Eur. J. Entomol. 117: 118-128, 2020 | DOI: 10.14411/eje.2020.013

Contribution of population-level phenotypic plasticity to the invasiveness of Zaprionus indianus (Diptera: Drosophilidae)Original article

Nicolás José LAVAGNINO, Marcos IMBERTI, Nicolás FLAIBANI, Victoria Estefanía ORTIZ, Juan José FANARA
Laboratorio de Evolución, Departamento de Ecología, Genética y Evolución, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, IEGEBA-CONICET, Ciudad Universitaria, Pabellón II, Buenos Aires 1428, Argentina; e-mails: nlavagnino@gmail.com, paicos@gmail.com, n.flaiba@gmail.com, vicrotas@gmail.com, juan_jose_fanara@yahoo.com

Zaprionus indianus is a species of fly native to the Afrotropical biogeographic region, which around twenty years ago invaded the American continent. Several studies have shown that local adaptation and phenotypic plasticity of an invasive species in its native range could favour the colonization of new environments. Zaprionus indianus is a holometabolous generalist polyphagous species that breeds and feeds on the fruits of several different species, which constitute different environments. In this context, we performed a comparative analysis of the phenotypic plasticity of morphological and life history traits in response to seven different breeding environments (i.e. different breeding fruits). The comparison was of native (Africa) vs. invaded range (South America) wild-derived populations. The population-level phenotypic plasticity values related to heterogeneity in different breeding environments for most traits analysed were higher for one of the native range population. This differentiation was also recorded for the ranking across breeding environments of developmental time and wing length mean phenotypic values. In addition, mean phenotypic values pooled across fruit treatments were larger for individuals from the invaded range, which suggests local adaptation. Results define a scenario in which, although not for all the populations analysed, phenotypic plasticity contributes to the invasiveness and local adaptation in native range population of Z. indianus.

Keywords: Diptera, Drosophilidae, Zaprionus indianus, phenotypic plasticity, life history traits, morphological traits, invaded range, native range

Received: September 24, 2019; Revised: March 11, 2020; Accepted: March 11, 2020; Published online: March 24, 2020  Show citation

ACS AIP APA ASA Harvard Chicago Chicago Notes IEEE ISO690 MLA NLM Turabian Vancouver
LAVAGNINO, N.J., IMBERTI, M., FLAIBANI, N., ORTIZ, V.E., & FANARA, J.J. (2020). Contribution of population-level phenotypic plasticity to the invasiveness of Zaprionus indianus (Diptera: Drosophilidae). EJE, 117, Article 118-128. https://doi.org/10.14411/eje.2020.013
Share...
Download citation
  • BibTeX (.bib)
  • Bookends (.ris)
  • EasyBib (.ris)
  • EndNote (.enw)
  • EndNote 8 (.xml)
  • ISI WoS (.isi)
  • Medlars (.medlars)
  • Mendeley (.ris)
  • MODS (.xml)
  • Papers (.ris)
  • RefWorks (.txt)
  • RefManager (.ris)
  • RIS (.ris)
  • MS Word (.xml)
  • Zotero (.ris)
    • Open full article

    References

    1. Acurio A.E. & Rafael V.L. 2009: Inventario taxonómico de Drosophilidae (Diptera) en el Parque Nacional Yasuni amazonia ecuatoriana. - Acta Amazon. 39: 713-718. Go to original source...
    2. Aluja M. & Mangan R.L. 2007: Fruit fly (Diptera: Tephritidae) host status determination: critical conceptual, methodological, and regulatory considerations. - Annu. Rev. Entomol. 53: 473-502. Go to original source...
    3. Balaguer L., Martínez-Ferri E., Valladares F., Pérez-Corona M.E., Baquedano F.J., Castillo F.J. & Manrique E. 2001: Population divergence in the plasticity of the response of Quercus coccifera to the light environment. - Funct. Ecol. 15: 124-135. Go to original source...
    4. Bates D., Maechler M., Bolker B. & Walker S. 2015: Fitting linear mixed-effects models using lme4. - J. Stat. Soft. 67: 1-48. Go to original source...
    5. Benítez Díaz E.A. 2015: Primer registro de la Mosca Africana del Higo, Zaprionus indianus (Diptera: Drosophilidae) en Paraguay. - Bol. Mus. Nac. Hist. Nat. Parag. 19: 100-110.
    6. Benjamini Y. & Hochberg Y. 1995: Controlling the false discovery rate: A practical and powerful approach to multiple testing. - J. Roy. Stat. Soc. (B) 57: 289-300. Go to original source...
    7. Bitner-Mathé B.C. & David J.R. 2015: Genetic variability and phenotypic plasticity of metric thoracic traits in an invasive drosophilid in America. - Genetica 143: 441-451. Go to original source...
    8. Blackburn T.M., Py¹ek P., Bacher S., Carlton J.T., Duncan R.P., Jaro¹ík V., Wilson J.R.U. & Richardson D.M. 2011: A proposed unified framework for biological invasions. - Trends Ecol. Evol. 26: 333-339. Go to original source...
    9. Callaway R., Pennings S.C. & Richards C.L. 2003: Phenotypic plasticity and interactions among plants. - Ecology 84: 115-128. Go to original source...
    10. Carreira V.J., Mensch J. & Fanara J.J. 2009: Body size in Drosophila: genetic architecture, allometries and sexual dimorphism. - Heredity 102: 246-256. Go to original source...
    11. Carreira V.P., Mensch J., Hasson E. & Fanara J.J. 2016: Natural genetic variation and candidate genes for morphological traits in Drosophila melanogaster. - PLoS ONE 11: e0160069, 26 pp. Go to original source...
    12. Castrezana S. 2007: New records of Zaprionus indianus Gupta, 1970 (Diptera, Drosophilidae) in North America and a key to identify some Zaprionus species deposited in the Drosophila Tucson Stock Center. - Dros. Inf. Serv. 90: 34-36.
    13. Castrezana S. 2011: New record on novel hosts for the drosophilid pest Zaprionus indianus. - Dros. Inf. Serv. 94: 1-2.
    14. Castro F.L. & Valente V.L.S. 2001: Zaprionus indianus invading communities in the southern Brazilian city of Porto Alegre. - Dros. Inf. Serv. 84: 15-17.
    15. Chassagnard M.T. & Kraaijeveld A.R. 1991: The occurrence of Zaprionus sensu stricto in the Paleartic region (Diptera, Drosophilidae). - Ann. Soc. Entomol. Fr. 27: 495-496. Go to original source...
    16. Chaves N.B. & Tidon R. 2008: Biogeographical aspects of drosophilids (Diptera, Drosophilidae) of the Brazilian savanna. - Rev. Bras. Entomol. 52: 340-348. Go to original source...
    17. Chun Y.J., Collyer M.L., Moloney K.A. & Nason J.D. 2007: Phenotypic plasticity of native vs. invasive purple loosestrife: a two-state multivariate approach. - Ecology 88: 1499-1512. Go to original source...
    18. Commar L.S., Galego L.G., Ceron C.R. & Carareto C.M.A. 2012: Taxonomic and evolutionary analysis of Zaprionus indianus and its colonization of Palearctic and Neotropical regions. - Genet. Mol. Biol. 35: 395-406. Go to original source...
    19. David J.R., Araripe L.O., Bitner-Mathé B.C., Capy P., Goñi B., Klaczko L.B., Legout H., Martins M.B., Vouidibio J., Yassin A. & Moreteau B. 2006a: Quantitative trait analysis and geographic variability of natural populations of Zaprionus indianus, a recent invader in Brazil. - Heredity 96: 53-62. Go to original source...
    20. David J.R., Araripe L.O., Bitner-Mathé B.C., Capy P., Goñi B., Klaczko L.B., Legout H., Martins M.B., Vouidibio J., Yassin A. & Moreteau B. 2006b: Sexual dimorphism of body size and sternopleural bristle number: a comparison of geographic populations of an invasive cosmopolitan drosophilid. - Genetica 128: 109-122. Go to original source...
    21. Davidson A.M., Jennions M. & Nicotra A.B. 2011: Do invasive species show higher phenotypic plasticity than native species and, if so, is it adaptive? A meta-analysis. - Ecol. Lett. 14: 419-431. Go to original source...
    22. De Toni D.C., Hofmann P.R.P. & Valente V.L.S. 2001: First record of Zaprionus indianus (Diptera, Drosophilidae) in the State of Santa Catarina, Brazil. - Biotemas 14: 71-85.
    23. Einhorn K.S. 2005: Growth and physiology of ash (Fraxinus excelsior) and beech (Fagus sylvatica) seedlings in response to a light gradient following natural gap formation. - Ecol. Bull. 52: 1-53.
    24. Fernandes Rodrigues D.R. & Araújo E.L. 2011: Ocorrência de Zaprionus indianus Gupta (Diptera: Drosophilidae) em frutos de juazeiro Ziziphus joazeiro Mart. (Rhamnaceae) no estado do Rio Grande do Norte. - Rev. Bras. Frut. 33: 1356-1358. Go to original source...
    25. Ferreira Mendes M., Berti Valer F., Aleixo Vieira J.G., Laner Blauth M. & Gottschalk M.S. 2017: Diversity of Drosophilidae (Insecta, Diptera) in the Restinga forest of southern Brazil. - Rev. Bras. Entomol. 61: 248-256. Go to original source...
    26. Fordyce J.A. 2006: The evolutionary consequences of ecological interactions mediated through phenotypic plasticity. - J. Exp. Biol. 209: 2377-2383. Go to original source...
    27. Forsman A. 2015: Rethinking phenotypic plasticity and its consequences for individuals, populations and species. - Heredity 115: 276-284. Go to original source...
    28. Fox J. & Weisberg S. 2019: An {R} Companion to Applied Regression. 3rd ed. Sage, Thousand Oaks, CA, 607 pp.
    29. Furtado I.S., Martins M.B. & Costa J.E. 2009: First record of Zaprionus indianus (Diptera: Drosophilidae) in the Urucu Petroleum Province in Amazonas, Brazil. - Dros. Inf. Serv. 92: 17-18.
    30. Galego L.G.C. & Carareto C.M.A. 2007: Analysis of the drosophilid Zaprionus indianus introduction in Brazil: Contribution of esterase loci polymorphisms. - Dros. Inf. Serv. 90: 79-84.
    31. Gianoli E. & Valladares F. 2012: Studying phenotypic plasticity: the advantages of a broad approach. - Biol. J. Linn. Soc. 105: 1-7. Go to original source...
    32. Goñi B., Fresia P., Calviño M., Ferreiro M.J. & Valente V.L.S. 2001: First record of Zaprionus indianus Gupta, 1970 (Diptera, Drosophilidae) in southern localities of Uruguay. - Dros. Inf. Serv. 84: 61-65.
    33. Goñi B., Martinez M.E., Techera G. & Fresia P. 2002: Increased frequencies of Zaprionus indianus Gupta, 1970 (Diptera, Drosophilidae) in Uruguay. - Dros. Inf. Serv. 85: 75-80.
    34. Gratani L., Meneghini M., Pesoli P. & Crescente M.F. 2003: Structural and functional plasticity of Quercus ilex seedlings of different provenances in Italy. - Oecologia 17: 515-521. Go to original source...
    35. Gupta J.P. 1970: Description of a new species of Phorticella and Zaprionus (Drosophilidae) from India. - Proc. Indian Natl. Sci. Acad. (B) 36: 62-70.
    36. Harrison X.A. 2014: Using observation-level random effects to model overdispersion in count data in ecology and evolution. - PeerJ. 2: e616, 19 pp. Go to original source...
    37. Holle S.G., Tran A.K., Burkness E.C., Ebbenga D.N. & Hutchison W.D. 2019: First detections of Zaprionus indianus (Diptera: Drosophilidae) in Minnesota. - J. Entomol. Sci. 54: 99-102. Go to original source...
    38. Hulme P.E. 2008: Phenotypic plasticity and plant invasions: Is it all Jack? - Funct. Ecol. 22: 3-7. Go to original source...
    39. Joshi N.K., Biddinger D.J., Demchak K. & Deppen A. 2014: First report of Zaprionus indianus (Diptera: Drosophilidae) in commercial fruits and vegetables in Pennsylvania. - J. Insect Sci. 14: 259, 4 pp. Go to original source...
    40. Karan D., Moreteau B. & David J.R. 1999: Growth temperature and reaction norms of morphometrical traits in a tropical drosophilid: Zaprionus indianus. - Heredity 83: 398-407. Go to original source...
    41. Kato C.M., Foureaux L.V., César R.A. & Torres M.P. 2004: Occurrence of Zaprionus indianus Gupta, 1970 (Diptera: Drosophilidae) in Minas Gerais State. - Ciência Agrotecnol. 28: 454-455. Go to original source...
    42. Kaufman S.R. & Smouse P.E. 2001: Comparing indigenous and introduced populations of Melaleuca quinquenervia (Cay.) Blake: Response of seedlings to water and pH levels. - Oecologia 127: 487-494. Go to original source...
    43. Lachaise D. & Tsacas L. 1983: Breeding-sites in tropical African drosophilids. In Ashburner M., Carson H.L. & Thompson J.N. (eds): Genetics and Biology of Drosophila. Academic Press, New York, pp. 221-332.
    44. Lasa R. & Tadeo E. 2015: Invasive drosophilid pests Drosophila suzukii and Zaprionus indianus (Diptera: Drosophilidae) in Veracruz, Mexico. - Fla Entomol. 98: 987-988. Go to original source...
    45. Lavagnino N.J., Carreira V.P., Mensch J., Hasson E. & Fanara J.J. 2008: Geographic distribution and hosts of Zaprionus indianus (Diptera: Drosophilidae) in North-Eastern Argentina. - Rev. Soc. Entomol. Argent. 67: 189-192.
    46. Lavagnino N.J., Imberti M., Ortiz V.E., Flaibani N. & Fanara J.J. 2019: Contrasting levels of genotype by environment interaction for life history and morphological traits in invasive populations of Zaprionus indianus (Diptera: Drosophilidae). - Insect Sci. [in press]. Go to original source...
    47. Leao B.F.D. & Tidon R. 2004: Newly invading species exploiting native host-plants: the case of the African Zaprionus indianus (Gupta) in the Brazilian Cerrado (Diptera, Drosophilidae). - Ann. Soc. Entomol. Fr. 40: 285-290. Go to original source...
    48. Loh R. & Bitner-Mathé B.C. 2005: Variability of wing size and shape in three populations of a recent Brazilian invader, Zaprionus indianus (Diptera: Drosophilidae), from different habitats. - Genetica 125: 271-281. Go to original source...
    49. Loh R., David J.R., Debat V. & Bitner-Mathé B.C. 2008: Adaptation to different climates results in divergent phenotypic plasticity of wing size and shape in an invasive drosophilid. - J. Genet. 87: 209-217. Go to original source...
    50. Maclean H.J., Kristensen T.N., Sørensen J.G. & Overgaard J. 2018: Laboratory maintenance does not alter ecological and physiological patterns among species: A Drosophila case study. - J. Evol. Biol. 31: 530-542. Go to original source...
    51. Markow T.A., Hanna G., Riesgo-Escovar J.R., Tellez-Garcia A.A., Richmond M.P., Nazario-Yepiz N.O., Ramírez Loustalot Laclette M., Carpinteyro-Ponce J. & Pfeiler E. 2014: Population genetics and recent colonization history of the invasive drosophilid Zaprionus indianus in Mexico and Central America. - Biol. Invas. 16: 2427-2434. Go to original source...
    52. Matesanz S., Gianoli E. & Valladares F. 2010: Global change and the evolution of phenotypic plasticity in plants. - Ann. N.Y. Acad. Sci. 1206: 35-55. Go to original source...
    53. Miner B.G., Sultan S.E., Morgan S.G., Padilla D.K. & Relyea R.A. 2005: Ecological consequences of phenotypic plasticity. - Trends Ecol. Evol. 20: 685-692. Go to original source...
    54. Morton J.F. 2013: Fruits of Warm Climates. Echo Point Books & Media, Vermont, 550 pp.
    55. Norry F.M., Vilardi J.C., Fanara J.J. & Hasson E. 1994: Courtship success and multivariate analysis of sexual selection on morphometric traits in Drosophila buzzatii (Diptera: Drosophilidae). - J. Insect Behav. 8: 219-229. Go to original source...
    56. Oliveira G.F., de Melo K.P.S., Garcia A.C.L. & Rohde C. 2009: First record of Zaprionus indianus (Diptera, Drosophilidae) in Fernando de Noronha, an Oceanic Island of Pernambuco State, Brazil. - Dros. Inf. Serv. 92: 18-20.
    57. Parker I.M., Rodriguez J. & Loik M.E. 2003: An evolutionary approach to understanding the biology of invasions: Local adaptation and general-purpose genotypes in the weed Verbascum thapsus. - Conserv. Biol. 17: 59-72. Go to original source...
    58. Pasini M.P.B. & Link D. 2011: Efficiency of different traps to capture Zaprionus indianus (Diptera: Drosophilidae) in fig orchard in Santa Maria county, Rio Grande do Sul state, Brazil. - Int. J. Agric. Sci. Soil Sci. 1: 349-354.
    59. Pigliucci M. 2001: Phenotypic Plasticity: Beyond Nature and Nurture. Johns Hopkins University Press, Maryland, 348 pp. Go to original source...
    60. Pimentel D. 2011: Biological Invasions: Economic and Environmental Costs of Alien Plant, Animal, and Microbe Species. CRC Press, Boca Raton, FL, 463 pp. Go to original source...
    61. Poppe J.L., Schmitz H.J., Grimaldi D. & Da Silva Valente V.L. 2014: High diversity of Drosophilidae (Insecta, Diptera) in the Pampas Biome of South America, with descriptions of new Rhinoleucophenga species. - Zootaxa 3779: 215-245. Go to original source...
    62. Renkema J.M., Miller M., Fraser H., Légaré J.P. & Hallett R.H. 2013: First records of Zaprionus indianus Gupta (Diptera: Drosophilidae) from commercial fruit fields in Ontario and Quebec, Canada. - J. Entomol. Soc. Ont. 144: 125-130. Go to original source...
    63. Reznick D. & Travis J. 1996: The empirical study of adaptation in natural populations. In Rose M.R. & Lauder G.V. (eds): Adaptation. Academic Press, San Diego, pp. 243-289.
    64. Ribeiro Barbosa M.R., Graciolli G. & Paiva F. 2012: Diptera, Drosophilidae, Zaprionus indianus Gupta, 1970: Distribution extension for the state of Mato Grosso do Sul, Brazil. - Check List 8: 175-176. Go to original source...
    65. Ricciardi A. & Atkinson S.K. 2004: Distinctiveness magnifies the impact of biological invaders in aquatic ecosystems. - Ecol. Lett. 7: 781-784. Go to original source...
    66. Richards C.L., Bossdorf O., Muth N.Z., Gurevitch J. & Pigliucci M. 2006: Jack of all trades, master of some? On the role of phenotypic plasticity in plant invasions. - Ecol. Lett. 9: 981-993. Go to original source...
    67. R Core Team 2016: A Language and Environment for Statistical Computing. URL: http://www.R-project.org/
    68. Roff D.A. 1992: The Evolution of Life Histories: Theory and Analysis. Chapman & Hall, New York, 535 pp.
    69. Rohlf F.J. 2001: tpsDig. 1.31 Version. URL: http://life.bio.sunysb.edu/morph/
    70. Sakai A.K., Allendorf F.W., Holt J.S., Lodge D.M., Molofsky J., With K.A., Baughman S., Cabin R.J., Cohen J.E., Ellstrand N.C. et al. 2001: The population biology of invasive species. - Annu. Rev. Ecol. Evol. Syst. 32: 305-332. Go to original source...
    71. Santos J.F., Rieger T.T., Campos S.R.C., Nascimento A.C.C., Félix P.T., Silva S.V.O. & Freitas F.M.R. 2003: Colonization of northeast region of Brazil by the drosophilid flies Drosophila malerkotliana and Zaprionus indianus, a new potential insect pest for Brazilian fruticulture. - Dros. Inf. Serv. 86: 83-125.
    72. Schlichting C.D. & Pigliucci M. 1998: Phenotypic Evolution: A Reaction Norm Perspective. Sinauer, Sunderland, 340 pp.
    73. Schmitz H.J., Valente V.L.S. & Hofmann P.R.P. 2007: Taxonomic survey of Drosophildae (Diptera) from mangrove forests of Santa Catarina Island, Southern Brazil. - Neotrop. Entomol. 36: 53-64. Go to original source...
    74. Seebens H., Blackburn T.M., Dyer E.E., Genovesi P., Hulme P.E., Jeschke J.M., Pagad S., Py¹ek P., Winter M., Arianoutsou M. et al. 2017: No saturation in the accumulation of alien species worldwide. - Nat. Comm. 8: 14435, 9 pp. Go to original source...
    75. Sexton J.P., McKay J.K. & Sala A. 2002: Plasticity and genetic diversity may allow saltcedar to invade cold climates in North America. - Ecol. Appl. 12: 1652-1660. Go to original source...
    76. Soto I., Corio C., Fanara J.J. & Hasson E. 2006: First record of Zaprionus indianus Gupta 1970 (Diptera, Drosophilidae) in Argentina. - Dros. Inf. Serv. 89: 13-14.
    77. Stearns S.C. 1992: The Evolution of Life Histories. Oxford University Press, Oxford, 249 pp.
    78. Tidon R., Leite D.F. & Leao B.F.D. 2003: Impact of the colonisation of Zaprionus indianus (Diptera, Drosophilidae) in different ecosystems of the Neotropical Region: 2 years after the invasion. - Biol. Conserv. 112: 299-305. Go to original source...
    79. Tsutsui N.D. & Suarez A.V. 2003: The colony structure and population biology of invasive ants. - Conserv. Biol. 17: 48-58. Go to original source...
    80. Valladares F., Sanchez-Gomez D. & Zavala M.A. 2006: Quantitative estimation of phenotypic plasticity: bridging the gap between the evolutionary concept and its ecological applications. - J. Ecol. 94: 1103-1116. Go to original source...
    81. Van der Linde K., Steck G.J., Hibbard K., Birdsley J.S., Alonso L.M. & Houle D. 2006: First records of Zaprionus indianus (Diptera: Drosophilidae), a pest species on commercial fruits from Panama and The United States of America. - Fla Entomol. 89: 402-404. Go to original source...
    82. Van Kleunen M., Dawson W., Schlaepfer D., Jeschke J.M. & Fischer M. 2010: Are invaders different? A conceptual framework of comparative approaches for assessing determinants of invasiveness. - Ecol. Lett. 13: 947-958. Go to original source...
    83. Van Timmeren S. & Isaacs R. 2014: Drosophila suzukii in Michigan vineyards, and the first report of Zaprionus indianus from this region. - J. Appl. Entomol. 138: 519-527. Go to original source...
    84. Vasconcelos A., Azevedo T. & Santos R.S. 2017: Primeiro registro de Zaprionus indianus Gupta (Diptera: Drosophilidae) no Estado do Acre, Brasil. - EntomoBrasilis 10: 60-63. Go to original source...
    85. Vilela C.R. 1999: Is Zaprionus indianus Gupta, 1970 (Diptera: Drosophilidae) currently colonizing the Neotropical region? - Dros. Inf. Serv. 82: 37-39.
    86. Vilela C.R., Teixeira E.P. & Stein C.P. 2001: Mosca africana do figo, Zaprionus indianus (Diptera: Drosophilidae). In Vilela E.F., Zucchi R.A. & Cantor F. (eds): Historico e Impacto das Pragas Introduzidas no Brasil. Holos, Ribeirao Preto, pp. 48-52.
    87. Yassin A., Araripe L.O., Capy P., Da Lage J.L., Klaczko L.B., Maisonhaute C., Ogereau D. & David J.R. 2008a: Grafting the molecular phylogenetic tree with morphological branches to reconstruct the evolutionary history of the genus Zaprionus (Diptera: Drosophilidae). - Mol. Phylogenet. Evol. 47: 903-915. Go to original source...
    88. Yassin A., Capy P., Madi-Ravazzi L., Ogereau D. & David J.R. 2008b: DNA barcode discovers two cryptic species and two geographical radiations in the invasive drosophilid Zaprionus indianus. - Mol. Ecol. Resour. 8: 491-501. Go to original source...
    89. Zenni R.D., Lamy J.B., Lamarque L.J. & Port A.J. 2014: Adaptive evolution and phenotypic plasticity during naturalization and spread of invasive species: implications for tree invasion biology. - Biol. Invas. 16: 635-644. Go to original source...
    90. Zuur A.F., Leno E.N., Walker N.J., Saveliev A.A. & Smith G.M. 2009: Mixed Effects Models and Extensions in Ecology with R. Springer, New York, 574 pp. Go to original source...

    This is an open access article distributed under the terms of the Creative Commons Attribution 4.0 International License (CC BY 4.0), which permits use, distribution, and reproduction in any medium, provided the original publication is properly cited. No use, distribution or reproduction is permitted which does not comply with these terms.


    Return to the content