JMIH 2016: Anolis conspersus Color variation and Habitat Use

Bright and early this morning, Christopher Peterson kicked off the anole talks of the day on the topic “Intraspecific color and habitat use variation in Anolis conspersus.” Christopher noted that on Grand Cayman there appear to be three color morphs for A. conspersus: brown, blue, and green and asked if color morph was correlated with habitat use. Christopher captured 309 lizards across the island, photographed them for color analysis, and took a large number of habitat measurements plus basic morphology of the lizards (mass, SVL). When analyzing the color data, however, he noticed that the picture was not so clear: many of the lizards had both blue and green coloration. Since these were not discrete groups, instead he analyzed body pattern, which appeared to be more discrete and showed the same geographical variation. In general, lizards on the East of the island were brown and spotted while the lizards on the West of the island were green/blue with vermiculated pattern.

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Using a complex logistic regression, Christopher analyzed the discretized character state with his habitat and morphological measurements. Disappointingly, he found no associations between morphology or habitat use with body patterns. He concluded that the variation in pattern and coloration is probably best explained by geographic location alone and that future genetic analyses may help clear up if this is a geographical cline with isolation by distance.

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JMIH 2016: Malarial Infection Rates Greater in Anolis carolinensis than Anolis sagrei in Central Florida

Cells infected with P. floridense (left and right) vs. healthy cells (middle)

Cells infected with P. floridense (left and right) vs. healthy cells (middle)

Brian Devlin, a graduate student from University of Central Florida, presented a poster on differential rates of malarial infection by Plasmodium floridense between two Anolis species in Central Florida. While both species exist in the area, A. sagrei is the more recent invader. Brian hypothesized that the infection rate would be higher in A. sagrei because A. carolinensis has coexisted with the parasite longer and might have developed some resistance to it.

Brian collected blood samples from both species and examined the cells under the microscopes to look for signs of malarial infection.  He actually found that the infection rates of P. floridense were significantly greater in A. carolinensis. Infection rate also did not correlate with SVL, sex, presence of tail autonomy, date or locality of the lizard. However, there is a higher rate of infection in the warmer months (May-July) possibly due to the in increase rainfall resulting greater mosquito presence. From these results, Brian hypothesized that the lower malarial infection rates in A. sagrei might have helped the species to outcompete A. carolinensis and successfully establish in Florida.

Brian's poster

Brian’s poster

 

JMIH 2016: Herp League Graduate Student Award Winner: Urban Habitat Partitioning by Two Common Species of Puerto Rican Anolis

Kristin Winchell, my fellow lab mate at the Revell lab, presented her work on the habitat use of two urban dwellers in Puerto Rico. Past studies have shown that Anolis cristatellus and Anolis stratulus vary in abundances and use different portions of the natural habitat. As early as 1964, Rand showed that A. stratulus was less abundant and perched higher on trees in forest habitat. Picture1However, we know very little about whether these patterns are maintained in urban areas where species have access to novel manmade structures. To address this, Kristin evaluated the habitat use of these two species across seven urban replicates and contrasted it to the available habitat. She found that urban A. stratulus uses more isolated perches with greater vegetative canopy and perches at higher portions of the habitat. Anolis cristatellus uses perches that are less isolated, shaded mostly by manmade canopy (i.e. buildings and houses) and at lower heights. When examining these patterns in a multidimensional space, she showed that A. cristatellus has expanded its urban niche through the use of manmade structures, while A. stratulus uses a subset of the natural portion of the habitat that A. cristatellus also uses.

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Her research shows that these two urban dwellers interact with the novel portions of the habitat differently. Anolis cristellus has expanded its niche towards manmade structures which has implications for adaptation to enhance stability and locomotion when using these structures as shown in some of her previous work (Winchell, et al, 2016).  Anolis stratulus uses the less available remnants of the natural habitat which may have implication for conservation if they become sparser as urbanization expands.

JMIH 2016: Comparative Phylogeography of Three Widespread Anolis species across the Puerto Rico Bank

Alexandra Herrera presented on using genetic population structure to understand how geographical processes have shaped genetic isolation of three widespread Anolis species on the Puerto Rican Bank. Geographical processes are an important event in shaping current populations and can lead to interesting patterns of diversification. However, these processes may not necessarily affect species similarly. In this study, Alexandra used a combination of nuclear genes and one mitochondrial gene to examine the population structure of these three anoles.

Evidence strongly suggests that populations of Anolis pulchellus were separated into two major clades through the formation of mountains. These two clades are made up of one cluster from south Puerto Rico and a cluster that includes both Northeast Puerto Rico and the Virgin Islands.

20160708_144935The diversification of the other two species corresponds with tectonic and sea level changes. For Anolis stratulus, divergence between populations in PR, Culebra, Vieques and the Virgins Islands occurred at the end of the Pliocene after the formation of the Virgin Passage.  These populations formed five clusters east PR, south PR, Virgin Islands, Vieques-Culebra and Peter-Norman islands.

20160708_145223For Anolis cristatellus the divergence between east PR and south PR with the Virgin Islands was estimated around the late Miocene-Pliocene transition when the Mona and Virgin Island passages formed. These populations formed 4 clusters east PR, south PR,  Virgin Islands and Carrot Rock-Peter-Culebra-Piñeiro.

20160708_145426This research shows that each species had a different diversification pattern and that they all occurred around the middle of late Pleistocene. Furthermore, geographical processes may affect species differently, leading to various patterns of population structures.

 

 

JMIH 2016: Phylogeography and Population structure of Anolis cristatellus

Quynh Quach presenting her Master's thesis work at JMIH.

Quynh Quach presenting her Master’s thesis work at JMIH.

Quynh Quach, a master’s student from the Revell Lab at U. Mass. Boston, presented her thesis research on “Phylogeography and Population Structure of Anolis cristatellus on the island of Vieques.” Before Quynh joined the Revell lab, former  post-doc Graham Reynolds and former Losos lab undergraduate Tanner Strickland looked at the phylogeography of Anolis cristatellus across Puerto Rico and the Virgin Islands using mitochondrial DNA (in review). Tanner’s work revealed that there was a mitochondrial break on the island of Vieques, just off the coast of Puerto Rico. The mitochondrial data suggested that there were two genetically different groups of A. cristatellus, one on the East and one on the West of Vieques. The only problem was, as we know, mtDNA patterns are not always supported by nuclear whole-genome DNA patterns. In addition, Tanner’s dataset only consisted of 9 samples from Vieques.

When Quynh joined the lab, she wanted to know more about this pattern. Would this division be supported by nuclear genome analyses? Were these lineages anthropogenically introduced? If not, what was the origin of these groups – historical allopatry followed by secondary contact or isolation by distance? So she set out to answer these questions by collecting 300 tail tips from across the island of Vieques, extracting and sequencing both mtDNA and nuclear DNA.

The mtDNA variation shows a strong geographic pattern.

The mtDNA variation shows a strong geographic pattern.

Quynh first constructed a mitochondrial phylogeny to verify the pattern observed by Tanner and Graham. The mtDNA analysis confirmed that there are 2 mtDNA clades on Vieques with strong geographic patterns. The island-wide pattern of mtDNA variation was not what we would expect if anthropogenic introduction were the cause since this would be unlikely to show such a clear East-West pattern with the small contact zone in the middle. So then how did this pattern arise?

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Analysis with K=2 shows two clear groups associated with the East and West.

To answer that question, Quynh next looked at nuclear DNA using RADseq. She sequenced 48 individuals: 5 from Virgin Islands, 6 from Puerto Rico, and 37 from Vieques, then de novo assembled the genome and called 16,808 SNP’s. She ran STRUCTURE and DAPC analyses on this data and found that the Virgin Island samples form 1 cluster and Puerto Rico and Vieques form a second cluster with 4.1% divergence between the groups. But she wondered, what if we look at just Vieques and specify K=2? When she did this with DAPC and saw a clear geographic pattern similar to what she found with the mtDNA. Finally, she tested whether this represented isolation by distance. She found that there was significantly reduced gene flow between geographically distant individuals, supporting this hypothesis as the most likely cause of the variation.

Lastly, Quynh emphasized that it is important to consider multiple genetic markers and not just rely on mtDNA results. Had the group stopped at their original mitochondrial analysis, they would have reached a very different conclusion.

JMIH 2016: The Effect of Incubation Moisture on Desiccation Rate

Corey Cates, a PhD student in the Warner Lab presented his latest
results on desiccation tolerance in Anolis sagrei. Desiccation tolerance
is resistance to water loss and is crucial for lizards especially in dry
habitats. Lizards have parchment-shell eggs that take up water from
the environment during incubation. Corey used two incubation conditions to
test whether desiccation tolerance changes throughout the lifetime of
a lizard and whether incubation moisture has an effect on desiccation
tolerance. His study site consists of four islands within the Tomoka River
in Tomoka State Park, Florida. Two of them
have little vegetation, arid climate and lizards lay their eggs in
dry substrate that consists of shells and rocks. The other two islands
have more vegetation cover, cooler climate and moist dark soil to
incubate the eggs. Corey collected individuals from all
islands and incubated their eggs under dry and wet conditions. He
found that desiccation tolerance is highly plastic: hatchlings that
were incubated under dry conditions show low desiccation rates,

matching rates for naturally incubated individuals.

Experimentally incubated individuals match desiccation rates of individuals sampled in the field

Experimentally incubated individuals match desiccation rates of individuals sampled in the field

He also found that desiccation rates decrease within the lifetime of an individual.

Desiccation rates decrease after releasing hatchlings on experimental islands

Desiccation rates decreased after hatchlings were released on experimental islands

He then released the hatchlings to measure survival. He found that desiccation rates are adaptive: individuals incubated under humid conditions had higher survival on mesic islands, and dry incubated individuals had higher survival on arid islands. Future research will focus on exploring the physiological mechanisms that lead to differences in desiccation tolerance, such as scale number and scale size. He will be continuing complementary research during his dissertation work to further explore the effects of incubation moisture on lizard phenotypes.

JMIH 2016: Evolution Isn’t Slow–Experimental Studies of Eco-Evolutionary Dynamics

For many years, biologists believed that evolution was a process that played out over vast stretches of geological time and would not be observable during field studies. More recent research, however, has begun to show that evolution can occur very quickly and that experiments in the field can address evolution in action. Tom Schoener, eminent professor at the University of California, Davis, shed light on our evolving view of how evolution occurs in his talk, “Eco-evolutionary Aspects of the Lizard Anolis sagrei in an Island Metapopulation” at JMIH 2016.

By introducing a novel predator, the curly-tailed lizard (Leiocephalus carinatus), which devours anoles, to a series of small islands in the Bahamas, Schoener and colleagues were able to observe evolutionary responses in A. sagrei in fewer than 10 years. By preying on A. sagrei, curly-tailed lizards induced behavioral changes in perch height, and created selection for relatively longer limbs that increase anoles’ ability to escape this predator. Curly-tailed lizards also caused a variety of ecological effects, including reducing anole populations and changing arthropod abundance, which may affect the future evolution of anoles on these islands. Ongoing monitoring shows that these anole populations seem to be rebounding and that different types of selection may be acting on hindlimb length.

A curly-tailed lizard (Leiocephalus carinatus) displays its namesake in Florida. Photo: Ianaré Sévi.

A curly-tailed lizard (Leiocephalus carinatus) displays its namesake in Florida. Photo by Ianaré Sévi.

Perhaps not surprisingly, many of the experimental islands were occasionally devastated by hurricanes which are becoming more frequent and more powerful in the Caribbean. While these extreme weather events interrupted some of Schoener’s planned research, they also provided a unique opportunity to study how hurricanes may cause natural selection. Schoener found that anoles which survived hurricanes had longer hindlimbs, and these lizards were better able to hold onto trees and other perches at high wind speeds, likely increasing survival of hurricanes by preventing lizards being blown out to sea! Taken together this body of research suggests that novel environmental changes, such as invasive species or increasingly extreme weather, exert selection on organisms and that we can observe these organisms evolving rapidly on ecological timescales.

JMIH 2016: Late-season Lizards Hatch More Quickly and Run Faster

Previous research in the Warner lab has shown that temperature during egg development influences fitness and performance in Anolis sagrei. In particular, a warmer incubation temperature increases sprint speed. The breeding season of A. sagrei spans from March through October, with lower temperatures early in the season and higher temperatures late in the season. Phil Pearson, a masters student in the Warner Lab, conducted an experiment to test whether embryos are developmentally adapted to their incubation temperature. He collected eggs from two temporally-separated cohorts and incubated them under two different temperatures, simulating seasonal temperature differences. He found that late season hatchlings had higher egg survival when incubated under late season temperature. Regardless of incubation temperature, late season embryos had higher sprint speed, larger body size and longer tails. This might compensate for the late start, since they are competing with early cohort individuals in the population.

Late season hatchlings have higher sprint speed regardless of incubation temperature

Late season hatchlings have higher sprint speed regardless of incubation temperature

Overall, this suggests that timing of oviposition has greater effect on morphology and performance than incubation temperature. Future analysis will show whether timing of oviposition affects survival. Phil released the hatchlings on small islands to measure fitness using a mark-recapture approach and will hopefully present his findings at future meetings.

 

JMIH 2016: Jonathan Losos, Distinguished Herpetologist

JMIH

The Joint Meeting of Ichthyologists and Herpetologists (JMIH) kicked off here in New Orleans yesterday. In the morning, Jonathan Losos got started with the first of many anole themed talks. Jonathan gave one the plenary addresses as the Herpetologists’ League’s “Distinguished Herpetologist” of 2016. As such, he joins a long list of accomplished herpetologists, including the first recipient of the honor in 1981, the great anole biologist (and Jonathan’s undergraduate advisor) Ernest E. Williams.

In Jonathan’s talk entitled “Known knowns and unknown unknowns: herpetological progress in fits and starts”, Jonathan started by paying homage to Ernest Williams. He managed to find slides from Ernest’s 1981 plenary address in which the perception at the time of anole biology was compared to a well-built building. All there was to know about anoles was known… or so people thought. In reality, the building looked more like this:

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E.E. Williams slide on the state of anole knowledge from 1981

The metaphorical building at the time was in fact only partially built, with bits and pieces of different areas more complete than others. Jonathan’s talk focused on the fact that despite over 3 decades of progress, so much is still unknown about anoles, including basic natural history of many species. And so Jonathan shared with us a few stories highlighting some surprising anole findings and remaining unknowns, featuring the work of his students from the past 20 years:

Genetic diversity:
One surprise finding over the past 30 years is that several anole species have deep 2016-07-07 11.07.29mitochondrial splits. Anolis oculatus, for example, on the tiny island of Dominica has 4 distinct lineages with as great as 10% mtDNA divergence (Malhotra and Thorpe 2000)! And they aren’t the only ones. Rich Glor and Jason Kolbe really broke this story open with their analysis of several anole species showing multiple mitochondrial lineages for each (Kolbe et al. 2007). This brings into question our estimates of diversity. If every species is actually 4+ species, have we underestimated diversity?

New Species:
According to Jonathan, it seems that the “dawn of anole discovery” peaked in the 1970’s – the last very distinctly different anole was discovered nearly 40 years ago. And yet just last month, Luke Mahler et al. published a record of a new species of anole discovered on the island of Hispaniola! Hispaniola has been intensely studied by anole biologists, making this all the more surprising. In honor of the naturalist that found the species in the wild, the authors named the new species Anolis landestoyi. This new species has a striking appearance, similar to a chameleon and to the Cuban “false chameleons” (Chamaeleolis clade of anoles), and brings up the question of whether there might be a seventh ecomorph.

Anolis landestoyi, photo by D. Luke Mahler

Anolis landestoyi, photo by D. Luke Mahler

Territoriality:
Highlighting the work of two other Losos Lab members, Alexis Harrison and Ambika Kamath, Jonathan talked about how little we know abut anole territoriality. Conventional knowledge says that males maintain polygynous territories and don’t move too far. But Ambika has shown in her dissertation work that male Anolis sagrei actually move quite a bit, and Alexis has shown that male Anolis carolinensis mate with females on opposite sides of their site, not just nearby females! In general, Jonathan commented that there is a large amount of work to be done still on social behavior in anoles.

Anolis proboscis (photo by Luke Mahler)

Anolis proboscis (photo by Luke Mahler)

Basic Biology:
Jonathan talked about the Anole Annals darling, Anolis proboscis. This understudied species sports a large nasal projection of unknown purpose. This odd species was thought to be extinct for many years until it was “rediscovered” only a few years ago. While sexual selection seems like an obvious cause of this structure (the females do not possess horns), its not clear what the males use it for. The obvious hypothesis, that it is used for male-male combat, is easily refuted by video demonstrating that this structure bends easily. Moreover, it appears that they can bend the horn, as seen in this video! The mystery of this structure’s function remains unsolved.

Finally, Jonathan talked about an interesting anecdote: that Anolis agassizi from Malpelo island seems to have a strange preference for the color orange, as described by Rand et al. (1975). Jonathan described a recent test of this preference replicating the Chuckles candy experiment (the experiment has been described here on Anole Annals) and confirming that  this species does, in fact, prefer the colors orange and yellow when it comes to Chuckles candy. He also showed a video of A. agassizi swooping in from afar to eat (attack?) an orange. Why are they so attracted to this color? Sounds like a project waiting to happen.

In short, Jonathan emphasized these two main points:
1. Natural history information is key; you need to know basic aspects of biology and natural history to dig into the deeper questions.
2. There are so many questions to be answered about anoles still, and room for all who want to join the party.

 

 

Anole Adventures in the Cayman Islands

A sagrei on bluff - Cayman Brac NH

A. sagrei on Cayman Brac.

As part of an ongoing study of Anolis sagrei, recently posted about here with additional links therein, I had the pleasure of joining Anthony Geneva and Shea Lambert on a trip to Cayman Brac. We later met up with Graham Reynolds and his undergraduate student Amy Castle on Little Cayman, and closed the trip out with two days on Grand Cayman. Spending time on all three Cayman Islands was a real treat, in large part because of visually stunning anoles like Anolis conspersus and Anolis maynardi. These two species have received a lot of attention on Anole Annals. Rather than rewrite what’s already been written, I’ve decided just to share some pictures from the team. If you’d like to learn more, click on the species names above and explore to your heart’s content. Enjoy!

A conspersus - hotel NH

A. conspersus on Grand Cayman.

A maynardi - Little Cayman SL

A. maynardi on Little Cayman. Photo by Shea Lambert.

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