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Phylogenetics and Paleontology at the Dawn of Evolutionary Biology

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Phylogenetics and Paleontology at the Dawn of Evolutionary Biology Evo Edu Outreach (2010) 3:576–584 DOI 10.1007/s12052-010-0280-7 ORIGINAL SCIENTIFIC ARTICLE Brocchi, Darwin, and Transmutation: Phylogenetics and Paleontology at the Dawn of Evolutionary Biology Stefano Dominici & Niles Eldredge Published online: 30 September 2010 # Springer Science+Business Media, LLC 2010 Abstract Giambattista Brocchi’s(1814) monograph (see death in 1827. The notion that new species replace older, Dominici, Evo Edu Outreach, this issue, 2010) on the extinct ones—in what today would be called an explicitly Tertiary fossils of the Subappenines in Italy—and their phylogenetic context—permeates these essays. Herschel’s relation to the living molluscan fauna—contains a theoret- (1830) discussion of temporal replacement of species and ical, transmutational perspective (“Brocchian transmuta- the modernization of faunas closely mirrors these prior tion”). Unlike Lamarck (1809), Brocchi saw species as discussions. His book, dedicated to the search for natural discrete and fundamentally stable entities. Explicitly anal- causes of natural phenomena, was read by Charles Darwin ogizing the births and deaths of species with those of while a student at Cambridge. Darwin’s work on HMS individual organisms (“Brocchi’s analogy”), Brocchi pro- Beagle was in large measure an exploration of replacement posed that species have inherent longevities, eventually patterns of “allied forms” of endemic species in time and in dying of old age unless driven to extinction by external space. His earliest discussions of transmutation, in his essay forces. As for individuals, births and deaths of species are February 1835, as well as the Red Notebook and the early understood to have natural causes; sequences of births and pages of Notebook B (the latter two written in 1837 back in deaths of species produce genealogical lineages of descent, England), contain Brocchi’s analogy, including the idea of and faunas become increasingly modernized through time. inherent species longevities. Darwin’s first theory of the Brocchi calculated that over 50% of his fossil species are origin of species was explicitly saltational, invoking still alive in the modern fauna. Brocchi’sworkwas geographic isolation as the main cause of the abrupt reviewed by Horner (1816) in Edinburgh. Brocchi’s appearance of new species. We conclude that Darwin was influence as a transmutational thinker is clear in Jameson’s testing the predicted patterns of both Brocchian and (1827) “geological illustrations” in his fifth edition of his Lamarckian transmutation as early as 1832 at the outset of translation of Cuvier’s Theory of the Earth (read by his his work on the Beagle. student Charles Darwin) and in the anonymous essays of 1826 and 1827 published in the Edinburgh New Philo- Keywords Giambattista Brocchi . John Herschel . Charles sophical Journal—which also carried a notice of Brocchi’s Darwin . Transmutation S. Dominici (*) Museo di Storia Naturale, Sezione di Geologia e Paleontologia, Università di Firenze, It was widely accepted among savants [i.e. in the via La Pira 4, 1830s], even in Britain, that some kind of natural 50121 Florence, Italy process, as yet unknown, must be responsible for the e-mail: [email protected] origin of new species (M. Rudwick 2008). N. Eldredge In this Special Issue of Evolution: Education and Division of Paleontology, Outreach devoted to the topic of phylogenetics, we explore The American Museum of Natural History, Central Park West @ 79th St, the early history of transmutational thinking, emphasizing New York, NY 10024, USA the expansion of systematics from its foundational role in Evo Edu Outreach (2010) 3:576–584 577 the recognition (and consequent classification) of natural that nature usually leaves us nothing but minute, nay groups of “allied forms,” into a full blown search for puerile, details on which to found our distinctions” (Elliot patterns of genealogical descent—a pursuit initially con- 1984). That claim was the very hallmark of Lamarck’s fined to studies of the fossil record and in particular to the transmutation. scientific explanation of the origin of the modern fauna (see Lyell, alluding to Lamarck, put the problem in the also Eldredge 2010, this issue). This paper augments the opening page of his second volume very well as he invited historical background that led to Darwin’s initial work on his reader to: transmutation while serving as unpaid ship’s naturalist Inquire first, whether species have a real and while on the HMS Beagle from 1831 to 1836 as initially permanent existence in nature; or whether they are (and in greater detail) described by Eldredge (2009a). A capable, as some naturalists pretend, of being indef- companion paper (Dominici 2010, this issue) provides for initely modified in the course of a long series of the first time an English translation of the key components generations (1832). of Brocchian transmutation as originally published in Brocchi (1814). For Lamarck, species are spatiotemporally neither Jean-Baptiste Lamarck stands as the premier icon of pre- discrete nor stable. The counter claim—that species indeed Darwinian transmutationism. Debates in Great Britain (and are discrete and stable—is of course a foundational tenet of elsewhere) in the 1820s and 1830s over “transmutation” the religiously based view of creation, the very opposite generally centered around Lamarck’s ideas (and to a lesser of transmutation. As the creationist natural philosopher extent those of his younger colleague Etienne Geoffroy St. Whewell (the man who coined the term “science”) put it, Hilaire). Those adamantly opposed to transmutation, like “Species have a real existence in nature and a transition William Whewell for example, generally focused on from one to another does not exist” (Whewell 1837; Hull attacking Lamarck (Whewell 1837). And Lyell’s(1832) 1973; [in a later edition “transition” is changed to second volume of his Principles of Geology (see Secord “transmutation”; Whewell 1858]). 1991), while quietly leaving the door open for some But there was a third alternative—a form of transmuta- explanation of the origin of species in natural causal terms tion that saw species as discrete and stable, yet connected eventually to be discovered a century or two down the road, through successions of births and deaths forming lineages was primarily a no-holds-barred attack on Lamarck’s ideas. of ancestry and descent of “allied forms”—all attributable to In like manner, those who favored transmutation tended natural causes. Thus descendant species replace extinct ones to agree explicitly with Lamarck—the best example lying through time. This line of thinking was fully naturalistic and with the “Edinburgh Lamarckians” (Secord 1991), such as scientific—a second, explicitly non-Lamarckian, form of Darwin’s mentor Robert Grant, the geologist Robert transmutation that was originated by the Italian geologist Jameson (who may have penned the pro-Lamarckian and paleontologist Giambattista Brocchi—whoseworkwas anonymous essay of 1826 printed in his own journal widely read, discussed and, in some quarters, admired— [Anonymous 1826]), and others. The standard storyline is especially in Edinburgh. that Darwin succeeded where Lamarck ultimately failed If it was Brocchi who pioneered this second, separate because, with natural selection, which Darwin discovered in line of transmutational thinking, it was none other than 1838, Darwin had a plausible mechanism to explain Charles Robert Darwin who was its final champion while evolution—one that has clearly stood the test of time. on HMS Beagle (1831–1836) and for a time, after he Rather than comparing putative causal mechanisms for arrived back in England. While in South America, Darwin transmutation, we suggest that examining competing documented the replacement of an extinct by a modern empirically based claims of natural biological pattern congeneric species—and also compared allopatric distribu- reveals not one but two forms of transmutational thinking tions of what he always saw as discrete living species in in Great Britain in the 1820s and 1830s. Lamarck’s central southern South America, culminating in his recognition of claim—a prediction, in effect—is that if transmutation is levels of replacement patterns in the Galapagos avifauna. “true,” we should ultimately expect to document constant Tellingly, neither man was an admirer of Lamarck. change within species, such that a perfect data set of fossils would reveal complete intergradation between “species” through time; and we would as well expect to see similar Giambattista Brocchi patterns of smooth intergradation among living species geographically. Lamarck (1809) wrote “Let me repeat that Giambattista Brocchi was born in Bassano del Grappa in the richer our collections grow, the more proofs do we 1772, training as a historian and a mineralogist. In 1814, find that everything is more or less merged into Brocchi published Conchiologia Fossile Subapennina, the everything else, that noticeable differences disappear, and second important monograph on Tertiary molluscan pale- 578 Evo Edu Outreach (2010) 3:576–584 ontology after Lamarck’s(1802–1809) work on the inver- Brocchi’s text] and the study of modern molluscan tebrate fossils of the Paris Basin, published between systematics without comparison with their fossilized ante- 1802 and 1806 (Rudwick 2005). Finding more living cedents, writing: analogs in the modern seas (over 50%, Brocchi claimed) I agree that wanting to describe all the shells of the along the Italian coast
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