Abstract
Throughout the Victorian era, the metaphor “living fossil” repeatedly crisscrossed social and scientific domains. The term existed in popular culture before and after Darwin’s Origin. Most notably, it also operated as two distinct scientific concepts, one introduced by Darwin and another in cultural evolutionists’ depiction of human living fossils. Serving in different ways, living fossils were typically aberrant, persistent and unchanging examples that contradicted an expectation of ongoing change and associated progress. We explore the development and relationships of living fossil applications, focusing principally on Darwin’s concept. In Origin, Darwin deployed living fossils as exceptions that prove the rule of his principles of natural selection and divergence. He structured a case for the causal adequacy of these principles to explain living fossils’ persistence, invariance, and taxonomic positions in gaps between other groups. As other natural historians began discussing living fossils and labeling new ones, Darwin’s concept endured, but was subject to perceivable variation; associations with natural selection or divergence varied greatly and attributes of his living fossil examples were sometimes ignored. Cultural evolutionists adopted a view that human societies developed over time in a unilinear succession of stages. In this view primitive groups, their implements, languages, and cultures, stopped evolving at different points in the past and persisted unchanged into the present. While Darwin’s concept and this anthropological concept were connected associatively to the evolution of languages and to themes of spatial isolation, prolonged stasis and disruption of expected progress, they inherited significantly different theoretical backgrounds and commitments.
Keywords: Concepts, Cultural evolution, Darwin, Living fossil, Metaphors
Hence we can see why all the species in the same region do at last, if we look to wide enough intervals of time, become modified; for those which do not change will become extinct.
Darwin (1859, p. 315)
Introduction
A scientific concept of living fossils has somehow survived for over 150 years, all the while varying and expanding its meanings (Lidgard and Kitchen 2023). Its use in present-day scientific publications is flourishing despite heated criticism that has led some critics to call for its abandonment (Casane and Laurenti 2013; Lidgard and Love 2018). The term living fossil circulates readily in modern popular parlance, journalism and literature, taking on ancillary meanings and hinting not at a corruption of an original idea, but rather the kinds of common imaginaries about time and directionality that also captivated a public in the 19th century Britain of Charles Darwin.
The Victorian era was a time infused with the idea of progress. However, a belief that progress was an enduring natural phenomenon had developed earlier, among philosophers and writers of the Enlightenment. This idea strengthened through the 18th century amid a complex societal mélange of political transformations, economic improvements, and the rising prominence of science and reason (Mokyr 2014; Tam and Meek Lange 2024). The Victorians saw progressive development in technological discoveries, political reforms, commercial prosperity and print culture (Claeys 2005; Frawley 2017). Darwin’s private and public writings from the 1830s onward show a commitment to changing conditions over geologic time and progress in the history of life (Hodge 2009, 2013a). Many of his contemporaries held similar views (Watson 1845; Ursillo 2023). Cultural evolutionists in the nascent discipline of anthropology adopted yet another view of progress: beginning in prehistory, all societies progress through universal stages of development, though some slow down and a few, as human living fossils, cease to advance and continue unaffected into the present. Here we consider how “living fossil” and some of its implied meanings—being neither extinct nor significantly changed—fit into this contrary flow of ideas about progress.
Understanding how Darwin developed and deployed living fossils is important because of the way it augments and coincides with our current knowledge of his theorizing. Much scholarship has been devoted to the arguments Darwin had structured by 1859 with the publication of On the Origin of Species, yet few studies have centered on his use of “living fossil” as a scientific concept.1 In Origin, Darwin exploited living fossil both as a metaphor and a scientific concept roughly akin to an exception that proves the rule of his two major theses, natural selection and the principle of divergence. It was one among many metaphors that played vital roles in Origin, both in clarifying ideas and creating “new, literal, meanings” (White et al. 2021, p. 175; see also Beer 2009). We emphasize how living fossils offered Darwin and other researchers a way to theorize about stasis and isolation against a backdrop of predictable change and progress, just as degeneration and atavism drew public attention for their defiance of progressive expectations. The phrase “living fossil” existed in the public sphere prior to 1859, crisscrossing between scientific and public domains (Carus 1818; Bird 1839; Dalton 1853). It was a metaphor occurring in literature, in colloquial expressions connoting something or someone being out of date, and in the strange phenomenon of living animals entombed in sediment or stone. Taking seriously Darwin’s framing of his scientific living fossil concept allows us to assess to what degree he gave voice to things already in the Zeitgeist (Radick 2009), or determined something precise in his own created theories. Darwin’s living fossil concept and the cultural evolutionists’ concept of human living fossils bear only partial similarity to the broader use of the metaphor outside of scientific publications. However, societal appearances of living fossil sometimes reveal connections to various prerequisites and characteristics of these two distinct scientific concepts.
What metaphors and concepts are and the ways they are used and understood entail an immense range of possible positions (Feest and Steinle 2012; Semino and Demjén 2017; Margolis and Laurence 2019; Garello 2024). Language is metaphorical, and metaphors are inescapable elements of science’s conceptual system (Reynolds 2022). Consequently, how metaphors and concepts figure in historical roles that traverse both scientific disciplinary boundaries and boundaries between scientific and social discourse touches not just historians, but also scholars in philosophy, linguistics, discourse and communication, cognitive psychology, and various scientific disciplines. Furthermore, metaphors sometimes serve as scientific concepts (Knudsen 2015), and concepts can be metaphorical; living fossil is an example. We therefore need to fashion a distinction for this study about domains of use. The term living fossil can be thought of as a container or vessel, its contents varying in accord with their uses in different domains (cf. Kindi 2012). Where living fossil pertains to Darwin, natural historians or cultural evolutionists in their respective scientific domains, we use the phrase scientific concept or simply concept. Where it pertains to speech or literature in the social domain, we use the phrase popular metaphor. Intriguingly, these varied uses and domains intertwined through much of the 19th century, consistent with what historian Robert Young described as a common context of biological, social and philosophical ideas, few parts of which were fully isolated (Young 1969, 1973; see also Dawson et al. 2004).
What connects these applications of the living fossil label? What do instances of imperfect similarities to Darwin’s concept suggest about the world in which he formalized his thinking? And what may have rendered other organismal living fossils different from human ones? Darwin’s scientific work must be read in relation to contemporary concerns of 19th century empire, politics, literature, and knowledge flow (Browne 1996, 2002; Sera-Shriar 2018). We show that while these two scientific concepts intersected in meaningful ways, they inherited different intellectual lineages, commitments, and consequences. Two perspectives emerge from our analysis: Darwin’s organismal living fossil concept with its ties to ecological and evolutionary processes, and the anthropological concept of static living fossil human populations with its ties to racial and ethnic hierarchies and Enlightenment stadial theories. The popular metaphor drew from both wells, with associations at the broadest level between, for example, anomaly and ancientness, and spatial isolation and discovery. Understanding how varied uses of living fossil in popular discourse and in distinct scientific living fossil concepts relate to the social and geopolitical world furthers discussion about the ways exceptions to expected trends, such as progress and extinction, were handled in science and broader culture.
Our approach embraces narrative structuring and interpretation (cf. Carr 2008; Lorenz et al. 2021), contextually grounded in historical actors, texts, discourses, metaphors, and scientific concepts, rather than other approaches that accentuate linguistics, Begriffsgeschichte (conceptual history), or a conceptual metaphor framework (Koselleck 2002; Kuukkanen 2008; Ifversen 2011; Anderson 2017; Garello 2024). The intended result is not to renounce these other approaches and the value of historical categories or semantic representations, but to better approximate the similarities and differences of what living fossil allowed diverse 19th century actors to theorize, defend, and suggest in both scientific and popular spaces. Numerous examples show scientific concepts changing over time, such as evolution (Bowler 1975), organism (Cheung 2006; Wolfe 2014), and gene (Griffiths 2002; Brigandt 2010). Metaphors too can undergo change, sometimes moving back and forth between scientific and social domains (Musolff 2014; Surman et al. 2014). We interpret how scientific concepts in different spaces accommodate change or are separated, and how they interact with uses in the public domain—here in the form of metaphors (Bal 2002; Surman et al. 2014; Arabatzis and Nersessian 2015). In comparing Darwin’s and cultural evolutionists’ scientific living fossil concepts, our approach focuses on disciplinary contexts and inherited backgrounds, and how these actors used concepts purposefully to meet different epistemic aims and perform inferential and explanatory roles (Brigandt 2012, 2020; Arabatzis 2019).2
We first examine living fossil as a popular metaphor that may have been known to Darwin during his evolutionary theorizing. We then consider shared knowledge and assumptions in scientific disciplines, as Darwin’s living fossil concept and the cultural evolutionists’ concept (to some extent) rely on them to contextualize and derive their meanings. We trace changes in Darwin’s ideas from the 1830s onward leading to his mature living fossil concept in the late 1850s and in Origin. Following on in the late 19th century with other naturalists’ variations on Darwin’s concept, we consider the varied guises of living fossils in literature, expeditions to find living relics, and, most acutely, theorizing in historical linguistics and Victorian anthropology that framed cultural evolutionists’ concept of human living fossils. In the final section, we position different applications of living fossil—together with their entangled characteristics—at the end of the 19th and the beginning of the 20th century. We see that the functions of living fossils in particular readings of evolutionism varied with how change and lack of change were perceived.
Living Fossil as a Popular Metaphor
It was only in the early 19th century that our current understanding of the word fossil became commonplace among natural historians (Forli and Guerrini 2022). Among the public at large, an older meaning of fossil as any noteworthy object dug out of the ground also lingered on through the century (Anonymous 1855). This older meaning acted as a metaphor in the story of “Merry the Miner,” a veritable living fossil trapped underground in a cave (Bird 1839). In another story, an old soldier recalls, “[w]e were covered from head to foot by a thick coating of filth, which grew hard, and converted us into living fossils” (Jonnes 1858, p. 180).
Far more frequently, living fossil acted as a metaphor that conveyed a prolonged continuity of stasis, irrespective of scientific, literary or social context. Living fossil was used—albeit sparingly—as early as 1712 by the Welsh naturalist Edward Lloyd (Lhwyd) in a Royal Society publication that referenced “the account of living fossil mussels” (Lhwyd 1712, p. 506). Another use is found in episodes of living frogs and toads miraculously wedged into stony encasements or large tree trunks (Bondeson 1999). This use is fairly literal in the sense of the older meaning of fossil, but becomes more metaphorical in the modern sense: paleontological fossils are of course dead, leaving only traces or preserved parts. In 1818, the prominent physiologist, naturalist, and philosopher Carl Gustav Carus wrote on the possibility of these “lebende Fossilien”—living creatures buried in stone for thousands of years—and suggested that such creatures had been asleep and buried, safe from the revolutions of the world above that had killed other plants and animals (Carus 1818, p. 143). The paleontologist William Buckland put the matter to an experimental test in 1825. He buried twelve toads sealed in glass underground and waited over a year (Buckland 1833). The toads died, and to his satisfaction the grim results dispelled any possibility that prior living fossil toads had survived in a completely enclosed stony tomb. According to Buckland, the toads in earlier reports likely found a small cavity as tadpoles, grew too large to leave, but survived via tiny cracks in the stone, enabling insects and air to make their way to them (Gordon 1894). Newspaper and magazine reports of the phenomenon of literal living fossils endured nonetheless: a live snake found in a block of coal (Anonymous 1817), more live toads in gravel sediments being dug out of the Erie Canal (Hinton 1832), even a live pterodactyl excavated in France (Presse grayloise 1856), which upon exhumation was said to have shaken its wings before “uttering a hoarse cry” and dying on the spot—though that one proved to be a hoax (Meagher 1856, p. 37).3 Toads in holes made the rounds in literature as well, with a living fossil toad appearing amongst a litany of curios in Alexandre Dumas’ Les Mohicans de Paris (1857). At the 1862 Great Exhibition of London, a lump of coal and a frog were displayed together, with an announcement that the frog had been found alive inside. William Buckland’s son Frank lambasted the Exhibition’s display of what was, in his view, clearly a hoax (Buckland 1862). William Darwin alerted his father of the matter, and Charles thought the possibility of any living frog being found in a completely closed space was quite incredible, recalling William Buckland’s experiment.4
Beyond the toads, living fossil as a popular metaphor loosely related to stasis took the form of scholars who became immersed in ancient history (Anonymous 1854), or who personified the past: “he was a living fossil of the age of Aristotle and Plato” (Winslow 1856, p. lxvi). Living fossils were workers who refused to adopt modern methods (Dalton 1853), church parsons who could not give up the old ways (Bainbridge 1858), or persons who were simply out of touch with the present: “We hope you have kept a journal detailing your observations of the ways and customs of living fossils. Have you discovered to what species of defunct animal Mr Hilton belongs—Megatherium or Dinotherium?” (Narcot and Narcot 1875, p. 204). Anomalous living fossil forms, those that did not fit neatly into taxonomic placements, captivated the imagination in fictional literature (Krzeminski 2021; Fallon 2024). A living fossil in these contexts was evocative of something being an irruption in time, characterized by its uncanny time-delayed manifestation, the way fossils themselves broadly function in literary fiction (Doane 2020). More common than specific living fossil animals in the public imagination were degenerate forms or atavistic returns from a previous era that both functioned as disruptors of an expectation of progress (Hurley 1996; Wagner 2020).
Darwin’s own use of living fossil is evocative. To fully understand his living fossil concept, however, more than a reading of stagnation against progress is necessary. We next examine the shared knowledge and assumptions in disciplines we now know as geology, paleontology, and taxonomy that were current when Darwin conducted the evolutionary theorizing leading up to Origin.
Shared Knowledge and Assumptions
Whether living fossil serves as a scientific concept or as a metaphor in common speech or prose, the phrase is fundamentally comparative and contextual. It only makes sense when the reader is aware of related things whose properties, by contrast, have significantly different values. Several prerequisites were important for Darwin’s living fossil concept. Some of these elements later informed other scientists’ variations on his concept, and likely also anthropologists’ sense of time and tempo of change in human prehistory. In the 19th century scientific domain, both a living fossil and its comparative set were the same sorts of entities (organisms, species, or other taxa for Darwin and other natural historians; human groups and relic particularities of their languages or cultures for cultural evolutionists). All these entities were compared together in the context of properties such as geologic age and duration, morphological likeness, taxonomic (or explicitly genealogical) relationships, linguistic relationships, cultural artifacts and so on. Living fossil as a metaphor in the public domain operated similarly, but did not draw on the same swathe of expert knowledge. It differed markedly in the scale, substance or number of properties being considered. For instance, labelling a politician a living fossil compared to other politicians or normal citizens may only refer to the politician’s unchanging attitudes amid progressive change in social circumstances of years or decades. Here, time is only crudely analogous to geologic time or stratigraphy, and attitudes are unlikely referents of paleontological properties.
A vast expanse of geologic time was one prerequisite for Darwin’s and cultural evolutionists’ mechanisms to function (Manias 2015; Dear 2016). By the early 19th century, many leading naturalists accepted a deep time history of the earth well beyond the literal Biblical chronology (Rudwick 2005). James Hutton’s uniformitarianism, the idea that present-day processes have operated similarly throughout the geologic past, was expanded by Charles Lyell and helped lay the foundations of Darwin’s evolutionary theorizing (Virgili 2007; Rudwick 2008). Georges Cuvier had made extinction of taxonomic lineages in the geologic past plausible, though some naturalists still maintained the continued existence of fossil forms in remote places. A mere five years before Charles Darwin was born, Thomas Jefferson sent Meriwether Lewis and William Clark on their Corps of Discovery Expedition with a request to look for a living form of what he called a mammoth and the giant ground sloth Megalonyx (Barrow 2009; Rowland 2009). With the encroachment of settlers further into the American interior, Jefferson could imagine that mammoths now existed in a restricted range: “Those parts still remain in their aboriginal state, unexplored and undisturbed by us, or by others for us. He [the mammoth] may well exist there as he did formerly where we find his bones” (Jefferson 1787, p. 54).5 Long spans of geologic time allowed extinctions to gradually eliminate lineages genealogically intermediate between the aberrant forms of Darwin’s living fossils and taxonomically adjacent groups that had not only survived, but had likely maintained or even increased their diversity. For cultural evolutionists, a long time interval of human prehistory complemented inferences of hindered or frozen rates of progress among globally distant peoples (Manias 2015).
Resolving the order and spatial correlation of geologic strata and the construction of a geological timescale were two more prerequisites. They permitted direct comparison of the relative ages and durations of taxonomic groups (Şengör 2016). British and continental European geologists documented sequences of strata from local outcrops to regional scales, establishing tentative global stratigraphic divisions by 1840 (Burchfield 1998). Similarly, by the 1830s, antiquarians and archeologists had begun to adopt a three-age system of periodization for human prehistory (Stocking 1991; Reybrouck 2012).
Advancement in the theory and method of classifying organisms was yet another prerequisite. The simplicity of the Linnaean classification system fostered its widespread use, yet as Linnaeus himself admitted, it was an artificial system. As late 18th century naturalists attempted to classify vastly different groups of organisms, they were perplexed by conflicting distributions of characters and implausible relationships (Rieppel 2010; Jenner 2022). Antoine-Laurent de Jussieu and others developed versions of an alternative method, the natural system, based on numerous diverse characters and bottom-up grouping of taxa into successive categories (Stevens 1994; Jenner 2022). By the 1820s, many competitors claimed theirs as the natural system (Endersby 2010). Among these, the quinarian system introduced by William Sharp Macleay became influential in early Victorian Britain (Macleay 1819). Its underlying tenets were developed further in the form of Darwin’s “lineage thinking” and evolutionary arguments (Winsor 2013; Novick 2019; Jenner 2022, pp. 50–58). Meanwhile, plant and animal taxa recorded in collections and catalogues, together with proposed higher-level taxonomies of living and fossil groups, established a comparative groundwork (Nicol 1849; Müller-Wille 2017). This groundwork enabled visual contrasts for different groups of organisms as graphs of geologic durations and relative changes in diversity over geologic time, and as trees of life (Archibald 2014; Sepkoski 2018; Sepkoski and Tamborini 2018).6
Notions of species transformism and a paleontological “succession of types” were prerequisites for some species concepts and for fostering an expectation of progressive development that would be set against Darwin’s living fossil concept (Darwin 1839, pp. 210–212; Tanghe 2017).7 Transformist ideas were being discussed in different European states by the early 1800s (Richards 2002a; Sloan 2019a; Corsi 2024). The anonymously published Vestiges of the Natural History of Creation brought an evolutionary scheme into wide public view (Chambers 1844). In 1852, Herbert Spencer anonymously authored “the development hypothesis,” which critiqued special creation and favored species transmutation (Spencer 1891). Despite lingering ambiguity about the respective influence of various forebearers on Darwin, the not infrequent perspective—and conceivably the expectation—of species transformism over geologic time was ever more evident (Richards and Ruse 2016; Tanghe 2017; Sloan 2019b; Douglas 2022). Darwin’s own acknowledgement of transmutationist forerunners, although decidedly wanting, finally appeared in Origin’s third edition and was expanded in the fourth edition (Darwin 1861, 1866; Corsi 2024).
Darwin’s Living Fossils as a Scientific Concept
Darwin used living fossils as exceptions that prove the rule of his principles of natural selection and divergence. How and why he arrived at that usage gives us a fuller picture of the ways his theorizing developed and by what means he structured arguments for his theory. He depicted living fossils in chapter four of Origin:
All fresh-water basins, taken together, make a small area compared with that of the sea or of the land; and, consequently, the competition between fresh-water productions will have been less severe than elsewhere; new forms will have been more slowly formed, and old forms more slowly exterminated. And it is in fresh water that we find seven genera of Ganoid fishes, remnants of a once preponderant order: and in fresh water we find some of the most anomalous forms now known in the world, as the Ornithorhynchus and Lepidosiren, which, like fossils, connect to a certain extent orders now widely separated in the natural scale. These anomalous forms may almost be called living fossils; they have endured to the present day, from having inhabited a confined area, and from having thus been exposed to less severe competition. (Darwin 1859, p. 107)
As we shall see, this is neither a complete nor an unambiguous definition. Here, we attempt to unravel the several elements linked to his concept and the attributes of his living fossil examples by also analyzing his surviving manuscripts, notes, and correspondence. His concept developed gradually alongside concerns about classification and what would become his keystone concepts of divergence and natural selection (Kohn 2009; Winsor 2013; Novick 2019; Love 2024). We situate indications of his thinking about living fossils and important changes in these other concepts in the historiography of two critical periods, from the end of the Beagle voyage to the mid-1840s, and from the early 1850s to 1859. We then consider how Darwin’s living fossil concept is contextualized by other concepts in Origin. The concept is also framed in part by the relevant attributes of taxa that he named explicitly as living fossils. We also reflect on the enduring nature of his living fossil concept after 1859, involving transformations of the concept and its application to taxa known to Darwin yet not designated by him as living fossils.
Classifying and Theorizing
When Darwin’s voyage on the Beagle ended in the fall of 1836, most British naturalists shared an image of nature as progressive (Bowler 1976). The fossil record provided empirical evidence of a rise in complexity from the deep past to the present, a pattern reminiscent of an older concept of the Chain of Being. Yet both zoological and botanical classifications, including quinarian ones, were largely atemporal. While some zoological classifications acknowledged the idea of a progressive scale among living organisms and related fossils, they did not necessarily imply evolution (Ursillo 2023). Evidence of these patterns in earth’s history seemed incontrovertible, yet as historian Peter Bowler noted, explanations differed from repeated divine interventions of separate creations to the “possibility that a process of diversification in time might explain why species fall into natural groups” (Bowler 2021, p. 178). Taxonomic analysis of Darwin’s Beagle collections of South American fossil mammal bones and Galapagos land birds by Richard Owen and John Gould indicated the likelihood of a succession of taxa of the same type in the same country, and of species transmutation (Ospovat 1981; Hodge 2009, 2013a).
By July 1837, Darwin had begun work on his transmutation notebooks. Over the next several years, he filled these notebooks with his early evolutionary theorizing, sifting through patterns of classification of organisms and temporal and geographic patterns of diversity that he sought to explain by some causal mechanism(s). From the outset he explored species propagation, variation and adaptation in relation to progress in the history of life. He allowed that rates of change among groups could vary and that certain lower species might not change at all if the conditions of their habitats remained constant (Hodge 2009). Species propagations, extinctions and gaps between taxa were imagined and revised in iterations of branching diagrams, embracing a tree of life as both metaphor and concept (Putten 2020). Darwin wrote of his first diagram in 1837: “The tree of life should perhaps be called the coral of life, base of branches dead, so that passages cannot be seen.”8 A few pages later, below a tree diagram he had sketched with labeled clusters of branches, he noted the gaps between clusters, also writing, “[t]hus genera would be formed.—bearing relation to ancient types.”9 Those passages would come to represent morphological and taxonomic gaps between clusters of species in surviving groups of living taxa. Darwin considered and rejected several evolutionary theories in the notebooks. None closely approached the initial version of natural selection that appeared after reading Malthus in late 1838, and that Darwin expounded later in his private 1842 Sketch and 1844 Essay (Darwin 1909; Kohn 1980).
Darwin’s transmutation notebooks also illuminate his intense engagement with classification of living and fossil organisms. Classification was both a basic tool and a goal of Victorian naturalists, and a vast community of hobbyists and scientists was naming and classifying organisms at a frenzied pace (Endersby 2010; Winsor 2013). The quinarian system developed by William Sharp Macleay, and revised in a distinct but related form by William Swainson, was often discussed by Victorian naturalists in the 1820s and 1830s (Macleay 1819; Novick 2016; Mursinna 2023). Macleay’s system reinforced the view that classification could uncover real groups of species and their real relationships rather than artificial constructs (and species nominalism). Richard Owen, Thomas Henry Huxley and Joseph Hooker all briefly expressed their interest, and Robert Chambers described the system as “a powerful additional proof of the hypothesis of organic progress by virtue of law” (Chambers 1844, p. 250). Macleay was committed to numerical symmetry and continuity within and between groups in organic nature (Novick 2016; Jenner 2022; Mursinna 2023). Taxa at a given hierarchical level of classification formed five circles of relationships, termed affinities. In many pre-evolutionary natural classifications, affinities were understood as a sort of essential similarity, an idea that later in the 19th century would be compared to homology (O’Hara 1991; Panchen 1994; Stevens 1994). In each circle, individual taxa were linked in a circular chain or series of affinity from one to the next, based on similarities among the majority of their characters. The circles represented taxonomic groups, each with parallel series, but clearly separated from other groups. At any one level the gap between each pair of adjacent circles was often bridged by osculating (kissing) taxa, aberrant in form but sharing a few characters that could indicate a degree of affinity with each adjacent circle (Ospovat 1981; Novick 2019). Osculant taxa would also have few species, an assertion soon confirmed by others. When no appropriate taxa existed to complete the chain of affinity within a circle or to show osculating links between circles, Macleay and other quinarians used placeholders and assumed that such forms would eventually be found (Jenner 2022). Macleay also recognized secondary relationships formed by analogies, insulated parts or characters that seemed to imply a similarity between different organisms or taxonomic groups that are otherwise quite different in the majority of their characters. But by the mid-1840s, analogies as useful characters and the supposed reality of numerical symmetries in natural classification were effectively decimated by the criticisms of ornithologist and systematist Hugh Strickland and others (Strickland 1840; Mursinna 2023).
Darwin had encountered Macleay’s work while at Cambridge and engaged at length with affinities, analogies, and osculant groups in the transmutation notebooks (Ospovat 1981; Di Gregorio 1996; Novick 2016). Darwin gathered just as much from Macleay’s underlying quinarian principles as he did from the particulars of quinarian patterns (Novick 2019). Principles such as continuity, a primary role for characters of affinity, and assessing variation and inferring relationships in the context of the entire organism were significant as Darwin moved toward temporalizing the natural system. Macleay’s reinforcement of previously recognized patterns in classifications was also important: distinct taxonomic groups were separated by gaps, and these gaps were wider at higher levels of classification (Winsor 2023).
As Darwin elaborated his tree of life concept through 1837 into 1838, he considered possible explanations for patterns of taxonomic discordance and diversity. A scant few pages after his inaugural sketch of a zoonomical system in Notebook B, he wrote, “?How is it that there come aberrant species in each genus (with well characterized parts belonging to each) approaching another.”10 Historian Aaron Novick described how Darwin sought to explain Macleay’s account of character variation among taxa, “Darwin argued that the oldest characters tend to vary least, and thus that the amount of variation in a part can be a guide to genealogical relationships” (Novick 2019, p. 266). Perhaps also inspired by quinarian views of continuity and the expectation that gaps ought to be filled, he questioned the nature of gaps in hierarchical groups: “the greater the groups the greater the gaps (or solutions of continuous structure) between them.—for instance, there would be great gap between birds and mammalia, Still greater between Vertebrate and Articulata, still greater between animals & Plants. But yet, besides affinities from three elements, from the infinite variations, & all coming from one stock & obeying one law, they may approach,—some birds may approach animals, & some of the vertebrates invertebrates.”11 He hypothesized that the varied taxonomic breadth of different genera involved some species adapting to changing circumstances by propagating with different rates of irregular branching, and other groups going extinct. Darwin pondered the expectation of intermediate steps between groups, perhaps still undiscovered, and wrote a first reference to the platypus Ornithorhynchus. He wrote that, “[w]e have not the slightest right to say there never was common progenitor to mammalia & fish. when there now exists such strange forms as ornithorhync[h]us.”12 Later in Notebook B, he referred to the pattern that aberrant and osculant groups have few species but nonetheless sometimes do form genera associated with gaps.13 The place of the platypus in the order of nature had been debated since the end of the 18th century. Naturalists puzzled over its seemingly conflicting characters: the beak and webbed feet of a duck, the fur and mammary glands of a mammal, and both the presence of a cloaca and the possibility that it laid eggs like a reptile or bird (Shaw 1799; Owen 1832). There was no fossil record to help with classification and no definitive proof of egg-laying until the late 19th century (Hall 1999; Rieppel 2023).
Darwin likely first read Macleay’s Horae Entomologicae (1819) in 1838 and first referred to him in Notebook C (Novick 2019). There Darwin presented a view of osculant groups whose temporal dimension profoundly transformed the quinarian systems of Macleay and his followers: “there can be no animal at present time having an intermediate affinity between two classes.—there may be some descendant of some intermediate link.—the only connection between two such classes will be those of analogy, which when sufficiently multiplied become affinity yet often retaining a family likeness, & this I believe the case.—Any animal really connecting the fish & mammalia must be sprung from some source anterior to giving off of these two families, but we see analogies between fish.”14 A concept of ongoing extinction, disrupting continuity and expected diversification and transformation, was critically important. The next few pages of Notebook C offer inferences that genera can be formed with the extinction of the ancestral species, that intermediate species also disappear in the process, and that an osculant genus may sometimes survive. In this confirmation of the roles of descent and extinction, temporalizing the natural system is an implicit idea (Novick 2019). In Notebook E, Darwin noted the confined habitat of Ornithorhynchus and the aberrant nature of Lepidosiren (lungfish).15 In early 1839, Owen described Lepidosiren annectans at a meeting of the Royal Society (Owen 1841). The complex character distribution of Lepidosiren and its relationships to both fishes and reptiles (including amphibians) was hotly debated (Milne Edwards 1841). Uncertainty of its position in classifications and the problem of evolution of different parts of organisms at different rates (mosaic evolution) continued for decades, even after recognition of lungfish fossil remains in the 1870s (Rieppel 2023).
In the 1840 essay “Observations upon the Affinities and Analogies of Organized Beings,” Hugh Strickland acknowledged the possibility of continuity evidenced by affinity at lower taxonomic ranks, but not at higher ranks such as classes. He wrote: “But if birds have a tendency to unite with mammals by means of Ornithorynchus, and Vertebrata with Molluska by means of Myxine [hagfish], then this approximation is not to be considered as a distinct principle, but only as an undetermined analogy or affinity” (Strickland 1840, p. 226). What is missing in this stance? Darwin’s appeal to different scales of deep time versus ordinary, experienced time provides a partial answer. As historian Peter Dear noted, “the creation of the categories by the coming-into-being of the kinds of organisms that instantiated them was always something that Darwin regarded, conceptually, as having occurred prior to an organism’s taxonomic placement” (Dear 2016, p. 7). Darwin had used a “thought experimental” form of reasoning throughout the notebooks, as James Lennox noted, “not as a method of defense for his theory, but as a means of exploring the ability of his developing ideas to explain facts that typically pose problems for transformist views of species origins” (Lennox 2008, p. 71). Darwin’s hypothesis allowed that gradually over vast amounts of time with species propagation, aberrant (osculant) groups would appear to have deviated from surviving, yet distantly related groups, with gaps due to past extinctions of connecting forms. In the section entitled “On the Evidence from Geology” in the Sketch of 1842, he hypothesized a very gradual introduction of a long series of new forms from a common ancestor, along with extermination of old forms. Even with an initial “perfect gradation” of forms, preservation and discovery of all the intermediates would be a near impossibility: “... it is certain all the numerous fossil forms fall in(to), as Buckland remarks, not present classes, families and genera, they fall between them: so is it with new discoveries of existing forms. Most ancient fossils, that is most separated (by) space of time, are most apt to fall between the classes—(but organisms from those countries most separated by space also fall between the classes (e.g) Ornithorhync[h]us?)” (Darwin 1909, p. 24).
A note dated April 1843 likely contains Darwin’s first written use of the phrase living fossil. He wrote: “Aberrant groups Hensleigh remarks that groups are aberrant because they are few in number—it is doubtful how far this holds with genera—I look at aberrant groups as ‘living fossils’ [...] If aberrant genera have few species proof of extinction for a genus wd not be less aberrant whether or not many species. as Ornithorynchus & Echidna.—Apteryx.”16 In 1844, he discussed with a few colleagues contrasts between aberrant groups like Ornithorhynchus with isolated distribution and typical genera that are distributed more widely, and their contrasting potentials for adaptation.17 Ultimately, a section of the 1844 Essay provides what Darwin called a genealogical explanation for resulting patterns in classifications of gaps and aberrant groups:
It follows, from our theory, that two orders must have descended from one common stock at an immensely remote epoch; and we can perceive when a species in either order, or in both, shows some affinity to the other order, why the affinity is usually generic and not particular [...] And generally, it may be observed in the writings of most naturalists, that when an organism is described as intermediate between two great groups, its relations are not to particular species of either group, but to both groups, as wholes. A little reflection will show how exceptions (as that of the Lepidosiren, a fish closely related to particular reptiles) might occur, namely from a few descendants of those species, which at a very early period branched out from a common parent-stock and so formed the two orders or groups, having survived, in nearly their original state, to the present time. (Darwin 1909, p. 212)
While Darwin returned to these matters intermittently during the decade or so after 1844, much of his time was given over to writing treatises on his geological observations and on living and fossil barnacles. His living fossil concept had not yet reached the fuller, mature sense reflected in his later notes and correspondence and in Origin.
Two Keystones
Changes in Darwin’s thinking about living fossils in the 1850s were intertwined with changes in his views on variation and classification, adaptation and natural selection, and branching and divergence (Browne 1980; Ospovat 1981; Kohn 2009; Love 2024). Darwin maintained in both the 1842 Sketch and the 1844 Essay that there is little variation among organisms, or that it is uncommon in the wild. Both variation and transmutation were then associated primarily with changes in physical conditions, as opposed to interrelations with other organisms (Ospovat 1981; Partridge 2018). The tendency for new species to diverge from a parent form lacked a clear explanation. Natural selection was not yet a process that operated constantly on abundant variation. Adaptation was clearly a part of his early consideration of species formation (Hodge 2009). However, it was not until the late 1850s in his revised concept of natural selection that the idea of relative adaptation in response to interactions with other organisms surpassed perfect adaptation in response to changing physical conditions (Ospovat 1981; Kampourakis 2023; Love 2024; but see also Hodge 2013a, 2013b).
The principle of divergence developed out of work on classification. Copious published morphological comparisons and classifications revealed a hierarchy of taxonomic clusters which could be of radically different sizes and geographic spreads. Even with the addition of fossils, gaps were present between clusters where one would expect intermediate forms showing gradual transitions, or might be occupied only by a species-poor aberrant group that shared a few affinities each with adjacent clusters. Darwin had been accumulating evidence toward explanations for these patterns occasionally since the 1840s. Communications with fellow naturalist George Waterhouse considered Richard Owen’s morphological work on Lepidosiren, Ornithorhynchus, and their intermediate links to larger taxonomic groups.18 Darwin likewise mused about the characters and intermediate status of ganoid fishes.19 He began considering information about the rarity and geographic ranges of intermediate versus typical forms at various taxonomic levels.20
Perhaps most importantly, Darwin drew from Owen’s concept of the archetype. Owen’s archetype was a transcendental model that united the members of a large taxonomic group (Sloan 2019a). Darwin reinterpreted Owen, famously writing on the back cover of his copy of Owen’s On the Nature of Limbs (1849), “I look at Owens Archetypus as more than ideal, as a real representation as far as the most consummate skill & loftiest generalizations can represent th parent-form of th Vertebrata.—I follow him that there is a created archetype, the parent of its class.”21 Even in the absence of any definitive fossil evidence, Darwin was able to use a form of imaginary genealogical reasoning together with a concept of continuing extinction to reinterpret Owen’s idealized archetype as a concrete ancestor (Jenner 2022). This reimagining informed his living fossil concept, especially for aberrant forms lacking a fossil record, like Ornithorhynchus and Lepidosiren.
In the fall of 1854 Darwin returned to work on his species theory and on a nascent idea from his earlier theorizing; the principle of divergence would result four years later. His studies of barnacles had shown him that variation was actually abundant in wild populations even in the absence of changes in physical conditions. As historian Janet Browne noted, given this new perspective, Darwin reconsidered “how speciation and extinction occurred when there were no geological or geographical changes necessarily invoked by the theory” (Browne 1980, p. 59). Focusing first on extinction, he looked to aberrant living genera and taxonomic statistics of variation for evidence, thinking that their smaller species numbers were related to their anomalous state, and that they might be less well adapted and thus more likely to have experienced extinctions. He solicited taxonomic lists from colleagues and calculated average numbers of species in aberrant versus normal genera, initially finding that their different averages supported his view. He also found that small normal genera had very localized distributions while small aberrant genera had scattered distributions. For the latter, he inferred the extinction of some species of a genus that had once occupied a broad geographic range.
Darwin remained committed to the idea—dating from past consideration of osculant groups in classification—that character associations and extinction of intermediate taxa, not species number alone, were of paramount importance for judging the nature of aberrance.22 Exchanging views with Joseph Hooker in late 1854, he wrote, “to take an example, Ornithoryhnchus & Echidna would not be less aberrant if each had a dozen [...] species instead of one. What would really make these 2 genera less anomalous, would be the creation of many genera & sub-families round & radiating from them on all sides. [...] if it is extinction which has made the genus anomalous, as a general rule, the same causes of extinction would allow the existence of only a few species in such genera.”23 Many of Darwin’s contemporaries shared a similar understanding of aberrant forms as a taxonomic pattern, but not the expectation of combined processes of divergence and extinction as mechanisms underlying the pattern.
Darwin broadened his calculations beginning in late 1854 and soon confirmed differentially higher numbers of varieties in large genera, with varieties as incipient species. He wrote that “[a]ssuming species approximately constant, if extinction has fallen near and around the aberrant genera, then creation has fallen on the typical and larger genera.”24 An elevated rate of species production could provide an explanation for divergence. Groups undergoing increase would be prone to having species that are closely allied to one another while groups declining through extinction would have more dissimilar species. Small newly formed local genera, as opposed to dying aberrant genera, could be the source of new species, but Darwin needed an ecological mechanism that would operate whether or not local conditions were changing, or where organisms were not being subjected to unsettling forces (Browne 1980; Kohn 1985). His first inkling of local divergence increasing diversity as a result from struggle began early in 1855 and is apparent in his ecological interpretation of the division of labor in late 1856. Darwin wrote that “[t]he advantage in each group becoming as different as possible, may be compared to the fact that by division of [...] labour most people can be supported in each country—Not only do the individuals of each group strive one against the other, but each group itself with all its members, some more numerous, some less, are struggling against all other groups, as indeed follows from each individual struggling.”25
After discovery of an error in his calculations in mid-1857, Darwin agonizingly revised them under the guidance of his neighbor and younger colleague John Lubbock. Between then and late 1858, he completed his principle of divergence.26 Divergence itself became always occurring and plainly beneficial—new species exploit divergent specializations, driven in part by competition for places in the economy of nature. With consistent natural selection, more species would be able to coexist by becoming increasingly diversified (Browne 1980; Ospovat 1981).
Thoughts of aberrant or osculant groups and hierarchical classification had not left him, as indicated in an undated note scribbled in the margin of Hooker and Thomson’s Flora Indica: “Like ancient fossils, there are living fossils” he wrote (Hooker and Thomson 1855, p. 207).27 Darwin mused on the abnormal character of geographically isolated organisms in another note from November 1857, writing that “[t]he abnormal character of F. W. [fresh water] Plants (as Hooker agrees with me is case) No No (there is no reason why more shd be abnormal than of land Plants—Marsh Plants; not connected for others cd get so easily adapted: there is reason, like in any isolated spot, that the F. W. Plants, shd belong to many genera) may all be compared to the remnants of broken races of man in mountainous recesses—they are interesting under this view as living fossils.”28
While preparing his big book, Natural Selection, Darwin in 1858 added a section to his “Natural Selection” chapter dealing with the principle of divergence, writing, “in the long run, more descendants from a common parent will survive, the more widely they become diversified in habits, constitution & structure so as to fill as many places as possible in the polity of nature, the extreme varieties & the extreme species will have a better chance of surviving or escaping extinction, than the intermediate & less modified varieties or species” (as cited in Stauffer 1975, p. 238). A few pages earlier he had imagined that if Ornithorhynchus had an advantage to vary and increase in numbers, it would have become greatly diversified like other Australian marsupials. Implicitly, as Ornithorhynchus was indeed one of those intermediate and less modified forms, it lacked said competitive advantage. In his chapter titled “Difficulties On The Theory,” Darwin wrote:
[O]ne ought to think of such animals as the Ornithorhynchus, which though an indisputed mammal, presents in its skeleton & other parts some few plain resemblances to reptiles & birds. [...] Look at the mud‐fish (Lepidosiren annectens), which is so intermediate in structure, that although the greatest living authority considers it to be certainly a fish, many highly competent judges class it as a reptile: if then there be any truth in our theory, it would not be ridiculous to suppose that the Lepidosiren could [...] be modified by natural selection into an ordinary fish, or into a reptile. The case is almost parallel with that often encountered by philologists: to one who knew no other language, dead or living, besides French & English, how absurd would the assertion seem, that evêque & bishop had both certainly descended from a common source, & could still be connected by intermediate links, with the extinct word ‘episcopus.’ (in Stauffer 1975, p. 384)
Here Darwin drew an analogy from philology to bolster his argument for divergence, extinction, and the formation of gaps over the course of time.29 Philology is a source from which historians have traced contributions to Darwin’s conceptualization of descent in nature and humans (Alter 1999; Richards 2002b; Radick 2008). By the early 19th century, philologists had independently developed concepts of historical changes in words and languages comparable to later biological concepts of descent with modification, extinction, and divergence from a common ancestor (van Wyhe 2005; Turner 2014). Living fossils could occur in languages as they did in the natural world (Schleicher 1863; Krischel and Fangerau 2013).
In the middle of June 1858, Darwin received the letter from Alfred Russel Wallace that precipitated his rushed preparation of an abstracted version of Natural Selection. Between that summer and November of 1859, Origin was completed. An indication of ecological conditions related to his living fossil examples is present in an undated note, probably part of his preparations, discussing confined and peculiar habitats and their relationship to geographic distributions over time:
It may be too hypothetical, but the very same cause which prevent complication wd tend to leave the inhabitants of such peculiar habitats in lower state of complication. They wd be like fossil, pr wd be allied to those intermedial form, which we must believe to be parent of present orders & Familia. Now considering the fewness of F.W. [fresh water] forms the proportion of Being in this condition is surprising but it is very easy to deceive myself. Ganoid Fish - Lepidosiren (Proteus?) - Ornithorhynchus? new Echidna. [...] Hence we may say F.W. productions are living fossils. [...] Fr water lakes & rivers so small area, perhaps too much disjoined.30
Darwin solicited comments on sections of his Origin manuscript from a few friends and colleagues and used them in revisions and corrections. In December 1858, Darwin responded to a letter from Hooker on plant distributions, and there introduced his living fossil concept: “It is that species inhabiting a very large area, & therefore existing in large numbers & which have been subjected to the severest competition with many other forms, will have arrived through natural selection, at a higher stage of perfection than the inhabitants of a small area.—Thus I explain the fact of so many anomalous or what may be called ‘living fossils’ inhabiting now only fresh-water, having been beaten out & exterminated in the sea by more improved forms; thus all existing Ganoid fishes are fresh-water as is Lepidosiren & Ornithorhynchus &c.”31 In a letter to Charles Lyell October 1859, Darwin also defended the idea that organisms living in simple and largely unaltered habitats might be sufficiently well-fitted to such conditions and thus not show any history of significant change or increase in complexity.32 With these seeds of an ecological mechanism in hand, Darwin developed them in Origin as fundamental parts of his evolutionary arguments.
Darwin’s Concept in Origin and in Letters
Darwin’s mature living fossil concept is not developed fully in Natural Selection (Stauffer 1975). The closest ties are found in the chapter “Difficulties on the Theory,” where he discussed the survival of intermediate forms Ornithorhynchus and Lepidosiren, whose affinities connect what are now distantly related groups. By contrast in Origin, Darwin’s concept and example organisms became vital exemplars of the fourth chapter, “Natural Selection,” rather than merely one of the difficulties addressed in subsequent chapters. This change may be interpreted as a shift of living fossils by Darwin from pattern concerns of classification to process concerns as he structured his case to establish a true cause, or vera causa, in explaining his theory.
We contend that in Origin, Darwin’s living fossil concept became inextricably coupled with his concepts of natural selection and the principle of divergence. In the first paragraph of “Circumstances favourable to Natural Selection” in chapter four, he argued, “for as all organic beings are striving, it may be said, to seize on each place in the economy of nature, if any one species does not become modified and improved in a corresponding degree with its competitors, it will soon be exterminated” (Darwin 1859, p. 102). This statement underwrites a view of ongoing change and progressive development, and at first acquaintance appears to be contradicted by examples of long-persisting and unchanging forms. Darwin would turn that apparent contradiction to his advantage.
Darwin selected Ornithorhynchus, Lepidosiren and ganoid fishes as living fossil examples, expounding the existence of natural selection and the competence of his theory to explain their aberrance, long survival and continued existence. His argument is about establishing the explanatory plausibility of his theory, akin to what historian and philosopher James Lennox termed “how possibly” narratives, “used to turn supposed counter-examples into possible consequences of the operations of the causes proposed by his theory” (Lennox 2008, p. 68).
While acknowledging that isolation sometimes favors formation of new species, Darwin made the case that small, disjointed areas protected by barriers or peculiar but stable physical conditions may persist over time. Their occupants may likewise persist, undergoing gradual change up to the point where they are well adapted to the uniform conditions. Smaller numbers of inhabitants in these isolated habitats would lower the probability of useful variation and the inhabitants would eventually experience less pressure to change and diversify under natural selection. Conversely, among the occupants of large, more variable areas the struggle for survival—competition to fill more variable places in nature—would be more severe. They would continually adapt, diversify and improve over time. Yet their entry into, and domination over, certain isolated habitats would be deterred by barriers or physical conditions, or by long-established inhabitants already highly adapted to the distinctive local conditions. Ornithorhynchus, Lepidosiren and ganoid fishes are all found exclusively in freshwater habitats, which he compared to the isolation of small islands. So it is, as Darwin wrote, that “[a]ll fresh-water basins, taken together, make a small area compared with that of the sea or of the land; and, consequently, the competition between fresh-water productions will have been less severe than elsewhere; new forms will have been more slowly formed, and old forms more slowly exterminated” (Darwin 1859, p. 107).33 The living fossil concept now exemplified an ecological and evolutionary process, not simply a pattern of aberrance in classification. In brief, a living fossil was for Darwin an exception that conforms to the plausible outcome of a causal rule, natural selection.
Darwin’s anomalous living fossil examples also conformed to the process of divergence in combination with extinction. He wrote that “like fossils, [they] connect to a certain extent orders now widely separated in the natural scale” (Darwin 1859, p. 107). Character affinities connecting Ornithorhynchus with mammals and birds, and Lepidosiren with fishes and lower reptiles (amphibians), had been widely discussed by Victorian naturalists. In the text of the 1859 edition of Origin, the position of ganoid fishes intermediate between major groups is not clearly stated. There was no doubt that the great fossil diversity of ganoids in the Paleozoic had dwindled to a few living taxa. Louis Agassiz had created the order Ganoides based on the presence of bony diamond-shaped scales, though ganoid affinities and proper member taxa were debated soon after (Agassiz 1833; Lütken 1871). Darwin commented in his abstract of a paper by Huxley that the affinities of ganoids related them to two major groups, cartilaginous and bony fishes (Huxley 1858).34 Darwin added this explanation in the supplement to the second edition of Origin (1860), clarifying this attribute of their living fossil status. Years later in The Descent of Man, Darwin noted that both ganoids and Lepidosiren, “serve to connect more or less closely the several great vertebrate classes,” referring then to fishes and amphibians (Darwin 1871, p. 204).
The final sections of chapter four in Origin are devoted to “Divergence of Character,” wherein Darwin illustrated the process of divergence with extinction in the book’s only figure, his metaphor and concept of the tree of life. Unlike the many previous tree of life diagrams (Pietsch 2012), Darwin offered a plausible cause for the hierarchy and arrangement of taxonomic groups. The original lineages variously branch, diverge, form clusters as higher taxa, or become extinct, save for one relatively unchanging line, lineage F. He wrote, “[i]t is worth while to reflect for a moment on the character of the new species F,14 which is supposed not to have diverged much in character, but to have retained the form of (F), either unaltered or altered only in a slight degree” (Darwin 1859, p. 124). Scholars with varied disciplinary perspectives have oftentimes interpreted lineage F as a living fossil (Stamos 2002; Gross 2006; Costa 2009; Reznick and Ricklefs 2009). Yet Darwin did not specifically name this lineage as such. In the diagram, the patterns of divergence, extinction and stasis (in lineage F) are conjectural and also representative of every level of taxonomic organization; the narrative of process implied in his diagram operates together with observable patterns in nature and the geologic record as explanation for his causal theory of evolution (Gross 2006; Priest 2018). It is conceivable that while the imagined lineage F did relate to his living fossil concept, Darwin may also have had in mind groups with long fossil records and relatively unchanging forms that he did not include under his concept. In part of chapter ten, “On the Geological Succession of Organic Beings,” Darwin returned to the diagram and wrote, “it is quite possible, as we have seen in the case of some Silurian forms, that a species might go on being slightly modified in relation to its slightly altered conditions of life, and yet retain throughout a vast period the same general characteristics. This is represented in the diagram by the letter F14” (Darwin 1859, pp. 331–332). Neither Ornithorhynchus nor Lepidosiren had a known fossil record, and Darwin was aware that ganoid fishes became prominent in the Devonian, not the earlier Silurian.35 Darwin may have been referring to Silurian occurrences of lineages like the living brachiopod Lingula or the cephalopod mollusk Nautilus. In chapter nine, “On the Imperfection of the Geological Record,” he referred to both, writing that “[s]ome of the most ancient Silurian animals, as the Nautilus, Lingula, &c., do not differ much from living species; and it cannot on my theory be supposed, that these old species were the progenitors of all the species of the orders to which they belong, for they do not present characters in any degree intermediate between them” (Darwin 1859, p. 124). As far as we are aware, Darwin never explicitly acknowledged Lingula or Nautilus as a living fossil.
The only other mention of living fossil in Origin comes in a forward-looking discussion near the end of chapter fourteen, “Recapitulation and Conclusion.” Here Darwin wrote that in the future, classifications will become genealogies, and that “we have to discover and trace the many diverging lines of descent in our natural genealogies, by characters of any kind which have long been inherited. [...] Species and groups of species, which are called aberrant, and which may fancifully be called living fossils, will aid us in forming a picture of the ancient forms of life” (Darwin 1859, p. 486). This particular sense of living fossil would later be taken up by anthropologists and ethnologists in comparative studies of so-called primitive and advanced human populations and of an assumed evolutionary hierarchy of different racial and ethnic groups. We will return later to the suggested prospect of pictures of ancient life and of taxonomic patterns as genealogies when we consider the cultural evolutionists’ living fossil concept.
Darwin designated other living fossils twice in letters written after 1859. The organisms labelled in both instances survive in circumstances with few inhabitants compared to much larger surrounding areas and with peculiar or harsh ecological conditions that offer protection from competitive struggle. Darwin had closely followed works describing blind cave organisms—crustaceans, insects, spiders, the fish Amblyopsis, the amphibian Proteus, and others—and corresponded with some of the authors of those publications. In the section “Use and disuse, combined with natural selection; organs of flight and of vision” of Origin’s chapter five, Darwin had hypothesized an explanation for these forms, writing that “[b]y the time that an animal had reached, after numberless generations, the deepest recesses, disuse will on this view have more or less perfectly obliterated its eyes, and natural selection will often have effected other changes, such as an increase in the length of the antennae or palpi, as a compensation for blindness” (Darwin 1859, pp. 138–139). He concluded on the following page that, “[f]ar from feeling any surprise that some of the cave-animals should be very anomalous, as Agassiz has remarked in regard to the blind fish, the Amblyopsis, and as is the case with the blind Proteus with reference to the reptiles of Europe, I am only surprised that more wrecks of ancient life have not been preserved, owing to the less severe competition to which the inhabitants of these dark abodes will probably have been exposed” (Darwin 1859, p. 140). An article written by the botanist and entomologist Andrew Murray (1860) criticized Darwin’s theory as an explanation for the gradual evolution of blind cave insects. Murray’s critique, that blind insects of the same genus were found in different caves on two continents but that related sighted species did not exist outside the caves, alarmed Darwin.36 He wrote to Lyell, “[t]he difficulty is that amongst the blind insects of the caves in distant parts of world, there are some of the same genus, & yet the genus is not found out of caves, or living in the free world. I have little doubt that like the fish Amblyopsis & like Proteus in Europe, these insects are ‘wrecks of ancient life’ or ‘living fossils’, saved from competition & extermination.”37 Darwin addressed the issue in the third edition of Origin: “[I]t is possible that the progenitor or progenitors of these several species may formerly have ranged widely over both continents, and have since (like the elephant on both continents) become extinct, excepting in their present secluded habitations” (Darwin 1861, p. 156).
Lastly, Darwin referred to the newly discovered gymnosperm plant Welwitschia as a living fossil in late 1861. Joseph Hooker at Kew Gardens had been sent specimens of a bizarre plant that grew where few others could survive in the Namib desert of southwest Africa. Hooker set about trying to determine affinities of its obscure vascular organization and pollen cone structure, engaging in discussions with his friend Darwin. Darwin wrote to Hooker, “I have looked in Lindley about Gnetum; what a curious form your new one must be,—what a fine living fossil, preserved from past times.”38 He followed this thought two months later in another letter, that “[y]our African plant seems to be a vegetable Ornithorhynchus,—indeed much more than that.”39 Like Lepidosiren and Ornithorhynchus, Welwitschia had no known fossil record. Both its discoverer, Frederic Welwitsch, and Hooker determined a whole host of anomalous character relationships, affinities potentially linking it to widely different groups: cycadophytes, conifers, and dicotyledenous flowering plant families Casuarinaceae, Proteaceae and Santalaceae. Hooker eventually referred to Welwitschia as close to the Gnetaceae, while speculating on the characters of its ovule that, “when we see this organ occurring in a hermaphrodite flower, it is easy to suppose that we have in Welwitschia a transition in function, as well as in structure, between the gymnospermous and angiospermous Dicotyledons” (Hooker 1863, p. 24).
We have defended a view that against the expectation of ongoing change and progressive development, Darwin used living fossils in support of his evolutionary theory. Living fossils were exceptions that prove the rule of his principles of natural selection and divergence through the actions and circumstances these principles entailed in the natural world. Darwin associated several attributes with his named living fossil examples relevant to processes of natural selection and divergence, among them, isolation and protection from competition and struggle, extinction over time forming gaps and taxonomically intermediate aberrant forms, small species numbers and likely declining diversity over geologic time. These attributes of his living fossil concept are not without some ambiguities. Only ganoid fishes had an empirical fossil record attesting to patterns of morphological sameness and declining diversity. The lack of a fossil record as a possible attribute of Darwin’s living fossil concept is not supported by the ganoids. It is also unclear whether Darwin considered all of the referenced blind cave organisms as intermediate taxonomically between larger, more distantly related groups. Another form of ambiguity arises through Darwin’s interactions with cultural evolutionism and implications of racial hierarchy in The Descent of Man (1871), which we consider in a later section. For now, having traced the circuitous development of Darwin’s living fossil concept and his named examples, we turn to how that concept endures, even in light of transformations through the work of other scientists.
Variations on Darwin’s Concept after 1859
Scientific concepts like Darwin’s living fossil are dynamic entities that can undergo transformation to broaden or narrow their properties and scope across time, authors, and contexts, yet still retain their epistemic function (Brigandt 2010; Surman et al. 2014). Variations of Darwin’s concept have persisted for a century and a half. They are part of a research agenda of questions applied to a vast menagerie of entities at many levels in the biological hierarchy (Lidgard and Love 2018; Lidgard and Kitchen 2023). After 1859, as scientists began discussing living fossils and labeling new ones, variations among the properties chosen to distinguish living fossils became apparent. Outside the sciences in the wider public domain, the popular metaphor living fossil and certain attributes of Darwin’s living fossil examples spurred renewed literary forays into little-known places and atavistic appearances.
Historian James Secord inquired about the circulation of knowledge, “[h]ow does it cease to be the exclusive property of a single individual or group and become part of the taken-for-granted understanding of much wider groups of people?” (Secord 2004, p. 655). Darwin’s concept endured well after 1859, but was subject to transformation in various ways as natural historians and others found a novel, advantageous role for living fossil—an alluring focal point for their work. Darwin’s concept was no longer proprietary in the sense that it became ineluctably social among members of a highly motivated community, whether or not they recognized the concept’s central role supporting Darwin’s arguments. Among the ensuing publications on living fossils, associations with natural selection and divergence together or separately were varied. Authors seldom referenced more than one of the attributes that Darwin weighed, if they appeared at all. Darwin’s case for transmutation and branching descent met with general acceptance among many scientists. Natural selection and its integrated causal role received a far more contentious reception (Hull 1983). This difference is apparent in the sorts of criteria that other scientists used in discussing living fossils. A few, including even the Reverend George Warington, defended the credibility of the process of natural selection in explaining living fossils, writng that “[i]solated localities—as islands, fresh-water lakes, caves, &c.—are ever found to present the greatest number of peculiar forms, often so resembling bygone types as to receive the name of ‘living fossils.’ While, if the comparative improved condition of the species generally be inquired after, it needs but to put the flora and fauna of an isolated and extended area into actual competition, the result speaks for itself” (Warington 1867, p. 56). The responsibility of natural selection for living fossils Ornithorhychus and Lepidosiren was repeated at times in scientific publications (Symonds 1864; Büchner 1868). August Weismann constructed a most unusual integration of natural selection with the living fossil concept in theorizing evolutionary transformations of larval developmental stages among different species of sphinx moths (Weismann 1882). He invoked natural selection as the cause of increasingly complex marking. He had discovered a rare larva, a living fossil—he wrote—with only slight marking. The other known species had more marking that he inferred had evolved to better camouflage them from predators (Beddard 1892).
Variations on Darwin’s scientific living fossil concept that focused primarily on divergence were more common, usually with little or no reference to struggle and competition, isolation and protection, or natural selection in general. These designations usually characterized a living fossil taxon as intermediate between recognized groups or implied a taxonomically deep branch in a genealogical sense based on shared affinities. Taxonomic, philosophical and popular science publications applied this manner of description to the platypus, echidna and lungfish (Büchner 1864, 1890; Tissandier 1875; Hopley 1882; Anonymous 1887; Lumholtz 1890; Clodd 1895). In a hypothesized phylogeny, paleozoologist Ludwig Rütimeyer named camels and Tragulus as living fossil end members of a basal group of ruminants leading to divergent lines of ungulate mammals (Rütimeyer 1866, 1898).40 The anomalous character relationships of the Philippine flying lemur Galeopithecus and of Brachyuromys and related Madagascar rodents resulted in more living fossil designations (Finn 1893; Major 1897). The tuatara, Sphenodon, was named a living fossil whose peculiar characters were used to determine that it was the only living representative of the Rhynchocephalia, an otherwise extinct order of reptiles (Anonymous 1892). Yet another example emphasized divergence and past extinction of related taxa: “The Hoatzin is more than to any other birds nearly related to the Fowls. But he diverges from them so far that he is regarded as the sole representative of a distinct group. All his near kindred have passed away, and he stands solitary, a living fossil, the only survivor of a number of families that have either disappeared, too primitive to hold their own, or have advanced to a higher organisation” (Headley 1895, p. 286).41
In assessments of Darwin’s principle of divergence and certain hypothesized intermediate forms, naturalists sought physical evidence from both living and fossil taxa, as for example among the possible relationships of Archeopteryx to birds and reptiles (Huxley 1868). Darwin’s work inspired new directions for naturalists who sought knowledge of both known and potential living fossils, including lungfish (Conant 1986; Rieppel 2023). Crinoidea, a class of marine echinoderms dredged up from the ocean’s depths, became central to such searches following naturalist and biogeographer Edward Forbes’ study of crinoid fossils and their relationship to living groups (Thomson 1873; Alaniz 2014). Linking the anomaly of deep sea forms to their isolation, protection from competition and struggle, and ancient history, Huxley suggested that “the things brought up will very frequently be zoological novelties; or, better still, zoological antiquities, which in the tranquil and little-changed depths of the ocean have escaped the causes of destruction at work in the shallows, and represent the predominant population of a past age” (Huxley 1873, p. 833). For individuals like Charles Wyville Thomson, professor of natural history at Queen’s University in Belfast, the deep-sea floor and its purportedly unchanging environment promised an ideal site to search for intermediate living fossil forms (Alaniz 2014; Pettitt 2020). Living crinoids dredged up from Norwegian fjords had suggested partial validation of Darwin’s theories and the stalked crinoid Rhizocrinus lofotensis boasted a morphology previously only found in extinct fossil forms. Forbes had determined a polarity of extinct cystoids and many living echinoderms, and this form was morphologically intermediate between the two (Rehbock 1983).42 Further discoveries of stalked crinoids confirmed for Thomson the possibility that living crinoids could be “missing links which might connect present with the past” (Pourtalès 1868; Thomson 1873, p. 80). While the findings of Thomson’s 1872 HMS Challenger expedition ultimately suggested that there was no smooth gradation or evidence of transitional forms that linked species in a way that would satisfy his interpretation of Darwin’s natural selection, the initial publication of results garnered large scientific and public interest in living fossils (Thomson 1880).
Another aspect of the scientific living fossil concept—surviving terrestrial geographic relicts—developed from growing recognition of geologically older floras that had been protected from adverse changes in ecological conditions and survived in isolated pockets in the wake of past glacial advances and retreats. James Geikie, for example, described isolated remnants of the Scandinavian flora thus: “[T]his flora is largely confined to the northern latitudes; but isolated colonies occur in our own mountains and in the mountains of middle and southern Europe. [...] [T]hey are, as it were, living fossils—monuments of former great physical and climatic changes” (Geikie 1887, p. 404). Similar floristic remnants were described as living fossils on several peaks of the Atlas mountain system in Algeria (Battandier 1894). Relatedly, a few elements of a former subtropical flora on Sakhalin Island were referenced as living fossils that had survived in protected pockets through the interval of glacial cooling (Krasnow 1896).
In the public domain, geographic relics flourished in popular tales of exploration. These were not the same as a scientific living fossil concept, but use of the popular metaphor appeared to draw parallels on some features. As with the Challenger expedition and its crinoids, naturalists’ travel accounts told tales of animals discovered in little-traveled places, and fictional encounters with prehistoric beasts made for pulpy literary fodder. The time-travel tales of H. G. Wells, and Pierre Boitard’s novel Paris avant les hommes (1861) considered encounters between denizens of different evolutionary stages. However, encounters with primeval relics were not confined to these tales alone. Several 19th century literary and natural historical works used the motif of isolated animals or habitats surviving beyond their presumed disappearance, then being discovered during an adventure (Fallon 2024). A much older tradition of the lost-world narrative can be found in medieval travel books and works from the Enlightenment (Sommer 2007). Still, from the 19th century onwards, European expeditions to very real but very faraway places gave new life to those narratives. And just underfoot, the subterranean domain held a privileged place in the Victorian mind. As naturalists exhumed relics and fossils from underground, they recast the geologic history of local neighborhoods to involve cave-dwelling hyenas and hippos (Buckland 1823; Sommer 2003). Popular works, such as Jules Verne’s Voyage au centre de la Terre (1864) saw European protagonists encountering Mesozoic ichthyosaurs and more in underground lost worlds.43 Arthur Conan Doyle”s The Lost World (1912) offered vindication for its protagonist who brings home a living pterodactyl as proof of the various living fossils encountered. Its unfortunate escape later on paints the discovery and return in a somber tone: “[T]here can be no doubt that somewhere out in the wastes of the Atlantic, the last European pterodactyl found its end” (Doyle 1912, p. 303).44 The possibility of rediscovering the past in distant places formed an associative space for imagining natural historical, anthropological, or fictitious living fossils. A commonality among these real or imagined tales was the difficult work required to access the spaces in which living fossils lurked.
As the 19th century neared its close, a few scientific claims of living fossil status appeared that were based on little more than the close resemblance of living forms to ancient relatives, some of which had been presumed extinct. Here a variant of Darwin’s scientific living fossil concept persisted, even though references to the principle of divergence or natural selection were weak, at best. In his extensive study of radiolarian protozoa, Ernst Haeckel argued that, “the number of these ‘living fossil’ forms is much greater than was previously supposed [...] Among the Miocene Radiolaria numerous species [...] are not to be distinguished from the corresponding still living forms” (Haeckel 1887, p. clxxiv). The definition of living fossils by prolific naturalist Richard Lydekker was just as limited:
For such survivors from a distant past we venture to suggest the title of ‘living fossils,’ seeing that for the most part they have but little in common with the dominant fauna of the greater part of the world; while their alliance with extinct types is of the most intimate kind. It is, of course, difficult to know where to draw the line in the use of such an arbitrary designation; but we shall endeavour to restrict the term either to types which, although still more or less abundantly represented at the present day, are of extreme antiquity, or to such as are now represented by comparatively few forms, living either in distant parts of the world or in the ocean depths, but which were abundant in past epochs. (Lydekker 1894, p. 153)
Lydekker added more living fossil taxa to the existing registry: the mollusks Pleurotomaria, Trigonia and Nautilus, the brachiopods Lingula, Crania and Discina, the crinoids Pentacrinus and Rhizocrinus, the Port Jackson shark Cestracion (now Heterodontus), the giant salamander Cryptobranchus (now Andrias), the opossum Didelphys, and a few others.
We found no statements showing that Darwin explicitly acknowledged the living fossil status of the additional taxa designated by others. While we do not claim that he thought none of them complied with his concept in Origin, his reflections on the ancient plant lineage of Ginkgo offer a possible clue to his thinking. Linnaeus named the genus, but English botanist James Smith inappropriately substituted the name Salisburia and placed it among the conifers (Linnaeus 1771; Smith 1797). In the 1850s, Hooker thought Salisburia shared affinities with cycads, which at that time were classified as conifers. Darwin was aware that Salisburia and its allies had an abundant fossil record extending back to the Jurassic at least.45 Then in a letter to Darwin in 1877, the paleobotanist Gaston de Saporta expressed the view that placing certain plants including some cryptogams and Araucaria and Ginkgo “among the conifers” was both “robust and persistent,” with these groups showing little or no variation over a vast expanse of geologic time.46 In his reply, Darwin referenced not living fossils, but rather the past statements of Thomas Huxley: “With respect to persistent types your cases are curiously analogous to those which occur in many groups of the animal kingdom & have been commented on by Huxley.”47 Huxley had previously argued, along with Lyell, that the fossil record did not support progressive improvement among successive forms over geologic time (Lyons 1993). Huxley’s concept of persistent types accepted transmutation of species, but at least initially did not countenance transitions between major groups that belonged to distinct morphological and developmental archetypes (Lyons 1995). Huxley wrote that “certain well marked forms of living beings have existed through enormous epochs, surviving not only the changes of physical conditions, but persisting comparatively unaltered, while other forms of life have appeared and disappeared. Such forms may be termed ‘persistent types’ of life; and examples of them are abundant enough in both the animal and the vegetable worlds” (Huxley 1859, p. 152). Over vast amounts of time, some forms persisted and others changed, both tendencies being compatible with Darwin’s theory. Consistent with our interpretation, Darwin apparently never wrote that Salisburia (Ginkgo) was a living fossil, despite the assertions of many subsequent authors.
By the end of the 19th century, a living fossil in scientific discussions was a burdened organism, indeed. For Darwin it emerged quite soberly from evidence of intermediacy in classification, as well as its seeming escape from competitive struggle and congested habitats. It operated, for him, as an exception that demonstrated the explanatory competence of his evolutionary theory. For others, however, living fossils became anomalies writ large, exemplars of earlier states of evolution, demonstrations of conserved traits, or relics bridging a time-gap to the past. In popular metaphors, many of these themes materialized with living fossils coming to characterize any out-of-time appearance. In anthropology, living fossils would emerge as a distinct scientific concept out of different theoretical backgrounds and intellectual commitments.
Humans as Fossils in a Cultural Anthropological Context
Encounter was a driving theme for the anthropological living fossil concept. Darwin had considered the aberrant living forms of remote Australia, as well as underground caves. His description of what we might call the conditions of existence and persistence for living fossils, both in an 1858 letter to Hooker, and ultimately in chapter four of Origin, centered on the importance of isolation and, consequently, protection from competition.48 In that same 1858 letter to Hooker, Darwin demonstrated the detrimental impact of evolving and existing in a small area. He noted how the New Zealand flora and fauna “yield to those of Europe.” Darwin was imagining the effects of competition and natural selection in large, connected spaces with plentiful natural variation, and during encounters between improved occupants of these large expanses and plausibly less improved occupants of smaller isolated territories (Tammone 1995; Paterson 2005). Importantly, he was doing so during a period of colonial expansion. Increasing exploration and contact were encouraged in part by the notion of distant colonized lands as so-called lost worlds, where populations of plants, animals and peoples had long endured in a primeval state (Stafford 2017). The use of scientific ideas such as natural selection as justification for social and political ideologies in the wake of Darwin’s writing, and his own relationship to racial theorizing in the context of colonialism, are important parts of an ongoing discussion (see below). Encounters between Europeans on the move and populations around the world, precipitated a boundary-breach which he characterized as potentially disastrous to isolated, less improved forms in the natural world, a view which was extended to people (Brantlinger 2003). In Origin, isolation of certain freshwater organisms from severe competition with occupants of the vast lands and oceans enabled largely unchanged forms to persevere through time. Encounters between large and small populations of people, cosmopolitan and remote, formed a real political backdrop for this thinking.
Beyond the phrase living fossil itself, and beyond isolation and relic populations, what was shared between anthropological and Darwinian living fossil concepts? Tracing the influences operating on the early discipline of anthropology is hardly simple. The varied subfields of anthropology precipitated out of a range of intellectual traditions: linguistics and philology, antiquarianism, ethnology, anatomy, geology, paleontology and more (Waitz 1863; Stocking 1991; Alter 1999; Manias 2013). Darwin wrote at length on the parallels between the evolution of species and languages in The Descent of Man (1882), even comparing crinoids to language (Radick 2008). Both Darwin and the cultural evolutionists used living fossil to assert categorization, something that innovative metaphors often do effectively (Glucksberg 2008). Still, the intellectual inheritance of the anthropological living fossil concept differs from Darwin’s natural historical one.
The lens applied to studies of human diversity before the mid-19th century was one of development. Prior to Origin, interest in human origins and racial diversity attracted views as divergent as those of polygenist Robert Knox and salvage ethnologist James Cowles Prichard (Sera-Shriar 2015). Prichard, founder of Britain’s Ethnological Society, defended a single origin of humanity as a monogenist, and drew attention to the risk of extinction for Indigenous populations. It was through ongoing colonial encounters that a concern over the fate of Indigenous cultures was fostered (Brantlinger 2003). Prichard’s “On the extinction of human races” (1839), read before the British Association for the Advancement of Science, spotlighted the precariousness of so-called primitive groups and the value of their cultural practices that might be lost through colonial domination (Bowler 1992). Darwin was present at that meeting, heard the paper, and was inspired to join the British Select Committee of Aborigines formed to solicit information on endangered people from travelers abroad (Gruber 1970; Richards 2017).
Prichard’s influence on Darwin’s early thinking about human races and their fates was manifold, and distinct from the contributions of later cultural evolutionists. Counter to Knox and American physical anthropologists such as Samuel George Morton, Prichard and Darwin shared a monogenist view and emphasized human societal development in a non-unilinear framework (Kuklick 2012; Sera-Shriar 2017; Douglas 2022). Prichard’s calls for preservation of endangered groups, largely via collection practices, influenced Darwin, and both men considered colonial presence as a precipitating cause of that endangering.49 Darwin’s early work and engagement with Prichard no doubt helped shape his views on the perilousness of Indigenous encounters with European colonial expansion. Still, Darwin’s later writings on human evolution exemplified in Descent held together the views of not just salvage ethnologists, but also cultural evolutionists, physical anthropologists, and archeologists (Richards 2017).
Through the mid-19th century different measures of relative progress of human groups were developed: numerals to express different quantities (Crawfurd 1863); technological development including animal and plant domestication, tools and weapons (Figuier 1861; Lubbock 1865; Pitt-Rivers 1906); comparative anatomy including brain case size (Nott and Gliddon 1854); and others. As anthropological pioneer Theodor Waitz articulated in 1863, the classification of humankind according to descent, not mere “resemblance,” proceeded upon three tracks: “the physical, the linguistic, and the historical stand-points,” and crucially, “the results obtained by no means agree” (Waitz 1863, p. 230). With the emergence of various methods in anthropology, especially in social and cultural evolution, a space was fashioned for considering humans as living fossils. Intellectual historian James Turner observed that “physical anthropology had thin connections to philology or antiquarianism [...] [i]n contrast cultural and social anthropology grew directly from philology and allied studies” (Turner 2014, p. 330). The scientific concept of living fossils in 19th century anthropology is associated more vividly with the work of social and cultural evolutionists than that of archeologists and physical anthropologists. Cultural evolutionists sometimes did draw upon knowledge from other emerging anthropological studies. For instance, from the 1830s through the 1850s, discoveries of stone then bronze then iron artifacts in stratigraphic sequence, of presumed human skeletal fragments associated with bones of extinct fossil animals, and of more complete skeletons found near the Neander River and in Brixham Cave established the great antiquity of humans and the apparent extinction of human-like Neanderthals (Rudwick 2008; Madison 2020; Qureshi 2020; Walker et al. 2021).
Imaginative approaches to studying humanity’s past took shape under a loose framework of social and cultural evolution. Rather than emerging from hypothesized processes of natural selection and divergence, the human living fossil emerged from social-historical views that in various ways echoed ideas from the Scottish Enlightenment (Reybrouck 2012; Manias 2015; Qureshi 2020). A tradition of “conjectural history” prominent in the late 18th century classified stages of societal history of humans without necessary reference to deep time (Wolloch 2011, p. 252). Typically, these stages were hunting (savage), pastoral (herdsmen), agricultural, and commercial (Meek 1976; Olson 2003). In turn, these were sometimes viewed as parallel to stages of individual development. The relevant theoretical claim was that human societies universally developed according to a fixed ladder-like, or stadial, pattern (Wolloch 2011; Irving-Stonebraker 2017). Moving on to the 19th century, cultural evolutionists inherited this background, and developed a concept of human prehistory in which societies developed over time in a more or less unilinear succession of stages, that certain human groups could bear the hallmarks of their primitive pasts, and that not all human groups progressed at the same rate.50 Seemingly primitive peoples encountered by Europeans were largely unchanged relics from a shared human past. In what became known as the comparative method, these contemporaries were proxies for humanity’s earlier stages of development, and their presumed static nature saw them labeled as living fossils (McNiven and Russell 2005; Turnbull 2022; Lucas 2023). This developmental perspective, as historian George Stocking called it, was far from monolithic (Stocking 1991). Cultural evolutionists were a fluid group, and no precise doctrine was expansive enough to encompass all of them.
The comparative method of these 19th century anthropologists looked back to the idea that different human societies followed a shared stadial pattern of development, while adding its own operational and racialist shifts (Manias 2015; Seth 2016; Irving-Stonebraker 2017). Foremost among the cultural evolutionists were John Lubbock and Edward B. Tylor in Britain, and Lewis Henry Morgan in America. In 1865, Lubbock made a stark connection between the relative evolutionary progress of animals and that of people. He wrote that “[i]n fact, Van Diermaner, [Tasmanian] and South American are to the antiquary what the opossum and the sloth are to the geologist” (Lubbock 1865, p. 336). Identifying human groups as proxies involved seeking living fossil vestiges within cultures that could give clues to the past. Tylor considered his so-called “survivals,” referring to rituals, irrational practices and habits, as evidence of earlier stages of development (Stocking 1991; Lucas 2023). That he gave them the moniker survivals resonates with the living fossil nature of these proxies: they had survived where other practices had gone extinct. Philologists’ genealogies of languages and presumed stasis and universality of cultural customs were invoked in a “rough scale of civilization,” with occasional reference to comparative physical form (Tylor 1871, p. 24; Stocking 1991; Turner 2014; Mandler 2020).
Tylor stated his argument on the development of human culture with intended clarity: “Its standard of reckoning progress and decline is not that of ideal good and evil, but of movement along a measured line from grade to grade of actual savagery, barbarism, and civilization. The thesis which I venture to sustain, within limits, is simply this, that the savage state in some measure represents an early condition of mankind, out of which the higher culture has gradually been developed or evolved, by processes still in regular operation as of old, the result showing that, on the whole, progress has far prevailed over relapse” (Tylor 1871, p. 28). Both Morgan and Tylor had an explicitly unilinear developmental framework, leaving open the conclusion that less advanced contemporaries were proxies for earlier stages of development, through which more advanced groups had already passed. Being stuck in, or relapsing into, an earlier phase was often considered less a measure of survival success, as in the case of living fossil organisms, but rather a developmental failing (Lubbock 1865; Darwin 1871; Kingstone 2018).51 From there, cultural evolutionism built its knowledge on the fundamental assertion that human groups were in fact actual relics from an earlier stage of societal development (Guha 1998; Seth 2016; Sera-Shriar 2018). The proffered assumption was that some primitive groups, their implements, languages and cultures, more or less stopped evolving at different points in the past and persisted unchanged into the present.
The cultural evolutionists’ framework of human living fossils certainly had elements in common with Darwin’s living fossil concept, yet the cultural evolutionists themselves cannot be read as strictly Darwinian (Kuklick 2012; Candea 2018). The two concepts are set in different theoretical backgrounds and contexts. As historian Peter Bowler has said of the cultural evolutionists’ perspective: “This was evolution—but it was not very Darwinian because its basic model of progress was a ladder leading towards the goal of modern industrial society, not a branching tree driven by divergent adaptations” (Bowler 2021, p. 137). Their views are often associated with the social evolutionism of Herbert Spencer (Spencer 1857; McNiven and Russell 2005). Natural selection was not widely taken up by them, and theories of variation and differentiation were not their focus (Murphree 1961; Stocking 1991). The relationships of the various cultural evolutionists to Darwinian principles is complex. Lubbock cited both Darwin and Herbert Spencer, drawing on their notions of progress, but Morgan and Tylor resisted reference to a Darwinian view of evolution (Murphree 1961).
Darwin never explicitly referred to any human group as living fossils. Still, his treatment of human origins and races in Descent did not align seamlessly with the developed concept of living fossils in the natural world described in Origin. There are notable similarities between his discussion of racial hierarchies and primitive extinctions, and the cultural evolutionists’ frameworks. Darwin’s posture in Descent owed much to cultural evolutionists; historian George Stocking counted nearly fifty references to exponents like Tylor and Lubbock (Stocking 1991). Yet he also drew on Prichard and the ethnologists (Browne 1996; Desmond and Moore 2011; Sera-Shriar 2015). How, then, should historians interpret his views on primitive groups, living fossils, and racial hierarchy, given these sometimes conflicting influences?
Historians have recognized that Darwin, despite his abolitionist views and other humanitarian postures, is not absolved from many of the racial hierarchical views that were prevalent in cultural evolutionist discourse. The full nature of his attitudes towards race are a subject of considerable study (Alter 2007; Radick 2008; Desmond and Moore 2011; Qureshi 2011; Manias 2013, 2015; Sera-Shriar 2015; Richards and Ruse 2016; Higgitt 2021; Milam and Seth 2021). His most famous reflections on Indigenous people derived from his encounter with the people of Tierra del Fuego during his Beagle voyage, and exposed his shock at what he described as “unbroken savages” (Desmond and Moore 2011; Seth 2016; Richards 2017).52 That fascination with the extraordinary difference he perceived contributed to his consideration of humanity’s former states. He wrote, “could our progenitors have been men like these?—men, whose very signs and expressions are less intelligible to us than those of the domesticated animals; men, who do not possess the instinct of those animals, nor yet appear to boast of human reason, or at least of arts consequent on that reason. I do not believe it is possible to describe or paint the difference between savages and civilized man. It is the difference between a wild and tame animal.”53 Such passages have led historians to emphasize that, though Darwin defended a monogenist position, his racialist thinking nonetheless identifies him as an “emotional polygenist,” shocked as he was by the difference (Richards 2017, p. 6). Abolitionist and humanitarian commitments did not preclude Darwin from thinking in racial hierarchical terms (Douglas 2022).
Even through his monogenist posture, Darwin expressed the differences between races in ways that could conceivably be interpreted as a semblance of living fossils, emphasizing the possible informative alikeness of so-called savages to former states of humanity, and reflecting on the dwindling numbers and ranges of those people. Entries in Darwin’s diary record encounters with natives during his brief times ashore in Australasia and South America. Although he did not explicitly charge the people he met with being relics from earlier states of humanity, he later came close in Descent (Darwin 1871; Bowler 1992; Reybrouck 2012). Isolation (often geographic) had sheltered certain groups from the competitive pressures that otherwise yielded cultural progress, and thus those groups retained a primitive state (Bowler 1992). He considered the gaps that would be created from extinctions, as well, writing that “[t]he break between man and his nearest allies will then be wider, for it will intervene between man in a more civilized state, as we may hope, even than the Caucasian, and some ape as low as a baboon, instead of as now between the negro or Australian and the gorilla” (Darwin 1871, p. 201). Perhaps here Darwin was thinking again with Prichard and the salvage ethnologists, lending their concerns over extinctions to ongoing disappearance of evidence for his views of the evolutionary history of humans.
Here, however, the similarities between the meaning of living fossil in Origin and that of the cultural evolutionists diverge. They stemmed from different intellectual inheritances, subjects, and motivations. The value-laden language of a natural historical living fossil suggested forms that were successful in their primitive states, and well-suited to their particular lifestyles; endangered human groups’ lifestyles were often seen as ill-suited. Similarly, while the living fossil savage was promulgated as an antithesis to the European (Richards 2017), for a platypus no such binary existed. Rather its affinities to other kinds gave it its status. These differences demonstrate the versatility of the deeper living fossil metaphor, and the nuanced meanings it raised in different contexts, even for the same individual.
Of course, it would be wrong to think that it was only in the case of human living fossils that cultural constructs applied. In the natural historical cases of non-human groups, often social corollaries were used to help interpret observed patterns and behaviors, and the living fossil concept broadly could be wielded with an emphasis on primitivism, stagnancy, survival, or capacity for enduring change in either context. Both can be similarly understood as having been informed by the social context and constructs of the day. And while only nominally Darwinian, the cultural evolutionists’ concept of a human living fossil formed a thematic bridge for zoological, paleontological, and ethnological conceptualization. The precondition of deep time for human history, too, provided a catalyst for relating biological, geological, and archeological natural sciences, while cultural studies of difference and inferred evolutionary stages facilitated that exploration (Grayson 1983; Van Riper 1993; Rudwick 2005; O’Connor 2007). References to living fossils abounded in this context, with naturalists at the nexus of comparative studies in human and natural sciences drawing on the living fossil’s conceptual weight. For example, in 1875, Clémence Royer, the unauthorized French translator of Darwin’s Origin and elected member of the Société d’Anthropologie de Paris, identified the orangutan as “intermediate between the group of walking bimanes and that of climbing quadrumanes,” and therefore neither “man nor monkey,” but a “sort of aberrant form and true living fossil” (Royer 1875, p. 624).54 Following the Neanderthal fossil discovery in the Neander Valley in 1856, the lines between the comparative anatomical and cultural evolutionary approaches to humanity’s development were blurred even further. The hierarchical view of racial progressions through time was applied to fossil Neanderthals in the mid-19th century, and since (Mulvaney 1966; McNiven and Russell 2005; Madison 2020).
In no other case was the application of living fossil status to a human group more consistent than that of the Indigenous Australians. Geographically distant from Europe, and already courting interest along with Australia’s aberrant animal forms, these Indigenous people were framed by European naturalists as so-called missing links in humanity’s evolutionary history (Stocking 1991; Turnbull 2022). Huxley’s lectures and subsequent Evidence as to Man’s Place in Nature (1863) drew connections between Neanderthal skull fragments and Indigenous Australian crania from the Hunterian collection in London. This connection threw “living savage races into the fossil gap” (Stocking 1991, p. 48). Huxley argued that it would not take much formal change to the Indigenous Australian brain case to make it identical with “that of the aberrant fossil” of Neanderthal (Huxley 1863, p. 180). Lyell critiqued Huxley’s suggestion, arguing that without geological evidence, one could not assume that “the appearance of what are called the inferior races of mankind has always preceded in chronological order that of the higher races” (Lyell 1863, p. 90). The collapse of time that living fossils accomplished, bringing the past into living color and therefore study in the present, was not itself a satisfactory argument for the true conditions and forms of earlier ages without sufficient fossil evidence. Lyell’s critique suggested that while comparisons were being drawn between living groups and Neanderthals, those comparisons were not adequate justification for a true chronology between them (McNiven and Russell 2005). The comparison seemed merely analogical rather than historical. Still, towards the end of the century, Tylor pushed his spectrum further in his consideration of the Tasmanian people, for whom he argued “the lowest of normal tribes may be claimed for them.” The Tasmanians, he maintained, “place us in full view of the state of a people in the paleolithic stage,” while their “Australian neighbors” occupied the space of “modern neolithic savages” (Tylor 1894, p. 152).
What were the implications of someone being a living fossil in a Victorian colonial and scientific framework? In Europe, certain ethnic and religious minorities including Basques and Jews were not immune from being labelled as living fossils (Lagarde 1873; Leroy-Beaulieu 1895). For less advanced societies at the outposts of empires, the prognosis was not hopeful. In 1881, Charles Darwin wrote to philosopher William Graham on the role of natural selection in the “progress of civilization,” such that “at no very distant date, what an endless number of the lower races will have been eliminated by the higher civilized races of the world.”55 Darwin himself was ambiguous on endangered groups’ capacity for improvement or change via the civilizing hand (Richards 2017). Whether or not there was possibility for those primitive groups to adapt, progress, or improve was tied to related political views and justification for radical imperial policies; extinction was regularly viewed as an inevitable outcome (McGregor 1993; Mandler 2020; Sepkoski 2020). Patrick Brantlinger called this extinction discourse “a powerful axis of ideas that has been hegemonic for countless European explorers, colonists, writers, artists, officials, missionaries, humanitarians, and anthropologists” (Brantlinger 2003, p. 190).
From the conceptual vista of a law of progressive societies, such as that put forward by Herbert Spencer, primitive groups were seen as having failed an evolutionary race, rarely as impressive survivors (Claeys 2005). Poor environments were sometimes blamed for the living fossil status of such groups, stymying their evolutionary progress (Bowler 2021). Yet isolation was often touted as the only means of survival for those groups, incurring less competition to both drive change and render them extinct. Robert Frazer described certain hill tribes of northern India as “living fossils of primeval times,” having been “driven by other invading races to the lofty mountain ranges, where, amid the dense forests, their descendants still live undisturbed” (Frazer 1896, p. 49). William Henry Flower’s view of the declining numbers of Bushmen (now San peoples) and related groups in Africa is similar (Flower 1889). These cases can be interpreted as human groups embodying something loosely akin to biological unfitness in the natural world, largely devoid of adaptation to modern existence. The savagery involved in the colonial enterprise of the 19th century, sweeping, as it did “from the Hottentots in Africa through the Veddahs, the Australians, the Tongans, the American Indians, down to the Fuegians at the tip of South America” characterized groups as living fossils at the same time that it precipitated discourse about their inevitable extinction (Stocking 1991, p. 153).
Living fossils in the Darwinian conception were isolated relics that had much to say about former states in evolutionary history. Purported living fossil humans functioned similarly. Still, a different emphasis arose from the lessons that crinoids or platypus could impart compared to relic languages or Indigenous Australians. By having emerged out of different theoretical contexts, the former were aberrant survivors and the latter were near-extinct vestiges. A belief in laws of progress underpinned both views, but their respective relation to possible future development and valorous persistence differed. While transformist thinking, adaptation to protective environments and successful evasion of competition informed the fates of organismal living fossils, extinction thinking largely tempered the imagined possibilities for human living fossils. Humans might not improve, or if they did it would be through supportive intervention. Whether living fossils courted admiration for their survival or disgust for their primitive persistence was context dependent. The social dimensions of the living fossils, therefore, could be quite cautionary. As literary scholar Jessica Krzeminski has argued, individual living fossils became an essential character in Victorian culture as their unchanging nature was itself considered monstrous (Krzeminski 2021). In the late 19th century, transformation—once a source of unease—was now an expectation for humankind and nature, and societal stagnation became a new locus of concern.
Placing Living Fossils in Context
By the end of the 19th century, the colloquial and literary use of the popular metaphor could be reconnected with Darwin’s scientific concept and the relation it bore to human evolution and linguistics. A short story from the end of the century encapsulates these entangled commitments. In 1898, Wendell Phillips Garrison, an American literary abolitionist and chronicler of Darwin’s work for young audiences, published a short satire set as a sequel to Jonathan Swift’s famed Gulliver’s Travels (1726). Garrison’s piece, The New Gulliver (1898), followed a philologist as he became stranded on the isle of the Houyhnhmns, that race of talking horses from Swift’s earlier imagination. Arguing that “if Cuvier could reconstruct an animal from a few bones, or from a single one,” a philologist “should be able to frame a grammar and even a vocabulary from materials not less scanty,” the philologist sets out to explore the local horse culture (Garrison 1898, p. 17). The language-adept Houyhnhmns surprise the protagonist with an enclosure containing remnants of a “troop of diminutive four-toed Houyhnhmns” (Garrison 1898, p. 25). These forms, which had come from “remote swamps,” and displaying no discernable language nor “arts or civic organization,” are quickly related by the author to “what Darwin would have called a ‘living fossil,’ that had endured so long ‘from having inhabited a confined area, and from having been exposed to less varied, and therefore less severe, competition’” (Garrison 1898, pp. 25, 32). They are later identified with Orohippus major, an Eocene fossil form known from Wyoming and described by paleontologist Othniel Charles Marsh. Garrison’s tale turns towards a consideration of the necessity of language for moral sense, with the Houyhnhmns and their diminutive living fossil forms becoming an increasingly thinly veiled analogy for 19th century discourse around human evolution, language development, and civilizational progress. In the end, the sea subsumes the island and the civilization on it. Garrison’s story drew a tight connection between literary, Darwinian, philological and anthropological uses of living fossil. His own use is not incidental—he cites Darwin’s own passage and draws attention to the former isolation of the four-toed equines. The New Gulliver is kaleidoscopic in its encapsulation of the ways in which the popular metaphor and elements of Darwin’s concept were co-constitutive at the end of the 19th century.
Into the 20th century, use of the popular metaphor and scientific use of variations on Darwin’s living fossil concept expanded, and continue today (Lidgard and Love 2018, 2023; Lidgard and Kitchen 2023). The cultural evolutionists’ living fossil concept also persisted, at least into the early 20th century. The perceived connection between Indigenous Australians and Neanderthals, for example, was resilient and continued to be cited as justification for placing Indigenous Australians as the lowest, and therefore earliest form of humanity (Sollas 1911; Sommer 2005; Bowler 2021). In 1927, anthropologists Baldwin Spencer and F. J. Gillen depicted a direct comparison between living fossil peoples and animals in Australia. They wrote: “Australia is the present home and refuge of creatures, often crude and quaint, that have elsewhere passed away and given place to higher forms. This applies equally to the aboriginal as to the platypus and kangaroo. Just as the platypus, laying its eggs and feebly suckling its young, reveals a mammal in the making, so does the Aboriginal show us, at least in broad outline, what early man must have been like” (Spencer and Gillen 1927, p. vii). However, shifts in cultural anthropology were underway in the early 20th century. Global comparative judgments on cultures informed by evolutionary rankings gradually fell out of favor, displaced in part by ethnographic field studies of the details of individual societies, and a framework of cultural relativism established by Franz Boas in American institutions (Stocking 1982; Candea 2018). While the core scientific program of the cultural evolutionists faltered, some aspects of racialism lingered (Teslow 2021; Caspari 2023). The stories of the Darwinian living fossil concept, the cultural evolutionists’ concept, and the popular metaphor, therefore diverge further in the periods after the focus of our paper, and certainly warrant their own telling in future studies.
Our historiographical intervention, so to speak, is a reinterpretation of living fossil, elevating its stature as an object of study in its own right. In the main, previous studies have underemphasized how Darwin’s development of a living fossil concept was integral through his stages of evolutionary theorizing, as we have shown here. It functioned in Darwin’s argumentative strategy for Origin as an explanatory concept and a “how possibly” narrative (Lennox 2008), inverting a presumptive counter-example (long-persisting and unchanging forms) to demonstrate the effectiveness of Darwin’s causal evolutionary processes. Darwin’s concept persevered as other 19th century natural historians took up living fossils in their research, but some of Darwin’s intentional properties were narrowed or splintered. This reinterpretation offers insight into both the construction and subsequent usage of the concept across scientific disciplines. Many, perhaps most, modern biologists and paleontologists use particular cross-cutting properties or criteria to categorize certain taxa as living fossils (Sterner 2023), often without much consideration of Darwin’s actual intentions and practice, or the elusive variations in meaning that began in the 19th century and proliferated thereafter (McNiven and Russell 2005; Lidgard and Love 2018; Mandler 2020; Lidgard and Kitchen 2023).
In our comparisons of living fossil in Darwin’s work, the cultural anthropologists’ work, and popularly, we illuminated how different readings of evolutionism coexisted in the last half of the 19th century. Possibilities for identifying and construing change and changelessness were pursued for divergent aims. Yet across domains, the appearances of living fossil also reveal an interconnectivity between change and changelessness that is not exclusively oppositional. While previous historical studies exposed ambiguities between Darwin’s description of evolution and living fossils in Origin, his views on human racial hierarchy, his connections to cultural evolutionists and his conflicts with their inherited, presupposed stadial theories, they have seldom considered nomadic transitions of the living fossil metaphor between social and scientific domains as part of a larger whole. Our study discerns how concepts and metaphors can be parts of different discursive fields that overlap every now and again. With living fossils, we show that concepts and metaphors transform and can be separated between different domains and contexts or can undergo lesser transitions while persisting nevertheless over time.
Our interpretations are largely structured around linear or normal time, but other possibilities exist. For example, consider Reinhart Koselleck’s theoretical framework of “layers of time,” denoting multiple passages of time that reveal different tempos of change “co-present in any given historical moment” (Zammito 2015, p. 199; Koselleck 2018). Living fossil in the 19th century operated as a counter to dominant synchronic and certain diachronic views of time, centered on an expectation of ongoing change and progressive development. All our actors are plausibly looking backward and invoking a long-term layer of time, even when their direct, intentional applications of a living fossil concept or metaphor are synchronic in the moment. Multiple layers of time were perceived directly as European explorers were contacting so-called backward societies (Koselleck 2002; Ankersmit 2021; Beriain et al. 2024) and cultural evolutionists framed the latter as changeless human living fossils. Our actors’ expectations also accord with the idea that development and progress had been universalized through a “temporalization” of history by the mid-19th century (Koselleck 2002; Pankakoski 2020, p. 77).
It is somewhat of a scholarly tradition to parse what in Darwin’s wider world may have influenced his own idea formation and what instead he may have influenced with his theorizing. We contend that Darwin’s living fossil concept contributed uniquely to his own concepts of natural selection and divergence. Broadly, living fossil concepts in the 19th century interacted, indicative of Darwin’s own developing ideas on the natural world and reflective of his cultural milieu. Versions of Darwin’s scientific living fossil concept have persisted and diversified in the scientific lexicon for more than a century and a half, as living fossil has also continued to swim metaphorically through popular parlance—neither of them remaining static over time.
Acknowledgements
Our research benefited greatly from the resources provided by the Darwin Correspondence Project (https://www.darwinproject.ac.uk/) and Darwin Online (https://darwin-online.org.uk/). John Adolfo and Jessica Krzeminski offered thoughts on living fossils in 19th century deep sea exploration and literature, respectively. Emily Kern, Alan Love and Lynn Nyhart shared comments on versions of the manuscript, and our reviewers provided a number of constructive suggestions. We are grateful to all.
Author Contributions
S. L. and E. K. wrote and reviewed the manuscript.
Funding
No funding was received to assist with the preparation of this manuscript.
Data Availability
No datasets were generated or analysed during the current study.
Declarations
Conflict of interest
The authors have no conflicts of interest to declare.
Ethical approval
Not applicable.
Footnotes
We acknowledge that the terms scientist and scientific were not used uniformly during the 19th century (Ross 1962). Throughout the text, we use these terms in their modern senses for consistency and clarity.
Philosophical discussions of concepts as mental representations, abilities, and definitionally bounded abstract objects (e.g., Margolis and Laurence 2019) are largely outside our intended focus here.
Pterodactyls belong to the Order Pterosauria, an extinct group of flying reptiles that lived during the Mesozoic Era.
From W. E. Darwin, 5 October 1862, Darwin Correspondence Project, “Letter no. 3789F,” accessed on 6 May 2024, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-3789F.xml. Also published in The Correspondence of Charles Darwin, vol. 24 (Supplement); To Frank Hurndall, 20 September 1881, Darwin Correspondence Project, “Letter no. 13345,” accessed on 6 May 2024, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-13345.xml.
Jefferson’s disbelief in extinction may have informed his insistence of mammoths’ existence elsewhere, and though there is evidence that later in his life he acquiesced to the possibility of extinction, there is disagreement about such a change in his views. His mammoths are now acknowledged as mastodons. See “Letter from Thomas Jefferson to President John Adams, 11 April 1823,” in Cappon (1971, p. 592) for Jefferson’s admittance of extinction, and Rowland (2009) for a critical reading of this change.
The fraught history of racial and ethnic classification before and during the formative years of anthropology is beyond our scope here, but we return to certain elements of the cultural evolutionists’ classifying and theorizing—the comparative method, conjectural history, and stadial theories—in a later section. See for instance Müller-Wille (2014) and Vartija (2021) for perspectives on this history of racial classification.
We set aside the very deep history of contending logical and genealogical concepts of species, and of early ideas bearing on species evolution. See for instance Richards (2010) and Sloan (2019a).
Darwin, C. R. 1837–1838. Notebook B: [Transmutation of species], 25. CUL-DAR121. Transcribed by Kees Rookmaaker. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. 1837–1838. Notebook B: [Transmutation of species], 36. CUL-DAR121. Transcribed by Kees Rookmaaker. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. 1837–1838. Notebook B: [Transmutation of species], 28. CUL-DAR121. Transcribed by Kees Rookmaaker. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. 1837–1838. Notebook B: [Transmutation of species], 42-43. CUL-DAR121. Transcribed by Kees Rookmaaker. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. 1837–1838. Notebook B: [Transmutation of species], 97. CUL-DAR121. Transcribed by Kees Rookmaaker. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. 1837–1838. Notebook B: [Transmutation of species], 126e. CUL-DAR121. Transcribed by Kees Rookmaaker. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. 1838.02–1838.07. Notebook C: [Transmutation of species], 201. CUL-DAR122. Prepared by Kees Rookmaaker. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. 10.1838–7.1839. Notebook E: [Transmutation of species], 66 and 168e. CUL-DAR124. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/). Macleay (1819) noted the difficulty of proving affinities of Ornithorhynchus between groups that seem only distantly related, leaving the problem to the future.
Darwin, C. R. 4.1843. “Aberrant groups.” CUL-DAR205.5.60. Transcribed by Kees Rookmaaker, corrections by John van Wyhe. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/). The living species of echidnas are now placed along with Ornithorhynchus in the Order Monotremata, the only egg-laying mammals. The flightless kiwi, Apteryx, could possibly have represented an ancestral flightless condition in birds. However, he later concluded that Apteryx had descended from flying ancestors, and the wings were abortive (Stauffer 1975).
Darwin, C. R. 1844.03.31. Swainson’s statement. CUL-DAR205.5.97. Transcribed by Christine Chua. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
From George Robert Waterhouse, [April 1844], Darwin Correspondence Project, “Letter no. 2026,” accessed on 5 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-2026.xml. Also published in The Correspondence of Charles Darwin, vol. 7 (Supplement).
Darwin, C. R. n.d. [Abstract of British Association meeting reports, 1846]. CUL-DAR74.110-115. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
To J. D. Hooker, 31 March [1844], Darwin Correspondence Project, “Letter no. 744,” accessed on 7 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-744.xml. Also published in The Correspondence of Charles Darwin, vol. 3; From H. C. Watson to J. D. Hooker, 12 April 1847, Darwin Correspondence Project, “Letter no. 1079,” accessed on 15 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-1079.xml. Also published in The Correspondence of Charles Darwin, vol. 4.
This transcription is from one accompanying a scan of the monograph in the Darwin library. https://www.biodiversitylibrary.org/page/33865499#page/2/mode/1up.
Darwin, C. R. 1851.12.04. It makes not the slightest difference in considering [Fuliala], or Stylops, or Sagitta, or Ornithorhynchus aberrant. CUL-DAR205.9.243. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/); From J. D. Hooker 5 December [1854], Darwin Correspondence Project, “Letter no. 1611,” accessed on 21 March 2024, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-1611.xml Also published in The Correspondence of Charles Darwin, vol. 5.
To J. D. Hooker, 11 [December 1854], Darwin Correspondence Project, “Letter no. 1612,” accessed on 10 December 2021, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-1612.xml. Also published in The Correspondence of Charles Darwin, vol. 5.
Darwin, C. R. 1854.11. Assuming species approximately constant if extinction has fallen. CUL-DAR205.5.147. Transcribed by Christine Chua. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. 1856.09.25. The advantage in each group becoming as different as possible may be. CUL-DAR205.5.171. Transcribed by Christine Chua. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
To J. D. Hooker 8 [June 1858], Darwin Correspondence Project, “Letter no. 2282,” accessed on 7 March 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-2282.xml, Also published in The Correspondence of Charles Darwin, vol. 7; To Asa Gray 5 September [1857], Darwin Correspondence Project, “Letter no. 2136,” accessed on 7 March 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-2136.xml. Also published in The Correspondence of Charles Darwin, vol. 6.
The note is part of Charles Darwin’s Library: https://www.biodiversitylibrary.org/item/105730#page/510/mode/1up.
Darwin, C. R. 1857.11. The abnormal Insects & Birds on islds (plants too easily diffused)—the fishes & shells. CUL-DAR205.9.334. Transcribed by Christine Chua. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Philology, the study of the historical development, structure, and relationships of languages, transformed rapidly in the 19th century, expanding from its classical roots to encompass a greater diversity of living and largely extinct languages, and attaining a degree of scientific and societal influence.
Darwin, C. R. n.d. Though time is most important element as giving many individuals. CUL-DAR205.3.194-196. Transcribed by Christine Chua. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
To J. D. Hooker, 24 December [1858], Darwin Correspondence Project, “Letter no. 2384,” accessed on 3 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-2384.xml. Also published in The Correspondence of Charles Darwin, vol. 7.
To Charles Lyell, 11 October [1859], Darwin Correspondence Project, “Letter no. 2503,” accessed on 7 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-2503.xml.
Darwin later defends the position that his case for living fossils is consistent with the process of natural selection in a letter to Charles Lyell. To Charles Lyell, 12 September [1860], Darwin Correspondence Project, “Letter no. 2915,” accessed on 5 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-2915.xml. Also published in The Correspondence of Charles Darwin, vol. 8.
Darwin, C. R. and Emma Darwin n.d. [Abstract of Quarterly Journal of the Geological Society of London, 1851-.] CUL-DAR75.18-28. Transcribed by Christine Chua. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. [1847]. Abstract of Miller, First impressions of England. CUL-DAR205.9.196. Transcribed by Christine Chua. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/).
Darwin, C. R. n.d. Abstract of Murray, Proceeding of Bot. Soc. of Edinburgh Nov. 10th—1859. CUL-DAR205.3.219. Transcribed by Christine Chua. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/.
To Charles Lyell, 10 January [1860], Darwin Correspondence Project, “Letter no. 2647,” accessed on 3 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-2647.xml. Also published in The Correspondence of Charles Darwin, vol. 8.
To J. D. Hooker, 23 October [1861], Darwin Correspondence Project, “Letter no. 3296,” accessed on 3 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-3296.xml. Also published in The Correspondence of Charles Darwin, vol. 9.
To J. D. Hooker, 18 [December 1861], Darwin Correspondence Project, “Letter no. 3346,” accessed on 6 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-3346.xml. Also published in The Correspondence of Charles Darwin, vol. 9.
Tragulidae is a small family of artiodactyls including mouse deer.
Hoatzins (Opisthocomus hoazin) have wing claws when young, an atypical digestive system, and other obscure character relationships.
Cystoidea is an extinct class of crinozoan echinoderms known from earlier periods of the Paleozoic.
Ichthyosauria is an order of extinct marine reptiles that lived during the Mesozoic Era.
Specimen evidence held weight for some naturalists in adjudicating claims of living fossil sightings, such as living Pleisiosaurus, in the 19th century. Lyell’s own interest in the topic was rebuffed by Richard Owen’s rejection of any claims without physical evidence. Others would invoke native lore as support for the existence of creatures otherwise thought extinct. The lasting differential weight given to these forms of support is evident in the controversial cryptozoology space, formalized in the 1950s. For more on these connections, see Fallon (2024).
From Oswald Heer, 28 September 1875, Darwin Correspondence Project, “Letter no. 10175,” accessed on 11 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-10175.xml. Also published in The Correspondence of Charles Darwin, vol. 23.
From Gaston de Saporta 16 December 1877, Darwin Correspondence Project, “Letter no. 11281,” accessed on 6 April 2024, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-11281.xml.
To Gaston de Saporta, 24 December 1877, Darwin Correspondence Project, “Letter no. 11287,” accessed on 11 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-11287.xml.
To J. D. Hooker, 24 December [1858], Darwin Correspondence Project, “Letter no. 2384,” accessed on 3 February 2023, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-2384.xml. Also published in The Correspondence of Charles Darwin, vol. 7.
Prichard considered climate in human extinctions, but both he and Darwin largely implicated European expansion. Darwin considered it an extreme development of otherwise common competition between tribes. Prichard wrote that “[w]herever Europeans have settled, their arrival has been the harbinger of extermination to the native tribes. Whenever the simple pastoral tribes come into relations with the more civilised agricultural nations, the allotted time of their destruction is at hand; and this seems to have been the case from the time when the first shepherd fell by the hand of the first tiller of the soil” (Prichard 1839, p. 496). And Darwin during his Beagle voyage wrote this: “[w]herever the European has trod, death seems to pursue the aboriginal…We may look to the wide extent of the Americas, Polynesia, the Cape of Good Hope and Australia, and we find the same result. Nor is it the white man alone that thus acts the destroyer. […] The varieties of man seem to act on each other in the same way as different species of animals—the stronger always extirpating the weaker” (Darwin 1839, p. 520).
The history of interactions among different intellectual streams in cultural and physical fields of anthropology is convoluted (Reybrouck 2012; Manias 2013; Turner 2014). We necessarily omit much of this history, including concerns over the frequency of diffusion (horizontal transfer) of words and customs among cultures (for more on this see Kressing 2013; Candea 2018).
In opposition to these generally progressive unilinear views were accounts of Greek and Roman decline, finally overcome with the Renaissance and its efforts to match the achievements of the ancients. Perhaps more significant for our analysis were biblically derived frameworks of relative degeneration, in which cultures had started in an advanced state and some had fallen into primitivism faster or further than others. For cultural evolutionists, however, primitive groups needed explanation for their static nature, or regression, not an accelerated descent.
Darwin, C.R. The Beagle diary of Charles Darwin (1831–1836), p. 426. Edited by Kees Rookmaaker and John van Wyhe. EH88202366. http://darwin-online.org.uk/.
Darwin, C.R. The Beagle diary of Charles Darwin (1831–1836), p. 774. Edited by Kees Rookmaaker and John van Wyhe. EH88202366. http://darwin-online.org.uk/.
Royer took a clearly racialist posture in the preface of her translation, drawing on natural selection as a mechanism by which races progressively supplant one another, resulting in a true inequality between “superior” and “inferior” races. Brownen Douglas (2022) argued that this was quite counter-Darwinian in its reliance on a law of progress. See also Bernasconi (2010).
To William Graham 3 July 1881, Darwin Correspondence Project, “Letter no. 13230,” accessed on 18 May 2024, https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-13230.xml.
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