Category: Classics from the Literature Page 2 of 4

On Sexual Selection in Anolis

"Fig. 33. Sitana minor. Male, with the gular pouch expanded (from Gunther's 'Reptiles of India')."

In The Descent of Man and Selection in Relation to Sex (1871), Darwin laid out his ideas about sexual selection. Darwin believed that sexual selection was distinct from natural selection; while natural selection operates through survival and fecundity, sexual selection operates through differences in mating success among individuals.

The puzzle that Darwin set out to solve was the existence of traits like the peacock’s tale or the elk’s antlers. These traits differ between sexes but aren’t related directly to mating. And, these traits appear to be quite costly. Darwin reasoned that these costly traits must benefit the bearer in some way related to mating success. Most likely, these benefits accrued during battle with other members of same sex for mates or during the courtship of the opposite sex. In both cases, having the costly trait benefitted an individual’s mating success beyond the incurred survival or fecundity cost.

Now, how does this discussion relate to anoles?

Species Richness Patterns in Caribbean Anolis III

Lineage accumulation curves (Fig. 2, Rabosky and Glor 2010) showing that Hispaniola (blue), Jamaica (purple), and Puerto Rico (orange) have reached speciation-extinction equilibrium. Cuba (red) is still gaining species.


Losos, J.B., and D. Schluter. 2000. Analysis of an evolutionary species-area relationship. Nature 408: 847-850.

Rabosky, D.L, and R.E. Glor. 2010. Equilibrium speciation dynamics in a model adaptive radiation of island lizards. Proc. Nat. Acad. Sci. 107: 22178-22183.

Losos and Schluter (2000) return to Caribbean anoles to test three hypotheses about the species-area relationship: (1) that there is an area threshold above which speciation surpasses immigration as a source of new species; (2) above the threshold size, speciation events per unit time should increase with island area; and (3) the slope of the species-area relationship should become steeper above the area threshold. Qualitatively similar to Losos (1996), this paper was novel in that a newly available, nearly complete, mt-DNA phylogeny allowed Losos and Schluter to reconstruct immigration and speciation events and to model whether species number has reached speciation-extinction equilibrium.

Species Richness Patterns in Caribbean Anolis II

Losos, J.B. 1996. Ecological and evolutionary determinants of the species-area relation in Caribbean anoline lizards. Philosophical Transactions of the Royal Society of London B 351: 847-854.

As alluded to previously,  MacArthur and Wilson (1967) did consider evolutionary processes when they developed their Theory of Island Biogeography. Specifically, in situ evolutionary diversification (i.e. speciation) may contribute substantially to the species diversity of an island and should be considered in any general attempt to model species-area relationships on islands. Building on Rand’s 1969 paper studying the ecological determinants of species richness in Caribbean anoles, Losos (1996) incorporates an evolutionary perspective into the Caribbean Anolis species-area story.

Species Richness Patterns in Caribbean Anolis I

This is the first of a series of posts that will review a number of papers that examined species richness patterns in anoles, starting with Rand’s 1969 paper described below and moving towards the present day understanding. Read on!

Species richness predictions from the Theory of Island Biogeography

 Rand, A.S. 1969. Competitive exclusion among anoles (Sauria: Iguanidae) on small islands in the West Indies. Breviora 319: 1-16.

The empirically observed species-area relationship (SAR) is one of the closest things to a law that we have in evolutionary ecology. All else equal, the larger the area, the more species will be in it. In a paper in 1963, and a book in 1967, MacArthur and Wilson (M&W) put forward the Theory of Island Biogeography (TIB) to explain the species-area relationship on islands. They observed that the number of species that inhabit an island scales positively with island area and negatively with distance of an island from the mainland species source. M&W argued that the SAR is governed by two ecological processes: colonization and extinction. Colonization probability increases with the size of an island and decreases with island distance from the mainland. Extinction probability increases as island size decreases because small islands support smaller population sizes and leave species are more vulnerable to fluctuations in abiotic and biotic environmental factors. Thus, the TIB predicts that large islands close to the mainland will be species rich, while small islands far from the mainland will be species poor. The TIB inspired thousands of papers: according to Google Scholar, the 1967 book has been cited 11,148 times and the 1963 paper has been cited 1,416 times.

Anole Classics: Albert Schwartz (1968) on Geographic Variation in Anolis distichus

I previously characterized Albert Schwartz as one of the five kings of Greater Antillean anole taxonomy for having described eight new species from the region.  Although Schwartz described the fewest species among the five kings, focusing on new species masks Schwartz’s even more important contributions to cataloguing geographic variation within species.  Schwartz’s career-spanning interest in biogeography and geographic variation resulted in a prolific history of describing subspecies in anoles and other taxa.  Anyone who’s looked at Schwartz and Henderson’s classic book on West Indian reptiles and amphibians is familiar with the irregular blobs that designate subspecies boundaries on the range maps for many of the region’s most geographically widespread species.  Many of these blobs were the result of Schwartz’s own efforts.  The pinnacle of Schwartz’s work on geographic variation may be his 1968 monograph on geographic variation in Anolis distichus.

Kings of Greater Antillean Anole Taxonomy V: Orlando Garrido

The last of the five kings described more new anoles than any of the others: Orlando Garrido.  Garrido is unique among the five in two ways.  First, he’s still alive, still active, and still making contributions to our understanding of anole diversity.  Second, he’s actually a citizen of a Greater Antillean country: Cuba.

Garrido is often recognized as Cuba’s greatest naturalist.  In addition to his impressive body of work with reptiles, he has made many other important contributions to our understanding of Cuban nature, including the spectacular “Field Guide to the Birds of Cuba.”  His successes  are a testament to how far science has come since Barbour’s time, when practicing science in the West Indies required a wealthy North American pedigree.  I’ve credited Garrido with a whopping 24 species, all from his native Cuba.

The Kings of West Indian Anole Taxonomy IV: Albert Schwartz

I’ve credited the fourth king of Greater Antillean anole taxonomy – Albert Schwartz – with describing eight Greater Antillean anole species.  The period during which Schwartz’s career overlapped with Williams’s and that of the fifth yet-t0-be-revealed king were the glory years of Greater Antillean anole taxonomy.  Over a little more than a decade in the late 1960s through the 1970s, these three figures described over 10 species, including some of the last new species discovered on Hispaniola and Jamaica.  The activities of these three key figures were highly synergistic; Schwartz and Williams often contributed to one another’s work and divvied up projects to mutual benefit (even though they never described an anole species together) and Schwartz was a junior coauthor with the fifth king on several species descriptions.

After graduating with a PhD from the University of Michigan, Schwartz spent the majority of his academic career at Miami Dade Community College, an institution known more for its massive enrollment than for its faculty’s contributions to systematics.  Early in his career, Schwartz worked primarily in Cuba, resulting in the description of three species, including two locally restricted species related to the Cuban crown-giant anole Anolis equestris (baracoae and smallwoodi) and a widespread trunk-ground species (jubar) that is the xeric forest counterpart to another widespread Cuban trunk-ground anole found primarily in mesic environments (homolechis).  Schwartz would later devote his attention to Hispaniola, ultimately describing five species from both Haiti and the Dominican Republic.  As was the case with Williams, many of the Hispaniola taxa that Schwartz described were unusual montane endemics (rimarum, fowleri, sheplani, and eugenegrahami).

The Kings of Greater Antillean Anole Taxonomy III: Ernest Williams

Following Barbour’s work, nearly a quarter century would pass before another Harvard man picked up the mantle of describing new anoles.  Among his many other contributions, Ernest Williams named 12 species of anoles from Jamaica, Cuba, and Hispaniola between 1959 and 1975.  By the time Williams came along, most of the abundant and widely-distributed anoles had already been described.  Many of the species Williams described are montane endemics (A. reconditusA. christopheiA. etheridgeiA. dolicocephalusA. occultusA. singularisA. insolitus) that might have been more difficult to access during previous generations of herpetological exploration in the West Indies.  The last Greater Antillean species he described – Anolis marcanoi – was among the first “cryptic” anole species to be recognized with the aid of molecular markers.  Even after his work describing new Greater Antillean anoles came to an end, Williams continued to describe new species of anoles from the mainland through the 1980s.  In his last publication in 1999 (published after his death in 1998), Williams called an end to the era of discovery in anoles.  For more on Williams, you can read the memorial published in the Harvard Gazette in 2009 by A. W. Crompton, Karel Liem and Jonathan Losos.

The Kings of Greater Antillean Anole Taxonomy II: Thomas Barbour

I previously introduced my mission to recognize the five anole systematists responsible for describing the majority of the anole species found in the Greater Antilles.  The first king of Greater Antillean anole taxonomy was the prolific E. D. Cope.  Cope was the last in a line of authors who described anole species that he’d never actually spent time with in the field (see also Duméril and Bibron).  The next king on my list, by contrast, was an avid field biologist and conducted field work in the West Indies throughout his career.

One Night in Antigua – Photos from a Layover with the Colossus Anolis leachii

Early morning sighting of a female Anolis leachii on a leaf covered in raindrops.

Sometimes, they say, it’s about the journey, not the destination. This makes me think of exciting layovers I’ve had in Anolis country. At any place where the layover is long enough to permit stepping outside of the airport, I like to pop out and see what kind of anoles I can find lurking around the terminals. On a few occasions, travel requires an overnight stay in an exciting place. One of the benefits of working in remote Lesser Antillean islands is that infrequent (and unpredictable) airline schedules typically mean spending a night or two in tiny islands to and from the actual destination. For years I’ve been working in Montserrat, a small island with an active volcano, where I try to follow the flighty bugger Anolis lividus as close to the volcano as I can get. On our way home from Montserrat, we usually spend a night in Antigua, where the charming giant, Anolis leachii can be found.

View of a raincloud and rainbow as we descended into Antigua in January 2009.

Like most anoles from the Lesser Antilles, this species is abundant. However, few species from the Lesser Antilles are as large as A. leachii. It is a member of the bimaculatus clade of large anoles from the Northern Lesser Antilles. According to Scwhartz and Henderson (1991), it can reach a snout-vent length of 123 mm. For anyone who works with crown-giants, this may not seem so large, but for an aficionado of the Lesser Antillean anoles, Anolis leachii is a relative behemoth! Its body ranges from yellow to blue and green, and it’s covered in a purplish vermiculation. The large eyering ranges from a sulfurous yellow to a deep orange. Hands down, this is one of the world’s most beautiful anoles. Imagine my excitement when I got to spend a night chasing these lizards through the wilds of Antigua Village, a cushy beach resort teeming with Anolis leachii and the smaller congener, A. wattsii. Here I offer some photos of this dazzling species, and a bit of its taxonomic history.

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