Distribution, Abundance, and Biological Characteristics of Groundfish off the Coast of Washington, Oregon, and California, 1977-1986

We compare results of bottom trawl surveys off Washington, Oregon, and California in 1977, 1980, 1983, and 1986 to discern trends in population abundance, distribution, and biology. Catch per unit of effort, area-swept biomass estimates, and age and length compositions for 12 commercially important west coast groundfishes are presented to illustrate trends over the lO-year period. We discuss the precision, accuracy, and statistical significance of observed trends in abundance estimates. The influence of water temperature on the distribution of groundfishes is also briefly examined. Abundance estimates of canary rockfish, Sebastes pinniger, and yellowtail rockfish, S. Jlavidus, declined during the study period; greater declines were observed in Pacific ocean perch, S. alutus, lingcod, Ophiodon elongatus, and arrowtooth flounder, Atheresthes stomias. Biomass estimates of Pacific hake, Merluccius productus, and English, rex, and Dover soles (Pleuronectes vetulus, Errex zachirus, and Microstomus pacificus) increased, while bocaccio, S. paucispinis, and chilipepper, S. goodei, were stable. Sablefish, Anoplopoma fimbria, biomass estimates increased markedly from 1977 to 1980 and declined moderately thereafter. Precision was lowest for rockfishes, lingcod, and sablefish; it was highest for flatfishes because they were uniformly distributed. The accuracy of survey estimates could be gauged only for yellowtail and canary rockfish and sablefish. All fishery-based analyses produced much larger estimates of abundance than bottom trawl surveys-indicative of the true catchability of survey trawls. Population trends from all analyses compared well except in canary rockfish, the species that presents the greatest challenge to obtaining reasonable precision and one that casts doubts on the usefulness of bottom trawl surveys for estimating its abundance. (PDF file contains 78 pages.)

Biological characteristics and mortality of western butterfish (Pentapodus vitta), an abundant bycatch species of prawn trawling and recreational fishing in a large subtropical embayment

The western butterfish (Pentapodus vitta) is numerous in the bycatch of prawn trawling and recreational fishing in Shark Bay, Western Australia. We have thus determined crucial aspects of its biological characteristics and the potential impact of fishing on its abundance within this large subtropical marine embayment. Although both sexes attained a maximum age of 8 years, males grow more rapidly and to a larger size. Maturity is attained at the end of the first year of life and spawning occurs between October and January. The use of a Bayesian approach to combine independent estimates for total mortality, Z, and natural mortality, M, yielded slightly higher point estimates for Z than M. This result indicates that P. vitta is lightly impacted by fishing. It is relevant that, potentially, the individuals can spawn twice before recruitment into the fishery and that 73% of recreationally caught individuals are returned live to the water.

Biological Characteristics and Fishery Assessment of Alaska Plaice, Pleuronectes quadrituberculatus, in the Eastern Bering Sea

Alaska plaice, Pleuronectes quadrituberculatus, is one of the major flatfishes in the eastern Bering Sea ecosystem and is most highly concentrated in the shallow continental shelf of the eastern Bering Sea. Annual commercial catches have ranged from less than 1,000 metric tons (t) in 1963 to 62,000 t in 1988. Alaska plaice is a relatively large flatfish averaging about 32 cm in length and 390 g in weight in commercial catches. They are distributed from nearshore waters to a depth of about 100 m in the eastern Bering Sea during summer, but move to deeper continental shelf waters in winter to escape sea ice and cold water temperatures. Being a long-lived species (>30 years), they have a relatively low natural mortality rate estimated at 0.20. Maturing at about age 7, Alaska plaice spawn from April through June on hard sandy substrates of the shelf region, primarily around the 100 m isobath. Prey items primarily include polychaetes and other marine worms. In comparison with other flatfish, Alaska plaice and rock sole, Pleuronectes bilineatus, have similar diets but different habitat preferences with separate areas of peak population density which may minimize interspecific competition. Yellowfin sole, Pleuronectes asper, while sharing similar habitat, differs from these two species because of the variety of prey items in its diet. Competition for food resources among the three species appears to be low. The resource has experienced light exploitation since 1963 and is currently in good condition. Based on the results of demersal trawl surveys and age-structured analyses, the exploitable biomass increased from 1971 through the mid-1980’s before decreasing to the 1997 level of 500,000 t. The recommended 1998 harvest level, Allowable Biological Catch, was calculated from the Baranov catch equation based on the FMSY harvest level and the projected 1997 biomass, resulting in a commercial harvest of 69,000 t, or about 16% of the estimated exploitable biomass.

Yellowfin Sole, Pleuronectes asper, of the Eastern Bering Sea: Biological Characteristics, History of Exploitation, and Management

Yellowfin sole, Pleuronectes asper, is the second most abundant flatfish in the North Pacific Ocean and is most highly concentrated in the eastern Bering Sea. It has been a target species in the eastern Bering Sea since the mid-1950's, initially by foreign distant-water fisheries but more recently by U.S. fisheries. Annual commercial catches since 1959 have ranged from 42,000 to 554,000 metric tons (t). Yellowfin sole is a relatively small flatfish averaging about 26 cm in length and 200 g in weight in commercial catches. It is distributed from nearshore waters to depths of about 100 m in the eastern Bering Sea in summer, but moves to deeper water in winter to escape sea ice. Yellowfin sole is a benthopelagic feeder. It is a longlived species (>20 years) with a correspondingly low natural mortality rate estimated at 0.12. After being overexploited during the early years of the fishery and suffering a substantial decline in stock abundance, the resource has recovered and is currently in excellent condition. The biomass during the 1980's may have been as high as, if not higher than, that at the beginning of the fishery. Based on results of demersal trawl surveys and two age structured models, the current exploitable biomass has been estimated to range between 1.9 and 2.6 million t. Appropriate harvest strategies were investigated under a range of possible recruitment levels. The recommended harvest level was calculated by multiplying the yield derived from the FOI harvest level (161 g at F = 0.14) hy an average recruitment value resulting in a commercial harvest of 276,900 t, or about 14% of the estimated exploitable biomass.

Biological characteristics and determination of population dynamic parameters of Sphyraena jello in the coastal waters of Province Bushehr- Persian Gulf

Sphyraena jello, Pick handle Barracuda, is amongst the highly valuable and main commercial fisheries resources in the southern waters of Iran. Given such an economically significant position, this study, being conducted in 2006-2007, attempts to investigate its biological habit and characteristics in Iran’s water of the Persian Gulf. For the sampling purpose, three major landings namely Bushehr, Deylam and Genaveh were selected to obtain samples from commercial catches. The sampling is composed of 655 males and 515 females during a twelve month period. By studying the feeding through the counting method, it is revealed that, Liza subviridis characterized by %42.8 and Sepia pharaonis by %8.4 made the highest and lowest stomach content respectively. The findings showed that male fish in smaller size will mature sooner than females’ specimen but this gender ratio or proportion was not significantly different except during October and September. Such a difference between male and female in different months could be originated from longer residing of female group in spawning ground compared to male group. The previous spawning lasted during September –October, and there was a peak of spawning in feeding in August. The lowest fat proportion for both male and female genders was reported 0.10 and 0.11 respectively in October; but the highest level of condition factor was reported to be 0.59 and 0.63 during November and June.

Biological characteristics and determination of population dynamics parameters of Nemipterus japonicus in the coastal waters of Province Bushehr, Persian Gulf

This study was conducted to determine biological characteristics and population dynamics parameters of threadfin bream (Nemipterus japonicus) in Persian Gulf (Bushehr Province), during November 2006 and October 2007. The minimum and maximum specimens were 75-273 mm FL and their weight was 7.6 - 351.9 g. Based on the exponential relationship between fork length and weight, slope (b) for individuals, males and females was 2.987321, 2.992546 and 3.007314, respectively. The emptiness value (V) was 45.6% and it shows that N. japonicus is a moderate feeder. The results of Fp indicates that crustacean with 78.2% are main foods, mollusca (27.7%), fishes (20.7%), polychaeta (19.2%) and Foraminifera (11.7%) were identified as minor foods and phytoplanktons (9.9%), nematoda(8.0%), echinodermata (2.3%) and sea weeds (0.3%) were random foods. The reproduction studies showed the spawning season extended within 2 peaks, from April- May and September and main spawning occurs in spring season.The mean absolute and relative fecundities were 472388±42633 and 3817±293 (X±SE), respectively. The maximum, minimum and mean of oocyte diameter were 0.448, 0.022 and 0.221mm (SE=0.071), respectively.The fork length at 50% maturity estimated to be 20.25 cm for females. The growth coefficient (K) , length infinity (L∞ ) and ɸ' was estimated 0.42/yr , 34.17 cm and 2.69, respectively. The coefficient of total mortality, fishing mortality, natural mortality and E was 1.37, 0.43, 0.94 and 0.31, respectively.

The study of biological characteristics and heavy metal determination of Cd, Pb, Fe, V, Ni in the muscle, liver and gonad tissues of kingfish Scomberomorus commerson in the coastal waters of the Hormozagan Province (Persian Gulf)

Biological studies and heavy metal (Ni, v, Fe, Pb,Cd) determination in liver, gonad and muscle of Scomberomorus commerson were carried out from Oct 2006 to Sept 2007 in Hormozgan coastal waters. 599 Samples were gutted for reproduction and nutrition studies, fork length and weight were measured to nearest cm and g respectively. Meanwhile 40 samples were also investigated for heavy metal studies. All specimens were collected from two major landing sites (Bandar Lengeh & Bandar Abbas). Minimums & maximum fork length & weight were 29, 128 cm & 235 and 15350 g respectively.Isometric growth was shown according to our study and b was estimated 2.9 (overall), 2.91(male) & 2.89 (female). The average relative gut length was 0.52± 0.007 and it was determined that S. commerson is a carnivorous. More than 99 percent of gut content was different teleost fishes. Gastro somatic index had two peaks in Nov & Jan (before spawning) and with a decreased trend in July, the spawning period. Occurrence of empty stomach was estimated % 65.77. Maximum amount of condition factor was in Dec. Spawning season was started from June. The average of Absolute & relative fecundity (to weight unit) was 1217149±179315 and 178.2±15.58 respectively. Lm50% was estimated 75 cm for females. Sex ratio was 0.97: 1 (male: female). Chi- Square test showed no significant difference (p>0.05). Maximum amount of hepatosomatic index was estimated in March.Metal concentrations were determined using either Flame Atomic Absorption Spectroscopy (for Fe) or Graphite Furnace Atomic Absorption Spectroscopy (for Pb,Cd,Ni and V). The mean concentration (μg/g dry weight)of Pb,Cd,Ni,V and Fe in the liver were 0.0309, 0.0268, 0.0672, 0.0077, 2.5159 in the gonad 0.0440 ,0.0295, 0.1096, 0.0000, 1.4449 and in the muscle 0.0244, 0.0324, 0.0656, 0.0128, 1.6138 respectively. The maximum metal concentrations were below the maximum permissible limits for human consumption recommended by the USEPA, WHO and the UK. The results of Kendall's Tau-b correlation coefficient were as follows: The Liver tissue: There were significant positive linear relationships between accumulation of V, Fe, and Pb with Fork length, Pb and Fe with weight, GSI with Pb, Cd, V and 109 Fe, and a negative linear relationships between HSI with accumulation of V and Fe, Fork length, weight and GSI. The Gonad tissue: There were significant positive linear relationships between GSI with accumulation of Pb, Cd, Fe, Fork length and weight, a negative linear relationship between HSI with Fork length, weight and GSI. The Muscle tissue: There were significant positive linear relationships between accumulations of V, Fe with Fork length and weight factors and as well as GSI with Cd, V, Fe, Pb, Fork length and weight,a negative linear relationship between HSI with Fork length, weight, Cd, Fe and GSI. The results of Mann-Whitney U tests (P≤0.05) show that there were significant differences between summer and autumn from heavy metal contents in the studied tissues point of view. The only exceptions were for Ni in the liver, gonads and muscle and as well as there were significant differences between male and female from heavy metal contents in the studied tissues. The only exceptions were for Pb in muscle, Ni in liver, gonad and muscle, V in muscle, and Cd and Fe in gonads.

The study of biological characteristics and determination of population dynamic parameters of sawtooth barracuda (Sphyraena putnamae) in the Persian Gulf (Hormozgun Province waters)

The biological characteristics and population dynamisms of Sphyraena putnamae, were studied in the northern Persian Gulf and Oman Sea restricted to Hormuzgan province waters within 13 months period, from November 2006 up to November 2007. Biometrical and anatomical measurements were carried out, and biological surveys were conducted on 486 specimens. On the other hand, the growth and mortality parameters were estimated by using 3096 samples. These samples were collected from 3 landings, namely Bandar Abbas, Bandar Lengeh and Bandar Jask. The measurements of the minimum and maximum Fork lengths and weights were 11.7 to 8.03 cm and 135.0 to 4140.0 g, respectively. The results indicated that this species, having the Relative Length of Gut, RLG=0.34±0.002, is strongly carnivorous (often fish-eater), proven by the fact that more than 98% of its stomach contents were fish pieces. Examining the changes in the index of stomach emptiness by the percentage of CV = 0.47% indicates that this fish is Moderate feeder. The level of feeding increased in March, before spawning and decreased in June and September, simultaneously with the spawning season. There are 2 peaks of reproduction or spawning seasons during the months of April-May and September, of which the prior is assumed as the main spawning. The sex ratio (M:F) was calculated 0.5:1.0(X2 =2.11), which did not show a significant difference with expected level of 1:1 (P>0.05). The average absolute and relative reproduction rates of Sphyraena putnamae is respectively as follows: 1866827.1±255448.9 and 1097.7±94.3. The highest and the lowest diameter of matured egg are from 200 to 750 μ, and its average diameter is 402.10 ± 0.190 μ. A parameter for Saw-tooth barracuda length measurement, Lm50, based on the Fork-length, was calculated as 54.01 cm. In other words, as far as the fisheries management is concerned, the fish whose lengths are less than 54.01 cm should not be caught. The calculated level of (R2) (correlations of total length & weight), indicated strong correlations between length and weight of this fish, and the obtained formula included W =0.007100 FL 2.9295 and reinforced this assumption. The “K” Index for this fish in 3 above mentioned landings (Jask, Bandar-Abbas and Bandar-Length) were 1.24, 0.37 and 0.46 per year, respectively and the FL index for the same landings were estimated as 129, 110 and 134 cm, respectively. The growth coefficient (MONRO) for the above mentioned regions were calculated as 3.601, 3.647 and 3.917, respectively; and in the surveyed regions there were no significant differences in populations. The Total mortality coefficient (Z) was calculated 0.76, 1.12 and 1.07 per year, the Natural mortality coefficient was 0.46, 0.63 and 0.70, and the Fishing mortality coefficient (rate) (F) was found to be 0.30, 0.49 and 0.37 per year. The value of the exploitation rate (E) is equal to 0.39 per year, indicating that this species is an under-exploited resource, and there is no excessive fishing pressure on the fish supply of this species in the afore-said regions. The highest level of exploitation was found for ‘Bandar Abbas’ fishing region and the lowest level of exploitation is in ‘Bandar Lengeh’ waters.

Determination of biological characteristics and blood groups of hilsa shad (Tenualosa ilisha) in the north of Persian Gulf (Bushehr coastal waters)

Hilsa shad, Tenuolosa ilisha, belongs to Clupeidae family and Alsinae subfamily is an euryhaline pelagic and anadromous species living in marine and freshwater waters. Regarding to study the biological characteristics of this species, this study was carried out in the Northern Persian Gulf within Bushehr province waters during years 2006-7. A total of 344 specimens were collected and transported to the laboratory for further different biological measurements consist of: reproduction (GSI, fecundity, maturity stages), feeding (stomach contents, food preference ...), aging; 58 morphometric and ٧ meristic measurements. The results indicate that minimum, maximum and mean body weight are 203, 953 and 481.35±147.64 g, respectively and this values for total length are 26.5, 45.1 and 35.99±3.98 cm. For food regimes, the most abundant phytoplankton foods belong to Bacillariophyta (83.74٪) and zooplanktons of Arthropoda (51٪). The mean RLG was measured about 1.53±0.22 and the mean feeding intensity was about 51.79±38.13. Also, the aging of studied fishes showed that they have 2 to 4 years and most of the samples had 2+ years. The reproduction studies showed that Indian shad is a batch spawner species. The average gonadosomatic index (GSI) is 1.64±1.382 and the main spawning season was determined for April-May.

Study on biological characteristics, sexual maturation and reproduction of Liza abu in the water of Khozestan Province

As the most of the fish resources are known and exploited, protecting their generation is of the greatest importance. Aquaculture is one of the efficient procedures in protecting and reviving fish resources and knowing about the reproductive cycle and gonads development has an important role in approaching this aim. Liza abu belongs to the family Mugilidae that according to its resistance to the environmental condition and its fast growth , can be introduced as a fish with economical value. As there is no scientific data on the reproductive biology of this species , study on the reproductive biology and gonad development is considered as the aim of this research . For this purpose , 360 samples of this species were investigated during the period from February 2007 to January 2008 in Khozestan Province . After studing morphological and histological characteristics of gonad specimen , they were prepared through histological method. Samples were prepared through usual histological method and studied under light microscope. According to the results, the maturity stages of male and female Liza abu were separated to six different successive stages. In ovaries , these stages were as follow : In stage І, the oocytes were small , this stage was observed from July to October . In stage ІІ, considerable growth was observed in the oocytes . This stage was observed from October to January . In stage III, due to vitellogenesis, the maximum growth was observed and three layers of theca, granullosa and follicle cells were visible. This stage was observed during January and February . In stage IV, migration of germinal vesicle was observed and due to hydration of the oocytes , their diameter was increased. The ovaries were yellowish and in maximum size and ovules could be easily observed with naked-eye . This stage was observed in February and March . In stage V, spawning occured. This stage was observed in April . In stage VI, ovaries consisted of immature and atretic oocytes and also empty follicles. This stage was observed in May and June. In testes , these stages were as follow : In stage I , the testes were small in size and contained the spermatogonia which were the only cellular components.This stage was observed in August and September . In stage II (maturing virgin ) , the spermatogonia and the primary spermatocytes were visible. This stage was observed in October . In stage III (developing), intensive spermatogenesis was occured and the primary and the secondary spermatocytes were the most visible cells during this stage .This stage was observed from November to January. In stage IV(developed), cells of all stages of spermatogenesis could be seen but the secondary spermatocytes and spermatids were in large number. This stage was observed from January to March. In stage V , the testes were filled with sperms. This stage was observed in March and April .In stage VI, residual spermatozoa and the spermatogonia were visible in the testes. This stage was observed from May to August. According to cyclic changes in GSI, sexual maturation in breeding begins in January and spawning occurs in April. The ova diameter ranged from 30.75 μ in stage I to 472.19 μ in stage IV. In this study , the sex ratio was 1:2.7, and male and female percentage were 27.02% and 72.98% respectively. This means that females predominate males. In this study absolute fecundity was calculated and changing between 30805.44 to 431247.3 was observed and absolute fecundity was calculated 111275.3 in average.

Study biological characteristics related with development and reproduction in Barbus capito on the southern coast of the Caspian Sea, Gilan Province

In the present research, a total of 207 pieces of fish from 25 sampling stations in Gilan Province coasts in the years 2001-2002 were biologically studied in terms of their growth and development, reproduction and feeding. The average length and weight of the fishes are increased, as they get older. The highest index of length and weight growth is observed in the years 1 to 2. As the age increases, gradient of length and weight growth diagrams decrease. In studying the relation between length and weight, it was observed that proportionate to the total length, the weight is increased progressively. The fatness coefficient index in the initial years of life and prior to maturity is higher than the post maturity period. As the age increases, the decrease of this index is observable. The fatness coefficient index rate is directly related to index of fullness. The highest Gonadosomatic Index is seen in the months of June and July, i.e. at the times of spawning; and the lowest index rate is observed in the months of November and December. The appropriate temperature for reproduction of these species is from 18 to 22 degree centigrade. The Gonadosomatic Index is higher in spring and summer seasons as compared with autumn and winter. Besides, as the fishes become aged, the amount of the said index increases in a manner that the gradient of it in the years to maturity is less than the maturity time and thereafter. Sexual maturity stages in different months are directly related to Gonadosomatic index, and increase as the age increases. The sexual ratio of male fishes to the female fishes in terms of number is plus one prior to maturity; about one at the time of maturity and minus after maturity. In general the frequency of male fishes as compared with female fishes in all group ages is approximately two times. The fecundity mean, and the diameter and the rate of eggs will substantially increase, as the Gonadosomatic index rises. The maturity age in the male fishes is 3 to 4 years and in female fishes is 4 to 5 years. The spawning of this species in rivers occurs repeatedly and in different time intervals, and do not take place once (Asyncronous). The Gastrosomatic index is directly related to index of fullness and will decrease, as the age increases. The index of fullness is relatively the months of April and May. The underlying reason is the need of the fishes to energy for reproduction. As the spawning time commences, the index of fullness moves down and the downward direction continues. After spa g mg and reduction of the volume of energy in the body, the index of fullness rises, and it will be substantially high until the beginning of fall. In fall and winter as it gets cold, the index of fullness moves downward and the body fat deposits are used. A correlation is shown between the changes in vacuity index and fullness indices. This means that as the fullness index rises, the vacuity index decreases, and vice versa. The Hepatosomatic index prior to the reproduction is at the highest amount and after spawning is at the lowest. No correlation is observed between the fullness and Hepatosomatic indices. In other words reproduction is an inherent and instinct originated matter; and its cycle goes on, alternately and in an orderly manner, upon completion of germinal cells, even when it coincides with reduction or stoppage of somatic cell growth. The rising trend of Hepatosomatic starts in August and will continue until the next July. The volume of fat around digestive tract is severely reduced in early spring and this trend will reach its apex in summer season. In the cold seasons, i.e. the fall and winter, the accumulation of fat around digestive tract increases. Consequently, a meaningful and inverse relation is observed between index of fullness, also the progress of sexual maturity stages and the volume of fat.

Environmental and biological characteristics of lagoons on the southwest coast of Ireland

Coastal lagoons are defined as shallow coastal water bodies partially separated from the adjacent sea by a restrictive barrier. Coastal lagoons are protected under Annex I of the European Habitats Directive (92/43/EEC). Lagoons are also considered to be “transitional water bodies” and are therefore included in the “register of protected areas” under the Water Framework Directive (2000/60/EC). Consequently, EU member states are required to establish monitoring plans and to regularly report on lagoon condition and conservation status. Irish lagoons are considered relatively rare and unusual because of their North Atlantic, macrotidal location on high energy coastlines and have received little attention. This work aimed to assess the physicochemical and ecological status of three lagoons, Cuskinny, Farranamanagh and Toormore, on the southwest coast of Ireland. Baseline salinity, nutrient and biological conditions were determined in order to provide reference conditions to detect perturbations, and to inform future maintenance of ecosystem health. Accumulation of organic matter is an increasing pressure in coastal lagoon habitats worldwide, often compounding existing eutrophication problems. This research also aimed to investigate the in situ decomposition process in a lagoon habitat together with exploring the associated invertebrate assemblages. Re-classification of the lagoons, under the guidelines of the Venice system for the classifications of marine waters according to salinity, was completed by taking spatial and temporal changes in salinity regimes into consideration. Based on the results of this study, Cuskinny, Farranamanagh and Toormore lagoons are now classified as mesohaline (5 ppt – 18 ppt), oligohaline (0.5 ppt – 5 ppt) and polyhaline (18 ppt – 30 ppt), respectively. Varying vertical, longitudinal and transverse salinity patterns were observed in the three lagoons. Strong correlations between salinity and cumulative rainfall highlighted the important role of precipitation in controlling the lagoon environment. Maximum effect of precipitation on the salinity of the lagoon was observed between four and fourteen days later depending on catchment area geology, indicating the uniqueness of each lagoon system. Seasonal nutrient patterns were evident in the lagoons. Nutrient concentrations were found to be reflective of the catchment area and the magnitude of the freshwater inflow. Assessment based on the Redfield molar ratio indicated a trend towards phosphorus, rather than nitrogen, limitation in Irish lagoons. Investigation of the decomposition process in Cuskinny Lagoon revealed that greatest biomass loss occurred in the winter season. Lowest biomass loss occurred in spring, possibly due to the high density of invertebrates feeding on the thick microbial layer rather than the decomposing litter. It has been reported that the decomposition of plant biomass is highest in the preferential distribution area of the plant species; however, no similar trend was observed in this study with the most active zones of decomposition varying spatially throughout the seasons. Macroinvertebrate analysis revealed low species diversity but high abundance, indicating the dominance of a small number of species. Invertebrate assemblages within the lagoon varied significantly from communities in the adjacent freshwater or marine environments. Although carried out in coastal lagoons on the southwest coast of Ireland, it is envisaged that the overall findings of this study have relevance throughout the entire island of Ireland and possibly to many North Atlantic coastal lagoon ecosystems elsewhere.