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Pattern Cladistics and the ‘Realism–Antirealism Debate’ in the Philosophy of Biology

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Abstract

Despite the amount of work that has been produced on the subject over the years, the ‘transformation of cladistics’ is still a misunderstood episode in the history of comparative biology. Here, I analyze two outstanding, highly contrasting historiographic accounts on the matter, under the perspective of an influential dichotomy in the philosophy of science: the opposition between Scientific Realism and Empiricism. Placing special emphasis on the notion of ‘causal grounding’ of morphological characters (sensu Olivier Rieppel) in modern developmental biology’s (mechanistic) theories, I arrive at the conclusion that a ‘new transformation of cladistics’ is philosophically plausible. This ‘reformed’ understanding of ‘pattern cladistics’ entails retaining the interpretation of cladograms as ‘schemes of synapomorphies’, but in association to construing cladogram nodes as ‘developmental-genetic taxic homologies’, instead of ‘standard Darwinian ancestors’. The reinterpretation of pattern cladistics presented here additionally proposes to take Bas Van Fraassen’s ‘constructive empiricism’ as a philosophical stance that could properly support such analysis of developmental-genetic data for systematic purposes. The latter suggestion is justified through a reappraisal of previous ideas developed by prominent pattern cladists (mainly, Colin Patterson), which concerned a scientifically efficient ‘observable/non-observable distinction’ linked to the conceptual pair ‘ontogeny and phylogeny’. Finally, I argue that a robust articulation of Antirealist alternatives in systematics may provide a rational basis for its disciplinary separation from evolutionary biology, as well as for a critical reconsideration of the proper role of certain Scientific Realist positions, currently popular in comparative biology.

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Notes

  1. Sokal and Sneath (1963) and Sneath and Sokal (1973) remain the definitive treatments of numerical taxonomy by its main proponents. Sneath’s “Thirty Years of Numerical Taxonomy” (1995) is also essential reading for understanding the posterior development of ‘phenetics’. Jensen (2008) is a recent reappraisal of the role that phenetics might continue to have in contemporary systematic biology.

  2. Despite its success in history and philosophy of science circles, the ultimate implications of Hull’s book were themselves the subject of strong criticism, not long after the volume was published, by some of the scientists involved. The book review “Lord of the Flies: The Systematist as Study Animal”, by the prominent cladists James Farris and Norman Platnick (Cladistics 5: 295–310 (1989)), was certainly not too sympathetic to Hull’s effort of analyzing communities of workers in systematic biology as groups whose ideas are subject to Darwinian selection processes. Farris and Platnick asked themselves “what Hull’s conclusions would be like had he been conversant with the history of the systematics of even a single group of organisms”, declaring at the end of their review that “in short, Hull has produced a flawed (and sometimes despicable) book that systematists will peruse mostly for the “gossip” it contains”. In virtue of his participation in the appearance of the cladistic viewpoint known in the literature as ‘transformed’ of ‘pattern cladistics’ (the main subject of this article), Platnick’s work is cited in several places across the piece. Farris’s interesting opinions about pattern cladistics at different stages of its development (see, for instance, Farris 1985; Kluge and Farris 1999) will only be mentioned briefly (see Note 30).

  3. Williams and Ebach (2007; Chap. 1, Sect. 1, “The Evolution of “Cladistics””) include a detailed reconstruction of the ways in which the word cladogram was used both by pheneticists and cladists, as well as by some ‘evolutionary systematists’, during the 1960s and 1970s (Hull’s account on the subject is included in Chap. 7, “Down with Cladism–Long Live Cladism”, of his 1988 book). The consequences of those usages are also reviewed by Williams and Ebach, within a rich, refreshing historiographic context. The importance of their book, particularly in virtue of its critical interplay with the discourse implicit in Hull’s volume (see also the Introduction in Williams and Ebach’s article included in the present compilation), is briefly considered in subsequent sections of this paper.

  4. Hull (1988) places his ‘baptism’ of ‘transformed cladistics’ at the banquet address presented during the first meeting of the Hennig Society (in Lawrence, Kansas, where the first Numerical Taxonomy Conference had taken place 13 years earlier). According to his own account, Hull made such reference ‘in an allusion to (a) remostrance he had received” from Farris (see Note 3). Hull adds, in justification to his analogy and in line with the main evolutionism-oriented, sociology of science-related argument of his book, that “just as a character can become modified successively through time in biological evolution, scientists can also gradually change their minds” (op. cit., p. 191).

  5. Hull (1988, pp. 246–247) mentions that disagreements between Nelson and two of his colleagues at the AMNH, Joel Cracraft and Niles Eldredge, resulted in the suspension of plans to write a joint book on the ‘principles of comparative biology’ and in the subsequent, separate publication of a text with Eldredge and Cracraft as authors (Phylogenetic Patterns and the Evolutionary Process, in 1980). Hull also makes reference to the publication of Phylogenetics: The Theory and Practice of Phylogenetic Systematics, by Edward Wiley (1981). Despite retaining some key agreements with the views of Nelson and Platnick, it is not difficult to see that the volumes authored by the latter systematists already departed from a ‘hard-core’ version of ‘transformed cladistics’. Although the three authors have preserved interests in theoretical aspects of systematics throughout their careers, I also maintain that neither of them has been significantly involved in published controversies concerning transformed cladistics beyond the early 1990s.

  6. In his “Philosophy and the transformation of cladistics revisited” Platnick (1985) stated that “one of the conceptual clarifications that has characterized cladistics since the time of Hennig (1966) concerns the relationship between cladograms and phylogenetic trees; whereas Hennig tended to view the two as identical, more recent workers have viewed cladograms not simply as trees but rather as sets of trees” (op, cit., p. 91). At this point in time, Platnick considered that this was one of the main points of agreement across the entire cladistics community (see Note 5). Such opinion seems to have played a role in his (and Nelson’s, according to Hull) ‘denial’ of the existence of ‘transformed cladistics’ (“when Nelson began to publish, he objected to cladism being referred to as a “school”. Cladism did not exist except in the minds of certain anti-cladists. Later, he and Platnick were equally insistent that pattern cladism did not exist”; Hull 1988, p. 238). Williams and Ebach (2007) certainly accept the ‘reality’ of pattern cladistics—more properly, though, they claim that pattern cladistics is the ‘true Cladistics’ (written with capital ‘C’; “what we understand Cladistics to be; in fact what we understand Systematics to be”; op. cit., p. 87).

  7. Hull (1988, Chap. 4) presents a useful discussion of the controversies surrounding the use of the term monophyly, by proponents of the three main systematic ‘schools’.

  8. After his remark on the ‘non-existence’ of transformed cladistics (see Note 6), Hull added the following: “One thing is certain: for not existing, (the conceptual modification of Hennig’s phylogenetic systematics) has garnered its share of names—canonical cladism, Cladism with a capital “C”, transformed cladism, methodological cladism, pattern cladism, natural order systematics, and modern cladism” (references to articles where these names are mentioned, according to Hull, have been omitted).

  9. With respect to Popper’s own judgements on evolutionary theory, Hull reminds the reader that, shortly after the introduction of the falsifiability principle, “Popper added Darwinism to his list of pseudo-scientific theories only to remove it still later” (Hull 1988, p. 342). Popper’s works and secondary literature about his philosophical views has been cited extensively by systematists for decades (see Note 10). The discussion about a/the ‘philosophical foundation of molecular systematics’ between Helfenbein and DeSalle (2005) and Faith (2006) is also a good source of Popper-related works. (See also the exchanges about Popper and systematics included in volume 50 of Systematic Biology (2001)).

  10. Nelson’s 1978 paper, “Ontogeny, phylogeny, paleontology and the biogenetic law” (critically revisited by Williams and Ebach 2003) undoubtedly shows a strong interest in Popper’s falsificationist principles. Oddly, Williams claims (personal communication) that the Popperian element in Nelson’s work is superfluous. Platnick and Gaffney’s three-part review of Popper’s books (and related publications) appeared within volumes 26 and 27 (corresponding to years 1977 and 1978, respectively) of Systematic Zoology.

  11. Hull (1988, p. 247) reports that “Popper was so impressed” with the reviews that Platnick and Gaffney had prepared about his work that “he wrote Platnick to compliment him on the understanding of his philosophical views that these reviews revealed”. However, like the prestigious philosopher of science Elliot Sober (whom has also never shown much sympathy for ‘transformed cladistics’; see, for instance, Sober 2000, p. 186), Hull has consistently stated that hypothetico-deductivism has no valid application in cladistics (Hull 1983, 1988, 1989). Olivier Rieppel’s recent criticisms of Popperianism in systematics are partially indebted to Hull’s views (see, for instance, Rieppel 2003, 2008; further comments on some of Rieppel’s important contributions to current philosophy of systematics are mentioned elsewhere in this article). It is worth noting that Williams and Ebach (2007, p. 113) also find no merit in advocating Popperian views in (C/c)ladistics (see Note 10).

  12. Chapter 11 in Williams and Ebach (2007; “Character Conflict”) is a very useful discussion of this subject, that simultaneously displays some of the conceptual/methodological characteristics associated to these authors’ historiographic treatment of (C/c)ladistics.

  13. Farris (1983) definitively solidified a specific interpretation of Popper’s notion of ‘explanatory power’ as the basis of the common usage of ‘parsimony’ in (automatized) cladistic analysis (and therefore its establishment as a key directive of ‘modern phylogenetic systematics’). Much of Arnold Kluge’s work (see his article in the present volume, for relevant references) is also concerned with this aspect. Besides the papers of his authorship cited in Note 11, Rieppel (2004) contains a lucid criticism of the notion of falsification as it has been used in (phylogenetic) systematics.

  14. This particular definition of Empiricism is based on the presentation to the section on ‘Scientific Realism’ in the useful introductory textbook by Balashov and Rosenberg (2002) on the philosophy of science.

  15. ‘Operationalism’ is the kind of Empiricism to which Sneath and Sokal make reference in their 1963 treatise (see, for instance, their elaborations on ‘operational homology’ in pp. 69–74). Pointing out to physicist Percy Bridgman as its initiator, Godfrey-Smith (2003, p. 30) defines operationalism as the view according to which scientists “should use language in such a way that all theoretical terms are tied closely to direct observational tests”.

  16. Named after Huxley (1940), the ‘New Systematics’—that is, the Evolutionary (Modern) Synthesis attempt to reform systematics by placing population-genetic assumptions at the base of classificatory practices—is treated (with approval) by Hull (1988; Chap. 3).

  17. The outstanding basic systematics textbook by Schuh (2000) uses this expression in its general discussion of ‘the schools of taxonomy’ (p. 8).

  18. The definition is taken from Statis Psillos’ fine defense of Scientific Realism. Psillos also describes Realism in terms of the realization of the fact that “successful scientific theories describe truly (or, near truly) the unobservable world (which) best explains why these theories are empirically successful” (Psillos 1999, p. 71).

  19. In the context of its upcoming appearance in the argument of the present paper, it is important to define the brand of Empiricism known as ‘Instrumentalism’—it is the view that advocates that ‘(scientific) theories are (nothing but) ‘instruments’ for prediction’. Psillos (op. cit., p. 17) additionally characterizes ‘eliminative instrumentalism’ (a stronger form of this type of empiricism) as the view that “theories should not aim to represent anything ‘deeper’ than experience, because, ultimately, there is nothing deeper than experience to represent”.

  20. As a philosopher, Brady worked out the key and most advanced arguments in support of pattern cladistics from a conceptual perspective (see main text). Patterson, on the other hand, was probably the most cautious systematist with respect to the import of philosophical arguments in cladistics (see Patterson 1982a, 1983). His influence was the main impulse for the development of cladistics in the United Kingdom. Unfortunately, both Brady and Patterson died when they were still active researchers (for a brief recollection of Patterson’s achievements, see Nelson 1998 and Forey et al. 2000).

  21. Patterson (1982b, p. 286) clearly saw the source of the disagreement between the two ‘viewpoints’ that, at the time, were about to split cladistics in two: “phylogenetic cladists appeal to the evolutionary process in order to justify their groups, and pattern cladists do not find this necessary”.

  22. Hull’s defense of the individuality thesis, including original citations, is developed in Chap. 6 (“Down with Darwinism—Long Live Darwinism”) of his 1988 book.

  23. In a review of the volume edited by Cracraft and Eldredge in 1979, Phylogenetic Analysis and Paleontology (Evolution 34: 822–823 (1980); the volume reviewed is the compilation where some of Gaffney’s work was published), Ghiselin noted that “I (…) wrote on the hypothetico-deductive method in phylogenetics and in evolutionary theory generally long before the cladists began to invoke it” and added that “the use of Popper’s refutationist philosophy has lately being seized upon as an ideological weapon by the cladists”. Walter Bock and Roger Miles have been also implied among the systematists responsible for bringing Popper’s ideas into systematics (see Helfenbein and DeSalle 2005, p. 273 and Williams and Ebach 2007, p. 112, respectively).

  24. The differences of opinion between Hull (1988) and Williams and Ebach (2007) on this subject are crucial (see Note 32). In my experience as a systematist, the label ‘pattern cladist’ still does function as a derogatory label. Mishler (1987), Felsenstein (2001) and Funk (2001) offer three additional (though not completely neutral), contextualized views on ‘pattern cladistics’.

  25. For an outstanding review of the semantic dimensions of ‘evolution’, see Robert Richards’ book The Meaning of Evolution (especially Chaps. 1 and 2). See also Brady (1994).

  26. The accounts on this issue presented by Hull (1988) and Williams and Ebach (2007) diverge widely.

  27. Among the unsympathetic reactions to pattern cladistics available in print (and besides those already noticed), Ridley (1986) and Dawkins (1986, Chap. 10) deserve brief mention. In my opinion, arguments presented in the former book are not philosophically sophisticated (E. O. Wiley’s judgement about Ridley’s work in The Quarterly Review of Biology (62:293–295 (1987)) is still worth reading, in virtue of his own statements about the subject). Dawkins’ opinions, on the other hand, are best understood as the result of a high degree of (neo) Darwinian entrenchment.

  28. This phrasing is evidently based on Stephen Jay Gould’s reference to a progressively non-pluralistic ‘Modern Synthesis’ (Gould 1983).

  29. The most succinct token of Hennig’s realism is probably the following: “transformation naturally refers to real historical processes of evolution” (Hennig 1966, p. 93). Again, consider that besides Hull’s and Williams and Ebach’s, multiple authors have provided their own (partially congruent, partially divergent) accounts of the separation between ‘phylogenetic systematics’ and ‘pattern cladistics’.

  30. In their criticism of the ‘three-taxon statement approach’ (see Williams and Ebach 2007, Chap. 12), Kluge and Farris (1999) practically assimilated phenetics to pattern cladistics. This exercise is historiographically important, for it represents the intention, from the side of two ‘faithful Hennigians’, to completely separate ‘correct phylogenetic systematic methodology’ from the ‘modernization’ of cladistic procedure attempted by two of the original pattern cladists (‘three-taxon statements’ were initially proposed in Nelson and Platnick (1991)).

  31. See the Introduction in Williams and Ebach’s article in the present compilation.

  32. Although Williams and Ebach’s book (2007) can be most adequately described as a continuation of the ‘historiographic space’ initially explored by Nelson and Platnick (1981), its main argumentative line responds directly to Hull’s intentions. This response is highly critical: “(Hull’s) account of the development of Phylogenetic Systematics has attracted much attention, yet his narrative seriously misrepresented progress in systematic biology during the 1900s, fictionalising the development of Cladistics and interpreting it from a neo-Darwinian point of view. Rather than an uninvolved and impartial historian, Hull seems to have held as an objective to demonise what he identifies as pattern cladistics (…). Hull embraces a transformational view of the world that in spite of his training as a philosopher and historian of science is an exceedingly narrow viewpoint (…). The history of Cladistics may be understood as more complex than Hull’s account, yet at the same time surprisingly straightforward in its achievements” (Williams and Ebach, op. cit., pp. 86–87).

  33. Accordingly, Williams and Ebach state that the situation which justified pursuing the reconfiguration of paleontology as a biological field of inquiry had to do with the position of privilege that the latter discipline held, for decades, in matters concerning systematics and evolutionary biology. They recall how the cause for this concern was eloquently pointed out by Patterson by quoting, in complete agreement, that “by about 1960 palaeontology had achieved such a hold on phylogeny reconstruction that there was common place belief that if a group had no fossil record, its phylogeny was totally unknown and unknowable” (Patterson 1987, p. 8; see also Williams and Ebach 2004, p. 687). Patterson’s opinion must be contextualized: systematics was experiencing the peak of the controversy between ‘molecules and morphology’ that followed the application of increasingly sophisticated sequencing technologies to the generation of nucleotidic data for cladistic (and non-cladistic, as well) analysis.

  34. Nelson interacted with Brundin during his postdoctoral visit in Stockholm. Patterson, in turn, knew about Hennig’s work through Brundin’s 1966 monograph, which was pointed out to him by Nelson when they coincided in London in 1967. See Williams and Ebach (2004) and Chap. 6 of their 2007 book for a thorough appraisal of Brundin’s role in the development of cladistics, and Hull (1988, mainly Chap. 4) for a less enthusiastic view about the importance of Brundin’s work in twentieth century systematics.

  35. A curious reference to ‘revolution’ can be found in Nelson and Patterson (1993) response to phylogenetic cladist Michael Donoghue’s review of Hull’s 1988 book (Biology and Philosophy 5:459–472 (1990)).

  36. This expression is used by Gee (2000, Chap. 4).

  37. A skeptic might ask, however, if metaphysical issues are really that important in comparative biology to start with. Without doubt, contemporary Darwinism places a strong emphasis in ontological concerns by attaching central importance to Darwin-Haeckel’s powerful metaphor of ‘the unique Tree of Life’. The importance of metaphysical issues vis-à-vis epistemology is critically examined in some of the pieces in the first section of the present volume.

  38. Here, I deliberately avoid compromising myself with a specific reconstruction of the chronology and reasons behind events corresponding to the onset of the ‘realist turn’. In my view, that would almost imply reconstructing the recent history of philosophy of science as a whole. The literature on Scientific Realism already cited provides a wealth of information highly relevant to the subject.

  39. It is noticeable, even amusing, that Shanahan (1992) implicitly assumes the separation between evolutionary biology and systematics. Without ever citing Brady, Shanahan accepts a sub-classification of the disciplines of comparative biology just like a pattern cladist would.

  40. I would argue that the reason why the R-AR debate has only been dealt with by just a few authors is one of the consequences of the aforementioned ‘hardening’ of process cladistics: the retention of Popper as the ‘ultimate source of philosophical grounding’ for systematic practice. This attitude has practically hindered the possibility of looking into the more recent period of discussion in philosophy of science as a source of elements to continue elaborating on the foundations of the discipline.

  41. It has been common to construe Scott-Ram’s 1990 book as a failed attempt to criticize transformed/pattern cladistics (see, for instance, the review by John Gittleman in The Quarterly Review of Biology 66:338 (1991) and the criticism published under the false name of “Aquiles Meo de la Torre” in Cladistics 7:201–212 (1991)). My own impression is that Scott-Ram’s analysis is much more subtle that Ridley’s, for example. Although I do not agree with the bulk of his conclusions, I appreciate Scott-Ram’s identification of a ‘theoretical’ versus a ‘descriptive attitude’ in systematics, his criticism of Popperianism, and his appeal to take “the rapidly expanding fields of molecular genetics and developmental biology” (op. cit., p. 184; italics mine) more seriously in comparative biology.

  42. The strong realist leanings of both authors is well known; nevertheless, it seems worth recalling that, for Boyd (1999)—who refers to pattern cladistics as ‘extreme cladism’ and to phylogenetic systematics (i.e. process cladistics) as ‘modest cladism’—quite simply “descent and ancestry are real” (op. cit., p. 183). Wilson’s volume also includes a contribution by phylogeneticist Kevin de Queiroz (1999), where a section on ‘realism and antirealism’ and ‘monism and pluralism’ is included. His standpoint—a straightforward realism with respect to species and higher taxa—might be considered representative of a large segment of the contemporary systematics community, not directly involved in philosophical and/or historiographic issues in comparative biology.

  43. Besides preserving the depth of understanding of scientific issues that would be expected from a practicing systematist, Rieppel’s extraordinary output during recent years on the philosophy of systematics has had more exposure to a wider audience (because of the journals in which it has been published) than Reif’s work.

  44. Rieppel’s early papers on the philosophy of systematics appeared in the late 1970s. His 1988 book (Fundamentals of Comparative Biology) has been essential reading in systematics courses in many countries since its publication.

  45. I refer here to those sources that, despite being actively employed by Hennig, have nevertheless been obscured by the increasingly narrow path that historiographies on cladistics have tended to take over the years.

  46. This is Olivier Rieppel’s own phrase (personal communication at the Mexico conference).

  47. It would be difficult to select a single, most representative quote of Rieppel’s ‘pattern cladistic stage’, but the period certainly extends well into the 1990s.

  48. Psillos (1999) and Godfrey-Smith (2003) provide brief and clear characterizations of these notions, central to the tension that has developed between R–AR in recent years. Godfrey-Smith (op. cit., p. 220) talks about the underdetermination of theories by evidence (UTE) in the following terms: “Empiricists argue that there will always be a range of alternative theories compatible with all our evidence. So we can never have good empirical grounds for choosing one of these theories over others and regarding it as representing how the world really is. If we have no empirical grounds for such a choice, then we have no grounds at all”. The pessimistic meta-induction is described in the same book (p. 239) as “an argument against some forms of scientific realism. The argument holds that theories have changed so much in the history of science that we should not have much confidence in our current theories. In the past, scientists have often very confident that their theories were true, but (…) they usually turned out to be wrong. So we should expect the same from our own current theories.” Finally, the ‘no miracles’ argument (NMA; Psillos 1999) corresponds to the very definition of Scientific Realism introduced in previous sections of this paper: “What needs to be explained, the explanandum, is the overall empirical success of science. NMA intends to conclude that the main theses associated with scientific realism, especially the thesis that successful theories are approximately true, offer the best explanation of the explanandum” (Psillos, op. cit., p. 71).

  49. Besides a number of specialized papers published in refereed philosophical journals, two edited volumes attest to the influence of van Fraassen’s work in contemporary philosophy of science—Churchland and Hooker (1985) and Monton (2007). See also van Fraassen (1989, 2002).

  50. This is van Fraassen’s original, rather elegant proposal of ‘constructive empiricism’: “Science aims to give us theories which are empirically adequate; and acceptance of a theory involves as belief only that it is empirically adequate. This is the statement of the antirealist position I advocate; I shall call it constructive empiricism” (van Fraassen 1980; Chap. 1.3).

  51. ‘Taxic’ versus ‘transformational homology’ is a widely used distinction in philosophy of systematics, roughly equivalent to the ‘pattern versus process’ contrast. I have decided not to use it much in this paper, though, to avoid duplicity of terminology when speaking about the ‘transformation of cladistics’. Besides Rieppel (1988a), Williams (2004) is highly recommended for further information on the history and meaning of the distinction.

  52. As a matter of fact, Rieppel’s output during his ‘pattern cladistics stage’ had already included papers specifically dealing with empiricism and the ‘reality’ of groups (see Rieppel 1988b, in particular). It is intriguing that he does not mention them very much in the context of his current discussions of instrumentalism and realism.

  53. “Today, one might still understand the “problem of homology” (…) to reside in the never-ending debate between the “historical” and “causal-analytical” schools (Spemann 1915); the former now residing in the discipline of numerical phylogenetics (phenetics), the latter resurrected and given the new name, “evolutionary developmental biology”” (Williams and Ebach 2005, p. 366). Notice how these authors see both numerically oriented ‘phylogenetics’ and ‘evo-devo’ as ‘Enemies of Homology’, without realizing the link between Patterson’s inclinations towards ontogeny as ‘positive’ evidence in systematics—under pattern cladistics assumptions—and Rieppel’s ‘causal grounding’—the ‘devo’ part in ‘evo-devo’. See Note 55 and the final two sections of this paper for further elaboration.

  54. Quentin Wheeler, a major contemporary phylogenetic cladist, has had this to say about developmental biology in general (significantly, in the part of his review of the Milestones in Systematics volume (Williams and Forey 2004) that corresponds to Nelson’s chapter): “The complex characters of evolution that we set out to explain and understand are no longer studied in detail and are sidelined for more rapidly analyzable data that lacks such inherent interest and distracts from what is most intellectually exciting. As Nelson suggests, we have much yet to learn from a returned emphasis on character analysis and a cladistic revolution to complete. It is worth noting, too, that one of the most promising interfaces for character analysis is with developmental biology, quite possibly an intellectually richer place to focus molecular work so as to bridge the continued gap between our understanding of genotype and phenotype” (Wheeler 2005, p. 17).

  55. Karl von Baer’s 1828 ‘scheme of the progress of development’ (in vertebrates) is a ‘proto-cladistic’ representation of hierarchical relationships between ontogenetic stages of development, where nodes do not correspond to ancestors, but to progressively (as one moves towards the tips of the hierarchy) less general developmental states in increasingly more particularized morpho-anatomical elements (‘parts’) of the groups of organisms involved. As stated in the main text, developmental-genetic evidence might in principle be susceptible of codification into matrices, which could then be analyzed in search for congruent patterns.

  56. Criticisms to van Fraassen’s notions on this subject (see, for instance, Godfrey-Smith’s 2003 argument about observability versus detectability) are put aside here, in order to treat them in a forthcoming, more detailed analysis of the relevance of ‘constructive empiricism’ for systematics. “Constructive empiricism now” (2001) contains some of van Fraassen’s main responses to these criticisms.

  57. It should be noticed that when the developmental-genetic aspect of the causal grounding of morphological characters is considered, literally thousands of ‘molecular characters’ might be involved (e.g. sequences of homeobox genes and their ‘targets’). However, causal grounding is not exhausted by linear gene sequences alone; it evidently involves expression patterns, developmental timing of such expression, regulatory interactions within ‘networks of developmental genes’ and morphogenetic processes (that involve cell- and tissue-level interactions). The connections between causal grounding, ‘epigenetics’ and ‘developmental systems theory’ are surely a subject that also deserves treatment elsewhere.

  58. As stated in Note 6, Williams and Ebach’s historiography firmly holds on to pattern cladistics individuation, erecting it to the status of ‘eucladistics’ (notwithstanding Nelson and Platnick’s early negation of the existence of a separation within cladistics). See, however, Ebach et al. (2008).

  59. This is particularly the case with respect to Hull’s position: “(…) I find realist philosophies more satisfying than the nonrealist views now being explicated by such philosophers as van Fraassen (…) and (Nancy) Cartwright” (Hull 1988, p. 468).

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Acknowledgments

The initial stages of this project were supported by a Kew Latin American Research Fellowship (KLARF) I received between 2004 and 2005. During this period, I presented versions of this paper in seminars at the Natural History Museum (London) and the Royal Botanic Gardens, Kew (United Kingdom), and at the 2005 Biennial Meeting of the International Society for the History, Philosophy and Social Studies of Biology (ISHPSSB), celebrated in Guelph (Canada). I greatly acknowledge the invaluable help of David Williams, Malte Ebach, Olivier Rieppel, Kirk Fitzhugh, Paula Rudall, Richard Bateman, Michael Frohlich, Dennis Stevenson, Elena Álvarez-Buylla, Carlos López Beltrán, Jorge González Astorga, Fernando Nicolalde Morejón, Ronald Amundson, Rasmus Winther and Thomas Reydon in a number of aspects related, in one way or another, to the final contents of this paper, aspects which would be impossible for me to describe in full detail here. I also appreciate very much the interest shown by Bas van Fraassen in the early stages of my project. Last, but not least, I especially thank Rasmus and Thomas for the kind patience they showed during the final stages of the preparation of this volume. All mis-interpretations or mis-representations of the information I gathered from my colleagues and friends remain entirely my responsibility.

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Vergara-Silva, F. Pattern Cladistics and the ‘Realism–Antirealism Debate’ in the Philosophy of Biology. Acta Biotheor 57, 269–294 (2009). https://doi.org/10.1007/s10441-009-9083-z

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