Comunidades bentónicas de charcas rocosas del norte de Portugal: variabilidad estacional y entre charcas
DOI:
https://doi.org/10.3989/scimar.03669.21APalabras clave:
charcas rocosas, intermareal rocoso, variabilidad estacional, bentos, Sabellaria alveolata, PortugalResumen
Investigamos la variabilidad estacional (invierno vs verano y entre estaciones) y espacial (entre charcas) de las comunidades bentónicas en charcas rocosas del nivel intermareal medio a lo largo de la costa de Viana do Castelo (Norte de Portugal). Además comparamos los parámetros físicos entre las charcas, incluyendo el tamaño, temperatura y posición a lo largo de la costa. Mientras que las charcas no difieren para ninguno de los parámetros físicos examinados, los resultados indican una clara diferencia estacional en la estructura de las comunidades, las cuales incluyen un total de 49 taxones de algas y 13 de animales. Estos resultados fueron inducidos por seis taxones más abundantes en invierno (el poliqueto formador de arrecifes Sabellaria alveolata, el alga coralinal articulada Corallina spp., el alga parda Bifurcaria bifurcata, el alga coralinal incrustante Litophyllum incrustans, el alga roja Chondracanthus acicularis y los caracoles ramoneadores Gibbula spp.) y cuatro taxones de algas más abundantes en verano (el alga parda invasiva Sargassum muticum, el alga verde Ulva spp., el quelpo Laminaria ochroleuca y el alga roja filamentosa Ceramium spp.). Estos datos contribuyen al conocimiento de las comunidades de charcas rocosas intermareales y tienen potenciales implicaciones en los programas de monitorización enfocados a medir las modificaciones ecológicas relacionadas con alteraciones naturales y antropogénicas y son útiles para identificar los procesos responsables de la variabilidad de estas comunidades.
Descargas
Citas
Airoldi L. 2000. Responses of algae with different life histories to temporal and spatial variability of disturbance in subtidal reefs. Mar. Ecol. Prog. Ser. 195: 81-92. http://dx.doi.org/10.3354/meps195081
Anderson M.J. 2001. A new method for non-parametric multivariate analysis of variance. Austral. Ecol. 26: 32-46.
Anderson M.J. 2006. Distance-based tests for homogeneity of multivariate dispersions. Biometrics 62: 245-253. http://dx.doi.org/10.1111/j.1541-0420.2005.00440.x PMid :16542252
Anderson M.J., Robinson J. 2003. Generalized discriminant analysis based on distances. Aust. N.Z. J. Stat. 45: 301-318. http://dx.doi.org/10.1111/1467-842X.00285
Andrew N.L., Mapstone B.D. 1987. Sampling and the description of spatial pattern in marine ecology. Oceanogr. Mar. Biol. Annu. Rev. 52: 39-90.
Araújo R., Sousa-Pinto I., Bárbara I., Quintino V. 2006. Macroalgal communities of intertidal rock pools in the northwest coast of Portugal. Acta Oecol. 30: 192-202. http://dx.doi.org/10.1016/j.actao.2006.04.002
Araújo R., Vaselli S., Almeida M., Serrão E., Sousa-Pinto I. 2009. Effects of disturbance on marginal populations: human trampling on Ascophyllum nodosum assemblages at its southern distribution limit. Mar. Ecol. Prog. Ser. 378: 81-92. http://dx.doi.org/10.3354/meps07814
Arenas F., Fernández C. 2000. Size structure and dynamics in a population of Sargassum muticum (Phaeophyceae). J. Phycol. 36: 1012-1020. http://dx.doi.org/10.1046/j.1529-8817.2000.99235.x
Arenas F., Fernández C., Rico J.M., Fernández E., Haya D. 1995. Growth and reproductive strategies of Sargassum muticum Yendo (Fensholt) and Cystoseira nodicaulis (Whit.) Roberts. Sci. Mar. 59: 1-8.
Arenas F., Viejo R.M., Fernández C. 2002. Density-dependent regulation in an invasive seaweed: responses at plant and modular levels. J. Ecol. 90: 820-829. http://dx.doi.org/10.1046/j.1365-2745.2002.00720.x
Astles K.L. 1993. Patterns of abundance and distribution of species in intertidal rock pools. J. Mar. Biol. Assoc. U.K. 73: 555-569. http://dx.doi.org/10.1017/S0025315400033105
Ayata S.D., Ellien C., Dumas F., Dubois S., Thiébaut E. 2009. Modelling larval dispersal and settlement of the reef-building polychaete Sabellaria alveolata: role of hydroclimatic processes on the sustainability of biogenic reefs. Contin. Shelf Res. 29: 1605-1623. http://dx.doi.org/10.1016/j.csr.2009.05.002
Baer J., Stengel D.B. 2010. Variability in growth, development and reproduction of the non-native seaweed Sargassum muticum (Phaeophyceae) on the Irish west coast. Estuar. Coast. Shelf Sci. 90: 185-194. http://dx.doi.org/10.1016/j.ecss.2010.08.011
Balestri E., Benedetti-Cecchi L., Lardicci C. 2004. Variability in patterns of growth and morphology of Posidonia oceanica exposed to urban and industrial wastes: contrasts with two reference locations. J. Exp. Mar. Biol. Ecol. 308: 1-21. http://dx.doi.org/10.1016/j.jembe.2004.01.015
Benedetti-Cecchi L. 2000. Priority effects, taxonomic resolution, and the prediction of variable patterns of colonisation of algae in littoral rock pools. Oecologia 123: 265-274. http://dx.doi.org/10.1007/s004420051013
Benedetti-Cecchi L. 2001. Variability in abundance of algae and invertebrates at different spatial scales on rocky sea shores. Mar. Ecol. Prog. Ser. 215: 79-92. http://dx.doi.org/10.3354/meps215079
Benedetti-Cecchi L., Cinelli F. 1992. Effects of canopy cover, herbivores and substratum type on patterns of Cystoseira spp. settlement and recruitment in littoral rockpools. Mar. Ecol. Prog. Ser. 90: 183-191. http://dx.doi.org/10.3354/meps090183
Benedetti-Cecchi L., Cinelli F. 1994. Recovery of patches in an assemblage of geniculate coralline algae: variability at different successional stages. Mar. Ecol. Prog. Ser. 110: 9-18. http://dx.doi.org/10.3354/meps110009
Benedetti-Cecchi L., Cinelli F. 1995. Habitat heterogeneity, sea urchin grazing and the distribution of algae in littoral rock pools on the west coast of Italy (western Mediterranean). Mar. Ecol. Prog. Ser. 126: 203-212. http://dx.doi.org/10.3354/meps126203
Benedetti-Cecchi L., Cinelli F. 1996. Patterns of disturbance and recovery in littoral rock pools: nonhierarchical competition and spatial variability in secondary succession. Mar. Ecol. Prog. Ser. 135: 145-161. http://dx.doi.org/10.3354/meps135145
Benedetti-Cecchi L., Nuti S., Cinelli F. 1996. Analysis of spatial and temporal variability in interactions among algae, limpets and mussels in low-shore habitats on the west coast of Italy. Mar. Ecol. Prog. Ser. 144: 87-96. http://dx.doi.org/10.3354/meps144087
Benedetti-Cecchi L., Bertocci I., Micheli F., Maggi E., Fosella T., Vaselli S. 2003. Implications of spatial heterogeneity for management of marine protected areas (MPAs): examples from assemblages of rocky coasts in the northwest Mediterranean. Mar. Env. Res. 55: 429-458. http://dx.doi.org/10.1016/S0141-1136(02)00310-0
Bertocci I., Maggi E., Vaselli S., Benedetti-Cecchi L. 2005. Contrasting effects of mean intensity and temporal variation of disturbance on assemblages of rocky shores. Ecology 86: 2061-2067. http://dx.doi.org/10.1890/04-1698
Breitburg D.L. 1984. Residual effects of grazing: inhibition of competitor recruitment by encrusting coralline algae. Ecology 65: 1136-1143. http://dx.doi.org/10.2307/1938321
Britton-Simmons K.H. 2004. Direct and indirect effects of the introduced alga Sargassum muticum on benthic subtidal communities of Washington State, USA. Mar. Ecol. Prog. Ser. 277: 61-78. http://dx.doi.org/10.3354/meps277061
Bulleri F., Bertocci I., Micheli F. 2002. Interplay of encrusting coralline algae and sea urchins in maintaining alternative habitats. Mar. Ecol. Prog. Ser. 243: 101-109. http://dx.doi.org/10.3354/meps243101
Caley M., Carr M., Hixon M.A., Hughes T.P., Jones G.P, Menge B.A. 1996. Recruitment and the local dynamics of open marine populations. Annu. Rev. Ecol. Syst. 27: 477-500. http://dx.doi.org/10.1146/annurev.ecolsys.27.1.477
Chapman A.R.O. 1990. Effects of grazing, canopy cover and substratum type on the abundances of common species of seaweeds inhabiting littoral fringe tide pools. Bot. Mar. 33: 319-326. http://dx.doi.org/10.1515/botm.1990.33.4.319
Cheshire A.C., Westphalen G., Wenden A., Scriven L.J., Rowland B.C. 1996. Photosynthesis and respiration of phaeophycean-dominated macroalgal communities in summer and winter. Aquat. Bot. 55: 159-170. http://dx.doi.org/10.1016/S0304-3770(96)01071-6
Clarke K.R. 1993. Nonparametric multivariate analyses of changes in community structure. Austral. J. Ecol. 18: 117-143. http://dx.doi.org/10.1111/j.1442-9993.1993.tb00438.x
Coelho H.S., Neves R.J.J., White M., Leitão P.C., Santos A.J. 2002. A model for ocean circulation on the Iberian coast. J. Mar. Syst. 32: 153-179. http://dx.doi.org/10.1016/S0924-7963(02)00032-5
Davison I.R., Pearson G.A. 1996. Stress tolerance in intertidal seaweeds. J. Phycol. 32: 197-211. http://dx.doi.org/10.1111/j.0022-3646.1996.00197.x
Desroy N., Dubois S.F., Fournier J., Ricquiers L., Le Mao P., Guerin L., Gerla D., Rougerie M., Legendre A. 2011. The conservation status of Sabellaria alveolata (L.) (Polychaeta: Sabellariidae) reefs in the Bay of Mont-Saint-Michel. Aquat. Conserv.: Mar. Freshwat. Ecosyst. 21: 462-471. http://dx.doi.org/10.1002/aqc.1206
Dethier M.N. 1982. Pattern and process in tidepool algae: factors influencing seasonality and distribution. Bot. Mar. 25: 55-66. http://dx.doi.org/10.1515/botm.1982.25.2.55
Dethier M.N. 1984. Disturbance and recovery in intertidal pools: maintenance of mosaic patterns. Ecol. Monogr. 54: 99-118. http://dx.doi.org/10.2307/1942457
Dethier M.N., Graham E.S., Cohen S., Tear L.M. 1993. Visual versus random-point percent cover estimations: "objective" is not always better. Mar. Ecol. Prog. Ser. 96: 93-100. http://dx.doi.org/10.3354/meps096093
Dias J.M.A., Gonzalez R., Garcia C., Diaz-del Rio V. 2002. Sediment distribution patterns on the Galicia-Minho continental shelf. Prog. Oceanogr. 52: 215-231. http://dx.doi.org/10.1016/S0079-6611(02)00007-1
Díez I., Santolaria A., Secilla A., Gorostiaga J.M. 2009. Recovery stages over long- term monitoring of the intertidal vegetation in the Abra de Bilbao area and on the adjacent coast (N. Spain). Eur. J. Phycol. 4: 1-14. http://dx.doi.org/10.1080/09670260802158642
Dubois S., Retière C., Olivier F. 2002. Biodiversity associated with Sabellaria alveolata (Polychaeta: Sabellariidae) reefs: effects of human disturbances. J. Mar. Biol. Assoc. U.K. 82: 817-826. http://dx.doi.org/10.1017/S0025315402006185
Dubois S., Commito J.A., Olivier F., Retière C. 2006. Effects of epibionts on Sabellaria alveolata (L.) biogenic reefs and their associated fauna in the Bay of Mont Saint-Michel. Estuar. Coast. Shelf Sci. 68: 635-646. http://dx.doi.org/10.1016/j.ecss.2006.03.010
Dubois S., Comtet T., Retière C., Thiébaut E. 2007. Distribution and retention of Sabellaria alveolata larvae (Polychaeta: Sabellariidae) in the Bay of Mont-Saint-Michel, France. Mar. Ecol. Prog. Ser. 344: 15-28.
Femino R., Mathieson A. 1980. Investigations of New England marine algae IV. The ecology and seasonal succession of tide pool algae at Bald Head Cliff, York, Maine, USA. Bot. Mar. 23: 319-332.
Fournier J., Etienne S., Le Cam J.B. 2010. Inter- and intraspecific variability in the chemical composition of the mineral phase of cements from several tube-building polychaetes. Geobios 43: 191-200. http://dx.doi.org/10.1016/j.geobios.2009.10.004
Fraschetti S., Terlizzi A., Benedetti-Cecchi L. 2005. Patterns of distribution of marine assemblages from rocky shores: evidence of relevant scales of variation. Mar. Ecol. Prog. Ser. 296: 13-29. http://dx.doi.org/10.3354/meps296013
Godet L., Fournier J., Jaffré M., Desroy N. 2011. Influence of stability and fragmentation of a worm-reef on benthic macrofauna. Estuar. Coast. Shelf Sci. 92: 472-479. http://dx.doi.org/10.1016/j.ecss.2011.02.003
Golléty C., Migné A., Davoult D. 2008. Benthic metabolism on a sheltered rocky shore: role of the canopy in the carbon budget. J. Phycol. 44: 1146-1153. http://dx.doi.org/10.1111/j.1529-8817.2008.00569.x
Guerra-García J.M., Cabezas M.P., Baeza-Rojano E., García-Gómez J.C. 2011. Spatial patterns and seasonal fluctuations of intertidal macroalgal assemblages from Tarifa Island, southern Spain: relationship with associated Crustacea. J. Mar. Biol. Assoc. U.K. 91: 107-116. http://dx.doi.org/10.1017/S0025315410001219
Guimaraens M.A., Coutinho R. 2000. Temporal and spatial variation of Ulva spp. And water properties in the Cabo Frio upwelling region of Brazil. Aquat. Bot. 66: 101-114. http://dx.doi.org/10.1016/S0304-3770(99)00070-4
Guiry M.D., Cunningham E.M. 1984. Photoperiodic and temperature responses in the reproduction of north-eastern Atlantic Gigartina acicularis (Rhodophyta: Gigartinales). Phycologia 23: 357-367. http://dx.doi.org/10.2216/i0031-8884-23-3-357.1
Hewitt J.E., Thrush S.F., Cummings V.J., Turner S.J. 1998. The effect of changing sampling scales on our ability to detect effects of large-scale processes on communities. J. Exp. Mar. Biol. Ecol. 227: 251-264. http://dx.doi.org/10.1016/S0022-0981(97)00274-8
Huggett J., Griffiths C.L. 1986. Some relationships between elevation, physico-chemical variables and biota of intertidal rock pools. Mar. Ecol. Prog. Ser. 29: 189-197. http://dx.doi.org/10.3354/meps029189
Huthnance J.M., Van Aken H.M., White M., Barton E.D., LeCann B., Coelho E.F., Fanjul E.A., Miller P., Vitorino J. 2002. Ocean margin exchange-water flux estimates. J. Mar. Syst. 32: 107-137. http://dx.doi.org/10.1016/S0924-7963(02)00034-9
Izquierdo J.L., Pérez-Ruzafa I.M., Gallardo T. 2002. Effect of temperature and photon fluence rate on gametophytes and young sporophytes of Laminaria ochroleuca Pylaie. Helgoland Mar. Res. 55: 285-292 http://dx.doi.org/10.1007/s10152-001-0087-6
Konar B., Foster M.S. 1992. Distribution and recruitment of subtidal geniculate coralline algae. J. Phycol. 28: 273-280. http://dx.doi.org/10.1111/j.0022-3646.1992.00273.x
Kooistra W.H.C.F., Joosten A.M.T., van den Hoek C. 1989. Zonation patterns in intertidal pools and their possible causes: a multivariate approach. Bot. Mar. 32: 9-26. http://dx.doi.org/10.1515/botm.1989.32.1.9
Lawton J.H. 1996. Patterns in ecology. Oikos 75: 145-147. http://dx.doi.org/10.2307/3546237
Lemos R.T., Pires H.O. 2004. The upwelling regime off the west Portuguese coast, 1941–2000. Int. J. Climat. 24: 511-524. http://dx.doi.org/10.1002/joc.1009
Levin S.A. 1992. The problem of pattern and scale in ecology. Ecology 73: 1943-1967. http://dx.doi.org/10.2307/1941447
Littler M.M., Kauker B.J. 1984. Heterotrichy and survival strategies in the red alga Corallina officinalis L. Bot. Mar. 27: 37-44. http://dx.doi.org/10.1515/botm.1984.27.1.37
Masterson P., Arenas F.A., Thompson R.C., Jenkins S.R. 2008. Interaction of top down and bottom up factors in intertidal rockpools: Effects on early successional macroalgal community composition, abundance and productivity. J. Exp. Mar. Biol. Ecol. 363: 12-20. http://dx.doi.org/10.1016/j.jembe.2008.06.001
McArdle B.H., Anderson M.J. 2001. Fitting multivariate models to community data: a comment on distance-based redundancy analysis. Ecology 82: 290-297. http://dx.doi.org/10.1890/0012-9658(2001)082[0290:FMMTCD]2.0.CO;2
Menge B.A., Ashkenas L.R., Matson A. 1983. Use of artificial holes in studying community development in cryptic marine habitats in a tropical rocky intertidal region. Mar. Biol. 77: 129-142. http://dx.doi.org/10.1007/BF00396310
Metaxas A., Scheibling R.E. 1993. Community structure and organization of tidepools. Mar. Ecol. Prog. Ser. 98: 187-198. http://dx.doi.org/10.3354/meps098187
Metaxas A., Hunt H.L., Scheibling R.E. 1994. Spatial and temporal variability of macrobenthic communities in tidepools on a rocky shore in Nova Scotia, Canada. Mar. Ecol. Prog. Ser. 105: 89-103. http://dx.doi.org/10.3354/meps105089
Neto A.I. 2000. Observations on the biology and ecology of selected macroalgae from the littoral of São Miguel (Azores). Bot. Mar. 43: 483-498. http://dx.doi.org/10.1515/BOT.2000.049
Pedersen M.F., Borum J., Fotel F.L. 2010. Phosphorus dynamics and limitation of fast- and slow-growing temperate seaweeds in Oslofjord, Norway. Mar. Ecol. Prog. Ser. 399: 103-115. http://dx.doi.org/10.3354/meps08350
Reed D.C., Laur D.R., Ebeling A.W. 1988. Variation in algal dispersal and recruitment: the importance of episodic events. Ecol. Monogr. 58: 321-335. http://dx.doi.org/10.2307/1942543
Roberts D.E., Smith A., Ajani P., Davis A.R. 1998. Rapid changes in encrusting marine assemblages exposed to anthropogenic point-source pollution: a beyond BACI approach. Mar. Ecol. Prog. Ser. 163: 213-224. http://dx.doi.org/10.3354/meps163213
Rubal M., Veiga P., Vieira R., Sousa-Pinto I. 2011. Seasonal patterns of tidepool macroalgal assemblages in the North of Portugal. Consistence between species and functional group approaches. J. Sea Res. 66: 187-194. http://dx.doi.org/10.1016/j.seares.2011.07.003
Sánchez I., Fernández C. 2005. Impact of the invasive seaweed Sargassum muticum (Phaeophyta) on an intertidal macroalgal assemblage. J. Phycol. 41: 923-930. http://dx.doi.org/10.1111/j.1529-8817.2005.00120.x
Sánchez I., Fernández C., Arrontes J. 2005. Long-term changes in the structure of intertidal assemblages after invasion by Sargassum muticum (Phaeophyta). J. Phycol. 41: 942-949. http://dx.doi.org/10.1111/j.1529-8817.2005.00122.x
Steen H. 2003. Intraspecific competition in Sargassum muticum (Phaeophyceae) germlings under various density, nutrient and temperature regimes. Bot. Mar. 46: 36-43. http://dx.doi.org/10.1515/BOT.2003.006
Steneck R.S. 1986. The ecology of coralline algal crusts: convergent patterns and adaptive strategies. Annu. Rev. Ecol. Syst. 17: 273-303. http://dx.doi.org/10.1146/annurev.es.17.110186.001421
Terlizzi A., Anderson M.J., Fraschetti S., Benedetti-Cecchi L. 2007. Scales of spatial variation in Mediterranean subtidal sessile assemblages at different depths. Mar. Ecol. Prog. Ser. 332: 25-39. http://dx.doi.org/10.3354/meps332025
Thompson R.C., Crowe T.P., Hawkins S.J. 2002. Rocky intertidal communities: past environmental changes, present status and predictions for the next 25 years. Environ. Conserv. 29: 168-191. http://dx.doi.org/10.1017/S0376892902000115
tom Dieck I. 1992. North Pacific and North Atlantic digitate Laminaria species (Phaeophyta): hybridization experiments and temperature responses. Phycologia 31: 147-163. http://dx.doi.org/10.2216/i0031-8884-31-2-147.1
Underwood A.J. 1997. Experiments in Ecology: their logical design and interpretation using analysis of variance. Cambridge University Press, Cambridge, UK.
Underwood A.J., Chapman M.G. 1996. Scales of spatial patterns of distribution of intertidal invertebrates. Oecologia 107: 212-224. http://dx.doi.org/10.1007/BF00327905
Underwood A.J., Petraitis P.S. 1993. Structure of intertidal assemblages in different locations: how can local processes be compared? In: Ricklefs R., Schutler D. (eds.) Species diversity in ecological communities. Univ. Chicago Press, Chicago, pp. 38-51. PMid :8480505
Underwood A.J., Skilleter G.A. 1996. Effects of patch-size on the structure of assemblages in rock pools. J. Exp. Mar. Biol. Ecol. 197: 63-90. http://dx.doi.org/10.1016/0022-0981(95)00145-X
Underwood A.J., Chapman M.G., Connell S.D. 2000. Observations in ecology: you can't make progress on processes without understanding the patterns. J. Exp. Mar. Biol. Ecol. 250: 97-115. http://dx.doi.org/10.1016/S0022-0981(00)00181-7
Valdivia N., Scrosati R.A., Molis M., Knox A.S. 2011. Variation in community structure across vertical intertidal stress gradients: how does it compare with horizontal variation at different scales? PLoS ONE 6(8): e24062. http://dx.doi.org/10.1371/journal.pone.0024062 PMid :21887371 PMCid:3161098
Van Tamelen P.G. 1996. Algal zonation in tidepools: experimental evaluation of the roles of physical disturbance, herbivory and competition. J. Exp. Mar. Biol. Ecol. 201: 197-231. http://dx.doi.org/10.1016/0022-0981(95)00182-4
Viejo R.M. 1997. The effects of colonization by Sargassum muticum on tidepool macroalgal assemblages. J. Mar. Biol. Assoc. U.K. 77: 325-40. http://dx.doi.org/10.1017/S0025315400071708
Wilson D.P. 1970. Additional observations on larval growth and settlement of Sabellaria alveolata. J. Mar. Biol. Assoc. U.K. 50: 1-31. http://dx.doi.org/10.1017/S0025315400000576
Wolfe J., Harlin M. 1988a. Tidepools in Southern Rhode Island, U.S.A. I. Distribution and seasonality of macroalgae. Bot. Mar. 31: 525-536. http://dx.doi.org/10.1515/botm.1988.31.6.525
Wolfe J., Harlin M. 1988b. Tidepools in Southern Rhode Island, U.S.A. II. Species diversity and similarity analysis of macroalgal communities. Bot. Mar. 31: 537-546. http://dx.doi.org/10.1515/botm.1988.31.6.537
Descargas
Publicado
Cómo citar
Número
Sección
Licencia
Derechos de autor 2012 Consejo Superior de Investigaciones Científicas (CSIC)
![Creative Commons License](http://i.creativecommons.org/l/by/4.0/88x31.png)
Esta obra está bajo una licencia internacional Creative Commons Atribución 4.0.
© CSIC. Los originales publicados en las ediciones impresa y electrónica de esta Revista son propiedad del Consejo Superior de Investigaciones Científicas, siendo necesario citar la procedencia en cualquier reproducción parcial o total.Salvo indicación contraria, todos los contenidos de la edición electrónica se distribuyen bajo una licencia de uso y distribución “Creative Commons Reconocimiento 4.0 Internacional ” (CC BY 4.0). Puede consultar desde aquí la versión informativa y el texto legal de la licencia. Esta circunstancia ha de hacerse constar expresamente de esta forma cuando sea necesario.
No se autoriza el depósito en repositorios, páginas web personales o similares de cualquier otra versión distinta a la publicada por el editor.