Maternal transmission of bacterial microbiota during embryonic development in a viviparous lizard.

IMPORTANCE: We investigated the presence and diversity of bacteria in the embryos of the viviparous lizard Sceloporus grammicus and their amniotic environment. We compared this diversity to that found in the maternal intestine, mouth, and cloaca. We detected bacterial DNA in the embryos, albeit with a lower bacterial species diversity than found in maternal tissues. Most of the bacterial species detected in the embryos were also found in the mother, although not all of them. Interestingly, we detected a high similarity in the composition of bacterial species among embryos from different mothers. These findings suggest that there may be a mechanism controlling the transmission of bacteria from the mother to the embryo. Our results highlight the possibility that the interaction between maternal bacteria and the embryo may affect the development of the lizards.

Habitat thermal quality for Gopherus evgoodei in tropical deciduous forest and consequences of habitat modification by buffelgrass.

Tortoises of the genus Gopherus evolved in North America and have survived major environmental challenges in the past 40 million years. However, this genus now faces multiple anthropogenic threats, such as the introduction of invasive plant species. Buffelgrass (Cenchrus ciliaris) is considered one of the greatest threats to arid and tropical ecosystems, where gopher tortoises inhabit, because the grass displaces native flora and fauna. Modification of the environment as a result of this invasive plant portends an alteration of the available thermal landscape. The aim of this paper is twofold: 1) to evaluate the thermal quality of the primary habitat of Gopherus evgoodei (tropical deciduous forest [TDF], and 2) determine the potential thermal changes due to habitat modification by buffelgrass. First, we obtained data on body temperature of active tortoises in semi-captivity. Second, we measured the operative environmental temperature during 5 years at three sites south of Sonora, Mexico that support G. evgoodei: a) a pristine TDF (Conserved-TDF); b) a forest patch surrounded by introduced buffelgrass pasture (Partial-TDF); and c) an introduced buffelgrass pasture area (Buffel-Pasture). Our results demonstrate that the intact microhabitats within the TDF provide G. evgoodei with high thermal quality at both spatial and temporal scales. However modified habitat by buffelgrass had higher operative temperatures for G. evgoodei than TDF. The thermal quality of the sites disturbed with buffelgrass can exceed the thermal requirements of G. evgoodei by up to 25 °C. Finally, we discussed potential collateral effects of habitat modification by invasion by buffelgrass.

Thermoregulation of Liolaemus aparicioi (Iguania: Liolaemidae) along a 1000 m elevational gradient in La Paz Valley, La Paz, Bolivia.

Lizard species have diverse behavioral and physiological responses to thermo-environmental conditions, which allow them to inhabit a broad range of latitudes and elevations. Because the availability of suitable thermal resources is limited and more variable at high-elevation environments than at lower elevations, we expect high-elevation lizards to be constrained in their thermoregulation relative to lizards at lower elevations by the fewer available thermal resources to reach optimal temperatures (colder environment). We studied the thermal biology of an endemic and Critically Endangered lizard, Liolaemus aparicioi, to assess its thermal responses along a 1000 m elevational gradient in La Paz Valley from May to August of 2015 (dry season). We took field body and microhabitat temperatures at capture sites (substrate and air above ground), and body size (snout-vent length and mass) of individuals at Taypichullo (3000 m asl), Gran Jardín de la Revolución Municipal Park (3500 m asl), and Taucachi (4000 m asl) localities. Operative temperatures were taken from calibrated models deployed in different available microhabitats. Preferred temperatures and thermal tolerance limits were determined in laboratory settings for lizards from each locality. Field body, microhabitat, and operative temperatures decreased with increasing elevation and differed between sexes. Lizards at the high elevation locality had the lowest thermoregulatory efficiency as compared with the mid and lower elevation localities. In laboratory measurements, while the preferred temperatures varied between sexes, pooled preferred temperatures and thermal tolerances were similar in all localities. Although thermal resources at high elevation can limit thermoregulatory possibilities in L. aparicioi, behavioral microhabitat use, time allocated to thermoregulation, and physiological adjustments seem to be possible strategies to counteract thermal costs along elevational gradients.

Thermoregulatory strategies of three reclusive lizards (genus Xantusia) from the Baja California peninsula, Mexico, under current and future microenvironmental temperatures.

The thermal quality of the habitat is key for the regulation of body temperature in terrestrial ectotherms and, therefore, permits them to carry out their fundamental biological activities. In thermally heterogeneous environments, ectotherms might follow different behavioral or physiological strategies to maintain their body temperature within biologically adequate boundaries, for which they depend on microhabitat selection. These aspects are, thus, relevant in the context of habitat degradation and land-use change. In this study, we characterized the thermal ecology of three lizard species (genus Xantusia) that differ in microhabitat use along the Baja California peninsula, Mexico. We made three predictions: (1) the three species will follow different thermoregulatory strategies according to habitat thermal quality; (2) the thermal requirements and tolerances of these species will match the environmental or microenvironmental thermal conditions; and (3) due to their habitat and range restriction, the species studied will be highly vulnerable to climate change. Our results indicate the existence of thermoregulatory mechanisms in Xantusia to face thermal heterogeneity, including behavioral thermoregulation by choosing different microhabitats, shifts in activity periods, and adaptation to particular high thermal quality microhabitats. Furthermore, despite their association to specific microhabitats and specialized physiology, the studied species will not be adversely affected by climate change, as the increased microenvironmental temperatures will lead to a higher habitat thermal quality and lower costs of thermoregulation. However, we do not discard other indirect adverse effects of climate change not considered in this study.

Comparative thermal ecology parameters of the mexican dusky rattlesnake (Crotalus triseriatus).

Montane habitats exhibit a high degree of thermal heterogeneity, and thus provide considerable thermoregulatory challenges for ectotherms. Comparative analyses provide an opportunity to understand how variation in abiotic factors (e.g., operative temperatures, thermal quality) can affect life history traits within species. We studied the thermal ecology of three populations of the rattlesnake Crotalus triseriatus inhabiting different volcanoes in the central region of Mexico using the Hertz et al. (1993) protocol. The average body temperature of dusky rattlesnakes from the three study sites was 22.4 °C; mean active body temperature was higher in site 2 than in sites 1 and 3, but no differences between females, males and juveniles nor an interaction among site and sex was found. The thermal quality was low in the three sites, particularly in sites 1 and 3. Thermoregulation accuracy statistically differed among populations: individuals from site 2 were more accurate thermoregulating, while individuals from site 1 were the least accurate. Compared to other snakes, dusky rattlesnakes can be considered as a eurythermic species, which can often be active at relatively low body temperatures.

Sex determination systems in reptiles are related to ambient temperature but not to the level of climatic fluctuation.

BACKGROUND: Vertebrates exhibit diverse sex determination systems and reptiles stand out by having highly variable sex determinations that include temperature-dependent and genotypic sex determination (TSD and GSD, respectively). Theory predicts that populations living in either highly variable or cold climatic conditions should evolve genotypic sex determination to buffer the populations from extreme sex ratios, yet these fundamental predictions have not been tested across a wide range of taxa. RESULTS: Here, we use phylogenetic analyses of 213 reptile species representing 38 families (TSD = 101 species, GSD = 112 species) and climatic data to compare breeding environments between reptiles with GSD versus TSD. We show that GSD and TSD are confronted with the same level of climatic fluctuation during breeding seasons. However, TSD reptiles are significantly associated with warmer climates. We found a strong selection on the breeding season length that minimises exposure to cold and fluctuating climate. Phylogenetic path analyses comparing competing evolutionary hypotheses support that transitions in sex determination systems influenced the ambient temperature at which the species reproduces and nests. In turn, this interaction affects other variables such as the duration of the breeding season and life-history traits. CONCLUSIONS: Taken together, our results challenge long-standing hypotheses about the association between sex determination and climate variability. We also show that ambient temperature is important during breeding seasons and it helps explain the effects of sex determination systems on the geographic distribution of extant reptile species.

Novel placental structure in the Mexican gerrhonotine lizard, Mesaspis viridiflava (Lacertilia; Anguidae).

The evolution of viviparity alters the physical relationship between mothers and offspring and the prevalence of viviparity among squamate reptiles presents an opportunity to uncover patterns in the evolution of placental structure. Understanding the breadth of this diversity is limited because studies of placental structure and function have emphasized a limited number of lineages. We studied placental ontogeny using light microscopy for an embryological series of the Mexican gerrhonotine lizard, Mesaspis viridiflava. This species develops an elaborate yolk sac placenta, an omphaloplacenta, which receives vascular support arising in a structure known only from other gerrhonotine lizards. A prominent feature of the omphaloplacenta is a zone of uterine and embryonic epithelial cell hyperplasia located at the upper shoulder of the yolk mass, often extending above the yolk mass. The omphaloplacenta covers more than one-half of the surface area of maternal-embryonic contact. The chorioallantoic placenta has a more restricted distribution because the allantois remains in the embryonic hemisphere of the egg throughout development and lies internal to the vascular support for the omphaloplacenta in areas where they overlap. The structural profile of the chorioallantoic placenta indicates a potential for respiratory exchange and/or hemotrophic nutritive transport, while that of the omphaloplacenta suggests that nutritive transfer is primarily via histotrophy. An eggshell is present in the earliest embryonic stages examined but regresses relatively early in development. Placental specializations of this species are consistent with a pattern of matrotrophic embryonic nutrition and have evolved in a unique lineage specific developmental pattern.

Inferring introgression using RADseq and DFOIL : Power and pitfalls revealed in a case study of spiny lizards (Sceloporus).

Introgression is now commonly reported in studies across the Tree of Life, aided by recent advancements in data collection and analysis. Nevertheless, researchers working with nonmodel species lacking reference genomes may be stymied by a mismatch between available resources and methodological demands. In this study, we demonstrate a fast and simple approach for inferring introgression using RADseq data, and apply it to a case study involving spiny lizards (Sceloporus) from northeastern México. First, we find evidence for recurrent mtDNA introgression between the two focal species based on patterns of mito-nuclear discordance. We then test for nuclear introgression by exhaustively applying the "five-taxon" D-statistic (DFOIL ) to all relevant individuals sampled for RADseq data. In our case, this exhaustive approach (dubbed "ExDFOIL ") entails testing up to ~250,000 unique four-taxon combinations of individuals across species. To facilitate use of this ExDFOIL approach, we provide scripts for many relevant tasks, including the selection of appropriate four-taxon combinations, execution of DFOIL tests in parallel and visualization of introgression results in phylogenetic and geographic space. Using ExDFOIL , we find evidence for ancient introgression between the focal species. Furthermore, we reveal geographic variation in patterns of introgression that is consistent with patterns of mito-nuclear discordance and with recurrent introgression. Overall, our study demonstrates that the combination of DFOIL and RADseq data can effectively detect introgression under a variety of sampling conditions (for individuals, populations and loci). Importantly, we also find evidence that batch-specific error and linkage in RADseq data may mislead inferences of introgression under certain conditions.

Behavioral and physiological polymorphism in males of the austral lizard Liolaemus sarmientoi.

Integrative behavioral studies show that the interplay between individual physiology and social behavior influences the ecology of the species, ultimately affecting individual fitness. Particularly in lizards, color polymorphism is associated with differential behaviors and reproductive strategies, which are evident in mature males during the mating season. Dominant males generally have greater endurance, higher body temperature, and larger bodies than submissive males, so they can acquire and defend larger territories and have greater access to females for mating. We studied whether the color morphs observed in males of one of the world's southernmost reptiles, Liolaemus sarmientoi, are related to behavioral variation during agonistic interactions, thermal physiology, morphology, and/or locomotor stamina. Liolaemus sarmientoi males exhibit three color morphs: red (RR), red-yellow (RY), and yellow (YY). These lizards exhibit subtle behavioral displays and we did not observe stamina differences among morphs. However, we found that RR males are more aggressive than YY males during agonistic encounters. In addition, greater body temperature change during trials, higher field body temperatures, and greater head sizes of RR males compared to RY or YY indicate that RR is a dominant morph, which may influence their ability to acquire and defend territory and tactics for achieving reproductive success.

Estimating the temporal and spatial extent of gene flow among sympatric lizard populations (genus Sceloporus) in the southern Mexican highlands.

Interspecific gene flow is pervasive throughout the tree of life. Although detecting gene flow between populations has been facilitated by new analytical approaches, determining the timing and geography of hybridization has remained difficult, particularly for historical gene flow. A geographically explicit phylogenetic approach is needed to determine the overlap of ancestral populations. In this study, we performed population genetic analyses, species delimitation, simulations and a recently developed approach of species tree diffusion to infer the phylogeographic history, timing and geographic extent of gene flow in lizards of the Sceloporus spinosus group. The two species in this group, S. spinosus and S. horridus, are distributed in eastern and western portions of Mexico, respectively, but populations of these species are sympatric in the southern Mexican highlands. We generated data consisting of three mitochondrial genes and eight nuclear loci for 148 and 68 individuals, respectively. We delimited six lineages in this group, but found strong evidence of mito-nuclear discordance in sympatric populations of S. spinosus and S. horridus owing to mitochondrial introgression. We used coalescent simulations to differentiate ancestral gene flow from secondary contact, but found mixed support for these two models. Bayesian phylogeography indicated more than 60% range overlap between ancestral S. spinosus and S. horridus populations since the time of their divergence. Isolation-migration analyses, however, revealed near-zero levels of gene flow between these ancestral populations. Interpreting results from both simulations and empirical data indicate that despite a long history of sympatry among these two species, gene flow in this group has only recently occurred.

Thermoregulation of two sympatric species of horned lizards in the Chihuahuan Desert and their local extinction risk.

Thermoregulatory studies of ectothermic organisms are an important tool for ecological physiology, evolutionary ecology and behavior, and recently have become central for evaluating and predicting global climate change impacts. Here, we present a novel combination of field, laboratory, and modeling approaches to examine body temperature regulation, habitat thermal quality, and hours of thermal restriction on the activity of two sympatric, aridlands horned lizards (Phrynosoma cornutum and Phrynosoma modestum) at three contrasting Chihuahuan Desert sites in Mexico. Using these physiological data, we estimate local extinction risk under predicted climate change within their current geographical distribution. We followed the Hertz et al. (1993, Am. Nat., 142, 796-818) protocol for evaluating thermoregulation and the Sinervo et al. (2010, Science, 328, 894-899) eco-physiological model of extinction under climatic warming. Thermoregulatory indices suggest that both species thermoregulate effectively despite living in habitats of low thermal quality, although high environmental temperatures restrict the activity period of both species. Based on our measurements, if air temperature rises as predicted by climate models, the extinction model projects that P. cornutum will become locally extinct at 6% of sites by 2050 and 18% by 2080 and P. modestum will become extinct at 32% of sites by 2050 and 60% by 2080. The method we apply, using widely available or readily acquired thermal data, along with the modeling, appeared to identify several unique ecological traits that seemingly exacerbate climate sensitivity of P. modestum.

Molecular phylogenetics and species delimitation of leaf-toed geckos (Phyllodactylidae: Phyllodactylus) throughout the Mexican tropical dry forest.

Methods and approaches for accurate species delimitation continue to be a highly controversial subject in the systematics community. Inaccurate assessment of species' limits precludes accurate inference of historical evolutionary processes. Recent evidence suggests that multilocus coalescent methods show promise in delimiting species in cryptic clades. We combine multilocus sequence data with coalescence-based phylogenetics in a hypothesis-testing framework to assess species limits and elucidate the timing of diversification in leaf-toed geckos (Phyllodactylus) of Mexico's dry forests. Tropical deciduous forests (TDF) of the Neotropics are among the planet's most diverse ecosystems. However, in comparison to moist tropical forests, little is known about the mode and tempo of biotic evolution throughout this threatened biome. We find increased speciation and substantial, cryptic molecular diversity originating following the formation of Mexican TDF 30-20million years ago due to orogenesis of the Sierra Madre Occidental and Mexican Volcanic Belt. Phylogenetic results suggest that the Mexican Volcanic Belt, the Rio Fuerte, and Isthmus of Tehuantepec may be important biogeographic barriers. Single- and multilocus coalescent analyses suggest that nearly every sampling locality may be a distinct species. These results suggest unprecedented levels of diversity, a complex evolutionary history, and that the formation and expansion of TDF vegetation in the Miocene may have influenced subsequent cladogenesis of leaf-toed geckos throughout western Mexico.

Landscape genetics of leaf-toed geckos in the tropical dry forest of northern Mexico.

Habitat fragmentation due to both natural and anthropogenic forces continues to threaten the evolution and maintenance of biological diversity. This is of particular concern in tropical regions that are experiencing elevated rates of habitat loss. Although less well-studied than tropical rain forests, tropical dry forests (TDF) contain an enormous diversity of species and continue to be threatened by anthropogenic activities including grazing and agriculture. However, little is known about the processes that shape genetic connectivity in species inhabiting TDF ecosystems. We adopt a landscape genetic approach to understanding functional connectivity for leaf-toed geckos (Phyllodactylus tuberculosus) at multiple sites near the northernmost limit of this ecosystem at Alamos, Sonora, Mexico. Traditional analyses of population genetics are combined with multivariate GIS-based landscape analyses to test hypotheses on the potential drivers of spatial genetic variation. Moderate levels of within-population diversity and substantial levels of population differentiation are revealed by FST and Dest. Analyses using structure suggest the occurrence of from 2 to 9 genetic clusters depending on the model used. Landscape genetic analysis suggests that forest cover, stream connectivity, undisturbed habitat, slope, and minimum temperature of the coldest period explain more genetic variation than do simple Euclidean distances. Additional landscape genetic studies throughout TDF habitat are required to understand species-specific responses to landscape and climate change and to identify common drivers. We urge researchers interested in using multivariate distance methods to test for, and report, significant correlations among predictor matrices that can impact results, particularly when adopting least-cost path approaches. Further investigation into the use of information theoretic approaches for model selection is also warranted.

Variación morfológica de la lagartija partenogenética Aspidoscelis rodecki (Squamata: Teiidae): implicaciones evolutivas y de conservación

Morphologic variation of the parthenogenetic lizard Aspidoscelis rodecki (Squamata: Teiidae): evolutionary and conservation implications. Post-formational divergence has been used for the recognition of new parthenogenetic species. Currently, the parthenogenetic lizard Aspidoscelis rodecki McCoy and Maslin 1962 is recognized as a single taxon that was derived from a single, parthenogenetically capable, hybrid. This lizard had been derived via hybridization between individuals of two gonochoristic species, Aspidoscelis angusticeps Cope 1878 and Aspidoscelis deppii Wiegmann 1834. The distribution of A. rodecki includes Isla Contoy and Isla Mujeres and the adjacent mainland of Quintana Roo, México. Previous studies have found post-formational divergence in genetic, chromatic and life-history characteristics among a continental population (Puerto Juárez) and an insular population (Isla Contoy). A meristic analysis was carried out to evaluate the morphological divergence among both populations of A. rodecki. We used 38 individuals from Puerto Juárez and 23 individuals from Isla Contoy. Nine meristic characters with discrimination value among species of the genus Aspidoscelis were used in both univariate (t-Student) and multivariate analyses (principal components and canonical variate analysis). According to both analyses, Puerto Juárez is meristically distinguishable from Isla Contoy. Both populations differ in five meristic characters and were a high correct classification in the canonical variate analysis: 97% of Puerto Juárez and 100% of Isla Contoy. A small sample from Isla Mujeres and a single specimen from Punta Sam (mainland) may represent different morphological groups. Due to the patterns of phenotypic variation, A. rodecki is considered as a single variable parthenogenetic species with high priority to conservation. The populations of A. rodecki have been extremely affected by the tourism developers. If the habitat of the parthenogenetic lizard (beach grasses) is allowed to stay, the expansion by the developers will not affect the survivorship of these populations. Nevertheless, the first sign of development is the total destruction of natural grasses that occurs on the beach, leaving only sand. There is a last chance to save the parthenogenetic lizard A. rodecki, but any effort will be useless without the support from the environmental authority of Mexico and cooperation from the developers. We suggest that Puerto Juárez and Isla Contoy receive separate management because they have unique portions of phenotypic variation of A. rodecki. The two lizard populations can be considered separate "Evolutionary Significant Units" (ESU). Rev. Biol. Trop. 56 (4): 1871-1881. Epub 2008 December 12.

La divergencia post-formación se ha utilizado para el reconocimiento de nuevas especies partenogenéticas. Actualmente, la lagartija partenogenética Aspidoscelis rodecki McCoy y Maslin 1962 es reconocida como una sola especie, que se originó de un híbrido partenogenético. Estudios previos han encontrado divergencia genética, en coloración y en características de historia de vida entre una población continental (Puerto Juárez) y una insular (Isla Contoy) en Quintana Roo, México. Se llevó a cabo un análisis merístico para evaluar la divergencia entre ambas poblaciones de A. rodecki. Se utilizaron 38 individuos de Puerto Juárez y 23 individuos de Isla Contoy. Se usaron nueve características merísticas y se realizaron análisis univariados (t de Student) y multivariados (análisis de componentes principales y análisis de variación canónica). De acuerdo a ambos análisis, Puerto Juárez es merísticamente distinguible de Isla Contoy. Ambas poblaciones difieren en cinco características merísticas y presentaron un alto porcentaje de clasificación en el análisis de variación canónica: 97% para Puerto Juárez y 100% para Isla Contoy. Una pequeña muestra de Isla Mujeres y un solo individuo de Punta Sam (continente) pueden representar otros grupos morfológicos. Con base en los patrones de variación fenotípica, A. rodecki es considerada como una sola especie partenogenética variable y prioritaria para la conservación. Debido a que cada población contiene una porción única de la variación de A. rodecki, Puerto Juárez e Isla Contoy merecen un manejo separado y se sugiere que cada una constituya una "Unidad Evolutiva Significativa" (ESU).

[Morphologic variation of the parthenogenetic lizard Aspidoscelis rodecki (Squamata: Teiidae): evolutionary and conservation implications]. / Variación morfológica de la lagartija partenogenética Aspidoscelis rodecki (Squamata: Teiidae): implicaciones evolutivas y de conservación.

Post-formational divergence has been used for the recognition of new parthenogenetic species. Currently, the parthenogenetic lizard Aspidoscelis rodecki McCoy and Maslin 1962 is recognized as a single taxon that was derived from a single, parthenogenetically capable, hybrid. This lizard had been derived via hybridization between individuals of two gonochoristic species, Aspidoscelis ungusticeps Cope 1878 and Aspidoscelis deppii Wiegmann 1834. The distribution of A. rodecki includes Isla Contoy and Isla Mujeres and the adjacent mainland of Quintana Roo, México. Previous studies have found post-formational divergence in genetic, chromatic and life-history characteristics among a continental population (Puerto Juárez) and an insular population (Isla Contoy). A meristic analysis was carried out to evaluate the morphological divergence among both populations of A. rodecki. We used 38 individuals from Puerto Juárez and 23 individuals from Isla Contoy. Nine meristic characters with discrimination value among species of the genus Aspidoscelis were used in both univariate (t-Student) and multivariate analyses (principal components and canonical variate analysis). According to both analyses, Puerto Juárez is meristically distinguishable from Isla Contoy. Both populations differ in five meristic characters and were a high correct classification in the canonical variate analysis: 97% of Puerto Juárez and 100% of Isla Contoy. A small sample from Isla Mujeres and a single specimen from Punta Sam (mainland) may represent different morphological groups. Due to the patterns of phenotypic variation, A. rodecki is considered as a single variable parthenogenetic species with high priority to conservation. The populations of A. rodecki have been extremely affected by the tourism developers. If the habitat of the parthenogenetic lizard (beach grasses) is allowed to stay, the expansion by the developers will not affect the survivorship of these populations. Nevertheless, the first sign of development is the total destruction of natural grasses that occurs on the beach, leaving only sand. There is a last chance to save the parthenogenetic lizard A. rodecki, but any effort will be useless without the support from the environmental authority of Mexico and cooperation from the developers. We suggest that Puerto Juárez and Isla Contoy receive separate management because they have unique portions of phenotypic variation of A. rodecki. The two lizard populations can be considered separate "Evolutionary Significant Units" (ESU).

Deep genealogical history without population differentiation: discordance between mtDNA and allozyme divergence in the zebra-tailed lizard (Callisaurus draconoides).

The peninsula of Baja California has a complex geological history that has strongly affected the regional biota. Genealogical histories of many species have revealed congruent patterns, which suggest that the peninsula was temporarily submerged at two locations. We sequenced a total of 1953 base pairs (bp) of the mitochondrial genome for 42 specimens of the zebra-tailed lizard (Callisaurus draconoides). The resulting maternal genealogy supports the former existence of a mid-peninsular seaway and a Plio-Quaternary seaway across the Isthmus of La Paz. In addition, a genealogical break is revealed in the vicinity of Loreto. This genealogical break may have resulted from prolonged submergence of the Loreto Basin during Pliocene. The mid-peninsular seaway may have occurred as early as late Miocene, at a time significantly earlier than previously hypothesized. Comparison with other genealogies and geological evidence suggests that current models on the evolution of Baja California's fauna are temporally shallow. The deep genealogical patterns of C. draconoides also disagree with the very limited population differentiation previously reported for allozyme markers, suggesting that maternal history may not be an appropriate approximation for population differentiation.