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Differences Between Marine and Freshwater Fish Larvae: Implications for Recruitment

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Differences Between Marine and Freshwater Fish Larvae: Implications for Recruitment AR-318 ICESJ.mar.Sci.,51:91 97. 1994 Differencesbetween marine and freshwaterfish larvae: implications for recruitment E. D. Houde Houde, E. D. Differences between marine and freshwater fish larvae: implications for recruitment. ICES J. mar.Sci., 51: 91-97. Differences in the dynamics (growth and mortality) and energetics properties of marine and freshwater fish larvae have important implications fordetermining the lifestages at which year-class size is established. Aftercorrecting fortemperature effects,marine fish larvae, which typically weigh less at hatch, can be shown to experience higher mortality rates, have higher metabolic requirements, and have longer larval stage durations than do freshwater fish larvae. Growth rates and growth efficiencies are similar for the two categories of larvae. The difference in body size between typically small marine and typically large freshwaterfish larvae is an important factor affecting their dynamics and energetics. Predicted mean survivorship to metamorphosis of a cohort of freshwater fish larvae is 5.30%, while that of marine larvae is only 0.12%. The probability of significant density-dependent regulation during the larval stage could be relatively high for marine species, based upon their life-history properties. The probability that episodic mortalities of larvae will have significantimpacts on recruitment may be higher for freshwater fish than for marine fish.Starvation mortality is more probable formarine larvae, primarily because of their small body size, associated high metabolic demands, and possibly higher ingestion requirements. Because expected mean survivorship of freshwater larvae is 44 times higher than that of marine larvae, juvenile-stage dynamics will be relatively important in controlling regulating recruitment levels of freshwater fishes.In contrast, properties of marine species indicate that larval-stage dynamics will have a greater in0uence on recruitment success. Key words: larval fish, dynamics, energetics, marine, freshwater, recruitment mechanisms. Received 7 April 1993; accepted 27July 1993. E. D. Houde: Chesapeake Biological laboratory, Center for Environmental and Estuarine Studies, The Unil'ersity of Maryland System. PO Box 38, Solomons, Maryland 20688, USA. Introduction ontogenies vary widely (Balon, 1981). Development during the larval stage may be brief or protracted, result­ Research on fishearly life stages has proliferated in recent ing in either short- or long-stage durations. Because of the years, particularly on marine species. The high fecundities variability in morphologies and body sizes of larvae, as of fishesensure high larval abundances, which arc subject well as stage durations, it is not surprising that many to: (I) density-independent controls that are believed to biological properties differ substantially among species be a primary cause of recruitment variability; and (2) (Miller et al., 1988; Pepin, I 991; Pepin and Myers, 1991). possible density-dependent regulation, which will tend The vital rates (mortality rates, growth rates, stage to stabilize recruitment levels (Cushing, 1974a,b; durations) and energetics of larvae are strongly affected Rothschild, I 986). Growth rates, mortality rates, stage by temperature (Houde, 1989a; Morse, 1989). Tempera­ durations, and energetics properties of teleost larvae, ture and body size probably are the main variables affect­ when compared across taxa and ecosystems (Houde and ing egg and larval stage dynamics (Pepin, 1991) and Zastrow, in press), have many similarities, but they also prey consumption rates (Mackenzie et al., 1990). Their show significantdifferences that have implications for the effectson larval-stage durations may have had important recruitment process, as well as for development of fishery consequences in the evolution of teleost life-history modes research and management plans. (Houde, I989a). Fishes in both marine and freshwater habitats exhibit There are taxa-specific as well as ecosystem-specific diverse morphologies and behaviors. This diversity differences in vital rates and energetics properties of fish extends to the larval stages in which morphologies and larvae (Houde and Zastrow, in press) that could affect 1054 3139I 94I 010091+07S08.00 0 © 1994 International Council forthe Exploration of theSea 92 £. D. Houde recruitment levels, in addition to influencing dynamics from marine or freshwater systems, acknowledging that and energetics of older life stages. Houde and Zastrow (in such characterization docs not recognize species which press) noted some probable differences in larval-stage arc atypical and whose life histories may deviate signifi­ properties of marine and freshwater fish larvae in a cantly from the norm. Differences between means were broader analysis across aquatic ecosystems and taxa. judged to be significant at a=0.10, rather than at 0.05. In the present paper. I critically compare averaged to increase the power of the tests to detect probable properties of larvae from the combined marine eco­ differences in these quite variable data. which often were systems with properties of larvae from freshwater based upon few observations. ecosystems and discuss the potential for regulation and Regression statistics relating G, z. D. and QO 2 to W0 control of recruitment during the larval and juvenile were calculated to dctermmc if these variables were de­ stages in each system. pendent upon weight at hatching. and if the dependence differedbetween freshwater and marine larvae. Expected numbers of metamorphosed survivors. at mean values of Methods vital rates and ontogcnic attributes. were calculated for fishes from freshwater and marine ecosystems, based Data on larval dynamics (vital rates) and energetics, upon the relationship: categorized by ecosystems and by taxonomic groups. -Z(log. wm•• -log. W0)] _ were compiled by Houde and Zastrow (1991, in press). Nm.,-Px N0e [---- ()) Those reports included data on both freshwater and G = marine tclcost larvae. but the analysis and interpretations where N m., number of metamorphosed survivors: did not focus on implications for recruitment in fresh­ N0 = initial number of larvae; and P= probability of water and marine ecosystems. Herc, the analysis is surviving "episodic" mortality events. extended explicitly to examine and compare the relation­ The ratio of N m•• rw=Nm••·MAR for equal N0 was calcu­ ships between size at hatching and vital rate parameters lated to indicate relative stage-specific survivorship. By in the two categories of larvae. Recruitment levels in varying Z. G, and Pin Equation (I). the changes in rates each case arc predicted for mean values of rates estimated or episodic loss probabilities were calculated that are during larval life. and simple simulations arc run to required to cquali,c survivorship at metamorphosis in the demonstrate the extent to which average dynamics in the marine and freshwater ecosystems. Effects of initial egg larval stage must vary to precipitate important changes in or larval numbers (i.e. variability in N 0), caused b) recruitment levels. differencesin mean fecundities between marine and fresh­ Most tcleost taxa for which published data were avail­ water fishes, and the consequences for survivorship at able arc included in the analysis. Exceptions were a few metamorphosis, were considered. Telcost fecundity data taxa of principally freshwater fisheswith unusually large. compiled by Winemiller and Rose (1992) were used for precocial (se11.1u Balon, 1981) hatchlings that have either this purpose. a curtailed or absent larval stage. e.g. Salmonidac, The potential for density-dependent regulation of Acipenseridac, lctaluridae. The variables that were con­ larval survival via growth-rate responses to initial larval = = sidered are: W0 dry weight at hatching (µg): W mei dry abundances was compared for marine and freshwaterfish = weight at metamorphosis (µg): G weight-specific larvae by simulating responses of N m., in Equation (I) to 1 = growth coefficient (µg µg ' day- ): Z instantaneous variability in Ci and N 0• In an example, Ci was reduced by 1 ): = mortality coefficient (day- D larval-stage duration 10, 25, or 50% per five-fold increase in N0• Then, the = (day): 1.c. days to grow from WO to W me,; QO2 weight­ expected survivorships were compared for marine and 1 1 specific oxygen uptake (µI 02 mg- h- ); I= weight­ freshwater larvae to determine their sensitivities to specific food mgcstion (day 1), i.e. fraction of larval body density-dependent growth. = weight consumed daily; and K 1 gross growth efficiency, definedas GI I. Temperature-adjusted mean values of each parameter Results for larvae from five marine ecosystems and freshwater. and from nine broad taxonomic groups. were estimated Marine fish larvae. on average. are smaller at hatch. (Houde and Zastrow. 1991, in press). Adjustments forthe suffer higher mortality rates, have higher weight-specific substantial temperature effects were accomplished via respiration rates, and have longer stage durations than do analysis of covariance. Mean values that arc reported here freshwater larvae (Table I). The I 0-fold difference in for data from pooled marine ecosystems and from frcsh­ mean weight at hatching potentially has important conse­ w�tcr systems are those compiled by Houde and Zastrow quences for larval-stage dynamics and recruitment. (in press). except for minor differences resulting from a Weight-specific growth rates do not differ significantly. few additions or changes to the earlier data set. These
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