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Sexual Dimorphism and Reproduction of the Amphipod Hyale Crassi

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Sexual Dimorphism and Reproduction of the Amphipod Hyale Crassi Zoological Studies 44(3): 382-392 (2005) Sexual Dimorphism and Reproduction of the Amphipod Hyale crassi- cornis Haswell (Gammaridea: Hyalidae) Kwok-Ho Tsoi and Ka-Hou Chu* Department of Biology, The Chinese University of Hong Kong, Shatin, N.T., Hong Kong, China (Accepted April 22, 2005) Kwok-Ho Tsoi and Ka-Hou Chu (2005) Sexual dimorphism and reproduction of the amphipod Hyale crassicor- nis Haswell (Gammaridea: Hyalidae). Zoological Studies 44(3): 382-392. In the present study, we explored the sexual maturation, sexual dimorphism, and fecundity of a hyalid amphipod Hyale crassicornis in the labora- tory. This amphipod passed through 11 to 21 instars, and the mean lifespan of males and females was 178 and 175 d, respectively. Males and females attained sexual competency between instars 5 and 7 at an age of 34~44 d, and between instars 5 to 8 at an age of 38~53 d, respectively. Precopulatory pairing behavior was the criterion used for the onset of sexual maturity. Nine of 10 characters studied exhibited sexual dimorphism. Transitions of growth phases can be revealed from the allometric relationships of the sexually dimorphic gnathopods 1 and 2 to body length. Male gnathopods 2 grew in 3 phases, while female gnathopods 1 and 2 grew in 2 phases. Each female generated a total of 2 to 17 (mean, 6) broods. Brood size ranged from 3 to 33 (mean, 13) offspring and was positively correlated with the body length of the female. Each female produced 16 to 290 offspring throughout its lifespan. http://zoolstud.sinica.edu.tw/Journals/44.3/382.pdf Key words: Amphipod, Hyale crassicornis, Hyalidae, Growth, Reproduction. S exual dimorphism is often strong in change in the allometric relationship between the amphipods (Barnard 1975). The sexes of gam- flagellar length of antenna 2 and body length is a marideans can usually be distinguished by differ- good indicator of sexual maturity in male ences in secondary sexual characters, including Orchomene nanus (Moore and Wong 1996). body size, gnathopods (Moore and Wong 1996), The onset of precopulatory pairing behavior and antennae (Barnard 1971, McLaughlin 1980). (involving males grasping and carrying females), Development of these secondary sexual charac- which can be induced by a series of pheromonal ters reflects sexual differentiation and maturation stimulations (Borowsky 1990), occurs in some (Hartnoll 1982). The growth of variable character amphipods once sexual maturity is attained. Y (a sexual character measurement) relative to Precopulatory pairing behavior indicates the matu- reference character X (body length measurement) rity of males and females. As sperm cannot be is expressed by the allometric equation, Y = aXb, stored in female amphipods, mature males pair in which the constant“a”represents the propor- with females until the completion of ovulation and tional factor and“b”the allometric coefficient fertilization (Borowsky 1990). Because this behav- (Huxley 1924). Any change in level of allometry ior is energetically costly (Calow 1979, Moore (allometric coefficient) indicates a variation in the 1986a) and highly risky (Strong 1973), males relative growth rate and hence reveals a growth- begin to pair with females only when ovulation is phase transition from the sexually immature phase about to occur following molting (Iribarne et al. to pre-mature or mature phases (Charniaux- 1995). A study of Corophium volutator showed Cotton 1957, Hartnoll 1978). For instance, a that mature males selected females on the basis *To whom correspondence should be addressed. Tel: 852-26096772. Fax: 852-26035391. E-mail: [email protected] 382 Tsoi and Chu -- Reproduction of Hyale crassicornis 383 of their time to receptivity rather than their body values as 6.5~8.2, and dissolved oxygen contents size (Forbes et al. 1996). as 4.9~7.0 mg/L. A 14:10 h light: dark regime at a Hyale crassicornis (Gammaridea: Talitroidea: light intensity of between 857 and 1212 lux was Hyalidae) is distributed in intertidal areas of Kiama, maintained. New South Wales, Cape Naturaliste and Western Australia (Barnard 1974). Information on the spa- Growth tial distribution of this amphipod species in Southeast Asia is still limited. In Hong Kong Twenty-five amphipods (including 9 males waters, particular Hyale species are recognized and 16 females) were successfully reared through- (Jiang and Zhou 1982, Moore 1986b, Ren 1994), out their lifespan and examined daily for molting. but these do not include H. crassicornis. However Parameters including body length (from the ros- we have collected this species from the intertidal trum to the posterodistal end of urosomite 3 in a zone in eastern waters of Hong Kong. It is com- naturally curved posture), length and articular monly found beneath rocks and pebbles, and is numbers of left antennae 2 (from the posterior end often associated with the seaweed Ulva lactuca. of the peduncle to the anterior tip of the flagellum), The present study reports the growth pattern and and propodus length of the left gnathopods 1 and reproductive biology of H. crassicornis. Such infor- 2 (the linear distance between 2 adjacent joints of mation will enhance our understanding of the biol- the dactylus and carpus) were measured 1 d after ogy of this species found in Southeast Asia. molting. For these measurements, the amphipod was placed on a glass slide and covered with a small sheet of thin plastic film. Each amphipod MATERIALS AND METHODS was examined for no more than 2 min to minimize any effect of hypoxia and environmental stress. Culture The image of the amphipod was recorded using a compound microscope (Nikon model SE, Nikon The amphipod H. crassicornis was collected Corporation, Tokyo, Japan) and a color camera set from Tolo Harbor, Hong Kong and reared in 3 out- (Teli® CCD CS 5110, Tokyo Electronic Industry, door culture tanks at the Marine Science Tokyo, Japan), and the various body parameters Laboratory, The Chinese University of Hong Kong. were measured to 10-2 mm with a video-based Each tank contained 3000 L of natural seawater image analyzing program (Quantimet 500C, Leica, with the turnover rate of ~2600 L/d. For the growth Wetzler, Germany). Data collections were limited study, ovigerous females were randomly collected to instar 15 for males and instar 18 for females from the tanks. No more than 2 freshly released because there were only 2 males and 3 females juveniles were taken from each female. Each which survived beyond these specified instars. juvenile was individually reared in a 100 ml glass beaker filled with 80 ml of artificial seawater Sexual maturation (Instant Ocean®, Aquarium Systems, Mentor, USA). Every 2 d, the water temperature, salinity, Another batch of newly hatched juveniles pH, and dissolved oxygen content of the experi- (including 11 males and 13 females) were collect- mental vessels were monitored, and about 40 ml ed and reared under standardized laboratory con- of the culture water was replaced with freshly pre- ditions as described above. Once an experimental pared and well-aerated artificial seawater. Each juvenile attained instar 4 or sexual differentiation, individual was fed a small piece (with a volume of another sexually mature individual of the opposite 1~2 mm3) of freeze-dried krill after water replace- sex and of similar size was collected from the out- ment. Since this amphipod was found to live and door culture tanks and introduced into the experi- graze on the green alga U. lactuca, a piece of thal- mental vessel for rearing with the juvenile. The lus (10 cm2) was also provided as shelter and a pair was fed with 2 pieces of krill flesh with a vol- supplementary food source for the amphipods. ume of 1 mm3. A small piece of U. lactuca thallus The thallus piece was replaced once it had been was provided as shelter and a supplementary food severely grazed. Any fecal matter, food residue, source. Maintenance was carried out as described exuviae, and undesirable algae were removed dur- above. ing the water exchange. Throughout the experi- Each pair of amphipods was examined daily mental period, the range of water temperature was for molting, precopulatory pairing, and juvenile recorded as 24~26°C, salinity as 30~33 ppt, pH release. The 1st occurrence of precopulatory pair- 384 Zoological Studies 44(3): 382-392 (2005) ing behavior and production of a clutch shortly and dissolved oxygen content as > 7.0 mg/L. In after the behavior indicated that such experimental total, 102 females carrying early (stage I) eggs amphipods (of either sex) had attained sexual were screened. Numbers of stage I eggs (Sainte- maturity. In total, 24 experimental amphipods, Marie et al. 1990, Jones and Wigham 1993) liber- including 11 males and 13 females were success- ated from the marsupium were counted under a fully reared until maturity. The 13 females were dissecting microscope, and the body lengths of continuously observed throughout their entire lifes- these females were measured. A regression line pan. Occurrences of all pairing behaviors and of brood size versus female body length was numbers of juveniles released from each brood obtained using the method of least squares. were recorded until the death of the females. Statistical analysis Fecundity The data were analyzed with linear regres- , A 3rd batch of ovigerous females was collect- sion, Student s t-test, two-factor analysis of vari- ed from the outdoor culture tanks for elucidating ance (ANOVA), and Tukey test according to Zar the relationship between fecundity and body (1996). The test statistics of the growth phase length. Water temperatures were recorded as lines was calculated based on the method 23~25°C, salinity as 32~33 ppt, pH values as 8~9, described by Somerton (1980). The computer Table 1. Lifespan and the total number of instars recorded in Hyale crassicornis (SD, stan- dard deviation) Male Individual 1 2 3 4 5 6 7 8 9 Mean SD Lifespan (d) 262 164 248 191 199 167 111 136 121 177.7 52.9 (instar) 19 13 15 15 16 15 12 12 12 14.3 2.3 Female Individual 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Mean SD Lifespan (d) 170 201 211 159 254 150 99 143 143 258 213 138 122 185 197 150 174.6 45.2 (instar) 17 17 18 18 21 15 13 15 16 18 21 13 18 20 17 11 16.8 2.8 Table 2.
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