We thank the Smithsonian Tropical Research Institute for help with obtaining collecting and export permits in Panama, and Javier Guevara for issuing permits for collecting and exporting specimens from Costa Rica. Browse Subject Areas? Click through the PLOS taxonomy to find articles in your field.
Abstract Natural selection often results in profound differences in body shape among populations from divergent selective environments. Introduction Numerous studies have documented adaptation to divergent natural selection regimes [1] — [8].
Materials and Methods Molecular Laboratory Methods and Analysis of Genetic Distance A primary purpose of this study is to determine how body shape evolves at different phylogenetic levels of divergence i. Download: PPT. Figure 1. Map of collection sites for Brachyrhaphis terrabensis , B. Geometric Morphometric Analyses We used a total of fish in the geometric morphometric analysis Appendix I : B. Results Effects of Predation Environment on Body Shape Consistent with the predictions in our first hypothesis, we found that body shape differed between predation environments both within Brachyrhaphis rhabdophora and between B.
Table 1. Results of mixed-repeated-measures MANOVA testing for interactions between combinations of species-group, predation-environment, size and index-variable. Table 2. Table 3. Figure 3. Morphological Divergence in Female Brachyrhaphis.
Figure 4. Morphological Divergence in Male Brachyrhaphis. Discussion The principal objective of our study was to test for divergent morphologies driven by predation environment in Brachyrhaphis fishes at two taxonomic levels in two phylogenetically sister lineages, and determine how much variation occurs within populations and species that have evolved in similarly divergent selective regimes.
Conclusions In conclusion, sister taxa Brachyrhaphis roseni and B. Supporting Information. Figure S1. Table S1. Table S2. Table S3. Table S4. Acknowledgments We thank P. References 1. Nosil P Divergent host plant adaptation and reproductive isolation between ecotypes of Timema cristinae walking sticks. American Naturalist — View Article Google Scholar 2. Nosil P Adaptive population divergence in cryptic color-pattern following a reduction in gene flow. Evolution — View Article Google Scholar 3.
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In morphology, the dietary shift from soft browsing to hard and fibrous grazing plant material is accompanied by profound changes in the skull and mandible, including the acquisition of high-crowned hypsodont molars, longer snout, and deeper mandible [ 44 — 46 ]. This pattern is present in equids, and also appeared several times among Pecora. Nonetheless, true grazing is restricted to a minority of species, most of them being properly defined as mixed-feeders consuming both grasses and soft material [ 44 ].
The data were obtained from images in lateral view taken from the scientific literature or directly from specimens see S2 File for full details , representing species.
On each image we recorded nine landmarks to register mandibular shape and analyzed shapes by means of geometric morphometrics see S3 File for details. The ungulate tree was assembled from literature [ 16 , 46 , 47 ].
The third real case pertains to extant lizards of the genus Anolis. The genus includes more than species distributed in the Neotropical region and the Caribbean. Insular anoles fall into six distinct ecomorphs which have been intensely studied as a classic example of convergent evolution [ 19 ]. The data include a species wide tree for Anolis lizards living on the main islands of the Greater Antilles, and 11 phylogenetic principal components extracted analyzing lizards body shapes [ 48 , 49 ] see Supplementary S4 File for the R code.
Six species do not fall into any ecomorph category and are therefore not expected to converge. The average tree size in the simulation experiments was The clades set to converge varied from 17 to 44 species average On average, the heights of the clades set to converge were These results indicate that search. The power of search. We found this usually depends on the balancing between the clade set to converge and its sister node, and the strong phenotypic autocorrelation between these clades because a given clade necessarily includes all of the descendants of its daughter node.
When the sister to the real mrca is made up of very few species search. Whichever exact mrca pair is identified, The Type I error the percentage of false positives remains remarkably low Table 1. Type II error of search. This is not surprising because under the state case species evolving under a single state appear randomly across the tree, hence the effect of f transformation is diffused rather than focusing on a single clade.
We found 47 instances of convergence among groups of three randomly selected species out of 1, simulations with phenotypes designed to evolve under the BM model. This means that the Type I error rate of search. By using the OU process to model convergence, we found search.
The corresponding figure for Type II error rate 4. When testing for convergence between clades, we found two instances of convergent morphological evolution, both pertaining the same clade, Barbourofelidae. The latter includes false saber-toothed cats of the genera Barbourofelis and Albanosmilus. They were found to be convergent on both Smilodontini and Homotheriini within machairodonts, which represent the true sabertoothed cats Fig 5.
It is noteworthy that search. The mean angle between barbourofelids and Smilodontini is The angle between their ancestors is This suggests that the two clades evolved along parallel trajectories.
The angle between barbourofelids and Homotheriini is The computational time was seconds. A The clades found to converge were Homotheriini orange and Barbourofelidae blue and Smilodontini green. Deformation grids are shown at the extremes of both axes. The silhouette for Homotherium was available for reuse under the Creative Commons Attribution 3.
The silhouettes for Smilodon and Barbourofelis are our own work. By using the mandibular shapes of Barbourofelis fricki and Smilodon fatalis as the ancestral states to all barbourofelids and machairodonts, respectively, the results are similar to those obtained without specifying phenotypes at the mrca nodes Table 2B , S3 File , and this may help explaining the good performance of search.
By performing the analysis collapsing machairodonts and barbourofelids under a single state, search. The computational time was 44 seconds.
This latter analysis performed by using convratsig [ 40 ] produced significant results for all the measures Table 3. The mean angle between all grazers collapsed under a single state is The mean angle between grazing artiodactyls and grazing perissodactyls is This same figure is 0.
Michael Keesey are available for reuse under the Creative Commons Attribution 3. By using search. The left columns represent the results obtained by applying search. The last two rightmost columns are the corresponding results pertaining to the metrics C1 and C5, retrieved from [ 22 ]. The Anolis species that cannot be ascribed to any ecomorphs are, unsurprisingly, not found to converge.
By using the C1 metric, Stayton [ 22 ] found 4 of 6 ecomorphs converging. By using the metric C5, convergence is found in 3 ecomorphs. Species not ascribed to an ecomorph were not found to converge for either of the metrics.
Evolutionary convergence has been the focus of many evolutionary studies [ 6 , 21 ]. Morphological convergence arises from adaptation to similar niches by different lineages, which can be separated geographically, phylogenetically and temporally [ 12 ], although different processes such as phylogenetic and developmental constraint, and even chance may produce the same pattern [ 50 — 53 ].
There are several methods available in the literature to test the hypothesis of morphological convergence. Most of them rely on the basic assumption that convergence implies stronger phenotypic resemblance than expected by phylogenetic distance. Although the method we propose here, search.
Our procedure to identify convergence between clades is at least as dependent on ancestral state estimation as many other approaches e. However, in search. We demonstrate search. Although the mrcas set to converge are not always found with precision under the automatic mode, the species actually set to converge are correctly identified up to The lower performance of seach.
We successfully applied search. The first real case study regards the evolution of mandibular shapes in felids. Intriguingly, Metailurini i. This means search.
We used the felid data to compare search. While both functions recognize the same pattern, search. Grazing adaptations in the mandible evolved independently in horses genus Equus and several bovid lineages, most notably among antelopes. We found evidence for convergent evolution between Equus and strictly grazing bovids, such as Bison , Bos , and Alcelaphus.
This is especially noteworthy considering that the paleontological tree we used includes a number of non-grazing equids, such as hipparionoid horses and browsing anchitheriine equids, plus several extinct rhinos and tapirs which were all browsers. This demonstrates the method was able to find convergence among grazers despite the effect of phylogeny and body size on mandibular shape variation [ 46 ].
The final real case pertains to Anolis ecomorphs. Compared to other statistical procedures used to test for morphological convergence, search. In addition, being much faster than alternative approaches, seach. There are several instances reported in literature of convergence between closely related clades and even single species with close phylogenetic proximity.
Caution must be applied to the choice of the ancestral phenotype in the presence of strong phenotypic drift. Two clades are selected to evolve convergent tip phenotypes simple convergence , convergent tip phenotypes starting from ancestors which are similar to each other convergence and parallel trajectories or no convergence.
At running the code, the user interactively chooses which kind of simulation to perform and how much the convergent clades have to be different phenotypically from the rest of the tree. The authors declare no conflict of interest. Alistair Evans, Roland Sookias and two anonymous reviewers provided insightful comments to an earlier version of this manuscript.
Browse Subject Areas? Click through the PLOS taxonomy to find articles in your field. Abstract Morphological convergence is an intensely studied macroevolutionary phenomenon. Funding: The authors received no specific funding for this work.
Materials and methods The method is based on phylogenetic ridge regression, RRphylo [ 28 ]. Howard DS: Reinforcement: origin, dynamics, and fate of an evolutionary hypothesis. Hybrid zones and the evolutionary process. Edited by: Harrison RG. Kirkpatrick M: Reinforcement during ecological speciation. Proc Biol Sci. Roff DA: The evolution of life histories: Theory and analysis.
Stearns SC: The evolution of life histories. Annals of the Entomological Society of America. Mol Phylogenet Evol. Journal of Insect Behavior. Bennet-Clark HC: Songs and the physics of sound production. Journal of Experimental Biology. Download references. This paper was greatly improved by critical reviews of anonymous referees. This work was supported financially by the undergraduate research fellowships of the Ewha University to students for morphological measurements, by an Ewha University research grant to Y-JW, and by an Ewha University research grant to JCC.
You can also search for this author in PubMed Google Scholar. YJ participated in the design of the study and the writing of the manuscript, as well as conducting all cricket sampling in the field and performing all statistical analyses. Y-JW participated in the design of the study and the writing of the manuscript, as well as conducting all morphological measurements.
JCC coordinated the entire project and participated in writing of the manuscript. In addition, all authors read and approved the final manuscript. Additional file 1: G. See the Method for definitions of morphological characters. See Table 1 for sample sizes. DOC 44 KB. Reprints and Permissions. Jang, Y. Convergent and divergent patterns of morphological differentiation provide more evidence for reproductive character displacement in a wood cricket Gryllus fultoni Orthoptera: Gryllidae.
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Abstract Background In ecological character displacement, traits involved in reproductive isolation may not evolve in arbitrary directions when changes in these traits are by-products of adaptation to an ecological niche. Results Except for mirror area, the number of teeth in a file, and ovipositor length of G. Conclusion Divergence-enhancing selection may be acting on mirror area as well as calling song characters, whereas local adaptation or clinal effects may explain variation in other morphological characters in sympatric populations of G.
Background Once the subject of controversy, character displacement, a pattern in which the difference between two species is accentuated in areas of sympatry and is reduced in areas of allopatry [ 1 , 2 ], is now recognized as a powerful force driving trait diversification and even speciation in sympatry [ 3 — 8 ]. Figure 1. Full size image.
Table 1 Collecting localities of G. Full size table. Results Male Morphological Characters of G. Table 2 The analyses of the multivariate general liner models on male morphological variables. Table 3 Post hoc analyses of pairwise comparisons of male morphological characters in G.
Figure 2. Figure 3. Table 4 The analyses of the multivariate general liner models on female morphological variables. Discussion The most obvious pattern of morphological variation in both G.
Conclusion The pattern of morphological differentiation in sympatry and allopatry adds one more line of evidence for reproductive character displacement in the acoustic communication of G.
Methods Study Species G. Morphological Measurements All morphological measurements were conducted on the first-generation offspring of the field-caught females.
Statistical Analysis Because of significant correlations among morphological characters, we used multivariate general linear models GLM to test whether far allopatric, near allopatric, and sympatric populations of G. References 1. Article Google Scholar 2.
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