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Chromosomal speciation and adaptive radiation of mole rats in Asia Minor correlated with increased ecological stress.

The evolutionary forces causing chromosomal speciation and adaptation are still enigmatic. Here we tested the Israel evolutionary model of positive association of diploid chromosome number (2n) and genetic diversity with aridity stress in subterranean mole rats, on a 30-times-larger scale in Asia Minor. We analyzed both karyotype and allozyme diversity across Turkey, based on 37 allozymic loci in 20 localities of the Spalax leucodon and 4 localities of the Spalax ehrenbergi superspecies. We found extensive chromosomal speciation in S. leucodon (2n = 38, 40, 50, 54, 60, and 62) and in S. ehrenbergi (2n = 52, 56, and 58), presumably representing from 14 to > 20 additional biological species. Genetic diversity indices were low, but, like the chromosome number (2n), positively correlated with aridity stress, increasing centripetally from the periphery toward geologically young, arid, and climatically unpredictable central Anatolia. Nei's genetic distance D across all populations averaged 0.174 (range 0.002-0.422), supporting, combined with 2n and ecogeography, the biological species status of most tested populations. Chromosome evolution is the basis of speciation and adaptation in Spalax; it provides both postmating reproductive isolation, as well as higher levels of recombination with increased 2n. A mathematical model shows that a Robertsonian fission of a single metacentric considerably increases haplotype diversity. This haplotype diversity may contribute to population adaptation to climatic stress and ecological unpredictability in space and time. The increase in diversity corroborates the nichewidth genetic-variation hypothesis.

Animals↗

Sexual isolation in Drosophila melanogaster: a possible case of incipient speciation.

It is generally believed that Drosophila melanogaster has no closely related species with which it can produce the viable and fertile hybrids that are essential for the genetic analysis of speciation. Following the recent report of molecular differentiation between a Zimbabwe, Africa, population and two United States populations, we provide evidence that strong sexual isolation exists between the D. melanogaster population in Zimbabwe and populations of other continents. In the presence of males of their own kind, females from most isofemale lines of Zimbabwe would not mate with males from elsewhere; the reciprocal mating is also significantly reduced, but to a lesser degree. The genes for sexual behaviors are apparently polymorphic in Zimbabwe and postmating reproductive isolation between this and other populations has not yet evolved. Whole chromosome substitutions indicate significant genetic contributions to male mating success by both major autosomes, whereas the X chromosome effect is too weak to measure. In addition, the relative mating success between hybrid and pure line males supports the interpretation of strong female choice. These observations suggest that we are seeing the early stages of speciation in this group and that it is driven by sexual selection. The genetic and molecular tractability of D. melanogaster offers great promise for the detailed analysis of this apparent case of incipient speciation.

Animals↗

Sexual selection and speciation in field crickets.

Recent theoretical work has shown that sexual selection may cause speciation under a much wider range of conditions than previously supposed. There are, however, no empirical studies capable of simultaneously evaluating several key predictions that contrast this with other speciation models. We present data on male pulse rates and female phonotactic responses to pulse rates for the field cricket Gryllus texensis; pulse rate is the key feature distinguishing G. texensis from its cryptic sister species G. rubens. We show (i) genetic variation in male song and in female preference for song, (ii) a genetic correlation between the male trait and the female preference, and (iii) no character displacement in male song, female song recognition, female species-level song discrimination, or female song preference. Combined with previous work demonstrating a lack of hybrid inviability, these results suggest that divergent sexual selection may have caused speciation between these taxa.

Animals↗

Nickel speciation of residual oil fly ash and ambient particulate matter using X-ray absorption spectroscopy.

The chemical speciation of Ni in fly ash produced from approximately 0.85 wt % S residual (no. 6 fuel) oils in laboratory (7 kW)- and utility (400 MW)-scale combustion systems was investigated using X-ray absorption fine structure (XAFS) spectroscopy, X-ray diffraction (XRD), and acetate extraction [1 M NaOAc-0.5 M HOAc (pH 5) at 25 degrees C]-anodic stripping voltammetry (ASV). XAFS was also used to determine the Ni speciation of ambient particulate matter (PM) sampled near the 400-MW system. Based on XAFS analyses of bulk fly ash and their corresponding acetate extraction residue, it is estimated that > 99% of the total Ni (0.38 wt %) in the experimentally produced fly ash occurs as NiSO4.xH2O, whereas > 95% of the total Ni (1.70 and 2.25 wt %) in two fly ash samples from the 400-MW system occurs as NiSO4.xH2O and Ni-bearing spinel, possibly NiFe2O4. Spinel was also detected using XRD. Acetate extracts most of the NiSO4.xH2O and concentrates insoluble NiFe2O4 in extraction residue. Similar to fly ash, ambient PM contains NiSO4.xH2O and NiFe2O4; however, the proportion of NiSO4.xH2O relative to NiFe2O4 is much greater in the PM. Results from this and previous investigations indicate that residual oil ash produced in the 7-kW combustion system lack insoluble Ni (e.g., NiFe2O4) but are enriched in soluble NiSO4.xH2O relative to fly ash from utility-scale systems. This difference in Ni speciation is most likely related to the lack of additive [e.g., Mg(OH)2] injection and residence time in the 7-kW combustion system.

Absorptiometry, Photon↗

Development and evaluation of an impactor for a PM2.5 speciation sampler.

A conventional impactor for a particle speciation sampler was developed and validated through laboratory and field tests. The speciation sampler consists of the following components: a PM2.5 conventional impactor that removes particles larger than 2.5 microns, an all-glass, coated honeycomb diffusion denuder, and a 47-mm filter pack. The speciation sampler can operate at two different sampling rates: 10 and 16.7 L/min. An experimental characterization of the impactor's performance was conducted. The impactor's collection efficiency was examined as a function of critical design parameters such as Reynolds number, the distance from the nozzle exit to the impaction plate, and the impaction substrate coating method. The bounce of particles larger than the cut point was successfully minimized by using a greased surface as the impaction substrate. Additionally, a series of field intercomparison experiments were conducted at both 10 and 16.7 L/min airflow. PM2.5 mass and SO4(2-) concentrations were measured and compared with the Federal Reference Method (FRM) and found to be in good agreement. Results of the laboratory chamber tests also indicated that the impactor's performance was in good agreement with the FRM.

Air Pollution↗

Nickel and sulfur speciation of residual oil fly ashes from two electric utility steam-generating units.

Representative duplicate fly ash samples were obtained from the stacks of 400- and 385-MW utility boilers (Unit A and Unit B, respectively) using a modified U.S. Environmental Protection Agency (EPA) Method 17 sampling train assembly as they burned 0.9 and 0.3 wt % S residual (No. 6 fuel) oils, respectively, during routine power plant operations. Residual oil fly ash (ROFA) samples were analyzed for Ni concentrations and speciation using inductively coupled plasma-atomic emission spectroscopy, X-ray absorption fine structure (XAFS) spectroscopy, and X-ray diffraction (XRD). ROFA deionized H2O extraction residues were also analyzed for Ni speciation using XAFS and XRD. Total Ni concentrations in the ROFAs were similar, ranging from 1.3-1.5 wt%; however, stack gas Ni concentrations in the Unit A were 0.990 microg/Nm3 compared with 0.620 microg/Nm3 for Unit B because of the greater residual oil feed rates employed at Unit A to attain higher 400-MW load conditions with a lower heating value oil. Ni speciation analysis results indicated that ROFAs from Unit A contain approximately 3 wt % NiSO4 x xH2O (where x is assumed to be 6 for calculation purposes) and appoximately 4.5 wt% of a Ni-containing spinel compound, similar in composition to (Mg,Ni)(Al,Fe)2O4. ROFAs from Unit B contain on average 2 wt% NiSO4 x 6 H20 and 1.1 wt% NiO. XAFS and XRD analyses did not detect any nickel sulfide compounds, including carcinogenic nickel subsulfide (Ni3S2) (XAFS detection limit is 5% of the total Ni concentration). In addition, XAFS measurements indicated that inorganic sulfate and organic thiophene species accounted for > 97% of the total S in the ROFAs. Unit A ROFAs contained much lower thiophene proportions because cyclone-separated ROFA reinjection is employed on this unit to collect and reburn the larger carbonaceous particles.

Air Pollution↗

A simple algorithm to infer gene duplication and speciation events on a gene tree.

MOTIVATION: When analyzing protein sequences using sequence similarity searches, orthologous sequences (that diverged by speciation) are more reliable predictors of a new protein's function than paralogous sequences (that diverged by gene duplication), because duplication enables functional diversification. The utility of phylogenetic information in high-throughput genome annotation ('phylogenomics') is widely recognized, but existing approaches are either manual or indirect (e.g. not based on phylogenetic trees). Our goal is to automate phylogenomics using explicit phylogenetic inference. A necessary component is an algorithm to infer speciation and duplication events in a given gene tree. RESULTS: We give an algorithm to infer speciation and duplication events on a gene tree by comparison to a trusted species tree. This algorithm has a worst-case running time of O(n(2)) which is inferior to two previous algorithms that are approximately O(n) for a gene tree of sequences. However, our algorithm is extremely simple, and its asymptotic worst case behavior is only realized on pathological data sets. We show empirically, using 1750 gene trees constructed from the Pfam protein family database, that it appears to be a practical (and often superior) algorithm for analyzing real gene trees. AVAILABILITY: http://www.genetics.wustl.edu/eddy/forester.

Algorithms↗

Comparison of comprehensive two-dimensional gas chromatography coupled with sulfur-chemiluminescence detector to standard methods for speciation of sulfur-containing compounds in middle distillates.

The monitoring of total sulfur content and speciation of individual sulfur-containing compounds in middle distillates is required for efficient catalyst selection and for a better understanding of the kinetics of the reactions involved in hydrotreament processes. Owing to higher resolution power and enhanced sensitivity, comprehensive two-dimensional gas chromatography (GCxGC) hyphenated to sulfur chemiluminescence detection (SCD) has recently evolved as a powerful tool for improving characterization and identification of sulfur compounds. The aim of this paper is to compare quantitatively GCxGC-SCD and various other methods commonly employed in the petroleum industry, such as X-ray fluorescence, conventional GC-SCD, and high-resolution mass spectrometry, for total sulfur content determination and speciation analysis. Different samples of middle distillates have been analyzed to demonstrate the high potential and important advantages of GCxGC-SCD for innovative and quantitative analysis of sulfur-containing compounds. More accurate and detailed results for benzothiophenes and dibenzothiophenes are presented, showing that GCxGC-SCD should become, in the future, an essential tool for sulfur speciation analysis.

Journal Article↗

Mitochondrial and allozyme genetics of incipient speciation in a landlocked population of Galaxias truttaceus (Pisces: Galaxiidae).

Galaxias truttaceus is found in coastal rivers and streams in south-eastern Australia. It spawns at the head of estuaries in autumn and the larvae spend 3 months of winter at sea before returning to fresh water. In Tasmania there are landlocked populations of G. truttaceus in a cluster of geologically young lakes on the recently glaciated Central Plateau. These populations have no marine larval stage and spawn in the lakes in spring. Speciation due to land locking is thought to be a frequent occurrence within Galaxias. To investigate the nature of the speciation event which may be occurring within lake populations of G. truttaceus we studied the mitochondrial DNA (mtDNA) and allozyme diversity of both lake and stream populations. Using the presence or absence of restriction sites recognized by 13 six-base restriction endonucleases, we found 58 mtDNA haplotypes among 150 fish collected from 13 Tasmanian and one south-east Australian mainland stream populations. The most parsimonious network relating the haplotypes by site loss or gain was starlike in shape. We argue that this arrangement is best explained by selection upon slightly beneficial mutations within the mitochondrial genome. Gene diversity analysis under Wright's island model showed that the populations in each drainage were not genetically subdivided. Only two of these stream haplotypes were found among the 66 fish analyzed from four lake populations. Despite the extreme lack of mtDNA diversity in lake populations, the observed nuclear DNA heterozygosity of 40 lake fish (0.10355) was only slightly less than that of 82 stream fish (0.11635). In the short time (3000-7000 years) that the lake fish have been landlocked, random genetic drift in a finite, stable-sized population was probably not responsible for the lack of mtDNA diversity in the lake populations. We infer the lake populations have probably experienced at least one, severe, but transitory bottleneck possibly induced by natural selection for life-history characters essential for survival in the lacustrine habitat. If speciation is occurring in the landlocked populations of G. truttaceus, then it may be driven by genetic transilience.

Animals↗

Nuclear gene genealogies reveal historical, demographic and selective factors associated with speciation in field crickets.

Population genetics theory predicts that genetic drift should eliminate shared polymorphism, leading to monophyly or exclusivity of populations, when the elapsed time between lineage-splitting events is large relative to effective population size. We examined patterns of nucleotide variation in introns at four nuclear loci to relate processes affecting the history of genes to patterns of divergence among natural populations and species. Ancestral polymorphisms were shared among three recognized species, Gryllus firmus, G. pennsylvanicus, and G. ovisopis, and genealogical patterns suggest that successive speciation events occurred recently and rapidly relative to effective population size. High levels of shared polymorphism among these morphologically, behaviorally, and ecologically distinct species indicate that only a small fraction of the genome needs to become differentiated for speciation to occur. Among the four nuclear gene loci there was a 10-fold range in nucleotide diversity, and patterns of polymorphism and divergence suggest that natural selection has acted to maintain or eliminate variation at some loci. While nuclear gene genealogies may have limited applications in phylogeography or other approaches dependent on population monophyly, they provide important insights into the historical, demographic, and selective forces that shape speciation.

Animals↗

The theory of speciation via the founder principle.

The founder principle has been used to explain many instances of rapid speciation. Advances from theoretical population genetics are incorporated into MAYR's original founder-effect genetic-revolution model to yield a newer model called the genetic transilience. The basic theoretical edifice lies upon the fact that founder event can sometimes lead to an accumulation of inbreeding and an induction of gametic disequilibrium. This, in turn, causes alleles to be selected more for their homozygous fitness effects and for their effects on a more stable genetic background. Selection occurring in multi-locus systems controlling integrated developmental, physiological, behavioral, etc, traits is particularly sensitive to these founder effects. If sufficient genetic variability exists in the founder population, such multilocus genetic systems can respond to drift and the altered selective forces by undergoing a rapid shift to a new adaptive peak known as the genetic transilience. A genetic transilience is, therefore, most likely to occur when the founder event causes a rapid accumulation of inbreeding without a severe reduction in genetic variability. The implications of this model are then examined for three aspects of the founder-effect genetic-transilience model: the attributes of the ancestral population, the nature of the sampling process used to generate the founders and the attributes of the founder population. The model is used to explain several features of the evolution of the Hawaiian Drosophila, and experimental designs are outlined to test the major predictions of the theory. Hence, this theory of speciation can be tested in the laboratory, using systems and techniques that already exist--a rare attribute of most models of speciation.

Animals↗

Using SINEs to probe ancient explosive speciation: "hidden" radiation of African cichlids?

Cichlid fishes of the east African Great Lakes represent a paradigm of adaptive radiation. We conducted a phylogenetic analysis of cichlids including pan-African and west African species by using insertion patterns of short interspersed elements (SINEs) at orthologous loci. The monophyly of the east African cichlids was consistently supported by seven independent insertions of SINE sequences that are uniquely shared by these species. In addition, data from four other loci indicated that the genera Tilapia (pan-African) and Steatocranus (west African) are the closest relatives to east African cichlids. However, relationships among Tilapia, Steatocranus, and the east African clade were ambiguous because of incongruencies among topologies suggested by insertion patterns of SINEs at six other loci. One plausible explanation for this phenomenon is incomplete lineage sorting of alleles containing or missing a SINE insertion at these loci during ancestral speciation. Such incomplete sorting may have taken place earlier than 14 MYA, followed by random and stochastic fixation of the alleles in subsequent lineages. These observations prompted us to consider the possibility that cichlid speciation occurred at an accelerated rate during this period when the African Great Lakes did not exist. The SINE method could be useful for detecting ancient exclusive speciation events that tend to remain hidden during conventional sequence analyses because of accumulated point mutations.

Base Sequence↗

Host-associated speciation in a coral-inhabiting barnacle.

Host specificity of symbionts is considered an important factor associated with sympatric speciation. Here, we examine host specificity and the degree of host-associated speciation in the barnacle Savignium milleporum, an obligate symbiont of the hydrocoral Millepora (the "fire coral"). Little morphological variability was revealed between barnacles collected from two morphs of the hydrocoral Millepora dichotoma (encrusting or branching) or from its congener Millepora platyphylla, but a molecular analysis revealed an unexpected pattern of DNA sequence divergence. The sequences of the 12S mitochondrial rDNA were nearly identical within each of the three barnacle populations (average sequence divergence <1%), and the sequences obtained for barnacles collected from the two different morphs of M. dichotoma differed considerably (ca. 9% average sequence divergence). However, S. milleporum collected from M. platyphylla were nearly identical to the barnacles from the branching M. dichotoma (<0.5% average sequence divergence). The pattern of speciation demonstrated by Savignium barnacles indicates the gradual colonization of similar hosts (i.e., sequential evolution), rather then "casual" colonization, as indicated for other systems. If this is indeed so, then symbiont phylogeny should roughly correlate with host phylogeny. Additionally, the data support the "rendezvous host" hypothesis, which invokes the opportunity of both sexes to meet as a major component for which selection favors the costly habit of host specificity.

Animals↗

Sex-related genes, directional sexual selection, and speciation.

Reproductive isolation and speciation can result from the establishment of either premating or postmating barriers that restrict gene flow between populations. Recent studies of speciation have been dominated by a molecular approach to dissect the genetic basis of hybrid male sterility, a specific form of postmating reproductive isolation. However, relatively little attention has been paid to the evolution of genes involved in premating isolation and genes generally involved in other sex-related functions (e.g., mating behavior, fertilization, spermatogenesis, sex determination). We have assembled DNA sequences from 51 nuclear genes and classified them based on their functional characteristics. The proportion of nonsynonymous to synonymous nucleotide substitutions were compared between Drosophila melanogaster, Drosophila simulans, and Drosophila pseudoobscura, as well as between Caenorhabditis elegans and Caenorhabditis briggsae. We found a high ratio of nonsynonymous to synonymous substitutions for sex-related genes (i.e., genes involved in mating behavior, fertilization, spermatogenesis, or sex determination). The results suggest that directional sexual selection has shaped the evolution of sex-related genes and that these changes have more likely occurred during the early stages of speciation. It is possible that directional selection becomes relaxed after reproductive isolation has been completed between more distantly related species (e.g., D. melanogaster and D. pseudoobscura). However, a saturation in the number of nucleotide substitutions since the time of species separation may mask any sign of directional selection between more distantly related species.

Animals↗

Effective population size may limit the power of laboratory experiments to demonstrate sympatric and parapatric speciation.

Laboratory experiments designed to elucidate the mechanisms of sympatric and parapatric speciation may have been handicapped by too small population sizes, although this possibility has seldom been discussed. In this paper we review the published records of sympatric and parapatric speciation experiments to test the relative importance of selection intensity applied, duration of experiment and effective population size. Our results show that among these factors only effective population size has had a general effect on the generation of assortative mating. Reduced interbreeding is less likely to develop in small populations where the selection process often seems to have been opposed by inbreeding depression or loss of genetic variation. This study demonstrates that the experimental evidence frequently used as an argument against sympatric and parapatric speciation models is not as strong as previously believed.

Animals↗

Speciation as a positive feedback loop between postzygotic and prezygotic barriers to gene flow.

Speciation is intimately associated with the evolution of sex-and-reproduction-related traits, including those affecting hybrid incompatibility (postzygotic isolation) and species recognition (prezygotic isolation). Genes controlling such traits are not randomly distributed in the genome but are particularly abundant on the sex chromosomes. However, the evolutionary consequences of the sex linkage of genes involved in speciation have been little explored. Here, we present simulations of a continent-island diploid model that examines the effects of reduced recombination using both autosomal and sex-linked inheritance. We show first that linkage between genes affecting postzygotic and prezygotic isolation leads to a positive feedback loop in which both are strengthened. As species recognition evolves, genes causing hybrid incompatibility will hitchhike along with those improving premating isolation, leading to stronger hybrid incompatibility and thus increased pressure for further preference divergence. Second, we show that this loop effect is generally enhanced by sex linkage, because recombination is eliminated in the heterogametic sex, leading to tighter effective linkage between the two classes of genes and because natural selection is more efficient at sex-linked loci, as recessive alleles are not masked by dominance in the heterogametic sex. Accordingly, hitchhiking can be important in promoting speciation and can also lead to increased postzygotic isolation through adaptive evolution.

Adaptation, Biological↗

Habitat preference and the marine-speciation paradox.

Marine organisms challenge the classical theories of local adaptation and speciation because their planktonic larvae have the potential to maintain high gene flow. The marine-speciation paradox is illustrated by contact zones between incipient species that are so large that allopatric divergence seems unlikely. For this reason any mechanism preventing sympatric larvae of two incipient species from coexisting in the same habitats can be a powerful promoter of speciation. The contact zone between two hybridizing taxa of mussel, Mytilus edulis and M. galloprovincialis, in Europe provides an excellent example. Although the zone itself extends over thousands of kilometres, the opportunities for interbreeding are considerably reduced by the small-scale mosaic structure of the zone, where local patches of each taxon alternate at scales of kilometres or less, in response to locally variable ecological factors. Habitat choice by settling larvae would be a less costly mechanism than post-settlement selection to maintain such a mosaic structure. Unfortunately the role of selective settlement has remained hypothetical because larvae could not be scored by classical genetic markers. PCR markers allowed us to study larvae and settlement in ecologically contrasting sites within the zone. We show that only a subset of the genotypes present in the plankton settle in some sites, and that the adults on these sites show the same genetic bias. Genetically based variation in pre-settlement processes therefore accounts for the ecological segregation observed, though it is not the only factor involved in limiting successful interbreeding. The present dataset also supports previous reports of partial spawning asynchrony.

Animals↗

Floral symmetry affects speciation rates in angiosperms.

Despite much recent activity in the field of pollination biology, the extent to which animal pollinators drive the formation of new angiosperm species remains unresolved. One problem has been identifying floral adaptations that promote reproductive isolation. The evolution of a bilaterally symmetrical corolla restricts the direction of approach and movement of pollinators on and between flowers. Restricting pollinators to approaching a flower from a single direction facilitates specific placement of pollen on the pollinator. When coupled with pollinator constancy, precise pollen placement can increase the probability that pollen grains reach a compatible stigma. This has the potential to generate reproductive isolation between species, because mutations that cause changes in the placement of pollen on the pollinator may decrease gene flow between incipient species. I predict that animal-pollinated lineages that possess bilaterally symmetrical flowers should have higher speciation rates than lineages possessing radially symmetrical flowers. Using sister-group comparisons I demonstrate that bilaterally symmetric lineages tend to be more species rich than their radially symmetrical sister lineages. This study supports an important role for pollinator-mediated speciation and demonstrates that floral morphology plays a key role in angiosperm speciation.

Biodiversity↗