Abstract
Mimicry and deception are two important issues in studies about animal communication. The reliability of animal signs and the problem of the benefits of deceiving in sign exchanges are interesting topics in the evolution of communication. In this paper, we intend to contribute to an understanding of deception by studying the case of aggressive signal mimicry in fireflies, investigated by James Lloyd. Firefly femmes fatales are specialized in mimicking the mating signals of other species of fireflies with the purpose of attracting responding males to become their prey. These aggressive mimics are a major factor in the survival and reproduction of both prey and predator. It is a case of deception through active falsification of information that leads to efficient predation by femmes fatales fireflies and triggered evolutionary processes in their preys’ communicative behaviors. There are even nested coevolutionary interactions between these fireflies, leading to a remarkable system of deceptive and counterdeceptive signaling behaviors. We develop here a semiotic model of firefly deception and also consider ideas advanced by Lloyd about the evolution of communication, acknowledging that deception can be part of the explanation of why communication evolves towards increasing complexity. Increasingly complex sign exchanges between fireflies evolve in an extremely slow pace. Even if deceptive maneuvers are played out time and time again between particular firefly individuals, the evolution of the next level of complexity—and thus the next utterance in the dialogue between species—is likely to take an immense amount of generations.
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Notes
We should add, however, that this interpretation was controversial. Cf. the discussion between Copeland and Lloyd (1983). Copeland claimed, on the basis of laboratory findings, that Lloyd’s interpretation of mimicry was not so secure, given that was based on a correlation study. He also argued that Photuris females completely change behavior from a virgin phase, where they signal for mating with conspecific males, to a post-mating phase, where their signal behavior changes to attract prey males from other species (see also Nelson et al. 1975). Then, how could mimicry by Photuris males be a reproductive strategy? Lloyd challenged these claims by considering, first, that the behavior of firefly communication, including aggressive mimicry, cannot be properly observed in fireflies confined in labs, as those studied by Copeland. By the way, we have here an interesting exchange about the relationship between lab and field studies of animal behavior. Secondly, he argued that Copeland had not been fair in the manner he addressed the quality and extent of his data about aggressive mimicry. Finally, he discussed Copeland’s argument that other mechanisms than mimicry might be involved in the behaviors observed by him, calling attention to the fact that this was true, but Copeland did not put forward any hypothetical alternative mechanisms. Lloyd also considered different hypotheses about why male Photuris mimicry might be aiming at mating with females that were no longer virgins, having switched to predatory behaviour, such as female selection of male phenotype, paternal investment, and even forced insemination of the already-mated Photuris females. Despite the controversies, the existence of aggressive mimicry and its consequences to the evolution of communication in fireflies are widely accepted, as it can be seen, for instance, in a recent paper by Lewis and Cratsley (2008, p. 313), who offer an overview of a number of recent works that, in their terms, “... provided considerable evidence supporting the idea that Photuris predation represents an important force shaping the evolution of bioluminescent signals in North American fireflies”.
It is striking to note the formal similarities with the arms race between code makers and code crackers in military intelligence in the history of warfare, cf. Singh (1999).
We will follow here the scholarly practice of citing from the Collected Papers of Charles Sanders Peirce by volume number and paragraph number, preceded by ‘CP’; the Essential Peirce, by volume number and page number, preceded by ‘EP’. References to the Annotated Catalogue of the Papers of Charles S. Peirce will be indicated by ‘MS’, followed by the manuscript number and pages. References to Writings of Charles S. Peirce: a Chronological Edition will be indicated by ‘W’, followed by page number.
It should be noted that “sign” here is ambiguous. It may refer to the sign vehicle specifically—in contrast to the object and the interpretant—or it may refer to the whole complex of sign vehicle, object, and interpretant. In most cases, the context makes it clear which use is intended.
Non-symbolic dicisigns may be—according to Peirce’s 10-fold sign classification—Dicent Sinsigns (e.g., the wind direction indicated by a weathercock, the object portrayed by a photograph) or Dicent Indexical Legisigns (e.g., a street cry whose tone and theme identifies the individual uttering it). (CP 2.254–265).
Here, we disagree with Terrence Deacon’s (1997) attempt at characterizing animal semiosis as iconic and indexical only, while claiming that it is a human privilege to process symbols. The Peircean symbol notion refers to simpler semioses than human semiotics, the specificity of which must be characterized by other means.
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Acknowledgments
Charbel N. El-Hani is thankful to the National Council for Scientific and Technological Development (CNPQ) for funding of research projects and research grants. He is also indebted to the Foundation of Research Support of the State of Bahia (FAPESB) for funding of research projects. João Queiroz thanks the National Council for Scientific and Technological Development (CNPQ) and the Foundation of Research Support of the State of Bahia (FAPESB) for funding of research projects and research grants. Frederik Stjernfelt is thankful to the Center for Semiotics, Aarhus University, for fine research conditions. All three authors are indebted to Kalevi Kull for his comments on the original manuscript.
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El-Hani, C.N., Queiroz, J. & Stjernfelt, F. Firefly Femmes Fatales: A Case Study in the Semiotics of Deception. Biosemiotics 3, 33–55 (2010). https://doi.org/10.1007/s12304-009-9048-2
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DOI: https://doi.org/10.1007/s12304-009-9048-2