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Predicting Amphipods' Brood Size Variation in Brackish Environments

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Predicting Amphipods' Brood Size Variation in Brackish Environments Journal of Experimental Marine Biology and Ecology L 248 (2000) 207±223 www.elsevier.nl/locate/jembe Predicting amphipods' brood size variation in brackish environments: an empirical model for Corophium multisetosum Stock, 1952 (Corophiidae) in Ria de Aveiro (NW Portugal) M.R. Cunhaa,* , M.H. Moreira a , J.C. Sorbe b aDepartamento de Biologia, Universidade de Aveiro, P-3810-193 Aveiro, Portugal bLaboratoire d'Oceanographie Biologique, UMR 5805 (CNRS-UB1), 2, rue du Prof. Jolyet, F-33120 Arcachon, France Received 13 September 1999; received in revised form 20 January 2000; accepted 27 January 2000 Abstract Data on fecundity of Corophium multisetosum from AreaoÄ (Ria de Aveiro, Portugal) are analysed by non linear regression to quantify the relationship between brood size (Ne ) and head length (Lh, in mm), water temperature (T, in degrees Celsius) and salinity (S, in psu). The aim of the analysis is to obtain a simple line Neh5 a 1 bL , in which the slope (b) and the y intercept (a) are functions of salinity and/or temperature on each sampling occasion. The equation Ne 5 2 (22.940 2 8.027S) 1 (289.431 1 18.171S 1 12.904T 2 0.368T ) Lh explains 64% of the vari- ability of brood size throughout the breeding period. The model predicts an optimal temperature around 188C and a very low fecundity at low salinities. The graphical comparison of the lines obtained by the model and by a usual linear regression illustrates its potential usefulness to predict fecundity changes. The authors suggest that the observed variation in the fecundity of other brackish-water amphipods can be described and predicted using similar models. 2000 Elsevier Science B.V. All rights reserved. Keywords: Amphipods; Brackish-water; Fecundity; Model 1. Introduction Invertebrates that release their offspring at advanced stages of development, or provide some kind of brood protection appear to enjoy ecological advantages where *Corresponding author. Tel.: 1351-234-370-785; fax: 1351-234-426-408. E-mail address: [email protected] (M.R. Cunha) 0022-0981/00/$ ± see front matter 2000 Elsevier Science B.V. All rights reserved. PII: S0022-0981(00)00164-7 208 M.R. Cunha et al. / J. Exp. Mar. Biol. Ecol. 248 (2000) 207 ±223 salinity undergoes pronounced changes (Kinne, 1970a). Amphipods are thus amongst the most successful animal groups in colonising brackish environments. Their spatial and temporal patterns of abundance are ultimately the consequences of the schedules of fecundity and survivorship that represent life history strategies. Overall the strategy adopted by an organism is a compromise allocation of energy to the various aspects of its life history, each of which contributes to total ®tness (Begon and Mortimer, 1986). The age of ®rst reproduction, reproductive effort (the proportion of the available resource input that is allocated to reproduction) and longevity are crucial aspects of life history schedules. The egg size, brood size and number of broods per female are the main traits that determine reproductive effort in amphipods (Sainte-Marie, 1991). Brood size in gammarideans is often reported as being proportional to body length of incubating females and this relationship is frequently summarised by a linear regression analysis of the raw or log-transformed data (e.g. Fish, 1975; Sheader, 1978; Fish and Mills, 1979; Murdoch et al., 1986). Differences between the mean brood size of two generations or ¯uctuations throughout the breeding period are usually ascribed to the variation in body length of the incubating females (e.g. Dauvin, 1988a,b; Beare and Moore, 1998). However, other studies have shown that this is not a rule. The scatter of values for females of the same size is often large, especially if samples over the entire breeding period are pooled, and sometimes no satisfactory explanation can be found for the variability in the number of embryos per brood, e.g. Echinogammarus obtusatus (Sheader and Chia, 1970). The slope of the regression line that represents the increase in brood size with increasing body length can be used as an index of fecundity (Sheader, 1978). Several authors have shown that the slope of the regression may change throughout the breeding period, which implies a temporal variation in the size-speci®c fecundity (Sheader, 1978, 1983; Fish and Mills, 1979; Naylor et al., 1988). Environmental factors such as latitude, temperature, photoperiod, oxygen concentration and food availability may be important in determining brood size, as it has been suggested in studies on several amphipod species (Kinne, 1959; Vlasblom, 1969; Fish and Preece, 1970; Nelson, 1980; Van Dolah and Bird, 1980; Sheader, 1983; Moore, 1986). In the present work, ®eld data on brood size of Corophium multisetosum are analysed in relation to the body size of incubating females and also to the temperature and salinity throughout the breeding period. These environmental factors, currently assessed in brackish-water studies, are easy to measure and can be used as simple indices of seasonal changes. The analysis aims to produce a mathematical equation allowing the prediction of changes in fecundity according to the seasonal variation of temperature and salinity. The term `fecundity' is used as a synonym for the number of embryos per brood. 2. Material and methods 2.1. Data The analysis hereafter is based on the specimens collected monthly over one year (May 1988±April 1989) in Areao.Ä This site, where C. multisetosum attains maximal M.R. Cunha et al. / J. Exp. Mar. Biol. Ecol. 248 (2000) 207 ±223 209 population densities (Queiroga, 1990), is located at the upper reaches of Canal de Mira, Ria de Aveiro (NW Portugal). Ten corer replicates (sampled area510 3 0.01 m2 ) were collected at low water of new moon spring tides and the samples were preserved in formalin. In the study area, the average depth at low water is always lower than 0.5 m and the tidal range varies between 0.2 and 1.0 m at neap and spring tides, respectively. Strati®cation of the water column was never observed. Water temperature and salinity were recorded at low and high water (early morning and beginning of the afternoon, respectively) near the bottom using a SCT meter (YSI model 33). Further details on the sampling methodologies, the seasonal variation of environmental factors and the macrobenthic community in the studied site are given by Cunha and Moreira (1995). Incubating females of C. multisetosum were later separated from the remaining fauna and the developmental stage of the embryos was assessed (Cunha et al., 2000a). The size of the females, expressed as head length, was measured to the nearest 1/60 mm in three replicates and the number of embryos per brood was counted only in those females with an undamaged marsupium. In order to minimise the error resulting from intramarsupial loss only females carrying embryos in an early developmental stage (F1 females with rounded embryos) were considered for the statistical analysis. Further details on the abundance, biomass, production, life history and reproductive biology of C. mul- tisetosum in the site studied are given by Cunha et al. (2000a,b). 2.2. Statistical analysis The signi®cance of the head length (Lh ), temperature (T ) and salinity (S) as sources of variation in the brood size (Ne )ofC. multisetosum was ®rst assessed by a simple factorial ANOVA (SPSS package). The relation between brood size and each factor was analysed separately and then the coef®cients of the general equation were estimated using a non-linear regression model (SPSS package). 3. Results 3.1. Data In Ria de Aveiro, C. multisetosum breeds throughout the year but in May, July and August only a few incubating females were present in the population (Table 1). An intense recruitment peak occurred during the autumn and a smaller peak in spring (Cunha et al., 2000a). The analyses hereafter are based on a sample of 217 F1 females with undamaged marsupium (Table 1). The diameter of the recently laid eggs varied from 0.30 to 0.35 mm regardless of the female size. Brood size varied from nine to 72 (global average 29.9) and the head length of incubating females ranged from 0.500 to 1.017 mm. The temporal variation in the average brood size and average head length of incubating C. multisetosum females showed disagreeing trends especially in the period from September to November (Fig. 1). The relationship between brood size (Neh) and head length (L ) obtained by a least-squares linear regression (GR) using the global data set was: 2 Neh5221.868 1 64.599L (R 5 0.183; n 5 217) (GR) 210 M.R. Cunha et al. / J. Exp. Mar. Biol. Ecol. 248 (2000) 207 ±223 Table 1 Water temperature (8C) and salinity (psu) data over the sampling period in AreaoÄ a Date Temperature Salinity Finc % F 1 n LW HW LW HW 16 May 1988 19.0 18.0 0.2 0.5 1 0.0 0 14 June 20.9 21.2 0.5 3.0 85 96.5 3 12 July 22.0 24.1 0.5 1.0 1 100.0 1 10 Aug 23.5 25.3 0.2 1.8 7 28.6 2 13 Sept 22.5 23.7 1.0 6.5 379 70.4 57 10 Oct 18.0 18.0 1.0 6.5 789 96.5 75 07 Nov 17.5 18.0 0.3 2.0 769 24.3 9 07 Dec 12.0 12.5 0.8 2.0 1726 40.6 13 05 Jan 1989 11.0 12.4 2.2 3.0 456 74.1 26 06 Feb 11.5 12.5 0.7 3.0 650 57.1 25 08 Mar 16.2 17.0 0.7 2.6 643 40.9 2 05 Apr 15.0 14.5 0.5 1.2 206 36.4 4 a The number of incubating females (Finc ), the percentage of females carrying embryos in a early developmental stage (F11 ), and the number (n) of F females examined for statistical analysis are also indicated for each month.
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