Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)

El pez león Pterois volitans ha invadido el Atlántico occidental causando una seria disminución en el reclutamiento de peces arrecifales, afectando poblaciones de competidores y no competidores de todas las tallas, lo cual ha alterado la complejidad de la red trófica y la estabilidad arrecifal. Esta...

Full description

Autores:
Torres Rodríguez, Javier Alfonso
Tipo de recurso:
Fecha de publicación:
2016
Institución:
Universidad Nacional de Colombia
Repositorio:
Universidad Nacional de Colombia
Idioma:
spa
OAI Identifier:
oai:repositorio.unal.edu.co:unal/75850
Acceso en línea:
https://repositorio.unal.edu.co/handle/unal/75850
Palabra clave:
Biología
Founder event, Pterois volitans, inbreeding, genetic drift, biological invasions, mitochondrial control region, microsatellites.
Evento fundador, Pterois volitans, endogamia, deriva génica, invasiones biológicas, región control mitocondrial, microsatélites.
Rights
openAccess
License
Atribución-NoComercial-SinDerivadas 4.0 Internacional
id UNACIONAL2_f42665eea2788935baf39da1729d26e0
oai_identifier_str oai:repositorio.unal.edu.co:unal/75850
network_acronym_str UNACIONAL2
network_name_str Universidad Nacional de Colombia
repository_id_str
dc.title.spa.fl_str_mv Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)
title Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)
spellingShingle Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)
Biología
Founder event, Pterois volitans, inbreeding, genetic drift, biological invasions, mitochondrial control region, microsatellites.
Evento fundador, Pterois volitans, endogamia, deriva génica, invasiones biológicas, región control mitocondrial, microsatélites.
title_short Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)
title_full Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)
title_fullStr Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)
title_full_unstemmed Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)
title_sort Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)
dc.creator.fl_str_mv Torres Rodríguez, Javier Alfonso
dc.contributor.advisor.spa.fl_str_mv Acero, Arturo
Betancur, Ricardo
dc.contributor.author.spa.fl_str_mv Torres Rodríguez, Javier Alfonso
dc.subject.ddc.spa.fl_str_mv Biología
topic Biología
Founder event, Pterois volitans, inbreeding, genetic drift, biological invasions, mitochondrial control region, microsatellites.
Evento fundador, Pterois volitans, endogamia, deriva génica, invasiones biológicas, región control mitocondrial, microsatélites.
dc.subject.proposal.eng.fl_str_mv Founder event, Pterois volitans, inbreeding, genetic drift, biological invasions, mitochondrial control region, microsatellites.
dc.subject.proposal.spa.fl_str_mv Evento fundador, Pterois volitans, endogamia, deriva génica, invasiones biológicas, región control mitocondrial, microsatélites.
description El pez león Pterois volitans ha invadido el Atlántico occidental causando una seria disminución en el reclutamiento de peces arrecifales, afectando poblaciones de competidores y no competidores de todas las tallas, lo cual ha alterado la complejidad de la red trófica y la estabilidad arrecifal. Esta invasión constituye una problemática prioritaria debido al potencial impacto que causa en el ambiente y al sector socioeconómico, por lo cual la aplicación de todas las herramientas científicas es urgente. En este estudio se evaluó la dinámica temporal de la diversidad genética de la invasión del pez león en dos localidades del Caribe (SA Caribe occidental y SM Caribe sur) a partir de dos marcadores moleculares, el primero consistió en un análisis de la región control mitocondrial de 432 individuos, el cual no reveló cambios temporales en ambas localidades, aunque se detectó la presencia del haplotipo H3 en un individuo en SA, lo cual obedece a una baja frecuencia de esta información. Se elaboraron y estandarizaron 10 loci tipo microsatélites que se amplificaron en 364 individuos, el análisis mostró un déficit de heterocigotos, lo cual puede estar relacionado con efectos de endogamia y deriva génica. El estudio reveló que la invasión se encuentra estructurada por seis stocks genéticamente diferentes (K = 6) de los cuales se encontraron cuatro en SA y cinco en SM, lo cual concuerda con variaciones espaciales y temporales relacionadas a diferencias en las tasas de reclutamiento de los individuos colonizadores de cada localidad.
publishDate 2016
dc.date.issued.spa.fl_str_mv 2016-08-25
dc.date.accessioned.spa.fl_str_mv 2020-03-04T21:39:28Z
dc.date.available.spa.fl_str_mv 2020-03-04T21:39:28Z
dc.type.spa.fl_str_mv Trabajo de grado - Maestría
dc.type.driver.spa.fl_str_mv info:eu-repo/semantics/masterThesis
dc.type.version.spa.fl_str_mv info:eu-repo/semantics/acceptedVersion
dc.type.content.spa.fl_str_mv Text
dc.type.redcol.spa.fl_str_mv http://purl.org/redcol/resource_type/TM
status_str acceptedVersion
dc.identifier.uri.none.fl_str_mv https://repositorio.unal.edu.co/handle/unal/75850
url https://repositorio.unal.edu.co/handle/unal/75850
dc.language.iso.spa.fl_str_mv spa
language spa
dc.relation.references.spa.fl_str_mv Ahrenholz DW y Morris J.A. 2010. Larval duration of the lionfish, Pterois volitans along the Bahamian Archipelago. Environmental Biology of Fishes, 88: 305-309. Albins MA y Hixon MA. 2008. Invasive Indo-Pacific lionfish Pterois volitans reduce recruitment of Atlantic coral-reef fishes. Marine Ecology Progress Series, 367: 233-238. Albins MA y Hixon MA. 2013. Worst case scenario: potential long-term effects of invasive predatory lionfish (Pterois volitans) on Atlantic and Caribbean coral-reef communities. Environmental Biology of Fishes, 96: 1151-1157. Arbeláez N, y Acero A. 2011. Presencia del pez león Pterois volitans (Linnaeus) en el manglar de la bahía de Chengue, Caribe colombiano. Boletín de Investigaciones Marinas y Costeras, 40 (2): 431-435. Arias-González JE, González-Gándara JC, Cabrera JL y Christensen V. 2011. Predicted impact of the invasive lionfish Pterois volitans on the food web of a Caribbean coral reef. Environmental Research, 111 (7): 917-925. Badii MH y Landeros J. 2007. Invasión de especies o el tercer jinete de Apocalipsis ambiental. International Journal of Good Conscience, 2(1): 39-53. Bandelt HJ, Forster P y Röhl A .1999. Median-joining networks for inferring intraspecific phylogenies. Molecular Biology and Evolution, 16 (1): 37-48. Barson NJ, Cable J, y Van Oosterhout C. 2009. Population genetic analysis of microsatellite variation of guppies (Poecilia reticulata) in Trinidad and Tobago: evidence for a dynamic source–sink metapopulation structure, founder events and population bottlenecks. Journal of Evolutionary Biology, 22 (3): 485-497. Bax N, Williamson A, Aguero M, Gonzalez E y Geeves W. 2003. Marine invasive alien species: a threat to global biodiversity. Marine policy, 27 (4): 313-323. Betancur R, Hines A, Acero A, Ortí G, Wilbur AE y Freshwater DW. 2011. Reconstructing the lionfish invasion: insights into Greater Caribbean biogeography. Journal of Biogeography, 38 (7): 1281-1293. Briggs JC. 2007. Marine biogeography and ecology: invasions and introductions. Journal of Biogeography, 34 (2): 193-198. Briggs JC. 2013. Invasion ecology: origin and biodiversity effects. Environmental Skeptics and Critics, 2 (3), 73-81. Butterfield JS, Díaz-Ferguson E, Silliman BR, Saunders JW, Buddo D, Mignucci-Giannoni A y Hunter ME. 2015. Wide-ranging phylogeographic structure of invasive red lionfish in the Western Atlantic and Greater Caribbean. Marine Biology, 162 (4): 773-781. Courtenay WR. 1995. Marine fish introductions in southeastern Florida. American Fisheries Society Introduced Fish Section Newsletter, 14 (1): 2-3. Darling ES, Green SJ, O’Leary JK y Côte IM. 2011. Indo-Pacific lionfish are larger and more abundant on invaded reefs: a comparison of Kenyan and Bahamian lionfish populations. Biological Invasions 13 (9): 2045-2051. DeWoody JA y Avise JC. 2000. Microsatellite variation in marine, freshwater and anadromous fishes compared with other animals. Journal of Fish Biology, 56 (3): 461-473. Dias P, Verheyen GR y Raymond M. 1996. Source-sink populations in Mediterranean blue tits: evidence using single locus minisatellite probes. Journal of Evolutionary Biology. 9 (6): 965-978. Earl DA. 2009. Structure Harvester v0. 3. [Internet]. 05 de Enero de 2016. Disponible en: <http://users. soe. ucsc. edu/~ dearl/software/struct_harvest>. Excoffier L, Smouse PE y Quattro JM. 1992. Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data. Genetics, 131 (2): 479-491. Excoffier L, Laval G y Schneider S. 2005. Arlequin (version 3.0): an integrated software package for population genetics data analysis. Evolutionary Bioinformatics, 1: 47-50. Evanno G, Regnaut S y Goudet J. 2005. Detecting the number of clusters of individuals using the software STRUCTURE: a simulation study. Molecular ecology, 14 (8): 2611-2620. Ferreira CE, Luiz OJ, Floeter SR, Lucena MB, Barbosa MC, Rocha CR y Rocha LA. 2015. First record of invasive lionfish (Pterois volitans) for the Brazilian coast. PLos ONE, 10(2): 1-5. Franco–Herrera A. 2005. Oceanografía de la ensenada de Gaira: el Rodadero, más que un centro turístico en el Caribe Colombiano, Bogotá: Fundación Universidad de Bogotá Jorge Tadeo Lozano. p. 56. Frankham R. 1995. Conservation genetics. Annual Review of Genetics, 29 (1): 305-327. Frankham R. 2005. Resolving the genetic paradox in invasive species. Heredity, 94: 385. Freshwater DW, Hines A, Parham S, Wilbur AE, Sabaoun M, Woodhead J, Akins JL, Purdy B, Whitfield PE y Paris CB. 2009. Mitochondrial control region sequence analyses indicate dispersal from the US East Coast as the source of the invasive Indo-Pacific lionfish Pterois volitans in the Bahamas. Marine Biology, 156: 1213-1221. Goudet J. 2001. FSTAT, a program to estimate and test gene diversities and fixation indices (version 2.9. 3). Lausanne University, Lausanne, Switzerland. Green SJ, Akins JL, Maljković A y Côté IM. 2012. Invasive lionfish drive Atlantic coral reef fish declines. PloS one, 7 (3):1-3. Guo SW y Thompson EA. 1992. Performing the exact test of Hardy–Weinberg proportion for multiple alleles. Biometrics 48 (2): 361-372. Haag CR, Riek M, Hottinger JW, Pajunen VI y Ebert D. 2006. Founder events as determinants of within-island and among-island genetic structure of Daphnia metapopulations. Heredity, 96 (2): 150-158. Hamner RM, Freshwater DW y Whitfield PE. 2007. Mitochondrial cytochrome b analysis reveals two invasive lionfish species with strong founder events in the western Atlantic. Journal of Fish Biology 71: 214-222. Holland BS. 2001. Invasion without a bottleneck: microsatellite variation in natural and invasive populations of the brown mussel Perna perna (L). Marine Biotechnology, 3 (5): 407-415. Hubisz MJ, Falush D, Stephens M, y Pritchard JK. 2009. Inferring weak population structure with the assistance of sample group information. Molecular Ecology Resources, 9 (5): 1322-1332. IGAC. 1986. San Andrés y Providencia. Aspectos Geográficos. Instituto Geográfico Agustín Codazzi. Bogotá. 156 p. Johnston MW, y Purkis SJ. 2015. Hurricanes accelerated the Florida–Bahamas lionfish invasion. Global change biology, 21 (6): 2249-2260. Kolbe JJ, Richard E, Glor RE, Rodríguez-Schettino L, Chamizo-Lara A, Larson A y Losos JB. 2004. Genetic variation increases during biological invasion by a Cuban lizard. Nature, 431, 177-181. Lande R. 1988. Genetics and demography in biological conservation. Science, 241 (4872), 1455-1460. Larkin MA, Blackshields G, Brown NP, Chenna R, McGettigan PA, McWilliam H y Higgins DG. 2007. Clustal W and Clustal X version 2.0.Bioinformatics, 23 (21): 2947-2948. Librado P y Rozas J. 2009. DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25(11): 1451-1452. Lindholm AK, Breden F, Alexander HJ, Chan WK, Thakurta SG. y Brooks R. 2005. Invasion success and genetic diversity of introduced populations of guppies Poecilia reticulata in Australia. Molecular Ecology, 14: 3671-3682. Luikart G, Sherwin WB, Steele BM y Allendorf FW. 1998. Usefulness of molecular markers for detecting population bottlenecks via monitoring genetic change. Molecular Ecology, 7(8): 963-974. Martínez-Viloria H, Gómez ÁR y Acero A. 2011. Presencia del pez león, Pterois volitans (Actinopterygii: Scorpaenidae) en el Departamento de la Guajira, Mar Caribe de Colombia. Boletín de Investigaciones Marinas y Costeras, 40 (2): 445-447. Mack RN, Simberloff D, Lonsdale MW, Evans H, Clout M y Bazzaz F. 2000. Invasiones Biológicas: Causas, Epidemiología, Consecuencias globales y Control. Tópicos en Ecología. 5:1-19. Meunier C, Tirard C, Hurtrez‐Boussès S, Durand P, Bargues MD, Mas‐Coma S y Renaud F. 2001. Lack of molluscan host diversity and the transmission of an emerging parasitic disease in Bolivia. Molecular Ecology, 10 (5):1333-1340. Miller SA, Dykes DD y Polesky H. 1988. A simple salting out procedure for extracting DNA from human nucleated cells. Nucleic acids research, 16 (3): 1215. Morris JA y Akins JL. 2009. Feeding ecology of the invasive lionfish in the Bahamian archipelago. Environmental Biology of Fishes. 86: 389-398. Morris JA y Whitfield PE. 2009. Biology, Ecology, Control and Management of the Invasive Indo-Pacific Lionfish: An Updated Integrated Assessment. NOAA Techn. Mem. NOS NCCOS 99. 57 pp. Nei M. 1987. Molecular Evolutionary Genetics Columbia University Press New York 512. O'reilly PT, Canino MF, Bailey KM y Bentzen P. 2004. Inverse relationship between FST and microsatellite polymorphism in the marine fish, walleye pollock (Theragra chalcogramma): implications for resolving weak population structure. Molecular Ecology, 13(7): 1799-1814. Peakall RO y Smouse PE. 2006. GENALEX 6: genetic analysis in Excel. Population genetic software for teaching and research. Molecular Ecology Notes, 6 (1): 288-295. Planes S y Lecaillon G. 1998. Consequences of the founder effect in the genetic structure of introduced island coral reef fish populations. Biological Journal of the Linnean Society, 63 (4):537-552. Pimentel D, Zuñiga R y Morrison D. 2005. Update on the environmental and economic costs associated with alien-invasive species in the United States. Ecological Economics. 52: 273-288. Posada D y Crandall KA. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics, 14 (9): 817-818. Prahl H y Erhardt H. 1985. Colombia: Corales y arrecifes coralinos. FEN Colombia. Fondo para la protección y medio ambiente “José Celestino Mutis”. Bogotá. Raymond M y Rousset F. 2003. A population genetic software for exact test and ecumenicism: GENEPOP, version 3.4. Heredity, 68: 248-249. Reed DH y Frankham R. 2003. Correlation between fitness and genetic diversity. Conservation Biology. 17: 230-237. Rius M, Turon X, Bernardi G, Volckaert FA y Viard F. 2015. Marine invasion genetics: from spatio-temporal patterns to evolutionary outcomes. Biological Invasions, 17 (3): 869-885. Roman J y Darling JA. 2007. Paradox lost: genetic diversity and the success of aquatic invasions. Trends in Ecology and Evolution, 22 (9): 454-464. Sawyer SA y Hartl DL. 1992. Population genetics of polymorphism and divergence. Genetics, 132 (4):1161-1176. Sax DF, Stachowicz JJ, Brown JH, Bruno JF, Dawson MN, Gaines SD, Grosberg RK, Hastings A, Holt RD, Mayfield MM, Mary I, O’Connor MI y Rice WR. 2007. Ecological and evolutionary insights from species invasions. Trends in Ecology and Ecology, 22 (9): 465-471. Schneider S, Roessli D y Excoffier L. 2000. Arlequin: a software for population genetics data analysis. User manual version, 2: 2496-2497. Schofield PJ. 2009. Geographic extent and chronology of the invasion of non-native lionfish (Pterois volitans [Linnaeus 1758] and P. miles [Bennett 1828]) in the Western North Atlantic and Caribbean Sea. Aquatic Invasions, 4: 443–449. Schultz TF, Fitzpatrick CK, Freshwater DW y Morris JA. 2013. Characterization of 18 polymorphic microsatellite loci from invasive lionfish (Pterois volitans and P. miles). Conservation Genetics Resources, 5 (3): 599-601. Sivasundar A, Bermingham E y Orti G. 2001. Population structure and biogeography of migratory freshwater fishes (Prochilodus: Characiformes) in major South American rivers. Molecular Ecology, 10 (2): 407-417. Sutherland WJ, Clout M, Côté IM, Daszak P, Depledge MH, Fellman L, Fleishman E, Garthwaite R, Gibbons DW, De Lurio J, Impey AJ, Lickorish F, Lindenmayer D, Madgwick J, Margerison C, Maynard T, Peck LP, Pretty J, Prior S, Redford KH, Scharlemann PW, Spalding M y Watkinson AR. 2010. A horizon scan of global conservation issues for 2010. Cell Press, 25: 1-7. Tajima F. 1989. Statistical method for testing the neutral mutation hypothesis by DNA polymorphism. Genetics, 123 (3): 585-595. Tamura K, Glen Stecher, Daniel Peterson, Alan Filipski y Sudhir Kumar. 2013. MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution, 30 (12): 2725-2729. Toledo-Hernández C, Vélez-Zuazo X, Ruiz-Diaz CP, Patricio AR, Mège P, Navarro M y Papa R. 2014. Population ecology and genetics of the invasive lionfish in Puerto Rico. Aquatic Invasions, 9 (2): 227-237. Torres-Rodriguez JA, Gonzales-Corredor JD y Acero A. Registro de un ejemplar de pez león (Pterois volitans) sin vestigios de aleta dorsal espinosa. En preparación. Triantafyllidis A, Abatzopoulos TJ, Leonardos J y Guyomard R. (2002). Microsatellite analysis of the genetic population structure of native and translocated Aristotle's catfish (Silurus aristotelis). Aquatic Living Resources, 15 (06): 351-359. Van Oosterhout C, Hutchinson WF, Wills DP y Shipley P. 2004. MICRO‐CHECKER: software for identifying and correcting genotyping errors in microsatellite data. Molecular Ecology Notes, 4 (3): 535-538. Vila C, Sundqvist AK, Flagstad Ø, Seddon J, Kojola I, Casulli A y Ellegren H. 2003. Rescue of a severely bottlenecked wolf (Canis lupus) population by a single immigrant. Proceedings of the Royal Society of London. Biological Sciences, 270 (1510): 91-97. Weir BS y Cockerham C. 1984. Estimating F-statistics for the analysis of population structure. Evolution, 38 (6): 1358-1370. Whitfield PE, Hare JA, David AW, Harter SL, Munoz RC y Addison CM. 2007. Abundance estimates of the Indo-Pacific lionfish Pterois volitans/miles complex in the Western North Atlantic. Biological Invasions, 9: 53-64. Willi Y, Van Buskirk J y Hoffmann AA. 2006. Limits to the adaptive potential of small populations. Annual Review of Ecology, Evolution, and Systematics, 37: 433-458.
dc.rights.spa.fl_str_mv Derechos reservados - Universidad Nacional de Colombia
dc.rights.coar.fl_str_mv http://purl.org/coar/access_right/c_abf2
dc.rights.license.spa.fl_str_mv Atribución-NoComercial-SinDerivadas 4.0 Internacional
dc.rights.spa.spa.fl_str_mv Acceso abierto
dc.rights.uri.spa.fl_str_mv http://creativecommons.org/licenses/by-nc-nd/4.0/
dc.rights.accessrights.spa.fl_str_mv info:eu-repo/semantics/openAccess
rights_invalid_str_mv Atribución-NoComercial-SinDerivadas 4.0 Internacional
Derechos reservados - Universidad Nacional de Colombia
Acceso abierto
http://creativecommons.org/licenses/by-nc-nd/4.0/
http://purl.org/coar/access_right/c_abf2
eu_rights_str_mv openAccess
dc.format.extent.spa.fl_str_mv 39p.
dc.format.mimetype.spa.fl_str_mv application/pdf
dc.publisher.department.spa.fl_str_mv Centro de estudios en Ciencias del mar-CECIMAR
dc.publisher.faculty.spa.fl_str_mv Facultad Caribe
dc.publisher.branch.spa.fl_str_mv Universidad Nacional de Colombia - Sede Caribe
institution Universidad Nacional de Colombia
bitstream.url.fl_str_mv https://repositorio.unal.edu.co/bitstream/unal/75850/1/80098976.2016.pdf
https://repositorio.unal.edu.co/bitstream/unal/75850/2/license.txt
https://repositorio.unal.edu.co/bitstream/unal/75850/3/license_rdf
https://repositorio.unal.edu.co/bitstream/unal/75850/4/80098976.2016.pdf.jpg
bitstream.checksum.fl_str_mv 9cb9eef1c1f5594754e57bd2bb7d263f
6f3f13b02594d02ad110b3ad534cd5df
217700a34da79ed616c2feb68d4c5e06
e56af216dcbf2748c308f8d948da7af5
bitstream.checksumAlgorithm.fl_str_mv MD5
MD5
MD5
MD5
repository.name.fl_str_mv Repositorio Institucional Universidad Nacional de Colombia
repository.mail.fl_str_mv repositorio_nal@unal.edu.co
_version_ 1806886514933628928
spelling Atribución-NoComercial-SinDerivadas 4.0 InternacionalDerechos reservados - Universidad Nacional de ColombiaAcceso abiertohttp://creativecommons.org/licenses/by-nc-nd/4.0/info:eu-repo/semantics/openAccesshttp://purl.org/coar/access_right/c_abf2Acero, Arturo663ab63d-7d1b-4316-8808-dd9d6031d4feBetancur, Ricardo63a1e14e-dd94-4953-be9e-70aa92b1b959Torres Rodríguez, Javier Alfonso127ef9e3-61c8-4467-a4f7-941a26fe84ed2020-03-04T21:39:28Z2020-03-04T21:39:28Z2016-08-25https://repositorio.unal.edu.co/handle/unal/75850El pez león Pterois volitans ha invadido el Atlántico occidental causando una seria disminución en el reclutamiento de peces arrecifales, afectando poblaciones de competidores y no competidores de todas las tallas, lo cual ha alterado la complejidad de la red trófica y la estabilidad arrecifal. Esta invasión constituye una problemática prioritaria debido al potencial impacto que causa en el ambiente y al sector socioeconómico, por lo cual la aplicación de todas las herramientas científicas es urgente. En este estudio se evaluó la dinámica temporal de la diversidad genética de la invasión del pez león en dos localidades del Caribe (SA Caribe occidental y SM Caribe sur) a partir de dos marcadores moleculares, el primero consistió en un análisis de la región control mitocondrial de 432 individuos, el cual no reveló cambios temporales en ambas localidades, aunque se detectó la presencia del haplotipo H3 en un individuo en SA, lo cual obedece a una baja frecuencia de esta información. Se elaboraron y estandarizaron 10 loci tipo microsatélites que se amplificaron en 364 individuos, el análisis mostró un déficit de heterocigotos, lo cual puede estar relacionado con efectos de endogamia y deriva génica. El estudio reveló que la invasión se encuentra estructurada por seis stocks genéticamente diferentes (K = 6) de los cuales se encontraron cuatro en SA y cinco en SM, lo cual concuerda con variaciones espaciales y temporales relacionadas a diferencias en las tasas de reclutamiento de los individuos colonizadores de cada localidad.The lion fish Pterois volitans has invaded the western Atlantic causing a serious decline in the recruitment of reef fish, affecting population’s competitors and non-competitors of all sizes, which has altered the complexity of the food web and reef stability. This invasion is a priority issue because of the potential impact caused on the environment and the socio-economic sector, for which the application of all scientific tools is urgent. In this study the temporal dynamics of the genetic diversity of the invasion of lionfish in two locations of the Caribbean (SA Western and Caribbean SM southern Caribbean) from two molecular markers was evaluated, the first was an analysis of the mitochondrial control region of 432 subjects, which revealed no temporal changes in both locations, although the presence of H3 haplotype was detected in an individual in SA, which is due to a low frequency of this information. 10 microsatellite loci were developed and standardized in 364 subjects, the analysis showed a deficit of heterozygotes, which may be related to effects of inbreeding and genetic drift. The study revealed that the invasion is structured by six different genetic stocks (K = 6) of which were found four in SA and five in SM, which is consistent with spatial and temporal variations related to differences in rates of recruitment of the colonizers of each locality.Maestría39p.application/pdfspaBiologíaFounder event, Pterois volitans, inbreeding, genetic drift, biological invasions, mitochondrial control region, microsatellites.Evento fundador, Pterois volitans, endogamia, deriva génica, invasiones biológicas, región control mitocondrial, microsatélites.Variación de la diversidad genética luego de un evento fundador en un pez invasor (Pterois volitans) (scorpaenidae)Trabajo de grado - Maestríainfo:eu-repo/semantics/masterThesisinfo:eu-repo/semantics/acceptedVersionTexthttp://purl.org/redcol/resource_type/TMCentro de estudios en Ciencias del mar-CECIMARFacultad CaribeUniversidad Nacional de Colombia - Sede CaribeAhrenholz DW y Morris J.A. 2010. Larval duration of the lionfish, Pterois volitans along the Bahamian Archipelago. Environmental Biology of Fishes, 88: 305-309. Albins MA y Hixon MA. 2008. Invasive Indo-Pacific lionfish Pterois volitans reduce recruitment of Atlantic coral-reef fishes. Marine Ecology Progress Series, 367: 233-238. Albins MA y Hixon MA. 2013. Worst case scenario: potential long-term effects of invasive predatory lionfish (Pterois volitans) on Atlantic and Caribbean coral-reef communities. Environmental Biology of Fishes, 96: 1151-1157. Arbeláez N, y Acero A. 2011. Presencia del pez león Pterois volitans (Linnaeus) en el manglar de la bahía de Chengue, Caribe colombiano. Boletín de Investigaciones Marinas y Costeras, 40 (2): 431-435. Arias-González JE, González-Gándara JC, Cabrera JL y Christensen V. 2011. Predicted impact of the invasive lionfish Pterois volitans on the food web of a Caribbean coral reef. Environmental Research, 111 (7): 917-925. Badii MH y Landeros J. 2007. Invasión de especies o el tercer jinete de Apocalipsis ambiental. International Journal of Good Conscience, 2(1): 39-53. Bandelt HJ, Forster P y Röhl A .1999. Median-joining networks for inferring intraspecific phylogenies. Molecular Biology and Evolution, 16 (1): 37-48. Barson NJ, Cable J, y Van Oosterhout C. 2009. Population genetic analysis of microsatellite variation of guppies (Poecilia reticulata) in Trinidad and Tobago: evidence for a dynamic source–sink metapopulation structure, founder events and population bottlenecks. Journal of Evolutionary Biology, 22 (3): 485-497. Bax N, Williamson A, Aguero M, Gonzalez E y Geeves W. 2003. Marine invasive alien species: a threat to global biodiversity. Marine policy, 27 (4): 313-323. Betancur R, Hines A, Acero A, Ortí G, Wilbur AE y Freshwater DW. 2011. Reconstructing the lionfish invasion: insights into Greater Caribbean biogeography. Journal of Biogeography, 38 (7): 1281-1293. Briggs JC. 2007. Marine biogeography and ecology: invasions and introductions. Journal of Biogeography, 34 (2): 193-198. Briggs JC. 2013. Invasion ecology: origin and biodiversity effects. Environmental Skeptics and Critics, 2 (3), 73-81. Butterfield JS, Díaz-Ferguson E, Silliman BR, Saunders JW, Buddo D, Mignucci-Giannoni A y Hunter ME. 2015. Wide-ranging phylogeographic structure of invasive red lionfish in the Western Atlantic and Greater Caribbean. Marine Biology, 162 (4): 773-781. Courtenay WR. 1995. Marine fish introductions in southeastern Florida. American Fisheries Society Introduced Fish Section Newsletter, 14 (1): 2-3. Darling ES, Green SJ, O’Leary JK y Côte IM. 2011. Indo-Pacific lionfish are larger and more abundant on invaded reefs: a comparison of Kenyan and Bahamian lionfish populations. Biological Invasions 13 (9): 2045-2051. DeWoody JA y Avise JC. 2000. Microsatellite variation in marine, freshwater and anadromous fishes compared with other animals. Journal of Fish Biology, 56 (3): 461-473. Dias P, Verheyen GR y Raymond M. 1996. Source-sink populations in Mediterranean blue tits: evidence using single locus minisatellite probes. Journal of Evolutionary Biology. 9 (6): 965-978. Earl DA. 2009. Structure Harvester v0. 3. [Internet]. 05 de Enero de 2016. Disponible en: <http://users. soe. ucsc. edu/~ dearl/software/struct_harvest>. Excoffier L, Smouse PE y Quattro JM. 1992. Analysis of molecular variance inferred from metric distances among DNA haplotypes: application to human mitochondrial DNA restriction data. Genetics, 131 (2): 479-491. Excoffier L, Laval G y Schneider S. 2005. Arlequin (version 3.0): an integrated software package for population genetics data analysis. Evolutionary Bioinformatics, 1: 47-50. Evanno G, Regnaut S y Goudet J. 2005. Detecting the number of clusters of individuals using the software STRUCTURE: a simulation study. Molecular ecology, 14 (8): 2611-2620. Ferreira CE, Luiz OJ, Floeter SR, Lucena MB, Barbosa MC, Rocha CR y Rocha LA. 2015. First record of invasive lionfish (Pterois volitans) for the Brazilian coast. PLos ONE, 10(2): 1-5. Franco–Herrera A. 2005. Oceanografía de la ensenada de Gaira: el Rodadero, más que un centro turístico en el Caribe Colombiano, Bogotá: Fundación Universidad de Bogotá Jorge Tadeo Lozano. p. 56. Frankham R. 1995. Conservation genetics. Annual Review of Genetics, 29 (1): 305-327. Frankham R. 2005. Resolving the genetic paradox in invasive species. Heredity, 94: 385. Freshwater DW, Hines A, Parham S, Wilbur AE, Sabaoun M, Woodhead J, Akins JL, Purdy B, Whitfield PE y Paris CB. 2009. Mitochondrial control region sequence analyses indicate dispersal from the US East Coast as the source of the invasive Indo-Pacific lionfish Pterois volitans in the Bahamas. Marine Biology, 156: 1213-1221. Goudet J. 2001. FSTAT, a program to estimate and test gene diversities and fixation indices (version 2.9. 3). Lausanne University, Lausanne, Switzerland. Green SJ, Akins JL, Maljković A y Côté IM. 2012. Invasive lionfish drive Atlantic coral reef fish declines. PloS one, 7 (3):1-3. Guo SW y Thompson EA. 1992. Performing the exact test of Hardy–Weinberg proportion for multiple alleles. Biometrics 48 (2): 361-372. Haag CR, Riek M, Hottinger JW, Pajunen VI y Ebert D. 2006. Founder events as determinants of within-island and among-island genetic structure of Daphnia metapopulations. Heredity, 96 (2): 150-158. Hamner RM, Freshwater DW y Whitfield PE. 2007. Mitochondrial cytochrome b analysis reveals two invasive lionfish species with strong founder events in the western Atlantic. Journal of Fish Biology 71: 214-222. Holland BS. 2001. Invasion without a bottleneck: microsatellite variation in natural and invasive populations of the brown mussel Perna perna (L). Marine Biotechnology, 3 (5): 407-415. Hubisz MJ, Falush D, Stephens M, y Pritchard JK. 2009. Inferring weak population structure with the assistance of sample group information. Molecular Ecology Resources, 9 (5): 1322-1332. IGAC. 1986. San Andrés y Providencia. Aspectos Geográficos. Instituto Geográfico Agustín Codazzi. Bogotá. 156 p. Johnston MW, y Purkis SJ. 2015. Hurricanes accelerated the Florida–Bahamas lionfish invasion. Global change biology, 21 (6): 2249-2260. Kolbe JJ, Richard E, Glor RE, Rodríguez-Schettino L, Chamizo-Lara A, Larson A y Losos JB. 2004. Genetic variation increases during biological invasion by a Cuban lizard. Nature, 431, 177-181. Lande R. 1988. Genetics and demography in biological conservation. Science, 241 (4872), 1455-1460. Larkin MA, Blackshields G, Brown NP, Chenna R, McGettigan PA, McWilliam H y Higgins DG. 2007. Clustal W and Clustal X version 2.0.Bioinformatics, 23 (21): 2947-2948. Librado P y Rozas J. 2009. DnaSP v5: a software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25(11): 1451-1452. Lindholm AK, Breden F, Alexander HJ, Chan WK, Thakurta SG. y Brooks R. 2005. Invasion success and genetic diversity of introduced populations of guppies Poecilia reticulata in Australia. Molecular Ecology, 14: 3671-3682. Luikart G, Sherwin WB, Steele BM y Allendorf FW. 1998. Usefulness of molecular markers for detecting population bottlenecks via monitoring genetic change. Molecular Ecology, 7(8): 963-974. Martínez-Viloria H, Gómez ÁR y Acero A. 2011. Presencia del pez león, Pterois volitans (Actinopterygii: Scorpaenidae) en el Departamento de la Guajira, Mar Caribe de Colombia. Boletín de Investigaciones Marinas y Costeras, 40 (2): 445-447. Mack RN, Simberloff D, Lonsdale MW, Evans H, Clout M y Bazzaz F. 2000. Invasiones Biológicas: Causas, Epidemiología, Consecuencias globales y Control. Tópicos en Ecología. 5:1-19. Meunier C, Tirard C, Hurtrez‐Boussès S, Durand P, Bargues MD, Mas‐Coma S y Renaud F. 2001. Lack of molluscan host diversity and the transmission of an emerging parasitic disease in Bolivia. Molecular Ecology, 10 (5):1333-1340. Miller SA, Dykes DD y Polesky H. 1988. A simple salting out procedure for extracting DNA from human nucleated cells. Nucleic acids research, 16 (3): 1215. Morris JA y Akins JL. 2009. Feeding ecology of the invasive lionfish in the Bahamian archipelago. Environmental Biology of Fishes. 86: 389-398. Morris JA y Whitfield PE. 2009. Biology, Ecology, Control and Management of the Invasive Indo-Pacific Lionfish: An Updated Integrated Assessment. NOAA Techn. Mem. NOS NCCOS 99. 57 pp. Nei M. 1987. Molecular Evolutionary Genetics Columbia University Press New York 512. O'reilly PT, Canino MF, Bailey KM y Bentzen P. 2004. Inverse relationship between FST and microsatellite polymorphism in the marine fish, walleye pollock (Theragra chalcogramma): implications for resolving weak population structure. Molecular Ecology, 13(7): 1799-1814. Peakall RO y Smouse PE. 2006. GENALEX 6: genetic analysis in Excel. Population genetic software for teaching and research. Molecular Ecology Notes, 6 (1): 288-295. Planes S y Lecaillon G. 1998. Consequences of the founder effect in the genetic structure of introduced island coral reef fish populations. Biological Journal of the Linnean Society, 63 (4):537-552. Pimentel D, Zuñiga R y Morrison D. 2005. Update on the environmental and economic costs associated with alien-invasive species in the United States. Ecological Economics. 52: 273-288. Posada D y Crandall KA. 1998. Modeltest: testing the model of DNA substitution. Bioinformatics, 14 (9): 817-818. Prahl H y Erhardt H. 1985. Colombia: Corales y arrecifes coralinos. FEN Colombia. Fondo para la protección y medio ambiente “José Celestino Mutis”. Bogotá. Raymond M y Rousset F. 2003. A population genetic software for exact test and ecumenicism: GENEPOP, version 3.4. Heredity, 68: 248-249. Reed DH y Frankham R. 2003. Correlation between fitness and genetic diversity. Conservation Biology. 17: 230-237. Rius M, Turon X, Bernardi G, Volckaert FA y Viard F. 2015. Marine invasion genetics: from spatio-temporal patterns to evolutionary outcomes. Biological Invasions, 17 (3): 869-885. Roman J y Darling JA. 2007. Paradox lost: genetic diversity and the success of aquatic invasions. Trends in Ecology and Evolution, 22 (9): 454-464. Sawyer SA y Hartl DL. 1992. Population genetics of polymorphism and divergence. Genetics, 132 (4):1161-1176. Sax DF, Stachowicz JJ, Brown JH, Bruno JF, Dawson MN, Gaines SD, Grosberg RK, Hastings A, Holt RD, Mayfield MM, Mary I, O’Connor MI y Rice WR. 2007. Ecological and evolutionary insights from species invasions. Trends in Ecology and Ecology, 22 (9): 465-471. Schneider S, Roessli D y Excoffier L. 2000. Arlequin: a software for population genetics data analysis. User manual version, 2: 2496-2497. Schofield PJ. 2009. Geographic extent and chronology of the invasion of non-native lionfish (Pterois volitans [Linnaeus 1758] and P. miles [Bennett 1828]) in the Western North Atlantic and Caribbean Sea. Aquatic Invasions, 4: 443–449. Schultz TF, Fitzpatrick CK, Freshwater DW y Morris JA. 2013. Characterization of 18 polymorphic microsatellite loci from invasive lionfish (Pterois volitans and P. miles). Conservation Genetics Resources, 5 (3): 599-601. Sivasundar A, Bermingham E y Orti G. 2001. Population structure and biogeography of migratory freshwater fishes (Prochilodus: Characiformes) in major South American rivers. Molecular Ecology, 10 (2): 407-417. Sutherland WJ, Clout M, Côté IM, Daszak P, Depledge MH, Fellman L, Fleishman E, Garthwaite R, Gibbons DW, De Lurio J, Impey AJ, Lickorish F, Lindenmayer D, Madgwick J, Margerison C, Maynard T, Peck LP, Pretty J, Prior S, Redford KH, Scharlemann PW, Spalding M y Watkinson AR. 2010. A horizon scan of global conservation issues for 2010. Cell Press, 25: 1-7. Tajima F. 1989. Statistical method for testing the neutral mutation hypothesis by DNA polymorphism. Genetics, 123 (3): 585-595. Tamura K, Glen Stecher, Daniel Peterson, Alan Filipski y Sudhir Kumar. 2013. MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution, 30 (12): 2725-2729. Toledo-Hernández C, Vélez-Zuazo X, Ruiz-Diaz CP, Patricio AR, Mège P, Navarro M y Papa R. 2014. Population ecology and genetics of the invasive lionfish in Puerto Rico. Aquatic Invasions, 9 (2): 227-237. Torres-Rodriguez JA, Gonzales-Corredor JD y Acero A. Registro de un ejemplar de pez león (Pterois volitans) sin vestigios de aleta dorsal espinosa. En preparación. Triantafyllidis A, Abatzopoulos TJ, Leonardos J y Guyomard R. (2002). Microsatellite analysis of the genetic population structure of native and translocated Aristotle's catfish (Silurus aristotelis). Aquatic Living Resources, 15 (06): 351-359. Van Oosterhout C, Hutchinson WF, Wills DP y Shipley P. 2004. MICRO‐CHECKER: software for identifying and correcting genotyping errors in microsatellite data. Molecular Ecology Notes, 4 (3): 535-538. Vila C, Sundqvist AK, Flagstad Ø, Seddon J, Kojola I, Casulli A y Ellegren H. 2003. Rescue of a severely bottlenecked wolf (Canis lupus) population by a single immigrant. Proceedings of the Royal Society of London. Biological Sciences, 270 (1510): 91-97. Weir BS y Cockerham C. 1984. Estimating F-statistics for the analysis of population structure. Evolution, 38 (6): 1358-1370. Whitfield PE, Hare JA, David AW, Harter SL, Munoz RC y Addison CM. 2007. Abundance estimates of the Indo-Pacific lionfish Pterois volitans/miles complex in the Western North Atlantic. Biological Invasions, 9: 53-64. Willi Y, Van Buskirk J y Hoffmann AA. 2006. Limits to the adaptive potential of small populations. Annual Review of Ecology, Evolution, and Systematics, 37: 433-458.ORIGINAL80098976.2016.pdf80098976.2016.pdfapplication/pdf852071https://repositorio.unal.edu.co/bitstream/unal/75850/1/80098976.2016.pdf9cb9eef1c1f5594754e57bd2bb7d263fMD51LICENSElicense.txtlicense.txttext/plain; charset=utf-83991https://repositorio.unal.edu.co/bitstream/unal/75850/2/license.txt6f3f13b02594d02ad110b3ad534cd5dfMD52CC-LICENSElicense_rdflicense_rdfapplication/rdf+xml; charset=utf-8811https://repositorio.unal.edu.co/bitstream/unal/75850/3/license_rdf217700a34da79ed616c2feb68d4c5e06MD53THUMBNAIL80098976.2016.pdf.jpg80098976.2016.pdf.jpgGenerated Thumbnailimage/jpeg4738https://repositorio.unal.edu.co/bitstream/unal/75850/4/80098976.2016.pdf.jpge56af216dcbf2748c308f8d948da7af5MD54unal/75850oai:repositorio.unal.edu.co:unal/758502024-07-05 23:10:33.252Repositorio Institucional Universidad Nacional de Colombiarepositorio_nal@unal.edu.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