computation through morphological computing, can contribute to machine learning and artificial intelligence, and how much on the other hand models and experiments in machine learning and robotics can motivate, justify, and inform research in computational cognitive science, neurosciences, and computing nature. We propose that one contribution can be understanding of the mechanisms of ‘learning to learn’, as a step towards deep learning with symbolic layer of computation/information processing in a framework linking connectionism with symbolism. As all natural systems possessing intelligence are cognitive systems, we describe the evolutionary arguments for the necessity of learning to learn for a system to reach human-level intelligence through evolution and development. The paper thus presents a contribution to the epistemology of the contemporary philosophy of nature. (shrink)

Evolutionary change is opportunistic, but its course is strongly constrained in several fundamental ways. These constraints (historical/phylogenetic, functional/adaptive, constructional/morphogenetic) and their dynamic relationships are discussed here and shown to constitute the conceptual framework of Constructional Morphology. Notwithstanding recent published opinions which claim that the discovery of constraints renders Neodarwinian selection theory obsolete, we regard the insights of Constructional Morphology as being entirely consistent with this theory. As is shown here in the case of the Hyracoidea, formal analysis (...) of the constraints which have framed the evolution of various characters extends our understanding of the evolution of a taxon. (shrink)

From the early nineteenth century, the successful use of fossils in stratigraphy oriented paleontology towards geology. The consequent marginalising of biological objectives was countered in the twentieth century by the rise of ‘Paläobiologie’, first in the German cultural area and only later, as ‘paleobiology’, in the anglophone world. Several kinds of paleobiological research flourished internationally after the Second World War, among them the novel field of ‘paleoecology’. Within this field there were attempts to apply functional morphology to the problematical (...) cases of fossil organisms, for which functions cannot be observed directly. This article describes the origins of the kind of functional inference for fossils that I proposed in 1961 as the method of ‘paradigms’. Here I summarize some of my ‘worked exemplars’, which were intended to show the paradigm method in action. These case-studies were all taken from the paleontologically important phylum of the Brachiopoda, but the method was claimed to have much wider implications for the interpretation of the fossil record in terms of adaptive evolution. This article takes the history of the paradigm method as far as the late 1960s. I hope to trace, in a sequel, its ambivalent fate during the 1970s and beyond, when for example Gould’s critique of ‘the adaptationist programme’ and the rise of computer-based quantitative methods for the evolutionary interpretation of the fossil record led to the relative eclipse of functional morphology in paleontology. (shrink)

Richard Owen has been condemned by Darwinians as an anti-evolutionist and an essentialist. In recent years he has been the object of a revisionist analysis intended to uncover evolutionary elements in his scientific enterprise. In this paper I will examine Owen's evolutionary hypothesis and its connections with von Baer's idea of divergent development. To give appropriate importance to Owen's evolutionism is the first condition to develop an up-to-date understanding of his scientific enterprise, that is to disentagle Owen's contribution (...) to the modernization of typology and morphology. I will argue that Owen's Platonic essentialism is rhetorical and incongruous. On the contrary, an interpretation of the archetype based on Aristotle's biological works makes possible a new conception of type, based on a homeostatic mechanism of stability. The renewal of morphology hinges on homological correspondences and a homeostatic process is also the origin of serial and special homology. I will argue that special homology shows an evolutionary orientation insofar as it is a typically inter-specific character while serial homology is determined through an elementary usage of the categories of developmental morphology. (shrink)

Evolutionary developmental biology (evo-devo) is considered a ‘mechanistic science,’ in that it causally explains morphological evolution in terms of changes in developmental mechanisms. Evo-devo is also an interdisciplinary and integrative approach, as its explanations use contributions from many fields and pertain to different levels of organismal organization. Philosophical accounts of mechanistic explanation are currently highly prominent, and have been particularly able to capture the integrative nature of multifield and multilevel explanations. However, I argue that evo-devo demonstrates the need for (...) a broadened philosophical conception of mechanisms and mechanistic explanation. Mechanistic explanation (in terms of the qualitative interactions of the structural parts of a whole) has been developed as an alternative to the traditional idea of explanation as derivation from laws or quantitative principles. Against the picture promoted by Carl Craver, that mathematical models describe but usually do not explain, my discussion of cases from the strand of evo-devo which is concerned with developmental processes points to qualitative phenomena where quantitative mathematical models are an indispensable part of the explanation. While philosophical accounts have focused on the actual organization and operation of mechanisms, properties of developmental mechanisms that are about how a mechanism reacts to modifications are of major evolutionary significance, including robustness, phenotypic plasticity, and modularity. A philosophical conception of mechanisms is needed that takes into account quantitative changes, transient entities and the generation of novel types of entities, feedback loops and complex interaction networks, emergent properties, and, in particular, functional-dynamical aspects of mechanisms, including functional (as opposed to structural) organization and distributed, system-wide phenomena. I conclude with general remarks on philosophical accounts of explanation. (shrink)

Exploring history pertinent to evolutionary developmental biology (hereafter, Evo-devo) is an exciting prospect given its current status as a cutting-edge field of research. The first and obvious question concerns where to begin searching for materials and sources. Since this new discipline adopts a moniker that intentionally juxtaposes ‘evolution’ and development’, individuals, disciplines, and institutional contexts relevant to the history of evolutionary studies and investigations of ontogeny prompt themselves. Each of these topics has received attention from historians and thus (...) there is both primary and secondary material from which to draw. For example, many historians have documented the historical trajectories of genetics and embryology, their split, and various relations (or lack thereof), especially in the first three decades of the 20th century. (shrink)

Morphological EvoDevo is a field of biological inquiry in which explicit relations between evolutionary patterns and growth or morphogenetic processes are made. Historically, morphological EvoDevo results from the coming together of several traditions, notably Naturphilosophie, embryology, the study of heterochrony, and developmental constraints. A special feature binding different approaches to morphological EvoDevo is the use of formalisms and mathematical models. Here we will introduce anatomical network analysis, a new approach centered on connectivity patterns formed by anatomical parts, with its (...) own concepts and tools specifically designed for the study of morphological EvoDevo questions. Riedl’s concept of burden is tightly related to the use of anatomical networks, providing a nexus between the evolutionary patterns and the structural constraints that shape them. (shrink)

Accounting for the evolutionary origins of morphological novelty is one of the core challenges of contemporary evolutionary biology. A successful explanatory framework requires the integration of different biological disciplines, but the relationships between developmental biology and standard evolutionary biology remain contested. There is also disagreement about how to define the concept of evolutionary novelty. These issues were the subjects of a workshop held in November 2009 at the University of Alberta. We report on the discussion and (...) results of this workshop, addressing questions about (i) how to define evolutionary novelty and understand its significance, (ii) how to interpret evolutionary developmental biology as a synthesis and its relation to neo-Darwinian evolutionary theory, and (iii) how to integrate disparate biological approaches in general. (shrink)

In late winter of 1864, Charles Darwin received two folio volumes on radiolarians, a group of one-celled marine organisms that secreted siliceous skeletons of unusual geometry. The author, the young German biologist Ernst Haeckel (fig. 1), had himself drawn the figures for the extraordinary copper-etched illustrations that filled the second volume.1 The gothic beauty of the plates astonished Darwin (fig. 2 ), but he must also have been drawn to passages that applied his theory to construct the descent relations of (...) these little known creatures. He replied to Haeckel that the volumes "were the most magnificent works which I have ever seen, & I am proud to possess a copy from the author."2 Emboldened by his own initiative in contacting the famous naturalist, Haeckel, a few days later, sent Darwin a newspaper clipping that described a meeting of the Society of German Naturalists and Physicians at Stettin, which occurred during the previous autumn. The article gave an extended and laudatory account of Haeckel's lecture defending Darwin's theory.3 Darwin immediately replied in his second letter: "I.. (shrink)

D’Arcy Thompson’s drawings showing coordinated differences between the shape of an individual of one species and the shape of an individual of another have been reproduced and discussed countless times. However, while they have been widely regarded as inspirational, their interpretation in causal terms has proved difficult, and there is as yet no consensus on this matter. Here, I approach these Thompsonian transformations from a particular angle, namely their dimensional insufficiency. I argue that this problem must be solved before the (...) issue of causality can be considered. This approach leads to the conclusion that Thompsonian transformations—or “morphological transformations”—have not taken place in evolution, and logically can never do so, because they involve the direct conversion of the adult form of one species into that of another. In contrast, “developmental transformations” do occur in the short term, within the lifetime of a single individual. And “evolutionary transformations” occur in the long term, in a context that can be described, following Scott Gilbert, as five-dimensional. I argue that these two kinds of transformation—developmental and evolutionary—have different causal agencies. I consider the possible nature of these agencies and, related to that, the way in which Thompson’s work connects with Darwinian evolutionary theory. (shrink)

Accounting for the evolutionary origins of morphological novelty is one of the core challenges of contemporary evolutionary biology. A successful explanatory framework requires the integration of different biological disciplines, but the relationships between developmental biology and standard evolutionary biology remain contested. There is also disagreement about how to define the concept of evolutionary novelty. These issues were the subjects of a workshop held in November 2009 at the University of Alberta. We report on the discussion and (...) results of this workshop, addressing questions about (i) how to define evolutionary novelty and understand its significance, (ii) how to interpret evolutionary developmental biology as a synthesis and its relation to neo-Darwinian evolutionary theory, and (iii) how to integrate disparate biological approaches in general. (shrink)

The early evolutionary history of echinoderms was reconstructed on the basis of structural-functional considerations and application of the quasi-engineering approach of ‘Konstruktions-Morphologie’. According to the presented evolutionary scenario, a bilaterally symmetrical ancestor, such as an enteropneust-like organism, became gradually modified into a pentaradial echinoderm by passing through an intermediate pterobranch-like stage. The arms of a pentaradial echinoderm are identified as hydraulic outgrowths from the central coelomic cavity of the bilateral ancestor which developed due to a shortening of the (...) body in length but widening in the diameter. The resulting pentaradial symmetry is a consequence of mechanical laws that dictate minimal contact surface areas among hydraulic pneumatic entities. These developed in the coelomic cavity (metacoel) in the bilaterally symmetrical ancestor, when from the already U-shaped mesentery with the intestinal tract two additional U-shaped bows developed directly or subsequently. During the subsequent development tensile chords of the mesentery ‘sewed’ the gut with the body wall first in three and secondly in five ‘seams’. During the direct development five ‘seams’ between tensile chords and body wall developed straightly. These internal tensile chords subdivide the body coelom into five hydraulic subsystems (‘pneus’), which eventually arrange in a pentaradial pattern. The body could then enlarge only between the tensile chords, which means that five hydraulic bulges developed. These bulges initially supported the tentacles and finally each of them enclosed the tentacle until only the feather-like appendages of the tentacles projected over the surface. The tentacles with their feathers were transformedinto the ambulacral system, and the bulges become the arms. These morphological transformations were accompanied and partly determined by specific histological modifications, such as the development of mutable connective tissues and skeletal elements that fused to ossicles and provided shape stabilization in form of a calcareous skeleton in the body wall. The organism resulted was an ancestral echinoderm (‘Ur-Echinoderm’) with an enlarged metacoel, stabilized by hydraulic pressure working againsta capsule of mutable connective tissue, skeletal elements and longitudinal muscles. In regard to these reconstructions, the body structure of echinoderms can be understood as a hydraulic skeletal capsule. (shrink)

Embryogenesis involves biochemical patterning as well as mechanical morphogenetic movements, both regulated by the expression of the regulatory genes of development. The reciprocal interplay of morphogenetic movements with developmental gene expression is becoming an increasingly intense subject of investigation. The molecular processes through which differentiation patterning closely regulates the development of morphogenetic movements are today becoming well understood. Conversely, experimental evidence recently revealed the involvement of mechanical cues due to morphogenetic movements in activating mechano-transduction pathways that control both the differentiation (...) and the active morphogenesis of fundamental events in embryonic tissue development. Here I will first focus on the central role the shape of biological structures of the molecular and mesoscopic cell scales plays in mechano-transduction. Then, after a short description of the genetic regulation of the Drosophila embryo mesoderm invagination at gastrulation—one of the best-understood morphogenetic movement of embryogenesis—I will detail the processes of differentiation and of active morphogenesis of Drosophila embryo gastrulation, which require mechano-transduction. Finally, I will explore the putative consequences in evolution of these mechano-transduction events. I speculate that the first ancient multicellular organisms might have emerged from primary morphological and differentiation patterns induced by primitive mechano-sensation effects allowed by mechano-transduction in response to their physical environment, such as touch by gravity or water flows, which might have acted as primitive motor–sensorial systems, thus conditioning the emergence of our primary animal ancestors. (shrink)

According to many biologists, explaining the evolution of morphological novelty and behavioral innovation are central endeavors in contemporary evolutionary biology. These endeavors are inherently multidisciplinary but also have involved a high degree of controversy. One key source of controversy is the definitional diversity associated with the concept of evolutionary novelty, which can lead to contradictory claims (a novel trait according to one definition is not a novel trait according to another). We argue that this diversity should be interpreted (...) in light of a different epistemic role played by the concept of evolutionary novelty—the structuring of a problem space or setting of an explanatory agenda—rather than the concept’s capacity to categorize traits as novel. This distinctive role is consistent with the definitional diversity and shows that the concept of novelty benefits ongoing investigation by focusing attention on answering different questions related to comprehending the origins of novelty. A review of recent theoretical and empirical work on evolutionary novelty confirms this interpretation. (shrink)

This paper examines a new challenge to neo-Darwinism, a movement known as process structuralism. The process structuralist critique of neo-Darwinism holds 1) that there are general laws in biology and that biologists should search for these laws; 2) that there are general forms of morphology and development and that biologists should attempt to uncover these forms; 3) that organisms are unified wholes that cannot be understood without adopting a holistic perspective; and 4) that no special, causal primacy should be (...) given to the genes in development and morphology. This paper places process structuralism in its historical context, examines the philosophical underpinnings of the movement, and discusses some of the evidence used to support its claims. (shrink)

A concept for a primitive angiospermous branch system is given in order to have a starting point for the derivation of the diverse and highly differentiated branch systems observed in contemporary angiosperms. Hitherto Troll's (1964, 1969) comparative study of the synflorescences in this plant group — developed out predominantly on herbaceous plants — was the most comprehensive and sophisticated treatment dealing with branch systems. Unfortunately, the work on tropical tree architecture by Hallé et al. (1978) has no reference to the (...) classical studies of Troll and his pupils. Thus Müller-Doblies and Weberling (1984) emphasized the high degree of terminological incompatibility between the two works. Angiosperms are seen as a monophyletic plant division. Consequently, the branch system of the first primitive angiosperms must be the starting point in the evolution of the abundant diversity of branch systems and growth forms of modern angiosperms. If it is accepted that primitive angiospermous shoots were terminated by a large flower, one may assume, that the reproductive end of the shoot was enriched by paracladia early in evolution, thereby developing a terminal inflorescence instead of a single flower. Thus the primitive shoot unit was divided into a basal vegetative region — the trophotagma, branching retardively — and the reproductive terminal region, the anthotagma, branching simultaneously. It is demonstrated through a selection of different examples, that the construction of such a system possesses the options for several modifications, enabling the evolution of the abundant diversity of branch systems which characterizes contemporary angiosperms. (shrink)

In plant morphology, most structures of vascular plants can easily be assigned to pre-established organ categories. However, there are also intermediate structures that do not fit those categories associated with a classical approach to morphology. To integrate the diversity of forms in the same general framework, we constructed a theoretical morphospace based on a variety of modalities where it is possible to calculate the morphological distance between plant organs. This paper gives emphasis on shoot, leaf, leaflet and trichomes (...) while ignoring the root. This will allow us to test the hypothesis that classical morphology (typology) and dynamic morphology occupy the same theoretical morphospace and the relationship between the two approaches remains a question of weighting of criteria. Our approach considers the shoot (i.e. leafy stem) as the basic morphological structural unit. A theoretical data table consisting of as many lines as there are possible combinations between different modalities of characters of a typical shoot was generated. By applying a principal components analysis (PCA) to these data it is possible to define a theoretical morphospace of shoots. Typical morphological elements (shoots, leaves, trichomes) and atypical structures (phylloclades, cladodes) including particular cases representing ‘exotic’ structures such as the epiphyllous appendages of Begonia and ‘water shoot’ and ‘leaf’ of aquatic Utricularia were placed in the morphospace. The more an organ differs from a typical shoot, the further away it will be from the barycentre of shoots. By giving a higher weight to variables used in classical typology, the different organ categories appear to be separate, as expected. If we do not make any particular arbitrary choice in terms of character weighting, as it is the case in the context of dynamic morphology, the clear separation between organs is replaced by a continuum. Contrary to typical structures, “intermediate” structures are only compatible with a dynamic morphology approach whether they are placed in the morphospace based on a ponderation compatible with typology or dynamic morphology. The difference in points of view between typology and continuum leads to a particular mode of weighting. By using an equal weighting of characters, contradictions due to the ponderation of characters are avoided, and the morphological concepts of continuum’ and ‘typology’ appear as sub-classes of ‘process’ or ‘dynamic morphology’. (shrink)

The importance of evolutionary parallelisms and their differences from evolutionary convergences have been historically underappreciated, as recently noticed in Gould's last book `The structure of evolutionary history'. In that book, Gould make an effort to distinguish and to reinterpret these concepts in the light of the new discoveries of the last decades on developmental biology and genetics, presenting the elegant metaphor of `Pharaonic bricks versus Corinthian columns'. In this paper I will briefly discuss these concepts, and will (...) argue that, despite the advances that have been made to define them in theory, it is rather hard to differentiate them in a practical phylogenetic context. In order to do so, I will provide some few examples from my own empirical studies on the last years of one of the most morphologically and taxonomically diverse groups of Vertebrates, the catfishes. (shrink)

Gebserian consciousness structures as expressed in human culture are readily understood as analogous to developmental dynamics in holistic biology, and eastern praxis and philosophy. Probably the most integral and fruitful western paradigm of whole-organism biology is that carefully developed in the Goethean tradition to incorporate an understanding of consciousness. This tradition profoundly understands our early childhood through adult development as a dynamic organic process in time, thus integrating the study of heterochrony and chronobiology with morphology, physiology and consciousness. In (...) addition to revealing compelling convergences with the classic work of Gebser on consciousness structures, this Goethian tradition reveals significant parallels with contemporary studies in the realm of biophysics, the science of complexity and eastern spiritual traditions which regard the body as a coherent field, manifesting dynamic, directed currents of development that can be traced and mapped out. These paradigms recognize the importance of recursive organizing principles that structure the biosphere's spatial dimensions and, in the temporal realm, structure long- and short-term evolutionary cycles. The marriage of Gebserian cultural philosophies with Goethean biology, the science of complexity and eastern praxis traditions leads to a simple, elegant and remarkably broad appreciation of the fundamental role of consciousness as an essential property of dynamic living systems. This synthesis helps address many of the difficult questions faced by students of consciousness in a relatively simple and straightforward manner, beginning with a focus on our own species and close mammalian relatives. Two main conclusions arise from this synthesis. First, although enjoying increased attention recently, the Gebserian and Goethean paradigms were far ahead of their time and so have been overlooked. Second, consciousness must be regarded as a primary biologically-embedded datum of life/existence and not merely an epiphenomenon associated with the function of localized or specific organs. (shrink)

Hybridisations between related species with divergent ontogenies can provide insights into the bases for evolutionary change in development. One example of such hybridisations involves sea urchin species that exhibit either standard larval (pluteal) stages or those that develop directly from embryo to adult without an intervening feeding larval stage. In such crosses, pluteal features were found to be restored in fertilisations of the eggs of some direct developing sea urchins (Heliocidaris erythrogramma) with the sperm of closely (Heliocidaris tuberculata) and (...) distantly (Pseudoboletia maculata) related species with feeding larvae. Such results can be argued to support the punctuated equilibrium model—conservation in pluteal regulatory systems and a comparatively rapid switch to direct development in evolution.1,2 Generation of hybrids between distantly related direct developers may, however, indicate evolutionary convergence. The ‘rescue’ of pluteal features by paternal genomes may require maternal factors from H. erythrogramma because the larva of this species has pluteal features. In contrast, pluteal features were not restored in hybridisations with the eggs of Holopneustes purpurescens, which lacks pluteal features. How much of pluteal development can be lost before it cannot be rescued in such crosses? The answer awaits hybridisations among indirect and direct developing sea urchins differing in developmental phenotype, in parallel with investigations of the genetic programs involved. BioEssays 26:343–347, 2004. © 2004 Wiley Periodicals, Inc. (shrink)

The evolutionary embryologist Gavin Rylands de Beer can be viewed as one of the forerunners of modern evolutionary developmental biology in that he posed crucial questions and proposed relevant answers about the causal relationship between ontogeny and phylogeny. In his developmental approach to the phylogenetic phenomenon of homology, he emphasized that homology of morphological structures is to be identified neither with the sameness of the underlying developmental processes nor with the homology of the genes that are in involved (...) in the development of the structures. De Beer’s work on developmental evolution focused on the notion of heterochrony, arguing that paedomorphosis increases morphological evolvability and is thereby an important mode of evolution that accounts for the origin of many taxa, including higher taxa. (shrink)

A 365 million year‐old trilobite moult‐carcass assemblage was described by Błażejowski et al. (2015) as the oldest direct evidence of moulting in the arthropod fossil record. Unfortunately, their suppositions are insufficiently supported by the data provided. Instead, the morphology, configuration and preservational context of the highly fossiliferous locality (Kowala Quarry, Poland) suggest that the specimen consists of two overlapping, queued carcasses. The wider fossil record of moulting actually extends back 520 million years, providing an unparalleled opportunity to study behaviour, (...) ecology and development in early animals. Taking cues from modern analogues, it is possible to quantify precise details about moulting behaviour to determine broad‐scale evolutionary trends, ontogenetic sequences and morphological selection pressures. In this review, we argue that this rich source of data has been underused in evolutionary studies, though has great potential for investigating the life history and evolution of arthropods in deep time. (shrink)

A method of phylogenetic reconstruction as proposed by a number of scientists of the Senckenberg Research Institute is discussed. The method is based on functional-morphological studies, the evolutionary adaptation principle of Bock and Von Wahlert (1965) and so-called model reconstruction. It is argued in this paper that direction of the adaptation process cannot be determined because of lack of knowledge about particular selective forces and that theories of model reconstruction are not open to contradiction in the sense of Popperian (...) falsification. Although it has been claimed that the method provides the only valid directional argument for morphoclines in cladistic studies, it remains unclear how to proceed when morphoclines show contradictory polarities. Moreover, it is doubtful whether polarities of morphoclines can be determined independently of phylogenetic hypotheses, and also whether the use of multistate morphoclines is methodologically valid. By relying on a particular evolutionary theory, i.e. the neo-Darwinian theory, and consequently assigning natural selection as the major agent of directional progress, the Senckenburg method of phylogenetic reconstruction restricts itself to microevolutionary change and, therefore, cannot be used when other hypotheses on the evolutionary process appear to explain the speciation process more plausibly, i.e. hypotheses on macroevolution. Furthermore, it is an unproved statement that evolution always proceeds according to the principle of economy. (shrink)

With the advent of evolutionary developmental research, or EvoDevo, there is hope of discovering the roles that the genetic bases of development play in morphological evolution. Studies in EvoDevo span several levels of organismal organization. Low-level studies identify the ultimate genetic changes responsible for morphological variation and diversity. High-level studies of development focus on how genetic differences affect the dynamics of gene networks and epigenetic interactions to modify morphology. Whereas an increasing number of studies link independent acquisition of (...) homoplastic or convergent morphologies to similar changes in the genomes, homoplasies are not always found to have identical low-level genetic underpinnings. This suggests that a combination of low- and high-level approaches may be useful in understanding the relationship between genetic and morphological variation. Therefore, as an empirical and conceptual framework, we propose the causality horizon to signify the lowest level that allows linking homoplastic morphologies to similar changes in the development. A change in a system below the causality horizon cannot be generalized. In more concrete terms, homoplastic morphologies cannot be reduced to the same change in gene regulation when that change occurs below the causality horizon; rather, a higher-level mechanism should be identified. (shrink)

How Darwin's “endless forms most beautiful” have evolved remains one of the most exciting questions in biology. The significant variety of bacterial shapes is most likely due to the specific advantages they confer with respect to the diverse environments they occupy. While our understanding of the mechanisms generating relatively simple shapes has improved tremendously in the last few years, the molecular mechanisms underlying the generation of complex shapes and the evolution of shape diversity are largely unknown. The emerging field of (...) bacterial evolutionary cell biology provides a novel strategy to answer this question in a comparative phylogenetic framework. This relatively novel approach provides hypotheses and insights into cell biological mechanisms, such as morphogenesis, and their evolution that would have been difficult to obtain by studying only model organisms. We discuss the necessary steps, challenges, and impact of integrating “evolutionary thinking” into bacterial cell biology in the genomic era. (shrink)

The different aspects of floral evolution—Angiosperm descent, floral morphology and pollination ecology—are discussed on the basis of the anthocorm theory of the angiospermous flower. Opposed ideas are critically compared and rejected mainly on account of several inconsistencies and flaws in old floral concepts. Floral evolution passed from a very early phase of dicliny, anemophily and aphananthy of the anthocorm to a phase of incipient entomophily soon associated with a partial sex reversal within the anthocorm. This second phase culminated in (...) the monoclinous and primarily zoophilous flower types, whereas other floral types, more particularly the primarily anemophilous, diclinous flowers, are the direct descendants of the anthocorms of the initial phase. The second phase gave rise to at least two morphologically different flower types and two separate semophyletic lines of floral evolution , in each of which the primary zoophily is almost always associated with the development of intrafloral semaphylls usually of androecial derivation. Early flower types were aphananthous and clearly anthocormoid and assumed the morphology and other characteristics of a true flower by contractions, condensations, readjustments, oligomerisations, reductions and modifications leading to the great variety of adaptive floral patterns.Divers aspects de l'évolution de la fleur — à savoir la descendance des Angiospermes, la morphologie de la fleur, et l'écologie de la pollination — sont discutés à partir de l'hypothèse florale de l'anthocorme. Les opinions opposées, après d'être comparées et critisées, sont rejetées principalement à cause des inconséquences et inconsistances des anciens théorèmes morphologiques concernant l'origine de la fleur. L'évolution de la fleur commençait avec une phase d'antan de diclinie, anémophilie et aphananthie, et s'avançait graduellement dans la direction d'une phase entomophilique initiale, bientôt accompagnée d'un renversement de sexe partiel à l'intérieur de l'anthocorm. La deuxième phase culminait vers les types florals monoclines avec zoophilie primaire pendant que des autres types florals, particulièrement les fleurs diclines avec anémophilie primaire, sont les descendants directs du stade initiale de l'anthocorme. A partir de la phase II , au moins deux types florals différents et aussi deux sémophylèses florals séparés se développaient. Dans chacun de ses phases III et III A la zoophilie primaire est presque toujours accouplée avec l'origine des sémaphylles intrafloraux le plus souvent dérivés des étamines. Les anciens types des fleurs ayant été aphanantique et au début encore nettement anthocormoïdes , graduellement prenaient la forme d'une des plusieurs types floraux adaptifs par une contraction de l'axe florale et par l'entassement, l'oligomérisation et la modification des pièces de la fleur.Die drei Aspekte der Evolution der Blüte — die Abstammungsgeschichte der Angiospermen, Blütenmorphologie und Bestäubungsökologie — werden, ausgehend von der Anthokormtheorie der Angiospermenblüte, eingehend besprochen. Entgegengesetzte Meinungen werden kritisch verglichen und hauptsächlich auf Grund mehrerer Strittigkeiten und Fehler der älteren blütenmorphologischen Axiomen abgelehnt. Die Evolution der Blüte fing an mit einer frühen Phase mit Diklinie, Anemophilie und Aphananthie des Anthokorms und schritt allmählich weiter in die Richtung von beginnender Entomophilie, bald mit einer partiellen Geschlechtsinversion innerhalb des Anthokorms zusammengehend. Diese zweite Phase kulminierte in den monoklinen und primär zoophilen Blütentypen, während andere Blütengestalten, besonders die primär anemophilen und diklinen Angiospermenblüten, die unmittelbaren Abkömmlinge der Initialphase des Anthokorms sind. Aus der zweiten Phase entwickelten sich wenigstens zwei morphologisch differenzierte Blütenformen und auch zwei geschiedene Blütensemophylesen , in jeder die primäre Zoophilie fast immer verbunden ist mit der Entwicklung von intrafloralen und meistens von Staubblättern angeleiteten Semaphyllen . Die frühen Blütentypen waren aphanantisch und anfänglich noch deutlich anthokormoid und nahmen allmählich die entgültige Gestalt von einer der vielen adaptiven Blütentypen an durch eine Verkürzung der Blütenachse, und durch ein Zusammendrängen, Oligomerisation und Abänderung der Blütenteile. (shrink)

When phylogenetic trees constructed from morphological and molecular evidence disagree (i.e. are incongruent) it has been suggested that the differences are spurious or that the molecular results should be preferred a priori. Comparing trees can increase confidence (congruence), or demonstrate that at least one tree is incorrect (incongruence). Statistical analyses of 181 molecular and 49 morphological trees shows that incongruence is greater between than within the morphological and molecular partitions, and this difference is significant for the molecular partition. Because the (...) level of incongruence between a pair of trees gives a minimum bound on how much error is present in the two trees, our results indicate that the level of error may be underestimated by congruence within partitions. Thus comparisons between morphological and molecular trees are particularly useful for detecting this incongruence (spurious or otherwise). Molecular trees have higher average congruence than morphological trees, but the difference is not significant, and both within- and between-partition incongruence is much lower than expected by chance alone. Our results suggest that both molecular and morphological trees are, in general, useful approximations of a common underlying phylogeny and thus, when molecules and morphology clash, molecular phylogenies should not be considered more reliable a priori. (shrink)

Dicotyledons are polarly organised in several ways. In plant morphology polarity, a principle allowing comparison of different plant structures has until yet not been studied. A division** of the vegetative plant in shoot and root as polar structures leads to the distinction of four instead of three basic organs: leaf, shoot axis, root axis and root cap together with the root hairs. The flower is also polarly organised, its poles are formed by the carpels and the stamens. The foliage (...) leaves are also polarly organised which is reflected by the morphological relationship of foliage leaf, stamen and carpel. The stamen uses the hypophyll*** as base of construction and the carpel uses the epiphyll**** as base of construction. Hypophyll and epiphyll are the two poles of the foliage leaf. Root and shoot, the polar entities of the vegetative plant and stamen and carpel, the polar entities of the generative plant, are morphologically correlated. Stamen and carpel can be understood as a combination of the basic organs of vegetative and generative parts of the plant. The basic organs of the generative plant are pollen grain and embryo sack with their gametophytes. The quantitative comparison of variable proportions is supplemented by a qualitative comparison of polarities. The result is, that the organisation type of the dicotyledons can yet be understood as constituted of morphological related parts.* in german Grundorgane. (shrink)

Konrad Lorenz, an ethologist, proposed that certain physical elements are perceived as cute and induce caretaking behavior in other individuals, with the evolutionary function of enhancing offspring survival. He called these features Kindchenschema, baby schema. According to his introspection, these include a large forehead, chubby round features, and chubby cheeks. Previous studies are limited to examining the effects of these facial features on perceived cuteness. However, other morphological factors may be related to perceived cuteness. This study uses Bayesian optimization, (...) one of the global sequential optimization methods for estimating unknown functions, to search for facial morphological features that enhance the perceptions of facial cuteness. We applied Bayesian optimization incorporating Gaussian process ordinal regression, which allows an estimation of the latent cuteness function based on evaluations using the Likert scale. A total of 96 preschool children provided the facial images used in this study. We summarized the facial shape variations using methodologies of geometric morphometrics and principal component analysis up to the third principal component, which we refer to as the face space. A total of 40 participants evaluated the images created by warping the average facial texture of the children's faces with randomly generated parameters in the face space. Facial traits related to perceived cuteness were estimated based on the averaged cuteness function. Perceived cuteness was linked to the relative lower position of facial components and narrower jawline but not to the forehead height. (shrink)

Morphology and its relevance for systematics is a promising field for the application of biosemiotic principles in scientific practice. Genital coupling in spiders involves very complex interactions between the male and female genital structures. As exemplified by two spider species,Nephila clavipesandNephila pilipes ssp. fenestrata, from a biosemiotic point of view the microstructures of the male bulb’s embolus and the corresponding female epigynal and vulval parts form the morphological zone of an intraspecific communication and sign-interpreting process that is one of (...) the prerequisites for sperm transfer. Hence these morphological elements are of high taxonomic value, as they play an essential role in mating and fertilization and consequently in establishing and preserving a reproductive community. Morphology clearly benefits from a biosemiotic approach, as biosemiotics helps to sort out species-specific morphological characters and to avoid problematic typological interpretations. (shrink)

Evolutionary investigations of human crania typically take a limited view of cranial diversity as they discount the possibility that human cranial variation could simply be due to the effects of random genetic drift, gene flow and mutation in favor of natural selection and developmental changes. Natural selection alone cannot explain similarities between patterns of cranial and molecular diversity observed in humans. It appears that the amount of phenotypic variance in the human cranium decreases at the population level as a (...) function of distance from Sub‐Saharan Africa much in the same way as observed for human molecular data.1 BioEssays 29:1185–1188, 2007. © 2007 Wiley Periodicals, Inc. (shrink)

The current Evolutionary Psychology Movement argues that the mind/brain cannot be understood except by conceiving of it as the product of design by natural selection. Cognitive science should proceed by reconstructing the adaptive pressures that shaped the mind and then looking for mental structure predicted by this these reconstructions. It is argued that this is not a practical solution, because our ability to reconstruct the evolutionary pressures that shaped the mind is exactly proportional to our preexisting understanding of (...) mental structure. EPM's criticisms of contemporary cognitive science are strikingly similar to the criticisms of behavioristic psychology made by the founders of ethology, Lorenz and Tinbergen, fifty years ago. But the ethologists drew the conclusion that psychology should investigate the mind's performance on ecologically valid tasks. This is shown to be a more tractable proposal for injecting an evolutionary perspective into cognitive science. In the second part of the paper it is argued that the concept of 'design' central to EPM is not fully naturalistic, in the sense that it envisages a process more like intelligent design than like a realistic model of evolution by natural selection. An enriched model of evolutionary psychology is outlined, modeled on recent 'evolutionary developmental biology' approaches to morphological evolution, and which we call 'evolutionary developmental psychology'. Like 'evo-devo' more generally, EDP recognizes that selection builds systems that develop in an environment, not phenotypes that are transmitted. The nature of evolutionary dynamics is a function of the nature of development. It is argued that EDP allows a more productive integration of the role of the environment in psychological development into evolutionary psychology than has so far been achieved by EPM. (shrink)

Many of the current comparisons of taxic phylogenetic and biological homology in the context of morphology focus on what are seen as categorical distinctions between the two concepts. The first, it is claimed, identifies historical patterns of conservation and variation relating taxa; the second provides a causal framework for the explanation of this conservation and variation. This leads to the conclusion that the two need not be placed in conflict and are in fact compatible, having non-competing epistemic purposes or (...) mapping the same extensions in the form of monophyletic groupings (see Roth, The biological basis of homology 1–26, 1988; Sluys, J Zool Syst Evol Res 34:145–152, 1996; Abouheif, Trends Ecol Evol 12:405–408, 1997; Brigandt, J Exp Zool 299:9–17, 2003, Biol Philos 22:709–725, 2007; Assis and Brigandt, Evol Biol 36:248–255, 2009). This article argues that moves in this direction miss the essential disagreement between these concepts as they have been developed in the context of the debate concerning the best concept for evolutionary investigation. We should rather see these concepts employing a common fundamental methodological approach to homology, but disagreeing about how to apply the methodology effectively. Both concepts employ class reasoning, which pursues homologies as units of generalization—more precisely, as sources of reliable and relevant group-bound information in the form of shared underlying causes. The dispute can be better understood by two poles that structure such reasoning: the need for a reliable basis for projections about the causal history of shared structures, and the desire to identify homologous characters with more informative and specific causal information relevant to generalizing about evolutionary processes. Judgments in favor of one or the other in turn have affected the scope or extension of these competing homology concepts. (shrink)

Organisms are self-producing and self-maintaining, or autopoietic systems. Therefore, the course of evolution and adaptation of an organism is strongly determined by its own internal properties, whatever role external selection may play. The internal properties may either act as constraints that preclude certain changes or they open new pathways: the organism canalizes its own evolution. As an example the evolution of feeding mechanisms in salamanders, especially in the lungless salamanders of the family Plethodontidae, is discussed. In this family a large (...) variety of different feeding mechanisms is found. The authors reconstruct this evolutionary process as a series of bifurcation points of either constraints or opportunities forming a sequence of preconditions for the formation of a high-speed projectile tongue characteristic of tropical salamanders. Furthermore, it is shown how parallel evolution of seemingly unrelated domains within an organism such as respiratory physiology, life history biology and pattern of ontogeny has rather direct relevance to the feeding biology, thus demonstrating that organisms always evolve as wholes. (shrink)