Author: Julian Velasco

Are There Solitary Mainland Anolis Lizards?

Anolis (Phenoacosaurus) heterodermus, a mainland anole that co-occurs with few or no other anole species

One of the most important questions in ecology and evolution is about the role of biotic interactions in driving phenotypic and behavioral changes across species. The insular Anolis species are a good model to address this kind of question due to their high abundance and pervasive ecological interactions across islands. Some insular species, however, live in isolation on small islands across the Pacific and Caribbean islands (21 species). These species have evolved similar morphologies across islands. For instance, Poe et al (1) found that body size evolved by exaptation (remember the classic Gould and Vrba 1982 paper) to colonize these small (and depauperate) islands successfully. By contrast, Poe et al. (1) showed that sexual size dimorphism (SSD) evolved by adaptation likely after island colonization to minimize intraspecific competition.

In brief, these solitary insular anoles evolved phenotypic (body size and SSD) traits by two different processes. Cool! But, what happens in mainland areas? Much work has been devoted to Caribbean species, but the mainland offers many more species and very little research has been conducted there to understand ecological and evolutionary processes. So, we decided to establish whether solitary ecology can be extended to mainland species or whether it is an island ecological phenomena.

The first problem that we had to resolve was trying to establish whether mainland species tend to live in geographical/ecological isolation as insular species. We adopted a novel concept in macroecology (the diversity field concept) developed by Mexican macroecologists (Hector Arita and Fabricio Villalobos see 2, 3) implemented here using extensive distributional information for almost all known Anolis species (377 spp), which I generated during my Ph.D. thesis (see 4 for an example using these maps). The diversity field concept allows us to establish how many species co-occur with a given species across its geographic range.

We calculated how many congeners can co-occur within the distributional area of each Anolis species using the range maps (see figure below). We divided mainland species into two groups: those co-occurring with few congeners (i.e., “solitary-like”, I had to say that his term did not like to reviewers, so we used a “species-poor” forms in the paper). Then, we test whether these “solitary-like” mainland species are different from other mainland species using a randomization approach. Our results revealed that “solitary-like” mainland species exhibit different traits from random mainland assemblages. These unique traits (i.e., uniform body size and greater SSD) suggest that solitary ecology from insular anoles can be extended to mainland settings.

 

field_diversity

Figure. Diversity fields for some Anolis species. Note that the diversity field is the set of richness values of co-occurring anoles inside each distributional area.

The next question was focused to establish whether similar (ecological and evolutionary) processes affected body size and SSD patterns in a similar way. We found that the phylogenetic position of body size and SSD shifts did not coincide and also with the evolutionary transitions to solitariness (i.e., reduced level of sympatry). We suggested that both traits are decoupled across the entire Anolis radiation and likely that both traits evolved exaptatively. In other words, it is possible to think that “solitary-like” species retained body size and SSD from their most recent common ancestors to facilitates the lonely life.

The paper is very short (less than 2500 words) and was published in the May number of Biology Letters(5).

 

Why Are Most Caribbean Anolis Species Endemic to a Single Island?

Available climatic space showing the position of each pixel predicted as presence from ecological niche modeling across all Caribbean islands.

One of the most interesting patterns in the insular anole radiation is the observation that the majority of species are single-island endemics (150 of out 166 species). This observation in the Caribbean anole lizards has been known from a while and several studies have attempted to establish the underlying causes of this striking pattern (e.g., 1, 2, 3).

In a recent study, as part of my PhD dissertation, I used a different approach to try to understand why most of these species are unable to colonize other islands. I used a recently developed conceptualization to link abundances and ecological niche requirements at coarse-grain scales; this approach has been developed in the lab of my advisor (see 4, 5, 6; but see 7, 8, 9 for discussions and counter-examples; this approach has been strongly debated in the literature in the last years).

We used ecological niche modeling -ENM- to predict species’ distributions across all Caribbean islands for each species with at least 10 occurrence records. We estimated the position of each pixel predicted as presence in the ecological space using Euclidean distances. In short, we characterize all pixels for a single species and calculated which of these were close to the niche centroid (which we assume as the best conditions for species presence) and which were close to the niche periphery (see figure above). We predicted that pixels predicted by ENM as presences within each native island will be more close to the niche centroid and those predicted as presences in other islands will be in the periphery of the niche.

 

We found that many species follow the predicted pattern; in other words, we found that the “best” niche conditions are in the native island regardless of climatic heterogeneity observed in each island and the “worst” niche conditions are outside native islands. We also used other metrics to corroborate our results. We interpreted these results as  instances of recent climatic niche conservatism (within lineages) and therefore this operates as a constraint in the ability of each species to colonize other islands (i.e. due to the low suitable climatic conditions for successful population establishment). We only gathered data for 70 species and therefore it will be necessary more data and more studies (including physiological experiments) to corroborate our assertions.

Also, we examined the pattern of realized climatic niche shifts across the anole radiation and we found evidence of several instances of climatic niche convergence. We concluded that anoles evolved to occupy different portions of the climate space and in several cases evolved quickly to occupy some portions of this space (e.g., cold climatic conditions) and recently most of these species likely adapted very well to climatic conditions in their. native islands.

The paper was published in Evolutionary Biology.

 

 

 

Is There a Crisis in Anolis Taxonomy?

A few days ago, I had a very interesting discussion in twitter with some colleagues (see here [1] and here [2]) about whether there is a decline and crisis of new taxonomists in amphibians and reptiles. I wonder if this situation might be the case fir Anolis lizards, or whether Anolis has its own tendency in species description. So, I decided to take the bull by the horns and I went to the Uetz Reptilia Database to see the numbers.

First, it seems that there is not an increase in species description for anoles across time. In early years, it was a bunch of new descriptions as also recently (Fig 1).

Figure1

However, if you check how many anole taxonomists have described species, an interesting pattern emerges. Very few people have described almost all of anole diversity (Table 1). Only 15 anologists have described all currently known species (400 according to the Uetz database), and more interesting is the fact that only five of them are still alive! (Gunther Köhler, Orlando Garrido, James McCraine, Steven Poe and Larry Wilson; Table 1).

Table 1. Ranking of anole taxonomists by number of species described.

Table 1. Ranking of anole taxonomists by number of species described.

Then, to establish if new authors (new anole taxonomists) are emerging in recent times, I made an accumulation curve of authors across years of description (Fig. 2). I considered only the senior author (Fig. 2, top) and all authors involved in the description  (Fig. 2, bottom). From these figures, it is possible to see, apparently, that very few new authors are emerging as anole taxonomist specialists.

Accumulation of authors by year. Only senior author (top), all authors involved in the description (bottom)

Accumulation of authors by year. Only senior author (top), all authors involved in the description (bottom)

This suggests, at least to me, that something is happening in anole taxonomy. As we discussed in twitter, this scarcity in taxonomists in recent times could be due to several factors: lack of interest in describing new forms, lack of funding resources, lack of access to comparative material (i.e., herpetological collections), lack of writing skills to generate scientific papers, or lack of adequate taxonomic training. In any case, I feel that anole taxonomy would be in crisis if new people, mostly from Latin America, are not interested in describing and revising new species in each country. But, why are very few people seem to be interested in describing new valid species nowadays? Is it perhaps that taxonomy is a discipline with little interest in academia? Does doing so serve to get a job? Is it not worth publishing papers in taxonomy or even describing new forms?

Although today there are a bunch of journals oriented to taxonomy and many online resources to access to primary literature (Sci-Hub [3], BHL[4], etc.), it seems that very few people are interested to tackle these problems in our iconic lizards.

I would like to see your opinions on this

References

[1] https://twitter.com/CrawfordAJ/status/713398834005561344

[2] https://twitter.com/CrawfordAJ/status/713459231983280130

[3] https://en.wikipedia.org/wiki/Sci-Hub

[4] http://www.biodiversitylibrary.org

Can Climatic Niche Traits Drive Cladogenesis in Anolis Lizards?

Anolis onca basking. Photo by Gabriel Ugueto.

The ecological niche is one of the most controversial concepts in ecology with a long history of debate about its definition and scope. Some authors have suggested that this concept should be abandoned (see for example McInerny & Etienne, 2012a,b,c), while others, including me, consider that this controversial history, and a plethora of definitions, should not preclude its use. However, it needs to be explicitly defined to known exactly we are talking about.

The ecological niche is understood as the set of environmental factors that allow a local population to persist without immigration from others sources (Hutchinson 1959). Some evolutionary ecologists have envisioned the ecological niche as a phenotypic extension of a species and therefore subject to natural selection and other evolutionary process. Based on this, in my doctoral dissertation, I evaluated whether the ecological niche (or more precisely, the climatic niche), as a population-level trait, can promote species diversification in Anolis lizards. Although there are several studies linking organismal and species’ level traits with speciation, there are few exploring this association in Anolis lizards.

Our study exploring this was recently published in the Journal of Biogeography (Velasco et al. 2016). Explicitly, we adopt  Hutchinson’s niche definition using only coarse-grain variables (or the Grinnellian niche dimension; see Soberón 2007). To do this, we compiled an extensive occurrence database for 328 species with help of several colleagues. Using climatic layers from WorldClim database, we calculated a set of niche metrics for species and clades, including mean niche position (based on PCA analyses), niche breadth (based on Mahalanobis distances) and occupied niche space (as a proxy of climatic niche diversity).

We compared how climatic niche breadth and occupied niche space differ among regions (insular vs. mainland) and clades (Fig. 1 & 2). Mainland species and clades tend to exhibit larger niches than Caribbean clades and species (Fig. 1).

figure 1

These differences were maintained after controlling for range size differences. We suggest that these differences are directly related to the available climatic space in each region. For instance,  Caribbean islands exhibit a limited climatic space in comparison with  mainland regions and therefore insular clades occupy all available climatic conditions in each island (Fig. 2). By contrast, mainland clades are more restricted to climatic conditions and therefore occupy only a portion of all available climatic conditions (Fig. 2).

figure 2

 

We correlated these niche metrics with species diversification using a calibrated time tree. We found that anole clades occupying warmer and drier areas diversified more than clades occupying very humid and colder areas. In addition, anole clades with narrow niches tend to speciate more than clades with widespread niches (Fig. 3). Our findings suggest that climatic specialization has played a strong role in anole diversification with differences among insular and mainland settings.

figure 3

It would be interesting to evaluate whether other traits (e.g., body size, geographical range size, or other Eltonian niche dimensions) also play a similar role on cladogenesis in Anolis lizards and evaluate their relative importance in diversification dynamics.

References

Hutchinson, G.E. (1957) Concluding remarks. Cold Spring Harbor Symposia on Quantitative Biology, 22, 415-427.

McInerny, G.J. & Etienne, R.S. (2012a) Ditch the niche ? is the niche a useful concept in ecology or species distribution modelling? J. Biogeogr., 39, 2096-2102.

McInerny, G.J. & Etienne, R.S. (2012b) Stitch the niche ? a practical philosophy and visual schematic for the niche concept. J. Biogeogr., 39, 2103-2111.

McInerny, G.J. & Etienne, R.S. (2012c) Pitch the niche ? taking responsibility for the concepts we use in ecology and species distribution modelling. J. Biogeogr, 39, 2112-2118.

Soberón, J. (2007) Grinnellian and Eltonian niches and geographic distributions of species. Ecology Letters, 10, 1115-1123.

Velasco, J. A., Martínez-Meyer, E., Flores-Villela, O., García A., Algar, A. C., Köhler, G. and Daza, J. M. (2016), Climatic niche attributes and diversification in Anolis lizards. J. Biogeogr., 43: 134-144. doi:10.1111/jbi.12627

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