And he’s not happy about it! Photo by Karen Cusick
Details on Daffodil’s Photo Blog.
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I’m currently reading a 274 page tome called “The Biology and Biodemography of Anolis carolinensis” by Robert E. Gordon. Dating back to 1956, this impressive piece of scholarship is Gordon’s Ph.D. thesis. Gordon collected the bulk of his data in biweekly nocturnal surveys of the demography and spatial ecology of two populations of green anoles. The surveys continued for over a year, and consequently, this document is filled with insights into these lizards’ ecology.
One sentence that caught my attention was this, from page 195:
Anolis activity is primarily diurnal, although movement and feeding were observed at night under conditions of bright moonlight.
We’ve had observations of anoles feeding at artificial lights before, but have any of you night-owl herpers observed something similar under natural light?
A figure from Gordon (1956). Can we please bring back this elegant asymmetric bar graph plotting style?
There are several previous posts concerned with lizards missing feet or limbs (1, 2, 3). At the risk of being monotonous, here is another. I caught this male green anole (Anolis carolinensis) in Auburn, AL this morning (6.24.16). He was sitting on someone’s porch railing at my apartment complex. In addition to his front left limb he was missing 2 fingers on the front right hand and one on the back left foot. He has no other signs of damage, however, not even any evidence of a regenerated tail. I think what sets this example apart from ones previously given is that the entire limb is missing. There is only a tiny nub of bone at the shoulder, and a small flap of skin. Interestingly, the tiny nub moves back and forth beneath the skin when he runs, as if the entire limb were still present and useful.
We (Warner Lab) have had lizards hatch without limbs in the past. We even had one (A. cristatellus) hatch this year with 6 limbs (three front right arms). It is possible that this carolinensis never developed this limb to begin with; however, the tiny flailing nub and flap of skin make me feel that is not the case. Anyhow, this guy seems fat and happy and still moves pretty fast, despite the handicap.
Shane Campbell-Staton giving his talk at Evolution 2016
We’ve heard about the effects of polar vortexes here on Anole Annals before. The infamous 2013/2014 event brought record-breaking snow and low temperatures to the Southern U.S., leaving people and animals both a little chilled. This created the perfect opportunity for Shane Campbell-Staton to investigate the effects of such extreme events on thermal tolerance of the native Carolina Anole, Anolis carolinensis. Shane also spoke about this at SICB earlier this year, and AA contributor Martha Muñoz covered the talk pretty thoroughly here on Anole Annals. Nevertheless, I’ll summarize some key points here in case you missed it.
An unlucky lizard during the polar vortex snow storms in the South.
Shane got lucky in the sense that he had measured thermal tolerance in August 2013 for populations affected by the polar vortex, 5 months before the event. Typically, the cold arctic air is tightly constrained around the North pole, but periodically the boundaries weaken and the cool air expands southward. These events are not regular, so Shane had no idea one was coming that winter or that it would extend so far south. It was serendipitous that his study populations, 3 in Texas and 1 in Oklahoma, were impacted by the extreme weather event. This species, particularly in the Southern portion of its range, is not used to low temperatures and reports came in of anoles dying off during the storm.
Air temperatures for January 5-7, 2014, compared to the 1981-2010 average. Map by NOAA Climate.gov
So Shane returned in August of 2014 and sampled again, curious as to how this cold impacted thermal tolerance. He found that tolerance to low temperatures, measured as critical thermal minimum (CTmin), was lower in some populations after the event! Even more, the difference was greatest in the Southernmost population (Brownsville, Texas). Shane returned again in the fall of 2014 to see if this effect persisted or if it was simply a plastic response to the event. He found that the populations sampled in 2014, and presumably their offspring, still had lower critical thermal minimums. This result suggests that the extreme cold weather had caused an evolutionary shift in cold tolerance via natural selection: only the animals that could tolerate the cold temperatures survived and passed on their cold-tolerance genes. Shane went on to conduct a common garden study to verify that the trait was not simply a plastic response. He found that the lower CTmin persisted in lab-reared animals: strong evidence that these shifts had a genetic basis.
Lastly, Shane looked at the functional genomics of cold tolerance. Using liver tissues to obtain transcriptomes (representing expressed genes), he found several gene modules associated with thermal tolerance including some associated with respiratory electron transport chain, lipid metabolism, carbohydrate metabolism, and angiogenesis/blood coagulation. He also found that the gene expression patterns in the Southern populations affected by the storm resembled the Northern populations that more regularly experience cool temperatures, indicating a common genetically based adaptive response across populations.
By Pavitra Muralidhar
Adaptive radiation is one of the most intriguing processes in evolutionary biology, and anoles are one of the well-studied examples of this process. Anoles have diversified into over 400 species across the Caribbean and Central America, and contain a multitude of highly divergent morphological and behavioral types. Thanks to an impressive history of research on this clade, we now know quite a lot about the phenotypic aspects of this adaptive radiation; however, we still don’t have a good understanding of the genetic mechanisms underlying this diversity of form, physiology, and behavior. The recent advent of next-generation sequencing, and thus the ability to quickly sequence entire genomes of non-model organisms, offers a tantalizing possibility for investigating the genetic basis of adaptive radiation in Anolis.
Tollis et al., in a lightning talk at Evolution, take advantage of these new genome-sequencing techniques to approach the genetics of adaptive radiation in Anolis. To understand the genetic mechanisms underlying the adaptive radiation of anoles, they preformed de novo genome sequencing on three Anolis species (Anolis frenatus, Anolis apletophallus, and Anolis auratus), chosen to capture different sub-groups of the Anolis phylogeny. With these data, and the published genome sequence of Anolis carolinensis, they looked for patterns in the rate of evolution compared to other vertebrate groups. They also looked within the Anolis genome to detect specific genetic regions associated with selection across the anole radiation.
Tollis et al. found that, in general, anoles appear to have a high rate of molecular evolution for a vertebrate species, which may parallel the high rate of phenotypic evolution seen in this clade. In addition, Tollis et al. looked for signatures of selection across the four Anolis genomes and identified regions associated with reproduction, olfactory reception, and limb development. This last category is of special interest, given that anoles are notorious for changes in limb morphology between species and that limb morphology is one of the key components of ectomorphs in the Greater Antilles. Tollis et al. have provided a great example of using new genetic tools to approach fundamental questions about the mechanisms underlying adaptive radiation.
The invasive brown anole A. sagrei is a territorially polygynous species, and male aggressive behavior is an important trait that affects male fitness. Aggressive behavior is quite variable across individuals and populations, and can differ based on intra- and inter-specific community context. As AA regulars know, A. sagrei is also a very successful invasive species; it has been established in southern Florida for decades, and has been steadily spreading north along the gulf coast, colonizing new regions of the US. Populations at the leading edge of the range expansion experience different biotic and abiotic environments than established populations, which can lead to different selective pressures and divergence in relevant traits. Invasive populations of A. sagrei thus provide a good opportunity to explore variation in aggressive display behavior across different ecological contexts.
Julie Wiemerslage decided to take that opportunity and explore the variation in aggressive behavior across different populations of A. sagrei. In her poster “Population Differences in Territorial Aggression in the Invasive Brown Anoles, Anolis sagrei” she proposes the following two hypotheses: 1) Lizards at the leading edge of the range expansion will be more aggressive, allowing them to outcompete other species in their new range 2) Lizards at the leading edge will be less aggressive, because population densities will be lower than areas with established populations.
To test these hypotheses, Wiemerslage collected male lizards from a) native populations, b) well-established invasive populations, and c) recent invasive populations and brought them to the lab for behavioral trials. For each population, she placed pairs of males together in a cage and quantified aggressive behavioral traits including pushups, head bobs, lunges, and dewlap flashes (don’t worry, the lizards were tethered so they couldn’t actually harm one another). She found that aggression was lowest in the leading edge populations, supporting hypothesis 2. Interestingly, the most aggressive populations were the well-established invasive populations, while individuals from the native range showed an intermediate level of aggression. The cause of this pattern is unclear, though Wiemerslage suggests that more information about these source populations (such as density, community composition) will improve our understanding of the factors affecting aggressive behavior.
We as a species are rapidly changing the global environment. The changes that get the most press are those related to climate, but we are also changing the structure of environments through land development. This leads to many important questions, one of which is whether or not the novel environments that we construct can drive evolutionary change. Kristin Winchell, a graduate student in Liam Revell’s lab at UMass Boston, has been addressing this question in the Puerto Rican lizard Anolis cristatellus, which is common in urban settings. Kristin hypothesized that urban environments should select for longer legs and greater surface area of lamellae (the morphological structures on anole toes that let them grip flat surfaces). Her reasoning was that long legs should allow animals to run faster, which should be beneficial in cities where perches and refuges are further apart than in dense natural forests. Greater surface area of lamellae should be beneficial for better grip of smooth man-made surfaces. Kristin compared morphological traits of multiple pairs of urban/natural environment populations and her hypotheses were supported. Not only that, but differences between populations were maintained in individuals developed under common garden conditions, consistent with a genetic basis of the differences. You can see these results in Kristin’s excellent recent paper in Evolution. Kristin also presented some new preliminary results that directly link the morphological changes she has observed to performance on man-made surfaces. Overall, Kristin’s work indicates that urban environments can be a potent force of rapid microevolutionary change and highlights that we are not only changing the abiotic landscape of the globe, but the evolutionary landscape as well.
Anolis sagrei. Photo by Alex Gunderson
The concept of trade-offs, that if you want to increase your performance in one function you have to decrease performance in another, is fundamental to ecology and evolution. However, detecting trade-offs and the underlying mechanisms that give rise to them is extremely difficult. In his talk, Bob Cox summarized years of research that he and his collaborators have done to understand life-history trade-offs in realistic ecological contexts using the brown anole (Anolis sagrei). Bob’s general approach is to experimentally manipulate the reproductive effort of individuals by removing ovaries and testes before releasing them onto cays in the Bahamas. He then estimates important ecological and physiological parameters such as survival, fat reserves, and immune function to see if he can detect trade-offs between reproductive effort and these other traits. In general, he has found that reproductive investment significantly decreases survival and physiological performance and that effects are often contingent upon factors such as the presence or absence of predators. Check out Bob’s website for a more information about his integration of experimental, ecological, and evolutionary studies to understand how trade-offs shape animal life-histories.
Researchers that are interested in ecological and evolutionary dynamics through time often make inferences about past patterns and processes using modern data, such as DNA sequences and geographic distributions of extant taxa. But this is not the only possible approach. Studies of extinct taxa and populations using fossils can provide direct measures of species distributions and abundances in the past, which are often impossible to accurately infer with modern data alone.
In her talk titled “Extinction biases and their ramifications on Caribbean lizard communities,” Melissa Kemp described her research using fossil data to characterize the former herpetofaunal community of several islands in the Caribbean. She explored the following questions linking extinction to community ecology: 1) how has extinction proceeded in the Caribbean lizard community? 2) what is the impact of species extinction on the whole community? 3) can we predict future patterns of extinction using fossil data?
To characterize past extinction patterns, Kemp measured species abundance and morphological traits of fossil remains through time in lizard communities in the Caribbean. She sought to determine whether certain taxa underwent more local extinctions, and whether extinctions were correlated with certain morphological traits. She also quantified community evenness to see how extinction events affect the whole lizard community. She found that one family, the Leiocephalidae, has gone extinct more often than others. Interestingly, in a four-species community in which Leiocephalidae went extinct, anoles went from relatively average abundance to becoming the dominant taxa, a pattern which continues to this day. Modern Leiocephalids have been shown to predate on anoles, so this community shift may have been a result of predator release. In addition, anole body sizes increased after Leiocephalid extinction, lending further support to the predator release conclusion.
After looking at historical patterns of extinction and diversity, Kemp explored whether fossil data might give us insight into current and future patterns of extinction. For example, are species that have gone extinct in some areas vulnerable to extinction in other parts of their range? And if so, what traits are causing this vulnerability? To address these questions, Kemp compared traits of extinct taxa to traits of modern successful introduced species, which are likely to have a very low risk of extinction. She found that extinct species tend to have different reproductive modes and habitats from introduced species, suggesting that these traits may have played a role in their extinctions. In addition, modern species with similar suites of traits as the extinct taxa may be more vulnerable to extinction in the future.
Kemp’s research shows that it’s not always best to leave the past behind. Fossil data enhances our understanding not only of extinct species, but of modern ecological and evolutionary processes as well.
Evolution 2016: Landscape Topography and Species Diversification in South American Frogs and Lizards
Species divergence is driven by a wide variety of forces, but two of the strongest predictors of speciation are the amount of time a lineage has persisted in a landscape, and the ability of lineages to move through a landscape. Lineages are more likely to diverge when they have occupied a landscape for a long time, and/or if their ability to move is restricted, thus limiting gene flow.
In his talk titled “Geographical factors promoting diversification of the northern Andes and Brazilian Cerrado regions: the case of frogs and Anole lizard species,” Carlos Guarnizo described his efforts to test whether these patterns hold true in both different landscapes and different taxa. He surveyed two herpetofaunal communities in two diversity hotspots in South America: frogs in the northern Andes mountain range and Anolis lizards in the Brazilian Cerrado. The montane Andean landscape is structurally complex and covers a range of altitudes, while the Cerrado region is a more uniform savannah-like environment, with intermediate structural complexity. Guarnizo used species distributions and genetic data to look at patterns of diversification across these landscapes to explore which landscape characteristics lead to higher levels of divergence and speciation.
He found that in both areas, topography was a strong predictor of divergence; specifically, more structurally complex landscapes led to higher levels of genetic divergence between sister lineages. These genetic breaks are also often deeper than previously realized, likely representing cryptic species. Despite these strong genetic splits, the niches occupied by sister taxa are generally well-conserved, lending support to the conclusion that landscape structure – rather than adaptive divergence – is responsible for the genetic divergence observed. Interestingly, in Andean frogs, Guarnizo found that the strongest genetic breaks did not occur across mountain peaks as previously thought. Instead, valleys appear to be the strongest geographic barrier to dispersal.
These cases show that landscape topography is a strong factor determining genetic divergence across different landscapes and taxa (including anoles), and may lead to high levels of cryptic speciation.