Evolutionary impacts of introgressive hybridization in a rapidly evolving group of jumping spiders (F. Salticidae, Habronattus americanus group).

Introgressive hybridization can be a powerful force impacting patterns of evolution at multiple taxonomic levels. We aimed to understand how introgression has affected speciation and diversification within a species complex of jumping spiders. The Habronattus americanus subgroup is a recently radiating group of jumping spiders, with species now in contact after hypothesized periods of isolation during glaciation cycles of the Pleistocene. Effects of introgression on genomes and morphology were investigated using phylogenomic and clustering methods using RADseq, ultraconserved elements (UCEs), and morphological data. We characterized 14 unique species/morphs using non-metric multidimensional scaling of morphological data, a majority of which were not recovered as monophyletic in our phylogenomic analyses. Morphological clusters and genetic lineages are highly incongruent, such that geographic region was a greater predictor of phylogenetic relatedness and genomic similarity than species or morph identity. STRUCTURE analyses support this pattern, revealing clusters corresponding to larger geographic regions. A history of rapid radiation in combination with frequent introgression seems to have mostly homogenized the genomes of species in this system, while selective forces maintain distinct male morphologies. GEMMA analyses support this idea by identifying SNPs correlated with distinct male morphologies. Overall, we have uncovered a system at odds with a typical bifurcating evolutionary model, instead supporting one where closely related species evolve together connected through multiple introgression events, creating a reticulate evolutionary history.

From the mountains to the coast and back again: Ancient biogeography in a radiation of short-range endemic harvestmen from California.

The harvestmen genus Calicina is represented by 25 short-range endemic species occurring in the western Sierra Nevada, Transverse and Coast Ranges of California. Our principal aim was to reconstruct the temporal and spatial biogeographic history of this arachnid lineage. We inferred a time-calibrated species tree for 21 of 25 described Calicina species using multiple genes and multilocus coalescent-based methods. This species tree was used as a framework for algorithmic biogeographic and divergence time analyses, and a phylogenetic canonical correlation analysis (CCA) was used to examine the relationship between morphological evolution and environmental variables. Species tree and biogeographic analyses indicate that high-elevation Sierran taxa are early-diverging in Calicina, with subsequent biogeographic "criss-crossing" of lineages from the Sierra Nevada to the Coast Ranges, back to the Sierra Nevada, then back to Coast Ranges. In both the Sierra Nevada and Coast Ranges, distantly-related parapatric lineages essentially never occur in sympatry. CCA reveals that in both the Coast Ranges and the Sierra Nevada, distant phylogenetic relatives evolve convergent morphologies. Our evidence shows that Calicina is clearly dispersal-limited, with an ancient biogeographic history that provides unique insight into the complex geologic evolution of California since the mid-Paleogene.

The power and perils of 'molecular taxonomy': a case study of eyeless and endangered Cicurina (Araneae: Dictynidae) from Texas caves.

Rapid development in karst-rich regions of the US state of Texas has prompted the listing of four Cicurina species (Araneae, Dictynidae) as US Federally Endangered. A major constraint in the management of these taxa is the extreme rarity of adult specimens, which are required for accurate species identification. We report a first attempt at using mitochondrial DNA (mtDNA) sequences to accurately identify immature Cicurina specimens. This identification is founded on a phylogenetic framework that is anchored by identified adult and/or topotypic specimens. Analysis of approximately 1 kb of cytochrome oxidase subunit I (CO1) mtDNA data for over 100 samples results in a phylogenetic tree that includes a large number of distinctive, easily recognizable, tip clades. These tip clades almost always correspond to a priori species hypotheses, and show nonoverlapping patterns of sequence divergence, making it possible to place species names on a number of immature specimens. Three cases of inconsistency between recovered tip clades and a priori species hypotheses suggest possible introgression between cave-dwelling Cicurina, or alternatively, species synonymy. Although species determination is not possible in these instances, the inconsistencies point to areas of taxonomic ambiguity that require further study. Our molecular phylogenetic sample is largest for the Federally Endangered C. madla. These data suggest that C. madla occurs in more than twice the number of caves as previously reported, and indicate the possible synonymy of C. madla with C. vespera, which is also Federally Endangered. Network analyses reveal considerable genetic divergence and structuring across caves in this species. Although the use of DNA sequences to identify previously 'unidentifiable' specimens illustrates the potential power of molecular data in taxonomy, many other aspects of the same dataset speak to the necessity of a balanced taxonomic approach.

Genealogical exclusivity in geographically proximate populations of Hypochilus thorelli Marx (Araneae, Hypochilidae) on the Cumberland Plateau of North America.

The issue of sampling sufficiency is too infrequently explored in phylogeographical analysis, despite both theoretical work and analytical methods that stress the importance of sampling effort. Regarding the evolutionary pattern of reciprocal monophyly, both the probability of recovering this pattern and the possible inferences derived from this pattern, are highly contingent upon the density and geographical scale of sampling. Here, we present an empirical example that relates directly to this issue. We analyse genetic structure in the southern Appalachian spider Hypochilus thorelli, using an average sample of 5 mitochondrial DNA (mtDNA) sequences per location for 19 locations. All sampled sites are reciprocally monophyletic for mtDNA variation, even when separated by geographical distances as small as 5 km. For populations separated by greater geographical distances of 20-50 km, mtDNA sequences are not only exclusive, but are also highly divergent (uncorrected p-distances exceeding 5%). Although these extreme genealogical patterns are most seemingly consistent with a complete isolation model, both a coalescent method and nested cladistic analysis suggest that other restricted, but nonzero, gene flow models may also apply. Hypochilus thorelli appears to have maintained morphological cohesion despite this limited female-based gene flow, suggesting a pattern of stasis similar to that observed at higher taxonomic levels in Hypochilus.

Phylogenetic utility and evidence for multiple copies of elongation factor-1alpha in the spider genus Habronattus (Araneae: Salticidae).

In the continuing quest for informative genes for use in molecular systematics, the protein-coding gene Elongation factor-1alpha (EF-1alpha) has rapidly become one of the most prevalent "single-copy" nuclear genes utilized, particularly in arthropods. This paper explores the molecular evolutionary dynamics and phylogenetic utility of EF-1alpha in the salticid spider genus Habronattus. As has been reported for other arthropod lineages, our studies indicate that multiple (two) copies of EF-1alpha exist in Habronattus. These copies differ in intron structure and thus in size, making it possible to easily separate PCR amplification products. We present data for an intronless EF-1alpha copy for three Habronattus species. The presence of nonsense mutations and generally elevated rates of amino acid change suggest that this copy is evolving under relaxed functional constraints in Habronattus. A larger taxon sample (50 species plus outgroups) is presented for an EF-1alpha copy that includes both intron and exon regions. Characteristics of both regions suggest that this is a functional, orthologous copy in the species sampled. Maximum-likelihood relative-rate comparisons show that exon third codon sites are evolving more than 100 times as fast as second codon sites in these sequences and that intron sites are evolving about twice as fast as exon third sites. In combination, the EF-1alpha data provide robust, species-level phylogenetic signal that is largely congruent with morphologically well supported areas of Habronattus phylogeny. The recovery of some novel clades, and the unexpected fragmentation of others, suggests areas requiring further phylogenetic attention.

Deep molecular divergence in the absence of morphological and ecological change in the Californian coastal dune endemic trapdoor spider Aptostichus simus.

Aptostichus simus is a trapdoor spider endemic to the coastal dunes of central and southern California and, on morphological grounds, is recognized as a single species. Mitochondrial DNA 16S rRNA sequences demonstrate that most populations are fixed for the same haplotype and that the population haplotypes from San Diego County, Los Angeles County, Santa Rosa Island, and Monterey County are extremely divergent (6-12%), with estimated separation times ranging from 2 to 6 million years. A statistical cluster analysis of morphological features demonstrates that this genetic divergence is not reflected in anatomical features that might signify ecological differentiation among these lineages. The species status of these divergent populations of A. simus depends upon the species concept utilized. If a time-limited genealogical perspective is employed, A. simus would be separated at the base into two genetically distinct species. This study suggests that species concepts based on morphological distinctiveness, in spider groups with limited dispersal capabilities, probably underestimate true evolutionary diversity.

A combined molecular approach to phylogeny of the jumping spider subfamily dendryphantinae (araneae: salticidae).

Four gene regions were sequenced for 30 species of jumping spiders, most from the subfamily Dendryphantinae, to investigate their molecular phylogeny and evolution. These are three regions from the mitochondria (ca. 560 bp of 16S plus adjacent tRNA, 1047 bp of cytochrome oxidase 1 (CO1), and 414 bp of NADH1 (ND1) and one region from the nuclear genome (ca. 750 bp of 28S). Parsimony and likelihood analyses of these gene regions separately and together support the monophyly of the dendryphantines as delimited previously by morphological characters. A group of elongate-bodied genera are placed as basal among the dendryphantines, and previously proposed relationships of Poultonella, Paraphidippus, and Sassacus vitis are confirmed. Comparison of overall rates of molecular evolution indicates striking differences across the gene regions, with highest divergence in ND1, CO1, 16S, and 28S in decreasing order. All four regions are characterized by both within- and among-site rate variation. Phylogenetic results from CO1 conflict conspicuously with phylogenetic results from the other genes and morphological data. Attempts to account for potential sources of this conflict (e.g., accommodating biased base composition, high homoplasy, within- and among-site rate variation, etc.) are largely unsuccessful.

Molecular insights into species phylogeny, biogeography, and morphological stasis in the ancient spider genus Hypochilus (Araneae: Hypochilidae).

The spider genus Hypochilus is currently restricted to cool, moist microhabitats in three widely separated montane regions of North America, providing an opportunity to study both deep (i.e., continental level) and shallow (within montane region) biogeographic history. Members of the genus also retain many plesiomorphic morphological characteristics, inviting the study of comparative rates of morphological evolution. In this paper, Hypochilus phylogeny and associated evolutionary problems are addressed using both new molecular (28S nDNA and CO1 mtDNA) and previously published (K. M. Catley, 1994, Am. Mus. Nov. 3088, 1-27) morphological data. Although the molecular data provide limited resolution of root placement within Hypochilus, most analyses are at least consistent with morphology-supported montane relationships of (Rockies (California, Appalachian)). The monophyly of Hypochilus species distributed in the California mountains is ambiguous, with several analyses indicating that this fauna may be paraphyletic with respect to a monophyletic Appalachian lineage. The montane regions differ in consistent ways in depths of both mitochondrial and nuclear phylogenetic divergence. Molecular clock analyses, in combination with arthropod-based mtDNA rate calibrations, suggest that the regional faunas are of different ages and that speciation in all faunas likely occurred prior to the Pleistocene. Limited intraspecific sampling reveals extraordinarily high levels of mtDNA cytochrome oxidase sequence divergence. These extreme divergences are most consistent with morphological stasis at the species level, despite preliminary evidence that Hypochilus taxa are characterized by fragmented population structures.

The origin of the word Stent.

In 1856, the English dentist Charles Stent developed a thermoplastic-like material for taking impressions of toothless mouths. This "Stent mass" was later used as a device or mould for keeping a skin graft in place; it was also used to provide support for anastomosis. A hundred years after the inventor's death in 1885, the word stent has been adopted all over the world in interventional radiology but today it is understood to mean percutaneous tubular structures that induce or maintain lumen patency. The true origin of the word stent is not found in many dictionaries. In most references, the wrong dentist is given credit for the discovery. Dictionaries also refer to the obsolete English and Scottish words stent and stint which mean, among other things, "to extend". The true origin of the word is therefore somewhat unclear.

Molecular phylogenetics at the population/species interface in cave spiders of the southern Appalachians (Araneae:Nesticidae:Nesticus).

This paper focuses on the relationship between population genetic structure and speciation mechanisms in a monophyletic species group of Appalachian cave spiders (Nesticus). Using mtDNA sequence data gathered from 256 individuals, I analyzed patterns of genetic variation within and between populations for three pairs of closely related sister species. Each sister-pair comparison involves taxa with differing distributional and ecological attributes; if these ecological attributes are reflected in basic demographic differences, then speciation might proceed differently across these sister taxa comparisons. Both frequency-based and gene tree analyses reveal that the genetic structure of the Nesticus species studied is characterized by similar and essentially complete population subdivision, regardless of differences in general ecology. These findings contrast with results of prior genetic studies of cave-dwelling arthropods that have typically revealed variation in population structure corresponding to differences in general ecology. Species fragmentation through both extrinsic and intrinsic evolutionary forces has resulted in discrete, perhaps independent, populations within morphologically defined species. Large sequence divergence values observed between populations suggest that this independence may extend well into the past. These patterns of mtDNA genealogical structure and divergence imply that species as morphological lineages are currently more inclusive than basal evolutionary or phylogenetic units, a suggestion that has important implications for the study of speciation mechanisms.

The effect of heterochromatin on synapsis of the sex chromosomes of Peromyscus (Rodentia, Cricetidae).

The pairing behavior of the sex chromosomes in male and female individuals representing seven species of Peromyscus was analyzed by electron microscopy of silver-stained zygotene and pachytene configurations. Six species possess submetacentric or metacentric X chromosomes with heterochromatic short arms. Sex-chromosome pairing in these species is initiated during early pachynema at an interstitial position on the X and Y axes. Homologous synapsis then progresses in a unidirectional fashion towards the telomeres of the X short arm and the corresponding arm of the heterochromatic Y chromosome. The distinctive pattern of synaptic initiation allowed a late-synapsing bivalent in fetal oocytes to be tentatively identified as that of the X chromosomes. In contrast to the other species, Peromyscus megalops possesses an acrocentric X chromosome and a very small Y chromosome. Sex-chromosome pairing in this species is initiated at the proximal telomeric region during late zygonema, and then proceeds interstitially towards the distal end of the Y chromosome. These observations suggest that the presence of X short-arm heterochromatin and corresponding Y heterochromatin interferes with late-zygotene alignment of the pairing initiation sites, thereby delaying XY synaptic initiation until early pachynema. The pairing initiation sites are conserved in the vicinity of the X and Y centromeres in Peromyscus, and consequently the addition of heterochromatin during sex-chromosome evolution essentially displaces these sites to an interstitial position.

Developing solutions for dental X-ray processors.

Six developing solutions suitable for automatic X-ray processors with high developer temperature have been tested during a period of 14 days without replenishment of the chemicals. The results are presented as blackening of dental test films with an image of an aluminium step wedge (penetrometer). Both speed and contrast could be measured in this manner. Freshly mixed solutions showed marked differences in processing of films where the developers Dürr, Readymatic (Kodak) and Tetenal (Mediquipe) were best. After 14 days in the processor, the developing effect was poor for most of the tested liquids, due to atmospheric oxidation. The best images, with the least radiation, will be obtained with freshly mixed chemicals suitable for the films used. Besides a lower radiation dose to the patient, due to a short exposure time, blurring caused by the patient's moving will be reduced. Other factors such as the pack and prices must also be taken into consideration in the choice of radiographic solutions.

Surgical treatment of nasopalatine duct cysts. A follow-up study.

In a clinical-radiological study, 43 operated and histopathologically verified incisive canal cysts were subjected to follow-up investigation. The clinical investigation revealed no subjective symptoms. Abnormalities of the sensibility to the anterior part of the palate were noted in four cases, hyposensibility in three cases and hypersensibility in the fourth. The radiological investigation showed that bone regeneration occurred 1 year postoperatively at the earliest. After 3 years, two-thirds of the patients showed complete bone regeneration. Large cysts (diameter greater than 10 mm) showed the same tendency for bone regeneration as small cysts. The five patients in whom no bone regeneration had occurred were subjected to reoperation. The histologic investigation of these patients revealed scar tissue.