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Background: Evolution of novelty is a central theme in evolutionary biology, yet studying the origins of traits with an apparently discontinuous origin remains a major challenge. Venom systems are a well-suited model for the study of this phenomenon because they capture several aspects of novelty across multiple levels of biological complexity. However, while there is some knowledge on the evolution of individual toxins, not much is known about the evolution of venom systems as a whole. One way of shedding light on the evolution of new traits is to investigate less specialised serial homologues, i.e. repeated traits in an organism that share a developmental origin. This approach can be particularly informative in animals with repetitive body segments, such as centipedes. Results: Here, we investigate morphological and biochemical aspects of the defensive telopodal glandular organs borne on the posterior legs of venomous stone centipedes (Lithobiomorpha), using a multimethod approach, including behavioural observations, comparative morphology, proteomics, comparative transcriptomics and molecular phylogenetics. We show that the anterior venom system and posterior telopodal defence system are functionally convergent serial homologues, where one (telopodal defence) represents a model for the putative early evolutionary state of the other (venom). Venom glands and telopodal glandular organs appear to have evolved from the same type of epidermal gland (four-cell recto-canal type) and while the telopodal defensive secretion shares a great degree of compositional overlap with centipede venoms in general, these similarities arose predominantly through convergent recruitment of distantly related toxin-like components. Both systems are composed of elements predisposed to functional innovation across levels of biological complexity that range from proteins to glands, demonstrating clear parallels between molecular and morphological traits in the properties that facilitate the evolution of novelty. Conclusions: The evolution of the lithobiomorph telopodal defence system provides indirect empirical support for the plausibility of the hypothesised evolutionary origin of the centipede venom system, which occurred through functional innovation and gradual specialisation of existing epidermal glands. Our results thus exemplify how continuous transformation and functional innovation can drive the apparent discontinuous emergence of novelties on higher levels of biological complexity. [ABSTRACT FROM AUTHOR]

Background: In the context of evolutionary arthopodial transformations, centipede ultimate legs exhibit a plethora of morphological modifications and behavioral adaptations. Many species possess significantly elongated, thickened, or pincer-like ultimate legs. They are frequently sexually dimorphic, indicating a role in courtship and mating. In addition, glandular pores occur more commonly on ultimate legs than on walking legs, indicating a role in secretion, chemical communication, or predator avoidance. In this framework, this study characterizes the evolutionarily transformed ultimate legs in Lithobius forficatus in comparison with regular walking legs. Results: A comparative analysis using macro-photography, SEM, μCT, autofluorescence, backfilling, and 3D-reconstruction illustrates that ultimate legs largely resemble walking legs, but also feature a series of distinctions. Substantial differences are found with regard to aspects of the configuration of specific podomeres, musculature, abundance of epidermal glands, typology and distribution of epidermal sensilla, and architecture of associated nervous system structures. Conclusion: In consideration of morphological and behavioral characteristics, ultimate legs in L. forficatus primarily serve a defensive, but also a sensory function. Moreover, morphologically coherent characteristics in the organization of the ultimate leg versus the antenna-associated neuromere point to constructional constraints in the evolution of primary processing neuropils. [ABSTRACT FROM AUTHOR]