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Remembering Lourdes Rodriguez Schettino

Lourdes Rodriguez Schettino with Ada Chamizo and Rich Glor in 1999.

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.

Lourdes Rodriguez Schettino in 1995.

 

 

Richard Etheridge, Founder of Anole Systematics, Passes Away

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.

Growth Rate in a Population of Anolis nebulosus from Tropical Dry Forest

Anolis nebulosus is an endemic species of Mexico. Its distribution occurs along the coasts of the Mexican Pacific, from Sinaloa to the Isthmus of Tehuantepec, but it enters the highlands a little bit in the Sierra Madre Occidental. At the end of the 80’s, very little was known about the natural history of populations of this species.

Our study was conducted at the Field Station Chamela, Jalisco México during a period from 1988-1990. It is believed that the growth rates of juveniles are restricted by available food resources. In animals that grow throughout the year, such as tropical lizards, it is believed that growth will be faster during the rainy season. Consequently, we wanted to know (a) if there are differences in growth rates between sexes of A. nebulosus; (b) whether growth rates vary among age classes, between dry and wet seasons, and between years; and (c) what extrinsic factors (e.g., environmental temperature, precipitation, and food availability) promote variation on growth rate of different age classes and sexes between seasons and years.

Our study shows the growth rate of males and females and feeding behavior; male grow faster than females; males reached sexual maturity at minimum SVL of 32 mm, while females at 35 mm of SVL; lizards live only one reproductive season, that is during rainy season–after that, they die. In 1989, juvenile males and females grew more in both seasons (rainy and dry) than adults; for 1990, there were no differences by season or between age classes.

The information that was presented about the growth of Anolis nebulosus in Ecology and Evolution  is part of the information I (ARB) have shared with my ex-graduate students, now doctors (Uriel Hernández-Salinas, Raciel Cruz-Elizalde, and Christian Berriozabal- Islands), as well as my colleague, Dr. Shai Meiri. This study opens a window on knowledge of the natural history of this fascinating species model.

 

Evolution of Claws in Greater Antillean Anoles

Much has been made about the role of convergent morphology in the partitioning of vertical habitat by Greater Antillean anole ecomorphs. In particular, the role of toepads in allowing anoles to utilize vertical habitat space has been the focus of abundant research. It is easy to see why, given the novelty of the toepad in anoles (and, of course, geckos). But when you look across the lizard tree, there plenty of arboreal lizards lack toepads entirely and instead rely on their claws for climbing.

Functional studies in other claw-bearing squamates provide some predictions for how claws may be related to habitat use. For example, studies have shown that highly curved claws perform better in performance trials of clinging strength (Zani 2000), and so we would expect that more arboreal species should have more highly curved claws. Yet claws remain a relatively neglected trait in anoles, despite the wealth of literature related to other aspects of their ecomorphology.

Is claw morphology related to habitat partitioning in anoles? Or is it really all about the toepad? In a recent publication, my coauthors and I set out to examine how claw morphology varies across the Greater Antillean anole radiation with these questions in mind. We measured the claws and toepads of 428 museum specimens across 57 species to test the prediction that claw morphology was related to habitat use in Greater Antillean anoles. Our sampling spanned every ecomorph present on each Greater Antillean island, as well as several non-ecomorph species.

First, we tested the prediction that arboreal species should have more curved claws than terrestrial species. This pattern has been observed in squamate reptiles broadly, but has not been tested within anoles. Unfortunately, only one truly terrestrial anole exists in the Greater Antilles, A. barbouri. Therefore, to increase our sampling, we included four terrestrial mainland species. As predicted, we found that terrestrial anoles had much flatter claws than arboreal species. Claw curvature was also positive correlated with perch height


Fig 1 PCA plot of first and second principal components for univariate claw measurements. Convex hulls are depicted for each Greater Antillean ecomorph and terrestrial species (both Greater Antillean and mainland) are labelled. Insets: (A) Superimposed means of geometric morphometric shape for terrestrial and arboreal species; (B) Boxplots of claw curvature measurements for arboreal and terrestrial species (*** P < 0.001).

Next, we tested the prediction that ecomorphs are differentiated by claw morphology. We found that ecomorph did predict claw morphology, but that differences were more pronounced for some ecomorphs. Notably, more terrestrial ecomorphs (trunk-ground) were differentiated from more arboreal ecomorphs (trunk and trunk-crown) in line with predictions. In the most extreme case, twig anoles appear to have dramatically reduced claws. Their claws were not only flatter, but also shorter and thinner relative to body size compared to all other ecomorphs. We can only speculate as to the reason that this might be the case. Perhaps claws are unimportant or an impediment in twig anole locomotion? Clearly, functional studies are required to fully understand the relationship between being a twig anole and claw morphology.

Finally, we tested for patterns of covariation, or phenotypic integration, between claw and toepad traits. Claw curvature was positively correlated with toepad width and lamella number, both traits that positively correlate with perch height and demonstrated to improve clinging performance in anoles. Other claw traits such as relative length and diameter did not correlate with toepad traits. This is perhaps unsurprising, as these traits also do not appear to be differentiated between more arboreal and terrestrial species. Given the developmental literature showing that terminal digit features are decoupled from the rest of the digit (Casanova and Sanz-Ezquerro 2007), our observed pattern of integration is unlikely to be the result of developmental linkage.

In conclusion, our data suggest that integration between claw curvature and toepad traits is driven by correlative selection due to structural habitat use. Our results strongly support the hypothesis that claws play an important role in the partitioning of vertical habitat within the adaptive radiations of anoles, and suggest that they are likely shaped by similar patterns of correlative selection that have driven the evolution of ecomorphs overall. Still, we have a lot to learn about how variation in claw shape influences actual locomotor performance in anoles. Claws are a trait just waiting for more research!

Casanova, J. C., and J. J. Sanz-Ezquerro. 2007. Digit morphogenesis: is the tip different? Dev. Growth Differ. 49:479–491.
Yuan, M. L., M. H. Wake, and I. J. Wang. 2019. Phenotypic integration between claw and toepad traits promotes microhabitat specialization in the Anolis adaptive radiation. Evolution 73:231–244.
Zani, P. A. 2000. The comparative evolution of lizard claw and toe morphology and clinging performance. Journal of Evolutionary Biology 13:316–325.

Where Do Anolis Lizards Lay Their Eggs?

Searching for anole eggs

In this southern city where the heat is on, and the parties last ‘til the break of dawn, residents are busy patrolling territories, showing off their physique, and courting each other. With all this activity going on, South Miami, FL surely takes the crown for the best people anole-watching destination. All these anoles surely have different stories for how they got to Miami, and some research has shed light on this issue. But what is unknown about anoles in this city is how they manage to proliferate when the temperature can be unforgiving to developing embryos (Sanger et al. 2018; Hall and Warner 2018).

Anolis lizards have attracted the attention of many anolologists that are mesmerized by their conspicuously beautiful dewlap displays, superb clinging abilities (Donihue et al. 2018; Winchell et al. 2018), and an urge to show off their physiology (Battles and Kolbe 2018). However, one aspect of their biology that is much less conspicuous is their nesting behavior. Even biologists that have devoted countless hours studying their behaviors know very little about where females lay eggs, what microhabitats they prefer, and how those nesting behaviors impact embryonic development. In this recently published study, we unveil the mystery of where these city slickers’ nests are located compared to their forest-dwelling counterparts.

A Web-Based Identification Key for Anolis

Can you identify this anole? If not, don’t despair. The Anolekey is here to help.

In 1995 Ernest Williams and collaborators published a landmark underappreciated paper describing their “computer approach” to identification of Anolis lizards (Williams et al. Breviora 502:1-47). These authors employed the data management and programming tool HyperCard to produce an application that matched a set of character scores from an unknown specimen to scores for the same characters in Williams’ vast database of Anolis. This “Anolis Handlist” was a stark and wonderful departure from the dichotomous keys standardly used by taxonomists and field biologists.

With this post I announce the release of a preliminary version of a new computer key to Anolis. This key operates similarly to the Handlist, but uses the Lucid platform. Dear readers of Anole Annals, I am writing to request your help to Beta-test this first version of the key. A few notes regarding use of the key:

–The key is accessible through http://www.stevenpoe.net/anole-key-and-collecting-guide.html. Use password “EEW” for access.

–Use of the key should be fairly intuitive (Figure 1), but will be most efficient if the manuscript describing the key (posted on the website) is accessed prior to use. The key is intended for use both in the field—where dewlap photos and obvious traits such as body size and locality are likely to be informative—and in identifying museum specimens. For this latter function of identifying preserved individuals, entering information for many characters may be critical for accuracy.

Figure 1. If you tell the key you have found an anole at 1650 meters elevation in the Eastern Andes of Ecuador with a body length of 90 millimeters, the key identifies Anolis fitchi and A. podocarpus as the most likely species. The “Differences” function suggests these two species may be distinguished by dewlap scalation.

–Trait data for every species of Anolis (n = 398; see justification for species sample at stevenpoe.net; Figure 2) were taken from 1000s of preserved specimens and from observation of 273 species of Anolis in life during fieldwork in 19 countries. Thirty-three characters are employed to identify species. Example conditions for each character are figured with photographs.

Figure 2. The key includes trait data for every species of Anolis.

–Dewlap photos for 248 species are included. A long-term goal is to include (preferably male) dewlap photos for every species. Please consider contributing dewlap photos that would enhance the key (see below).

–Some other anole-related items on the website are pertinent to the key, or perhaps of additional interest. These items include an Anolis species list, Type descriptions for all recognized species of Anolis, justification for recognized species, photo guides to a few local anole faunas, and a manuscript describing how to catch anoles.

–Male dewlap photos needed: agassizi, alayoni, alfaroi, allisoni, altavelensis, altitudinalis, alumina, anatoloros, anchicayae, anfiloquiae, annectens, antioquiae, argillaceus, armouri, baracoae, barbouri, bellipeniculus, bimaculatus, birama, biscutiger, blanquillanus, bonairensis, brasiliensis, bremeri, breslini, brunneus, caceresae, capito, caquetae, carlostoddi, centralis, charlesmyersi, chlorodius, chrysolepis, chrysops, clivicola, cobanensis, confusus, cristifer, cryptolimifrons, cupeyalensis, cuprinus, cyanopleurus, cyanostictus, cymbops, darlingtoni, desechensis, dissimilis, distichus [d. distichus: New Providence Island, Bahamas], divius, dracula, eladioi, ernestwilliamsi, euskalerriari, favillarum, forresti, fugitivus, fungosus, gingivinus, gonavensis, griseus, guafe, guazuma, haetianus, haguei, heteropholidotus, huilae, ignigularis, imias, immaculogularis, inderenae, inexpectatus, insolitus, juangundlanchi, kahouannensis, kreutzi, laevis, lamari, landestoyi, lividus, longiceps, loveridgei, luciae, lucius, macilentus, macrophallus, maia, menta, meridionalis, microtus, mirus, monensis, monteverde, morazani, muralla, nasofrontalis, neblininus, nelsoni, nicefori, oligaspis, onca, oporinus, paravertebralis, phyllorhinus, pigmaequestris, pijolense, planiceps, pogus, porcus, prasinorius, properus, propinquus, pseudopachypus, pseudotigrinus, purpurgularis, pygmaeus, ravitergum, rejectus, rivalis, roatanensis, roosevelti, rubribarbaris, rubribarbus, ruibali, ruizii, sabanus, santamartae, schwartzi, shrevei,singularis, solitarius, spectrum, squamulatus, strahmi, tandai, tenorioensis, terueli, tetarii, tigrinus, toldo, townsendi, triumphalis, umbrivagus, vanidicus, vaupesianus, vescus, vicarius, villai, wattsi, wellbornae, wermuthi, whitemani.

–To close, a caveat and a plea for help: This is a first version. In spite of my best efforts over 10+ years of work on this key, there undoubtedly are many errors in scoring for many species. I certainly have missed some reported invasions, failed to measure an especially large specimen of some species, and neglected to record a comprehensive range of trait values for some species, among other failings. My fervent hope is that colleagues will join me in vetting this key via its use in the field and in museum studies. The current version of the key will not ensure accurate identification of all specimens of anoles. But hopefully this key is a starting point that may achieve some utility as a preliminary guide to the species of this group. Future versions, with the help of colleagues, may eventually serve as an authoritative guide to anole identification.

–For questions and comments regarding the key and submission of dewlap photos: kunayalae@yahoo.com.

 

 

 

New Research Project on Anolis pogus on St. Maarten

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.

  This new research, executed by Aeres University student Joost Merjenburgh, will provide insight into the status of the vulnerable species and hopefully prevent its extinction.The bearded anole is an important endemic species to St. Maarten, which means that the species is native and can only be found here. In the past, the bearded anole occurred on Anguilla and possibly on Saint-Barthélemy, but has vanished on both islands for unknown reasons.

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.

Anolis Newsletter VII is Now Published!

Following on from the last Anolis Symposium, held at the Fairchild Tropical Botanic Gardens, Miami FL, from 17-18 March 2018, we solicited submissions for the next issue of the Anolis Newsletter. We now finally have them all together! The newsletter represents an informal forum for the presentation of data and discussion of theory relevant to anoles. It serves three functions: to allow investigators to inform others of their current and future research; to provide an outlet for speculation and theoretical musings perhaps inappropriate for publication in more formal venues; and to give an opportunity to present data and ideas that otherwise might never be distributed. As with previous newsletters, there is a general request that nothing said herein be quoted without the authors’ express permission.

Spanning 333 pages, including 47 separate articles, I can now bring you the Anolis Newsletter VII!

Click this link here to access a PDF copy.

As always, you can also find links to PDF’s of all editions of the Anolis Newsletters (1972-2019) at the dedicated page on the Losos Lab website:
https://sites.wustl.edu/losos/home/anolis-newsletters-2/

Here’s a rundown of the articles included in Edition VII:

Yasel U. Alfonso and Dennis D. Ávila
Ectoparasite-host relationships: a case study of Anolis lizard ecomorphs on southeastern  Cuba

Yasel U. Alfonso, Javier Torre, and Daniel J. Paluh
The bizarre false-chameleons (clade Chamaeleolis, Anolis) from Cuba

Andrew C. Battles
How do anoles respond to urbanization? A summary of Ph.D. research on ecology and thermal biology in Anolis in Miami, FL

Winter A. Beckles
Variation in habitat lighting may mediate the persistence of dewlap color polymorphism
in South Florida bark anoles

Tom W. Brown
Conservation concerns for loyal lizards: Habitat specificity, site fidelity, a localised territory and in-situ growth rates for Anolis bicaorum (Squamata; Dactyloidae), endemic to Isla de Utila, Honduras

Stephanie L. Clements, Shantel Catania, and Christopher A. Searcy
Non-native species dominate herpetofaunal community patterns in both native and non-native habitat patches in urban Miami-Dade County

Rachel E. Cohen
Anole behavioral neuroendocrinology.

Jens De Meyer, Colin M. Donihue, Daniel Scantlebury, Julienne Ng, Rich E. Glor, Jonathan B. Losos, and Anthony J. Geneva
Protocol for setting up and rearing a successful lizard room

Colin M. Donihue and Anthony Herrel
A report on Anolis nubilus from the now rat-free island of Redonda

J. Sean Doody, Sean Sullivan, and Deby Cassill
Plasticity in hatching of anoles

Claire Dufour
Ongoing and future research on Anolis

A. C. Echternacht
Anolis research in the Echternacht Lab

Sean T. Giery and James T. Stroud
Geographic variation in trophic ecology of the Brown anole (Anolis sagrei): species-rich
communities are composed of more diverse populations

Joshua M. Hall, Timothy S. Mitchell, and Daniel A. Warner
The brown anole (Anolis sagrei) as a model for studying life-history adaptation to seasonality

Emma A. Higgins, Adam C. Algar, and Doreen S. Boyd
Revealing controls on abundance and microhabitat use of Anolis lizards in a changing
island landscape using airborne remote sensing

Michele A. Johnson
The potential for large-scale behavioral studies: A call to Anolis field biologists

Janson Jones
Beneath the Spanish moss: Growing up with Anolis in Florida. A photographic naturalist’s perspective

Masakado Kawata
Collaborative research projects on Anolis lizards in Cuba

Oriol Lapiedra
The ecological and evolutionary consequences of behavior in a changing Planet

Oliver Ljustina and James T. Stroud
Little evidence for size-structured habitat use in a diverse Anolis community

Michael L. Logan
Using transplant experiments to understand adaptation and speciation in anoles

Jonathan B. Losos
Anolis research in the Losos Lab

Inbar Maayan
A case study of character displacement and phylogeography of Jamaican anoles

D. Catalina Mantilla, Samuel D. Morales, Rafael Parra-Medina, and James T. Stroud
Histopathology of large epidermal cysts on the invasive Puerto Rican Crested Anole
(Anolis cristatellus) in Miami, Florida, USA

Gregory C. Mayer and Tony Gamble
Using archival DNA to elucidate anole phylogeny

Gregory C. Mayer and Jonathan B. Losos
Anoles not found

Walter E. Meshaka, Jr
The winds of stability: A south Florida residential Anolis assemblage over time

Walter E. Meshaka, Jr
What are the ecological costs and benefits to northern geographic expansion by a
successful anole?

Tim Mitchell
A call for more long-term studies of plasticity in anoles

Caitlin C. Mothes, James T. Stroud, Stephanie L. Clements, and Christopher A. Searcy
Predicting the invasion dynamics of anoles (and other lizards) using ecological niche
modeling

Julienne Ng, Daniel J. MacGuigan, Audrey L. Kelly, and Richard E. Glor
Do male-male interactions drive changes in dewlap size?

Stuart V. Nielsen, Aaron M. Bauer, and Tony Gamble
Natural History Note: Anolis sagrei foraging on a patch of obvious prey

Joanna Palade, Kenro Kusumi, Alan Rawls, and Jeanne Wilson-Rawls
Satellite cells demonstrate expanded musculoskeletal potential

Ivan Prates
Using mainland anole genomes to understand habitat shifts through time

Ashley M. Rasys, James D. Lauderdale, and Douglas B. Menke
Establishment of genome editing methods in Anolis sagrei

Liam J. Revell
Can we detect differences in the rate of discrete character evolution between clades of
anoles?

M. Christine Rose-Smyth
Role of a sweet-toothed anole (Anolis conspersus) in orchid pollination

James T. Stroud
Using introduced anoles as natural experiments in ecology and evolution

James T. Stroud
Thoughts on the ecology and evolution of anoles; insights from 5 years of meandering
strolls

Lindsey Swierk
Ongoing research on the ecology and behavior of Anolis aquaticus

Sarin (Putter) Tiatragul and Daniel A. Warner
Beating the Heat: nest characteristics of anoles across suburban and forest habitats in
South Miami

Javier Torres
Sterility in odd-looking Anolis mestrei (Dactyloidae) living in sympatry with trunkground
anoles

Nathan W. Turnbough
Are anole appetites altering ambient ant assemblages?

Kamau Walker, Andrew C. Battles, and James T. Stroud
Inter-specific predation bteween two eco-morphologically similar Anolis lizards

Kristin M. Winchell
Urban habitats: A natural experiment perfect for anoles

Amber N. Wright
Some thoughts on the use of experimental enclosures for studying anoles

Cindy Xu, Elizabeth D. Hutchins, Minami A. Tokuyama, Jason M. Newbern, and Kenro
Kusumi
Identifying molecular and cellular mechanisms of tail regeneration in anoles

Testis Size Evolves Faster than Sperm Size across Anoles

Sperm of Anolis sagrei. Picture by Ariel Kahrl.

Anyone who has worked with anoles in the field can tell you that during the breeding season, you can see these lizards mating pretty frequently. In fact, brown anoles are so promiscuous they have one of the highest rates of multiple paternity in vertebrates! In addition to this, females can store sperm for months at a time, which suggests that this group probably experiences a lot of sperm competition. We know from other groups that sperm competition can lead to the evolution of lots of interesting and diverse traits, but we know relatively little about how sperm competition targets these traits in anoles (though see Klaczko et al’s 2015 paper about the rate of genital evolution in anoles). 

Some of our previous work has shown that sperm size and sperm count are correlated with reproductive success in the lab, but it’s unclear how these important components of male reproductive success evolve. To explore this, we collected data on testis size (as a proxy for sperm production/sperm count), and sperm morphology (measuring the sperm head, midpiece and tail) for 26 species of anoles. We hypothesized that if testis size had a higher rate of evolution than sperm morphology, sperm competition likely targets sperm count over sperm size.

Rates of evolution of testis size, body size and sperm morphology next to an anole sperm.

In fact, we found that testis size evolves much faster than sperm morphology, confirming our hypothesis. We also found that within the sperm, the midpiece (which contains the mitochondria) evolves 2-3 times faster than the rest of the sperm cell. This is maybe unsurprising, considering that the midpiece size is correlated with reproductive success in brown anoles, and is often associated with sperm longevity and sperm velocity in other species.

 

You can read the pre-print from JEB below!

Kahrl, AF, Johnson, MA, and Cox, RM. 2019.Rapid evolution of testis size relative to sperm morphology suggests that postcopulatory selection targets sperm number in Anolis lizards. Journal of Evolutionary Biology.

https://onlinelibrary.wiley.com/doi/abs/10.1111/jeb.13414

New Book on Lizard Behavior Appears Next Month

Name that cover-gracing anole!

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!

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