In a somewhat autobiographical romp through the history of species delimitation, David Hillis, in a recently published article in the Journal of Herpetology, details the state-of-the-field in terms of phylogenetic and species delimitation, detailing both the many advances that have been made over the last few decades, but also pointing out where things are out of whack and need some recalibration. There’s much more to the article than the figure above, but that’s a good place to start!
Author: Jonathan Losos Page 14 of 130
Professor of Biology and Director of the Living Earth Collaborative at Washington University in Saint Louis. I've spent my entire professional career studying anoles and have discovered that the more I learn about anoles, the more I realize I don't know.

Anolis landestoyi. Photo by Miguel Landestoy.
As reported by Science Daily:
Elevation matters when it comes to climate change, deforestation and species survival
- Date:
- February 25, 2019
- Source:
- University of Toronto
- Summary:
- A study examining the impact of deforestation on lizard communities in the Dominican Republic demonstrates differing outcomes at different elevations. In the lowlands, deforestation reduces the number of individuals, but not which species occur in an area. In the highlands, it’s the opposite. When the forest is cut down at higher elevations, the newly created pastures become filled with species found in the warmer lowlands. But locally adapted mountain lizards cannot survive as temperature rises.
University of Toronto student George Sandler was shocked to see the rainforest floor suddenly come to life around him, as if in a scene from an Indiana Jones movie.
“The forest floor started rustling around me,” says Sandler, “as dozens of crabs emerged from holes and crevices. Some were huge, the size of dinner plates. I even spotted a hermit crab climbing up a tree, lugging its heavy shell along with it.”
But Sandler wasn’t in the field to study crabs. He was in the Dominican Republic to take a census of the region’s Anolis lizard species for a study on the effects of deforestation being conducted by researchers Luke Mahler, Luke Frishkoff and collaborators. In the Caribbean nation, deforestation is the main form of natural habitat loss as residents cut down rainforest in order to produce charcoal, as well as create pastures for livestock and farmland for crops.
It is no surprise that deforestation has a profound effect on biodiversity; scientists have been studying this problem around the globe for decades. What is surprising is the difficulty they still face in making detailed predictions about which species survive, especially in relation to other factors such as climate change and natural local conditions.
Now, using the data collected in the census, the research team has discovered details about how Anolis lizards are being affected by the loss of their habitat.
“When it comes to predicting the effects of deforestation,” says Mahler, “elevation matters.”
Mahler is an assistant professor in the Department of Ecology & Evolutionary Biology (EEB) in the Faculty of Arts & Science at the University of Toronto. Frishkoff led the research while he was a postdoctoral fellow in Mahler’s lab at U of T and is lead author of the paper describing their findings, published today in Nature Ecology & Evolution; he is currently an assistant professor at the University of Texas at Arlington. Sandler and researchers from the National Museum of Natural History in Santo Domingo were also co-authors.
Mahler and Frishkoff analyzed populations of lizards in both lowland and highland regions affected by deforestation. Generally, the lowlands are warmer than the highlands due to altitude; also, forest canopy blocks direct sunlight, making forests at any altitude cooler than their immediate surroundings.
“It turns out that deforestation changes lizard communities in fundamentally different ways in the lowlands as compared to the highlands,” says Mahler. “In the lowlands, deforestation reduces the number of individuals, but not which species occur in an area. In the highlands, it’s the opposite.”
“When the forest is cut down at higher elevations,” says Frishkoff, “the newly created high elevation pastures become filled with species we saw down in the warmer lowlands. But, the locally adapted mountain lizards cannot survive.”
The invasion into the highlands by lowland-dwelling lizards was made possible by a combination of human activity and natural factors; i.e. deforestation and elevation respectively. Thanks to the altitude, the temperature of deforested fields in the highlands was comparable to the temperature of forested lowlands.
As it is in many regions around the world, the problem of deforestation in the Dominican Republic is dire. In 2016, Mahler announced the discovery of a previously unknown chameleon-like Anolis lizard on the island of Hispaniola. In the paper describing the discovery, Mahler and his co-authors recommended that the new species, dubbed Anolis landestoyi, be immediately classified as critically endangered because the lizard was threatened by illegal clear-cutting in the region.
Unlike the crabs that crowded around Sandler in the rainforest, the lizards were more elusive and difficult to survey. In order to obtain accurate counts, the students employed a technique known as mark-resight.
“We hiked out to our designated plots,” says Sandler, who was an undergraduate student while conducting the field work and is currently an EEB graduate student at U of T. “Then we walked around looking for lizards. We carried a paint spray gun filled with a non-toxic, water soluble paint — a different colour for each of the six observation periods. If we saw a lizard we would note the species, if it had any paint on it already, and the colour of the paint. Then we would spray the lizard with the paint gun we were carrying, a task that was a little tricky with some of the more skittish species!”
Paint on a lizard indicated that it had already been counted; and the number of unpainted lizards that were observed during each period allowed the researchers to calculate how many lizards were going uncounted.
“It’s not your typical summer job,” says Mahler. “Each survey is essentially a game in which you try to find all the lizards in an area and zap them with paint. It’s a messy affair, but we get great data from it.”
“Our results help us better understand the likely consequences of climate change and how it will interact with human land-use,” says Frishkoff.
For lowland forest Anolis lizards, deforestation just means a decline in abundance or relocating to the highlands. But for highland species, the situation is more critical. Unlike their lowland cousins, they have reached high ground already and in the face of deforestation have nowhere to go — a situation facing more and more species around the world.
“Our data suggest that while many lowland Anolis species might not be seriously affected by deforestation and the gradual warming brought about by climate change,” says Frishkoff, “the opposite is true for the unique mountain lizard species which do not tolerate land-use change well, and which are already on the top of the island.
“Land-use and climate change are a double whammy for these species. If we cut down the mountain forests these lizards have nowhere left to go. Gradual warming might push species up slope, but when you’re already at the top of the mountain, you can’t move any higher.”
Story Source:
Materials provided by University of Toronto. Note: Content may be edited for style and length.
Journal Reference:
- Luke O. Frishkoff, Eveling Gabot, George Sandler, Cristian Marte, D. Luke Mahler. Elevation shapes the reassembly of Anthropocene lizard communities. Nature Ecology & Evolution, 2019; DOI: 10.1038/s41559-019-0819-0
Our recent mention of Richard Etheridge’s death reminded us that we’d never noted the passing of Lourdes Rodriguez Schettino nearly two years ago, in March of 2017.
Lourdes was an expert on the lizards of Cuba, particularly the iguanid lizards, about which she truly wrote the book. She also was the lead author on a lovely book on Cuban herpetology (below).
Sadly, I am unaware of any obituaries of Lourdes in English, but the Instituto de Ecología y Sistemática, where she worked for many years, has a nice summary of her life’s work, and there is a brief notice in Revista Colombiana de Ciencia Animal.
Word has belatedly reached Anole Annals’ ears that, sadly, Richard Etheridge passed away last month at the age of 89. Richard was not only a titan of lizard systematics, but the founder of anole systematics. Amazingly, the classic worked that is the foundation of all that has followed in anole systematics was his never-published PhD dissertation. Bobby Espinoza and Kevin de Queiroz wrote a biography of Richard’s career that was published a decade ago in Copeia and that, happily, is freely available.
This just in from the Daily Herald, the self-proclaimed leading for St. Maarten and the northeastern Caribbean:
Research ongoing on status of endemic Bearded Anole Lizard
Bearded anole with a dorsal stripe, dark bands and a turquoise spot around the eyes foraging on St. Maarten. (Joost Merjenburgh photo)
COLE BAY–Nature Foundation, in collaboration with Reptile, Amphibian and Fish Conservation Netherlands RAVON, is researching the local status of the bearded anole (Anolis pogus) lizard, also known as the Anguilla Bank bush anole.
The research is aimed to better understand the current abundance, distribution and habitat preference of the bearded anole, as presently only little data is available. The data will provide insight about the population trend of the species and are needed for its International Union for Conservation of Nature (IUCN) red-list re-assessment. With this information measures can be made to conserve and protect the species.
Merjenburgh is investigating a total of 100 80-square-metre location plots each. All bearded anole are counted and measurements are taken of tree canopy cover and habitat type. Until now the bearded anole population seems to be well distributed on the island. The bearded anole seems to be quite abundant when searching in the right habitat and is mostly absent in residential areas.
Bearded anoles have a special appearance which can differ between individuals. Some individuals have a turquoise-coloured spot around the eye, others have a dorsal stripe or dark bands across the neck and body, and some have all characteristics. The reason there is so much difference in pattern and coloration is that they use their appearance for camouflage and communication purposes.
The bearded anole can also use the dewlap, a flap of skin that hangs beneath the lower jaw, to communicate. Although both genders have a dewlap, the one on the female is usually much smaller and less colourful than the one on the male. The male extends his dewlap mainly to attract females during mating season or to confront other males when they enter his territory, explained Merjenburgh.
Their diet, like other anoles, consists mainly of small arthropods – for example, cockroaches, crickets and spiders are their favourite foods – and occasionally flowers and fruits.
This species also falls prey to predators sometimes. One of those predators is the mongoose, which is an introduced species. The mongoose eats every native reptile species on the island and is probably the reason the only native snake, the leeward island racer, has probably become extinct on St. Maarten, said Merjenburgh.
A new book on lizard behavior has an anole gracing the cover (who can guess the species?) and an all-star cast of contributors, including a number familiar to AA faithful (see Table of Contents below). The book is due out in a month, but you can go ahead and pre-order it. Sadly, it’s priced in the stratosphere: $159 for the 400 page volume. Let’s hope for a paperback edition!
From the archives. One of the greatest <i>Anole Annals</i> posts of all times, because why not?
“You’ve gotta see this!” my fiancé Mark called to me one morning. He was outside, which could mean only one thing: a wildlife encounter was underway. Living in a semi-rural neighborhood in Florida, you never knew what you would see, from a mated pair of Sandhill Cranes walking down the street with their young, to Gopher Tortoises excavating burrows in the front yard.
I walked downstairs to the concrete area under our elevated house where Mark was staring at something on the ground. I looked down to see a frog (Cuban Treefrog) with the tail of an A. carolinensis protruding from its gullet.
“I knew that lizard,” Mark said forlornly.
I don’t know how long this has been the case, but you can download it, or chapters within it, at the Biodiversity Heritage Library.
Can anyone advise our correspondent in North Carolina who writes about the anole above with enormous growths that may be endolympathic sacs out of control:
Maybe you have some ideas about this growth, which I originally thought were calcium storage seen in anoles, but this just looks like it is going to pop any second. I bred this girl about 2.5 years ago and she usually lives with my boss unless he goes on vacation, which is when I take care of her. I also still have her Dad and sister (long story and I’ll never do it again!). I let the other eight (!) babies that I raised and Mom, who was used to the wild, go. This girl has always been very hyper and green all the time and has laid a bunch of eggs this past spring and started getting these pockets (thought calcium). But now the one side is sooo big! She gets almost the same care as my two (Dad and sister), except maybe a little less sunlight (mine are at a window). I was worried it is an abscess or parasite or something. I asked someone at the NCSU vet school and he wasn’t sure. Oh, she hasn’t been eating well, either, the last few days.

For some reason, this obituary of Ernest Williams is going around the internet again, 9+ years after it was published in the Harvard Gazette.
Ernest Edward Williams

Faculty of Arts and Sciences — Memorial Minute
At a Meeting of the Faculty of Arts and Sciences on May 19, 2009, the following Minute was placed upon the records.
Ernest Williams was a man of many contrasts. Biology at Harvard in the third quarter of the last century was full of outsized personalities—titans in the field with strong opinions and no reservations about expressing them. In such company, Williams appeared a wallflower, seemingly wishing to be anywhere but in the midst of their arguments. Yet, one-on-one, Williams had an incisive wit and a dry sarcasm—discussions with him were always stimulating and provocative as he never missed a chance to challenge one’s thinking, sometimes quite pointedly.
To some, Williams’s work came across as old-fashioned. His subject, systematics — the study of the evolutionary relationships of species—is among the oldest in science, and his papers — florid and opinionated and, above all, long—recalled an approach to scholarship no longer in vogue. Yet much of his work was boldly innovative; some papers are still widely cited, and in several cases his work was well ahead of its time, presaging approaches to the study of evolutionary biology that were not to catch on for several decades.
Ernest Edward Williams was born January 7, 1914, in Easton, Pennsylvania, the only child of middle-aged parents. Like many boys, particularly of that time, he grew up loving nature and spent many hours capturing salamanders and other creatures. After attending Lafayette College, Williams joined the Army, serving in Europe during World War II. Upon his return, Williams entered graduate school at Columbia University, where he was the last graduate student of the great anatomist William King Gregory.
Williams’s doctoral thesis focused on the structure of the neck vertebrae of turtles and how variation among species reflects their evolutionary heritage. The work demonstrated the combination of careful attention to detail with the ability to interpret results in the broader context that was to characterize Williams’s career. More than fifty years later the work is still foundational in understanding the evolution of turtle diversity.
In 1950, after completing his degree, Williams moved to Harvard, where he initially served as a laboratory coordinator for the anatomy course of the legendary paleontologist Alfred Sherwood Romer, then subsequently was appointed as an assistant professor and made coordinator of a General Education course on evolution. The Museum of Comparative Zoology’s Curator of Herpetology, Arthur Loveridge, retired in 1957, and Williams was appointed to take his place. In 1970 Williams rose to the rank of professor and in 1972 became Alexander Agassiz Professor of Zoology.
Williams initially focused on continuing his work on turtle systematics, leading to a series of publications including a still-important treatise published with Loveridge in 1957. Williams soon realized, however, that the museum’s collections were inadequate for the detailed analysis he conceived, which required large samples from many populations. This recognition that the museum’s herpetological collections were wide in scope, but lacking in depth, led Williams in two directions. First, it compelled him to work greatly to expand the Herpetology Department’s holdings, ultimately leading to a quadrupling of the department’s collections (to more than 300,000 specimens) by the time he retired as curator in 1980, making the Museum of Comparative Zoology (MCZ) one of the greatest herpetological repositories in the world. Second, it led Williams’s attention to focus on lizards in the genus Anolis, a very species-rich group from the Caribbean and Central and South America. A previous curator of herpetology and director of the MCZ, Thomas Barbour, had extensively collected anoles in the Caribbean; Williams, whose focus was much more evolutionarily-oriented than most systematists of the day, recognized that this group could be a model for studying large-scale evolutionary and biogeographical phenomena.
And, indeed, they were, and still are. Williams recognized that anoles have diversified for the most part independently on each of the major islands of the Caribbean (Cuba, Hispaniola, Jamaica, and Puerto Rico); most remarkably, the end-result of these evolutionary radiations has been very similar, with the same set of habitat specialists — which Williams labeled “ecomorphs,” a term now widely employed in evolutionary biology — evolving independently on each island. Such convergence of entire faunas is a rare phenomenon and Anolis has become a textbook example.
Williams’s work on anole evolution synthesized a wide variety of fields, including biogeography, functional morphology, population genetics, behavior, and ecology. Yet, Williams was a systematist by training, with little background in most of these areas. The primary means by which Williams orchestrated this broad-based investigation of anole diversity was his ability to identify the best organismally minded graduate students in Harvard’s biology department, regardless of their specific interests. As a result, the list of Williams’s graduate students reads like a Who’s Who in ecology and evolutionary biology.
At a time when the MCZ’s curators had little say in curricular matters, Williams pioneered a highly popular course on vertebrate biology. This course, which Williams taught for many years, helped keep organismal biology alive at Harvard and was a crucial step in the creation of the Department of Organismic and Evolutionary Biology (OEB).
Williams was not the most gentle-hearted of advisors. Gruff and very critical, he had high standards, expected students and colleagues to meet them, and was not shy about letting them know when they had not. Words of praise were not handed out liberally, but were cherished when received. With undergraduates, however, Williams showed a different side, being supportive and encouraging when needed and available at any time for discussion and advice. A number of Harvard undergraduates who worked with him have gone on to become evolutionary biologists, and several have continued to work on anoles, in many cases following up on ideas he initially conceived.
Williams remained active after his retirement from the Harvard faculty in 1984, continuing his evolutionary and systematic studies. Eventually, he moved back to his native Pennsylvania, where he died in 1998, taking his encyclopedic font of knowledge with him. Nonetheless, his spirit and ideas live on in the form of his many academic descendants, which include many prominent active scientists, several members of the U.S. National Academy of Sciences, a high-ranking U.S. government official, and the current curator of herpetology of the MCZ.
Respectfully submitted,
A. W. Crompton
Karel F. Liem
Jonathan Losos, Chair