Revisiting the classification of squamate adhesive setae: historical, morphological and functional perspectives.

Research on gecko-based adhesion has become a truly interdisciplinary endeavour, encompassing many disciplines within the natural and physical sciences. Gecko adhesion occurs by the induction of van der Waals intermolecular (and possibly other) forces between substrata and integumentary filaments (setae) terminating in at least one spatulate tip. Gecko setae have increasingly been idealized as structures with uniform dimensions and a particular branching pattern. Approaches to developing synthetic simulacra have largely adopted such an idealized form as a foundational template. Observations of entire setal fields of geckos and anoles have, however, revealed extensive, predictable variation in setal form. Some filaments of these fields do not fulfil the morphological criteria that characterize setae and, problematically, recent authors have applied the term 'seta' to structurally simpler and likely non-adhesively competent fibrils. Herein we briefly review the history of the definition of squamate setae and propose a standardized classificatory scheme for epidermal outgrowths based on a combination of whole animal performance and morphology. Our review is by no means comprehensive of the literature regarding the form, function, and development of the adhesive setae of squamates and we do not address significant advances that have been made in many areas (e.g. cell biology of setae) that are largely tangential to their classification and identification. We contend that those who aspire to simulate the form and function of squamate setae will benefit from a fuller appreciation of the diversity of these structures, thereby assisting in the identification of features most relevant to their objectives.

The same but different: setal arrays of anoles and geckos indicate alternative approaches to achieving similar adhesive effectiveness.

The functional morphology of squamate fibrillar adhesive systems has been extensively investigated and has indirectly and directly influenced the design of synthetic counterparts. Not surprisingly, the structure and geometry of exemplar fibrils (setae) have been the subject of the bulk of the attention in such research, although variation in setal morphology along the length of subdigital adhesive pads has been implicated to be important in the effective functioning of these systems. Adhesive setal field configuration has been described for several geckos, but that of the convergent Anolis lizards, comprised of morphologically simpler fibrils, remains largely unexplored. Here, we examine setal morphology along the proximodistal axis of the digits of Anolis equestris and compare our findings to those for a model gecko, Gekko gecko. Consistent with previous work, we found that the setae of A. equestris are generally thinner, shorter, and present at higher densities than those of G. gecko and terminate in a single spatulate tip. Contrastingly, the setae of G. gecko are hierarchically branched in structure and carry hundreds of spatulate tips. Although the splitting of contacts into multiple smaller tips is predicted to increase the adhesive performance of a fiber compared to an unbranched one, we posited that the adhesive performance of G. gecko and A. equestris would be relatively similar when the configuration of the setal fields of each was accounted for. We found that, as in geckos, setal morphology of A. equestris follows a predictable pattern along the proximodistal axis of the pad, although there are several critical differences in the configuration of the setal fields of these two groups. Most notably, the pattern of variation in setal length of A. equestris is effectively opposite to that exhibited by G. gecko. This difference in clinal variation mirrors the difference in the direction in which the setal fields of anoles and geckos are peeled from the substrate, consistent with the hypothesis that biomechanical factors are the chief determinants of these patterns of variation. Future empirical work, however, is needed to validate this. Our findings set the stage for future comparative studies investigating the functional morphology of these convergent adhesive apparatuses. Such investigations will lead to an enhanced understanding of the interactions between form, function, and environment of fibril-based biological adhesive systems.

Biogeographic links between southern Atlantic Forest and western South America: Rediscovery, re-description, and phylogenetic relationships of two rare montane anole lizards from Brazil.

Data on species ranges and phylogenetic relationships are key in historical biogeographical inference. In South America, our understanding of the evolutionary processes that underlie biodiversity patterns varies greatly across regions. Little is known, for instance, about the drivers of high endemism in the southern montane region of the Atlantic Rainforest. In this region, former biogeographic connections with other South American ecosystems have been invoked to explain the phylogenetic affinities of a number of endemic taxa. This may also be the case of the montane anole lizards Anolis nasofrontalis and A. pseudotigrinus, known from few specimens collected more than 40years ago. We combine new genetic data with published sequences of species in the Dactyloa clade of Anolis to investigate the phylogenetic relationships of A. nasofrontalis and A. pseudotigrinus, as well as estimate divergence times from their closest relatives. Based on newly sampled and previously overlooked specimens, we provide a taxonomic re-description of those two taxa. Our phylogenetic analysis recovered six main clades within Dactyloa, five of which were previously referred to as species series (aequatorialis, heterodermus, latifrons, punctatus, roquet). A sixth clade clustered A. nasofrontalis and A. pseudotigrinus with A. dissimilis from western Amazonia, A. calimae from the Andes, A. neblininus from the Guiana Shield, and two undescribed Andean taxa. We therefore define a sixth species series within Dactyloa: the neblininus series. Close phylogenetic relationships between highly disjunct, narrowly-distributed anoles suggest that patches of suitable habitat connected the southern Atlantic Forest to western South America during the Miocene, in agreement with the age of former connections between the central Andes and the Brazilian Shield as a result of Andean orogeny. The data also support the view of recurrent evolution (or loss) of a twig anole-like phenotype in mainland anoles, in apparent association with the occurrence in montane settings. Our findings stress the value of complementary genetic sampling efforts across South American countries to advance studies of mainland anole taxonomy and evolution.

Intersexual chemo-sensation in a "visually-oriented" lizard, Anolis sagrei.

While the conspicuous visual displays of anoles have been studied in great depth, the possibility that these lizards may also interact through chemical signalling has received hardly any consideration. In this study, we observed the behaviour of male brown anoles (Anolis sagrei) when introduced into an environment previously inhabited by female conspecifics, and compared it to when they were introduced into an untreated environment. The males in our tests exhibited significantly more elaborate display behaviour (i.e., greater number of dewlap extensions and head-nods) and a significantly greater number of tongue extrusions while in the cage formerly occupied by females than when placed in the untreated, control cage. The absolute numbers of tongue extrusions, however, were relatively low in comparison to average tongue-flick rates of 'true' chemically-oriented lizards. Our results strongly suggest that the males were capable of detecting chemical cues left behind by the females. These observations provide the first evidence of intersexual chemo-sensation in an anole lizard.