Anolis cristatellus on a smooth, vertical substrate in Puerto Rico (photo by K. Winchell)
In urban areas, the number of natural substrates (e.g. trees) is reduced. In their place are novel manmade substrates (e.g. walls, metal gates). These surfaces undoubtedly have different properties relevant to anole locomotion: they are smoother, harder, and (in the case of walls) much broader and flatter compared to natural surfaces in a forest. In urban areas lizards still use these substrates at high frequency, but do they do so effectively? Kolbe and co-authors began to dive into this complex topic in their recent publication, “City slickers: poor performance does not deter Anolis lizards from using artificial substrates in human-modified habitats” (Kolbe et al. 2015).
The relationship between habitat use, morphology, and performance for anoles has been extensively studied in natural environments (reviewed in Losos 2009). Urban environments add new dimensions to this area of research. Resource distribution and abundances differ drastically compared to natural areas. For example, the distribution of available perches and what they are made of in urban habitats is very different from a forest. Moreover, the properties of these resources differ drastically as well: urban substrates are smoother, broader, and have different thermal properties, to start. Understanding these differences in habitat use and how they influence performance and, ultimately, adaptive responses in anoles is the topic of ongoing collaborative research that I (K. Winchell) and the Kolbe lab have been conducting.
An actual perch used by Anolis cristatellus – neither smooth nor flat! (photo by K. Winchell)
In the real world, lizards cling to everything from smooth, flat concrete walls to rough, rounded tree trunks. So why is it that most studies on cling force in anoles focus on clinging to smooth flat substrates? Does cling force differ if the substrate is rounded or rough? Jason Kolbe sought to answer this question in his recent publication, “Effects of Hind-Limb Length and Perch Diameter on Clinging Performance in Anolis Lizards from the British Virgin Islands” (Kolbe 2015).
We know that morphology impacts performance in anoles and that performance varies with environment. For example, sprint speed is correlated with limb length, but this relationship depends on the diameter of the substrate (e.g. Losos and Sinervo, 1989). We also know a little about clinging performance in anoles. Greater cling force is correlated with larger toepads and more lamellae on smooth flat surfaces (Irschick et al., 1996; Zani 2000; Elstrott and Irschick, 2004), but adhesion on rougher surfaces may be influenced by claw and toe morphology (Zani 2000).
There appears to be an unexplored interaction between substrate properties and clinging ability that involves more than just toepad characteristics. Specifically, Kolbe points out that claws can increase clinging ability by digging into the perch or simply by increasing friction on the surface. Limbs can also increase friction via the application of compression forces to the substrate. In other words, cling force can be increased, particularly on rough surfaces, by using muscular force to grasp rather than relying on van der Waals forces from the toepads, which are more effective on smooth flat surfaces.
Anolis cristatellus (photo by KMW)
Anolis stratus (photo by KMW)
Anole species used in this study: Anolis cristatellus (left) and Anolis stratulus (right). Photos by K. Winchell.
Kolbe investigated this further by looking at the interaction between limb length and clinging ability on perches of different diameters with Anolis cristatellus and Anolis stratulus from the British Virgin Islands. Specifically, he hypothesized that cling force should increase as the ability of a lizard to obtain a firm grasp on a substrate improves (i.e. when it can wrap its limbs around the substrate). This ability to form a secure grasp is dependent on both the diameter of the perch and on lizard limb length.
In collaboration with the Conservation Biology course taught by Dr. Karen Beard here at Utah State University, where I am a Ph.D. student, I have been involved in gathering life history data on ~400 species of reptiles that have been introduced outside of their native ranges for an analysis of how life history traits (e.g., diet, fecundity, longevity) interact with other factors to influence the likelihood of successful establishment. Appendix A of Fred Kraus’ 2009 book Alien Reptiles and Amphibiansis the source of the species list we are using, and included in this analysis are 26 species of Anolis. This is where you come in.
First, we coded all anoles as (i) sexually-dichromatic, (ii) diurnal, (iii) non-venomous, (iv) oviparous, (v) omnivores that lack (vi) temperature-dependent sex determination and (vii) parthenogenesis. Is anyone aware of any exceptions to these seven generalizations?
Second, we searched for data on clutch size, clutch frequency, incubation time, and longevity. The Anole Classics section of this site and the Biodiversity Heritage Library were particularly useful. After conducting what I feel to be a pretty thorough literature scavenger hunt, I am forced to conclude that some of these data simply do not exist at the species level for all of the species we’re interested in, or are not explicitly stated in a way that is obvious to a non-anole-expert. Of course, there is a lot of literature, including many books that I don’t have access to, and there are also lots of credible observations that don’t get published. I’m hoping that some of the readership here can help fill in at least some of the blanks in the table below. As one member of the team, I did not collect all of the data that are filled in myself, nor have I personally vetted every value, so if you spot an error please do point it out.
Two important points:
Many environmental factors obviously influence the life history parameters of our beloved and wonderfully plastic reptiles, so we appreciate that many of these values would be better represented by ranges and are dependent on latitude, altitude, climate, and many other factors. Where a range is published, we are using its median value.
I should also emphasize that, because of the large size of this study and the diversity of taxa included (ranging in size from giants like Burmese Pythons, Nile Crocodiles, and Aldabra Tortoises to, well, anoles and blindsnakes), it is more important for the data to reflect the relative values of these life history parameters across all anoles (and all reptiles) than it is to specifically and precisely represent all known variation within a given species of anole.
Without further ado (for your enjoyment, and because I know from my own blog that nobody reads posts lacking pictures, I’ve embedded an image of each species):
Species
Median clutch size
Median clutches per year
Incubation time (days)
Maximum longevity (months)
A. aeneus
2
A. baleatus
A.bimaculatus
2
43
84
A. carolinensis
1.15
6
41.5
65
A.chlorocyanus
1
18
A. conspersus
1
A. cristatellus
2.5
18
83
A. cybotes
1
18
45
A. distichus
1
16
45.5
A. equestris
1
1
48
149
A. extremus
A. ferreus
1
18
A. garmani
1.5
18
67
A. grahami
1
A. leachii
A. lineatus
A. lucius
1
3.5
60
A. marmoratus
2
50
A. maynardi
A. porcatus
1
18
63.5
A. pulchellus
1
A. richardii
1
A. sagrei
2
20
32
22
A. stratulus
A. trinitatis
2
50
A. wattsi
1
Thanks in advance. I think this is a great blog and I hope to post something more interesting on here soon.
