Reproduced from: sci-hub.tw/10.1007/s12304-018-9337-8

Biosemiotics
https://doi.org/10.1007/s12304-018-9337-8

Agency, Meaning, Perception and Mimicry: Perspectives from the Process of Life and Third Way of Evolution

R. I. Vane-Wright^1,2

dickvanewright@btinternet.com

¹ Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, UK
² Durrell Institute of Conservation and Ecology, School of Anthropology and Conservation, University of Kent, Canterbury CT2 7NR, UK

Received: 12 May 2018 /Accepted: 8 November 2018/Published Online: 29 November 2018
© Springer Nature B.V. 2018

Abstract

The concept of biological mimicry is viewed as a ‘process of life’ theory rather than a ‘process of change’ theory—regardless of the historical interest and heuristic value of the subject for the study of evolution. Mimicry is a dynamic ecological system reflecting the possibilities for mutualism and parasitism created by a pre-established bipartite signal-based relationship between two organisms – a potential model and its signal receiver (potential operator). In a mimicry system agency and perception play essential, interconnected roles. Mimicry thus describes emergent biologically meaningful relationships based on synergy, and is not an object-based theory. Biosemiotics offers a particularly valuable discipline for analysing the dynamics and nuances of mimicry systems, and can thus pave the way for a better and more complete understanding of how mimicry has evolved in the past, and how it might evolve in the future—presented here with special reference to the need for an integrated, ‘third way of evolution’ approach to biological relativity. A revised definition of mimicry is proposed.

Keywords

Mimicry, Agency, Perception, Definition, Process of life, Synergism

En Avant

In physics there are only happenings, no doings … the agency that arises with life brings value, meaning and action into the universe … With agency, values have emerged … With value comes meaning … meaning in the sense of mattering to some entity.
Kauffman (2008: 72, 74)

Almost 40 years ago I offered the following reasonable (but flawed) definition of mimicry. Reasonable because it has been said to be a “widely used functional definition” (Dalziell and Welbergen 2016: 612; also Jamie 2017); flawed for reasons I will explain below:

Mimicry involves an organism (the mimic) which simulates signal properties a second living organism (the model) which are perceived as signals of interest by a third living organism (the operator), such that the mimic gains in fitness as a result of the operator identifying it as an example of the model
Vane-Wright (1980:4)

One of the problems with definitions coined for natural phenomena is that even reasonable ones have a ‘shelf-life’. Samuel Butler once described definition as the act of “enclosing a wilderness of idea within a wall of words”. And the problem with language is its cultural basis and bias – as discussed in various ways by Whorf (1942), Wittgenstein (1953), von Bertalanffy (1973), Bohm (1980) – and many others. Words and language have a strong and often limiting ‘top down’ influence on what we attend to and how we interpret our sensory input (perception), and thus our understanding or ‘creation’ of reality. Similarly, current understanding of a phenomenon strongly influences how and what words we use to describe it, with risks of circular reasoning. Our apperception is also constrained by our wider culture – as demonstrated, for example, by Goodwin (1994: 28 et seq.) for the arguably significant influences of Christian values and mythology on supposedly ‘objective’ Darwinism, Weismannism and neo-Darwinism.

But all is not lost. From time to time scientific understanding (‘normal science’) undergoes ‘turns’, changes, or even revolutions (Kuhn 1970). Even so, we do not always remember to update our definitions – or refuse to, as science often runs for long periods with more than one theory addressing the same ‘reality’ (with the possible exception of psychology, none more so than in the field of organic evolution). This can readily lead to misunderstandings and disagreements, not based on different notions of reality per se but subjective cultural differences, represented by those walls of words.

A recent book over 500 pages long includes more than 2000 references on the subject of mimicry, crypsis and adaptive resemblances in general (Quicke 2017). Nevertheless, this remarkable compendium has just one index entry for semiotics. The purpose of this paper is to explore what mimicry might be – and the ways in which it might evolve in light of the contemporary movement called ‘the third way of evolution’ (for a review, see Kull 2017). It will come as no surprise to readers of this journal that many of the changes from normal (more accurately dominant) evolution science have been anticipated within the discipline of biosemiotics. But before attempting this, I present a partial summary of certain developments relevant to this current “scientific turn in evolutionary theory” (Kull 2017:645).

Origins of Life

Influenced by Erkki Haukioja (see below) I have quite recently contended that, before trying to understand the processes of organic evolution (change), ideally we should enquire into existence – “the nature of life itself – the system that is to evolve” (Vane-Wright 2014a). The existence of life poses at least three interrelated questions –establishing what properties a system must have if it is to be considered living, the origin of such a system, and the processes entailed in continuation of such a living system.

I am not going to comment on the origins of living systems other than to suggest there are only two fundamental possibilities – either by divine intervention, or by spontaneous generation (Vane-Wright 2014a; I am not considering ‘artificial life’). Both means of origination could have happened more than once. An additional issue, from an Earth-centric viewpoint, is whether life was divinely created or arose spontaneously on our planet – or elsewhere, and somehow spread here subsequently. For my part I am content to accept that life as we know it, including our own, originated here on planet Earth by some form of spontaneous generation approximately four billion years ago – and, even if life originated in this way many times, extinction of all but one primal lineage has occurred, such that all living organisms now known to us form a monophyletic group derived from the last “universal common ancestor” (Theobald 2010). This all-encompassing group can be referred to as superdomain Biota (Vane-Wright, 2014).

Process of Living Theory (POL) Versus Change Theory

…“modern evolutionary theory actually consists of two parts – an explicit part concerning change and an implicit part concerning existence – the basic phenomena in the domain of POL [process of living] are seen in the light of the neo-Darwinian theory which emphasizes change. This is a potential cause of much confusion in evolutionary biology because the most serious problems actually seem to lie in the domain of POL and not in the domain of the theory of change.”

Some of the ideas encapsulated in the above quote from Haukioja (1982: 358) arguably go back to Immanuel Kant, and were richly explored in various ways by von Uexküll (1926). However, the modern form of this idea lies with Haukioja’s insight which, though in my opinion profound, has been almost totally ignored. This may be due to the unusual terms in which he presents his arguments – I think it fair to say his essay is largely ‘a difficult read’ (and apparently it was extremely difficult to write: Erkki Haukioja pers. comm. 2014).

Of course, widespread disregard might suggest the issue has no substance. However, a fundamentally similar thesis has been more recently expressed in a totally different manner by the well-known palaeontologist Niles Eldredge – in a work which, unlike most of his books, is also little-known and rarely cited. Thus he writes “According to the popular press … every bit of animal behaviour …boils down to passing genes along to the next generation. Forgotten in the process is the simple fact that animals need to eat simply to live” (Eldredge 2004:16). And more directly: “Is life about evolution, or is life about living?” (Eldredge 2004: 43). The same thread can also be found in von Bertalanffy (1973: 160): “…fundamental problems are ‘swept under the carpet’… selection, competition, and ‘survival of the fittest’ already presuppose the existence of self-maintaining systems; they therefore cannot be the result of selection” [emphases original].

Thus we see that Dobzhansky’s famous dictum “Nothing in biology makes sense except in the light of evolution” has been internalised well – thinking about life in other ways now rarely finds favour with mainstream evolutionary biologists! In this general context it is notable that Shapiro (2011: 127) points to the conflation of the problems posed by the origin of life, evidence for the reality of evolution, and what the nature of evolutionary change might actually be, “in the minds of the lay public (and also of scientists)”.

Haukioja (1982) considered various questions about basic biological phenomena as unwarranted projections of gene-centred neo-Darwinian orthodoxy. His analysis was based on developing a minimalist process of living theory (‘POL’) separate from a body of theory about the processes of change. This led him to a number of different interpretations to those derived from population genetics and ‘selfish gene’ theory. For example, challenging problems under neo-Darwinism for traits such as the persistence of sexual reproduction (Maynard Smith 1978), iteroparity versus semelparity (Ranta et al. 2000), and artificial versus natural selection (Haukioja 1982: 371 et seq.) simply fall away when viewed in the context of POL. Perhaps most radical (at the time) was a demonstration that this also applies to sociobiology: “it seems to be a completely open question whether kin-selection has any appreciable role in real life” (Haukioja 1982:368).

There are parallels here between Haukioja (1982) and the synergism hypothesis of Peter Corning (1983) – yet another related body of theory often neglected despite its independent introduction as ‘synergistic selection’ by no less a figure than John Maynard Smith (1982; Maynard Smith and Szathmáry 1995). According to Corning (2005; 2018; Corning and Szathmáry 2015), synergism at all levels in living systems creates emergent phenomena that have real consequences for evolution (including mimicry: see below). Or as Kauffman (1995: 207, 208) has stated: “Each organism lives in the niche created by the artful ways of other organisms” while “The coevolution of organisms alters both the organisms themselves and the ways organisms interact”. In this sense Affifi (2018: 22) asserts “Evolution is always co-evolution.”

Haukioja (1982:365–366) also pointed to other critical differences. POL emphasises the dynamic continuity or cessation of lineages through time, whereas evolution addresses the types of entities that proceed. Likewise, from a POL perspective, genes are tools for transporting individuals in time, not the reverse (or as Eldredge 2004:30 put it, “an egg is a hen’s way of making another hen”). Evolution is not any sort of ‘goal’ for an organism – and genes do not have independent existence other than as parts of whole, functioning organisms. Genetic similarity between parent and offspring, viewed from POL, is not the reason for reproduction: organisms reproduce in this way simply because there is no better strategy for long-term (geological) survival – and we study only those organisms that exist. In theory at least, different genes will have separate roles in maintaining organism functionality (sustaining the life of the individual) from those affecting the physiology of reproduction, and again those influencing the (ecologically appropriate) regulation of reproduction – and this needs to be acknowledged, at least in certain analyses (Haukioja 1982).

Haukioja’s analysis challenges us to consider what it is that changes. The overarching need of the individual organism is to stay alive. But no living system can last forever – at the very least, accidental death will eventually occur. Thus, as above, only those organisms that successfully produce offspring will continue to be represented through time (cf. Eldredge 2004:36 – “in the long term, without reproduction, life would have ceased billions of years ago”). This is where evolutionary change occurs. So it is not individuals that evolve (although they develop, grow, and undergo many changes throughout their individual lives – and some of these processes can and do have consequences for their offspring: e.g. Jablonka and Lamb 2005; Shapiro 2011; Noble 2013), in the sense of change through geological time, nor is it ‘species’ (taxa). What changes over successive time intervals is the collective “mean kind” (Haukioja 1982: 363) of the ancestor-descendant lineage. This can be construed as the process described by Bernhard Rensch and Julian Huxley as anagenesis – the gradual or stepwise “organisational modification” (Remane 1952, as translated by Zachos and Hossfeld 2010) across successive generations of a population through time – regardless of whether reproduction is sexual or asexual, whether or not speciation (cladogenesis) occurs, and whether or not the change is regarded as ‘progressive’ (although it must, overall, be adaptive).

Compare this with Hennig’s (1966: Fig. 4) diagram labelled “species cleavage”, which shows a sexually-reproducing line, with genealogical (tokogenetic) relationships indicated between successive generations of individuals, splitting as a result of an extrinsic barrier to become two separate lineages (nascent species). However, this cladogenetic process does not necessarily involve anagenesis (even though it will normally result in subsequent change due to various processes, including shifting patterns of coevolution as well as genetic drift sensu lato and natural selection). Consequently, maintaining life (POL) including reproduction, changes in the ‘mean kind’ of lineages through time (anagenesis), and separation of lineages through time (cladogenesis) should not be conflated under one theory – or at least, not one theory that interprets everything “in the light of evolution” acting at only one privileged level (e.g. genes, as in the case of conventional neo-Darwinism). In this spirit Noble (2012, 2015) has proposed we adopt “biological relativity”, and accept that there is no privileged, unique or final level of causation (e.g. DNA) – on the contrary, there is a need for integration of all relevant causal levels (including of course the molecular genetic level).

Over periods of time greater than the average lifespan of the individual organism, later members of a single lineage may come to be very different (e.g. in phenotype, genotype, and/or ecological relationships, etc., including shifts in metabolic and developmental interaction patterns due to holobiosis: Doolittle and Booth 2017) when compared with their progenitors. The significance of this for mimicry theory is that the processes that govern functional mimicry systems as part of the dynamic of staying alive should not be conflated with the processes affecting the origin of mimicry within a lineage, or subsequent changes to that mimicry system thereafter (evolution). But before looking at this from the ‘third way’ perspective it is necessary to say something about what the third way means: its origins, current status, and what it might become.

The Third Way of Evolution: Origins, Current Status, and Possible Future

Nature cannot be understood by pretending it is simple
Elton (1966:xi)

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The mechanist, starting from the physico-chemical standpoint, interprets the living thing by analogy with a machine. The vitalist, on the other hand, supposes a guiding entelechy, which summons order out of chaos; he thus adopts a dualistic attitude. The elements of truth in both these views are recognized, and their opposition is resolved in the organismal approach to the living creature. This approach is conditioned by the belief that the vital co-ordination of structures and processes is not due to an alien entelechy, but is an integral part of the living system itself. Arber (1954:100).

Origins

The third way approach to evolution has multiple origins, far too disparate and complex to review in any comprehensive fashion here. It is convenient to see in Immanuel Kant’s writing about self-organisation the origins of POL theory, leading eventually to the insights of Humberto Maturana and Francisco Varela. Jean-Baptiste Lamarck introduced the idea of evolution through the action or agency of the whole organism, and Alfred Wallace and Charles Darwin the idea of natural selection in an essentially ecological vision of evolution. The brief light of the organic selectionists (Douglas Spalding, Conwy Lloyd Morgan, Henry Osborn, James Baldwin, Edward Poulton), in my view the first wave of third way thinking, was soon displaced by the atomism of the mutationists – in the arena of mimicry Reginald Punnett being a leading example. Mutationism and natural selection were then brought together in what became neo-Darwinism, notably through the work of Ronald Fisher, John Haldane, Sewall Wright and, so it is often said, Ernst Mayr and Theodosius Dobzhansky – although there is a wider tradition certainly in the writings of these last two authors concerning the role of the whole organism, its purposiveness and behaviour. Meanwhile the work of Charles Elton and other ecologists led to a systems view of life (in which Ludwig von Bertalanffy was a major influence), culminating perhaps with James Lovelock’s vision of Gaia and the dynamic self-organising interconnectedness of the entire biosphere. Two more lines of thinking, via a revival of organic selectionism in the form of Conrad Waddington’s genetic assimilation, led to epigenetics and ‘evo-devo’ (within which movement Brian Goodwin was very significant) on the one hand, and synergism and the work of Peter Corning and the later work of John Maynard Smith on the other. Yet another strand was introduced by physicists Max Planck and Erwin Schrödinger seeking the foundation of order in living systems in both pre-existing order and in disorder. Considerations of disorder led to the introduction of thermodynamics to biological thinking (e.g. the work of Ilya Prigogine), while the need to consider pre-existing order paved the way for discovery of the genetic code and both a hardening of preformationist/reductive genetics on the one hand, but also a revitalisation of epigenetic and cell physiological approaches on the other (for example in Stuart Kauffman’s ideas on work cycles). Yet another line of thinking led from Jakob von Uexküll to Thomas Sebeok and the emergence of biosemiotics. By the end of the twentieth century there was a veritable maelstrom of seemingly different and potentially irreconcilable scientific approaches to the nature of life and its evolution, with a mood of uncertainty and disarray well captured, for example, by Dupré (1995). Many of these strands of thought are superbly summarised and synthesised in the work of Capra and Luisi (2014) and, most recently, Corning (2018). But there was also a clear need, if not for synthesis as such, of simply ‘bringing together’ (long ago beautifully presaged by botanist Agnes Arber, quoted above).

The “Third Way” Website

In May 2014 physiologist Denis Noble, geneticist James Shapiro and Neo-Lamarckian Raju Pookottil launched a website entitled The Third Way (2014). It first appeared with a manifesto stating that “The vast majority of people believe that there are only two alternative ways to explain the origins of biological diversity. One way is Creationism. .. The other way is Neo-Darwinism, which has elevated Natural Selection into a unique creative force that solves all the difficult evolutionary problems. Both views are inconsistent with significant bodies of empirical evidence and have evolved into hard-line ideologies. There is a need for a more open “third way” of discussing evolutionary change based on empirical observations.” [At the time of writing the current manifesto says much the same, but fails to make explicit what the ‘third way’ is.]

With the strapline “evolution in the era of genomics and epigenomics”, this website is pronounced “open to established scholars in the sciences, philosophy, history and related humanities who have published work related to The Third Way”. Scholars are invited to post brief statements of their interests, including short autobiographical notes and references to their key works. While the site managers make it clear that they do not “necessarily endorse any of them”, the aim being “simply to make new thinking about evolution available in one place on the web”, it is made explicit that “the website and scientists listed … do not support or subscribe to … inscrutable divine forces or supernatural intervention, whether they are called Creationism, Intelligent Design, or anything else.”

By mid-2018 some 64 scholars had entries (sad to say, one of them already deceased) – a small fraction of the numerous researchers who would currently subscribe to some or even many of the views expressed. However, what is perhaps most striking is how difficult it is to pigeon-hole almost any of those listed into single, traditional scientific disciplines. The sheer discipline diversity is also notable: physics, biophysics, molecular biology, physiology, genetics, genome biology, neo-Lamarckism, plasticity, virology, microbiology, epigenetics, development, parasitology, niche construction, behaviour, semiotics, psychobiology, complex systems, integrative biology and philosophy of science.

Possible Future

Several of the ‘third way’ protagonists played a significant role in organising the 3-day joint meeting of the Royal Society and British Academy entitled “New trends in evolutionary biology: biological, philosophical and social science perspectives” held in London, November 2016 (see Kull 2017 for a review). In future it seems possible that this movement, for want of a better word, might spark a coherent ‘third way synthesis’, perhaps a twenty-first century successor to the twentieth century ‘synthetic theory’ – but as yet that seems a long way off, and possibly not even the intention at all – perhaps accepting or even embracing Dupré’s (1995) “disunity of science”. And of course, in true Kuhnian tradition, there is great resistance to change from the majority still operating in the dominant paradigm, often said to be “just fine” (e.g. Williams 2015). For now the only commonality is that of scholars who, while passionately interested in evolution as part of the natural sciences, are significantly, even deeply dissatisfied with the reductionism of neo-Darwinian orthodoxy. So there is as yet no single Third Way perspective – and perhaps may never be. If there is a current common thread it is to do with self-organisation and the multiplicity of different factors acting at different levels of organisation that are at play in the unfolding of life. And this is celebrated: “The divergences and multiplicity of ideas, opinions and theories on this website are necessary since many fields of evolutionary biology remain relatively unexplored.” And as already noted, it has also found a name: biological relativity (Noble 2012). Briefly stated, biological relativity avoids “the assumption that there is a privileged scale at which biological functions are determined” (Noble 2012: 55), and requires “integration of a variety of mechanisms of evolutionary change that must interact, rather than the single mechanism postulated by the Modern Synthesis” (Noble 2015:7).

The Role of Agency in Life and its Evolution

A shift into a new niche or adaptive zone is, almost without exception, initiated by a change in behaviour. Mayr (1963:604)

The idea that the behaviour of organisms is a major factor in evolution goes back at least to the often unfairly criticised J.-B. Lamarck. More modern expressions can be found in the writings of at least one of the ‘architects’ of neo-Darwinism, Ernst Mayr (as quoted above), and whole works have been devoted to the role of behaviour in evolution (e.g. Piaget 1979; Plotkin 1988a; Vane-Wright 2014b).

As briefly outlined above, a major thread among ‘third way’ approaches to organic evolution concerns a number of ideas related to the manifest autonomy of whole organisms (Ruiz-Mirazo and Moreno 2012) – going back to Kant, and reaching its modern zenith in related concepts such as autopoiesis (e.g. Maturana and Varela 1980; Thompson 2007), teleonomic selection (Corning 2014b), synergism (Corning 1983, 2005, 2018) and systems theory (e.g. Corning 2014a; Capra and Luisi 2014). Autonomy is also recognised in such notions as value, purposiveness, intentionality, affordance and meaning when applied to living organisms. Here I just want to focus on another of these ideas, that of agency – or simply ‘doings’ to borrow a word from the Kauffman epigraph at the head of this paper.

Agency, the capacity for action, is a key property in considering a phenomenon like mimicry, as it reflects how the autonomous directed activity of an organism is essential for its ability to perceive and thus interpret signals (see below), while at the same time it also indicates the means by which the organism then carries out informed (and sometimes misdirected or misinformed) responses to signals by literally doing some-thing within its own existential environment (von Uexküll’s Umwelt; see Tønnessen et al. 2016). As pointed out by Jones (2017), that organisms have agency does not necessarily infer they also have intentionality. Even so, with respect to the choices or options open to an organism in pursuing its own, individual but also lineage-constrained life-of-meaning (this last in the sense of Goodwin 2007), agency is one of the essential characteristics of life.

However, it must be admitted that the notion of agency remains quite challenging to define, other than as a broad-brush metaphor. Barandiaran et al. (2009: 1), noting that “most current researchers assume an intuitive and unproblematic notion of agency” suggest the following short definition: “an agent is an autonomous organization [cf. “self maintaining automata” of Haukioja 1982: 360] capable of adaptively regulating its coupling with the environment according to the norms established by its own viability conditions” (Barandiaran et al. 2009: 8). The authors then unpack this notion with respect to three essential factors: “a) a system must define its own individuality, b) it must be the source of activity in its environment (interactional asymmetry) and c) it must regulate this activity in relation to certain norms (normativity)” (Barandiaran et al. 2009: 8). This seems very close to my own hitherto intuitive, un-articulated under-standing of the term (as in the previous paragraph) – and is clearly a POL statement, not an evolutionary one.

In this context the in-depth review of Tønnessen (2015) concerning the terms agent and agency as used within the biosemiotics community, is notable. Tønnessen (2015: 14) concludes that “most … appear to agree that core attributes of an agent include goal-directedness, self-governed activity, processing of semiosis and choice of action, with these [four] features being vital for the functioning of the living system in question.” I my view all of these attributes are instantiated by the signal receiver (operator in sense of Vane-Wright 1976) in a functional mimicry system.

The most complete ‘third way of evolution’ treatment of agency that I have seen is that of Walsh (2015). In a chapter entitled “Object and agent: enacting evolution” he makes the case that evolution is an ecological phenomenon that affects lineages as “a consequence of individual organisms’ purposive engagement with their conditions of existence … in pursuit of [their] particular way of life” (Walsh 2015:208). Walsh (2015: 209) then goes on to cite Ingold (1986) with approval in stating that organisms are agents that participate actively in their own evolutionary process (cf. Kull 2000), and Ingold (1989: 208) again to the effect that we need to replace population genetics (an object based theory) with a relationships theory in which the organism is enmeshed as “a creative agent within a total field of relations whose transformations describe a process of evolution”. Similar ideas can be found in the writings of many other, essentially ‘third way’ thinkers (e.g. Whitehead 1929; Bohm 1980; Ho 2017; Affifi 2018). Kauffman (2008: e.g. 263) makes a similar argument in a different way, insisting that biology cannot be reduced to physics because physics cannot accommodate agency, a fundamental “expression of life”.

Walsh (2015) then draws a critical distinction between ‘object theories’ [comparable to Kauffman’s notion of physics?] and ‘agent theories’ to identify why organisms “have disappeared from [conventional] evolutionary theorising” (Walsh 2015:214;cf. Bruce 2014). This leads ineluctably to the conclusion that neo-Darwinism cannot just be tinkered with or simply ‘extended’ (as proposed e.g. by Pigliucci and Müller 2010) to accommodate organismal and relational approaches, but needs to be replaced or at least complemented by a wholly separate theory of evolution built around autonomy, purpose, action and agency, all within a nuanced environmental context. This argument can be compared with Noble’s (2012) call for “biological relativity”, and his suggestion that the neo-Darwinian theory “requires either extension or replacement” (Noble 2015). Walsh proposes to name this new or additional theory “Situated Darwinism”– but whatever it may be called, his whole argument underlies the centrality of agency for a theory of evolution that has the capacity to address ecological relationships of the kind instantiated in the theory of mimicry, and asserts that neo-Darwinism, at least in its extreme gene-centred form, is constitutionally incapable of doing so. In my opinion this again reflects Haukioja’s (1982) view that the conflation of change theory with POL theory lies at the heart of all these difficulties.

Most Vertebrates Don’t like Being Stung

Consider the classic ‘armchair’ scenario for development of aversion-based behaviour in young or naïve insectivorous vertebrates endeavouring to prey on bees for the first time. After a few encounters the expectation is that many or even most vertebrates will come to associate the stinging sensation with one or more conspicuous features of the insect, such as buzzing, colour pattern, even smell (Guilford et al. 1987; Siddall and Marples 2011). Then by remembering this association, they will have learned to avoid bees in future – at least for significant periods.

During the latter half of the 19th and first half of the 20th centuries extreme scepticism, even outright rejection (e.g. Piepers 1913 for an egregious example) regarding the reality of mimicry was often voiced – in part as a reaction against the almost fanatical enthusiasm for the subject displayed by Edward Poulton and his associates. So it is perhaps surprising that, in the context of mimicry, the works of Jane and Lincoln Brower (e.g. Brower et al. 1960; Brower and Brower 1962) were among the first well-designed experimental investigations to provide convincing empirical evidence that some vertebrates do find bees largely unacceptable, and can learn to reject them on sight (and/or sound, and/or smell). They also showed, by the same token, frequent rejection of putative mimics.

The learned aversion to bees offers a good example of semiosis: a reliable, signal-based communication system operating between two organisms to facilitate a mutually beneficial relationship. We can reasonably suggest, echoing Kauffman (2008), that this system has value and meaning because it matters to both entities: the vertebrate and the bee. Neither is injured if the insectivore, based on previous bad experience, declines to attack other insects with the same (aposematic) sound, smell and/or coloration. Of course, this raises issues about the mechanisms that lie behind this process. For example, if those individual bees necessary for the initial ‘training’ process die or are severely debilitated, is kin selection necessary or not for the evolution of the aposematic signals in the first place?

However, ignoring the kin selection issue in the context of this paper (and mindful of Haukioja 1982), it is evident that as soon as such a semiotic relationship is established (signal generator, and a ‘tuned’ receiver that interprets the signal and acts accordingly), immediately there is the possibility (even if not the reality) that a third organism, by simulating the relevant sign well enough for the receiver to interpret it in the standard (normative) way, can either become part of that synergy, or parasitize the established communication channel for its own antagonistic or defensive purposes. In the next section I consider the case of a toad, a bee and a robber-fly, as investigated by Brower et al. (1960), together with another classic case, that of cleaner-fish mimicry.

Intention, Agency and Meaning in Functional Mimetic Relationships

Brower et al. (1960) investigated a system involving a putative mimic, the predatory robber fly Mallophora bomboides, which has a very similar appearance and sound to that of the bumblebee Bombus pensylvanica (as B. americanorum), a putative model with which it shared the same Florida habitat. Their laboratory experimental design was based on a locally occurring toad, Bufo (Anaxyrus) terrestris, as predator, and included the use of dragonflies as edible prey controls. The overall goal was to find evidence to test whether the similarity was or was not mimetic, and if it was, whether the system represented aggressive mimicry, Batesian mimicry, or both.

Naïve toads, on presentation of intact Bombus pensylvanica, attacked – but after no more than two experiences, they then rejected further bees on sight alone – thus demonstrating that the toads could rapidly associate the signal pattern of the bumblebee with unacceptable prey. In another series of presentations toads did not form any aversion to bees from which the stings had been removed. Likewise, except in one instance after receiving a bite on the nose, naïve toads found Mallophora bomboides acceptable prey. But those toads that had had two experiences of intact bees thereafter not only avoided B. pensylvanica but also the robber flies. Thus the authors offered convincing evidence that, with respect to a locally occurring vertebrate insectivore, the bumblebee is aposematic – and acts as a model for the robber fly, which can be regarded as a Batesian mimic.

With respect to aggressive mimicry, Brower et al. (1960: 353) observed that the robber flies did prey “extensively” on the bumblebees (but they also ate a wide variety of other insects). Although they were unable to carry out any experimentation on this aspect of the bee/fly system, they speculated that it was possible that the similarity of the flies to the bees might also “in part be [due to] the visual selection resulting from the defensive behavior of the models towards the mimics which attack them”.

If aggressive mimicry does occur in this system, even if only occasionally, then the system is subject to the actions of two very different operators (sensu Vane-Wright 1976; cf. Boppré et al. 2017). Moreover, these actions fall into two classes sharply distinguished in almost all classifications of mimetic phenomena – aggressive and protective (Batesian). As pointed out by Dalziell and Welbergen (2016: 611), it is not helpful to label this, as has been done in some other cases, “Batesian-aggressive mimicry” or any other compound term of this type. Rather, this serves to remind us that mimicry classes are not labels that can be applied to whole species, or even individual organisms, but ultimately relate instead to a specific, momentary interaction (a Whiteheadian ‘event’: Casati and Varzi 2015) where the signal mediated dynamic confers an instantaneous advantage on the mimic. Thus in any specific event “an organism must either fulfil the role of mimic, or model, not both … logically, models and mimics can be defined but only as interactive elements – they are not ‘labels’ to be applied in a fixed or permanent way to individuals, populations or species” (Vane-Wright 1981: 38). To which I might now add, sensu Walsh (2015), they are not objects.

What the Brower et al. (1960) study also demonstrates is that intentionality, agency and meaning are all part of functional mimetic relationships – even if intentionality is not conscious. But this qualification may not always apply, as some cleaner fish mimics appear to demonstrate.

Cleaner fish (wrasse: LABRIDAE) and their aggressive mimics (blennies: BLENNIIDAE) have been extensively investigated (e.g. Eibl-Eibesfeldt 1959; Wickler 1963; Losey 1972; Côté and Cheney 2005). While the biology of the various fish is varied and complex in detail, the basic cleaner-fish-schema is simple to comprehend: cleaner wrasse remove ectoparasites and epidermal debris from a wide variety of other fish – a ‘service’ often provided at specific locations among coral reefs, so-called ‘cleaning stations’. The supposed mutualism is based around the distinct shape, colour pattern and ‘dancing’ behaviour of the cleaners, which provide a multicomponent signal that fish in need of cleaning (‘clients’) recognise and accept. To be cleaned effectively a client on arrival at a station must stop, and then open its mouth and gills to permit the cleaners to go about their work. Certain scale-predator blennies are able to invade this semiotic system through simulation of both colour-pattern and behaviour. If a fish then mistakenly offers itself for cleaning to such a blenny, its only ‘reward’ will be to have pieces bitten from its fins.

An interesting example of how the dynamics of such a cleaner system can be affected by the behaviour of the mimic and its potential receivers is offered by the fangblenny Plagiotremus rhinorhynchos, a facultative mimic of the bluestreak cleaner Labroides dimidiatus (Côté and Cheney 2005). Unlike most organisms involved in mimicry, the fangblenny does not have a fixed signal, but can choose to adopt mimetic coloration and behaviour according to its ecological circumstances. If feeding at or near a cleaning station with abundant cleaner wrasse, the fangblenny adopts the characteristic ‘electric blue stripe’ pattern of Labroides dimidiatus and acts as an aggressive mimic. However, if cleaners are absent from a reef, or the fangblennies are working among fish shoals in open water, they switch to one or other of two different colour patterns, thus becoming relatively inconspicuous and less likely to invite attack when engaged in scale predation. Both feedings strategies are equally effective according to Côté and Cheney (2015). Thus in this case adopting a mimetic pattern becomes a (seemingly intentional) choice, effected by the agency and behavioural and physiological plasticity of the whole organism. For certain this plasticity is made possible in part by information encoded within the genome, but the whole organism, not just its genes, is responsible for effective deployment. That this variable dynamic is open to multifold evolutionary change through time must be expected – but that will not affect the dynamic at any particular moment. Thus, again, the process-of-living dynamic and the evolutionary-change dynamic should not be conflated.

Another Look at the Definition of Mimicry

Mimicry is a logical necessity arising from the signal-based interactions that we know (believe) to exist in nature, regardless of whether or not there are any real examples. Mimicry theory describes the possibilities for the evolution of parasitism and mutualism inherent in the establishment of communication. A concept of communication which does not embrace perception is, I believe, impossible. Vane-Wright (1981:37)

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Common to all examples of mimicry … is the deception of the signal-receiver by a counterfeit signal that carries a quite specific meaning for the receiver. Wickler (1968:241)

In light of all the above, together with various criticisms I received in 1981 (notably from Robinson 1981 and Endler 1981), new insights from Dalziell and Welbergen (2016), and some suggestions received from Kalevi Kull (pers. comm.), I offer a revised definition or summary description of mimicry:

Mimicry is a functionally tripartite ecological relationship in which a living organism (the mimic), by simulation of signal properties (i.e. the properties used in the interpretation process of the receiver) of another living organism (the model) that have specific meaning for a third living organism (the receiver), becomes misinterpreted by the receiver, which, in accordance with its perception of the similarity, acts as if the mimic were an example (or instance) of the model, resulting in at least some momentary advantage to the mimic.

The question of whether or not the model must be a living organism, and whether or not there has to be an advantage to the mimic, seem to divide opinion. In my view non-living models generate only cues, not signals, and a notion of mimicry in which misinterpretation by the receiver has no ecological consequences seems trivial. The following explanatory notes relate, in part, to changes from my earlier definition (Vane-Wright 1980: as quoted at the beginning of this paper). First, the concepts included:

1. Functionally tripartite – involving three objectively different organisms that can belong to one, two or three species (totally conjunct, three partially conjunct, and disjunct systems of Vane-Wright 1976).
2. Living – mimicry describes signal mediated interactions between living organisms
   – stones, foetid odours, etc. as ‘models’ are thus excluded (e.g. most examples of “cue mimicry”: Jamie2017; see also Zabka and Tembrock 1986).
3. Ecological – mimicry relates to ‘economic’ relationships between organisms embedded within ecosystems where their individual Umwelten overlap.
4. Simulation – mimicry involves simulation of signals, not cues (cf. Maynard Smith and Harper 2003; “signal mimicry” of Jamie 2017).
5. Specific meaning – mimicry is dependent on the existence of a prior or independent synergistic bipartite relationship between two organisms mediated by a co-evolved (‘honest’) signal generated by one organism (‘signal transmitter 1’ of Wickler 1965) and received and ‘understood’ by the other (the ‘receiver’), such that both organisms benefit (mutualism) (cf. Vane-Wright 1976). Thus the primary (honest) signal has specific meaning for the receiver (within its Umwelt), and is an example of synergism in the sense of Corning (Corning 2018).
6. Model, and Perception – the “percept” of signal transmitter 1 as an internalized representation of the model is something which could then be said to be in “the mind of the receiver” (Dalziell and Welbergen 2016 Fig. 2; 614) or “the brain’s model of its world” (Friston et al. 2012: Fig. 1), or simply an element of the receiver’s “mental representation of the world” (Coon 1983). According to Gregory (1972:11) “Perception is not determined simply by stimulus patterns; rather it is a dynamic searching for the best interpretation of the available data.” This led Gregory (1980) to contend that perceptions are like the hypotheses of science, and Friston et al. (2012) to propose that related actions of a receiver can be regarded “as performing experiments that confirm or disconfirm those hypotheses”. The idea that action is necessary for perception can be traced back to Jean Piaget’s cyclical assimilation and accommodation schemata for understanding cognition, recently reformulated as the organism’s need for “mastery of the laws of sensorimotor contingencies” (Di Paolo et al. 2014). In a general mimicry framework Vane-Wright (1976: 30) asserted that “The nature of the signal-receiver in a mimetic system comprises two fundamental properties, the ability to receive information (sensor function), and the drive to react to the information received (motor function).” The second element would have been better formulated as the drive to act upon its interpretation of the sensory input (motor function), reflecting the receiver’s agency in pursuit of a life of meaning. To this needs to be added the insight of Affifi (2018: 23) that “sensori-motor coordination is … not a temporal process where sense precedes action. It is instead a deep and continuous integration of each.” Thus the “sensorimotor world—a body-oriented world of perception and action—is none other than von Uexküll’s original notion of an Umwelt” (Thompson 2007: 59).
7. Mimic – mimicry then represents the possibility of another organism (‘signal transmitter 2’ of Wickler 1965, or the ‘mimic’) gaining an advantage by effective signal simulation whereby it elicits the same percept in the brain of the receiver as signal transmitter 1 (now in this context the ‘model’). This then presents the potential for synergistic (mutualistic) or antagonistic (parasitic) relationships created by the honest signal-mediated synergy between the model and receiver – “the possibilities for the evolution of parasitism and mutualism inherent in the establishment of communication” (Vane-Wright 1980, and above).
8. Act – if the percept of the model evoked by the mimic results in the receiver (or in this context the operator sensu Vane-Wright 1976) acting “similarly to both of them” (Wickler 1965: 519), this results in functional mimicry if the mimic thereby gains some advantage. As Wickler (1965, 1968) indicates, the normal response of the receiver to the model is always advantageous to the mimic, as otherwise the mimetic signal (or interaction) would not be preserved (this last being an evolu-tionary argument, involving stabilizing selection). In somewhat confusing contrast, although I think they mean the same, Dalziell and Welbergen 2016 (611: Fig. 1) state that “as a result of perceiving the similarity between the mimic and the model – [the receiver] changes its behaviour and benefits the mimic”. The actions of the receiver are also critically important in another context (see under (4) below).

The next section addresses concepts excluded. In line with Haukioja’s (1982) strategy of separating process of living theory from change theory, key exclusions relate to evolution. This is not to say that mimicry systems are not the product of evolution (in my view they are), or that mimicry systems are not suitable for the study of evolution (almost ideal I would suggest, as Darwin, Poulton and others long insisted). Quite simply, this exclusion is in the same spirit as that of pattern cladists who assert that, if we wish to use cladistics as a tool to study evolution, it is best if our analyses are as independent of evolutionary models and assumptions as possible (e.g. Brady 1985).

1. Fitness. According to my original definition (Vane-Wright 1980, as quoted above), the advantage to the mimic leads to an increase in “fitness” (intended in the neo-Darwinian sense, thus implying natural selection). Thus according to Dalziell et al.(2015: 643), in the context of vocal mimicry in birds, “a vocalisation is mimetic if the … resemblance between the mimic and the model … confers a selective advantage on the mimic.” While this will typically be the case in an evolving lineage (reflecting “the possibilities for the evolution of parasitism and mutualism” as above), I now consider fitness or selective advantage as unnecessary ballast for a purely functional definition (cf. Dalziell and Welbergen 2016). But as I have already briefly noted above, for the resemblance to be something other than trivial, there has to be some, at least momentary advantage to the mimic in any event when functional mimicry occurs. Ironically, I argued against Robinson (1981) for excluding fitness, natural selection and evolution (Vane-Wright 1981) – but now consider myself to have been in error – caused by an infatuation with the history of science more than the biology itself. In short, neither considerations of the origin of mimicry nor its subsequent evolution should be included within the definition, which relates to a life process (ecological interaction), not to evolution (theory of change).
2. Signals “of interest”. Properly defined, signals are necessarily ‘of interest’, and so this qualifier is not required.
3. Deception (Maynard Smith and Harper 2003: 86) is not a necessary requirement for mimicry. This view is contra Wickler (1965, 1968, 2013) and Boppré et al. (2017), but corresponds to my former position (Vane-Wright 1976, 1981, 1991), and is affirmed by Dalziell and Welbergen (2016), even though they incorrectly assert that deception was part of my earlier characterisation (Vane-Wright 1980). Mimetic relationships can be mutualistic (in which case they are not ‘deceptive’ in Wickler’s sense) or parasitic (in which case in some sense they are deceptive – but see Vane-Wright 1991 on ‘deception’– one of those “wall” words I strongly suspect).
4. Learning is not a necessary requirement for the definition of mimicry (Dalziell and Welbergen 2016). This is in disagreement with Boppré et al. (2017). Dalziell and Welbergen (2016: 612) argue that the “definition of mimicry [should be] indepen-dent of the mechanism of acquisition”. Acquisition implies evolution, which is why (as above) I would now exclude learning from the definition. However, despite the critique of Plotkin (1988b) regarding poverty of empirical evidence, learning seems unquestionably important in the evolutionary dynamics of many and, ultimately, perhaps all mimicry systems – and, following a sensorimotor approach, arguably key to perception too (Di Paolo et al. 2014).
5. However, learning can be argued to be essential for the existence (evolution) of the primary bipartite model/receiver system – unless we also allow for instinctive (innate) signalling systems (which in terms of the original organic selectionists are merely advantageous patterns of learned or imitative behaviour that have become fixed by processes akin to Waddingtonian genetic assimilation – cf. Baldwin 1902, also Affifi 2018 on “hardened habits”). Thus in the wasp-moth system explored by Boppré et al. (2017) the percepts of the wasps not to engage with look-alikes (e.g. nest mates) are thought to be instinctive, and so the authors excluded this system involving hyper-accurate protective simulation of wasps by moths from mimicry sensu stricto (in their sense, treating the system instead under the rubric masquerade). Again, with respect to tripartite mimicry, on further reflection it now seems to me unnecessary to reject this system from mimicry simply because we presume the wasps are acting instinctively. However, Boppré et al. (2017) argue that the difference between learned and instinctive behaviour on the part of the receiver can lead to marked differences in evolution, and Dalziell and Welbergen (2016: 612) also point to learning as likely fundamental to the evolutionary dynamics of a mimicry complex. I fully concur with these evolutionary arguments.
6. Convergence is not necessary for a system to be considered mimetic (Wickler 1965, 2013) – automimicry offers an excellent example (Brower et al. 1967). Convergence is an evolutionary concept which should therefore in any event be excluded from the definition, as argued above.

Dalziell and Welbergen (2016) also suggest that ideas such identification, or misidentification should be excluded from the definition of mimicry. Here we have a terminological problem. Wickler (2013) not unreasonably associates such terms with a static taxonomic rather than a dynamic ethological mindset. But more generally the word identify has the sense of recognise. This was always my intention, not that birds would be exercising some sort of folk taxonomy, but instead be reacting to mimics according to their acceptance (recognition or not) of the mimic as an example of the model. There could be a need for a term here (perhaps differentiate, as in Losey 1972). Thus Dalziell and Welbergen (2016), while rejecting identification employ “classification” instead, with much the same meaning of assign to a class or category. But in formal taxonomy this is more or less exactly what is meant by identification, whereas classification in taxonomy implies the alternative basic meaning of classify, i.e. to create or formulate classes or categories (Vane-Wright 2017).

However, given the perception criterion formally introduced by Dalziell and Welbergen (2016), and Wickler’s (2013) view that the receiver “just responds in the same way to what it perceives to be one and the same stimulus”, we should perhaps cease using terms based on (mis)identification and (mis)classification. If we need go deeper into perception we should arguably ‘dissect’ it in the sense of hypotheses and active hypothesis testing, as already discussed above. Even so, it is notable that some make a distinction between “accurate perception, whereby our interpretation matches the objective nature of the object or stimulus … and mistaken or false perception, where we in some way mis-interpret what is presented to the senses” (Gross 1987:78).

In Conclusion: Investigations into the Phenomena of Mimicry from POL and Third Way Perspectives

Objects are far more than patterns of stimulation: objects have pasts and futures; when we know its past or can guess its future, an object transcends experience and becomes an embodiment of knowledge and expectation without which life of even the simplest kind is impossible. Gregory (1972:8)

If mimicry is restricted, as above, to a tripartite relationship between three living organ-isms mediated by two signs (one of them honest, the other honest or dishonest) that occurs as a momentary event between two of them (in the Now in the sense of Kull 2018), then by analogy with Gregory (above) we can divide the study of mimicry into three programmes: investigations into past, present, and future. Past and future can be seen as two different approaches that entail change (evolution theory), while studying mimicry as restricted above becomes an investigation into processes of living (POL theory).

In studying the past, a major interest will always be the individual origin of different mimicry systems. In this genetics necessarily plays a significant role. For example, the evolution of mimicry has been widely discussed as a two-part process (Turner 1983, 1984). The first step depends on co-occurrence and some fortuitous similarity (brought about by a change of behaviour, range, mutation, etc. etc.) of the proto-mimic to the potential model, sufficient to confer an advantage on the former. If greater advantage is to be had by improvement of the initial similarity, this will most likely come about by classic natural selection involving increased fitness of better mimics (see below). The process of improvement should stop when no further increase in advantage occurs (thus orthogenesis is not expected).

However, this is a convergence model. Before a particular system is assessed with respect to its origin, it is necessary to know about the phylogenetics of the lineages to which the given life-cycles belong. Thus in the case of automimicry as cited above, there is no need for a two-step process. In some cases of Müllerian mimicry the same is likely to be true, insofar as whole clades may evolve and diversify into numerous daughter lineages, but apparently maintain their original (plesiomorphic) aposematic signal(s). Robinson and Vane-Wright (2018: 698, 712) refer briefly to the Indo-Pacific blue-tiger complex among several genera of milkweed butterflies, which could well be such a case.

If mimicry is seen as a purposive adaptation then, with respect to evolutionary dynamics, a primary focus must continue to be the presupposition of some advantage in the normal processes of life of the mimic and/or its reproduction relative to other individuals not so adapted, or not so well adapted. Thus Gilbert Waldbauer and his co-workers, in a series of increasingly sophisticated field trials involving experimentally altered putative mimics (male Callosamia promethea moths) demonstrated very strong selective advantage in favour of mimicry (e.g. Sternberg et al. 1977).

In attempting to further assess future as well as past evolutionary trajectories for a given mimicry system, fields such as game theory, population genetics and phylogenomics (e.g. Kunte et al. 2014; Timmermans et al. 2017) will continue to be very important. However, these approaches will be greatly improved by better understanding of the ecological dynamics that occur in the Now – if only to better appreciate the variables and constraints that affect a given system, and base evolutionary modelling on more realistic, even if still necessarily simplifying assumptions.

As noted above, the most likely means of origin of mimicry and its subsequent evolution, the two-step process and “modest proposals” of Turner (1983, 1984, respectively) remain entirely plausible. Thus, borrowing words from Shapiro (2011: 144), “inventions that survive purifying selection and prove useful are subject to microevolutionary refinement, perhaps by the kind of processes envisaged in conventional theories”. However, for understanding the process dynamics of mimicry, it is still necessary to acknowledge purpose, agency and meaning (teleonomy: Corning 2014b) – ideas largely if not entirely expunged from extreme forms of neo-Darwinism.

Maran (2017) has laid out an extensive and rich exploration of mimicry in terms of biosemiotics. As many biologists associated with the Third Way embrace agency, purpose and meaning as essential for making sense of evolution, biosemiotics becomes a key method for analysing the life processes entailed in functional mimicry and related phenomena. However, if we accept Haukioja’s distinction, it appears that many commentators on mimicry (myself included) have in the past conflated, and many still continue to muddle together processes of living theory with processes of evolution theory. In much the same way that life must come before organic evolution, and in order to understand the latter it would be best to understand the former first, it would likewise be best if our attempts to understand the ways in which mimicry can come about and then evolve are based, in future, on a better understanding of mimicry and its dynamics, simply as a life process.

Reducing all to a single message, I would say that mimicry is an emergent phenomenon arising from the primary establishment of a coevolved, synergistic semiotic relationship between two organisms when a third organism, by simulating the signal, is welcomed to the party – or becomes an undesirable gate-crasher. Pace Gregory above, it is an agent-based process of life theory of dynamic relationships, not an object theory.

Finally, as suggested to me by Michael Boppré, I need to emphasise that this is a preliminary discussion of mimicry seen as a purely ecological dynamic (POL theory) in the context of ‘third way’ evolutionary thinking (biological relativity). I have done this with special reference to the inter-linked concepts of agency, meaning and perception, and the crucial role of whole organisms. This debate will need clarification, and also extension to all types of adaptive resemblances, notably those included under masquerade and crypsis. This paper is intended merely as a stimulus.

Acknowledgements

This is unfinished work of 50 years standing. Anything good here I owe to others; all mistakes, misunderstandings, omissions and other shortcomings are mine. Conrad Waddington, David Bohm, Willi Hennig, Ernst Mayr, Jean Piaget, Colin Patterson, Richard Gregory, Brian Goodwin, Andrew Packard, Erkki Haukioja, Fritjof Capra, Stuart Kauffman, Terence Deacon, Peter Corning, Patrick Bateson, Denis Walsh, Kalevi Kull, Wolfgang Wickler, Michael Boppré – and many many others – some known to me in person, some only from their writing (a few cited here), have all been very influential. Currently I need to read more John Dewey, Jakob von Uexküll, Thomas Sebeok and Evan Thompson. I am very grateful to the library staff of the Natural History Museum, London, for assistance, to Kalevi Kull and anonymous reviewers who helped me improve the manuscript, and to Timo Maran and Karel Kleisner for their kind invitation – and exceptional patience. This paper is respectfully dedicated to my late friend and mentor Lincoln Pierson Brower (1931–2018), who did so much to inspire and foster my interest in mimicry – and in milkweed butterflies.

Compliance with Ethical Standards

Conflict of Interest The author has no conflict of interest with respect to the contents of this paper.

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