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Orchids are the most diverse family of angiosperms, with over 25 000 species, more than mammals, birds and reptiles combined. Tests of hypotheses to account for such diversity have been stymied by the lack of a fully resolved broad-scale phylogeny. Here, we provide such a phylogeny, based on 75 chloroplast genes for 39 species representing all orchid subfamilies and 16 of 17 tribes, time-calibrated against 17 angiosperm fossils. A supermatrix analysis places an additional 144 species based on three plastid genes. Orchids appear to have arisen roughly 112 million years ago (Mya); the subfamilies Orchidoideae and Epidendroideae diverged from each other at the end of the Cretaceous; and the eight tribes and three previously unplaced subtribes of the upper epidendroids diverged rapidly from each other between 37.9 and 30.8 Mya. Orchids appear to have undergone one significant acceleration of net species diversification in the orchidoids, and two accelerations and one deceleration in the upper epidendroids. Consistent with theory, such accelerations were correlated with the evolution of pollinia, the epiphytic habit, CAM photosynthesis, tropical distribution (especially in extensive cordilleras), and pollination via Lepidoptera or euglossine bees. Deceit pollination appears to have elevated the number of orchid species by one-half but not via acceleration of the rate of net diversification. The highest rate of net species diversification within the orchids (0.382 sp sp−1 My−1) is 6.8 times that at the Asparagales crown.
1. Introduction
Orchids form the largest family of flowering plants, with over 880 genera and 25 000 species; they comprise roughly 8% of all vascular plant species and grow in a wide range of habitats worldwide [1,2]. Essentially all temperate orchids are terrestrial, but most orchids inhabit tropical forests and over 80% of those are epiphytes. All orchids rely on fungi for germination and carbon capture in the protocorm stage, and in many taxa this mycorrhizal association remains obligate for life [3]. Orchids in more than 30 small mycoheterotrophic lineages have lost all photosynthetic capacity and rely entirely on fungi for energy [4]. Finally, orchids display extraordinary floral diversity, with striking adaptations to different pollinators among close relatives, partitioning of individual pollinators by precise placement of pollen packets (pollinia) on different parts of their bodies, and extreme convergence and divergence among crossable taxa. The evolution of pollinia, specialization on individual pollinators or mycorrhizal fungi, pollination via deceit, euglossine bees or Lepidoptera, epiphytism per se or associated traits such as CAM photosynthesis, and predominant distribution in the tropics (and especially in extensive cordilleras) have all been proposed as drivers of the extraordinary species richness of orchids [5–18].
A well-resolved, strongly supported, time-calibrated phylogeny is fundamental to any attempt to test such hypotheses and assess the impact of individual traits and the history of geographical and ecological spread on net rates of species diversification. Over the past two decades, molecular phylogenetics has greatly advanced our understanding of orchid relationships. Investigators have used sequences of one to five plastid loci, segments of nuclear ribosomal DNA, or single mitochondrial or low-copy nuclear genes to identify five orchid subfamilies and their relationships to each other, identifying Apostasioideae as sister to all other orchids, Vanilloideae or Cypripedioideae next divergent, and the two largest subfamilies, Orchidoideae and Epidendroideae, sister to each other [19–25]. These advances have overturned many classic views of orchid relationships based on morphology alone [7,26] and led to a new phylogenetic classification of the orchids [1,2], including 17 tribes and 44 subtribes of orchidoids and epidendroids. Molecular phylogenetic studies to date have, however, failed to agree in the placement of Cypripedioideae and Vanilloideae, or to resolve and strongly support relationships among many of the 13 tribes of subfamily Epidendroideae [25–28], which comprises around 80% of all orchid species.
To clarify relationships across orchids and test theories about the impact of various traits on their net rate of species diversification, we adopted a phylogenomic approach, using massively parallel sequencing to amass data on a large fraction of the coding regions present in the plastid genome, including a far greater number of characters per taxon than any prior broad-scale study of orchid phylogeny. We extended this phylogeny using a supermatrix approach to include representatives of 40 of 43 orchid subtribes, and calibrated this tree against the ages of several angiosperm fossils to produce a new timeline for orchid evolution, identify points in the family's history at which the rate of net species diversification accelerated significantly and determine whether such accelerations are correlated significantly with characters that have been proposed as likely drivers of orchid speciation.
2. Material and methods
(a) Phylogenetics
We aligned data for 75 plastid genes from 39 species representing all orchid subfamilies and 16 of 17 tribes, as well as 73 species stratified across all monocot orders and 23 placeholders for major groups of eudicots and basal angiosperms employed as outgroups (electronic supplementary material, table S1). A maximum-likelihood (ML) analysis of the plastome data was conducted using RAxML v. 8.0.9 using the automatic bootstrap option [29]; we compared results obtained from unpartitioned data with those using an optimal partitioning of the 75 loci identified using AIC in PartitionFinder v. 1.1.1 [30]. We conducted a similar analysis of a second dataset that added three plastid genes (atpB, psaB and rbcL) for 162 orchid species, including placeholders for all five orchid subfamilies, 18 of 19 tribes and 40 of 43 subtribes, representing all but 0.4% of described orchid species (electronic supplementary material, table S2).
(b) Time calibration
We calibrated the supermatrix tree against time using branch lengths based on atpB, psaB and rbcL in a Bayesian framework using BEAST v. 1.8.0 [31] (see electronic supplementary material). Seventeen fossils were used as calibration priors, with offsets corresponding to their minimum estimated ages (electronic supplementary material, table S3). All fossil priors were assigned a lognormal distribution (s.d. = 2), accounting for uncertainty in both absolute fossil age estimation and phylogenetic placement. Priors were also placed on the crowns of rosids, magnoliids and Caryophyllales + asterids. Due to a lack of fossils easily attributed to these clades, normal priors were placed on them with mean offsets and 95% confidence intervals mirroring the posterior ages from the exponential clock analysis of Bell et al. [32], with the expectation that the true ages of these nodes would be captured by these wide, conservative priors. Uniform priors were placed on the root node and the stem of Illicium following [33]. Two chains of 100 million generations were run. Tree files from these independent chains were combined after removing 25% as burn-in to construct the maximum clade credibility chronogram.
(c) Rates of net species diversification
To test for significant shifts in net diversification regimes across lineages, we used the Bayesian approach implemented in BAMM v. 2.0 using Bayes factors [34] (see the electronic supplementary material). Analyses were conducted on a chronogram limited to Orchidaceae and other lineages of Asparagales, for Poisson rate priors of 0, 0.2, 0.4, 0.6, 0.8, 1, 5 and 10. For each prior value, two independent MCMC chains of 50 million generations were run; after removing 15% as burn-in, we analysed the output to identify the single best shift configuration with the highest posterior density and the maximum shift credibility configuration. The relative probabilities of less likely shift configurations were also calculated for each value of the Poisson prior.
(d) Correlates of net diversification rates
Based on existing hypotheses, we assessed correlations between six sets of character states and apparent rates of speciation and extinction in orchid lineages using BiSSE [35]. Rationales for these hypotheses are given in the Discussion. Sources of phenotypic data are provided in the electronic supplementary material. BiSSE cannot calculate likelihoods on unresolved tips representing more than 190 taxa, so all tips were down-weighted by a factor of 25, with small clades rounded up to 1 [36].
Gains and losses of each character-state were mapped onto the chronogram using MP and ML. For each character, an unconstrained model for diversification was compared to models where speciation (λ), extinction (μ) and character-state transition rates (q) were individually constrained (λ0 = λ1, μ0 = μ1, q01 = q10). Likelihoods of the constrained models were compared to the unconstrained model, with significance of likelihood scores assessed with ANOVA tests. Net rates of diversification for different character-states were calculated as λ − μ. For individual traits, we measured the advantage in net diversification per million years within lineages conferred by a character-state as measures the advantage in net diversification per million years per lineage across lineages. ξ is a heuristic measure, given that the realized shifts in character-states (i.e. in births of lineages marked by those states) depends on the standing number of lineages with each alternative state at any one time. Previous studies have used ζ as the rate of net species diversification; essentially, it measures the average rate of diversification within lineages marked by a particular character-state, such as red flowers. We are coining the term 'across-lineage diversification' for ξ; it essentially measures the rate at which new lineages marked by, say, red flowers arise via mutation of character-states.
We conducted a phylogenetically unstructured comparison of differences among tropical terrestrial clades, tropical epiphytic clades and temperate terrestrial clades in net diversification rate D = (ln S)/T, where S = number of species in a clade and T = stem age of that clade [36].
3. Results
ML analysis of 77 880 aligned bases (including 31 691 informative characters, of which 9315 were informative within orchids) produced a fully resolved tree in which 26 of 38 nodes within Orchidaceae have bootstrap support values more than or equal to 98% (figure 1). The same topology, with negligible differences in branch lengths and support values, was recovered with an analysis based on the 50 partitions identified by PartitionFinder; for simplicity, we focus here on the tree obtained from the unpartitioned data. This is the first fully resolved, strongly supported backbone phylogeny for Orchidaceae, including representatives of all tribes except the small mycoheterotrophic Gastrodieae (six genera, 70 species). This phylogeny places Apostasioideae sister to all other subfamilies, then Vanilloideae, then Cypripedioideae sister to Orchidoideae and Epidendroideae, and resolving relationships among the tribes of the latter. The exceptionally short branches along the spine of Epidendroideae may help explain why previous analyses based on only one or a few genes have been unable to resolve and strongly support many of the intertribal relationships within that subfamily (figure 1). The plastome tree strongly supports the positions of two previously unplaced subtribes, with Collabiinae sister to Podochileae and Dendrobiinae sister to Malaxideae (figure 1). As expected, the mycoheterotrophic taxa examined (Corallorhiza, Neottia, Rhizanthella) all exhibited accelerated rates of nucleotide substitution. Rhizanthella has one of the most highly reduced plastomes of any angiosperm sequenced to date, with only 59 910 bp and 33 genes [37], and shows a huge acceleration of nucleotide substitution (figure 1).
Our supermatrix analysis indicates that the highly specialized Rhizanthella (which flowers underground and has fleshy fruits) diverged from its closest relatives 31 Mya (figure 2; electronic supplementary material, figure S2). This analysis largely preserved relationships among subfamilies, tribes and subtribes seen in the plastome study, but greatly increased the numbers of species and subtribes represented. The supermatrix tree identifies the previously unplaced subtribe Agrostophyllinae as sister to the unplaced genus Coelia, with both sister to Epidendreae, and Calypsoeae sister to all three lineages (figure 1; electronic supplementary material, figure S1).
Orchids appear to have diverged from the common ancestor of all other members of Asparagales ca 112 Mya, and extant orchid lineages began diverging from each other 90 Mya. Orchids diverged from the common ancestor of all other Asparagales ca 112 Mya, and extant orchid lineages began diverging from each other 90 Mya (see electronic supplementary material, figure S1, and figure 2 for ages ±95% confidence intervals). The stem age of Vanilloideae is 84 My; of Cypripedioideae, 77 My; and of Orchidoideae and Epidendroideae, 64 My, at the dawn of the Tertiary. The eight tribes and three previously unplaced subtribes of so-called 'upper' epidendroids—the clade spanned by Arethuseae and Cymbidieae—diverged rapidly from each other between 37.9 and 30.8 Mya (figure 2). A series of short branches also separate four of the five tribes of the 'lower' epidendroids (Nervilieae, Tropidieae, Sobralieae and Triphoreae). Our age estimates push back the stem and crown ages of Orchidaceae by 7 and 10 My, respectively, relative to the estimates by Gustafsson et al. [24], but are supported by a much wider set of fossil calibration points.
(a) Evolution of characters hypothesized to affect net diversification
Pollinia characterize Orchidoideae and Epidendroideae and evolved no later than 64 Mya (electronic supplementary material, figure S2a). Epiphytism appears to have evolved once at the base of the upper epidendroids, no later than 35 Mya, and to have been lost at least three times, in Bletiinae, Calypsoeae and Arethusinae (electronic supplementary material, figure S2b). A very few cases of epiphytism are also known from scattered species of Cypripedioideae (e.g. Paphiopedilum, Phragmipedium) and Orchidoideae (e.g. Dispersis, Eurystyles, Pseudoeurystyles) [2]. CAM photosynthesis appears to have arisen at least four times, in the upper epidendroids (with several subsequent losses and reappearance in Epidendreae minus Ponerinae), in Sobralieae, and in scattered species of Vanilleae (electronic supplementary material, figure S2c). Orchids initially had tropical distributions, lost them in ancestral Orchidoideae, secondarily regained them in Cranichideae, and within the latter again lost them in Pterostylidinae and Cranichidinae (electronic supplementary material, figure S2d).
Deceit pollination evolved at least once, in the stem group of all orchids except Apostasioideae, and then was lost in Codonorchideae, Goodyerinae–Spiranthinae–Cranichidinae, Neottieae, Sobralieae–Triphorieae, Tropideae, Thelasinae, Podochilinae, Agrostophyllinae, Bletiinae, Ponerinae, Aerangidinae–Angraecinae, Cymbidiinae, Catasetinae and Zygophyllinae–Stanhopeinae–Coeliopsidinae–Maxillariinae (electronic supplementary material, figure S2e). Deceit appears to involve Hymenoptera in cypripedioids, and many orchidoids in Orchideae and especially Diurideae, except for fungus gnats in subtribe Acianthinae. Dipterans are involved in pollination of many species of the pleurothallid alliance and others, whereas oil-gathering Centris bees appear to be duped by many Oncidiinae. Pollination by male euglossine bees appears to have arisen at least twice, in Catasetinae and Zygophyllinae–Stanhopeinae–Coeliopsidinae, in subtribes that evolved 16–22 Mya, with a later loss in Maxillariinae (electronic supplementary material, figure S2f). Pollination by Lepidoptera appears to have arisen at least five times, in Disinae and Orchidinae, Collabinae, Eriinae, Epidendreae and allies, and Angraecinae–Aerangidinae, with losses in Bletiinae and Pleurothallidinae–Ponerinae (electronic supplementary material, figure S2g).
Invasion of the Andes at more than 1300 m elevation appears to have arisen in at least 10 lineages, including Vanilleae, Orchidinae, Goodyerinae–Spiranthinae–Cranichidinae, Malaxideae, Calypsoeae, Epidendreae, Angraecinae, and Cymbidieae minus Cymbidiinae (electronic supplementary material, figure S2h). Invasion of the New Guinea Highlands at more than 1000 m elevation occurred in at least 15 lineages (electronic supplementary material, figure S2i). Although MP and ML reconstructions suggest many fewer invasions, deep sharing of a distribution in the New Guinea Highlands is impossible given their recent uplift ca 12 Mya [38].
(b) Rates of net diversification within orchids and their close relatives
Apparent rates of speciation and extinction at the crown of Asparagales are λ = 0.088 sp sp−1 My−1 and μ = 0.031 sp sp−1 My−1, respectively. A wide range of Poisson priors—from 0.6 to 10.0—each identified significant shifts of net diversification within Asparagales as sampled, all within the orchids, with one acceleration in the orchidoids, a second acceleration in the upper epidendroids, a nested deceleration in Agrostophyllinae plus Calypsoeae, and a further nested acceleration in Laeliinae–Pleurothallidinae–Ponerinae (figure 3). Initial rates of speciation and extinction in the last, most rapidly diversifying clade—endemic to the Neotropics—are λ = 0.510 My−1 and μ = 0.128 My−1. Rates of net species diversification per million years (λ—μ) in that clade were thus 6.8 times that at the Asparagalean crown; that across the orchidoids, 1.3 times that rate; and that at the base of the upper epidendroids, 2.9 times that rate. Low rates of net diversification persisted in the lower epidendroids, coupled with a drop by two-thirds in the speciation rate. Low values of the Poisson prior (0.0–0.4) led to two inferred shifts in net diversification, in the orchidoids and the upper epidendroids. This same set of shifts was identified as much less likely (52–82%) than the four-shift set emerging from Poisson priors of 0.6, 0.8, 1.0, 5 and 10.
(c) Correlates of net diversification rate
Evolution of pollinia significantly accelerated both speciation and extinction rates, as predicted, and resulted in a 4.9% higher rate of net diversification per million years within lineages (table 1). Only one transition to pollinia occurred, so rates of net diversification across lineages per million years caused by initiation of new lineages with pollinia present or absent was quite low (0.5%). Epiphytism significantly accelerated both speciation and extinction rates relative to the terrestrial habit, yielding a 8.8% edge in net diversification per million years within lineages; q01 and q10 did not differ significantly, so ξ = 0.0%. Tropical distributions significantly accelerated speciation and extinction rates relative to extratropical distributions, resulting in a 2.1% higher rate in net diversification per million years within lineages and 1.9% across lineages. In orchids, this last difference, however, appears to be driven primarily by habit, not latitude per se. A phylogenetically unstructured comparison among subtribes and similar taxonomic units showed that the average net diversification rate did not differ significantly between extratropical and tropical terrestrial lineages (0.115 ± 0.057 My−1 versus 0.118 ± 0.061 My−1, p > 0.92), and that the average rate for tropical epiphytic lineages (0.266 ± 0.077 My−1) was significantly greater than that for both temperate and tropical terrestrial lineages (p < 0.001) (table 2). CAM photosynthesis accelerated speciation and extinction rates relative to C3 photosynthesis, yielding a 20.3% advantage in net diversification per million years within lineages and an 11.8% disadvantage across lineages (ξ = −11.8%, table 1). This calculation overstates CAM's disadvantage across lineages, because CAM characterizes a minority of lineages (electronic supplementary material, table S4), and the actual flux between CAM and non-CAM lineages depends on both ξ and the relative abundance of CAM and non-CAM taxa. Tropical distributions significantly increased speciation and extinction rates, and boosted diversification rates within lineages by 2.1% and across lineages by 1.9%.
Deceit pollination—via mimicry of food sources, nesting sites or potential mates—significantly increased speciation and extinction rates but resulted in a small decrease in net diversification within lineages (ζ = −2.2%), balanced by a comparable increase across lineages (ξ = 1.9%). Provision of chemical allurants for male euglossine bees had no significant impact on λ or μ but increased net diversification rates within lineages by 5.2% with a smaller negative effect (−2.0%) across lineages (table 1). Pollination by Lepidoptera increased speciation and extinction rates within lineages, albeit non-significantly, resulting in a large positive effect (7.1%) on net diversification within lineages and a negligible effect on diversification across lineages (table 1). Life in extensive tropical cordilleras—as exemplified by the Andes and New Guinea Highlands—significantly increased speciation and extinction rates, yielding a large 24.9% advantage in net diversification rate within lineages, but also an apparently large 21.3% disadvantage across lineages (table 1). However, the latter effect is overstated, because most lineages did not invade these cordilleras (electronic supplementary material, table S4) and because diversification within and across montane lineages is underestimated by overestimates of the age of montane lineages in New Guinea (see above). Epiphytism and pollination via deceit, euglossine bees or Lepidoptera accelerated net diversification by large amounts, but their overall effect is less than that of CAM photosynthesis and (especially) life in extensive tropical cordilleras. A 5% advantage in net diversification per million years translates, via compound interest, into a 165% advantage in species numbers in 20 My; a 15% advantage translates into a 1536% advantage over the same period.
4. Discussion
The evolution of pollinia should accelerate speciation by permitting precise placement of pollen and allowing specialization on individual pollinators (e.g. moths versus bees) or parts thereof [6], and by increasing the importance of genetic drift, perhaps resulting in an alternation of drift with strong selection on sexual characteristics [16]. In addition, pollinia allow very small numbers of variants to produce large numbers of offspring, promoting speciation from small numerical bases (perhaps due to highly inefficient pollination), which might otherwise lead to demographic collapse and extinction.
Epiphytism should accelerate speciation in several ways. First, epiphytism is a key innovation that allows the invasion of a new adaptive zone—the branches and boles of trees—largely unoccupied by other vascular plants. Second, epiphytism should help generate and maintain high levels of plant diversity because the habitable surface of bark, branches and twigs is much greater than the ground occupied by a forest; because variation in insolation and humidity within tree crowns allows local, fine-scale niche partitioning; and because altitudinal and topographic variation in fog deposition and evaporation rates creates a range of conditions that can be partitioned at broader spatial scales and may serve to isolate populations and foster local speciation [12–15,36]. Third, epiphytism is associated with high rainfall and humidity, and thus often with tropical montane conditions; the latter can provide large-scale barriers to gene flow (e.g. deep valleys, high ridges) that can isolate populations at larger spatial scales and further accelerate speciation [13,36]. Finally, the tiny seeds associated with epiphytism can provide occasional long-distance dispersal, permitting genetic differentiation to proceed in parallel at many sites along the length of extensive montane areas, such as the Andes [36]. Indeed, epiphytism in Bromeliaceae—the angiosperm family with the second largest number of epiphytic species after Orchidaceae—is associated with an acceleration of speciation rates by 2.5- to 5.3-fold relative to terrestrial lineages [36]. Almost all bromeliads have a Neotropical distribution, so they show no confounding of latitude with epiphytism. Tropical distributions should increase rates of speciation relative to those outside the tropics as a result of greater habitat area, more stable climates, lack of glaciation, and greater opportunity for coevolution of plants and their mutualists and specialized herbivores [17]. Within orchids, temperate versus tropical latitudes per se appear to have had no significant effect on diversification, but the epiphytic habit appears to have accelerated net diversification rates by 8.8% per million years (table 1). Gravendeel et al. [12] used a phylogenetically unstructured analysis to show that epiphytic orchid genera contain more species, on average, than terrestrial genera. Our results generalize their findings and are the first rigorous demonstration, for orchids, that epiphytism accelerates differentiation when phylogeny at various levels is taken account.
CAM photosynthesis had no significant effect on net diversification in bromeliads—in which it occurs both in terrestrial lineages on dry sites and in epiphytic lineages [36]—but it did accelerate speciation, extinction and net diversification in orchids, perhaps because it is so closely tied to epiphytism in orchids [15] and may permit invasion of the most exposed perches, as well as drier forests at lower elevations.
Deceit pollination characterizes one-third of all orchids [9,39]. Mimicry of potential mates is much less common than mimicry of food or nesting sites, but is thought likely to accelerate speciation because subtle changes in floral morphology or volatile compounds can attract different pollinators and lead to reproductive isolation [9], and often results in high efficiency of pollen transfer to conspecifics [40]. We propose that pollination via deceit inevitably involves a density-dependent advantage of new mimetic morphs that are rare relative to models; such negative density-dependence is a potentially strong but previously overlooked force that could favour high rates of diversification. Pollination by male euglossine bees should accelerate speciation by allowing small chemical changes in the allurants provided by orchids to attract bees that are reproductively isolated from each other and by permitting partitioning of individual bee species by placing pollinia on different parts of their bodies [6,18]. Pollination by Lepidoptera often involves nectar storage in corolla tubes or spurs whose lengths can be easily modified and, in so doing, lead to the rapid recruitment or evolution of different pollinators and, ultimately, to accelerated rates of speciation. Floral spurs are not, however, always associated with nectar production [41], so that the argument favouring the evolution of progressively longer spurs and pollinator mouthparts [42] does not necessarily hold, and is difficult to test given the possibility that some orchids (e.g. Disa) use empty spurs as a deceitful means of attracting pollinators.
This study shows that orchids are remarkably species-rich partly as a result of three likely accelerations of net diversification rates, apparently driven in part by the acquisition of pollinia, epiphytism, tropical distributions, CAM photosynthesis, pollination via deceit, male euglossine bees or Lepidoptera, and life on extensive tropical cordilleras. It must be recognized, however, that shifts in net diversification are scale-dependent. Our sampling is well suited to detect shifts in diversification at tribal or subtribal levels within orchids, but inadequate to identify such shifts within genera. Nevertheless, our findings provide the first quantitative support for several earlier hypotheses regarding the genesis of orchid diversity and identify specific points in orchid evolution where these factors played a role. They show that multiple factors—several of them interconnected—have contributed to orchid diversification.
The first significant acceleration of net diversification is at the base of Orchidoideae, one node removed from that at which pollinia evolved (figure 3). The second acceleration coincides with the origin of epiphytism in the upper epidendroids and, less precisely, with the origin of several montane clades in the Andes. A deceleration of diversification occurs in Epidendreae–Agrostophyllinae–Calypsoeae, as expected because this group does not consist of montane epiphytic lineages: Calypsoeae are temperate terrestrials, most Agrostrophyllinae are epiphytes from lowland tropical forests, and—uniquely within tribe Epidendreae—early-divergent Bletiinae have regained the terrestrial habit. The final acceleration coincides with the later divergent elements of tribe Epidendreae and a large concentration of Andean taxa pollinated via deceit or Lepidoptera. Pleurothallids, the most diverse element of this clade, are largely pollinated by dipterans. Low rates of net diversification persisted in the lower epidendroids, which are marked (as are orchidoids) by a lack of epiphytism but, unlike orchidoids, largely lack pollination via deceit, euglossine bees or Lepidoptera. We suspect that this last distinction accounts for low diversification in the lower epidendroids and the displacement of the first acceleration of diversification from the orchidoid–epidendroid crown to the orchidoid crown.
The synergistic effects of epiphytism and life in extensive, topographically complex, tropical cordilleras on geographical speciation at small spatial scales are likely to have had quite large effects on orchid diversity, given the very large number of species in certain clades centred on the Andes (e.g. Cymbidieae minus Cymbidiinae, Epidendreae) and the New Guinea Highlands (e.g. Dendrobiinae), as well as the high local diversity, narrow endemism and rapid spatial turnover of orchid species in these areas [43,44]. The effects of epiphytism per se, CAM photosynthesis and life in tropical cordilleras cannot be teased apart using current techniques, and will always be hard to separate given the causal links among these traits. Surprisingly, epiphytism appears to have had a stronger effect than tropical distribution in accelerating species diversification in orchids. There is no doubt that sexual selection and floral diversification have had a large impact on orchid diversification (table 1). The fact that approximately one-third of all orchids engage in deceit pollination—a mechanism almost unknown in other plants—is especially striking, and suggests that the evolution of such pollination may have increased orchid diversity by 50% over what it otherwise would have been, given non-deceit pollination in the remaining two-thirds of all orchids. Yet deceit, considered alone, appears to have augmented orchid diversity not by accelerating net diversification, but simply by adding more species at roughly the same rate.
The fact that orchids show no significant acceleration of diversification relative to other Asparagales among the three earliest divergent subfamilies (figure 3) implies that the defining characteristics of the orchids as a whole—the column, mycoheterotrophic germination, minute seeds lacking endosperm and a specialized labellum—did not, by themselves, accelerate diversification. But these traits almost surely acted in concert with the evolutionary triggers identified above to promote high levels of speciation. The ability of pollinia to allow small numbers of variants (based on limited pollination or recent mutation) to produce large numbers of offspring, promoting speciation from small numerical bases, depends on the production of large numbers of (therefore) small seeds, and such seeds would be favoured by mycoheterotrophic germination [45]. The dispersal of pollinia and extreme specialization on individual pollinators or pollinator body parts clearly were facilitated by the orchid column and labellum [6]. Evolution of epiphytism was surely favoured by the possession of tiny, dust-like seeds that could settle on twigs and branches regardless of orientation.
The extent and resolution of our phylogenetic reconstruction is inadequate currently to detect the likely diversifying influence of repeated, small-scale adaptive radiations in pollinators or mycorrhizal fungi. Within Platanthera, Disa, Corycium and Drakaea, closely related species often diverge in pollinators or the placement of pollinia on the same pollinator, suggesting that adaptive radiation in pollinators or pollinia position may be an important driver of diversification among closely related orchids [6–8,46]. By contrast, the evidence for narrow specialization of closely related orchids on different mycorrhizal fungi is limited to mycoheterotrophic taxa; photosynthetic species often overlap strongly in their fungal partners [46]. Pollinators but not mycorrhizal fungi are likely to provide a mating barrier between orchid species.
Finally, invasion of extensive tropical cordilleras per se should accelerate speciation, extinction and net diversification, given the abundance of natural barriers to gene flow, dynamic shifts in the location of favourable habitats, and the possibility for geographical speciation proceeding in parallel along the lengths of mountain chains [13,14,17,36]. Invasion of such cordilleras as the Andes and the New Guinea Highlands had by far the largest positive effect on net species diversification within lineages (see above).
We suggest that three additional factors likely to contribute to orchid diversification should be explored, as follows.
(a) Dispersal, time and familial range
The small seeds of orchids and the long time since their initial diversification almost surely fostered greater diversity by permitting them to spread to every continent and most latitudes. Across angiosperm families, species richness and age are unrelated, but families with broader geographical and latitudinal ranges are more diverse [47]. Analyses of orchid historical biogeography (Givnish et al., in preparation) suggest that intercontinental dispersal in orchids is relatively uncommon, but alone appears to have greatly increased orchid species number over what it would have been had orchids remained on a single continent.
(b) Karyotypic evolution
Chromosomal evolution is often overlooked as a diversifying influence in orchids. Yet orchids show a remarkable amount of polyploid and aneuploid variation in chromosome number across subfamilies and within many of the largest genera (e.g. Bulbophyllum, Dendrobium, Epidendrum, Malaxis, Pleurothallis s.l.) [48]. The combination of small seeds, early mycoheterotrophy and pollinia in orchids allows demographic recovery from repeated bottlenecks in effective population size; such bottlenecks are likely to fix mutations via drift, and fixation of karyotypic mutants would quickly generate post-mating isolation barriers.
(c) Limited gene flow or local pollinators in montane epiphytes
A number of epiphytic groups from tropical cordilleras (e.g. Bulbophyllum, Lepanthes) show substantial genetic differentiation among populations even though most temperate orchids surveyed do not (see data of Phillips et al. [49]). The evolution of dozens of closely related Teaguiea species over only a few kilometres in Andean Ecuador [50] and rapid geographical turnover of species identity with distance in Andean epiphytic orchids more generally [43,44] are both consistent with gene flow over only short distances. Whether this is a result of rapid rainout of seeds over a short distance in wet tropical cordilleras, or limited physiological tolerance or dispersal ability in small, soft-bodied, desiccation-intolerant dipterans and other weak-flying pollinators specific to individual species or clades, is currently unknown. The roles of limited dispersal and species ranges of such pollinators, and of intermittent population bottlenecks and drift on chromosome number and speciation in epiphytic orchids, deserve further study.
Data accessibility
All data are presented or cited in the electronic supplementary material.
Authors' contributions
T.J.G. and K.M.C. conceived the study; M.A., K.M.N., W.M.W., N.H.W., J.L.-M. and K.M.C. obtained the molecular data; S.P.L. and L.E. contributed biogeographic data; D.S., A.Z., W.J.D.I., R.K., S.J.H. and T.J.G. participated in data analysis; M.A.C. and M.T.K.A. provided critical plant material; T.J.G. drafted the manuscript; all authors provided comments and final approval.
Competing interests
We have no competing interests.
This research was supported by grant no. DEB-0830836 to T.J.G. from the NSF Assembling the Tree of Life (AToL) Program; collection efforts in Chile were supported by grant ICM P02-005 to M.T.K.A.
We thank Joel McNeal, Erica Wilson and Laura Williams for access to additional plant material.
- Received June 29, 2015.
- Accepted July 27, 2015.
- © 2015 The Author(s)
Published by the Royal Society. All rights reserved.


text:
– As noted before, the February 4th WWE RAW did a strong 3.54 rating with 4.80 million viewers.
The show opened huge with a 3.79 quarter rating with CM Punk's promo. Ryback vs. Antonio Cesaro lost 526,000 viewers. Rey Mysterio's promo with The Shield video along with a Trish Stratus Hall of Fame video gained 373,000 viewers. Santino Marella vs. Jack Swagger lost 302,000 viewers. Cody Rhodes vs. Alberto Del Rio and then a Big Show/Del Rio interview segment at 9pm gained 243,000 viewers for a 3.64 quarter rating.
Mysterio vs. Daniel Bryan lost 93,000 viewers. Kane and Bryan arguing plus another Big Show segment in his hotel room lost 87,000 viewers. Kane vs. Sheamus lost another 167,000 viewers. The MizTV segment with Paul Heyman and Brock Lesnar gained 600,000 viewers at 10pm for a 3.82 quarter rating – the high point of the show.
Wade Barrett vs. Randy Orton lost 683,000 viewers. Chris Jericho vs. CM Punk gained 137,000 viewers. Bruno Sammartino's Hall of Fame video gained 34,000 viewers. Big Show and Del Rio's brawl plus the Brad Maddox promo and brawl with The Shield, John Cena, Ryback and Sheamus gained 255,000 viewers for a 3.63 overrun rating – a weak gain for the overrun.
Source: Wrestling Observer Newsletter

subset: commoncrawl


text:
Mad Men premiered Sunday night and lesbians across American rejoiced.
We will have plenty to say about the marvelous women of Mad Men this season, but for now, let's look at Episode 1's L-word moment.
Don Draper (Jon Hamm), who is determined to be a faithful father after Betty (January Jones) issued an ultimatum last season, spent some quality time with daughter Sally (Kierman Shipka) last night.
Sally has been acting out a bit because Daddy's gone a lot. In fact, she ruined his valise by smashing the clasp with a hammer. But when Betty tells Don about it, she jokes, "… she's taken to your tools like a little lesbian."
Wait, what? Two conventional Sixties parents had an exchange that so casually dropped the word "lesbian" in reference to their daughter? Really?</>
I'm not quite old enough to remember words that were in casual usage in 1963, but I do know that I never even heard the word "lesbian" until college — maybe later. The two girls in my dorm who walked down the hall holding hands were "homos." Athletic girls were "tomboys" and older unmarried women were "spinsters," even when two of them lived together. Then again, I was raised in the Bible Belt by Southern Baptist parents and went to Baylor. I didn't exactly live in a progressive world.
Still, in 1963, Stonewall had not happened and homosexuals were far from accepted. Mad Men's storyline about Sal's struggles with being a closeted gay man indicates that even in a New York ad agency, homosexuality is a taboo. And last season, when the normally outspoken Joan (Christina Hendricks) hears a confession of love from her roommate Carol, she simply pretends not to understand.
My initial reaction was that Mad Men goofed. But a comment I read on the fabulous Mad Men fansite Basket of Kisses made me reconsider. As the commenter pointed out, Betty attended Bryn Mawr, one of the Seven Sisters colleges. She doubtless had lesbian classmates, whether she regarded them favorably or not. Perhaps Betty's background gives credence to her comment about Sally.
Does anyone know if "lesbian" was used casually in the early Sixties? Or was Betty's remark anachronistic? Do you care if Mad Men time shifts details for the sake of the story?


text:
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Have been going there for 20years have always had great experience. Everyone always very friendly and knowledgable. lots of great stuff in stock. Go...
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I have used Mr Ed's for ten years at two different houses. I have recomended them to several of my friends, and would recomend them to any body that a...
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Would be nice if I could get someone to answer your phone when I call....
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Aqua Scape Specialties
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Sunshine Pools Inc
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brett k
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Delta Pools
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Horrible experience - BEWARE - I hired Chip at the beginning of May because my pool was leaking. Chip said he would order a new liner and it would tak...
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Mr Ed's Swimming Pool Service
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they cleaned and filled out pool with our water softened water. instead of compensating for the low calcium level, they just assumed it was a certain ...
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Walt Clark
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Al Shirley & Sons Inc
Winchester, VA
Laurie S
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you guys have been awesome your professionalism has been to the utmost. I am extremely happy with the work you have done. we are really pleased with ...
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Great people who know what they are doing. Friendly family owed business. Good honest service and fair prices ....
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Pool service claimed I needed a new motor on polaris pump. Cost was $350. It took about 15 minutes to remove my existing motor and then another 30 m...
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The best and friendliest Vic is Awsome...
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Joe is a great guy! We got our pool installed yesterday, and it looks great! Thanks Joe!...
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Sparkling Pool Service
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East Tennessee Pool & Spa
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Over charge and left without the pool even working or cleaned. Sent boys to open the pool but moved slower than you could imagine. No pride in job. Wi...
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Phoenix Spas Corporation
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text:
We dream view this image of the three brothers together, so ... you see them side by side, it transports us to the wonderful image of the belly of her mother when she was expecting a baby. Patience and love are the feelings that filled the environment, we were all very expectantly, once achieved we hugged making with parents and it was exciting to see in the viewfinder of the camera this memory for a lifetime.
Up,up, and away! He was such a champion sleeper. Sometimes you get super lucky and work with a baby that makes your job a bit easier. Mak THE explorer.
This photo is a 2016 version of an iconic image Holy Mother, we call them saints because we photographed babies enter the world with all the innocence in the purest state, full of tenderness and have an angelic beauty that radiate pure light. The star of this picture is called Ignacio and his mother Nanu!
This little sweetheart's mother wanted to incorporate her wedding gown into an image for her newborn session. As she may never actually wear the gown in the future, this image is a precious memory for her parents.
This little sweetheart was somewhat alert for her portrait session, but she loved being wrapped up. Her parents loved my angora bunny bonnet, and we all thought that she looked just perfect as a snuggly little bunny!
This is the beautiful Baby Juliana, nestled sweetly amongst the flowers, butterflies, and jewels. A portrait fit for a princess.
This is my little client Eva and her father. She is 12 days on this picture. Father was very creative during the Photosession and we took a lot of amazing pictures in that day.
This is little Oliver, just a few weeks new. Trying out his new toes and feet in mommy's arms.
@etrolid @tinytoes_photography
This is gorgeous baby Harlo! The image was taken in my home based studio, using natural light. He was such a perfect little sleepy baby to photograph :)
This image was taken during a newborn session in my home based studio using natural light. The love between these three was amazing, and I love their expressions in this image.
This image was a composite on my carousel horse using mom and dad to help:) Mom LOVES carousel horses and when she saw mine as a prop in the studio she had to use it for their session!
This image was a composite image. I felt like having a little play around and being a bit creative. The raft is home made, and the image was taken in my home based studio. It was a cloudy day, so I used my Jinbeii continuous light. I then used a gorgeous digital backdrop :)
These adorable twin boys were exactly like peas in a pod! Aren't they the cutest??
All of 25 days of age but posed like 5 day old newbie <3
The parents of small Stern really wanted, just as they have done for his sister, also make a photo shoot for his 1st birthday.
He was a lovely cheerful man who has laughed the entire shoot.
Great to photograph these children.
Sweet little Carmen.
She was born with 35weeks and such a strong girl.
Came home from the hospital verry soon.
Carmen did a great job on her newborn session.
she's my youngest daughter, Olivianne Natalie. She is 3th years age. I take this picture when golden hours moment , September 2016. This is one of my favorite photoshoot.
Play to create is one of the principles of Holy Mother Photographers, play to be children again and to imagine beyond making is what we love to do. Every detail is thought out and makes the digital retouching allows us to tell a story of fantasy. Felipe is the magician full of magic and joy this cirquense moment.
One of my best friends is a Harry potter fan, so we decided to make a Harry potter setup with her gorgeous new daughter Quinn! I had the hat and owl specially made and my amazing husband hand crafted the curved bench for me!
Oh the beautiful baby Aivlee! She's one of my hello baby milestone planners and this was a custom set up for her 6 month photoshoot. She's totally gorgeous in this soft pink set up.
Meet newborn Kate Scarlett
This beautiful girl was Born earlier and was so small.
The shoot went great.
Kate slept trough the whole photoshoot.
This little girl stole my heart!


text:
Henry Bloom, the owner of a Hanover Township massage parlor, will be tried in Northampton County on prostitution and related charges, Judge Richard D. Grifo ruled Wednesday.
Bloom's attorney, Lawrence Fox, had asked that Bloom be tried elsewhere because of what he termed extensive publicity about the case. He had also asked that certain items of evidence seized during the execution of a search warrant at The Body Clinic, 1452 Roselawn Drive, not be admitted into evidence. He also wanted certain statements Bloom made to police before he was arrested not to be introduced during the trial.


text:
Hi everyone,
I'm creating my own OS from scratch, and I'm first producing an installation CD so that I can easily install my kernel on a hard drive.
So far, I've written a CD-based bootstrap loader in NASM that finds a program within the
root directory of the CD, loads it into a known area in memory, and executes it (this uses the ISO-9660 specification). I'll more-than-likely just remain in 16-bit mode to run my installer, and switch to an installer that runs within the kernel once I get far enough (i.e. once I actually have a kernel!).
Now, my plan is to make the CD work like a Windows installation CD; I want it to display a "Press any key to boot from the CD..." prompt, and 'skip' to the next boot device if a key isn't pressed within a certain time limit. The only thing is, I'm not sure what I need to do to skip to the next boot device?
Thanks in advance!
P.S. Please could any example code be written in NASM or pseudo-code? :-)


text:
The table is set for our twin bill 2011 Key West Literary Seminar: THE HUNGRY MUSE: An Exploration of Food in Literature. Panelists have been apportioned to one of two (or in some cases, both) sessions. Here's how the menu is shaping up:
Session One: January 6 – 9, 2011
Diana Abu-Jaber, Roy Blount Jr., Frank Bruni, Billy Collins, Jason Epstein, Jonathan Gold, Darra Goldstein, Daniel Halpern, Madhur Jaffrey, Judith Jones, Harry Mathews, Bich Minh Nguyen, Molly O'Neill, Julia Reed, Ruth Reichl, Calvin Trillin, and Kevin Young
Session Two: January 13 – 16, 2011
Elizabeth Berg, Roy Blount Jr., Kate Christensen, Billy Collins, John T. Edge, Adam Gopnik, Gael Greene, Jane Hirshfield, Madhur Jaffrey, Mark Kurlansky, David Mas Masumoto, Nicole Mones, Molly O'Neill, Michael Ruhlman, and Calvin Trillin
We may add a few more panelists over the summer. Speakers and session assignments will be announced as they're added to the roster– right here on Littoral, and also on our Speakers page, where you can see all of our current speakers, with links to their biographies, bibliographies, and other information from around the Web.
You are welcome to attend either or both sessions of the Seminar, which run from Thursday to Monday. If you are interested in registering for the Seminar, we urge you to act soon, as seats do fill quickly. We will hold your space for a deposit of $100. Click here to register.


text:
Bass Pro Shops has teamed up with Griffin International to deliver a series of outdoor enthusiast games, coming to Xbox 360, PC, and Wii within the coming year. The fishing game, called The Strike, is coming out later this year. You can study up on that title by reading our recent The Strike preview. It wasn't until just now at the Electronic Entertainment Expo that we were able to see its partner in crime, The Hunt. If you couldn't tell by the name, this game is all about hunting. And with the Bass Pro Shops license, you can expect authentic in-game gear to go along with the light gun peripheral when the game releases early in 2010.
The Hunt largely follows the same model as The Strike. For the casual gamer, a series of quick play features exist. The one I got a look at was an on-rails shooter at a leisurely pace. All you have to do is point and shoot at the adorable wildlife. That includes everything from cuddly rabbits to hulking bears. With a couple dozen animals in the game, there's plenty to aim your sights at. Other mini-games include the likes of skeet shooting and ATV racing.
For the more avid hunter there's the career mode. There you can build up your hunter and purchase new weapons and gear like shotguns, crossbows, buck scents and more. Out in the woods, each region offers up a series of hunts that can be tackled in any order. Legendary animals can be hunted. Special tasks can be completed. A GPS can be pulled up to locate missions or figure out where you are should you get lost in the large levels.
Before you can move on to the next area, you'll have to earn enough reputation points to unlock it. These are earned by completing tasks, though you won't need to pass them all to move on. It's all designed to be non-linear and allow players to move at their own pace doing what they like.
As a Bass Pro Shop game, certain concessions have been made in the name of safety and realism. You can't ever run with your gun up since that is a big no-no in real life. You'll also find yourself penalized if you start hunting does instead of the big bucks. It's all about recreating the hunting experience, even if that means you aren't running around shooting everything like a maniac. This realism can also be a big positive for avid hunters. Finding a big buck isn't a matter of random luck. Masking your scent and sitting in camouflage is a good start. If you don't think you're in a good spot, it might be a good idea to start looking for deer trails to follow or fresh droppings.
The Xbox 360 peripheral wasn't ready to be shown, but I did get my hands on the Wii peripheral. This gun casing holds both the Wii remote and the Nunchuk, housed in a plastic holster that is shaped a bit like a figure-8. It feels good in your hands and has a weight to it. There's even a bolt to pull back each time you want to reload, which is a pretty cool feature.
The Hunt is due out in early 2010 on PC, Xbox 360 and Wii. It will have the same features across all platforms.


text:
Invasive Lionfish speared by Diver
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text:
While unprecedented attention is being focused on developments in Central America, Mexico, and most of South America, not many in Washington seem to be aware that there is serious trouble brewing in Puerto Rico.
One who is aware is Louisiana's Sen. J. Bennett Johnston, who said: ''Puerto Rico could become a dramatically incendiary issue that could virtually destroy our Caribbean policy, our image, and our credibility, and bring us untold political miseries. All the elements are there for that to happen.''
The basic element, according to Senator Johnston, is that a large and growing number of Puerto Ricans are being led to believe that ''a request for statehood is a guarantee of statehood.''
Puerto Rico's pro-statehood governor, Carlos Romero Barcelo, when asked recently what he would do if Congress did not act, answered: ''The real answer to that is that I don't think that will ever happen, because it will mean that Congress would be negating democracy itself.''
As US citizens, Romero explained, Puerto Ricans have ''a constitutional, inherent right'' to statehood. Congress and the nation ''have a moral commitment to the people in Puerto Rico'' to approve it.
But it isn't just Romero. Mr. Johnston notes that Puerto Ricans ''can be made to feel that they have been misled, lied to, deceived, because every president since Eisenhower has said, 'Your status is up to you.' ''
Clearly, it isn't up to the Puerto Ricans. The decision whether to admit Puerto Rico would be made in Washington, not in San Juan. And, to say the least, it would not be an easy matter for Congress.
Puerto Rico would be, by far, the poorest state: Its per capita income is one-half that of Mississippi, one-third the national per capita income. Several federal studies indicate that major changes would be required to convert the island into a state-like economy. Puerto Rico is exempt from federal taxes.
Puerto Rico's industrialization, its once-famous Operation Bootstrap, was based precisely on the present unique relationship to the US. Thus there is not only the question of whether Puerto Rico could withstand the additional weight of federal taxation, but whether it could accelerate its economic growth so as to close the enormous gap with the other 50 states.
Island statehooders respond that Congress would approve a post-statehood 20 -year economic transition period, gradually phasing in federal taxation and phasing out several existing benefits, such as the federal rum tax ''rebate'' now received by the island treasury.
Governor Romero has also suggested that the US take over Puerto Rico's public debt, now about $8 billion.
Congress would have to deal with a sensitive cultural issue. Puerto Rico, according to island statehooders, would become the ''Hispanic state,'' retaining its own distinct cultural identity. In a book titled ''Statehood Is for the Poor ,'' Governor Romero wrote: ''As I have said so often, our language and our culture are not negotiable!''
However, in admitting into the union four territories with large non-English-speaking populations - Louisiana, Oklahoma, New Mexico, and Arizona - Congress did require that English be declared the official language. Would Congress not require the same of Puerto Rico?
Puerto Rico would have more representation in Congress than 27 states: no fewer than seven representatives and the two senators. Would the House reapportion itself or increase its size, something it has not done in 70 years?
Needless to say, there are important political implications in all this, especially in view of the fact that Puerto Ricans historically have voted straight party ticket. Thus presumably the entire congressional delegation would belong to the same party.
Finally, there is little in recent history to suggest that Congress would act speedily on statehood. Hawaii and Alaska waited a half century after being declared ''incorporated territories,'' which in fact meant a commitment to eventual statehood.
Yet President Reagan assured Puerto Ricans this year that ''Congress and the people of this country would welcome Puerto Rican statehood.'' President Carter, in a 1978 message read on the island, said that if Puerto Rico selected statehood, ''it will be yours.''
The danger is this: A Puerto Rican petition for statehood would require Congress to face and unravel a series of highly complex economic, political, and cultural issues without precedent in US history. So, to the degree that local and US political leaders convince the Puerto Ricans that statehood is there for the asking, they are leading the island and the US into stormy waters.


text:
This 21 acre property sits west of Plymouth, IN 46563 right off of Interstate 30 in Marshall County. The property has potential for a future building site or small retreat area. This piece offers some recreational opportunities such as ATV trails, hiking trails, and others. Wildlife such as whitetail deer, turkeys, birds, and other small game are...21 Acres : $70,000
Rose Rd, 21 Acre Parcel, West
Plymouth : Marshall County : IN


text:
25 Cryptic Crosswords and 'How To Do Cryptics' tutorial.
A collection of 25 quality crosswords for cryptic crossword fans, plus 'How To Do Cryptics' tutorial for those eager to learn or hone their skills.
We have graded our cryptic crosswords according to how long they take to complete. This issue has been graded level 2. The level 1 crosswords are the quickest to solve, level 2 crosswords take a bit more effort and to solve the level 3 crosswords you have to really go up a gear and remember to think laterally!
This interactive book is packed with original features, including a handy Clue Bar which displays intersecting clues for easy reference. The Toggle Button allows you to navigate effortlessly between 'Across' and 'Down' and the Auto-Check function is a useful tool for highlighting letters entered incorrectly. There are unlimited Word/Letter Hints to ensure you're never stuck and your progress on each puzzle is saved automatically for future completion. You can clear and solve each crossword time and time again.
Brought to you by Lovatts Media Group, creators of the popular Magic Touch Puzzles series.


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Show how much you care for your loved one with this gorgeous heart ring. Made from sterling silver, it features a cutout heart that has been adorned with sparkling cubic zirconia. It's a dazzling expression of love.
Every girl deserves to feel like a princess. With this stackable ring, you'll feel special every day. This romantic ring features a silver tiara set with a sparkling cubic zirconia atop a sterling silver band. Perfect as a special gift to wear alone or in a beautiful stack.
Angel wing series rings each face is different, it took off the black mystery veil, let us see the most noble and dignified side of it. Like every woman is looking forward to a beautiful metamorphosis, it finally completed the transformation, and with a more noble gesture to come to your side, brings the brilliancy to you.
This ring shows a retro and elegant feeling in terms of shape and color aspects, three-dimensional part of the side looks like a pair of angel wings. It's finely textured and intricately sculpted to resemble lace. Wear this delicate piece every day. It is said that your IQ will be zero when you fall in love with someone, do not lay down your dignity in front of love, love never menial, may it bring you more confidence.
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Harry Potter and the Goblet of Fire (2005) BRRip 720p
13,839,720 views Harry, Ron and Hermione enter their fourth year at Hogwarts. After having a strange dream of the Dark Lord and his Death Eaters...
Harry Potter and the Half-Blood Prince (2009) 720p
12,809,170 views Scarred by his experience at the Ministry of Magic with Voldemort, Harry is reluctant to return to Hogwarts...
Harry Potter and the Deathly Hallows: Part 1 (2010) BluRay 720p
10,992,760 views The film begins with Harry Potter ( ) and Hermione Granger ( ) starting their search for Voldemort's Horcruxes...
Alice in Wonderland (2010) 480p
10,115,670 views Alice Kingsleigh (Mia Wasikowska) is the daughter of Charles Kingsleigh (Marton Csokas), a wealthy man who planned to find profitable shipping routes through the world in the 19th century...
Mr. Turner (2014) BRRip 720p
2,306,840 views Mr. Turner explores the last quarter century of the great if eccentric British painter J.M.W. Turner (1775-1851). Profoundly affected by the death of his father, loved by a housekeeper he takes for granted and occasionally exploits sexually, he forms a close relationship with a seaside landlady with whom he eventually lives incognito in Chelsea, where he dies. Throughout this, he travels, paints, stays with the country aristocracy, visits brothels, is a popular if anarchic member of the Royal Academy of Arts, has himself strapped to the mast of a ship so that he can paint a snowstorm, and is both celebrated and reviled by the public and by royalty.
Enchanted (2007) 360p
2,197,420 views The beautiful princess Giselle is banished by an evil queen from her magical, musical animated land and finds herself in the gritty reality of the streets of modern-day Manhattan. Shocked by this strange new environment that doesn't operate on a "happily ever after" basis, Giselle is now adrift in a chaotic world badly in need of enchantment. But when Giselle begins to fall in love with a charmingly flawed divorce lawyer who has come to her aid - even though she is already promised to a perfect fairy tale prince back home - she has to wonder: Can a storybook view of romance survive in the real world?
Upside Down (2012) BluRay 720p
2,131,340 views Look up towards the sky and rub your eyes because you won't believe what you see: cities, forests, and oceans with their own inverted gravity...
Chicken Run (2000) BRRip 360p
1,126,230 views Having been hopelessly repressed and facing eventual certain death at the chicken farm where they are held, Rocky the rooster and Ginger the chicken decide to rebel against the evil Mr. and Ms. Tweedy, the farm's owners. Rocky and Ginger lead their fellow chickens in a great escape from the murderous farmers and their farm of doom.
The Last Samurai (2003)
805,570 views In the 1870s, Captain Nathan Algren, a cynical veteran of the American Civil war who will work for anyone, is hired by Americans who want lucrative contracts with the Emperor of Japan to train the peasant conscripts for the first standing imperial army in modern warfare using firearms. The imperial Omura cabinet's first priority is to repress a rebellion of traditionalist Samurai -hereditary warriors- who remain devoted to the sacred dynasty but reject the Westernizing policy and even refuse firearms. Yet when his ill-prepared superior force sets out too soon, their panic allows the sword-wielding samurai to crush them. Badly wounded Algren's courageous stand makes the samurai leader Katsumo...
The Love Punch (2013)
712,780 views Richard and Kate are a divorced couple who have an amicable relationship. Richard who's about to retire learns that his company's assets have been frozen because it's under investigation and that includes the pension fund. When the owner goes out of the country, Richard decides to pursue him and Kate goes with him. When they learn the man doesn't care about the employees, they decide to get the money some other way; by stealing the diamond he gave his girlfriend. So they follow them and Kate gets close to the girlfriend.


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Opera North
by Stuart Sweeney
August 23, 2012 -- Barbican, London, UK
"Carousel" is one of my favourite musicals, ranking alongside "Oklahoma", with superb songs and having more dramatic weight than many, although raising concerns that are discussed later. Opera North's production is full of vitality and visual interest to set alongside the wonders of the score. Apart from the most well known numbers such as "You'll Never Walk Alone" and "June is Busting Out All Over", songs such as "Mister Snow" and "When the Children Are Asleep" have an easy, tuneful grace that I find delightful.
I particularly enjoyed the songs performed by the male members of the cast: Eric Greene as Billy, the rough-neck central character, Michael Rouse as Jigger, the thief and killer who leads Billy to disaster, and Joseph Shovelton as Enoch, the upstanding contrast to the ill-fated pair; all three command the stage and their voices suit the songs admirably. Claire Butler as Carrie makes the most of her role's humour and sings in a bright, winning style. Gillene Herbert as Julie achieves a strong emotional connection with the audience, and as Nettie, Elena Ferrari commands the stage; however with both singers I found their operatic vibrato too great for the songs.
The dance and movement direction provide heaps of energy to the show. The ensemble dance by Kay Shepherd, and the "ballet" and "June is busting ..." sections by Kim Brandstrup fit seamlessly together. In the extended duet, Alex Newton as Julie's daughter and Nicholas Cass-Beggs make the most of Brandstrup's choreography – with its lifts and swings effectively conveying the dreams of the 15-year old girl. Anthony Ward provides arresting, stylised sets accommodating the different scenes.
The production opens with the wonderful "Carousel Waltz", but with a newly added, potted history of the childhood of Billy Bigelow, the fairground barker at the heart of the narrative. This attempts to provide some background for his character, but it passes too quickly to make an impact. Presumably it has been included to counter a problematic aspect of "Carousel" - the attitudes to women on display. Two sections of the show highlight this aspect. The words of the lovely song "What's the Use of Wondering" include:
Oh, what's the use of wond'ring
Something made him the way that he is,
So, Julie belongs to Billy and it doesn't matter how he behaves in general and to her. Secondly there are the two instances where Billy hits his wife and later his daughter; in response to her daughter, Julie says, ""Yes, Darling, I believe that it IS possible for somebody to hit you, good and hard, and for it not to hurt." This romantic approach to domestic violence is disturbing, especially when it comes from the most sympathetic character in the show. But many books, films and productions from 50 years ago and more contain casual racism and other attitudes unacceptable today. However, you don't have to agree with its thesis to admire an art work, although this may well lessen your emotional involvement. I would prefer that "Carousel" didn't espouse this acceptance of woman as victim, but in Opera North's fine production, the love story and the wonderful score combine to overcome this shortcoming.
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text: At last Printit4u will now be able to offer all customers full colour personalisation on the front of our promotional notebooks Stunning full colour imagery can now be printed on the Tuscon and Matra medium Ivory notebooks from our Castelli diary and notebook collection. From colour logos to full colour covers we can print high definition images. For further information please contact printit4u


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pbbthhh bologna wtf is that???
pbbthhh bologna wtf is that???
Exaggeration but it's still pretty damn funny
I watched this 14 times before having a heart attack.I will not sue you but only because this is so damn funny.
wtf bolnga
so true
why is it spelled bologna?!?!
Straight to the point
Short and sweet :).


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Qigong Massage DVD with Dr. Yang, Jwing Ming
This DVD is an excellent introduction to the art of massage and to the acupressure points and channels in the human body. It presents the fundamental techniques and theories of Qigong massage that therapists can use to enhance their skills, and deepen their knowledge and application of Qi (energetic) healing.
Not only for therapists! Novices will find the instruction practical and useful for recovering from fatigue, aches, and pains, tension, and more. A special section on Self-Massage is presented to help those who wish to enhance massage treatment benefits, or who lack a partner or therapist.
This DVD contains 70 minutes of Self-Massage instruction, and 120 minutes of basic Two-Person whole body massage techniques. Additional DVD-only content offers a complete introduction to the concepts of Qi and Qigong, and provides a glossary of Chinese Qigong terminology.


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Amanda Seyfried: Urth on Earth!
Amanda Seyfried keeps it casual as she and a pal stop at Urth Caffe after a workout on Thursday (July 12) in West Hollywood, Calif.
The 26-year-old actress also stopped by a Fred Segal boutique, where she did a little shopping before grabbing lunch.
PHOTOS: Check out the latest pics of Amanda Seyfried
The next day, Amanda was spotted out for a walk with her adorable pet pooch Finn! She brought a bottle of water along to make sure both she and her pup stayed hydrated.
15+ pictures inside of Amanda Seyfried out keeping busy…


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Founded in 2008, ACP aims to ease the transition from the military to the civilian workforce. ACP is the only nonprofit organization engaged in national corporate career counseling for our returning military.
More than 65 of America's finest companies support ACP. We are also proud to work with a variety of best-in-class veteran service organizations, professional groups and military transition programs.
ACP's Citizens Mentoring Program enables individuals who are not employed by ACP's Partner Companies to participate as Mentors. Citizens Mentors support ACP veterans interested in diverse industries from Nonprofit to Public Service to Small Business.
The ACP Women's Veteran Mentoring Program is an initiative focused on connecting female veterans with entrepreneurs and female business leaders for one-on-one yearlong mentorships.
ACP has a bipartisan Advisory Council, comprised of prominent Americans from the military and business communities. We're grateful to these members for their help in building ACP's programs.
ACP has an independent Board of Directors who meet quarterly at ACP's New York City offices and provide strategic advice.
ACP is proud to report that our most recent Audited Financials show that 98% of every dollar goes directly toward our veteran programs.
ACP publishes an Annual Report to showcase the organization's impact on the veteran community.


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Welcome to Southwinds...your new home. Quiet and friendly community with 1 and 2 bedrooms conveniently located near Bellevue Offutt AFB. We offer everything you could possibly need including a full size washer and dryer in every apartment, a fully equipped kitchen with microwave, as well as walk in and extended closets. Easy access to restaurants and shops, and great schools for all ages nearby. Online payment and applications for your convenience. We welcome you and your pets here at Southwinds so make the move you will not regret!
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In times of transition, we all need a place to stay, a place to call home. Why hassle with buying, moving, setting up and then having to move, store, sell or discard furniture when you are done with it, when you can rent furniture from CORT? Cost-effective, convenient and hassle-free, furniture rental is the smart, flexible alternative for times of transition.
Whether you're a corporate commuter, student, stager, or serving in the military, furniture rental is an ideal solution. From several months to several years and everything in between, CORT has the lease term options you need for your stay.
Personalize your own space by choosing your furniture by the room or by the piece. With CORT's Move-In Ready packages, a trained professional will pick furniture to match your needs and budget.
Your prompt, professional delivery includes a complete furniture package setup down to the last pillow, lampshade, or spoon. CORT Furniture Rental provides a furnished life without compromise or commitment.
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Petitioning for an Appeal.
1000 signatures needed!
Nationals Member for Mildura Peter Crisp will be launching a petition today - protesting the sentence handed to 37-year-old Brandon Osborn.
Osborn was charged for the murder of Mildura Rural City Council worker Karen Belej - with the Crown accepting a guilty plea for manslaughter.
Mr Crisp believes the sentence was too lenient and doesn't give proper justice for Karen and her supporters.
But the clock is ticking with the judicial process only allowing 28 days for an appeal to be lodged - meaning locals have until September 18th to sign the petition.
If the petition is successful and the process is completed, it will be referred to as "Karen's Law".
Mr Crisp says violence of any form is a social issue that needs to be addressed.
For more information, or details on how to sign, click the link below:


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From Denmark:
What is a truly normal blood sugar profile for a non-diabetic healthy, non-obese individual? I know the numbers for the preprandial blood sugar (70-90 mg/dl [3.9-5.0 mmol/L]). What I would like to know is some more about the effects of the phase I insulin secretion. Is it true that a truly normal blood sugar profile rarely goes above 120 mg/dl [6.7 mmol/L] and returns to baseline within at most an hour after peaking?
Why are diabetics (missing the phase I response in most cases) advised to eat a "normal" (rice, bread,...) diet, when they almost surely reach the 180-200 postprandial level even with injected insulin? Are postprandial blood sugars in this range dangerous in the long term?
If possible it would be nice if you could provide some references to the medical literature concerning normal blood sugar profiles.
Published figures for normal fasting blood sugar levels do vary slightly with the laboratory depending on whether the report is for serum or whole blood, on the average delay before the sample is entered into the process and on idiosyncrasies in the instrumentation. In general though a figure of 70 to 95 mg/dl would be typical; but you'll find a slightly wider range if you look in Normal Regulation of Blood Glucose at the EndocrineWeb.com web site.
The normal response to a load depends of course on the nature of the carbohydrate; but in a standardised glucose tolerance test the peak would be somewhere between 30 and 60 minutes and the ultimate return to fasting levels nearer two hours after the peak. The assay is such a common one that there is really no literature on what is strictly normal and I think that your best reference figures for this would be from your nearest hospital laboratory which nowadays express standards in terms of 95% confidence limits or the mean +/- 2 standard deviations. You have to remember too that not all of these samples would have been normal.
For diabetics the overall efficacy of their blood sugar control and vulnerability to complications is usually expressed by the Hemoglobin A1c level; but increasingly and for subjects who are on a NovoLog or Humalog/Glargine regimen with the short acting insulin given immediately after the meal with the dose calibrated to the pre-meal blood sugar and the 'carbs' actually consumed it has become common, at least occasionally, to also check the two hour post prandial glucose level. If this is consistently above 200mg/dl then the insulin dose or the 'carbs' eaten can be adjusted in the expectation of even better control.
Original posting 29 Jan 2004
Posted to Blood Tests and Insulin Injections
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Last Updated: Tuesday April 06, 2010 15:09:54
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