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Age Determination and Seasonal Growth of the Dogwhelk Nucella Freycineti(Deshayes)

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Age Determination and Seasonal Growth of the Dogwhelk Nucella Freycineti(Deshayes) Benthos Research Vol.55,No.2:43-51(2000) BENTHOS RESEARCH The Japanese Association of Benthology Age Determination and Seasonal Growth of the Dogwhelk Nucella freycineti(Deshayes) Kei Kawai Marine Biodiversity Laboratory,Faculty of Fisheries,Hokkaido University,Minato3-1-1, Hakodate,Hokkaido041-8611,Japan Abstract:Growth patterns in the shell and soft parts of the dogwhelk Nucella freycineti were examined. Growth rings in the shells were investigated by X-ray photography and were shown to form annually.In snails that hatched in summer,the first rings formed in the first winter.The shell growth equation was determined based on the age character,population age structure,seasonal growth rate and size frequency distribution of the population.The maximum shell growth occurred in summer.The life span was esti mated to be>11years.Low physiological activity and/or starvation induced by low temperature might be main factor(s)causing the formation of annual rings,although age might also influence ring forma tion.Total body weight increased in late summer in males and from late summer to winter in females. Gonad weight showed a similar trend in each sex.After the reproductive season,shell growth occurred in summer,and body weight increased from summer to winter.These growth characteristics might promote higher survival and fitness. Key words:annual ring,energy allocation,gender,growth,Nucella spans(Crisp1984;Tamai1988). INTRODUCTION Energy allocation between growth and reproduc tion has also been a central topic in life history studies. Growth has received considerable attention in life his Many reseachers have studied the relationship among tory studies,and the starting point of any comprehen reproduction,shell growth and soft part growth in gas sive analysis of life history is the aging of individuals tropods(e.g.Webber1970;Stickle1973).Differences in in a population(e.g.Roff1992;Stearns1994).Gastro reproduction and growth between males and females pods and bivalves have been aged using rings on their have also been studied.The synchronized reproductive shell surfaces(e.g.Uno1962;Williamson&Kendall output periods in males and females in Nucella lapillus 1981;Goshima et al.1991;Kawai et al.1993).How (see Feare1970a)and Thais lamellosa(see Stickle1973) ever,many animals have no such distinct characters.In suggest that in both species,males and females have such cases,the size frequency distribution is commonly similar body growth patterns.However,males and fe divided into each age class using the arithmetic prob males of the dogwhelk N.freycineti have completely ability method described by Harding(1949)(Goodwin different reproductive cycles(Kawai&Nakao1993). 1978;Gardner&Thomas1987;Staikou et al.1988),but Therefore,males and females may have different this method is not applicable for animals with long life growth patterns and allocate energy differently. The objectives of this research were to find shell characters that can be used for aging N.freycineti,to Received April14,2000:Accepted September22,2000 *Corresponding author.Present address:Research Cen determine its growth equation,to examine the growth ter for the Pacific Islands,Kagoshima University, patterns of soft parts and shells in males and females, Kourimoto1-21-24,Kagoshima890-8580,Japan. and to discuss different energy allocation patterns 43 Kawai between males and females,and between shells and soft To ascertain the reliability of the bands as growth parts. markers,the time and cumulative frequency of their formation were determined by observing the shell edges.To estimate the shell length at the time of band MATERIALS AND METHODS formation,a linear regression equation of shell length against shell diameter was calculated.Since males and Sampling was carried out monthly at Usujiri(41•K57'N, females of Neptunea arthritica had different reproduc 140•K58'E),Hokkaido,Japan from September1987to tive cycles (Fujinaga1985)but similar shell growth September1991.Details of the study site are given in patterns(Fujinaga1987),I assumed that the shell Kawai&Nakao(1993).To examine the ring formation, growth of Nucella freycineti is also similar between about30dogwhelks were randomly collected every males and females. month at area A(Fig.1)during spring tide from To compensate for the problem in the use of the August1987to September1988.After each shell length arithmetic probability method for long life-span ani was measured with a caliper,the shell was cut from the mals,I divided the monthly size-frequency distributions second spiral rib in the body whorl vertically against into each age class based on the age distribution in the the columella.The inside of the shell,including the field sample determined by ring analysis,and the sea apex,was removed with a small grinder.About390 sonal growth rate of each age class.The age snails treated in this manner were filed using sand distribution study was carried out monthly from De paper, steeped in0.1N Hydrochloric acid for12hours, cember1987to December1988and from April1989to and then photographed using X-ray photography September1992at70stations(area B)(Fig.1)using a (CMB-2,Softex Inc.).Bands in the photos were identi 1x1m2quadrat.Six subtidal stations were observed fied and the shell diameter of each band was measured. from December1987to December1988,but eliminated in the latter survey because no dogwhelks were observed at these stations.After measuring their shell lengths in the field,they were returned to their original positions in the field.To determine the age distribution,all dogwhelks at64stations were collected in September 1992and aged by the methods mentioned earlier.To de termine the seasonal changes in mean shell length in each age class,samples were collected from15stations (area C)(Fig.1)in June,September,December1991 and March1992.Ages in each sample were determined by the method descrived above.The size frequency dis tributions in area B were divided into each age class using a computer program(Tsutsumi&Tanaka1987), based on the population age structure in September1992 and the seasonal mean shell length of each age class in area C.To construct the growth curve,mean shell lengths at successive growth bands were fitted to the von Bertalanffy equation,which was modified by Moreau(1987)as follows: Lt=L•‡(1-exp(-k(t/12t0)+A sin (2ƒÎ(t/12-ts)))), where Lt,is the shell length(mm)at time t(month), L•‡is the theoretical maximal shell length(mm),k is a growth constant,t0is the theoretical time(month) when the shell length equals zero,A is ck/2ƒÎ,c is a con Fig.1.Map showing the study areas(A,B and C)and ap stant of fluctuation in monthly growth,and ts is the proximate height in meter from M.T.L.Black circles time(month)from t0to the beginning of the first fluc show the sampling stations. tuation of shell growth.Parameters of the equation 44 Œû‘¸/AB•ç•œd•œƒS'.'•ç‡_..‹œ‡_‚ïƒÌ"6'•A•E•E.•‰•B.•E•E•c.—•9..'‚•99/Om440•œHokkaido•œ•`“±;}.•E42•E•œ/ŽOUsujiri•E‚ñ.q•E•E'“K•E4•E.•E42.•E44.'Ž¢•œ•A•c•`•`6Š••A•E•E•••E•œŽOƒÁ•A•œ•œ..•E•E!tq‘Ï•E...Šs•E•E•B'•œ"•œ'50m'•E.•œ•B•B•E•B50..•œ./.ƒ{8.ƒ{'•‘‹¿•œƒÁ•E’Ê•Eˆñ‰T'•E.c•ˆ.!•œ•A•E'•œ•‰!•œ.'•E•Emáo•œ.•œ.•œ•œmle/“ñ7•œ•œ•œ Growth in Nucella freycineti were calculated by a computer program using the sim 100 plex method(Okumura1986). Using the formula describing the relationships δ among shell length,total soft part dry weight and 呂75 ・∈ gonad in each sex(Kawai1993),and the von Bertalanffy equation determined in this research, 畜 monthly dry weights of each part in each age classwere ε5・ calculated.The sizesat maturity of males and females 皇 ・ § were15mm and19mm,respectively(Kawai&Nakao 1993).For snails smaller than these sizes,the equation §25 for the sex-undetermined classwas used. L/ O ASONDJFMAMJJAS Month RESULTS Rings as an age character and age determination Fig.3.Diagram showing the proportion(%)of the ani mals with a new band at the shell margin from August Growth rings were clearly observed in the X-ray photos 1987to September1988. (Fig.2).White-colored rings were distinguished from black-colored parts.About5%of the specimens had formed a new ring by August(Fig.3).The frequency of 40Jun.1990 dogwhelks with a new ring steeply increased from No- 30 vember,reaching100%in February. The liner regression equation between shell length 20 (SL)and shell diameter(SD)was SL=1.48SD-0.97 10 (r2=0.97,n=50).The maximum number of rings ob 0 served was eleven.The mean(±SD)shell lengths for ring#1,2,3,4,5,6,7and8were6.01±2.20,12.05 40Sep .1990 ±1.20,16.40±1.23,19.64±2.01,23.99±3.01,25.90 30 ±2 .20,27.01±3.05,and28.12±2.10mm,respectively. The mean shell length of the newly settled dogwhelks in毒諜、伽._、.藤 ω20 冠 ';弓10 も0 .≡ 逼40D。 、.1990 230 ∈ ≡20, 10 0 40Mar ,1991 30 20 10 0 12340 Fig.2.Growth bands in an adult Shelllength(mm》 Numbers show each growth band,andNucella freycineti.d7corresponds to Fig.4.Size-frequency distribution of shell length at sta the diameter of the7th ring formation. tion C from June1990to March1991. 45 Kawai Table1.Mean shell length(mm•}SD)in each age class February in the field was5.25•}1.53mm and was not from June1990to March199 1.MonthAgeJun.Sep.Dec.Mar.17.72•}2.5410.54•}4.14212.75•}2.1915.0217.91•}0.3916.67•}1•B15318.64•}3.7219.22•}4.2719.35•}2.0621.27•}5.00421.82•}4.6922.97•}5.2523.43•}4.6424.55•}3.35524.46•}3.4725.13•}6.0324.58•}3.5227.58•}8.07626.71•}5.5228.62•}9.1027.87•}4.2727.81•}1.11727.91•}6.5431.69•}6.9528.64•}4.8733.01•}7.05828.70•}2.4835.61937.85 significantly different from the mean shell length of ring#1(U-test,P>0.05),suggesting that the first ring recognized in the X-ray photos formed in the first winter for newly hatched snails.
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