STATUS OF GIANT CLAM RESOURCES OF THE PHILIPPINES

Field surveys were conducted to determine distribution and abundanceof giant dam resources in the Philippines. All seven known speciesof giant clams were recorded with Tridacna crocea, T. maximaand T. squamosa occurring the most frequently. The larger speciesT. derasa, T. gigas, Hippopus hippopus and H. porcellanus wererelatively rare. The reduction of giant dam stocks in the Philippines is dueto the uncontrolled exploitation of this resource which is primarilygathered for shellcraft and as supplementary diet in coastalvillages. (Received 26 February 1988; accepted 6 July 1988)     

Growth, summer cohort output, and observations on the reproduction of brook silverside, Labidesthes sicculus (Cope) in the Kawartha Lakes, Ontario

The brook silverside, Labidesthes sicculus (Cope), is unique in Canadian waters, as it completes its life cycle in 1 year. Previous studies based on scale ageing had suggested the species was an “annual,” but we confirmed this for the first time by otolith analysis. Growth rates from both back-calculation, and the Gompertz model, indicated an asymptote near the end of the summer, and average summer growth rates of 0.77 and 0.70 mm/day, respectively. The Gompertz model gave the best fit (n = 201, r = 0.744) with an L∞ of 85.4 mm TL, and instantaneous growth rate, g, of 0.0264. Back-counting daily growth increments allowed us to show that broods of young fish were produced throughout the summer, from late May to mid August, with maximum hatch taking place in mid-July. The species is a “batch” (serial) spawner, with only a fraction of the eggs ripening in the ovary and being released at as yet undetermined intervals. Eggs of the larger immature fish in the first summer developed from 0.05 to 0.21 mm in diameter (preserved) by fall, and in mature fish of the following spring and summer, developed to 1.2 mm in diameter (preserved), 1.4 mm fresh, at spawning. Attached to each egg was a filament averaging 2.0 cm in length, adhesive in nature, and presumably for attachment to vegetation. The egg also had microscopic hairs on its surface. No evidence was found to support temperature-dependant sex determination, nor were embryos or sperm found in the ovaries of spawning females, unlike Labidesthes sicculus vanhyningi, (the southern subspecies) which has internal fertilization. The Canadian species possesses a genital papilla through which the eggs were released, and an apparently much smaller male genital papilla than the southern subspecies.     

Qualitative models of the life cycles of Calanoides acutus,Calanus propinquus,and Rhincalanus gigas

Summary The life cycle of Calanoides acutus, Calanus propinquus, and Rhincalanus gigas was analyzed in samples collected by five expeditions from 1963 to 1984 in the Scotia and Weddell Seas. The result of this analysis, and of published data suggest that C. acutus and C. propinquus have a one year life cycle. Males of C. acutus are present in the population for a short period of time during late winter and fertilization occurs in deep waters. C. propinquus mates in the surface and males are present in the population throughout the summer. Part of its population may remain in the surface waters in winter. R. gigas has a more flexible, one or two year life cycle. Late summer spawning may occur in this species.This is contribution No. 95 of the Alfred-Wegener-Institut für Polar- und Meeresforschung     

Egg size in relation to fertilization dynamics in free-spawning tropical reef fishes

In marine invertebrates that spawn by simply releasing their gametes into the water (free-spawning), fertilization success likely is often limited by low sperm concentrations, due to dispersion of mates and dispersal of gametes by water movements. Production of large, low density eggs might be advantageous when sperm concentrations consistently are low, because large target size might increase egg/sperm encounters, and more low than high density eggs could be produced per clutch. Although average fertilization success in the labrid Thalassoma bifasciatum is 95% in both group spawns (in which multiple males compete for fertilizations by producing large quantities of sperm) and pair (mono-male) spawns, it is slightly lower in pair spawns, due to low level sperm limitation that arises because pair-spawning males release near the minimum number of sperm necessary for maximum fertilization. I examined whether variation in egg size and content in T. bifasciatum and other free-spawning fishes is related to variation in spawning mode, to assess whether compensatory production of large, low-density eggs might be contributing to high fertilization success in pair spawns. I found no difference between the volume or density of eggs of (1) pair- and group-spawning females of T. bifasciatum, or (2) pair-and group-spawning congeneric species of labrids, scarids, and serranids, or (3) labrids and scarids with vigorous, rapid spawning movements (which could turbulently diffuse gamete clouds) and those with slow movements. Further, egg density does not decline with increasing egg volume among those fishes. Assuming that egg size can affect fertilization success, then sperm limitation seems unlikely to represent a significant problem for pair-spawning T. bifasciatum, probably because mates place their vents close together during gamete release. The situation regarding sperm limitation in other fishes, and effects of environmentally generated water turbulence on it, are less clear. Interspecific variation in the size and content of these fishes' eggs may relate to provisioning of offspring for different larval life-histories.     

Enzyme-linked immunosorbent assay (ELISA) used in quantification of reproductive output in the pacific oyster,Crassostrea gigas,in Korea

Polyclonal antibody developed from egg proteins of the Pacific oysters, Crassostrea gigas, was used in quantification of the eggs. Quantity of the egg protein was measured using ELISA and a weight-normalized gonadosomatic index (GSI) was then established. Histology indicated that the oysters collected from Goseong Bay, Korea, in 2000 initiated gametogenesis as early as late February and spawned mostly in late June to early July when water temperature reached 23–25 °C. A second spawning peak was also observed in late August and the spawning activity continued to late October. Oyster egg proteins could be detected by ELISA in all months except March and November; a few oysters collected during December and January contained a measurable amount of eggs. The highest monthly mean GSI, 0.423, was observed in mid-June when the oysters were ready to spawn. The maximum GSI recorded in this study was 0.667. A positive correlation was found between oyster size and fecundity; the number of eggs increased as oyster biomass increased (r²=0.7497). Our data suggest that oysters in Goseong Bay produce 3–196 million eggs during the spawning season. The immunological method developed in this study was sensitive enough to measure a small quantity of eggs and considered to be method of choice for studying bivalve reproduction.     

Egg Trading in the Chalk Bass,Serranus tortugarum,a Simultaneous Hermaphrodite

The simultaneously hermaphroditic fish, Hypoplectrus nigricans, engages in a form of reciprocal spawning called “egg trading”, where one partner gives up eggs for fertilization in exchange for the opportunity to fertilize those of the partner. Since egg trading appears to be important in the evolutionary maintenance of hermaphroditism in H. nigricans, an attempt was made to determine if another species, Serranus tortugarum, behaved in a similar fashion. S. tortugarum do trade eggs, and the phenomenon seems to be widespread in simultaneously hermaphroditic serranids. However, the cues for spawning are much subtler than in H. nigricans and other simultaneously hermaphroditic serranids. This trait appears to be an evolutionary response to the occurrence of heavy sperm competition in the form of streaking.     

The annual reproductive pattern of the Antarctic clam, <Emphasis Type="Italic">Laternula elliptica</Emphasis> from Marian Cove,King George Island

The annual reproductive cycle of the Antarctic soft-shelled clam, Laternula elliptica, in Marian Cove, King George Island was studied over a 2-year period from February 1998 to January 2000. Annual changes in the gametogenesis were investigated by measuring the percentage of area occupied by oocytes in a follicle [follicle index (FI)] and the oocyte size. In 1998, the monthly mean FI increased significantly from October to November, peaked in December, and decreased rapidly from December to January. In February and March 1999, degenerated eggs were observed in the spent follicles. Degeneration and resorption of residual eggs by phagocytosis occurred mostly in February and March in both 1998 and 1999, although the resorption process was observed year-round. The histology indicated that complete vitellogenic growth of L. ellpitica at Marian Cove takes at least a year and the clams spawn annually during the austral summer. The ripening and subsequent spawning of clams at Marian Cove in 1998 and 1999 coincided with the algal blooming (September–October 1998 and December and January 1999–2000) suggesting that in coastal Antarctica food supply is a crucial factor that governs gonad maturation and subsequent spawning along with the water temperature.     

Comparison of quantitative gonad maturation scales in a temperate oyster (Crassostrea gigas) and a sub-tropical oyster (Crassostrea corteziensis)

Quantitative scales to evaluate maturity of female gonads were compared for a temperate oyster, Crassostrea gigas and a tropical oyster, Crassostrea corteziensis grown in the same locality. C. gigas had well-defined maturation and spawning periods and a resting phase in winter; in C. corteziensis mature individuals occurred most of the year and there were several spawning peaks. Of the quantitative scales used here, average oocyte diameter and gonad coverage area, much used for C. gigas, and ovary maturity index, less used, were inadequate to distinguish the reproductive pattern of C. corteziensis, since they both skewed the degree of maturation in vitellogenic stages in favor of C. gigas. Maximum oocyte diameter and maximum cytoplasm area were different among species at vitellogenic stages and also in previtellogenic stages. Nucleus:cytoplasm ratio was significantly different in previtellogenic and spawned stages between C. gigas and C. corteziensis. The Index of gonadal development was skewed in favor of C. gigas in postvitellogenic stage. The only scale that was comparable between species was reproductive potential, but it also was one of the most laborious. Other ordinal scales commonly used, such as a visual external evaluation of the gonad, only classified correctly a quarter of the stages.     

Temperature influence on the spawning performance of artificially-matured Japanese eel, Anguilla japonica, in captivity

We studied the influence of temperature on the spawning performance of artificially matured Japanese eels, Anguilla japonica, in captivity. We used routine hormone injections to bring females and males to maturity in separate aquaria. We recorded the behavior of three pairs of such hormone-treated matured eels in an aquarium (2 replicates) at four temperatures: 14, 18, 22, and 27°C, respectively. They became active and frequently left the bottom swimming in the water column, and spawning events occurred. Females released eggs in the water column around the activity peaks. Males preceded females in reaching activity peaks (presumably the timing of sperm ejection and egg release), possibly resulting in the low fertilization we observed in this experiment. Males and females returned back to the aquarium bottoms and became quiet after spawning. On several occasions, male-female or female-female pairs were observed to ‘cruise together’ in the water column for several to tens of seconds prior to egg releasing, but no courtship behavior indicative of spawning such as pairing and chasing was observed in the eels in our study. Our results suggest that 18–22°C might be the thermal preference for spawning for Japanese eels, which approximates the temperature range of the 500 m deep water layer around the Mariana Islands seamount area, the presumed spawning site for the Japanese eel.

Sexual reproduction of Acropora reef corals at Moorea, French Polynesia

Little information is available on reproductive processes among corals in isolated central Pacific reef regions, including French Polynesia. This study examined the timing and mode of sexual reproduction for Acropora reef corals at Moorea. Spawning was observed and/or inferred in 110 Acropora colonies, representing 12 species, following full moon periods in September through November 2002. Gamete release was observed and inferred in four species of Acropora between 9 and 13 nights after the full moon (nAFM) in September 2002. Twelve Acropora spp. spawned gametes between 5 and 10 nAFM in October 2002, with six species spawning 7 nAFM and four species spawning 9 nAFM. In November 2002, spawning of egg and sperm bundles was observed and inferred in 27 colonies of Acropora austera, 6 nAFM. These are the first detailed records of spawning by Acropora corals in French Polynesia.     

Reproduction and distribution of two species of Goniastrea (Scleractinia) from the Great Barrier Reef province

The reproductive biology of Goniastrea aspera at Magnetic Island was compared with that of a very similar sympatric species, G. favulus as reported by Kojis and Quinn at Heron Island. The development of gametes was similar in both species, but there was no evidence for an adolescent protandrous period of development in G. aspera such as that recorded for G. favulus at Heron Island. Other reproductive differences between the two species were found in egg size and the mode of spawning. The eggs of G. aspera are smaller and more numerous than those of G. favulus. Goniastrea aspera expelled buoyant packets of eggs and sperm, while G. favulus had sticky sinking eggs which were released separately from the sperm. The spatial pattern of the two species was examined on the reef flat at Magnetic Island to determine whether the observed differences in spawning behaviour and egg buoyancy might have an effect on egg retention and the distribution of adult colonies. The results of this comparison failed to detect any difference in the degree of aggregation of the two species. This is not the result which would be expected if sticky sinking eggs helped retain developing larvae in the vicinity of the adult. These results, together with evidence from a wide range of coelenterates and observations on the larvae of G. aspera point to post spawning larval behaviour as the most likely factor in determining where these species will settle.     

Paralarvae of the jumbo squid,Dosidicus gigas

Abstract. The rhynchoteuthion stage of the jumbo squid, Dosidicus gigas, has morphological characteristics similar to the paralarvae of the purpleback squid, Sthenoteuthis oualaniensis, making it difficult to determine the locations, seasons, and conditions where spawning of jumbo squid occurs. In this study, 180 paralarvae of D. gigas were collected off the west coast of the Baja California peninsula and identified by sequencing a 369-bp fragment of the cytochrome oxidase I gene. Of these, 77 specimens, 0.8–6.0 mm in mantle length, were described. Indices based on morphometric ratios were used to determine whether the shapes of different body structures were reliable for identifying the paralarvae. For two ratios, the results appear to discriminate members of D. gigas from those of S. oualaniensis. Additionally, eye or intestinal photophores were not observed in any paralarvae of D. gigas. Morphological and morphometric information provided a valuable basis for the discrimination and identification of these two species.     

Spawning behavior of Arctic charr (Salvelinus alpinus): Spawning synchrony,vibrational communication,and mate guarding

A mismatch in synchrony between male and female gamete release in external fertilizers can result in reduced or failed fertilization, sperm competition, and reduced paternity. In Arctic charr (Salvelinus alpinus), males can adopt either a guard or sneak tactic resulting in both pre‐ and postcopulatory competition between males with alternative reproduction tactics. Here, spawning behavior of free‐living Arctic charr was video‐recorded, and their reproductive behavior was analyzed. From evaluating 157 spawning events, we observed that females mainly spawned with a guarding male and that the female and the guarding male synchronized timing of gamete release under sperm competition. Although sneakers spawned with higher synchrony than the guarding male in single‐male spawning events, the average sneaker released his milt less synchronized with the female than the guarding male under sperm competition. Approximately 50% of the recorded spawning events occurred under sperm competition, where each event included an average of 2.7 males. Additionally, sneakers were more exposed to sperm competition than guarding males. An influx of males, in close proximity to the female, occurred during the behavioral sequences leading up to egg release, but this influx seemed not dependent on egg release, suggesting that something else than gonadal product attracts sneaker males to the spawning female. Just before and during the actual release of gametes, the spawning couple vibrates their bodies in close contact and it seems likely that this vibrational communication between the spawning couple, which results in a larger amplitude sound wave than seen under regular courting, reveals time of gamete release to sneaker males. Thus, vibrational communication may enable synchrony between the guarding male and the female, and this might be traded against the cost of higher detectability from surrounding sneaker males, eavesdropping in close proximity.     

Effects of salinity variations on the rates of photosynthesis and respiration of the juvenile giant clam (Tridacna gigas,Bivalvia, Cardiidae)

Giant clams are in symbiotic relationship with the photosynthesizing zooxanthellae and dwell in the shallow coral reefs of the Indo-Pacific region where they may be exposed to fluctuating salinities during high levels of precipitation. This study evaluated the effects of reduced salinities (18, 25, and 35‰ as control) on the rates of photosynthesis and respiration in juvenile Tridacna gigas for 14 days. At 18‰, total mortality was recorded after day 4, so no photosynthetic measurements were subsequently conducted for this treatment. However, results showed no significant differences in the photosynthetic rates among treatments. Respiration rates were significantly increased at both low salinities resulting in low P_g/R ratios. After 14 days of exposure, photosynthetic parameters at 25‰ were not significantly different from the control which suggests that juvenile T. gigas may be able to acclimate to osmotic stress and adjust its photophysiology following reductions in salinity of up to 25‰.     

Host hybridization as a potential mechanism of lateral symbiont transfer in deep‐sea vesicomyid clams

Deep‐sea vesicomyid clams live in mutualistic symbiosis with chemosynthetic bacteria that are inherited through the maternal germ line. On evolutionary timescales, strictly vertical transmission should lead to cospeciation of host mitochondrial and symbiont lineages; nonetheless, examples of incongruent phylogenies have been reported, suggesting that symbionts are occasionally horizontally transmitted between host species. The current paradigm for vesicomyid clams holds that direct transfers cause host shifts or mixtures of symbionts. An alternative hypothesis suggests that hybridization between host species might explain symbiont transfers. Two clam species, Archivesica gigas and Phreagena soyoae, frequently co‐occur at deep‐sea hydrocarbon seeps in the eastern Pacific Ocean. Although the two species typically host gammaproteobacterial symbiont lineages marked by divergent 16S rRNA phylotypes, we identified a number of clams with the A. gigas mitotype that hosted symbionts with the P. soyoae phylotype. Demographic inference models based on genome‐wide SNP data and three Sanger sequenced gene markers provided evidence that A. gigas and P. soyoae hybridized in the past, supporting the hypothesis that hybridization might be a viable mechanism of interspecific symbiont transfer. These findings provide new perspectives on the evolution of vertically transmitted symbionts and their hosts in deep‐sea chemosynthetic environments.     

Sperm allocation pattern during a reproductive season in the copulating marine cottoid species, <Emphasis Type="Italic">Alcichthys alcicornis</Emphasis>

The sperm allocation pattern of a copulating marine cottid fish, Alcichthys alcicornis, was investigated. A total of 86 mating events using six males were conducted in aquarium tanks over 10 days, and in 36 of them, spermatozoa were collected using a false copulation method. Males released 3–8 × 10⁸ spermatozoa in early events, with the number decreasing gradually during subsequent mating events. This sperm allocation was represented as an “early investment and tapering” pattern. It was discussed why males have significantly higher sperm release in early spawning events. The reproductive behavior consists of spawning and subsequent copulation. Spermatozoa have the ability to fertilize eggs from multiple clutches, and in earlier produced clutches the level of sperm competition should be relatively low. In addition, if early spawn happens to be the first spawn with a female, spermatozoa that are released into the water column after spawning are responsible for fertilizing the female’s first clutch. The probability of this occurring should decrease dramatically as the season progresses, due to the highly synchronous seasonal spawning of females. All of these factors should select for high sperm numbers in early ejaculates. Based on such reproductive ecology of A. alcicornis we hypothesize that this sperm allocation pattern is an adaptive reproductive strategy in response to egg availability and sperm competition occurring within the ovarian cavity.     

Spawning behaviour ofCheimerius nufar in captivity

Synopsis The santerCheimerius nufar is widespread in tropical and subtropical waters of the western Indian Ocean and forms an important component of linefish catches along the east coast of southern Africa. Observations of spawning behaviour in captivity have revealed that spawning occurs during spring over a period of four months. Mating takes place at sunrise and may continue for up to 105 min (mean duration = 60 min). During spawning males become dark with prominent white markings. They become very aggressive and set up territories. Females remain a uniform silvery-pink. Mating occurs between males and females of similar sizes, culminating in egg and sperm release near the surface. Individual fish spawned up to 14 times during a morning. Streaking occurred throughout the season, with a second male joining a spawning pair and releasing gametes simultaneously. Slinger,Chrysoblephus puniceus, a dominant fish on offshore reefs in the same region, interfered with spawning throughout the season and were observed to eat eggs when they were released. The spawning strategy ofC. nufar is similar to protogynous species in several respects, indicating that this functional gonochorist may not conform to current theoretical predictions.     

Reproduction of the introduced oyster Crassostrea gigas (Bivalvia: Ostreidae) cultured on rafts in Spain

The oyster Crassostrea gigas was introduced in Spain for aquaculture purposes; however, until now, it is not known whether populations are established in the wild, being necessary to define whether this species is spawning and which environmental variables trigger this process. The influence of environmental parameters on the reproduction of C. gigas was evaluated from January 2008 to October 2009 with oysters grown on a raft in the Ría de Arousa (Galicia, NW Spain). Temperature and chlorophyll a are directly correlated to sexual maturation. Oysters can mature at temperatures below 14°C. The temperature necessary for spawning differs between the sexes, requiring a temperature above 15°C for males and 18°C for females. Females had a single massive spawn between June and September, while males had partial spawning from May to December with two peaks, one in May–September and another in October–December, with the second peak more pronounced. The first spawning peak is related to high temperatures and concentrations of chlorophyll a, and the second spawning peak is mainly related to the food availability in the water. The spawning asynchrony may be impeding establishment of wild C. gigas populations in Spain.     

Reproductive behavior of the Atlantic silverside,Menidia menidia (Pisces,Atherinidae)

Synopsis The spawning behavior of the Atlantic silverside,Menidia menidia, was studied at two sites on the North Edisto River estuary in South Carolina. Prespawning schools moved back and forth along the shoreline as the time of high tide approached. Spawning runs took place in the upper intertidal zone at high tide.Spawning fish deposited their eggs on three types of substrates: 1) the lower stems of cordgrass plants,Spartina alterniflora, 2) detrital mats, and 3) exposed cordgrass roots along erosional scraps. Spawning behavior during egg deposition and fertilization was similar for all three substrates. Females released eggs during a rapid fluttering motion of the posterior half of the body. A similar movement accompanied release of sperm by males. Behavior of fish just prior to spawning insured deposition of gametes at locations that provided protection from thermal and drying stress during development. Eggs were deposited at mean intertidal elevations of 1.8 and 1.5 meters above mean low water (MLW) at respective study sites. They were exposed to the atmosphere for approximately ten hours between successive high tides.During spawning runs in which eggs were deposited at the base of cordgrass plants, ambient dissolved oxygen concentrations of the water in the spawning area were sometimes reduced to < 1.0 mg. 1^–1. Spent fish, apparently incurring an oxygen debt while spawning, formed a nonschooling aggregation offshore from the spawning zone.The recurrent use of specific spawning substrates for egg deposition resulted in an uneven distribution of spawning runs along the shoreline at each study site.Contribution No. 409 to the Gulf Breeze Environmental Research Laboratory. Contribution No. 377 to the Belle W. Baruch Institute for Marine Biology and Coastal Research.     

Induction of meiotic gynogenesis in Atlantic cod,Gadus morhua (L.)

Gynogenesis is one of several chromosome‐manipulating techniques used in fish. In gynogenesis the male does not contribute to the genetic material of the offspring, and the sperm cells act only as stimulators in order for the egg to start development. This technique has several applications, both in aquaculture and in biological research: gynogenetic fish may be used as a step in the production of all‐female populations, the production of isogenetic‐ and inbred lines, revealing of the sex determination mechanism, construction of genetic maps, and testing of environmental vs genetic control of different traits. The aim of this study was to develop a simple protocol for production of gynogenetic cod (Gadus morhua L.) for further use in aquaculture research. Various milt dilutions and UV‐irradiation doses were tested, in order to inactivate the sperm without destroying its ability to induce egg development. This was followed by pressure treatment of the eggs shortly after ‘fertilization’ to suppress the completion of meiosis II, and thereby restoring diploidy. A dose of 9000 erg mm^?2, followed by a 5‐min pressure treatment (58.6 MPa) 180 min‐degrees after fertilization gave 100% gynogenetic larvae. Histologically, sexual differentiated fish were all females, possibly confirming female homogamety in Atlantic cod. No particular signs of reduced growth, survival or enhanced deformity rates were observed after the fish had reached the juvenile phase. Mortality was, however, high during the egg and larval stages. This protocol has made capable the production of gynogenetic cod juveniles in significant amounts using relatively simple means; the next step will be to elaborate on the technique in order to produce mitotic gynogenetic (double haploid) individuals, which are 100% homozygous.