text:
- Research article
- Open Access
Mitogenomic phylogenetic analyses of the Delphinidae with an emphasis on the Globicephalinae
© Vilstrup et al; licensee BioMed Central Ltd. 2011
- Received: 18 November 2010
- Accepted: 10 March 2011
- Published: 10 March 2011
Previous DNA-based phylogenetic studies of the Delphinidae family suggest it has undergone rapid diversification, as characterised by unresolved and poorly supported taxonomic relationships (polytomies) for some of the species within this group. Using an increased amount of sequence data we test between alternative hypotheses of soft polytomies caused by rapid speciation, slow evolutionary rate and/or insufficient sequence data, and hard polytomies caused by simultaneous speciation within this family. Combining the mitogenome sequences of five new and 12 previously published species within the Delphinidae, we used Bayesian and maximum-likelihood methods to estimate the phylogeny from partitioned and unpartitioned mitogenome sequences. Further ad hoc tests were then conducted to estimate the support for alternative topologies.
We found high support for all the relationships within our reconstructed phylogenies, and topologies were consistent between the Bayesian and maximum-likelihood trees inferred from partitioned and unpartitioned data. Resolved relationships included the placement of the killer whale (Orcinus orca) as sister taxon to the rest of the Globicephalinae subfamily, placement of the Risso's dolphin (Grampus griseus) within the Globicephalinae subfamily, removal of the white-beaked dolphin (Lagenorhynchus albirostris) from the Delphininae subfamily and the placement of the rough-toothed dolphin (Steno bredanensis) as sister taxon to the rest of the Delphininae subfamily rather than within the Globicephalinae subfamily. The additional testing of alternative topologies allowed us to reject all other putative relationships, with the exception that we were unable to reject the hypothesis that the relationship between L. albirostris and the Globicephalinae and Delphininae subfamilies was polytomic.
Despite their rapid diversification, the increased sequence data yielded by mitogenomes enables the resolution of a strongly supported, bifurcating phylogeny, and a chronology of the divergences within the Delphinidae family. This highlights the benefits and potential application of large mitogenome datasets to resolve long-standing phylogenetic uncertainties.
- Sister Taxon
- Killer Whale
- High Posterior Density
- Codon Site
- Bayesian Phylogenetic Analysis
The mitochondrial genome is typically non-recombining, has a relatively high substitution rate, and has a smaller effective population size than the nuclear genome [1]. These properties can increase the probability of congruence between the mitochondrial gene tree and the species tree, helping to resolve relationships between recently divergent taxa [2]. Under some conditions, however, phylogenetic analyses of mitochondrial DNA (mtDNA) sequence data can fail to resolve the relationships among taxa into a fully bifurcating tree. Theoretical and empirical studies suggest that greater phylogenetic resolution and bootstrap support for inter-specific nodes should be achievable by increasing the amount of sequence data [3–7]. Additionally, including sequence data from more than one gene will reduce the influence of any variation between genes in phylogenetic signal due to selection or the effects of stochastic lineage sorting [8].
However, not all polytomic relationships are 'soft', and some cannot be resolved by adding sequence data [9]. In some cases even complete mitochondrial genomes have failed to resolve multifurcating relationships (e.g., [10]). Although unlikely [11], a 'hard' molecular polytomy could be the result of a true simultaneous speciation event into multiple daughter species [9, 12–14]. Such events could occur when multiple populations of an ancestral species become simultaneously isolated during periods of rapid environmental change [12]. Simultaneous adaptive radiation into multiple species could also occur in sympatry due to intra-specific competition and assortative mating if species occupy a narrow niche width [15].
The use of high-throughput sequencing and pooled tagging methods [16] has the potential to generate large amounts of manifold-coverage sequence data for large numbers of samples quickly and at a relatively low cost, allowing improved resolution of phylogenetic relationships to be achieved routinely (e.g., [17]). Such an approach can allow the differentiation between rapid and simultaneous cladogenesis and, when combined with additional tests, can determine if a molecular polytomy reflects a true species polytomy (e.g., [18–20]).
Previous phylogenetic studies using a variety of markers have suggested an episodic rapid rate of speciation within the Delphinidae family [21–28]. This has resulted in several long-standing uncertainties in the phylogenetic relationships within this group. Analysis of sequences of the cytochrome b gene (1,140 bp) [21] led to a number of suggested taxonomic revisions from previous classifications based on morphology [29]. However, a number of these revisions had poor support and required further analysis, e.g., the inclusion of Grampus griseus (Risso's dolphin), and removal of Orcinus orca (killer whale) from the Globicephalinae subfamily, and the grouping of O. orca with the genus Orcaella into a proposed Orcininae subfamily. The genus Lagenorhynchus was found to be polyphyletic, with L. albirostris and L. acutus removed from the remaining four congeners and not found to be closely related to each other [21]. There was high support for paraphyly of the genera Tursiops and Stenella [21], which has been subsequently strengthened by mitogenomic and multi-locus analyses [25, 28]. More recent studies, which included both mtDNA and nuclear DNA (nuDNA), suggested an additional revision of the placement of Steno bredanensis (rough-toothed dolphin) with Orcaella [24, 26]; however, another multi-locus study grouped S. bredanensis with the Sotalia genus, consistent with previous classifications [27]. These more recent analyses using both nuDNA and mtDNA [24, 26, 27] have failed to confirm all of the revisions suggested by LeDuc et al. [21] and have not produced consistent estimates of relationships within this family. For example the ordering of the branches containing the species O. orca and L. albirostris, and the subfamilies Delphininae, Globicephilinae, and Lissodelphininae has differed considerably among studies [24, 26, 27]. In addition to being highly variable, the placement of L. albirostris has typically been one of the most weakly supported [21, 26, 27].
Here we estimate the phylogeny of Delphinidae using complete mitogenomes generated using high-throughput sequencing. This is the first time complete mitogenome sequences have been published for five of the species, and mitogenome sequences from a further three species have previously only been used as outgroup species for an intra-specific phylogenetic study on the killer whale [17]. In total, mitogenome sequences from 17 of the 37 extant species within Delphinidae were included in the analyses. This case study on a rapidly radiating group is one of the first to test the power of large sequence datasets produced by parallel-tagged high-throughput sequencing in combination with improved analytical techniques to address long-standing phylogenetic uncertainties. Specifically we test three revisions suggested by LeDuc et al. [21] regarding the placement of O. orca, G. griseus, and L. albirostris, and a further suggested revision regarding the placement of S. bredanensis [24]. We ultimately test the hypothesis that these uncertainties result from a true species polytomy caused by simultaneous speciation.
Results and Discussion
Samples used in study, including outgroup and Delphininae sequences from Genbank (accession numbers are given), and multiple specimens per species used in some analyses.
Common name
Genbank Acc. No.
Lagenorhynchus albirostris
White beaked dolphin
Ref. [46]
Feresa attenuata
Pygmy killer whale
This study
Feresa attenuata
Pygmy killer whale
This study
Peponocephala electra
Melon-headed whale
This study
Peponocephala electra
Melon-headed whale
This study
Globicephala macrorhynchus
Short-finned pilot whale
Ref. [17]
Globicephala macrorhynchus*
Short-finned pilot whale
This study
Globicephala macrorhynchus*
Short-finned pilot whale
This study
Globicephala macrorhynchus
Short-finned pilot whale
This study
Globicephala melas*
Long-finned pilot whale
This study
Globicephala melas
Long-finned pilot whale
Ref. [17]
Globicephala melas
Long-finned pilot whale
This study
Pseudorca crassidens
False killer whale
This study
Pseudorca crassidens
False killer whale
This study
Pseudorca crassidens
False killer whale
Ref. [17]
Grampus griseus
Risso's dolphin
Ref. [28]
Orcaella brevirostris
Irrawaddy dolphin
This study
Orcaella heinsohni*
Australian snubfin dolphin
This study
Orcaella heinsohni*
Australian snubfin dolphin
This study
Orcaella heinsohni*
Australian snubfin dolphin
This study
Orcaella heinsohni*
Australian snubfin dolphin
This study
Orcaella heinsohni
Australian snubfin dolphin
This study
Orcinus orca
Killer whale
Ref. [17]
Orcinus orca
Killer whale
Ref. [17]
Steno bredanensis*
Rough-toothed dolphin
This study
Stenella attenuata
Pantropical spotted dolphin
Ref. [28]
Sousa chinensis
Indopacific humpbacked dolphin
Ref. [28]
Tursiops truncatus
Common bottlenose dolphin
Ref. [28]
Stenella coeruleoalba
Striped dolphin
Ref. [28]
Delphinus capensis
Long-beaked common dolphin
Ref. [28]
Tursiops aduncus
Indian Ocean bottlenose dolphin
Ref. [28]
Monodon monoceros
Ref. [46]
Phocoena phocoena
Harbor porpoise
Ref. [46]
Inia geoffrensis
Amazon river dolphin
Ref. [46]
Lipotes vexillifer
Yangtze river dolphin
Ref. [47]
Phylogenetic analyses
The p-values of the approximately unbiased test and weighted or unweighted Kishino-Hasegawa and Shimodaira-Hasegawa tests are provided as well as the bootstrap and Bayesian posterior probabilities of the selected topology.
Our analyses strongly support the placement by LeDuc et al. [21] of Grampus griseus within the Globicephalinae subfamily, a result that now finds wide support from a range of different markers [24, 26, 27]. In contrast to LeDuc et al.'s [21] revisions we were able to reject the suggested grouping of Orcinus orca with Orcaella: the proposed Orcininae subfamily. Instead we find that O. orca is sister taxon to the rest of the Globicephalinae subfamily, which also contains the Orcaella genus. We were also able to reject the placement of Steno bredanensis in Globicephalinae as proposed by Caballero et al. [24] and supported by McGowen et al. [26]. Traditionally, and based on morphology alone, S. bredanensis has been placed in the subfamily Stenoninae with the genera Sousa and Sotalia [29]. Sousa chinensis has since been moved to the Delphininae subfamily and our analyses suggest that S. bredanensis is sister taxon to the rest of the species within this clade. Membership of these subfamilies appears to be supported by shared derived morphological characteristics, however, a thorough cladistic morphological analysis at the genus level remains lacking.
Consistent with geographical distribution, morphological differences, and a high number of fixed differences in the mtDNA control region [33], we found a deep phylogenetic divergence between Orcaella brevirostris and O. heinsohni (Figure 1). This divergence is dated to the Pliocene (5.3-2.6 Mya; Figure 1), a period characterised by elevated cyclical fluctuations in sea temperatures and sea levels with an overall trend of cooling temperatures and an increasing west-to-east sea surface temperature (SST) difference across the equatorial Pacific [34–37]. Our data also indicate a rapid radiation of the other extant species within the Globicephalinae subfamily during this period of climatic variation.
Caveats and recommended future work
The inclusion of nuDNA loci in some of the recent studies [24, 26, 27] may explain the incongruencies noted above. The mitogenome is a single, maternally inherited, haploid locus and might not wholly reflect the underlying pattern of population divergence and lineage formation (e.g., [38]). Nuclear genes can have greater resolving power for deep-level phylogenetic inference [39] and, when combined with whole mitogenomes, they have also been able to improve resolution in cases of apparent incomplete lineage sorting due to rapid radiations [40]. Historical hybridization is another potential source of incongruence between mtDNA and nuDNA trees [2, 12]. This is a particularly important factor to consider if the Biological Species Concept rather than the Phylogenetic Species Concept is being used to define species boundaries, as reproductive isolation is a key criterion for species status under the former [41].
A recent study based on the control regions of the two pilot whale species, Globicephala melas and G. macrorhynchus, found no support for reciprocal monophyly [42]. It was suggested that this could be due to incomplete lineage sorting or a lack of sufficient data. We found complete lineage sorting with high monophyletic support for all species for which we had multiple representatives, including the two pilot whale species (Figure 1). However, we had small sample numbers from a limited geographical range for all taxa. Including single or low numbers of representatives of each species assumes that the individuals sequenced and included in the phylogenetic analyses are representative of a reciprocally monophyletic clade [41]. However, increased sampling may reveal instances of paraphyly or polyphyly [41]. Increased sampling and mitogenome sequencing, such as conducted on the killer whale [17], is needed to confirm complete lineage sorting for the remaining species included in this study. Increased taxon sampling, such as the inclusion of the species within the Lissodelphininae, may also further reduce phylogenetic error [7, 43, 44].
We have used complete mtDNA genome sequences to resolve the phylogenetic relationships within the Delphinidae with high support. Future studies could apply similar methods to resolve para- and polyphyletic genera, e.g., the four Lagenorhynchus species within the subfamily Lissodelphininae [45]. Our study further supports previous empirical and theoretical studies [3–7], that increasing sequence data can improve phylogenetic resolution, even in species known to have undergone rapid radiation in the recent past, and can help to discriminate between hard and soft polytomies. However, some clades will remain unresolved, most likely due to simultaneous speciation events [12]. With current high-throughput sequencing techniques, animal mitogenomes can be sequenced relatively quickly and at low cost, and are an attractive candidate for future phylogenetic analyses, particularly if combined with current analytical tools that can aid in the reconstruction of challenging phylogenies.
Sample collection, DNA extraction, amplification and sequencing
Epidermal tissue was obtained by remote biopsying of free-ranging animals [46] and stored in 20% dimethyl sulphoxide (DMSO) saturated with salt [47].
Genomic DNA was extracted from epidermis using the Qiagen DNeasy (Qiagen DNeasy, Valencia, CA, USA) kit following the manufacturer's guidelines. The mitochondrial genome was amplified in 3-5 overlapping amplicons (dependent on DNA preservation) using previously published long-range PCR primers [17] and primers designed specifically for this study (see Additional file 3). Additional sequencing primers were also designed for gap filling using conventional Sanger sequencing at the commercial service offered by Macrogen (Seoul, South Korea). Each 25 μl PCR contained 1 μl extracted DNA, 1× PCR buffer, 2 mM MgSO4, 0.4 μM of each primer, 100 nM mixed dNTPs and 0.1 μl High Fidelity Platinum Taq (Invitrogen, Carlsbad, California). PCR amplifications were performed using an MJ Thermocycler with a 4 minute activation step at 94°C, followed by 35 cycles of 94°C for 30 seconds, 62°C for 30 seconds, 68°C for 6 minutes 30 seconds, followed by a final extension period of 72°C for 7 minutes. For amplifications under 6 kb in size the extension time was reduced to 3-5 minutes.
The amplified PCR products were purified using an Invitek PCRapace purification kit (PCRapace, Invitek, Berlin, Germany) and quantified using a NanoDrop spectrophotometer (NanoDrop Products, Wilmington, DE) to determine DNA concentration used to balance and pool amplicons in equimolar ratios. Length of fragment, ratio of fragment lengths per individual, and DNA concentration was taken into account when balancing the samples. Samples were either individually tagged according to Meyer et al. [16] and built into shotgun sequencing libraries following the manufacturer's instructions (454 Life Sciences, Branford, CT), or grouped into sets of 8-10, where within each sample set individual libraries were made to contain a different 10 bp multiplexing identifier (MID) tag, allowing libraries to be combined prior to emulsion PCR. Sequencing libraries were quantified by qPCR [16] and pooled at equimolar concentrations. Library pools were divided among regions on GS FLX sequencing runs, using either LR70 or Titanium chemistry (454 Life Sciences). Sequencing data was parsed into individual extractions and identifier tags were removed using a custom tag-removal Perl script (M. Rasmussen, unpublished, University of Copenhagen).
Phylogenetic analyses
Sequences were assembled using gsMapper (Roche Applied Science, Indianapolis, IN, USA) and aligned by eye using SE-AL v2.0a11 (A. Rambaut, University of Oxford), while Geneious (Biomatters Ltd., Auckland, New Zealand) and Sequencher v4.8 (Genes Code Corporation, Ann Arbor, MI) were used to check coverage and sequence reliability. Conspicuous indels, base position differences, and differences in homopolymeric regions were double-checked and sequences with higher coverage were generally given preference. In order to ensure that the data was not affected by the erroneous incorporation of nuclear pseudogenes (numts) we visually assessed the recovered sequences for the presence of stop codons or frame-shift mutations in the aligned protein-coding genes. We observed no evidence that numts might be present in the data. This may be explained by a combination of (a) the general difficulty with PCR amplifying long amplicons, requiring relatively high levels of template for successful amplication, and thus (b) the fact that mtDNA template copy numbers are much higher than those for nuDNA templates, leading to preferential mtDNA over nuDNA amplification.
A total of 35 mitogenome sequences were used in the analyses, representing 21 species (Table 1). Of these, 18 mitogenomes were amplified and sequenced for this study and 17 mitogenomes attained from Genbank [17, 30, 48, 49], which included 4 outgroup sequences (narwhal, Monodon monoceros; harbor porpoise, Phocoena phocoena; Yangtze river dolphin, Lipotes vexillifer; Amazon river dolphin, Inia geoffrensis). The 18 generated sequences consist of 5 species (pygmy killer whale, Feresa attenuata; melon-headed whale, Peponocephala electra; Irrawaddy dolphin, Orcaella brevirostris; Australian snubfin dolphin, Orcaella heinsohni; rough-toothed dolphin, Steno bredanensis) whose mitogenomes had not been previously sequenced prior to this study.
A single representative mitogenome from each of the 21 species was used for the initial Bayesian phylogenetic analysis and divergence date estimation. This step was taken so that a speciation prior could be used for the tree topology and node times. Sequences were aligned and the 2 rRNA and 12 protein-encoding genes (excluding ND6) were used to form a data set comprising 13,958 sites. Stop codons were removed from all genes and the control region was excluded from analysis due to saturation, repetitive sequences, and alignment ambiguities. The resulting alignment was divided into four partitions: first, second and third codon sites of the protein-coding genes (3,792 bp per partition), and rRNA genes (2,582 bp). A comparison of Bayesian information criterion values in Modelgenerator [50] were used to find the optimal time-reversible substitution model for each partition. This criterion has been found to perform well in relation to other criteria used in evolutionary model selection [51]. The selected models were GTR+I+G for first and third codon sites, HKY+I+G for second codon sites, and TN93+I+G for the rRNA partition. In all cases, rate variation among sites was modelled using a gamma distribution with six categories [52]. There was little variation in the base frequencies across taxa (see Additional file 4).
The Bayesian phylogenetic analysis was performed using BEAST v1.6 [53]. An uncorrelated lognormal relaxed-clock model was used to allow rate variation among branches [54]. A Bayes-factor analysis indicated that this model received decisive support in comparison to a strict-clock model. The four data partitions shared the same relaxed clock but were allowed to have different relative rates. An exponential prior with a mean of 1/3 was used for the standard deviation of the lognormal distribution of rates, and a Yule prior was specified for the tree topology and relative divergence times. To enable the estimation of absolute divergence times in the tree, four calibrations based primarily on fossil calibrations, along with estimated divergence dates from published studies [27, 28, 55] (see Additional file 5), were incorporated into the analysis. The calibrations were implemented in the form of uniform prior distributions for the ages of the four nodes, and monophyly was enforced on the clades defined by these four nodes.
Posterior distributions of parameters, including the tree topology and divergence times, were estimated by Markov chain Monte Carlo (MCMC) sampling. Samples were drawn every 5,000 MCMC steps over a total of 50,000,000 steps. The first 10% of samples were discarded as burn-in. Convergence to the stationary distribution and acceptable mixing were investigated using the diagnostic software Tracer v1.5 (Rambaut and Drummond, 2007, University of Oxford). From the set of posterior samples, the tree with the highest product of clade credibilities was identified and the branch lengths were rescaled to match mean posterior estimates.
Additional phylogenetic analyses were performed in order to examine the effect of data partitioning. First, the analysis was repeated without data partitioning, so that the protein-coding genes and the rRNA genes shared the same substitution model and mean evolutionary rate. Second, analyses were performed on three data sets in which sites were randomly assigned to the four data partitions mentioned earlier. Randomisation of sites (sampling without replacement) was performed using the Java application SiteSampler v1.1 [56] Support for the different partitioning schemes was examined by assessing Bayes factors, calculated using a harmonic-mean estimator in the software Tracer v1.6 [57, 58].
A second set of phylogenetic analyses was performed on the full dataset (35 mitogenomes), including multiple representatives per species, coding missing data as N and using data partitioning for each codon position, rRNA, and control region. For each partition, the best model of molecular evolution that was compatible with models implemented in MrBayes 3.12 [59] was selected using the Bayesian information criterion. The selected models were TN93+I+G for the first and second codon sites, HKY+I+G for third codon sites and the control region, and HKY+I+G for the rRNA genes. Similar models were recovered after RY-recoding a hypervariable part of the control region (nucleotides 15556-15588 according to the Globicephala macrorhynchus, Genbank accession number HM060334, reference mitogenome). Bayesian phylogenetic analysis was performed using the 5-partition datasets (with and without RY-recoding of the control region) using MrBayes 3.12 [59]. Posterior distributions of parameters were estimated using two independent MCMC analyses, each comprising one cold and three heated chains. Samples from the posterior were drawn every 1,000 steps over a total of 10 million steps, which appeared to be sufficient to keep the average standard deviation of split frequencies below the critical value of 0.01. The first 25% of samples were discarded as burn-in. A majority-rule consensus tree was constructed from the posterior sample of trees. A supplemental set of analyses was performed with MrBayes 3.12 and PhyML 3.0 [60] using an unpartitioned dataset under a GTR+I+G model as selected using ModelTest [61] and Akaike Information Criterion, with and without RY-recoding of the hypervariable segment of the control region. All analyses yielded identical tree topologies and similar node support values. For maximum-likelihood analyses, the strength of the phylogenetic signal was assessed via non-parametric bootstrapping with 250 pseudo-replicates. In addition, using Consel 0.1 k [62] and site-wise likelihood values recovered from PhyML analyses, levels of statistical support for alternative topologies were evaluated from the p-values of approximately unbiased tests, weighted or unweighted unilateral Kishino-Hasegawa and Shimodeira-Hasegawa tests, and bootstrap and Bayesian posterior probabilities for the selected topologies (Table 2). All trees were drawn using Dendroscope [63].
The authors thank the Danish Council for Independent Research: Natural Sciences, and the Danish National Research Foundation and the Systematics Research Fund from the Linnean Society for generous funding. We thank Bill Perrin, Sebastián Duchene Garzon, and Eric Archer, four anonymous reviewers, and the editors for critical input and comments on the manuscript. We also thank Morten Rasmussen for bioinformatics support, and we thank Uko Gorter for whale illustrations.
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This article is published under license to BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.


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Frequently Asked Questionsfor the 5 Day Detox
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subset: commoncrawl


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Oration honouring Patrick O'Flaherty
There is a sense, no slight sense, in which Patrick O'Flaherty's life track follows that of his great academic subject, Dr. Johnson. So the public orator must, for these brief moments and on this great stage, be his Boswell.
Johnson came out of a small community to university, took up teaching, took up journalism, took up writing, took up criticism and biography, and wrote his masterwork, The Dictionary of the English Language.
O'Flaherty came into St. John's from Long Beach, a small place of settlement adjacent to Northern Bay, first to school and then on to university. Unlike his illustrious predecessor who did not, O'Flaherty took his degree with great distinction and extraordinary rapidity: BA at 20, MA at 21, PhD at 24.
Like Johnson he too became a teacher, first at the University of Manitoba, then at Memorial and, as his students will attest, taught with great power, skill and learning. For 30 years he was a regular on CBC, as well as in the local and national press. He has published two novels and two collections of short fiction.
He started his writing career as a critic of the work of Dr. Johnson and then moved on to the literary history of Newfoundland and Labrador. But his masterwork is the three-volume history of Newfoundland of which the final volume has just arrived from the printers for a launch next week.
Now that is only a citation of achievements and does not really get at the substance of his contribution to this our university, to this our country.
Permit a dilation on the real matter of the man. Patrick O'Flaherty has undergone several transformations in the course of his career and been remarkably influential in the development of Newfoundland studies in its broadest form. Starting out as a literary critic (of Samuel Johnson), he became a literary historian (of Newfoundland) and then a historian (also of Newfoundland). In each of these stages of his career he distinguished himself by the quality of his analysis and of his prose. But he saw his role in the academy as more than that of a first-rate scholar for he challenged his students, developed new programs, served as editor to his colleagues, and found an audience beyond the university in his journalism.
His pioneering study of Newfoundland writing, The Rock Observed opened a range of opportunity not just in terms of texts to be read but of studies to be done. O'Flaherty developed the first undergraduate and graduate courses in Newfoundland literature and steadily students began to work in the field, producing dissertations and articles.
This, in its turn, fed the local writers and can be seen as having stimulated the current lively literary scene.
His next scholarly enterprise was to propose the creation of a journal, Newfoundland Studies, to showcase the new scholarship related to work on this place. He not only founded the journal but also served as one of its editors until his retirement. Determined that it would cover all aspects and would be accessible to the general reader, he had set himself a difficult task. That the journal is now firmly established and well-regarded both inside and outside the academic world says much for his academic initiative, the standards he set and the example he provided.
What is unquestionably his major work is his history of Newfoundland. Despite remarkable and groundbreaking work by historians in the 20th century none of them had taken on the task of writing a general history perhaps because Prowse's monumental work remained, for over a century, the definitive study.
O'Flaherty not only immersed himself in all the recent scholarship but also revisited the primary sources. He took the work of his colleagues in history and geography, economics and folklore and, while treating that work with critical respect, was able, because of the depth of his own research, to challenge some of their findings.
Among other things, this resulted in a re-assessment of William Coaker – regarded by many Newfoundlanders as something of a secular saint – and of A.B. Morine – regarded by those same Newfoundlanders as an unregenerate rogue.
Heresy, but then this was the man who, earlier, had questioned the Newfoundland credentials of our greatest poet, E. J. Pratt.
Historian, literary critic, editor, public scholar, novelist, journalist, Patrick O'Flaherty has taken his learning from the campus to the country; he has had a major impact on the development of Newfoundland and Labrador studies and, in all his writing, displayed a remarkable capacity for scholarship and for the presentation of that scholarship in superb and striking prose.
Recognized by the arts community in the Newfoundland Arts Hall of Honour, recognized nationally by the Order of Canada, vice-chancellor, I ask the university to recognize Patrick O'Flaherty by conferring upon him the degree of doctor of letters, honoris causa.
Shane O'Dea
Public orator


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Mary Nuttall
Growing up in Sandy, Utah, Mary Nuttall at the age of ten began a serious study of ballet at Jacqueline's School of Ballet. Mary attended Regional Dance America where she was an honor student and was awarded scholarships to RDA's Choreography Conference five times and the International Ballet Competition's School twice. She was a selected participant for Prix de Lausanne and a finalist at Youth America Grand Prix where she was awarded scholarships to John Cranko School in Stuttgart Germany and National Ballet of Canada's Ballet School in Toronto, Canada. She attended the John Cranko School as well as Netherlands Dance Theater training programs. Mary joined Utah Regional Ballet in 2009. She has toured Europe with URB and has performed many roles including White Swan, Sugar Plum Fairy and Ucanogos. Mary's Choreography has been performed at Regional Dance America, Utah Valley University and in Utah Regional Ballet's Choreography Design Concert where she was awarded first place in the professional division. Mary has her Bachelor of Fine Arts in Dance with an Emphasis in Ballet from Utah Valley University and has been teaching at Jacqueline's School of Ballet for four years.


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|Policy status:
|In Force
|Date Effective:
|Date Amended:
1994; 2003; 2006
|Policy Type:
|Economic Instruments>Fiscal/financial incentives>Grants and subsidies, Regulatory Instruments>Auditing
|Policy Target:
|Industry>Industrial processes, Transport, Buildings>Building Type>Industrial, Buildings>Building Type>Residential, Buildings>Building Type>Non-residential
|Ministry of Employment and the Economy
|EAP is funded through the Energy Aid Scheme. In the period 1992-2009 the total amount of financial support provided was EUR 26.5M, while the total cost of audits reached EUR 60.6M. In 2008-2011 subsidies for energy auditing were about EUR 8M.
The purpose of an energy audit is to analyse the energy use of the facility being audited, to work out the potential for energy savings and present a profitability calculation of saving proposals. Energy audits are also one important element in the implementation of energy efficiency agreements to figure out energy savings measures and energy savings potentials.
Energy audit subsidies for service buildings and of industrial and energy production facilities have been granted since 1992. The actual energy audit programme started in 1994. The Ministry of Employment and the Economy (MEE) provides subsidies for carrying out the energy auditing of the commercial and public buildings and industrial and energy sectors (since 2014 according to the restrictions based on EED Article 8 mandatory energy audits for large companies). The MEE also supports municipalities to carry out audits concerning promotion of renewable energy use in the municipality district (Renewable Energy Municipal Audit).
Audit programme includes promotion of audit activities, development and monitoring, the training and qualification of energy auditors, and the quality assurance of audits. The results of energy audit activities have been monitored through a separate monitoring system since 1994.
|Related policies:
|Energy Aid Scheme
Last modified: Wed, 17 Jun 2015 15:32:27 CEST


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hanbleceya Section
Hanbleceya and its leaders fined for investment law violations
by Jack Mayne Two people involved in Hanbleceya, the now defunct Normandy Park mental health treatment and housing program, and the ownership corporation have been fined assessed costs totaling $18,500... [Read more]
After two years, Hanbleceya will be leaving Normandy Park by July 12
Kerry Paulson of Hanbleceya. by Jack Mayne Kerry Paulson, managing partner of adult care home Hanbleceya says July 12 will be the organization's last day in Normandy Park. Hanbleceya bills itself as... [Read more]
No license needed for Hanbleceya's mental health homes in Normandy Park
by Jack Mayne Hanbleceya is operating four of five facilities in Normandy Park without licenses because there are no health care licenses for their facilities either from the State of Washington or from... [Read more]
'Bigots of Normandy Park' Website Taken Down After 8 Days
by Jack Mayne A website aimed at discrediting the Normandy Park Cares organization – which has been opposed to Hanbleceya, a for-profit mental health business operating illegally according to the DSHS... [Read more]
Hanbleceya Defends Normandy Park Group Homes; Residents Seek Regulation
A Hanbleceya home in Normandy Park as seen in a car's side rear-view mirror on Tuesday, July 31, 2012. Photo by Scott Schaefer. by Jack Mayne An organization that treats people suffering from chronic... [Read more]
'Group Home Task Force' Meets to Discuss Hanbleceya Concerns
A task force consisting of Normandy Park residents, elected officials, mental health service providers and law enforcement held its first meeting on May 30th at Normandy Park City Hall to discuss concerns... [Read more]


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Good morning Lovelies! I can't believe we are in February already! Roll on the spring!
I'm on me jolly's at the moment and hoping this has posted on time, I scheduled it before I went so fingers crossed? I'm in Malaysia!
So onto my post! Here are my samples for Show us your Twinchies
My first is Centura pearl card, I used adirondack pink sherbet ink through a heart and swirl mask by Splodgaway, the stamp is a freebie from a mag!
My second is Linen card, I stamped with Versamark using a Stamp-em-up stamp set I swiped over with pan pastels, I used the 2 smallest heart Dies and decoupaged them for some dimension! I also lightly inked all the edges with Black soot DI!


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I recall exactly when we started telling our eldest DI conceived child about his conception. He was four years old and we were on holiday in France. He crept into our bed one morning and started asking about how babies grow and where they come from. I took a very deep breath and launched in on a semi-prepared speech about babies being made with a sperm from a daddy and an egg from a mummy, but that sometimes daddies needed to borrow sperm from another man. The term 'borrow' seems very odd today but we had taken this language from the only book available at that time to help parents with the task of sharing donor conception information, Robert Snowden's The Gift of a Child, first published in 1984.
From that day on, regularly and when an appropriate 'hook' to hang information on occurred, we would repeat and elaborate on the story of how we added to our family (I already had a son from my first marriage). Will's younger sister Zannah picked up the information by association and apparent osmosis, although she did need some clarifications as she clearly thought that I had met 'the nice man' Daddy had borrowed some sperm from. But it wasn't really until the book My Story arrived in our house when the children were 8 and 5 that the penny dropped and Will started asking many more questions. When I later studied child development I understood that the brain makes a developmental leap at this sort of age, allowing a child to understand concepts and information in a more sophisticated way, so it was not surprising that donor conception had not been fully understood until this point. What was clearly true for both children, and we checked this out with them later, was that neither recalled ever having been 'told' about how they came into the family. They accepted the information as simply part of their story, and have continued to do so without problem.
When the five families who started DC Network (then DI Network) came together in 1992/3, it was clear from talking to the others that they had shared the instinct that starting to 'tell' early was going to be most helpful for their children. Subsequent exploration showed that this was supported by research with adoptive families. DC Network has never had any reason to doubt that telling as a process, started under age 5, by parents who are comfortable and confident with their decisions, is the way that children integrate the information most easily. But some parents get stuck. The reasons for this are rarely completely clear but can be rooted in some sense of shame and stigma from one parent or the other and/or a deep fear of rejection by an adored, often only child. Their intentions are often very good but years pass, the child becomes 7, 8 or 9 and parents, often full of anxiety and aware of the recommended starting age of under 5, wonder if they have now left it too late.
To my mind there is no such thing as too late. Starting under five is ideal. Parents get used to the language in stages and children don't remember a time when they didn't know. At seven, eight or above, particularly if they have never talked with anyone about DC, parents may be rusty and awkward in their style and language, BUT good preparation including practice with what they are going to say and strong motivation in knowing this is the right thing to do, can overcome a lot. DC Network is always happy to give personal support and guidance to parents who feel they could do with some help or to refer to counsellors or other well-informed support organisations.
Even telling for the first time in early teenage years can be done well if this is the time that parents have a strong incentive or motivation to do it, prepare themselves well and recognise that they may have to go through some rocky moments. Developmentally it's not a great time. Young people are beginning to separate from parents, identify much more strongly with peers and generally question everything. Having what can feel like having the rug pulled from under their feet…because of course they need parents to be strong and solid to push against at this time…is not ideal. It can be done, but best before adolescence kicks in…or possibly leave till late teenage years, but this is more controversial.
And what about much later on? Over the last couple of years Walter and I have been having an increasing number of peer support sessions with couples who, in their middle age, have come to the conclusion that their DC children should be told about their beginnings. Without exception they are thoughtful people, often very successful in their own careers and with children between about 18 and 30 whom they love dearly. There are differences in each family's story but what is true for them all is a realisation of how wrong secrecy about donor conception has been – they were all told by their clinics not to tell – and a wish to respect their children by being open with them now. The fear they all share is that of turmoil, change in the often extremely good relationships they have and ultimately rejection. Again, good preparation is the key. They cannot prepare their children for the revelation but they can prepare themselves. The principles of this preparation are given in the Telling and Talking booklet 17+ (available from DC Network as a download or printed booklet) but all the couples we have seen have enjoyed the reassurance and support of talking about their dilemma face to face with people who are also DC parents, albeit those who defied the advice and told early. Walter and I have admired their willingness to re-visit old losses and sorrows (an important part of the preparation) and put themselves through a deeply uncomfortable time for the benefit of their adult children. It takes courage to do this.
So the answer to the question in the title, 'Does it matter when you tell' is both yes and no. Starting early means less difficulty and discomfort for both parents and children. Getting the first words out, at whatever age, is rarely easy but once started there is always a hook to hang further information on and children don't remember the early stuff anyway. Leaving it till later comes with some risks but, at the risk of boring you, preparation is definitely the way to go and that's where all four of the Telling and Talking booklets come in. Not that just reading a book can make everything alright. It can't…but they are designed to give confidence to parents and provide the structural bones and language to help parents do what feels right in their family. And for those in the UK there are Preparation for DC Parenthood and Telling and Talking workshops held regularly by DC Network.
Does Will remember that first conversation in France? I know he doesn't but we certainly do!


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17 Sep 2016
16 Aug 2016
11 Sep 2013
15 Dec 2011
I am actress, still trying to make my foothold strong in Bollywood. I had made a debut in Bollywood in 2004 but that went unnoticed and I moved on to south industry.
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List of Sabyasachi Chakraborty movies
Sabyasachi Chakraborty is one of the most talented actors of Bengal whose acting prowess is well known all over India.Sabyasachi was born on 8th September, 1956 and his parents Jagadish Chandra Chakraborty and Monica Chakraborty used to call him by the name Benu. In 1975, he passed Higher Secondary Examination from Delhi's Andrews College. He also completed his B.Sc from the famous Hansraj College of Delhi. After this , he cleared AMI examination from Delhi in 1978. Apart from acting , his love for nature and forests are well known. He is a very passionate about wild life photography. In 1986 , he got married and has two sons.He made his debut in Tapan Sinha'a film " Antardhan" in 1992. After this, he has worked in several famous television serials, most ...By Joy» read full biography
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Sabyasachi was in consideration for a role in Lagaan at the time director Ashutosh Gowarikar was in the process of casting, but finally it didnt materialize.By Making things happen :)
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This is for the third time that Parambrata Chattopadhyay is in the role of Topshe in a Feluda thriller within a span of four years. The first time we saw him as Topshe in Bombayer Bombete, and then in Kailashe Kelenkari and now the third time in Tintorettor Jishu. The Satyajit Ray story centers on a painting by famous Italian painter of the 16th century. The painting was of a Jesus by Tintoretto. It was theft and smuggled by an art dealer, And to het the painting back Feluda (Sabyasachi Chakraborty) and Topshe (Parambrata Chatterjee) along with Jatayu (Bibhu Bhattacharya) goes to Hong Kong.
Q. Sabyasachi Chakraborty is our Feluda these days. He has done one telefilm (Bakshya Rahashya) and three feature films (Bombayer Bombete, Kailashe Kelenkari, Tintorettor Jishu) all directed by Sandip Ray. A. In Bakshya Rahashya Feluda, alias Prodosh Chandra Mitter was a cerebral sleuth. But when Sandip Ray first brought gen-next Feluda in celluloid he made his private investigator more action oriented than a thoughtful detective. So, Sabyasachis portrayal of Feluda is Bombaier Bombete become the target of comparison with the earlier actor Soumitra Chatterjee.
Parthasarathi Joarddar
Paran Bandyopadhyay
Biplab Chattopadhyay
Soumitra Chatterjee
Kharaj Mukherjee
Raja Sen
Sandip Ray
Shantilal Mukhopadhyay
Rajesh Sharma
Prabhat Roy
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LG Announces Quick Voice – Apple Siri & Samsung S-Voice Rival Introduced to Optimus Smartphones
After revealing it will not be joining the tablet race this year, leaving the market to the ASUS / Google alliance and Microsoft and its partners to enjoy the spoils, part of Korean tech maker LG's focus on smartphones will include the introduction of a Siri-a-like voice assistant software called Quick Voice.
The Quick Voice speech app is said to work much the same as its rivals – say a command or ask a question and like a little pocket Jeeves, Quick Voice will track down an answer from the web and speak the answer, hopefully without much delay (iPhone 4S… *cough*). There will also be the usual talk to text for messaging and app launching and you'll even be able to use "in app" searches such as hitting up YouTube to find that "Call Me Maybe" video all the kids are talking about.
The software is set to go live with LG smartphones such as the Optimus VU and LTE II in the States but with Apple's Siri and Samsung's S-Voice already giving it verbal worldwide we imagine a widespread roll out will reach other markets soon enough and we'd imagine the Quick Voice app will surface on some familiar gadgets such as the recently released Optimus 4X HD.
Let us know your thoughts on our comments below or via our @Gadget_Helpline Twitter page or Official Facebook group.
Tags: Android, Apple, Galaxy SIII, iPhone, LG, Optimus, Quick Voice, S Voice, samsung, Siri, smartphone, Tablets, Voice


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Dental health has improved dramatically with falling rates of tooth decay and attention has now shifted to the needs of an ageing population, with an increased emphasis upon aesthetics – that is, having a full set of sparkling white teeth. Without proper protection, exposure to these may risk the health of the dental technologist.
Dental lab sandblasters are used to increase bond strength by divesting the casting investment and increasing effective surface area as well as polishing the surface of castings. Sandblasters that are used in dental laboratories are enclosed and often operate through a vacuum or via dental compressor. Dental laboratory sandblasters are operated via foot pedal and often feature gloves to work through, this keeps everything contained. Sandblasting media can be Aluminum Oxide, glass beads or Silicone Carbide grit. Be sure to choose a dental laboratory sandblaster that employs proper dust retraction.
The processes involved in building any kind of dental prosthesis have a number of stages where there is a potential health hazard to the dental technologist. First, the dentist makes a mould of the patient's mouth or teeth which the dental technologist uses to make a plaster model. This is used to make up a metal framework for the prosthesis onto which porcelain is overlaid, using a sandblasting technique. After baking the metal-porcelain layers, there is grinding to achieve the required shape.
This series of operations encompasses not just exposure to chemical hazards, which will be discussed elsewhere, but also the generation of dust particles of various compositions and sizes. Research has shown us that the dust from the types of materials that are used in dental technology contain between 54-70 per cent of respirable particles (that is, particles of less than 5 microns in size that enter the lungs). A particular concern is the silica content of this dust, which can reach 30 per cent and may exceed maximum recommended levels during the sandblasting and grinding processes.
Another is exposure to dusts from heavy metals like the cobalt-chromium-molybdenum alloys. Both exposures may lead to a lung condition known as pneumoconiosis, of which several cases have been reported as being linked to dental technology.
There have also been a number of cases of silicosis, a well-known and very serious occupational lung disease that is cause by inhaling respirable crystalline silica dust. These have been highlighted in the United States Center for Disease Control's publication Morbidity and Mortality Weekly Reports (MMWR). Traditionally, silicosis is associated with the mining, quarrying and ceramics industries.


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It's that time of the month again where every beauty blogger makes a list of beauty products they've been loving recently and who am I not to join in? It may be 2nd May but I always say better late than never! If I was to write a list of everything I've been using a lot or really enjoying, it would be a very long list! So I'm going to share with you the products I have actually FINISHED, because you all know as a beauty blogger how hard it is to finish a product before you move onto the next one to try ;) So here goes...
Nivea Daily Essentials Refreshing Facial Cleansing Wipes; Nivea Pure and Natural Lip Balm; Montagne Jeunesse Mud Pac Dead Sea
I recently review a liquid make-up remover, but that doesn't mean I don't use make-up wipes. The Nivea Daily Essentials Refreshing Facial Cleansing Wipes were recently on offer where I work, and I'm never one to spend much on face wipes (it seems pretty pointless to me, they just go in the bin!), so I thought I'd try them out. I was very impressed. One side of the wipe is a smooth surface, whilst the other side has little bumps. This is really great as the wipe is slightly exfoliating and really works well at getting all your make-up off!
Another Nivea product I can't get enough of is the Nivea Pure and Natural Lip Balm. Made with milk and honey, this product is extremely moisturising to your lips and is really smooth to apply. I prefer a lip balm in the form of a stick as I find it easier to apply and is quick and simple. I find this one quite hard to get hold of sometimes but when I see it, I stock up.
I've always enjoyed taking some time out to pamper myself, and if you like to do this too then you need to try Montagne Jeunesse Mud Pac. This face mask is an ultra deep pore cleanser, which is exactly what I need as I'm quite prone to blackheads :( I use this once a week and I often find I can get two applications out of one of these little packs!
There are so many great things I can say about MAC Studio Moisture Fix. As I have dry skin I find that my make-up can look really cakey when I apply my powder. After a couple of squirts of this the cakiness has gone! Magic. It is super moisturising, very lightweight and absorbs into your skin quickly. And it even has SPF 15 in it, what more could you want?
Again going down the dry skin route, I need to moisturise my body after every shower. Garnier Intensive 7 Day Nody Lotion for Extra Dry Skin is great and is really god on all those extra dry areas like your elbows and ankles. I do find that it can take quite a while to absorb into my skin but I don't mind that as when it has, the results are fantastic.
I bought the Laura Mercier Secret Concealer on a whim on a shopping trip with my cousin before christmas. it was just by chance that the assistant tried it on me and I was amazed that it actually managed to cover the bags under my eyes, which is a pretty hard task for any concealer as they are quite big! It is definitely not light weight, it actually applies quite thick so if your looking for something light then this isn't for you. But if you want a product that covers your dark eyes then you should give this a try.
MAC Prep + Prime Skin is also another little wonder product. I must admit I am hopeless with remembering to use primer, but after using this one I find it much easier to apply my foundation. It gives me a smoother surface to work on and I also find it evens out the redness in my skin. It's a little pricey, but you need such a small amount, mines lasted me forever!
And last but by no means least, it's the every popular Benefit they're real Mascara. I tried the Bad Gal mascara but didn't think much of it, but this one is a whole other kettle of fish. I love the rubber wand, I find them much easier to work with, they give volume and cover the entire lash. I'm actually quite sad that mine has run out, but I did just get a free sample in this months Elle magazine :)
What did you use up last month?


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Guy who wants to receive a*** from a woman
Something I have been looking for for ages. I want a woman to pound my ass with a strapon.
I had my fingers in my a****** today as I wanked off. I'm happy to be shared by a dominating couple. The woman can f*** me and then only if she wants her guy can f*** me too.
Bend me over, spank me. Treat me rough, slap my b**** so I know you're in charge and then insert your strapon or d*** into my ass and ram it n hard, f*** me rough. The other one can come round and I'll lick her p**** or suck on her strapon or his c***.
Right now I need to be g********* and spit roasted by a bunch of girls ' guys!!!


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October is the treasurer of the year,
And all the months pay bounty to her store:
The fields and orchards still their tribute bear,
And fill her brimming coffers more and more.
But she, with youthful lavishness,
Spends all her wealth in gaudy dress,
And decks herself in garments bold
Of scarlet, purple, red, and gold.
...She lives her life out joyously,
Nor cares when Frost stalks o'er her way
And turns her auburn locks to gray.
Paul Laurence Dunbar
I began by reading the group The Iroqouis Legend of the Corn Husk Doll which is a really neat little story.Then we set to work!
There's a really simple how-to in this book. They're fairly easy to make, and they take very little materials, as well as cost effective (as long as you have a fields of dried corn right out your back door like I do. Though I will warn you, be prepared to get muddy!)
I'll be adding my little gathering lady to our fall nature table at home :)
Have you ever made them?


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Click here to edit subtitle
Official 2014 Shirt
Car Show Details
Contact Us
...The West has a large event, the East has a large event, but the Midwest has not seen a large Subaru Enthusiast gathering like this.....until now...
What is Scoobapalooza?
Simply put, Scoobapalooza is the new Midwest SUBARU Enthusiast Event. Bringing together Subaru owners from Kansas, Iowa, Illinois, Missouri, Arkansas, Indiana, and any other surrounding areas.
So what can I expect out of this Scoobapalooza?
For 2014, we are excited to announce the return of the Meet, Greet, & Show. Along with the main gathering, the All Subaru Car Show will of course be happening as well. Also, we are really stoked to bring you the exclusive Scoobapalooza Autocross, sponsored by RacingLine USA.
What's the Schedule for Saturday?
8:00AM-9:30AM - The gates will be open for Scoobapalooza Autocross Participants
9:00AM - The gates will be open for all Scoobapalooza participants
9:00AM-12:00PM - Car Show Registration will be available
9:00AM-2:00PM - Raffles will be available to enter
2:30PM - Awards will begin
4:00PM - 2014 Scoobapalooza Closes and everyone must be out of the Family Arena Parking Lot
How do I get registered to participate in the All Subaru Car Show?
Pre registration for the All Subaru Car Show is available NOW! be sure to check out the car show details
Can I be involved in both the All Subaru Car Show and Scoobapalooza Autocross?
For the 2014 Scoobapalooza Event, we will be able to offer a limited amount of entries for BOTH the All Subaru Car Show and Scoobapalooza Autocross. If you do have an interest in participating in BOTH of the events, you will FIRST need to contact us at
[email protected]
so that we can be sure to gather your information and assist you with proper preregistration.
Are there any discounted accommodations for 2014?
You bet! Check out the
page for all of the details on the discounted rooms for Scoobapalooza participants!
What's this I hear about a t-shirt?
Check out the
Official Shirt page
for all the details!
There will be a limited amount of shirts available during the Meet, Greet, & Show and they will be at a higher costs than the preorder.
Shirts will
be shipped out to participants this year
Will there be refunds if it rains?
No refunds. Scoobapalooza is a rain or shine event.
Can I bring my own items to sell at this event?
Yes, but
if you are a registered vendor. If you are not a registered vendor you will not be allowed to distribute any items at the event. If you are found to be selling or distributing any information/product without being on the approved list, you will be asked to leave the venue.
My group/company/team wants to become a sponsor and/or support this event, how do we make that happen?
First, thank you for considering becoming a part of this event! The process towards sponsorship is easy! Just
Contact Us
to get more information regarding the sponsor tier levels. We look forward to hearing from you.
What can't I bring to the Saturday 'Meet, Greet, and Show'
Weapons/drugs of any kind,
BBQ pits, animals, alcohol;
Both the Venue and the Scoobapalooza staff reserve the right to deem any items/actions as harmful/dangerous and therefore unfit for the venue and/or Scoobapalooza Event. If you have a question about something please ask.
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- Case report
- Open Access
Popliteal lymph node dissection for metastatic squamous cell carcinoma: a case report of an uncommon procedure for an uncommon presentation
- Basem B Morcos1Email author,
- Sameh Hashem2 and
- Firas Al-Ahmad1
© Morcos et al; licensee BioMed Central Ltd. 2011
- Received: 20 July 2011
- Accepted: 15 October 2011
- Published: 15 October 2011
Lymph node metastasis from cutaneous squamous cell carcinoma is uncommon. The popliteal fossa is rarely involved with metastasis. Popliteal lymph node dissection is uncommonly performed and not frequently discussed in the literature. We present a case of squamous cell carcinoma of the heel with popliteal and inguinal metastasis. This is followed by a description of the relevant anatomy of the popliteal fossa and the technique of popliteal lymphadenectomy.
- Squamous cell carcinoma
- Popliteal metastasis
- Popliteal dissection
Regional lymph node (LN) metastasis is an important prognostic factor for skin cancers like melanomas and squamous cell carcinomas (SCC) [1]. The usual nodal basins involved are the cervical, axillary and inguinal regions. However, metastasis can involve other areas, like the epitrochlear and popliteal regions. The use of preoperative lymphoscintigrapy (LS) has allowed the detection of interval or in-transit LN, which can also be involved with metastatic disease [1].
Skin cancers of the lower limb below the knee mainly metastasise to the groin. Metastasis to the popliteal LN is much less common. In melanoma, the incidence of popliteal metastasis ranges from 0.3% to 7% [2–4]. For SCC the incidence is not known, but appears to be less than these figures. Lymph nodes in this region should be managed as those in any other nodal basin. They should be looked for, and if involved, they should be resected. Due to the fact that popliteal node metastasis is uncommon and that many surgeons lack the experience in popliteal dissection, this issue is infrequently discussed in the literature. The wider adoption of sentinel lymph node (SLN) biopsy for skin cancer is causing more surgeons to deal with LN in this region. We present a case of SCC with popliteal LN metastasis. This is followed by a review of the relevant anatomy and the surgical technique of popliteal lymphadenectomy.
Case presentation
In 2008, a 41-year-old lady developed an ulcer on her left heel. This was excised by a general surgeon at another hospital. Pathological examination revealed a 3.8 cm, moderately differentiated SCC; the margins were involved. No further treatment was offered to the patient. A year later, in June 2009, the tumor recurred and was excised again; the margins were negative this time.
Popliteal lymph node dissection
In general there are 2 steps to popliteal LN dissection. The first step is adequate exposure of the diamond shaped fossa. This should include careful identification and preservation of the neurovascular structures. Once that is done, dissection of the fat pad should be performed thoroughly.
Subsequently, the popliteal fossa is entered by opening the popliteal fascia vertically in the midline. The first structure to appear is the Tibial nerve, which crosses vertically and disappears between the 2 heads of Gastrocnemius. Lateral to it is the Peroneal nerve, which courses laterally along the biceps femoris tendon and then turns obliquely towards the fibula. Both nerves should be retracted laterally with vessel loops to allow adequate and safe exposure of the vessels and the LNs. Dissection deep to the Tibial nerve will reveal the popliteal vein. The popliteal artery passes deep and slightly medial to the vein. It is the deepest structure in the popliteal fossa and lies on the femur and capsule of the knee joint. The LNs are contained within the fatty tissue that is found superficial, alongside and deep to the popliteal vessels. LN dissection should include, therefore, all the fatty tissue in the popliteal fossa till reaching the posterior aspect of the knee joint. It should be noted that the popliteal fossa is wider in the deeper part than in the superficial part. Adequate retraction of the Gastrocnemius muscle heads, therefore, is essential to ensure adequate exposure and dissection of all the LNs.
After securing hemostasis, the fascia is approximated and the wound is closed over a negative suction drain. A knee splint or back slab is preferably applied at this stage and left in for the first few postoperative days. Postoperative Lymphoedema is expected in some patients [4]. The incidence of lymphoedema is expected to increase dramatically with the addition of inguinal dissection and or radiotherapy.
We describe an unusual case. The incidence of LN metastasis from SCC is relatively uncommon. It ranges from 0.5% to 16% [7], with most series reporting a rate of < 5% [8]. Usually this occurs in tumors with poor prognostic factors such as larger size, poor differentiation, presence of perineural invasion or recurrent tumors. Popliteal metastasis is quite rare. In a series of 1, 118 patients, with localized squamous cell carcinoma of the extremity, treated at M.D. Anderson Hospital and Tumor Institute in Houston, Texas, only 16 patients (1.4%) developed recurrence [9]. Of the 106 treated patients who had regional or distant metastasis, only 2 patients had popliteal metastasis, and both had inguinal metastasis as well.
Although SLN biopsy is mainly indicated for melanomas, it is gaining wider application in the management of other skin tumors, like SCC [10]. Still, the indications for SLN biopsy in SCC are not clear. Its' use in lower extremity tumors will allow the detection of more patients with popliteal LN metastasis, many of those will probably be micrometastases.
The procedure of popliteal lymphadenectomy was first described by Karakousis in 1980. Most later descriptions are based on it. Knowledge of the relevant anatomy and the surgical technique of popliteal LN dissection is important to avoid injury to important structures and to allow adequate LN dissection.
Popliteal metastasis from skin SCC is quite uncommon. It usually occurs in patients with advanced or recurrent tumors. Appropriate management is not clear, but surgical dissection, in the absence of distant metastasis, is indicated since there is no known effective adjuvant treatment for skin SCC.
Written informed consent was obtained from the patient for publication of this Case report and any accompanying images. A copy of the written consent is available for review by the Editor-in-Chief of this journal.
Authors' Affiliations
- McMasters KM, Chao C, Wong SL, Wrightson WR, Ross MI, Reintgen S: Interval sentinel lymph nodes in melanoma. Arch Surg. 2002, 137 (5): 543-7. 10.1001/archsurg.137.5.543. discussion 547-9View ArticlePubMedGoogle Scholar
- Hatta N, Morita R, Yamada M, Takehara K, Ichiyanagi K, Yokoyama K: Implications of popliteal lymph node detected by sentinel lymph node biopsy. Dermatol Surg. 2005, 31 (3): 327-30.View ArticlePubMedGoogle Scholar
- Menes TS, Schachter J, Steinmetz AP, Hardoff R, Gutman H: Lymphatic drainage to the popliteal basin in distal lower extremity malignant melanoma. Arch Surg. 2004, 139 (9): 1002-6. 10.1001/archsurg.139.9.1002.View ArticlePubMedGoogle Scholar
- Thompson JF, Hunt JA, Culjak G, Uren RF, Howman-Giles R, Harman CR: Popliteal lymph node metastasis from primary cutaneous melanoma. Eur J Surg Oncol. 2000, 26 (2): 172-6. 10.1053/ejso.1999.0765.View ArticlePubMedGoogle Scholar
- Barrasa Shaw A, Merle FS, Diana CF, Manez JC, Albadalejo CV: Popliteal lymphadenectomy on sentinel lymph node melanoma metastasis. Clin Transl Oncol. 2006, 8 (3): 218-20. 10.1007/s12094-006-0014-z.View ArticlePubMedGoogle Scholar
- Karakousis CP: The technique of popliteal lymph node dissection. Surg Gynecol Obstet. 1980, 151 (3): 420-3.PubMedGoogle Scholar
- Cherpelis BS, Marcusen C, Lang PG: Prognostic factors for metastasis in squamous cell carcinoma of the skin. Dermatol Surg. 2002, 28 (3): 268-73. 10.1046/j.1524-4725.2002.01169.x.PubMedGoogle Scholar
- Joseph MG, Zulueta WP, Kennedy PJ: Squamous cell carcinoma of the skin of the trunk and limbs: the incidence of metastases and their outcome. Aust NZJ Surg. 1992, 62 (9): 697-701. 10.1111/j.1445-2197.1992.tb07065.x.View ArticleGoogle Scholar
- Ames FC, Hickey RC: Metastasis from squamous cell skin cancer of the extremities. South Med J. 1982, 75 (8): 920-3. 10.1097/00007611-198208000-00005. 932View ArticlePubMedGoogle Scholar
- Ross AS, Schmults CD: Sentinel lymph node biopsy in cutaneous squamous cell carcinoma: a systematic review of the English literature. Dermatol Surg. 2006, 32 (11): 1309-21. 10.1111/j.1524-4725.2006.32300.x.PubMedGoogle Scholar
This article is published under license to BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.


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The lavish Montsorrel estate in Palm Beach looked lovely but had no takers for 20 months. On Friday, the seller's patience was rewarded when packaging king Nelson Peltz, 44, agreed to pay $18 million in reportedly the most expensive real-estate sale ever in the eastern United States.
Set on seven acres, Montsorrel has 608 feet of private ocean frontage and a heated saltwater swimming pool. The two-story 35,000-square foot mansion has 14 main rooms, eight bedrooms and 10 full bathrooms. It has heated marble floors in the bathrooms and hidden electrical winches to ease chandelier cleaning. Terraces are protected in stormy weather by hydraulically operated arms that lift the travertine marble floor at an angle.


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The glossary of wine-speak is something that is constantly evolving, but once you get your head around some of the better-known phrases (and even some of the more left-of-centre ones too!) you'll be able to confidently weave your way into a serious, in-depth wine discussion at any occasion and in any company. Below is a brief foray into the language of wine, which will hopefully allow you to manage some of those awkward wine-related moments with more confidence.
Acidity: Acidity is a vital component when it comes to bringing freshness to wine. Wines with excessive acidity will taste sour and tart whereas wines lacking acidity can seem dull and lifeless. Racy and crisp = good acidity. Flabby = bad.
Aroma: The smell of a young wine, hopefully full of lovely fresh (primary) fruit.
Balance: The holy grail for winemakers. Wines should be in balance, with the acidity, alcohol, fruit and tannins all coming together in harmony.
Barnyard: There's no denying that if a wine smells like a barnyard it's going to be tricky to be seen as a positive. Yet historically it was! In more modern times if your wine smells of a barnyard it is more than likely to be slightly faulty or have been produced in a winery with less than acceptable hygiene standards.
Body: Body refers to the complete sense the wine purveys on the palate. Wines with more concentrated fruit flavours, higher alcohol and ripeness will be full bodied and softer wines with less concentrated fruit will most likely be described as light bodied.
Chunky: Not often heard, but when thrown about refers to some wines that are perhaps carry a bit of extra weight.
Clunky: When a wine is out of balance with too much acid and not enough fruit and tannins, or also a wine with too much fruit and lacking acidity. Simply put, a wine that is out of balance.
Closed: If a wine is described as closed it could mean a couple of things. Hopefully it means that with a bit of air it might start to open up and show itself off a bit better but it could also mean that it's a bit young and could benefit from some time in the cellar.
Corked: Often misunderstood. Cork taint is a fault and a corked wine will smell mouldy and of damp cardboard. It is the result of an unclean or faulty cork. When the cork and wine come into contact this musty smell is imparted upon the wine.
Delicate: Wines which are lighter in body, with a softer personality, normally less fruity and more perfumed.
Delicious: Modern term for wines which are quite simply delicious. We got you there!
Earthy: A term quite often used to try and make excuses for under-ripe wines. Can be positive though, and some older wines can develop a lovely earthy character.
Elegant: In a nutshell, elegant wines are seamlessly constructed with a degree of subtlety rather than being big, jammy and full-bodied.
Fat: Some people like fat wines, some people even have fat wine fetishes, and some people feel you need a knife and fork to get through fat wines. Fat wines have a plumpness to them, and spades of concentrated fruit with lower acidity, giving a sense of opulence and oomph!
Finish (and Length): Not when the bottle is empty but more how the wine ends up on the palate. If the flavour lingers for a while and has layers of fruit, you may say 'Wine Z has a nice long complex finish'. Wines that have a short finish will usually be cheaper simpler wines but can still have 'a short but pleasant finish'.
Flabby (what fat wines can quickly become): A wine lacking in acidity that looks flat and unflattering.
Flashy: Commonly wines that attract a hefty price tag, that have probably seen a lot of new oak and will have lots of ripe, rich fruit.
Focused: When a wine has a nose and fruit profile that is really precise and nicely defined.
Funky: Nothing to do with James Brown, funky is loved by some and loathed by others. Generally used to describe a wine that had some slightly different aromas and flavours, which are neither fruity nor faulty!
Herbaceous: Often wines, and some specific varieties, have a lovely herbaceous character, ranging from thyme to lavender and fennel. If a wine is overtly herbaceous it may lack a bit of ripeness.
Intensity: Something for wine makers to strive for. Wines with intensity of fruit balanced with acidity, alcohol and tannins is what we all want to drink and what wine makers look to capture when making wines.
Lean: Even though they might lack the profile of some of their fuller-bodied friends, lean wines, with their lighter framework and more streamlined personality lean wines can still provide oodles of drinking pleasure.
Linear: A wine with nicely defined acidity which gives the wine drive and focus.
Masculine: A full bodied wine with concentrated fruit, high tannins or a dose of new oak would fall into the masculine category.
Minerality: Minerality is more often than not used to describe wines with a flinty, slately, steeliness (but is often wrongly used to describe wines which have racy acidity).
Nose: The nose is wine-speak for the smell.
Oxy (Oxidized): Oxy is used if a wine has been exposed to oxygen and has deteriorated due to the damaging effect oxygen can have on the structure of a wine. Some wines on the other hand are made with a bit of oxidative handling and are intended to have an oxidized character (think Amontillado Sherry and Tawny Port)
Oaky: Oak should always been in balance when used in winemaking. When used judiciously, oak adds a lovely toasty character to white wine and helps build in complexity. With reds it adds layers of flavour and helps the wine soften out before bottling.
Plump: A plump wine is one that has concentration of sumptuous primary fruit and that isn't over extracted and fat but more… plump!
Pretty: Pretty wines are making a comeback! Petty will refer to a wine that is elegant, normally with a nice floral character that charms the palate rather than making it work too hard.
Silky: Velvety and silky can be used interchangeably to describe a bottle that has a lovely mouth feel, and with a plushness of fruit that is usually fairly plump but not quite fat.
Smokey: Smokiness can be derived from a period of oak maturation. If a wine has a smoky character in its youth it may well have spent some time in highly toasted new oak barrels and could benefit from some in time in the cellar before you pop open the next bottle
Steely: When a wine has higher acidity it will come across as being steely.
Tannins: Tannins are mostly found in red wines and are the cause of the drying sensation you feel on your gums. High, unripe tannins can be aggressive and unpleasant, but round, soft tannins help bring structure to a wine.
Yummy: Perhaps not a technical wine tasting term but let's face it, sometimes a wine is just yummy and demands that you drink it up in good company and with a smile on your face.
So there you have it: 35 introductory pieces of wine-speak that will not only build your knowledge of the glossary of wine, but hopefully give you more confidence when it comes to selecting your next bottle. If nothing else, hopefully it will at least it give you something to chat about when you are next cornered by a sommelier at a dinner party!
^You must be a Vintage Cellars Wine Club member and present your membership card at time of purchase to quality for points, Cellar Shares or any other 'Member Bonus' offers. Cellar Shares are mailed regularly throughout the year summarising your points for the transaction period.
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