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Grammar and vocabulary notes follow for the aid of Spanish students. De repente vio al lobo, que era enorme, delante de ella. Vieron la puerta de la casa abierta y al lobo tumbado en la cama, dormido de tan harto que estaba.

Muchachita is a diminutive form of muchacha , a word for girl. The diminutive is formed here using the suffix -ita. The diminutive form can indicate that the girl is small or can be used to indicate affection.

Abuelita , a form of abuela or grandmother, is another diminutive found in this story. In that case, it is probably being used as a term of affection rather than referring to her size.

The name of the story itself is another diminiutive; a caperuza is a hood. The sole exception was an Ibo tale, which grouped with European variants of Little Red Riding Hood, thus endorsing the collector's belief that the story is not of local origin, but an Ibo oral translation of the western fairy tale [50].

The tale was subsequently modified to create a novel redaction that spread across central and southern societies on the continent, and even as far as Antigua.

Although bootstrap and posterior support for this clade was relatively modest, it is remarkable that the phylogenetic signal in this tradition was sufficiently strong to be detected by all three analyses, despite the massive cultural and human upheavals that occurred during the forced displacement of African populations during the slave trade.

Since there is no evidence to suggest they share a more recent common ancestor with The Wolf and the Kids or Little Red Riding Hood, they cannot be classified as members of either international type.

One intriguing possibility raised in the literature on this topic that would be consistent with these results is that the East Asian tales represent a sister lineage that diverged from ATU and ATU before they evolved into two distinct groups.

A more detailed exposition of this theory has been set out by the Sinologist Barend J. Noting that the The Tiger Grandmother encompasses a spectrum of more ATU like variants and more ATU like variants, Haar argues that the East Asian tales represent an ancient autochthonous tale type that is ancestral to the other two.

On the basis of qualitative comparisons among these and other Asian tales, he conjectures that the tale originated in China and spread westwards to the Middle East and Europe between the twelfth and fourteenth century, a period during which there were extensive trade and cultural exchanges between east and west.

At some unspecified later point, the tale type split into the lineages that gave rise to Little Red Riding Hood and the Wolf and the Kids.

Although it is tempting to interpret the results of the analyses in this light, there are several problems with this theory.

Of course, as mentioned previously, literary evidence about the origins of oral tales can be unreliable and biased toward Europe.

However, at the very least, the existence of ATU in first century Europe means that the putative Asian ancestral tale type would have to had to have spread west long before the opening of trade routes in the twelfth to fourteenth centuries, as suggested by Haar [51].

Second, if ATU and ATU are more closely related to each other than they are to the East Asian tales, they would be expected to share derived characters i.

However, there is not a single characteristic shared by these two tale types that does not also occur in the East Asian group.

If that were the case, we would expect earlier versions of ATU and ATU to be more similar to the East Asian tales than later variants, as original elements of the story would be lost or substituted as each tradition evolved.

However, this prediction is contradicted by the available chronological data on the tales' histories. In ATU , this test first appears in a version of the fable recorded in the fourteenth century [46] , and is lacking in the original version.

An almost identical episode occurs in variants of The Wolf and the Kids and is also present in the African tales , in which the wolf drinks something or cuts his tongue to smooth out his voice.

However, it does not appear in any recorded versions prior to the publication of the Grimms' Children and Household Tales in [1]. The latter trait has excited particular interest among folklorists, since it occurs in the oral tale The Story of Grandmother and not in Little Red Riding Hood where the girl gets eaten.

To investigate the evolution of these similarities more rigorously, the ancestral states of the traits discussed above were reconstructed on the tale phylogenies see Methods for details.

The results are shown in Table 1 below. An alternative — and, to the best of this author's knowledge, novel — explanation for the relationship of the East Asian tales to ATU and ATU is that the former is derived from the latter two, rather than vice versa.

This hypothesis would account for the finding that important traits shared by the East Asian tales and Little Red Riding Hood and The Wolf and the Kids are not ancestral, suggesting that they were borrowed instead.

Given the number and striking nature of these resemblances, it seems unlikely that they could have evolved independently. Borrowing is also consistent with patterns of conflicting signal in the NeighbourNet graph, which appear to be especially prevalent around the East Asian group.

This impression is confirmed by a comparison of taxon-specific delta scores and Q-residuals, which are higher on average for the East Asian tales than other tales.

The average delta score of the East Asian tales is 0. To investigate this hypothesis further, another set of analyses were carried out in which the East Asian tales were removed from the data along with the characters that were only present in this group.

It was reasoned that if these tales evolved by blending together elements of ATU and ATU then their removal should result in a more phylogenetically robust distinction between these two groups.

This prediction was tested by maximum parsimony bootstrapping and Bayesian inference. For reference, consensus trees derived from both analyses are presented in the Supporting Information, together with a NeighbourNet graph excluding the East Asian tales Figures S1 , S2 and S3.

Thus, both analyses indicate that the East Asian tales are a source of conflicting signal in the data, in line with the hybridisation hypothesis.

While on current evidence this appears to be the best available explanation for the relationships between the East Asian group and ATU and ATU , questions remain about how, where and when the latter two tale types were adopted and combined.

Based on the similarities described above, it seems likely to have occurred sometime between the origin of the lineage leading to Little Red Riding Hood and The Story of Grandmother, but before the publication of Perrault's classic tale in Given the antiquity and wide geographic diffusion of The Wolf and the Kids, it is certainly plausible that ATU would have also reached China by this time, perhaps between the twelfth and fourteenth centuries, i.

Given the current state of the evidence, such scenarios are necessarily speculative. However, the digitisation and translation of an ever increasing number of folklore collections from Asia, as well as other regions, promise to yield a wealth of new data with which to investigate these questions more thoroughly in the future.

In the meantime, this case study has shown that phylogenetic methods provide powerful tools for analysing cross-cultural relationships among folktales that can be used to classify groups based on common ancestry, reconstruct their evolutionary histories, and identify patterns of contamination and hybridisation across traditions.

While these goals are clearly of crucial importance to comparative studies of folklore, they also have potentially exciting applications in other fields too.

As previous researchers have pointed out, the faithful transmission of narratives over many generations and across cultural and linguistic barriers is a rich source of evidence about the kinds of information that we find memorable and motivated to pass on to others [9] [53] [54].

In the present case, stories like Little Red Riding Hood, The Tiger Grandmother and The Wolf and the Kids would seem to embody several features identified in experimental studies as important cognitive attractors in cultural evolution.

Equally, these methods could be applied to explore how tales are influenced by cultural, rather than psychological, selection pressures.

Such an analysis might address whether local modifications of different tale-types exhibit consistent patterns, and see if they covary with specific ecological, political or religious variables.

Future work on these questions promises to generate important insights into the evolution of oral traditions, and open new lines of communication between anthropologists, psychologists, biologists and literary scholars.

Cladistic analysis employs a branching model of evolution that clusters taxa on the basis of shared derived evolutionarily novel traits. To search for the most parsimonious tree MPT , the present analysis employed an efficient tree-bisection-reconnection algorithm implemented by the heuristic search option in PAUP 4 [57] , carrying out 1, replications to ensure a thorough exploration of tree-space.

The RI is a measure of how well similarities among a group of taxa can be explained by the retention of shared derived traits on a given tree [34].

A maximum RI of 1 indicates that all similarities can be interpreted as shared derived traits, without requiring additional explanations, such as losses, independent evolution or borrowing.

As the contribution of these latter processes increase, generating similarities that conflict with the tree, the RI will approach 0.

Maximum parsimony bootstrapping is a technique for measuring support for individual clades [58]. It involves carrying out cladistic analyses of pseudoreplicate datasets generated by randomly resampling characters with replacement from the original matrix.

Support for the clades returned by the original analysis is then estimated by calculating the frequency with which they occur in the most parsimonious trees obtained from the pseudoreplicates.

The bootstrap analyses reported here were carried out in PAUP 4 [57] using heuristic searches of 1, replicates. Bayesian inference proceeds by calculating the likelihood of the data given an initially random tree topology, set of branch lengths and model of character evolution, and iteratively modifies each of these parameters in a Markov Chain Monte Carlo MCMC simulation.

Moves that improve the likelihood of the data are always accepted, while those that do not are usually rejected although some may occasionally be accepted within a certain threshold so as to avoid getting trapped in local optima.

Unlike the trees output by a cladistic analysis, which are based on a single optimality criterion i. The Bayesian approach has been found to be particularly effective when there is wide variance in the amount of evolution that has occurred in different regions of the character data or tree, since it explicitly incorporates these parameters i.

The Bayesian analyses reported here were carried out in MrBayes 3. Two analyses were carried out simultaneously, each using four MCMC chains that were run for 1 million generations.

Log likelihood values for the remaining trees in each sample were then graphed as a scatterplot to check that the two runs had converged.

As with the other two methods, NeighbourNet clusters taxa into hierarchically nested sets. However, unlike cladistics and Bayesian inference, it does not employ a strict branching model of descent with modification, and as such these sets can overlap and intersect with one another.

Accordingly, it is claimed that NeighbourNet is better able to capture conflicting signal in a dataset resulting from borrowing and blending among evolutionary lineages [35].

The method involves calculating pairwise distances between the taxa based on the character data, and generating a series of weighted splits that are successively combined using an agglomerative clustering algorithm.

Where the splits are highly consistent, the diagram will resemble a branching tree-like structure. Incompatible splits, on the other hand, produce box-like structures that lend a more latticed appearance to the network.

The extent of reticulation in the folktale network was quantified using the delta-score and Q-residual score [36] , [61]. Quartets are scored from 0 to 1 according to how resolved the splits between each pair of taxa are, with values closer to 0 being more tree-like and values closer to 1 more reticulate.

The estimation of the delta score includes a normalisation constant, whereas Q-residuals had to be normalised by rescaling all between-taxa distances in the network so that they average 1.

The NeighbourNet analysis and calculation of d-scores and Q-residulas were carried out in SplitsTree v4. Character states were reconstructed in the putative last common ancestor of ATU and ATU tales through parsimony analysis and Bayesian inference.

Next, the evolutionary history of each character was reconstructed on the MPTs by minimising the total number of changes required by each tree.

The parsimony analyses were carried out in the software program Mesquite, using the Character Trace module [62]. The analysis was carried out in MrBayes 3.

Majority rules consensus of the most parsimonious trees returned by cladistic analyses of the data with East Asian tales removed. The numbers beside the nodes represent bootstrap support values for each clade.

Maximum clade credibility tree returned by a Bayesian analysis of the data with the East Asian tales removed. Numbers represent posterior probabilities associated with each clade.

NeighbourNet graph of the data with East Asian tales removed. The average delta score on the Network was 0.

Conceived and designed the experiments: JT. Performed the experiments: JT. Analyzed the data: JT.

Wrote the paper: JT. Browse Subject Areas? Click through the PLOS taxonomy to find articles in your field. Abstract Researchers have long been fascinated by the strong continuities evident in the oral traditions associated with different cultures.

The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript Competing interests: The author has declared that no competing interests exist.

Download: PPT. Figure 1. Map of the approximate locations from which tales were sourced. Results The cladistic analysis returned equally most parsimonious trees MPTs.

Figure 2. Majority-rules consensus of the most parsimonious trees returned by the cladistic analysis of the tales. Figure 3. Maximum clade credibility tree returned by the Bayesian phylogenetic analysis of the tales.

Figure 4. Split graph returned by the NeighbourNet analysis of the tales. Discussion Comparative cross-cultural studies of folklore have long been dogged by debates concerning the durability and integrity of oral traditions.

Table 1. Methods Phylogenetic Reconstruction Cladistic analysis employs a branching model of evolution that clusters taxa on the basis of shared derived evolutionarily novel traits.

Ancestral state reconstruction Character states were reconstructed in the putative last common ancestor of ATU and ATU tales through parsimony analysis and Bayesian inference.

Supporting Information. Figure S1. Figure S2. Figure S3. Table S1. List of tales used in the analyses and their sources. File S1. List of characters.

File S2. Data matrix. Author Contributions Conceived and designed the experiments: JT. References 1. Thompson S The Folktale.

New York: Dryden. Grimm W Preface. Children's and Household Tales. London: George Bell. Journal of Folklore Research 1— View Article Google Scholar 5.

Copenhagen: Rosenkilde and Bagger. Human Nature-an Interdisciplinary Biosocial Perspective — View Article Google Scholar 8. Sugiyama MS Cultural variation is part of human nature.

Human Nature — View Article Google Scholar 9. London: Routledge. A Classification and Bibliography. Helsinki: FF Communications. Parts I—III.

Helsinki: Folklore Fellows Communications. Journal of Folklore Research — View Article Google Scholar The Journal of American Folklore — PLoS Biol 9: e Science — Trends Genet — Nature — Roos T, Heikkilä T Evaluating methods for computer-assisted stemmatology using artificial benchmark data sets.

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