Edward Hitchcock’s Pre-Darwinian (1840) “Tree of Life”

The “tree of life” iconography, representing the history of life, dates from at least the latter half of the 18th century, but evolution as the mechanism providing this bifurcating history of life did not appear until the early 19th century. There was also a shift from the straight line, scala naturae view of change in nature to a more bifurcating or tree-like view. Throughout the 19th century authors presented tree-like diagrams, some regarding the Deity as the mechanism of change while others argued for evolution. Straight-line or anagenetic evolution and bifurcating or cladogenetic evolution are known in biology today, but are often misrepresented in popular culture, especially with anagenesis being confounded with scala naturae. Although well known in the mid 19th century, the geologist Edward Hitchcock has been forgotten as an early, if not the first author to publish a paleontologically based “tree of life” beginning in 1840 in the first edition of his popular general geology text Elementary Geology. At least 31 editions were published and those between 1840 and 1859 had this “paleontological chart” showing two trees, one for fossil and living plants and another for animals set within a context of geological time. Although the chart did not vary in later editions, the text explaining the chart did change to reflect newer ideas in paleontology and geology. Whereas Lamarck, Chambers, Bronn, Darwin, and Haeckel saw some form of transmutation as the mechanism that created their “trees of life,” Hitchcock, like his contemporaries Agassiz and Miller, who also produced “trees of life,” saw a deity as the agent of change. Through each edition of his book Hitchcock denounced the newer transmutationist hypotheses of Lamarck, then Chambers, and finally Darwin in an 1860 edition that no longer presented his tree-like “paleontological chart.”     

The roots of phylogeny: how did Haeckel build his trees?

Haeckel created much of our current vocabulary in evolutionary biology, such as the term phylogeny, which is currently used to designate trees. Assuming that Haeckel gave the same meaning to this term, one often reproduces Haeckel's trees as the first illustrations of phylogenetic trees. A detailed analysis of Haeckel's own evolutionary vocabulary and theory revealed that Haeckel's trees were genealogical trees and that Haeckel's phylogeny was a morphological concept. However, phylogeny was actually the core of Haeckel's tree reconstruction, and understanding the exact meaning Haeckel gave to phylogeny is crucial to understanding the information Haeckel wanted to convey in his famous trees. Haeckel's phylogeny was a linear series of main morphological stages along the line of descent of a given species. The phylogeny of a single species would provide a trunk around which lateral branches were added as mere ornament; the phylogeny selected for drawing a tree of a given group was considered the most complete line of progress from lower to higher forms of this group, such as the phylogeny of Man for the genealogical tree of Vertebrates. Haeckel's phylogeny was mainly inspired by the idea of the scala naturae, or scale of being. Therefore, Haeckel's genealogical trees, which were only branched on the surface, mainly represented the old idea of scale of being. Even though Haeckel decided to draw genealogical trees after reading On the Origin of Species and was called the German Darwin, he did not draw Darwinian branching diagrams. Although Haeckel always saw Lamarck, Goethe, and Darwin as the three fathers of the theory of evolution, he was mainly influenced by Lamarck and Goethe in his approach to tree reconstruction.     

The Charms of Nature: Darwin on Meaning and Value

I review George Levine’s provocative and highly original book Darwin Loves You. Levine, whose “home discipline” is English Literature, offers a compelling interpretation of Darwin’s works, evaluating their content and Darwin’s prose style to identify a distinctly Darwinian attitude toward nature as a source of meaning and value. Levine believes that Darwin exemplifies the capacity to feel “enchantment” about the natural world, suggesting that, if Darwin’s example were followed, a “Darwinian re-enchantment of the world” would be brought about. This would offer a secular, non-supernatural basis for purpose, meaning, and value. I conclude with a few critical remarks about the scope and cogency of Levine’s proposal.     

Don’t Call it “Darwinism”

Evolutionary biology owes much to Charles Darwin, whose discussions of common descent and natural selection provide the foundations of the discipline. But evolutionary biology has expanded well beyond its foundations to encompass many theories and concepts unknown in the 19th century. The term “Darwinism” is, therefore, ambiguous and misleading. Compounding the problem of “Darwinism” is the hijacking of the term by creationists to portray evolution as a dangerous ideology—an “ism”—that has no place in the science classroom. When scientists and teachers use “Darwinism” as synonymous with evolutionary biology, it reinforces such a misleading portrayal and hinders efforts to present the scientific standing of evolution accurately. Accordingly, the term “Darwinism” should be abandoned as a synonym for evolutionary biology.     

Joseph Hooker Takes a “Fixed Post”: Transmutation and the “Present Unsatisfactory State of Systematic Botany”, 1844–1860

Joseph Hooker first learned that Charles Darwin believed in the transmutation of species in 1844. For the next 14 years, Hooker remained a “nonconsenter” to Darwin’s views, resolving to keep the question of species origin “subservient to Botany instead of Botany to it, as must be the true relation”. Hooker placed particular emphasis on the need for any theory of species origin to support the broad taxonomic delimitation of species, a highly contentious issue. His always provisional support for special creation waned during the 1850s as he lost faith in its expediency for coordinating the study of plant geography, systematics and physiology. In 1858, Hooker embraced Darwin’s “considerable revolution in natural history,” but only after Darwin had carefully molded his transmutationism to meet Hooker’s exacting specifications.     

Brocchi,Darwin, and Transmutation: Phylogenetics and Paleontology at the Dawn of Evolutionary Biology

Giambattista Brocchi’s (1814) monograph (see Dominici, Evo Edu Outreach, this issue, 2010) on the Tertiary fossils of the Subappenines in Italy—and their relation to the living molluscan fauna—contains a theoretical, transmutational perspective (“Brocchian transmutation”). Unlike Lamarck (1809), Brocchi saw species as discrete and fundamentally stable entities. Explicitly analogizing the births and deaths of species with those of individual organisms (“Brocchi’s analogy”), Brocchi proposed that species have inherent longevities, eventually dying of old age unless driven to extinction by external forces. As for individuals, births and deaths of species are understood to have natural causes; sequences of births and deaths of species produce genealogical lineages of descent, and faunas become increasingly modernized through time. Brocchi calculated that over 50% of his fossil species are still alive in the modern fauna. Brocchi’s work was reviewed by Horner (1816) in Edinburgh. Brocchi’s influence as a transmutational thinker is clear in Jameson’s (1827) “geological illustrations” in his fifth edition of his translation of Cuvier’s Theory of the Earth (read by his student Charles Darwin) and in the anonymous essays of 1826 and 1827 published in the Edinburgh New Philosophical Journal—which also carried a notice of Brocchi’s death in 1827. The notion that new species replace older, extinct ones—in what today would be called an explicitly phylogenetic context—permeates these essays. Herschel’s (1830) discussion of temporal replacement of species and the modernization of faunas closely mirrors these prior discussions. His book, dedicated to the search for natural causes of natural phenomena, was read by Charles Darwin while a student at Cambridge. Darwin’s work on HMS Beagle was in large measure an exploration of replacement patterns of “allied forms” of endemic species in time and in space. His earliest discussions of transmutation, in his essay February 1835, as well as the Red Notebook and the early pages of Notebook B (the latter two written in 1837 back in England), contain Brocchi’s analogy, including the idea of inherent species longevities. Darwin’s first theory of the origin of species was explicitly saltational, invoking geographic isolation as the main cause of the abrupt appearance of new species. We conclude that Darwin was testing the predicted patterns of both Brocchian and Lamarckian transmutation as early as 1832 at the outset of his work on the Beagle.     

Dimetrodon Is Not a Dinosaur: Using Tree Thinking to Understand the Ancient Relatives of Mammals and their Evolution

The line of descent that includes all living mammals extends back in time over 300 million years. Many of the ancient relatives of mammals that fall along this line are very different in appearance from living mammals and are frequently mistaken for reptiles such as dinosaurs. This misconception is reinforced by the fact that these animals are often referred to as “mammal-like reptiles,” a term reflecting outdated methods for classifying organisms. In reality, these ancient mammal-relatives, known as synapsids, are more closely related to living mammals than they are to any reptiles. Evolutionary trees, which depict patterns of descent from common ancestors among organisms, are very useful for understanding why this is the case and for reconstructing the evolutionary histories of many of the unique characters found in mammals. Here, I provide an introduction to evolutionary trees and their implications for understanding the relationships between mammals, synapsids, and reptiles. This is followed by a review of synapsid diversity and a discussion of how evolutionary trees can be used to investigate when in synapsid history different mammalian characteristics first appeared.     

Separated at Birth: The Interlinked Origins of Darwin’s Unconscious Selection Concept and the Application of Sexual Selection to Race

This essay traces the interlinked origins of two concepts found in Charles Darwin’s writings: “unconscious selection,” and sexual selection as applied to humanity’s anatomical race distinctions. Unconscious selection constituted a significant elaboration of Darwin’s artificial selection analogy. As originally conceived in his theoretical notebooks, that analogy had focused exclusively on what Darwin later would call “methodical selection,” the calculated production of desired changes in domestic breeds. By contrast, unconscious selection produced its results unintentionally and at a much slower pace. Inspiration for this concept likely came from Darwin’s early reading of works on both animal breeding and physical ethnology. Texts in these fields described the slow and unplanned divergence of anatomical types, whether animal or human, under the guidance of contrasting ideals of physical perfection. These readings, it is argued, also led Darwin to his theory of sexual selection as applied to race, a theme he discussed mainly in his book The Descent of Man (1871). There Darwin described how the racial version of sexual selection operated on the same principle as unconscious selection. He thereby effectively reunited these kindred concepts.     

The Preface to Darwin’s <Emphasis Type="Italic">Origin of Species</Emphasis>: The Curious History of the “Historical Sketch” *

Almost any modern reader’s first encounter with Darwin’s writing is likely to be the “Historical Sketch,” inserted by Darwin as a preface to an early edition of the Origin of Species, and having since then appeared as the preface to every edition after the second English edition. The Sketch was intended by him to serve as a short “history of opinion” on the species question before he presented his own theory in the Origin proper. But the provenance of the “Historical Sketch” is somewhat obscure. Some things are known about its production, such as when it first appeared and what changes were made to it between its first appearance in 1860 and its final form, for the fourth English edition, in 1866. But how it evolved in Darwin’s mind, why he wrote it at all, and what he thought he was accomplishing by prefacing it to the Origin remain questions that have not been carefully addressed in the scholarly literature on Darwin. I attempt to show that Darwin’s various statements about the “Historical Sketch,” made primarily to several of his correspondents between 1856 and 1860, are somewhat in conflict with one another, thus making problematic a satisfactory interpretation of how, when, and why the Sketch came to be. I also suggest some probable resolutions to the several difficulties. How Darwin came to settle on the title “Historical Sketch” for the Preface to the Origin is not certain, but a guess may be ventured. When he first submitted the text to Asa Gray in February 1860 he called it simply “Preface Contributed by the Author to this American Edition” (Burkhardt et al., eds., vol. 8, 1993, p. 572; the collected correspondence is hereafter cited as CCD). In fact he had thought of it as being properly called a Preface much earlier, perhaps as early as 1856, as will be seen in what follows. It came to be called “An Historical Sketch of the Recent Progress of Opinion on the Origin of Species” only in the third English edition, April 1861. This is the title it retained thereafter, with the exception of an addition to the title in the sixth English edition, “Previously to the Publication of the First Edition of this Work” (Peckham, 1959, pp. 20, 59). The word “sketch,” on the other hand was one of two words Darwin commonly used in private correspondence to refer to the book that would later become the Origin, the other word being “Abstract,” and both signifying that Darwin thought of the work as being a resume rather than a full-fledged study (e.g., letter to J.D. Hooker, May 9 1856, CCD vol. 6 p. 106; letter to Baden Powell January 18 1860, CCD vol. 8 p. 41; letter to Lyell 25 June 1858, CCD v. 7, 1991, pp. 117–8; letter to Lyell May 1856, CCD, v. 6 p. 100). The most likely source of the title “Historical Sketch” for Darwin’s Preface is Charles Lyell’s Principles of Geology in which, beginning with the third edition (1834), Lyell added titles to his chapters, calling chapters 2–4 “Historical Sketch of the Progress of Geology” (Secord, in Lyell [1997], p. xlvii; for other uses by Lyell of this expression, cf. Porter, 1976, p. 95; idem 1982, p. 38; and Lyell, 1830 [1990], p. 30). Further parallels between Lyell’s Introduction and Darwin’s “Historical Sketch” in terms of content and strategy are suggested below.     

Against reduction

In Molecular Models: Philosophical Papers on Molecular Biology, Sahotra Sarkar presents a historical and philosophical analysis of four important themes in philosophy of science that have been influenced by discoveries in molecular biology. These are: reduction, function, information and directed mutation. I argue that there is an important difference between the cases of function and information and the more complex case of scientific reduction. In the former cases it makes sense to taxonomise important variations in scientific and philosophical usage of the terms “function” and “information”. However, the variety of usage of “reduction” across scientific disciplines (and across philosophy of science) makes such taxonomy inappropriate. Sarkar presents reduction as a set of facts about the world that science has discovered, but the facts in question are remarkably disparate; variously semantic, epistemic and ontological. I argue that the more natural conclusion of Sarkar’s analysis is eliminativism about reduction as a scientific concept.     

Morphological hemiplasies in cladistic studies of phylogeny,with examples from birds

Cladograms usually include many evolutionary reversions, parallelisms, and convergences united under the term homoplasy. Recently, it has become evident that molecular traits that look like homoplasy may be in fact true homologies. The processes of independent sorting of genes, which provide the basis of these events, were recently termed hemiplasy. The present study demonstrates theoretical possibility of independent manifestation of true homology (synapomorphy) in remote lineages of phylogenetic tree, as morphological characters are analyzed; this phenomenon is similar in manifestation (but not in nature) to hemiplasy. In the case of “morphological hemiplasy,” characters that appeared only once in evolution may formally be treated by a cladist as parallelisms, since they occur in remote lineages of phylogenetic tree. It is proposed that “morphological hemiplasy” is responsible for a number of uncertain cases in avian phylogeny. Examples are provided by the ducklike bill in Paleogene Presbyornithidae and extant Anatidae and Anseranatidae (Anseriformes), the apomorphic structure of the flying apparatus in Paleogene Jungornithidae (Apodiformes) and extant Trochilidae, and “independent” development of a highly-modified zygodactyl foot in Zygodactylidae and Pici.     

The PhyloCode is fatally flawed, and the “Linnaean” System can easily be fixed

Promoters of the PhyloCode have mounted an intensive and deceptive publicity campaign. At the centerpiece of this campaign have been slogans such as that the Linnaean System will “goof you up,” that the PhyloCode is the “greatest thing since sliced bread,” and that systematists are “afraid” to propose new names because of “downstream consequences.” Aside from such subscientific spin and sloganeering, proponents of the PhyloCode have offered nothing real to back up claims of greater stability for their new system. They have also misled many into believing that the PhyloCode is the only truly phylogenetic system. The confusion that has been fostered involves several discrete arguments, concerning: a new “method” of “designating” names, rank-free taxonomy, uninomial nomenclature, and issues of priority. Claims that the PhyloCode produces a more stable nomenclature are false, as shown with the example of “paleoherbs.” A rank-free system of naming requires an annotated reference tree for even the simplest exchanges of information. This would be confusing at best and would cripple our ability to teach, learn, and use taxonomic names in the field or in publications. We would be confronted by a mass of polynomial names, tied together only by a tree graphic, with no agreed name (except a uninomial, conveying no hierarchy) to use for any particular species. The separate issue of stability in reference to rules of priority and rank can be easily addressed within the current codes, by implementation of some simple changes, as we will propose in this article. Thus there is no need to “scrap” the current Linnaean codes for a poorly reasoned, logically inconsistent, and fatally flawed new code that will only bring chaos.     

“Critical size” in the yeastCandida utilis as influenced by physicochemical parameters of cultivation

The “critical size” of cells of budding yeasts was demonstrated to be influenced by physicochemical parameters of cultivation (temperature, aeration), type of substrate under conditions of nonlimited growth, “history” of the inoculum and its genealogical age, in addition to medium composition.     

The Extended Synthesis: Something Old, Something New

The eclipse of Darwinism began to end in the 1980s and hangs in the balance today. We need an Extended Synthesis, using “extension” metaphorically. We must extend back in time to recover important aspects of Darwinism that were set aside, and then lost during neo-Darwinism, then move forward beyond neo-Darwinism to encompass new data and concepts. The most comprehensive framework for the Extended Synthesis is the Major Transitions in Evolution. The Extended Synthesis rests comfortably within a philosophical perspective in which biology does not need to be connected with other areas of science in order to justify itself. I am attracted to an older concept in which biology needs a covering law to connect it with the rest of the natural sciences. Darwin implicated a “higher law,” but did not specify it. If we can elucidate that law, the Extended Synthesis will become the Unified Theory of Biology called for by Brooks and Wiley 25 years ago.     

The Origin of the Vertebrate Eye

In his considerations of “organs of extreme perfection,” Charles Darwin described the evidence that would be necessary to support the evolutionary origin of the eye, namely, demonstration of the existence of “numerous gradations” from the most primitive eye to the most perfect one, where each such tiny change had provided a survival advantage (however slight) to the organism possessing the subtly altered form. In this paper, we discuss evidence indicating that the vertebrate eye did indeed evolve through numerous subtle changes. The great majority of the gradual transitions that did occur have not been preserved to the present time, either in the fossil record or in extant species; yet clear evidence of their occurrence remains. We discuss the remarkable “eye” of the hagfish, which has features intermediate between a simple light detector and an image-forming camera-like eye and which may represent a step in the evolution of our eye that can now be studied by modern methods. We also describe the important clues to the evolutionary origin of the vertebrate eye that can be found by studying the embryological development of our own eye, by examining the molecular genetic record preserved in our own genes and in the genes of other vertebrates, and through consideration of the imperfections (or evolutionary “scars”) in the construction of our eye. Taking these findings together, it is possible to discuss in some detail how the vertebrate eye evolved.     

Gene sharing and genome evolution: networks in trees and trees in networks

Frequent lateral genetic transfer undermines the existence of a unique “tree of life” that relates all organisms. Vertical inheritance is nonetheless of vital interest in the study of microbial evolution, and knowing the “tree of cells” can yield insights into ecological continuity, the rates of change of different cellular characters, and the evolutionary plasticity of genomes. Notwithstanding within-species recombination, the relationships most frequently recovered from genomic data at shallow to moderate taxonomic depths are likely to reflect cellular inheritance. At the same time, it is clear that several types of ‘average signals’ from whole genomes can be highly misleading, and the existence of a central tendency must not be taken as prima facie evidence of vertical descent. Phylogenetic networks offer an attractive solution, since they can be formulated in ways that mitigate the misleading aspects of hybrid evolutionary signals in genomes. But the connections in a network typically show genetic relatedness without distinguishing between vertical and lateral inheritance of genetic material. The solution may lie in a compromise between strict tree-thinking and network paradigms: build a phylogenetic network, but identify the set of connections in the network that are potentially due to vertical descent. Even if a single tree cannot be unambiguously identified, choosing a subnetwork of putative vertical connections can still lead to drastic reductions in the set of candidate vertical hypotheses.     

Darwin on Aristotle

Charles Darwin's famous 1882 letter, in response to a gift by his friend, William Ogle of Ogle's recent translation of Aristotle's Parts of Animals, in which Darwin remarks that his “two gods,” Linnaeus and Cuvier, were “mere school-boys to old Aristotle,” has been thought to be only an extravagantly worded gesture of politeness. However, a close examination of this and other Darwin letters, and of references to Aristotle in Darwin's earlier work, shows that the famous letter was written several weeks after a first, polite letter of thanks, and was carefully formulated and literally meant. Indeed, it reflected an authentic, and substantial, increase in Darwin's already high respect for Aristotle, as a result of a careful reading both of Ogle's Introduction and of more or less the portion of Ogle's translation which Darwin says he has read. Aristotle's promotion to the pantheon, as an examination of the basis for Darwin's admiration of Linnaeus and Cuvier suggests, was most likely the result specifically of Darwin's late discovery that the man he already knew as “one of the greatest ... observers that ever lived” (1879) was also the ancient equivalent both of the great modern systematist and of the great modern advocate of comparative functional explanation. It may also have reflected some real insight on Darwin's part into the teleological aspect of Aristotle's thought, indeed more insight than Ogle himself had achieved, as a portion of their correspondence reveals. This revised version was published online in July 2006 with corrections to the Cover Date.     

On Darwin's 'metaphysical notebooks'. I: teleology and the project of a theory.

Huxley's essay On the Reception of the 'Origin of Species' brings us close to the issue of cause and of why- and how-questions in the understanding of the living world. The present contribution, which is divided into two parts, reviews the problem of Teleology as conceived by Huxley and re-examines Darwin as the author who revealed the existence of a 'foundations problem' in the explanation of an entire realm of nature, i.e., the problem of explaining such realm in terms of its own, specific legality, or iuxta sua propria principia. In the first part the enquiry is mainly focused on the secularization of natural history after Paley; in the second part it is mainly focused on the desubjectivization of the inquiry into natural history after Erasmus Darwin and Lamarck. The second part will be published in the next issue of Rivista di Biologia/Biology Forum. In the first part below an analysis is made of Notebooks M and N. The author disputes the correctness of conceiving them only as the works where Darwin envisages the 'metaphysical' themes later to become the subject of The Expression of the Emotions. He suggests to conceive of them also as the works where Darwin defines the terms of the general project of his own, peculiar evolutionary theory. The author then outlines the intellectual progress of Darwin from the inosculation to the transmutation hypotheses. Darwin's reading of Malthus appears to be analytically decisive, because it offers him the vintage point to attack the metaphysical and theological citadels on the morphological side. Darwin is thus able to re-consider Erasmus' comprehensive zoonomic project, by displacing it, however, from the old idea of the scala naturae to the new one of the "coral of life", and by emphasising the distinction between "the fittest" and "the best" vs. the tradition of Natural Theology.