Costa rica

We need scarcely add that the contemplation in natural science of a wider domain than the actual leads to a far better understanding of the actual.' (p. 267,The, Nature of the Physical World.)x PREFACE evolutionary theory was thus chiefly retrogressive, the mighty body of Mendelian researches throughout the world has evidently out- grown the fallacies with which it was at first fostered.As a pioneer of genetics he has done more than enough to expiate the rash polemics of his early writings.To treat Natural Selection as an agency based independently on its own foundations is not to mimimize its importance in the theory of evolution.On the contrary, as soon as we require to form opinions by other means than by comparison and analogy, such an indepen- dent deductive basis becomes a necessity.This necessity is particu- larly to be noted for mankind; since we have some knowledge of the structure of society, of human motives, and of the vital statistics of this species, the use of the deductive method can supply a more intimate knowledge of the evolutionary processes than is elsewhere possible.In addition it will be of importance for our subject to call) attention to several consequences of the principle of Natural Selection!which, since they do not consist in the adaptive modification of specific I forms, have necessarily escaped attention.The genetic phenomena of I dominance and linkage seem to offer examples of this class, the future ' investigation of which may add greatly to the scope of our subject.No efforts of mine could avail to make the book easy reading.I have endeavoured to assist the reader by giving short summaries at the ends of all chapters, except Chapter IV, which is summarized conjointly with Chapter V.Those who prefer to do so may regardChapter IV as a mathematical appendix to the corresponding part of the summary.The deductions respecting Man are strictly in- separable from the more general chapters, but have been placed together in a group commencing with Chapter VIII.I believe no one will be surprised that a large number of the points considered demand a far fuller, more rigorous, and more comprehensive treat- ment.It seems impossible that full justice should be done to the subject in this way, until there is built up a tradition of mathematical work devoted to biological problems, comparable to the researches upon which a mathematical physicist can draw in the resolution of special difficulties.

@book{doi105962bhltitle27468,
    author = "Fisher, Ronald Aylmer",
    title = "The genetical theory of natural selection",
    year = "1930",
    booktitle = "Clarendon Press eBooks",
    abstract = "We need scarcely add that the contemplation in natural science of a wider domain than the actual leads to a far better understanding of the actual.' (p. 267,The, Nature of the Physical World.)x PREFACE evolutionary theory was thus chiefly retrogressive, the mighty body of Mendelian researches throughout the world has evidently out- grown the fallacies with which it was at first fostered.As a pioneer of genetics he has done more than enough to expiate the rash polemics of his early writings.To treat Natural Selection as an agency based independently on its own foundations is not to mimimize its importance in the theory of evolution.On the contrary, as soon as we require to form opinions by other means than by comparison and analogy, such an indepen- dent deductive basis becomes a necessity.This necessity is particu- larly to be noted for mankind; since we have some knowledge of the structure of society, of human motives, and of the vital statistics of this species, the use of the deductive method can supply a more intimate knowledge of the evolutionary processes than is elsewhere possible.In addition it will be of importance for our subject to call) attention to several consequences of the principle of Natural Selection!which, since they do not consist in the adaptive modification of specific I forms, have necessarily escaped attention.The genetic phenomena of I dominance and linkage seem to offer examples of this class, the future ' investigation of which may add greatly to the scope of our subject.No efforts of mine could avail to make the book easy reading.I have endeavoured to assist the reader by giving short summaries at the ends of all chapters, except Chapter IV, which is summarized conjointly with Chapter V.Those who prefer to do so may regardChapter IV as a mathematical appendix to the corresponding part of the summary.The deductions respecting Man are strictly in- separable from the more general chapters, but have been placed together in a group commencing with Chapter VIII.I believe no one will be surprised that a large number of the points considered demand a far fuller, more rigorous, and more comprehensive treat- ment.It seems impossible that full justice should be done to the subject in this way, until there is built up a tradition of mathematical work devoted to biological problems, comparable to the researches upon which a mathematical physicist can draw in the resolution of special difficulties.",
    url = "https://doi.org/10.5962/bhl.title.27468",
    doi = "10.5962/bhl.title.27468",
    openalex = "W2004778468",
    references = "darwin2009the, doi101017cbo9780511693946006, doi101017cbo9780511702884, doi101038033529a0, doi101111j136523111908tb02141x, doi1023071929022, doi1023074345450, doi105962bhltitle121292, doi105962bhltitle61004, doi105962bhltitle87899, openalexw2163836228"
}

@article{odonald1970polymorphic,
    author = "O'Donald, Peter and Pilecki, Christine",
    title = "Polymorphic Mimicry and Natural Selection",
    year = "1970",
    journal = "Evolution",
    url = "https://doi.org/10.2307/2406813",
    doi = "10.2307/2406813",
    number = "2",
    openalex = "W4243948369",
    pages = "395",
    volume = "24"
}

The natural color pattern of individuals of the unpalatable and mimetic butterfly Heliconius erato was altered to a unique nonmimetic pattern. When returned to natural populations, the nonmimetic individuals remained for shorter periods of time and received more wing damage indicative of predator attacks than did the controls. The results indicate that Müllerian mimicry was functioning to protect the butterflies from predation.

@article{benson1972natural,
    author = "Benson, Woodruff W.",
    title = "Natural Selection for Müllerian Mimicry in Heliconius erato in Costa Rica",
    year = "1972",
    journal = "Science",
    abstract = "The natural color pattern of individuals of the unpalatable and mimetic butterfly Heliconius erato was altered to a unique nonmimetic pattern. When returned to natural populations, the nonmimetic individuals remained for shorter periods of time and received more wing damage indicative of predator attacks than did the controls. The results indicate that Müllerian mimicry was functioning to protect the butterflies from predation.",
    url = "https://doi.org/10.1126/science.176.4037.936",
    doi = "10.1126/science.176.4037.936",
    number = "4037",
    pages = "936-939",
    volume = "176"
}

@misc{benson1972natural1,
    author = "Benson, W. W",
    title = "Natural selection for Mllerian mimicry in Heliconus erato in Costa Rica",
    year = "1972",
    howpublished = "Science, v. 176, p. 936-939",
    note = "talkorigins\_source = {true}; raw\_reference = {Benson, W. W., 1972, Natural selection for Mllerian mimicry in Heliconus erato in Costa Rica: Science, v. 176, p. 936-939.}"
}

Journal Article Why male butterflies are non-mimetic: natural selection, sexual selection, group selection, modification and sieving Get access JOHN R. G. TURNER JOHN R. G. TURNER 1Department of Ecology and Evolution, Division of Biological Sciences, State University of New York, Stony Brook, New York 11794, U.S.A Search for other works by this author on: Oxford Academic Google Scholar Biological Journal of the Linnean Society, Volume 10, Issue 4, December 1978, Pages 385–432, https://doi.org/10.1111/j.1095-8312.1978.tb00023.x Published: 14 January 2008 Article history Accepted: 01 November 1977 Published: 14 January 2008

@article{doi101111j109583121978tb00023x,
    author = "Turner, John R.",
    title = "Why male butterflies are non-mimetic: natural selection, sexual selection, group selection, modification and sieving*",
    year = "1978",
    journal = "Biological Journal of the Linnean Society",
    abstract = "Journal Article Why male butterflies are non-mimetic: natural selection, sexual selection, group selection, modification and sieving Get access JOHN R. G. TURNER JOHN R. G. TURNER 1Department of Ecology and Evolution, Division of Biological Sciences, State University of New York, Stony Brook, New York 11794, U.S.A Search for other works by this author on: Oxford Academic Google Scholar Biological Journal of the Linnean Society, Volume 10, Issue 4, December 1978, Pages 385–432, https://doi.org/10.1111/j.1095-8312.1978.tb00023.x Published: 14 January 2008 Article history Accepted: 01 November 1977 Published: 14 January 2008",
    url = "https://doi.org/10.1111/j.1095-8312.1978.tb00023.x",
    doi = "10.1111/j.1095-8312.1978.tb00023.x",
    openalex = "W1978034514",
    references = "doi1023072989703"
}

NATURAL SELECTION ON COLOR PATTERNS IN POECILIA RETICULATA

@article{doi101111j155856461980tb04790x,
    author = "Endler, John A.",
    title = "NATURAL SELECTION ON COLOR PATTERNS IN POECILIA RETICULATA",
    year = "1980",
    journal = "Evolution",
    abstract = "NATURAL SELECTION ON COLOR PATTERNS IN POECILIA RETICULATA",
    url = "https://doi.org/10.1111/j.1558-5646.1980.tb04790.x",
    doi = "10.1111/j.1558-5646.1980.tb04790.x",
    openalex = "W2006099940",
    references = "doi10100797814615695655, doi1010160022519375901113, doi101111j155856461975tb00851x, doi101146annureves08110177000545, doi1023071378997, doi1023071437762, doi1023071788577, doi1023072401303, doi105962bhltitle69899, doi105962bhltitle87588, openalexw1486180449, openalexw2151993477"
}

No abstract available. ECA Estudios Centroamericanos, Vol. 42, No. 466-467, 1987: 547-555.

@article{solísrivera1987costa,
    author = "Solís Rivera, Luis Guillermo",
    title = "Costa Rica",
    year = "1987",
    journal = "ECA: Estudios Centroamericanos",
    abstract = "No abstract available. ECA Estudios Centroamericanos, Vol. 42, No. 466-467, 1987: 547-555.",
    url = "https://doi.org/10.51378/eca.v42i466-467.8428",
    doi = "10.51378/eca.v42i466-467.8428",
    number = "466-467",
    openalex = "W4401634131",
    pages = "547-555",
    volume = "42"
}

Frequency-dependent selection on warning color can maintain narrow hybrid zones between unpalatable prey taxa. To measure such selection, we transferred marked Heliconius erato (Lepidoptera: Nymphalidae) in both directions across a 10-km-wide hybrid zone between Peruvian races differing in color pattern. These experimental H. erato were released at four sites, along with control H. erato of the phenotype native to each site. Survival of experimental butterflies was significantly lower than that of controls at two sites and overall. Most selection, measured as differences in survival, occurred soon after release. Selection against foreign morphs was 52% (confidence limits: 25-71 %) and was probably due to bird attacks on unusual warning-color morphs (more than 10% of the recaptures had beak marks). Since only three major loci determine the color-pattern differences, this suggests an average selection coefficient of 0.17 per locus, sufficient to maintain the narrow clines in H. erato.

@article{doi101111j155856461989tb04237x,
    author = "Mallet, James and Barton, Nick",
    title = "STRONG NATURAL SELECTION IN A WARNING-COLOR HYBRID ZONE",
    year = "1989",
    journal = "Evolution",
    abstract = "Frequency-dependent selection on warning color can maintain narrow hybrid zones between unpalatable prey taxa. To measure such selection, we transferred marked Heliconius erato (Lepidoptera: Nymphalidae) in both directions across a 10-km-wide hybrid zone between Peruvian races differing in color pattern. These experimental H. erato were released at four sites, along with control H. erato of the phenotype native to each site. Survival of experimental butterflies was significantly lower than that of controls at two sites and overall. Most selection, measured as differences in survival, occurred soon after release. Selection against foreign morphs was 52\% (confidence limits: 25-71 \%) and was probably due to bird attacks on unusual warning-color morphs (more than 10\% of the recaptures had beak marks). Since only three major loci determine the color-pattern differences, this suggests an average selection coefficient of 0.17 per locus, sufficient to maintain the narrow clines in H. erato.",
    url = "https://doi.org/10.1111/j.1558-5646.1989.tb04237.x",
    doi = "10.1111/j.1558-5646.1989.tb04237.x",
    openalex = "W2332616442",
    references = "benson1972natural, doi101007bf02986626, doi101038hdy196642, doi101038hdy197987, doi101086409995, doi101093biomet5212225, doi101093genetics754733, doi101098rstb19850066, doi101111j109583121986tb01772x, doi101111j109583121987tb00435x, doi101111j155856461967tb00126x, doi101111j155856461969tb03516x, doi101126science1443615183, doi101146annureves16110185000553, doi1023072407738, doi1023072989703, doi105962p203311, openalexw3133798068"
}

Hybrid zones can yield estimates of natural selection and gene flow. The width of a cline in gene frequency is approximately proportional to gene flow (sigma) divided by the square root of per-locus selection (square root of s). Gene flow also causes gametic correlations (linkage disequilibria) between genes that differ across hybrid zones. Correlations are stronger when the hybrid zone is narrow, and rise to a maximum roughly equal to s. Thus cline width and gametic correlations combine to give estimates of gene flow and selection. These indirect measures of sigma and s are especially useful because they can be made from collections, and require no field experiments. The method was applied to hybrid zones between color pattern races in a pair of Peruvian Heliconius butterfly species. The species are Müllerian mimics of one another, and both show the same changes in warning color pattern across their respective hybrid zones. The expectations of cline width and gametic correlation were generated using simulations of clines stabilized by strong frequency-dependent selection. In the hybrid zone in Heliconius erato, clines at three major color pattern loci were between 8.5 and 10.2 km wide, and the pairwise gametic correlations peaked at R approximately 0.35. These measures suggest that s approximately 0.23 per locus, and that sigma approximately 2.6 km. In erato, the shapes of the clines agreed with that expected on the basis of dominance. Heliconius melpomene has a nearly coincident hybrid zone. In this species, cline widths at four major color pattern loci varied between 11.7 and 13.4 km. Pairwise gametic correlations peaked near R approximately 1.00 for tightly linked genes, and at R approximately 0.40 for unlinked genes, giving s approximately 0.25 per locus and sigma approximately 3.7 km. In melpomene, cline shapes did not perfectly fit theoretical shapes based on dominance; this deviation might be explained by long-distance migration and/or strong epistasis. Compared with erato, sample sizes in melpomene are lower and the genetics of its color patterns are less well understood. In spite of these problems, selection and gene flow are clearly of the same order of magnitude in the two species. The relatively high per locus selection coefficients agree with "major gene" theories for the evolution of Müllerian mimicry, but the genetic architecture of the color patterns does not. These results show that the genetics and evolution of mimicry are still only sketchily understood.

@article{doi101093genetics1244921,
    author = "Mallet, James and Barton, Nick and Lamas, Gerardo and Santisteban, James Godde Maria S. and Muedas, M and Eeley, H",
    title = "Estimates of selection and gene flow from measures of cline width and linkage disequilibrium in heliconius hybrid zones.",
    year = "1990",
    journal = "Genetics",
    abstract = {Hybrid zones can yield estimates of natural selection and gene flow. The width of a cline in gene frequency is approximately proportional to gene flow (sigma) divided by the square root of per-locus selection (square root of s). Gene flow also causes gametic correlations (linkage disequilibria) between genes that differ across hybrid zones. Correlations are stronger when the hybrid zone is narrow, and rise to a maximum roughly equal to s. Thus cline width and gametic correlations combine to give estimates of gene flow and selection. These indirect measures of sigma and s are especially useful because they can be made from collections, and require no field experiments. The method was applied to hybrid zones between color pattern races in a pair of Peruvian Heliconius butterfly species. The species are Müllerian mimics of one another, and both show the same changes in warning color pattern across their respective hybrid zones. The expectations of cline width and gametic correlation were generated using simulations of clines stabilized by strong frequency-dependent selection. In the hybrid zone in Heliconius erato, clines at three major color pattern loci were between 8.5 and 10.2 km wide, and the pairwise gametic correlations peaked at R approximately 0.35. These measures suggest that s approximately 0.23 per locus, and that sigma approximately 2.6 km. In erato, the shapes of the clines agreed with that expected on the basis of dominance. Heliconius melpomene has a nearly coincident hybrid zone. In this species, cline widths at four major color pattern loci varied between 11.7 and 13.4 km. Pairwise gametic correlations peaked near R approximately 1.00 for tightly linked genes, and at R approximately 0.40 for unlinked genes, giving s approximately 0.25 per locus and sigma approximately 3.7 km. In melpomene, cline shapes did not perfectly fit theoretical shapes based on dominance; this deviation might be explained by long-distance migration and/or strong epistasis. Compared with erato, sample sizes in melpomene are lower and the genetics of its color patterns are less well understood. In spite of these problems, selection and gene flow are clearly of the same order of magnitude in the two species. The relatively high per locus selection coefficients agree with "major gene" theories for the evolution of Müllerian mimicry, but the genetic architecture of the color patterns does not. These results show that the genetics and evolution of mimicry are still only sketchily understood.},
    url = "https://doi.org/10.1093/genetics/124.4.921",
    doi = "10.1093/genetics/124.4.921",
    openalex = "W2110474028",
    references = "doi105962p203311"
}

The classification of reproductive isolating barriers laid out by Dobzhansky and Mayr has motivated and structured decades of research on speciation. We argue, however, that this classification is incomplete and that the unique contributions of a major source of reproductive isolation have often been overlooked. Here, we describe reproductive barriers that derive from the reduced survival of immigrants upon reaching foreign habitats that are ecologically divergent from their native habitat. This selection against immigrants reduces encounters and thus mating opportunities between individuals from divergently adapted populations. It also reduces the likelihood that successfully mated immigrant females will survive long enough to produce their hybrid offspring. Thus, natural selection against immigrants results in distinctive elements of premating and postmating reproductive isolation that we hereby dub "immigrant inviability." We quantify the contributions of immigrant inviability to total reproductive isolation by examining study systems where multiple components of reproductive isolation have been measured and demonstrate that these contributions are frequently greater than those of traditionally recognized reproductive barriers. The relevance of immigrant inviability is further illustrated by a consideration of population-genetic theory, a review of selection against immigrant alleles in hybrid zone studies, and an examination of its participation in feedback loops that influence the evolution of additional reproductive barriers. Because some degree of immigrant inviability will commonly exist between populations that exhibit adaptive ecological divergence, we emphasize that these barriers play critical roles in ecological modes of speciation. We hope that the formal recognition of immigrant inviability and our demonstration of its evolutionary importance will stimulate more explicit empirical studies of its contributions to speciation.

@article{doi10155404428,
    author = "Nosil, Patrik and Vines, Tim and Funk, Daniel J.",
    title = "PERSPECTIVE: REPRODUCTIVE ISOLATION CAUSED BY NATURAL SELECTION AGAINST IMMIGRANTS FROM DIVERGENT HABITATS",
    year = "2005",
    journal = "Evolution",
    abstract = {The classification of reproductive isolating barriers laid out by Dobzhansky and Mayr has motivated and structured decades of research on speciation. We argue, however, that this classification is incomplete and that the unique contributions of a major source of reproductive isolation have often been overlooked. Here, we describe reproductive barriers that derive from the reduced survival of immigrants upon reaching foreign habitats that are ecologically divergent from their native habitat. This selection against immigrants reduces encounters and thus mating opportunities between individuals from divergently adapted populations. It also reduces the likelihood that successfully mated immigrant females will survive long enough to produce their hybrid offspring. Thus, natural selection against immigrants results in distinctive elements of premating and postmating reproductive isolation that we hereby dub "immigrant inviability." We quantify the contributions of immigrant inviability to total reproductive isolation by examining study systems where multiple components of reproductive isolation have been measured and demonstrate that these contributions are frequently greater than those of traditionally recognized reproductive barriers. The relevance of immigrant inviability is further illustrated by a consideration of population-genetic theory, a review of selection against immigrant alleles in hybrid zone studies, and an examination of its participation in feedback loops that influence the evolution of additional reproductive barriers. Because some degree of immigrant inviability will commonly exist between populations that exhibit adaptive ecological divergence, we emphasize that these barriers play critical roles in ecological modes of speciation. We hope that the formal recognition of immigrant inviability and our demonstration of its evolutionary importance will stimulate more explicit empirical studies of its contributions to speciation.},
    url = "https://doi.org/10.1554/04-428",
    doi = "10.1554/04-428",
    openalex = "W2153863517",
    references = "doi10103835077075, doi101111j155856461989tb04237x"
}

@incollection{crossref2007costa,
    title = "Costa Rica – Costa Rica",
    year = "2007",
    booktitle = "Index of North and South American Constitutions 1850 to 2007",
    url = "https://doi.org/10.1515/9783110968002.189",
    doi = "10.1515/9783110968002.189",
    openalex = "W2494154018",
    pages = "189-193"
}

Biological evolution is a fact--but the many conflicting theories of evolution remain controversial even today. In 1966, simple Darwinism, which holds that evolution functions primarily at the level of the individual organism, was threatened by opposing concepts such as group selection, a popular idea stating that evolution acts to select entire species rather than individuals. George Williams's famous argument in favor of the Darwinists struck a powerful blow to those in opposing camps. His Adaptation and Natural Selection, now a classic of science literature, is a thorough and convincing essay in defense of Darwinism; its suggestions for developing effective principles for dealing with the evolution debate and its relevance to many fields outside biology ensure the timelessness of this critical work.

@book{doi1015159781400820108,
    author = "Williams, George C.",
    title = "Adaptation and Natural Selection",
    year = "2008",
    booktitle = "Princeton University Press eBooks",
    abstract = "Biological evolution is a fact--but the many conflicting theories of evolution remain controversial even today. In 1966, simple Darwinism, which holds that evolution functions primarily at the level of the individual organism, was threatened by opposing concepts such as group selection, a popular idea stating that evolution acts to select entire species rather than individuals. George Williams's famous argument in favor of the Darwinists struck a powerful blow to those in opposing camps. His Adaptation and Natural Selection, now a classic of science literature, is a thorough and convincing essay in defense of Darwinism; its suggestions for developing effective principles for dealing with the evolution debate and its relevance to many fields outside biology ensure the timelessness of this critical work.",
    url = "https://doi.org/10.1515/9781400820108",
    doi = "10.1515/9781400820108",
    openalex = "W2020289104"
}

Wing pattern evolution in Heliconius butterflies provides some of the most striking examples of adaptation by natural selection. The genes controlling pattern variation are classic examples of Mendelian loci of large effect, where allelic variation causes large and discrete phenotypic changes and is responsible for both convergent and highly divergent wing pattern evolution across the genus. We characterize nucleotide variation, genotype-by-phenotype associations, linkage disequilibrium (LD), and candidate gene expression patterns across two unlinked genomic intervals that control yellow and red wing pattern variation among mimetic forms of Heliconius erato. Despite very strong natural selection on color pattern, we see neither a strong reduction in genetic diversity nor evidence for extended LD across either patterning interval. This observation highlights the extent that recombination can erase the signature of selection in natural populations and is consistent with the hypothesis that either the adaptive radiation or the alleles controlling it are quite old. However, across both patterning intervals we identified SNPs clustered in several coding regions that were strongly associated with color pattern phenotype. Interestingly, coding regions with associated SNPs were widely separated, suggesting that color pattern alleles may be composed of multiple functional sites, conforming to previous descriptions of these loci as "supergenes." Examination of gene expression levels of genes flanking these regions in both H. erato and its co-mimic, H. melpomene, implicate a gene with high sequence similarity to a kinesin as playing a key role in modulating pattern and provides convincing evidence for parallel changes in gene regulation across co-mimetic lineages. The complex genetic architecture at these color pattern loci stands in marked contrast to the single casual mutations often identified in genetic studies of adaptation, but may be more indicative of the type of genetic changes responsible for much of the adaptive variation found in natural populations.

@article{doi101371journalpgen1000796,
    author = "Counterman, Brian A. and Araújo-Pérez, Félix and Hines, Heather M. and Baxter, Simon W. and Morrison, Clay and Lindstrom, Daniel P. and Papa, Riccardo and Ferguson, Laura and Joron, Mathieu and ffrench‐Constant, Richard H. and Smith, Christopher P. and Nielsen, Dahlia M. and Chen, Rui and Jiggins, Chris D. and Reed, Robert D. and Halder, Georg and Mallet, Jim and McMillan, W. Owen",
    title = "Genomic Hotspots for Adaptation: The Population Genetics of Müllerian Mimicry in Heliconius erato",
    year = "2010",
    journal = "PLoS Genetics",
    abstract = {Wing pattern evolution in Heliconius butterflies provides some of the most striking examples of adaptation by natural selection. The genes controlling pattern variation are classic examples of Mendelian loci of large effect, where allelic variation causes large and discrete phenotypic changes and is responsible for both convergent and highly divergent wing pattern evolution across the genus. We characterize nucleotide variation, genotype-by-phenotype associations, linkage disequilibrium (LD), and candidate gene expression patterns across two unlinked genomic intervals that control yellow and red wing pattern variation among mimetic forms of Heliconius erato. Despite very strong natural selection on color pattern, we see neither a strong reduction in genetic diversity nor evidence for extended LD across either patterning interval. This observation highlights the extent that recombination can erase the signature of selection in natural populations and is consistent with the hypothesis that either the adaptive radiation or the alleles controlling it are quite old. However, across both patterning intervals we identified SNPs clustered in several coding regions that were strongly associated with color pattern phenotype. Interestingly, coding regions with associated SNPs were widely separated, suggesting that color pattern alleles may be composed of multiple functional sites, conforming to previous descriptions of these loci as "supergenes." Examination of gene expression levels of genes flanking these regions in both H. erato and its co-mimic, H. melpomene, implicate a gene with high sequence similarity to a kinesin as playing a key role in modulating pattern and provides convincing evidence for parallel changes in gene regulation across co-mimetic lineages. The complex genetic architecture at these color pattern loci stands in marked contrast to the single casual mutations often identified in genetic studies of adaptation, but may be more indicative of the type of genetic changes responsible for much of the adaptive variation found in natural populations.},
    url = "https://doi.org/10.1371/journal.pgen.1000796",
    doi = "10.1371/journal.pgen.1000796",
    openalex = "W2047862598",
    references = "doi105962p203304"
}

Mimicry--when one organism (the mimic) evolves a phenotypic resemblance to another (the model) due to selective benefits--is widely used to illustrate natural selection's power to generate adaptations. However, many putative mimics resemble their models imprecisely, and such imperfect mimicry represents a specific challenge to mimicry theory and a general one to evolutionary theory. Here, we discuss 11 nonmutually exclusive hypotheses for imperfect mimicry. We group these hypotheses according to whether imperfect mimicry reflects: an artifact of human perception, which is not shared by any naturally occurring predators and therefore is not truly an instance of imperfect mimicry; genetic, developmental or time-lag constraints, which (temporarily) prevent a response to selection for perfect mimicry; relaxed selection, where imperfect mimicry is as adaptive as perfect mimicry; or tradeoffs, where imperfect mimicry is (locally) more adaptive than perfect mimicry. We find that the relaxed selection hypothesis has garnered the most support. However, because only a few study systems have thus far been comprehensively evaluated, the relative contributions of the various hypotheses toward explaining the evolution of imperfect mimicry remain unclear. Ultimately, clarifying why imperfect mimicry exists should provide critical insights into the limits of natural selection in producing complex adaptations.

@article{doi101086673758,
    author = "Kikuchi, David W. and Pfennig, David W.",
    title = "Imperfect Mimicry and the Limits of Natural Selection",
    year = "2013",
    journal = "The Quarterly Review of Biology",
    abstract = "Mimicry--when one organism (the mimic) evolves a phenotypic resemblance to another (the model) due to selective benefits--is widely used to illustrate natural selection's power to generate adaptations. However, many putative mimics resemble their models imprecisely, and such imperfect mimicry represents a specific challenge to mimicry theory and a general one to evolutionary theory. Here, we discuss 11 nonmutually exclusive hypotheses for imperfect mimicry. We group these hypotheses according to whether imperfect mimicry reflects: an artifact of human perception, which is not shared by any naturally occurring predators and therefore is not truly an instance of imperfect mimicry; genetic, developmental or time-lag constraints, which (temporarily) prevent a response to selection for perfect mimicry; relaxed selection, where imperfect mimicry is as adaptive as perfect mimicry; or tradeoffs, where imperfect mimicry is (locally) more adaptive than perfect mimicry. We find that the relaxed selection hypothesis has garnered the most support. However, because only a few study systems have thus far been comprehensively evaluated, the relative contributions of the various hypotheses toward explaining the evolution of imperfect mimicry remain unclear. Ultimately, clarifying why imperfect mimicry exists should provide critical insights into the limits of natural selection in producing complex adaptations.",
    url = "https://doi.org/10.1086/673758",
    doi = "10.1086/673758",
    openalex = "W2233326040",
    references = "doi101098rspb19940102, doi101098rspb19970022, doi101111j109583121981tb01842x"
}

Carbonate communities: The activity of anaerobic methane oxidizing microbes facilitates precipitation of vast quantities of authigenic carbonate at methane seeps. Here we demonstrate the significant role of carbonate rocks in promoting diversity by providing unique habitat and food resources for macrofaunal assemblages at seeps on the Costa Rica margin (400-1850 m). The attendant fauna is surprisingly similar to that in rocky intertidal shores, with numerous grazing gastropods (limpets and snails) as dominant taxa. However, the community feeds upon seep-associated microbes. Macrofaunal density, composition, and diversity on carbonates vary as a function of seepage activity, biogenic habitat and location. The macrofaunal community of carbonates at non-seeping (inactive) sites is strongly related to the hydrography (depth, temperature, O2) of overlying water, whereas the fauna at sites of active seepage is not. Densities are highest on active rocks from tubeworm bushes and mussel beds, particularly at the Mound 12 location (1000 m). Species diversity is higher on rocks exposed to active seepage, with multiple species of gastropods and polychaetes dominant, while crustaceans, cnidarians, and ophiuroids were better represented on rocks at inactive sites. Macro-infauna (larger than 0.3 mm) from tube cores taken in nearby seep sediments at comparable depths exhibited densities similar to those on carbonate rocks, but had lower diversity and different taxonomic composition. Seep sediments had higher densities of ampharetid, dorvilleid, hesionid, cirratulid and lacydoniid polychaetes, whereas carbonates had more gastropods, as well as syllid, chrysopetalid and polynoid polychaetes. Stable isotope signatures and metrics: The stable isotope signatures of carbonates were heterogeneous, as were the food sources and nutrition used by the animals. Carbonate δ13Cinorg values (mean = -26.98‰) ranged from -53.3‰ to +10.0‰, and were significantly heavier than carbonate δ13Corg (mean = -33.83‰), which ranged from -74.4‰ to -20.6‰. Invertebrates on carbonates had average δ13C (per rock) = -31.0‰ (range -18.5‰ to -46.5‰) and δ15N = 5.7‰ (range -4.5‰ to +13.4‰). Average δ13C values did not differ between active and inactive sites; carbonate fauna from both settings depend on chemosynthesis-based nutrition. Community metrics reflecting trophic diversity (SEAc, total Hull Area, ranges of δ13C and δ15N) and species packing (mean distance to centroid, nearest neighbor distance) also did not vary as a function of seepage activity or site. However, distinct isotopic signatures were observed among related, co-occurring species of gastropods and polychaetes, reflecting intense microbial resource partitioning. Overall, the substrate and nutritional heterogeneity introduced by authigenic seep carbonates act to promote diverse, uniquely adapted assemblages, even after seepage ceases. The macrofauna in these ecosystems remain largely overlooked in most surveys, but are major contributors to biodiversity of chemosynthetic ecosystems and the deep sea in general.

@article{doi101371journalpone0131080,
    author = "Levin, Lisa A. and Mendoza, Guillermo and Grupe, B and Gonzalez, Jennifer P. and Jellison, Brittany and Rouse, Greg W. and Thurber, Andrew R. and Warén, Anders",
    title = "Biodiversity on the Rocks: Macrofauna Inhabiting Authigenic Carbonate at Costa Rica Methane Seeps",
    year = "2015",
    journal = "PLoS ONE",
    abstract = "Carbonate communities: The activity of anaerobic methane oxidizing microbes facilitates precipitation of vast quantities of authigenic carbonate at methane seeps. Here we demonstrate the significant role of carbonate rocks in promoting diversity by providing unique habitat and food resources for macrofaunal assemblages at seeps on the Costa Rica margin (400-1850 m). The attendant fauna is surprisingly similar to that in rocky intertidal shores, with numerous grazing gastropods (limpets and snails) as dominant taxa. However, the community feeds upon seep-associated microbes. Macrofaunal density, composition, and diversity on carbonates vary as a function of seepage activity, biogenic habitat and location. The macrofaunal community of carbonates at non-seeping (inactive) sites is strongly related to the hydrography (depth, temperature, O2) of overlying water, whereas the fauna at sites of active seepage is not. Densities are highest on active rocks from tubeworm bushes and mussel beds, particularly at the Mound 12 location (1000 m). Species diversity is higher on rocks exposed to active seepage, with multiple species of gastropods and polychaetes dominant, while crustaceans, cnidarians, and ophiuroids were better represented on rocks at inactive sites. Macro-infauna (larger than 0.3 mm) from tube cores taken in nearby seep sediments at comparable depths exhibited densities similar to those on carbonate rocks, but had lower diversity and different taxonomic composition. Seep sediments had higher densities of ampharetid, dorvilleid, hesionid, cirratulid and lacydoniid polychaetes, whereas carbonates had more gastropods, as well as syllid, chrysopetalid and polynoid polychaetes. Stable isotope signatures and metrics: The stable isotope signatures of carbonates were heterogeneous, as were the food sources and nutrition used by the animals. Carbonate δ13Cinorg values (mean = -26.98‰) ranged from -53.3‰ to +10.0‰, and were significantly heavier than carbonate δ13Corg (mean = -33.83‰), which ranged from -74.4‰ to -20.6‰. Invertebrates on carbonates had average δ13C (per rock) = -31.0‰ (range -18.5‰ to -46.5‰) and δ15N = 5.7‰ (range -4.5‰ to +13.4‰). Average δ13C values did not differ between active and inactive sites; carbonate fauna from both settings depend on chemosynthesis-based nutrition. Community metrics reflecting trophic diversity (SEAc, total Hull Area, ranges of δ13C and δ15N) and species packing (mean distance to centroid, nearest neighbor distance) also did not vary as a function of seepage activity or site. However, distinct isotopic signatures were observed among related, co-occurring species of gastropods and polychaetes, reflecting intense microbial resource partitioning. Overall, the substrate and nutritional heterogeneity introduced by authigenic seep carbonates act to promote diverse, uniquely adapted assemblages, even after seepage ceases. The macrofauna in these ecosystems remain largely overlooked in most surveys, but are major contributors to biodiversity of chemosynthetic ecosystems and the deep sea in general.",
    url = "https://doi.org/10.1371/journal.pone.0131080",
    doi = "10.1371/journal.pone.0131080",
    openalex = "W1577451703",
    references = "doi101126science2264677965, doi10120197814200374493, doi1015159780691239477, doi105860choice380926, openalexw12294379"
}

@incollection{bergoeing2017natural,
    author = "Bergoeing, Jean Pierre",
    title = "Natural Shape Types in Costa Rica",
    year = "2017",
    booktitle = "Geomorphology and Volcanology of Costa Rica",
    url = "https://doi.org/10.1016/b978-0-12-812067-5.00002-2",
    doi = "10.1016/b978-0-12-812067-5.00002-2",
    openalex = "W2572293231",
    pages = "23-43"
}

Biological mimicry has served as a salient example of natural selection for over a century, providing us with a dazzling array of very different examples across many unrelated taxa. We provide a conceptual framework that brings together apparently disparate examples of mimicry in a single model for the purpose of comparing how natural selection affects models, mimics and signal receivers across different interactions. We first analyse how model–mimic resemblance likely affects the fitness of models, mimics and receivers across diverse examples. These include classic Batesian and Müllerian butterfly systems, nectarless orchids that mimic Hymenoptera or nectar‐producing plants, caterpillars that mimic inert objects unlikely to be perceived as food, plants that mimic abiotic objects like carrion or dung and aggressive mimicry where predators mimic food items of their own prey. From this, we construct a conceptual framework of the selective forces that form the basis of all mimetic interactions. These interactions between models, mimics and receivers may follow four possible evolutionary pathways in terms of the direction of selection resulting from model–mimic resemblance. Two of these pathways correspond to the selective pressures associated with what is widely regarded as Batesian and Müllerian mimicry. The other two pathways suggest mimetic interactions underpinned by distinct selective pressures that have largely remained unrecognized. Each pathway is characterized by theoretical differences in how model–mimic resemblance influences the direction of selection acting on mimics, models and signal receivers, and the potential for consequent (co)evolutionary relationships between these three protagonists. The final part of this review describes how selective forces generated through model–mimic resemblance can be opposed by the basic ecology of interacting organisms and how those forces may affect the symmetry, strength and likelihood of (co)evolution between the three protagonists within the confines of the four broad evolutionary possibilities. We provide a clear and pragmatic visualization of selection pressures that portrays how different mimicry types may evolve. This conceptual framework provides clarity on how different selective forces acting on mimics, models and receivers are likely to interact and ultimately shape the evolutionary pathways taken by mimetic interactions, as well as the constraints inherent within these interactions.

@article{anderson2020natural,
    author = "Anderson, Bruce and de Jager, Marinus L.",
    title = "Natural selection in mimicry",
    year = "2020",
    journal = "Biological Reviews",
    abstract = "Biological mimicry has served as a salient example of natural selection for over a century, providing us with a dazzling array of very different examples across many unrelated taxa. We provide a conceptual framework that brings together apparently disparate examples of mimicry in a single model for the purpose of comparing how natural selection affects models, mimics and signal receivers across different interactions. We first analyse how model–mimic resemblance likely affects the fitness of models, mimics and receivers across diverse examples. These include classic Batesian and Müllerian butterfly systems, nectarless orchids that mimic Hymenoptera or nectar‐producing plants, caterpillars that mimic inert objects unlikely to be perceived as food, plants that mimic abiotic objects like carrion or dung and aggressive mimicry where predators mimic food items of their own prey. From this, we construct a conceptual framework of the selective forces that form the basis of all mimetic interactions. These interactions between models, mimics and receivers may follow four possible evolutionary pathways in terms of the direction of selection resulting from model–mimic resemblance. Two of these pathways correspond to the selective pressures associated with what is widely regarded as Batesian and Müllerian mimicry. The other two pathways suggest mimetic interactions underpinned by distinct selective pressures that have largely remained unrecognized. Each pathway is characterized by theoretical differences in how model–mimic resemblance influences the direction of selection acting on mimics, models and signal receivers, and the potential for consequent (co)evolutionary relationships between these three protagonists. The final part of this review describes how selective forces generated through model–mimic resemblance can be opposed by the basic ecology of interacting organisms and how those forces may affect the symmetry, strength and likelihood of (co)evolution between the three protagonists within the confines of the four broad evolutionary possibilities. We provide a clear and pragmatic visualization of selection pressures that portrays how different mimicry types may evolve. This conceptual framework provides clarity on how different selective forces acting on mimics, models and receivers are likely to interact and ultimately shape the evolutionary pathways taken by mimetic interactions, as well as the constraints inherent within these interactions.",
    url = "https://doi.org/10.1111/brv.12564",
    doi = "10.1111/brv.12564",
    number = "2",
    openalex = "W2982527475",
    pages = "291-304",
    volume = "95",
    references = "doi10103714088000, doi101093acprofoso97801985286090010001, doi101098rspb19790081, doi101111j109636421860tb00146x, doi105860choice432194, doi105962bhltitle59991, doi105962bhltitle82303, doi107208chicago97802261186970010001, openalexw12294379, openalexw1594469865"
}

@incollection{peterman2021costa,
    author = "Peterman, Keith and Cordes, Matthew",
    title = "Costa Rica: Natural Capital",
    year = "2021",
    booktitle = "ACS Symposium Series",
    url = "https://doi.org/10.1021/bk-2021-1382.ch018",
    doi = "10.1021/bk-2021-1382.ch018",
    openalex = "W4293166939",
    pages = "211-223",
    references = "doi101126scienceaaz8901, doi1023073983623"
}

The constraints in traditional music style transfer algorithms are difficult to control, thereby making it challenging to balance the diversity and quality of the generated music. This paper proposes a novel weak selection-based music generation algorithm that aims to enhance both the quality and the diversity of conditionally generated traditional diffusion model audio, and the proposed algorithm is applied to generate natural sleep music. In the inference generation process of natural sleep music, the evolutionary state is determined by evaluating the evolutionary factors in each iteration, while limiting the potential range of evolutionary rates of weak selection-based traits to increase the diversity of sleep music. Subjective and objective evaluation results reveal that the natural sleep music generated by the proposed algorithm has a more significant hypnotic effect than general sleep music and conforms to the rules of human hypnosis physiological characteristics.

@article{doi103390math11153345,
    author = "Huang, Wenkai and Zhan, Feng",
    title = "A Novel Probabilistic Diffusion Model Based on the Weak Selection Mimicry Theory for the Generation of Hypnotic Songs",
    year = "2023",
    journal = "Mathematics",
    abstract = "The constraints in traditional music style transfer algorithms are difficult to control, thereby making it challenging to balance the diversity and quality of the generated music. This paper proposes a novel weak selection-based music generation algorithm that aims to enhance both the quality and the diversity of conditionally generated traditional diffusion model audio, and the proposed algorithm is applied to generate natural sleep music. In the inference generation process of natural sleep music, the evolutionary state is determined by evaluating the evolutionary factors in each iteration, while limiting the potential range of evolutionary rates of weak selection-based traits to increase the diversity of sleep music. Subjective and objective evaluation results reveal that the natural sleep music generated by the proposed algorithm has a more significant hypnotic effect than general sleep music and conforms to the rules of human hypnosis physiological characteristics.",
    url = "https://doi.org/10.3390/math11153345",
    doi = "10.3390/math11153345",
    openalex = "W4385420925",
    references = "doi101007s10682021101269"
}

Flight was a key innovation in the adaptive radiation of insects. However, it is a complex trait influenced by a large number of interacting biotic and abiotic factors, making it difficult to unravel the evolutionary drivers. We investigate flight patterns in neotropical heliconiine butterflies, well known for mimicry of their aposematic wing color patterns. We quantify the flight patterns (wing beat frequency and wing angles) of 351 individuals representing 29 heliconiine and 9 ithomiine species belonging to ten color pattern mimicry groupings. For wing beat frequency and up wing angles, we show that heliconiine species group by color pattern mimicry affiliation. Convergence of down wing angles to mimicry groupings is less pronounced, indicating that distinct components of flight are under different selection pressures and constraints. The flight characteristics of the Tiger mimicry group are particularly divergent due to convergence with distantly related ithomiine species. Predator-driven selection for mimicry also explained variation in flight among subspecies, indicating that this convergence can occur over relatively short evolutionary timescales. Our results suggest that the flight convergence is driven by aposematic signaling rather than shared habitat between comimics. We demonstrate that behavioral mimicry can occur between lineages that have separated over evolutionary timescales ranging from <0.5 to 70 My.

@article{doi101073pnas2300886121,
    author = "Page, E and Queste, Lucie M. and Rosser, Neil and Salazar, Patricio A. and Nadeau, Nicola J. and Mallet, James and Srygley, Robert B. and McMillan, W. Owen and Dasmahapatra, Kanchon K.",
    title = "Pervasive mimicry in flight behavior among aposematic butterflies",
    year = "2024",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "Flight was a key innovation in the adaptive radiation of insects. However, it is a complex trait influenced by a large number of interacting biotic and abiotic factors, making it difficult to unravel the evolutionary drivers. We investigate flight patterns in neotropical heliconiine butterflies, well known for mimicry of their aposematic wing color patterns. We quantify the flight patterns (wing beat frequency and wing angles) of 351 individuals representing 29 heliconiine and 9 ithomiine species belonging to ten color pattern mimicry groupings. For wing beat frequency and up wing angles, we show that heliconiine species group by color pattern mimicry affiliation. Convergence of down wing angles to mimicry groupings is less pronounced, indicating that distinct components of flight are under different selection pressures and constraints. The flight characteristics of the Tiger mimicry group are particularly divergent due to convergence with distantly related ithomiine species. Predator-driven selection for mimicry also explained variation in flight among subspecies, indicating that this convergence can occur over relatively short evolutionary timescales. Our results suggest that the flight convergence is driven by aposematic signaling rather than shared habitat between comimics. We demonstrate that behavioral mimicry can occur between lineages that have separated over evolutionary timescales ranging from <0.5 to 70 My.",
    url = "https://doi.org/10.1073/pnas.2300886121",
    doi = "10.1073/pnas.2300886121",
    openalex = "W4392167888",
    references = "doi101038s4158602407263w, doi101073pnas1907847116, doi101093beheco83239, doi101093bioinformaticsbty633, doi101111j109636421860tb00146x, doi101146annureven26010181002235, doi101186s128590171934z, doi1018637jssv067i01, doi1018637jssv082i13, doi1018901012641, openalexw3133798068"
}

@misc{crossrefNonecosta,
    title = "Costa Rica - Environment \& Natural Resources",
    year = "None",
    booktitle = "Foreign Law Guide",
    url = "https://doi.org/10.1163/2213-2996\_flg\_com\_057121",
    doi = "10.1163/2213-2996\_flg\_com\_057121",
    openalex = "W4251418147"
}