What Makes Anolis Communities Complete?

One of my favorite graphic representations of a typical anole community is the one where all ecomorphs are hanging out together in a tree and a scrub next to said tree. Each ecomorph has its structural microhabitat place and they are all spaced out evenly across the tree to represent competition. Originally the figure was published by Williams (1983) and then modified later on. Arriving on the Greater Antilles, one thus expects to promptly be able to say hi to all these ecomorphs at the next best tree. Well, from my personal experience, I can tell you that this is unfortunately not the case.

Idealized representation

Localities where all ecomorphs are found together are scarce, and all of them are famous, having served as field sites for the most groundbreaking of anole discoveries. But what about the rest of them? Something must prevent the co-occurrence of ecomorphs in all these other places. This was noted before: Losos (2009) remarked that all utilized structural microhabitats exploited by all ecomorphs are present throughout the islands, so “complete” ecomorph communities should also be able to occur everywhere.

A common explanation for the absence of certain  “functional types” (= Anolis ecomorphs) from local communities is a process that is called “filtering.” Modern community assembly theory distinguishes two such types of filters: 1. Biotic interaction filters and 2. Environmental filters.

 Filters

Biotic filtering involves competitive exclusion: For anoles this phenomenon caused ecological speciation which led to the convergent evolution of the ecomorph communities. But biotic filtering should not be expected to occur at this stage of the radiation: Different ecomorphs are not competing for the same structural microhabitat niche in different localities. This leaves environmental filtering. In our study recently published in Ecology and Evolution, on which I am reporting here, we tested whether environmental filtering could be a possible explanation for the absence of ecomorphs in local communities.

First, we modeled Anolis ecomorph community completeness by constructing environmental niche models for each ecomorph (the sum of species belonging to that ecomorph) on each island. These models were then overlaid for all ecomorphs per island.  ECC map

 The map for ecomorph community completeness shows a very patchy distribution of areas where all ecomorphs are expected to occur. Comparisons of environmental niches among these islands revealed that only Hispaniola and Cuba have their complete Anolis ecomorph communities occurring in a similar bioclimatic parameter space.

This patchiness could be explained by elevation for all islands except Jamaica: the Anolis community completeness map strongly resembles the topographic relief of the Greater Antilles. Looking more closely into the climatic parameters, Jamaica has much lower daily and annual temperature ranges which are also not related to the island’s elevation, whereas in the rest of the Greater Antillean islands, they are. Occurrence probability of ecomorphs seems to be coupled to environmental parameters, which explains why some ecomorphs are “filtered out“ in some locations: they do not encounter a favorable environment there.

Since I mentioned initially that filtering relates to “functional types” (not species), the filtering must be a result of certain functional properties of the Anolis ecomorphs’ phenotype. We wanted to take the study a step further and actually investigate one (among many) possible functional trait: body mass.

Advice Needed: Field Sites for A. sagrei in Florida

Anolis sagrei. Photo by Janson Jones.

I’m planning an in-depth behavioral study of Anolis sagrei for the summer and need your help finding suitable field sites in Florida.

My ideal location would have the following traits:

– Abundant A. sagrei in an area large enough to support at least 50 adult males

– Relatively open understory

– Not heavily trafficked by people (I’d like to minimize the frequency of behavioral trials being disrupted by inquisitive passersby), but still safe to work in

– Management receptive to researchers

Does anyone know of protected areas, biological or agricultural field stations, or other underutilized green spaces that might fit the bill? I’m open to locations throughout the state.

Thanks in advance for any suggestions!

Exploring the Anolis Y Chromosome

Sex chromosomes have historically been identified by inspecting chromosome spreads under a light microscope and looking for a morphologically distinct or heteromorphic pair of chromosomes – typically and X and Y or a Z and W. However, heteromorphic sex chromosomes are absent in many animal groups, particularly fish, amphibians, and lizards, making it difficult to determine whether a species with genetic sex determination has an XY or ZW system. As a consequence, the study by staustinreview.com of sex chromosome evolution in clades in which cryptic or homomorphic sex chromosomes are prevalent has been hampered by a lack of identified sex chromosomes in these groups. New methods are needed to find the sex chromosomes in these species and increase our understanding of homomorphic sex chromosome biology, the evolution of sex determining systems, and patterns of sex chromosome evolution overall.

David Zarkower and I have a paper in press at Molecular Ecology Resources that uses high-throughput DNA sequencing to identify sex-specific genetic markers as a means to reveal sex chromosome systems in species that lack heteromorphic sex chromosomes. We are using a newly developed DNA sequencing technique called restriction site associated DNA sequencing or RAD-seq. RAD-seq sequences the DNA flanking very specific DNA sequences (restriction enzyme recognition sites) scattered throughout the genome, generating tens of thousands of genetic markers. RAD-seq is a powerful technique for exploring genetic variation in ‘nonmodel’ species because it does not require a fully sequenced genome, requires relatively modest sequencing capacity, and can detect even minor genetic differences among individuals. We are using RAD-seq to 1) identify sex-specific molecular markers (i.e., bits of DNA found in individuals from one sex but not the other), and 2) using these markers to determine whether a species has XY or ZW sex chromosomes. Species with male-specific markers will have an XY system while species with female-specific will have a ZW system.

We are interested in using RAD-seq to screen various vertebrate species for sex chromosomes, but first wanted to validate the technique using a species with a known sex-determining mechanism. We chose the green anole (Anolis carolinensis) because its X and Y chromosomes are small and homomorphic. Therefore A. carolinensis sex chromosomes should provide a rigorous test of this technique and success with Anolis suggests there may be broad utility using this technique in other groups with homomorphic sex chromosomes.

We performed RAD-seq on seven male and ten female A. carolinensis and recovered one male-specific molecular marker. We confirmed that the marker was male-specific using PCR and also found that this genetic marker is conserved in some additional Anolis species, confirming homology among the Y chromosomes of these species (Anolis sex chromosome homology has been discussed previously on Anole Annals 1, 2). These results highlight the potential utility of RAD-seq as a tool to discover the sex chromosome systems of large numbers of species in a rapid, cost-effective manner.

PCR validation of the male-specific RAD-seq marker in Anolis carolinensis.

PCR validation of the male-specific RAD-seq marker in Anolis carolinensis.

In addition to learning about Anolis sex chromosomes the male-specific molecular marker we identified can be used to sex individuals of many Anolis species using a simple PCR-based assay, particularly species in the A. carolinensis group and in the Norops clade. This enables identification of an individual’s sex prior to the onset of secondary sexual characteristics, for example in embryos, thereby aiding developmental studies of sexually dimorphic phenotypes. The importance of sexual dimorphism to Anolis ecology and evolution has been examined previously (1, 2, 3, 4), but there is certainly much more to learn, particularly about how sexually dimorphic traits develop and evolve. The ability to sex Anolis embryos is an important step to advance this research.

Phylogenetic relationships among sampled species illustrating the sex-specific amplification of the gene rtdr1y in selected anole species. The autosomal gene kank1 was used as an internal positive control in all reactions. Bands labelled with ‘NS’ are nonspecific PCR products.

Phylogenetic relationships among sampled anoles illustrating the sex-specific amplification of the gene rtdr1y in selected anole species. The autosomal gene kank1 was used as an internal positive control in all PCR reactions. Bands labelled with ‘NS’ are nonspecific PCR products.

Nine Caribbean Skinks Petitioned for Inclusion on Endangered Species List

A while back, we reported on a monograph Hedges and Conn that described an enormous number of new skink species (35) from the Caribbean. Now efforts are being made to prevent some of these species from going extinct. The Center for Biological Diversity has just filed a petition with the U.S. Fish and Wildlife Service asking that nine Caribbean skink species be placed on the Endangered Species List. Those species are: Culebra Skink (Spondylurus culebrae), Mona Skink (Spondylurus monae), Monito Skink (Spondylurus monitae), Lesser Virgin Islands Skink (Spondylurus semitaeniatus), Virgin Islands Bronze Skink (Spondylurus sloanii), Puerto Rican Skink (Spondylurus nitidus), Greater Saint Croix Skink (Spondylurus magnacruzae), Greater Virgin Islands Skink (Spondylurus spilonotus) and Lesser Saint Croix Skink (Capitellum parvicruzae).

A press release from the CBD explains all:

“The Center for Biological Diversity filed a formal petition today seeking Endangered Species Act protection for nine newly identified species of skinks found only in Puerto Rico and the Virgin Islands. These rare lizards with smooth skins are on the knife’s edge of extinction due to introduced predators and habitat destruction. Reptiles around the globe are in the midst of an extinction crisis with roughly 1 in 5 species considered endangered or at risk of disappearing.

Puerto Rican skink
Puerto Rican skink photo © Puerto Rico Wildlife/Alfredo Colón (alfredocolon.zenfolio.com). Photos and maps are available for media use.

“Time is running out for these lizards,” said Collette Adkins Giese, a Center biologist and lawyer focused on protecting reptiles and amphibians. “The Caribbean is home to extremely rare animals found nowhere else in the world, but too many have already gone extinct. To save these skinks, we need to get them protected under the Endangered Species Act.”

Scientists recently recognized the nine petitioned skinks, along with dozens of others on Caribbean islands. The scientists initiated their study after finding unusually large genetic differences among populations of these skinks on different islands in the Caribbean. All of the newly identified endemic Caribbean skinks are near extinction (or already extinct) due to introduced predators like mongooses and cats, as well as large-scale habitat destruction for development and agriculture.

This loss is alarming because reptiles play important roles as predators and prey in their ecosystems and they’re valuable indicators of environmental health. The animals in today’s petition will reap life-saving benefits from the Endangered Species Act, which has a 99 percent success rate at staving off extinction for species under its care.

“Skinks have a slow-moving curiosity and are not adapted to fast predators such as the mongoose, introduced by humans,” said Dr. Blair Hedges of Pennsylvania State University, the lead author of the 2012 study that recognized the petitioned species. “The survival of these skinks depends on the special measures of protection that only the Endangered Species Act can provide.”

Although reptiles have been around for hundreds of millions of years and survived every major extinction period, now, due largely to human impacts, they’re dying off at up to 10,000 times the historic extinction rate. About 20 percent of reptiles in the world are endangered or vulnerable to extinction. Within the Caribbean, scientists estimate that reptiles have levels of endangerment that are at or near the highest levels worldwide.

The Center was joined in its petition for these nine skinks by Dr. Renata Platenberg, an ecologist specializing in Caribbean reptiles.

Background
The petitioned-for Caribbean skinks, which can grow to be about 8 inches long, are unique among reptiles in having reproductive systems most like humans, including a placenta and live birth. They have cylindrical bodies, and most have ill-defined necks that, together with their sinuous movements and smooth, bronze-colored skin, make them look like stubby snakes.

Four of the species for which we petitioned are found within the territory of Puerto Rico: the Culebra skink (Culebra and the adjacent islet of Culebrita), Mona skink (Mona Island), Monito skink (Monito Island) and Puerto Rican skink (Puerto Rico and several of its satellite islands). The remaining five are found in the Virgin Islands: the Greater St. Croix skink (St. Croix and its satellite Green Cay), Lesser St. Croix skink (St. Croix), Greater Virgin Islands skink (St. John and St. Thomas), Lesser Virgin Islands skinks (St. Thomas and two adjacent islets, several British Virgin Islands) and Virgin Islands bronze skink (St. Thomas and several of its islets, several British Virgin Islands).

Eight of the nine petitioned-for species fall within the genus Spondylurus, and one falls within the genus Capitellum. The genus Spondylurusincludes what are now known as the Antillean four-lined skinks because of the four major dark stripes on their back and sides. Skinks in the genus Capitellum are called the Antillean small-headed skinks and have small feet and short heads, lacking dark dorsolateral stripes.”

The entire petition can be downloaded from the CBD’s website.

Mindo Amphibian and Reptile Book Now Available in US

While on the theme of Ecuador from yesterday’s post on 25 newly described Ecuadorian herps…Word has just reached AA‘s ears that the fabulous Amphibians and Reptiles of Mindo is now available in the U.S. You can order it from Eagle Mountain Publishing, which also sells a lot of great old reptile and amphibian books at rock bottom prices.

In celebration of this great event, we’re re-printing the review posted a few months ago:

The team at Tropical Herping has done it again! This time, a fabulous, lavish, luscious, information-packed guide to the spectacular herpetofauna of Mindo Parish, Ecuador. Originally available online, the book is now available in print. I had the privilege of writing the foreword, appended below. More information is available on the TH website, as well as an order form.

Foreword:

Small in size, but a global giant in biodiversity, Ecuador is awash in all manner of fauna and flora. Birds, butterflies, trees—the country is a hotspot for just about everything. But no group of organisms is more beautiful, more charismatic, more scientifically captivating than Ecuador’s reptiles and amphibians. The Amazon rainforest dominates the attention of the public, but other parts of the country, especially the mountainous regions, are just as biologically rich. One such area is the small parish of Mindo in Pichincha Province, home to 102 species of creepy crawlies. And what an ensemble! Brilliant colors, toxic skin and venom, sweet serenades, menacing looks, gorgeous displays—this region is an encyclopedia of herpetology in just 268 square kilometers.

Field guides play an essential role in making the fauna and flora of an area widely accessible. They are at the front line of nature education and conservation, the place where the fruits of scientific exploration are distilled, synthesized, packaged, and presented to the public at large. Since the time of Roger Tory Peterson, field guides have played another role, being a venue for beautiful, yet accurate, scientific illustration, allowing readers to not only understand the identifying marks of each species, but also to appreciate them esthetically.

Despite its bountiful herpetofauna, until now no field guides existed for Ecuador’s amphibians and Reptiles. The Tropical Herping team has brilliantly stepped into this void, producing a guide to the herps of Mindo that hopefully will serve both as a model of how guides should be produced and an inspiration to the production of similar efforts elsewhere in Ecuador and beyond. The Amphibians and Reptiles of Mindo is particularly notable in three respects. First is the breadth and depth of information provided for each of Mindo’s species. These authors know their fauna in exquisite detail and have synthesized that knowledge in a clear and lucid manner. The inclusion of frog calls, recorded by the authors themselves, is an added bonus bridging the paper and digital eras. Second, the public often does not understand the connection between scientific research and the information presented in field guides, magazine articles and nature documentaries. Unlike most field guides, The Amphibians and Reptiles of Mindo makes this link crystal clear, providing citations so that readers know where to turn to learn more. Indeed, especially impressive is the fact that the authors did a great deal of field work themselves to round out knowledge of these species, presenting that information for the first time here. Finally, third, the book is simply beautiful. The photographs are simply stunning and the maps and other illustrations lovely as well.

The publication of The Amphibians and Reptiles of Mindo could not come at a better time. The Mindo region is a microcosm for all that ails the natural world. Deforestation, habitat fragmentation, pollution, overharvesting—all are threats. Mindo has one thing going for in its favor—it has become a nature vacation travel destination, providing jobs and economic rationale for preserving natural habitats. But, ecotourism can be a two-edged sword, as people and development are drawn to the area with potentially negative consequences. Mindo has the opportunity to show how responsible stewardship can be mutually beneficial to man and nature, and this lovely book shows what is at stake. Three cheers for the three authors of this magnificent volume. Long live the herpetofauna of Mindo!

Ecuadorian Anoles on BBC News

otongae

During the last five years, herpetologists at the Museo de Zoología QCAZ, Pontificia Universidad Católica del Ecuador (PUCE), have discovered and described 35 new species of amphibians and reptiles, some of which are anoles. BBC news recently posted a photographic article on this work, which was funded by the Ecuadorian government and PUCE. Anolis otongae and A. podocarpus are some of the recently discovered species featured in that article.

The Museo de Zoología QCAZ also maintains ReptiliaWebEcuador, a website on Ecuadorian reptiles with tons of information in Spanish, including pictures, maps, free downloads, and more. Visit us if you want to know more about Ecuadorian anoles.

Film on Haiti Herpetological Exploration to Premiere at Film Festivals

httpv://www.youtube.com/watch?v=OChvJfMG9aI#t=86

We’ve reported previously on the expedition to Haiti led by Blair Hedges of Penn State that led to the rediscovery of A. darlingtoni as well as many other important herpetological finds. Now a film about this expedition, the sad state of Haiti’s environment and efforts to protect it and its fauna is being showcased at several film festivals around the world.

Extinction in Progress premieres at the Environmental Film Festival in Washington, D.C. on March 19th and subsequently will be shown at festivals in Seoul (South Korea), Torino (Italy), Zaragoza (Spain), Goías (Brazil) and Prizren (Kosovo).

extinction in progress

Yes, that’s Anolis rupinae!

The Green Anole, Newsletter of the LBJ Wildflower Center Young EcoExplorers

Winter2008

The Green Anole is the quarterly children’s newsletter written for EcoExplorer members of the Lady Bird Johnson Wildflower Center in Austin, Texas.

Reconstructing the History of Anole Sex Chromosomes

Gorman_Dominica_1965_Anolis_oculatus

George Gorman in Dominica

In the 1960s and 70’s evolutionary cytogenetics experienced a remarkable burst of interest and scholarship. Thanks largely to the efforts of George Gorman (at right) and others working at the Museum of Comparative Zoology, anoles played a central role in this research (some historical detail has previously been posted on AA). Among their findings was the occurrence of heteromorphic sex chromosomes, sex chromosomes that are visibly distinguishable from each other under a microscope, in several Anolis species but not others. Furthermore, Gorman and colleagues discovered that those Anolis species with heteromorphic sex chromosomes all had male heterogamety, with some having an XX/XY system while others had an XXXX/XXY system. Chromosomes from nearly 100 Anolis species were examined during this period and about 1/3 of those species had heteromorphic sex chromosomes. Interest in chromosome evolution waned in the 1980’s as DNA sequence data became increasing accessible, but there has been a recent resurgence thanks, in part, to sex chromosomes.

Anole – Gecko Habitat Overlap

Although it may seem paradoxical given their natural history, I often see nocturnal geckos basking during the day. They usually occupy trunks between 1-3m, which is prime trunk-ground and trunk ecomorph territory. In Miami we have two of the former (A. sagrei and A. cristatellus), and one of the latter (A. distichus). Here is a photo fresh from today showing such overlap in habitat use. I have yet to see any interactions between anoles and geckos in Miami, has anyone else observed any here or elsewhere?

Brown anole Anolis sagrei and gecko (Hemidactylus sp., probably mabouia) sharing habitat space_caption

Cuban brown anole (Anolis sagrei – left) and gecko (Hemidactylus sp., most likely H. mabouia – right) sharing habitat space

Of note – this was taken 23rd Feb 2014 in south Miami (FL) at roughly 1:30pm. It’s hot and sunny.

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