The Santini Sisters Survive Sex and Dating

I couldn't help but either laugh or groan according to the predicament Victoria "Vickie" and Josephine "Josie" got themselves into.

The only problem was there was no real demarcation between them except for there presence. There were some link breaking issues as well, but it was only in a few places. It is implied that a sequel is in the works and I for one look forward to reading more. Mar 14, Karen rated it liked it. Michele Taylor rated it it was ok Apr 14, Pongo rated it did not like it Jul 18, Angela Myers rated it liked it Jun 22, Alicia Jackson rated it really liked it Apr 30, Amy Sandrin rated it it was amazing Jun 18, B Butterfield rated it really liked it Sep 14, Denise Dunne rated it really liked it Jun 22, Lisa marked it as to-read Mar 09, Joanie marked it as to-read Mar 12, Jill marked it as to-read Mar 18, Julia marked it as to-read Jul 01, Trish Necaise added it Dec 12, KimberlyDawn marked it as to-read Jan 02, Lk Moore added it Mar 23, Kellie Demarsh marked it as to-read May 22, Laura Ballantyne marked it as to-read Sep 11, Mary Gilgannon is another author who blows me away with the variety and depth of her writing.

She writes historical romance and romantic fantasy full of emotion and compelling conflict.

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Colleen has worked as a stand-up comedienne, Hollywood script girl and private investigator, and is both smart and witty. It took me about six months. This is the first in a series. The second book comes out next October. The book has humor, romance, a little bit of mystery, a beautiful setting and characters I hope readers will want to know better. Facebook Email Print Twitter Reddit. Leave a Reply Cancel reply Enter your comment here Fill in your details below or click an icon to log in: Email required Address never made public. If two fossils from the same taxon with different age estimates are included in an analysis, the older fossil has the potential to be recovered as a direct ancestor and would be considered a sampled ancestor under our model.

These parameters are commonly used to describe diversification processes. Note, that the time of origin is a model parameter as opposed to the previous application of the FBD model Heath et al. Here, we allow the oldest fossil to be the direct ancestor or sister lineage to all other samples because there is no prior evidence ruling those scenarios out.

We analyzed a data set from Ksepka et al. The morphological data matrix used here samples 36 fossil species we excluded Anthropornis sp. UCMP due to absence of the formal description for this specimen and 19 extant species we treated the Northern, Southern, and Eastern Rockhopper penguins as three distinct species for purpose of the analysis.

The original matrix contained characters. We excluded outgroup taxa Procellariiformes and Gaviiformes because including them would violate the model assumptions: We excluded characters that became constant after excluding outgroup taxa, resulting in a matrix of characters. The morphological characters included in the matrix ranged from two- to seven-state characters.

The molecular alignment comprises the nuclear recombination-activating gene 1 RAG-1 , and the mitochondrial 12S, 16S, cytochrome oxidase 1 CO1 , and cytochrome b genes. Each region is represented by more than sites with sites in total.

The Santini Sisters Survive Sex and Dating by Cindi Myers

Some regions are missing for a few taxa. The morphological data-set was originally developed to resolve the phylogenetic placement of fossil and extant penguins in a parsimony framework. Thus, efforts were focused on parsimony-informative characters. Though some apomorphic character states that are observed only in a single taxon are included in the data-set, no effort was made to document every possible autapomorphy.

Thus, such characters can be expected to be undersampled. As with essentially all morphological phylogenetic data-sets, invariant characters were not scored. We updated the fossil stratigraphic ages—previously summarized in Ksepka and Clarke —to introduce time intervals for fossil samples as presented in online Supplementary Material SM , Table 1 available on dryad at http: For fossil species known from a single specimen, fossil stratigraphic ages represent the uncertainty related to the dating of the layer in which the fossil was found.

For fossils known from multiple specimens, the ages were derived from the ages of the oldest and youngest specimens. We do not model ordered characters and treated 34 characters that were ordered in the Ksepka et al. Evolution of morphological characters has a different nature from evolution of DNA sequences and, therefore, requires different assumptions. In contrast to molecular evolution models, we do not know the number of states each character can take and the number of states is not constant for different characters. We consider two ways to approach this problem.

First, we can assume that the number of possible states for a character is equal to the number of different observed states. Typically, one would count the number of different states in the data matrix for the character. Here, we obtained the number of observed states from the larger data matrix used in Ksepka et al. Having the number of observed characters for each character, we partition the morphological data matrix in groups of characters having the same number of states and apply a distinct substitution model of the corresponding dimension to each partition.

Another approach is to treat all the characters as evolving under the same model. The model dimension in this case is the maximum of the numbers of states observed for characters in the matrix. Another difference comes from the fact that typically only variable characters are recorded. Thus, the second adjustment to the model accounts for the fact that constant characters are never coded. This model is called the Mkv model Lewis We compared a model which assumed no variation in substitution rates of different morphological characters with a model using gamma distributed rates with a shared shape parameter for all partitions.

We also compared a strict clock model and an uncorrelated relaxed clock model Drummond et al. We completed a model selection analysis comparing different combinations of the assumptions for the Lewis Mk model, morphological substitution rates, and FBD model assumptions by running eight analyses of morphological data with different model settings. We then estimated the marginal likelihood of the model in each analysis using a path sampling algorithm Baele et al. The traditional model selection tool is a Bayes factor, which is the ratio of the marginal likelihoods of two models: Following this logic, the model that provides the best fit is the model with the largest marginal likelihood.

The model combinations with marginal likelihoods are described in Table 2. For most of the bifurcation events in trees from the posterior samples of the penguin analyses only one lineage survives, whereas another lineage goes extinct. This suggests a non-neutrality in the evolution of the populations. To assess whether the FBD model, which assumes all lineages in the tree develop independently, fits the data analyzed here, we performed the posterior predictive analysis following Drummond and Suchard under model 8 in Table 2.

The idea of such an analysis is to compare the posterior distribution of trees to the posterior predictive distribution Gelman et al. The posterior predictive distribution can be approximated by the sample of trees simulated under parameter combinations drawn from the original posterior distribution. For this analysis, we considered tree statistics which can be grouped into two categories: For the total-evidence analysis of the penguin data set, we chose the substitution and clock models for morphological character evolution with the largest marginal likelihood from the model comparison analysis analysis 8 in Table 2.

This model suggests that the data are partitioned in groups of characters with respect to the number of observed states. Each partition evolves under the Lewis Mkv model and the substitution rate varies across characters according to a Gamma distribution shared by all partitions. The morphological clock is modeled with an uncorrelated relaxed clock model with log-normal distributed rates. A separate log-normal uncorrelated relaxed clock model is assumed for the molecular data alignment. Each branch is assigned a total clock rate drawn from a log-normal distribution and this rate is scaled by a relative clock rate for each gene so that relative clock rates for five partitions sum up to one.

We also ran the same analysis under the strict molecular clock. We also analyzed this data set under the birth—death model without sampled ancestors Stadler In addition to analyzing the full data set, we performed a separate analysis of only living penguins and the crown fossils, to examine the effect of ignoring the diversification of fossil taxa along the stem lineage.

Crown fossils were selected if the fossil lineage was a descendant of the MRCA of all extant species with a posterior probability greater than 0. Thus, this analysis includes six fossils listed in Table 1 and all living penguins. For this analysis we did not condition on recording only variable characters i. First, we summarized the posterior distribution of full trees using summary methods from Heled and Bouckaert An MSACC tree is a tree from the posterior sample that maximizes the product of posterior clade probabilities. Here, a clade denotes two types of objects.

The first type is a monophyletic set of taxa with a bifurcation node as the MRCA. The second type is a monophyletic set of taxa with a two-degree sampled node as the MRCA. This can happen when one of the taxa in the group is a sampled ancestor and it is ancestral to all the others in the clade. Then this taxon will be the MRCA of the whole group assuming it is an ancestor to itself.

Second, we removed all fossil lineages from the posterior trees thereby suppressing two-degree nodes and then summarized the resulting trees which are strictly bifurcating with a maximum clade credibility tree with common ancestor ages. To assign a common ancestor age to a clade, we consider a set of taxa contained in the clade and find the age of the MRCAs of these taxa in every posterior tree including the trees where these taxa are not monophyletic and take the mean of these ages.

The tree prior parameterization, clock, and substitution models used for eight analyses of penguin morphological data with marginal likelihoods for model testing. Posterior probabilities of fossils to be sampled ancestors for eight models summarized in Table 2. The posterior predictive analysis of the penguin data for branch length and tree-shape statistics indicating no significant difference in the posterior and posterior predictive tree distributions for model 2 in Table 2. The plots do not show obvious discrepancy in the posterior and posterior predictive distributions of these statistics.

The distributions of the tree imbalance statistics almost coincide. The numbers at the bases of clades show the posterior probabilities of the clades. The filled circles represent sampled ancestors. Fossils with positive evidence of being sampled ancestors are shown in red gray in printed version. Fossils Paraptenodytes antarcticus and Palaeospheniscus patagonicus both appear around the same time and have the same prior probabilities of 0.

Penguin reconstructions used with permission from the artists: The evidence for fossils to be sampled ancestors. The samples above the shaded area i. The posterior probabilities of most clades including fossils are low, reflecting the large uncertainty in the topological placement of the fossil taxa, whereas many clades uniting extant taxa receive substantially higher posterior probabilities. We calculated Bayes factors for all fossils to be sampled ancestors assuming the prior probability that a fossil is a sampled ancestor is defined by the tree prior model conditioned on the number of sampled extant and fossil species and assigned sampling intervals.

Adding comparative morphological and molecular data to the sample size and sampling intervals provides positive evidence that the fossil taxa representing the species Palaeospheniscus patagonicus and Icadyptes salasi are sampled ancestors. Eretiscus tonnii, Marplesornis novaezealandiae, Paraptenodytes antarcticus , and Pygoscelis grandis show positive evidence to be terminal samples Fig.

Due to the large uncertainty in the topological placement of fossil taxa, the relationships displayed in the summary tree are not the only ones supported by the posterior distribution.

Thus, in some cases an alternate topology cannot be statistically rejected and a careful review of the entire population of sampled trees is required to fully account for this. Below, we summarize the features of the MSACC topology that differ from previous estimates of penguin phylogeny, keeping this uncertainty in mind.

The maximum clade credibility tree of extant penguins with common ancestor ages. The numbers at the bases of clades show the posterior probabilities of the clades after removing fossil samples. Allowing fossils to represent ancestors yields several interesting results. Although there is no evidence in comparative data to support an ancestral position for Spheniscus muizoni Fig.

The Santini Sisters Survive Sex and Dating

The ancestral position is consistent with the morphological data set: Furthermore, at least for the discrete characters sampled, it does not exhibit apomorphies providing direct evidence against ancestral status. This fossil taxon was inferred as the sister taxon to Eudyptes by several previous studies Hospitaleche et al. Our results indicate a 0. Presumably the position of M. Most of the clades along the stem receive very low posterior probabilities, which is not unexpected given that some stem penguin taxa remain poorly known, with many morphological characters unscorable.

These clades correspond to the large polytomies from Ksepka et al. Of particular note is the placement of Palaeospheniscus bergi and Palaeospheniscus biloculata on a branch along the backbone of the tree leading to all modern penguin species. Palaeospheniscus penguins share many synapomorphies with crown penguins and only a single feature in the matrix presence of only the lateral proximal vascular foramen of the tarsometatarsus contradicts this possibility and supports their forming a clade with E.

The posterior distribution of our analysis supports a clade containing the three Palaeospheniscus species and E. Overall, the estimated clades with large posterior probabilities greater than 0.

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Low posterior probability values are due to the sparse morphological data and complete lack of molecular data for many early stem taxa. Several species such as Palaeeudyptes antarcticus are based on very incomplete fossils, in some cases a single element, and so we view the relationships estimated for deep stem penguins as incompletely established for the time being.

Despite the high degree of uncertainty in the phylogenetic relationships of fossil species, the overall support for the general scenario placing most fossil penguins along the stem with a recent appearance of crown penguins is strong. To better describe, measure, and visualize the topological uncertainty of total-evidence analyses, methods similar to Billera et al.

We estimated the divergence dates for extant penguins Fig. In general, the estimates are younger than those reported in previous studies: Their estimate of the age of the MRCA of the extant penguins was Most recently, Jarvis et al. Notably, this last date can be considered applicable to the crown only if Aptenodytes or Pygoscelis is the sister taxon to all other penguins, otherwise this date applies to a more nested node, implying an older age for the crown.

We assert that this is the best constrained estimate of the age of the penguin crown clade to date, because it avoids potential pitfalls related to the use of secondary calibrations, samples all extant species, and most importantly includes all reasonably complete fossil taxa directly as terminals or sampled ancestors. This final point is crucial, not only because including fossils as terminals has been shown influence phylogenetic accuracy under many conditions e.

The small gap between our Moreover, our results suggest many extant penguin species are the product of recent divergence events, with 13 of 19 sampled species splitting from their sister taxon in the last 2 myr. Penguins have a relatively dense fossil record compared to other avian clades, with thousands of specimens known from four continents and spanning nearly the entirety of their modern day geographical range. If crown penguins originated at 20—50 Ma as implied by previous studies, it would require major ghost lineages Clarke et al.