Microsatellite variation and evolution in the Peromyscus maniculatus species group

Variation at 12 pure-repeat dinucleotide microsatellites from Peromyscus maniculatus was analyzed for samples of all species in the P. maniculatus species group and P. leucopus. Except for one locus (Pml08) that amplified a product only for P. maniculatus, these microsatellites yielded reliable estimates of variation across these species; per-locus polymorphism and allele-size distribution were not significantly different among or between any of the species sampled from mainland populations. Significantly lower levels of variation and the distribution of alleles in the two populations of the insular-endemic P. sejugis were consistent with the expectation of substantial founder effect and suggest a lack of recent gene flow between these populations. Phenetic analyses of genetic distances based on shared allele frequencies uniformly produced well-supported trees that were entirely concordant with the a priori corroborated relationships within the P. maniculatus species group; this result was not obtained with analyses of a genetic distance computed from differences in allele sizes. The microsatellite data do not support the hypothesis that P. sejugis should be considered conspecific with P. maniculatus but yield a strongly supported sister-group association between P. sejugis and P. keeni.     

Integrating coalescent and ecological niche modeling in comparative phylogeography

Understanding the factors that contribute to the formation of population genetic structure is a central goal of phylogeographic research, but achieving this goal can be complicated by the stochastic variance inherent to genetic processes. Statistical approaches to testing phylogeographic hypotheses accommodate this stochasticity by evaluating competing models of putative historical population structure, often by simulating null distributions of the expected variance. The effectiveness of these tests depends on the biological realism of the models. Information from the fossil record can aid in reconstructing the historical distributions of some taxa. However, for the majority of taxa, which lack sufficient fossils, paleodistributional modeling can provide valuable spatial-geographic data concerning ancestral distributions. Paleodistributional models are generated by projecting ecological niche models, which predict the current distribution of each species, onto a model of past climatic conditions. Here, we generate paleodistributional models describing the suitable habitat during the last glacial maximum for lineages from the mesic forests of the Pacific Northwest of North America, and use these models to generate alternative phylogeographic hypotheses. Coalescent simulations are then used to test these hypotheses to improve our understanding of the historical events that promoted the formation of population genetic structure in this ecosystem. Results from Pacific Northwest mesic forest organisms demonstrate the utility of these combined approaches. Paleodistribution models and population genetic structure are congruent across three amphibian lineages, suggesting that they have responded in a concerted manner to environmental change. Two other species, a willow and a water vole, despite being currently codistributed and having similar population genetic structure, were predicted by the paleodistributional model to have had markedly different distributions during the last glacial maximum. This suggests that congruent phylogeographic patterns can arise from incongruent ancestral distributions. Paleodistributional models introduce a much-needed spatial-geographic perspective to statistical phylogeography. In conjunction with coalescent models of population genetic structure, they have the potential to improve our understanding of the factors that promote population divergence and ultimately produce regional patterns of biodiversity.     

Blood group complexity: the Pm locus in Peromyscus maniculatus1

The Pm blood group system in Peromyscus maniculatus was originally described by Rasmussen as consisting of two co-dominant alleles, Pm^A and Pm^B, responsible for the antigenic characteristics A and B. This paper describes two additional antisera, anti-C and anti-X, which expand this system and show it to be serologically complex. Anti-X is comprised of antibodies directed against at least two antigenic characteristics X‘ and X“. Four and possibly six alleles are described by the four antisera. Different subspecies exhibit different spectra of alleles at the Pm locus. A segregation distortion is evidenced.     

Mitochondrial diversification of the Peromyscus mexicanus species group in Nuclear Central America: biogeographic and taxonomic implications

The mountain mice of the Peromyscus mexicanus group currently encompass six known species; however, the limits between species remain uncertain, with two considered monotypic and the other four having multiple associated subspecific names. Based on the most comprehensive sampling of the group throughout its distribution in Nuclear Central America, we used data of the mitochondrial cytochrome b gene to assess its genetic diversity, phylogeny, and main biogeographic and diversification patterns. Our mitochondrial phylogeny only partially reflects the current taxonomy of the group, in agreement with some of the taxonomically recognized species. Specifically, our phylogenetic results show that the group is highly structured, including four main clades with genetic distances ranging from 11 to 8.6%. A remarkable level of differentiation is found at a more local level, defined as 15 different lineages with high nucleotide and haplotype diversity (π = 0.068, h = 0.99), and with divergence and genetic distance values (p‐uncorrected = 9.9–2.4%; K2P = 10.8–3.0%) similar to values observed between species within Peromyscus. Accordingly, we propose that the reference name P. mexicanus is polyphyletic and should be restricted to the mountains of central Mexico west of the Isthmus of Tehuantepec. We suggest to limit the other five recognized specific names to equal number of lineages, as monophyletic, and to revalidate three junior synonyms: Peromyscus salvadorensis, P. nicaraguae and P. tropicalis. The Isthmus of Tehuantepec, the Motagua‐Polochic‐Jocotán fault system, the Maya Highlands and the Honduras Depression are examples of geographic features that are likely associated with the differentiation of main lineages. Some other lineages may represent candidate species, hence the need to review the taxonomic status of the entire P. mexicanus group.     

Analyzing niche stability and biogeography of Late Ordovician brachiopod species using ecological niche modeling

Gauging the potential impacts of environmental change on the geographic distributions of species is a central area of modern biogeographic analysis, often involving complex models of species–environment interactions. The geographic distribution of fossil species can also provide a framework to test the impact of environmental change on biogeography and ecological niches of species, yet few paleontological analyses have attacked this question in deep time. Herein we present a quantitative biogeographic analysis to examine the stability of ecological niches and geographic ranges of rhynchonelliform brachiopods during an interval of sea level change preserved in Upper Ordovician strata of the Cincinnati Arch.The intensive sampling, excellent preservation, and numerous prior paleoecological and sedimentological analyses within the tri-state region of Kentucky, Indiana, and Ohio provide a robust framework for detailed paleobiogeographic study. Quantitative biogeographic modeling methods incorporating GIS (Geographic Information Systems) are utilized in order to spatially analyze the geographic ranges of brachiopod species of the Corryville and Mt. Auburn Formations of the C3 (uppermost Maysvillian) depositional sequence.This study employs the ecological niche modeling program GARP (Genetic Algorithm using Rule-set Prediction) to predict the geographic distribution of eight brachiopod species during three time slices within the C3 sequence. This method estimates a species’ geographic range by modeling the ecological niche of the species based on a set of known species occurrence data coupled with environmental data inferred from sedimentologic proxies. Once environmental tolerances for a species are modeled; the species is predicted to occur wherever its preferred set of environmental conditions occurs within the study region.Distributional patterns were reconstructed for three time slices during the C3 sequence. Recovered range predictions were quantitatively analyzed for evidence of temporal range changes. Results indicate that average species range within the study area decreased and species tracked their preferred niche with high fidelity during the transition from the early to middle portions of the C3 depositional sequence, an interval of rapid relative sea level change. However, during the transition from the middle to late portions of the sequence, gradual shallowing within the basin and development of discontinuous habitat patches correlates with niche evolution of five of the eight species modeled. The average area a species occupied within the basin increased during this interval, but there is a mixed response including both increases and decreases in range size within the study group. In general, the species that exhibit niche evolution increased their geographic range size while those that continue to track their niche with high fidelity experience a decrease in geographic range size. During the latter half of the C3 sequence, previously continuous habitats become fragmented, thereby isolating individual populations and providing a mechanism for niche evolution. The rate of sea level change and the corresponding fragmentation of previously continuous habitats into isolated patches appear to be the primary controls on both mean geographic range size and relative degree of niche evolution.     

Pliocene–Pleistocene ecological niche evolution shapes the phylogeography of a Mediterranean plant group

Estimating species ability to adapt to environmental changes is crucial to understand their past and future response to climate change. The Mediterranean Basin has experienced remarkable climatic changes since the Miocene, which have greatly influenced the evolution of the Mediterranean flora. Here, we examine the evolutionary history and biogeographic patterns of two sedge sister species (Carex, Cyperaceae) restricted to the western Mediterranean Basin, but with Pliocene fossil record in central Europe. In particular, we estimated the evolution of climatic niches through time and its influence in lineage differentiation. We carried out a dated phylogenetic–phylogeographic study based on seven DNA regions (nDNA and ptDNA) and fingerprinting data (AFLPs), and modelled ecological niches and species distributions for the Pliocene, Pleistocene and present. Phylogenetic and divergence time analyses revealed that both species form a monophyletic lineage originated in the late Pliocene–early Pleistocene. We detected clear genetic differentiation between both species with distinct genetic clusters in disjunct areas, indicating the predominant role of geographic barriers limiting gene flow. We found a remarkable shift in the climatic requirements between Pliocene and extant populations, although the niche seems to have been relatively conserved since the Pleistocene split of both species. This study highlights how an integrative approach combining different data sources and analyses, including fossils, allows solid and robust inferences about the evolutionary history of a plant group since the Pliocene.     

Mutualistic mimicry enhances species diversification through spatial segregation and extension of the ecological niche space

Species richness varies among clades, yet the drivers of diversification creating this variation remain poorly understood. While abiotic factors likely drive some of the variation in species richness, ecological interactions may also contribute. Here, we examine one class of potential contributors to species richness variation that is particularly poorly understood: mutualistic interactions. We aim to elucidate large‐scale patterns of diversification mediated by mutualistic interactions using a spatially explicit population‐based model. We focus on mutualistic Müllerian mimicry between conspicuous toxic prey species, where convergence in color patterns emerges from predators' learning process. To investigate the effects of Müllerian mimicry on species diversification, we assume that some speciation events stem from shifts in ecological niches, and can also be associated with shift in mimetic color pattern. Through the emergence of spatial mosaics of mimetic color patterns, Müllerian mimicry constrains the geographical distribution of species and allows different species occupying similar ecological niches to exist simultaneously in different regions. Müllerian mimicry and the resulting spatial segregation of mimetic color patterns thus generate more balanced phylogenetic trees and increase overall species diversity. Our study sheds light on complex effects of Müllerian mimicry on ecological, spatial, and phylogenetic diversification.     

Collinearity in ecological niche modeling: Confusions and challenges

Ecological niche models are widely used in ecology and biogeography. Maxent is one of the most frequently used niche modeling tools, and many studies have aimed to optimize its performance. However, scholars have conflicting views on the treatment of predictor collinearity in Maxent modeling. Despite this lack of consensus, quantitative examinations of the effects of collinearity on Maxent modeling, especially in model transfer scenarios, are lacking. To address this knowledge gap, here we quantify the effects of collinearity under different scenarios of Maxent model training and projection. We separately examine the effects of predictor collinearity, collinearity shifts between training and testing data, and environmental novelty on model performance. We demonstrate that excluding highly correlated predictor variables does not significantly influence model performance. However, we find that collinearity shift and environmental novelty have significant negative effects on the performance of model transfer. We thus conclude that (a) Maxent is robust to predictor collinearity in model training; (b) the strategy of excluding highly correlated variables has little impact because Maxent accounts for redundant variables; and (c) collinearity shift and environmental novelty can negatively affect Maxent model transferability. We therefore recommend to quantify and report collinearity shift and environmental novelty to better infer model accuracy when models are spatially and/or temporally transferred.     

Adding ecology into phylogeography: ecological niche models and phylogeography in tandem reveals the demographic history of the subalpine warbler complex

Capsule: This study documents evidence of interglacial refugia during the Last Interglacial for birds in the Mediterranean region, and emphasizes the importance of the Last Interglacial on the geographic distribution and genetic structure of Mediterranean species.

Aims: We focused on the historical biogeography of the subalpine warbler complex: Subalpine Warbler Sylvia cantillans and Moltoni’s Warbler Sylvia subalpina; we tested if this Mediterranean bird complex shared a similar demographic fate as the present-day widespread species in the temperate zones of Europe, through the late Quaternary glacial-interglacial cycles.

Methods: An ecological niche model was developed to predict the geographic distribution of the subalpine warblers under the past (the Last Interglacial and the Last Glacial Maximum) and the present bioclimatic conditions. Additionally, Bayesian Skyline Plot analysis was used to assess effective population size changes over the history of the subalpine warbler complex.

Results: During the Last Glacial Maximum, the subalpine warblers almost reached their current distribution in the Mediterranean region; yet, unlike the widespread temperate bird species, they survived the Last Interglacial in allopatric refugia in the Mediterranean region.

Conclusion: A unique biogeographic pattern was revealed, indicating the importance of the Last Interglacial on current distributional patterns and demographic histories of common bird species in the Mediterranean region. This study suggests that Mediterranean biogeography is far more complex than previously assumed, and so deserves further study and more attention.     

Southward Pleistocene migration of Douglas-fir into Mexico: phylogeography, ecological niche modeling, and conservation of 'rear edge' populations

? Poleward Pleistocene plant migration has been an important process structuring modern temperate and boreal plant communities, but the contribution of equatorward migration remains poorly understood. Paleobotanical evidence suggests Miocene or Pleistocene origin for temperate 'sky island' plant taxa in Mexico. These 'rear edge' populations situated in a biodiversity hotspot may be an important reserve of genetic diversity in changing climates. ? We used mtDNA sequences, cpDNA sequences and chloroplast microsatellites to test hypotheses of Miocene vs Pleistocene colonization of temperate Douglas-fir in Mexico, explore geographic patterns of molecular variation in relation to Pleistocene climate history using ecological niche models, and assess the taxonomic and conservation implications. ? We found strong evidence for Pleistocene divergence of Douglas-fir in Mexico (958 thousand yr before present (ka) with the 90% highest posterior density interval ranging from 1.6 million yr before present (Ma) to 491 ka), consistent with the southward Pleistocene migration hypothesis. Genetic diversity was high and strongly partitioned among populations. Spatial patterns of molecular variation and ecological niche models suggest a complex late Pleistocene history involving periods of isolation and expansion along mountain corridors. ? These results highlight the importance of southward Pleistocene migration in establishing modern high-diversity plant communities and provide critical insights into proposals to conserve the unique biodiversity of Mexican Douglas-fir and associated taxa.     

Comparative phylogeography of three codistributed stomatopods: origins and timing of regional lineage diversification in the Coral Triangle

The Indonesian-Australian Archipelago is the center of the world's marine biodiversity. Although many biogeographers have suggested that this region is a "center of origin," criticism of this theory has focused on the absence of processes promoting lineage diversification in the center. In this study we compare patterns of phylogeographic structure and gene flow in three codistributed, ecologically similar Indo-West Pacific stomatopod (mantis shrimp) species. All three taxa show evidence for limited gene flow across the Maluku Sea with deep genetic breaks between populations from Papua and Northern Indonesia, suggesting that limited water transport across the Maluku Sea may limit larval dispersal and gene flow across this region. All three taxa also show moderate to strong genetic structure between populations from Northern and Southern Indonesia, indicating limited gene flow across the Flores and Java Seas. Despite the similarities in phylogeographic structure, results indicate varied ages of the genetic discontinuities, ranging from the middle Pleistocene to the Pliocene. Concordance of genetic structure across multiple taxa combined with temporal discordance suggests that regional genetic structures have arisen from the action of common physical processes operating over extended time periods. The presence in all three species of both intraspecific genetic structure as well as deeply divergent lineages that likely represent cryptic species suggests that these processes may promote lineage diversification within the Indonesian-Australian Archipelago, providing a potential mechanism for the center of origin. Efforts to conserve biodiversity in the Coral Triangle should work to preserve both existing biodiversity as well as the processes creating the biodiversity.     

Adding more ecology into species delimitation: ecological niche models and phylogeography help define cryptic species in the black salamander (Aneides flavipunctatus)

Being able to efficiently and accurately delimit species is one of the most basic and important aspects of systematics because species are the fundamental unit of analysis in biogeography, ecology, and conservation. We present a rationale and approach for combining ecological niche modeling, spatially explicit analyses of environmental data, and phylogenetics in species delimitation, and we use our methodology in an empirical example focusing on Aneides flavipunctatus, the black salamander (Caudata: Plethodontidae), in California. We assess the relationships between genetic, environmental, and geographic distance among populations. We use 11 climatic variables and point locality data from public databases to create ecological niche models. The suitability of potential contact zones between parapatric lineages is also assessed using the data from ecological niche modeling. Phylogenetic analyses of portions of the mitochondrial genome reveal morphologically cryptic mitochondrial lineages in this species. In addition, we find that patterns of genetic divergence are strongly associated with divergence in the ecological niche. Our work demonstrates the ease and utility of using spatial analyses of environmental data and phylogenetics in species delimitation, especially for groups displaying fine-scaled endemism and cryptic species.     

A new and widely distributed species of Pseudoarachniotus

Summary A new species ofPseudoarachniotus, P. hyalinosporus, is described and illustrated. A total of 54 isolates of this species was studied and their growth was compared on several media.Although certain differences appeared in colonial coloration and growth characteristics, all strains were similar in 2 fundamental characteristics, viz., type of ascospore and type of gametangial initials and their subsequent development. The initials in early stages are often so closely appressed one to another that they give the appearance of a single structure. Further development takes place by the elongation of the ascogonium which forms a loop around the male initial at right angles to the linear plane of that initial. The ascospores ofP. hyalinosporus are small, about 2.2 × 3.3µ, thin-walled, oval, hyaline to very pale yellow, and smooth, thereby differing fromPseudoarachniotus reticulatus which has globose, echinulate-reticulate ascospores, andP. roseus andP. citrinus which have large, thick-walled, oval to oblate ascospores. Furthermore, the colonies ofP. roseus are red, those ofP. citrinus are yellow, and inP. hyalinosporus colonies are white with pale yellow asciferous areas and usually with a great deal of green pigment both above and reverse. Of the 54 strains of this new species, 25 were isolated from rodent lungs, 9 from dung, 19 from soil and one from a tinea pedis condition.Pseudoarachniotus hyalinosporus was found in two collections from India and it appears to be widely distributed in California.Supported in part by Botany Department Research Grant # 1344, University of California, Los Angeles, California.A portion of this study was made possible through laboratory facilities provided by Dr.B. Samantarai, Ravenshaw College, Cuttuck, Orissa, India.     

An exhaustive analysis of heuristic methods for variable selection in ecological niche modeling and species distribution modeling

Ecological niche models and species distribution models are used in many fields of science. Despite their popularity, only recently have important aspects of the modeling process like model selection been developed. Choosing environmental variables with which to create these models is another critical part of the process, but methods currently in use are not consistent in their results and no comprehensive approach exists by which to perform this step. Here, we compared seven heuristic methods of variable selection against a novel approach that proposes to select best sets of variables by evaluating performance of models created with all combinations of variables and distinct parameter settings of the algorithm in concert. Our results were that—except for the jackknife method for one of the 12 species and fluctuation index for two of the 12 species—none of the heuristic methods for variable selection coincided with the exhaustive one. Performance decreased in models created using variables selected with heuristic methods and both underfitting and overfitting were detected when comparing their geographic projections with the ones of models created with variables selected with the exhaustive method. Using the exhaustive approach could be time consuming, so a two-step exercise may be necessary. However, using this method identifies adequate variable sets and parameter settings in concert that are associated with increased model performance.     

Sixty polymorphic microsatellite markers for the oldfield mouse developed in Peromyscus polionotus and Peromyscus maniculatus

We isolated and characterized 60 novel microsatellite markers from the closely related oldfield mouse (Peromyscus polionotus) and deer mouse (Peromyscus maniculatus) for studies of conservation, ecological, quantitative and population genetics. We assessed all 60 markers in a wild population of Peromyscus polionotus rhoadsi (N = 20) from central Florida and found an average of nine alleles per marker and an observed heterozygosity (H_O) of 0.66 (range = 0.00–1.00). These polymorphic markers contribute to the growing number of genomic resources for Peromyscus, an emerging model system for ecological and evolutionary research.