Fire Increases Genetic Diversity of Populations of Six-Lined Racerunner.

Wildfires are highly variable and can disturb habitats, leading to direct and indirect effects on the genetic characteristics of local populations. Florida scrub is a fire-dependent, highly fragmented, and severely threatened habitat. Understanding the effect of fire on genetic characteristics of the species that use this habitat is critically important. We investigated one such lizard, the Six-lined Racerunner (Aspidoscelis sexlineata), which has a strong preference for open areas. We collected Six-lined Racerunners (n = 154) from 11 sites in Highlands County, FL, and defined 2 time-since-last-fire (TSF) categories: recently burned and long unburned. We screened genetic variation at 6 microsatellites to estimate genetic differentiation and compare genetic diversity among sites to determine the relationship with TSF. A clear pattern exists between genetic diversity and TSF in the absence of strong genetic differentiation. Genetic diversity was greater and inbreeding was lower in sites with more recent TSF, and genetic characteristics had significantly larger variance in long unburned sites compared with more recently burned sites. Our results suggest that fire suppression increases variance in genetic characteristics of the Six-lined Racerunner. More generally, fire may benefit genetic characteristics of some species that use fire-dependent habitats and management efforts for such severely fragmented habitat will be challenged by the presence of multiple species with incompatible fire preferences.

Fire alters patterns of genetic diversity among 3 lizard species in Florida Scrub habitat.

The Florida Sand Skink (Plestiodon reynoldsi), the Florida Scrub Lizard (Sceloporus woodi), and the Six-lined Racerunner (Aspidoscelis sexlineata) occur in the threatened and fire-maintained Florida scrub habitat. Fire may have different consequences to local genetic diversity of these species because they each have different microhabitat preference. We collected tissue samples of each species from 3 sites with different time-since-fire: Florida Sand Skink n = 73, Florida Scrub Lizard n = 70, and Six-lined Racerunner n = 66. We compared the effect of fire on genetic diversity at microsatellite loci for each species. We screened 8 loci for the Florida Sand Skink, 6 loci for the Florida Scrub Lizard, and 6 loci for the Six-lined Racerunner. We also tested 2 potential driving mechanisms for the observed change in genetic diversity, a metapopulation source/sink model and a local demographic model. Genetic diversity varied with fire history, and significant genetic differentiation occurred among sites. The Florida Scrub Lizard had highest genetic variation at more recently burned sites, whereas the Florida Sand Skink and the Six-lined Racerunner had highest genetic variation at less recently burned sites. Habitat preferences of the Florida Sand Skink and the Florida Scrub Lizard may explain their discordant results, and the Six-lined Racerunner may have a more complicated genetic response to fire or is acted on at a different geographic scale than we have investigated. Our results indicate that these species may respond to fire in a more complicated manner than predicted by our metapopulation model or local demographic model. Our results show that the population-level responses in genetic diversity to fire are species-specific mandating conservation management of habitat diversity through a mosaic of burn frequencies.

Phenotypic population divergence in terrestrial vertebrates at macro scales.

Phenotypic divergence between populations, i.e. how much phenotypes within a species vary geographically, is critical to many aspects of ecology and evolution, including eco-geographical trends, speciation and coexistence. Yet, the variation of divergence across species with different ecologies and distributions and the relative role of adaptive causes remains little understood. We predict that genetic control vs. phenotypic plasticity of traits, geographical distance and (assuming adaptation) environmental differences should explain much of the phenotypic variability between populations. We tested these predictions with body sizes of 1447 populations in 98 terrestrial vertebrate species. Population phenotypic variability differs strongly across species, and divergence increases with increasing levels of clade-typical phenotypic plasticity, the area covered by populations and body size. Geographical distance and environmental dissimilarity are similarly important predictors of divergence within species, highlighting a potential role for biotic and environmental conditions. Increased availability of phylogeographical and ecological data should facilitate further understanding of population divergence drivers at broad scales.

When Rensch meets Bergmann: does sexual size dimorphism change systematically with latitude?

Bergmann's and Rensch's rules describe common large-scale patterns of body size variation, but their underlying causes remain elusive. Bergmann's rule states that organisms are larger at higher latitudes (or in colder climates). Rensch's rule states that male body size varies (or evolutionarily diverges) more than female body size among species, resulting in slopes greater than one when male size is regressed on female size. We use published studies of sex-specific latitudinal body size clines in vertebrates and invertebrates to investigate patterns equivalent to Rensch's rule among populations within species and to evaluate their possible relation to Bergmann's rule. Consistent with previous studies, we found a continuum of Bergmann (larger at higher latitudes: 58 species) and converse Bergmann body size clines (larger at lower latitudes: 40 species). Ignoring latitude, male size was more variable than female size in only 55 of 98 species, suggesting that intraspecific variation in sexual size dimorphism does not generally conform to Rensch's rule. In contrast, in a significant majority of species (66 of 98) male latitudinal body size clines were steeper than those of females. This pattern is consistent with a latitudinal version of Rensch's rule, and suggests that some factor that varies systematically with latitude is responsible for producing Rensch's rule among populations within species. Identifying the underlying mechanisms will require studies quantifying latitudinal variation in sex-specific natural and sexual selection on body size.

The phylogeny of a species-level tendency: species heritability and possible deep origins of Bergmann's rule in tetrapods.

One of the most widely recognized generalizations in biology is Bergmann's rule, the observation that, within species of birds and mammals, body size tends to be inversely related to ambient temperature. Recent studies indicate that turtles and salamanders also tend to follow Bergmann's rule, which hints that this species-level tendency originated early in tetrapod history. Furthermore, exceptions to Bergmann's rule are concentrated within squamate reptiles (lizards and snakes), suggesting that the tendency to express a Bergmann's rule cline may be heritable at the species level. We evaluated species-level heritability and early origination of Bergmann's rule by mapping size-latitude relationships for 352 species onto a tetrapod phylogeny. When the largest available dataset is used, Bergmann's rule shows significant phylogenetic signal, indicating species-level heritability. This represents one of the few demonstrations of heritability for an emergent species-level property and the first for an ecogeographic rule. When species are discretely coded as showing either Bergmann's rule or its converse, parsimony reconstructions suggest that: (1) the tendency to follow Bergmann's rule is ancestral for tetrapods, and (2) most extant species that express the rule have retained this tendency from that ancient ancestor. The first inference also generally holds when the discrete data or size-latitude correlation coefficients are analyzed using maximum likelihood, although the results are only statistically significant for some versions of the discrete analyses. The best estimates of ancestral states suggest that the traditional adaptive explanation for Bergmann's rule-conservation of metabolic heat-was not involved in the origin of the trait since that origin predates the evolution of endothermy. A more general thermoregulatory hypothesis could apply to endotherms and some ectotherms, but fails to explain why salamanders have retained Bergmann's rule. Thus, if thermoregulation underlies the origin of a Bergmann's rule tendency, this trait may have been continuously maintained while its cause changed. Alternatively, thermoregulation may not underlie Bergmann's rule in any tetrapod group. The results also suggest that many extinct groups not included in our analyses followed Bergmann's rule.

Sensitivity of intraspecific latitudinal clines of body size for tetrapods to sampling, latitude and body size.

Recent studies have shown that most tetrapod groups (mammals, birds, chelonians, amphibians) show general intraspecific tendencies for increasing body size with latitude, whereas squamates (lizards and snakes) show an intraspecific tendency towards decreasing body size with latitude. Here I evaluate whether these size trends are general by using independent contrasts analysis to investigate the dependence of intraspecific size-latitude relationships (r), and the magnitude alone of size-latitude relationships ([r]), for tetrapod vertebrates, on sample size, range of latitudes sampled, average latitude sampled, and body size. Range of latitudes sampled, average latitude sampled, and body size did not influence body size-latitude relationships (r) or the magnitude alone of body size-latitude relationship ([r]). Sample size did not influence size-latitude relationships (r), but did influence the magnitude alone of size-latitude relationships ([r]), possibly indicating increased precision of estimating size-latitude relationships with increased sampling. In short, intraspecific size-latitude relationships are similar for species of different sizes, occurring at different latitudes, sampled over different latitudinal ranges, and differing in number of populations sampled (though magnitude alone is influenced by sample size). These results suggest that intraspecific size-latitude trends are general, and biologically significant (i.e., are not artifacts of sampling), thus deserving explanation.

Bergmann's rule in nonavian reptiles: turtles follow it, lizards and snakes reverse it.

Bergmann's rule is currently defined as a within-species tendency for increasing body size with increasing latitude or decreasing environmental temperature. This well-known ecogeographic pattern has been considered a general trend for all animals, yet support for Bergmann's rule has only been demonstrated for mammals and birds. Here we evaluate Bergmann's rule in two groups of reptiles: chelonians (turtles) and squamates (lizards and snakes). We perform both nonphylogenetic and phylogenetic analyses and show that chelonians follow Bergmann's rule (19 of 23 species increase in size with latitude: 14 of 15 species decrease in size with temperature), whereas squamates follow the converse to Bergmann's rule (61 of 83 species decrease in size with latitude; 40 of 56 species increase in size with temperature). Size patterns of chelonians are significant using both nonphylogenetic and phylogenetic methods, whereas only the nonphylogenetic analyses are significant for squamates. These trends are consistent among major groups of chelonians and squamates for which data are available. This is the first study to document the converse to Bergmann's rule in any major animal group as well as the first to show Bergmann's rule in a major group of ectotherms. The traditional explanation for Bergmann's rule is that larger endothermic individuals conserve heat better in cooler areas. However, our finding that at least one ectothermic group also follows Bergmann's rule suggests that additional factors may be important. Several alternative processes, such as selection for rapid heat gain in cooler areas, may be responsible for the converse to Bergmann's rule in squamates.

Is Bergmann's Rule Valid for Mammals?

Bergmann's rule states that, within species of mammals, individuals tend to be larger in cooler environments. However, the validity of the rule has been debated. We examined the relationship between size and latitude as well as size and temperature within various species of mammals. We also tested the idea that smaller mammals follow Bergmann's rule more strongly than larger mammals, as expected if heat conservation is the cause of the rule. When all studies were included, the percentage of species showing a positive correlation between size and latitude was significantly >50% (78 of 110 species). Similarly, the percentage of species showing a negative correlation between size and temperature was significantly >50% (48 of 64). Analyses using only significant studies or only studies that sampled extensively also support Bergmann's rule. The size-latitude and size-temperature trends were consistent within all orders and most families for which data are available. We did not find support for the hypothesis that smaller mammals conform more strongly to Bergmann's rule than larger mammals. Thus, we found broad support for Bergmann's rule as a general trend for mammals; however, our analyses do not support heat conservation as the explanation.