Aspects of the biology and population dynamics of freshwater mussels in Lake Kariba and Lake McIlwaine

The essential aim of this study was to provide a broad foundation of biological knowledge upon which a programme of mussel utilization and management could be built. Results of the study are presented in three main sections. Part 1 describes the stock of Lake Kariba and Lake McIlwaine; part 2 describes various aspects of the breeding biology of the three species; and part 3 presents the results of morphological, biochemical and age analyses - aspects which are used for initial standing crop and production calculations. The final discussion concludes the thesis with a general examination in ecological terms of the factors which have influenced the development and nature of the mussel faunas of the two lakes under consideration.

Reproductive biology of blue marlin (Makaira nigricans) in the western Pacific Ocean

The reproductive biology of blue marlin (Makaira nigricans) was assessed from 1001 fish (ranging from 121 to 275 cm in eye-to-fork length; EFL) caught by Taiwanese offshore longliners in the western Pacific Ocean from September 2000 to December 2001 and from 843 gonad samples from these fish, The overall sex ratio of the catch was approximately 1:1 dur ing the sampling period, but blue marlin are sexually dimorphic; females are larger than males. Reproductive activity (assessed by histology), a gonadosomatic index, and the distribution of oocyte diameters, indicated that spawning occurred predominantly from May to September. The estimated sizes-at-maturity (EFL50) were 179.76 ±1.01 cm (mean ±standard error) for females and 130 ±1 cm EFL for males. Blue marlin are multiple spawners and oocytes develop asynchronously. The proportion of mature females with ovaries containing postovulatory follicles (0.41) and hydrated oocytes (0.34) indicated that the blue marlin spawned once every 2–3 days on average. Batch fecundity (BF) for 26 females with the most advanced oocytes (≥1000 μm), but without postovulatory follicles, ranged from 2.11 to 13.50 million eggs (6.94 ± 0.54 million eggs). The relationships between batch fecundity (BF, in millions of eggs) and EFL and round weight (RW, kg) were BF = 3.29 × 10 –12 EFL5.31 (r2 = 0.70) and BF = 1.59 × 10–3 RW 1.73 (r2= 0.67), respectively. The parameters estimated in this study are key information for stock assessments of blue marlin in the western Pacific Ocean and will contribute to the conservation and sustainable yield of

Reproductive biology, spawning season, and growth of female rex sole (Glyptocephalus zachirus) in the Gulf of Alaska

Rex sole (Glyptocephalus zachirus) have a wide distribution throughout the North Pacific, ranging from central Baja California to the western Bering Sea. Although rex sole are an important species in the commercial trawl fisheries off the U.S. West Coast, knowledge of their reproductive biology is limited to one study off the Oregon coast where ovaries were analyzed with gross anatomical methods. This study was initiated to determine reproductive and growth parameters specific to rex sole in the Gulf of Alaska (GOA) stock. Female rex sole (n=594) ranging in total length from 166 to 552 mm were collected opportunistically around Kodiak Island, Alaska, from February 2000 to October 2001. All ovaries were analyzed by using standard histological criteria to determine the maturity stage. Year-round sampling of rex sole ovaries confirmed that rex sole are batch spawners and have a protracted spawning season in the GOA that lasts at least eight months, from October to May; the duration of the spawning season and the months of spawning activity are different from those previously estimated. Female rex sole in the GOA had an estimated length at 50% maturity (ML50) of 352 mm, which is greater than the previously estimated ML50 at southern latitudes. The maximum age of collected female rex sole was 29 years, and the estimated age at 50% maturity (MA50) in the GOA was 5.1 years. The von Bertalanffy growth model for rex sole in the GOA was significantly different from the previously estimated model for rex sole off the Oregon coast. This study indicated that there are higher growth rates for rex sole in the GOA than off the Oregon coast and that there are differences in length at maturity and similarity in age at maturity between the two regions.

Biology and assessment of the painted sweetlips (Diagramma pictum (Thunberg, 1792)) and the spangled emperor (Lethrinus nebulosus (Forsskål, 1775)) in the southern Arabian Gulf

The population biology and status of the painted sweeplips (Diagramma pictum) and spangled emperor (Lethrinus nebulosus) in the southern Arabian Gulf were established by using a combination of size-frequency, biological, and size-at-age data. Transverse sections of sagittal otoliths were characterized by alternating translucent and opaque bands that were validated as annuli. Comparisons of growth characteristics showed that there were no significant differences (P>0.05) between sexes. There were well defined peaks in the reproductive cycle, spawning occurred from April to May for both species, and the mean size at which females attained sexual maturity was 31.8 cm fork length (LF) for D. pictum and 27.6 cm (LF) for L. nebulosus. The mean sizes at first capture (21.1 cm LF for D. pictum and 26.4 cm LF for L. nebulosus) were smaller than the sizes for both at first sexual maturity and those at which yield per recruit would be maximized. The range of fishing-induced mortality rates for D. pictum (0.37−0.62/yr) was substantially greater than the target (Fopt=0.07/yr) and limit (Flimit=0.09/ yr) estimates. The range of fishing-induced mortality rates for L. nebulosus (0.15/yr to 0.57/yr) was also in excess of biological reference points (Fopt=0.10/yr and Flimit=0.13/yr). In addition to growth overfishing, the stocks were considered to be recruitment overfished because the biomass per recruit was less than 20% of the unexploited levels for both species. The results of the study are important to fisheries management authorities in the region because they indicate that both a reduction in fishing effort and mesh-size regulations are required for the demersal trap fishery.

Reproductive biology of female Rikuzen sole (Dexistes rikuzenius)*

The annual ovarian cycle, mode of maturation, age at maturity, and potential fecundity of female Rikuzen sole (Dexistes rikuzenius) from the North Pacific Ocean off the coast of Japan were studied by 1) histological examination of the gonads, 2) measurement and observation of the oocytes, and 3) by otolith aging. The results indicated that ovulation occurs from September to December and peaks between September and October. Vitellogenesis began again soon after the end of the current season. Maturity was divided into eight phases on the basis of oocyte developmental stages. Mature ovaries contained developing oocytes and postovulatory follicles but no recruiting oocytes, indicating that this species has group-synchronous ovaries and is a multiple spawner. Almost all females matured first at an age of 1+ year and spawned every year until at least age 8+ years. Potential fecundity increased exponentially with body length and the most fecund fish had 15 times as many oocytes as the least fecund fish. Potential fecundity and relative fecundity were both positively correlated with age from 1 to 6+ years, but were negatively correlated, probably because of senescence, in fish over 7 years. These results emphasize that the total productivity of a D. rikuzenius population depends not only on the biomass of females older than 1+ but also on the age structure of the population.

Reproductive biology of carpenter seabream (Argyrozona argyrozona) (Pisces: Sparidae) in a marine protected area

The carpenter seabream (Argyrozona argyrozona) is an endemic South African sparid that comprises an important part of the handline fishery. A three-year study (1998−2000) into its reproductive biology within the Tsitsikamma National Park revealed that these fishes are serial spawning late gonochorists. The size at 50% maturity (L50) was estimated at 292 and 297 mm FL for both females and males, respectively. A likelihood ratio test revealed that there was no significant difference between male and female L50 (P>0.5). Both monthly gonadosomatic indices and macroscopically determined ovarian stages strongly indicate that A. argyrozona within the Tsitsikamma National Park spawn in the astral summer between November and April. The presence of postovulatory follicles (POFs) confirmed a six-month spawning season, and monthly proportions of early (0−6 hour old) POFs showed that spawning frequency was highest (once every 1−2 days) from December to March. Although spawning season was more highly correlated to photoperiod (r = 0.859) than temperature (r = −0.161), the daily proportion of spawning fish was strongly correlated (r= 0.93) to ambient temperature over the range 9−22oC. These results indicate that short-term upwelling events, a strong feature in the Tsitsikamma National Park during summer, may negatively affect carpenter fecundity. Both spawning frequency and duration (i.e., length of spawning season) increased with fish length. As a result of the allometric relationship between annual fecundity and fish mass a 3-kg fish was calculated to produce fivefold more eggs per kilogram of body weight than a fish of 1 kg. In addition to producing more eggs per unit of weight each year, larger fish also produce significantly larger eggs.

The Bay Scallop, Argopecten irradians, Massachusetts Through North Carolina: Its Biology and the History of Its Habitats and Fisheries

This article covers the biology and the history of the bay scallop habitats and fishery from Massachusetts to North Carolina. The scallop species that ranges from Massachusetts to New York is Argopecten irradians irradians. In New Jersey, this species grades into A. i. concentricus, which then ranges from Maryland though North Carolina. Bay scallops inhabit broad, shallow bays usually containing eelgrass meadows, an important component in their habitat. Eelgrass appears to be a factor in the production of scallop larvae and also the protection of juveniles, especially, from predation. Bay scallops spawn during the warm months and live for 18–30 months. Only two generations of scallops are present at any time. The abundances of each vary widely among bays and years. Scallops were harvested along with other mollusks on a small scale by Native Americans. During most of the 1800’s, people of European descent gathered them at wading depths or from beaches where storms had washed them ashore. Scallop shells were also and continue to be commonly used in ornaments. Some fishing for bay scallops began in the 1850’s and 1860’s, when the A-frame dredge became available and markets were being developed for the large, white, tasty scallop adductor muscles, and by the 1870’s commercial-scale fishing was underway. This has always been a cold-season fishery: scallops achieve full size by late fall, and the eyes or hearts (adductor muscles) remain preserved in the cold weather while enroute by trains and trucks to city markets. The first boats used were sailing catboats and sloops in New England and New York. To a lesser extent, scallops probably were also harvested by using push nets, picking them up with scoop nets, and anchor-roading. In the 1910’s and 1920’s, the sails on catboats were replaced with gasoline engines. By the mid 1940’s, outboard motors became more available and with them the numbers of fishermen increased. The increases consisted of parttimers who took leaves of 2–4 weeks from their regular jobs to earn extra money. In the years when scallops were abundant on local beds, the fishery employed as many as 10–50% of the towns’ workforces for a month or two. As scallops are a higher-priced commodity, the fishery could bring a substantial amount of money into the local economies. Massachusetts was the leading state in scallop landings. In the early 1980’s, its annual landings averaged about 190,000 bu/yr, while New York and North Carolina each landed about 45,000 bu/yr. Landings in the other states in earlier years were much smaller than in these three states. Bay scallop landings from Massachusetts to New York have fallen sharply since 1985, when a picoplankton, termed “brown tide,” bloomed densely and killed most scallops as well as extensive meadows of eelgrass. The landings have remained low, large meadows of eelgrass have declined in size, apparently the species of phytoplankton the scallops use as food has changed in composition and in seasonal abundance, and the abundances of predators have increased. The North Carolina landings have fallen since cownose rays, Rhinoptera bonsais, became abundant and consumed most scallops every year before the fishermen could harvest them. The only areas where the scallop fishery remains consistently viable, though smaller by 60–70%, are Martha’s Vineyard, Nantucket, Mass., and inside the coastal inlets in southwestern Long Island, N.Y.

Rangia and Marsh Clams, Rangia cuneata, R. flexuosa, and Polymesoda caroliniana, in Eastern México: Distribution, Biology and Ecology, and Historical Fisheries

Rangia and marsh clams, Rangia cuneata, R. flexuosa, and Polymesoda caroliniana, occur in brackish waters along México’s eastern coast from the northern State of Tamaulipas to the southern State of Campeche. The clams were important to the prehispanic people in the southern part of the State of Veracruz, where they were used as food and as construction material. In modern times, they are harvested for food. The fishermen wade in shallow water and harvest the clams in soft sediments by hand. Annual landings of whole clams during a recent 5-yr period, 1998–2002, were 1,139–1,695 t. The only area with a substantial ongoing clam fishery is in the Lower Papaloapan River Basin, including Alvarado Lagoon, where as many as 450 fishermen are licensed harvesters. This fishery for the Rangia and marsh clams is the most important clam fishery along México’s Gulf Coast.

The Northern Rockfish, Sebastes polyspinis, in Alaska: Commercial Fishery, Distribution, and Biology

The northern rockfish, Sebastes polyspinis, is the second most abundant rockfish in Alaska, and it supports a valuable trawl fishery. Little information is available, however, on either the biology of this species or its commercial fishery. To provide a synopsis of information on northern rockfish in Alaska, this study examined data for this species from commercial fishery observations in 1990–98 and from fishery-independent trawl surveys in 1980–99. Nearly all the commercial catch came from bottom trawling, mostly by large factory-trawlers, although smaller shore-based trawlers in recent years took an increasing portion of the catch in the Gulf of Alaska. Most of the northern rockfish catch in the Gulf of Alaska was taken by a directed fishery, whereas that of the Aleutian Islands predominantly came as discarded bycatch in the Atka mackerel fishery. In both regions, most of the catch was taken from a number of relatively small and discrete fishing grounds at depths of 75–150 m in the Gulf of Alaska and 75–175 m in the Aleutian Islands. These grounds, especially in the Gulf of Alaska, are on shallow rises or banks located on the outer continental shelf, and often are surrounded by deeper water. Five fishing grounds were identified in the Gulf of Alaska, and eleven in the Aleutian Islands. One fishing ground in the Gulf of Alaska, the “Snakehead” south of Kodiak Island, accounted for 46% of the total northern rockfish catch in this region. Analysis of the survey data generally revealed similar patterns of geographic distribution as those seen in the fishery, although some of the commercial fishing grounds did not stand out as areas of special abundance in the surveys. The surveys also found two areas of abundance that were not evident in the fishery data. Relatively few juvenile northern rockfish were caught in any of the surveys, but those taken in the Gulf of Alaska tended to occur more inshore and at shallower depths than adults. Individual size of northern rockfish was substantially larger in the Gulf of Alaska than in the Aleutian Islands according to both fishery and survey data. Analysis of age data from each region supports this, as Gulf of Alaska fish were found to grow significantly faster and reach a larger maximum length than those in the Aleutian Islands. Sex ratio in the Gulf of Alaska was nearly 50:50, but females predominated in the Aleutian Islands by a ratio of 57:43. In both regions, size of females was significantly larger than males.

Quahogs in Eastern North America: Part I, Biology, Ecology, and Historical Uses

The northern quahog, Mercenaria mercenaria, ranges along the Atlantic Coast of North America from the Canadian Maritimes to Florida, while the southern quahog, M. campechiensis, ranges mostly from Florida to southern Mexico. The northern quahog was fished by native North Americans during prehistoric periods. They used the meats as food and the shells as scrapers and as utensils. The European colonists copied the Indians treading method, and they also used short rakes for harvesting quahogs. The Indians of southern New England made wampum from quahog shells, used it for ornaments and sold it to the colonists, who, in turn, traded it to other Indians for furs. During the late 1600’s, 1700’s, and 1800’s, wampum was made in small factories for eventual trading with Indians farther west for furs. The quahoging industry has provided people in many coastal communities with a means of earning a livelihood and has provided consumers with a tasty, wholesome food whether eaten raw, steamed, cooked in chowders, or as stuffed quahogs. More than a dozen methods and types of gear have been used in the last two centuries for harvesting quahogs. They include treading and using various types of rakes and dredges, both of which have undergone continuous improvements in design. Modern dredges are equipped with hydraulic jets and one type has an escalator to bring the quahogs continuously to the boats. In the early 1900’s, most provinces and states established regulations to conserve and maximize yields of their quahog stocks. They include a minimum size, now almost universally a 38-mm shell width, and can include gear limitations and daily quotas. The United States produces far more quahogs than either Canada or Mexico. The leading producer in Canada is Prince Edward Island. In the United States, New York, New Jersey, and Rhode Island lead in quahog production in the north, while Virginia and North Carolina lead in the south. Connecticut and Florida were large producers in the 1990’s. The State of Campeche leads in Mexican production. In the northeastern United States, the bays with large openings, and thus large exchanges of bay waters with ocean waters, have much larger stocks of quahogs and fisheries than bays with small openings and water exchanges. Quahog stocks in certifi ed beds have been enhanced by transplanting stocks to them from stocks in uncertified waters and by planting seed grown in hatcheries, which grew in number from Massachusetts to Florida in the 1980’s and 1990’s.

The Fisheries for Mangrove Cockles,Anadaraspp., from Mexico to Peru, With Descriptions of Their Habitats and Biology, the Fishermen’s Lives, and the Effects of Shrimp Farming

This paper provides the first description of the mangrove cockle, Anadara spp., fisheries throughout their Latin American range along the Pacific coast from Mexico to Peru. Two species, A. tuberculosa and A. grandis, are found over the entire range, while A. similis occurs from El Salvador to Peru. Anadara tuberculosa is by far the most abundant, while A. grandis has declined in abundance during recent decades. Anadara tuberculosa and A. similis occur in level mud sediments in mangrove swamps, comprised mostly of Rhizophora mangle, which line the main-lands and islands of lagoons, whereas A. grandis inhabits intertidal mud flats along the edges of the same mangrove swamps. All harvested cockles are sexually mature. Gametogenesis of the three species occurs year round, and juvenile cockles grow rap-idly. Cockle densities at sizes at least 16–42 mm long ranged from 7 to 24/m2 in Mexico. Macrofaunal associates of cockles include crustaceans, gastropods, and finfishes. The mangrove swamps are in nearly pristine condition in every country except Honduras, Ecuador, and Peru, where shrimp farms constructed in the 1980’s and 1990’s have destroyed some mangrove zones. In addition, Hurricane Mitch destroyed some Honduran mangrove swamps in 1998. About 15,000 fishermen, including men, women, and children, harvest the cockles. Ecuador has the largest tabulated number of fishermen, 5,055, while Peru has the fewest, 75. Colombia has a large number, perhaps exceeding that in Ecuador, but a detailed census of them has never been made. The fishermen are poor and live a meager existence; they do not earn sufficient money to purchase adequate food to allow their full health and growth potential. They travel almost daily from their villages to the harvesting areas in wooden canoes and fiberglass boats at low tide when they can walk into the mangrove swamps to harvest cockles for about 4 h. Harvest rates, which vary among countries owing to differences in cockle abundances, range from about 50 cockles/fisherman/day in El Salvador and Honduras to 500–1,000/ fisherman/day in Mexico. The fishermen return to their villages and sell the cockles to dealers, who sell them mainly whole to market outlets within their countries, but there is some exporting to adjacent countries. An important food in most countries, the cockles are eaten in seviche, raw on the half-shell, and cooked with rice. The cockles are under heavy harvesting pressure, except in Mexico, but stocks are not yet being depleted because they are harvested at sizes which have already spawned. Also some spawning stocks lie within dense mangrove stands which the fishermen cannot reach. Consumers fortunately desire the largest cockles, spurning the smallest. Cockles are important to the people, and efforts to reduce the harvests to prevent overfishing would lead to severe economic suffering in the fishing communities. Pro-grams to conserve and improve cockle habitats may be the most judicious actions to take. Preserving the mangrove swamps intact, increasing their sizes where possible, and controlling cockle predators would lead to an increase in cockle abundance and harvests. Fishes that prey on juvenile cockles might be seined along the edges of swamps before the tide rises and they swim into the swamps to feed. Transplanting mangrove seedlings to suitable areas might increase the size of those habitats. The numbers of fishermen may increase in the future, because most adults now have several children. If new fishermen are tempted to harvest small, immature cockles and stocks are not increased, minimum size rules for harvestable cockles could be implemented and enforced to ensure adequate spawning.

Biology and Fishery for Atlantic Thread Herring, Opisthonema oglinum, along the North Carolina Coast

Thread herrings, Opisthonema spp., are small, nearshore, pelagic clupeid fishes that form dense, surface schools in tropical to subtropical coastal waters. Ecologically, thread herrings form an important forage base for many large, predatory fishes (Finucane and Vaught, 1986). Commercially, thread herrings are targeted by artisanal to moderate-sized seine fisheries off the coasts of Ecuador and Peru (Patterson and Santos, 1992), Costa Rica (Stevenson and Carranza, 1981), Venezuela, the continental margins of the Caribbean, the Gulf of Mexico, and near the islands of Cuba, Hispaniola, Puerto Rico, Jamaica, and Trinidad (Reintjes, 1978). Most of the catch is reduced to fish meal and fish oil (Patterson and Santos, 1992), although minor quantities are used for human consumption (Reintjes, 1978).

Biology and Management of Deepwater Snappers of the Hawaiian Archipelago

Commercial and recreational deepwater (100-400 m) bottom-fishing in Hawaii targets a multispecies group of lutjanid snappers. Relatively little is known about the life history of these species. Research in Hawaii and elsewhere in the tropical Pacific suggests that most of the species are slow growing, long lived, and have a relatively high age at sexual maturity. Stock assessment is difficult because of the multispecies nature of the fishery. However, recent analysis of commercial fishery data indicates that some of the species may currently be overexploited. Research is underway to determine the efficacy of management measures such as minimum-size limit changes or seasonal and spatial fishery closures to maintain optimal spawning biomass.

Population Biology and Life History of the North American Menhadens, Brevoortia spp.

Four recognized species of menhaden, Brevoortia spp., occur in North American marine waters: Atlantic menhaden, B. tyrannus; Gulf menhaden, B. patronus; yellowfin menhaden. B. smithi; and finescale menhaden, B. gunteri. Three of the menhaden species are known to form two hybrid types. Members of the genus range from coastal waters of Veracruz, Mex., to Nova Scotia, Can. Atlantic and Gulf menhaden are extremely abundant within their respective ranges and support extensive purse-seine reduction (to fish meal and oil) fisheries. All menhaden species are estuarine dependent through late larval and juvenile stages. Depending on species and location within the range, spawning may occur within bays and sounds to a substantial distance offshore. Menhaden are considered to be filter-feeding, planktivorous omnivores as juveniles and adults. Menhaden eggs, immature developmental stages, and adults are potential prey for a large and diverse number of predators. North American menhadens, including two hybrids, are hosts for the parasitic isopod, Olencira praegustator, and the parasitic copepod, Lemaeenicus radiatus. Although the data are quite variable, a dome-shaped Ricker function is frequently used to describe the spawner-recruitment relationship for Atlantic and Gulf menhaden. Each of these species is treated as a single stock with respect to exploitation by the purse-seine reduction fishery. Estimates of instantaneous natural (other) mortality rates are O.45 for Atlantic menhaden and 1.1 for Gulf menhaden.

Biology and Potential Use of Pacific Grenadier, Coryphaenoides acrolepis, off California

Grenadiers (family Macrouridae) are the most abundant fish on most continental slope areas worldwide. Off California the Pacific grenadier, Coryphaenoides acrolepis, occurs in relatively large numbers and may have marketing potential. This repon provides information on the biology of the species and catch results from a number of scientific cruises. Catch data on several other species found together with Pacific grenadier, panicularly sablefish, Anoplopoma fimbria, are also given. The fish were caught with a bottom trawl (15 trawls), and with free-vehicle longline gear (117 sets). The latter was a hook and line system in which the gear was dropped to the seafloor untethered to the fishing vessel, and floated to the surface, with the catch, when detachable weights were automatically released. Sablefish dominated longline catches in depths of 200-600fm (334-1,098m), while Pacific grenadier was most abundant between 600 and 1,OOOfm (1,098-1,830m). Best trawl catches of Pacific grenadier were made at depths between 615 and 675fm (1,125 and 1,235 m) and at 760fm (1,391 m). Ripe females were absent from our samples, but spent females were found during the entire year with highest numbers in the spring and early summer. Only one larva was found despite extensive sampling with plankton nets. Pacific grenadier was found to have good edible qualities by a taste-test panel, although the protein content (15 percent) and flesh yield (24 percent) were significantly lower than those of other fishes. A second species, the giant grenadier, Albatrossia pectoralis, was found to have exceptionally poor eating qualities and even lower protein content.