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Review: Diapause in Crustaceans and Its Ecological and Biogeographical Implications: an Example from Patagonian Ephemeral Pools

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Review: Diapause in Crustaceans and Its Ecological and Biogeographical Implications: an Example from Patagonian Ephemeral Pools Anales Instituto Patagonia (Chile), 2013. 41(2):43-50 43 REVIEW: DIAPAUSE IN CRUSTACEANS AND ITS ECOLOGICAL AND BIOGEOGRAPHICAL IMPLICATIONS: AN EXAMPLE FROM PATAGONIAN EPHEMERAL POOLS REVISIÓN: DIAPAUSA EN CRUSTÁCEOS Y SUS IMPLICANCIAS ECOLÓGICAS Y BIOGEOGRÁFICAS: UN EJEMPLO EN LAS POZAS PATAGÓNICAS Patricio De los Ríos-Escalante1,2 & Cinthia Robles3 ABSTRACT The production of diapause eggs is a strategy for survival adopted by microcrustaceans living under adverse environments. This strategy also, aids dispersion and colonization in new environments. In the long term, this colonization process can enhanced genetic isolation as the first step in the process of species evolution. The present study is: (a) wider review of the diapause process in inland water crustaceans and its ecological, evolutionary and biogeographical implications, and (b) it’s application to microcrustacean assemblages in inland waters in southern Patagonia. In the literature the diapause strategy was described from anostracans, cladocerans and copepods as microcrustaceans, and amphipoda in malacostracan crustaceans. High crustacean species richness was recorded from Southern Patagonia, this region having species of southern South American and Sub-Antarctic islands origin. Their spatial distribution is probably explained by the presence of migratory birds. Also, many habitats of the microcrustaceans are ephemerals, that would be studied under metapopulation and metacommunities view point. Key words: diapause, colonization, metapopulations, metacommunities, anostracans, cladocerans, copepods. RESUMEN Los microcrustáceos tienen como estrategias del ciclo de vida la generación de huevos en estado de diapausa bajo condiciones ambientales adversas, esta estrategia provee facilidades para la dispersión que generan procesos de colonización los que estimulan procesos de aislamiento genético como primer 1 Escuela de Ciencias Ambientales, Facultad de Recursos Naturales, Universidad Católica de Temuco, Casilla 15-D, Temuco, Chile. [email protected] 2 Núcleo de Estudios Ambientales, UC Temuco. 3 Escuela de Pedagogía Media, Facultad de Educación, Universidad Católica de Temuco, Casilla 15-D, Temuco, Chile. Received: Dic. 13, 2012 Accepted: Mar. 05, 2014 44 P. DE LOS RÍOS & C. ROBLES paso de procesos de especiación. El presente estudio es una revisión sobre los procesos de diapausa en crustáceos de aguas continentales y sus implicancias ecológicas, evolutivas y biogeográficas y se aplicará esto a la realidad de aguas continentales de la Patagonia. La literatura describe la presencia de anostracos, cladóceros y copépodos como microcrustáceos y anfípodos como malacostracos y todos estos grupos tienen diapausa como estrategia. La literatura describe alta riqueza de crustáceos en el sur de la Patagonia, y esta región tiene especies del sur de Sudamérica y de islas sub-Antárticas y es probable que la presencia de aves migratorias que explicarían la distribución espacial de estas especies. De igual modo, muchos habitats de microscrustáceos son lagunas temporales que podrían ser explicadas bajo el punto de vista de metapoblaciones y metacomunidades. Palabras clave: diapausa, colonización, metapoblaciones, metacomunidades, anostracos, cladó- ceros, copépodos. INTRODUCCIÓN that generate resting eggs. These subsequently hatch when the environmental conditions are optimal (De Microcrustaceans such as Branchiopoda, los Ríos-Escalante, 2010). From a biogeographical Copepoda and Ostracoda have diapause egg pro- view point, Patagonia has species from Southern duction- the generation of restings eggs - as a life South America and Sub-Antarctic islands (Menu- strategy for species perpetuation under adverse Marque et al. 2000; Pugh et al. 2002; Dos Santos environments, specifically drying of their habitats et al. 2008). Within this scenario, the aim of the (Alekseev, 2007a;b;2010;Alekseev et al. 2007; present study is to review diapause strategies of Muñoz, 2009). This life strategy has many ecological, microcrustaceans in general with emphasis on bio- systematic and evolutionary implications. Resting geographical and ecological implications and apply eggs, can be spread by agents such as wind and other this to a study of microcrustacean (Branchiopoda, fauna and initiate the colonization process in new Copepoda, and Amphipoda) species distribution in habitats or habitats where microcrstaceans have been Patagonian inland waters. been reduced or disappeared from. This could have impacts on metapopulations and metacommunities BRANCHIOPODA (Altermatt, 2008). This spreading and colonization dynamics would have evolutionary and systematic implicantions as, if the resting eggs colonize new The branchiopods described for Patago- habitats with environmental heterogeneity, selection nia include Anostracans (families Artemiidae and pressure would eliminate all individuals other than Branchinectidae), and Cladocerans (families Daph- those with optimal genes for survival and reproduc- nidae, Bosminidae, Chydoridae), with both endemic tion, and as a consequence would create genetic and cosmopolitan species (De los Ríos-Escalante, isolation as the first step in the speciation process 2010). Both Anostracan families Artemiidae and (De los Ríos & Zúñiga, 2000; Gajardo et al. 1995; Branchinectidae are present in saline and freshwater 2004; Ebert et al. 2002). This scenario, over a wide lakes and pools respectively (De los Ríos-Escalante, spatial and temporal scale could create zones where 2010). The Artemiidae is represented by the species certain taxa could originate, spread and colonize new, Artemia persimilis (Piccinelli & Prosdocimi, 1968) more distal habitats as the first step in a potential , found in two saline lakes in the Magallanes region, speciation process, that could be described using Laguna Amarga (51° 00’ S: 72° 48’ W) in Torres biogeographical techniques, such as track analysis del Paine National Park and Laguna de los Cisnes or endemicity parsimony analysis (Morrone, 2004; (53° 14’ S: 70° 23’ W) close to Porvenir on Tierra 2006). del Fuego (De los Ríos-Escalante & Gajardo, 2010). In Patagonia (38-54° S) there are many The population in Laguna Amarga is permanent ephemeral pools (formed in spring, and disappearing and practically exclusive to the zooplankton of this over the other seasons), that would contain crustaceans habitat (Campos et al., 1996), whereas the popula- REVISIÓN : DIAPAUSA EN CRUSTÁCEOS 45 tion in Laguna de los Cisnes depends on variations (Brady, 1875), B. michaelseni (Mrázek, 1901), B. in salinity- it can be present at high salinity levels, poppei (Mrázek, 1901) and Parabroteas sarsi (Ekman, but if the salinity decreases, the population disap- 1905). Amongst the Cyclopoids there are mainly pears (De los Ríos-Escalante & Gajardo, 2010). For endemic species from Southern South America (De Branchinectidae, the species Branchinecta gaini and los Ríos-Escalante, 2010), such as Acanthocyclops B. granulosa are reported from the Magallanes re- michaelseni (Mrázek, 1901), A. vernalis (Fisher, gion at 53 and 51° S respectively, B. vuriloche from 1853), Eucyclops ensifer Kiefer, 1936, Eucyclops the Aysen region at 45° S (Rogers et al. 2008) and serrulatus (Fisher, 1851), Macrocyclops albidus juvenile specimens in coastal ponds in the Araucania (Jurine, 1820), Mesocyclops araucanus (Löffler, region at 38° S (De los Ríos-Escalante et al. 2010). 1962), Metacyclops mendocinus (Wierzejski, 1892), Branchinecta are found in ephemeral pools present Microcyclops anceps (Richard, 1897), Paracyclops only in spring, and this genus is probably confined to fimbriatus chiltoni (Thomson, 1882) and Tropo- oligotrophic and low conductivity inland waters (De cyclops prasinus meridionalis Kiefer, 1927. Only los Ríos et al. 2008). The cladocerans have species the species Mesocyclops logisetus (Thiébaud, 1912) with cosmopolitan distibutions and endemic species is distributed among all geographical gradients in (De los Ríos-Escalante, 2010) such as Ceriodaphnia tropical and subtropical South America. There are dubia Richard, 1894, Daphnia ambigua Scourfield, no reported studies of diapause in South American 1947, D. dadayana, Paggi, 1999, D. pulex Leydig, inland water copepods, the available information is 1860; Scapholeberis exsipinifera (Nicolet, 1849); for New Zealand copepods (Cough et al., 2001; Simosa exspinosa (De Geer, 1778), Neobosmina Hall & Burns, 2001; Banks, 2007; Taylor 2010), chilensis (Daday, 1902), Ilyocryptus spinifer Her- and for Antarctic population of B. poppei (Jiang et rick, 1884; Alona poppei Richard, 1897, A. pul- al. 2012). Nevertheless, the presence of the above- chella King, 1853, A. quadrangularis (O.F. Müller, mentioned species in ephemeral pools would indicate 1776), A. clathratula (G.O. Sars, 1896), Alonella the presence of diapause life strategies in copepods excise (Fisher, 1854), Camptocercus rectirrostris (De los Ríos-Escalante, 2010). (Schoedler, 1862), Pleuroxus aduncus (Jurine, 1820), Chydorus sphaericus (O.F. Müller, 1785), AMPHIPODS Leydigia leydigi (Schoedler, 1863) and Paralona nigra (G.O. Sars, 1862). The cladocerans from the The amphipods reported from Southern northern hemisphere that produce diapause resting Patagonia included species reported for southern eggs have been studied (Altermatt, 2008; Muñoz, South America (De los Ríos-Escalante et al., 2012) 2009) and although a life strategy of sexual and such as Hyalella costera González & Watling, asexual generations is reported, this situation has not 2001, H. chiloensis González & Watling, 2001, been reported
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