MARINE ECOLOGY PROGRESS SERIES Vol. 168: 127-134, 1998 Published July 9 Mar Ecol Prog Ser Combined effects of salinity and temperature on juvenile survival, growth and maturation in the polychaete Hydroides elegans Jian-Wen Qiu*,Pei-Yuan Qian Department of Biology, The Hong Kong University of Science and Technology, Clear Water Bay, Kowloon, Hong Kong ABSTRACT- Effects of salinity (15 to 35%) and temperature (15 to 30°C) on the survival and growth of juven~lepolychaete Hydroides elegans (Haswell),and on their subsequent reproduction, were exam- ined in 2 laboratory experiments. Within the experimental range, temperature did not influence sur- vivorship. However, low salinity reduced survivorship: at 15x0,all juveniles died within 8 d; at 2076.. about 30Y0 of the individuals died within 8 d, but almost all remaining individuals survived throughout the following 17 d of the experiment. At hlgher salinities, over 95 % of the individuals survived through- out the experiment. Juveniles were more vulnerable to low salinity (20%) wikn 1 d of settlement than at older ages. Both low temperature and low salinity led to slower growth and subsequently to a longer time to maturation. At t25%oand t20°C, first spawning occurred on Day 16 of the experiment. More treatments were found to have mature worms on Day 25 than on Day 16 of the experiment. Tubes of reproductive individuals were usually longer than 1.2 cm. The male to female ratio varied among treat- ments from n 2 !o ? 0 and appeared to be independent of salinlty and temperature conditions. Aver- age fecundity ranged from 1100 to 9050 oocytes per female and seemed unaffected by temperature. Average fecundity was similar at salinities 225% but was lower at the lowest survival salinity (20%). This is the first study to report on growth and maturation of H. elegans under controlled laboratory con- ditions. It provides information to explain the settlement and population changes of this species in the field. The results also support the ideas that early developmental stages are more sensitive to environ- mental stress than late juveniles and adults, and that juveniles are most vulnerable at the onset of benthic life. KEY WORDS: Polychaete . Hydroides - Growth Fecundity. Maturation - Salinity. Temperature INTRODUCTION mal sexual development may require less stressful conditions (Nias et al. 1993). Experiments using the early life stages of Hydroides This study follows on from Qiu & Qian (1997) to test elegans (Qiu & Qian 1997) suggest that low salinity in the developmental sensitivity of juvenile Hydroides summer could limit embryonic and larval develop- elegans to salinity and temperature and to determine if ment, and eventually suppress the recruitment of this the results support the hypothesis that field settlement species in Hong Kong waters. However, in addition to patterns can be explained by the sensitivity of life-his- the embryonic and larval stages, the juvenile period is tory traits to environmental stress. Since the juveniles often a critical stage where heavy mortality occurs. of H. elegans are well adapted to food limitation (they Such high mortality is especially evident at the onset of can survive in 0.22 pm filtered seawater for >20 d; juvenile life due to vulnerability to stresses such as Qian et al. unpubl. data), this study focused on the sen- predation, competition, desiccation, low salinity, and sitivity of j.uvenile survival, growth, and reproductive deposition of sediment (Gosselin & Qian 1997). Fur- parameters (i.e. whether stressed conditions will lead thermore, being able to grow under stress does not to unbalanced sex ratio, decreased oocyte production guarantee a population's reproductive success, as nor- or delayed reproduction) to salinity and temperature, 2 important environmental factors that exhibit seasonal changes in Hong Kong. O Inter-Research 1998 Resale of full article not permitted 128 Mar Ecol Prog Ser 168 127-134, 1998 MATERIALS AND METHODS food conditions, H. elegans matured in about 2 wk post-settlement. Therefore, starting from Day 10 of the Rearing procedures. Juveniles were developed experiment, cultures were checked daily for mature from gametes fertilized in the laboratory in April worms by filtering about 1 1 culture medium from each (Expt I) and August (Expt IT) 1997 Procedures for fer- beaker through a 25 pm mesh. Any oocyte, embryo, or tllization and larval culture were adopted from Had- larva retained on the mesh would indicate spawning fleld et al. (1994) and described in Qiu & Qian (1997). within the last 24 h. When the flrst spatvnlng occurred Competent larvae were induced to settle in Petri on the 16th day of the experiment, survivors in all dishes (diameter, 5 cm) using 105 cells n~l-'Isochrysis dishes were enumerated. Tubes of the survivors were galbana as food and the leachate of the bryozoan video recorded, then the worms in 1 of the 3 replicates Bugula neritina, which has been shown to effectively were detached and placed individually in Petri dishes induce H. elegans larvae to settle (Bryan et al. 1998). containing 10 m1 seawater to examine reproductive In each dish, 25 to 40 out of 100 larvae metamor- status. Individuals not releasing gametes and those phosed within 24 h. Larvae that did not metamor- releasing sperm and oocytes were recorded and the phose within 24 h were decanted. Juveniles settling number of oocytes produced by each female (fecun- too close to each other and excess juveniles were dity) was enumerated. Since, on Day 16, mature worms scraped off, leaving only 15 to 20 individuals per dish were found only at 225?~.,salinity and 220°C, other in Expt I and 15 to 30 in Expt TT. The setup of the 2 treatments were chcckcd daily until mature worms experiments is detailed below. were found at the lowest temperature (15°C) This hap- Expt I: Metamorphosis to maturation. Expt I, com- pened on Day 25, at which time the experiment was menced on 21 April 1997, was designed to examine terminated. Survivors in the 2 remaining dishes in how salinity and temperature affect survival, growth each treatment were rnl~nte?,their tl~heswere \rides and maturation of newly metamorphosed juveniles. recorded, and they were then detached to determine The experimental ranges of saljnity (15 to 35%) and reproductive status. temperature (15 to 30°C) were comparable to those Expt 11: Age-dependent tolerance to low salinity. encountered in the eastern part of Hong Kong waters Expt I revealed that at 20%0salinity, about 30'ic,of the (EPD 1996). The setup followed a 2-factor design, worms died within the first 8 d of the experiment, but using 4 temperature levels (15, 20, 25, 30°C) and 5 afterwards virtually all remaining individuals sur- salinity levels (15, 20, 25, 30, 35%) for a total of 20 vived to the termination of the experiment (Day 25). treatments. Sallnities were obtained by diluting artifi- This suggests that younger juveniles may be more cial seawater (35%) with deionized water. Each of vulnerable to low salinity than older juveniles. Alter- the 20 treatments consisted of 3 replicate dishes, each natively, the lower mortality after the first 8 d of containing 15 to 20 juveniles that had metamor- Expt I might be due to acclimation of juveniles to low phosed within the previous 24 h Each Petri dish was salinity. Therefore, a second experiment, commenced tied to a plastic net (mesh size: 1 X 1 cm) and placed on 10 August 1997, was conducted to test whether in a polyethylene beaker containing 1.8 1 seawater. A age is a determining factor in the vulnerability to low total of 943 juveniles with an average Initial tube salinlty in juvenile Hydroides elegans. H. elegans length of 0.65 + 0.12 mm (n = 30) were used. The competent larvae began metamorphosing at 6 h and beakers were placed in biological incubators (Powers reached a peak about 10 to 14 h after introduction Scientific SD33SE) at the 4 designated temperatures. into Bugula neritina leachate. At 24 h, larvae that had During the experiment, seawater was changed every not metamorphosed were decanted by transferring 3 d and Isochrysis galbana was added to the cultures dishes into new culture media containing no larvae. daily to obtain an initial concentration of 2 X 105 cells At 36 h, juveniles were used for the salinity tolerance ml-l. experiment and aged as 1 d old for convenience, On the 8th day of the experiment, the num.ber of sur- although they represented juveniles which had meta- vivors in each of the d~sheswas counted and the morphosed for 12 to 30 h Juveniles of other ages (2 lengths of the survivors were measured. Since the to 10 d) were obtained by reanng the same batch of shape of a Hydro~deselegans tube is often irregular, juveniles at 35%0 salinity, 25"C, and 105 cells m1 ' direct length measurement under a microscope is 1sochry.sis galbana until the day of the experiment. impossihle. Therefore, H. elegans tubes were recorded Petrl dishes (diameter 5 cm) containing 1, 2, 5, and using a video camera mounted on a dissecting micro- 10 d old juveniles were each placed in a beaker con- scope (magnification:6.3~) (Qian & Pechenik in press). taining 0.6 1 seawater (salinity: 15, 20, and 355:,) and Tube Images were then digitized and lengths were I. galbana at a concentration of 105 cells ml-' (3 repli- measured using an image analysis software (Optimas catedtreatment, each containing 15 to 30 indiv~du- 5.2). A preliminary study showed that under ample als). Different salinitles were obtained by dilutlng Qiu R Qlan: Salinity, tempelrature and juvenile polychaete 129 artificial seawater (35%) with deionized water. The 4 all died) ranged from 0.19 to 1.01mm d-'.