Seed production and growth of Modiolus capax Conrad (Bivalvia: Mytilidae) in laboratory conditions.
|Subject:||Fishery management (Methods)|
Mungaray, Miguel Robles
Serrano-Guzman, Saul Jaime
|Publication:||Name: Journal of Shellfish Research Publisher: National Shellfisheries Association, Inc. Audience: Academic Format: Magazine/Journal Subject: Biological sciences; Zoology and wildlife conservation Copyright: COPYRIGHT 2007 National Shellfisheries Association, Inc. ISSN: 0730-8000|
|Issue:||Date: Dec, 2007 Source Volume: 26 Source Issue: 4|
|Topic:||Computer Subject: Company growth|
|Product:||Product Code: 9108581 Fisheries Management Programs NAICS Code: 92614 Regulation of Agricultural Marketing and Commodities|
|Geographic:||Geographic Scope: United States Geographic Code: 1USA United States|
ABSTRACT The complete ontogeny and postlarval growth of Modiolus
capax are described for the first time. Spawning was easily achieved by
a gradual increase in water temperature (l[degrees]C/h) from
26[degrees]C to 31-32[degrees]C after exposing the breeders to air for 6
h. The veliger stage (103 [+ or -] 8 [micro]m) was reached 23-24 h after
fertilization and the low umbo stage (165 [+ or -] 9 [micro]m) on the
fifth day of culture; growth thereafter was temperature-dependent. At
28.5 [+ or -] 0.5[degrees]C, competent pediveligers (275 [micro]m) were
first observed on the tenth day of culture and on the twelfth day at
26.5 [+ or -] 0.5[degrees]C, but survival was twice (42%) that obtained
at 28.5[degrees]C. Most larvae settled between 290 and 310 [micro]m;
metamorphosis took 3-4 days. For settlement, several suspended
artificial substrata were tested; ropes of twisted anchovy fishing nets
captured the larger percentage. However, the use of a downwelling system
was the most efficient strategy for the settlement of M. capax over
artificial substratum. After metamorphosis, the shell hinge slowly moved
toward a more posterior position, and at 440 [micro]m the disconch had
adventitious serrated hairs. Seed growth was exponential and
temperature-dependent. At 28.0 [+ or -] 0.5[degrees]C, growth ranged
from 26-127 [micro]m [day.sup.-1], and from 24-63 [micro]m [day.sup.-1]
at 24.8 [+ or -] 1.7[degrees]C. In both cases, survivorship from
pediveliger to seed was approximately 80%.
KEY WORDS: Modiolus capax, mussel, seed production
Mytilus spp, the most thoroughly studied and exploited mussel genus, does not occur in the Eastern Tropical Pacific or in the Gulf of California (Keen 1971, Brusca 1980). In these subtropical and tropical regions, Garcia-Cubas and Reguero (1987) reported 27 mussel species; most are, however, scarce throughout their geographic distribution or are smaller than 40 mm. The fat horse mussel Modiolus capax (Conrad, 1837) stands out as a potential candidate for culture operations because of its large size, gregarious habits, and wide geographic distribution. Modiolus capax can reach over 100 rum in shell length, forms dense mussel beds in the intertidal and subtidal down to 46 m, and is found from Santa Cruz, CA to Paita, Peru, including the Galapagos Islands and the Gulf of California, where it is especially abundant.
To assess the feasibility of culturing M. capax, Aguirre-Hinojosa and Buckle Ramirez (1991) tested a number of strategies to collect seed from the wild, but with generally poor results. Under laboratory conditions, Orduna-Rojas and Farfan (1991) induced the spawning and reared the larvae of M. capax up to the pediveliger stage. In this work, we report the successful production of seed and subsequent growth under laboratory conditions.
MATERIALS AND METHODS
For the two culture experiments, adult mussels 5.5-7.0 cm in length were collected by scuba diving in San Carlos, Sonora, Mexico (27[degrees]56'N, 11l[degrees]03'W). Their transport to the laboratory in Bahia Kino, 2 h away by car, was carried out in ice coolers or inside seawater wet jute bags. Upon arrival, the mussels were cleared of epibionts, placed in groups of 20 organisms in Nestier boxes suspended inside a 2-ton fiberglass tank filled with filtered seawater (10 [micro]m, 35 [+ or -1] 1 [per thousand], 26[degrees]C), and let to depurate for 48-72 h before the inducement of spawning.
Because of advancement of the spawning season (late August and September), a low number of animals were expected to spawn; thus 60-80 mussels were tested in each trail. Sudden heat shock trails to induce spawning failed but it was easily achieved by a gradual increase in water temperature (1[degrees]C/hr) from 26[degrees]C to 30-31[degrees]C after 6 h of air exposure, as recommended by Orduna-Rojas and Farfan (1991). The operation took place inside a 3.5-ton fiberglass tank filled with filtered (5 [micro]m) seawater. Once completed, the mussels were removed from the tank, and an air line was introduced to provide gentle aeration to the spawned products.
The veliger stage was reached 23-24 h later. The larvae were recovered on a 60-[micro]m mesh sieve, washed, resuspended in a 10-L bucket to draw several aliquots for counting and measuring, and transferred to a 3.5-ton fiberglass tank (no repetitions) provided with filtered seawater (5 [micro]m, 35 [+ or -] 1 [per thousand]), continuous aeration, and a mixture of two microalgae.
In the first experiment, the number of D larvae recovered was 1.5 [+ or -] 0.25 x [10.sup.6], and 1.2 [+ or -] 0.20 x [10.sup.6] in the second experiment. In both culture experiments, a mixture of Isochrysis galbana (TISO) and Chaetoceros calcitrans (CCAL) was provided in a 1:1 or 2:1 ratio, according to availability, at a concentration of 35 x [10.sup.6] cell [mL-.sup.-1] up to the fourth day, and twice this concentration until the ninth day. Every other day, the water of the rearing containers was changed, and all the larvae retained on an appropriate mesh sieve were rinsed, resuspended, sampled for density and growth assessment, and returned to a clean container with fresh rearing medium. On the other days, we checked water temperature, salinity, dissolved oxygen, and pH along with microalgae concentration to replenish the consumed cells.
On the tenth day of culture, a shift was made from static (batch) culture to a flow-through mode, at which time the larvae were presented to the artificial substrata. During the first experiment, the pediveligers (300 [+ or -] [10.sup.3]; 276 [+ or -] 20 [micro]m) remained in the 3.5-ton fiberglass tank and were fed with a 2-4 L [min.sup.-1] flow of filtered seawater containing the TISO-CCAL microalgae mixture at 125 [+ or -] 43 x [10.sup.3] cell [mL.sup.-1]. On the 25th day of culture, the water flow was increased to 8-12 L [min.sup.-1] and maintained until the conclusion of the experiment (Day 55). In this experiment, we tested several substrata: plastic disc (20 cm in diameter) stacked in modules of 20, monofilament fishing nets inside vexar bags, 1/4" yellow polypropylene rope wound around PVC frames (0.5 x 0.5 m), and ropes of twisted anchovy nets also wound around PVC frames. To leach out chemicals and for conditioning, the collectors were kept in running seawater for two weeks and weighed before their introduction into the culture system. Five of each of the artificial collectors was maintained in suspension inside the rearing tank. For the first two weeks, seed collectors were checked daily, and every other day thereafter. Spat samples (n [greater than or equal to] 15) for growth assessment were obtained every other day. Partial renovation of the culture medium (1/2-1/3 of total volume) was made every 4-6 days. The incoming and outgoing water flow and microalgae concentrations were monitored once per day, along with water temperature, salinity, and pH, which remained at 28.5 [+ or -] 0.5[degrees]C, 35 [+ or -] 1 [per thousand], and 8.2 [+ or -] 0.1 pH units, respectively.
In the second experiment, the pediveligers (550 [+ or -] 20 x [10.sup.3]; 268 [+ or -] 15 [micro]m) were equally divided in two downwelling systems. We used 1-m-diameter sieves with a mesh size of 180 [micro]m seated inside rectangular fiberglass tanks (1.5 x 1.0 x 2.0 m) irrigated with a seawater flow of 1-2 L [min.sup.-1] containing the TISO-CCAL microalgae mixture at 124 [+ or -] 39 x [10.sup.3] cell [mL.sup.-1]. From Day 28 of culture until the end of the experiment (61 days), water flow rate was increased to 8-12 L [min.sup.-1]. For settlement, a 0.5 x 0.5 m PVF frame, around which 20 m of preweighed wet tope of twisted anchovy fishing net was wound, was placed on top of each sieve. Every other day, sieves and fiberglass tanks were cleaned, and a spat sample was taken for observation and growth assessment (n [greater than or equal to] 15); once per day, incoming and outgoing water flow and microalgae concentration were checked, as well as water temperature, salinity, and pH. Throughout the experiment, these variables averaged 24.8 [+ or -] 1.7[degrees]C, 35 [+ or -] 1 [per thousand], and 8.14 [+ or -] 0.1 pH units, respectively.
Settlement was assessed at the very end of the experimental periods. To estimate settlement on the plastic discs, we counted the spat on each side of all 20 discs from three modules. The spat on the walls of the fiberglass tanks was scraped, and their number assessed by weight. Settlement on the remaining artificial collectors was also estimated by weight. At the end of both experiments, 15 samples of spat each containing 50-150 organisms were weighed ([+ or -] 0.001 g), and the average weight was used to assess the number of spat settled on the surface of the culture tank and on each seed collector; spatfall and seed morphometric relationships were later assessed in several subsamples.
The unfertilized egg diameter ranged from 70-90 [micro]m and averaged 73.4 [+ or -] 7 [micro]m. Early trochophore larvae (92.5 x 87.5 [micro]m) developed approximately 9 h after fertilization (AF), and larvae with straight hinges and well-developed ciliated velum developed in 23-24 h. The average shell length (greatest distance parallel to the shell hinge) of D-shaped larvae was 103 [+ or -] 8 [micro]m, and that of low umbo veligers, first observed the fifth day of culture, averaged 165 [+ or -] 9 [micro]m (Fig. 1a to 1b). Early knobby umbo veligers, eyeded pediveligers, and competent pediveligers measured 210 [+ or -] 10, 236 [+ or -] 10 and 292 [+ or -] 17 [micro]m, respectively (Fig. 1c to 1d).
The effect of temperature on larval growth was evident by the rime the larvae reached the knobby umbo stage. During the first experiment, water temperature was 28.5 [+ or -] 0.5[degrees]C, and these larvae were observed for the first time on the seventh day of culture, one day sooner than in the second experiment when the water temperature dropped to 26.5 [+ or -] 0.5[degrees]C. The lag for the competent pediveliger stage was even longer; these larvae were first observed on Days 10 and 12 of culture, respectively. The estimated daily larval growth rate at 28.5[degrees]C was 18.4 [+ or -] 0.44 [micro]m [day.sup.-1], and at 20.5[degrees]C it was 15.4 [+ or -] 0.28 [micro]m [day.sup.-1] (Fig. 2a). In contrast, larval survivorship was lower at the higher water temperature (20% and 47.5%, respectively).
Most larvae settled between 290 and 310 [micro]m, although pediveligers as large as 350 [micro]m were observed swimming. Metamorphosis took 3-4 days. Abundant epibiosis over the shell was observed in over 80% of the spatfall > 320 [micro]m on Day 13 of culture at 28.5[degrees]C and on Day 16 ar 26.5[degrees]C. After metamorphosis, there was a slow displacement of the hinge toward the posterior end and the shell (Fig. 1e to 1g), and at 440 [micro]m, the disconch had adventitious serrated hairs (Fig. 1h to 1i); spat larger than 1,000 [micro]m had hairs over most of the shell (Fig. 1j to 11).
In both experiments, survivorship from pediveliger to spat was approximately 80% (230 [+ or -] 15 x [10.sup.3] and 350 [+ or -] 25 x [10.sup.3], respectively). In the first experiment, most settlement (60%) occurred on the bottom of the culture vessel and on the ropes made of anchovy fishing nets (25 [+ or -] 8%). Settlement on the 1/4" polypropylene ropes and monofilament fishing nets was 7 [+ or -] 3% and 5 [+ or -] 3%, respectively. The plastic discs contained 3 [+ or -] 1% of the seed, of which most were located on the upper side and periphery of the discs. Based on these results, in the second experiment we used a downwelling system and only ropes made of twisted anchovy fishing net for settlement, after which practically all the spat settled on the ropes.
After settlement, growth was exponential (Fig. 2b). During this growth phase, there was a larger difference in water temperature between the two experiments (28.0 [+ or -] 0.5[degrees]C and 24.8 [+ or -] 1.7[degrees]C). Spat size at the end of the first experiment ranged from 2,280-6,300 [micro]m, and growth rates, estimated at 10-day intervals, ranged from 26-127 [micro]m [day.sup.-1]; at the end of the second experiment, the spat measured 1,770-3,180 [micro]m, and growth varied from 24-63 [micro]m [day.sup.-1]. In both experiments, the daily growth in shell length (ShL) was best described ([r.sup.2.sub.adj] > 0.96) by the equation ShL = a x [e.sup.(b x Day of culture)]; the estimated parameters are in Figure 2b. The relative instantaneous growth indexes for the total length of each experiment (T) estimated as K (Log Sh[L.sub.f] - Log Sh[L.sub.i])/T (Wilson 1977) were 0.027 and 0.017, respectively.
The morphometric relationships of ShL versus height and depth of larvae and spat were linear; the relationship between ShL and total weight of spat (1,000-6,300 [micro]m) was exponential. Plots and equations are in Figure 2c to 2d. Roughly, larval shell height was 81 [+ or -] 5% of ShL, and shell depth was 58 [+ or -] 9% of ShL. In spatfall [greater than or equal to] 500 [micro]m, shell length and height tended to be similar; thereafter, shell height was 69 [+ or -] 7% of ShL and shell depth was 55 [+ or -] 4% of ShL.
[FIGURE 1 OMITTED]
There are more than 100 Modiolinidae species (OBIS 2006, Soot-Ryen 1955). Larvae from plankton samples have been described for Modiolaria adriatica (L), M. discolor (L), M. marmorata (Forbes), M. nigra (Gray), Modiolus demissus (Dillwyn), and M. modiolus (L) (reviewed by Chanley 1970). Larval development under laboratory conditions has also been reported for M. demissus (Loosanoff et al. 1906, Chanley & Andrews 1971) and M. modoiolus (de Schweinitz & Lutz 1976) and three other species: Modiolara impacta, Xenostrobus pulex (Redfearn et al. 1986), and M. capx (Orduna Rojas & Farfan 1991). The earlier studies were devised as aids for identification of bivalve larvae in plankton samples; thus, they are very thorough in shell morphology, but with the exception of the work describing M. modiolus, they provide no information on the chronology of the different ontogenic stages. As for the M. capax study, larval settlement was not reached. Here, we report the complete ontogeny and postlarval growth of M. capax under laboratory conditions.
The early ontogeny observed in this work was in agreement with that previously reported by Orduna Rojas and Farfan (1991). Briefly, from 24-28[degrees]C, the extrusion of the polar bodies occurred within 10 min AF, first cleavage occurred ar 1 h AF, morula stage at 3 h AF, cilliated gastrula at 7 h AF, early trochophore at 10 h AF, late trochophore 20 h at AF, and D shaped-larvae with well-developed velum developed at 23-24 h AF. Most studies on mussel larval development do not provide details on the chronology of the embryonic stages; if, however, the straight hinge larvae hatch in 23-24 h, the timing for the embryonic stages are likely to be similar to those observed in M. capax and Perna viridis (Tan 1975, Sreenivasan et al. 1988).
[FIGURE 2 OMITTED]
At both experimental temperatures (28 and 26.5[degrees]C), M. capx low umbo veliger was reached on Day 5 of culture, after which growth was temperature-dependent. At 28[degrees]C, the knobby umbo, eyed, and pediveliger larvae were observed for the first time on Days 7, 8, and 9 of culture, respectively; at 26.5[degrees]C, these stages were reached 1-2 days later, and 3-8 days later at 24[degrees]C--the experimental temperature used by Orduna Rojas and Farfan (1991). Given the increase in growth rate with increased water temperature, and because M. capax is a tropical and subtropical mussel, perhaps the pediveliger stage is attainable in 8 days if larvae are cultured at 30[degrees]C to 31[degrees]C. However, based on the observed lower larval survival at higher temperatures, the near-optimum culture temperature is most likely around 27[degrees]C to 27.5[degrees]C. The larval development of most temperate mytilids is 16-26 days long (Redfearn et al. 1986, Lutz & Kennish 1992, Satuito et al. 1994, Toro et al. 2004), but in tropical mytilids such as Perna perna and P. virids, competent pediveligers can be obtained in 9-11 days of culture at 31[degrees]C (Siddall 1978, 1979, Nair & Appukutan 2003); the near-optimum temperature for larval culture or settlement, however, is lower.
Modiolus capax eggs, D-shaped larvae, and competent pediveligers are relatively large (73 [+ or -] 5, 103 [+ or -] 8 and [greater than or equal to] 270 [micro]m, respectively). For most mytilids, the average egg ranges from 50-70 [micro]m in diameter and hatch as D-shaped larvae smaller than 90 [micro]m, and their competent pediveligers are usually smaller than 270 [micro]m (Losanoff et al. 1966; Chanley & Andrews 1971, Redfern et al. 1986, Lutz & Kennish 1992, Nair & Appukutan 2004). Perna indica and Modiolus modiolus are mussels with large metamorphosing pediveliger--such as M. capax--which also spawn relatively large eggs (Kuriakose 1980, Appukutan et al. 1984, Appukutan et al. 1988, de Schweinitz & Lutz 1976). In general, the M. capax larval shell is almost symmetrical in longitudinal profile and has low and knobby umbones. In most larvae, shell length is 27-38 [micro]m longer than shell height, but in plantigrades [greater than or equal to] 500 [micro]m, shell length and height tend to be similar. This change in shape is concurrent with the displacement of the hinge toward a more posterior position. Another distinct characteristic of the spatfall are periostracal serrated hairs, first present over the disconch and later over most of the shell. Perhaps this latter event marks a change from postlarval life to the juvenile stage, when the mussels are no longer capable of detaching and pursue another site for settlement. Periostracal hairs are characteristic of the genus Modiolus. Among their functions are camouflage, protection against desiccation, fouling by byssus threads produced by conspecifics, and predation from encrusting and boring organisms. In burrowing forms, the hairs also serve to stabilize the shell in soft substrata (Dixon et al. 1995). Besides these functions, the periostracal hairs of M. capax adults, as well as their byssus threads, act as 'seed collectors'; in the wild it is infrequent to find mussel spat elsewhere.
Given a choice, M. capax prefers dark and fibrous artificial substrates for settlement. Similar results were reported by Aguirre Hinojosa and Buckle Ramirez (1992) for M. capax in the wild. They analyzed the annual pattern of M. capax settlement in Bahia de los Angeles, B.C., Mexico, using two types of artificial substrates in suspension: rubber strings and ropes of twisted anchovy fishing net. Most of the spat was found on the latter; in general, however, settlement was very poor. This could be partially explained by the tendency of M. capax to settle to the bottom, not on suspended structures. In our first experiment, even though the water column of the culture vessel was practically filled with artificial collectors, 60% of the larvae settled on the bottom of the tank and 23% settled on the anchovy fishing net collectors. In contrast, in the downwelling system, >90% of the larvae settled over the fishing net ropes. This result also suggests that the scant settlement found on hanging artificial substrates in the wild as well as over natural mussel bed--as found in population studies carried out in Bahia de los Angeles, B.C., and La Paz, B.C.S. (Garza-Aguirre & Buckle-Ramirez 1989, Ochoa Baez 1987)--is most likely due primarily to poor larval or postlarval survivorship. In both locations, the number of seed over the mussel beds was <20% of the total population, an extremely low recruitment rate when compared with those reported worldwide for natural populations of Mytilus-like species (Curiel-Ramirez Gutierrez 2000, Caceres-Martinez 1994, Dare & Davis 1975).
The growth rate of M. capax seed was exponential and temperature-dependent. At 28.0 [+ or -] 0.5[degrees]C, growth ranged from 26-127 [micro]m [day.sup.-1], and at 24.8 [+ or -] 1.7[degrees]C from 24-63 [micro]m day-1. These rates are high and comparable with those reported for M. galloprovincialis maintained in laboratory conditions (Satuito et al. 1994, Curiel-Ramirez Gutierrez 2005). They also agree with the growth assessed in recently settled M. capax on suspended artificial collectors in Bahia de los Angeles, B.C. (Aguirre Hinojosa & Buckle Ramirez 1992). Growth of spatfall settled over mussel beds is usually slower; in Bahia de La Paz B.C.S., the estimated relative instantaneous growth index of M. capax settled in late summer is 0.015 (Ochoa Baez 1987), which is consistent with the index calculated in this study (K = 0.017) for the seed raised at lower temperatures.
The authors thank Alfredo Herrera Mesina and Francisco Hoyos Chaires for allowing us to conduct this research in the facilities of the Sonora State Oyster Center (Centro Ostricola del Estado de Sonora; COES). Special thanks are due to the COES technical staff, and to Leticia Monrroy Lara and Carlos Ceballos for their assistance in rearing the larvae.
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CLAUDIA FARFAN, (1) * MIGUEL ROBLES MUNGARAY (2) AND SAUL JAIME SERRANO-GUZMAN (2)
(1) Departamento de Acuicultura, Centro de Investigacion Cientifica y de Educacion Superior de Ensenada, Km 107 Carretera Tijuana-Ensenada, Ensenada, Baja California, 22860, Mexico; (2) Centro Ostricola dei Estado de Sonora (COES). Bahia Quino, Sonora, Mexico
* Corresponding author. E-mail: email@example.com.
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