Year: 2013 Page 13 of 37

Global Warming Effects On Tropical Lizards

Martha Munoz starting the all-anole morning with a comparison of the thermal niches of different species of Hispaniolan cybotoid anoles

Martha Munoz starting the all-anole morning with a comparison of the thermal niches of different species of Hispaniolan cybotoid anoles

The symposium “The Biological Impacts of Tropical Climate Warming for Ectothermic Animals,” was recently (Aug. 1-3) held in San Juan Puerto, and it was a great success. In a two-part post, I will provide a brief summary. Today will focus on four talks on the second morning, all of which focused on Anolis. In the next post, I will review the rest of the symposium.

Martha Muñoz began the day by talking about the thermal biology of cybotoid anoles (members of the cybotes species group) in Hispaniola. These species show a remarkable elevational range from sea level to over 3000 meters. Martha pointed out that in this respect, Hispaniola is a much better place to look at questions related to elevation than Puerto Rico, a comment greeted with jeering from much of the crowd. Nonetheless, she scoffed at the discussion of the “high elevation” A. gundlachi at 850 m. Why, A. shrevei, in Hispaniola doesn’t even occur that low! In any case, what Martha showed is that despite the great thermal differences in habitats at different elevations, the cybotoids maintained approximately the same body temperature at all sites and have the same preferred temperatures and critical thermal maxima. Clearly, they are using thermoregulatory behavior to buffer their thermal physiology from selection in different environments and, indeed, field observations show that high elevation species do bask more. However, anoles can’t thermoregulate at night, and there is where adaptive differentiation occurs: high elevation anoles can withstand lower temperatures than lower elevation species. To clinch the deal, Martha measured the temperatures lizards experience at night. Indeed, the species at high elevation experience temperatures that would kill low elevation species.

Luisa Otera showing a slide of her collaborator, George Gorman, in his cowboy salad days

Luisa Otera showing a slide of her collaborator, George Gorman, in his cowboy salad days

Luisa Otera spoke next on the “Effects of recent climate warming on the reproductive phenology of Puerto Rican Anolis lizards.” Luisa revisited sites at which George Gorman had examined A. cristatellus 40 years ago. Gorman had found that at higher elevations, female reproduction tapered off in the winter, whereas at lower elevations, they continue reproducing year-round. Her prediction was that with higher temperatures, reproduction should be extended in the winter at high elevations. For the most part, this prediction was confirmed, though surprisingly not so at the sea-level site.

Most surprising, in a new twist, Luisa found that female reproduction could vary over a very short spatial scale. In particular, if a lizard has a territory in the open with a lot of sun, it can breed year round, whereas it’s neighbor under the shade of trees a few meters away may not be able to do so in the winter. Perhaps this explains the contrary finding at the sealevel sites: greater tree cover may have actually made conditions worse.

Luisa pointed out that warming isn't always bad--for some lizards, it allows them to extend their breeding seasons

Luisa pointed out that warming isn’t always bad–for some lizards, it allows them to extend their breeding seasons

gunderson

Gunderson’s data show that even lizards with body temperatures outside of their preferred range are still quite active

Alex Gunderson spoke next on “Behavioral responses to climate change: natural selection on the thermal physiology of Anolis sagrei.” Perceptive readers will note that these three talks focused sequentially on the trunk-ground anoles of three different islands. Coincidence? You be the judge. In any case, in a very thought-provoking talk, Alex pointed out that much of the literature predicting the response of species to global warming focuses on the effect that higher temperatures will have on the time in which lizards can be active, which affects factors like food acquisition. However, Gunderson note that although activity time is usually treated as a binary variable—a lizard is either active or it isn’t—his extremely detailed behavioral data (299 focal observations), indicate that, in fact, the effect of temperature on activity is continuous rather than binary. Indeed, lizards engage in all major activities—eating, mating, fighting—at temperatures substantially outside (mostly below) their “preferred temperatures.” This finding calls for a re-thinking of how we model the effects of climate change on lizard populations—they may be forced to be active at temperatures they’re not so happy about, but they will do more than stay in their hidey-holes.

Next, Michael Logan reprised his talk on the “Rapid evolution in response to climate change: natural selection on the thermal physiology of Anolis sagrei” which he gave at the Evolution meetings five weeks previously. But here he had twice as much time to speak and correspondingly gave greater details. Since I’ve reported on the talk previously, I’ll just summarize here: in a very cool experiment, he moved brown anoles from a shady habitat to a much hotter one. Before doing so, he measured the performance curves of each lizard (i.e., how their ability to sprint was affected by temperature). His prediction was that individuals that could sprint at higher temperatures would be favored by natural selection in the new habitat. And sure enough, they were! By contrast, another population in a shaded habitat experienced no selection on thermal performance. If thermal sensitivity of sprinting is a heritable trait—a big if, Mike noted—this strong selection could suggest that populations might be able to adapt very rapidly to warming climates.

Anoles on the rocks, so to speak

After a wonderful trip to Puerto Rico for the recent Thermal Ecology meeting mentioned here on Anole Annals and so heavily attended by anolologists, we had the opportunity to visit some of the natural forests that the country had to offer.

Riparian habitat in the forest by the El Verde Field Station, Puerto Rico

Riparian habitat in the forest by the El Verde Field Station, Puerto Rico

Whilst in El Verde National Park, we were regaled with stories of local Anolis advancing to the ground and using riparian habitat despite what their ecomorph classification might suggest. Given the recent AA interest in aquatic anoles (1,2,3), I thought a short note on this may be appreciated. Apologies for the deceivingly melodramatic title; alas it was literal, not figurative.

An adult male A. evermanni perched on a boulder surrounded by fast flowing water

An adult male A. evermanni perched on a boulder surrounded by fast flowing water

Anolis evermanni, a trunk-crown ecomorph, has been known to use boulders along one of the streams for the past two decades or so. With great anticipation, whilst marching through the forest spotting copious numbers of Anolis gundlachi, we were en route to our final destination to find out! Upon reaching the stream, which incidentally offered some beautiful tropical scenery accompanying the break in the canopy, we were not disappointed to find A. evermanni dotted all around the waterway!

I assure you there is an anole there - this wasn't just an excuse for a rest...!

I assure you there is an anole there – this wasn’t just an excuse for a rest…!

An adult male A. evermanni displaying

An adult male A. evermanni displaying

Back in 1990, Jonathon Losos postulated that this shift in microhabitat from trees to boulders forced a change in locomotor strategy. Whilst anoles are able to travel continuously in a forest, by travelling down a tree, along the ground and then up another, the structural heterogeneity presented by these riparian boulders meant that jumping needed to be more commonly adopted. He posited that the difference in thermal strategy of A. evermanni and A. gundlachi, a heliotherm and a thermoconformer respectively, would affect their likelihood of using these boulders along the highly sunny stream. Although A. gundlachi were observed present along the shaded edge, they rarely ventured further out. After some thought on site, this prompted a brief hypothesis by a couple of us; when the sun began to disappear, would the larger A. gundlachi displace the A. evermanni on the boulders?

This may take some imagination, but that blur to the right of the central vine - I assure you that's a boulder-loving A. gundlachi!

This may take some imagination, but that blur to the right of the central vine – I assure you that’s a boulder-loving A. gundlachi!

After a couple of hours of enjoying the forests of El Verde, we returned to the field station. As we were leaving and the sun was beginning to calm, I spotted our first A. gundlachi out on a stream boulder followed shortly after by a handful of A. stratulus. This would seem to offer a cool behavioural research opportunity for someone that enjoys sitting in the sun by a river watching lizards…(can’t be that bad a gig, can it?).

A. stratulus also getting in on the gig

A. stratulus also getting in on the action

A New Confirmation Of Pair Bonding In Anolis Limifrons

Monogamy, or the formation of stable pair bonds between males and females for reproductive purposes, is thought to be relatively rare across animals. While social pair formation is observed (commonly in birds and occasionally in reptiles), genetic assessments of parentage have revealed that mating fidelity is infrequent. Social monogamy is therefore not equivalent to genetic monogamy. However, the reasons for the persistence of social monogamy despite promiscuous mating remain unclear.

Sleepy lizards are the best known example of pair-bonding in lizards

Sleepy lizards are the best known example of pair-bonding in lizards (photo by J. Todd Kemper)

A new paper by Alexis Harrison revisits one of the only examples of social pair-bonding known from anoles–a population of Anolis limifrons in the La Selva Biological Station in Costa Rica. While most anoles are polygynous, with the territory of one male overlapping the territories of several females, Talbot (1979) noticed that 70% of adult A. limifrons in La Selva were found in pairs of a single male and female in close proximity to each other. However, such pair bonding has not been documented in any other population of the species, making La Selva an intriguing outlier.

A pair of Anolis limifrons

A pair of Anolis limifrons (photo by Jason Weigner)

Hueyfest: A Symposium Honoring Ray Huey

Ray Huey and friends at last year's World Herpetological Congress in Vancouver

Ray Huey and friends at last year’s World Herpetological Congress in Vancouver

Learn about Ray’s storied past.

Ray Huey has been a pioneer in the field of physiological ecology and evolution. Building on the work of Ruibal, Rand,Williams and others (as he always stresses), Ray was instrumental in making anoles a model for understanding thermal biology, integrating behavior, physiology, evolution and,  most recently, conservation biology. And then there’s Ray’s other side. Who else could get away with using a Rolling Stone‘s album in the title of a paper?

A symposium in Ray’s honor will be held in Seattle on Friday, October 4th and is open to anyone, but attendance is limited, so register today. All the details are available on the fest’s website. Whether you attend or not, check out the Hueyblog and add your own tributes and reminiscences.

Eye Color In Anolis Of The Guadeloupe Archipelago

The geographic variation of the highly variable anole Anolis marmoratus from the Archipel de la Guadeloupe (France) has focused interest on the process of speciation resulting from divergent selective pressures. Nice detailed analyses such as that recently published in Molecular Ecology confirmed that differences in body color seem to correlate with environmental characteristics. Within the large diversity of form and color of anoles of the different islands of the archipelago, one phenotypic feature that appears to be variable is the color of the scales around the eye.Untitled
For two years, the team FORCE (UMR 7205 Museum National d’Histoire Naturelle/CNRS, Paris, France) in collaboration with La Direction des collections of the Museum National d’Histoire Naturelle (Paris, France) has been conducting different studies of the behaviors and forms of the anoles of the different islands of the archipelago. The contrasted orange color of the scales around the eye of males Anolis desiradei is remarkable and could probably play a role within the context of behavioral ecology of these anoles. At La désirade, these anoles are living in the same habitat as the endangered Iguana delicatissima. We also recorded some populations of males of Anolis marmoratus on the Basse-Terre with marked black scales around the eyes. In Marie-Galante, the scales around the eye are blue, green or yellow while the head is more often blue or green. The scales around the eye of females of all of the studied populations are often white or pale yellow.Untitled1

We are now measuring the diversity of this phenotypic trait to test various hypotheses of the role of these colors in the communication between the individuals within the selected populations of Anolis from different islands.

More Remembrances Of Ken Miyata

Ken Miyata's handiwork on display outside David Wake's office

Ken Miyata’s handiwork on display outside David Wake’s office

Recently, a chapter of Ken Miyata’s thesis on the ecology of Ecuadorian anoles was published in the Bulletin of the Museum of Comparative Zoology, along with remembrances of Ken–who died 30 years ago–by Jerry Coyne, Chuck Crumly, Ray Huey, Eric Larson, Greg Mayer, and B Wu.

David Wake knew Ken Miyata, too, and here’s what he had to say: “Ken did an undergraduate honor’s thesis with me in MVZ.  He was far ahead of his time — we had no digital database but he wanted to do detailed mapping of some species so he selected Batrachoseps attenuatus and then laboriously went through the large MVZ collection.  He made a pin for each locality and on the pin recorded the MVZ catalogue number (or first in a series in the case of multiple specimens). Then he researched the exact locality, often going to field notes.  The result is still on the wall outside my office!  From time to time someone suggests taking it down, but it has now gained the status of historical document!  And it is a constant reminder to me of Ken and his enthusiasms and diligence.”

Anoles Link Spatially Distinct Terrestrial Food Webs – Part 1 Of 2

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LIke all the anoles in our study, a considerable fraction of A. equestris’ diet was derived from the flow of allochthonous resources into its habitat.

A. sagrei, probably the most common vertebrate in Florida perch low on trees making occasional forays to the ground to feed.

A. sagrei, probably the most common vertebrate in Florida perch low on trees making occasional forays to the ground to feed.

The ecological importance of small, terrestrial insectivores such as litter frogs and small geckos is a topic that I’ve been curious about for years. While my dissertation research does not include anything about it, I am still quite curious about how these small, diverse and abundant vertebrates fit into ecosystems. Anoles possess all of the attributes that seem to predispose them to strong interactions and soon after arriving in Miami to begin my Ph.D., I decided to launch a small, side-project using tried-and-true food web tools, stable isotopes and gut content analysis to try to illustrate if and how these small, rather inconspicuous predators might affect ecosystem structure and function. The results of this study were just published online in Functional Ecology.

Basically, we found that anoles couple adjacent food webs by consuming insects that move across habitat boundaries. While food web linkages are a potentially important ecological dynamic and our study yielded some unique findings, there are other bits of information for those more generally interested in anole biology. Therefore, I’ve decided to break this post into two parts. Part 1 deals with our primary findings and contextualizes them within current understanding of food web ecology linkages. It’s sort of a geeky treatment of the subject. Part 2 will illustrate some of the other data that we’ve collected that were not dealt with explicitly in the paper that will be of interest to, I suspect, AA readers.

Bidirectional trophic linkages couple canopy and understory food webs

Sean T. Giery,Nathan P. Lemoine, Caroline M. Hammerschlag-Peyer, Robin N. Abbey-Lee, and Craig A. Layman

1.  Cross-system resource flux is a fundamental component of ecological systems. Allochthonous material flows generate trophic linkages between adjacent food webs, thereby affecting community structure and stability in recipient systems.

2.  We investigated cross-habitat trophic linkages between canopy and understory food webs in a terrestrial, wooded, ecosystem in South Florida, USA. The focal community consisted of three species of Anolis lizards and their prey. We described interspecific differences among Anolis species in the strength and routing of these cross-habitat flows using stable isotope analysis, stomach content analysis, and habitat use data.

3.  All three Anolis species in this study consumed different prey, and occupied vertically distinct arboreal habitats. Despite these differences, carbon isotope and stomach content analysis revealed strong integration with understory and canopy food webs for all Anolis species. Modes of resource flux contributing to the observed cross-habitat trophic linkages included prey movement and the gravity-driven transport of detritus.

4.  Our study shows that terrestrial systems are linked by considerable bidirectional cross-system resource flux. Our results also suggest that considering species-specific interactions between predator and prey are necessary to fully understand the diversity of material and energy flows between spatially separated habitats.

MAp

The study system was dominated by St. Augustine grass and isolated Ficus trees.

Some basics – The community was composed of four anole species, Anolis sagrei, A. distichus, A. carolinensis*, and A. equestris. The study site was recently featured in AA. Generally, the purpose of the study was to describe variation among species in resource use using stomach contents, habitat use, and stable isotope analysis. But based on some initial observations and a bit of stable isotope data, we had considered that there might be a role for anoles in ecosystems via linking spatially distinct food webs. That is, anole diets might be sourced, in part, by primary production originating outside their respective microhabitats. Basically, we knew that anoles occupy distinct arboreal habitats, but when we examined the stomach contents of each, we found that some prey were from habitats spatially distinct from the ones used by each anole species (e.g., How do terrestrial grasshoppers get inside a canopy giant anole such as A. equestris?),which spawned a more in-depth investigation. Additionally, some initial stable isotope data strongly supported the same interpretation – that is, anole diets might be at least partially derived from allochthonous resources.

Ecological And Population Data On Some Little Known Ecuadorian Anoles

Ken Miyata–naturalist, fly-fisherman, and photographer extraordinaire–died tragically young 30 years ago at the age of 32. Among the many items of unfinished business was his gargantuan thesis, Patterns of Diversity in Tropical Herpetofaunas, 787 pages in length and entirely unpublished. The dissertation ranges far and wide over topics herpetological and ecological–check out the Table of Contents at the bottom of the post. Over the years, Ernest Williams tried to talk a number of scientists into guiding some of the chapters into print, but the task was too large and so it has remained shelf-bound.

Anolis peraccae. Photo by Luke Mahler.

Anolis peraccae. Photo by Luke Mahler.

After a recent trip to Ecuador, I happened to be looking at the thesis for other reasons (parts of it were incorporated into the description of A. lyra by Poe et al. in 2009) and came across Chapter 2. This multi-part section includes separate studies on the habitat use of three anole species at the Río Palenque field station (A. chloris, A. festae and A. peraccae) and population biology and dynamics of two other species elsewhere (A. boettiger and A. gemmosus). Miyata argued that little was known of the population biology of South American anoles. Thirty years on, the situation isn’t all that different.

Anolis chloris. Photo by Luke Mahler.

Anolis chloris. Photo by Luke Mahler.

As a result, the data presented by Miyata in 1983 are still very relevant today and deserve wider circulation. And for that reason, we decided to publish a lightly-edited version of parts Chapter 2 in the Bulletin of the Museum of Comparative Zoology, appropriate given that Miyata was a grad student in the Museum (this paper, like all other Bulletins of the MCZ, is available online). In addition, in an online supplementary file, a number of his friends) Jerry Coyne, Chuck Crumly, Ray Huey, Eric Larson, Greg Mayer and B Wu) provide reminiscences of Ken.

The paper ranges widely over matters of anole ecology, behavior, and population biology, providing data on five species for which almost nothing exists in the literature. The paper’s findings are summarized in the abstract:

Anolis festae. Photo by Luke Mahler.

Anolis festae. Photo by Luke Mahler.

Little is known about the ecology and natural history of South American anoles. This study reports the results of a variety of different studies on several relatively common species of Ecuadorian Anolis. In part I, habitat use and population density are compared among three species of Anolis that occur in sympatry at a number of sites in Ecuador. The three species—A. chloris, A. festae, and A. peraccae—are roughly the same body size. These species perch primarily on tree trunks, and A. chloris perches substantially higher than the other two species, which are similar in perch height. Large differences from one year to the next were observed both in mean perch height and in population densities.

Anolis gemmosus. Photo by Jonathan Losos

Anolis gemmosus. Photo by Jonathan Losos

In Part II, natural history, growth rates, and population densities are reported for two little known Anolis species, A. bitectus and A. gemmosus. Although the two species are from nearby regions and are similar in microhabitat use, they show more differences than similarities in most aspects of their biology. The species have similar ranges in active body temperatures, but A. bitectus is thermally passive, whereas A. gemmosus appears to thermoregulate. Populations of A. gemmosus tend to remain constant through time, whereas A. bitectus undergoes moderate population fluctuations. Both species exhibit little sexual size dimorphism, but in A. bitectus females are larger, and in A. gemmosus males are larger. Anolis bitectus has a fairly high characteristic growth rate, whereas that of A. gemmosus is quite low.

Microsoft Word - Table of Contents entire thesis.docxMicrosoft Word - Table of Contents entire thesis.docx

Evidence Suggests That Cloacas Carry Risk: Venereal Disease and Lizard-Parasite Coevolution

Anolis cristatellus wileyae - Does this pair need to worry about Cyrtosomum infections?

Anolis cristatellus wileyae – Does this pair from St. Thomas need to worry about Cyrtosomum infections?  Read on.

I wanted to write a few posts about parasites because hey – anoles have some really neat parasites! First up are the nematodes in the genus Cyrtosomum, which have been reported from several anoles and some other lizard taxa like Sceloporus and Cyclura. You might remember Cyrtosomum because C. penneri was the worm that AA-contributor Gerrut Norval and his colleagues used to infer that the Taiwanese population of A. sagrei originated from Florida and not Hawaii, something that wasn’t clear from sagrei molecular data (cool!).

Aside from pinworms (we’ll get to these another day), Cyrtosomum species are probably the most widespread nematode parasites in anoles. They’re really small (difficult to see without magnification), and occur in really large numbers in the lizards’ large intestines (many hosts have worm burdens of 100-200). Until recently, we knew very little about their life history – we knew that they could multiply within a single host individual via infective larvae, but we didn’t know how they moved between host individuals. Several authors (including Norval et al.) noted that Cyrtosomum species are only found in adult lizards, and suggested the possibility that these are sexually transmitted parasites.

Okay, let’s just take a moment… Worms, from sex. Yep.

Gabe Langford and his students tested this hypothesis in C. penneri, and

A Dearth Of Anole Talks At The Ecological Society Of America Meetings

In contrast to their ubiquity at the evolution meetings [1,2], anoles are barely in attendance at the enormous Ecological Society of America meetings. Last year there were but three (including the famous “Kitty Cam” talk), and this year the dearth is worse, with but a single talk this week in Minneapolis. So, here it is, by Rich Glor and part of the symposium on “Coexistence of Closest Relatives: Synthesis of Ecological and Evolutionary Perspectives”:

Patterns of Anolis lizard coexistence across a complex island landscape

Thursday, August 8, 2013: 3:40 PM

M100EF, Minneapolis Convention Center

Richard E. Glor , Biology, University of Rochester, Rochester, NY

Background/Question/Methods

In species rich adaptive radiations, complex patterns of species coexistence result from speciation and natural selection acting over evolutionary time. In Anolis lizards, similar communities of 4-6 arboreal microhabitat specialists that partition available perches by size and height have evolved independently on each of the four largest Caribbean islands. These well-studied communities of anole ‘ecomorphs,’ however, account for only a fraction of anole species diversity. Most anole diversification has occurred subsequent to microhabitat specialization, resulting in complexes of closely-related species that belong to the same ecomorph but appear to be allopatrically or parapatrically distributed in different macrohabitats. We use integrative analyses that combine thousands of anole species occurrence records from public biodiversity databases, GIS environmental data, molecular phylogenetic and population genetic data, and data on phenotypic variation to test three predictions about anole species co-occurrence across the island of Hispaniola: (1) local communities tend to be comprised of microhabitat specialists that evolve relatively early in anole radiations, (2) closely-related or incipient species tend to be allopatrically or parapatrically distributed in distinct macrohabitats, and (3) habitat filtering in extreme habitats leads to communities in which species that are more closely related to one another than expected by chance.

Results/Conclusions

Our analyses support the prediction that local communities tend to be comprised of phenotypically distinct microhabitat specialists that evolved early in anole radiations and are relatively distantly related. Our analyses also support the prediction that closely-related species tend to be allopatrically or parapatrically distributed in distinct macrohabitats. This result, together with a growing body of molecular population genetic data, suggests that both geographic and ecological processes play an important role in recent anole speciation events, and dictate patterns of co-occurrence in closely related anole species. Finally, we find preliminary support for the importance of habitat filtering by recovering evidence for a latitudinal gradient in the phylogenetic composition of local communities whereby communities at the highest altitudes tend to include more closely-related species than communities at lower altitudes. This pattern is particularly evident in the Cordillera Central, where communities of closely related and ecologically unusual species tend to occur at the highest altitudes.

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