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Seasonal Variation in Dry Weight and Elemental Composition of the Early Developmental Stages of Petrolisthes Laevigatus

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Seasonal Variation in Dry Weight and Elemental Composition of the Early Developmental Stages of Petrolisthes Laevigatus Helgol Mar Res (2013) 67:167–177 DOI 10.1007/s10152-012-0313-4 ORIGINAL ARTICLE Seasonal variation in dry weight and elemental composition of the early developmental stages of Petrolisthes laevigatus (Gue´rin, 1835) (Decapoda: Porcellanidae) in the Seno de Reloncavı´, southern Chile P. Gebauer • K. Paschke • A. Barrı´a • K. Anger Received: 9 January 2012 / Revised: 21 May 2012 / Accepted: 5 June 2012 / Published online: 21 June 2012 Ó Springer-Verlag and AWI 2012 Abstract In the Seno de Reloncavı´, southern Chile, conditions. In summer, by contrast, plankton productivity seasonal changes in dry weight (DW) and elemental and temperatures are generally high, allowing for fast lar- composition (CHN) were studied in embryo (initial val growth and development. This coincides with minimal embryonic stage), newly hatched zoeae, and newly settled biomass and energy contents both at hatching and settle- megalopae of a porcelain crab, Petrolisthes laevigatus. ment. In conclusion, our data suggest that seasonal patterns Samples were taken throughout the seasons of egg laying in the biomass of early developmental stages of P. lae- (March-December), hatching (August-February), and set- vigatus may reflect phenotypic variability as an adaptive tlement (October–February). Values of DW and CHN per response to predictable variations in environmental condi- embryo or larva, respectively, were consistently minimum tions, allowing this species to reproduce in temperate in the middle of each season and maximum near its regions with marked seasonality in water temperature and beginning and end. Patterns of seasonal variation in early plankton productivity. embryonic biomass may thus be carried over to larvae at hatching and, possibly, to the settlement stage. Such carry- Keywords Bioenergetic traits variability Á Embryo Á over effects may be selectively advantageous, as zoeae Larvae Á Elemental composition Á Petrolisthes laevigatus Á released at the beginning or near the end of the hatching Seasonal variability season face conditions of poor planktonic food availability in combination with low winter temperatures or decreasing temperatures at the end of summer (enforcing long devel- Introduction opment duration). Hence, an enhanced female energy allocation into egg production may subsequently translate Population demographics and dynamics of marine inver- to enhanced yolk reserves remaining at hatching, allowing tebrates are consequences of pre- and post-settlement for a larval development under unfavourable winter processes (Connell 1985; Gaines and Rougharden 1985; Grosberg and Levitan 1992; Gime´nez 2004). Analysis of life-history traits is thus critical for an understanding of Communicated by Heinz-Dieter Franke. these aspects of population biology. Life-history patterns P. Gebauer (&) Á A. Barrı´a have generally been considered as adaptive, because they Centro I-MAR, Universidad de Los Lagos, are subject to environmental pressures of evolutionary Casilla 557, Puerto Montt, Chile selection towards an optimization of the survival of prog- e-mail: [email protected] eny (Roff 1992; Stearns 1992; Levin and Bridges 1995). K. Paschke However, both physical and biological conditions can Instituto de Acuicultura, Universidad Austral de Chile, modify some traits, indicating phenotype plasticity (Pfennig Casilla 1327, Puerto Montt, Chile et al. 2010). Intraspecific variations in various life-history traits have been found, for instance, in crustaceans and other K. Anger Biologische Anstalt Helgoland, Stiftung Alfred-Wegener-Institut marine invertebrates (Hines 1986a, b; Anger 2001, 2006). fu¨r Polar- und Meeresforschung, 27498 Helgoland, Germany Among these intraspecifically variable traits, offspring size 123 168 Helgol Mar Res (2013) 67:167–177 has been considered as particularly important, because it long duration of the periods of egg laying, larval hatching, affects developmental time, survival in the plankton, and and megalopal settlement, females and early developmen- the success of recruitment (Miner et al. 2005; Marshall and tal stages (embryos, zoeae, megalopae) face seasonally Keough 2006; Moran and McAlister 2009). Such variations variable environmental conditions. in offspring size have been studied mainly across the lati- In a temperate environment, as Seno de Reloncavı´, with tudinal range of distribution of species, whose populations marked seasonality in temperature, ranging between 4 °C may live under different environmental conditions (Brante (August) and 17 °C, (January) in the intertidal zone et al. 2004; Ouellet and Plante 2004; Rotllant et al. 2004; (Gebauer et al. 2007) or between 9 °C (July) and 17 °C Bas et al. 2007; Lardies et al. 2010; Weiss et al. 2010). (February) in the water column, while primary productivity Seasonal variations in environmental factors such as fluctuates approximately between 1 mg C m-3 h-1 (win- temperature and food supply have been identified, in gen- ter) and 23–12 mg C m-3 h-1 (spring and fall) (Iriarte eral, as important causes of intrapopulational variability in et al. 2007), the occurrence of embryos, early larvae, and reproductive traits of marine invertebrates (Ghiselin 1987). settlers of P. laevigatus throughout extended periods raises However, such variations have not extensively been stud- the question whether these early developmental stages ied in decapod crustaceans. Seasonal differences have been differ intra-annually in biomass, elemental composition, found, for instance, in the number and volume of anomuran and/or quality (expressed in terms of energy content). If and brachyuran crab eggs (Sampedro et al. 1997; Bas et al. these aspects show variability, this might indicate an 2007), in the elemental composition of shrimp and crab adaptive strategy to face the seasonally or latitudinally embryos (Paschke 1998; Bas et al. 2007; Urzu´a et al. 2012) variable conditions. and in the biochemical composition of blue crab embryos (O’Leary Amsler and George 1984). Furthermore, seasonal variations have been identified in the nutritional vulnera- Materials and methods bility of first-stage larval shrimp (Crangon crangon; Paschke et al. 2004) and porcelain crab (Petrolisthes Throughout the periods of egg laying, hatching, and set- laevigatus; Gebauer et al. 2010). tlement, porcelain crabs (P. laevigatus) were regularly The species P. laevigatus (Gue´rin 1835) is a filter- collected from the intertidal zone of Pelluhuı´n, Seno de feeding crab that typically inhabits mid-upper rocky Reloncavı´, Chile (41°450S, 72°200W). Females with mature intertidal zones on the Pacific coast of South America. Its gonads were manually collected and transported to the geographic range extends from the south of Peru (12°S) to laboratory, where they were kept in aquaria. These were southern Chile (46°S) (Retamal 1981). Near the southern placed in a water bath connected to an open system with limit of this range, in the Seno de Reloncavı´ (41°450S, circulating coastal seawater (9.5–15 °C, corresponding to 72°200W), females with early eggs are present from March field temperatures during the period of collection; constant to January–February, whereas final embryonic stages are 31 psu), together with males, remaining unfed for maxi- mainly found between August and February (Gebauer et al. mally five days, until eggs were laid (females were checked 2007). The embryonic development may last up to daily). Every month (from March to December), samples approximately five months (Gebauer et al. 2007). Female of embryos in initial stage (meaning less than 1 day of P. laevigatus are able to reproduce at least twice during embryonic development) were removed from five females each reproductive season (Gebauer, personal observation). and prepared for later analyses (see below). In females, In southern-Central Chile, the eggs of P. laevigatus in an carapace length (CL) was measured, and all embryos were initial embryonic stage have a volume of about 0.23 mm3 counted. For thirty embryos from each female, embryonic and a dry weight of ca 88 lg (Lardies and Wehrtmann length and width were measured under a stereo micro- 1996). The release of the larvae extends for 7 months, from scope, and embryo volume was calculated (Valde´s et al. August to February (Gebauer et al. 2007), with different 1991). These embryos were subsequently divided into five cohorts showing differential nutritional vulnerability replicate samples for further determinations of dry weight (Gebauer et al. 2010). The nutritional vulnerability index (DW) and elemental composition (carbon, hydrogen, (NVI; Gebauer et al. 2010) fluctuates between 2.2 and 1.7 nitrogen; collectively CHN). in August and October, respectively, indicating high values From August to February, females with embryos in a of point of reserve saturation and low values of point of no final stage of embryonic development were regularly col- return, representing high dependence on food of early lected from the same intertidal habitat. They were subse- larval stage (Gebauer et al. 2010). The settlement period of quently kept for maximally 5 days in individual aquaria this species extends over six months, from October to without food (10–15 °C; field temperatures during the March, showing both inter- and intra-annual variations in period of collection; constant 31 psu), until Zoea I larvae settler abundance (Gebauer et al. 2007, 2011). Due to the hatched. The water was changed daily and checked for the 123 Helgol Mar Res (2013) 67:167–177 169 presence of larvae. Thirty newly hatched larvae obtained beginning and the
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