Differences in visual signal design and detectability between allopatric populations of anolis lizards

We tested the prediction of the sensory drive hypothesis using four allopatric populations of the lizard Anolis cristatellus from two distinct environments (i.e., mesic and xeric conditions). For each population, we measured habitat light characteristics and quantified signal design by measuring the spectral and total reflectance and transmittance of the dewlap. We used these data to calculate dewlap detectability using an empirically based model of signal detection probability. We found that populations from mesic and xeric conditions occupy two distinct habitats with respect to light intensity and spectral quality and that dewlap design has diverged between populations in a way that increases signal detectability in each habitat. The major difference in dewlap design was in total reflectance and transmittance, making dewlaps from xeric habitats darker and dewlaps from mesic habitats brighter. Furthermore, dewlap detection decreased significantly when a dewlap from a xeric habitat is detected under the spectral conditions of a mesic habitat. The converse is true for a dewlap from a mesic habitat. We propose that sensory drive has promoted divergence in dewlap design in distinct habitat light conditions, and we discuss the possibility that selection might promote early stages of reproductive isolation as a by-product of selection on dewlap design to distinct habitat light conditions.     

Sensory response patterns and the evolution of visual signal design in anoline lizards

 The evolutionary relationship between visual system response and visual signal design was investigated in four species of anoline lizards which occupy distinctly different habitats. Anoles display with motion patterns of a colorful throat fan called the dewlap. We assessed signal visibility by recording evoked potentials from the optic tectum in response to a moving stimulus flag (a dewlap-like stimulus), and, in one species, by testing behavioral response. The motion pattern, intensity and spectral quality of the stimulus flag, and the background against which it was viewed, were independently manipulated. In all cases, high-velocity motion patterns with a high percentage of brightness contrast between stimulus and background produced the greatest response. Differences in spectral quality between stimulus and background (color contrast) had no effect on tectal responses, but did influence the behaviorally measured detection probability. Using habitat light data we estimated the visibility of the dewlap of each species in different natural habitats. Each species' dewlap was highly visible in its own habitat, but some were much less visible in the habitats of some other species. Habitat light conditions and visual system response properties appear to have constrained the evolution of dewlap design, in at least some of the species. Accepted: May 1998     

Habitat light and dewlap color diversity in four species of Puerto Rican anoline lizards

Closely related species often have signals that differ dramatically in design. The evolution of such differences may be important in the process of speciation. Selection for signal detectability under different habitat conditions has been proposed as a mechanism leading to the evolution of signal diversity. We examined dewlap color in four closely related species of Anolis lizards that occupy habitats with different light conditions. Initially, we tested the hypothesis that lizards choose specific light conditions within each habitat in which to signal. We rejected this hypothesis for all four species. We next calculated the detectability of the dewlap color of all four species at display locations in each habitat. If selection for detectability under the different light conditions explained the divergence in signal design, the occupant of a given habitat was predicted to have the highest signal detectability in that habitat. However, the rank order of detectability of the four dewlap colors was nearly the same in all four habitats. We concluded that divergent selection for signal detectability does not, by itself, explain the evolution of dewlap color diversity. We hypothesize that the evolution of dewlap color diversity results from simultaneous selection for multiple functions of dewlap color.     

Microhabitat use, diet, and performance data on the Hispaniolan twig anole, Anolis sheplani: pushing the boundaries of morphospace

Caribbean Anolis lizards are often cited as a textbook example of adaptive radiation. Similar morphologies (ecomorphs) have originated in similar ecological settings on different large islands in the West Indies. However, relatively little is known about one of the morphologically most specialized and divergent ecomorphs: the twig anoles. Here, we investigate aspects of morphology, dewlap size, locomotor and bite performance, structural habitat and diet of the poorly known twig anole, Anolis sheplani from Hispaniola. Few observations have previously been made of this species in its natural habitat, and few quantitative data on its natural history are available. A. sheplani is an extreme twig anole with respect to its morphology, performance capacities, and ecological niche. Males and females of this species do not differ from each other in body dimensions, performance or habitat use, but males do have a bigger dewlap than females. We present data for 25 individuals and compare them with data for other Greater Antillean anoles. It becomes apparent that twig anoles constitute a large component of the morphological, functional, and ecological diversity of Anolis lizards. Small twig anoles such as A. sheplani appear to be pushing the boundaries of morphospace and are thus crucial in our understanding of the evolution of phenotypic diversity.     

Lizards speed up visual displays in noisy motion habitats

Extensive research over the last few decades has revealed that many acoustically communicating animals compensate for the masking effect of background noise by changing the structure of their signals. Familiar examples include birds using acoustic properties that enhance the transmission of vocalizations in noisy habitats. Here, we show that the effects of background noise on communication signals are not limited to the acoustic modality, and that visual noise from windblown vegetation has an equally important influence on the production of dynamic visual displays. We found that two species of Puerto Rican lizard, Anolis cristatellus and A. gundlachi, increase the speed of body movements used in territorial signalling to apparently improve communication in visually 'noisy' environments of rapidly moving vegetation. This is the first evidence that animals change how they produce dynamic visual signals when communicating in noisy motion habitats. Taken together with previous work on acoustic communication, our results show that animals with very different sensory ecologies can face similar environmental constraints and adopt remarkably similar strategies to overcome these constraints.     

Climate‐related environmental variation in a visual signalling device: the male and female dewlap in Anolis sagrei lizards

Animals communicate using a variety of signals that differ dramatically among and within species. The astonishing dewlap diversity in anoles has attracted considerable attention in this respect. Yet, the evolutionary processes behind it remain elusive and have mostly been explored for males only. Here, we considered Anolis sagrei males and females to study signal divergence among populations. First, we assessed the degree of variation in dewlap design (size, pattern and colour) and displays by comparing 17 populations distributed across the Caribbean. Second, we assessed whether the observed dewlap diversity is associated with variation in climate‐related environmental conditions. Results showed that populations differed in all dewlap characteristics, with the exception of display rate in females. We further found that males and females occurring in ‘xeric’ environments had a higher proportion of solid dewlaps with higher UV reflectance. In addition, lizards inhabiting ‘mesic’ environments had primarily marginal dewlaps showing high reflectance in red. For dewlap display, a correlation with environment was only observed in males. Our study provides evidence for a strong relationship between signal design and prevailing environmental conditions, which may result from differential selection on signal efficacy. Moreover, our study highlights the importance of including females when studying dewlaps in an evolutionary context.     

A new species of dactyloid anole (Iguanidae, Polychrotinae, Anolis) from the southeastern slopes of the Andes of Ecuador

We describe a new species of Anolis from the southeastern slopes of the Andes of Ecuador, province of Zamora-Chinchipe, Parque Nacional Podocarpus. It belongs to (1) the aequatorialis species-group by being of moderate to large size with narrow toe lamellae, and (2) the eulaemus sub-group by having a typical Anolis digit, in which the distal lamellae of phalanx II distinctly overlap the proximal scales of phalanx I. The new species is most similar morphologically to Anolis fitchi but differs from it mainly by having a dewlap with longitudinal rows of 2-5 granular, minute scales separated by naked skin (longitudinal rows of one or two keeled, large scales separated by naked skin in Anolis fitchi) and a vertically shorter dewlap (longer dewlap in Anolis fitchi).     

Do interspecific interactions between females drive shifts in habitat use? A test using the lizards Anolis carolinensis and A. sagrei

An important goal in evolutionary ecology is to understand how and why coexisting closely related species partition habitat among themselves. Although studies of interspecific interactions typically focus on males, interactions between females may also play an important role in shaping habitat use within multi‐species communities. The green anole (Anolis carolinensis) exhibits a wide range of habitat use in south‐eastern Louisiana, but its observed habitat use is restricted and altered in areas where it occurs with the introduced Anolis sagrei. We staged interactions between these two species in the laboratory to test the hypothesis that A. sagrei dominate A. carolinensis in contests over shared habitat. We examined whether species identity, bite force, dewlap size, and body size affected the outcome of interspecific interactions between both males and females, and tested the prediction that bite force and size would be the most important determinants of interaction outcomes in both sexes. In male interspecific interactions, we found that individuals with relatively larger dewlaps tended to score higher on aggressive behaviours regardless of species identity, and that interactions consisted of signalling and rarely escalated to physical combat. However, we found that A. sagrei females achieved higher aggressive scores than A. carolinensis females in almost all cases, lending support to the notion that female interspecific behaviour is probably more important than male behaviour in driving changes in habitat use. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 843–851.     

Are subspecies of Anolis lizards that differ in dewlap color and pattern also genetically distinct? A mitochondrial analysis

Subspecies of Anolis lizards are often defined on the basis of geographic variation in the color and pattern of the dewlap, an extensible throat fan considered central to species recognition and sexual selection. Among the most impressive examples of this phenomenon are two species of trunk anoles found across Hispaniola and the Bahamas: Anolis distichus is divided into 16 subspecies with dewlap colors ranging from deep wine red to pale yellow while Anolis brevirostris is divided into three subspecies with dewlaps ranging from pale yellow to orange. Limited sampling of allozyme data indicates some genetic divergence among subspecies and suggests that they may deserve recognition at the species-level. Our goal here is to use more comprehensive geographic sampling of mtDNA haplotypes to test whether the five subspecies of A. distichus and three subspecies of A. brevirostris that occur in the Dominican Republic correspond with genetically distinct populations that may warrant recognition under the general lineage concept. We obtain an aligned dataset of 1462bp comprised of the genes encoding ND2 and adjacent tRNAs from 76 individuals of A. distichus from 28 localities and 12 individuals of A. brevirostris from five localities. We find that haplotypes sampled from each Dominican subspecies of A. distichus form well-supported and deeply divergent clades (>10% uncorrected sequence divergence). Strong concordance between mtDNA haplotype structure and previously diagnosed phenotypic variation in traits central to interspecific communication (i.e., the dewlap) leads us to hypothesize that each of the presently recognized Dominican subspecies of A. distichus and A. brevirostris deserves elevation to full species status under the general lineage concept.     

Visual pigments and optical habitats of surfperch (Embiotocidae) in the California kelp forest

We studied the optical microhabitat use and visual pigment variation among a group of closely related teleosts (surfperch: Embiotocidae) living along the nearshore central California coast. We employed a diver-operated spectroradiometer to record the optical microhabitat use of eight surfperch species in Monterey Bay. and microspectrophotometry to measure visual pigment absorbance for nine surfperch species. Species were dichromatic with mixtures of A1- and A2-based visual pigments exhibiting extensive maximum absorbance (lambda(max)) variation across species: 455-482 nm for SWS cones and 527-546 nm for LWS cones. Interspecific variation in sidewelling irradiance measurements (mean lambdaFmaxs) significantly accounted for 63% of the variation in surfperch LWS visual pigments and 83% of the interspecific variation in SWS visual pigments using a phylogenetically-corrected regression technique. Optimality models for maximizing relative photon capture of background radiance demonstrate that the LWS cone lambda(max) values are tuned for maximizing photon capture of the species-specific horizontal visual field, while the SWS cone lambda(max), are well offset from the dominant background radiance. This study is one of the first to demonstrate species-specific differences in habitat usage at microhabitat scales accounting for differences in photoreceptor peak absorbance among closely related, sympatric species.     

Dewlap and other variation in the lizards Anolis distichus and A. brevirostris (Reptilia: Iguanidae)

The dewlap of male anoline lizards is an important component of the mate recognition system. Much attention has focussed on the role dewlap colour plays in the interactions between species of Anolis but little has been paid to the effects of intraspecific variability in this feature. Patterns of intrasubspecific variability in this feature are described for several subspecies of Anolis distichus and A. brevirostris . In most cases variability in dewlap colour appears unrelated to the amount of electrophoretic variability and the level of population differentiation, although there is some evidence for large scale geographic subdivision within three of the taxa examined. Female choice and the implications for various species concepts are discussed.     

Climatic niche shift predicts thermal trait response in one but not both introductions of the Puerto Rican lizard Anolis cristatellus to Miami, Florida, USA

Global change is predicted to alter environmental conditions for populations in numerous ways; for example, invasive species often experience substantial shifts in climatic conditions during introduction from their native to non-native ranges. Whether these shifts elicit a phenotypic response, and how adaptation and phenotypic plasticity contribute to phenotypic change, are key issues for understanding biological invasions and how populations may respond to local climate change. We combined modeling, field data, and a laboratory experiment to test for changing thermal tolerances during the introduction of the tropical lizard Anolis cristatellus from Puerto Rico to Miami, Florida. Species distribution models and bioclimatic data analyses showed lower minimum temperatures, and greater seasonal and annual variation in temperature for Miami compared to Puerto Rico. Two separate introductions of A. cristatellus occurred in Miami about 12 km apart, one in South Miami and the other on Key Biscayne, an offshore island. As predicted from the shift in the thermal climate and the thermal tolerances of other Anolis species in Miami, laboratory acclimation and field acclimatization showed that the introduced South Miami population of A. cristatellus has diverged from its native-range source population by acquiring low-temperature acclimation ability. By contrast, the introduced Key Biscayne population showed little change compared to its source. Our analyses predicted an adaptive response for introduced populations, but our comparisons to native-range sources provided evidence for thermal plasticity in one introduced population but not the other. The rapid acquisition of thermal plasticity by A. cristatellus in South Miami may be advantageous for its long-term persistence there and expansion of its non-native range. Our results also suggest that the common assumption of no trait variation when modeling non-native species distributions is invalid.     

A genomic assessment of species boundaries and hybridization in a group of highly polymorphic anoles (distichus species complex)

Delimiting young species is one of the great challenges of systematic biology, particularly when the species in question exhibit little morphological divergence. Anolis distichus, a trunk anole with more than a dozen subspecies that are defined primarily by dewlap color, may actually represent several independent evolutionary lineages. To test this, we utilized amplified fragment length polymorphisms (AFLP) genome scans and genetic clustering analyses in conjunction with a coalescent‐based species delimitation method. We examined a geographically widespread set of samples and two heavily sampled hybrid zones. We find that genetic divergence is associated with a major biogeographic barrier, the Hispaniolan paleo‐island boundary, but not with dewlap color. Additionally, we find support for hypotheses regarding colonization of two Hispaniolan satellite islands and the Bahamas from mainland Hispaniola. Our results show that A. distichus is composed of seven distinct evolutionary lineages still experiencing a limited degree of gene flow. We suggest that A. distichus merits taxonomic revision, but that dewlap color cannot be relied upon as the primary diagnostic character.     

Dawn and dusk "chorus" in visually communicating Jamaican anole lizards

A key hypothesis explaining the existence of dawn and dusk choruses in acoustically communicating animals centers on the need to advertise continued territorial occupancy after and before a period of nocturnal inactivity. If this hypothesis is correct, it follows that similar dawn and dusk choruses should occur in territorial animals exploiting other signal modalities. Adult male Anolis lizards defend territories by using elaborate head-bobbing displays and extensions of a throat fan or dewlap. Males are inactive at night and return to their territories at dawn, remaining there until dusk. I quantified the production of visual displays as a function of time of day for four species on the island of Jamaica: Anolis lineatopus, Anolis sagrei, Anolis grahami, and Anolis opalinus. All exhibited dawn and/or dusk peaks in display behavior. These patterns have remarkable parallels with the dawn and dusk choruses reported for many acoustically communicating animals.     

Competition and community structure in diurnal arboreal geckos (genus Phelsuma) in the Indian Ocean

In this study, we investigate community structure in day geckos (genus Phelsuma ) in the Indian Ocean. Much of what we know about communities of diurnal arboreal lizards comes from studies of Caribbean Anolis . Phelsuma in the Indian Ocean are ecologically similar to Anolis but not closely related. Using field observations and an experiment, we test three hypotheses for Phelsuma communities, all derived from work on Anolis : (1) Phelsuma species richness will be correlated with the diversity of available perches, (2) sympatric species will partition their habitat use, shifting their habitat use depending on which other species of Phelsuma are present, and (3) experimentally removing individuals of one species will lead to changes in the microhabitat use and/or abundance of sympatric congeners. We find support for all three hypotheses. We also describe some unique aspects of Phelsuma communities, such as partitioning of palm vs. non-palm trees. This study identifies some potentially general features of diurnal arboreal lizard communities, and suggests that some aspects of community assembly might be repeatable.     

Habitat light, colour variation, and ultraviolet reflectance in the Grand Cayman anole, Anolis conspersus

Data from a diversity of sources are consistent with the hypothesis that the Grand Cayman anole, Anolis conspersus, is descended directly from Anolis grahami of Jamaica. Although the two species have remained morphologically similar, coloration in A. conspersus has changed considerably from that of its ancestor. The most dramatic difference is seen in dewlap colour, where A. conspersus has evolved a blue and highly UV-reflective dewlap from the ancestral orange-and-yellow colour state. In addition, variation in normal (non-metachrosis) dorsum coloration in A. grahami populations is limited to shades of green (olive, emerald, teal), whereas in A. conspersus dorsum coloration varies from green to blue and to brown. This increased colour variation occurs despite Grand Cayman being a small, relatively featureless island only 35 km in length. Results of this study suggest that ambient light differences associated with precipitation-related vegetation structure may have played an important role in the evolution of A. conspersus body colour variation. Evidence is presented to show how geological, ecological, and physiological factors could have interacted to select for a short wavelength-reflective dewlap from a long wavelength-reflective precursor following the colonization of Grand Cayman from Jamaica by A. grahami between 2 and 3 Mya.     

Not All Signals are Equal: Male Brown Anole Lizards (Anolis sagrei) Selectively Decrease Pushup Frequency Following a Simulated Predatory Attack

The use of conspicuous communication signals often increases a signaler's risk of predation. Many species communicate with a repertoire of signals that may differ in their conspicuousness to predators. Few studies have examined the ability of prey to selectively decrease the use of individual signals in their displays under heightened predation risk. Here, I examined the behavior of male brown anole lizards (Anolis sagrei) in response to a simulated predatory attack from a model kestrel. This species communicates with three major visual signal types, the head‐bob, pushup, and dewlap extension, which vary in their motion and spectral characteristics. I predicted that lizards would decrease frequencies of the dewlap extension and pushup following the attack, but not the head‐bob. Males modulated their use of individual signals by decreasing pushup rates, but not head‐bob rates. Decreases in dewlap frequency were marginally significant. One explanation for these results is that lizards decrease frequencies of signal types based partly on their conspicuousness. The energetic cost of each signal type may be an equally important factor that determines the signaler's response to predators, particularly if a predatory attack is perceived as imminent.     

Spectral composition and visual foraging in the three‐spined stickleback (Gasterosteidae: Gasterosteus aculeatus L.): elucidating the role of ultraviolet wavelengths

Visual signalling can be affected by both the intensity and spectral distribution of environmental light. In shallow aquatic habitats, the spectral range available for visually mediated behaviour, such as foraging, can reach from ultraviolet (UV) to long wavelengths in the human visible range. However, the relative importance of different wavebands in foraging behaviour is generally unknown. Here, we test how the spectral composition of ambient light influences the behaviour of three‐spined sticklebacks (Gasterosteus aculeatus) when foraging for live cladoceran Daphnia magna. Although paying particular attention to the UV waveband, we measured the foraging preferences of sticklebacks for prey presented under four different spectral conditions. These conditions selectively removed UV (UV–), short‐wave (SW–), mid‐wave (MW–) or long‐wave (LW–) light from the entire spectrum. The absence of UV and long wavelengths strongly reduced prey attractiveness for G. aculeatus compared with conditions without short‐wave and mid‐wave light. To control for potential light habitat preferences in the main experiment, we conducted a further choice experiment without prey stimuli. Fish in these trials did not discriminate significantly between the different spectral conditions. When comparing both experiments, it was observed that, although filter preferences for MW– and LW– conditions were virtually consistent, they differed at shorter wavelengths, with a reduced preference for UV– conditions and, at the same time, an increased preference for SW– conditions in the presence of prey. Thus, prey choice seems to be strongly affected by visual information at the short‐wave end of the spectrum. The foraging preferences were also mirrored by the chromatic contrast values between prey and the experimental background, as calculated for each lighting condition using a series of physiological models on stickleback perception. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 359–368.     

Phylogeny,paraphyly and ecological adaptation of the colour and pattern in the Anolis roquet complex on Martinique

Martinique is an environmentally heterogeneous island with a complex geological history. It is occupied by a solitary anole, Anolis roquet, showing marked geographical variation in colour and other features. Phylogenetic analysis of a segment (1 kb) of the mitochondrial cytochrome b gene across the Anolis roquet series in the southern Lesser Antilles and at 63 localities of Anolis roquet in Martinique indicate that A. roquet is paraphyletic as A. extremus (Barbados) is nested within the Martinique populations. Moreover, divergent phylogenetic lineages exist within Martinique (max. 10.6% uncorrected pairwise), and these lineages are closely associated with the geological history of this complex island. However, objective quantification of the spectroradiometric analysis of hue by delta analysis, together with analysis of the colour pattern, indicate that they are primarily determined by adaptation to environmental conditions, irrespective of these phylogenetic lineages. There is remarkable convergence in hue and pattern in both extreme xeric (dark chevrons on a dull, generally grey/brown, background), and montane conditions (black reticulation and non-UV white spots on a bright, saturated green background). Moreover, parallel trends occur between Martinique and other Lesser Antillean anoles, which further argues for adaptation (increase in green saturation in montane areas and higher levels of UV on the dewlap of some Atlantic forms). As an exception, there are two specific situations where anoles from different lineages look different. These are (i). in the low-altitude regions of the northwest where the northwestern and central lineages make contact, and (ii). in the far south of the island where the southern and central lineages meet.