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Concept: Chameleon

328

Stretching elastic tissues and using their recoil to power movement allows organisms to release energy more rapidly than by muscle contraction directly, thus amplifying power output. Chameleons employ such a mechanism to ballistically project their tongue up to two body lengths, achieving power outputs nearly three times greater than those possible via muscle contraction. Additionally, small organisms tend to be capable of greater performance than larger species performing similar movements. To test the hypothesis that small chameleon species outperform larger species during ballistic tongue projection, performance was examined during feeding among 20 chameleon species in nine genera. This revealed that small species project their tongues proportionately further than large species, achieving projection distances of 2.5 body lengths. Furthermore, feedings with peak accelerations of 2,590 m s(-2), or 264 g, and peak power output values of 14,040 W kg(-1) are reported. These values represent the highest accelerations and power outputs reported for any amniote movement, highlighting the previously underestimated performance capability of the family. These findings show that examining movements in smaller animals may expose movements harbouring cryptic power amplification mechanisms and illustrate how varying metabolic demands may help drive morphological evolution.

Concepts: Evolution, Metabolism, Biology, Organism, Species, Genus, Tongue, Chameleon

181

Fluorescence is widespread in marine organisms but uncommon in terrestrial tetrapods. We here show that many chameleon species have bony tubercles protruding from the skull that are visible through their scales, and fluoresce under UV light. Tubercles arising from bones of the skull displace all dermal layers other than a thin, transparent layer of epidermis, creating a ‘window’ onto the bone. In the genus Calumma, the number of these tubercles is sexually dimorphic in most species, suggesting a signalling role, and also strongly reflects species groups, indicating systematic value of these features. Co-option of the known fluorescent properties of bone has never before been shown, yet it is widespread in the chameleons of Madagascar and some African chameleon genera, particularly in those genera living in forested, humid habitats known to have a higher relative component of ambient UV light. The fluorescence emits with a maximum at around 430 nm in blue colour which contrasts well to the green and brown background reflectance of forest habitats. This discovery opens new avenues in the study of signalling among chameleons and sexual selection factors driving ornamentation.

Concepts: Fluorescence, Ultraviolet, Organism, Species, Skull, Fluorescent lamp, Genus, Chameleon

151

Amniotes possess variability in sex determining mechanisms, however, this diversity is still only partially known throughout the clade and sex determining systems still remain unknown even in such a popular and distinctive lineage as chameleons (Squamata: Acrodonta: Chamaeleonidae). Here, we present evidence for female heterogamety in this group. The Malagasy giant chameleon (Furcifer oustaleti) (chromosome number 2n = 22) possesses heteromorphic Z and W sex chromosomes with heterochromatic W. The panther chameleon (Furcifer pardalis) (2n = 22 in males, 21 in females), the second most popular chameleon species in the world pet trade, exhibits a rather rare Z1Z1Z2Z2/Z1Z2W system of multiple sex chromosomes, which most likely evolved from W-autosome fusion. Notably, its neo-W chromosome is partially heterochromatic and its female-specific genetic content has expanded into the previously autosomal region. Showing clear evidence for genotypic sex determination in the panther chameleon, we resolve the long-standing question of whether or not environmental sex determination exists in this species. Together with recent findings in other reptile lineages, our work demonstrates that female heterogamety is widespread among amniotes, adding another important piece to the mosaic of knowledge on sex determination in amniotes needed to understand the evolution of this important trait.

Concepts: Human, Male, Chromosome, Sex, Reptile, Gamete, Sex-determination system, Chameleon

18

One of the most distinctive traits found within Chamaeleonidae is their split/cleft autopodia and the simplified and divergent morphology of the mesopodial skeleton. These anatomical characteristics have facilitated the adaptive radiation of chameleons to arboreal niches. To better understand the homology of chameleon carpal and tarsal elements, the process of syndactyly, cleft formation, and how modification of the mesopodial skeleton has played a role in the evolution and diversification of chameleons, we have studied the Veiled Chameleon (Chamaeleo calyptratus). We analysed limb patterning and morphogenesis through in situ hybridization, in vitro whole embryo culture and pharmacological perturbation, scoring for apoptosis, clefting, and skeletogenesis. Furthermore, we framed our data within a phylogenetic context by performing comparative skeletal analyses in 8 of the 12 currently recognized genera of extant chameleons.

Concepts: Evolution, Biology, Developmental biology, Greek loanwords, Chameleon, Veiled Chameleon, Chamaeleo, Chameleons

9

Lizards and snakes exhibit colour variation of adaptive value for thermoregulation, camouflage, predator avoidance, sexual selection and speciation. Furcifer pardalis, the panther chameleon, is one of the most spectacular reptilian endemic species in Madagascar, with pronounced sexual dimorphism and exceptionally large intraspecific variation in male coloration. We perform here an integrative analysis of molecular phylogeography and colour variation after collecting high-resolution colour photographs and blood samples from 324 F. pardalis individuals in locations spanning the whole species distribution. First, mitochondrial and nuclear DNA sequence analyses uncover strong genetic structure among geographically restricted haplogroups, revealing limited gene flow among populations. Bayesian coalescent modelling suggests that most of the mitochondrial haplogroups could be considered as separate species. Second, using a supervised multiclass support vector machine approach on five anatomical components, we identify patterns in 3D colour space that efficiently predict assignment of male individuals to mitochondrial haplogroups. We converted the results of this analysis into a simple visual classification key that can assist trade managers to avoid local population overharvesting.

Concepts: DNA, Evolution, Biology, Sexual dimorphism, Color, Population genetics, Chameleon, Furcifer

5

The Malagasy giant chameleons (Furcifer oustaleti and Furcifer verrucosus) are sister species that are both broadly distributed in Madagascar, and also endemic to the island. These species are also morphologically similar and, because of this, have been frequently misidentified in the field. Previous studies have suggested that cryptic species are nested within this chameleon group, and two subspecies have been described in F. verrucosus. In this study, we utilized a phylogeographic approach to assess genetic diversification within these chameleons. This was accomplished by (1) identifying clades within each species supported by both mitochondrial and nuclear DNA, (2) assessing divergence times between clades, and (3) testing for niche divergence or conservatism. We found that both F. oustaleti and F. verrucosus could be readily identified based on genetic data, and within each species, there are two well-supported clades. However, divergence times are not contemporary and spatial patterns are not congruent. Diversification within F. verrucosus occurred during the Plio-Pleistocene, and there is evidence for niche divergence between a southwestern and southeastern clade, in a region of Madagascar that shows no obvious landscape barriers to dispersal. Diversification in F. oustaleti occurred earlier in the Pliocene or Miocene, and niche conservatism is supported with two genetically distinct clades separated at the Sofia River in northwestern Madagascar. Divergence within F. verrucosus is most consistent with patterns expected from ecologically mediated speciation, whereas divergence in F. oustaleti most strongly matches the patterns expected from the riverine barrier hypothesis.

Concepts: DNA, Gene, Evolution, Species, Phylogenetics, Clade, Madagascar, Chameleon

3

The trade in wildlife and keeping of exotic pets is subject to varying levels of national and international regulation and is a topic often attracting controversy. Reptiles are popular exotic pets and comprise a substantial component of the live animal trade. High mortality of traded animals raises welfare concerns, and also has implications for conservation if collection from the wild is required to meet demand. Mortality of reptiles can occur at any stage of the trade chain from collector to consumer. However, there is limited information on mortality rates of reptiles across trade chains, particularly amongst final consumers in the home. We investigated mortality rates of reptiles amongst consumers using a specialised technique for asking sensitive questions, additive Randomised Response Technique (aRRT), as well as direct questioning (DQ). Overall, 3.6% of snakes, chelonians and lizards died within one year of acquisition. Boas and pythons had the lowest reported mortality rates of 1.9% and chameleons had the highest at 28.2%. More than 97% of snakes, 87% of lizards and 69% of chelonians acquired by respondents over five years were reported to be captive bred and results suggest that mortality rates may be lowest for captive bred individuals. Estimates of mortality from aRRT and DQ did not differ significantly which is in line with our findings that respondents did not find questions about reptile mortality to be sensitive. This research suggests that captive reptile mortality in the home is rather low, and identifies those taxa where further effort could be made to reduce mortality rates.

Concepts: Reptile, Question, Squamata, Pet, Chameleon, Wildlife, Amniote, Tuatara

2

Chameleons have a diagnostic thermal protection that enables them to live under various conditions. Our developed special radiative control therefore is inspired by the chameleon thermal protection ability by imitating its two superposed layers as two pigment particles in one coating layer. One particle imitates a chameleon superficial surface for color control (visible light), and another particle imitates a deep surface to reflect solar irradiation, especially in the near-infrared region. Optical modeling allows us to optimally design the particle size and volume fraction. Experimental evaluation shows that the desired spectral reflectance, i.e., low in the VIS region and high in NIR region, can be achieved. Comparison between the measured and calculated reflectances shows that control of the particle size and dispersion/aggregation of particle cloud is important in improving the thermal-protection performance of the coating. Using our developed coating, the interior temperature decreases and the cooling load is reduced while keeping the dark tone of the object.

Concepts: Light, Sun, Thermodynamics, Electromagnetic spectrum, Infrared, Visible spectrum, Chameleon, Emissivity

1

Much of our current state of knowledge concerning sex chromosome evolution is based on a handful of ‘exceptional’ taxa with heteromorphic sex chromosomes. However, classifying the sex chromosome systems of additional species lacking easily identifiable, heteromorphic sex chromosomes is indispensible if we wish to fully understand the genesis, degeneration, and turnover of vertebrate sex chromosomes. Squamate reptiles (lizards and snakes) are a potential model clade for studying sex chromosome evolution as they exhibit a suite of sex determining modes yet most species lack heteromorphic sex chromosomes. Only 3 (out of 203) chameleon species have identified sex chromosome systems (all with female heterogamety, ZZ/ZW). This study uses a recently developed method to identify sex-specific genetic markers from restriction site-associated DNA sequence (RADseq) data, which enables the identification of sex chromosome systems in species lacking heteromorphic sex chromosomes. We used RADseq and subsequent PCR validation to identify an XX/XY sex chromosome system in the veiled chameleon (Chamaeleo calyptratus), revealing a novel transition in sex chromosome systems within the Chamaeleonidae. The sex-specific genetic markers identified here will be essential in research focused on sex-specific, comparative, functional and developmental evolutionary questions, further promoting C. calyptratus' utility as an emerging model organism. This article is protected by copyright. All rights reserved.

Concepts: DNA, Gene, Evolution, Species, Chromosome, Squamata, Chameleon, Veiled Chameleon

0

While many animals display different colour patterns that signal different messages, some species use various tactics to separate between colour and pattern displays. The common chameleon (Chamaeleo chamaeleon) is capable of rapidly changing and separating among displays of colour patterns and ornaments. We used chameleons to study the contextual role of separation among colour and pattern displays. Specifically, we studied the predominant white badge, which is composed of multiple parts, during different seasons and in different social contexts. We hypothesized that the badge contains important information about the sender and, therefore, would be present during important social contexts. We carried out a series of trials to document the presence/absence of the badge and found that the badge is individually specific and reflects body size. We also revealed that the badge remained fixed throughout other body colour changes, but was replaced by other colour patterns during mating behaviour. During social encounters, additional dark patches delineating the badge appeared, presumably amplifying its signal. Thus, we suggest that the badge constitutes an important feature in intraspecific communication, and is possibly employed to display quality. However, the replacement of the badge by other displays during courtship suggests that during important social events like mating, chameleons transmit exclusive information that is not broadcast by the badge. Our findings demonstrate the importance of separation between colour patterns, and the alternative use of intraspecific colour patterns for specific social contexts in chameleons.

Concepts: Separation, Chameleon, Veiled Chameleon, Chamaeleo, Chameleons, Jane Arden, Jack Bond, Chamaeleo chamaeleon