Founder Effect Speciation Lives! New Experimental Results Revive Mayr’s Theory

Today’s post is only tangentially related to anoles, but it’s about a new paper that seems to have received relatively little attention, so I thought it worth writing about. The idea of founder effect speciation goes back to the writings of Ernst Mayr and historically has been very important in the development of ideas about how new species originate. However, in recent years FE speciation has fallen on hard times. Theorists have claimed it to be highly unlikely, lab experiments have failed to find much support for it. More than a few evolutionary biologists have declared the idea dead and buried.

As an aside, why talk about FE speciation in these pages? The answer is simple—at least a few anoles (e.g., the green anole, A. carolinensis, and the festive anole, A. sagrei) have routinely colonized islands in the Caribbean, and very likely many of these colonizations involve the arrival of a single, impregnated, female. If the FE speciation occurs, these Caribbean anoles might be a good place to look for it. Moreover, a recent experimental study on A. sagrei (of which I was an author) reported that founder effects could have persistent effects on morphology, at least over the several-year span of the study.

And that leads us to the study in question, by Daniel Matute of the University of Chicago (and soon to be faculty at the University of North Carolina). In a truly gargantuan experiment on laboratory fruitflies just published in the Journal of Evolutionary Biology, Matute showed that reproductive isolation can, in fact, evolve as a result of extreme and persistent founder effects. The extent of this study is truly mind-boggling. A founder effect was induced by taking a single male and female fruit fly and putting them in a vial. Then, from their eggs, a single male and female were randomly chosen to form the second generation. This was continued for 30 generations. Sounds like a lot of work, right? Well, catch this: Matute started this experiment with not a single vial containing two flies, but with 1000 vials in which he replicated the experiment–I’ve never heard of such a massive experiment (though some Drosophila-savvy friends say I need to read the literature more). Now, admittedly many of the populations went extinct very quickly because of the intense inbreeding—80% were gone by generation 5 and only 12% lasted the full 30 generations. But, still that’s a lot of Drosophila TLC.

Degree of reproductive isolation (as measured from mate choice trials). The red histogram is the distribution of reproductive isolation between founder effect populations and the parental population; blue is between individuals from parental populations. 100 out of 123 surviving founder effect populations had reproductive isolation values greater than zero.

Degree of reproductive isolation (as measured from mate choice trials). The red histogram is the distribution of reproductive isolation between founder effect populations and the parental population; blue is between individuals from parental populations. Approximately 100 out of 123 surviving founder effect populations had reproductive isolation values greater than zero.

Of the 123 surviving lines, 100 of the lines showed some degree or reproductive isolation (i.e., flies preferred to mate with members of their own population rather than with members of the parental population), and in 3 of the lines, in which 80% of the matings were with their own kind, this degree of evolution of reproductive isolation was found to be statistically significant. Note, too, that even though the degree of reproductive isolation (RI) was only statistically significant in those three lines, the mean degree of reproductive isolation of all  FE lines from the parental (red line in figure to right) was greater than the degree of isolation in almost all parental x parental crosses. Or, looked at another way, a substantial number of FE lines evolved greater RI than seen in any of the parental crosses.

A number of perspectives can be taken on these findings. A conservative interpretation is that, at least very occasionally (0.3% out of 1000 initial founder events; 2.4% of 123 surviving populations), founder effects followed by very small population sizes for 30 generations can lead to the evolution of significant amounts of reproductive isolation. Given that the primary architects of FE speciation theory (Mayr, Templeton, others) have always said that FE speciation is a rare event, this result will be seen by many as supporting their position. Ardent proponents of founder effect speciation will go a step further and argue that the experiment provides at least suggestive evidence that founder effects can not infrequently lead to the evolution of enhanced reproductive isolation, given the relatively large number of populations with high degree of RI (see figure above). On the other hand, detractors will no doubt argue that the extremely stringent conditions imposed in the experiment, especially the maintenance of a population size of two for 30 generations, is both unrealistic of conditions likely to occur in nature and doesn’t closely model the theoretical ideas put forward by Mayr, Templeton, and others.

Although no doubt various camps will view these results in different ways, if nothing else, this is the first glimmer of support for FE speciation in a long time; it will be interesting to see whether the paper succeeds in putting founder effects back on the speciation playing field.

Blue-Eyed Agamids

In flipping through field guides to look at agamid dewlaps, I came across a couple of South-East Asian Agamid lizards with blue eyes. Given our recent discussions of blue-eyed anoles, I thought I’d point out these lizards as well. All information is from L. Lee Grismer’s Lizards of Peninsular Malaysia, Singapore, and their Adjacent Archipelagos
(and here’s an AA review of the book).

Aphaniotis fusca (Dusky/Peninsular Earless Agama): more fascinating than the blue eyes themselves is the variation among individuals in iris colour–A. fusca irises can be blue, orange, or brown. It appears that blue irises are restricted to adult males, but not all adult males have blue eyes. Further, the inside of these lizards’ mouths (the oral mucosa, to be precise) are also bright blue, similar to  Anolis onca.

Aphaniotis fusca (photo from www.fieldherpforum.com)

Aphaniotis fusca (photo from www.fieldherpforum.com)

Gonocephalus liogaster (Orange-Ringed Anglehead Lizard): hands-down winner for lizard with the most dramatic eyes, adult males of this species have a bright blue iris surrounded by a bright orange ring.

Gonocephalus liogaster (photo by Robert Yeoh)

Gonocephalus liogaster (photo by Robert Yeoh)

 

Anolis Photo Contest Voting Deadline This Sunday

Will this lovely photo by Diana Troya take home the gold?

Will this lovely photo by Diana Troya take home the gold?

With nearly 500 votes cast, it’s coming down to the wire. Get your vote in before the Sunday, 6 p.m. deadline!

Urban Anole Phylogenetic Study: You Can Help!

A. cristatellus on a rooftop in Mayagüez, PR

A. cristatellus on a rooftop in Mayagüez, PR

Urbanization poses a major challenge for many species, altering natural environments in ways that few animals can tolerate. Despite this, some species persist and even thrive in urban areas. In my research in the Revell lab at UMass Boston, I’ve been studying adaptation in response to urbanization in Anolis cristatellus, the Puerto Rican crested anole. However, among anoles urban tolerance is by no means restricted to A. cristatellus. We suspect that readers of this blog have probably observed many different anole species occupying and thriving in urban areas.  Consequently, we would like to ask for your help in gathering some information on this topic.

Anolis Allisoni Fight

This is a video from the YouTube channel of the thetravelholics that I stumbled upon a while ago showing two male A. allisoni fighting.

Notice:

1.The male that had the upper hand turned brown while the losing male remained blue throughout the fight.

2.When the male turned brown some of the skin on the back of his neck remained blue, this is possibly an example of selective color change.

3.Both males had prominent shoulder patches and black patches behind the eyes.

Help Identify a Mystery Lizard

mystery_lizardWe found the lizard depicted above in the herpetological collection at the University of Kansas. We have no information about where it is from or who collected it. Can anybody help us identify what species it is?

What Exactly Is A Dewlap?

Now that’s a dewlap! Photo by Niranjan Sant

All of us that observe and study Anolis (and Sitana) know that dewlaps make our favourite lizards special. Dewlaps can be key for species identification, and we pay a lot of attention to variation in their size and colour. But what exactly constitutes a dewlap?

I’m currently compiling a list of all lizard species with dewlaps, and have run into the problem of how to define a dewlap. Our trusted friend Wikipedia defines a dewlap thus:  a longitudinal flap of skin that hangs beneath the lower jaw or neck of many vertebrates. Helpful, perhaps, but only slightly, as the difference between a small dewlap and a puffed-out throat is not always obvious.

So we’re turning to you, dear reader, to help us figure out what a dewlap is. I think the easiest way to do so is by showing you a number of pictures of lizards and asking you to deliver a verdict on whether these lizards are in fact dewlapped (photos from various internet sources). Feel free to offer an alternative definition also, if you’d like.

1. Rock Agama

Kenyan Rock Agama

 

 

 

 

 

2. Calotes calotes from Sri Lanka

Green Forest Lizard (Calotes calotes)

 

 

 

 

 

 

 

 

3. Panther chameleon

panther-chameleon-01

 

 

 

 

 

 

 

4. Varanus griseus

varanus_griseus_8202

 

 

 

 

 

 

5. Iguana iguana

Stowaway – Can You Identify This Unfortunate Traveller?

Anolis sagrei (male) 266

An Anolis sagrei male from my study area in southwestern Taiwan.

Working with the brown anole (Anolis sagrei) is an eye-opener to the ability of some species to disperse far beyond the barriers that limit their natural dispersal potential. Anolis sagrei from Florida managed to reach Taiwan, most likely along with some nursery or agricultural products. They have also managed to reach Singapore, also I suspect along with some nursery products.

unknown snake

Can you identify this snake for us please?

Recently, an instance of a non-anole long-distance traveler came to my attention, and I would like to ask if anyone can help us identify the species involved. A friend told me about a snake that he obtained from someone, who got it from a person who imported wood from South America. Apparently, as the importer was about to start processing the wood he received, the snake slithered out of it. To me, the snake looks like a neotropical whipsnake (Masticophis mentovarius). However, it traveled by ship, meaning it could have gotten onboard at any port where the ship might have stopped. Thanks.

Re-Branding The Brown Anole

Doesn’t get much more festive than this guy!

I’m currently preparing a grant proposal featuring Anolis sagrei for a philanthropic foundation with broad goals that extend well beyond biology. The people reading this proposal will thus have diverse backgrounds, probably mostly non-biological and certainly non-herpetological. In reading a draft of the proposal, a friend remarked “This lizard needs a new name for this grant!” And, indeed, on reflection perhaps “brown anole” is not the most exciting name for the general public.

So, who wants to help re-brand A. sagrei? Of course, an obvious possibility is Sagra’s Anole, since the species was, after all, named after Mssr. Ramon de la Sagra*. But…that’s not really that exciting either. My friend (a biologist) suggested “the tramp anole,” using “tramp” in the ecological sense of a weedy species that is able to colonize and establish in a wide variety of habitats. A possibility, for sure, but maybe some of the other connotations aren’t so good. His second suggestion, though,  merit’s consideration. Turns out that “sagre” in Italian means “a festival.” And what lizard could be more of a partier than the ever-flashing, always rollicking, A. sagrei? So, I put it to you, anole world, what if we start referring to this species as “the festive anole”?

*It’s an interesting story how the species ended up A. sagrei, and not A. sagrai, but that’s a tale for another time.

Another Blue Anole!

There’s been discussion about blue anoles previously on Anole Annalsbut I’ve come across yet another on the internet. The anole in the above picture  is a species from the Sierra de Bahoruco in the Dominican republic. Apparently it has not been formally described yet and so does not yet have a name, but from the looks of it I would say it was related to A.alliniger or singularis. This actually leaves Puerto Rico as the only island without a blue trunk-crown anole. I have found one other picture of this species on Flickr (though it does not allow for posting on other web pages).

Below is another photo from the Anolis Contact Group, of yet another new species that appears to be related to singularis.

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