Bermuda as an Evolutionary Life Raft for an Ancient Lineage of Endangered Lizards

Oceanic islands are well known for harboring diverse species assemblages and are frequently the basis of research on adaptive radiation and neoendemism. However, a commonly overlooked role of some islands is their function in preserving ancient lineages that have become extinct everywhere else (paleoendemism). The island archipelago of Bermuda is home to a single species of extant terrestrial vertebrate, the endemic skink Plestiodon (formerly Eumeces) longirostris. The presence of this species is surprising because Bermuda is an isolated, relatively young oceanic island approximately 1000 km from the eastern United States. Here, we apply Bayesian phylogenetic analyses using a relaxed molecular clock to demonstrate that the island of Bermuda, although no older than two million years, is home to the only extant representative of one of the earliest mainland North American Plestiodon lineages, which diverged from its closest living relatives 11.5 to 19.8 million years ago. This implies that, within a short geological time frame, mainland North American ancestors of P. longirostris colonized the recently emergent Bermuda and the entire lineage subsequently vanished from the mainland. Thus, our analyses reveal that Bermuda is an example of a “life raft” preserving millions of years of unique evolutionary history, now at the brink of extinction. Threats such as habitat destruction, littering, and non-native species have severely reduced the population size of this highly endangered lizard.     

Tree Rings and Ecology

Tree rings have long been recognized as a useful indicator of past environmental changes. Besides traditional applications in studies of climate change and archaeology, the technique of dendrochronology has been widely used in answering ecological questions. This has been partly reflected by the broad topics on tree rings and ecology presented at the 7th International Conference on Dendrochronology （ICD）, which was held in Beijing, June 11-17 2006. At this conference, Dr Henri Grissono-Mayer, the plenary speaker for the theme, defined dendroecology as the science that uses tree rings, dated to their exact year of formation, to analyze temporal and spatial relationships between living organisms and their environment.     

Communicating Phylogeny: Evolutionary Tree Diagrams in Museums

Tree of life diagrams are graphic representations of phylogeny—the evolutionary history and relationships of lineages—and as such these graphics have the potential to convey key evolutionary ideas and principles to a variety of audiences. Museums play a significant role in teaching about evolution to the public, and tree graphics form a common element in many exhibits even though little is known about their impact on visitor understanding. How phylogenies are depicted and used in informal science settings impacts their accessibility and effectiveness in communicating about evolution to visitors. In this paper, we summarize the analysis of 185 tree of life graphics collected from museum exhibits at 52 institutions and highlight some potential implications of how trees are presented that may support or hinder visitors’ understanding about evolution. While further work is needed, existing learning research suggests that common elements among the diversity of museum trees such as the inclusion of anagenesis and absence of time and shared characters might represent potential barriers to visitor understanding.     

On the Ecology of Coexistence in Parthenogenetic and Bisexual Lizards of the Genus Cnemidophorus

In the center of their distribution in North America, membersof a complex of parthenogenetic and bisexual lizards of thegenus Cnemidophorus may be found side by side with one or moreof the others in a bewildering variety of patterns and ecologicalassociations. Exactly why so many congeneric species are foundliving together and exactly what their ecological, physiologicaland geographical requirements are remains virtually unknown.Here 1 summarize all reported cases of sympatry among six speciesof Cnemidophorus and discuss in detail specific examples ofsome of these interactions based on several years of field collectionsand observations. In addition I present the first field experimentalstudy designed to understand the ecological relationships ofsympatry in Cnemidophorus.     

进化生态学的产生与发展

进化论、遗传学与生态学是在不同时期发展起来的独立学科,然而,随着科学的发展,们越来越认识到它们之间存在着密切的关系。Petrusewicz(1959)(见Shvarts,1977)曾写过《达尔文进化论是一种生态理论》一书,强调进化在本质上是一生态过程。Hutchinson(1965)写了一部题名为《生态学舞台和进化节目》一书,认为生态的布景可以作为进化过程的舞台。但是,最早提出进化生态学这一概念的则是Orians(1962)。进化生态学从产生到现在虽然只有20多年的历史,但已成为生态学研究的重要领域。下面从几个方面谈谈进化生态学的产生与发展。     

Generalization in Ecology and Evolutionary Biology: From Hypothesis to Paradigm

We argue that broad, simplegeneralizations, not specifically linked tocontingencies, will rarely approach truth in ecologyand evolutionary biology. This is because mostinteresting phenomena have multiple, interactingcauses. Instead of looking for single universaltheories to explain the great diversity of naturalsystems, we suggest that it would be profitable todevelop general explanatory frameworks. A frameworkshould clearly specify focal levels. The process orpattern that we wish to study defines our level offocus. The set of potential and actual states at thefocal level interacts with conditions at thecontiguous lower and upper levels of organization,through sets of many-to-one and one-to-manyconnections. The number of initiating conditions andtheir permutations at the lower level define thepotential states at the focal level, whereas theactual state is constrained by the upper-levelboundary conditions. The most useful generalizationsare explanatory frameworks, which are road maps tosolutions, rather than solutions themselves. Suchframeworks outline what is understood about boundaryconditions and initiating conditions so that aninvestigator can pick and choose what is required toeffectively understand a specific event or situation. We discuss these relationships in terms of examplesinvolving sex ratio and mating behavior, competitivehierarchies, insect life-histories and the evolutionof sex.     

Adaptive Divergence in Experimental Populations of Pseudomonas fluorescens. IV. Genetic Constraints Guide Evolutionary Trajectories in a Parallel Adaptive Radiation

The capacity for phenotypic evolution is dependent upon complex webs of functional interactions that connect genotype and phenotype. Wrinkly spreader (WS) genotypes arise repeatedly during the course of a model Pseudomonas adaptive radiation. Previous work showed that the evolution of WS variation was explained in part by spontaneous mutations in wspF, a component of the Wsp-signaling module, but also drew attention to the existence of unknown mutational causes. Here, we identify two new mutational pathways (Aws and Mws) that allow realization of the WS phenotype: in common with the Wsp module these pathways contain a di-guanylate cyclase-encoding gene subject to negative regulation. Together, mutations in the Wsp, Aws, and Mws regulatory modules account for the spectrum of WS phenotype-generating mutations found among a collection of 26 spontaneously arising WS genotypes obtained from independent adaptive radiations. Despite a large number of potential mutational pathways, the repeated discovery of mutations in a small number of loci (parallel evolution) prompted the construction of an ancestral genotype devoid of known (Wsp, Aws, and Mws) regulatory modules to see whether the types derived from this genotype could converge upon the WS phenotype via a novel route. Such types—with equivalent fitness effects—did emerge, although they took significantly longer to do so. Together our data provide an explanation for why WS evolution follows a limited number of mutational pathways and show how genetic architecture can bias the molecular variation presented to selection.UNDERSTANDING—and importantly, predicting—phenotypic evolution requires knowledge of the factors that affect the translation of mutation into phenotypic variation—the raw material of adaptive evolution. While much is known about mutation rate (e.g., Drake et al. 1998; Hudson et al. 2002), knowledge of the processes affecting the translation of DNA sequence variation into phenotypic variation is minimal.Advances in knowledge on at least two fronts suggest that progress in understanding the rules governing the generation of phenotypic variation is possible (Stern and Orgogozo 2009). The first stems from increased awareness of the genetic architecture underlying specific adaptive phenotypes and recognition of the fact that the capacity for evolutionary change is likely to be constrained by this architecture (Schlichting and Murren 2004; Hansen 2006). The second is the growing number of reports of parallel evolution (e.g., Pigeon et al. 1997; ffrench-Constant et al. 1998; Allender et al. 2003; Colosimo et al. 2004; Zhong et al. 2004; Boughman et al. 2005; Shindo et al. 2005; Kronforst et al. 2006; Woods et al. 2006; Zhang 2006; Bantinaki et al. 2007; McGregor et al. 2007; Ostrowski et al. 2008)—that is, the independent evolution of similar or identical features in two or more lineages—which suggests the possibility that evolution may follow a limited number of pathways (Schluter 1996). Indeed, giving substance to this idea are studies that show that mutations underlying parallel phenotypic evolution are nonrandomly distributed and typically clustered in homologous genes (Stern and Orgogozo 2008).While the nonrandom distribution of mutations during parallel genetic evolution may reflect constraints due to genetic architecture, some have argued that the primary cause is strong selection (e.g., Wichman et al. 1999; Woods et al. 2006). A means of disentangling the roles of population processes (selection) from genetic architecture is necessary for progress (Maynard Smith et al. 1985; Brakefield 2006); also necessary is insight into precisely how genetic architecture might bias the production of mutations presented to selection.Despite their relative simplicity, microbial populations offer opportunities to advance knowledge. The wrinkly spreader (WS) morphotype is one of many different niche specialist genotypes that emerge when experimental populations of Pseudomonas fluorescens are propagated in spatially structured microcosms (Rainey and Travisano 1998). Previous studies defined, via gene inactivation, the essential phenotypic and genetic traits that define a single WS genotype known as LSWS (Spiers et al. 2002, 2003) (Figure 1). LSWS differs from the ancestral SM genotype by a single nonsynonymous nucleotide change in wspF. Functionally (see Figure 2), WspF is a methyl esterase and negative regulator of the WspR di-guanylate cyclase (DGC) (Goymer et al. 2006) that is responsible for the biosynthesis of c-di-GMP (Malone et al. 2007), the allosteric activator of cellulose synthesis enzymes (Ross et al. 1987). The net effect of the wspF mutation is to promote physiological changes that lead to the formation of a microbial mat at the air–liquid interface of static broth microcosms (Rainey and Rainey 2003).Open in a separate windowFigure 1.—Outline of experimental strategy for elucidation of WS-generating mutations and their subsequent identity and distribution among a collection of independently evolved, spontaneously arising WS genotypes. The strategy involves, first, the genetic analysis of a specific WS genotype (e.g., LSWS) to identify the causal mutation, and second, a survey of DNA sequence variation at specific loci known to harbor causal mutations among a collection of spontaneously arising WS genotypes. For example, suppressor analysis of LSWS using a transposon to inactivate genes necessary for expression of the wrinkly morphology delivered a large number of candidate genes (top left) (Spiers et al. 2002). Genetic and functional analysis of these candidate genes (e.g., Goymer et al. 2006) led eventually to the identity of the spontaneous mutation (in wspF) responsible for the evolution of LSWS from the ancestral SM genotype (Bantinaki et al. 2007). Subsequent analysis of the wspF sequence among 26 independent WS genotypes (bottom) showed that 50% harbored spontaneous mutations (of different kinds; see Open in a separate windowFigure 2.—Network diagram of DGC-encoding pathways underpinning the evolution of the WS phenotype and their regulation. Overproduction of c-di-GMP results in overproduction of cellulose and other adhesive factors that determine the WS phenotype. The ancestral SBW25 genome contains 39 putative DGCs, each in principle capable of synthesizing the production of c-di-GMP, and yet WS genotypes arise most commonly as a consequence of mutations in just three DGC-containing pathways: Wsp, Aws, and Mws. In each instance, the causal mutations are most commonly in the negative regulatory component: wspF, awsX, and the phosphodiesterase domain of mwsR (see text).To determine whether spontaneous mutations in wspF are a common cause of the WS phenotype, the nucleotide sequence of this gene was obtained from a collection of 26 spontaneously arising WS genotypes (WS_A-Z) taken from 26 independent adaptive radiations, each founded by the same ancestral SM genotype (Figure 1): 13 contained mutations in wspF (Bantinaki et al. 2007). The existence of additional mutational pathways to WS provided the initial motivation for this study.

TABLE 1

Mutational causes of WS

Genetic Tests for Ecological and Allopatric Speciation in Anoles on an Island Archipelago

From Darwin''s study of the Galapagos and Wallace''s study of Indonesia, islands have played an important role in evolutionary investigations, and radiations within archipelagos are readily interpreted as supporting the conventional view of allopatric speciation. Even during the ongoing paradigm shift towards other modes of speciation, island radiations, such as the Lesser Antillean anoles, are thought to exemplify this process. Geological and molecular phylogenetic evidence show that, in this archipelago, Martinique anoles provide several examples of secondary contact of island species. Four precursor island species, with up to 8 mybp divergence, met when their islands coalesced to form the current island of Martinique. Moreover, adjacent anole populations also show marked adaptation to distinct habitat zonation, allowing both allopatric and ecological speciation to be tested in this system. We take advantage of this opportunity of replicated island coalescence and independent ecological adaptation to carry out an extensive population genetic study of hypervariable neutral nuclear markers to show that even after these very substantial periods of spatial isolation these putative allospecies show less reproductive isolation than conspecific populations in adjacent habitats in all three cases of subsequent island coalescence. The degree of genetic interchange shows that while there is always a significant genetic signature of past allopatry, and this may be quite strong if the selection regime allows, there is no case of complete allopatric speciation, in spite of the strong primae facie case for it. Importantly there is greater genetic isolation across the xeric/rainforest ecotone than is associated with any secondary contact. This rejects the development of reproductive isolation in allopatric divergence, but supports the potential for ecological speciation, even though full speciation has not been achieved in this case. It also explains the paucity of anole species in the Lesser Antilles compared to the Greater Antilles.     

Evolutionary Changes in Dopaminergic Modulation of Courtship Behavior in Cnemidophorus Whiptail Lizards

Preoptic dopamine release is integral to the display of copulatory behaviors in male mammals and birds. However, while the anatomical distributions of the dopamine synthesizing enzyme tyrosine hydroxylase are similar among vertebrates, evolutionary changes in the functional role of dopamine are poorly understood. In this study, we tested whether a dopamine D1 receptor agonist would facilitate the display of courtship and copulatory behaviors in two related Cnemidophorine lizards (Cnemidophorus inornatus and Cnemidophorus uniparens). Cnemidophorus lizards offer a unique system to study evolutionary changes in functionality because ancestral (e.g., C. inornatus) and descendant (e.g., C. uniparens) species can be studied in parallel. Cnemidophorus uniparens is an all-female, parthenogenetic species and is the triploid descendant of the sexual and diploid species C. inornatus. Here we report that in castrated male C. inornatus and ovariectomized C. uniparens a dopamine D1 agonist increased the proportion of individuals mounting and decreased the latency to mount. Moreover, there was a species difference in sensitivity to the agonist: Mounting was elicited at a lower dose in C. uniparens than in C. inornatus. One possible explanation for this heightened sensitivity in the triploid parthenogen is that, by virtue of the increased ploidy, the parthenogen has elevated levels of D1 receptor in limbic brain areas modulating courtship behavior.